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Vol. 7, No. 1

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•OTANiCAL GARDEN

MICHIGAN BOTANIST

J a n u a ry, 1 968

THE MICHIGAN BOTANIST— Published by the Michigan Botanical Club four
times per year: January, March, May, and October. Second-class postage
paid at Ann Arbor, Michigan.

Subscriptions: $3.00 per year. Single copies: $.75.

A11 back issues are available, at the following prices per volume:

Vol. 1 (2 numbers, at $1.00 each): $2.00
Vol. 2-5 (4 numbers, at $.50 each): $2.00
Vol. 6- (4 numbers, at $.75 each): $3.00

Subscriptions (from those not members of the Michigan Botanical Club) and
all orders for back issues should be addressed to the business and circulation
manager, who will assume that subscriptions received prior to the last number
of a volume are intended to begin with the first number of that volume unless
specified to the contrary.

Articles and notes dealing with any phase of botany in the Upper Great
Lakes Region may be sent to the editor in chief. The attention of authors pre¬
paring manuscripts is called to “Information for Authors” (Vol. 6, p. 202; re¬
prints available from the editor).

Editorial Board

Edward G. Voss, Editor in chief

Herbarium, The University of Michigan, Ann Arbor, Michigan 48104.

Laura T. Roberts, Business and Circulation Manager
1509 Kearney Rd. , Ann Arbor, Michigan 48104.

Alexander H. Smith
Warren P. Stoutamire
W. H. Wagner, Jr.

Charles Barclay
Rogers McVaugh
C. Marvin Rogers

THE MICHIGAN BOTANICAL CLUB

Membership in the Michigan Botanical Club is open to anyone interested in
its aims: conservation of all native plants; education of the public to appreciate
and preserve plant life; sponsorship of research and publication on the plant
life of the State; sponsorship of legislation to promote the preservation of
Michigan native flora and to establish suitable sanctuaries and natural areas;
and cooperation in programs concerned with the wise use and conservation of
all natural resources and scenic features.

Dues are modest, but vary slightly among the chapters. In all cases, they
include subscription to THE MICHIGAN BOTANIST.

President: W. H. Wagner, Jr., Botanical Gardens, University of Michigan, Ann
Arbor, Michigan 48105

Vice President: William F. Hopkins, 6210 Cowell Rd., Brighton, Michigan 48116
Recording Secretary: Mary Cooley, 703 S. Forest, Ann Arbor, Michigan 48104
Corresponding Sec.: Margaret Buswell, 19204 Plainview, Detroit, Mich. 48219
Treasurer: Charles Buswell, 19204 Plainview, Detroit, Michigan 48219
Corresponding Secretary, Southeastern Chapter:

Catherine Hamilton, 9591 Grandmont, Detroit, Michigan 48227
Secretary, Huron Valley Chapter:

Ursula R. Freimarck, 704 Madison PL, Ann Arbor, Michigan 48103

1968

THE MICHIGAN BOTANIST

3

RARE OLD PUBLICATIONS IN MICHIGAN HERBARIA

Rogers McVaugh

University of Michigan Herbarium, Ann Arbor

The term “ herbarium’ ’ today usually brings to mind a collection
of pressed and dried plant specimens, sometimes the property of an
individual but usually belonging to a museum, a college or university,
or some other public institution. Herbarium specimens of vascular
plants are now usually mounted on sheets of paper of a standard size,
and filed individually, but in some desired order, in wooden or metal
cases where they may be available for study and research.

The original herbarium, or “Hortus Siccus, ” however, as it began
to develop in Europe in the 16th and 17th Centuries, was a much less
pretentious affair. Almost every botanist had his own herbarium,
consisting primarily of plants that he himself had collected locally,
with the addition of such duplicates as he could obtain from his friends
and correspondents. These personal herbaria were smaller than most
institutional herbaria today, and many of them differed from modern
collections in one important respect: The mounted specimens, glued
or otherwise affixed to paper, were assembled into books of a conven¬
ient size, often bound in leather or other hard covers and shelved like
any other book. Some of these early herbaria, beautifully and sumptu¬
ously bound, are today among the botanical treasures of the world’s
great museums. The practice of binding herbaria into books, however,
did not recommend itself to botanists who wanted to study particular
groups of plants. The specimens in the bound volumes were not inter¬
changeable, which meant that anyone who wanted to study all the willows
(for example) in several different herbaria, first had to locate all the
specimens in each individual volume, then assemble the volumes in
some way to compare them. The system, in other words, was neither
open-ended nor convenient, and the custom of preserving herbaria in
books fell into disuse about the beginning of the 19th Century, except
as students and amateur botanists began their collections in this way. 1

Interestingly enough, however, there is a lineal descendant of the
bound herbarium-volume which has persisted through the 19th Century
and well into our own. This is the so-called flora exsiccata , a set of
dried plants of one particular region or one particular plant group,
distributed as a unit. Such [plantae] exsiccatae, or dried plants, were

lone of the most interesting such collections now in the Herbarium of the
University of Michigan is the seven-volume set comprising the original herbar¬
ium of Douglass Houghton, the first State Geologist of Michigan. Most of the
specimens are from the vicinity of Troy, New York, collected in 1829 when
Houghton was 20 years old and a student of Amos Eaton’s at the Rensselaer
Institute; some were collected the following year, when Houghton was an assistant
professor at the same place, and a few during 1831, when he was surgeon and
botanist on the first Schoolcraft expedition to the sources of the Mississippi River.

4

THE MICHIGAN BOTANIST

Vol. 7

often collected in many duplicate sets, and sold or exchanged by the
collectors. In recent years the Plantae Exsiccatae Grayanae were sent
out in about 100 equivalent sets of 100 specimens at a time, each speci¬
men accompanied by a printed label bearing information about the bib¬
liographic history of the species, together with full data relative to the
place of collection and the habitat, and pertinent information about the
plant itself. The more recent sets of exsiccatae have customarily been
sent to the recipients in the form of individual herbarium specimens,
but a number of the earlier ones were distributed as units, and were
intended as such; sometimes they were bound by the senders into books,
sometimes not; often they were accompanied by printed title page and
index, sometimes by a few paragraphs or pages of explanatory text, and
(most important to taxonomists today) the printed labels with the speci¬
mens sometimes included descriptions of new varieties or species.

The International Code of Botanical Nomenclature now specifies
that anyone may describe and name a plant that has previously been
unknown to science, providing that he complies with certain regulations
in the Code, but providing especially that he distribute to the botanical
public a printed description of the plant he intends to name as new.
Names may not be validated by announcing them at a public meeting,
or by sending out handwritten or typed descriptions— only by distribu¬
tion of printed matter. The reader will see at once where this is
leading: Most printed matter today (and in the 19th Century as well),
was distributed in the form of books, pamphlets, and periodical litera¬
ture. As such, the material was customarily given a place on the
shelves of some library, and thus remained permanently available for
consultation. But why should not one of the bound sets of exsiccatae
be considered a book? It has a title page, date, author, printed text,
and index; it has been widely distributed to the botanical public. It
differs from a conventional book only in having dried plant specimens
pressed between the pages and forming an integral part of the volume
as a whole. Why then does the Code of Nomenclature specifically pro¬
vide that “on or after 1 Jan. 1953” the publication of new names may
not be effected by the distribution of such exsiccatae ?

The answer is not hard to find. These collections of dried plants
are not really equivalent to books, even though they may be bound in
book form. At the time of distribution they are hardly ever repre¬
sented by more than 100 copies, and usually by fewer than this. This
means that even at first they do not reach a very wide botanical audi¬
ence. By the time they are a century old, most of the copies will
have been broken up and distributed as individual specimens through
herbaria; or some or all of the specimens lost from the pages, or
destroyed. Of the three publications discussed below, none is known
to be represented now by more than two or three unbroken sets. Fin¬
ally it may be said that botanists have never consistently classed
exsiccatae as literary contributions, but rather as collections of speci¬
mens accompanied by printed matter. Thus for the most part exsic-

1968

THE MICHIGAN BOTANIST

5

catae are not listed in published bibliographies, library catalogues,
and the like, and many botanists are hardly aware of their existence.
The Code rightly discourages the practice of publishing new names for
plants in this way, just as it condemns publication in ephemeral media
such as daily newspapers and seed catalogues. On the other hand, it
is necessary to take into account the fact that in the past many new
names were first put before the public by the distribution of printed
matter accompanying just such sets of dried plants. To invalidate all
these names now would mean that some well-known and long-accepted
plant names would have to be changed, and for this reason the older
names published in this way are still considered validly and effec¬
tively published. Two such names well known to Michigan botanists
are Panicum xanthophysum and Calamagrostis inexpansa , both pub¬
lished by Asa Gray in the work described below.

Because of the great rarity of most of these sets of dried plants,
and because botanical activity in Michigan has been of recent develop¬
ment as compared to that in the great botanical centers of Europe and
even in the eastern part of the United States, it is interesting to be
able to report here upon the existence in Michigan herbaria of three
complete exsiccatae , dating respectively from 1805-1814, 1826-1827,
and 1834-1835.

1. NORTH AMERICAN GRAMINEAE AND CYPERACEAE,

by Asa Gray

Some years ago Rickett and Gilly2 reported upon this, Asa Gray’s
second botanical publication. They noted that a “ bound copy of Part I
is preserved in the library of the New York Botanical Garden . . . and
a bound copy is in the herbarium of the Philadelphia Academy of Na¬
tural Sciences.” They seem to have seen a bound copy of Part II only
in the Philadelphia Academy. They noted that some additional speci¬
mens, from Part I as well as from Part II, are scattered through the
herbarium of the New York Botanical Garden, and that a similar “ dis¬
persed set is in the Gray Herbarium.” They commented that the emi¬
nent agrostologist Trinius “had access to a set of this work and was
one of the first to use the names published there” [i.e. about 1841].
With respect to additional sets of Gray’s exsiccatae , Rickett and Gilly
said merely “Other bound or dispersed sets are undoubtedly in exist¬
ence.”

It was therefore a matter of unusual interest that a complete set
was found in the herbarium of Wayne State University (WUD3), where
it forms part of a collection of ten volumes formerly the property of
H. P. Sartwell (1792-1867) of Penn Yan, New York. Sartwell was one

2Rickett, H. W., and C. L. Gilly. Asa Gray’s earliest botanical publications
(1833-1836). Bull. Torrey Bot. Club 69:461-470. 1942.

3 WUD is the internationally accepted abbreviation for the herbarium of
Wayne State University, as listed in Index Herbariorum, 5th edition, p. 56 (1964).

6

THE MICHIGAN BOTANIST

Vol. 7

of Gray’s correspondents, and indeed collected several sets of speci¬
mens that were distributed in 1834 and 1835 among the other Gramineae
and Cyperaceae. There is evidence among the other specimens in
Sartwell’s herbarium that he corresponded with Dennis Cooley, who
from 1827 until his death in 18 60 lived in Macomb County, Michigan,
and there are a few specimens indicating that Sartwell himself col¬
lected near Detroit and Windsor; but I have not been able to establish
any other connection between Sartwell and the State (or Territory) of
Michigan. Dr. C. M. Rogers, through whose kindness I have been able
to examine the collections, informs me that there is apparently no
record of how this part of Sartwell’s herbarium came to Wayne, pre¬
sumably long after his death.

50. Calamagrostis inexpansa ( sp . nov .) panicula oblonga laxiuscu-
la, floribus approximate appressis ; glumis subequalibus
oblongn-lanceolatis acutissimis, inferiore perianthio j
longioribus ; valvula inferiori superiore J majori, seta
(recta) supra basin exserta vix tnqtiante ; pilis glumas
subdupio brevioribus ; processu barbato.

Hab. Pen Yan, New-York. JDr« H. P. Sartwell.

j.

28. Panicum xanthophysum (sp. nov.) culmo (simplici aut ba-
si ramoso) panicula foliisque glabris; vaginis villosis, fo¬
lds lato-lanceolatis valde nervosis acutissiniis, basi cilia-
tis ; panicula simpliciusculsi paucifl ra, ramis erectis vel
adpressis ; spiculis glol)oso-t)bovatis pubescenhbus ; glu-
sna inferiore flosculis sub duplo brevit>re, superiore eos-
dem SBquante 9-nervi ; flore masculo bivalvi hermaph-
rodilum glabfum obovatum sub-iequante.

Ob8. — P. latifolio aflini8.

Hab. — Pine Plains, near Oneida Lake, New-York.

Fig. 1. The labels bearing the original descriptions of Calamagrostis inexpansa
and Panicum xanthophysum. These seem to have been printed in vertical strips
with a margin of about one-half inch on each side, then cut apart as indicated by
the ink lines drawn at the bottom.

Fig. 2. (facing page). The title page of the Wayne State University copy of the
first volume of Gray’s North American Gramineae and Cyperaceae , bearing
Sartwell’s stamp.

1968

THE MICHIGAN BOTANIST

7

-Sci'ttofU.

NORTH AMERICAN

ORA MINE JE AND CYPERACEJE,

BY ASA GRAY, M. D.

5PJMB® I.

NEW-YORK :

PRINTED BT J. POST, 101 JOHN STREET.

8

THE MICHIGAN BOTANIST

Vol. 7

The Wayne copy of Gray’s Gramineae and Cyperaceae consists of
two bound volumes, each stamped “ H. P. Sartwell” on the title page.
The two are quite differently bound, and slightly different in size; the
pages in Part I, of a good white paper, measure about 37 x 25.5 cm,
whereas those in part II, of a thinner and slightly brownish paper,
measure about 35.5 x 26.5 cm. Each volume includes 100 specimens,
each on a separate page with its own printed label, the labels num¬
bered consecutively 1-100, and 101-200. Part I includes in addition a
title page, dedication page, a one-page introduction and explanation of
the work, and a page bearing the index to nos. 1-100. Part 2 includes
in addition to the specimens a title page identical with that of Part I
(except for the change from I to II and in the date from 1834 to 1835);
a one-page index to nos. 101-200; and a printed correction-slip per¬
taining to the names of three species in Part I.

As the paper on which the introductory pages are printed is dif¬
ferent in both parts from the paper on which the plants are mounted,
it seems that the printing, and the assembling of the specimens, were
carried on separately. Rickett and Gilly note that at least some of
the sets seem “to have been issued unbound and probably unmounted.”
These authors quote Gray as offering to have several copies of Part I
bound up himself, probably in the fall of 1834, for distribution to
Torrey and others. There is nothing in Sartwell’s copy of Part I to
show whether or not he received a bound copy from Gray, but it is
probable that his copy of Part II was presented to him in this way.
On the blank page preceding the title page is an inscription in Gray’s
hand: “Dr. H. P. Sartwell from the author”; furthermore there is a
hand-written note from Gray to Sartwell pasted on the page after the
index. Gray had acknowledged in Part I the help he had received from
Sartwell in assembling the sets for this flora exsiccata , but he still
needed additional material of some species. He therefore indicated
by pencilled notations near the labels in Part II the species that he
wanted. His note read: “Of those marked + I wish more speci¬
mens [;] those + ! I very much need.” He did not hesitate to make
his wishes known; of 100 species in Part II, 43 are marked “+” and
9 others are marked “ + !”.

It is an interesting commentary upon the herbarium practices of
the time that although Sartwell evidently kept the specimens as a unit,
and thought of them as books (in one note written 22 February 1837
he mentioned “Gray’s grassbooks 1 & 2”), he loaned individual speci¬
mens to his correspondents by the simple expedient of cutting out
pages and sending the mounted specimens, carefully re -inserting and
taping the cut pages when they were returned. In this way he sent a
number of specimens for study to Professor James Hadley of Fair-
field, New York, Gray’s former Professor of Chemistry and Materia
Medica and the person to whom Gray had dedicated the Gramineae and
Cyperaceae. Sartwell was also much interested in the genus Carex ,
and in Part II the specimens of this genus bear a number of annota-

1968

THE MICHIGAN BOTANIST

9

tions by specialists to whom he submitted them.

The specimens in the North American Gramineae and Cyperaceae
were mainly from central New York, where Gray collected between
1828 (when he was not yet 18 years old), and 1835. 4 In the published
work 71 species (29 in Part I, 42 in Part II) are labelled as from the
“ Western part of New-York, ” but presumably most of these came
from between Utica and Oneida Lake, in what would now be considered
the central part of the state. Gray’s correspondents Crawe and Sart-
well sent him specimens from a little farther west and northwest, but
still not really from western New York.

About two-thirds of the species in Part I are from New Jersey,
where Gray collected in the pine -barrens and near the Atlantic coast
in 1833. One grass in this same Part, Koehleria cristata, was col¬
lected for Gray in “Michigan territory” by a friend, Dr. N. W. Fol-
well, who lived in New York.4

In Part II about four-fifths of the species are labelled as from
western New York or simply from Utica, from near Oneida Lake, or
from Oneida County. Presumably these represented Gray’s own col¬
lections. Many of the others were credited to Sartwell, or to Dr. J. B.
Crawe of Watertown, New York, Gray’s acquaintance and one-time field
companion.

The new names and new combinations published in Parts I and II
are listed by Richett & Gilly; actually they are not very numerous.
Of interest to Michigan botanists are the following, including those of
the two species of grasses mentioned above, both described for the
first time in the North American Gramineae and Cyperaceae , Part I,
early 1834:

Calamagrostis inexpansa A. Gray, Gram. & Cyp. 20. 1834.

Panicum xanthophysum A. Gray, Gram. & Cyp. 28. 1834.

The new combination Muhlenbergia sylvatica (Torr.) Torr. in A.
Gray was validly published in the same work (p. 13), as was also the
new name Vilfa vaginiflora Torr. in A. Gray (p. 3) [ Sporobolus vagini-
florus (Torr. in A. Gray) Wood].

2. SAULES DE LA SUISSE, by N.-C. Seringe

Nicolas-Charles Seringe (1776-1858), of French birth, lived in
Switzerland for many years, and is known chiefly for his collaboration
with A. P. De Candolle in the preparation of treatments of certain plant
families for the Prodromus Systematis Naturalis Regni Vegetabilis
(1824-1830). In 1815 he published at Berne an account of the willows

4For an account of Asa Gray’s earlyyearsin central New York, his training
in medicine, and the beginnings of his interest in botany, see the first chapter
(pp. 1-18) of Asa Gray , by A. Hunter Dupree (Harvard University Press, Cam¬
bridge, 1959). N. Wright Folwell, Gray’s classmate in medicine, is mentioned
on p. 19.

10

THE MICHIGAN BOTANIST

Vol. 7

of Switzerland,5 a book of a hundred pages, and his own first publica¬
tion of any size. It is not generally known that 10 years before this
he had begun publication of an account of Swiss willows in another
form. This earlier publication, of which presumably only a very
small number of copies was issued, bears the title “Saules de la
Suisse. ” It is excessively rare today even in Switzerland; John Briquet,
the biographer of Swiss botanists, seems to have known of the first
three fascicles only. 6

In 1952 the University of Michigan bought from a European book¬
seller a set of “Saules de la Suisse” that seems to be complete in 7
separately bound fascicles (“Cahiers”). Each fascicle measures about
24 x 33 cm, and is saddle -stitched, with a tough flexible heavy paper
cover bearing a printed enumeration of the species and varieties that
are included. The whole work treats 88 consecutively numbered taxa
(most of which today would be called “forms”). At least two (nos.
39 and 40) are specimens of fungi that parasitize leaves.

One of the earlier biographers of Seringe referred to the “ Saules
de la Suisse” as “a curious work, in which dried specimens take the
place of pictures,” and this states the case fairly enough. Each fasci¬
cle includes, for every species or variety treated, one or more small
but carefully selected and prepared specimens of the plant itself.
Often different growth stages are represented by specimens collected
at different times; often flowering and fruiting specimens are both in¬
cluded; sometimes vegetative shoots with stipules, bark specimens, or
even the characteristic galls formed on certain species are shown.

In Cahiers 1-5 each specimen is mounted on a page together with
a small printed label bearing the name of the taxon and the number
assigned to it by Seringe, e.g. “5. Salix aurita. Smith.” Other infor¬
mation about the specimens, including some bibliographic references,
some descriptive matter, details of habitat, uses, etc., of the plant,
and place and date of collection, is gathered together on an introduc¬
tory page of printed text. The introductory pages are headed “ ler
CAHIER,” “ 2me CAHIER,” etc., and each is dated at bottom.

Summary of Cahiers 1

- 5

Cahier

Date

Included taxa

1

Dec 1805

1-6

2

Dec 1805

7-11

3

Dec 1805

12-18

4

Dec 1806

19-27

5

Jan 1808

28-44

In the fifth Cahier, for the first time, Seringe proposed a new
species, Salix kanderiana. It is his no. 42, described briefly and con-

5Essai d’une Monographie des Saules de la Suisse. 100 pp., 8vo. , 2 plates.
Berne, 1815.

6 Briquet, John. Bull. Soc. Bot. Suisse 50 A: 437. 1940.

1968

THE MICHIGAN BOTANIST

11

trasted with Salix acuminata. Like the others mentioned below, this
name seems to have been validly published here, several years before
it appeared in the “Essai” of 1815.

In the 6th and 7th Cahiers, the style is changed a little. The
dates (1809 and 1814, respectively) are printed on what corresponds
to a table of contents pasted to the front cover. There is no separate
introductory page in addition to the list on the front cover, but the
labels that accompany the specimens are more or less expanded to
include the kind of information that made up the preliminary text in
fascicles 1-5. Thus the label for no. 47, Salix serpillifolia Willd.,
lists French and German vernacular names, and localities at which
the plant occurs, and includes an 8-line description contrasting this
species with another. Two new species ( Salix grandifolia , no. 55, and
S. uliginosa> no. 60), are named and described in Cahier 6.

The work ends with nos. 45-60 in Cahier 6, and 61-88 in Cahier
no. 7. With the issue of this last fascicle Seringe seems to have con¬
cluded the work; evidently he was already planning his “ Essai” which
was to be published in 1815, for the last line on the “ contents” page
of no. 7 reads: “ Des remarques sur les 7 cahiers de cette collec¬
tion vont etre publiees.”

3. HORTUS SICCUS LONDINENSIS, by Mariano Lagasca

Mariano Lagasca (1776-1839) was one of the ablest botanists of his
time. Colmeiro said that he was “the leading botanist of our century,
and for many years the only one who upheld in the scientific world the
honor of Spanish botany.” Unfortunately he was one of the victims of
the Spanish counter-revolution of 1823, in which he lost most of his
possessions including his manuscripts, his books, and his herbarium.
He left Spain and emigrated to England, where he lived for a number
of years under difficult circumstances. Perhaps in an effort to sup¬
plement his income, he planned to issue a hortus siccus including the
common flowering plants and ferns growing within a radius of 20 miles
around London. Assisted by his two eldest sons, he hoped to prepare
a number of parts each containing 25 plants, and sell 30 sets of each
part. The price was fixed at one pound per part. As originally an¬
nounced, it was intended to publish one part every other month, or six
each year, but this was too ambitious an undertaking. The first set of
25 plants, with the accompanying text, seems to have been delivered in
December, 1826; the second, third, and fourth sets are all dated 1827.
From a note at the beginning of part 3, that fascicle appears to have
been published about the first of April; part 4 was promised for the
first of July. The first “volume” was completed with part 4, and no
more was ever issued. (Lagasca stated in the preface to the first
part, that “the work may be bound in volumes of four parts without
any injury to the specimens.”)

Apparently no complete copy of the Hortus Siccus Londinensis has

12

THE MICHIGAN BOTANIST

Vol. 7

survived in England. Three of the four fascicles came many years
ago into the possession of the British botanist G. Claridge Druce, who
published a note with an enumeration of all the specimens.7 The speci¬
mens themselves, with the rest of Druce’ s large herbarium, were ulti¬
mately bequeathed to Oxford University. Druce stated that as far as
he could determine there was no copy of Lagasca’s work in the Brit¬
ish Museum (Natural History). In recent years I have inquired about
it, but I have never found any indication that there may be another
copy at any of the great English herbaria.

Perhaps not surprisingly, however, there proved to be three com¬
plete sets in Mexico City. Lagasca, of course, had many ties with
Vicente Cervantes and the other Spanish botanists in Mexico, as well
as friends and acquaintances among political and religious figures of
the day. The late Professor Faustino Miranda very kindly arranged
an exchange by which one copy came from Mexico to the University
of Michigan.

Each of the four fascicles of Lagasca’s work begins with a title
page, and on the following page a pasted-in “Contenta” slip bearing
a neatly printed list of the species included in that particular part.
The full title, not a particularly long one for the time, is as follows:

Hortus Siccus Londinensis ; or, a Collection of dried specunens of
Plants , growing wild within twenty Miles round London , named on the
Authority of the Banksian Herbarium, and other original Collections .

The first fascicle includes an additional, prefatory page, follow¬
ing the title page, announcing and describing the intended publication.
Lagasca stated that the “chief aim of its Publication is to assist be¬
ginners, and to gratify those lovers of the Science, who, from their
other pursuits, cannot devote themselves to the laborious employment
of collecting, drying and arranging in a Herbarium, the Plants of their
native country.”

Although the original intention was to issue the sets in parts of
25 species each, a note in the third fascicle announced that the size
was being increased to 30 species per part. Thus the first three
parts include a total of 80 species. Part 4 contains 40 species (81-
110, and a “ Supple mentum,” nos. 111-120). No new species are de¬
scribed in the work; as Lagasca said in the preface, “In a Country
like England, abounding in all that is necessary for instruction,” the
work might have been superfluous “were its object the advancement
of Botanical Science.” Most of the species are the ones that were
common around London, and so well known to botanists. Druce, how¬
ever, pointed out that several of the plants represented the first known
records for the county of Middlesex, even though none was very rare.

Each specimen in the work is accompanied by its own printed
label. An interesting feature of these labels is the fact that Lagasca

7LaGasca and his ‘Hortus Siccus Londinensis’. Jour. Bot. Brit. For. 46:
163-169. 1908 (with appended biographical and bibliographical notes, pp. 169-

170, by James Britten).

1968

THE MICHIGAN BOTANIST

13

included on them not only the name of the plant, and information about
its distribution in the London area, but also what he understood to be
its distribution in Spain. His own primary research interest was in
the Spanish flora, and apparently even here in London it was occupy¬
ing a part of his time and thought.

None of the three works described above can be said to have in¬
fluenced the Science of Botany to any great extent; few taxonomists
today have ever heard of any of them. Nevertheless each has its own
peculiar interest. Gray’s Gramineae and Cyperaceae and Seringe’s
Saules are of some importance nomenclaturally because of the few
new names that were published in them. All three are of historical
interest because they are illustrative of a method of publication that
was superficially attractive, but was eventually abandoned because of
its inefficiencies. All of them are of very real importance, more¬
over, because they provide factual information about plant distribution
in certain limited geographical areas, at a period when rapid changes
in all habitats were beginning to come about because of man’s activity.

Publications of interest

HANDBOOK ON ORCHIDS. Brooklyn Botanic Garden, Brooklyn, N.Y. 11225, 1967.
81 pp. $1.25. Enough of our readers are interested in orchids to justify calling
attention to this handbook, actually the latest issue (Vol. 23, No. 2) of the Brook¬
lyn Botanic Garden’s quarterly Plants & Gardens. Among the two dozen arti¬
cles and features on orchids is one, “Growing Native Orchids,” by our own
Michigan orchid man, Fred Case.

THE MUSHROOM HANDBOOK. By Louis C. C. Krieger, with a new preface and
appendix on nomenclatural changes by Robert L. Shaffer. Dover Publications,
New York, 1967. vii + 560 pp. + 32 col. pi. $3.50. A facsimile (but slightly
enlarged) reproduction of the 1936 Macmillan edition of a handbook first pub¬
lished in 1935 by the New York State Museum, with the scientific names brought
up to date in an appendix but no changes in the original thorough introduction,
keys, descriptions, or bibliography. Although originally intended as a popular
guide to the higher fungi of New York state, it is a useful work in Michigan as
well. Although the publishers are to be congratulated on an excellent printing
job, their own blurb on the back cover fails to cite the correct number of pages
or of figures (there is no fig. 13, it having originally been a forest type map of
New York) and is guilty of considerable exaggeration in referring to Krieger
as “America’s foremost mycologist.”

14

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NOTES ON THE GENUS SUILLUS (BOLETACEAE)

Alexander H. Smith and Harry D. Thiers
University Herbarium and Department of Botany,

The University of Michigan, Ann Arbor, and
Department of Biology, San Francisco State College,

San Francisco, California

Since first publishing on Suillus (Smith & Thiers, 1964) we have
continued to gather data toward an eventual monograph of the group
for North America. As we are well aware, there is still much work
to be done before we can claim to have a reasonable understanding
of the taxa in this group for this continent. With vast areas of
“Suillus country” still very inadequately sampled we do not as yet
have sufficient knowledge of the extent of gene exchange or the mech¬
anism by which it occurs. All that can be said at present is that
obviously gene exchange is going on at a rapid rate. One is tempted
to assume it occurs through hybridization but this has not been es¬
tablished experimentally. That previously undescribed taxa can still
be found in a state such as Michigan, in which so much collecting has
been done, only re-emphasizes the need for more intensive as well as
extensive field work.

During the past season three species were collected which deserve
special mention. One was a heavy fruiting of a generally rare species,
S. albidipes; the second, the discovery in Michigan of the fungus we
believe Frost discovered in New England and which was published in
Peck (1889) as Boletus unicolor ; and the third appears to be a Euro¬
pean species which has long but improperly been regarded as a syno¬
nym of Suillus granulatus.

1. Suillus albidipes (Peck) Singer.

During the fall of 1967 an unusual fruiting of this species oc¬
curred in a pine plantation near Hell, Michigan, in Livingston County.
Dr. Bruce Fralick, the owner, planted there a variety of pines and
other conifers. The fruiting was first discovered by Florence Hoseney
and Ruth Zehner in the course of general collecting in and near the
Pinckney Recreation Area. On Oct. 28 Smith made a special effort to
estimate the number of basidiocarps present by pacing off and count¬
ing the number on five transects through the particular area involved
in the fruiting, which was roughly ten acres of white pine mixed with
other conifer species. The rough calculation indicated that on that
morning about 5,000 basidiocarps of Suillus albidipes in various stages
of development were present in the area. To our knowledge this is
the largest fruiting of this species on record. It has also furnished
reliable data on variation in the species, as well as data on the fruit¬
ing pattern.

Most of the fruiting occurred in circles marked by the outermost

1968

THE MICHIGAN BOTANIST

15

branches of each tree (the trees are not yet pole size); anywhere from
10-15 basidiocarps occurred under almost every tree whether it was
white pine or some other species. In other words, proximity to the
tree was more likely the result of ecological factors than a genetic
affinity of the fungus for the species of tree. Where there were open
spaces caused by cutting Christmas trees, the basidiocarps were more
scattered singly or in clusters. In these open areas many of the older
basidiocarps were whitish from fading but the characteristic pale vi-
naceous- cinnamon to cinnamon-ochraceous colors were present on
young caps beside the old ones. The false veil was regularly well
developed, varied from white to vinaceous buff, and in age all signs
of it were often obliterated. Usually, however, the marginal area of
the pileus retained a patchy appearance from the veil remnants. Col¬
ored glandular dots were readily visible over the apical region of the
stipes at maturity. Pileus size ranged from 4 to 18 cm, a range that
dispels any illusion that might have existed in some quarters that S.
albidipes is a “small” species. However, the most remarkable fea¬
ture of this fruiting was the extreme constancy of the characters
which distinguish the species — color of pileus, nature and color of veil,
white stipe at first, and small spores.

Suillus granulatus was fruiting in less abundance at the same time,
with young material ample, and no false veil was seen on any of the
basidiocarps at any stage of development. Suillus luteus was also abun¬
dant in the same area but no estimate was made of the number of ba¬
sidiocarps seen; they were almost as numerous as those of S. albidipes.
Suillus brevipes was conspicuous by its absence. As one might imagine,
however, the ground in the part of the plantation studied was practically
paved with basidiocarps of the various species of Suillus present.

We have no data on the precipitation pattern which preceded this
heavy fruiting. Local showers in September, however, must have been
the influencing factor for in the Stinchfield Woods of the University’s
School of Natural Resources, there was practically no fruiting of fungi
at the time of greatest abundance at the Fralick plantation. However,
two weeks later after heavy rains, the usual fungi including the Suillus
species discussed above, were there in abundance. The two areas are
less than ten air miles apart. The moral for mushroom hunters is
obvious: If you do not find your quarry in one woods, try another.

2. Suillus unicolor (Frost in Peck) Kuntz. Fig. 1.

Pileus 4-7 cm broad, plano-convex, expanding to nearly plane, thinly slimy-
viscid, perfectly glabrous or rarely with a few reddish dots near the margin,
color “empire yellow” to lemon-color and constant for the life of the basidio-
carp. Context mild, odor not distinctive, when cut pale to bright lemon-yellow,
with KOH giving a flush of red and finally bluish-fuscous (the color change on
the pileus surface similar).

Tubes decurrent, shallow (2-3 mm in young specimens, 5-6 mm in mature
ones), gelatinous, bright yellow, readily separable from pileus; pores minute,
bright yellow, not staining when bruised.

16

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Vol. 7

Stipe 4-6 cm long, 8-10 mm thick, solid, narrowed downward to a subradi¬
cating base, within lemon-yellow in the apex and olivaceous in the base, surface
lemon-yellow but staining dull olivaceous where handled, near apex with raised
areas of glandules but the dots conclorous with remainder of the surface or in
age finally brownish; copious basal mycelium pale yellow.

Spores 7-9 x 2.6-3 p , smooth, in profile narrowly inequilateral, in face
view elongate-subfusoid to nearly oblong, pale yellowish in KOH, pale tawny in
Melzer’s solution. Basidia 4-spored. Pleurocystidia in fascicles, when re¬
vived in KOH with vinaceous brown content and vinaceous brown encrusting
debris around the cluster, individual cystidia cylindric to narrowly clavate,
34-47 x 5-9 p. Pileus cutis of appressed hyaline narrow tubular gelatinous
hyphae lacking clamp connections.

Scattered under an aspen-birch-spruce stand near edge of swampy area,
northwest corner of Montmorency County, Michigan, July 26, 1967, Smith 74648.

In the dried state the glandular dots are very conspicuous on the
stipe. The yellow of the pileus is preserved in drying. Frost did not
mention the stipe as staining olive for his Boletus unicolor. We are
assuming that this character was overlooked. There was no veil along
the pileus margin and were no fibrils on the pileus, so it is not likely
that this species represents a variant of S. hirtellus. S. subaureus is
closest but the stipe in it has a ferruginous to fulvous zone in the
base when cut, and the color changes on the surface are to brownish.
Also the basal mycelium is white, and the pileus has appressed to-
mentum over the surface to some extent at least.

3. Suillus lactifluus (Wither, ex S. F. Gray) comb. nov.

Leccinum lactifluum S. F. Gray, Nat. Arr. Brit. Pis. 1: 647. 1821.

Pileus 4-10 cm broad, obtuse to convex, expanding to plane, surface slimy,
color pinkish buff to more or less pinkish cinnamon, in age darker reddish cinna¬
mon, glabrous, in age streaked to some extent beneath the slime or finally dis¬
tinctly virgate; margin sterile and overhanging the edge as a thin membrane
0.5-1. 5 mm wide (as in species of section Leccinum of Leccinum). Context
watery pallid but with lemon-yellow areas especially near the tubes, dull lilac-
blue with KOH, red in NH4OH, olive with FeS04; odor slight, taste acid.

Tubes pale yellow (“Baryta yellow”), when young and just forming duller,
soon beaded with numerous droplets of a milklike latex, these evaporating by
maturity, adnate to short decurrent or in intermediate stages slightly depressed,
up to 8 mm deep. Pores minute, 2-3 per mm at first but becoming 1-2 per mm
at maturity, staining brownish where bruised (cinnamon buff).

Stipe 2-5 cm long, evenly enlarged downward or in age narrowed downward,
virgate within with lemon-yellow streaks, staining cinnamon to vinaceous cinna¬
mon in the base; surface pale lemon-yellow overall at first, very inconspicuously
punctate above but the ornamentation concolorous with remainder of the surface
and not darkening by maturity in spite of the fact that the surface stains brown
from handling, colored dots show slightly on the dried specimens.

Spore deposit cinnamon when air dried. Spores 7-9 x 3 p, in profile sub¬
oblong to slightly inequilateral, in face view oblong or nearly so, yellowish in
KOH, in Melzer’s solution dingy yellowish.

Basidia 4-spored. Pleurocystidia in clusters with vinaceous brown en¬
crusting material when revived in KOH and also many cystidia with colored con¬
tent, hyaline when fresh but turning vinaceous with KOH; individual cystidia cy¬
lindric to narrowly clavate, 34-45 (50) x 6-9 p. Caulocystida in massive clust¬
ers, 50-90 x 7-12 p, when revived in KOH smooth and hyaline for the most part,

1968

THE MICHIGAN BOTANIST

17

bi)
**— <

Suillus unicolor (Smith 74648), x 1.

18

THE MICHIGAN BOTANIST

Vol. 7

but with the content finely colloidal and in KOH red only in the content in the tip
of the cell, smooth, pedicel flexuous, some surface hyphae with reddish content
in KOH also, and with reddish debris between the cells; the red pigment in some
clusters of cystidia slowly fading on standing in KOH. Epicutis of pileus of gela¬
tinous hyphae 2-5p diam. , hyaline to yellowish in KOH. Clamp connections absent.

Under white pine, University of Michigan Botanical Gardens, Washtenaw
County, Oct. 10,1967, Florence Hoseney and Ruth Zehner collectors (Smith 75165),

Singer (1967) Taf. VIII, fig. 10, under S. granulatus y illustrates the
above species admirably except for the slightly colored stipe orna¬
mentation. He illustrates droplets of latex. We have seen this on S.
granulatus in North America, but never so copiously. The species we
have described here is not S. granulatus in the usual sense— a species
with a spotted stipe. S. lactifluus has a thin sterile membranous
margin with no other signs of any development that might be con¬
strued as a veil. The lack of glandular dots on the stipe when fresh
was constant in several baskets full of basidiocarps. The pileus was
slimy as in S. brevipes. The latter, however, has a smoother stipe
which remains white a much longer time.

The situation in regard to this species is most peculiar histori¬
cally. It happens, however, that the early descriptions of Boletus
lactifluus describe our material about as perfectly as can reasonably
be expected— including the sterile marginal membrane of the pileus.
The stipe is not spotted in young to mature specimens. However,
Pilat, under the name Ixocomus granulatus , gives an excellent illus¬
tration-including the brown stains on the stipe. Singer (1947) stated
11 Leccinum lactifluum (With, ex Fr.) S. F. Gray is Suillus granulatus
(L. ex Fr.) Kuntze ssp. typicus.”

We cannot accept Singer’s statement. As Pilat (1954) has clearly
shown (pi. 23) the stipe is not glandular dotted (Fries underlined punc-
tato-granuloso in his description). Hence it is simply not justifiable
mycologically or nomenclaturally to accept S. lactifluus as the type
subspecies of S. granulatus . Our observations on the abundant collec¬
tions made at the University Botanical Gardens verify the existence of
a constant taxon in the characters the early workers described. Since
it is distinct at the same level as other commonly recognized Suillus
species such as S. brevipes , we recognize it here as an autonomous
species related to the latter.

LITERATURE CITED

Peck, Charles H. 1889. Boleti of the United States. Bull. N.Y, State Mus.
8. pp. 73-66.

Pilat, Albert, and Otto Usak. 1954. Mushrooms. Bijl, Amsterdam 342 pp.
120 pi.

Singer, Rolf. 1947. The Boletoideae of Florida. The Boletineae of Florida with
notes on extralimital species III Am. Midi Nat. 37: 1-35.

Smith, Alexander H. , and Harry D. Thiers. 1964. A Contribution Toward a
Monograph of North American Species of Suillus. Ann Arbor, privately pub¬
lished. 116 pp. and 46 pi.

1968

THE MICHIGAN BOTANIST

19

OBSERVATIONS ON THE BOLBITIACEAE.

IV. A NEW GENUS OF GASTROMYCETOID FUNGI

Roy Watling

Royal Botanic Garden, Edinburgh, Scotland

Whilst curating dried material of the Bolbitiaceae and so called
gastromycetoid fungi connected with this agaric family housed at the
University of Michigan Herbarium, Ann Arbor, during 1965, several
collections of an undescribed member of the group were examined;
these specimens possessed characters uniting the genera Bolbitiusi
Conocybe , and Galeropsis (in its widest sense). Later that same year
whilst attending a showing of colour transparencies taken by members
of the NorthAmerican Mycological Association, H. E. Schweitzer showed
what I took to be this same fungus; from information later given to me
by the photographer, it agreed in all respects macroscopically with the
taxon described below. Dr. Clark Rogerson of the New York Botanical
Garden, learning of my proposed monographic treatment of the bolbi-
tiaceous fungi, sent for my attention two ‘ off beat’ bolbitii. One turned
out to be a member of the Conocybe lactea -crispa complex, but the
second collection turned out to be the fungus under consideration; how¬
ever, this time habitat notes and colour transparency accompanied the
dried material. This last collection suggested that all the notes avail¬
able to me should be drawn together and this then confirmed the au¬
tonomy of the taxon.

The recognition of this taxon and its placing in a new genus pose
several problems, but these must be deferred to a much more lengthy
discussion of the Galeropsis complex; the purpose of this communica¬
tion is to place on record this unique fungus in order that its true af¬
finities and distribution may be ascertained. From the herbarium col¬
lections which have been made available— and there is little doubt that
other collections exist elsewhere hiding under unnamed collections of
species of Bolbitius or under no name at all— the taxon appears to be
fairly widespread in North America west of the Appalachians and east
of the Rockies. However, the speed with which the carpophore turns
into a gelatinous mass makes the collecting of this fungus difficult.
The collections which are to hand have only been found due to patient
and slow collecting often in areas under continual surveyance or at the
other extreme to very chance collecting.

It is proposed to describe this fungus in a new genus, Gastrocybe,
because of the gastromycetoid nature of the basidia, the almost sym¬
metrically placed basidiospores, and the reduction of the pileus to a
gelatinous mass within a short time after full elongation of the stipe.

GASTROCYBE Watling, gen. nov.

Pileus primo glandiformis vel ellipticus, vel cylindrico-ovoideus
vix vel numquam expansus, unctus vel viscidus denique gelatinosus et
informis. Stipes aequalis vel subaequalis, vulgo bulbillosus, albido-

20

THE MICHIGAN BOTANIST

Vol. 7

hyalinus dein pallidus, fragilis, gracilis, saepe elongatus. Lamellae
adnatae vulgo ventricosae ad faciem venosae vel intervenosae, ferru-
gineae. Caro tenuis, albida vel pallida. Cellulae cuticulae pilei pyri-
formes vel spheropedunculatae. Basidia et sterigmata aliquantam
crassitunicatae. Basidiosporae fere symmetrica, Cheilocystidia copiosa;
pleurocystidia nulla.

Pileus ellipsoid-campanulate or conic, hardly or never expanding,
greasy to viscid rapidly becoming reduced to a gelatinuous mass, pig¬
mented. Stipe white or hyaline, hollow, fragile and slender often very
long. Gills strongly forked, branched and interveined, fused to form a
fairly regular honey-comb pattern within the cavity formed between the
pileus context and the stipe surfaces. Pileus ‘cuticle’ hymeniform.
Basidia strongly thickened and spores borne almost symmetrically upon
prominent thickened sterigmata, cheilocystidia present, specialised;
pleurocystidia absent.

Type species: G. lateritia, by original designation.

Derived from the Greek gaster, gastros , belly, referring to the
cavity in which the basidiospores are produced, plus cybe, head, re¬
ferring to the pileus.

Gastrocybe lateritia Watling, sp. nov.

Pileus 5-25 (30) mm altus / 2- 10 (15) mm latus, ellipticus vel cy-
lindrico-ovoideus, vix vel numquam expansus, viscidus, membranaceus,
striatus, fragilis, denique gelatinosus et informis. Stipes 50-100 (130)/
1-1.5 (3) mm, albidus, subaequalis, vulgo sub-bulbillosus, flexuosus vel
erectus, laevis vel ad basim albopruinosus. Lamellae ferrugineae, ad
faciem venosae vel intervenosae, ramosae vel bifurcatae. Caro tenuis
albida vel pallida. Basidiosporae late ellipsoideae, poro germinativo
(9.5) 10.5-12 (14) x (6) 7-8 ju, laeves. Cheilocystidia diversiformia,
plerumque lecythiformia vel capitata.

Pileus 5-25 (30) mm high x 2-10 (15) mm broad, ellipsoid-cam¬
panulate to conic, long, subacute, not expanding or hardly significantly
so, greasy, tacky or viscid, membranous, pellucid-striate to disc, es¬
pecially in immature specimens, then rapidly reduced to a gelatinous
mass which adheres to the apex of the stipe, either retaining former
shape or becoming amorphous, uniform or slightly darker at disc, light
chocolate brown, darkening at maturity and in dried material more dark
chestnut, even bay, polished. Stipe 50-100 (130) mm x 1-1.5 (3) mm,
pure white, shining, hardly discolouring, equal, flexuous or straight,
slightly attenuated upwards, dry, smooth or sparsely pruinose (more
so towards base) obscurely striate, fragile, easily collapsing, hollow.
Gills rich rust-brown, well-formed with acute margin, branched,
forked and interveined, the branches fusing with adjacent gill plates,
thus forming a fairly regular honeycomb structure with smaller ‘ lo-
cules’ nearer the margin of the pileus. Flesh white in stipe, hardly
discernible in pileus; smell and taste not noted. Basidiospores broadly
ellipsoid, (9.5) 10.5-12 (14) x (6) 7-8 j a, smooth with a very prominent

1968

THE MICHIGAN BOTANIST

21

germ-pore and broad apiculus, rich honey brown under the micro¬
scope. Basidia 4-spored, commencing thick -walled or becoming thick¬
ened at maturity, ca. 26 p long and up to 13.5 j u broad, tapered abruptly
at base; sterigmata prominent and wide with mamillate apex whenform-
ing. Cheilocystidia capitate, irregularly lecythiform with ± long (up to
6 i u long x 1.75 ju), ± flexuous neck, head up to 4 p in diameter, body
7.5 /d broad; pleurocystidia absent, those cystidia developing at junction
of interveins being cheilocystidia. Caulocystidia absent or present at
stipe apex; pilocystidia flexuous, filamentous, irregularly distributed on
the stipe, more frequent at base. Pileocystidia absent and pileus ‘cuti¬
cle’ consisting of palisade of irregularly pyriform, hyaline cells up to
18 p broad and covered on the outer surface with a mucilaginous deposit.

In grass lawns, gregarious or solitary.

The genus differs from Galeropsis as typified by G. desertorum
Vel. in the hymeniform epicutis, the colour of the basidiospores, the
‘deliquescence’ of the pileus, and the very fragile stature.

Material examined: Old Botanical Gardens, Ann Arbor, Washtenaw
Co., Mich. 29 VIII 1947, Mains (MICH); Laramie, Albany Co., Wyoming,
12 VII 1950, R. Solheim (MICH); cemetery, Ithaca, Gratiot Co., Michi¬
gan, 29 VIII 1957, V. Potter (MICH); Ithaca, Gratiot Co., Michigan, 31
VIII 1957, V. Potter (MICH, type); Peoria, Peoria Co., Illinois, H. E.
Schweitzer; Ogden, Weber Co., Utah, 27 VII 1965, C. Rogerson (NY);
also probably (photograph only) a collection by Kauffman, Ann Arbor,
Washtenaw Co., Michigan, date ? .

It would be premature to discuss the affinities of this fungus but
it is significant to draw parallels with other members of the Bolbi-
tiaceae. First of all, it appears that although the fungus is agaricoid
in general aspect it does not seem to actively discharge its spores;
it must be considered therefore a gastromycete in the traditional sense
(cf. Weraoa nivalis Smith, 1965, p. 395). This is based not on actual
field observations but on the anatomy of the basidium and the position
of the undischarged basidiospores on the basidium. Potter was unable
to procure a spore -print and there is no evidence of a deposit on the
paper to which the Michigan, Utah, and Wyoming specimens have ad¬
hered on drying. The thick wall to the basidium apex and elsewhere
in some examples, the thickened sterigmata, and the almost symmetric
positioning of the basidiospores lead only to one conclusion.

It has been impossible to obtain complete sections of the pileus
cuticle and context because they soon disorganise as described above,
to leave only a few pyriform or elliptic cells distributed in mucilage.
This therefore resembles in all respects that found in Bolbitius vitel-
linus where it may become one and a half or even twice as thick as
the hymeniform epicutis. There is every reason to suppose that it is
chemically similar to this layer in B. vitellinus and a similar one
found in Suillus spp., although it was impossible to test the dried ma¬
terial with either Alcian blue or Periodic acid Schiff’s reagents. This
phenomenon is widespread in members of the Bolbitiaceae although it

22

THE MICHIGAN BOTANIST

Vol. 7

20p

1968

THE MICHIGAN BOTANIST

23

is never as well-developed as it is in species of Bolbitius ; in fact, it
is because this character is correlated with other details that it is used
to separate Bolbitius and Conocybe. Even in Conocybe coprophila where
the pileus is distinctly viscid the thickness of the pellicle is never
greater than the thickness of the hymeniderm and never becomes re¬
duced to a gelatinous mass; in C.tenera the mucilage layer is quite thin
and soon destroyed giving to the fresh specimens of this taxon a
greasy-shiny character. It is characteristic of members of the C.
lactea group (Sect. Candidae) for the pileus to become tacky and soft
when mature and this same group parallels the taxon under considera¬
tion in that the pileus is often long and slender, cylindrical and only
slightly expanding; the stipe is also white or hyaline and less distinctly
ornamented than in the Conocybe tenera group (i.e., Conocybe , sect.
Conocybe) or in the C. silignea group (Sect. Pilosellae ) (see Watling,
1965). It appears the mucilage is a mucopeptide or mucopolysaccharide.

We have no knowledge as yet as to how the spores are dispersed;
it does not seem to be a mechanism dependent on scavenging insects
as in the Phallales, for none of the collectors mentioned an odour and
there is little doubt that of all the three persons involved Potter would
have noticed an odour had there been one— not only had he a good sense
of smell but most important the specimens were growing gregariously
over several feet of lawn.

H. E. Schweitzer (personal communication) informs me that the
fungus first appears in the early morning, frequently before the sun
rises, a common feature of many species of Coprinus and probably
the reason why records of many are lacking in local floras, etc. The
pileus of G. lateritia is reduced to a mucilaginous mass within one
hour of the stipe reaching its greatest length. As soon as the sun
becomes fairly high in the sky the whole carpophore collapses amongst
the grass and rapidly shrivels. Dr. C. Rogerson reported to me that
his specimens appeared after a lawn consisting mainly of Kentucky
bluegrass ( Poa p ra tens is) had been irrigated in mid-summer. The notes
accompanying Potter’s material also support Schweitzer’s and Roger¬
son’ s observations.

I am grateful to Prof. A. H. Smith for allowing me to study the
specimens housed at the University of Michigan Herbarium, Ann Arbor,
and for arranging for me to study under a grant from the National
Science Foundation.

Fig. 1. Gastrocybe lateritia Watling. A, a, a\ and a": Potter 3654, 31 VIII 1957,
Michigan (type); B & b: Potter, 29 VIII 1957, Michigan; C: Schweitzer, VII 1965,
Illinois, Ci, same specimen one hour later; D, d, & dT: Rogerson, 27 VTI 1965,
Utah; E & e: Mains, 29 VIII 1947, Michigan; F: Kauffman, ?, Michigan; G & g:
Solheim, 12 VII 1950, Wyoming. A, B, C, Ci, D, E, F, & G: habit sketches;
a, b, d, e, & g: basidiospores; aT & d’: cheilocystidia; an: basidium. (C, C^,
& D from colour transparencies; F from photograph; A & B from colour sketches;
E & G from dried material.)

24

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Vol. 7

REFERENCES CITED

Smith, Alexander H. 1965. New and unusual Basidiomycetes with comments on
hyphal and spore wall reactions with Melzer’s solution. Mycopath. Mycol.
Appl. 26: 385-402.

Watling, Roy. 1965. Observations on the Bolbitiaceae 2. A conspectus of the
family. Notes Roy. Bot. Gard. Edinb. 26: 289-323.

FUNARIA FLAVICANS, A MOSS NEW TO CANADA

Howard Crum and Wilfred Botham
Herbarium, The University of Michigan, Ann Arbor,
and R. R. 1, Cottam, Ontario

Although Funaria flavicans Mx. is no more than a weed in the
southeastern United States, it is progressively less common north¬
ward, and its recent discovery in southern Canada is a distinct sur¬
prise. It was collected on the east side of Point Pelee in Essex
County, Ontario, on sand near Lake Erie where wood had been burned
( Wilfred Botham 8, May 7, 1967, MICH). Point Pelee is not far from
southeastern Michigan, and the known distribution of the species else¬
where to the south indicates that F. flavicans may be confidently ex¬
pected in Michigan as well.

It grows on soil (or occasionally brick or stonework, often in
burned-over places, in gardens, pastures, woods, roadsides, and rail¬
road grades) from New York to Oklahoma and south to the Gulf of
Mexico. (Specifically, it is thus far known from Ala., Conn., Dela.,
Ga., Ill., Ind., Ky., La., Md., Miss., Mo., N. Y., N. C., Ohio, Okla.,
Ont., Penn., S. C., Tenn., Va., Washington D. C., W. Va.).

Although similar in a general way to the ubiquitous Funaria hy-
grometrica , F. flavicans is easily recognized by its setae which are
scarcely hygroscopic, its nearly straight and only slightly asymmetric
capsules becoming only moderately plicate on drying, and its inner
peristome of emarginate or bluntly bilobed segments about one -quarter
as long as the teeth of the outer peristome.

1968

THE MICHIGAN BOTANIST

25

MICHIGAN PLANTS IN PRINT
New Literature Relating to Michigan Botany

This section lists new literature relating to Michigan Botany under four
categories: A. Maps, Soils, Geography, Geology (new maps and selected bulle¬
tins or articles on soils and geology as these may be of use to field naturalists
and students of plant distribution); B. Books, Bulletins, etc., and C. Journal
Articles (listing, respectively, all separate publications and articles in other
periodicals which cite Michigan specimens or include research based on plants
of wild origin in Michigan; —not generally including work on cultivated plants nor
strictly economic aspects of forestry, conservation, or agriculture); D. History,
Biography, Exploration (travels and lives of persons with Michigan botanical con¬
nections). When the subject matter or relation to Michigan is not clear from the
title, annotations are added in brackets. Readers are urged to call to the editor’s
attention any titles (1960 or later) which appear to have been overlooked— es¬
pecially in less well known sources.

A. MAPS, SOILS, GEOGRAPHY, GEOLOGY

The following topographic maps for Michigan have been published by the
U.S. Geological Survey, Washington, D.C. 20242, since the previous listing in
our January, 1967, issue. A11 are 7 l/2-minute quadrangles (scale of 1:24,000
or about 2 1/2 inches to a mile). Maps are supplied with green overprint show¬
ing wooded areas unless request is made to the contrary. Standard topographic
quadrangle maps are $ .50 each, and maps in the 1:250,000 series are $ .75
each, with a 20% discount on orders totalling $20.00 or more.

Following the name of the quadrangle, the county or counties in which it
primarily lies are added in brackets in the list below:

Ann Arbor East [Washtenaw]

Ann Arbor West [Washtenaw]
Aurelius [Ingham]

Damon [Ogemaw]

Dimondale [Eaton]

Eaton Rapids [Eaton]

Hamburg [Livingston & Washtenaw]
Lansing North [Clinton]

Lansing South [Ingham]

Maple Rapids [Clinton]

Ogemaw Springs [Ogemaw]

Oregon [Monroe, plus Ohio]
(Replaces earlier Point Place)
Perrinton [Gratiot]

Pinckney [Livington & Washtenaw]

Pompeii [Gratiot]

Riley [Clinton]

Riverdale [Gratiot]

Rose City [Ogemaw]

St. Helen [Roscommon]

St. Helen NE [Ogemaw]

St. Helen NW [Roscommon]

St. Johns North [Clinton]

St. Johns South [Clinton]

Selkirk [Ogemaw]

South Lyon [Washtenaw, Livingston,
& Oakland]

Sumner [Gratiot]

Vestaburg [Montcalm]

Wacousta [ Clinton]

Slightly revised editions of the Milwaukee (NK 16-2) and Hancock (NL 16-
1,2) sheets in the 1:250,000 series have been issued.

Bayley, R. W., C. E. Dutton, and C. A. Lamey. “1966” [1967]. Geology of the
Menominee Iron-bearing District Dickinson County, Michigan and Florence
and Marinette Counties, Wisconsin. U.S. Geol. Surv. Prof. Pap 513. 96 pp. +
4 pi. in pocket. $2.50 (U.S. Gov. Printing Off.). [Text largely on bedrock geol¬
ogy but with useful general information on the area (162 square miles), which
includes the botanically interesting Quinnesec region. Three 1:24,000 topo¬
graphic maps show the usual surface features, plus bedrock, including out¬
crops.]

Hough, Ashbel F. 1967. Twenty -five -year results of a red pine provenance study.
For. Sci. 13: 156-166. [Includes map of Lake States showing “average annual

26

THE MICHIGAN BOTANIST

Vol. 7

accumulation of normal average daily temperatures above 50 °F (‘growth de¬
grees’) and average January temperatures.”]

Wier, Kenneth L. 1967. Geology of the Kelso Junction Quadrangle Iron County,
Michigan. U.S. Geol. Surv. Bull. 1226. 47 pp. + 3 pi. in pocket. $1.50 (U.S.
Gov. Printing Off.). [Devoted chiefly to subsurface features, but with some
information on surficial geology and 1:24,000 map including outcrops.]

Wiitala, S. W., T. G. Newport, and E. L. Skinner. 1967. Water Resources of
the Marquette Iron Range Area, Michigan. U.S. Geol. Surv. Water-Supply
Pap. 1842. 142 pp. + 4 pi. in pocket. $1.50 (U.S. Gov. Printing Off.). [In

addition to data on ground-water supply, there is considerable information on
lakes and streams in the 610 square mile area covered (one-third of Marquette
Co., including all of the Carp and parts of the Chocolay, Michigamme, and Es-
canaba river basins); includes 1:96,000 map (not topographic) showing surface
geology.]

Wyman, Donald, and Harrison L. Flint. 1967. Plant Hardiness Zone Maps. Ar-
noldia 27: 53-56 + map. $ .10. [ Text discusses such maps and compares

Arnold Arboretum map with that of USDA (noted in Mich. Bot. 6: 27. 1967).
Color map indicates boundaries of 3 hardiness zones in Michigan, total of 10
in U.S. and southern Canada.]

B. BOOKS, BULLETINS, SPECIAL REPORTS, AND MISCELLANEOUS

Baxter, Dow V. 1967. Disease in Forest Plantations: Thief of Time. Cran-
brook Inst. Sci. Bull. 51. 251 pp. $8.50. [Based in large part on experience
from the Saginaw Forest, Washtenaw Co.; see review on p. 30.]

Braun, E. Lucy. 1967. The Monocotyledoneae Cat-tails to Orchids. Ohio State
Univ. Press, Columbus. 464 pp. $10.00. [See review in Mich. Bot. 6: 184.
1967. Michigan is mentioned in the distribution of several species, particu¬
larly Disporum maculatum\ and Trillium cemuum is drawn from a Michigan
specimen.]

Hotchkiss, Neil. 1967. Underwater and Floating- Leaved Plants of the United
States and Canada. U.S. Dep. Interior, Bur. Sport Fish. & Wildl. , Resource
Publ. 44. 124 pp. $ .65 (U.S. Gov. Printing Off.). [A guide, with rough sketches
but no keys, to plants of both fresh and marine waters. Sometimes difficult
problems (as in Isoetes and Sparganium ) are avoided by only mentioning the
species, and other times by over-enthusiastic “lumping” (most botanists would
hesitate, e.g. , to lump Ranunculus gmelinii and R. flabellaris , Potamogeton
berchtoldii and P. lateralis , P . foliosus and P. hillii). Michigan is specifically
mentioned in the distribution of four species.]

(Huron Mountain Wildlife Foundation). 1967. Report of Huron Mountain Wildlife
Foundation— 1955-1966 with Bibliography and Related Material. 96 pp. [In¬
cludes chronological list of botanical and other scientific work done in the
Huron Mountains 1925-1966, annotated bibliography, and texts of reports by
Aldo Leopold (1938), Michigan Natural Areas Council (1960-1962), and other
documents related to concern of the Huron Mountain Club for the scientific
values of its approximately 18,000 acres of wild land in Marquette Co.]
Johnson, Von J. 1966. Seasonal Fluctuation in Moisture Content of Pine Foliage.
North Central For. Exp. Sta. , U.S. For. Serv. Res. Note NC-11. [4] pp. [One
of sources of data was jack and red pine needles from Manistee National For¬
est in Michigan.]

Lowe, Josiah L. 1966. Polyporaceae of North America The Genus Poria. St.
Univ. Coll. For., Syracuse Univ., Tech. Publ. 90. 183 pp. $1.50 (Dept. Forest
Extension, State Univ. College of Forestry, Syracuse, N.Y. 13210). [Many of
the species are said to occur in Michigan; no detailed distribution data.]
Reid, Derek A. 1965. A Monograph of the Stipitate Stereoid Fungi. Nova Hed-
wigia Beih. 18. 382 pp. + 48 pi. $25.00 [Two species of Cotylidia are cited

from Michigan localties.]

1968

THE MICHIGAN BOTANIST

27

Thompson, Paul W. 1967. Vegetation and Common Plants of Sleeping Bear.
Cranbrook Inst. Sci. Bull. 52. 47 pp. $1.50. [ See review, Mich. Bot. 6: 175.
1967]

Wilce, Joan Hubbell. 1965. Section Complanataof the Genus Lycopodium. Nova
Hedwigia Beih. 19. 233 pp. + 40 pi. $15.00. [Several species and hybrids in
Michigan, supported by full citations and general distribution maps. Repeated
apologies for use of an alleged invalid sectional name are unnecessary, as
Section Complanata was published by Victorin 10 years before the requirement
of a Latin diagnosis.]

C. JOURNAL ARTICLES

Allison, Leonard N. 1967. Beggar- Ticks cause mortality among fingerling Coho
Salmon. Progressive Fish-Culturist 29: 113. [Fruit of Bidens cernuus em¬
bedded in gills and oral cavity of fish at Platte River Trout Rearing Station at
Honor.]

Anderson, Gerald W. , and David W. French. 1965. Western gall rust in the Lake
States. For. Sci. 11: 139-141. [Found on jack pine in Hiawatha National For¬
est in Upper Peninsula.]

Baxter, Dow V. “1966” [1967*]. Fungus development from field to forest in

pine plantings of 1904 and later. Pap. Mich. Acad. 51: 149-156. [Fungus es¬
tablishment in plantings in Washtenaw Co.]

Bailey, Robert E. “1966” [1967*]. Postglacial pollen sequence in a bog of the
Loesell Field Laboratory of Eastern Michigan University. Pap. Mich. Acad.
51: 167-174. [Study in Ypsilanti Tp. , Washtenaw Co.]

Barnes, Burton V. 1967. Indications of possible mid-Cenozoic hybridization in
the aspens of the Columbia Plateau. Rhodora 69: 70-81. [Includes compari¬
sons with Michigan hybrids.]

Boneham, Roger F. 1967. Devonian Tasmanites from Michigan, Ontario, and
northern Ohio. Pap. Mich. Acad. 52: 163-173. [Michigan locations for these
fossils of green algae are cited in Alpena, Antrim, and Monroe counties]

Brown, Robert Thorson. 1967. Influence of naturally occurring compounds on
germination and growth of jack pine. Ecology 48: 542-546. [Effects of ex¬
tracts of 56 plant species “commonly associated with jack pine” on germina¬
tion of seeds from “northern Michigan” (Upper Peninsula). Salix pellita , one
species which gave nearly complete inhibition, would (if correctly identified)
be not only a significant range extension but also an unusual habitat if com¬
monly associated with jack pine in the U.P.]

Brunett, Fel V. 1966. An archaeological survey of the Manistee River Basin:
Sharon, Michigan to Sherman, Michigan. Mich. Archaeol. 12: 169-182. [In¬
cludes general observations on vegetation (also soils and history) of the val¬
ley.]

Church, George L. 1967. Pine Hills Elymus. Rhodora 69: 330-351. [Includes
results of experimental crossings of Michigan Hystrix with Illinois Elymus .]

Church, George L. 1967. Taxonomic and genetic relationships of eastern North
American species of Elymus with setaceous glumes. Rhodora 69: 121-162.
[Includes a key to eastern N. A. species of Elymus and citation of two repre¬
sentative specimens of E , wiegandii from Michigan.]

Colingsworth, Roscoe F. , Matthew H. Ho hn, and Gary B. Collins. 1967. Post¬
glacial physicochemical conditions of Vestaburg Bog, Montcalm County, Mich¬
igan, based on diatom analyses. Pap. Mich. Acad. 52: 19-30.

♦Although no new taxa of plants (only of insects and of fossil echinoids), for
which exact date of publication must often be known, are described in Vol. 51 of
the Papers of the Michigan Academy, the attention of bibliographers is called to
the fact that mailing to most Academy members was on May 4, 1967— although
authors did receive reprints prior to that date.

28

THE MICHIGAN BOTANIST

Vol. 7

Cooke, Wm. Bridge. 1962. A taxonomic study in the “Black Yeasts.” Myco-
path. Mycol. Appl. 17: 1-43. [Microstroma juglandis cited from Carya in
Michigan.]

Crum, Howard, and Norton G. Miller. 1967. Three propaguliferous Pohlias
from Michigan. Bryologist 70: 118-119.

Culberson, William Louis, and Chicita F. Culberson. 1967. A new taxonomy
for the Cetraria ciliaris group. Bryologist 70: 158-166. [ Generalized distri¬
bution maps include occurrence in Michigan of 3 of the 4 species.]

Davis, H. A , Albert M. Fuller, and Tyreeca Davis. 1967. Contributions toward
the revision of the Eubati of eastern North America. Castanea 32: 30-37.
[This first installment of a revision of Rubus subgenus Rubus (“Eubatus ”
nom. illeg.) includes a key to sections and a treatment of section Hispidi, with
references to type collections oiR.plus, R. kalamazooensis , R. compos , and
R. distinctus in Michigan (of which only R. plus is maintained).]

Dissing, Henry, and Morten Lange. 1967. Notes on the genus Helvella in North
America. Mycologia 59: 349-360. [Type of H. connivens sp. nov. is from
Cross Village, Emmet Co.; other specimens of this and of H. stevensii (in¬
cluding type) cited from Michigan.]

Forman, Richard T. T. 1967. New and uncommon Wisconsin mosses. Bryolo¬
gist 70: 115-117. [Michigan records for several of the species are referred
to.]

Getz, Lowell L. 1961. Temperatures in different vegetation types in southern
Michigan. Jack-Pine Warbler 39: 132-147. [ Data on maximum and minimum
temperatures at different levels throughout the year in six vegetation types in
Mud Lake Bog area, Washtenaw Co.]

Gilliam, Jeanne A., and Ronald O. Kapp. 1967. A post-Wisconsin pollen sequence
from Vestaburg Bog, Montcalm County, Michigan. Pap. Mich. Acad. 52: 3-17.

Graham, Shirley A. 1966. The genera of Araliaceae in the southeastern United
States. Jour. Arnold Arb. 47: 126-136. [A footnote by C. E. Wood mentions
occurrence oiOplopanax on Isle Roy ale.]

Imshaug, Henry A., and Irwin M. Brodo. 1966. Biosystematic studies on Leca-
nora pallida and some related lichens in the Americas. Nova Hedwigia 12: 1-
59 + 12 pi. [includes full citations of many Michigan collections and distribu¬
tion maps.]

Kapoor, B. M., and J. R. Beaudry. 1966. Studies on Solidago. VII . The taxo¬
nomic status of the taxa Brachychaeta, Brintonia, Chrysoma, Euthamia, Oli-
goneuron, and Petradoria in relation to Solidago. Canad. Jour. Genet. Cytol.
8: 422-443. [Includes chromosome count of 2n = 18 for 5. canadensis var.
canadensis from Sugar Island.]

Kawano, Shoichi, and Hugh H. litis. 1966. Cyto taxonomy of the genus Smilacina
(Liliaceae) II. Chromosome morphology and evolutionary considerations of
New World species. Cytologia 31: 12-28. [Material of S. racemosa from
Misery Bay, Houghton Co., represents the octopioid race of that species.]

King, Dennis R, and George S. Hunt. 1967. Effect of carp on vegetation in a
Lake Erie marsh. Jour. Wildlife Managem. 31: 181-188. [ Includes vegeta¬
tion map of study area in Monroe Co.]

Laing, Charles C. 1967. The ecology of Ammophila breviligulata. II. Genetic
change as a factor in population decline on stable dunes. Ecology 77: 495-
500. [One of the two sites from which experimental material was obtained
was Little Sable Point, Oceana Co. ]

Lentz, Paul L. 1967. Delineations of forest fungi. Several species of Deutero-
mycetes and a newly described Botryobasidium. Mycopath. Mycol. Appl. 32:
1-25. [Oidium simile cited from Barry Co.]

Levesque, Lucien, et Pierre Dansereau. 1966. Etudes sur les violettes jaunes
caulescentes de l’est de l’Amerique du Nord. I. Taxonomie, nomenclature,
synonymie et bibliographie. Nat. Canad. 93: 489-569. [A massive bibliography

1968

THE MICHIGAN BOTANIST

29

of literature citations, with generalized distribution maps showing V. eriocarpa
throughout Michigan and V. pubescens absent from western 3/4 of Upper Penin¬
sula. Both taxa in fact range throughout Michigan, as documented by Russell’s
maps published in 1965.]

Lombard, Frances F. , and Robert L. Gilbertson. 1966. Poria luteoalba and some
related species in North America. Mycologia 58: 827-845. [Two Michigan
collections of P. eupora cited.]

Milanez, Adauto I., and E. S. Beneke. “1966” [1967*]. An unusual Saprolegnia
from the Gull Lake area. Pap. Mich. Acad. 51: 175-182. [Plants close toS.
diclina from Kalamazoo and Barry counties.]

Paterson, R. A. 1967. Benthic and planktonic phycomycetes from northern
Michigan. Mycologia 59: 405-416. [Data from Grand Traverse Bay and Stur¬
geon Bay, Lake Michigan, and Douglas Lake.]

Pringle, James S. 1967. Taxonomy of Gentiana, section Pneumonanthae, in
eastern North America. Brittonia 19: 1-32. [Includes citation of representa¬
tive specimens and localities on distribution maps for 6 species in Michigan;
two Marquette Co. specimens appear to be hybrids G. linearis x rubricaulis . ]
Schneider, G. , D. P. White, and R. L. Harlan. “1966” [ 1967*] . Soil moisture
regime under old growth hardwoods. Pap. Mich. Acad. 51: 13-21. [Study in
Tuomey Woodlot, MSU, Ingham Co.]

Smith, Alexander H. 1965. New and unusual basidiomycetes with comments on
hyphal and spore wall reactions with Melzer’s solution. Mycopath. Mycol.
Appl. 26: 385-402. [Xerocomus truncatus Singer from Emerson (Chippewa
Co.) described.]

Stone, William J. H., and Ralph J. Green, Jr. 1967. The epiphytology of spear¬
mint rust in Indiana. Mycopath. Mycol. Appl. 31: 17-26. [One of 18 collec¬
tions of Puccinia menthae studied came from 4n M. arvensis x 4n M. sylvestris
from an unspecified Michigan locality.]

Thieret, John W. 1966. Synopsis of the genus Calamovilfa. Castanea 31: 145-
152. [C. longifolia var. magna mapped around Lakes Huron and Michigan

(omitting Lake Superior station); var. longifolia adventive in southwestern
Michigan.]

Urie, Dean H. “1966” [1967*]. Forest cover in relation to water yields on out-
wash sand soils. Pap. Mich. Acad. 51: 3-11. [Studies in jack pine, red pine,
oak, and lowland hardwood fojests in Manistee Co.]

Witmer, S. W. 1964. Butomus umbellatus L. in Indiana. Castanea 29: 117-119.

[Mentions occurrence in 7 counties of Michigan.]

Zahner, Robert, and John R. Donnelly. 1967. Refining correlations of water
deficits and radial growth in young red pine. Ecology 48: 525-530. [Data
from plantations near Pellston.]

D. HISTORY, BIOGRAPHY, EXPLORATION

Anderson, Edgar. 1965. David M. Gates, the new director. Missouri Bot. Gard.
Bull. 53(9): 1-7. [Sketch of Michigan-trained physicist with biological inter¬
ests, including two photos of ecological field work in Michigan.]

Buell, Murray F. 1963. Eminent ecologist for 1963 [William S. Cooper] . Bull.
Ecol. Soc. Am. 44: 108-109. [ Biographical sketch of an Alma College gradu¬
ate, author of classical ecological treatise on vegetation of Isle Roy ale.]
Dana, Samuel T. 1967. A Final Word, pp. ix-xi in Disease in Forest Planta¬
tions: Thief of Time, by Dow V. Baxter, Cranbrook Inst. Sci. Bull. 51. [A

tribute to the late Prof. Baxter; see also Smith, below.]

Evers, Robert A. 1965. Arthur Gibson Vestal 1888-1964. Trans. Illinois
Acad. 58: 77-81. [Vestal developed his interest in botany as a youth near
Pentwater, Michigan; he spent the summer of 1913 working with H. A. Gleason
at the University of Michigan Biological Station, near which “Vestal’s Bog”
bears his name.]

30

THE MICHIGAN BOTANIST

Vol. 7

Graustein, Jeannette E. 1967. Thomas Nuttall Naturalist, Explorations in Amer¬
ica 1808-1841. Harvard Univ. Press, Cambridge. 481 pp. $11.95. [Biography
of naturalist who first seriously studied the Michigan flora; chapter 5 includes
his journey through the Upper Great Lakes in 18 10— on which further informa¬
tion is provided in Mich. Bot. 6: 81-94. 1967.]

Martin, W. E. 1965. Arthur Gibson Vestal 1888-1964. Bull. Ecol. Soc. Am. 46:

27-29. [Includes list of major publications; cf. Evers, above.]

Smith, Alexander H. 1967. Dow V. Baxter (1898-1965). Mycologia 59: 565-567.

[ Obituary of University of Michigan professor, authority on wood-decaying
fungi, collector in Michigan, Alaska, and elsewhere.]

Sparrow, Frederick K. 1964. Robert M. Johns, 1928-1963. Mycologia 56: 309.
[ Brief sketch of 1958 Michigan Ph.D. whose mycological work in Michigan
centered on Phycomycetes.]

Taylor, Mrs. J. Lee, and Mrs. Robert Turner, eds. 1967. The Nut Jar A Cook¬
book. Mich. Nut Growers Assoc. 132 pp. $2.50. [Includes a history of nut
culture in Michigan; see review in Mich. Bot. 6: 178. 1967.]

Thackrey, Donald E. (ed.). 1967. Research Definitions and Reflections. Essays
on the Occasion of The University of Michigan’s Sesquicentennial. Univ. of
Mich., Ann Arbor. 207 pp. $1.75. [Includes essays on fungi by F. K. Sparrow
and on algae by W. R. Taylor, the latter a general historical discussion of algal
research at the U. of M.]

Welch, Winona H. 1964. Truman G. Yuncker 1891-1964. Taxon 13: 189-192.
{Dr. Yuncker was a native of Michigan and graduate of MSU.]

Review

DISEASE IN FOREST PLANTATIONS: THIEF OF TIME. By Dow V. Baxter.
Cranbrook Institute of Science (Bulletin 51), Bloomfield Hills, Michigan. 1967.
xviii + 251 pp. $8.50.

Those of us who remember back to the era of the 30’ s and the “CCC” days
somehow received the impression that all one had to do to grow trees was to
plant them and then wait for the crop to mature. If this idea still persists in
anyone’s mind, Dr. Dow Vawter Baxter’s posthumus book should dispel the last
vestiges of it. In fact after reading the book, one might wonder why any trees
in a plantation survive. However, for anyone with a healthy interest in trees, I
recommend this work as fascinating, not only because it is full of Baxter’s
superb photographs, but it is also written in his characteristically engaging
style. Baxter, as his students in forest pathology were keenly aware, preferred
to approach the problems of forest disease from an ecological standpoint, and
this is the most valuable aspect of Disease in Forest Plantations— it focuses on
the ecological aspects of having large areas of planted trees all the same age
and all the same species, and their vulnerability to attack. I heartily recom¬
mend the book to all naturalists.

—Alexander H. Smith

Editorial Notes

The October number (Vol. 6, No. 4) was mailed October 25, 1967.

THANKS TO REVIEWERS. One of the chief functions of the members of
the Editorial Board is to review manuscripts submitted for publication, consid¬
ering botanical content, clarity of presentation, and suitability for this journal.
Reviewers of individual papers are traditionally anonymous, but the editor takes
this opportunity to express his appreciation not only to members of the Editorial
Board (named on the inside front cover) but also the following, who have served
as reviewers for one or more articles submitted for volumes 4-6: Drs. Burton
V. Barnes, Charles B. Beck, W. S. Benninghoff, John E. Cantlon, Howard Crum,
Margaret B. Davis, Francis C. Evans, Henry Imshaug, Peter B. Kaufman, Robert
L. Shaffer, Helen V. Smith, Otto T. Solbrig, Ronald L. Stuckey, H. W. Vogelmann.

Publications of Interest

SPRING FLORA OF MINNESOTA Including Common Cultivated Plants. By
Thomas Morley. Department of Botany,” University of Minnesota, Minneapolis,
1966. 283 pp. , wire-bound. $3.50 plus postage, from Nicholson Hall Book¬

store, Univ. of Minnesota, Minneapolis 55455. From its concise, lucid intro¬
duction to its above-average glossary, this little flora has aroused a more
enthusiastic response from the editor than any other he has seen in a long
time. It is both sensible and up to date in regard to taxonomic and nomencla-
tural problems, and includes the common cultivated plants as well as the
native and naturalized species which ordinarily bloom before June 7 in
Minnesota (a total of 850 species). Keys and helpful additional comments
are included, but no illustrations (except for those preceding the glos¬
sary, a county map, and a map of Minnesota vegetation types). Minnesota resi¬
dents are extremely fortunate to have such a fine volume available. Michigan
botanists will note the omission of such familiar spring species of our flora
as Cardamine douglassii, Tiarella cordifolia , Trillium erectum, Acer pensyl-
vanicum. The more northwestern flora, earlier cutoff date, and absence of
illustrations make this flora perhaps a little less useful in Michigan than
Fassett ’s excellent Spring Flora of Wisconsin. But the more modern text, in¬
clusion of cultivated plants, and greater number of grasses and sedges are
decided advantages. We look forward eagerly to the anticipated hardcover
edition contemplated by the Universiy of Minnesota Press!

HARVEST WITHOUT PLANTING Eating and Nibbling Off the Land. By Erika
E. Gaertner. Published by the author, Chalk River, Ontario, 1967. 66 pp.

$2.00 from the author or from Donald F. Runge, Ltd., 243 Pembroke St. W. ,
Pembroke, Ontario. Those who like to nibble off the land will doubtless find
some new recipes in this original compilation— unless they already have di¬
rections for Boiled Beaver Tail, Braised Muskrat in Tomato Sauce, Birch
Cheesecake, Elderberry Fritters, Milkweed Bud Salad, Cat- tail Pollen Pan¬
cakes, Crayfish Bisque, and such delicacies.

CONTENTS

Rare Old Publications in Michigan Herbaria

Rogers McVaugh . 3

Notes on the Genus Suillus (Boletaceae)

Alexander H. Smith & Harry D. Thiers . 14

Observations on the Bolbitiaceae.

IV. A New Genus of Gastromycetoid Fungi

Roy Watling . 19

Funaria flavicans, A Moss New to Canada

Howard Crum & Wilfred Botham . 24

Michigan Plants in Print . 25

Review— Disease in Forest Plantations: Thief of Time . 30

Editorial Notes . 31

Publications of Interest . 13, 31

( On the cover: Hedgehog or Medusa-head fungus , Hericium
erinaceus, photographed on a white oak trunk at the Rose Lake
Wildlife Experiment Station , Clinton Co., Michigan, by Peter
Stettenheim, September 27, 1966. The white threads across
the bottom are spider webs laden with spores from the fungus .)

LIBRARY

* ?

THE

mar 26 1968

Vol. 7, No. 2

NEW YORK

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MICHIGAN BOTANIST

March, 1968

THE MICHIGAN BOTANIST-Published by the Michigan Botanical Club four times per

year: January, March, May, and October. Second-class postage paid at Ann Arbor,

Michigan.

Subscriptions: $3.00 per year. Single copies: $.75.

All back issues are available, at the following prices per volume:

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Articles and notes dealing with any phase of botany relating to the Upper Great Lakes
Region may be sent to the editor in chief. The attention of authors preparing manuscripts
is called to “Information for Authors” (Vol. 6, p. 202; reprints available from the editor).

Editorial Board

Edward G. Voss, Editor in Chief

Herbarium, The University of Michigan, Ann Arbor, Michigan 48104

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1968

THE MICHIGAN BOTANIST

35

A PRELIMINARY STUDY OF CLA VARIADELPHUS

IN NORTH AMERICA

Virginia L. Wells and Phyllis E. Kempton
Anchorage, Alaskal

INTRODUCTION

It was our original intention to include Alaskan species of Clav ariadelphus
in a series of papers concerning Alaskan fungi, the first two of which have al¬
ready been published (1965, 1967). But it was soon apparent that our species
could not be properly placed until a critical examination of all North American
species of this genus was undertaken. This paper is the result of our four-year
study of the Clav ariadelphus flora of North America.

Clav ariadelphus was originally erected by Donk (1933) to separate three
species, C. pistillaris (type), C. truncatus, and C. ligula, from the other clavarioid
fungi. The genus received little attention until Doty (1948) included it in his
key, further delimiting it to those species which turn green with FeS04. Corner
(1950) divided the genus into three subgenera: Eu-Clav ariadelphus, Cantharel-
lopsis, and Typhulopsis. The species Corner placed in subgenus Typhulopsis
(C. junceus and C. fistulosus ) are not under consideration in this paper because
we were unable to obtain sufficient collections to examine and we feel that a
more critical study of them is necessary to determine their correct generic assign¬
ment.

Corner’s other subgenera, Eu-Clav ariadelphus and Cantharellopsis , are
based mainly on the size, shape, and degree of apical sterility or fertility of the
fruitbodies and, in our opinion, are artificial divisions. We believe that the genus
Clav ariadelphus is composed of two distinct series of species with some overlap¬
ping microscopic and macroscopic features, and that the genus should be divided
by a more basic character— the shape of the spores. We are therefore proposing
that the genus be divided into two sections: Section Clav ariadelphus, with
spores two to two-and-a-half times as long as broad; and Section Ligula, with
spores three or more times as long as broad.

Because we have had only limited use of standard color charts, most of the
colors given in the descriptions are our own color concepts. Where color terms
are enclosed in quotation marks, they represent Ridgeway’s colors used by other
authors or collectors.

All Alaskan collections are stored in the Wells-Kempton (W/K) private her¬
barium at Anchorage, Alaska. The following abbreviations are used in citing col¬
lections from other herbaria:

CINC— University of Cincinnati

DAOM— Dept, of Agriculture, Ottawa: Mycological Herbarium

MICH— University of Michigan

1327 Fifth Ave. (VLW) & 1020 H St. (PEK).

36

THE MICHIGAN BOTANIST

Vol. 7

RHP— Ronald H. Petersen, private herbarium
TENN— University of Tennessee
NCU— University of North Carolina
RM— University of Wyoming.

ACKNOWLEDGMENTS

We wish to thank the following people for their generosity in supplying specimens
and information critically needed for this study: Dr. J. Walton Groves and Dr. J. A. Parma-
lee of the Canada Department of Agriculture, Dr. Ronald H. Petersen and Dr. L. R. Hesler of
the University of Tennessee, Dr. C. R. Leathers of Arizona State University, Dr. J. B. Boze¬
man of the University of North Carolina, Dr. R. L. Shaffer of the University of Michigan,
Dr. Clark T. Rogerson of the New York Botanical Garden, Dr. Margaret Fulford of the
University of Cincinnati, and Dr. W. G. Solheim of the University of Wyoming. We are
particularly indebted to Dr. A. H. Smith of the University of Michigan for his encourage¬
ment, advice, and general assistance throughout the research and preparation of this paper.
We also wish to acknowledge financial support from the National Science Foundation
(Grant GB 4634) for field work and publication expenses during the latter part of this
study.

MACROSCOPIC CHARACTERS

Most species of Clavariadelphus are rather large in comparison with other
simple clavarioid fungi. The size of the fruitbody is of some taxonomic value in
distinguishing between species but, of course, is not absolute. The shape of ma¬
ture fruitbodies, however, is of more constant value as a distinguishing feature. All
young fruitbodies are typically cylindrical and obtuse to acute at the apex; but
with age, some species become clavately inflated in the upper part, others be¬
come apically- truncated, and one species (C. mucronatus) is both truncated and
mucronate. In all species the hymenial surface is smooth but becomes longitu¬
dinally rugulose in age. All are typically simple and unbranched although they
may have short branches or irregularities on or near their apices, and may grow
connately if in close contact during development.

The basic colors of all species are yellow to buff to alutaceous, tan, or
vinaceous-brown but shades of orange, pink, green, or lilac may occur and are
sometimes constant enough to be used as diagnostic features at the species level.
The bases of the fruitbodies are pallid to whitish but the color difference be¬
tween the upper and lower parts is gradual and of no specific value. As far as is
now known, only one species (C. pistillaris ) stains with age or bruising.

The color of the fresh context is white to pallid ochraceous and more or
less unchanging except for C. pistillaris which becomes vinaceous to brown when
cut or bruised. The flesh of all young fruitbodies is firm and solid at first, be¬
coming softer and more floccose with maturity, and occasionally collapses to
produce a hollow or perforation at the apex in extreme age. When dried, the
texture of the fruitbodies is fragile-brittle and specimens are easily crushed or
shattered in storage if not adequately protected.

As far as we know, no species of Clavariadelphus has an odor distinctive
enough to be used as an identifying character but taste, bitter or sweet, is a dis¬
tinctive and constant feature in some species.

1968

THE MICHIGAN BOTANIST

37

We believe that the spore deposit color, whether white or ochraceous, is an
important diagnostic character at the species level and is essential in cases where
other differentiating features such as spore size, fruitbody color, habitat, etc. are
nearly identical. We have not been able to detect any change in spore deposit
color, either fading or darkening, after drying or storing over a period of years.

We consider it important to know the following macroscopic features of
fresh specimens in order to properly place species of Clavariadelphus : spore de¬
posit color, color and color-changes of fruitbody and context, habit and habitat,
size and shape of the fruitbody, and the taste.

MACROCHEMICAL REACTIONS

The chemical solutions employed during this study were 10% FeS04, 15%
KOH, 28% NH4OH, and 2% phenol.

In testing fresh material, solutions were applied directly to the various
parts of the fruitbody. With dried material, small sections of hymenium and
flesh were first dipped in either methanol or isopropyl alcohol to act as a wet¬
ting agent, then placed on a glass slide and a drop of the testing solution applied.
The reactions to the above-mentioned chemicals are included in the descriptions
of the species, but some additional data are worth mentioning.

Doty (1948), in his key to the genera of clavarioid fungi, used the green re¬
action to FeSCG as a generic character of Clavariadelphus. We have found this
reaction to be constant in the material we have studied although, in contrast to
Doty’s thirty seconds, we found that it may take from thirty seconds to three
minutes for the reaction to be visible. The shade and intensity of the color var¬
ies considerably, not only between species, but also between individual fruit-
bodies of the same species. The color may be pale to deep forest green, olive, or
gray-green. On dried material, we found the reaction to be strongest on the con¬
text of the fruitbody and frequently intense enough to cause the mounting med¬
ium to turn green.

Except for C. unicolor, the surface of all truncated species in section
Clavariadelphus turned pink to red with KOH. However, one anomalous collec¬
tion of either C. truncatus or C. borealis {Smith 46253, MICH), from which no
spore deposit had been taken, gave a surprising negative KOH reaction. Sub¬
sequent examination showed all the fruitbodies in that collection to be sterile.
The deviation of chemical reaction appears to be related to spore production but
the problem requires further research.

To date, phenol has not produced a color change when applied to dried
material, but fresh fruitbodies of some species do stain strongly vinaceous on all
parts where phenol is applied. The phenol reaction may prove to be a valuable
character, but further testing on fresh specimens will be necessary before the ex¬
tent of its usefulness can be fully determined.

DISTRIBUTION, HABITAT, AND HABIT

We feel that much of North America has not been investigated sufficiently
to permit establishment of actual distributional patterns for many species of
Clavariadelphus. However, with the exception of C. unicolor which is known

38

THE MICHIGAN BOTANIST

Vol. 7

primarily from southeastern United States, all North American species appear to
be more common in northern regions or at high altitudes in southern regions-
While the fruiting season varies from one part of the continent to another, all
species are reported as autumnal.

The members of section Ligula are characterized by their densely gregari¬
ous manner of growth on matted needles and debris of coniferous trees. On the
other hand, those in section Clavariadelphus do not share either a common habi¬
tat or manner of growth. Some species show a preference for predominantly
deciduous woods while others have been found only in coniferous forests. Their
manner of growth can be described variously as solitary, gregarious, cespitose, or
subfastigiate. Some of the species in this latter section are little known, and fur¬
ther collecting may give more habit and habitat information for them than is
now recorded.

MICROSCOPIC CHARACTERS

Most of the microscopic characters of Clavariadelphus species are rather
monotonous in their similarity. For example, spores in those of both sections
are hyaline with yellow refractive oil drops or a yellow granulose content
(KOH), inamyloid, unicolorous in cotton blue, smooth, thin-walled, and rela¬
tively large, mostly over 8/ilong. However, the shape of the spores is distinctive
enough to divide the genus into the two sections we are proposing. In section
Clavariadelphus, the spores are ellipsoid, broadly ellipsoid, or oblong, mostly
equilateral, and are about two to two-and-a-half times as long as they are broad.
In section Ligula, the spores are narrowly ellipsoid to “boletoid,” the longer ones
depressed on the dorsal side and ventricose on the ventral side giving them a
characteristic “sway-backed” appearance in profile, and they are typically three
to five times as long as they are broad.

Since spore deposits were seldom present with herbarium collections used
for this study, we were concerned as to whether spores taken from dried speci¬
mens would measure differently from spores of a deposit. After numerous ex¬
aminations of our own collections, we concluded that there is no appreciable
difference in size between the two; nor could we find any difference in spore
size between those mounted in water from a fresh deposit and a KOH mount
from the same deposit after it had been dried.

Corner (1950) used the thickening in depth of the hymenium as a generic
character for Clavariadelphus. In connection with the “C. pistillaris ” he ex¬
amined, he reported, “hymenium gradually thickening from 100/uto 300ju. deep
in old specimens, very compact, with many included spores.” In the various
species we have examined, there is a constant proliferation of basidia for the full
life of the fruitbody, with old basidia collapsing after spore discharge and being
replaced by new basidia in various stages of development, but we have found no
evidence that this contributes to a measurable increase in the depth of the
hymenium and we rarely found spores trapped within the hymenium.

The basidia are narrowly clavate, usually filled with a yellow refractive
granulose content or oil drops, and are very long— a character Donk (1933) used
in separating Clavariadelphus from Clavaria. In our examinations, we have

1968

THE MICHIGAN BOTANIST

39

discovered that the basidia vary in length as much as 20 p to 60/ul from one area
of the fruitbody to another, a factor undoubtedly contributing to the disparity
in basidial measurements noted throughout the literature.

The longest basidia occur on the upper one-half to two-thirds of the fruit-
body, or just below the margin of the truncation of some species. The shortest
basidia occur on the apex of some species, on the margin of some truncated
species, or near the base of the fruitbody, and are frequently distorted or abort¬
ed. The sterigmata are straight on the longer basidia but tend to be malformed
on the shorter basidia in that they may branch, grow confluently, or be obtuse
and peg-like (Fig. 1). Corner (1950) used the term “caulocystidia” to describe
the sterile clavate cells he found near the base of the “C. pistillaris ” he ex¬
amined. Similar cells are frequently present near the base of all Clavariadelphus
fruitbodies but we prefer to call them “caulobasidia” because they are not
always sterile (Fig. 2).

The basidia are predominantly 4-spored, but some 2-spored basidia are
nearly always present in the hymenium, and 1- or 3-spored basidia are not un¬
usual near the fruitbody base or on the marginal area of some truncated species.
The basidia collapse after discharging the spores but we have never seen them be¬
come secondarily septate as reported by Petersen (1967) in Clavulina.

We consider cystidia to be absent from species in this genus even though
there are some sterile elements present on the margins of some truncated species
which might qualify for the terms “pseudoparaphyses” or “cystidioles” in the
sense of Singer and Gamundi (1963). But since the margin of the truncation is a
transitional zone between the fertile hymenium of the sides and the sterile sur¬
face of the apex, one should expect to find an immixture of fertile and sterile
elements in this area.

The subhymenium is composed of hyaline to yellow, narrow, interwoven
hyphae which tenaciously cling to the hymenial elements and join them to the
tramal hyphae. We feel that it is the presence or absence of the subhymenium,
or the degree to which it is developed, that plays a vital part in shaping the fruit¬
body. On the sides, where the subhymenium is strongly developed, there is little
lateral expansion. But in some species where the subhymenium is weakly or ir¬
regularly developed near the apex, that area may expand thus becoming clavate
or lobed or nearly convoluted. On truncated fruitbodies where the subhymeni¬
um is absent over the apex, there is relatively unrestricted lateral growth.

This constricting influence of the subhymenium is readily demonstrated
when attempts are made to crush out the hymenium under a cover slip. Pressure
causes the hymenial elements to become arranged into “fans” or “rosettes” with
their bases firmly tied together by the subhymenial hyphae. Further pressure
may result in the hymenial elements breaking off above their connection with
the subhymenium, leaving the subhymenium more or less intact. The bond be¬
tween the subhymenium and the trama is not as strong and pressure on the cover
slip will cause the subhymenium to separate from the tramal hyphae.

All tramal hyphae are hyaline to yellowish, or occasionally with oleaginous
content, smooth, and inamyloid. Initially they are narrow, moderately thick-
walled and arranged more or less longitudinally parallel, but with age they

40

THE MICHIGAN BOTANIST

Vol. 7

10

1968

THE MICHIGAN BOTANIST

41

become inflated, thin-walled, and interwoven in the center of the fruitbody
while remaining parallel in the base and under the subhymenium along the sides
of the fruitbody.

The mycelial hyphae occur in fascicles which macroscopically appear as
whitish strands binding the substratum into clumps, pads, or mats. These vege¬
tative hyphae have thin to thick walls which are either smooth or decorated with
yellow crystals as seen when mounted in KOH or water. The crystals are of vary¬
ing shapes and distribution: They may be up to 7.5/i long, aciculate, and lying at
various angles across the hyphae; irregularly angular to subangular, 1 .5-4.5/lx diam,
and arranged in rosette-like clusters; or they may be scattered with no particular
pattern and of both sizes and shapes. At times they may become dislodged from
the walls, leaving sections of the hyphae bare. We have not found either the
shape or the distribution of the crystals to be of any taxonomic value at the
species level.

Clamp connections are abundantly present in all parts of the fruitbody and
the mycelium, and occur as three different types: 1. Thin-walled clamp connec¬
tions in which the diameter of the clamp is approximately the same as the diam¬
eter of the connected hypha (Fig. 3). This type is the most common. 2. “In¬
flated” clamp connections in which the diameter of the clamp is two to four
times that of the connected hypha, and in which the walls are thick (1-1. 5/4 ) and
yellow refractive as seen when mounted in KOH (Fig. 4). These are most fre¬
quently found in the basal regions of the fruitbody and on the mycelium.
3. “Medallion” clamp connections in which the clamp branch forms a free arc or
loop over the septum (Fig. 5). This type may be found scattered throughout the
upper part of the fruitbodies but is less common than the preceding types.

We are including ten species in this paper, four of which are new. It is in¬
teresting to note that of these four, two have previously been described under
names of older species: C. cokeri was described as Clavaria pistillaris by Coker
(1923) and C. mucronatus as Clavaria unicolor by Doty (1944).

CLA VARIADELPHUS Donk

Fruitbodies typically simple (exceptionally with one or several short
branches at the apex), small to relatively large (2-20 cm tall), clavate, subventri-
cose, ligulate or turbinate, or rugose to irregularly contorted; apices acute to ob¬
tuse, truncate, or occasionally mucronate; yellow, ochraceous, alutaceous, tan,
or buff but some species with red, pink, orange, olive, gray, green, or lilac also
present, color sometimes changing markedly with age or bruising, base pallid but
not sharply delimited; surface smooth becoming longitudinally rugose with age;

Figs. 1-15. Clavariadelphus. 1. Basidia with malformed sterigmata. 2. Caulobasidia. 3. Com¬
mon thin-walled clamp connections. 4. Inflated clamp connections. 5. “Medallion” clamp
connections. 6. Spores of C. subfastigiatus. 7. Tubular to clavate terminal cells on sterile
apex of C. unicolor . 8. Flexuous to branched terminal cells on sterile apex of C. unicolor.
9. Spores of C. lovejoyae. 10. Clavate to subventricose terminal cells on sterile apex of C.
lovejoyae. 11. Spores of C. borealis. 12. F usoid-ventricose terminal cells on sterile apex of
C. borealis. 13. Spores of C. mucronatus. 14. Spores of C. ligula. 15. Spores of C.
sachalinensis. Figs. 3, 4, 5, 6, 9, 11, 13, 14, and 15, scale A. Figs. 1,2, 7, 8, 10, and 12, scale B.

42

THE MICHIGAN BOTANIST

Vol. 7

hymenium covering entire fruitbody except the base or absent also on the apex
of some truncated species; flesh firm becoming spongy and floccose, whitish,
sometimes staining when cut, green with FeSC^; habit gregarious, solitary, or
subfastigiate, in deciduous or coniferous woods. Spores white or ochraceous in
deposit, oblong, broadly ellipsoid, or narrowly ellipsoid to boletoid, smooth,
thin-walled, hyaline with yellowish granular content or yellowish oil drops, in¬
amyloid. Basidia predominantly 4-spored but 2-spored ones usually also present,
long and narrowly clavate. Cystidia none. Hyphae thin to moderately thick-
walled, smooth to obscurely roughened (incrusted with crystals on mycelium),
oleiferous hyphae sometimes present, clamp connections constantly present.
Type species: Clavariadelphus pistillaris (Fr.) Donk.

KEY TO THE SECTIONS

A. Spores oblong, ellipsoid, or broadly ellipsoid with the length two times

the width or shorter . Section Clavariadelphus

A. Spores narrowly ellipsoid to “boletoid” with the length three times the

width or longer . Section Ligula

Section CLAVARIADELPHUS

KEY TO THE SPECIES

1. Spore deposit ochraceous; fruitbodies truncate; in coniferous woods . . . 1. C. truncatus

1. Spore deposit white; fruitbodies clavate, truncate, or irregularly flat¬
tened; solitary, gregarious, or fastigiate; in deciduous or coniferous

woods . 2

2. Fruitbodies clavate; scattered to gregarious in deciduous woods;
flesh and hymenium staining vinaceous-brown with age or bruising;

taste often bitter . 2. C. pistillaris

2. Not as above . 3

3. Fruitbodies fascicled to subfastigiate at the base, clavate, subclavate, or

irregularly flattened; in coniferous woods . 4

3. Fruitbodies truncate; solitary or gregarious; in coniferous or deciduous

woods . 5

4. Fruitbodies pallid at first becoming light cinnamon or brownish-or¬
ange, the apex obtuse . 3. C. subfastigiatus

4. Fruitbodies pink at first becoming alutaceous, the apex acute at first

but becoming obtuse or flattened and antler-like . 4. C. cokeri

5. Fruitbodies pink to lilac at first becoming alutaceous; in deciduous

woods; typically in southeastern United States . 5. C. unicolor

5. Not as above . 6

6. Apex of fruitbody red; spores 11-17/i long . 6. C. lovejoyae

6. Apex of fruitbody yellow-orange; spores 9-1 ljii long . 7. C. borealis

1. Clavariadelphus truncatus (Quel.) Donk, Rev. Niederl. Homobas. Aphyll.
2:73. 1933.

Clavaria truncata Quelet, Enchir. Fung. p. 221. 1886.

Craterellus pistillaris Fries, Epicr. Syst. Mycol. p. 534. 1838.

Not Clavaria truncata Lovejoy, Bot. Gaz. 50:385. 1910.

1968

THE MICHIGAN BOTANIST

43

The following macroscopic description is adapted from Leathers’ (1955) de¬
scription with additional notes provided by Dr. A. H. Smith.

Fruitbody 9-15 cm tall, 4-6 cm diam. at the apex, simple, turbinate, or cla-
vate with a broadly convex apex which becomes truncated and often slightly de¬
pressed at disc with age, margin of the truncation recurved, gradually attenuated
downwards to a usually bulbous (1-2.5 cm diam.) but not sharply delimited base
immersed in a mat of white mycelium; pinkish-brown “onion skin pink” to “vi-
naceous tawny,” drying deep vinaceous-brown “hessian brown,” apex bright to
golden yellow when fresh “capucine yellow” to “orange buff,” drying to “clay
color;” surface smooth at the base becoming rugulose upward, at times almost
veined; context white, drying to pale cream “pinkish buff,” solid, spongy becom¬
ing fibrillose on drying, green in ferric sulphate, odor none, taste very sweet.
Spore deposit pale ochraceous.

Sparsely gregarious to cespitose on conifer needles, Aug.-Oct.

The following microscopic data are from our examination of Leathers 409
(MICH) labeled Clavaria truncata:

Spores 9-1 1.5 x 6-7p, broadly ellipsoid, hyaline with yellow oil drops, walls
smooth and thin. Hymenium present only on the sides of the fruitbody. Basidia
narrowly clavate, 92-115 x 10.5-1 2. 5/ll with sterigmata up to 7.5/i long, mostly
4-spored but 1- and 2-spored ones also present. Cystidia none. Subhymenium
interwoven of yellowish hyphae 2.3-3/u diam. Truncate apex sterile or with a few
scattered basidia; epicutis a hymeniform layer of hyaline to yellowish, thin-
walled, clavate to ventricose cells 50-80 x 10-17/u. Tramal hyphae yellowish,
smooth, 4. 5-7. 5 p diam, parallel in the base and under the hymenium, interwoven
in the center of the fruitbody. Clamp connections present.

Material studied: MICHIGAN: Leathers 409 (MICH).

Observations: Since we have never seen fresh material of Clavariadelphus
truncatus, we have relied heavily on Leathers’ (1955) macroscopic description.
Unfortunately, the collections from which Leathers drew his description no
longer include spore deposits but he was emphatic in stating that the deposits
were ochraceous.

Except for the spore deposit color, C. truncatus appears to be identical with
C. borealis. However, this close relationship between species is not unusual in
Clavariadelphus. A similar situation occurs between C. pistillaris (Fr.) Donk and
C. pistillaris sensu Corner (1950) and again, between C. ligula and C. sachalin-
ensis, although in these latter two cases there is also a difference in spore size.

Further field work will be necessary before the geographical range of C.
truncatus can be charted. It is possible that many collections we examined dur¬
ing the course of this study were actually C. truncatus but since spore deposits
were lacking, these collections have not been cited in this paper. It is also possi¬
ble that further field work may show differences of which we are now unaware
between C. truncatus and C. borealis.

44

THE MICHIGAN BOTANIST

Vol. 7

2. Clavariadelphus pistillaris (Fr.) Donk, Rev. Niederl. Homobas. Aphyll.

2:73; 1933.

Clavaria pistillaris Linnaeus ex Fries, Syst. Mycol. 1 'All . 1821.

Not Clavariadelphus pistillaris sensu Corner, Ann. Bot. Mem. 1:279-281.

1950.

Fruitbody 6-15 (-20) cm tall, 8-30 (-45) mm diam. near the apex, simple,
cylindrical becoming subventricose to clavate, apex usually rounded and fre¬
quently inflated with age, gradually narrowing below to a rather deeply inserted
base; at first whitish with a yellow apex, soon becoming ochraceous to ochrace-
ous-brown overall, staining brown to vinaceous-brown with age or handling, base
remaining whitish at all ages but not sharply delimited; moist to dry, unpolished,
smooth becoming longitudinally rugulose; context white, staining brown to
vinaceous-brown when cut, firm becoming floccose, odor none, taste mild to
bitter.

Chemical reactions: Fresh material: FeSOq— all parts greenish; KOFI— all
parts negative or hymenium yellow to orange; phenol— all parts dark vinaceous-
brown. Dried material: FeSOq— all parts dark green; KOH— all parts negative.

Spore deposit white. Spores (7.5-) 9-13.8 (-16) x 4.5-7.5/d, ellipsoid to ob¬
long, hyaline with yellow oil drops, walls thin and smooth. Hymenium present
on sides and apex of the fruitbody. Basidia narrowly clavate, (60-) 75-130 x
7.5-13 (-15 )p with sterigmata up to llju long, mostly 4-spored but 2-spored
ones also frequently present. Cystidia none. Subhymenium interwoven of
hyaline to yellowish hyphae 2-4.5 (-5.5 )p diam. Tramal hyphae hyaline to
yellowish, 3-14p diam., walls smooth to obscurely roughened, thin to slightly
thickened, parallel in the base and adjacent to the hymenium, interwoven in the
center of the fruitbody. Clamp connections present.

Gregarious to scattered in predominantly deciduous woods, autumn to
early winter.

Material studied: United States: ALASKA: Fairbanks, Wells & Kempton 8/15/64-10,
8/18/64-8, 8/26/65-14, 8/26/65-15, 8/26/65-21 (W/K); Palmer, E. Wiebe 8/28/67-22 (W/K).
CALIFORNIA: Trinity Co., Lanphere 12-6-62 (MICH); Crescent City, Smith 55916
(MICH); San Francisco, Whited 20 (MICH). IDAHO: Nordman, Smith 54566 (MICH).
MAINE: Madawaska Lake, Bigelow 4095 (MICH). MICHIGAN: Pinckney, Smith 64185
(MICH); Oakland Co., Smith 7012 (MICH); Haven Hill, Smith 64300 (MICH); Jackson Co.,
Shaffer 2604 (MICH); Pellston, Thiers 4394 (MICH). NORTH CAROLINA: Macon Co.,
Rogerson C266 (RHP), Petersen C948 (RHP); Transylvania Co., Petersen C475 (RHP);
Thomas Ridge, A. J. S[harp] 8205 (TENN); Cataloochee, Bain 8396 (TENN); Linville
Falls, Totten 5759 (NCU); Chapel Hill, Coker 13960 (NCU), Burnham 68 (NCU), Honeycutt
14068 (NCU); Highlands, Jones 13101 (NCU). OREGON: Grants Pass, Smith 55799,
56045, 56046 (MICH). TENNESSEE: Knox Co., S. L. Wallace 6543 (TENN); Nales Creek,
Hesler 22939 (TENN). VERMONT: Windham Co., Shaffer 3510 (MICH). VIRGINIA:
Mt. Lake, Meyer 14706 (TENN). WASHINGTON: Mt. Rainier Nat. Park, Smith 31663
(MICH). Canada: NOVA SCOTIA: Kentville, Harrison 95904, 111226 (DAOM). QUEBEC:
Drayton F5713 (DAOM); Groves and Pennoyer 54146 (DAOM).

Observations: In this paper, we are using Donk’s (1933) concept of C.
pistillaris because the spore size he gives for this species ([8-] 10-16 x [4-] 5-7. 5p)
is in agreement with that of collections found in North America.

1968

THE MICHIGAN BOTANIST

45

We could not accept Corner’s (1950) concept of C. pistillaris because he
gave a much broader spore size (12-15 x 7-1 Oju), and there is a strong indication
(Cotton and Wakefield, 1919) that the spore deposit of British material is ochra-
ceous whereas all North American collections are reported to have a white spore
deposit.

During the course of this study, we examined several collections which ob¬
viously are related to C. pistillaris but which we could not describe in this paper
because of the lack of adequate field notes and spore deposits.

3. Clavariadelphus subfastigiatus Wells & Kempton, sp. nov.

Fructificatio 4-9 cm alta, 5-11 (-25) mm diam. ad apicem, simplex, crescens in fasci-
culis basibus subfastigiatis, cylindrica demum subventricosa vel subclavata, interdum com-
planata; diluta cinnamomea, ad basim pallens; apex obtusus, fertilis; laevis demum longitud-
inaliter rugulosa; contextus albidus, firmus demum floccosus, sapor initio amarus demum
nullus. Sporae in cumulo albae, 8.5-10.7 (-12) x 5.6-6.5/i, late ellipticae. Cystidia nulla.
Fibulae praesentes. In sylvis coniferae. TYPUS: Wells et Kempton 10/8/66-3. Salmon
River Campground, Mt. Hood, Oregon (W/K).

Fruitbody 4-9 cm tall, 5-1 1 (-25) mm diam. at the broadest part, occuring
in clusters, simple or occasionally connate at the base, cylindrical becoming sub-
ventricose to subclavate, occasionally flattened, apices obtuse; primordia pallid
flesh-color becoming light cinnamon to dark ochraceous-flesh or brownish-or¬
ange, paler toward the base, apices concolorous; moist to dry, unpolished,
smooth becoming rugulose, base pallid pruinose, with rather stout white rhizo-
morphs; context whitish, slowly and slightly vinaceous when cut or bruised,
solid, longitudinally fibrous at first becoming subfloccose, odor none, taste
slightly to moderately bitter at first but soon fading to mild!

Chemical reactions: Fresh material: FeS04— all parts weakly green; KOH—
hymenium forest green fading to yellow-green, context yellow; NH4OH— hymeni-
um weakly green with fumes, yellowish with solution; phenol— all parts dark vina¬
ceous. Dried material: FeS04— all parts blackish-green; KOH— all parts negative.

Spore deposit white. Spores 8.5-10.7 (-12) x 5.6-6.5/i, ellipsoid, hyaline
with one to several yellow oil drops, walls thin and smooth (Fig. 6). Hymenium
present on sides and apex of the fruitbody. Basidia narrowly clavate, 84-112
(-131) x (7.5-) 8.5-1 lp with sterigmata up to 9.2p long, mostly 4-spored but
some 2-spored ones usually also present. Cystidia none. Subhymenium 25-7 5p
thick, interwoven of hyaline to yellow hyphae 2. 5-3. 5 (-ll)p. diam. Tramal
hyphae hyaline to yellowish, 3-7.5 (-9.5 )p diam., walls smooth to obscurely
roughened, thin to slightly thickened, longitudinally parallel in the base and
under the hymenium, interwoven in the center of the fruitbody. Clamp con¬
nections present.

Gregarious in clusters of 6-20 fruitbodies, rarely singly, on well decayed
humus of coniferous woods, Oct.

Material studied: OREGON: Salmon River Campground, Mt. Hood, Wells & Kempton
10/8/66-3 (Type), 10/8/66-4 (W/K).

Observations: A few of the fruitbodies in the type collection have acute,
shriveled, and blackened apices which we attribute to adverse weather condi¬
tions, probably freezing, but the majority have normally rounded and concolor¬
ous apices.

46

THE MICHIGAN BOTANIST

Vol. 7

C. subfastigiatus seems to be most closely related to C. cokeri from which
it differs in color, slightly broader spores, and possibly in the structure of the
subhymenium. The depth of the subhymenium is given in the description of C.
subfastigiatus because it appeared to be broader and more distinctive than in any
other Clavariadelphus species we have examined. We feel that more collections
will have to be studied before this subhymenial structure can be evaluated as a
taxonomic feature.

4. Clavariadelphus cokeri Wells & Kempton, sp. nov.

Fructificatio usque ad 20 cm alta, simplex, crescens in fasciculis basibus confertae et
penitus insertae, cylindrica demum anguste clavata, interdum complanata; apex subacutus
demum obtusus, interdum brevis ramosus; rosea demum alutacea; laevis demum longitudina-
liter rugulosa; contextus albidus, firmus demum tloccosus, sapor leviter piperatus. Sporae in
cumulo albae, 9.4-11.3 x 4.7-5 (-5.6)//, oblongae vel subellipticae. Cystidia nulla. Fibulae
praesentes. In sylvis coniferae. TYPUS: W. C. Coker R. 27, September 6, 1919, Redding,
Connecticut (NCU).

Fruitbody up to 20 cm tall, occurring in clusters with the bases crowded
and deeply inserted, cylindrical and subacute at the apices at first, becoming nar¬
rowly clavate with an obtuse apex, sometimes flattened or branched like an ant¬
ler; smooth becoming rugosely channeled; dull rosy when young becoming rosy-
buff to alutaceous, the tip deepest colored, base whitish; context white, firm be¬
coming floccose, odor none, taste pleasant but mildly peppery.

Chemical reactions: Dried material: FeSOq— very obscurely greenish (re¬
action nearly negative); KOH— all parts negative.

Spore deposit white. Spores 9.4-11.3 x 4.7-5 (-5.6)/i (Coker measured
them 7.4-10 x 3. 8-4. 5 [-4.8] /i), oblong to subellipsoid, hyaline with yellow gran-
ulose content or with numerous yellow oil drops, walls thin and smooth. Hyme-
nium present on the sides and apex of the fruitbody. Basidia narrowly clavate,
(55-) 70-95 x (5.6-) 7-8.5) u, 4-spored, rarely 2-spored. Cystidia none. Sub¬
hymenium interwoven of yellow hyphae 2.8-4 p diam. Tramal hyphae hyaline
to yellowish, 5. 5-8. 5 /i diam., longitudinally parallel under the hymenium, inter¬
woven in the center of the fruitbody. Clamp connections present.

In clusters under hemlock, Sept.

Material studied: CONNECTICUT: Redding, Sept. 6, 1919, Coker R. 27 (Type)
(NCU).

Observations: The above macroscopic description was adapted from Cok¬
er’s field notes included with the collection. The type collection is ample but in
poor condition. The hyphae are so collapsed that accurate measurements are
difficult to obtain and the collection has been severely attacked by another fun¬
gus at some time or other during its storage. The fruitbodies are now covered
with innumerable alien spores but, fortunately, the spore deposit is clean. We at¬
tribute the weak FeS04 reaction to the fact that the collection appears to have
been dipped in some chemical solution which may have interfered with the reac¬
tion. However, Coker’s excellent notes, the good spore deposit, Miss Eaton’s
fine illustration (Coker 1932, PI. 23), plus the collection, poor though it is, com¬
bine to provide a good concept of this taxon.

1968

THE MICHIGAN BOTANIST

47

C. cokeri is very close to C. subfastigiatus but can be distinguished by its
different colors and narrower spores. Both species occur in clusters in coniferous
woods and both appear to be rare. The geographical range for neither is known
at this time.

5. Clavariadelphus unicolor (Rav. ex Berk.) Corner, Ann. Bot. Mem. 1:284-285.

1950.

Craterellus unicolor Ravenel ex Berkeley, Grevillea 1 : 148. 1873.

Craterellus corrugis Peck, Bull. Torrey Bot. Club 26:69. 1899.

Clavaria pistillaris var. unicolor f. A, Coker, Jour. Elisha Mitchell Sci. Soc.

63:60. 1947.

Clavaria unicolor (Rav. ex Berk.) Doty, Ore. Monogr. Bot. 7:23. 1944.

Fruitbody 2.5-6 (-8) cm tall, 2-4 (-5) cm diam. at the apex, simple, clavate
to obclavate with a rather short, equal to tapering base (5-8 mm diam.), soon
flaring upward rather abruptly to become turbinate with a truncated apex which
is frequently margined with an elevated, furrowed ridge, apex sometimes collaps¬
ing with age to form a cavity; pale ochraceous with tints of flesh pink, violet or
red-brown (“tawny” to “ochraceous-tawny”), becoming darker and somewhat
rufescent on drying (“brownish-vinaceous,” “Rood’s brown” to “Natal brown”),
paler toward the base and over the apex; smooth becoming longitudinally rugose
or nearly cerebriform near the margin of the truncation; context white, firm be¬
coming floccose, odor and taste not recorded.

Chemical reactions: Dried material: FeSOq— all parts green; KOH— all
parts negative.

Spore deposit white. Spores (8-) 9.4-11 (-13) x (4.2-) 4. 5-5. 5 (~6)p, ellip¬
soid to subovate, hyaline with yellow oil drops or granular content, walls thin
and smooth. Hymenium present on the sides and sometimes over the apex of the
fruitbody. Basidia narrowly clavate, (60-) 70-95 x 8.5-1 \p with sterigmata up to
9.5p long, mostly 4-spored but 2-spored ones also frequently present. Cystidia
none. Subhymenium interwoven of hyaline to yellowish hyphae 2Ap diam.
The truncate apex fertile or sterile: If fertile, the hymenium continued from the
sides but the subhymenium not differentiated over the apex. If sterile, the epi-
cutis of the truncation of two general types with intermediate forms: 1. A
hymeniform layer of yellow, tubular to narrowly clavate terminal cells 37-75 x
3-7 (-9.5)/u(Fig. 7); or 2. An interwoven layer of yellow, filamentous, flexuous,
branched, terminal hyphae with cells 18-40 (-75) x 3-6 (-7.5)jU (Fig. 8), some¬
times arising from a vertically oriented subcutis. Tramal hyphae hyaline to yel¬
lowish, 3.5-12 (-17)ft diam.; walls smooth to obscurely roughened, thin to slight¬
ly thickened; hyphae longitudinally parallel in the base and adjacent to the
hymenium, interwoven in the center of the fruitbody; oleiferous hyphae fre¬
quently present. Clamp connections present.

Gregarious in predominantly deciduous woods, usually in southeastern
United States, Nov.-Dee.

Material studied: TENNESSEE: Timberlake Road, Hesler 22240 (TENN); New
Hopewell, Hesler 10091 (TENN), Hesler & A. J. Sfharpj 7094 (TENN); Clinton Pike, Hesler
6544 (TENN); Ball Camp Pike, Hesler 1729 (TENN). NORTH CAROLINA: Chapel Hill,
Reasoner 1994 (NCU), Couch 3885 (NCU), Coker & C. R. Bell 14062, 14064 (NCU).

48

THE MICHIGAN BOTANIST

Vol. 7

Observations: The distribution of C. unicolor is interesting. Of all the her¬
barium material we examined the only collections we saw of it were from Ten¬
nessee and North Carolina. The original collection came from South Carolina,
Murrill (1940) reported it from Florida, and Coker (1947) added Georgia to the
list. Since there is so much evidence that C. unicolor is a southeastern United
States species, it is surprising that it occured even once as far north as Massachus-
sets— Peck’s Craterellus corrugis( 1899). We have not seen Peck’s type collection
but from his description and because Clavariadelphus unicolor is such a distinc¬
tive species, even in the dried condition, we feel that Burt (1914) was correct in
placing C. corrugis in synonomy with C. unicolor Rav. ex Berk.

Doty’s (1944) report of C. unicolor from the Pacific Coast is erroneous.
The species he described is included in this paper as C. mucronatus and while
there is some superficial resemblance, C. unicolor is readily distinguished by its
broader spores, different structure over the apex, and its habitat in deciduous
woods.

As far as we now know, C. unicolor is the only truncated species in Clavari¬
adelphus in which the truncation can be either completely sterile or completely
fertile. We have not studied sufficient material to report conclusively on the
overall percentage of fertility vs. sterility, but in one relatively large collection
we found it to be about 50-50.

6. Clavariadelphus lovejoyae Wells & Kempton, nom. nov.

Clavaria truncata Lovejoy, Bot. Gaz. 50:385. 1910.

Following is Fovejoy’s original description:

“Pileate tops bright red, shading into reddish orange at top of stipe to dull
flesh color at its base: ends truncate, convex to plane to somewhat concave,
0.5-3 cm. broad, smooth: whole plant to within a few centimeters of base of
stipe covered with a white bloom, persisting in dried specimens: flesh creamy,
spongy: stipe longitudinally grooved to base, 3-10 cm. long: spores white, 14 x
Ip.

“Habitat: Humus soil under balsam and spruce trees; gregarious and cespi-
tose, 4-6 in a group; Foxpark, alt. 2900 meters, August 8, 1909, no. 66.”

The following description is from our examination of the type collection:

Collection good, composed of 8-10 fruitbodies and a generous amount of
substratum including white to tannish mycelium. No spore deposit. Dried fruit-
bodies up to 10 cm tall, up to 15 mm diam. at the apex, subclavate to truncate,
base usually inflated; apex dull red to vinaceous red-brown, sides splotched with
vinaceous red-brown near the apex, buff to grayish-brown below, base pallid tan
(a few specimens have small, bright red spots just below the apex which look like
lacquer or nail polish). The white bloom mentioned by Fovejoy is not now par¬
ticularly discernable.

Chemical reactions: Dried material: FeS04— context dull green; KOH—
hymenium discolors solution to pink.

Spores of two general sizes and shapes: 1. Mostly (9.4-) 10.7-13 (-13.8) x
5-6 p, broadly ellipsoid to oblong (Fig. 9); 2. Rather frequently 15-17 x (5.3-)

1968

THE MICHIGAN BOTANIST

49

6-7/i, ellipsoid but compressed at the waist making them appear “peanut” shaped;
both types hyaline with yellow oil drops or granulose content, walls thin and
smooth (Fig. 9). Hymenium present only on the sides of the fruitbody. Basidia
narrowly clavate, (54-) 75-94 x (7.7-) 10-12. 3/^ , mostly 4-spored but 2-spored
ones also present. Cystidia none. Subhymenium interwoven of hyaline to yel¬
lowish hyphae 1.5-3jU diam. Truncation sterile or with scattered basidia near the
margin; epicutis a hymeniform layer of yellowish cells of varied shapes: mostly
tubular and occasionally apically branched, 30-61.5 x 3-6 (-7.5)jU; but may also
be clavate to subventricose with rounded or subacute apices, 37-52 x 3.7-7.5M
(Fig. 10). Tramal hyphae hyaline to yellowish, smooth, 3-15 p diam. longitudi¬
nally parallel in the base and under the hymenium, interwoven in the center of
the fruitbody. Clamp connections present.

Material studied: WYOMING: Foxpark, Lovejoy 66, (Type) (RM).

Observations: Most authors have placed Fovejoy’s fungus in synonymy
with Clavariadelphus truncatus (Quel.) Donk, but we consider it quite separate
from that species because of its white spore deposit and the reddish fruitbodies
which are still distinctively colored in the dried state. Fovejoy, herself, was
aware of Fries’ Craterellus pistillaris which is now widely accepted as a synonym
of Clavariadelphus truncatus, but she maintained that in her collection of twenty
specimens found in entirely different localities, not one had the coloring of that
species.

The fact that the spores are of two sizes in C. lovejoyae is a deviation from
other species we have studied in this genus. We note that Doty (1944) also ob¬
served this phenomenon when he examined the type collection. It will require a
study of additional collections to determine whether or not this duo-spore size is
a constant caracter for C. lovejoyae.

Presently, we consider C. lovejoyae to be most closely related to C. borealis
but distinct from it by the reddish colored fruitbodies and the larger spores.

7. Clavariadelphus borealis Wells & Kempton, sp. nov. (Fig. 16).

Fructificatio 4.5-14 cm alta, 1.5-5 cm diam. ad apicem, simplex, subclavata mox turbi-
nata; apex truncatus, sterilis, luteo-aurantiacus;latera laevia demum longitudinaliter rugulosa,
obscure alutacea; basi pallens; contextus aibidus, firmus demum floccosus, sapor dulcis vel
amaro-dulcis similis saccharine. Sporae in cumulo albae, 9-11.3 x 4.7-6.6/i, late ellipticae.
Cystidia nulla. Fibulae praesentes. Dispersa, gregaria, vel raro subcespitosa in sylvis conif-
erae. TYPUS: Wells et Kempton 8/24/66-10, Fairbanks, Alaska (W/K).

Fruitbody 4.5-14 cm tall, 1.5-5 cm diam. at the apex, simple, obtusely cylin¬
drical to subclavate, broadening above to become turbinate with age, apex round¬
ed at first but soon truncate, usually with an obtuse, puckered ridge around the
margin of the truncation, disc sometimes perforated; primordia pallid, very soon
yellowish over the apex, finally yellow-orange on the apex and dark alutaceous of¬
ten with a tint of lilac on the sides, paler toward the base; moist to dry, unpolished,
smooth becoming longitudinally rugose near the apex, base white mycelioid and
with a fine network of white rhizomorphs; context white, unchanging or only
faintly pinkish to brownish when cut or bruised, firmly stuffed becoming floccose
in the upper part, odor none, taste sweet to sweetly bitter as of saccharine.

50

THE MICHIGAN BOTANIST

Vol. 7

Chemical reactions: Fresh material: FeS04— all parts pale green; KOH—
surface of apex and sides vivid pink to red, base and context negative; NH4OH—
same as KOH; phenol— all parts negative. Dried material: FeS04— all parts green;-
KOH— hymenium and surface of truncation pinkish red, context negative.

Spore deposit white. Spores 9-1 1.3 x 4. 7-6. 6ft, ellipsoid, hyaline with yel¬
low oil drops or granulose content, walls thin and smooth (Fig. 1 1). Hymenium
present only on the sides of the fruitbody. Basidia narrowly clavate, 82-105
(-1 16) x 9-1 1 (-13)fi, with sterigmata up to 10.7ft long, mostly 4-spored but some
2-spored ones usually also present. Cystidia none. Subhymenium interwoven of
hyaline to yellowish hyphae 1.5-3. 5 (4.6)ft diam. Truncation sterile or with
scattered basidia near the margins; epicutis of two general types with intermedi¬
ate forms: 1. A hymeniform layer of yellow to subhyaline, tubular to clavate
cells 22-48 x 4-22ft, or of fusoid-ventricose cells 41-86 x 11-1 8fi (Fig. 12); and
2. an interwoven layer of yellow, filamentous, flexuous hyphae 1.8-4. 6ft diam.
Tramal hyphae hyaline to yellowish, 2.8-12 (- 1 6)ft diam, walls smooth to ob¬
scurely roughened, thin to slightly thickened, longitudinally parallel in the base

Fig. 16. Clavariadelphus borealis, x 1. (W/K 8/20/67-20)

1968

THE MICHIGAN BOTANIST

51

and under the hymenium, interwoven in the center of the fruitbody ; oleiferous
hyphae sometimes present. Clamp connections present.

Scattered, gregarious, or rarely subcespitose in mixed, but predominantly
coniferous, woods, Aug.-Oct.

Material studied: (All collections are in W/K and were collected by Wells and Kemp-
ton unless otherwise stated.) ALASKA: Near Anchorage, 8/28/59-3, 8/24/64-4, 8/16/65-2,
9/13/65-9, 9/20/65-7, 8/15/66-13, 8/29/66-14, 8/29/66-15, 8/20/67-20, 9/6/67-6, 9/26/67-7.
Near Fairbanks, 8/17/64-2, 8/24/65-9, 8/26/65-16, 8/24/66-10 (Type). Chitina, 9/4/66-5.
Haines, 9/9/67-25. OREGON: Mt. Hood area: Still Creek, E. Wiebe 9/24/66-1, 10/7/66-1,
10/7/66-2; Hood River Meadows, 10/11/66-1.

Observations; The range of C. borealis in North America is not known at the
present time. A few collections of it were found in Oregon during the season of
1966, and although it is not actually common in Alaska, it can be found every
year in the south-central and interior regions. Because of the strong similarity of
this species to C. truncatus it is likely that collections without spore deposits
have been placed in herbaria under that name.

Section LIGULA Wells & Kempton, sect. nov.

Sporae anguste ellipticae, 3-5plo longiora quam latiora. TYPUS: Clavariadelphus ligula
(Fr.) Donk.

KEY TO THE SPECIES

1. Apices of fruitbodies sterile, truncate and the truncation usually with a
small, acute, central mucro; whitish at first, remaining pallid over the
truncation with the sides becoming alutaceous to brownish-flesh , . . 8. C. mucronatus

1. Apices of fruitbodies fertile, rounded to rugose, variously inflated or
laterally flattened to ligulate; fruitbodies yellowish at first becoming
buff to alutaceous, apices more-or-less concolorous or with tints of yel¬

low or green . 2

2. Spore deposit white; spores (10-) 12-15 (-18)/i long . 9. C. ligula

2. Spore deposit ochraceous; spores (15-) 16-24 (-28)/i long .... 10. C. sachalinensis

8. Clavariadelphus mucronatus Wells & Kempton, sp. nov.

Fructificatio (2-) 3-6 (-8) cm alta, (5-) 10-15 (-25) mm diam. ad apicem, simplex,
anguste clavata demum anguste turbinata; apex acutus autem mox truncatus, sterilis, ple-
rumque mucronatus, mucrone parvo, acuto, centrali; alba, immutabilis ad apicem, latera ful-
vescens vel ex brunneo incarnata, laevia demum longitudinaliter rugulosa; contextus albidus,
firmus demum floccosus, sapor indistinctus. Sporae in cumulo albae, (10. 5-) 11-15 (-16.5)
x 3.5-4.7|/, anguste ellipticae. Cystidia nulla. Fibulae praesentes. Gregaria in colonis in syl-
vis coniferae. TYPUS: Wells et Kempton 9/9/67-2, mile 40, Haines cut-off road, Alaska
(W/K).

Fruitbody (2-) 3-6 (-8) cm tall, (5-) 10-15 (-25) mm diam. at the apex,
simple, narrowly clavate and acute at the apex at first but soon truncate with
the truncation plane to shallowly depressed and typically with a small, acute,
central mucro, disc sometimes perforated with age; white or nearly white at first,
remaining white or pale colored over the truncation with the sides becoming tan

52

THE MICHIGAN BOTANIST

Vol. 7

to light brownish-flesh (sides “pale pinkish-buff 5 becoming “cinnamon-buff’ to
“vinaceous-buff ’), base more or less concolorous; surface smooth becoming lon¬
gitudinally rugulose on the sides, unpolished, base with scattered white strigosity
and with scanty white mycelial mold throughout the substratum; context white,
unchanging when cut or bruised, firm becoming floccose, sometimes hollowed in
the upper part, odor and taste not distinctive or slightly bitter to metallic.

Chemical reactions: Fresh material: FeS04— all parts greenish; KOH— all
parts negative; phenol— hymenium brownish, context negative. Dried material:
FeS04— all parts green; KOH— all parts negative.

Spore deposit white. Spores (10. 5-) 11-15 (16.5) x 3.5-4.7jit, narrowly el¬
lipsoid, straight or the longer ones slightly “sway-backed” in profile, hyaline
with yellow oil drops or granulose content, walls thin and smooth (Fig. 13). Hy¬
menium present only on the sides of the fruitbody. Basidia narrowly clavate,
49-75 x 5.3-7.5/Lt, with sterigmata up to 7.5/i long if 4-spored, up to 9.5ju on
some 2-spored basidia. Cystidia none. Subhymenium interwoven of hyaline to
yellowish hyphae 1.5-3.5jU diam. Truncation sterile; epicutis a thin layer of yel¬
lowish, interwoven, filamentous hyphae 2-3. 7ju diam. which become appressed
and very cohesive with age. Tramal hyphae hyaline to yellowish, (2.5-) 3-11.3/4
diam., walls smooth to obscurely roughened, thin to slightly thickened; hyphae
longitudinally parallel in the base and adjacent the hymenium, interwoven in the
center of the fruitbody. Clamp connections present.

Gregarious in colonies on coniferous debris, Sept.-Nov.

Material studied: ALASKA: Mile 40, Haines cut-off road, Wells & Kempton 9/9/67-2
(Type), 9/9/67-3 (W/K). OREGON: Still Creek, Mt. Hood Nat. Forest, Smith 28485 (MICH).
WASHINGTON: Spanaway Park, Mt. Rainier Nat. Park, Bigelow 49509 (MICH); Klikitat
Co., woods above Trout Lake, Wm. B. Cooke 9618, 9620 (CINC); Gov’t. Mineral Springs,
Thos. McGinnis 11/5/67-1 (W/K).

Observations: This species is closely related to C. ligula. According to
Doty (1944), who confused it with C. unicolor, it may occur at the same time
and in the same vicinity as C. ligula. Although we were unable to obtain Doty’s
collections for study, his description and comments are so decidedly in line with
our concept of C. mucronatus that we do not hesitate to refer the fungus he de¬
scribed in his work to this species.

C. mucronatus is readily distinguishable from C. ligula by its sterile, trun-
cate-mucronate apex and paler colors, and from C. unicolor by its narrower
spores, coniferous habitat, truncate-mucronate apex, different colors and gen¬
erally smaller size. It is true that in dried material large fruitbodies of C. mu¬
cronatus may bear some superficial resemblance to small fruitbodies of C. uni¬
color, but their microscopic characters are entirely different.

In habit and habitat, C. mucronatus is like both C. ligula and C. sacha-
linensis in that it grows densely gregarious on debris of a variety of coniferous
trees. Doty, (1944, as Clavaria unicolor ) reports it on needle debris of Picea
engelmanii and Pseudotsuga taxifolia. Our Alaskan collections were on debris
under Tsuga heterophylla with C. sachalinensis in the vicinity. Although C.
mucronatus is usually found in large numbers, it may be rare, and possibly also
confined to the northwestern part of North America.

1968

THE MICHIGAN BOTANIST

53

The Bigelow collection cited above (49509, MICH) was covered with conid-
iospores from a species determined by Dr. Clark T. Rogerson of the N. Y. Bo¬
tanical Garden as (personal communication) “a species of Penicillium belonging
to the A symmetrica section. . .probably secondary, not a parasite.” Doty (1944)
noted that the two W. Bridge Cooke collections from Washington were immature.
Although we were able to do a sufficient amount of microscopic work on these
collections, they are in rather poor condition, possibly the result of having been
dried in a plant press.

9. Clavariadelphus ligula (Fr.) Donk, Rev. Niederl. Homobas. Aphyll. 2:73.

1933.

Clavaria ligula Fr., Syst. Mycol. 1:477-478. 1821.

Fruitbody 2-5.5 (-7) cm tall, 3-12 (-25) mm diam. at the apex, simple, cyl¬
indrical to clavate often becoming irregularly flattened or subspathulate to lig-
ulate with age, apex subacute to obtuse becoming rugose to irregularly inflated;
“pale ochraceous-buff ’ to “pale salmon” or “vinaceous-buff,” apex concolorous
to slightly darker, typically unicolorous and near “chamois” when dried; smooth
becoming rugulose, base white tomentose to strigose with abundant white my¬
celium binding the substratum; context white, unstaining, firm becoming floc-
cose in the apex, odor none, taste none to slightly bitter or metallic.

Chemical reactions: Dried material: FeS04— all parts green; KOH— all
parts negative.

Spore deposit white. Spores (10-) 12-15 (-18) x (2.8-) 3-4.5 (-5)ft, narrow¬
ly ellipsoid, straight or the longer ones slightly “sway-backed” in profile, hyaline
with yellow oil drops or granulose content, walls thin and smooth (Fig. 14). Hy-
menium present on sides and apex of the fruitbody. Basidia narrowly clavate,
(40-) 50-75 (-86) x 6.5-9.5/i with sterigmata up to long, mostly 4-spored but

2- spored ones also frequently present. Cystidia none. Subhymenium interwoven
of hyaline to yellowish hyphae 2-3. 5jU diam. Tramal hyphae hyaline to yellowish,

3- 9. 5/i diam., walls smooth to obscurely roughened, thin to slightly thickened;
hyphae longitudinally parallel in the base and adjacent to the hymenium, inter¬
woven in the center of the fruitbody. Clamp connections present.

Gregarious to subcespitose in colonies on coniferous debris, July-Oct.
(-Nov.).

Material studied: United States: CALIFORNIA: Sumner, Lanphere 10460 (MICH).
IDAHO: Priest Lake, R. H. Petersen C1866, C1877, C1896, C1960 (RHP), Smith 71055
(MICH); Payette Lakes, Smith 45132 (MICH), Bigelow 46882 (MICH). MAINE: Guerette,
Bigelow 3706 (MICH); Madawaska Lake, Bigelow 3485 (MICH). MICHIGAN: Tahquame-
non Falls, Leathers 101, 444 (MICH); Carson City, V. Pfotter] 10250, 10260 (MICH). ORE¬
GON: Mt. Hood, Bear Springs, Ore. Mycol. Soc. 49495 (MICH). PENNSYLVANIA: Buck
Hill Falls, Delafield Sept. 20, 1919 (NCU). WASHINGTON: Stevens Co., Cooke 22875,
22876, 22877 (DAOM); Longmire, Smith 49370 (MICH). Canada: MANITOBA: Victoria
Beach, Bisby F7132 (DAOM), Buller and Bisby F7133 (DAOM). NOVA SCOTIA: Kent-
ville, Harrison 95816, 95902, 111220 (DAOM). ONTARIO: Bell’s Corners, Elliott 54287
(DAOM); Bond Lake, Bell 7421 (TENN); Tilsonburg, Dearness 15256 (TENN); Lake Tim-
agami, Smith 4864-A (MICH). QUEBEC: Stevenson 95815 (DAOM). Sweden: Uppland,
Melderis 65949 (DAOM); Helsingland, Romell 72062 (DAOM). Switzerland: Favre Sept.
12, 1946 (MICH).

54

THE MICHIGAN BOTANIST

Vol. 7

Observations: Corner (1950) states that C. ligula is little known. We be¬
lieve that part of the problem stems from the fact that many authors have con¬
sidered it to be merely a small version of C. pistillaris. It is, however, easily dis¬
tinguished from C. pistillaris by its narrower spores, coniferous woods habitat,
color, and lack of staining, as well as its generally smaller size.

The occurrence of C. ligula in North America may also have been confused
by Coker (1923) who published an illustration of a pink C. ligula and described it
as, “When fresh dull pink, soon fading to leather color or with tints of fawn or
buff added.” We admit that we have never seen C. ligula in fresh condition, but
our examination of the Buck Hill Falls collection illustrated shows that any pink
color which might have existed has disappeared with drying, or with age, as Cok¬
er indicated. Since the spores measure 12.3-16 x 3-4. 6jU, this collection now
agrees both macroscopically and microscopically with our concept of C. ligula.

Coker (1923) drew his microscopic data for C. ligula from a collection
taken in the Adirondacks, New York ( Murrill 70, NCU). He reported the spores
were long, narrow, 15-18.5 x 4.5-5jU, and that there were no clamp connections
present. Although this collection is now in poor condition with most of the hy-
menium gone, we were able to ascertain that clamp connections are present, and
the few spores seen measured 16.9-20 x 4.6-6M. The long spores indicate that
Murrill’s collection is probably C. sachalinensis rather than C. ligula.

We have not been able to distinguish C. ligula from C. sachalinensis in dried
collections without a microscopic examination and we do not know whether
they can be distinguished from one another in the field. However, no real prob¬
lem exists in differentiating them. In C. ligula, the spore deposit is white and the
spores are rarely over 15/i long; in C. sachalinensis, the spore deposit is ochrace-
ous and the spores are usually well over 15/i long.

The range of C. ligula in North America is not known but all the collections
we examined came from northern United States and southern Canada, which
may indicate that it is limited to a rather narrow geographical zone. In addition,
field notes included with collections always mentioned a coniferous habitat and
most often, specifically mentioned pine. We doubt that C. ligula is restricted to
pine debris, but the point is open for further study.

10. Clavariadelphus sachalinensis (Imai) Corner, Ann. Bot. Mem. 1:282. 1950.

(Fig. 17).

Clavaria sachalinensis Imai, Trans. Sapporo Nat. Hist. Soc. 11(2): 73.

1930.

Fruitbody 2-7 (-15) cm tall, 3-15 (-35) mm diam. at the apex, simple or oc¬
casionally briefly branched above, cylindrical to subclavate becoming clavate to
subventricose, apex acute to obtuse becoming rugose to irregularly inflated,
rarely perforated in extreme age; light yellow to buff becoming dull yellow to
alutaceous, apex concolorous or yellow-green to olive-green; unpolished, smooth
becoming rugulose, base strongly white pruinose to substrigose and with abun¬
dant white mycelium binding the substratum; context whitish, unstaining, firm
becoming floccose in the upper part, odor none, taste none or slightly bitter.

1968

THE MICHIGAN BOTANIST

55

Chemical reactions: Fresh material: FeS04— all parts green; KOH— all
parts negative; phenol— all parts negative to obscurely brownish. Dried material:
FeS04— all parts dark green; KOH— all parts negative.

Spore deposit ochraceous. Spores variable in length, (13-) 16-24 (-28) x
4-6jU, narrowly ellipsoid, straight in face view, usually curved and appearing
“sway-backed” in profile, hyaline with yellow oil drops, walls thin and smooth
(Fig. 15). Hymenium present on sides and apex of the fruitbody. Basidia nar¬
rowly clavate, (56-) 60-90 (-115) x (7.5-) 8.5-11 (-13)jU with sterigmata up to 9.2ju
long, mostly 4-spored but 2-spored ones usually also present. Cystidia none. Sub-
hymenium interwoven of hyaline to yellowish hyphae 2-4. 5/i diam. Tramal
hyphae hyaline to yellowish, 3. 5-7. 5 (-ll)jU diam., walls smooth to obscurely
roughened, thin to slightly thickened, longitudinally parallel in the base and un¬
der the hymenium, interwoven in the center of the fruitbody. Clamp connec¬
tions present.

Gregarious to densely gregarious in colonies on debris of coniferous trees,
July-Oct.

Material studied: United States: ALASKA: (All collections are in W/K and were
collected by Wells and Kempton.) Anchorage, 8/4/60-1, 9/12/60-4, 8/16/65-1, 9/25/65-4,
8/18/66-3; 1600 ft. elev., 9/10/64-9, 9/16/65-13, 8/11/66-3; Bird Creek, 8/31/59-6, 8/6/64-
2, 8/27/64-7,8/18/65-8,9/20/65-12,8/11/66-20, 8/18/66-1, 8/18/66-7, 8/29/66-6, 9/11/66-
1, 7/20/67-19, 8/2/67-10; Chitina, 9/4/66-6, 9/5/66-12; Fairbanks, 8/23/65-4, 8/23/65-5,

Fig. 17. Clavariadelphus sachalinensis, x 1. (W/K 8/16/65-1)

56

THE MICHIGAN BOTANIST

Vol. 7

8/24/65-8,8/24/65-12, 8/25/65-2, 8/26/65-1, 8/26/65-2, 8/26/65-3; Haines, 9/9/67-4, 9/10/
67-24; Homer, 9/2/67-4; Hope, 9/22/66-9, 9/22/66-10; Ingram Creek, 9/20/65-11; Skilak
Lake, 8/9/65-3, 7/25/65-4, 9/8/65-3, 9/8/65-4. COLORADO: Grand Lake, Paul & Vir¬
ginia Shope 15612 (TENN), 53207 (DAOM). IDAHO: Payette Lakes, Smith 58371 (MICH);
Priest River Exp. Forest, Smith 70821 (MICH); Upper Priest River, Smith 71489, 74177
(MICH); McCall, Smith 59172, 59785 (MICH); Nordman, Smith 54567 (MICH). MAINE:
Guerette, Bigelow 3071, 3271, 3447, 3606 (MICH); Smith 3596 (MICH). MICHIGAN:
Wilderness State Park, Thiers 3228 (MICH), Smith 63355 (MICH). NEW MEXICO: Moun¬
tains near Santa Fe, Barrows 234, 440, 487 (MICH). OREGON: Mt. Hood area, H. V. Smith
& R. Keene 10/8/66-1 (W/K), Petersen 10/8/66-2 (W/K), Wiebe 11/5/66-1 (W/K), Phelps
11/6/66-1 (W/K), Ore. Myc. Soc. 10/10/66-2, 10/10/66-3, 10/10/66-4 (W/K). Canada:
BRITISH COLUMBIA: Prince George, Ziller 21598 (DAOM); Crescent Spur, Engelhardt
30839 (DAOM); Silverton, Pole Blight Reserve, Ziller 35242 (DAOM). QUEBEC: St. Au-
bert, Groves & Jackson 16807 (DAOM); Baie St. Paul, Elliott & Kendrick 53550 (DAOM).

Observations: Corner (1950) correctly suspected that C. sachalinensis oc¬
curred in North America. In fact, we have found it to be widespread on this con¬
tinent and extremely common in Alaska. It is also rather variable but we have
not been able to correlate differing features such as size, color, taste, spore
length, etc. into any pattern dependable enough to divide the species into varie¬
ties or forms.

In addition to the taxonomic characters given for this species, we found a
curious feature which is so constant that it almost has taxonomic value— a paucity
of either mature or immature spores in mounts of sectioned material. It has often
been necessary to examine several fruitbodies from a single collection before any
spores could be found. This condition was occasionally evident in sectioned ma¬
terial of C. ligula, but never to the degree exhibited by C. sachalinensis.

Since C. sachalinensis is so common in Alaska, we undertook a simple field
project which we hoped would shed some light on the sporulation pattern and
the longevity of the fruitbodies. On August 11, 1966, we took a collection of
variously sized fruitbodies from the first colony to appear and tagged about 30
of the remaining 100-150 fruitbodies. Thereafter, we took collections every
7-10 days and compared tagged specimens from this colony with others in the
vicinity. The fruitbodies were examined while fresh before a deposit was at¬
tempted, examined again in the fresh condition after a deposit was obtained, and
a final examination of the same material was made after it had been dried. On
September 1 1, 1966, we took the last collections when signs of a light frost were
evident. The results were as follows:

1. Mature and immature spores were rare to absent in all examinations of sec¬
tioned material.

2. Heavy spore deposits were obtained from all but the very smallest fruitbodies
in the first sampling.

3. We found no correlation between the age of the fruitbody and the size and
shape of the spores:

1st week— 20.6-28 x 4. 2-5. 6 ft
2nd week— 17.8-24.4 x 4.7-6 jU
3rd week— 23.4-28 x 4. 7-5. 6 jit
4th week— 20.6-26 x 3.7-5.6jU

4. There was no correlation between basidial size and spore size; i. e. fruitbodies
with large basidia did not necessarily produce large spores.

1968

THE MICHIGAN BOTANIST

57

5. The hymenium did not thicken appreciably in depth with age.

6. Old fruitbodies frequently became spotted with brown, or browned from the
tips downward after a light frost.

7. None of the specimens were invaded by larvae or attacked by fungal parasites
though both were present on other fungi in the vicinity.

Although this study shed little light on the problem of why there is such a
dearth of spores in mounted sections of C. sachalinensis, we do know that this
species can produce spores constantly over a period of at least thirty days.

EXCLUDED SPECIES AND VARIETIES

Clavariadelphus fistulosus (Fr.) Corner. We have not seen authentic material of
this species.

Clavariadelphus junceus (Fr.) Corner. We have not seen sufficient material.

Clavariadelphus maricolus (Kauff.) Corner. According to Dr. A. H. Smith (per¬
sonal communication), the type collection is unsatisfactory for study.

Clavariadelphus pistillaris (Fr.) Donk var. americanus Corner. This variety was
based on literature and we have seen no specimens which could be referred to it.

LITERATURE CITED

Burt, E. A. 1914. The Thelephoraceae of North America. II. Craterellus. Ann. Missouri
Bot. Gard. 1:327-350.

Coker, W. C. 1923. The Clavarias of the United States and Canada. Univ. North Carolina
Press, Chapel Hill. 209 pp.

Coker, W. C. 1947. Further notes on clavarias, with several new species. Elisha Mitchell
Sci. Soc. 63: 43-67.

Corner, E. J. H. 1950. A Monograph of Clavaria and Allied Genera. Ann. Bot. Mem. 1.
740 pp.

Cotton, A. D., & E. M. Wakefield. 1919. A revision of the British Clavariae. Brit. Mycol.
Soc. Trans. 6: 164-198.

Donk, M. A. 1933. Revision der Niederltfndischen Homobasidiomycetae-Aphyllophoraceae
II. Meded. Bot. Mus. Utrecht 9. 278 pp.

Doty, M. S. 1944. Clavaria, the Species Known from Oregon and the Pacific Northwest.
Ore. Monogr. Bot. 7. 91 pp.

Doty, M. S. 1948. A preliminary key to the genera of clavarioid fungi. Bull. Chicago Acad.
8: 173-178.

Leathers, C. R. 1955. The genus Clavaria Fries in Michigan. Ph. D. thesis, Univ. of Mich.,
Ann Arbor. 344 pp.

Murrill, W. A. 1940. Craterellus unicolor Berk. & Rav. in Florida. Mycologia 32: 415-416.
Peck, C. H. 1899. New species of fungi. Bull. Torrey Bot. Club 26: 63-71.

Petersen, R. H. 1967. Notes on clavarioid fungi. VI. Two new species and notes on the
origin of Clavulina. Mycologia 59: 39-46.

Singer, R., & I. J. Gamundi. 1963. Paraphyses. Taxon 12: 147-150.

Wells, V. L., & P. E. Kempton. 1965. Togaria aurea in Alaska. Mycologia 57: 316-318.
Wells, V. L., & P. E. Kempton. 1967. Studies on the fleshy fungi of Alaska. I. Lloydia 30:
258-268.

58

THE MICHIGAN BOTANIST

Vol. 7

ODD AND UNUSUAL DISCOMYCETES FROM MICHIGAN, I

Nancy Jane Smith

Department of Botany, The University of Michigan, Ann Arbor

Among Michigan’s more unusual cup-fungi, Peziza proteana (Boud.) Seaver
is noteworthy because of the variability of the ascocarps in their general appear¬
ance. Two forms of this species are recognized: f. proteana in which the asco¬
carps have the typical cup-shaped appearance, and f. sparassoides (Boud.) Korf
(Mycologia 48: 714. 1956). In the later the individual ascocarps are typically
fused at their bases and are one-sided to spathulate; this gives the cluster the ap¬
pearance of a Sparassis- type fruit-body. The unusual morphology is mirrored in
its common name of “cabbage-head fungus.” For many years the relationship
between the two forms was not understood and the sparassoid form was placed
with the Tubers and the Helvellas.

According to Dr. Korf, the sparassoid form is considered to be an excellent
edible fungus by a number of collectors. Although this species has been found in
Michigan on several occasions and is known from many parts of the United
States and Europe, it is considered a rare fungus. On October 6, 1967, Florence
Hoseney found an excellent fruiting of this fungus containing both forms. This
fruiting occured where a diseased elm had been removed and the stump burned
at the University of Michigan Botanical Gardens. The following is a description
of Mrs. Hoseney’s collection.

Macroscopic features: f. proteana.

Apothecia densely gregarious on burnt ground, reaching 9 cm in diameter,
5 cm in depth, regularly cup-shaped, split on one side or sometimes petaloid;
hymenium often radially wrinkled from the area of attachment of the apotheci-
um, pallid to light brown or often lilac tinted; outer surface of the ascocarp con-
colorous with the hymenium or slightly darker except paler near the base, ap¬
pearing mealy in older specimens; margin often splitting in age, often darker in
color than the hymenium; consistency very fragile.

Macroscopic features: f. sparassoides (Fig. 1).

Clumps up to 20 cm in diameter; apothecia commonly split down one side,
fused at or near the base. Similar to the above in other respects.

Microscopic characters (the same for both forms):

Spores hyaline, unicellular, 2-guttulate, nearly oblong, often with apical
truncate projections, remainder of ornamentation composed of irregular ridges
and warts, staining fairly readily in cotton blue, 10.5-12 x 4.5-6 p.

Asci 8-spored, cylindric, operculate, 182-230 x 7.8-1 Oft; wall, especially
around the operculum, blue in Melzer’s reagent; paraphyses septate, 3-4/iat the
base, 4-lp wide at the apex, narrowly clavate.

1968

THE MICHIGAN BOTANIST

59

Outer layer of the apothecium composed of large more or less globose cells
and with some interwoven hyphae; middle layer of interwoven hyphae and in¬
terior to this a layer of inflated cells, the latter merging into the hymenium.

ACKNOWLEDGMENTS

The author wishes to acknowledge financial aid for her studies on Discomycetes in
the form of a National Science Foundation Graduate Fellowship. In addition she wishes to
thank A. H. Smith, Director of the University of Michigan Herbarium, for the privilege of
studying the collections and for the use of the photograph.

Fig. 1. Peziza proteana f. sparassoides, xVz. (Photo A. H. Smith)

60

THE MICHIGAN BOTANIST

Vol. 7

ASEXUAL REPRODUCTION IN THE BURNING BUSH,
EUONYMUS A TROPURPUREUS

Lytton J. Musselman
Department of Botany and Zoology,

University of Wisconsin, Rock County Campus, Janesville

The burning bush, Euonymus atropurpureus Jacquin (Celastraceae) is a
shrub or small tree found in various habitats throughout its range from Ontario
to Montana south to eastern Virginia, Alabama, and Tennessee (Fernald, 1950).

Fig. 1. Rhizome and aerial stems of Euonymus atropurpureus (Musselman 1611) in winter
condition, from Laona Township, Winnebago Co., Illinois (T29N, R11E). Arrow indicates
level of soil surface. Note heavy browse by rabbits on stem at extreme right.

1968

THE MICHIGAN BOTANIST

61

In southern Wisconsin and northern Illinois, it is most commonly found along
streams (Fell, 1955; Hartley, 1966). According to Curtis (1959), in Wisconsin
it reaches its highest frequency value in the southern wet forest community. I
have collected it in the Illinois-Wisconsin state line area in mesic woods, on a
dry exposed roadside (Musselman 1613), and in river bottom forests. Specimens
are deposited in the herbarium of the University of Wisconsin Rock County
Campus, Janesville, Wisconsin.

The genus Euonymus contains many species which reproduce asexually
from the stem. Bailey (1942) lists Euonymus obovatus, E. nanus, E. patens, and
E. radicans as rooting from the stem or spreading by runners. Braun (1961) re¬
cords the commonly planted European species Euonymus europaeus as escaping
and reproducing by runners from the stem. However, neither Bailey nor Braun
record any form of asexual reproduction for E. atropurpureus. McNair (1930)
failed to mention any form of asexual reproduction in E. atropurpureus in an
anatomical investigation of several species of Euonymus.

I have observed vigorous asexual reproduction by well developed rhizomes
in every stand of E. atropurpureus I have studied. The rhizomes arise 1 to 15
centimeters below the soil surface and grow horizontally producing numerous
small roots and occasional aerial stems. The rhizomes are white with a soft cor¬
tex that is easily peeled from the hard woody stele. The rhizomes may be up to
three centimeters in width and produce numerous buds in addition to the roots
and aerial stems.

Excavations in stands of E. atropurpureus in the Illinois-Wisconsin state
line area reveal a complex intertwining network of rhizomes suggesting that it is
possible that one stand could be a clone consisting of a single genotype arising
from a single seed. Such stands contain stems that are tallest and oldest in the
center with shorter, younger stems at the periphery indicating that the plants in
the center may have given rise to surrounding stems by rhizomes.

ACKNOWLEDGMENT

I wish to thank Mr. Steven Houge for photographic assistance.

LITERATURE CITED

Bailey, L. H. 1942. The Standard Cyclopedia of Horticulture. Macmillan Co., New York.
Vol. 1.

Braun, E. L. 1961. The Woody Plants of Ohio. Ohio State Univ. Press, Columbus. 362 pp.
Curtis, J. T. 1959. The Vegetation of Wisconsin. Univ. Wisconsin Press, Madison. 657 pp.
Fell, E. W. 1955. Flora of Winnebago County, Illinois. Nature Conservancy, Washington.
207 pp.

Fernald, M. L. 1950. Gray’s Manual of Botany, ed. 8. American Book Co., New York, lxiv
+ 1632 pp.

Hartley, T. G. 1966. The flora of the “Driftless Area.” Univ. Iowa Stud. Nat. Hist. 21(1):
1-174.

McNair, G. T. 1930. Comparative anatomy within the genus Euonymus. Univ. Kan. Sci.
Bull. 19: 221-272.

62

THE MICHIGAN BOTANIST

Vol. 7

SEASONAL CHANGE IN ASAND PRAIRIE IN
VAN BUREN COUNTY, MICHIGAN

Dan Pokora

C. C. Adams Center for Ecological Studies, Department of Biology,
Western Michigan University, Kalamazoo

Seasonal change was one of the first aspects of community organization to
be recognized and described, but few phenological reports have been published.
The information presented here is the result of a study of aspection on a sand
area dominated by prairie plants. The study took place in the spring, summer,
and fall of 1966.

STUDY AREA

The study area lies about two miles southwest of Mattawan, Van Buren
County, Michigan, in the SE lA of Sect. 22, R 13 W, T 3 S. It consists of a nar¬
row strip lying southeast of the New York Central Railroad, between Drape Ave.
and West Robinson Ave. A seldom-used wheel-track runs through the south¬
western half of the area. Total length of the area is about 4,000 feet and the
width is about 150 feet. Elevation is mostly 820-840 feet above mean sea level.
Part of the area is sand that may have been exposed in construction of the em¬
bankment on which the railroad is located; another part may have been an old
railroad bed. No major disturbance to the area sampled has taken place during
the three-year period in which the area has been intensively studied. Local resi¬
dents cannot remember any major fire or disturbance in the area for at least six
years.

METHODS

In June of 1965, sixteen four square meter plots were placed randomly
throughout the study area. Each plot was divided into 16 1 square meter quad¬
rats, thus making a total of 256 1 square meter quadrats. All vascular plants
rooted in the quadrats were noted so that frequencies could be determined.

The area was visited once a week in the early part of the study (starting
March 1, 1966) and at least twice a week during the middle and later part of the
study (May to Nov., 1966).

Following Ahshapanek’s method (1962), I departed from the traditional
method of dividing the year into four rigid three-month seasons. Instead, my
seasons were based on phenological data, especially that of the time of flowering.
Thus the seasons and dates are not static, but change from year to year depend¬
ing on the environmental conditions present. In 1966 I defined them as follows:
hiemal, Nov. 20-May 3; vernal, May 3-June 7; aestival, June 7-Aug. 25; and au¬
tumnal, Aug. 25-Nov. 20.

Three phenological stages were used in this study. They were flowering,
first appearance of fruit, and time of distribution of seed. At the time of flower¬
ing the heights of the plants were measured.

Of the sixty-one species so far found on the study area, the twenty-four spe¬
cies with the highest frequencies were given formal consideration.

1968

THE MICHIGAN BOTANIST

63

COMPOSITION OF VEGETATION

Five species, Andropogon scoparius, Helianthus occidentals, Poa com-
pressa, Rosa Carolina, and Panicum oligosanthes, were found to have a frequency
of 50 per cent or higher. Three species had frequencies between 40% and 50%.
They were Rutnex acetosella, Tradescantia ohiensis, and Panicum perlongum.
The other species in Table I had frequencies of 4.2 to 39%.

Application of Raunkiaer’s life form system indicated that the area was
dominated by hemicryptophytes (77%), followed by therophytes (11.4%), geo-
phytes (5.7%), and nanophanerophytes (5.7%). This classification is based on
Ennis’s list of life forms (1928).

WEATHER

Temperature and precipitation data were determined from daily records of
the U.S. Weather Bureau Station at Kalamazoo State Hospital, published in Cli¬
matological Summaries for Michigan, 1966. This weather station is about four¬
teen miles from the study area and thus only approximates the conditions there.

Precipitation in 1966 was about three inches above the long-term average
of approximately thirty-five inches and 1965 was about equally wet. January,
February , June, July, and September were months of below-average precipitation;
March, April, May, and August, months of above-average precipitation. Tem¬
peratures showed no major departures from normal. In 1966, the last spring
frost was May 10 (average, May 4) and the first autumn frost Oct. 6 (average,
Oct. 14).

SEASONAL CHANGE

The 24 most frequent species, their times of flowering, their average heights
during flowering, their times of fruiting, and their dates of seed dispersal can be
found in Table I.

The hiemal season ran from Nov. 20 to May 3. During this time most of
the activity of growth had ceased. The dried stems of Andropogon scoparius and
Andropogon gerardi were still standing in early November. In fact they were
sometimes present in the next summer’s growth. An occasional green rosette of
pussy-toes or some goldenrod could be seen. Frost tended to hasten the deterio¬
ration of vegetation. As Weaver (1958) discovered in the Iowa prairie, the living
population of this prairie was in a dormant state and was found almost entirely
within the protecting soil.

The vernal season ran from May 3 to June 7. None of the 24 species of
plants was found in flower in the early part of May and the prairie retained its
characteristic hiemal appearance except for a few green leaves of bluestem
( Andropogon scoparius and Andropogon gerardi ), Poa compressa, Poa pratensis,
Fragaria virginiana, and Phlox pilosa. During the latter part of the month, two of
the 24 species studied were in flower. They were Rumex acetosella and Poa
pratensis.

Early June found four species of plants in flower. They were Fragaria
virginiana, Poa compressa, Rosa Carolina, and Phlox pilosa. Fragaria virginiana
was one of the first species to bloom. Its white flowers could be seen sporadically

64

THE MICHIGAN BOTANIST

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throughout the prairie. Poa pratensis, which flowered during the vernal season,
and Poa compressa were now taller than all the other species. They tended to be
quite conspicuous throughout the growing season. By late July Poa pratensis had
turned brown and remained this way until early November. This was not true of
Poa compressa which stayed green until October.

TABLE I. List in order of their appearing in flower, their date of flowering, height at time
of flowering, and date of seed dispersal for species on a sand area in Van Buren County.
Note: number in parentheses indicates number of plants used to determine avg. height.

Species

Date first
found in
flower

Avg. height of
plant at time
of flowering

Date of first
seed forma¬
tion

Date of first
seed dis¬
persal

Rumex acetosella

May 23

12.6 cm (10)

June 4

July 1

Poa pratensis

May 23

27.2 cm (10)

June 17

July 1

Rosa Carolina

June 4

21.5 cm (10)

July 17

Poa compressa

June 4

17.4 cm (10)

June 29

July 17

Fragaria virginiana

June 4

11.3 cm (10)

June 17

July 1

Phlox pilosa

June 4

22.0 cm (10)

June 29

July 15

Tradescantia ohiensis

June 10

37.8 cm (10)

July 1

July 17

Euphorbia corollata

June 17

19.4 cm (10)

July 17

July 21

Melilotus officinalis

June 17

71.1 cm (10)

June 29

July 1

Cyperus schweinitzii

June 17

27.5 cm (10)

July 15

July 17

Specularia perfoliata

June 27

28.3 cm ( 5)

July 1

July 1

Panicum perlongum

July 1

18.7 cm (10)

July 17

July 29

Panicum oligosanthes

June 29

32.1 cm (10)

July 15

Aug. 1

Erigeron strigosus

July 1

44.7 cm (10)

July 17

Hypericum perforatum

July 1

21.7 cm (10)

July 21

Aug. 14

Oenothera rhombipetala

July 15

32.5 cm (10)

Aug. 1

Aug. 14

Aster azureus

July 15

82.0 cm ( 5)

Aug. 3

Helianthus occidentalis

July 21

90.0 cm ( 5)

Aug. 14

Solid ago speciosa

July 21

80.0 cm (10)

Aug. 14

Artemisia caudata

July 24

88.3 cm ( 5)

Aug. 1

Andropogon scoparius

Andropogon gerardi

Aug. 14

Aug. 14

78.8 cm ( 5)

120.0 cm (10)

Aug. 14

Aug. 3

Aug. 14

Rubus sp.

Pteridium aquilinum

July 15

1968

THE MICHIGAN BOTANIST

65

The aestival season was by far the most active growing period on the area.
It began about June 7 and ran till about August 25. It was during the aestival
period that many of the characteristic prairie species were seen. The first con¬
spicuous flower to be seen in the early part of this period was spiderwort ( Tra -
descantia ohiensis ). The next two species to appear in the prairie during the
aestival season were Euphorbia corollata and Specularia perfoliata. Euphorbia
corollata or flowering spurge stayed in flower throughout the aestival season,
often bearing fruit and flowers simultaneously.

Toward the end of June and the beginning of July the introduced weed
Hypericum perforatum began to bloom. The middle of July saw Panicum oligos-
anthes and Aster azureus in flower. With the addition of these species to the list,
the community now had many of the characteristics of a relic prairie. In late
July one Solidago and Helianthus occidentals appeared in flower.

During August the last massive growing stage took place on the prairie with
the full growth of Andropogon scoparius and Andropogon gerardi. These were
two of the last species to flower and finished their life cycles in August.

The autumnal season from Aug. 25 to Nov. 20 saw the plants of the prairie
reach the end of their growing season, disperse their seeds, and collapse to rest
and wait for the next yearly cycle to begin. Plants that dispersed their seeds at
this time were Oenothera rhombipetala, Hypericum perforatum, and Panicum
oligosanthes.

STRATIFICATION

In any community there is a continual fight for survival among its species.
The prairie is no exception, even though the observer may see an orderly se¬
quence of events as he watches the prairie throughout the year. Even though
there is this continual fight for survival on the prairie there is also an example of
co-existence with a large number of plants making complementary use of the
environment. In this area, as in prairie areas previously studied (Weaver and Fitz¬
patrick, 1934; Curtis, 1959; Brewer, 1965), the smaller plants flower in early
spring and early summer before the larger plants flourish and take the majority
of the sunlight and nutrients. This form of stratification would also be favor¬
able to pollination both by wind and insects. It is interesting to note that those
plants which deviate most from this trend are weeds and not prairie species
(Table I).

ACKNOWLEDGMENTS

I am grateful to Richard Brewer for his guidance and suggestions and for his critical
reading of the manuscript and to Richard W. Pippen for his help in verifying some plants
and for making the Hanes Herbarium at Western Michigan University accessible to me.
Edward G. Voss also aided in plant identification. Plant names follow Gleason (1952).

LITERATURE CITED

Ahshapanek, D. 1962. Phenology of a native tail-grass prairie in central Oklahoma. Ecology
43: 135-138.

Brewer, R. 1965. Vegetational features of a wet prairie in southwestern Michigan. Occas.
Papers Adams Ctr. Ecol. Studies 13: 1-16.

66

THE MICHIGAN BOTANIST

Vol. 7

Curtis, J. T. 1959. The Vegetation of Wisconsin. Univ. Wisconsin Press, Madison, 657 pp.
Ennis, B. 1928. The Life Forms of Connecticut Plants and their Significance in Relation to
Climate. Conn. Geol. Nat. Hist. Surv. Bull. 43. 100 pp.

Gleason, H. A. 1952. The New Britton and Brown Illustrated Flora of Northeastern United
States and Adjacent Canada. New York. 3 vol.

Weaver, J. E., & T. J. Fitzpatrick. 1934. The prairie. Ecol. Monogr. 4: 110-295.

Weaver, J. E. 1958. Native grassland of southwestern Iowa. Ecology 39: 733-750.

ANNOUNCEMENT

SOME COMMON MUSHROOMS OF MICHIGAN'S
PARKS AND RECREATION AREAS

By Alexander H. Smith & Helen V. Smith

Special Publication No. 1, Michigan Botanical Club
Third Printing (1968) available soon.

This popular pocket-size publication (80 pp., 55 photos) is
available for $1.25 including postage and handling from Miss Barbara
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Almost 8,000 copies have been sold to date.

Proceeds from the sale of this non-technical guide to common
and conspicuous mushrooms of our vacation areas are being used for
the purchase of plant sanctuaries.

1968

THE MICHIGAN BOTANIST

67

THE DUCKWEEDS OF MINNESOTA

Howard L. Clark 1 and John W. Thieret
Department of Biology,

University of Southwestern Louisiana, Lafayette
INTRODUCTION

The duckweeds (Lemnaceae) contribute an often considerable and con¬
spicuous part of the aquatic vegetation .of Minnesota, a state (“The Land of
10,000 Lakes”) with about 4000 square miles of water area. Duckweeds occur
throughout the state, sometimes in such incredible myriads as to suggest that
numerically they are the most abundant of Minnesota’s higher plants. Hicks
(1937) estimated that “Compact floating mats of duckweeds. . .contain from
100,000 to 200,000 plants per square yard of water surface (genus Spirodela ),
300,000 to 800,000 per square yard (genus Lemna), and 1,000,000 to 2,000,000
per square yard (genus Wolffia)” Lrom these figures we estimate that, in a
Wolffia-covered pond 50 feet wide which we saw near Baxter, Cass County,
Minnesota, there were about 315,000,000 individual Wolffia plants, a figure
about IVi times the population of the United States.

The materials upon which our study is based are from two sources:
(1) exsiccata from the herbaria of the University of Minnesota, Minneapolis
(MIN) and Duluth (DUL), and the University of California, Berkeley (UC),
totalling 244 sheets; and (2) our 127 Minnesota collections preserved in 50%
methanol (LAL). In many of the herbarium specimens there was more than one
species of duckweed per collection; a few included all five members of the
Lemnaceae known from Minnesota. Most of our own collections contained
more than one species. Our collections, from 41 of Minnesota’s 87 counties,
formed the basis for about 40% of the dots on our distribution maps (figs. 7, 8,
9, 10). A complete list of exsiccata and pickled specimens examined by us is
available upon request.

During July and August 1967, while at the University of Minnesota Lorest-
ry and Biological Station at Lake Itasca,* 2 we made, separately, several duckweed¬
collecting trips that totalled about 2700 miles and that took us through a con¬
siderable portion of the state. Some areas proved to be good collecting sites;
others, especially the northwest and the northeast, were the source of relatively
few specimens.

LLOWERING OL DUCKWEEDS

Although it has become axiomatically accepted by many authors that
most Lemnaceae very rarely flower, some never, because of the family’s phylo¬
genetic loss of morphological complexity and their adeptness at vegetative repro¬
duction, we feel that this “axiom” is questionable. About 8% of the collections
examined by us contained fertile plants. Although this seems like a small
percentage, it must be taken into account that these collections were made, in

IPresent address: Department of Botany, Duke University, Durham, North Carolina.

2During this time, HLC held a stipend for summer research under NSF grant GB3390.

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respect to their fertility, randomly. Is it not possible that the flowers of most
duckweed species are as seasonal and ephemeral as are those of many other angio-
sperms, as for example maples or oaks or buttercups or catalpas? Consider, if
you will, what percentage of oak specimens, collected at random times of the
year, would be in flower. We know of no one who would limit his collection of
Lemnaceae to fertile plants only, as is usually the case with most other angio-
sperms. Duckweed flowers are, to say the least, not the most conspicuous and
showy, but they can often be found with diligent searching— equivalent to that
necessary to find the first trillium of the season as the winter snows have melted.
Schleiden (1839) observed, for three consecutive years, the annual production of
flowers of three species of Lemna in the vicinity of Berlin and concluded that
(translation ours), “After my observations, I must maintain that these three spe¬
cies of Lemna flower at least as often as other phanerogams and that it is solely
the collector at fault [“. . .es nur Schuld der Suchenden ist. . .”] because flowers
have been so seldom found.”

It is also entirely possible that some species that have only rarely been
found fertile, such as Spirodela polyrhiza, flower only in response to extreme
environmental conditions rather than to photoperiodic and other seasonal changes.
The excellent studies by Hillman (1959a, 1959b), Kandeler (1955, 1963, 1964,
1966), Landolt (1957), and others of the physiology and experimental control
of flowering in Lemna are now leading to an understanding of these processes,
not only in Lemnaceae but in other plants as well. By application of the results
of these studies of plants in carefully controlled environments, perhaps more
can be learned of the phenology, such as it may be, of duckweeds in their natu¬
ral habitats in Minnesota.

SYSTEMATIC SECTION

The duckweed flora of Minnesota consists of five species in three genera:
Spirodela polyrhiza, Lemna minor, L. trisulca, Wolffia Columbiana, and W. punc¬
tata. Three additional species have been reported to occur in the state: Lemna
gibba (Daubs, 1965), L. obscura (Daubs, 1965), andL. perpusilla (Gleason, 1952;
Gleason & Cronquist, 1963; Lakela, 1965; Moore and Tryon, 1964; Moyle &
Hotchkiss, 1945; Muenscher, 1944). We exclude these from the state’s flora;
they are discussed below under L. minor.

As an aid in the understanding of various terms used in the keys and de¬
scriptions, we have included, in place of a glossary and as fig. 6, a labelled dia¬
grammatic drawing of a duckweed. (Throughout this paper the term “frond” is
used to refer to a single segment of a duckweed plant.)

1. Frond with 1-many roots, much flattened, only the lenticular turions
rootless; reproductive pouches 2

2. Frond with 2-many roots and 4-many nerves, broadly obovate to

ovate . SPIRODELA POLYRHIZA

2. Frond with 1 root and 1-3 nerves

3. Margin of frond entire; nerves 3; upper surface of frond emersed;

fronds not in chainlike clones . LEMNA MINOR

3. Margin of frond dentate, serrate, or erose; nerve 1, with occa¬
sionally 2 obscure lateral nerves; frond submersed or, only when
fertile, most of the upper surface emersed; fronds usually in chain¬
like clones . LEMNA TRISULCA

1968

THE MICHIGAN BOTANIST

69

1. Frond without roots, spherical to hemispherical or ellipsoid; reproduc¬
tive pouch 1

4. Upper surface of frond convex, or a small portion slightly flattened
and roughened with several very minute central papillae; apex blunt,
not raised; frond not punctate; cells of frond uniformly inflated . . .

. WOLFFIA COLUMBIANA

4. Upper surface of frond smooth, flattened, with a raised acute apex
(best seen in side view); frond usually brown-punctate, especially
when dead; cells inflated in lower portion of frond, progressively
smaller and more compactly arranged toward upper surface. .WOLFFIA PUNCTATA

SPIRODELA

1. Spirodela polyrhiza (L.) Schleiden (figs, la, lb, 7)

Fronds solitary, or few attached by short deciduous stipes, usually 3-7 mm
long, 3-6 mm wide, occasionally somewhat larger, broadly obovate to almost
ovate, asymmetric, basal portion usually somewhat depressed dorsally and flatter
than the rest of the frond; dorsal surface glossy dark green to medium green, of¬
ten suffused with red and with a prominent red dot at the centrum, from which
radiate 5-16 nerves, margin red; ventral surface reddish purple to pale red; roots
3 or, more often, numerous.

Flowers have very rarely been observed on plants of this species. We have
found flowers on one of our Minnesota collections (Clearwater Co., small pond
just west of Lake Itasca P. 0. on Minn. 31 to Zerkel, Vi mile east of the Missis¬
sippi River, 23 July 1967, Clark and Williams 2025, LAF). The flowers in this
collection were invariably borne in the right-hand pouch of the third frond of a
clone of four fronds, the first being a germinated turion. No fruit could be found
in subsequent collections from the site. This species overwinters and survives
periods of drouth usually by the production of small, starch-filled, rootless
fronds— the turions (fig. la) (the “winter buds” or “hibernacula” of some auth¬
ors). When conditions are again favorable for growth, as in the spring, each
turion expels a small bubble of gas that causes it to rise to the surface of the wa¬
ter, where it germinates quite rapidly. Jacobs (1947) gives an excellent account
of Spirodela polyrhiza turions based upon his Ph.D. work at the University of
Minnesota.

Spirodela punctata Meyer (inch S. oligorrhiza (Kurz) Hegelm.) can reason¬
ably be expected to appear in the Minnesota flora, since this primarily Asiatic
species has been collected in widely separated localities in both the United States
and Canada (Daubs, 1965; McClure, 1964, p. 5). Its first observed United States
introduction was in Missouri in 1930. It is distinguished from S. polyrhiza by the
presence of 2-5 roots (rarely 1), narrowly obovate shape, absence of a red dot on
the upper surface, and generally smaller size. It differs from Lemna minor also
by having a red dorsal upper margin and being generally more asymmetric.

LEMNA

2. Lemna minor L. (figs. 2a, 2b, 8)

Syn.: L. obscura (Austin) Daubs; L. perpusilla of Minnesota references, not
Torr.

Fronds solitary, or few attached by short deciduous stipes, the stipes hyaline
and not appearing continuous with the dorsal surface; fronds elliptic to obovate,

70

THE MICHIGAN BOTANIST

Vol. 7

Fig. la. Spirodela polyrhiza, dorsal view, clone of two fronds.

Fig. lb. Spirodela polyrhiza, dorsal view, germinating turion.

Fig. 2a. Lemna minor, dorsal view, clone of two fronds.

Fig. 2b. Lemna minor, ventral view, germinating turion.

Fig. 3a. Lemna trisulca, dorsal view, clone of several tronds.

Fig. 3b. Lemna trisulca, dorsal view, clone of two tronds, the larger, with a spathe and flow¬
ers produced from the left-hand reproductive pouch.

(The vertical lines represent a length of 1 mm.)

1968

THE MICHIGAN BOTANIST

71

symmetrical to slightly asymmetrical, more asymmetrical when fertile, 2-4 mm
long, 1.1-2. 5 mm wide, green to pale green, often suffused with red, margins
very narrow, hyaline; nerves 3, somewhat obscure, root 1.

This species is quite variable and exhibits a large diversity of vegetative
forms, most of which are ecological or seasonal in origin.

Lemna gibba L. has been reported from Minnesota by Daubs (1965) on the
basis of a collection {Moyle 2331; cited incorrectly as 2321 by Daubs) made in
Douglas County (specimen in herb. UC; duplicate at the Lake Itasca Biological
Station). This collection, in flower and fruit, has been examined by us and is
clearly L. minor, as the seeds are solitary and shallowly ribbed and the fruit is not
winged. The seeds and fruit in this collection are typical of all fertile specimens
of L. minor examined by us. Lemna gibba normally has 2-6 seeds (rarely 1) per
fruit, the seeds are distinctly ribbed, and the fruit is winged.

Lemna obscura is being placed in synonymy by us, as we find that the
characteristics which are supposed to distinguish this species from L. minor (i.e.,
smaller size, amount of convexity, amount of pigmentation, and relative promi¬
nence of air spaces) intergrade freely in natural habitats. These characteristics
also have been observed to vary greatly in plants in axenic and other cultures
under a wide range of controlled environmental conditions (Landoldt, 1957;
McClure, 1964; Harrison, 1964; Spooner, 1967; and others). We found no fruit,
even on fronds that compared well vegetatively with descriptions of L. obscura,
definitely having the “broad shoulders” that are also, according to Daubs, defini¬
tive for this species. We have also attempted to examine the only fruiting collec¬
tion of L. obscura (Florida, Godfrey 55745, UC and FSU) cited by Daubs, but
the curators of the herbaria cited were not able to locate the specimens. Clark
(1968) examined many Louisiana specimens of Lemna which, vegetatively, fit
well Daubs’ description of L. obscura, a “species” that Daubs credits to Louisiana.
However, the fruits and seeds of these specimens resemble those of L. minor.
Thus we have been unable to verify Daubs’ assertion that L. obscura is distinguish¬
able from L. minor by its fruits and seeds. The senior author has found, during
the preparation (in collaboration with Gene J. Gonsoulin) of a paper on the fruits
and seeds of North American Lemnaceae, that these structures are more variable
than published reports would indicate.

Lemna minor and L. obscura can reportedly be separated on the basis of
their comparative flavonoid chemistry (McClure, 1964), but even this technique
may not be completely reliable. Ball, Beal, and Flecker (1967) have recently
shown that chromatographic pattern variation in two species of Spirodela can be
induced by controlled variation of culture conditions.

Careful examination of all available Minnesota Lemna collections, includ¬
ing many identified as L. perpusilla, and abundant living material of Minnesota
duckweeds, has convinced us that L. perpusilla, although reported to be a part of
the Minnesota flora, does not occur in the state. All Minnesota specimens that
we have seen identified as L. perpusilla have been annotated by us as L. minor.
Those authors who have ascribed L. perpusilla to Minnesota have used charac¬
teristics of the root cap to distinguish this species among Minnesota entire-
margined Lemna. The shape of the root cap in these plants is highly variable and
is, in our opinion, a feature of little taxonomic value. Lemna, in axenic culture

72

THE MICHIGAN BOTANIST

Vol. 7

STIPE

REPRODUCTIVE POUCH

CENTRUM REGION

ROOT SHEATH
ROOT CAP
ROOT

LATERAL NERVE
■CENTRAL NERVE

■APEX

Fig. 4a. Wolffia columbiana, dorsal view, clone of two tronds.

Fig. 4b. Wolffia columbiana, lateral view, clone of two tronds.

Fig. 5a. Wolffia punctata, dorsal view, clone of two fronds.

Fig. 5b. Wolffia punctata, lateral view, clone of two fronds.

(The horizontal lines represent a length ot 1 mm.)

Fig. 6. Diagrammatic drawing of a single duckweed frond.

1968

THE MICHIGAN BOTANIST

73

(McClure, 1964, p. 57, sub L. obscura), produced root caps that varied “from ob¬
tuse to acute even within a single clonal culture.” True L. perpusilla is clearly
distinguished from L. minor by lateral wings on the root sheath, distinct apical
and usually central dorsal papillae, and clear differences in flower and fruit, the
spathe being open and much reduced and the seed narrow and strongly ribbed.
The spathe of L. minor almost completely surrounds the immature flowers, being
irregularly ruptured at its apex by the developing flowers, and the seed is smooth
to very shallowly ribbed. Lemna perpusilla produces flowers and fruits more
commonly than most other species of North American Lemnaceae, so these char¬
acteristics are usually not difficult to observe when the species is collected.

3. Lemna trisulca L. (figs. 3a, 3b, 9)

Fronds usually 3-30, attached in chainlike clones, each frond with a stipe
up to 14 mm long, the stipe continuous with the body of the frond; fronds sub¬
mersed except when in flower, 4-10 mm long (not including the stipe), 1.4-4 mm
wide, pale green or green, occasionally reddish; apical margin thin, dentate, ser¬
rate, or sometimes erose; central nerve 1, continuous through the centrum into
the stipe, occasionally 2 obscure lateral nerves in robust fronds; root 1, often
deciduous.

Identification, particularly of dried specimens, can be somewhat more dif¬
ficult when the plants are in flower. In flowering condition, the fronds are short¬
er, thicker, and suffused with red, and float on the surface of the water. These
fertile fronds (fig. 3b) may be distinguished from L. minor by the non-entire
margins, by the blunt continuous stipe (the stipe of L. minor being pointed and
well differentiated from the frond), and by the more lateral position of the re¬
productive pouches. Flowers are common but not abundant on Minnesota speci¬
mens of L. trisulca; fruits are uncommon.

WOFFFIA

4. Wolffia Columbiana Karsten (figs. 4a, 4b, 10)

Fronds globose to broadly ellipsoid, 0.7-1. 4 mm long, truncate at the soli¬
tary reproductive pouch; dorsal surface convex, often with a few minute papillae;
frond floating at the surface of the water, with only a small portion of the dorsal
surface of the frond emersed, no clear distinction between dorsal surface and
sides; cells not punctate; inner cells of frond uniformly large and somewhat in¬
flated.

Several Minnesota collections have been found in flower, but no fruits
were present in these. Seeds evidently belonging to Wolffia sp. were found in
one collection (Blue Earth Co., Cottonwood Fake, Medo Twp., 18 Sept. 1958,
Moore and Bernardini 24307, MIN), but no flowers or attached fruits could be
found on the plants.

5. Wolffia punctata Grisebach (figs. 5a, 5b, 10)

Fronds ellipsoid to narrowly ellipsoid, 0.6-1. 2 mm long, truncate at the
solitary reproductive pouch; dorsal surface smooth, flat or slightly convex, with

74

Vol. 7

THE MICHIGAN BOTANIST

Fig. 7. Spirodela polyrhiza, documented distribution in Minnesota.

Fig. 8. Lemna minor, documented distribution in Minnesota.

Fig. 9. Lemna trisulca, documented distribution in Minnesota.

Pig. 10. Wolffia, documented distribution in Minnesota: circles— W. Columbiana, triangles—
W. columbiana and W. punctata together.

1968

THE MICHIGAN BOTANIST

75

a raised acute apex; frond floating at the surface of the water with the entire,
flattened, dorsal surface emersed, and with a clear distinction between this sur¬
face and the sides; cells usually uniformly punctate; inner cells of frond pro¬
gressively smaller and less inflated from ventral to dorsal surface, causing the up¬
per 1/3 of the frond to be quite darker than the lower portion.

This species has not yet been reported in flower in Minnesota. We saw no
collection that consisted of W. punctata alone; in every collection of it at least
one other duckweed— W. Columbiana— was present. In contrast, several collec¬
tions contained W. Columbiana as the only representative of Wolffia.

ACKNOWLEDGMENTS

We are grateful to Dr. Gerald B. Ownbey and Dr. Paul H. Monson for loan of speci¬
mens; to Dr. Ernest O. Beal for several valuable suggestions; and to Mr. Gene J. Gonsoulin
for assistance with the illustrations.

LITERATURE CITED

Ball, G. A., Jr., E. O. Beal, & E. A. Flecker. 1957. Variation of chromatographic spot pat¬
terns of two species of clonal plants grown under controlled environmental conditions.
Brittonia 19: 273-279.

Clark, Howard L. 1968. The Lemnaceae of Louisiana. M.S. thesis, Univ. Southwestern
Louisiana.

Daubs, Edwin Horace. 1965. A Monograph of Lemnaceae. Illinois Biol. Monogr. 34. 118 pp.
Gleason, Henry A. 1952. The New Britton and Brown Illustrated Flora of the Northeast¬
ern United States and Adjacent Canada. Vol. 1. New York Botanical Garden, New York.
482 pp.

Gleason, Henry A., & Arthur Cronquist. 1963. Manual of Vascular Plants of Northeastern
United States and Adjacent Canada. Van Nostrand, Princeton N. J. 810 pp.

Harrison, Dale E. 1964. The Taxonomic Significance of the Effect of Nutrient Media, Pho¬
toperiod, and Light Intensity on the Morphological Features of the Genus Spirodela
Schleid. M.S. thesis, North Carolina State Univ.

Hicks, L. E. 1937. The Lemnaceae of Indiana. Am. Midi. Nat. 18: 774-789.

Hillman, W. S. 1959a. Experimental control of flowering in Lemna. I. General methods.

Photoperiodism in L. pepusilla [sic] 6746. Am. Jour. Bot. 46: 466-473.

Hillman, W. S. 1959b. Experimental control of flowering in Lemna. II. Some effects of
medium composition, chelating agents and high temperatures on flowering in L. perpusilla
6746. Am. Jour. Bot. 46: 489-495.

Jacobs, Don L. 1947. An ecological life-history of Spirodela polyrhiza (greater duckweed)
with emphasis on the turion phase. Ecol. Monogr. 17: 437-469.

Kandeler, R. 1955. Uber die Bliitenbildung bei Lemna gibba L. I. Kulturbedingungen und
Tageslangenabhangigkeit. Zeitschr. Bot. 43: 61-71.

Kandeler, R. 1963. Die Aufhebung der photoperiodischen Steuerung bei Lemna gibba. Ber.
Deutsch. Bot. Ges. 75: 431-442.

Kandeler, R. 1964. Zweifache Wirkung von Bikarbonat auf die Lichtsteuerung der Bliiten-
bildung von Lemna gibba. Ber. Deutsch. Bot. Ges. 77: 140-142.

Kandeler, R. 1966. Trennung zweier Dunkelrotwirkungen bei der Lichtsteuerung der
Sprossvermehrung von Lemna gibba. Zeitschr. Pflanzenphysiol. 54:161-173.

Lakela, Olga. 1965. A Flora of Northeastern Minnesota. Univ. Minn. Press, Minneapolis.
541 pp.

Landoldt, E. 1957. Physiologische und okologische Untersuchungen an Lemnaceen. Ber.
Schweiz. Bot. Ges. 67: 271-410.

McClure, Jerry Weldon. 1964. Taxonomic Significance of the Flavonoid Chemistry and
the Morphology of the Lemnaceae in Axenic Culture. Ph.D. thesis, Univ. Texas.

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Moore, John W., & R. M. Tryon, Jr. 1946. A Preliminary Check List of the Flowering
Plants, Ferns and Fern Allies of Minnesota. Department of Botany, University of
Minnesota, Minneapolis. 94 pp. [mimeographed]

Moyle, John B., & Neil Hotchkiss. 1945. The Aquatic and Marsh Vegetation of Minnesota
and its Value to Waterfowl. Minn. Dep. Conserv. Tech. Bull. 3. 122 pp.

Muenscher, Walter Conrad. 1944. Aquatic Plants of the United States. Comstock Publ. Co.,
Ithaca, N. Y. 374 pp.

Schleiden, M. J. 1839. Prodromus monographiae lemnacearum oder conspectus generum
atque specierum. Linnaea 13: 385-392.

Spooner, William Edward. 1967. Local Population Variation in Spirodela polyrhiza (L.)
Schleid. (Lemnaceae). M.S. thesis, North Carolina State Univ.

Publications of Interest

THE THIRD WAVE. . .America’s New Conservation. U. S. Department of the Interior Con¬
servation Yearbook No. 3. [1967?] 128 pp. $2.00 (Supt. of Documents, Government

Printing Office, Washington 20402). While the Department of the Interior Yearbooks
will never seriously rival Sierra Club publications, this is a colorful and significant report
on federal conservation activities— despite the inevitable political overtones. The color pic¬
ture essay “Man Was Here” is particularly moving, in a depressing sort of way. Michigan
readers may regret the omission of Isle Royale from the color essay “A Parade of National
Parks,” but there are several photos from the Kalamazoo Nature Center in the concluding
chapter.

OUTDOORS USA. The Yearbook of Agriculture 1967. 408 pp. Advertised by the Depart¬
ment of Agriculture as “Big as all outdoors in scope, this is a handbook of conservation,
consumer’s guide to outdoor recreation, and primer on beautification.” The aim of the
Interior and Agriculture yearbooks is clearly to place the federal government on the
side of the angels regarding conservation. Botanical Club members will find much of inter¬
est in this volume, even though many would prefer a little less stress on “recreation” and a
little more on wild lands. Available for $2.75 from the Superintendent of Documents
(Washington 20402) or without charge from one’s congressmen.

BIBLIOGRAPHY & NATURAL HISTORY. Essays Presented at a Conference Convened in
June 1964 by Thomas R. Buckman. University of Kansas Publications Library Series 27.
1966. 148 pp. $5.00 (Univ. of Kansas Libraries, Lawrence, Kansas 66044). There are
enough identified botanical bibliophiles among our subscribers to justify calling attention
to this collection of essays ranging from W. T. Steam’s excellent “The Use of Bibliogra¬
phy in Natural History” (which includes among other topics a fine presentation on Lin¬
naeus and his “binomial system” for literature citations) to Jerry Stannard’s extensively
documented “Early American Botany and its Sources.” Illustrated botanical works in
China and Japan and a study of Linnaeus are among the several other subjects covered
in this volume.

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77

THE SPRING BEAUTIES ( CLAYTONIA ) IN MICHIGAN

Edward G. Voss

Herbarium, The University of Michigan, Ann Arbor

Confusion over our two species of Claytonia has existed ever since (as well
as before) Asa Gray began his illustrious botanical career. In his incomplete auto¬
biography, Gray wrote that in the spring of 1828 [at the age of 17 and three
years before receiving his M. D. degree] , while serving his medical apprenticeship
with Dr. J. F. Trowbridge of Bridgewater, New York,

. . .1 sallied forth one April day into the bare woods, found an early specimen of a
plant in flower, peeping through dead leaves, brought it home, and with Eaton’s
“Manual” without much difficulty I ran it down to its name, Claytonia Virginica.
(It was really C. Caroliniana, but the two were not distinguished in that book.) I was
well pleased, and went on, collecting and examining all the flowers I could lay hands
on; and the rides over the country to visit patients along with my preceptor, Dr.
Trowbridge, gave good opportunities. I began an herbarium of shockingly bad
specimens.!

Thus did Claytonia launch the botanical career of the man who was to become
the leading botanist of the nation; the author of numerous textbooks, manuals,
floras, and monographs; and the first professor to be appointed to the faculty of
the University of Michigan after it was reorganized in Ann Arbor in 1837.

The original Asa Gray Bulletin, published for the Gray Memorial Botanical
Chapter of the Agassiz Association,* 2 carried several short notes on Claytonia
e. g., in Nos. 5 & 6. 1894) and it was even suggested that C. virginica “be selected
as the Chapter flower.” (Asa Gray Bull. 3(8): 6. 1895.) When this selection was
proposed, it was observed that “Either species would serve our purpose, but [C.
virginica] is most common. The range of both is limited.”

The ranges are indeed limited, and have not been well understood in the
Great Lakes region. The data here presented on the occurrence of the two spe¬
cies in Michigan (with a few other observations) will add another dimension to
the studies cited under “References” on the taxonomy and cytology of Claytonia
which seem to indicate a resurgence of interest in the genus during the past few
years. The map in Fig. 1, indicating the location of Michigan counties, will aid in
understanding the geographic remarks.

The more northerly wide-leaved spring beauty, Claytonia caroliniana, al¬
though described as long ago as 1803, was not always recognized by early bota¬
nists or was considered to be a variety [var. latifolia Torr.] of C. virginica, the
more southern narrow-leaved spring beauty. Both species have, however, been

lSee pp. 14-15 in “Autobiography,” Chapter I of Letters of Asa Gray, ed. by Jane
Loring Gray (Macmillan, 1893). Gray’s memory evidently lapsed slightly when recording
these autobiographical notes, for the two species were clearly distinguished in Eaton’s Manu¬
al (both 3rd ed., 1822, and 4th ed., 1824), with “caroliniana Mx.” as a footnote on C.
spatulata, the name used by Eaton.

2See Harley H. Bartlett, History of the Gray Memorial Botanical Association and the
Asa Gray Bulletin, Asa Gray Bull. N. S. 1: 3-22. 1952. Botanists from Michigan were par¬
ticularly active in the Gray Memorial Botanical Chapter.

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admitted in all editions of Gray’s Manual, since 1848, and in almost all other
manuals published during the same period. But some collectors, originally fa¬
miliar with only C. virginica, have been slow to realize that another species also
occurs in this region. The first published report of C. caroliniana from Michi¬
gan3 appears to have been by Coleman in 1874, but the species was not included
in the Wheeler and Smith list of 1881. By 1892, both species were included in
the Beal and Wheeler list, and both were in Beal’s “Michigan Flora” of 1905.
The latter two lists properly indicated that C. caroliniana was restricted to the
northern part of the state, but erred in giving the range of C. virginica as “through¬
out,” thus obscuring the distinctive and essentially complementary patterns of the
two species (cf. Figs. 4 & 5). Early reports of C. virginica from northern Michi¬
gan are all almost certainly errors for C. caroliniana. I know of no previously
published records for C. virginica north of St. Clair and Kent counties which are
supported by specimens.

At the time of a previous summary (Voss, 1957) of the distribution of
Claytonia in the state, there appeared to be a belt one to three counties wide
across the middle of the Tower Peninsula, from which neither species was known.
North of this strip was only the wide-leaved species; south of it, only the nar¬
row-leaved one. Such a distinction in ranges seemed “too good to be true,” and I
suggested that “Further collecting in the two or three tiers of counties in the
center of the Tower Peninsula, between the present known ranges of the two
species, should be profitable.”

Following my own advice without delay, in' 1957 I pursued spring beau¬
ties, particularly across the middle of the state west of Saginaw Bay. On the
major excursion for this purpose (May 3-5), Thomas R. Detwyler was a most en¬
thusiastic and stimulating companion in the field, and we explored the “Thumb”
area southeast of Saginaw Bay the following spring. Numerous other students
and colleagues accompanied other trips during those years and the subsequent
ones, and to all of them I am grateful for the sharp eyes and willingness to look
into “just one more woodlot before dark” which have resulted in over 100 col¬
lections of Claytonia since 1957. (If this genus was so poorly represented in
herbaria from certain parts of the state, it could be assumed that other spring
plants were likewise under-represented, so that general collecting was also done
on all field trips.)

Plants of Claytonia are usually so abundant in the field that one readily
sees a great range of variability.4 To some extent, this may be associated with
extraordinary cytological diversity, as recently summarized by Fewis, et al.

3The early Michigan lists referred to are as follows:

Coleman, N. 1874. Catalogue of Flowering Plants of the Southern Peninsula of Michigan,
with a Few of the Crytogamia [sic] . Kent Sci. Inst. Misc. Publ. 2. 49 pp.

Wheeler, Charles F., & Erwin F. Smith. 1881. Michigan Flora. Rep. St. Hort. Soc. Mich.

10: 427-529. [Reprinted with different title page and repaged 1-105.]

Beal, W. J., & C. F. Wheeler. 1892. Michigan Flora. Rep. St. Bd. Agr. Mich. 30: 471-650.
[Reprinted and repaged 1-180.]

Beal, W. J. 1905 [“1904”]. Michigan Flora. Rep. Mich. Acad. 5: 1-147.

4As early as 1876, C. F. Wheeler (Bot. Gaz. 2: 65) noted considerable variation in C.
virginica, presumably at Hubbardston, Michigan.

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79

(1967b). Chromosome numbers in C. virginica throughout its range vary from
2n = 12 to ca. 191. Numbers may vary within a single population— and even in a
single individual (Lewis, 1962). Artificial hybridization between plants known to
differ in chromosome number has been successful in producing offspring (Roth-
well, 1959, p. 355). Higher polyploids of C. virginica were found in the area of
St. Louis, Missouri, to be separable morphologically from lower polyploids by the
broader leaves of the former (Lewis, et al., 1967a).

A thorough study of chromosomal behavior in Michigan Claytonias would
be highly desirable. Buds preserved from plants in the remarkable ravine in
Oceana County (described below) where both species occur were examined
through the kindness of W. H. Wagner, Jr., but the material was too old on May
3, 1964, for meiotic figures.5 I would be happy to supply as detailed directions

Fig. 1. Michigan, indicating location of counties.

5 Lewis (1967, p. 109) reports that meiosis in C. caroliniana may occur under the
snow very early in the spring.

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as possible to the sites where both (or either) species occur, to anyone with a de¬
sire to carry out for this region— but involving two species— the kind of extensive
study done for C. virginica alone by Lewis, et al. (1967a) in Missouri or Rothwell
and Kump (1965) in the New York area. The only Michigan counts given by
Lewis, et al. (1967b) are from Berrien County: 2n = 24, 25, 26.

CLAY TONI A VIRGINICA

Claytonia virginica L. (Lig. 2) ranges from Georgia, Louisiana, and eastern
Texas north to Nova Scotia, New Brunswick, southern and western New England,
southern Quebec and Ontario, the Lake States (becoming absent northward in
Michigan, Wisconsin, and Minnesota), Nebraska, and Oklahoma. One of 15
North American species in the genus, it is most closely allied to C. caroliniana,
the only other species in the states east of the Great Lakes, from which it is
normally distinguished readily by the shape of the cauline (stem) leaves. These
are linear to narrowly elliptical in C. virginica, characteristically about 13-17
times as long as broad, with no definite petiole. The basal leaves, arising directly
from the underground corm, are often larger and broader.

Normally there are two opposite cauline leaves, but aberrant individuals
with three leaves are occasional, as also in C. caroliniana; and rarely there may be
a flowering axillary branch (e. g., 5656, Tuscola Co.; 9267, Allegan Co.; 7911,
Clare Co.; 1 1445, Ottawa Co.; & McVaugh 1 1224, Branch Co.).

Fig. 2. Claytonia virginica, showing characteristic linear leaves. Photographed by James S.
Pringle at the Cootes’ Paradise Santuary of the Royal Botanical Gardens, near Hamilton,
Ontario.

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81

Broad-leaved plants of C. virginica may be mistaken for C. caroliniana, es¬
pecially if one is not familiar with the characteristic rhombic or diamond-shaped
blades of the latter. Davis recognizes C. virginica f. robusta (Somes) Palmer &
Steyermark as such a large elliptic- to lanceolate-leaved form, with a range from
Indiana to Iowa and Missouri. What could surely be referred to this form was
found with the normal form by M. F. Tessene on a trip with me in Auglaize
County, Ohio (floodplain forest along Auglaize River by Buckland Quarry, ca. 4
mi. NW of Wapakoneta, April 23, 1967. 12408, MICH, OS). The leaves of the
Ohio plants range up to 18 mm broad by 95 mm long (some slightly narrower
ones to 120 mm long), or less than 6 times as long as broad. Other robust plants
are discussed below under certain records for C. caroliniana.

In Michigan, C. virginica is extremely local beyond T20N; i. e., the north¬
ern edges of Mason, Lake, Osceola, Clare, Gladwin, and Arenac counties, or ap¬
proximately 44° 10* N. lat. The only collections from north of this line (see
Fig. 4) are as follows:

ROSCOMMON CO.: common in low deciduous woods, NViNWVi sec. 17, SV2 Richfield Tp.,
ca. 5.5 mi. SW of St. Helen, May 14, 1961, with Richard C. Crickmer. (9813) Carpinus
caroliniana and Viola fimbriatula, both at the northern edge of their range in the Lower
Peninsula, were in the same woods.

Fig. 3. Claytonia caroliniana, showing characteristic wide leaves. Photographed by Fred¬
erick W. Case, Jr., near Lewiston, Michigan, May 2, 1964.

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ALCONA CO.: abundant in young beech-maple woods near center of S edge of sec. 26, T27N,
R5E, ca. 1 mi. SE of Curran, May 10, 1957, with R. M. Schuster. (3533).

MACKINAC CO.: Round Island-discussed below under “Mixed Sites.”

DELTA CO.: occasional in deciduous woods near E edge of secs. 26-35, T41N, R23W, ca. 7
mi. NW of Gladstone, May 24, 1958. (5965) John H. Beaman first spotted the plants at this
site, which remains the only one known on the mainland of the Upper Peninsula.

CLA YTONIA CA R O LIN I ANA

Claytonia caroliniana Michx. (Fig. 3) ranges from Prince Edward Island,
Newfoundland, and Gasp£ west across Quebec (cf. Doyon & Lavoie, 1966),
Ontario, including the north side of Lake Superior, and the northern portions of
Michigan and Wisconsin, to northeastern Minnesota (Cook and St. Louis coun¬
ties), south through New England and New York to Pennsylvania and in the Ap¬
palachian Mountains to eastern Kentucky, North Carolina, Tennessee, and
Georgia; also in the Ozark Mountains of northwestern Arkansas, according to
Davis (1966).* * * * * 6 A map of the distribution, naturally incomplete but better than
many statements on this species, was published by Braun (Rhodora 39: 202.
1937), who has stated to me (December, 1953) that she does not believe the
southwestern Ohio reports (Schaffner, Ohio Jour. Sci. 38: 213. 1938). 7 This
map included only one Michigan record, on the Keweenaw Peninsula; Fig. 5 covers
this state more thoroughly.

The cauline leaves of C. caroliniana typically have a distinctive broad, more
or less diamond-shaped or rhombic to somewhat ovate blade and a definite petiole.
The total length (including petiole) is characteristically about 3-5 times the width,

^Not fitting into this pattern is a collection cited by Davis (1966) as “Hanover, Jef¬

ferson Co., Indiana, Young (MICH)”; I am more certain that the locality is wrong than the

identification, but question both. The printing on the label reads at the top “Herb, of A. H.

Young” and at the bottom “Hanover, Ind., 18 ” and is in a style common on labels of the
last quarter of the previous century. The bottom line is, I think, merely the address of the

collector; in addition to the name “Claytonia caroliniana” there has been written “La

Fayette” on the label, and this is presumably the locality. The date is given as April 1 877 —
the year when Young moved to Lafayette (cf. Natl. Cycl. Am. Biogr. 2: 397 & 22: 17). There
are only two plants, so far out of the otherwise known range of C. caroliniana that I would
echo the skepticism of Deam (FI. Indiana, pp. 1044-1045. 1940) regarding Indiana records
and suggest that these broad-, but scarcely rhombic-leaved plants may be only forms of C.
virginica. Jones (Am. Midi. Nat. 34: 279. 1945) similarly disposed of Illinois records of C.
caroliniana. See also footnote 7.

7Such reports, and others, of C. caroliniana from areas south and west of what is
apparently its range are presumably based on large, wide-leaved forms of C. virginica, as
suggested in the previous footnote. Whether these result from genetic or ecological factors,
or both, is not certain. The Bartlett collection of C. virginica f. robusta cited by Davis
(1966, p. 196) from Marion Co., Indiana, came from Little Eagle Creek, April 24, 1904,
and bears the notation by the collector: “Ecological form developed where the natural soil
(black forest loam) is covered with 4-5 inches of drifting sand.” There is a single plant,
with cauline leaves 20 x 108 and 21 x 113 mm and one basal leaf 27 x 150 mm. Broad¬
leaved plants from southern Minnesota have been questioned by Morley (Spring FI. Minn.,
p. 105. 1966). A collection from Oceana Co., Mich. (9391, with T. R. Detwyler, near center
sec. 25, Benona Tp., May 14, 1960) from beech-maple woods only 2 miles south of the ra¬
vine where both species occur resembles f. robusta, the largest cauline leaves being 15-17
mm broad and about 168 mm long. Some of the plants from Round Island (12438) are
similarly robust.

1968

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83

and this ratio is much more consistent than any absolute measurements (see under
“Distinguishing Characters,” below). Plants from higher elevations in the south¬
ern Appalachians have recently been recognized as var. spatulifolia (Salisb.)
Lewis (1966); these have much smaller and proportionally somewhat narrower
leaves than typical var. caroliniana ; judging from Lewis’ figures, they may run
close to 8 times as long as broad, but as they are much shorter than those of
equal width in C. virginica there should be no confusion. Lurther investigation of
small-leaved plants might be fruitful. Very small-leaved plants were found by L.
O. Gaiser in 1960 in a woods near Thedford, Ontario (see below); the leaves of
some, though not all, of these resemble Lewis’ figures of var. spatulifolia. Simi¬
lar very small-leaved individuals have been collected, generally with larger ones,
at several Michigan localities (e.g., 3427, Ogemaw Co.; 3529, Alcona Co.; 6099,
Baraga Co.; 1 1456, Oceana Co.; & Parmelee 2221.1 , Houghton Co., MSC).

Cytologically, C. caroliniana also displays extensive aneuploidy and poly¬
ploidy; it has known 2n numbers ranging from 16 to 38 (Lewis, 1967; Lewis, et
al., 1967b). Perhaps it would show a greater range than this if it were investigated
as thoroughly as C. virginica, although its more restricted geographic distribution
suggests less chromosome diversity as well.

In Michigan, C. caroliniana is definitely northern, evidently ranging a bit
farther south near Lake Michigan and Lake Huron than in the interior (Lig. 5).
The southernmost localities are discussed below under “Mixed Sites.” The spe¬
cies was listed by Billington (Pap. Mich. Acad. 4: 96. 1925), in addition to C.
virginica, from “Climax forest” at Warren Woods, in Berrien County, the south-
westernmost county in the state. Warren Woods is not generally a location for
northern plants; quite the contrary, it is the site for several species of southern
affinity known from few if any other places in Michigan (e. g., Diarrhena ameri-
cana, Uniola latifolia, Carex davisii, Trillium recurvatum, Chaerophyllum pro-
cumbens). Billington’s collections labeled as C. caroliniana consist of 5 speci¬
mens, on two sheets (WUD, BLH), both collected May 18, 1921. The cauline
leaves range from 9.5 to 14.5 mm broad, from 58 to 114 mm long, and from
5.5 to 12 times as long as broad. Insofar as lower bracts are present (the WUD
sheet lacks one raceme), they are herbaceous, supposedly a character for C. vir¬
ginica. 1 consider these plants to be robust forms of that species. On May 2,
1964, I searched Warren Woods for Claytonias, with Ronald L. Stuckey and
Chester W. Laskowski. We looked especially on the slopes and ravines, where a
colony of C. caroliniana might possibly be sheltered, hiking completely around
the upland area mapped by Billington as “Climax Lorest,” and angling across
the upland. Normal C. virginica was everywhere. Some plants with slightly wider
leaves than usual were found (1 1442, 1 1443, 1 1444), particularly in the narrow
spaces between buttresses at the bases of large trees, but none really resembled C.
caroliniana in leaf shape and they have all since been confirmed by R. J. Davis as
C. virginica. Their leaves range from 8.5 to 15.5 mm broad, from 65 to 97 mm
long, and from 4 to 10 times as long as broad.

DISTINGUISHING CHARACTERS

Leaf shape seems to be the best characteristic distinguishing Claytonia
caroliniana from C. virginica, and is the only one given in the key by Davis

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(1966). Other supposed differences sometimes mentioned in manuals do not
seem, in my experience, consistent enough to be useful. There is apparently a
slight phenological difference, as discussed under “Mixed Sites” below, but it is
of little use in distinguishing species as there is great overlap. The dividing point
in leaf shape has sometimes been stated as 6 times as long as broad; that is,
plants with leaves more than 6 times as long are C. virginica; those with leaves
less than 6 times as long are C. caroliniana. I find somewhat better discrimination
at 8 times as long as broad (Voss, 1966) and the same figure is given by Davis
(1966). Almost always, individuals are not near that dividing line, but there are
occasional borderline plants and a certain amount of intuition may be needed to
interpret ambiguous specimens.

The cauline leaves of Michigan plants of C. virginica apparently range
from (6-) 8.5 to 19 (-36) times as long as broad. In some very unusual individuals,
as cited above from Warren Woods, they may be as short as 4-5 times as long as

Fig. 4. Distribution of Claytonia virginica in Michigan, based on herbarium specimens ex¬
amined. (A few collections are not mapped, where the additional dots would only add con¬
gestion in already well-collected areas.)

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85

broad, but these are more elliptical or oblong than the characteristic shape of C.
caroliniana. They range from 1.5 to 14 (-18) mm broad and from 24 to 140
(-168) mm long. The racemes bear (1-) 6-16 (-22) flowers. The seeds are about
2-2.5 mm long. The bract below the lowest pedicel is scarious or often partly
herbaceous, rarely entirely herbaceous. Different stems from the same corm will
vary widely in the degree of herbaceousness of the bract.

In Claytonia caroliniana, our specimens have leaves which run from 2.5 to
6.5 (-7.8) times as long as broad (or very rarely as much as 8.8 times as long, but
these leaves with definite typical wide blade and unusually long petiole). The
width ranges from (2.3-) 3.5 to 30 mm, and the length from 12 to 111 (-132)
mm. The racemes bear (1-) 4-9 (-13) flowers. The seeds range 1.7-2. 3 (-2.5) mm
long. The bract below the lowest pedicel is, as in the other species, usually scar¬
ious or partly herbaceous, rarely entirely herbaceous.

In short, leaf shape appears to be the only useful distinguishing character.
The seeds of Michigan specimens run, if anything, slightly opposite the size

Fig. 5. Distribution of Claytonia caroliniana in Michigan, based on herbarium specimens ex¬
amined. (As in Fig. 4, a few collections not mapped.)

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difference indicated by Davis (1966) but the distinction is insignificant as the
seeds are much more variable than his simple “2 mm” vs. “2.5 mm” would im¬
ply. The bract character used by Fernald (Man., ed. 8. 1950) is wholly unreliable
in this region. Some plants of C. virginica do produce more flowers than any of
C. caroliniana, but except for those few individuals the number of flowers in an
inflorescence is not significant. The flowers are sometimes said to be smaller in
C. caroliniana , but I have not noticed any distinction in the field; in fact, this spe¬
cies has the largest flowers I have seen on any herbarium material, as much as 27
mm across (Chippewa Co., rich upland maple woods of sections 28 & 33, T48N,
R2E, Sugar Island, May 13, 1961. Hiltunen 2796, WUD). C. virginica may have
a little wider ecological amplitude, at least in southern Michigan (where C. caro¬
liniana does not grow), where it is found often in floodplain woods and thickets
as well as upland sites; but as discussed below, the two species may grow under
identical conditions.

MIXED SITES

Ordinarily, Claytonia caroliniana and C. virginica “seldom grow in the
same immediate area” (Davis, 1966, p. 295). In those regions where both spe¬
cies are found, they are usually separated by such factors as habitat or elevation.
Although most of his collecting was apparently conducted where C. caroliniana
does not occur, Rothwell(1959, p. 357) stated, “In none of the localities studied
were C. virginica and C. caroliniana found to be growing together, and in no case
were populations discovered which appeared to be intermediate between the 2
species.” Raymond (1949) suggested a distinct ecological separation in Quebec.
On the other hand, Uttal (1964) reported C. virginica and C. caroliniana in two
different but adjacent habitats in Virginia, with an apparent hybrid population
on “the border between the two parental habitats.”

Because the two species generally grow apart from each other, mixed col¬
lections are almost non-existent in herbaria. Davis (1966) implies that a collec¬
tion from Truro, Nova Scotia (Prince & Atwood 859, GH, DS), may have been
mixed, though segregated onto two sheets, for he cites the same number under
both species. At the northwest edge of its range, C. caroliniana is found, as is C.
virginica, near Duluth, St. Louis County, Minnesota, but the two are evidently
not in the same woods (cf. Lakela, FI. NE Minn., pp. 170 & 436, 1965). From
that region, C. caroliniana ranges northeastward, north of Lake Superior, into
Cook County (Butters & Abbe, Rhodora 55: 148. 1953) and the Canadian Lake-
head; while C. virginica ranges only southward— in Wisconsin, north to Barron,
Lincoln, and Kewaunee counties (Fassett, Spring FI. Wis., ed. 3, p. 54. 1957;
Seymour, FI. Lincoln Co. Wis., p. 227. 1960).

The zone of overlapping ranges extends eastward from Minnesota and Wis¬
consin across Michigan and Ontario. Here, both species sometimes grow in the
same woods, in exactly the same habitats— so close together, in fact, that the
numerous fragile stems of the two species may become thoroughly intermixed as
they rise through the ground from adjacent corms. In more numerous instances,
the two species may grow within a few yards or at least a mile of each other.

The only observation reported by Davis (1966, p. 295) where the two

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species grow near each other was at Newfound Gap in the Great Smoky Mountains
National Park, where he found C. virginica blooming earlier on the east side than
C. caroliniana on the west side. From this he generalized that when the two do
grow “in the same immediate area, . . . C. virginica blooms earlier.” My field ex¬
perience is just the opposite. When the two species have been found growing to¬
gether, in exactly the same situation, in Michigan and Ontario, C. virginica
reaches its peak a little later (but see notes on the Ontario site, below). The dif¬
ference is not great, and I judge only on the basis of mature plants, not having
observed which species actually begins to bloom first in these sites. Plants of C.
caroliniana in mixed populations have more mature flowers and fruit, and fewer
buds, than those of C. virginica. Described below are the station in Ontario and
the Michigan counties where the two species grow within a mile of each other,
including sites where they are actually mixed.

ONTARIO: LAMBTON CO.: Early in 1964, just a year before her death, L. 0.
Gaiser wrote me of a woods near Thedford, in which both species grew, and sent
me for examination all of her collections made there since 1960. (Her first col¬
lections, made April 27, 1959, are in the herbarium of the University of Guelph.)
Specimens were first collected at this site by the late Newton Tripp, a naturalist
of nearby Forest, on May 11, 1917 (specimens also now housed at Guelph). Dr.
Gaiser relocated the woods, on the south side of Ridge Road, ca. lA mi. W of
Cone. 6, Bosanquet Tp., or about 3 miles west of Thedford. Her careful survey of
Lambton County failed to reveal any other locations for C. caroliniana, although
C. virginica is common throughout the county. Dr. Gaiser’s impression was that
C. virginica was the less common species in this woods and occurred on higher
ground.8

Large specimens of C. virginica were collected “on the higher ground, under
beech tree. . .towards eastern edge of woods” May 4, 1960 (Gaiser 2576T, DAO,
MICH); the label includes the notations “Found this species only on this higher
ground. Flowering nearly over.” Indeed, the plants are definitely more mature
than those of C. caroliniana (Gaiser 2578T, DAO, MICH) collected the same
date “in the low ground. . .not on high ground.” Also collected at the same
time were plants of C. caroliniana (2580T, DAO, MICH) with small petals and
very small leaves (e. g., 3 x 15, 4 x 15, 5 x 23 mm). These were said to have
come from low ground along with other C. caroliniana. Her May 12, 1961, col¬
lections, on the contrary, indicate a less advanced state for C. virginica (2954T,
DAO, MICH), which is labeled: “NB On the higher ground, not with C. caro¬
liniana. ” Considerable variation is displayed in the size of the leaves of C. caro¬
liniana (2950T, DAO, MICH) collected on the same date; they range from 6 x 24
to 13 x 45 mm.

On May 27, 1966, Kenneth E. Guire and I relocated the woods while en
route to the Michigan Botanical Club spring campout on the Bruce Peninsula. We

8Dr. Gaiser’s observations are taken from A Survey of the Vascular Plants of Lamb¬
ton County, Ontario, by Lulu O. Gaiser, compiled by Raymond J. Moore (Plant Res. Inst.,
Canada Dep. Agr., 1966. 122 pp.); from the labels with her specimens; and from letters
dated March 6, March 13, and May 2, 1964.

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Vol. 7

were too late, however, for Claytonias, and after much searching found only a
few very old and shriveled leaves of C. virginica. Was it too late even for that
much evidence of C. caroliniana? (We did find large colonies of the green violet,
Hybanthus concolor, reported by Gaiser only from another location in the
county.) I returned to the woods on May 6, 1967, with W. H. Wagner, Jr., David
M. Hines, and M. F. Tessene, with better luck. We found Claytonias very scarce
except toward the west end of the woods, where both species were common, es¬
pecially C. virginica. C. caroliniana was in the northwest part of the woods, with
C. virginica alone extending southward, without significant changes in elevation
or exposure and with some dense colonies of both species mixed together. C.
caroliniana (12410) appeared to be slightly earlier phenologically; at least it sel¬
dom had unopened buds, while plants of C. virginica (12409) generally still had
some unopened buds. However, the oldest immature fruits appeared roughly to
be of the same age in the two species, suggesting that they may have begun
flowering at about the same time but that C. virginica (averaging, incidentally, a
few more flowers per raceme) would last longer.

This Thedford site is only about 20 miles farther south than the southern¬
most stations for C. caroliniana in Sanilac County, Michigan, just to the west
across the southern end of Lake Huron. But I have been unable to find both
species closer than two or three miles from each other in that county.

OSCEOLA COUNTY, MICHIGAN: When serious pursuit of Claytonias was be¬
gun with T. R. Detwyler in early May of 1957, Osceola County, in the west-cen¬
tral Lower Peninsula, was the first in which we recognized both species. Extensive
exploration of the numerous woodlots in the northern part of the county on
May 5 revealed either no Claytonia or only one of the species in each. Apparently

Fig. 6. The narrow4eaved Claytonia virginica and broad-leaved C. caroliniana, growing to¬
gether in the ravine (described in text) in Oceana Co., Michigan, May 3, 1964.

1968

THE MICHIGAN BOTANIST

89

similar-looking woods, on what seemed to be the same moraine, would not nec¬
essarily have the same species, however, and hoped-for correlations with habitat
did not materialize. On the east side of U.S. Highway 131, for example, in beech-
maple woods back of Maple Hill Cemetery ca. 2 miles northeast of LeRoy, C.
caroliniana was common. A mile south, on the west side of the highway, C. vir-
ginica was abundant in beech-maple-hemlock woods. In Highland Township, C.
virginica was common at the edge of a grazed lowland elm-maple woods at the N
edge of the NE14 of Sec. 29. Half a mile north, in the SE/4 of sec. 20, C. carolini¬
ana was locally abundant in alow elm-maple woodland; and a half-mile west, it was
frequent in low aspen-elm woods in the NW!4 of sec. 29. Nearby, in a fine mature
beech-maple-hemlock stand in the SE:A of sec. 19, C. caroliniana was abundant.

In 1957, thought was not given to possible phenological differences. The
evidence from 4 collections of C. virginica and 7 of C. caroliniana made on the
same date in northern Osceola County (all from different woods) is that C. vir¬
ginica is definitely later.

CLARE COUNTY: On May 4, 1957, our last collecting site before dusk— and a
quick search for a campsite— was ca. 4 mi. NW of Harrison in a grazed upland
maple woods in the SWXA of sec. 14, Greenwood Township, where we had been
directed by a nearby farmer in response to our desperate search for a good wood-
lot in otherwise rather bleak terrain. Here, C. caroliniana (3445) was abundant
and we included in the collection a narrow-leaved plant bearing 6 stems from a
single corm in the midst of a colony of wide-leaved ones. No other plants with
narrow leaves were seen, and these were perhaps a little less linear than usual for
C. virginica; we assumed them to be aberrant C. caroliniana, being unaware at the
time that the two might grow so close together. But they have since been con¬
firmed as C. virginica by R. J. Davis. This woods should be examined again
more thoroughly.

Somewhat farther south in Clare County, along Highway M-l 15 ca. 4 mi.
NW of Farwell, chiefly in the NE14 of sec. 17, Surrey Township, is a beech-maple
woods in which Richard C. Crickmer and I found both species on May 12, 1961.
Oddly enough, only C. virginica (971 1) was on the north side of the highway and
only C. caroliniana (97 07) on the south side. If once they grew beside each other,
the highway must have destroyed the area of overlap. The plants of C. caroliniana
were more advanced, even starting to turn yellow while those of C. virginica were
yet green.

MUSKEGON COUNTY: While both species have not been found together, de¬
spite search in many woods in the northwest part of the county, there remains the
possibility of a mixed site in Muskegon County. On May 14, 1960, T. R. Det-
wyler and I found large plants of C. virginica common in a low maple woods in
the NE/4 of sec. 13, White River Township, ca. IVi mi. NW of Montague— well
within its range. While both species of Erythronium, E. albidum and E. ameri-
canum, were in this woods, we did not find the other Claytonia. Not until May 3,
1964, did C. caroliniana turn up as far south as Muskegon Co., when R. L.
Stuckey, C. W. Laskowski, and I found it in deciduous woods in the NE14 of
sec. 12 (11470) and the EV2 of sec. 11 (11472), White River Tp.— both stations
only about a mile from C. caroliniana— and near the E edge of sec. 1, about IVi
mi. north (1 1464).

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THE MICHIGAN BOTANIST

Vol. 7

OCEANA COUNTY: On May 15, 1957, about 6:30 p.m., James S. Wilson and I
decided to delay our search for a campsite long enough for a quick look into a
beech-maple woods about 2 Vi miles west of Shelby. On the north side of a ravine
we found C. virginica (3747), mostly with a few flowers left; on the south side,
C. caroliniana (3745), mostly just past flowering; and along the bottom of the
ravine an assortment of plants (3746) which at the time we thought might in¬
clude some hybrids. (Later segregation into the two species was made by R. J.
Davis.) This woods was the first recognized station in Michigan for the occur¬
rence of both species, and on May 3, 1964, a return visit, with R. L. Stuckey and
C. W. Laskowski, was made at a more favorable time of day. The meeting point
of the two species is the second east-west ravine south of the county road (and
paralleling it). The ravine expands westward into the wide valley of Stony Creek,
which flows from north to south. On the west-facing slope of the upland, facing
the valley of Stony Creek, there is C. caroliniana, which is also on the south or
north-facing, slope of the ravine and the upland thence southward. On the north,
or south-facing, slope of the ravine and thence northward is only C. virginica.
Both species occur at the bottom of the ravine. Plants bearing leaves which ap¬
peared to include shapes of both species in one compact group of stems (Fig. 6)
were painstakingly excavated, revealing two corms of C. virginica (1 1455) and one
corm of C. caroliniana (11456) nearly or quite touching each other. It was here
that we realized that a phenological difference exists when the two species do
grow under identical conditions. Young buds were very scarce on C. caroliniana
while they were common on C. virginica.

Fig. 7. Claytonia virginica and C. caroliniana (note pair of broad leaves in lower center, be¬
hind peduncle and large linear leaves of C. virginica ), growing together on Round Island,
Mackinac Co., Michigan, May 29, 1967.

1968

THE MICHIGAN BOTANIST

91

If hybridization does occur between the two species, it may be taking place
in this ravine. Certain plants (e. g., 11459 and some of 3746) are of ambiguous
leaf shape when considered by themselves, but would not be too far out of line
for either species (especially C. virginica ) if found in otherwise pure stands. None
of them have been annotated as possible hybrids by Davis, but in the light of
Uttal’s Virginia report (1964) I think further study is warranted.

ROUND ISLAND (MACKINAC CO.): In the herbarium of the Cranbrook Insti¬
tute of Science is a sheet bearing two specimens labeled as C. virginica, collected
on “hardwood plateau” of Round Island, in the Straits of Mackinac, between
Mackinac Island and Bois Blanc Island, in June of 1941 by Marjorie T. Bingham.
In 1957,1 felt that these must be aberrant individuals of C. caroliniana, especially
since the lowest bract was scarious as sometimes stated for that species in manu¬
als. The site seemed far out of range for C. virginica (which was unknown from
the neighboring mainland in any direction) although the leaves were quite linear
(all at least 15 times as long as broad). Only a visit to Round Island9 could cli¬
max my decade of Claytonia- hunting, and on May 29, 1967, the opportunity
came, through the kindness of John W. Childs, of Wayne State University, and
his wife.

The high interior of the island, above the strong shoreline of post-glacial
Lake Nipissing and the thickets below it, rises a hundred feet above the Straits—
twice the estimate of Potzger,9 and is heavily wooded with beech ( Fagus grandi-
folia), birch ( Betula papyrifera), and sugar maple (Acer saccharum), the latter in
an almost pure stand northward. The trees are mostly only of medium size and
the whole upland has rather a park-like aspect. Shrubs such as red-berried elder
(Sambucus pubens ) are heavily browsed. On the calm, sunny day when we were
there, the forest floor was carpeted with a wide assortment of spring wildflowers,
the most abundant being Clay tonia caroliniana— which was apparently responsible
for the fragrance which perfumed the air. It seemed surprising that Mrs. Bingham
had not collected this species when she was on the island, but perhaps in June she
was too late, and could find only the slightly later C. virginica in suitable condi¬
tion to collect. For toward the north end of the upland, we did find C. virginica
(12440), thoroughly intermixed with C. caroliniana (12439) and, as usual, less
far advanced (Fig. 7). The plants first drew our attention by their white flowers
and large size. Subsequently, some pale pink-flowered C. virginica and some
white-flowered C. caroliniana were also found. One colony of very robust plants
(12438) included individuals resembling C. virginica f. robusta of farther south;
they had narrowly elliptical leaves up to 17 x 1 17 mm, though more typically 8 x
150 to 9 x 160 mm. Although C. caroliniana was abundant throughout the upland
of Round Island, we found C. virginica with it only in a very limited area near the
north side. In considerable spring collecting on the mainland to the south, in Em¬
met and Cheboygan counties around the University of Michigan Biological Sta¬
tion, C. virginica has never been found; the nearest known location is almost 100
miles south of Round Island, in Alcona County.

9The island, scarcely IV2 miles long and in federal ownership, is now completely un¬
inhabited, the lighthouse at its north end (across from Mackinac Island) having been
abandoned since the brief botanical report, based on two days of August collecting, by J. E.
Potzger: The Vegetation of Round Island (Straits of Mackinac), Michigan. Butler Univ. Bot.
Stud. 6: 116-122. 1944. The Round Island Quadrangle {1Vi\ U. S. Geol. Surv., 1964) pro¬
vides an up-to-date map of the island.

92

THE MICHIGAN BOTANIST

Vol. 7

IN CONCLUSION

The collections of the past 1 1 seasons have shown that the ranges of our two
species of Claytonia do meet, indeed overlap, across the middle of the state; that
both species, neither one clearly more so than the other, are extremely variable
morphologically and in flower color, which ranges from white to deep pink; that
leaf shape seems to be the only consistent distinguishing character— and even this
is occasionally ambiguous; and that when both species grow together, C. virginica
reaches the peak of its flowering later by perhaps a few days— no longer— than C.
caroliniana.

This report documents in some detail the distributions of the two species in
Michigan, with special reference to the area of overlap and to unusual occurrences,
with some additional data from Ontario and neighboring states. I hope that, as a
result of these data, botanists in the region will become interested in pursuing the
important as yet unanswered questions: Why are the ranges what they are? Given
that climatic or historical factors may explain the general ranges of the species,
then why in the zone of overlap does one woods usually have only one of them
while a nearby woods has only the other? Do the two species in fact hybridize?
What normally prevents hybridization when the two grow together? What is the
range of cytological diversity which they exhibit and is it correlated in any way
with the geographical range or with morphological variability? Are there some
unrecognized but better or more consistent characters than leaf shape to dis¬
tinguish the species and to identify hybrids?

Research on the “Claytonia problem” is barely begun in the local flora, even
though we know very much more about the occurrence of the species than we did
12 years ago!

ACKNOWLEDGMENTS

I am grateful to many persons who have aided the search for knowledge about spring
beauties during the course of this work: to the late Dr. Lulu O. Gaiser of Crediton, Ontario,
who wrote me of the Thedford woods and lent specimens from Lambton County, some of
which were retained for the University of Michigan Herbarium and the others returned after
Dr. Gaiser’s death to Raymond J. Moore to be incldded with her collections left to him and
placed in the herbarium of the Canada Department of Agriculture, Ottawa (DAO); to Dr. Ray
J. Davis of Idaho State University who kindly examined over 200 collections of Claytonia
from the University of Michigan Herbarium late in 1964; to the numerous residents of Michi¬
gan who have granted me permission to explore their woodlots(and to those from whom, be¬
cause of circumstances, permission could not be sought); to the many students, botanical col¬
leagues, and other friends, some of them previously mentioned, who joined in the search; to
those who have provided photographs to illustrate this report; and to the curators of herbaria
where specimens were examined: Michigan State University (MSC), Cranbrook Institute of
Science (BLH), Wayne State University (WUD), Isle Roy ale National Park (IRP); and the Uni¬
versity of Michigan Biological Station (UMBS). The distribution maps are based on speci¬
mens in these herbaria, in addition to the University of Michigan Herbarium (MICH) and my
personal herbarium (at Mackinaw City). Specimen collection numbers have been cited when¬
ever it seemed advisable to be specific, for the benefit of any who may want to examine par¬
ticularly interesting records. Except when otherwise indicated, all cited numbers are mine
and all collections are in the University of Michigan Herbarium (with occasional duplicates
distributed elsewhere).

1968

THE MICHIGAN BOTANIST

93

REFERENCES

Davis, Ray J. 1966. The North American perennial species of Claytonia. Brittonia 18: 285-
303.

Davis, Ray J., & Richard G. Bowmer. 1966. Chromosome numbers in Claytonia. Brittonia
18: 37-38.

Doyon, Dominique, & Victorin Lavoie. 1966. La distribution de quelques especes vegetales
dans la region de Quebec et leur cadre phytosociologique. Nat. Canad. 93: 797-821.

Lewis, Walter H. 1962. Aneusomaty in aneuploid populations of Claytonia virginica. Am.
Jour. Bot. 49: 918-928.

Lewis, Walter H. 1966. Claytonia caroliniana var. spatulifolia (Salisbury) Lewis, stat. nov.
(Portulacaceae). Ann. Missouri Bot. Gard. 53: 258-259.

Lewis, Walter H. 1967. Cytocatalytic evolution in plants. Bot. Rev. 33: 105-115.

Lewis, Walter H., YutakaSuda, & Bruce MacBryde. 1967a. Chromosome numbers of Clay¬
tonia virginica in the St. Louis, Missouri, area. Ann. Missouri Bot. Gard. 54: 147-152.

Lewis, Walter H., Royce L. Oliver, & Yutaka Suda. 1967b. Cytogeography of Claytonia
virginica and its allies. Ann. Missouri Bot. Gard. 54: 153-171.

Raymond, Marcel. 1949. Le Claytonia virginica L. dans le Quebec. Nat. Canad. 76:201-204.

Rothwell, Norman V. 1959. Aneuploidy in Clay tonia virginica. Am. Jour. Bot. 46: 353-360.

Roth well, Norman V., & Joseph G. Kump. 1965. Chromosome numbers in populations of
Claytonia virginica from the New York metropolitan area. Am. Jour. Bot. 52: 403-407.

Uttal, Leonard J. 1964. A hybrid population of Claytonia in Virginia. Rhodora 66: 136-139.

Voss, Edward G. 1957. Observations on the Michigan flora-VI. Distribution records of
some Angiosperms new, rare, or misinterpreted in the state. Brittonia 9: 83-101.

Voss, Edward G. 1966. Two kinds of beauties. Mich. Audubon Newsl. 14(1): 7-8.

NEW LOCALITIES FOR MOSSES RARELY FOUND
IN MICHIGAN'S UPPER PENINSULA

Maynard C. Bowers

Department of Biology,

Northern Michigan University, Marquette

As a result of field studies during the course in Algae and Bryophytes
taught at the Northern Michigan University Field Station and personal research, I
have found new localities for mosses rarely collected in the Upper Peninsula of
Michigan. These mosses are Mnium andrewsianum Steere, Mnium pseudopunc-
tatum Br. and Sch., and Schistostega pennata Hook, and Tayl. Each of these is
reported from but one Michigan locality in Darlington (1964).

Mnium andrewsianum Steere as first reported by Mazzer and Sharp (1963)
was new to the United States and they described its locality as a bog along the
Melstrand Road about eight miles east of Munising, Alger County. My collection
was made in a swamp near Ross Lake, Schoolcraft County.

Mnium pseudopunctatum Br. and Sch. was collected for the second time in
the Upper Peninsula during a visit to the Grand Sable Dunes along Lake Superior
west of Grand Marais, Alger County. Another collection was made in a bog of

94

THE MICHIGAN BOTANIST

Vol. 7

Calla palustris L., Wild Calla, along the Dead River at the north edge of Marquette,
Marquette County. The previous records for this species in Michigan are: Lowry
(1948), Lake City, Missaukee County; Darlington (1964), Cecil Bay, Emmet
County (Sharp); and for the Upper Peninsula, Crum (1964), Mackinac Island,
Mackinac County (Wynne).

Schistostega pennata Hook, and Tayl. was first discovered in Michigan by
Nichols (1933) in the cave behind Scott Palls approximately ten miles west of
Munising, Alger County. Geologically, this is just into the limestone area of the
eastern half of the Upper Peninsula. Knowing that similar caves exist in lime¬
stone cliffs along Lake Superior, I looked for this moss and subsequently found
it in the caves along Powell Point approximately two miles west of Munising.

While none of the above collections add significantly to the range of the
mosses, they do give more complete knowledge as to their distribution pattern in
Michigan. The voucher specimens for these new records are in the herbarium of
M. C. Bowers.

LITERATURE CITED

Crum, H. 1964. Mosses of the Douglas Lake Region of Michigan. Mich. Bot. 3: 3-12; 48-63.
Darlington, H. T. 1964. The Mosses of Michigan. Cranbrook Institute of Science, Bull 47.

212 pp.

Lowry, R. J. 1948. A cytotaxonomic study of the genus Mnium. Mem. Torrey Bot. Club
20(2): 1-42.

Mazzer, S. J., & A. J. Sharp. 1963. Some moss reports for Michigan. Bryologist 66: 68-69.
Nichols, G. E. 1933. Notes on Michigan bryophytes.— II. Bryologist 36: 69-78.

MICHIGAN PLANTS IN PRINT
New Literature Relating to Michigan Botany

B. BOOKS, BULLETINS, SEPARATE PUBLICATIONS

Barneby, Rupert C. 1964. Atlas of North American Astragalus. Mem. N.Y. Bot. Card. 13.
1188 pp. in 2 vols. $35.00. [Distribution maps in this monograph include 3 Michigan
stations for A. canadensis var. canadensis and 9 for A. neglectus, the latter also including
a Farwell name, based on Lapeer County specimens, in synonymy. 1

Dansereau, Pierre, & Gerard Pageau. 1966. Distribution Geographique et £cologique du
Betula alleghaniensis. Mem. Jard. Bot. Montreal 58. 56 pp. [Distribution map unique in
employing different symbols for different habitats where yellow birch occurs; a number
of Michigan stations included. A peripheral station in Michigan is said to be Moran in
“Tahquamenon” county (there is no such county; Moran is in Mackinac Co.). Phyto-
sociological data from 35 stations, including 3 in Michigan: Tahquamenon Falls, Wolfs
Bog in Cheboygan Co., and Highland “State Park” (i. e., State Recreation Area-which is
in Oakland Co., not Wayne as stated). 1

Kreisel, Hanns. 1967. Taxonomisch-Pflanzengeographische Monographic der Gattung
Bovista. Nova Hedw. Beih. 25. 244 pp. + 70 figs. DM80 or $20.00. [Michigan specimens
are cited for several species of these puffballs, and the very generalized distribution
maps includes ranges which pass through this region.]

Editorial Notes

Upon the expiration of terms of members of the Editorial Board of THE MICHIGAN
BOTANIST, we take this opportunity to express appreciation to the retiring members,
Charles Barclay, Dr. C. M. Rogers, and Dr. Warren P. Stoutamire, and to welcome three new
members: Dr. Virginia L. Bailey, chairman of the Department of Biological Sciences, Detroit
Institute of Technology; Dr. Burton V. Barnes, associate professor of forestry, The University
of Michigan; and Dr. Richard Brewer, director of the Charles C. Adams Center for Ecological
Studies and associate professor of biology, Western Michigan University, Kalamazoo.

In this issue, we introduce some new type faces— the first such change since the
journal began in 1962. Reactions from readers, whether pro or con, will be welcomed. Is the
type as readable as before, or more so, or less so?

Good black and white prints of botanical subjects, suitable for use on the cover of
THE MICHIGAN BOTANIST, are always welcome and may be submitted to the editor at
any time.

The January number (Vol. 7, No. 1) was mailed January 4, 1968.

“Winter Wildflowers"

The Michigan Botanical Club has undertaken a new project to culminate in Special
Publication No. 2, on the winter appearance and uses of wild plants. Photographs (no
smaller than 5 x 7) of artistic merit, showing in general a single species of wild plant, are
needed-those with distinctive and recognizable inflorescences, berries, etc., that might be
noticed on a winter hike or used in “winter bouquets.” Photographs of a few effective ar¬
rangements of dried native materials are also wanted. Especially desired are specific direc¬
tions from persons who have successfully employed different techniques of preparing wild
winter plants for use— such as drying with various agents, spraying to prevent disintegration,
embedding, glycerin treating, arranging pictures or greeting cards, and so forth.

Anyone with doubts as to the identification of his material should save a specimen for
checking. Credit will, of course, be given to all whose photographs and suggestions are ac¬
cepted for inclusion by the booklet committee. All materials should be submitted to the
committee chairman. Dr. Helen V. Smith, 1766 Glenwood Rd., Ann Arbor, Michigan 48104.

Errata

In addition to a few obvious omissions, alterations, or transpositions of letters, the
following recent errata may be noted:

Vol. 6, p. 21, line 23, for “1. 51” read: p. 51

p. 31, line 2 from bottom, for “26 years” read: 27 years
p. 91, line 16, for “173” read: 1753
Vol. 7, p. 28, line 9 from bottom: for “Ecology” read: Am. Midi. Nat.

CONTENTS

A Preliminary Study of Clavariadelphus in North America

Virginia L. Wells & Phyllis E. Kempton . 35

Odd and Unusual Discomycetes from Michigan, I

Nancy Jane Smith . 58

Asexual Reproduction in the Burning Bush,

Euonymus atropurpureus

Lytton J. Musselman . 60

Seasonal Change in a Sand Prairie in Van Buren
County, Michigan

Dan Pokora . 62

Announcement . 66

The Duckweeds of Minnesota

Howard L. Clark & John W. Thieret . 67

Publications of Interest . 76

The Spring Beauties ( Claytonia ) in Michigan

Edward G. Voss . 77

New Localities for Mosses Rarely Found in
Michigan’s Upper Peninsula

Maynard C. Bowers . 93

Michigan Plants in Print . 94

Editorial Notes . 95

“Winter Wildflowers” . 95

(On the cover: Woods at the south edge of Ypsilanti,
Washtenaw Co., Michigan, carpeted in the spring
with Claytonia virginica, the narrow-leaved Spring Beauty.
Photo by A. H. Smith.)

3

Ti 3

THE

Vol. 7, No.

MICHIGAN BOTANIST

May, 1968

THE MICHIGAN BOTANIST-Published by the Michigan Botanical Club four times per

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Region may be sent to the editor in chief. The attention of authors preparing manuscripts
is called to “Information for Authors” (Vol. 6, p. 202; reprints available from the editor).

Editorial Board

Edward G. Voss, Editor in Chief

Herbarium, The University of Michigan, Ann Arbor, Michigan 48104

Laura T. Roberts, Business and Circulation Manager
1509 Kearney Rd., Ann Arbor, Michigan 48104

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1968

THE MICHIGAN BOTANIST

99

THE STATUS AND DISTRIBUTION OF
GENTIAN A LINEARIS AND G. RUBRICAULIS

IN THE UPPER GREAT LAKES REGION1

James S. Pringle

Royal Botanical Gardens, Hamilton, Ontario.

During the preparation of my monograph on Gentiam, section Pneumo-
nan thae, in eastern North America (Pringle, 1967a), it became evident that
especially interesting problems in plant geography and evolutionary botany were
associated with two closely related species, G. linearis Froel. (Narrow-leaved or
Bog Gentian) and G. rubricaulis Schwein. (Red-stemmed Gentian). These are
illustrated in Figures 1 and 2; their ranges are shown in Figure 3. Descriptions of
these and related species have previously been published (Pringle, 1967a). Details
of their distribution have remained obscure, in part because they have often
been confused with each other and with several related species, and in part
because, until recently, few specimens had been collected from large portions of
their ranges. The distinctness of G. rubricaulis from G. linearis and hence its
proper taxonomic status have also remained in question.

This paper deals with the problems mentioned above, and with the nature
of the isolation which permits these species to retain their distinctness. It is
intended in part to lay the groundwork for a later paper on the origin, relation¬
ships, and migrational history of G. rubricaulis, which will report the results of
studies currently in progress. The origin of G. rubricaulis is a problem of particu¬
lar interest, because this species is restricted to territory glaciated during the
Wisconsin period, with no outlying populations indicating a survivium from
which it could have spread as the glaciers receded.

In my studies of Gentiana, I have examined the specimens in about eighty
herbaria, and have observed and collected plants in the field in many parts of
North America. This study of G. linearis and G. rubricaulis has included field
trips to many localities in northern Michigan, including all extensive areas of
granitic soils in the Upper Peninsula, as indicated by Martin’s maps (1957), to
adjacent portions of Wisconsin, and to a number of areas north and east of Lake
Huron in Ontario.

As Figure 3 indicates, G. linearis is primarily an eastern species. It was
apparently first collected in the Lake Superior region by Loring, who encoun¬
tered it between the Pic River and Sault Ste. Marie in Ontario, probably about
the middle of the nineteenth century. It does not appear to have been collected
in this region again until 1935, when it was discovered along the Batchawana
River in Ontario ( Taylor et al.; specimens cited below). It was first collected in
Michigan in 1942 {Bingham). The occurrence of G. linearis in the Lake Superior
region has remained in question up to the present, since few specimens of it had

^Contribution from the Royal Botanical Gardens, Hamilton, Ontario, No. 8, and
from the University of Michigan Biological Station.

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THE MICHIGAN BOTANIST

Vol. 7

Pig. 1. Gentiana linearis. Fig. 2. G. rubricaulis. Upper parts of flowering stems, x Vi (above),
and corollas slit longitudinally and pressed, with connate anthers separated, x 1 (below).

1968

THE MICHIGAN BOTANIST

101

Fig. 3. Distribution of Gentiana linearis and G. rubricaulis.

102

THE MICHIGAN BOTANIST

Vol. 7

been collected in this region until the present study was undertaken, and
most of these had been deposited in smaller herbaria. Fernald (1950) included
the Lake Superior region in its range;2 Gleason and Cronquist (1963), however,
indicated that its range extended westward only to New York and West Virginia.
The westernmost station2 mapped by Gillett in 1963 was in the eastern part of
northern Ontario. The present paper documents the occurrence of G. linearis in
the Lake Superior region, and includes the results of a study of the extent of its
occurrence in this region, the size and vigor of its populations, and the ecologic
basis of its restricted distribution.

The following list includes all the specimens of G. linearis I have encoun¬
tered which were collected west of Sault Ste. Marie. Specimens of other species,
and of G. linearis from other regions, have previously been listed (Pringle,
1967b). Abbreviations for herbaria follow Index Herbariorum, ed. 5, with the
additions of FORD, for the herbarium of the Ford Forestry Center, Michigan
Technological University, and HMC, for the herbarium of the Huron Mountain
Club.

MICHIGAN: BARAGA CO.: Haypress Dam meadow, T50N R30W, Bourdo 15 Aug
1961 (FORD, MSC, UWM); along Huron Bay-Peshekee Road, center of Sec. 26, T50N
R31W, Pringle 808 (HAM, MICH). MARQUETTE CO.: shore, Ives Lake, Bingham July
1942 (BLH); Yellow Dog Plain, in former river bed, T50N R28W, Hyypio 292 (MSC, TEX);
about small, drying lake in Yellow Dog Plain, Sec. 9, T50N R29W, Stoutamire 3129 (BLH,
HMC); along Marquette Co. Road 607, 1.0 mi s of jet with road to Lake Arfelin pub he
access, Sec. 27, T49N R30W, Pringle 399 (DAO, HAM, MICH); by road to Lake Arfelin
public access nr crossing of Ephraim Creek, Pringle 389 (HAM, MICH); s shore Ives Lake,
Pringle & Harris 400 (HAM, HMC, MICH); just n of Swanzy Lake, Old Swanzy, Pringle 426
(DAO, HAM, MICH); shore of Harvey Lake, Sec. 31, T47N R25W, Pringle 837 (ACAD,
DAO, HAM, MICH, OS, WUD).

ONTARIO: ALGOMA or THUNDER BAY DIST.: Pic to Sault, Loring no date (GH).
ALGOMA DIST.: Batchawana Falls, Taylor et al. 1335 (CAN, DAO, MT, TRT, US); Batcha-
wana River, Taylor et al. 1336 (WASH), 1338 (GH); w shore of small lake situated between
Otter Lake and Hwy. 17, Lake Superior Provincial Park, Cowell 281 (TRT); drained beaver
lake beside old logging road, Lost Lake, Lake Superior Provincial Park, Cowell 301 (TRT).
THUNDER BAY DIST.: Thunder Cape, Sibley Peninsula, w side at lake level, Love & Love
6953a (WIN).

G. linearis is highly localized in Michigan and northwestern Ontario. With¬
in some limited areas, however, it is quite abundant, and forms sizeable popula¬
tions. It is, for example, a common species in the highlands of Marquette and
Baraga Counties northwest of Champion, Michigan, where it not only occurs in
relatively stable communities but also soon colonizes newly available sites. In
such areas G. linearis is obviously a well-established component of the native
flora. Its restricted distribution in the Upper Great Lakes region does not appear
to be due to recent arrival, nor does the species appear to be a relict which is
being eliminated by changing conditions. Instead, it appears that a limitation of

2

And also Minnesota; Minnesota reports, however, appear in all cases to have been
based on misidentifications or failure to distinguish between G. linearis and G. rubricaulis.
3

Of G. linearis sensu stricto— Gillett’s G. linearis ssp. linearis.

1968

THE MICHIGAN BOTANIST

103

suitable habitat is largely responsible for its restricted distribution. The probable
nature of this limitation is discussed later.

Some botanists have assumed that G. linearis and G. mbricaulis intergrade
in the Great Lakes region [although Fernald (1935) called G. mbricaulis “one of
the most distinct of species”] . Among the objectives of my study has been to
determine to what extent, if any, these species do intergrade, and what kind of
isolation permits them to retain their distinctness. Breeding experiments indicate
that hybridization between G. linearis and G. mbricaulis is genetically possible.
It remains to be seen whether such hybrids will be fertile; however, their fertility
may reasonably be predicted in view of the similarity of the chromosomes of G.
linearis and G. mbricaulis (Rork, 1949; Pringle, in press) and the general interfer¬
tility among species in this group. There appear to be no seasonal or mechanical
barriers to their hybridization in nature. Their blooming seasons overlap where
they are geographically sympatric, and both species are pollinated by bumble¬
bees. (Plants of these species are self-fertile, but pollination depends on an
outside agent.) The anthers and stigmas of the two species are similar in position.

Despite the absence of intrinsic isolating mechanisms, G. linearis and G.
mbricaulis appear hardly ever to interbreed in nature. Two specimens from
Marquette County, Michigan, one from Ives Lake and one from Gwinn, have
previously been cited as possibly being of hybrid origin (Pringle, 1967a). The
Ives Lake specimen [Bingham July 1942 (BLH)] was collected in an area nota¬
ble for its diversity of habitats (Hagenah, 1963). Both G. linearis and G.
mbricaulis have been collected at nearby localities. In general the specimen is
characteristic of G. linearis , but its involucral leaves are wider toward the base
than is usual in this species, and they appear to have enveloped the calyces
somewhat. These traits suggest possible introgression of genetic material from G.
mbricaulis. In 1965, however, the small population at Ives Lake consisted only
of entirely characteristic G. linearis, with narrow, spreading involucral leaves
[Pringle and Harris 400 (HAM, HMC, MICH)] .

A specimen collected 16 Aug 1908 by Harrison and Harrison “in shade
along Halfway Creek northeast of township road, Gwinn” (GH)4 seems very
likely to be a hybrid between G. linearis and G. mbricaulis. This specimen
consists of two stems bearing marcescent corollas and ripe fruits. The leaves, like
those of G. linearis , increase only slightly in length and width toward the sum¬
mits of the stems, and appear to have been similar to leaves of G. linearis in
texture and color. The involucral leaves, although no wider than the involucral
leaves of occasional plants of G. linearis, are widest toward their bases, and they
evidently enveloped to a degree the bases of the inflorescences, as do the leaves
of G. mbricaulis. The calyx lobes are similar in size and shape to those of G.
mbricaulis. The corolla lobes are as long as those of G. mbricaulis, but are
strongly cucullate toward the outer edge, like those of G. linearis. This specimen
is like G. linearis in having had ripe fruits as early as August 16, and the seeds, in

In previous papers (Pringle 1967a, b) I erroneously cited the Harrisons’ 7 Aug 1908
collection from Tamarack Street, Gwinn (GH) as the possible hybrid. Specimens in this
collection are entirely characteristic of G. mbricaulis, as were all the plants in the Tamarack
Street population in 1965 [Pringle 394 (HAM, MICH)] .

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being relatively short and elliptic rather than oblong, are like those of G. linearis.
This specimen was probably collected along the creek just northeast of its cross¬
ing by what is now Highway M-35. Considerable building and recreational use of
the remaining open land have taken place along this part of Halfway Creek since
1908. In searches along this portion of the creek in 1965, and along relatively
undisturbed stretches further upstream in 1967, I was unable to find any gen¬
tians. Evidently no extensive population of intermediates has become estab¬
lished.

Gillett (1963) viewed G. linearis and G. rubricaulis (as G. linearis ssp.
linearis and ssp. rubricaulis) as two subspecies with contiguous, rather than
overlapping, ranges. Specimens from the vicinity of Sudbury, Ontario, were said
to be intermediate, indicating intergradation where the ranges met. For this
reason he considered the two taxa best recognized as two subspecies of a single
species. From correspondence with Dr. Gillett, and from searches of the herbaria
he used, it seems most likely that the specimens involved are Watson 1909 and
1914 (TRT), from Timagami Provincial Forest, Nipissing District. The leaves and
calyx lobes of these specimens are proportionately wider than is common in G.
linearis and are narrowly elliptic rather than strictly linear. Otherwise these
specimens appear typical of G. linearis. Such leaves occur occasionally through¬
out the range of G. linearis, especially on plants in shaded sites. These specimens
should probably be interpreted as representative of variation within G. linearis s.
str., rather than indicative of intergradation with G. rubricaulis.

I have found no other specimens at all intermediate between G. linearis
and G. rubricaulis. Moreover, G. linearis in its outlying populations in the Fake
Superior region does not appear to differ in any morphologic aspect from the
species in other parts of its range. Evidently, therefore, the situation is neither
one of clinal intergradation from G. linearis extremes in the East to G. rubri¬
caulis extremes in the Upper Great Fakes region nor one of extensive hybridiza¬
tion.

Even if the few specimens cited above do indicate that hybridization be¬
tween G. linearis and G. rubricaulis has occurred, this hybridization would
scarcely constitute by itself an adequate basis for classifying these two taxa as a
single species. There is a remarkable amount of interfertility within Gentiana,
and hybridization is not uncommon, even among highly diverse species. (See
Figure 7 and Pringle 1965, 1966, 1967a.) Indeed, considering how frequently
some Gentiana species hybridize, the distinctness between G. linearis and G.
rubricaulis seems especially sharp.

The morphologic differences between G. linearis and G. rubricaulis appear
to be comparable to those among other generally recognized species of Gentiana
in North America. The descriptions of the eastern North American species of
Gentiana, section Pneumonanthae, in my recent monograph (Pringle, 1967a)
provided a convenient basis for an objective approach to this matter. The
thirteen species descriptions in this paper closely parallel one another in format,
and were designed to deal with all of the readily discernible traits of gross
morphology in which any of these species differ from any others. Based on
observations of several thousand specimens, these descriptions indicate the para-

1968

THE MICHIGAN BOTANIST

105

meters of variation in about forty traits of presence or absence, size, shape,
color, puberulence, and coherence of floral and vegetative organs. Dissimilarity
values indicating the extent of the differences between any two species were
obtained by comparing the two species trait by trait, following the monograph
descriptions. Values were derived by adding points as follows: 0, if there was no
appreciable difference in the trait being considered; 1, if there was a slight to
moderate difference in the extremes reached, but considerable overlapping of
the range of variation; 2, if there was a greater difference in the extremes but
still considerable overlapping; 3, if the extremes and means were distinctly dif¬
ferent but overlapping could still be found; 4, if there were moderate differ¬
ences, with no overlapping in the ranges of variation; and 5, for extreme differ¬
ences. A value of 53 was obtained in contrasting G. linearis with G. mbricaulis.
Values for other comparisons of closely related species include: G. clausa - G.
andrewsii (exclusive of forms derived through introgression), 35; G. saponaria -
G. catesbaei, 37; G. alba - G. villosa, 57; G. clausa - G. saponaria , 60. Comparing
less closely related species produced divergence values of 70 for G. autumnalis -
G. puberulenta, 97 for G. autumnalis - G. villosa, and 1 14 for G. autumnalis - G.
andrewsii. In contrast, a comparison of the two varieties of G. andrewsii resulted
in a dissimilarity value of only 10. In summary, then, both the sharpness of the
distinction between G. linearis and G. rubricaulis and the extent of the differ¬
ences between them appear to justify their recognition as two species, following
the treatments in the commonly used manuals for northeastern North America.

As Mayr (1942) and Levin (1967) have pointed out, a clinal gradient
may be expected between races which have differentiated gradually, without a
history of complete separation. In contrast, sympatric, closely related entities
would be expected to remain relatively distinct if they had spread from different
centers; just how distinct would depend on how different were the ecologic
conditions to which they had become adapted while they were isolated geo¬
graphically. Sharp delimitation from related taxa may also be expected of spe¬
cies which have arisen abruptly, as through interspecific hybridization, if they
have been isolated genetically (as with allopolyploid hybrid derivatives) or other¬
wise from related species since their origin. The sharp distinction between G.
linearis and G. rubricaulis is contrary to what one would expect if the latter were
merely an ecotype which had differentiated gradually as G. linearis spread west¬
ward. From a phylogenetic viewpoint of plant classification, the recognition of
G. rubricaulis as a species may be further supported if current studies indicate
that the origin and development of G. mbricaulis were distinct from the history
of G. linearis.

Since the isolation of G. linearis from G. mbricaulis in the Upper Great
Lakes region seems to be due primarily to extrinsic factors, their ecologic posi¬
tions have been compared. Figure 4 shows a typical G. linearis habitat along
Ephraim Creek, northwest of Champion, Michigan. This is an open, wet valley
among rocky hills which are vegetated largely by white spruce [Picea glauca
(Moench) A. Voss] . The gentians occur in a community dominated by grasses
and sedges, above the level of standing water, below the spruces. Other habitats
include moist places along road cuts in this area. In the sandy out wash plains to

106

THE MICHIGAN BOTANIST

Vol. 7

the southeast, the gentians occur around kettles, again among grasses and sedges
in a fringe between the flooded communities below and the dry hills above. In
all cases, the sites are open or partially shaded, constantly moist but never
inundated.

G. linearis is most abundant in the Appalachians and in the highlands of
the northeastern states and Quebec. In these regions the soils are predominantly
strongly acid, and have developed from granite, gneiss, and similar crystalline
rocks with very low base contents. West of the Appalachians, granitic soils are
far less common, and are largely restricted to northern latitudes, where rocks of
the Canadian Shield are near the surface. South of Lake Superior, G. linearis
occurs only in northwestern Michigan, where outlying portions of the Canadian
Shield are located. From Figure 5, which shows the distribution of G. linearis in
northwestern Michigan in relation to geologic factors, it is evident that G.
linearis in this region is closely associated with granitic soils. It occurs in areas of
lithosols and ground till derived from underlying granite; moraines which, from
their positions in relation to areas of granitic rock and directions of glacial
movement, may be assumed to be largely granitic in content; and out wash plains
directly “downstream” from such granitic moraines.

Like G. linearis, G. rubricaulis grows in open or partially shaded sites
which are consistently moist but not inundated. Probably its most frequent
habitats are grass-sedge meadows along the sandy shores of large lakes. It also

Fig. 4. Habitat of Gentiana linearis along Ephraim Creek, northwest of Champion, Mar¬
quette County, Michigan.

1968

THE MICHIGAN BOTANIST

107

occurs in moist meadows and thickets along streams, and occasionally, especially
in northwestern Michigan and Wisconsin, where it is most abundant, in swamps
and bogs and around kettles in outwash plains. G. rubricaulis grows in both
alkaline and moderately acid soils, but not ordinarily in soils of granitic origin.
In the area mapped in Figure 5, G. rubricaulis is associated with moraines de¬
rived from the largely non-granitic Huronian rocks, with outwash from such
moraines, and with sandy lacustrine deposits of mixed, often highly calcareous,
content.

North of the Great Lakes region, G. linearis likewise appears to be closely
associated with granitic areas, while G. rubricaulis is associated with non-granitic
areas (Figure 6). G. linearis occurs only in the eastern part of this region, where
soils of granitic origin are predominant. G. rubricaulis occurs in the central and
western parts. Although the bedrock in the region between the Upper Great
Lakes and the James Bay Lowlands is the prevailingly granitic Canadian Shield,
there are many areas, some of them extensive, with non-granitic soils. North of
Lake Huron there is the extensive Clay Belt, a region of deep lacustrine deposits
in the bed of glacial Lake Barlow-Ojibway. The origin of these sediments, at
least in the western and central parts of the Clay Belt, was primarily the calcar¬
eous rocks of the James Bay Lowlands. Another large lake bed is that of glacial
Lake Agassiz, north and west of the Lake of the Woods, also with largely
non-granitic sediments. The granite of the Canadian Shield has been abundantly
disrupted west of the Clay Belt and the James Bay lowlands by basic intrusives,
so that even in areas of lithosols non-granitic soils may prevail locally. Dutilly et
al. (1954) have observed that in areas in the upper reaches of the Albany and
Attawapiskat watersheds where they collected G. rubricaulis , the presence of
such species as Thuja occidentalis L. and Ribes hudsonianum Michx. indicated
that the soils were not strongly acid.

The distributional data discussed above indicate that G. linearis and G.
rubricaulis probably remain as sharply distinct as they do where they are geo¬
graphically sympatric primarily because they grow in localities which differ in
soil types. The effectiveness of adaptations to contrasting soil types as an isolat¬
ing mechanism has been amply demonstrated (Mayr, 1947; Muller, 1952; Riley,
1952).

These gentians are restricted not only to certain conditions of soil chemis¬
try but also to certain conditions of moisture supply, drainage, and openness.
Consequently, populations of these species, especially G. rubricaulis , are general¬
ly small and widely separated. This discontinuity of distribution doubtless en¬
hances the effect of ecologic isolation. The effectiveness of distances of as little
as a few hundred feet as barriers to gene flow between populations of insect-pol¬
linated plant species has been demonstrated by Cook (1962). Keener (1966) has
provided an example, from the Appalachian shale barrens, of sympatric, interfer-
tile species of Clematis which appear to have remained sharply distinct almost
entirely because they occur in small, scattered populations. Because of the simi¬
larly discontinuous distribution of G. linearis and G. rubricaulis, the effects of a
successful hybridization between them would probably remain confined to the
population in which it occurred, paralleling the situation among species of Phlox
observed by Levin (1967, p. 106).

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THE MICHIGAN BOTANIST

Vol. 7

Direction of glacial movement, as
indicated by striae

o s 10 15 miles

I - S - L- T - 1

o io 20 kilometers

# Gentiana linearis
O Gentiana rubricaulis

I Baraga Co.

/V

_ _ I - ^

Bedrock (where overlain by transported
material)

Granite

Limestone and dolomite
Other rocks

Areas of outcrops and lithosols
iESpll Granitic

Non-granitic

Glacial and postglacial deposits

EZ

Outwash and sandy lacustrine deposits
Clayey lacustrine deposits
Dune-swale complex

Other areas - moraines and ground
till

Marquette
Dickinson Co.

Fig. 5. Distribution of Gentiana linearis
Michigan, in relation to geologic factors.

and G. rubricaulis in three counties in northern
Geology from Martin (1957).

1968

THE MICHIGAN BOTANIST

109

The distribution of G. linearis is paralleled to various degrees by those of
certain other species which are relatively common in the Appalachians, the
Laurentides, the Madawaska Highlands of eastern Ontario, and other parts of
northeastern North America, and also occur sporadically westward to the Lake
Superior region. Among these species are Potamogeton confervoides Reichenb.,
Oxalis montana Raf., and Aster nemoralis Ait. From field observations, habitat
notes in manuals, and correlations of the distribution patterns of these species
with geologic information, it appears that at least in some cases the rarity of
these species westward may be due largely to their being restricted to granitic or
strongly acid soils.

There are also a number of pairs of closely related species or varieties with
ranges overlapping in the Great Lakes region, of which one member is associated
with granitic or acid soils, while the other is found in calcareous soils. In some
cases, distribution patterns indicate that the oxylophyte has spread into the

Fig. 6. Distribution of Gentiana linearis and G. rubricaulis north of the Great Lakes, in
relation to geologic and soil factors. Information on geology and soils largely from Hills
(1960, 1962), Hills and Morwick (1944), and the Atlas of Canada (1958).

110

Vol. 7

THE MICHIGAN BOTANIST

Great Lakes region from the largely granitic Appalachians, while the calciphyte
has spread from a non-granitic trans-Appalachian survivium (cf. litis, 1965, pp.
149-151). Interfertility between the two members of a pair is known in some of
these cases, and may reasonably be expected in others. Therefore, it seems
probable that the ecologic isolation which appears to be largely responsible for
the distinctness of G. linearis and G. rubricaulis has a number of parallels. One
such parallel has become evident through my studies of Gentiana. G. clausa Raf.
and G. andrewsii Griseb., two similar but distinct species with ranges which
overlap from Vermont to Ohio, also appear to be isolated from each other by
adaptations to granitic and nongranitic soils, respectively. Other such pairs in¬
clude Lilium camdense L. and L. michiganense Farw. [L. canadense ssp. mich-
iganense (Farw.) Boivin & Cody] ; Viola blanda Willd.and V. incognita Brainerd;
and Prunus pumila L. var. cuneata (Raf.) Bailey [P. susquehanae Willd.] and var.
pumila. Detailed studies of the distribution of these species in relation to geo¬
logic and soil factors might well reveal similar ecologic isolation.

Fig. 7. Hybridization among Gentiana spp. of eastern and central North America. Abbrevia¬
tions are the first three letters of the specific epithets of G. affinis Griseb., G. alba Muhl., G.
andrewsii Griseb., G. austromontana Pringle & Sharp, G. autumnalis L., G. catesbaei Walt.,
G. clausa Raf., G. decora Pollard, G. linearis Froel., G. pennelliana Fern., G. puberulenta
Pringle, G. rubricaulis Schwein., G. saponaria L., and G. villosa L. Positions of symbols
represent approximations of morphologic similarity and presumed phyletic affinity, diverg¬
ing from probable primitive species at the upper left, with solitary flowers, open corollas,
large corolla lobes, and prolonged, subequally bifid corolla appendages. Heavier lines indi¬
cate natural hybridization known from five or more localities; lighter lines, natural hybridi¬
zation known or suspected from fewer than five localities; broken lines, only artificial
hybridization known.

1968

THE MICHIGAN BOTANIST

111

SUMMARY

Gentiana linearis Froel. and G. rubricaulis Schwein. both occur in the Upper Great
Lakes region. Although these species are closely related, they do not intergrade. They are
evidently ecologically isolated, G. linearis growing in soils derived from granite and similar
low-basic rocks, but G. rubricaulis in non-granitic soils. The sharpness and the extent of
their differences seem comparable to those among other generally recognized species of
Gentiana, and, along with features of the distribution patterns of these species, indicate that
G. rubricaulis probably cannot be interpreted merely as an ecotype of G. linearis which
differentiated as G. linearis spread northwestward during Wisconsin glacial recession.

ACKNOWLEDGEMENTS

Preliminary versions of the manuscript were read by Prof. Douglas R. Lindsay, of
Lakehead University, and Dr. S. Galen Smith, of the Wisconsin State University, White-
water, to whom I am grateful for their valuable comments and suggestions. I am also
grateful to the many curators and librarians who have made specimens and literature availa¬
ble, and to Dr. Eric A. Bourdo, Jr., Mr. and Mrs. William P. Harris, Jr., and the Huron
Mountain Wildlife Foundation for their hospitality and assistance while I was collecting in
northern Michigan in 1965.

LITERATURE CITED

Atlas of Canada. 1958. Canada Department of Mines and Technical Surveys, Ottawa. 110
maps.

Cook, S. A. 1962. Genetic system, variation, and adaptation of Eschscholtzia californica.
Evolution 16: 278-299.

Dutilly, A., E. Lepage, & M. Duman. 1954. Contribution a la flore du versant occidental de
la baie James, Ontario. Contr. Arct. Inst. Cathol. Univ. 5F. 144 pp.

Fernald, M. L. 1935. Critical plants of the Upper Great Lakes region of Ontario and
Michigan. Rhodora 37: 197-222, 238-262, 272-301, 324-341.

- . 1950. Gray’s Manual of Botany, ed. 8. Am. Book Co., New York, lxiv + 1632 pp.

Gillett, J. M. 1963. The Gentians of Canada, Alaska and Greenland. Research Branch,
Canada Dep. Agr. Publ. 1180. 99 pp.

Gleason, H. A., & A. Cronquist. 1963. Manual of Vascular Plants of Northeastern United
States and Adjacent Canada. D. Van Nostrand Co., Princeton, lii + 810 pp.

Hagenah, D. J. 1963. Pteridophytes of the Huron Mountains, Marquette County, Michigan.
Mich. Bot. 2: 78-93.

Hills, G. A. 1960. The soils of the Canadian Shield. Agr. Inst. Rev. 15(2): 41-43.

- . 1962. Soil-vegetation relationships in the Boreal Clay Belts of eastern Canada. In: W.

K. W. Baldwin, ed. Report on Botanical Excursion to the Boreal Forest Region in
Northern Quebec and Ontario. Canada Dep. Northern Affairs and Natl. Resources,
pp. 39-53.

- , and F. F. Morwick. 1944. Reconnaissance soil survey of parts of northern Ontario.

Ont. Soil Surv. Rep. 8. 56 pp + 3 maps.

litis, H. H. 1965. The genus Gentianopsis (Gentiana ceae): Transfers and phytogeographic
comments. Sida 2: 129-154.

Keener, C. S. 1966. A biosystematic study of Clematis subsection Integrifoliae (Ranun-
culaceae). Ph.D. thesis, North Carolina State University at Raleigh, vi + 144 pp.

Levin, D. A. 1967 Variation in Phlox divaricata. Evolution 21: 92-108.

Martin, H. M. 1957 Map of the Surface Formations of the Northern Peninsula of Michigan.

Mich. Dep. Conservation, Geol. Surv. Div., part of Publ. 49.

Mayr, E. 1942. Systematics and the Origin of Species, from the Viewpoint of a Zoologist.
Columbia University Press, New York. Republished 1964 by Dover Publications, New
York. xviii+ 334 pp.

- . 1947. Ecological factors in speciation. Evolution 1: 263-288.

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Muller, C. H. 1952. Ecological control of hybridization in Quercus: A factor in the mechan¬
ism of evolution. Evolution 6: 147-161.

Pringle, J. S. “1964” [1965] . Preliminary reports on the flora of Wisconsin No. 53. Genti-
ana hybrids in Wisconsin. Trans. Wis. Acad. 5 3: 273-281.

- . “1965” [1966]. Hybridization in Gentiana (Gentianaceae) : A resume of J. T. Curtis’

studies. Trans. Wis. Acad. 54: 283-293.

- . 1967a. Taxonomy of Gentiana, section Pneumonanthae, in eastern North America.

Brittonia 19: 1-32.

- . 1967b. List of specimens examined in the preparation of the paper: Taxonomy of

Gentiana, section Pneumonanthae, in eastern North America. Department of Botany,
University of Tennessee, Knoxville, [i] +44 pp., Xerox.

- . In press. Documented plant chromosome numbers 1968:1. Sida.

Riley, H. P. 1952. Ecological barriers. Am. Nat. 86: 23-32.

Rork, C. L. 1949. Cytological studies in the Gentianaceae. Am. Jour. Bot. 36: 687-701.

ANNOUNCEMENT

SOME COMMON MUSHROOMS OF MICHIGAN'S
PARKS AND RECREATION AREAS

By Alexander H. Smith & Helen V. Smith

Special Publication No. 1, Michigan Botanical Club
Third Printing (1968) available soon.

This popular pocket-size publication (80 pp., 55 photos) is
available for $1.25 including postage and handling from Miss Barbara
L. Bowen, 1003 Brooks, Ann Arbor, Michigan 48103. Make remit¬
tances payable to Michigan Botanical Club; write for quantity rates.
Almost 8,000 copies have been sold to date.

Proceeds from the sale of this non-technical guide to common
and conspicuous mushrooms of our vacation areas are being used for
the purchase of plant sanctuaries.

1968

THE MICHIGAN BOTANIST

113

TERATOLOGICAL STAMENS AND CARPELS
OF A WILLOW FROM NORTHERN MICHIGAN

Warren H. Wagner, Jr.

Botanical Gardens, The University of Michigan, Ann Arbor

The study of monstrosities has always intrigued biologists for their possi¬
ble bearing on our understanding of evolution and growth. Collectors of plants
in the Great Lakes region occasionally hear of aberrant specimens, even in the
newspapers. I wish to describe and discuss here an example of a phenomenon
which has previously been reported for flowers of willows, the sex organs of
which, stamens and carpels, show more or less intergradation. These specimens
provide fine demonstrations of the correspondence of parts in the two types of
organs, and it is my hope that readers will be encouraged to make further
searches for and analyses of such phenomena.

On May 22, 1954, along a roadside about 2 miles south of Germfask,
Schoolcraft Co., in the Upper Peninsula of Michigan, Dr. E. G. Voss collected a
specimen (1934) of the long-beaked willow, Salix bebbiana Sarg., which proved
to have monstrous flowers. It was growing with normal staminate and pistillate
willows of the same species; subsequent efforts to rediscover the plant in ques¬
tion have failed. Dr. Voss gave me some dried flowers which I cleared in 5
percent solution of sodium hydroxide to make the observations to be reported
below. By gently staining the specimens and mounting them in diaphane on a
microscope slide I could make out the corresponding parts more definitely.

THE NATURE OF PLANT TERATOLOGIES

Before describing the floral organs of this collection, I should make some
general comments on the nature of monstrous developments in plants. One may
well ask “Of what scientific interest are these freaks?” This is a fair question and
one which deserves an answer, even if it is only a tentative one. It should be
pointed out that there are all sorts of peculiar formations possible, and they are
due to a multitude of causes, some of which we understand to some extent and
some which remain entirely obscure. In 1964 the Ann Arbor News reported a
plant of evening primrose, Oenothera biennis, discovered by a fourth-grade stu¬
dent, the photograph of which showed a massive club of flowers nearly a foot
across. This phenomenon, which is found occasionally not only in evening prim¬
rose but also in such other plants as thistles, dandelions, mullein, sumacs, and
blackberries, is known as “fasciation.” The growing tip of the plant simply
expands in one dimension as it grows, becoming broader and broader. So far as I
know, the cause of this type of monstrosity is not yet known, but the condition
in the familiar cockscomb (a cultivar of Celosia argentea) of the garden is con¬
trolled genetically and the monstrous development is fixed.

Fasciation is but one of a series of growth irregularities known in plants.
For example, especially in ferns, there are numerous hybrids in which the parts
develop asymmetrically and irregularly. In these, the abnormal shape appears to
be related to a failure of the hereditary factors that control form to interact with

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harmony (Wagner, 1962). The “doubled” flowers of many garden plants also
have a genetic basis; in these, (as is illustrated especially well by the roses) many
of the stamens have become transformed into petals, so that instead of having
only five petals, the wild condition, the flower may have dozens of petals.

Sometimes the peculiarities of development are controlled by purely en¬
vironmental or extraneous factors. There are certain experimental tobacco hy¬
brids (Whitaker, 1924) which produce tumor-like outgrowths that are genetic in
origin, but similar structures can also be induced artificially in normal plants by
applying certain plant hormones (Whitaker, 1924; Brown & Gardner, 1937). As
we have seen in the past decade, the use of weed killers may produce much
altered growth patterns in plants if they survive. Strangely shaped plant speci¬
mens may also result from damage caused by mowing or by cattle or other
domestic animals stepping upon the new, embryonic shoots (cf. Wagner, 1966).
The possible causes for abnormal growth seem to be numerous. The resulting
abnormalities range from organs which are nearly typical, all the way to com¬
pletely disorganized plant tissues or tumors. For our purposes here, we shall
confine our discussion to those abnormal plant parts which, although deformed,
still bear some resemblance to the normal condition and can still be defined
approximately as “leaves,” “stems,” or “flowers.”

When we say “causes" we still have to recognize the superficiality of
referring to such vague concepts as “genetic factors” or “environmental factors”
as real causes. The basic questions involve what actually happens or what is upset
in the process of forming the plant or its parts. The study of monstrous develop¬
ment is known as “teratology,” and in recent years the subject has been fairly
unpopular in botany.1 Part of the unpopularity is due to our discouraging lack of
clear understanding of plant development (all the way from genic action to
tissue and organ formation).

There have been wild speculations about monstrosities. It is an old idea in
biology that teratologies represent “throw-backs” or “atavisms” which show
what ancestors were like. Heslop-Harrison (1952), who is one of the recent
authors who grappled with this idea, concluded that “abnormalities may have
significance in interpreting homologies of organs, but cannot be regarded as
atavistic” (italics mine). This will be the basis of the interpretation of the willow
flowers to be given below.

To illustrate how teratology may be used to clarify what are homologous
organs, we can consider the familiar anomalies in roses in which floral parts are
formed intermediate between sepals and petals, and between petals and stamens.
Such specimens show that the floral appendages all belong to the same general
category as the leaf. There is nothing particularly atavistic about these intermedi¬
ate structures; indeed, all they show is that you can get intergrades between
existing organs, not some organs of some ancestor of the distant past.

Other teratologies of roses are especially interesting when they involve the

medicine (perhaps as a result of the thalidomide tragedy which disfigured many
hundreds of children in Europe), the study of teratology and related subjects seems to have
undergone a renaissance, and it is possible that the same will happen in botany, especially as
a result of newer experimental techniques for studying plant development.

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growth of the whole flower; one of the theories of the flower is that it is a
determinate shoot, i.e., that it corresponds to a vegetative shoot but its leaves
become the floral parts and when the female organs are produced it stops grow¬
ing. Occasionally in roses the shoot “grows through”: after producing the floral
parts it continues to grow, but now produces ordinary foliage leaves. In such
specimens we have the unusual picture of a rose flower borne along a shoot, with
normal foliage leaves below the flower and then coming out of the top of the
flower!

Occasionally such abnormalities have been used to demonstrate homolo¬
gies and to help resolve controversies about the nature of certain reproductive
organs. The seed-bearing structure of the ginkgo tree, for example, has long been
subject to speculation— is it a modified “sporophyll,” i.e., fertile leaf, or is it a
naked stem? Structures intermediate between the fan-shaped foliage leaves and
stalk-like seed structures have been discovered which tend to support the inter¬
pretation that the “ovuliferous structure” of ginkgo is, in fact, a much-modified
sporophyll (cf. illustrations in Zimmerman, 1959, pp. 402-403).

The umbrella-shaped spore-bearing organs in the cones of scouring rushes
and horsetails, Equisetum, have also been debated. However, in 1939 Tschudy
reported a complete series of structures from “sporangiophores” (sporecase-bear-
ing organs) to ordinary vegetative sheath leaves, which demonstrated quite clear¬
ly the correspondence of these parts. He found these in teratological cones of
Equisetum telmateia, a western species. More recently, Dr. R. L. Hauke in his
investigations of Michigan Equisetum discovered a splendid example of this phe¬
nomenon in E. x litorale, a hybrid horsetail intermediate between common field
horsetail, E. arvense, and the water horsetail, E. fluviatile. A hybrid colony
covers a large part of an acre in Monroe Co., Michigan, not far from the village of
Milan. There are hundreds of specimens with this abnormality. In this hybrid
horsetail colony, the peculiarity of forming intermediate organs is probably
genetically fixed, and maintained by vegetative reproduction.

THE MONSTROUS WILLOW FLOWERS

When we think of the flowers of willow we think first of the catkins, the
“pussy willows”— clusters of dozens of much reduced florets and hairy parts.
Typically a whole willow plant is either male, the catkins made up of all stam-
inate flowers, or female, the catkins of all pistillate flowers. This arrangement as¬
sures cross-pollination. Interspecific hybridization is frequent and this causes
one of the problems of identifying willows. Willow crosses are common enough
to lead to some confusion between the species.

The willow flowers are borne in the axils of hairy scales or bracts. The
individual flower lacks sepals or petals, but there are usually one or more yellow¬
ish, shiny glands which produce nectar. These are shown at the bases of the three
staminate flowers at the upper left in Figure 1. The stamens of the male flower
number two in the majority of species but may be one or as many as twelve in
some. They are made up of long, delicate filaments topped by an anther of four
pollen sacs. Pollination is accomplished by insects, or by wind, or both. The
female flower is basically similar to the male flower in being borne in the axil of

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a bract along the catkin and possessing one or more glands. Instead of having
separate stamens, however, it has a single pistil, this a more or less stalked ovary
made up of two or four carpels topped by the two- or four-lobed stigma. The
pistil matures into a small capsule from which are produced numerous tiny black
seeds, these floated on the wind by masses of long hairs. Even though the pollen-
or seed-producing flowers lack obvious sepals or petals, the anatomical studies of
Fisher (1928) led to the conclusion that the gland is the remainder of an ancient
perianth, reduced now to a mere papilla and functioning secondarily as a
nectary.2

The Voss collection of Salix bebbiana differs from the normal form of
the species in having both types of sex organs in the same catkin. The sex
organs also show all degrees of intergrades in their characteristics. There is
nothing distinctive about the bracts, wlrich are typically yellowish and slightly
pilose, nor the glands, which are solitary, one per flower. No attempt was made
to estimate the quantitative incidence of the different forms of flowers present,
but Figure 1 illustrates the kinds of organs discovered.

In order to understand this series we can arbitrarily separate organs into
the following forms:

1. Normal male flowers. These are are the three illustrated at the upper left,
and show the basic structure of the stamens. A very long uniformly narrow
filament is topped by the anther with its four pollen sacs. The latter, when
mature, split longitudinally, releasing the pollen. The filament runs directly
into the “connective” which is the extremely narrow band of sterile tissue
connecting the pollen sacs. The top of the connective is seen merely as a
depression between the lobes of the anther.

2. Intermediate form I. These organs differ from typical stamens primarily in
the development of the connective. Now the connective is expanded, both
laterally and terminally. The pollen sacs become more or less separated and
moved around toward one side. There is also a tendency for the filament to
become shorter and broader. An obvious stigma is still not present, the apex
of the connective being represented only by a slightly projecting but naked tip.

3. Intermediate form II. The organ is almost perfectly transitional between

2

It is interesting to note that in the related genus Populus, including aspens, poplars,
and cottonwoods, there is a much more obvious perianth, the so-called “cup-like disk.” In
this genus wind-pollination is the rule. The plants, as in willows, are either male or female,
but in his investigations on Michigan aspens, Dr. Burton V. Barnes found, as have previous
workers (e.g., Erlanson & Hermann, 1928), a number of plants with a few to many bisexual
flowers containing both normal pistils and normal stamens. Teratological organs were also
found in Populus and included what appeared to be mainly fusions between organs-pistil,
filament, and anther fused together; anthers attached to the ovary wall; or 2-3 anthers more
or less joined (Barnes, pers. comm.).

- ►

Fig. 1. Free-hand drawings of flowers and separate sex organs from an unusual plant of Salix
bebbiana. Specimens marked with Roman numerals are examples of different stages of
intermediacy between stamens and carpels. All from one catkin of Voss 1934.

1968 THE MICHIGAN BOTANIST _ U7

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stamen and carpel. Now the pollen sacs (or ovules?) have moved entirely to
one face of the connective, which itself has become much expanded and
more or less inrolled at the margins. Also the beginnings of a stylar elongation
are obvious and a more or less rudimentary stigma occurs. However, carpel-
lary closure is little, if at all, evident.

4. Intermediate form III. Now the organs are obviously more carpel-like than
stamen-like. They differ, however, from carpels in the following ways: (a)
Although the style and stigma appear to be normally formed, fusion is still
not complete in the ovary region so that the ovary is like a tube with a
longitudinal opening along one side, (b) The ovules (or deformed pollen
sacs?) are borne near the base of the opening and are irregular in size and ar¬
rangement.

5. Carpels. These are like those of typical pistillate flowers of Salix beb-
biana , with one very important exception. In these anomalous flowers, the
basic separateness of the organs characteristic of staminate flowers is re¬
tained, so that the “female flower” (i.e., one without any obviously male
structures) appears to have two pistils. The ovary is normally composed of
two carpels which themselves are fused into a compound ovary. It seems
remarkable that the carpels have not fused with each other but have folded
their margins together and have closed upon themselves.

Fig. 2. Diagrammatic illustration of the changes illustrated by the intergradient organs
between stamens and carpels. Background guidelines to emphasize the corresponding parts.

1968

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119

Generally speaking, any given flower will have sex organs of approximately
the same type but there is some irregularity in this respect, as shown in the
drawings of whole flowers, those with two sex organs from the same flower
together.

SIGNIFICANCE OF THE MONSTROUS WILLOW FLOWERS

As early as a half-century ago, Worsdell (1915, pp. 190, 191; pi. xlvi)
commented on similar teratological conditions in willows found in England.
What the morphogenetic significance of these abnormalities may be can only be
speculated upon at this time. Perhaps they result from hybridization, or perhaps
from upsets in the genetic factors which control whether a particular appendage
is going to be a male organ or a female organ. It is conceivable that much can be
learned about the developmental processes of floral organs by detailed studies of
such teratological forms as these, and by comparing different examples of them.
I find it especially interesting that in the present material I observed no fusions
of parts at all, including even the most carpel-like structures, the ones which
would have been most expected to fuse. This is in striking contrast to what
Worsdell reported in “Salix cinerea var. monodelpha,, (op. cit.). In his words
“. . . in the normal male catkins the stamens frequently become united, and this
is constantly the case in the variety monodelpha. These facts clearly prove the
close alliance which exists between stamens and carpels.”

It seems to me that the major significance of the collection described here
is in showing what Worsdell called “the close alliance which exists between
stamens and carpels.” The organs are, it is true, very different from one another
in their typical expressions, and it is hard to visualize the corresponding parts.
Yet in the present study we seem to have excellent demonstration of the ho¬
mologies of parts. The stages illustrate graphically to what part of the stamen
each of the parts of the carpel corresponds— the stigma, the style, the ovary, the
ovules, and the stalk, as shown diagrammatically in Figure 2. It makes evident
how greatly reduced the anther connective is, when we realize that this simple
and narrow band of tissue joining the pollen sacs is homologous to the massive
walls of the ovary, as well as its style and stigma.

If the living remains of primitive flowering plants were not so complete, as
shown in the studies of the late I. W. Bailey and his students of Harvard Univer¬
sity, we might doubt that the simple anther connective could correspond to the
leaf-like walls of the carpel. There are living genera of primitive flowering plants
in which the stamen is normally more or less leaf-like, and the pollen sacs are
borne upon the reduced blades, including genera in such families as Degener-
iaceae, Himantandraceae, and Magnoliaceae. Families like these in which the
“connective” is more or less blade-like also possess such other primitive features
as wood without vessels or with primitive vessels; flowers with a spiral cone-like
arrangement of parts, the parts numerous and of indefinite number; the carpels
also essentially leaf-like; and the pollen of a primitive form with only a single
germination furrow. For students of Great Lakes plants, the families Ranun-
culaceae, Berberidaceae, Lauraceae, and the genus Liriodendron in the Mag¬
noliaceae are local plants of fairly primitive floral organization.

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Normal stamens and carpels of willows, on the other hand, are among the
most specialized forms we know, but a comparison of the Stage I intermediates
described and figured above show a remarkable likeness to some of those of the
“ranalian” families mentioned. The stamens of primitive flowering plants have a
similarly shorter filament, broader and readily visible connective, and pointed tip
that extends beyond the pollen sacs.

It should prove interesting to discover more specimens of willows like
these. Besides the general teratological questions already presented above, there
are others. For example, what of the anatomical changes involved in the transi¬
tion from stamens to carpels? The microscopic structure of the walls of the
normal pollen sacs is very different from that of the ovules. What are the inter¬
mediates like? How do the integuments correspond, if at all, to parts of the
pollen sacs? How does pollen formation intergrade with the formation of the
embryo sac? Such questions would be difficult to answer from the dried material
with which we worked here; and for this reason we should attempt to find (and
mark!) more examples. This means that we must examine each willow specimen
carefully with a good hand lens at the time of flowering. That Salix bebbiana
may be disposed toward abnormalities is already suggested in the reports by
Emma Cole (1901) from Grand Rapids and by Mr. and Mrs. Hanes (1947) from
Kalamazoo County that in this species staminate and pistillate flowers occur
together.

LITERATURE CITED

Brown, N. A. and Gardner, F. E. 1937. Indoleacetic acid galls of a secondary type. Phyto¬
pathology 27: 1110.

Cole, Emma J. 1901. Grand Rapids Flora. Grand Rapids, Mich. 170 pp.

Erlanson, Eileen W., & Frederick J. Hermann. 1928. The morphology and cytology of
perfect flowers in Populus tremuloides Michx. Pap. Mich. Acad. 8: 97-110.

Fisher, Mary J. 1928. Morphology and anatomy of the flowers of the Salicaceae. Am. Jour.
Bot. 15: 307-326; 372-394.

Hanes, Clarence R., & Florence N. Hanes. 1947. Flora of Kalamazoo County, Michigan.
Schoolcraft, Mich. 295 pp.

Heslop-Harrison, J. 1952. A re-consideration of plant teratology. Phyton 4: 19-34.
Moquin-Tandon, A. 1841. Elements de teratologie vegetale. Paris. 403 pp.

Tschudy, Robert H. 1939. The significance of certain abnormalities in Equisetum. Am.
Jour. Bot. 26:744-749.

Wagner, W. H., Jr. 1962. Irregular morphological development in hybrid ferns. Phyto¬
morphology 12: 87-100.

- . 1966. Illustrations of transient fern forms. Am. Fern Jour. 56: 101-107.

Worsdell, W. C. 1915. The principles of plant teratology. Roy. Soc. London. 2 vol.

Whitaker, T. W. 1924. The occurrences of tumors on certain Nicotiana hybrids. Jour.
Arnold Arb. 15: 144.

Zimmermann, W. 1959. Die Phylogenie der Pflanzen. ed. 2. Gustav Fischer, Stuttgart. 777

pp.

1968

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121

A PRELIMINARY REPORT ON THE DISTRIBUTION OF
GYMNOSPERMS IN MICHIGAN

Edward G. Voss

Herbarium, The University of Michigan, Ann Arbor

In the course of preparing a concise account of the gymnosperms of Michi¬
gan, 1 have become aware how poorly represented most of these plants are in the
herbaria of the state. Distribution maps based on currently known specimens
may be quite misleading compared to the actual ranges of the species concerned.
Hence, a series of preliminary county-distribution maps is here presented, in the
hope that it will inspire botanists in the field this summer to improve, by making
good collections, the basis for future maps. Only native (not planted) individuals
should be collected for this purpose.

Reasons for the poor representation of gymnosperms (and, for that mat¬
ter, of most trees) in herbaria probably include an erroneous assumption that the
plants are so common and conspicuous that sufficient specimens must already
have been collected, as well as the fact that making good specimens sometimes is
difficult. While large woody plants are conspicuous through a long season, find¬
ing and reaching collectible portions bearing flowers or fruit (or cones) is not
always easy; and herbarium curators generally frown on sterile specimens of
plants. Spruces ( Picea spp.) and hemlock ( Tsuga canadensis ) make particularly
discouraging collections since when dry they shed their needles, which must be
placed in a packet or envelope on the herbarium sheet. Cones tend to make
specimens large and bulky compared to those of many plants, giving the collec¬
tor another excuse to ignore conifers.

Specimens will be more valuable the farther they come from the range
suggested by the maps. An additional record, in other words, from an adjacent
county is much less significant than one from a hundred miles farther south than
previous specimens. Extension of known limits of range should give collectors a
special challenge. The more significant a record is, the more justified a sterile
specimen may be. Ordinarily, young cones on the branchlets, or older separate
cones carefully marked or tagged to make clear the specimen with which they
belong, add immensely to the value of herbarium material; unless quite small or
green, cones should not be pressed with the rest of the specimen. As with any
large plant, the collector should record on the label the overall stature of the tree
or shrub from which the specimen came, as well as the date, exact location,
habitat, and his name.

The distribution maps include a dot in each county from which a specimen
has been examined.1 In addition, certain islands or island groups have been

The maps are based on specimens examined in herbaria at the following institutions:
University of Michigan, Michigan State University, Wayne State University, Cranbrook In¬
stitute of Science, University of Michigan Biological Station, and Grand Valley State Col¬
lege. When additional herbaria have been checked, naturally some improvements in the maps
will result.

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mapped separately from the mainland of their respective counties: Isle Royale
(Keweenaw Co.), Drummond Island (Chippewa Co.), Mackinac, Round, and Bois
Blanc Islands (Mackinac Co.), the Beaver Islands (Charlevoix Co.), and the
Manitou Islands (Leelanau Co.). During the period 1963-1966, Paul 0. Rudolf
and collaborators issued a series of “Research Notes” from the Lake States (now
North Central) Lorest Experiment Station, on the botanical and commercial
ranges of several conifer species: LS-15 (jack pine), LS-62 (red pine), LS-63
(white pine), LS-73 (white spruce), LS-74 (black spruce), NC-16 (balsam fir),
NC-17 (tamarack). In several instances, these are based on information which
indicates more extensive ranges than the maps here; major range extensions are
mentioned below, with the Research Note cited merely by number.

Identification keys and some supplementary remarks have been included
in this report. Unless otherwise specified, all references to cones are to the
female cones.

KEY TO FAMILIES OF MICHIGAN GYMNOSPERMS

1. Leaves opposite or whorled, scale-like or needle-like . CUPRESSACEAE

1. Leaves alternate or in clusters, needle-like or linear

2. Fruit red and berry-like; leaves flattened, with strongly decurrent
bases, persistent, appearing 2-ranked, all green on both sides (may be

yellower beneath) . TAXACEAE

2. Fruit a dry woody cone; leaves flattened or not, but if so, not as
above (i.e., not decurrent, readily falling when dry, not 2-ranked,
and/or with distinct white lines) . PINACEAE

TAXACEAE

1 . Taxus canadensis Marsh. Map 1. Ground-hemlock; Yew

The only native species of the family in this area. Various cultivated yews
have similar fruit and needles (flat, + uniform yellow-green beneath, with decur¬
rent bases) to those of the native plant— even though some of them are upright,
like small trees. Our native species is a spreading monoecious shrub (most other
species are dioecious), with attractive red “fruit.” The latter consists of a single
seed surrounded by a fleshy cuplike structure (the aril). The bright red, pulpy
aril is edible, but the seed is reputed to be poisonous, as is the foliage, at least to
some animals. (The seed, even if it contains a poisonous principle, may pass
safely through the digestive tract if not chewed.) The foliage is reported to have
been used by several Indian tribes in the Michigan region and Quebec in the
making of a beverage. It is heavily browsed by deer and moose in Michigan. The
European yew ( T . baccata L.) is often cultivated for ornament and is apparently
much more poisonous.

PINACEAE
Key to the Genera

1. Leaves needle-like, all or mostly grouped in definite clusters on short
shoots

2. Leaves deciduous, crowded and numerous on short lateral shoots
(alternate leaves on new twigs); cones less than 2 cm long

1. LARIX

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THE MICHIGAN BOTANIST

123

2. Leaves evergreen, in clusters of (normally) 2 or 5 ; cones more than 2

cm long . 2. PINUS

1. Leaves flattened or 4-sided, alternate (spiraled), not in definite clusters

3. Leaves persistent on dry branches, sessile, separating cleanly from an

orbicular leaf -scar without any raised projection, the twig hence basi¬
cally smooth; cones erect, 3. 5-6. 5 cm long, the scales falling at ma¬
turity from persistent central axis . 3. ABIES

3. Leaves readily falling from dry branches, leaving persistent peg-like
bases (sterigmata), the twig hence very rough; cones pendent, (1)

1.3-6 cm long, falling entire at maturity

4. Leaves flattened, rounded at apex, distinctly short-stalked in ad¬
dition to the persistent narrow base . 4. TSUGA

4. Leaves + 4-sided, acute or sharp-pointed, sessile on the persistent

peg-like base . 5. PICEA

1 . LARIX

L. laricina (DuRoi) K. Koch Map 2. Larch; Tamarack

The only native species of the genus in eastern North America. The
European L. decidua is sometimes planted; it differs in its larger cones, with
pubescent scales.

2. PINUS

A detailed study, largely on the ecology and silviculture of the two important timber
species, is represented by the following reference: K. W. Horton & G. H. D. Bedell. 1960.
White and Red Pine. Canada Dep. Northern Affairs & NatL Resources. Forestry Branch,
Bull. 124. 185 pp.

Key to the Species

1. Needles usually 5 in a cluster, + triangular in section; membranous
sheath surrounding base of each needle cluster deciduous; cones cylin¬
drical, at least twice as long as wide . . 1. P. strobus

1. Needles usually 2 in a cluster, + semicircular in section; membranous
sheath surrounding base of each needle cluster + persistent; cones
short-ovoid, much less than twice as long as wide

2. Needles ca. 8-15 cm long, straight; bark of older branches and trunk

reddish; cones generally deciduous, subterminal on the branchlets ... 2. P. resinosa
2. Needles ca. 1.5-3 (6) cm long, + twisted; bark of older branches and
trunk dark gray to black; cones generally long-persistent, lateral on
the branchlets . 3 .P. banksiana

1. P. strobus L. Map 3. White Pine

The needles of white pine are lighter in color, softer in texture, finer and
less stiff than those of our other two species. The familiar and important white
pine is the official state tree of Michigan— a fitting designation, for this was the
backbone of the lumber industry, particularly in the last quarter of the 19th
century, when Michigan led the nation in lumber production. Here were the
finest stands of this species in the world: trees attaining at their best 5-7 feet in
diameter and 150-200 feet in height, although usually smaller.

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2. P. resinosa Aiton Map 4. Red Pine

This stately tree is frequently called by the misleading name “Norway
Pine”; it grows naturally only in northeastern North America. A somewhat
similar European species, P. nigra (Austrian Pine), is occasionally planted. Its
bark is gray or darker in contrast to the very distinctive reddish flaky bark of P.
resinosa; and the stiff needles do not break cleanly upon bending, compared to
those of P. resinosa , which snap sharply.

In some ways red pine is ecologically intermediate between white pine and
jack pine, growing with the former on better soils and with the latter on light
sandy soils, although white and jack pines seldom grow together. The range
extends south to Van Buren County on the west side of the state and to Gratiot
County in the central part (LS-62).

3. P. banksiana Lamb. Map 5. Jack Pine

The needles are + abruptly obtuse to acute but blunt at the apex. In the
similar P. sylvestris (Scots Pine; Scotch Pine), the needles average a bit longer
than in P. banksiana, and they tend to taper to sharp tips as well as being more
silvery in aspect. The curved cones of P. banksiana tend to point forward toward
the ends of their branches and are long-persistent, while the more readily decidu¬
ous cones of P. sylvestris are + horizontal or reflexed, pointing toward the bases
of their branches. The bark of P. sylvestris is a distinctive orange-brown, notice¬
able especially on the upper part of the trunk. While the latter species is widely
planted, it does not often become established on its own. Old plantations seen
from highways, as in Roscommon and Crawford counties, may give the impres¬
sion of stands of a native tree.

The range of jack pine extends south near Lake Michigan to Berrien
County and the south end of the lake (LS-15). It has been estimated that jack
pine predominates in over a million acres of forest in Michigan, over half of it in
the northern Lower Peninsula, where certain restricted areas of young growth
are the home of our most famous bird, the Kirtland’s or “Jack Pine” Warbler,
which breeds nowhere else in the world.

Figs. 1-3. Distributions of Michigan gymnosperms: (1) Taxus canadensis ; (2) Larix laricina\
(3) Pinus strobus.

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125

3. ABIES

Abies balsamea (L.) Mill. Map 6. Balsam Fir

The only species of the genus in our region. The young twigs vary from
densely to sparsely puberulent. .The needles usually appear + 2-ranked, in flat¬
tened sprays, and are usually rounded to slightly notched or 2-toothed at the
apex, 0.8-2. 5 (3.2) cm long. However, especially on older cone-bearing branches,
the needles may curve upward, giving a bushy spruce-like appearance to the
branches, and may be quite acute. Such needles tend to be shorter and thicker
than the flatter, longer needles of sterile lower branches. But all are readily
recognized by the smooth round leaf scars. This species is reported south to
Oceana, Newaygo, and Mecosta counties (NC-16).

This aromatic plant is a popular Christmas tree since the needles do not
fall readily as do those of the spruces. The blunt fragrant needles are used in
balsam pillows, and the “Canada balsam” of the microscopist is distilled from
the bark and needles. Resin-filled pustules are characteristic of the bark of this
species, and the cones are often heavily resinous.

4. TSUGA

Tsuga canadensis (L.) Carr. Map 7. Hemlock

The only species of the genus in our region. The young twigs are + densely
pubescent. The short needles, mostly 6-13 (16) mm long, in flat sprays, and the
small cones, mostly 13-22 mm long, are characteristic of this species. Even
shorter needles occur on the upper side of the twigs, closely appressed and thus
displaying conspicuously the prominent white lines of the under surface. This
characteristic is not easily seen on dried specimens, because the needles are so
readily deciduous, but it is a useful field mark. The needles of hemlock are
minutely toothed toward the rounded apex and have distinct petioles about 0.5
mm or a little longer.

Hemlock inner bark is purple, and the bark has been the main source of
natural tannin for the leather industry. This genus is not native in Europe, and
should not be confused with the poison-hemlock of Socratic fame. The latter is a
herb in the Umbelliferae, Conium maculatum L.

Figs. 4-6. Distributions of Michigan gymnosperms: (4) Pinus resinosa; (5) Pinus banksiana ;
(6) Abies balsamea.

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5. PICEA

Key to the Species

1. Young branchlets mostly + densely fine-pubescent; leaf bases below the

peg-like sterigmata + obscure on 1-year-old twigs; cones rather globose,

(1) 1. 5-2.8 cm long, the scales with slightly irregular or erose margins,

woody and rigid at maturity . 1 .P. mariana

1. Young branchlets glabrous; leaf bases prominent below the sterigmata
on 1-year-old twigs, appearing to cover them with ridges and grooves;
cones cylindrical, (2.5) 3-6 cm long, the scales nearly or quite entire,
thinner and more leathery . 2. P. glauca

1. P. mariana (Mill.) BSP. Map 8. Black Spruce

The sterigmata tend to be widely spreading in this species, even at right
angles to the twig, while in the next they tend to be strongly ascending. The
needles are quite variable in size and shape and sharpness. It has been reported
that the needles of seedlings of white spruce are minutely denticulate, while
those of black spruce are entire; there is no such distinction on mature needles,
and it should be carefully checked on seedlings.

This species is the usual host of dwarf mistletoe, Arceuthobium pusillum,
although the parasite has occasionally been found on white spruce in the vicinity
of the Straits of Mackinac (Beaver Island; Wilderness State Park, Emmet Co.;
Bois Blanc Island; and Drummond Island). Black spruce is reported south on the
west side of the state to Ottawa and Kent counties (LS-74).

2. P. glauca (Moench) A. Voss Map 9. White Spruce

The needles may be up to 18 mm long, usually averaging a little longer
than those of the black spruce (see above). This species is reported south to
Newaygo, Mecosta, and Bay counties (LS-73). Picea abies, the Norway spruce,
with drooping branchlets and cones at least twice as large as white spruce, is
often planted as an ornamental.

CUPRESSACEAE

In some manuals, this family has been included in the Pinaceae.

Figs. 7-9. Distributions of Michigan gymnosperms: (7) Tsuga canadensis; (8 ) Picea mariana;
(9) Picea glauca.

1968

THE MICHIGAN BOTANIST

127

Key to the Genera

1. Mature cones ca. 8-15 mm long, brown and + woody, the elongate
scales distinct; leaves all opposite and appressed, scale-like, dimorphic (a
pair of opposite strongly keeled leaves at right angles to adjacent pair of

flat leaves); plant a tree . 1. THUJA

1. Mature cones up to 8 (10) mm long, green or bluish, fleshy and
berry-like; leaves opposite or whorled, needle-like or if scale-like and
appressed then quite uniform (none keeled); plant a tree with needle¬
like leaves or a shrub . 2. JUNIPERUS

1. THUJA

Thuja occidentalis L. Map 10. Arbor Vitae; Northern White-cedar

In addition to this one species in our area, there is another North
American one in the Pacific Northwest. This is the characteristic tree of “cedar”
swamps. The youngest seedlings have opposite or whorled flattened needle-like
leaves, but the first branches have the characteristic scale-like leaves. The flat
unkeeled leaves (i.e., those on top and bottom of twigs in contrast to the keeled
leaves straddling the sides) have a + prominent resin gland near the apex; this is
especially evident on the lower leaf.

The true cedar ( Cedrus ) of the Old World is quite different; it has foliage
somewhat like that of Larix, but evergreen, and the cone scales fall separately
like those of Abies.

Fig. 10. Distribution of Thuja occidentalis in Michigan.

2. JUNIPERUS

The female cones are berry-like and dark bluish, often glaucous. The vola¬
tile oil they contain is the source of the distinctive flavor of gin. One often sees
on Juniperus colorful orange galls caused by rust fungi of the genus Gymn-
osporangium, an important disease of apples and related plants.

Key to the Species

1. Leaves in whorls of 3, all awl-like, articulated at the base, not decur¬
rent; cones on very short, scale-covered peduncles in the axils of awl¬
like leaves . 1 .J. communis

1. Leaves mostly opposite, some or all scale-like (awl-like leaves when
present often whorled but not articulated, decurrent at base); cones

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apparently terminal on short, scale-covered peduncles or branchlets like
the branchlets from which they arise

2. Plant an erect small tree with central trunk, growing in southern

Michigan; fruit on + straight, ascending peduncles (or branchlets) ... 2. J. virginiana
2. Plant a prostrate trailing shrub, growing in northern Michigan; fruit

usually on + arched or recurved peduncles . 3.J. horizontal is

1. J. communis L. Map 11. Common or Ground Juniper

Most of our plants are var. depressa Pursh, a + decumbent form whose
large cup- or saucer-shaped mats are characteristic of dunes and sandy places.
Some plants in the vicinity of the Sleeping Bear dunes (Leelanau Co.) are un¬
usually tall and robust. The upper surface of the needles is slightly concave and
is generally marked with a conspicuous white stripe the entire length. Various
forms of the species are cultivated for ornament.

2. J. virginiana L. Map 12. Red-cedar

This species is said to have 3-5 seeds in each berry-like cone, while the next
has only 1 or 2. Both species may have sharp awl-like leaves on the young
growth and seedlings (or sometimes after injury), while the leaves on old growth
are scale-like and overlapping. Red-cedar is probably more common and wide¬
spread in the state now than it was before the clearing and exposure of the
landscape and subsequent spread by seeds eaten by birds. It is often planted.

3. J. horizontalis Moench Map 13. Creeping Juniper

I can see no consistent difference in leaf shape by which to distinguish this
species from the preceding; both have a wide range of scale shapes and awl-like
leaves, chiefly the former. Sterile specimens for which the collector carelessly
omitted from the label any statement regarding the stature of the plant may
usually be distinguished by the definitely one-sided aspect of the trailing branch¬
es of J. horizontalis— which also grows in Michigan only north of the native range
of J. virginiana. Where the two species overlap (in Wisconsin and Maine) hybrids
are reported. A very blue-glaucous form grows on Mt. Bohemia, Keweenaw
County, and the species is found elsewhere on rock in the northwestern Upper
Peninsula and on Isle Royale. In the remainder of the state, it is confined to
sandy shores and dunes of the Great Lakes.

Figs. 11-13. Distributions of Michigan gymnosperms: (11) Juniperus communis; (12)
Juniperus virginiana; (13) Juniperus horizontalis.

1968

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129

REVIEW OF THE STATUS OF SOME
GREEN ALGAE IN THE GENUS COELASTRUM

Mason G. Fenwick
Department of Biological Sciences,

Northern Illinois University, DeKalb

The genus Coelastrum was established by Naegeli in 1849 when he de¬
scribed some plants which he obtained from pools in peat diggings at Zurich. He
established two species, Coelastrum sphaericum and C. cubicum, indicating C.
sphaericum as the type. Approximately 20 species are known for the genus and
have been summarized in a key by Ventatamaran and Goyal (1962). However,
there is doubt regarding the validity of some of these taxa. Our studies (1966)
have shown that some forms tend to be polymorphic and it is not possible to
delimit all species solely on the basis of morphology.

The coenobium of C. sphaericum illustrated by Naegeli (1849) could easily
be considered identical to coenobia (Figs. 4, 7, 8, 12) which we obtained from
our most recent cultures1 of C. proboscideum v. dilatatum Vischer2 and those
which Vischer (1927) lists as varieties of C. proboscideum. These same cultures
also produced many coenobia (Figs. 3, 5, 9, 13) which have the same appearance
as Naegeli’s C. cubicum. Some plants (Figs. 6, 10) resemble C. scabrum Reinsch.

Rayss as early as 1915 had indicated that some forms are polymorphic
when she recognized “sphaericum” and “cubicum” forms in her cultures of C.
proboscideum. Earlier, Chodat (1902) had described C. proboscideum as having:
“Cenobes comme ceux du C. sphaericum, mais chaque cellule termine'e par une
trompe plus marquee; la variability est extreme dans cette espece, dont plusieurs
e'tats correspondent a s’y meprende au C. sphaericum . ” Julia Snow (1903) also
recognized the difficulty in defining the taxa when she stated: “A specimen was
found agreeing in every respect with Coelastrum cubicum Nag., which produced
typical coenobia of Coelastrum proboscideum, so that the species of Coelastrum
cubicum is to be questioned.” Wille (1919) obtained “sphaericum” plants and
some forms which he considered to be intermediate between C. sphaericum and
C. cubicum from germinated resting spores of C. cubicum. The writer (1962) has
also germinated resting spores of a Coelastrum collected from Lake Huron and
has obtained from a single coenobium “sphaericum” or “proboscideum,” “cubi¬
cum” and “intermediate” types.

The “cubicum” growth form is not often found in collections and some
records of its occurrence are listed below. Korschikov (1953) in his paper on the
algae of the area of the North Donets Biological Station near Kharkov, Ukraine,
illustrated the plant (Fig. 1) and remarked: “I have serious doubts regarding its
independence— and more detailed description of this species should be made.”
He also mentioned that the forms he personally repeatedly observed happened
to be eight-celled or, more rarely, sixteen-celled coenobia. The writer has identi¬
fied “cubicum” plants (Fig. 18) from material collected in a beach pool at the

^rown in modified Bristol’s solution, minus nitrogen, with 10 grams glucose added
per liter.

^Indiana University Culture Collection, No. 282.

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1968

THE MICHIGAN BOTANIST

131

sandy margin of Douglas Lake, Michigan and also in collections (Figs. 21, 22)
taken from a bog habitat at Port Radium, N.W.T., Canada. We have also de¬
scribed coenobia from these areas which might be defined as being intermediate
in morphology between the “sphaericum” or “proboscideum” forms and “cubi-
cum” forms (Figs. 17, 24, 25). As mentioned earlier in this paper we found
coenobia in our cultures of C. proboscideum v. dilatatum which could be identi¬
fied with published figures representing C. proboscideum, C. cubicum, C.
sphaericum and C. scabrum. (It is interesting to note the similarity between our
“scabrum” form (Fig. 6) and Korschikov’s “cubicum” form (Fig. 2).) The
“sphaericum - proboscideum,” “cubicum” and “scabrum” forms which we have
described above apparently occur in nature only in habitats such as bogs and
pools which are low in available nitrogen. Similar forms produced in our nitro¬
gen deficient media would indicate that this is true.

Further investigations involving culture studies are necessary to determine
the range of phenotypic plasticity ofother members of the genus.

LITERATURE CITED

Chodat, R. 1902. Algues vertes de la suisse. Pleurococcoides-chroolepoides, Mater, pour la
Flore Crypt. Suisse [Beitr. Krypt. Schweiz] 1 (3): 1-373.

Fenwick, M. G., L. O. Hansen, & D. L. Lynch. 1966. Polymorphic forms of Coelastrum
proboscideum Bohn. Trans. Am. Micr. Soc. 85: 579-581.

Fenwick, M. G. 1962. Some interesting algae from Lake Huron. Trans. Am. Micr. Soc. 81:
72-76.

Korschikov, A. A. 1953. Viznacinik prisnovodnih vodorostej Ukrainskoj. R.S.R. 5, Pidklas
Protokovi (Protococcineae) Ukr. Akad. Nauk., Kiev. 436 pp.

Nageli, Carl. [1849] . Gattungen einzelliger Algen, physiologisch und systematisch bear-
beitet. 139 pp., Pis. 1-8.

Rayss, Tscharna. 1915. Le Coelastrum proboscideum Bohl. Etude de planctologie experi-
mentale suivie d’une revision des Coelastrum de la Suisse. Mater, pour la Flore Crypt.
Suisse [Beitr. Krypt. Schweiz] 5 (2): 1-65, Pis. 1-20.

Snow, Julia. 1903. The plankton algae of Lake Erie, with special reference to the Chloro-
phyceae. Bull. U. S. Fish. Comm. 1902: 369-394.

Ventataraman, G. S., & S. K. Goyal. 1962. A new species of Coelastrum (Coelastrum palli
sp. nov.) from India. Revue Algol. N. S. 6: 120-125.

Vischer, Wilhelm. 1927. Zur Biologie von Coelastrum proboscideum und einigen andern
Grunalgen. Verh. Naturf. Ges. Basel 38: 368-415.

Wille, N. 1919. Algologische Notizen XXVI. Das Keimen der Aplanosporen bei der Gattung
Coelastrum Nagl. Nytt Mag. Naturvid. 56: 23-27.

^ 1

Fig. 1. Korschikov’s “cubicum” form from the Ukraine.

Fig. 2. An 8-celled form which Korschikov named “cubicum.”

Figs. 3, 5, 9, 13. “Cubicum” forms from our cultures.2

Figs. 4, 7, 8, 12. Forms from our cultures2 which resemble Naegeli’s fig. of C. sphaericum.
Figs. 6, 10. “Scabrum-like” forms from our cultures.

Fig. 17. Plant intermediate between “proboscideum” or “sphaericum” forms and “cu¬
bicum” form, from Sedge Pool, Douglas Lake, Michigan.

Fig. 21, 22. “Cubicum” forms from Port Radium, N.W.T., Canada.

Fig. 18. “Cubicum” form from Sedge Pool, Douglas Lake, Michigan.

Figs. 16, 19, 11. “Sphaericum” or “proboscideum” - “cubicum” intermediate forms from
our cultures.2

Figs. 20, 24, 25. Intermediates from Port Radium, N.W.T., Canada.
aBristol’s minus nitrogen with 10 gm. glucose added per liter.

132

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Vol. 7

BRYOPHYTES NEW TO MICHIGAN

Howard Crum and Norton G. Miller
Herbarium, The University of Michigan, Ann Arbor,
and Department of Botany and Plant Pathology,
Michigan State University, East Lansing

During the past year a concentration of bryological effort both in the field
and in the laboratory has resulted in numerous additions to the moss flora of the
state, some of them of unusual interest. In addition to the nine species and two
varieties listed below, several records of interest will be published elsewhere by
Dr. Wolfgang S. G. Maass, who collected Sphagna in the vicinity of Douglas Lake
and also in the Upper Peninsula last summer, by Dr. Paul L. Redfearn, who
taught at the Gull Lake Biological Station and collected mosses in the southern
part of the state last summer, and by Mr. Samuel J. Mazzer who has recently
found at least one moss new to the state. We have seen and confirmed all of their
additions to the flora.

MUSCI

Brachythecium curtum (Lindb.) Limpr. Cheboygan Co.— Reeses Bog,
north of Burt Lake, G. E. Nichols 157, June-Aug. 1920 (MICH); Gorge of Carp
Creek, north of Burt Lake, R. R. Ireland, Jr., July 1961 (US).— Very likely all
American reports of B. starkei (Brid.) BSG can be referred here. Though closely
related to B. starkei, B. curtum has a distinctive aspect because of loosely spread¬
ing, more or less complanate, and more gradually acuminate leaves. The range
can be confirmed, insofar as we have studied it, as northern and central Europe
and eastern North America from Nova Scotia to Minnesota and Michigan and
southward to North Carolina.

Callier { gidium pseud ostramineum (C. M.) Grout var. plesiostramineum
(Ren.) Grout. Delta Co.— In dryer parts of Care x lasiocarpa association, in beach
pool, Big Bay de Noc, H. A. Gleason, Jr. 2424, Aug. 23, 1939 (MICH).— Ap¬
parently known otherwise only from two Alaskan localities.

Dicranella cerviculata (Hedw.) Schimp. Alger Co.— On moist sandstone,
Pictured Rocks, A. J. Sharp, July 22, 1955 (US). On moist sand, ledge of cliff,
Pictured Rocks at Miner’s Castle, TV. G. Miller 4391, with Pogonatum capillare,
Aug. 6, 1967 (MSC).— Circumpolar; Alaska to British Columbia, Labrador to
Michigan and New Jersey.

Dicranella schreberiana (Hedw.) Schimp. var .robusta Schimp. ex Braithw.
Cheboygan Co.— On wet, sandy shore, Douglas Lake, G. E. Nichols, June-Aug.
1920 (MICH). Gorge of Carp Creek, Frances E. Wynne 2692, July 24, 1942
(CAN). Mackinac Co.— Bois Blanc Island, Frances E. Wynne 2566, July 8, 1942
(CAN). Ontonagon Co.— On bank along Lake Superior, Porcupine Mountains, G.
E. Nichols & W. C. Steere, Aug. 20-27 , 1935 (MICH). Presque Isle Co.— On bank

1968

THE MICHIGAN BOTANIST

133

of small river running into Lake Huron at Hammond Bay, W. C. Steere 3166,
July 17, 1941 (MICH).— Specimens have been seen from England, Sweden, and
Denmark, as well as British Columbia and Alberta and numerous localities in the
East, from Maine to Michigan and Ontario, south to New York and New Jersey.

Fissidens bushii (Card. & The'r.) Card. & The'r. Branch Co.— Cold water, E1.
B. Mains, April 7, 1913 (MICH). Eaton Co.— On alluvial soil, swampy forest
along Grand River below the dam, Fitzgerald Park, just northwest of Grand
Ledge, N. G. Miller 3859, April 9, 1967 (MSC). Van Buren Co.— On clay in
hayfield, South Haven, Irma Schnooberger 54, Aug. 20, 1937 (MICH). Washten¬
aw Co.— On soil, Saginaw Forest, west of Ann Arbor, Florence Hoseney, April
14, 1967 (MICH).— An endemic of eastern North America, from Maine to Ontar¬
io and Michigan, south to Florida and Texas; commonly confused withF. taxi-
folius Hedw.

Grimmia pilifera P.-Beauv. Iron Co.— On granite rock, Horse Race Rapids,
Evelyn Wagoner, Flora Kubsch & Margaret Feigley, Aug. 18, 1948 (MICH).—
Widely distributed in eastern North America, also found in British Columbia,
Mexico, and Japan.

Isopterygium distichaceum (Mitt.) Jaeg. & Sauerb. Eaton Co.— On moist
earth in open woods, west of Fitzgerald Park, just northwest of Grand Ledge,//.
T. Darlington 8, May 1951 (MSC).— This species is rather uncommon but widely
distributed in eastern North America; we have seen specimens from Illinois,
Maine, North Carolina, New Jersey, and Tennessee. Dr. R. R. Ireland, Jr., who
has recently monographed the genus, kindly determined this specimen.

Plagiothecium latebricola BSG. Muskegon Co.— On decayed log (probably
of Tsuga ), Norton Township Park, A. /. Gebben, May 29, 1964 (MICH).— An
apparently rare species of modest appearance, probably easily overlooked in the
field or discarded without naming in the laboratory. Northern and central Eu¬
rope; Massachusetts to Ontario and Wisconsin, south to New Jersey.

Scopelophila ligulata (Spruce) Spruce. Alger Co.— On sandstone outcrop,
Pictured Rocks, Miner’s Castle, R. R. Ireland, Jr. 4525, Aug. 2, 1961 (US). On
moist sandstone, wall of gorge some distance below Tannery Falls, L. E. Ander¬
son 20053, Aug. 6, 1967 (MICH).— A rare, yet widely scattered species, usually
found on copper-bearing rocks. We have seen specimens from Switzerland,
Japan, Mexico, and Arizona, as well as Georgia, Illinois, North Carolina, and
Tennessee. The species has also been recorded from Central and South America,
the Azores, and many parts of Asia and Asia Minor.

Zygodon viridissimus (Dicks.) Brid. Luce Co.— On bark of sugar maple,
Upper Falls of the Tahquamenon River, L. E. Anderson 20052, Aug. 8, 1967
(MICH). Presque Isle Co.— On elm in hardwood swamp near Clinton Lake,/,. E.
Anderson & H. Crum, July 1967 (MICH).— Widespread in Europe, Washington to
California, Mexico, and eastern North America from Maine to North Carolina,
this species is decidedly disjunct in northern Michigan.

134

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THE MICHIGAN BOTANIST

HEPATICAE

Scapania saxicola Schust. Alger Co. -Growing with Anthelia juratz-
kana, Solenostoma sphaerocarpum, Mielichhoferia mielichhoferi, and Primula
mistassinica over face of damp, north-facing sandstone cliff above Lake
Superior, Pictured Rocks, east of Miner’s Castle, N. G. Miller 4384, Aug. 6, 1967
(MSC).— This diminutive liverwort was easily spotted in the field because of
abundant dark-red gemmae at the tips of both dorsal and ventral leaf lobes. The
gemmae of Scapania glaucocephala, perhaps the closest relative of S. saxicola,
are, by contrast, distinctly brownish. Dr. R. M. Schuster has kindly confirmed
the identification of this new state record; the species is otherwise known only
from Wisconsin and Minnesota.

DISTRIBUTIONAL HISTORY OF BUTOMUS UMBELLATUS
(FLOWERING-RUSH) IN THE WESTERN LAKE ERIE
AND LAKE ST. CLAIR REGION 1

Ronald L. Stuckey

Herbarium, The Ohio State University, Columbus

and

The Franz Theodore Stone Laboratory, Put-in-Bay, Ohio

Introduced marsh and aquatic flowering plants in North America present
challenging subjects for plant ecologists and phytogeographers, particularly if
there is an attempt to discover the probable distributional history of these
plants. In eastern North America, Butomus umbellatus L., the flowering-rush
(fig. 1), is becoming a classic example of an introduced species with a document¬
ed record of its occurrence and time of spread. Earl L. Core (1941) summarized
the distributional history of B. umbellatus in North America. The species is
introduced from Europe where the plants grow in marshes and along borders of
streams and lakes from Italy northward to Norway. In North America it was first
observed about 1897 along the St. Lawrence River in Quebec where the first
specimens were obtained in 1905. Core cited and mapped the subsequent collec¬
tion records for the St. Lawrence River, northern New York and Vermont, and

^Contribution from the Faculty of Organismic and Developmental Biology, The Ohio
State University.

1968

THE MICHIGAN BOTANIST

135

the western end of Lake Erie. Core’s paper can give the impression that the
species has spread southwestward into Lake Erie from the St. Lawrence River
valley. However, after a reinvestigation and examination of the distributional
data from B. umbellatus in Lake Erie and adjacent areas, I would like to suggest
an alternative view.

Core points out that in 1930 Oliver A. Earwell collected B. umbellatus in
Brownstown township and at River Rouge along the Detroit River in Wayne
County, Michigan. These are the oldest specimen records for the Lake Erie
region. Earwell (1938) wrote: “This species has long been naturalized in the
vicinity of Detroit and southward. I am informed on good authority that it was
well established before the present century and that it covered acres of marshes
at River Rouge before the operations of the Ford Motor Co. in this region
reclaimed the marshland and destroyed the better part of the stand.” On his
specimen from River Rouge ( O . A. Farwell 8759, MICH) he wrote, “Has been
here since before 1918!!!” According to Gaiser (1949) Farwell’s specimen 8742
(GH) bears the note, “In 1918 it covered acres of land in River Rouge that have

Fig. 1. Butomus umbellatus L. (Flowering-Rush) at Winous Point, Sandusky Bay, Ottawa
County, Ohio. Photographed July 1966 by Richard M. Lowden.

136

THE MICHIGAN BOTANIST

Vol. 7

since been filled in.” Farwell (1938) went on to say, “There is nothing to prove
or disprove the belief that the plant was brought over by Cadillac’s party more
than three centuries ago and that it has been here ever since.” From the rapidity
with which B. umbellatus has spread in the St. Fawrence River valley since 1900,
we can infer that if the species had been in the Detroit River 300+ years ago, it
would probably have had a wide distribution in the Fake Erie region before the
beginning of the twentieth century. It may have been for this reason that Marie-
Victorin (1939) did not accept Farwell’s undocumented statement. Marie-
Victorin (1938, p. 556-557) continued to suggest the St. Fawrence River as the
site of the original or earliest introduction; from there B. umbellatus entered the
“Great Fakes by way of Lake Erie.” However, FarwelTs statement that the
species was in the River Rouge marshes before 1918 bears further consideration.
In fact it is more plausible that the River Rouge locality represents a second
separate introduction of B. umbellatus in North America, but not one three
centuries old. There are no known stations for the flowering-rush between the
St. Lawrence River and the Detroit River during the period 1895 and 1918.
Gaiser (1949) and Witmer (1964), both of whom reviewed the records of the
total known distribution of B. umbellatus in North America, gave no informa¬
tion on its occurrence along Lake Ontario. The earliest records reported by
Gaiser (1949) for the eastern end of Lake Erie are 22 July 1941, from Erie
County, Pennsylvania ( Miss M. C. Wright s.n., CM), and 17 August 1948, from
Welland County, Ontario ( B . Miller s.n., MCM2). However, by 1956 flowering-
rush was known from several additional stations about the Canadian eastern end
of Lake Erie, as well as from five locations on the north side of Lake Ontario as
mapped by Montgomery (1956).

If we take Farwell’s date of before 1918 as the earliest record for the
occurrence of flowering-rush in the River Rouge area, we can trace its distribu¬
tion throughout the western end of Lake Erie and perhaps into Lake St. Clair
(fig. 2). From the River Rouge locality, B. umbellatus apparently spread south¬
ward along the Detroit River by the early 1930’s and into Lake Erie by 1932
where it was first found at Little Cedar Point near Toledo (Lucas County, Ohio).
It reached the larger United States islands in western Lake Erie by 1936, Kings¬
ville on the north shore of the lake by 1944, East Harbor (Ottawa County, Ohio)
by 1946, and the north shore of Pelee Island by 1948. In recent years the species
has been spreading in western Lake Erie about Sandusky Bay, along mouths of
rivers, and inland in marshy roadside ditches which are generally from one to
two miles from the lake shore. The species reached the east shore of Lake St.
Clair by 1934. An interpretation of an independent distributional history of B.
umbellatus at the western end of Lake Erie and Lake St. Clair appears more
plausible than an interpretation which brings the species from the St. Lawrence

J

In 1963, the herbarium of McMaster University (MCM), aside from a teaching
collection, was transfered to the Royal Botanical Gardens, Herbarium (HAM), Hamilton,
Ontario, Canada (J. S. Pringle. The Gard. Bull. Royal Bot. Gard. 21(3): 9-24. 1967). How¬
ever, Miller’s specimen is not among the B. umbellatus holdings at HAM today, nor is it in
the teaching herbarium at MCM (fide J. S. Pringle, in lift., 18 March 1968).

1968

137

THE MICHIGAN BOTANIST

Fig. 2. Known distribution of Butomus umbellatus L. in the Detroit River, Lake St. Clair,
and western end of Lake Erie based on herbarium specimens cited in this paper, in Core
(1941), and in Gaiser (1949). Each date represents the oldest known collection from that
locality. The stippled circle represents the locality for the 1918 occurrence reported by
Farwell. The dashed and dotted lines enclose approximate areas of the plants’ spread by the
date indicated.

138

THE MICHIGAN BOTANIST

Vol. 7

River across Lake Ontario, Niagara Falls, and eastern Lake Erie, as might be
implied by some of the earlier literature.

The idea that the spread of B. umbellatus in western Lake Erie has come
from a single introduction at River Rouge and from later established colonies is
perhaps here emphasized too strongly. Our information, based primarily on data
from herbarium specimens, is far from complete. We do not know how many of
the plant-records within the Lake Erie-Lake St. Clair region themselves represent
separate introductions or escapes from cultivation. According to Gaiser (1949)
there are apparently several undocumented accounts as to how and when the
plants arrived on the east side of Lake St. Clair. One hypothesis is introduction
by seed purchased at Toledo. It therefore may not be necessary to explain a
natural migration across this lake. The records cited from Washtenaw and Oak¬
land Counties, Michigan, probably represent escapes from cultivation unrelated
to the species’ spread about the shores of Lake Erie and Lake St. Clair. Further¬
more, flowering-rush probably has been in certain locations long before anyone
obtained a voucher herbarium specimen. Even today there are certain additional,
perhaps long-occupied, stations which botanists have not visited when flower¬
ing-rush plants were blooming.

The mechanisms by which B. umbellatus spreads under natural conditions
present an intriguing problem about which we have little documentation. Ridley
(1930, p. 232-233) summarized previous observations and reported that the
seeds generally float for one day after falling from the plant. If they become
overgrown with mycelium, they float a fortnight. The seeds may also drift along
the bottom of a river in silt, or the seedlings may float in the spring. In the Lake
Erie-Lake St. Clair region, the seeds, rootstalks, and bulblets doubtless have been
moved around by water currents. Marie-Victorin (1938) emphasized the produc¬
tion of numerous pea-sized bulblets as a means of dispersal in the St. Lawrence
River. Wild animals, such as muskrats, may also aid in the dispersal of B. umbel¬
latus (Gaiser, 1949). The plant’s dispersal mechanisms bear further critical obser¬
vation and study.

While studying the flora about the southwestern end of Lake Erie during
the summers of 1966 and 1967, 1 obtained additional records for the occurrence
of B. umbellatus. Since Core’s study (1941), this species has spread about San¬
dusky Bay where it is now known to occur on both sides of the Bay in the
roadside ditches at both the new and old Bay Bridges and in the marshes at
Winous Point at the head of Sandusky Bay. A few plants were found in ponds
and marshes behind the low sandy beach on the shore of Lake Erie at the east
edge of Port Clinton and at Magee Marsh (Crane Creek State Park) in the south¬
western corner of Lucas County. In several places the marshy ditches along state
route 2 from Port Clinton westward in Ottawa County contain numerous plants.
However, plants were not found at the Touissant Creek Wildlife Area (SE V* Sec.
8, TIN, R15E, Carroll Twp., Ottawa Co.) about 4.5 miles north of Oak Harbor
on state route 19, nor were any plants found at the Resthaven Wildlife Area
(Castalia Prairie), an extensive tract of marshes, ditches, and shallow ponds to
the north and west of Castalia, Erie County. These two localities have been
visited and studied frequently by me and/or by field botany classes from The
Stone Laboratory. The flowering-rush is known from locations one and one-half

1968

THE MICHIGAN BOTANIST

139

Fig. 3. Map of the Bass Island region in western Lake Erie showing locations where Butomus
umbellatus L. was observed during the summer of 1967. Solid ovals are at locations where
flowering specimens were found; diamonds are at locations for f. vallisneriifolius\ open ovals
are at locations where flowering rush was sought but not found. Place names and localities
are shown on the map by numbers as listed below.

1

Alligator Bar

10 Green Island

19

Put-in-Bay

2

Airport Point

11 Haunck’s Pond

20

Rattlesnake Island

3

Ballast Island

12 Honey Point

21

Smith’s Pond

4

Buckeye Island

13 Lighthouse Point

22

South Dock

5

East Point

14 Middle Bass Island

23

Squaw Harbor

6

Fisher’s Pond

15 Monument Bay

24

Starve Island

7

Fishery Bay

16 North Bass Island

25

South Bass Island

8

Fox’s Marsh

17 Peach Point

26

Sugar Island

9

Gibraltar Island

18 Perry’s Monument

27

Terwilliger’s Pond

140

THE MICHIGAN BOTANIST

Vol. 7

miles from the former locality ( Stuckey 4624 ) and three and one-half miles from
the latter locality (, Stuckey 3129). The species has not been observed in the
Portage River west of Oak Harbor to New Rochester or in the Maumee River
southwest of Maumee to the Indiana state line. Field studies have not been
conducted along these two rivers below these two towns. On the Erie Islands,.#.
umbellatus has not been discovered at several suitable but intensively studied
sites (fig. 3), such as Terwilliger’s Pond and Squaw Harbor on South Bass
Island, Haunck’s Pond (Wood, 1966) and Fisher’s Pond on Middle Bass Island,
the upper end of Fox’s Marsh on North Bass Island, and the east shore of Fox’s
Pond at the south end of Pelee Island.

The sterile underwater form with long limp leaves (B. umbellatus F. f.
vallisneriifolius (Sagorski) Gluck) was not reported from the Erie Islands by Core
(1941), but it was mentioned by Gaiser (1949) who quoted a communication
from Professor Jacob Verduin stating that such a colony occurred at Alligator
Bar, Gibraltar Island. During my survey of the aquatic plants of the Erie Islands
in the summer of 1967, several locations for forma vallisneriifolius were dis¬
covered. The plants generally grew in quiet water from one to two feet deep
along the gravelly or rocky dolomite shores. The locations of this underwater
form are mapped in figure 3.

The following specimen records further document the distribution of B.
umbellatus for the western Fake Erie-Fake St. Clair region in addition to those
cited in Core (1941) and Gaiser (1949).

MICHIGAN: MACOMB CO.: Marshes along Lake St. Clair, Jul 1937, Bingham s.n.
(BLH); 3 mi s e of Mt. Clemens, 17 Sep 1950, Rogers 7287 (ALBC, BLH, MSC, WUD).
MONROE CO.: Lake Erie beach, Erie Twp., 14 Sep 1937, Campbell s.n. (MICH); Raisin
River near Monroe, 17 Aug 1946, Thompson 5-296 (BLH); barrier beach, muck soil, Pointe
Mouillee State Game Area, 16 Jul 1949, McDonald 5173 (MSC); marsh, floating bog, same
locality, 10 Aug 1949 , McDonald 5364 (MICH); wet, lakeshore marsh, Sec. 14, T8S, R8E,
12 Jul 1958, Hiltunen & Hayes 1512 (WUD.); local, edge of drying swale, (land fill) with
Typha, etc., s side of Raisin River near its mouth, Monroe, 29 Sep 1962, Voss 11030
(MICH, MSC). OAKLAND CO.: Edge of lagoon, Cranbrook Estate, Bloomfield Twp., 26
Jun 1956, Hall s.n. (BLH); scattered along the shore in shallow water up to 6n deep,
Lakeville Lake, in the small cove adjacent to the Department of Conservation fishing site, 24
Jul 1966, Mazzer 4104 (MICH). WASHTENAW CO.: One plant seen in flower, SW Va Sec.
26, T2S, R6E, Ann Arbor Twp., at e edge of Ann Arbor at Geddes Pond along Huron River,
16 Jun 1965, Stuckey 2665 (MICH). WAYNE CO.: “Abundant,” edge of water, bank of the
Detroit River, Wyandotte, 29 Jun 1932, Grassl 51 1 0 (BLH, DAO, MICH); Detroit, Jul 1933,
Ackley s.n. (MICH); roadside ditch, Pointe Mouillee State Game Area, 13 Sep 1949,
McDonald 5643 (MSC); a “wet prairie,” wet heavy clayey soil, at the intersection of
Campau & Pt. Mouillee Rds. near Pt. Mouillee Game Reserve, 1 Sep 1951, Hall 1371 (BLH).

OHIO: ERIE CO.: Marsh at Carp Pond, Kelleys Island, 7 Jul I960, Secoy 135 (OS);
roadside ditch, route 2, 2 mi n w of Venice, Margaretta Twp., 18 Aug 1962, E. M. Herrick
s.n. (OS); locally abundant in shallow water or muddy bottom of Carp Pond, north-central
shore of Kelleys Island, 30 Jun 1966, Stuckey 2973 (OS); local in marsh along Sandusky
Bay at e end of new bridge, Margaretta Twp., at the s city limits of Bay View, 27 Jul 1966,
Stuckey 3129 (SMU); common in roadside ditch along state route 269, Margaretta Twp., ca.
Vi mi e of the town of Baybridge, 8 Jul 1967, Stuckey 4262 (OS); common in wet soil
throughout lower portions of Carp Pond, north-central shore of Kelleys Island, 15 Jul 1967,
Stuckey 4416 (OS). LUCAS CO.: Mud Hats at Reno Beach, 5 Sep 1944 , Bartley s.n. (OS); a
few plants in marsh behind low sandy beach at Magee Marsh (Crane Creek State Park), Sec.

1968

THE MICHIGAN BOTANIST

141

12 s e corner of Jerusalem Twp., 21 Aug 1967 , Stuckey 5305 (OS). OTTAWA CO.: Hatch¬
ery Bay [Fishery Bay], South Bass Island, 20 Aug 1936, Tiffany s.n. (OS) [First known
specimen record for the Erie Islands, but not cited by Core (1941)] ; edge of water, route 2
at route 240 at n end of Bay Bridge, Danbury Twp., 18 Aug 1962, E. M. Herrick s.n.( OS);
occasional and scattered in wet depression near dikes at Winous Point, w end of Sandusky
Bay, Bay Twp., ca. 4 mi s w of Port Clinton, 22 Jun 1966, Stuckey 2940 (FTSL); Honey
Point, North Bass Island, 5 Jul 1965, D. Albright 47 (OS); a few plants along gravel shore at
South Dock, s shore of North Bass Island, 4 Jul 1961 ,4182; occasional in shallow water at
Smith’s Pond, s e corner of North Bass Island, 4 Jul 1967, 4186; one small colony in
roadside ditch near Sandusky Bay, along old state route 2 just n of new Bay Bridge, s w
corner of Danbury Twp., ca. 1 mi e of Gypsum, 8 Jul 1967, 4260; occasional on thin, wet
soil over dolomite rock at Airport Point (opposite Starve Island) s e shore of South Bass
Island, 13 Jul 1967, 4350; common in roadside marsh along causeway in East Harbor State
Park at East Harbor, n side of Danbury Twp., 14 Jul 1967, 4430; a few plants in shallow
water of pond behind low sandy beach at w edge of Lakeview Park at e edge of Port
Clinton, Portage Twp., 19 Jul 1967, 4498; common in roadside ditch along state route 2, SE
lA Sec. 19, T7N, R16E, Erie Twp., ca. 2 mi n of Lacarne, 24 Jul 1967 ,4593; occasional in
roadside ditch along state route 2, SW lA Sec. 32, T8N, R15E, Carroll Twp., ca. 6 mi n of
Oak Harbor, 24 Jul 1961,4624, allTC L. Stuckey (OS).

ONTARIO: ESSEX CO.: Shore of river, Puce, 25 Jul 1936, Rabl s.n. (TRT); Wind¬
sor, 14 Aug 1936 , Brown s.n. (TRT); swamp, at 2 sites, from 15 to 20 plants were observed.
Canard River, 4 mi n of Amherstburg, 23 Jul 1948, Frankton et al. 713 (DAO); marshy field
at mouth of Sturgeon Creek, Loc. 74, Mersea Twp., 3 mi s e of Leamington, 18 Jun 1953,
Shields 1330 (HAM, TRT). marshy field near mouth of Big Creek, 482, Cone. II, Malden
Twp., 4% mi s e Amherstburg, 29 Jun 1953, Shields 1461 (TRT); on Hy no. 18, VA mi e of
La Salle, 21 Jul 1953, Montgomery & Shumovich 828 (OAC); on no. 18 highway, Linden
Beach, 16 Sep 1953, Montgomery & Shumovich 942 (OAC); edge of canal, near Thames R.
lighthouse, 16 Sep 1953, Montgomery & Shumovich 962 (OAC); on Hy #18, 2 mi s of
Amherstburg, 16 Sep 1953, Montgomery & Shumovich 952 (OAC); no. 18 highway at
Cedar Beach, 16 Sep 1953, Montgomery & Shumovich 943 (OAC); in ditch near edge of
creek #424, Sturgeon Creek about 3 mi s e of Leamington, 23 Jun 1954, Soper 5996 (TRT);
on Hy no. 18, IV2 mi e of Oxley at creek, 16 Sep 1956, Montgomery & Shumovich 946
(OAC); common along river shore, Ruscom River, Rochester Twp., 8 Aug 1962, Bassett &
Koyama 4240 (DAO). KENT CO.: Plentiful, ditch leading into dredge cut, Mitchell’s Bay,
Lake St. Clair, 12 Jun 1936, Doan s.n. (TRT); shallow water at edge of cattail marsh, Dover
Twp., Mitchell’s Bay, 7 Aug 1947, Cook 137 (UWO); 2 ft water, Lot l,Conc. VIII, Dover
Twp., 25 Jul 1948, Soper & Dale 4061 (DAO, TRT); wet ditch, Paincourt to Mitchell Bay,
10 Jul 1950, /. F. Clavert 61a (HAM); in shallow water at edge of ditch, Lot 9, Cone. XII,
Dover Twp., along ditch 1 mi s w of Mitchell Bay, 14 Aug 1950, Shields 314 (CAN, HAM,
TRT); in shallow water at edge of creek, Lot 13, Cone. XIV, Dover Twp., #29a, Rankin
Creek ca. VA mi n of Mitchell Bay, 17 Aug 1950, Shields 352 (TRT); in shallow water near
mouth of creek, #29b, Rankin Creek, ca. VA mi n of Mitchell Bay, 18 Aug 1950, Shields
364 (TRT); common in water-filled roadside ditches to Lake St. Clair, 8 mi w of Chatham,
14 Jul 1960, Mulligan & Frankton 2479 (DAO).

The following collections of B. umbellatus L. f. vallisneriifolius (Sagorski)
Gluck were taken during the summer of 1967 by the author. Specimens are in
The Ohio State University Herbarium (OS) and duplicates are elsewhere as
noted.

Buckeye Island: Occasional in water about 1 foot deep over dolomite rock, n
side of the island, 29 Jun, 4128; occasional in water about 1 foot deep over dolomite rock, s
side of the island, 29 Jun ,4183.

North Bass Island: Occasional in water about VA feet deep along s shore of the
island, about half way between South Dock and Honey Point, 1 Aug, 4757.

142

THE MICHIGAN BOTANIST

Vol. 7

South Bass Island: One small colony in about 2 feet of water in Fishery Bay at
Peach Point, 24 Jun, 4091 (MICH, SMU); occasional in water about 2 feet deep just w of
Perry’s Monument in Monument Bay, 11 Jul, 4297; locally common in quiet water about 1
foot deep, n e corner of Monument Bay along n w shore of the island, 11 Jul, 4302; locally
common in quiet water about 1 foot deep over dolomite rock along s shore, ca. IVz mi from
Lighthouse Point, 13 Jul, 4343 ; occasional in water about 1 foot deep along dolomite shore,
s e side of East Point, ca. V* mi s of Buckeye Island, 11 Aug, 4963.

ACKNOWLEDGMENTS

The field study was conducted as part of the teaching and research program at The
Franz Theodore Stone Laboratory, Put-in-Bay, Ohio, primarily during the summer of 1967.
Although many individuals are to be thanked for their help, I am particularly grateful to (1)
The Ohio Academy of Science and The Ohio Biological Survey for financial support through
which it was possible to employ respectively Mr. Harold Harlan and Mr. Ronnie Johnson as
field assistants, and to whom thanks are extended for valuable assistance, (2) Dr. Loren S.
Putnam, director, The Franz Theodore Stone Laboratory, who made the facilities of the
laboratory available for this study during the field season, and (3) my students in the
aquatic flowering plants course who helped survey the aquatic habitats.

Herbarium specimens cited have been compiled from (1) The Michigan Flora Project
records in the University of Michigan Herbarium, Ann Arbor (including data from specimens
at ALBC, BLH, MICH, MSC, and WUD), (2) specimens in The Ohio State University Herbar¬
ium (OS) and the herbarium at The Franz Theodore Stone Laboratory (FTSL), and (3)
information supplied from the Royal Botanical Gardens Herbarium (HAM), Hamilton,
Ontario, by Dr. James S. Pringle, from the National Museum of Canada (CAN), Ottawa,
Ontario, by Dr. James H. Soper, from the Research Branch of the Canada Department of
Agriculture (DAO), Ottawa, Ontario, by Dr. William J. Cody, and from the University of
Toronto (TRT), Ontario, by Dr. James E. Cruise. I have examined all cited specimens at
FTSL, MICH, and OS.

LITERATURE CITED

Core, Earl L. 1941. Butomus umbellatus in America. Ohio Jour. Sci. 41:79-85.

Farwell, Oliver A. 1938. Notes on the Michigan Flora. VII. Pap. Mich. Acad. 23:123-134.
Gaiser, L. O. 1949. Further distribution of Butomus umbellatus in the Great Lakes region.
Rhodora 51: 385-390.

Marie-Victorin, Frere. 1938. Phytogeographical problems of eastern Canada. Am. Midi. Nat.
19: 489-558.

- . 1939. Le Butomus umbellatus etait-il deja en Amerique au XVIIe siecle? (Abstract).

Ann. ACFAS 5: 108.

Montgomery, F. H. 1956. The introduced plants of Ontario growing outside of cultivation
(Part I). Trans. Royal Can. Inst. 31: 91-102.

Ridley, Henry N. 1930. The Dispersal of Plants throughout the World. L. Reeve & Co.,
Ashford, England. 744 pp.

Witmer, S. W. 1964. Butomus umbellatus L. in Indiana. Castanea 29: 117-118.

Wood, Duke Willard. 1966. Vascular Plant Flora of Haunck’s Pond, Middle Bass Island,
Ottawa County, Ohio. M.Sc. thesis, The Ohio State University. 59 pp.

1968

THE MICHIGAN BOTANIST

143

THE CANTHARELLACEAE OF MICHIGAN

Alexander H. Smith

University Herbarium and Department of Botany,

The University of Michigan, Ann Arbor

The Cantharellaceae have been studied in some detail in recent years
(Corner, 1966; Petersen, 1968, in press) but many problems are still unsolved
and much difference of opinion remains to be resolved. The family as I recognize
it consists of fungi with basidiocarps essentially trumpet-shaped to vase-shaped
or with the pileus, if one can be said to be present, lobed and/or lopsided in its
development. If the last mentioned condition prevails the basidiocarps are often
found in a compound cluster. The hymenophore may be the smooth under¬
surface of the pileus, or this surface may be variously wrinkled to vein-like from
radially arranged folds or wrinkles, or it may be in the form of fold-like lamellae
often with numerous cross veins. The spores may be white to buff or pinkish in
deposit and smooth to slightly ornamented. The pigment may be either in the
wall or the cell content. Depending on the species, spore shape varies from
globose to nearly oblong. In some the wall stains blue in cotton blue— lactic acid
medium following heating, and this dye often causes the ornamentation, if such
is present, to stand out rather clearly. Cystidia are rare in the family but known
for some American species (see Petersen, 1968, in press). The hymenophoral
trama is of loosely interwoven hyphae and hence unspecialized, as one would
expect in view of the configuration of the hymenophore. The pileus trama is also
typically unspecialized as to type especially in the cuticular region.

In the study presented here I have made an effort to be sure that all the
features emphasized in the descriptions were found on a single collection or even
on a single basidiocarp in order to avoid confusing similar-appearing taxa. This
procedure of course cannot be applied absolutely, especially where ranges of size
are concerned, but it does furnish a valid point for an interpretation of diversity
in the family. Wells and Kempt on (1968) found this method very useful in
Clavariadelphus, and it has been found to be effective in the three genera treated
here.

The major characters of value in taxonomy, as it now appears, are the
color of the spore deposit, presence or absence of clamp connections, configura¬
tion of the hymenophore, solid versus hollow stipe, spore ornamentation, and
location of pigments. Using these features, along with the pigmentation of the
basidiocarp generally, brings out certain interesting points relative to the evolu¬
tion of the group.

First, a white spored “line” of species can be traced from Clavariadelphus
to both Craterellus and Cantharellus and along with it we find increasing com¬
plexity in the configuration of the hymenophore. Secondly, an almost exactly
parallel series can be arranged leading into both genera for the species with
colored spores, only here in Craterellus it seems to end in species like C. fallax
with practically smooth hymenium whereas in Cantharellus the line with colored

144

THE MICHIGAN BOTANIST

Vol. 7

spores ends (as far as the family is concerned) in taxa which have fairly well-
formed gills, some with acute edges at maturity. The white-spored line at present
appears to connect readily with the Hygrophoraceae, which family features a
waxy hymenophore with thick gills and relatively long basidia. The line with
colored spores connects, though not as smoothly, with the genus Clitocybe sensu
lato of the Tricholomataceae of the Agaricales. The possibility of Gomphus
connecting up to Scutiger in the Polyporaceae should be carefully explored since
here is a true tendency for the production of wide pores and rather firm context
of the basidiocarp (see G. kauffrmnii).

I regard the family as primitive among the fleshy fungi largely because of
the relatively simple basiciocarps, the long basidia, and the facts that the latter
often bear more than four spores and that occasionally one observes lateral
sterigmata on basidia in some species.

Much more work needs to be done on the North American species, but it
is pointless to study herbarium specimens for which essential data are lacking,
because correlations of macroscopic and microscopic features cannot be made
accurately. The present account is given to place my own observations on record
in the hope they will serve as a guide to, and also facilitate, future work.

The color terms in quotation marks are from R. Ridgeway, Color Stand¬
ards and Color Nomenclature (Washington, D. C., 1912). The specimens cited
are deposited in the Herbarium of the University of Michigan.

The field work has been carried out over the years since 1933 with finan¬
cial aid from the University of Michigan Herbarium, from E. B. Mains (now
retired), from the Faculty Research Fund of the University of Michigan, and
from personal contributions.

Key to Genera

1. Hymenium vein-like to merulioid (of wide shallow pores) and spore wall

faintly colored buff to slightly browner (as seen under a microscope) . Gomphus

1. Hymenium smooth to vein-like or obtusely folded radially into rudi¬
mentary lamellae (with or without cross veins); spores if colored with

the pigment in the protoplasm . 2

2. Clamp connections absent; basidiocarps typically dark colored;

hymenium smooth to radially wrinkled or vein-like . Craterellus

2. Clamp connections present; hymenium typically yellowish or orange

and smooth to lamellate . 3

3. Basidiocarps cylindric to clavate, lacking a pileus (but almost pileate iri

C. truncatus); hymenium smooth or wrinkled . Clavariadelphus ^

3. Basidiocarps pileate and hymenophore usually more or less lamellate

(nearly smooth in some bright colored species) . Cantharellus

GOMPHUS S. F. Gray, Nat. Arr. Brit. PL 1: 638, 1821.

Basidiocarps fleshy, when young truncate-clavate with an even margin at
first but growing out irregularly to produce a lobed and wavy pileus which may
be lopsided or vase-shaped; pigmentation various; hymenophore varying from
smooth to radially wrinkled to merulioid with the depressions broad and shal-

^See Wells & Kempton (1968) for a treatment of the North American species.

1968

THE MICHIGAN BOTANIST

145

low or in extreme cases with poorly defined pores; spore wall slightly colored
as seen in groups of spores under the microscope.

Type species: Gomphus clavatus

Key to Species

1. Basidiocarps with dark colors dominant; hymenophore cinereous to

violaceous-drab or violet-brown . 2

1. Basidiocarps with bright colors; hyenophore pallid, yellowish or pale

orange . 3

2. Spores roughened; clamp connections present . 1. G. clavatus

2. Spores smooth; clamps absent . 2. G. pseudoclavatus

3. Pileus floccose-squamulose; typically solitary to gregarious . 3. G. floccosus

3. Pileus coarsely squamose and in large fruitings many of the basidiocarps

compound (merismoid) . 4. G. bonari

SECTION GOMPHUS

Basidiocarps with somber colors and hymenophore mostly as radial
wrinkles and veins often forming late and mostly near the pileus margin.

1 . Gomphus clavatus (Fr.) S. F. Gray, Nat. Arr. Brit. PI. 1 : 638. 1 821 .

Cantharellus clavatus Fries, Syst. Myc. 1 : 322. 1821.

Figs. 1-3.

Pileus 3.5-10 (15) cm broad, at first scarcely differentiated from the stipe
(the basidiocarps resembling those of Clavariadelphus truncatus in shape), soon
the margin growing outward irregularly to produce a lobed to lopsided structure,
finally irregularly funnel-shaped as the margin becomes uplifted; surface dry and
glabrous, unpolished to velvety, in age at times minutely scaly; color at first dull
vinaceous to purplish but fading to sordid brown to dingy alutaceous (“dark
purple-drab” when young, soon fading through “light russet-vinaceous,” “rus-
set-vinaceous,” avellaneous, or finally “clay-color” to “tawny olive”— dingy yel¬
low-brown). Context thick in the disc, thinner (about 5 mm) in the extended
margin, whitish to pale buff (“cinnamon-buff’); odor and taste not distinctive.

Hymenophore in the form of numerous low crowded frequently forked or
anastomosing ridges and with thick veins connecting the ridges, at times almost
poroid in appearance, decurrent almost to the base of the stipe, color purple to
purple-vinaceous fading to avellaneous or more vinaceous and when dusted with
spores with an ochraceous overtone.

Stipe 4-10 cm long, 8-30 mm thick below, frequently compound and
expanding upward into the pileus, sometimes many fused at the base into a
fleshy mass, solid, finally hollowed somewhat, purple-drab to avellaneous or
dingy alutaceous above, whitish below from a thin coating of mycelium.

Spore deposit pale alutaceous; spores 10-13 x 5-6.5 p narrowly ellipsoid to
narrowly ovoid, the outer wall warty to wrinkled, nearly hyaline in KOH, pale
dingy tawny in Melzer’s but soon merely pale ochraceous; in profile usually
with a distinct suprahilar depression.

Basidia 4- to 6-spored, 60-80 (90) x 7-9 p, narrowly clavate, hymenium
dull orange-brown when revived in KOH, orange-yellow in Melzer’s. Cystidia
none. Filamentous elements present in hymenium and 3-4.5 p wide (possibly
young basidia?).

146

THE MICHIGAN BOTANIST

Vol. 7

Gill trama with an interwoven central strand which is merely a slight
extension of the context of the pileus, the subhymenial zone yellowish in KOH
but fading, hymenopodial region only slightly more loosely floccose than the
central strand. Context of pileus of compactly interwoven thin-walled, smooth
hyaline hyphae 4-10 p in diam., and the cells mostly uninflated; epicutis of
pileus of upright filaments (a trichodermium) 40-80 x 2.5-6 \ u, brownish in mass
when revived in KOH but soon hyaline and hyaline when isolated under the
microscope, forming a very compact turf. Clamp connections regularly present
at the septa.

Habit, habitat, and distribution: Scattered, gregarious or cespitose, often
in arcs or rings, common in cold wet conifer forests in the fall in the Upper
Peninsula, not as abundant in the Lower Peninsula. We have one record of the
species from an elm-soft maple swamp at Sharon Hollow, Washtenaw Co. (Smith
62896); late summer and fall.

Observations: Further studies of Smith 62896 from the same locality are
needed as the basidiocarps were old when found. The rough spores and clamps,
however, exclude C. pseudoclavatus which is the species it was mistaken for
when collected. C. clavatus is a choice edible species in spite of its appearance.
The basidiocarps soon become riddled with larvae, however, so they should be
sorted when collected. Discard those with “pin-holes” in the flesh.

2. Gomphus pseudoclavatus (Smith) Corner, Monogr. Cantharelloid Fungi, p.
124. 1966.

Cantharellus pseudoclavatus Smith, Mycologia 39:505. 1947.

Pileus 3-8 (10) cm broad, truncate, the margin soon growing out and
becoming spreading to uplifted, often developing on one side only, pileus finally
flabelliform to funnel-shaped, the margin usually very wavy and lobed in age,
surface dry and unpolished or in age appressed fibrillose with the fibrils arranged
in fascicles at times, color near “vinaceous-buff,” “cinnamon-buff,” or “avel-
laneous” (pale alutaceous to grayish) when young, often a purplish tint pervad¬
ing along the margin, finally sordid alutaceous and remaining this color in drying
or becoming drab. Context thick in the disc but thin in the extended margin,
white to whitish; odor mild, taste slightly fungoid.

Hymenophore variable in form and color, more or less in the form of
fold-like gills moderately close to subdistant, decurrent, narrow, and connected
by numerous cross veins to such an extent as to give a shallowly poroid appear¬
ance, purplish vinaceous at first, with an ochraceous sheen as spores mature.

Stipe 2-6 cm long, 1-2 cm thick at apex, solid at first, pallid within,
enlarged upward, surface colored more or less like the pileus, unpolished to
tomentose, base white mycelioid.

Spore deposit pale ochraceous; spores 9-12 x 5-6.5 p, ovate to ellipsoid,
smooth, thin-walled, pale yellow in Melzer’s, nearly hyaline in KOH.

Basidia 50-80 x 9-11 (12) p, 4- to 8-spored, typically flexuous, when
revived in KOH the sections appearing pale bister, individual basidia nearly hy¬
aline but with a highly refractive amorphous content. Cystidia none.

1968

THE MICHIGAN BOTANIST

147

Subhymenium with elements approximately parallel to the basidia but
more interwoven as the loosely interwoven hymenopodial zone is approached,
hymenopodium of loosely interwoven hyphae; central strand of compactly inter¬
woven hyphae, pale bister in Melzer’s. Pileus with epicuticular hyphae hyaline in
KOH, appressed and 3-6 ju diam. Pileus context of hyphae similar to those of
epicutis or somewhat broader (4-9 p diam.), rather compactly interwoven,
smooth and thin-walled. Clamp connections absent.

Habit, habitat, and distribution: Cespitose-gregarious in oak-hickory
woods after heavy rains in late summer, rare. Known to date from Washtenaw
County, Michigan, and northern California.

Observations: In the field this species is easily mistaken for G. clavatus but
the smooth spores and lack of clamps at once distinguish it. It has a fleshy
carpophore and when immature it reminds one of Clavariadelphus truncatus.
Hence it appears to be properly placed in Gomphus. It is a much thicker fleshier
fungus than Cantharellus sinuosus Fries, if one judges it by the Friesian descrip¬
tions. To my knowledge the edibility of G. pseudoclavatus has not been tested,
but it would indeed be a surprise if the species were poisonous.

Section EXCAVATUS (Smith & Morse) Corner,

Monogr. Cantharelloid Fungi, p. 113. 1966.

Cantharellus section Excavatus Smith & Morse, Mycologia 39:500. 1947.

Turbinellus Earle, Bull. N. Y. Bot. Gard. 5 : 407. 1909.

Type species: Gomphus floccosus
3. Gomphus floccosus (Schw.) Singer, Lloydia 8: 140. 1945.

Cantharellus floccosus Schweinitz, Trans. Am. Philos. Soc. 4: 153. 1832.

Fig. 4.

Pileus 5-10 (15) cm broad, basidiocarp 8-20 cm high, truncate when
young, soon becoming hollowed in the center from the breaking up of the tissue
into floccose scales, the margin ascending at first but finally spreading and then
the pileus trumpet-shaped, the interior (upper) surface innately scaly with the
scales appressed toward the margin, more recurved in the tube and orange to
reddish orange but finally yellow, the spaces between also orange to yellow.
Context moderately thick, thin in old pilei, white or pallid, unchanging, fibrous;
odor and taste not distinctive.

Hymenophore with fold-like ridges which frequently fork or anastomose,
in age the configuration merulioid, decurrent almost to the base of the stipe in
an irregular manner, pale buff to more yellowish.

Stipe short and not sharply distinct from the pileus, solid at first but
becoming hollow from the apex down as the cavity forms in the pileus, surface
whitish and unpolished, tapered to the base which is usually deeply sunken in
the humus.

Spore deposit ochraceous; spores 12-15 x 6-7.5 p, outer wall slightly
wrinkled, wall slightly yellowish in KOH, in Melzer’s ochraceous tawny, in face
view narrowly elliptic, in profile often with a distinct suprahilar depression,
slightly yellowish in KOH.

Vol. 7

148 THE MICHIGAN BOTANIST

Basidia 52-60 x 10-12 p, clavate, hyaline in KOH, yellow in Melzer’s
Cystidia none. Hymenophoral trama interwoven and the hyphae about like those
of pileus context and merging with them, hyaline in KOH, yellowish in Melzer’s.
Clamp connections absent.

Habit, habitat, and distribution: Late summer and fall in cold habitats in
the northern part of the state under conifers, rare.

Observations: Some people eat and enjoy this species without suffering
distressing after-effects, but others are not so fortunate. Cases of poisoning by it
are usually of the mild type: the patient is not ordinarily hospitalized.

4. Gomphus bonari (Morse) Singer, Lloydia 8: 140. 1945.

Cantharellus bonari Morse, Mycologia 22: 291. 1930.

Pileus 3-12 cm broad, truncate when young, in age broadly vase-shaped,
margin finally spreading and undulating and/or lobed, the disc soon depressed
and the surface broken into thick more or less erect scales which fill the central
depression, scales orange at the tips blending to lemon-yellow at the base and
giving the entire pileus an orange-yellow color, fading in age to pale tan (“pink¬
ish-buff”) or becoming this color when dried. Context white, firm, tapering to
margin, relatively thin; odor and taste not distinctive.

Lamellae obtuse, very narrow, in the form of radially disposed subdistant
decurrent folds or by maturity somewhat merulioid, primary folds decurrent far
down the stipe, milk-white when fresh becoming dingy cream-color and when
dried pale dingy brownish.

Stipe 2-8 cm long, 10-20 mm thick, solid, glabrous, white, enlarged up¬
ward into the pileus, branching (up to 13 pilei seen attached to a common base),
many pilei remaining aborted, becoming hollowed from the apex downward.

Spore deposit weakly ochraceous. Spores 10-12(14) x 5-6 p, subellipsoid,
smooth or outer wall slightly wrinkled, ochraceous tawny in Melzer’s, in KOH
weakly yellowish.

Basidia 2- to 6-spored, 44-70 x 7-8 p, narrowly clavate with flexuous
pedicels, hyaline in KOH. Cystidia none. Hymenophoral trama of interwoven
hyaline hyphae lacking clamp connections. Pileus trama hyaline in KOH, the
hyphae similar to those of the hymenophoral trama.

Habit, habitat, and distribution: Not known for certain in the state, but on
some occasions clusters of undeveloped strongly scaly basidiocarps arising several
on a common stipe have been found. I now suspect these of belonging to this
species. Abundant material of this species has been collected in central Idaho.

Observations: In Idaho and Oregon I did not recognize the mature stage of
this species but misidentified it with C. floccosus. Perhaps the important ques¬
tion to ask is whether or not G. bonari is just a scaly merismoid variant of G.
floccosus. Dr. Ronald Petersen (pers. comm.), however, has found evidence that
at least one variant of G. floccosus in the southeast can be distinguished, and he
has found a significant difference in the spore ornamentation between G. bonari
and G. floccosus. It is likely that much of the northern and western material I
have previously placed in G. floccosus, is merely the mature stage of G. borani.

1968

THE MICHIGAN BOTANIST

149

CRATERELLUS Persoon, Myc. Eur. 2: 4. 1825.

The species I place here generally trend toward dark colors such as date
brown, gray, fuscous, or blackish; and clamp connections are absent. The basidi-
ocarp is more funnel-shaped than in Cantharellus but this is a relative feature. If
the genus can be maintained at all it must be on the absence of clamps combined
with the weak development of the hymenophore and the dark colors.

Type species: C. cornucopioides

Key to Species

1. Spore deposit salmon-buff . 10. C. fallax

1. Spore print white to dingy ochraceous . 2

2. Spore deposit pale buff . 9. C. cornucopioides

2. Spore deposit white . 3

3. Odor sickening sweetish . C. foetidus

3. Odor fragrant to lacking . 4

4. Hymenophore smooth or practically so; pileus watery brown fresh ... 5. C. caly cuius

4. Hymenophore becoming wrinkled to veined; pileus blackish brown

to bluish fuscous moist . 5

5. Cuticular hyphae disarticulating . 6. C. caeruleofuscus

5. Cuticular hyphae not disarticulating to an appreciable extent .

. 7. Craterellus cinereus var. multiplex

5. Craterellus calyculus (Berk. & Curt.) Burt, Ann. Missouri Bot. Gard. 1: 339.
1914.

Stereum calyculus Berk, et Curt. Jour. Bot. Hooker 1 : 238. 1849.

Pseudo craterellus calyculus (Berk. & Curt.) Reid, Persoonia 2: 124. 1962.

Pileus about 1 cm broad, plane with a wavy margin, disc perforated, moist
and hygrophanous, watery brownish fading to gray and near avellaneous as
dried, when faded the surface uneven to slightly scrupose.

Hymenium smooth to slightly sulcate, drying smooth, gray drying avellan¬
eous (with a tinge of pinkish), decurrent down the upper part of the stipe.

Stipe about 3 cm long and 4 mm at apex, narrowed to 2 mm at base, dingy
brownish below, concolorous with hymenium above, apical region cream-buff to
pale dingy pinkish-buff as dired.

Spore deposit white (but thin); spores 7-9 x 4. 5-5. 5 p, ellipsoid, smooth,
hyaline in KOH, yellowish in Melzer’s, thin-walled.

Basidia 33-46 x 8-9 p, relatively short-clavate, 4-, 6- (8-) spored, hyaline to
yellowish in both KOH and Melzer’s. Cystidia none.

Subhymenium of interwoven hyphae 5-9 p in diam, and with dingy yel¬
low-brown content, walls thin and smooth. Context hyphae scarcely distinct
from those of the subhymenium, content dingy pale yellow in KOH, yellow in
Melzer’s, the cells uninflated and 5-10 p in diam., epicutis of pileus merely of
compactly interwoven hyphae similar to those of context. Clamp connections
absent.

Habit, habitat and distribution: Solitary on a mound of earth formed
around an uprooted tree in a beech-maple forest, Garden Peninsula, Mich., Aug.
17, 1961. Smith 63943.

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THE MICHIGAN BOTANIST

Vol. 7

Observations: I use this name provisionally. Both Corner (1966) and Burt
(1914) indicate that the spores are colored, and Corner placed the species in
Pseudocraterelliis. A thin deposit of a typically buff-spored species often appears
white; hence I hesitate to emphasize the presumed difference. On the other hand
the cuticular hyphae did not disarticulate readily in the mounts I made which is
against my proposed identification. I simply take this means of placing the data
from my collection on record in the hope of stimulating further study on the
species involved. It might even be the “true” C. sinuosus.

6. Craterellus caeruleofuscus sp. nov.

Fig. 5.

Pileus (1) 3-8 cm latus, infundibuliformis, ad marginem lobatus, saepe fimbriatus,
caeruleofuscus demum griseobrunneus, glaber, levis vel rugulosus. Contextus tenuis; sapor
subamarus; odor nullus. Lamellae venosiformis, confertae, vel subdistantes, furcatae,
atrocaeruleae. Stipes 5-8 cm longus, 4-12 mm crassus, cavus, fuscus. Sporae in cumulo
albidae, 7-9 x 5-6 [d. Typus: Smith 42484 (MICH).

Pileus (1) 3-8 cm broad, infundibuliform and the hollow extending into
the stipe as in C. fallax, rarely compound with one pileus forming within an¬
other, margin very irregular to lobed or compound-lobed, the extreme edge at
times somewhat fimbriate to toothed, more or less “fuscous” when moist (bluish
black), fading to wood-brown or avellaneous in the manner of an hygrophaneous
species, glabrous, smooth to wrinkled; context thin and membranous, taste dis¬
agreeable, odor none.

Lamellae poorly formed, narrow and fold-like, close to subdistant, decur¬
rent, almost vein-like, with much forking and with obtuse edges, in dried spec¬
imens mostly disappearing except for near the pileus margin, bluish fuscous and
waxy when fresh, fading to a dull wood-brown, in drying becoming ochraceous
buff to pale dingy ochraceous or pallid.

Stipe 5-8 cm long, 4-12 mm at apex, equal or narrowed downward, hol¬
low, fuscous or nearly so, hymenium decurrent far down the stipe and the latter
smooth in this portion; stipe surface below the hymenium dingy and unpolished,
hymenium drying with a dingy ochraceous tone.

Spore deposit white; spores 7-9 x 5-6 ju, in face view elliptic to subovate, in
profile obscurely inequilateral to elliptic, thin-walled, smooth, hyaline in KOH
and merely yellowish to hyaline in Melzer’s.

Basidia mostly 4-spored (rarely 2-spored), 48-60 x 6.5-8 /r, narrowly clav-
ate and flexuous, slightly yellowish in Melzer’s, in KOH singly nearly hyaline but
sections of the hymenium brownish. Pleurocystidia and cheilocystidia absent.

Subhymenium of closely interwoven pallid brownish (in KOH) hyphae 4-5
Id diam. (almost as broad as the basidia), the hymenopodial zone of very loosely
interwoven but somewhat diverging hyphae extending principally from the cen¬
tral strand of the trama to the denser subhymenial area; central strand (where
gills are present) a loosely interwoven mass of hyphae similar to those of the
subhymenium and hymenopodium, brownish in KOH, the pigment located in
the wall. Epicutis of pileus of appressed non-gelatinous hyphae 5-7 ju in diam.
with the walls very slightly brownish in KOH (viewed singly) and somewhat
enlarged at the septa (to 8/i in hyphae 6 ju in diam.), hyphae readily breaking at

1968

THE MICHIGAN BOTANIST

151

the septa into units of 2-3 cells long, sparsely branched and frequently septate;
hyphae of the context similar to those of the epicutis but more interwoven;
clamp connections absent.

Habit, habitat, and distribution: Scattered to gregarious on Sphagnum in
bogs, Tahquamenon Falls State Park, September. Smith 42484.

Observations: This is close to Craterellus sinuosus Fries, and in fact I
would place it there except for the fact that Corner gives the spore print color
for that species as pale ochraceous and the spores as 10-13 x 7.5-9 q; the colors
he describes are also different.

7. Craterellus cinereus var. multiplex (Smith) comb. nov.

Cantharellus cinereus f. multiplex Smith, Pap. Mich. Acad. 38: 54. 1953.

Fig. 6.

Basidiocarps compound, 6-10 cm broad, 8-12 cm high, pilei 1^1 cm broad,
centrally to laterally stipitate (often merely eccentric), more or less infundibuli-
form and deeply perforated, glabrous or nearly so, faintly zoned, becoming
uneven to somewhat squamulose, the margin pallid and often crenate, central
part smoky fuscous to “mummy-brown.1’ Context thin, concolorous with sur¬
face; taste slightly disagreeable, odor faint and scarcely distinctive.

Famellae decurrent, fold-like but becoming more distinctly lamellate by
maturity, close to subdistant, edges obtuse, deep cinereous slowly becoming a
paler bluish gray.

Stipe compound and variable in width and length but up to 1 cm thick,
hollow, surface smooth, grayish, darker in the cortex.

Spore deposit white; spores 8-11 x 5-6 q, yellowish in Melzer’s, elliptic to
ovate in face view, in profde elliptic, or the ventral line straighter than the dorsal
line (or sometimes a little concave), thin-walled, sterigmal appendage very incon¬
spicuous.

Basidia 4-, 6-, or 8-spored, (45) 60-80 x 9-1 1 q, clavate, walls flexuous, in
sections orange to reddish in Melzer’s, in KOH pale bister to nearly hyaline.
Pleurocystidia and cheilocystidia none.

Subhymenium of compactly arranged hyphae more or less parallel to
the basidia, brownish in KOH, 3-5 q diam., hymenopodial area more inter¬
woven but equally as compact as the distinctly interwoven central area which is
more or less bister in KOH. Context of pileus bister in KOH in sections but
individual hyphae hyaline, thin-walled, equal to slightly inflated and 6-15 q
diam.; epicutis of pileus of compactly interwoven hyphae with some separate
filaments projecting or grouped to form fascicles. Clamp connections absent.

Habit, habitat, and distribution: Cespitose on the ground in mixed pine
and hardwoods, Colonial Point, Burt Fake, Michigan, Aug. 8, 1951. Smith
37531.

Observations: The gills are finally as well-formed as in Cantharellus, which
only emphasizes the close relationships among all taxa of this family. We have no
information on the edibility of this species.

152

THE MICHIGAN BOTANIST

Vol. 7

8. Crater ellus foetidus sp. nov.

Fig. 7.

Pileus 3-7 cm latus, infundibuliformis, aquose griseus demum cinereus, hygrophanus,
subsquamulosus, ad marginem undulatus. Contextus griseus, graveolens, sapor mitis.
Lamellae venosae, furcatae, confertae, cinereae, in siccatis subochraceae. Stipes 3-6 cm
longus, 1-2.5 cm crassus, cavus, fragilis, cinereus. Sporae in cumulo albidae, 8-10 (11) x 5-6
M- Fibulae nullae. Typus: Smith 62498 (MICH).

Pileus 3-7 cm broad, infundibuliform to very irregular, the hollow of the
funnel continued into the stipe, surface hygrophanous to subhygrophanous,
watery gray moist, fading to ashy gray, uneven or near the margin squamulose to
almost reticulate, in age the margin sometimes splitting and often quite wavy,
color of dried pilei pale drab. Context very fragile, watery gray; taste mild or
nearly so, odor sickening sweetish; KOH and FeS04 no reaction.

Lamellae vein-like to lamellate, narrow, much-branched and anastomosing,
close, in age often compound to nearly poroid, glaucous-cinereous to bluish
gray, drying pale dull cinnamon.

Stipe 3-6 cm long, 1-2.5 cm at apex, narrowed downward, hollow, fragile,
glaucous-cinereous like the gills (where covered by hymenium) but grayer and
naked below this line.

Spore deposit white; spores 8-10 (11) x 5-6 p, ellipsoid, smooth, thin-
walled, non-amyloid (yellowish in Melzer’s), sterigmal appendage very incon¬
spicuous.

Basidia 50-70 x 8.5-10 p, clavate, hyaline in KOH, 4-, 6-, and 8-spored,
narrowly clavate to flexuous to straight walls. Pleurocystidia and cheilocystidia
none. Subhymenium of elements about parallel with the basidia, 4-6 p diam.,
septa numerous, more interwoven toward the very loosely interwoven hymeno-
podial zone; central strand of compactly interwoven hyphae hyaline individually
but bister as sections are revived in KOH; hyphae of pileus context 6-14 q diam.,
hyaline, thin-walled, smooth, compactly interwoven; epicuticular hyphae more
compactly arranged but otherwise similar to those of the context. Clamp con¬
nections none.

Habit, habitat and distribution: Cespitose under oak, July 4, 1960,
Pinckney Recreation Area, Washtenaw County, Mich. Smith 62498 (type). Also
from Burt Lake, Smith 63890 and 63885. Not common.

Observations: The sickening-sweetish odor is a pronounced feature of fresh
collections, but is lost in drying. The species is most closely related to Craterellus
cinereus var. multiplex but the colors are different, the odor of fresh material is
different, and in C. foetidus there is a pronounced tendency of the hymeno-
phore to appear shallowly poroid. The question of whether this species should
be placed in Gomphus is pertinent but in that genus the species have solid stipes
at first. Coker (1919) described this fungus under the name C. sinuosus Fries.

9. Craterellus cornucopioides Pers., Myc. Eur. 2: 5. 1825.

Pileus 2-7 cm broad at widest point, infundibuliform when expanded and
in stature somewhat resembling C. fallax, with a perforated disc connecting to
the hollow apex of the stipe, surface wrinkled to smooth but unbroken, dark

1968

THE MICHIGAN BOTANIST

153

drab to blackish and fading to dark cinereous, margin slightly reddish brown
when fading. Context thin, rather brittle; taste mild, odor sweetly fragrant;
FeS04 no reaction ; KOH no distinctive reaction.

Hymenium smooth but becoming veined near pileus margin (rarely with
poorly formed gills), bluish drab developing vinaceous-buff tones and dingy
ochraceous as dried, decurrent to near base of basidiocarp.

Stipe 2-3 cm long (the part without hymenium), 3-4 mm at base and
evenly enlarged upward, hollow finally nearly to base, blackish, dark gray when
dried, pallid at base when fresh but lacking any obvious mycelial coating.

Spore deposit “cartridge-buff” (darker ochraceous as air-dried); spores
8-1 1 x 5-6.5 p, somewhat ovoid, smooth, thin-walled, hyaline in KOH and nearly
so in Melzer’s. Basidia 44-62 x 9-12 p, 4-spored, content weakly brownish
ochraceous in Melzer’s. Pleurocystidia none. Hyphae of pileus cutis interwoven,
4-10 /a diam., frequently septate but not disarticulating, content in Melzer’s
showing small droplets not distinctively colored. Clamp connections absent.

Gregarious under conifers, Aug. 7, 1967, Montmorency County, Mich.
Smith 74836.

Observations: The basidiocarps are not as thin as those of C. fallax but do
resemble specimens of C. cornucopioides collected by R. Shaffer in France. The
spores of Shaffer’s material were not pinkish in deposit. Burt (1914) reports
studying a specimen of C. sinuosus from Romell in which the hymenium dried
somewhat chamois colored and has spores 10-12 x 6-7 p, but C. sinuosus is not
described as similar to C. cornucopioides in stature. It is very likely true that
many of the septa in the cuticular hyphae of all these species are secondary, but
the cells do not disarticulate (separate), and so it is not obvious: 1) which septa
are secondary since clamps are absent, and 2) which septa differ chemically since
none of the cells disarticulate. Hence my collection is not a Pseudocraterellus.
Dr. Ronald Petersen is at present reviewing the status of the latter genus so it
will not be discussed further here. C. ochrosporus Burt, because of its large
spores, is close.

10. Crater ellus fallax sp. nov.

Fig. 8.

Pileus 2-6 (8) cm latus, infundibuliformis, siccus, scabrosus, subatratus. Contextus
fragilis; sapor mitis. Hymenium leve vel rugosum, cinereum vel fuscum. Stipes 4-10 cm
longus, cavus, deorsum attenuatus, cinereus vel subatratus. Sporae 11-14 x 7-9 p, leves.
Fibulae desunt. Typus: Smith 64571 (MICH).

Basidiocarp 2-6 (8) cm broad across the top, 4-8 (10) cm high, trumpet¬
shaped, the flaring margin entire to wavy or lobed or frayed, often rather deeply
incised when old and becoming quite ragged; upper surface and the interior of
the trumpet dry and unpolished and scabrous to merely rough or varying to
slightly squamulose, when wet fading as if hygrophanous, blackish when wet but
fading to dingy grayish brown or wood-brown and drying dark wood-brown to
cinnamon-drab. Context very thin, moderately brittle; odor and taste not dis¬
tinctive or the odor faintly fragrant.

Hymenophore smooth to rugose -wrinkled, never with a pattern of well-
formed forking veins near the pileus margin, fuscous-glaucous to dark cinereous

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and with a buff to salmon-buff tinge as spores mature, dark cinereous to buffy
cinereous as dried.

Stipe if distinct with only a short zone lacking hymenium and this area
appearing unpolished rather than waxy, hollow, tapered down to the base.

Spore deposit salmon buff in a good deposit. Spores 11-14x7-9 p, broad¬
ly elliptic in face view, in profile subelliptic (the ventral line in optical section
nearly straight), smooth, nearly hyaline in KOH, pale yellowish in Melzer’s, wall
thin.

Basidia 60-80 x 7-9 p, 2- and 4-spored, narrowly clavate to clavate, often
only 3 p thick at the basal cross wall, sections of hymenium bister in KOH (but
single cells hyaline or nearly so), in Melzer’s dark dingy yellow-brown in sections
from amorphous debris adhering to the hyphal walls. Cystidia none.

Subhymenium basically of interwoven much-branched hyphae with cells
3-9 p diam. and often of irregular shape, hyphae dingy yellow-brown in Melzer’s.
Basidia fasiculate. Hymenopodial zone scarcely distinct, intergrading with pileus
context. Pileus context of interwoven hyphae mostly 6-9 p diam. and with cells
uninflated but also some hyphae present with inflated cells to 9-15 p but narrow
hyphae 3-5 p diam. also present. Context hyphae dingy dark yellow-brown in
Melzer’s from amorphous debris but the cell walls themselves also dingy yellow-
brown in Melzer’s. Epicutis of closely appressed hyphae of the same type as
those making up the context of the pigmentation more pronounced. Clamp
connections none.

Habit, habitat, and distribution: Scattered, gregarious to cespitose on the
ground in deciduous and mixed forests throughout the state, common in the
summer and early fall especially during wet seasons.

Observations: This is the species that has long passed as Craterellus cornu-
copoides in this country. I have hesitated for years to describe it as a different
taxon, but Corner in his recent monograph sets forth an extremely ambiguous
concept in which the spores are described as white at first drying ochraceous,
and includes many variants based on spore size. It seems that the species is so
well known in Europe that critical studies correlating microscopic features with
macroscopic characters are lacking. But one feature seems clear, Europeans do
not regard C. cornucopioides as having a salmon-tinged spore deposit.

CANTHARELLUS Fries, Syst. Myc. 1:316. 1821.

Chanterel Earle, Bull. N. Y. Bot. Gard. 9: 407. 1909.

Pileus funnel-shaped, in species with narrow stipes with the disc often
perforated and the hollow continuous with that of the stipe. Lamellae absent
(the hymenium smooth) to present as conspicuous folds or in age with lamellae
having fairly acute edges, if the hymenium is smooth or nearly so the color is
yellow or orange or pinkish; stipes solid or hollow and not typically merismoid
in Michigan species. Spores smooth, non-amyloid, white to yellowish or pinkish
in deposits. Clamp connections present.

Type species: Cantharellus cibarius

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THE MICHIGAN BOTANIST

155

Key to Species

1. Stipe hollow or soon becoming so in the upper half . 2

1. Stipe solid and not becoming hollow . 8

2. Spore deposit white . 3

2. Spore deposit yellowish to pinkish . 5

3. Mycelium at base of stipe mustard yellow . 16. C. subperforatus

3. Stipe naked at base or with scant pallid mycelium . 4

4. Lamellae more or less poroid by maturity; basidiocarp very fragile

. 17. C. convolvalatus

4. Lamellae distant and never poroid; basidiocarp pliant . 18. C. tubaeformis

5. Spore 8-11 x 7-9 jU, subglobose . 11. C. sphaerosporus

5. Spores 9-13 x 6.5-8 p, ovate to elliptic . 6

6. P ileus blackish brown to date-color . 12. C. infundibuliformis

6. P ileus if yellow-brown at first soon yellow to orange and

hymenium bright colored from the first . 7

7. Lamellae poorly formed or hymenial surface smooth at first . 14. C. lutescens

7. Lamellae distinct at an early stage of development . 13. C. minor

8. Lamellae rudimentary . 15. C. la ter i Hus

8. Lamellae distinct at an early stage . 9

9. Basidiocarp cinnabar-red overall when young and fresh . 20. C. cinnabarinus

9. Not colored as in above choice . 10

10. Watery lilac in stipe apex when fresh . 19. C. septentrionalis

10. Not as above . 11

11. Gills pallid when young; spore deposit with a salmon-color tinge ....

. 22a. C. cibarius var. pallidifolius

11. Lamellae yellow to orange when young and fresh . 12

12. Pileus bright orange and spore deposit “orange-buff ’ . 21. C. friesii

12. Pileus egg-yellow to paler, and spore deposit weakly ochraceous

. 22. C. cibarius var cibarius

1 1. Cantharellus sphaerosporus Peck, Bull. Torrey Bot. Club 25: 323. 1898.

Pileus 1.5-3 cm broad, piano-depressed with an incurved margin, becoming
piano-depressed or finally somewhat funnel-shaped, margin thin and undulating
to wavy or crenate in age, disc becoming perforated, glabrous, subhygrophanous,
yellowish gray when moist, fading to grayish and in age or when dried pale drab,
surface becoming lacerated at times in age. Context thin, yellowish to grayish;
odor slightly fragrant, taste mild.

Lamellae decurrent, narrow, close to distant, edges obtuse, often forking
near pileus margin, grayish ochraceous to avellaneous, when dried with a buff
tone over the gray.

Stipe 3-6 cm long, 3-6 mm thick, equal or nearly so, hollow, pliant, gla¬
brous, dull to bright ochraceous orange, when dry ochraceous tones still showing
but clouded with gray Spore deposit pale yellow (near “warm buff”); spores
subglobose, 8-11 x 7-9 p, smooth, hyaline in KOH, yellowish in Melzer’s, thin-
walled.

Basidia 56-74 x 9-1 1 p, 4- or 6-spored, nearly hyaline in KOH, yellowish in
Melzer’s (in groups dingy yellow-brown), walls flexuous, in KOH bister in sec¬
tions but nearly hyaline individually. Pleurocystidia and cheilocystidia none.

Subhymenium of interwoven hyphae with cells enlarged up to 15 p at
times and much-branched; this layer merging into the context and central strand
which are of similar hyphae only slightly more compactly interwoven, hyphae

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5-9 /jl diam., and with few inflated cells, content (in KOH) slightly brownish as
seen under an oil immersion lens, bister in sections of the hymenium, dingy
yellow-brown in Melzer’s. Epicutis of hyphae similar to those of context. Clamp
connections present.

Habit, habitat, and distribution: Gregarious to scattered in a cold Thuja
bog, Burt Lake, Cheboygan County, Michigan, Oct. 14, 1960. Rare in the state.

Observations: The combination of subglobose spores, yellowish tones of
the pileus, bright orange stipe, and pale yellow spores distinguishes this species in
the stirps Infundibuliformis. We have no data on its edibility, but would rate it
not recommended because of its close relationship to C. infundibuliformis .

12. Cantharellus infundibuliformis Fries, sensu Smith & Morse, Mycologia 39:
533. 1947.

Fig. 9.

Pileus 2-3.5 cm broad, convex, depressed or expanding to broadly infundi-
buliform with irregular wavy margin or margin sulcate at times, surface glabrous,
moist and hygrophanous, color evenly blackish brown (“mummy-brown”) to
dark dingy yellowish brown (“bister”) before fading to avellaneous or ochrace-
ous-gray; disc often perforated in young specimens. Context watery and thin;
odor faintly fragrant, taste mild.

Lamellae distant, narrow, decurrent, forked, edges obtuse and even, color
“pale brownish-drab” (pale violaceous-brown) and glaucous (rather waxy in ap¬
pearance).

Stipe 3-6 mm thick at apex, narrowed downward, glabrous, naked, lemon-
yellow, with a whitish pith, sometimes tinged with umber in apical region, basal
mycelium when visible pallid.

Spore deposit “pale pinkish-buff” (ochraceous); spores 9-12 x 6.5-8 yu,
broadly ovate to broadly elliptic varying to subglobose, thin-walled, smooth,
pale yellowish in Melzer’s, sterigmal appendage distinct but small.

Basidia 50-65 x 8.5-1 1 /a, 2-, 4-, or rarely 6-spored, in sections of hymeni¬
um pale bister in KOH, single basidia nearly hyaline, sterigmata “Clavulina-like”
(with prominent curved sterigmata); wall somewhat flexuous in lower two-
thirds. Pleurocystidia and cheilocystidia none but some septate filaments found
in the hymenium and some evidence indicating indeterminate growth of the
hymenium present.

Subhymenium of frequently branched hyphae 5-8 yu in diam., arranged
approximately parallel to the basidia and causing the base of the hymenial area
to be rather dense and cellular; hymenopodial zone on the other hand of very
loosely interwoven hyphae presenting an “open” appearance in sections, central
strand of the trama of closely interwoven hyphae of about the same diameter
and type as make up the pileus context. Content of pileus of thin-walled smooth
hyphae brownish in mass but hyaline individually (in KOH), 6-15 /i broad and
the cells either uninflated or only slightly inflated, compactly interwoven.
Epicutis of pileus of hyphae like those of the context but more compactly

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157

interwoven and the cells both slightly narrower and less inflated. Clamp connec¬
tions regularly present at the septa of the hyphae in the pileus and lamellae.

Habit, habitat, and distribution: Gregarious on Sphagnum, Tahquamenon
Falls State Park, Luce County, Michigan, August 22, 1955, Smith 50289. This
species is almost as common as C. tubaeformis in Michigan and is often confused
with it.

Observations: The collection cited above, as close as I could discern, is
identical with material I collected in England. Hence Corner’s statement (with a
question mark) that the spores of this species are white indicates to me that both
C. tubaeformis and C. infundibuliformis may occur there. This species is not
recommended for the table.

13. Cantharellus minor Peck, Rep. N. Y. St. Mus. 23: 122. 1872.

Figs. 12, 14.

Pileus (1) 2-4 (8) cm broad, broadly convex to nearly plane at first, margin
incurved, at times the disc sharply depressed from the beginning, expanding to
shallowly infundibuliform and with a wavy and irregular margin, disc often
perforated (especially in age) and in large pilei the perforation broad and merely
an extension of the follow of the stipe, in young pilei the surface minutely
squamulose to fibrillose scurfy or practically glabrous, in age more often uneven
than scaly, moist and hygrophanous, bright orange-yellow (“capucine-yellow”),
duller when faded (“pale yellow-orange” or more dingy), grayish when dried.
Context soft and fragile, concolorous with pileus; odor slight but fragrant, taste
somewhat unpleasant.

Lamellae distinct and well formed, decurrent, narrow, subdistant, mostly
forked once near pileus margin, becoming conspicuously intervenose “ochrace-
ous salmon” varying to “vinaceous-cinnamon” (salmon-yellow becoming dingy
yellowish vinaceous), edges obtuse.

Stipe 3-6 cm long, 4-10 thick, central or eccentric, hollow, glabrous,
“capucine-yellow” or darker orange moist, gradually fading to dingy yellowish,
not discoloring when handled.

Spore deposit “ochraceous-salmon” (pinkish); spores 9-13 x 6-8 p, broadly
ellipsoid, smooth, thin-walled, hyaline in KOH and yellowish in Melzer’s. Basidia
4- and 6-spored, 70-90 x 8-11 p. Pleurocystidia and cheilocystidia none.

Subhymenium of much branched hyphae 5-9 p diam., oriented approxi¬
mately parallel to the basidia; hymenopodium not separated from the central
strand (both of interwoven hyphae). Epicutis of pileus hardly distinguishable
from the context, both of interwoven hyphae 5-9 p diam. with thin smooth
walls and practically hyaline in KOH, merely yellowish in Melzer’s. Clamp con¬
nections present.

Habit, habitat, and distribution: Gregarious to cespitose or scattered on
humus in the oak-aspen association of central Michigan, common and abundant
during wet seasons in July and August. It is often found along old logs almost
reduced to humus.

Observations: This is the species I (1953) previously tried to identify as
Cantharellus lutescens. On the basis of Corner’s work I am convinced this was an

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error. Peck described C. minor with a stipe that was hollow, a pileus egg yellow
and gills yellowish. The Michigan material fits this description well except that
the spores are larger and the pileus apparently more squamulose though the
latter character is quite variable in the material I have seen. Because of the
discrepancy in spore size, Peck’s name is used here provisionally. It should be
remembered, however, that most Cantharellus species with hollow stipes have
large spores.

14. Cantharellus lutescens Fries, Syst. Myc. 1: 320. 1821.

Fig. 10.

Pileus 2.5-6 cm broad, at first with the disc depressed and the margin
sharply decurved (but not incurved) and frayed to serrate (under a hand lens at
first), in age variously lobed and undulating or crenate and extreme edge serrate
to ragged; piano-depressed to infundibuliform in age and disc often becoming
perforated; color when young dingy yellow-brown (“Saccardo’s umber”), gradu¬
ally becoming a brighter yellow (“ochraceous-buff ’ to “ochraceous-salmon”), at
times the margin slightly virgate with dark lines ending in point-like dark
squamules, not hygrophanous. Context thin and pliant; odor (of large numbers
of basidiocarps) somewhat fragrant, taste not distinctive.

Hymenophore smooth to shallowly wrinkled or in age more like folds but
not gill-like, decurrent down the stipe in age, with a waxy sheen, “ochraceous-
orange” becoming “ochraceous-salmon” as spores mature.

Stipe 5-8 cm long, 4-6 mm thick at apex, 3-4 mm at the base, hollow,
uneven, glabrous, more or less ochraceous-orange, base with thin whitish
mycelium.

Spore deposit “ochraceous-salmon” or slightly paler (pinkish); spores 9-12
x 6.5-8 fi, ovate to elliptic in face view, in profile subovate to obscurely inequi¬
lateral, dingy yellowish in Melzer’s, yellowish hyaline in KOH, smooth, thin-
walled, sterigmal appendage minute and oblique.

Basidia 4- to 6-spored, 48-70 x 9-12 ju, hyaline in KOH, dingy yellowish in
Melzer’s, flexuous, narrowly clavate. Cystidia none.

Subhymenium of interwoven hyphae 5-10 ju in diam., and intergrading
with hyphae of context; context hyphae interwoven, 6-12 ju diam., branching
sparse to moderate and septa not numerous, walls slightly thickened (0.5-0. 7 n),
dingy yellowish in Melzer’s. Epicutis of pileus of radial hyphae more frequently
branched but otherwise similar to those of the context, appressed. Clamp con¬
nections present; rarely 2 occur at a single septum.

Habit, habitat, and distribution: Gregarious to scattered on wet moss in
bogs and cold springy habitats, common in late summer and early fall in north¬
ern Michigan. The description is of Smith 22149, from near Burt Fake, Aug. 20,
1964.

Observations: The spores are broad, as in the C. infund ibuliformis stirps,
and the aspect of the basidiocarp is also similar, but the hymenophore is basical¬
ly nearly emooth to undulating or finally radiately wrinkled. The configuration
of the hymenophore is actually a very constant feature. I have observed the
species in marked localities over a 20-year period and the range of variation is

1968

THE MICHIGAN BOTANIST

159

clearly limited as specified above. The pink in the spore deposit may not show in
thin deposits.

1 5. Cantharellus lateritins (Berk.) comb. nov.

Craterellus later itius Berkeley, Grevillea 1: 147. 1873.

Fig. 11.

Pileus 2. 5-5. 5 cm broad, plane to slightly depressed or finally broadly
infundibuliform, marginal portion typically undulating and arched, more or less
glabrous but appearing minutely squamulose in the disc on drying, surface actu¬
ally uneven, “zinc-orange” to “capucine-yellow” (bright orange-yellow), hy-
grophanous and fading to pale buff (“light ochraceous-buff ’ or “capucine-buff”)
when dried uniformly pale buff. Context moderately thick, 4-7 mm in disc and
tapered to the margin, soft and moderately brittle, white to pale buff, near the
margin concolorous with the surface; odor mild when fresh, after drying resem¬
bling that of Cantharellus cibarius, taste slightly bitter but soon fading.

Lamellae crowded, narrow, irregularly forking, no more than 0.5 mm
broad and up to 0.7 mm high, decurrent, edges obtuse and entire, “capucine-
buff' to “ochraceous buff” (orange-yellow to pale ocher-yellow), at times “pale
ochraceous-salmon,” in young specimens the hymenial surface may be nearly
smooth.

Stipe 1.7-4. 5 cm long, 10-16 mm thick at apex, 5-8 mm thick near base,
straight or curved, terete or compressed, solid, in age often channeled by larvae,
fleshy-fibrous, concolorous with lamellae or slightly darker.

Spore deposit pinkish-ochraceous; spores 7-9 x 4. 5-5. 5 n, ovoid to ellip¬
soid, hyaline in KOH, yellowish in Melzer’s, thin-walled, smooth.

Basidia 4- and 6-spored, 36-52 x 7-9 /jl , narrowly clavate, flexuous, yellow¬
ish in Melzer’s, in KOH hyaline but with highly refractive banded content (re¬
sembling a strand of coagulated protoplasm). Cystidia 50-60 x 3.5-5 q, flexuous-
cylindric with apex tapered to a subacute point, numerous (the young basidia
were clavate at apex). Content similar to that of basidia and basidioles.

Subhymenium showing the candelabrum-type branching and hymeno-
podial zone of loosely interwoven hyphae, the central strand if one is
present a slight projection downward of the context of the pileus. Context of
pileus of interwoven hyaline thin-walled hyphae not greatly inflated and vary
compactly arranged, yellow in KOH and slightly more so in Melzer’s. Epicutis of
pileus a loosely interwoven layer of hyphae having slightly thickened walls and
the cells readily disjointing and littering the mount with fragments 1-3 cells long
and the cells of various degrees and patterns of inflation but always 2 times or
more longer than broad and 3-8 /i in diam. Clamp connections present.

Habit, habitat, and distribution: Gregarious on soil in deciduous woods,
Warren Woods, Berrien County, Aug. 16, 1958. R. B. Howard and R. L. Shaffer
2064.

Observations: This is the only record from Michigan which we have of this
species. Because of the pinkish tint of the hymenophore in older specimens we
assume the spore-print would have had a tint of salmon in it but this was not
verified on the collection cited. The statement on spore color is the only

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THE MICHIGAN BOTANIST

Vol. 7

character in the above description taken from the literature. The diagnostic char¬
acters are the disarticulating cuticular hyphae and their slightly thickened walls,
the reduced type of hymenophore as compared with C. cibarius, and the gener¬
ally southern distribution. The presence of cystidia may be a very important
character as such structures are not typical of the genus but I should like to
verify it from additional collections. The absence of a fragment odor when fresh
may also be significant. “Craterellus cantharellus ” is the same but since the
epithet cantharellus cannot be transferred, it becomes necessary to use Berk¬
eley’s epithet. Burt (1914) studied Berkeley’s type. Since C. odoratus is de¬
scribed as having a hollow stipe, Shaffer’s collection can hardly be placed there.

16. Cantharellus subperf oratus sp. nov.

Pileus 2.5-6 cm latus, plano-depressus, ad marginem recurvatus, demum infundibuli-
formis, atrospadiceus demum sordide luteobrunneus, glaber, rugulosus. Contextus tenuis.
Odor fragrans. Sapor mitis. Lamellae angustae, decurrentes, distantes, caeruleofuscae
demum ochraceotinctae. Stipes 4-6 cm longus, 4-8 mm crassus, cavus, sursum subfuscus,
deorsum sordide aurantio-ochraceus. Mycelium laete olivaceoluteum. Sporae 8-11 x 7-9 p.
Typus: Smith 62936 (MICH).

Pileus 2.5-6 cm broad, piano-depressed at first, the margin somewhat in¬
curved, finally recurved, expanding to broadly infundibuliform and the margin
becoming sulcate to wavy or even or lobed, sometimes becoming perforated in
the center in a rather inconspicuous manner, blackish brown (“mummy-brown”)
when young and fresh, slowly becoming “snuff-brown” (dingy yellow-brown)
before fading to ochraceous gray, surface glabrous, smooth or becoming wrin¬
kled in age. Context thin and pliant; odor faintly fragrant, taste mild; no reac¬
tion to KOH.

Lamellae narrow, decurrent, distant, edges obtuse, many formed near
pileus margin, bluish drab toned ochraceous finally from spores but drab to
hair-brown when dried.

Stipe 4-6 cm long, 4-8 (10) mm thick at apex, equal or variously enlarged
at apex and/or base, hollow, often compressed in age, naked, dull ochraceous-
orange below, clouded with the gill color toward the apex; mycelium mustard
yellow.

Spore deposit white; spores 8-11 x 7-9 p, broadly ovate to elliptic or
subglobose, smooth, thin-walled, yellowish in Melzer’s, sterigmal appendage dis¬
tinct.

Basidia 6- and 8-spored, 60-90 x 9-12 p, with long flexuous pedicels,
yellowish in Melzer’s, lacking granules in the interior, sections bister in KOH but
single basidia nearly hyaline. Pleurocystidia and cheilocystidia absent.

Subhymenium compact, the hyphae frequently septate, the elements
mostly about parallel with the basidia but toward hymenopodium more inter¬
woven; hymenopodial area of loosely interwoven hyphae (the area as seen in
revived sections appearing nearly empty), the central strand of compactly inter¬
woven hyphae similar to those of the context; hyphae of pileus context inter¬
woven, mostly of uninflated cells 6-11 p diam., the walls thin and smooth,
hyaline, or faintly colored when compactly arranged. Epicutis of pileus of
hyphae like those of the context, appressed-interwoven. Clamp connections
present.

1968

THE MICHIGAN BOTANIST

161

Habit, habitat, and distribution: Gregarious on moss and duff in a cold
Abies-Thuja bog, Burt Lake, Cheboygan County, Aug. 18, 1960. Smith 62936.

Observations: The distinguishing features of this species are the mustard
yellow mycelium, white spore deposit, presence of clamp connections, and the
fact that the gills dry without any yellowish tones. It is most closely allied to C.
tubaeformis Fr. sensu Smith. Nothing is known of its edibility but I would not
recommend it.

17. Cantharellus convolvulatus sp. nov.

Fig. 13.

Pileus 3-7 cm latus, irregulariter infundibuliformis, ad marginem lobatus et convolvu¬
latus, perforatus, glaber, hygrophanus, luteo-brunneus demum ochraceus, deinde ochraceo-
griseus. Contextus fragilis, sapor mitis; odor nullus. Lamellae decurrentes angustae (ven-
osae), vinaceo-griseae. Stipes 2-5 cm longus, 6-12 mm crassus, fragilis, cavus, glaber, sordide
aurantiacus. Sporae in cumulo albidae, 9-13 x 6. 5-8. 5 jU, late ellipsoideae. Typus: Shaffer
2518 (MICH).

Pileus 3-7 cm broad, irregularly funnel-shaped with a crinkled to irregular
lobed and wavy margin, perforate at disc in age, surface glabrous, moist, hy-
grophanous, the margin translucent striate at times or remaining opaque, color
“buckthorn-brown” (a dingy yellow-brown) to dingy ochraceous when moist,
fading to dingy grayish ochraceous and drying avellaneous, to wood-brown.
Context very fragile, concolorous with surface; taste mild, odor none; KOH and
FeS04 both entirely negative.

Lamellae decurrent, foldlike, intervenose to irregularly poroid or com¬
pound poroid (with small pores grouped within larger ones— the hymenium very
irregular in appearance, “vinaceous-fawn” to “wood-brown” in color but near
avellaneous when dried, remaining readily visible on the dried specimens.

Stipe 2-5 cm long, 6-12 mm thick at apex, very brittle, equal or either
slightly narrowed or slightly enlarged downward, soon hollow, surface glabrous
and dull orange, cortex orange.

Spore deposit white; spores 9-12 x 6.5-8 n, thin-walled, smooth, oval to
broadly ellipsoid, greenish hyaline and granular in KOH, merely pale yellowish
to hyaline in Melzer’s.

Basidia 2- and 4-spored, 50-65 x 9-12 jU, narrowly clavate with somewhat
flexuous walls, frequently becoming secondarily septate. Pleurocystidia and
cheilocystidia none but some filaments present which become 2-5 septate and
with the cells sometimes inflated to 10-15 ju, walls of hymenial elements smooth
and pale brownish in KOH (nearly hyaline individually).

Subhymenium of subparallel to interwoven hyphae 7-9 /jl diam. terminat¬
ing in a basidium, often 2-5 septate below the terminal cell, some inflated hyphal
cells present near the central tramal strand and these 9-15 /jl diam. and often
irregular in shape, the walls of subhymenial elements pallid brownish as seen in
sections. Central area of gill trama of interwoven thin-walled hyphae with prac¬
tically hyaline walls, some hyphae uninflated and others with cells inflated 12-20
H , mostly 8-13 /jl in diam., branching rather frequent. Epicutis of pileus of
appressed-interwoven hyphae only slightly more compactly arranged than those
of the context but all the hyphae 6-11 jU diam., smooth and lacking appreciable

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pigmentation (though sections in KOH are brownish), merely yellowish in
Melzer’s. Clamp connections present regularly on hyphae of context and gill
trama; also at base of basidium.

Habit, habitat, and distribution: Gregarious at edge of a bog in an oak-
hickory woods, Pinckney Recreation Area, Washtenaw County, Aug. 5, 1960. R.
L. Shaffer 25 18 (type).

Observations: This Cantharellus is characterized by being very fragile, yet
the hyphae do not disarticulate; the hymenium becomes shallowly poroid by
maturity and is more or less avellaneous in color. The white spore deposit
distinguishes it from C. friesii in the sense of the latter in this work.

18. Cantharellus tubaeformis Fries, Syst. Myc. 1 : 3 19. 1821.

Fig. 15.

Pileus 1-3 (5) cm broad, convex to plane or broadly depressed and with an
arched incurved margin when young, margin finally spreading or uplifted and
becoming crenate to variously lobed, occasionally somewhat funnel-shaped in
age, surface moist and more or less uneven, at times with radial ridges terminat¬
ing in scabrous points, sometimes quite rough, sometimes practically glabrous,
usually not perforated in the disc at first but frequently becoming so in age,
moist, color “Saccardo’s-umber” to “sepia” (dark sordid yellowish brown), at
times more or less sordid ochraceous. Context thin and membranous, fragile,
yellowish to avellaneous; odor and taste not distinctive.

Famellae decurrent, narrow and with obtuse edges, subdistant to distant,
forked near pileus margin and variously veined, yellowish gray to “avellaneous”
often pale drab in age, creamy ochraceous-buff when dried.

Stipe 3-6 cm long, 3-7 mm thick, stuffed but becoming hollow and flabby,
subequal, terete or compressed, smooth or furrowed, glabrous, dark to pale
ochraceous above, usually whitish at the base.

Spore deposit white to creamy white; spores (8) 9-11 x 6-8 \ u, ellipsoid to
ovoid, smooth, thin-walled, hyaline in KOH, yellowish in Melzer’s (rarely pale
tawny).

Basidia 4-, 6-, or 8-spored, clavate, 3.5-5 \i at base, 9-12 ju at apex, hyaline
individually in KOH, yellowish in Melzer’s. Pleurocystidia and cheilocystidia
absent.

Subhymenium of the candelabrum-type, the “pedicels” making up the
hymenopodial zone, central strand of interwoven thin-walled smooth hyphae
5-1 1 p diam. and cells uninflated; hyphae of pileus context similar. Epicutis of
pileus of appressed hyphae like those of the context. Clamp connections present.

Habit, habitat, and distribution: Gregarious to scattered on Sphagnum in
bogs or on other mosses in cold springy areas, late summer to late fall; common
through the state wherever the right habitat exists.

Observations: This is the concept proposed in Smith and Morse (1947) and
the one I shall continue to use until it is clearly shown that such a white-spored
Cantharellus does not exist in Europe. For further comment see C. infundibuli-
formis. This species is not recommended for the table. Some in this group cause
mild cases of poisoning and the taxomony of the group is in such a state that it
is hard to be sure which variants are causing the trouble.

1968

THE MICHIGAN BOTANIST

163

19. Cantharellus septentrionalis sp. nov.

Pileus 2-3 cm latus, convexus, siccus, laete luteus, demum ad centrum pallidus et
subsquamulosus. Contextus luteus, tactu subfulvus. Odor fragrans. Sapor subacris. Lamellae
angustae, bifurcatae, subdistantes, luteae. Stipes 2-3 cm longus, 6-10 mm crassus, sursum
intus aquose-lilaceus, deorsum pallidus, extus sordide luteus. Sporae in cumulo citrinae, 8-10
x 6-6.5 p. Typus: Smith 67052 (MICH).

Pileus 2-3 cm broad, convex with an incurved margin, surface dry and
matted-fibrillose, “antimony-yellow” when fresh, soon pallid-canescent over the
disc, when mature diffracted scaly over all but disc, the squamules very minute
but giving pileus a rough appearance (not scaly as in G. floccosus). Context
yellow under the cuticle, paler in central area, staining orange-tawny when
bruised; odor fragrant, taste slightly peppery; with FeS04 watery grayish in stipe
apex, KOH no reaction.

Lamellae narrow, folklike, forked, decurrent, “buff-yellow” or paler and
brighter, edges obtuse and even.

Stipe 2-3 cm long, 6-10 mm at apex, solid, watery-lilac in apex when cut,
pallid below, solid, surface dry and unpolished, a duller paler yellow than the
gills and staining orange-tan where bruised.

Spore deposit lemon-yellow; spores 8-10 x 6-6.5 p, smooth, inamyloid, in
face view elliptic, in profile slightly allantoid, hyaline in KOH. Basidia 10-12 p
broad at apex, 4-spored, hyaline in KOH, yellowish in Melzer’s. Pleurocystidia
and cheilocystidia none. Subhymenium of more or less interwoven hyphae con¬
necting to a poorly defined central strand. Pileus cutis of appressed hyphae 5-8 p
diam. hyaline in KOH and Melzer’s, walls smooth and thin, content not distinc¬
tive in Melzer’s. Clamp connections present.

Habit, habitat, and distribution: Gregarious to cespitose, Tahquamenon
Falls State Park, Luce County, Aug. 7, 1963. Smith 67052.

Observations: This species is close to C. cibarius but has narrow fold-like
ridges for gills, the context at the apex of the stipe is lilaceous, and the spore
deposit when fresh is lemon-yellow. Its edibility has not been tested, but it
would be surprising if it were poisonous.

20. Cantharellus cinnabarinus Schweinitz, Trans. Am. Philos. Soc. 2: 4. 1832.

Fig. 16.

Pileus 1-5 (7) cm broad, plano-convex, expanding to shallowly infundibuli-
form, margin incurved at first, in age spreading to uplifted and variously crenate,
lobed or wavy, at times in age the margin very irregular, often unpolished when
young, soon naked, glabrous, color evenly cinnabar-red to reddish orange, fading
rapidly in sunlight and finally often nearly pallid, dried specimens pale buff.
Context thin toward the pileus margin, flushed pinkish at first but soon pallid;
taste slowly burning acrid, odor not distinctive when fresh but dried specimens
for a time at least faintly fragrant as in C. cibarius\ inky green with Melzer’s on
pileus, FeS04 no reaction or finally watery vinaceous, KOH no change.

Lamellae close to subdistant, varying to distant, long-decurrent, narrow,
edges obtuse, conspicuously intervenose or at times shallowly poroid, con-
colorous with pileus when fresh but soon paler, when dried more ochraceous
than the pileus, not staining when injured.

164

THE MICHIGAN BOTANIST

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Spore deposit delicately pinkish (in a thick deposit); spores 7-9 x 4. 5-5. 5
ju, elliptic in face view, in profile subelliptic or ventral line straight to slightly
concave, thin-walled, smooth, hyaline in KOH and merely yellowish hyaline in
Melzer’s.

Basidia 4- or 6-spored, 38-56 x 8-10 jit, narrowly clavate, hyaline in KOH
or with yellowish content, yellowish in Melzer’s. Pleurocystidia and cheilo-
cystidia none.

Subhymenium and hymenopodial areas of equally densely interwoven
hyphae, merging gradually into the central tramal strand, pale dingy yellowish
brown in sections in KOH but individual hyphae hyaline, thin-walled, smooth,
cells not appreciably inflated; hyphae of the pileus context compactly inter¬
woven, yellowish in sections (in KOH), hyaline individually, walls thin and
smooth, the cells equal to slightly enlarged either in the midportion or at the
septa; epicutis of pileus of appressed hyphae like those of context, local areas of
pileobasidia developing on raised areas. Clamp connections present.

Habit, habitat, and distribution: Solitary to gregarious in dry woods of oak
and hickory or beech-maple, especially along woodland roads, summer, common
during wet seasons.

Observations: The pilei show a marked tendency to develop patches of
hymenium on the surface or even aborted secondary pilei and as a result may
present a rather irregular appearance. The pinkish spore deposit, acrid taste, and
cinnabar color are distinctive. From C. friesii it differs particularly in color and
in the much more irregular pileus. It is edible but not usually found in sufficient
quantity for a meal.

21. Cantharellus friesii Quelet, Champ. Jura et Vosges 1: 191. 1874.

Pileus 8-30 mm broad, broadly convex with an inrolled margin, becoming
infundibuliform with a wavy-crenate margin, surface unpolished, opaque, when
moist “capucine-orange” (bright orange), fading to egg-yellow (never as red as in
C. cinnabarinus). Context thin, more or less concolorous with surface; taste
slowly acrid, odor fragrant (as in C. cibarius ); FeS04 dull vinaceous; KOH no
reaction.

Famellae narrow, close, forking, decurrent, edges obtuse, egg-yellow be¬
coming paler, strongly intervenose.

Stipe 1.5-3 cm long, 5-7 mm thick, solid, equal, pallid within, surface
unpolished and approximately concolorous with the pileus except for the whit¬
ish base.

Spore deposit “orange-buff”; spores 9-1 1 x 4. 5-5. 5 ju, smooth, nearly hya¬
line in Melzer’s thin-walled, elliptic to ovate in face view, in profile oblong to
slightly concave on the ventral line, nearly hyaline in KOH.

Basidia 40-60 x 9-12 /i, 4- to 6-spored, and the extra sterigmata at times
lateral near the apex, narrowly clavate, the walls more or less flexuous, hyaline
to yellowish in KOH, yellowish in Melzer’s, secondary septa rare. Pleurocystidia
and cheilocystidia none.

Subhymenium of interwoven hyphae 6-9 \x in diam., hymenopodium
loosely interwoven and hyphae generally extending laterally toward the hymeni-

1968

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165

um, yellowish in Melzer’s, thin-walled, hyphal cells mostly uninflated but up to
12 /jl or more in diam.; central strand of more compactly interwoven hyphae but
of the same type. Epicutis of pileus of appressed interwoven hyphae 6-12 ji
broad and with free hyphal ends projecting singly or in groups but end cells not
especially differentiated. Context of interwoven thin-walled, smooth more or
less hyaline hyphae. Clamp connections present on context tramal hyphae and at
base of basidium.

Habit, habitat, and distribution: Gregarious under hardwoods on moss and
litter, Sharon Hollow, Washtenaw Co., August 4, 1960. Smith 62787.

Observations: This species truly is closely related to C. cibarius but is
smaller, brighter orange over the pileus, has a dull vinaceous color change with
FeS04 and the spores are “orange-buff’ in a good deposit. It differs from C.
minor in having a solid stipe.

22. Cantharellus cibarius Fries, Syst. Myc. 1 : 318. 1821. var cibarius

Fig. 18.

Pileus (2) 3-8 (10) cm broad, broadly obtuse to plano-convex when young,
the margin incurved and irregular, expanding to plane with a wavy margin,
shallowly depressed or finally broadly infundibuliform, margin crenate to wavy
or variously irregular and often lobed, at times deeply sinuate on one side,
surface unpolished at first from a hoary coating, soon naked, not hygrophanous,
pale to rich egg-yellow at maturity, slowly paler in age on exposure to light,
staining greenish fuscous where touched with Melzer’s reagent. Context thick,
firm, yellowish to pallid; odor faintly fragrant, when dried odor decidedly fra¬
grant, taste slightly peppery.

Lamellae decurrent, close to subdistant, at times distant in age, narrow,
edges obtuse and even when young, subacute in age, egg-yellow or paler finally
when dried typically orange-buff, very slowly staining more ochraceous when
injured or the stains becoming visible only on the dried specimens.

Stipe 3-8 cm long, 5-15 mm thick at apex, equal or narrowed toward the
base, solid, center pallid, cortex yellowish, glabrous, concolorous with pileus or
paler at base, slowly bleaching on standing, slowly staining ochraceous when
injured, or stains showing only on dried specimens.

Spore deposit pale ochraceous; spores 8-10 x 4. 5-5. 5 ju, in face view ellip¬
tic, in profile view slightly to obscurely allantoid or subelliptic, hyaline in KOH,
yellowish in Melzer’s smooth, thin-walled.

Basidia 4- to 6-spored, 48-67 x 8-10 n, narrowly clavate, pedicel flexuous,
hyaline in KOH, yellowish in Melzer’s. Pleurocystidia and cheilocystidia none
except for fdaments 50-60 /jl long and 3.5-5 \ u. wide scattered in the humenium
(immature basidia?).

Subhymenium of hyphal cells oriented about parallel with the basidia and
3-5 /i diam.; hymenopodial zone not sharply distinct from the interwoven cen¬
tral strand, the hyphae of the latter hyaline individually, intricately interwoven,
walls thin and smooth; hyphae of pileus context similar, hyphal cells not greatly
inflated (5-12-15 p wide), cuticle of pileus merely of more compactly arranged
hyphae, above this a loose poorly formed epicutis of narrow (4-6 ju) thin-walled

166

THE MICHIGAN BOTANIST

Vol. 7

non-gelatinous hyphae with the end-cells slightly clavate, hyaline in KOH. Clamp
connections present.

Habit, habitat, and distribution: Cespitose to gregarious in deciduous
woods, especially along old roads in the woods; from the fourth of July on into
early autumn in Michigan. Common.

Observations: On the basis of specimens collected and studied in England I
regard this as the type variety. Its diagnostic features are the slightly peppery
taste developing slowly, the delayed staining to ochraceous when injured, fra¬
grant odor especially when dried, buff colored spore deposit and the overall
egg-eyllow coloration.

Edibility: This is one of the world’s highly prized edible species. In our
region, however, it is often difficult to find it free from insect larvae.

22a. Cantharellus cibarius var. pallidifolius var. nov.

Fig. 17.

Pileus 6-12 (15) cm latus, lateus; lamellae pallidae; sporae in cumulo salmonae.
Typus: Smith 62993 (MICH).

Pileus 6-12 (15) cm broad, broadly convex with an incurved margin, rather
irregular in outline, expanding to broadly convex, or shallowly depressed with a
spreading to uplifted margin, margin lobed, wavy and crenate in age and present¬
ing a very irregular appearance; surface dry and unpolished from a thin coating
fibrils, glabrescent in age and not hygrophanous, pileus essentially buff-yellow
to egg-yellow, about pinkish-buff when dried, staining ochraceous brown readily
when bruised. Context thick, white, firm; odor fragrant, taste mild but slowly
becoming peppery; FeS04 vinaceous brown; KOH intensifying the yellow of
the pileus.

Lamellae close to subdistant, narrow, decurrent, edges obtuse, many fork¬
ing or anastomosing and at times hymenophore practically poroid, color pallid
when young but gradually becoming egg-yellow at least along margin and rather
rusty ochraceous when dried.

Stipe 4-8 cm long, 1-3 cm thick, equal or narrowed downward, solid, white
within, soon staining rusty ochraceous when injured, surface whitish and un¬
polished, gradually becoming yellow in aging and staining rusty ochraceous
where injured.

Spore deposit ochraceous-salmon (flushed pink); spores 7-10 x 4.5-6 p,
oblong to elliptic in face view, slightly allantoid in profile, smooth, thin-walled,
hyaline in KOH, yellowish in Melzer’s.

Basidia (40) 60-90 x 7-9 g, narrowly clavate, mostly 4-spored, hyaline in
KOH, yellowish in Melzer’s. Pleurocystidia and cheilocystidia none.

Subhymenium and hymenopodium one unit— with candelabrum-like
branching of the hyphae; central strand of gill trama of compactly interwoven
hyphae merging with the hyphae of the pileus context; pileus context of inter¬
woven hyphae with thin smooth walls, the cells 5-9 (12) p broad and mostly
uninflated; pileus cutis of compactly appressed and interwoven hyphae like
those of the context; epicutis a loose wheft of hyaline thin-walled hyphae 4-7 p
diam., the end-cells merely obtuse. Clamp connections present.

1968

THE MICHIGAN BOTANIST

167

Habit, habitat, and distribution: Gregarious to scattered under young
beech-maple second-growth, Pellston, Aug. 23, 1960. Smith 62993. Common in
the state, from midsummer to late fall.

Observations: This is a variant with a large heavy basidiocarp which stains
readily, has pallid gills in fresh young rapidly developing pilei, and has a pink tint
to the spore deposit.

Edibility: Tills is the best of the variants of this species found in the state.
It is also more frequently found free of insect larvae.

LITERATURE CITED

Burt, E. A. 1914. The Thelephoraceae of North America. II. Craterellus. Ann Missouri Bot.
Card. 1: 327-350.

Coker, W. C. 1919. Craterellus, Cantharellus and related genera in North Carolina; with a
key to the genera of gill fungi. Jour. Elisha Mitchell Sci. Soc. 35: 24-48.

Corner, E. J. H. 1966. A monograph of Cantharelloid fungi. Oxford Univ. Press. 255 pp.
Petersen, Ronald, in press. Notes on Cantharelloid fungi I. Gomphus S. F. Gray and some
clues to the origin of the ramarioid fungi. Jour. Elisha Mitchell Sci. Soc.

Smith, Alexander H. 1953. New and rare agarics from the Douglas Lake Region and Tah-
quamenon Falls State Park, Michigan, and an account of the North American species
of Xeromphalina. Pap. Mich. Acad. 38: 53-87.

Smith, Alexander H., & Elizabeth Morse, 1947. The genus Cantharellus in the western
United States. Mycologia 39: 497-534.

Wells, Virginia, & Phyllis Kempton. 1968. A preliminary study of Clavariadelphus in North
America. Mich. Bot. 7: 35-57.

168

THE MICHIGAN BOTANIST

Vol. 7

Fig. 1. Gomphus clavatus xl Smith 54454

1968

THE MICHIGAN BOTANIST

169

/

Fig. 2. Gomphus clavatus xl Smith 72549

170

THE MICHIGAN BOTANIST

Vol. 7

Fig. 3. Gomphus clavatus xl Smith 62896

1968

THE MICHIGAN BOTANIST

171

Fig. 4. Gomphus floccosus xl Smith 70616

172

THE MICHIGAN BOTANIST

Vol. 7

Fig. 5. Craterellus caeruleofuscus xl Smith 42572

1968

THE MICHIGAN BOTANIST

173

Fig. 6. Craterellus cinereus var. multiplex xl Smith 37531

174

THE MICHIGAN BOTANIST

Vol. 7

Fig. 7 . Craterellus foetidus xl Smith 62498 (type)

1968

THE MICHIGAN BOTANIST

175

Fig. 8. Craterellus fallax xl Smith 64571 (type)

Fig. 9. Cantharellus infundibuliformis xl Smith 17626

1968

THE MICHIGAN BOTANIST

177

Fig. 10. Cantharellus lutescens xl Smith 75250

178

the MICHIGAN BOTANIST

Vol. 7

Fig. 11. Cantharellus hteritius

xl

Smith 9966

1968 _ THE MICHIGAN BOTANIST 1 79

Fig. 12. Cantharellus minor xl Smith 74950

Fig. 13. Cantharellus convolvulatus xl Shaffer 251 8 (type)

180

Vol. 7

THE MICHIGAN BOTANIST

Fig. 14. Cantharellus minor xl Smith 36985

1968 _ THE MICHIGAN BOTANIST

181

Fig. 15. Cantharellus tubaeformis xl Smith 67743

182

THE MICHIGAN BOTANIST

Vol. 7

Fig. 16. Cant hare llus cinna barinus xl Smith 32847

Fig. 17. Cantharellus cibarius var. pallidifolius xl Smith 62993 (type)

~.v.

1968

THE MICHIGAN BOTANIST

183

Fig. 18. Cantharellus cibarius var. cibarius xl Smith 28098

184

THE MICHIGAN BOTANIST

Vol. 7

yjature education

MAKING RUBBER DUPLICATES OF MUSHROOMS

Richard A. Mortemore
Kensington Metropolitan Park,
Milford, Michigan

The preparation of latex mushroom models can be a rewarding experience
for the avid as well as for the casual mycologist. All too soon after being
collected, most mushrooms begin to deteriorate. Unless the collector is also a
photographer, the rich colors and symmetrical shapes become only a memory
until the next season. With a little practice and artistic ability, a mushroom can
be duplicated in latex within a matter of hours.

The materials required in this process are rather inexpensive. A good grade
of plaster of paris, liquid latex, liquid green soap (available at pharmacies),
cardboard, and oil paints or water colors are needed.

Select the mushroom to be duplicated with care. Make sure that it is not
too soft and not perforated with too many worm holes. Carefully remove all
foreign material such as dirt, grass, and leaves. This may be done with the use of
forceps, water, or a stream of air. If the cap of the mushroom is sticky, such as
that of Boletus luteus, thoroughly wash and dry it or the detail in the plaster will
be poor.

Using a sharp knife, separate the cap from the stem. This step is not
necessary for all species. A few, such as stinkhorns and morels, may be cast in
one piece. (Fig. 1)

After separation of the mushroom cap, form two small cardboard boxes:
one to contain the cap and the other to hold the stem horizontally. Do not form
these containers excessively large or you will waste a great deal of plaster.

Mix your plaster to a creamy consistency and pour a small amount into
your container for the cap. Place the cap, £77/ (or pore) side up, into the plaster.
Make sure that a layer of plaster is between it and the bottom of the container.
Add plaster until it reaches just to the edge of the gills. Pour plaster into the
other container and immerse the stem halfway. Allow both to dry thoroughly.
(Fig. 2) If the container in which the plaster was mixed is thoroughly washed
before the plaster has a chance to harden, you will eliminate the problems of
removing the accumulation of caked plaster.

After the plaster is dry, brush a small amount of green soap on the ex¬
posed plaster surface surrounding the mushroom. This allows the plaster halves
to separate easily. Next, mix a new batch of plaster and pour it in the containers
to completely cover the cap and the stem. Allow this to dry thoroughly. (Fig. 3)

Using a knife, remove the cardboard container and carefully pry apart the
plaster halves. Allow the exposed plaster surfaces to dry for at least thirty
minutes, then remove the mushroom from the plaster. (Fig. 4) You may find
that in certain species, especially boletes, part of the cap will stick to the plaster.
This may be removed when dry, with the aid of a toothbrush.

1968

THE MICHIGAN BOTANIST

185

lijipP mpu

I

!

186

THE MICHIGAN BOTANIST

Vol. 7

* igs. 1-5. Steps in the preparation of a latex

mushroom model, as described in the text.

the plain'd ST % ^ °f

poured through these holes sn jP St St®m' T e llcluid iatex wil1 be

Plaster halves'* each ^together Zh theTf * le3St 3/8" P,aCe the
rubberband will ensure that the hJ 16 10 6S uPwards- Masking tape or a

5) th3t the halves rema"! together to prevent leakage. (Fig.

1968

THE MICHIGAN BOTANIST

187

Pour the latex into the cast until the cavity is filled. Allow it to remain in
the cast for several minutes, then pour the excess back into the latex container.
Allow the rubber in the cast to dry for at least two hours. Repeat the addition of
latex until an adequate thickness of rubber is attained. Prior to the removal of
the latex mushroom from the plaster, add a small amount of talcum powder to
the cavity. This will prevent any wet spots from sticking together.

Again using your knife, carefully pry the plaster halves apart. Trim away
any excess rubber with a pair of small scissors. Join the cap and the stem
together by using a small amount of the latex as a glue.

The easy part of the duplicating process will now be completed, for unless
you are artistic you may find the painting rather time consuming. Water colors
or oils, depending on your preferences, may be used. Several thin coats of paint
will look more realistic than one thick application.

When you succeed in your painting, a shiny finish to the cap may add a
touch of realism. This is accomplished by a brief spraying of clear Krylon. If a
dull finish is needed, spray with Krylon Matte.

Fig. 6. Several completed latex models of mushrooms.

188

THE MICHIGAN BOTANIST

Vol. 7

MICHIGAN PLANTS IN PRINT
New Literature Relating to Michigan Botany

This section lists new literature relating to Michigan Botany under four categories: A.
Maps, Soils, Geography, Geology (new maps and selected bulletins or articles on soils and
geology as these may be of use to field naturalists and students of plant distribution); B.
Books, Bulletins, etc., and C. Journal Articles (listing, respectively, all separate publications
and articles in other periodicals which cite Michigan specimens or include research based on
plants of wild origin in Michigan ;-not generally including work on cultivated plants nor
strictly economic aspects of forestry, conservation, or agriculture); D. History, Biography,
Exploration (travels and lives of persons with Michigan botanical connections). When the
subject matter or relation to Michigan is not clear from the title, annotations are added in
brackets. Readers are urged to call to the editor’s attention any titles (1960 or later) which
appear to have been overlooked-especially in less well known sources.

A. MAPS, SOILS, GEOGRAPHY, GEOLOGY

Lotan, James E., & Stephen G. Shetron. 1968. Characteristics of drumlins in Leelanau
County, Michigan. Pap. Mich. Acad. 53: 79-89. [Data on glacial drift.]

Redmond, C. E., & C. A. Engberg. 1967. Soil Survey of Arenac County, Michigan. U. S.
Dep. Agr. 142 pp. + 24 twice-folded map plates + [8] folded tables & indexes. [The
folded map plates include complete aerial photography for the county at a scale of
about 3n = 1 mile, with boundaries of soil types overprinted; hence an indispensible
aid for field work in the county. Fieldwork completed in 1954; series numbers are
now omitted from Soil Surveys. $2.25 from Govt. Printing Office, Washington, D.C.]
Weber, Herman L., et al. 1966. Soil Survey of Grand Traverse County, Michigan. U. S. Dep.
Agr., Series 1958, No. 34. 141 pp. + 92 once-folded map plates + [6] folded tables &
indexes. [The folded map plates include complete aerial photography for the county
at a scale of about 4n = 1 mile, with boundaries of soil types overprinted. Like the
other recent soil surveys which include such photography, indispensible to anyone
wanting quickly to survey the wooded and wild areas of the county. $4.75 from
Govt. Printing Office, Washington, D. C.]

B. BOOKS, BULLETINS, SEPARATE PUBLICATIONS

Ostrom, Arnold J. 1967. Forest Area in Michigan Counties, 1966. North Central For. Exp.
Sta., U. S. For. Serv. Res. Note NC-38. 4 pp. [Includes table with figures for all land,
non-commercial, and commercial forest area by counties.]

Ostrom, Arnold J. 1967. Forest Cover Types by Counties, Michigan, 1966. North Central
For. Exp. Sta., U. S. For. Serv. Res. Note NC-41. 4 pp. [Includes table showing, for
each county, total commercial forest land area (excluding parks and other reserved
areas) and acreage in each of 6 major forest types.]

Stone, Robert N. 1967. Forest Stand-Size Trends in Upper Michigan, 1955-1966.- North
Central For. Exp. Sta., U. S. For. Res. Note NC-39. 2 pp. [Table of acreages based on
1963-1966 survey ; general comparisons with 1946-1957 data.]

C. JOURNAL ARTICLES

Bassett, I. J. 1967. Taxonomy of Plantago L. in North America: Sections Holopsyllium
Pilger, Palaeopsyllium Pilger, and Lamprosantha Decne. Canad. Jour. Bot. 44:
565-577. [ P . cordata cited from Lansing and shown at two stations in Michigan on
distribution map.]

Catana, A. J., Jr. 1967. Forests of the Harvey N. Ott Preserve. Am. Midi. Nat. 78: 496-507.
[Quantitative study in Calhoun Co.]

Laing, Charles C. 1967. The ecology of Ammophila breviligulata. II. Genetic change as a
factor in population decline on stable dunes. Am. Midi. Nat. 77: 495-500. [This
corrects erroneous citation in Mich. Bot. 7: 28. 1968.]

1968

THE MICHIGAN BOTANIST

189

La Roi, George H. 1967. Ecological studies in the boreal spruce-fir forests of the North
American taiga. I. Analysis of the vascular flora. Ecol. Monogr. 37: 229-253. [Based
on data from stations across the continent, all Canadian except for one black spruce
stand and one white spruce-fir stand in Marquette Co. (located only by latitude and
longitude).]

Lewis, Walter H., Royce L. Oliver, & Yutaka Suda. 1967. Cytogeography of Claytonia
virginica and its allies. Ann. Missouri Bot. Gard. 54: 153-171. [Chromosome counts
for material from Berrien Co. given as 2n = 24, 25, & 26.]

Milanez, Adauto I., & Everett S. Beneke. 1968. New records of aquatic Phycomycetes from
Michigan. Pap. Mich. Acad. 53: 11-22. [Records from Kalamazoo and Barry coun¬
ties.]

Porsild, A. E. 1966. Contributions to the flora of southwestern Yukon Territory. Natl. Mus.
Canad. Bull. 216 (Contr. Bot. IV): 1-86. [159 distribution maps generally show no
stations beyond Alaska and Canada, except apparently for Poa canbyi and Phacelia
franklinii on Isle Royale.]

Pringle, James S. 1967. The fringed gentians— observations on their life histories in Ontario.
Wood Duck 21: 14-17. [Includes observations on Waugoshance Point (Wilderness
State Park, Emmet Co., Michigan).]

Roscher, John P. 1967. Alga dispersed by muskrat intestinal contents. Trans. Am. Micr. Soc.
86: 497-498. [13 species of viable algae recovered from cultures from muskrats
trapped in Kalamazoo Co.]

Schuyler, Alfred E. 1967. A taxonomic revision of North American leafy species of Scirpus.
Proc. Acad. Nat. Sci. Phila. 119: 295-323. [Cites chromosome counts for S. atro-
cinctus (n = 34) from Gladwin Co. and S. cyperinus (n = 33) from Cheboygan Co.]

Thomson, John W. 1967. The lichen genus Baeomyces in North America north of Mexico.
Bryologist 70: 285-298. [B. rufus is cited from Alger and Iron counties and Isle
Royale and mapped at these locations.]

Publications of Interest

/

PICTORIAL HISTORY OF MICHIGAN: The Early Years. By George S. May. William B.
Eerdmans Publ. Co., Grand Rapids, 1967. 239 pp. $15.00. Essentially a picture book
pertaining to Michigan before the 20th Century; the text is limited to helpful captions. A
number of the illustrations are reproductions of old maps, there is a section on “The Age
of the Lumberman,” and other features of natural history interest are shown. Douglass
Houghton is mentioned, as a geologist, but botanists might wish for more about the
earliest botanical work in the state (Zina Pitcher, Abram Sager, John Wright are nowhere
mentioned). The volume is fortunately well indexed, and is handsomely printed in a
large format; it is highly fitting that the printing, typography, and binding were all done
in Michigan.

THE LICHENS OF OHIO Part I. Foliose Lichens. By Conan J. Taylor. Biol. Notes 3, Ohio
Biological Survey, Columbus, 1967. 151 pp., plastic bound. $5.00 from Mailing Room,
OSU, 242 W. 18th Ave., Columbus, Ohio 43210. This work, which should be quite
useful in at least southern Michigan, is thoroughly illustrated with 120 photographs of
the species, and includes keys, descriptions, maps showing distribution (by counties) in
Ohio, and instructions for collecting and studying lichens. There is a checklist (with
nomenclatural equivalents) of the lichens of Ohio. Part II, completing the work, is
expected within a year.

190

THE MICHIGAN BOTANIST

Vol. 7

Review

A FIELD GUIDE TO WILDFLOWERS of Northeastern and North-central North America.
By Roger Tory Peterson and Margaret McKenny. Houghton Mifflin Co., Boston. 1968. 420
pp. $4.95.

“For some years we have been looking forward to the promised publication” of this
field guide-so we wrote in the very first issue of this journal (p. 44) six years ago! At last,
the long overdue volume is out, and it will be warmly welcomed by many wildflower en¬
thusiasts. The illustrations are very good, though the exact shades of the colored ones often
leave something to be desired. All grasses, sedges, rushes, and pondweeds are omitted but
otherwise the selection is unusually complete for a truly pocketsize book (nearly 1300
species, both native and alien). Inevitably there are points at which one might differ over
the selection of species which the average layman “is most likely to encounter.” Hieracium
florentinum is one of the most widespread and conspicuous weeds of roadsides and fields
throughout northern Michigan, but it is not included. The thousands of visitors to Michigan’s
shoreline parks and beaches are surely far more likely to encounter and wonder about such
showy plants as Ids lacustris, Cirsium pitched, and Anemone multifida than they are the ex¬
tremely rare and local orchids Cypripedium passerinum and Tdphora trianthophora, but the
latter are included and the former locally common ones are omitted.

A certain impatience with professional botanists is evident, to the point of inaccu¬
racy. Much is made (p. x and p. 90) of the fact that Gleason’s New Britton and Brown has
but 6 species of Antennada while Fernald’s edition of Gray’s Manual includes 32; it seems
unfair not to add that at least 13 of the latter are reported only from Newfoundland or
other portions of Canada specifically excluded from the range covered by Gleason. There
is a great amount of useful identification information (some of it a bit subjective) and, as
noted, a surprisingly large number of species in this compact guide. It is an excellent buy at
$4.95. Having made this point, with emphasis, let me go on to deplore th e arrangement of
this material which might be described as nearly chaotic.

There is no key or outline of any sort whatsoever. The fundamental grouping is by
color (not always of the flowers), with subdivisions often headed by such characters as
“Loose clusters or sprays of dangling flowers” or “Loose clusters of dangling bells” (the lat¬
ter for Prenanthes!). Surely the vagueness of this sort of terminology is no improvement over
the few essential botanical terms. And what would Roger Tory Peterson say about a bird
guide arranged by color (of, presumably, adult male plumage); or Roger Conant about a
guide based on snake and frog color; or A. B. Klots to a butterfly guide arranged by color?
From this standpoint, this number of the Field Guide series is distinctly retrograde. If color
is so useful for identification, then why did the authors not simply provide a key based on
color instead of technical characters?

The trouble with using color is that it simply doesn’t work for too many of the
plants. The authors have had to include a great many species under two or more colors, thus
consuming valuable space. And there remain dozens of additional species of which white or
other color forms are known but which for some reason are not included. Countless internal
contradictions could be cited. For example, Polygonum lapathifolium is listed only under
pink, though said to be “more often greenish white.” Tritium erectum is included in two
places under red and is said to be occasionally “greenish, white, etc.,” but it is listed under
no other color. Castilleja coccinea appears only under red, although the flowers are cor¬
rectly described there as “greenish-yellow.” Acorus calamus is shown under both yellow and
green, at the latter point said to have “no spathe,” although the figure clearly shows the long
spathe and the very next sentence refers to the “stem that bears the spathe.”

Fortunately there is a list of families with brief diagnoses in the front of the book,
each with a silhouette symbol which is rather cleverly done and which, together with family
name, appears with the species. So the effect of color-arrangement is somewhat overcome
and the reader is likely to gain some understanding of the families after all. Those who do
not mind leafing through a couple hundred pages to find related species which differ slightly
in color will no doubt be delighted with this otherwise highly recommended volume.

-E. G. V.

PROGRAM NOTES

Sept. 27-29, 1968: Fall Campout, Southeastern Chapter, Michigan

Club. Sleeper State Park is the place.

Botanical

Editorial Notes

The large size of this and some other issues results from the generosity of
authors (and their institutions or grants) who enable us to maintain our editorial
policy that all pages in excess of 10 in any article must be subsidized and that
subsidy of shorter articles (or of even the first 10 pages of longer ones) en¬
courages prompt publication. Subscription income alone would allow four issues
of approximately 32 pages each per year.

Good black and white prints of botanical subjects, suitable for use on the
cover of TF1E MICHIGAN BOTANIST, are always welcome and may be submit¬
ted to the editor at any time.

The March number (Vol. 7, No. 2) was mailed March 13, 1968.

Erratum

Vol. 7, p. 89, line 2 from bottom: for “C. caroliniana ” read C. virginica.

Publications of Interest

THE KOMAROV BOTANICAL INSTITUTE. 250 Years of Russian Research. By Stanwyn
G. Shetler. Smithsonian Institution Press, Washington, D. C., “1967” [1968]. 240 pp.
$5.95, distributed by Random House, Inc. The relevance of this volume to the Great
Lakes region is perhaps remote, but those interested in the history of botany, particular¬
ly at institutions which emphasize plants of northern latitudes, will find it a welcome
alternative to much digging around in the Russian language. This is a popular account,
fully illustrated, of the history of a rich tradition of botanical scholarship, chiefly in
taxonomy, over two and a half centuries and involving several institutions founded by
Peter the Great, culminating in the presentday Komarov Botanical Institute of the
Academy of Sciences of the USSR. The history of the monumental 30-volume Flora
SSSR is given in some detail, and there is material on the vast library, herbarium (nearly
6 million specimens), and other facilities as well as on the men who built up the
Botanical Garden and Botanical Museum which in 1931 were merged into the Botanical
Institute: Siegesbeck, Trinius, C. A. Meyer, Regel, Ruprecht, Maximowicz, Komarov,
Schischkin, and others.

LIMNOS The Magazine of the Great Lakes Foundation. Vol. I, No. 1, Spring 1968. Pub¬
lished four times a year, included with membership ($5.00) in the Great Lakes Founda¬
tion, 2200 North Campus Blvd., Ann Arbor, Michigan 48105. The first number of this
new periodical solicits articles and illustrations “dealing with discoveries or advances in
our knowledge of the Great Lakes”; and it includes articles on research by submarine in
Lake Michigan, water pollution, topography of the floor of Lake Michigan, the coho
salmon, and Great Lakes water levels.

CONTENTS

The Status and Distribution of Gentiana linearis and G. rubri-
caulis in the Upper Great Lakes Region

James S. Pringle . 99

Teratological Stamens and Carpels of a Willow from Northern
Michigan

Warren H. Wagner, Jr . 113

A Preliminary Report on the Distribution of Gymnosperms in
Michigan

Edward G. Voss . 121

Review of the Status of Some Green Algae in the Genus Coela-
strum

Mason G. Fenwick . . . 129

Bryophytes New to Michigan

Howard Crum & Norton G. Miller . 132

Distributional History of Butomus umbellatus (Flowering-Rush)
in the Western Lake Erie and Lake St. Clair Region

Ronald L. Stuckey . 134

The Cantharellaceae of Michigan

Alexander H. Smith . 143

Nature Education Feature-

Making Rubber Duplicates of Mushrooms

Richard A. Mortemore . 184

Michigan Plants in Print . 188

Publications of Interest . 189, 191

Review— A Field Guide to Wildflowers . 190

Program Notes . 191

Editorial Notes . . . 191

(On the cover: Dune grass, Elymus mollis Trin.,
on the shore of Lake Superior at Whitefish Point, Michigan.

Photographed July 20, 1967, by John S. Russell.)

THE

Vol. 7, No. 4

MICHIGAN BOTANIST

BOTANICAL GARDEN

October, 1968

THE MICHIGAN BOTANIST-Published by the Michigan Botanical Club four times per

year: January, March, May, and October. Second-class postage paid at Ann Arbor,

Michigan.

Subscriptions: $3.00 per year. Single copies: $.75.

All back issues are available, at the following prices per volume:

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with the first number of that volume unless specified to the contrary.

Articles and notes dealing with any phase of botany relating to the Upper Great Lakes
Region may be sent to the editor in chief. The attention of authors preparing manuscripts
is called to “Information for Authors” (Vol. 6, p. 202; reprints available from the editor).

Editorial Board

Edward G. Voss, Editor in Chief

Herbarium, The University of Michigan, Ann Arbor, Michigan 48104

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1509 Kearney Rd., Ann Arbor, Michigan 48104

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THE MICHIGAN BOTANICAL CLUB

Membership in the Michigan Botanical Club is open to anyone interested in its aims:
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Dues are modest, but vary slightly among the chapters and with different classes of
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Secretary, Huron Valley Chapter:

Ursula R. Freimarck, 704 Madison PI., Ann Arbor, Michigan 48103

1968

THE MICHIGAN BOTANIST

195

SEX DETERMINATION IN MOSSES

Dale H. Vitt

Herbarium, The University of Michigan, Ann Arbor

Genetically determined dioecism is one of the mechanisms making cross¬
fertilization obligatory in plants and animals. In fungi this is accomplished by
heterothallism with many species being bipolar, that is, with one set of genes
controlling compatability, whereas other species are tetrapolar, with two sets of
genes controlling compatability. It has been found recently (Raper, 1966) that
in tetrapolar basidiomycetes one set of genes, “aa,” controls the formation of
clamp connections and a second pair, “bb,” controls the migration of nuclei. In
the latter case, if fungi containing only the “aa” pair of alleles are mated, clamp
connections are formed, but there is no nuclear migration and thus no fertiliza¬
tion.

In many higher plants cross-fertilization and incompatability are deter¬
mined by a multiple allelic system. If the pollen and the female system contain
identical alleles, the pollen tube will not grow.

In accord with these incompatability mechanisms, dioecism may operate
at either the haploid or diploid level and thus prevent self-fertilization. Dioecism
is under the control of separate systems in haploid and diploid types of organ¬
isms (see figure 1). These two types of sex control systems may be distinguished
on both genetical and morphological grounds. Haploid dioecism occurs in many
algae and bryophytes and diploid dioecism, in many higher plants and animals.

The control of sex and consequently dioecism in some higher plants and
most animals is known to be by X and Y chromosomes. In these cases, the males
are heterogame tic (XY) and the female homogametic (XX). In this type of
system the Y chromosome is of considerable importance in determining the
phenotype of the individual. However, other mechanisms are known, such as the
balance concept of sex in Drosophila where the ratio of X’s to autosomes deter¬
mines the sex. If the ratio is below one, then the fly is a male; if it is above one,
it is a female. In this case the Y chromosome plays a relatively minor role in
determining the phenotype. A third condition is found in some Lepidoptera,
birds, and reptiles. If the animal contains XX it is a male, and if the genotype is
XO [no Y chromosome] it is a female. Here, then, the heterogametic sex is the
female.

In haploid plants, such as bryophytes and algae, the sex mechanism differs
from that of diploid plants in that both differential segments always behave as
units of heredity. In diploid plants there are two types of combinations possible
in the sporophyte generation, the XX and the XY; whereas in haploid plants
there is only one type possible, the XY. In diploid organisms the X chromosome
undergoes recombination in the homogametic sex; this never takes place in
haploid organisms as the X of female plants always pairs with the Y of the male
plants.

196

THE MICHIGAN BOTANIST

Vol. 7

PHENOTYPIC EXPRESSION OF SEX IN MOSSES

In bryophytes, dioecious species result when spores of two types produce
two different plants: the male containing only the Y chromosome and the
female containing only the X. This situation is not analogous to those higher
plants where a dioecious species is composed of male plants containing XY and
female plants containing XX. In bryophytes the XY condition is asexual and not
male as in diploid organisms.

Among diploid organisms the differential segment of the Y chromosome
never undergoes recombination, and many geneticists regard this as the reason
for the erosion and reduction in size of the Y chromosome during the course of
evolution. In haploid systems neither the X nor the Y undergoes recombination,
and yet the X chromosome is not eroded and reduced in size. From this com¬
parison of evidence, lack of recombination cannot explain the small size of the Y
chromosome.

Sex in mosses is shown phenotypically in several ways: (1) structure and
position of sex organs on the plant, (2) size of the individual plants, (3) size of
spores, and (4) type of gametes produced. The position of the sex organs can be
summarized as follows. Plants that are bisexual (fig. 2) are termed monoecious.
However, in mosses several different arrangements are possible. If the archegonia
and antheridia are in separate inflorescences, but on the same plant, the species is
called autoecious (figs. 3-6). Included in this type of arrangement are (a)
gonioautoecious (figs. 3, 4, 6), with the antheridia axillary or budlike below the

DIPLOID DIOECISM

XX

XY

IN

HAPLOID DIOECISM

Diploid

Sporophytes

Meiosis

XX X Y

1! II

Haploid

Gametophytes

9

cr

X Y

9 cr

XX

XY

Fertilization

XY

Fig. 1. A comparison of haploid and diploid dioecism. (Modified from Lewis, 1961)

1968

THE MICHIGAN BOTANIST

197

archegonia and (b) cladautoecious (fig. 5), with antheridia and archegonia on
separate branches. Many mosses, such as the common Mnium medium BSG, have
the antheridia in the same inflorescence as the archegonia (fig. 11). If these are
mixed together, the term synoecious is used. However, if the antheridia and
archegonia are in the same inflorescence but not mixed, the term paroecious is
used. Frequently the genera Poly trichum and A trichum have antheridia and

Fig. 2. Monoecious plants with male buds of Orthotrichum speciosum. Fig. 3. Gonio-
autoecious plants of Drepanocladus sp. with male buds below seta of capsule. Fig. 4.
Closeup of male buds of Drepanocladus sp.; the antheridia enclosed within the buds. Fig. 5.
Cladautoecious condition of unknown species; male bud on separate branch from female,
which is represented by the sporophyte. Fig. 6. Gonioautoecious condition of Orthotrichum
speciosum; male buds at arrows.

198 _ THE MICHIGAN BOTANIST _ Vol. 7

archegonia on the same plant in separate inflorescences and also mixed together;
this condition is referred to as polyoecious or heteroecious.

Of more important consideration here are the dioecious species. In some
cases, such as Mnium affine Bland, ex Schwaegr. and A trichum undulatum
(Hedw.) P.-Beauv. (figs. 8, 9), the male and female plants are undifferentiated. In
Orthotrichum lyellii Hook. & Tayl. (fig. 7), the male plants are slightly smaller
than the female plants. In some species of Macromitrium, Holomitrium, and
Eurhynchium the male plants are greatly reduced to what are called dwarf males
(figs. 10, 12). These are, in fact, very small male plants which grow attached to
the stems, leaves, or rhizoids of the larger female plants.

In addition to the above manifestations, many dioecious species, especially
in the family Orthotrichaceae, phenotypically express sex in the form of
“heterospory” or as will be explained later, anisospory. In this family, which is
mainly tropical, there are three large genera, Ortho trichum, Macromitrium, and
Schlotheimia, in addition to a few smaller genera. In this group, anisospory
appears to be correlated with a reduction in the Y chromosome and often a
reduction in the size of the male plants.

Fig. 7. Slight sexual dimorphism of Orthotrichum lyellii with male plant on right. Fig. 8.
Male and female plants of similar size of A trichum undulatum. Fig. 9. Male inflorescence on
left and sporophyte arising from female plant on right of A trichum undulatum .

1968

THE MICHIGAN BOTANIST

199

By means of spore counts, size frequency has been obtained for the spores
of many species. Figures 13 through 16 illustrate a few of these counts. Species
in which there are two sizes of spores show a bimodal peak. In Macromitrium
there seems to be a distinct correlation between anisospory and the occurrence
of dwarf males. The homosporous species of Macromitrium are all monoecious,
and as seen in figures 13 and 19 their spores are fairly constant in size (compare
this with figs. 14, 17, 18, which illustrate anisosporous spores). In Orthotrichum
there seems to be one dioecious species that has spores of two sizes, namely O.
lyellii Hook. & Tayl. (fig. 16), whereas at least one other dioecious species, O.
obtusifolium Brid. (fig. 15), appears to be homosporous.

Thus in mosses there are four types of dioecism:

Homosporous with no morphological dimorphism e.g. Orthotrichum
obtusifolium Brid.

Homosporous with dwarf males e.g. Eurhynchium pulchellum
(Hedw.) Jenn.

Anisosporous with slight dimorphism of plants e.g. Orthotrichum
lyellii Hook & Tayl.

Anisosporous with dwarf males e .g. Macromitrium species

However, in no case does anisospory appear with monomorphic dioecious spe¬
cies. This is significant in that both characteristics may be advanced.

HETEROSPORY AND ANISOSPORY

Heterospory can be defined as the production of two sizes of spores by
one plant. In all higher plants, the micro- and megaspores are produced in
separate sporangia. It follows that all products of a single meiotic division will be

Fig. 10. Cauline dwarf male on stem of female plant of Holomitrium sp. Fig. 11. Synoecious
condition of Mnium medium.

200

THE MICHIGAN BOTANIST

Vol. 7

Fig. 12. Dwarf males of
Eurhynchium pulchellum.

Fig. 13 Fig. 14

Fig. 15

Fig. 16

Fig. 13. Spore size distribution of monoecious (homosporous) species of Macromitrium (M.
ferriei, M. incurvum, & M. holomitrioides) . Fig. 14. Spore size distribution of dioecious
(anisosporous) species of Macromitrium (M. comatum & M. prolongatum). Fig. 15. Spore
size distribution of three dioecious species: Orthotrichum obtusifolium, Eurhynchium
pulchellum, & Schwetschkeopsis fabronia. Fig. 16. Spore size distribution of Orthotrichum
lyellii from three localities.

1968

THE MICHIGAN BOTANIST

201

all microspores or megaspores. This is the case in the heterosporous species of
Selaginella and Isoetes as well as the heterosporous Filicales, Marsiliaceae, and
Salviniaceae. In all these cases the gametophytes are endosporic.

In mosses and hepatics the micro- and megaspores are produced in the
same sporangium and both types of spores result from the same meiotic division,
two of the products being microspores and two, megaspores. In no case have
sporangia of two sizes been found. Greguss (1964) reported size differences in
sporangia of Fontinalis antipyretica Hedw. and Brachymenium mexicanum
Mont.; however, I can find no evidence of size differences in spores or sporangia
in these two species. As opposed to the case in true heterospory, bryophyte
gametophytes are exosporic. Thus the mechanism of sex and differential spore
size, as well as development, is not the same in higher plants as in bryophytes.
Since the condition in higher plants is termed heterospory, the condition in
bryophytes requires a new term, anisospory.

The above evidence allows speculation that the anisosporous type of spore
development could not have given rise to the heterosporous type. Instead it
seems probable that the anisosporous species of mosses have evolved parallel to
the higher plants with heterospory.

Fig. 17. Spores of Macromitrium comatum. Fig. 18. Enlarged view of spores of M. com-
atum. Fig. 19. Spore of Orthotrichum obtusifolium. Fig. 20. A probable instance of chrom¬
osome doubling (autopolyploidy) in a dioecious species, Orthotrichum lyellii.

202

THE MICHIGAN BOTANIST

Vol. 7

DIMORPHISM OF SEX CHROMOSOMES

Evidence available at present indicates that dimorphism of spores and male
plants is connected with dimorphism of sex chromosomes. In 1909, Douin
found that the spores of Sphaerocarpus germinate in tetrads giving rise to two
male and two female plants in groups. He concluded that sex differentiation
takes place in spore mother cell differentiation. Allen (1917), working with the
liverwort Sphaerocarpus donnellii, correlated a chromosome difference with the
dioecism. He found 14 autosomes of equal size and two unequal chromosomes
in diploid sporophyte cells. This was the first time that sex chromosomes of any
kind were reported. Early workers also found some chromosomes in mosses to
be completely heterochromatic. Heitz (1928) called the larger of these the M
and the smaller the m. In 1944, Ernst-Swarzenbach concluded that the dioecism
of Macromitrium was controlled by an XY mechanism. Ramsay (1966) showed
by photographs that at the meiotic division of the spore mother cells there is a
segregation of a large and a small chromosome which form a dimorphic bivalent
at metaphase.

Working with hepatics, Tatuno (1941) termed the large heterochromatic
chromosome H and the smaller h. Yano (Anderson, 1964) found that out of 94
dioecious species studied, 90 had only one H chromosome. He has postulated
that in monoecious species there are two H chromosomes and that these species
are either aneuploids or polyploids. Furthermore, in Mnium, Lowry (1948)
found that monoecious taxa may originate from dioecious ones by autopoly¬
ploidy. Heitz (Schuster, 1966) found the following correlations:

Chromosome

Species

Percent

number

studied

Monoecious Dioecious

n= 9

33

27 73

n=18

14

100 (93)? 0 (7)

SUMMARY

Evidence indicates that the evolution of sex as a means of out-breeding has evolved in
numerous ways. In those organisms which contain sex chromosomes, a relationship exists
between heteropycnosis and sex chromosomes. These heteropycnotic chromosomes can best
be referred to as H and h chromosomes. Furthermore, in many monoecious species of
bryophytes the pairing of these is H/H and h/h, not H/h.

At least in Macromitrium, dioecism is associated with morphologically different
chromosomes; however, anisospory exists in Orthotrichum lyellii with no morphological
difference in the sex chromosomes. Some species of bryophytes have two H chromosomes
and if these species are polyploids, they are monoecious. This polyploidy may be auto¬
polyploidy (fig. 20). One can speculate that dioecism is primitive and monoecism is derived
from it. In accordance with this, anisospory is probably an advanced character occurring
only in species in which polyploidy has not taken place.

ACKNOWLEDGMENTS

I wish to thank Dr. Howard Crum and Mr. Joseph Ammirati for their critical reading
of the manuscript. I also would like to thank Mr. Louis Martonyi and Mr. Frank Anderson
for assistance and use of the photographs.

1968

THE MICHIGAN BOTANIST

203

LITERATURE CITED

Allen, C. E. 1917. A chromosome difference correlated with sex differences in Sphaero-
carpus. Science 46: 466-467.

Anderson, L. E. 1964. Biosystematic evaluations in the Musci. Phytomorphology 14: 27-51.
Douin, C. I. 1909. Nouvelles observations sur les Sphaerocarpus. Revue Bryol. 36:37-41.
Emst-Swarzenbach, Marthe. 1944. La sexualite et la dimorphisme des spores des mousses.
Revue Bryol. Lichenol. 14: 105-113.

Greguss, P. 1964. The phylogeny of sexuality and triphyletic evolution of the landplants.
Acta Univ. Szeged., N.S. 10: 1-50.

Heitz, E. 1928. Das heterochromatin der moose. I. Jahrb. Wiss. Bot. 69: 762-818.

Lewis, K. R. 1961. The genetics of bryophytes. Trans Brit. Bryol. Soc. 4: 111-130.

Lowry, R. J. 1948. A cytotaxonomic study of the genus Mnium. Mem. Torrey Bot. Club
20: 1-42.

Raper, J. R. 1966. Genetics of Sexuality in Higher Fungi. Ronald Press, New York. 283 pp.
Ramsay, Helen P. 1966. Sex chromosomes in Macromitrium. Bryologist 69: 293-311.
Schuster, R. M. 1966. The Hepaticae and Anthocerotae of North America. Vol. I. Columbia
Univ. Press. N.Y. 802 pp.

Tatuno, S. 1941. Zytolhgische Untersuchungen fiber die lebermoose von Japan. Jour. Sci.
Hiroshima Univ., B. (Div. 2) 4: 73-187.

MOSQUITO POLLINATION OF

HABENARIA OBTUSATA (ORCHID ACEAE)1

Warren P. Stoutamire

Department of Biology, University of Akron, Akron, Ohio

In early July, 1912, Miss Ada Dietz, working in Reese’s Bog at the Univer¬
sity of Michigan Biological Station (Cheboygan Co., Michigan), saw a mosquito
whose head bore two yellow objects resembling pollen masses. This was reported
to John Dexter who also visited the bog and captured half a dozen female
mosquitoes with from 1 to 4 pollen masses on their eyes. The objects were
pollinia of Habenaria obtusata (Banks ex Pursh) Richards, which was flowering
in the vicinity. Dexter gathered pollen-free mosquitoes and flowering plants of
the orchid, enclosed both in an aquarium, and in a few days the insects were
carrying orchid pollinia on their eyes in the same position as those caught in the
bog (Dexter, 1913). The mosquitoes were not identified and the behavior of the
insects on the flowers apparently was not observed. This report of mosquitoes as
orchid pollinators was the first of several involving this plant. Specimens of the
insects have been observed carrying the pollinia in the Athabaska and Great
Slave Lake areas (John W. Thomson, pers. comm.; Raup, 1930), in the vicinity
of Churchill (Hocking et al., 1950; Hocking, 1953), and near Bear Lake by

^Research supported by NSF grant GB5784X. I wish to thank the members of the
Huron Mountain Wildlife Foundation for the use of their facilities during part of this work.

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Porsild (Twinn et al., 1948). The identified mosquitoes were all females in the
genus Aedes. West and Jenkins (1951) reported that mosquitoes caged with
Habenaria obtusata containing P32 fed on the orchids and became radioactive,
but mosquitoes also visited five other species treated with P32 (Salix, Ledum,
Rhododendron, Dryas, and Rubus species) and fed on radioactive nectar in these
flowers as well.

Many observations of mosquitoes resting or feeding on flowers are re¬
corded in the literature; Hocking (1953) and Haeger (1955) list plants distri¬
buted among about 30 plant families. Mosquitoes feed on sugar secretions from
both floral and extrafloral nectaries as well as on aphid honeydew. Nectar is an
important part of the diet of tundra mosquitoes, 45-95% of the weight of fed
insects being due to this (Hocking, 1953). Tropical and subtropical mosquitoes
also take flower nectar, and species of the genera Toxorhynchites (Megarrhinus)
(Jenkins & Carpenter, 1946, Jenkins, 1949) as well as Liponeura and Apistomyia
(Porsch, 1958) have the proboscis so modified that the insects are probably
incapable of sucking blood and may rely entirely on other liquids as energy
sources. In spite of numerous observations of mosquitoes on flowers, these

Fig. 1-2. Face and submedian section of Habenaria obtusata flower.

1-labellum, n-nectary, o-ovary, a-appendage, v-viscidium of pollinium, p-pollen mass in
anther locule, s-stigma.

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205

insects appear to be transient visitors to most flowers and are of little impor¬
tance as pollen transfer agents. To date we know of only one plant whose pollen
has repeatedly been found on mosquitoes and that plant is the subject of this
paper.

Habenaria obtusata has a general nearctic distrubution, occurring from
Labrador and the New England states westward to the Aleutian Islands and
British Columbia. The closely related or conspecific H. parvula (Platanthera
parvula in European usage), occurring from eastern Siberia to Scandinavia, com¬
pletes the circumpolar distribution. To my knowledge no insect visitors to the
latter orchid have been reported. Habenaria obtusata inhabits cool coniferous
forest areas where midsummer soil temperatures rarely exceed 15° C. In Mich¬
igan it often is rooted in moist humus which may be very wet during the early
part of the growing season. Plants also colonize well drained sandy humus under
pines and spruce near Lake Superior. Seedlings are subterranean for several
months, and then produce a single leaf during the first season of growth above
ground. Lateral droppers are produced which give rise to new plants near the
parental axis the following season, as is characteristic of most members of this
subfamily Orchidoideae. The plants do not form wide-spreading clones as do H.
flava, H. clavellata, and Orchis rotundifolia by the growth of droppers several
centimeters away from the parental axis. When plants reach flowering size they
usually produce one leaf and a few-flowered inflorescence 10-20 cm in height
(Fig. 3). The individual whitish-to-greenish flowers are galeate, with the hood
usually held horizontally and the labellum (“lip”) pendent or recurved below
(Fig. 1). The nectar spurs in Michigan plants are 7-10 mm long, about 5 mm of
which contains nectar at the time of anthesis. I can detect no odor to the
flowers, although Raup stated that they were scented.

The stigmatic surface is between the 2 half-anthers and directly above the
nectary opening (Fig. 2). The base of the labellum bears a ligulate appendage
which projects backward, partially closing the nectary lumen. The shape of the
appendage and the degree of obstruction differ from plant to plant, and also
change as flowers age. Entrance to the nectary is possible from the sides, where
canal-like passages are formed between the adjacent walls of the nectary and
labellum appendage. The adhesive gland (viscidium) of each pollinium is situated
to the side of these canals. An insect inserting its proboscis through one of the
canals will contact the adjacent viscidium.

I had the opportunity to observe three flowering colonies of this orchid in
Michigan during July, 1967, and to watch for insect visitors. The first observa¬
tions were made in a Thuja-Picea woodland near the shore of Albany Bay,
Mackinac Co., on July 3. Air temperature was 12°C between 2 and 4 P.M., and
mosquitoes were present in large numbers but lethargic. No insect visitors of any
kind were attracted to the 20 plants under observation during this period. The
greatest plant feeding activity of mosquitoes in Manitoba (Twinn et al., 1948), in
Florida (Haeger, 1955), and in Denmark (Nielsen & Grieve, 1950) occurs before
the dusk and after the dawn mating swarms. Twinn observed mosquitoes visiting
flowers of Habenaria obtusata at dusk but not at other times. This periodic
feeding and the low air temperature possibly account for failure to observe the
process in the field during this limited period.

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1968

THE MICHIGAN BOTANIST

207

Mosquitoes from this site were collected and caged with inflorescences
using a glass box made of 3.5 x 4.5" slide glasses taped at the edges. The insects,
in a room temperature of 20-25°C, alighted on the inflorescences within a few
minutes after the 7 PM caging. Two pollinia were removed from flowers before
equipment was ready for accurate observation. Several insects were followed
through the pollinium removal process during the following hour. An insect
alights on any projecting part of the inflorescence but especially on the lateral
sepals, rests for an indefinite period ranging from a few seconds to several
minutes, and then begins probing the surface of the flower with the proboscis.
This probing involves the front and rear surfaces of the sepals and petals, and
continues until the insect is perched across the face of the flower parallel to the
labellum. At this point the proboscis is directed to the region of the nectary
opening. The insect appears to be poorly oriented for finding the nectary and
the searching movements appear awkward. The mosquito attempts to force the
proboscis into the nectary but is inhibited by the obstructing appendage. When a
proboscis is inserted in one of the lateral canals and pushed into the spur (Fig.
4), the compound eye adjacent to the viscidium contacts this structure, and the
pollinium is attached either during feeding or as the head is withdrawn. A
mosquito maintains this feeding position for a minute or longer before with¬
drawing from the nectary. The pollinium, attached to an eye, is withdrawn from
its half-anther (Fig. 5), and the insect immediately attempts to remove the
attached pollinium with one or both front legs. Because of the lateral approach
to the nectary only one pollinium is removed per visit. An insect with a pol¬
linium attached to one eye rests for indefinite periods and then visits other
flowers. The pollinium projects forward parallel to the proboscis, and inhibits
the insertion of the proboscis into other nectaries as it strikes the surrounding
structures. Repeated attempts to insert the proboscis usually seem to result in
the proboscis being inserted in the opening opposite to the one from which the
first visit was made because of the obstructing attached pollinium on that side.
Clusters of pollen grains (massulae) are left on the stigmatic surface during the
probing movements or during the insertion of the proboscis in the alternate
opening. Several insects removed two pollen masses, one on each eye, after
several hours of confinement with the inflorescenes (Fig. 6). Dexter found
mosquitoes carrying 3 or 4 pollinia, but insects observed in this study removed a
maximum of 2. The presence of one or more pollinia on a mosquito appears to
increase its searching and probing activity on flowers. This increased activity also

Fig. 3. Natural stand of Habenaria obtusata. Plants are 13-16 cm tall.

Fig. 4. Aedes intrudens inserting proboscis in nectary

Fig. 5. Aedes after feeding, with freshly attached pollinium on right compound eye.

Fig. 6. Aedes with two pollinia.

Fig. 7. Resting Aedes with 2 pollinia. Viscidium can be seen attached to lower part of com¬
pound eye.

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increases the chances of pollen massulae being left on stigmas with concomitantly
increased setting of fruit.

A small collection of mosquitoes flying in the study area were identified as
Aedes intrudens, A. canadensis, and A. fitchii 2 Of 36 specimens examined, 1
carried a pollinium but was unfortunately lost before identifications were made.

A collection of approximately 100 mosquitoes was made in another col¬
ony of Habenaria obtusata growing in a Thuja bog near St. Martin’s Point,
Mackinac Co. The flying msects were Aedes vexans, A. fitchii, and A. intrudens,
of which 1 of A. vexans carried a pollinium on the lower side of the left
compound eye. A collection of similar size was made in a Tsuga-Thuja bog near
Conway Creek, Marquette Co., on July 17 after most of the orchids had ceased
flowering. Aedes intrudens, A. diantaeus, A. canadensis, and A. fitchii were
flying, and one female of A. intrudens or diantaeus carried a pollinium on its
right eye. (The specimen cannot be precisely identified because females of these
two species are very difficult to separate.) Two to 3 hours were spent at each of
these bogs watching for visitors during daylight hours. Mosquitoes were active
but not visiting the orchids.

Flowering plants of Habenaria obtusata were brought to Akron, Ohio, well
south of the range of the species, and females of Aedes intrudens were captured
locally. These were caged with the orchid, and within a few minutes an insect
was probing a flower and shortly removed a pollinium. Of 10 insects caged, 4
removed 6 pollinia from an inflorescence over a 12-hour period. One was
attached to a left eye, one to a right eye, and 2 insects carried pollinia on both
eyes. Mosquitoes from populations which never come in contact with this orchid
were attracted to it and functioned in a normal manner.

DISCUSSION

Species of Aedes have long been known to visit and to remove pollinia
from Habenaria obtusata but no other insect visitors to the orchid have so far
been recorded. The labellar appendage appears to be a morphological adaptation
to insects with the general head dimensions of mosquitoes, and the species seems
to be unique among flowering plants in this regard. It appears strange that close
associations between other plants and mosquitoes have not been reported in
view of the prevalence of both mosquitoes and flowering plants in all the tem¬
perate land masses. Such associations may well exist but perhaps have not yet
been observed because Culicine research has been largely anthropocentric. The
physical discomfort attendant on mosquito-flower studies under natural condi¬
tions also tends to inhibit observation.

Mosquitoes are relatively primitive dipterans; compared with many other
groups of this order they are inactive and spend much of their time resting on
vegetation or in protective cavities. Their slow movements when alighted and
their long legs with short hairs suggest that they are inefficient carriers of the
non-sticky types of pollen. They are present in large numbers in many habitats,
however, and additiional plant-mosquito relationships can be expected.

2

Mosquito identifications by Dr. Carl Venard, Ohio State University, whose assistance
is gratefully acknowledged.

1968

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209

A considerable amount of information is available concerning mosquito
feeding stimuli. Airborne factors activating mosquitoes include rise in tempera¬
ture, increase in C02, increase in moisture, and presence of volatile organic
compounds (Clements, 1963). Mosquitoes are very sensitive to temperature
changes, especially in the range of 29-30°C, and can detect and react to a
temperature gradient of 0.05°C/cm (Bates, 1949). Some of the dipteran-
attr acting heavy textured Araceae have a considerable temperature rise at
anthesis(Meeuse, 1966) but this is thought to be associated with volatilization of
attracting compounds rather than as an attracting mechanism in itself. The
immediate reaction of mosquitoes to warmth might serve either as a short-range
attractant or in orienting them on a flower. It is highly unlikely that the small
and thin textured flowers of Habenaria obutsata could produce enough tempera¬
ture rise to attract the insects, because of the very rapid thermal radiation from
the flower and the accompanying very high respiration rate required for an
effective temperature rise. Very small temperature changes in the thicker tissues
of the center of the flower could serve to orient the mosquito in front of the
column, however, and flowers should be examined with this in mind.

Chemical mosquito attractants have been intensively investigated, and of
the many compounds tested, one consistently initiates feeding reactions. Carbon
dioxide -air mixtures from such diverse sources as burning candles, human breath,
and gas cylinders induce mosquitoes to fly upwind and to orient themselves in
relation to the source of carbon dioxide (Wright, 1968). Carbon dioxide is one
of the respiration products of plants, and it is possible that the flowers of
Habenaria obtusata have capitalized on this as a short-range feeding stimulant.
Mosquitoes become more active as relative humidity increases and local addition
of water vapor to the air may also affect feeding responses of mosquitoes on the
flowers. The orchids grow on the forest floor, often in very humid places, or
about the sides of depressions where Sphagnum mdRhodobryum mosses abound.
The relative humidity in such areas is consistantly higher than on the surrounding
higher ground. Local addition of more water vapor in such habitats would seem
to be an ineffectual feeding stimulant. Little can be said of the volatile organic
compounds which attract biting mosquitoes, since most of the work done with
mosquito-attracting compounds involves animal skin secretions or their bypro¬
ducts. The flowers have little or no odor detectable to me, but organic com¬
pounds may be produced to which mosquitoes react.

Mosquito reactions to color are equivocal, but the insects generally prefer
surfaces of low reflectivity. If Aedes visits to Habenaria usually take place in the
dim light of dawn or dusk in an already heavily shaded environment, flower
color probably is of little importance to the insects. These flowers are white to
light green in color and are not differentiated from their surroundings in this
respect.

Repeated observation of mosquitoes in forested areas of northern Mich¬
igan and adjacent Ontario indicates that insects spend much of their time
clinging to vegetation, especially the undersides of leaves, to thin stems of herbs,
and to any convenient projecting object. The erect inflorescence of Habenaria
obtusata seems eminently suitable for such perching behavior, with its radially

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projecting flowers, each of which provides convenient footholds. Mosquitoes are
always plentiful in the vicinity of flowering plants and can be expected to alight
on the projecting inflorescences as a matter of course. Short range orienting
mechanisms could come into play after such contact. Limited observation of the
behavior of caged insects in the laboratory suggests that this may be the case.
Female mosquitoes perched on flowers for indefinite periods before starting the
probing reaction. This is very different behavior from their direct attack and
feeding on mammals.

Flowers of Habenaria obtusata are modified in two respects which appear
to represent adaptations to mosquito pollinators. The first is a short nectary
which is compatible, lengthwise, with the mosquito proboscis; and the second is
the labellum appendage at the entrance to the nectary. Other North American
species of Habenaria also bear calli on the face of the labellum, but they are
further from the nectary orifice, and the flowers are either larger in size or
appear to have internal dimensions incompatible with those of mosquitoes.
Hocking et al. (1950) reported twice seeing mosquitoes feeding on Habenaria
hyperborea (L.) R. Br. flowers, but pollinia were not removed. In some flowers
of this species the labellum at anthesis does not separate completely from the
galea but remains curved forward and upward as the flower expands. This pro¬
duces two lateral openings in the flower, one over each viscidium. Plants vary in
the degree of flower opening; the labellum in some becomes more or less com¬
pletely expanded and straight. The degree of opening in some plants also
increases as flowers age. The partially opened flowers of Habenaria dilatata
(Pursh) Hook, must also be entered laterally . Habenaria dilatata bears white and
very fragrant flowers and is visited and pollinated by Hymenoptera and
Lepidoptera, pollinia being attached to the visitor’s proboscis. The green or
cream colored flowers of the odorless or ill-smelling Habenaria hyperborea have
shorter nectaries than H. dilatata (3-7.5 vs. 5-10 mm) and appear to be of the
proper size for mosquito visits. I have never seen insects on the spikes of this
species, although colonies have been watched. Pollinia can often be found
attached to sepals and petals and to surrounding vegetation, where they have
obviously been detached by visitors. The visitors have not been identified, but
the restricted lateral opening of the flower, the dull color, and the short spur
suggest that mosquitoes could be effective pollinators. The poUinium length of
Habenaria hyperborea (1.5 mm) is the same as that for H. obtusata.

Pollinia attached to weak fliers, such as Aedes species, may inhibit move¬
ment of the insects. Fresh insects were not weighed during the investigation but
the average weight of 3.1 1 mg given for unfed Aedes communis (Hocking, 1953)
can be used as an approximation of the weight of A. vexans, which is of similar
size. The average individual weight of 10 Habenaria obtusata pollinia (refrig¬
erated for 9 months in sealed vials in this laboratory) is 0.06 mg. On this basis
each pollinium attached to a visiting Aedes vexans will add about 2% to the
weight of the insect. This appears to be very little in comparison with the 45%
and greater weight increase which a freshly fed insect supports in flight. The
position of the pollinia on the eyes may have considerably greater effect than
their weight, however. One or two such bulky objects, projecting forward near

1968

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211

the antennae may radically change the maneuverability of flying mosquitoes. We
do not know whether they in any way shorten the life of the insect, but they do
interfere with feeding at other Habenaria flowers.

All identified pollinia-carrying mosquitoes have been females. Twinn made
several collections in the Churchill area and found from 2 to 33% of the female
mosquitoes captured carried pollinia. Hocking (1953) gives some very high per¬
centage figures for female mosquitoes which are presumed to have visited
Habenaria obtusata, the figures increasing from less than 10% shortly after
flowering begins to over 70% 25 days later. These derived figures are based on
reasoning given in the 1950 paper (p. 76-77). Most mosquitoes netted in man’s
vicinity will have a higher percentage of females than is normal for the popula¬
tion, and this may erroneously suggest a preponderance of female mosquitoes
carrying pollinia. It is probably significant, however, that neither Hocking,
Twinn, nor I have found pollinia on male individuals which were also present.
Male mosquitoes bear much larger antennae than females, and these structures
may constitute a mechanical barrier to effective penetration of the flowers by
males. Carbon dioxide, warmth, and increase in relative humidity all stimulate
female mosquitoes to bite but the effect of these changes on male mosquitoes
has not been reported. The orchid flowers may attract and/or stimulate only
female insects. If this is the case, it is unusual because male mosquitoes visit
other flowers, no sexual selection being operative.

If other mosquito attracting flowers are discovered, they may exhibit an
association of characters similar to the ones present in this orchid. Bright color
appears to be of little significance. Nectar will be produced or suggested and, if
present, will be within proboscis-reach of the insect. The flowers may be odor¬
less and they may be held on erect or spreading peduncles which are in the flight
path of mosquitoes. The flowers may be relatively small. Such plants will be
most likely to occur in mosquito-rich habitats and plants with these attributes
should be examined for possible mosquito adaptation.

ADDENDUM: Since this article went to press, Mrs. F. W. Case observed a population of the
Geometrid moth Xanthorhoe munitata Hubner (det. J. H. Newman) in the vicinity of
Habenaria obtusata plants in Big Creek Swamp, Crawford Co., Michigan. Two moths
captured on flowers of the orchid carried pollinia on their eyes in the same position as the
objects are attached to/ledes mosquitoes. An assessment of the role played by Geometrids
requires more observation and should be a sanguine endeavor.

LITERATURE CITED

Bates, M. 1949. The Natural History of Mosquitoes. Macmillan Co., New York. 379 pp.
Clements, A. N. 1963. Physiology of Mosquitoes. Pergamon Book, Macmillan Co., New
York. 393 pp.

Dexter, J. S. 1913. Mosquitoes pollinating orchids. Science 37: 867.

Haeger, J. S. 1955. The non-blood feeding habits of Aedes taeniorhynchus (Diptera,
Culicidae) on Sanibel Island, Florida. Mosquito News 15: 21-26
Hocking, B. 1953. The intrinsic range and speed of flight of insects. Trans. Roy. Ent. Soc.
Fond. 104: 223-345

- , W. R. Richards, & C. R. Twinn. 1950. Observations on the binonomics of some

northern mosquito species (Culicidae: Diptera). Canad. Jour. Res. (D) 28: 58-80.

212

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THE MICHIGAN BOTANIST

Jenkins, D. W. 1949. Toxorhynchites mosquitoes of the United States. Proc. Ent. Soc.
Wash. 51: 225-229.

- , & S. J. Carpenter. 1946. Ecology of the tree hole breeding mosquitoes of nearctic

North America. Ecol. Monogr. 16: 31-47.

Meeuse, B. J. D. 1966. The voodoo lily. Sci. Am. 215: 80-88.

Nielsen, E. T. & H. Greve. 1950. Studies on the swarming habits of mosquitoes and other
nematocera. Bull. Ent. Res. 41: 227-258.

Porsch, O. 1958. Alte Insektentypen als Blumenausbeuter. Osterr. Bot. Zeit. 104: 115-164.

Raup, H. M. 1930. The pollinization of Habenaria obtusata. Rhodora 32: 88-89.

Twinn, C. R., B. Hocking, W. C. McDuffie, & H. F. Cross. 1948. A preliminary account of
the biting flies at Churchill, Manitoba. Canad. Jour. Res. (D) 26: 334-357.

West, A. S., & D. W. Jenkins. 1951. Plant feeding habits of northern mosquitoes studied
with radioisotopes. Mosquito News 11: 217-219.

Wright, R. H. 1968. Tunes to which mosquitoes dance. New Scientist 37: 694-697.

STUDIES ON THE HYDNUMS OF MICHIGAN. I.
GENERA PHELLODON, BANKER A, HYDNELLUM

Kenneth A. Harrison
Research Station, Kentville, Nova Scotia

The University of Michigan Herbarium contains an outstanding collection
of the hydnums of North America. Its foundation was laid by C. H. Kauffman
while making his studies of the Agaricaceae of Michigan and since has been
greatly enlarged through the collections made by A. H. Smith. In the last two
years I have had the opportunity to study these collections. This is the first of
two papers based on them and gives keys and descriptions for the identification
of all genera and species found to date in the state with details on a few species
that can be expected when collecting is expanded into counties not explored as
yet. Hydnums are exacting in their growth requirements and many kinds fruit
only in restricted areas, or during favorable seasons after long intervals of time.
It takes years to build up the necessary collections for even a modest flora of
any region and this makes the yearly records from Michigan uniquely valuable. It
is hoped that this paper will stimulate interest in these rare fungi so that addi¬
tional favorable collecting areas can be located and a more complete under¬
standing of the requirements of the species obtained. It is of interest to point
out that the distributions given in these papers are the most complete for any
comparable area in North America but the more common species have been
recorded from only 10 of the 83 counties in the state. In the paragraphs on
material examined, the collections cited are from Michigan unless otherwise
stated.

1968

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213

KEY TO GENERA OF THE HYDNACEAE sensu lato

1. Sporophores resupinate only. (Genera not included in this key, but
check the following for resupinate forms of some pileate species.)

I. Sporophores stipitate, pileate or composed of branched tubercles with

long teeth hanging from the branches, some species with resupinate forms . 2

2. Lignicolous or on cones . 3

2. Terrestrial or on rotted woody material or duff . 7

3. Sporophores ungulate, perennial, woody, on conifers . Echinodontium

3. Sporophores not ungulate, annual . 4

4. Sporophores pileate-resupinate, spores non-amyloid . Steccherinum

4. Sporophores pileate or tuberculate-branched, spores amyloid . 5

5. Sporophores on cones usually of Pinus spp., stipe eccentric, hairy* . Auriscalpium

5. Sporophores on logs or living trees . 6

6. Sporophores branched and branches with spines or spines hanging

from a central fleshy mass of tissue, flesh amyloid in part at least . Hericium

6. Sporophores conchate, pileate or resupinate . Gloiodon

7. Sporophores small (-4 cm), context soft, cottony, concrescent,
hymenium on blunt spines or folds, spines not terete in cross section;

spores white in a deposit . Sistotrema

7. Sporophores usually over 4 cm, context fleshy fibrous, firm to fragile,

hymenium covering spines; spores brown or white in deposit . 8

8. Spores brown . 9

8. Spores white . 10

9. Context fleshy , firm to fragile, growth determinate . Hydnum

9. Context tough, fibrous, growth indeterminate . Hydnellum

10. Context tough, fibrous, growth indeterminate . Phellodon

10. Context fleshy, firm to fragile, growth determinate . 11

II. Spores smooth . Dentinum

11. Spores echinulate . Bankera

*Beenakia Reid, a somewhat similar genus from Australia and New Zealand, grows on tree
ferns and has brown non-amyloid spores.

PHELLODON Karst., Rev. Mycol. 3 (9): 19. 1881. Medd. Soc. Fauna FI. Fenn.

6: 15. 1881.

Calodon Quel, in Cke. &Quel., Clav. Hymen. 196. 1878, pro parte.

Basidiocarps terrestrial, arising from a tomentose pad, pileate and stipitate.
Hyphal system monomitic, hyphae colored or with walls containing dark pig¬
ment granules, septate; septa without clamp connections. Spines decurrent, sub¬
ulate, terete in cross section, context of parallel hyphae densely packed. Spores
subglobose, hyaline, echinulate.

Type species; Phellodon niger (Fr.) Karst.

Quelet and Cooke in 1878 proposed the genus Calodon, characterized by a
fibrous context and containing species with both white and brown spores, but
did not validate it with a description. Phellodon Karsten (1881) was proposed
for hydnums with white spores and spines regardless of consistency of the con¬
text. Later in 1881 Karsten accepted Calodon Quel, in place of his own genus
and validly published it. Phellodon as originally described was well defined, has
priority, and is generally accepted. The spores have ornamentation in the form
of very fine processes. In the North American species studied, these processes

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appear to be pointed but as there are 15 to 16 on the circumference of a spore 4
I u in diameter, it is almost impossible to verify this without the aid of an oil
immersion lens.

KEY TO SPECIES OF PHELLODON

1.

1.

3.

Pileus strongly zonate . 3. P. tomentosus

Pileus not strongly zonate . 2

2. Stipe slender, usually under 4 mm in diam . 3

2. Stipe stout, usually over 4 mm in diam . 4

Pileus brownish black, spines stain brownish on bruising, amyloid-like

inclusions occurring in surface hairs . 2. P. melaleucus

Pileus bluish black, spines stain blackish on bruising, no amyloid-like inclusions observed

in surface hairs (not found in Mich.) . P. atratus

4. Spines not over 2 mm long, pileus tomentose, tan, becoming gray
and finally brown, flesh not black and hyphae not dark in Melzer’s reagent
. 4. P. confluens

4. Spines up to 4mm long . 5

5. Pileus deep purplish black when young, black to brownish in age; flesh

and surface hyphae dark in Melzer’s reagent . 1. P. niger var. niger

5. Pileus whitish when young to dark brown in age; flesh dark but surface

hyphae not dark in Melzer’s reagent . 1. P. niger var. alkoniger

1. Phellodon niger (Fries) Karst., Rev. Mycol. 3: 19. 1881. var. niger.

Hydnum nigrum Fries, Syst. Myc. I; 404. 1821.

Hydnellum nigrum Karst., Medd. Soc. Fauna FI. Fenn. 5: 41. 1879.

Calodon niger Quel., Ench. Fung. 191. 1886.

Pileus: 5-7.5 cm broad, often concrescent, convex when very young,
becoming plane and finally depressed, often scrobiculate on discs especially
when pilei are fused; surface velvety to strigose, tomentose, or tomentum
matting in age, pallid gray, becoming violet-black to dark brownish gray in age at
the center, margin thick becoming thin, light gray. Context duplex, upper layer
soft and concolorous with surface, lower layer and context of stipe firm, black;
no odor; taste mild; when dried with the odor of fenugreek (sweetly fragrant).

Spines: 2.5-4 mm long, light gray, darker when bruised.

Stipe: Variable depending on depth of duff, 2-6 x 0. 5-3.0 cm, several
arising from a common pad of mycelium, grayish, with a thick coating of
mycelial felt that may surround several stipes.

Chemical reactions: Not tested fresh. KOH turns context blue-green and
the dark incrusting granules in the hyphae dissolve, the pigment leaching into the
solution. The granules are black in both water and Melzer’s reagent.

Spores and Hymenium: Spores 4.2-5 x 4. 2^1. 7 p, globose, subglobose,
echinulate, apiculate, white in mass. Basidia 30-35 x 5-5.3 /x, clavate, sterigmata
3.5 p long.

Tramal features: Surface layer of hyphae 3. 2-4. 5 ju, dark because of
numerous incrusting granules, loosely interwoven, inner layer of hyphae 3. 2-4. 5
/x, parallel, with a heavy coating of granules; hyphae of spines 2. 5-3. 2 p, parallel,
sparsely branched from below septa.

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215

Habit, Habitat, and Distribution: Solitary, gregarious, or cespitose and
often concrescent, in mixed woods of conifers and hardwoods - often found
along with var. alboniger. It is known in Michigan from Alger, Washtenaw, and
Emmet Counties. We know of one collection from Wyoming and one from
Manitoba. It is generally found eastward to the Atlantic seaboard in Canada and
the United States.

Material examined: Mains Sept. 1, 1932. Harding Aug. 1, 1949. Smith
Oct. 8, 1931, Oct. 10, 1934,33080, 38677.

Discussion: In the herbarium the surface hyphae of var. niger show
amyloid-like granules whereas these are lacking in similar hyphae of var. albo¬
niger. For further comment see var. alboniger.

Phellodon niger var. alboniger (Peck) K. Harrison, Stip. Hyd. N. S. 22. 1961.
Figs. 1,2.

Phellodon alboniger (Peck) Banker, Mem. Torrey Bot. Club 12: 167. 1906.

Hydnum albonigrum Peck, Rep. N.Y. State Mus. 50: 1 10. 1897.

Calodon alboniger (Peck) Seeler, Rhodora 44: 168. 1942.

Pileus: 3-9 cm broad, convex to plane or sometimes depressed, surface
tomentose to smooth, grayish white with the center gray, soon becoming dull
brownish; margin thick, blue gray, becoming thin and lighter, soft and taking a
fingerprint readily under pressure, darker when bruised. Context duplex, upper
layer pale dull brown or concolorous with surface layer, in the stipe firm and
black; odor slight at first, strong when drying; taste mild to slightly acrid.

Spines: 2.5^4 mm long, stout, light grayish, decurrent, sometimes in cool
weather arranged concentrically.

Stipe: 4-10 x 1-2 cm, light brown becoming dark brown, felted, bulbous,
lower part up to 3 cm thick from a water-absorbing layer of mycelium.

Chemical reactions: Not tested fresh; with Melzer’s reagent showing dark
incrusting granules coating the hyphae in the darker layers of the pileus and
stipe, some of the hyphae next to the dark layer show incrusting granules,
hyphae of the surface layer smooth; KOH turns flesh blue-green and the
incrusted granules dissolve producing a dark olivaceous color.

Spores and Hymenium: Spores white in mass, globose to subglobose,
mostly 4.5-5 n, a few 4.5-5 .3 x 4.5-5 ju, coarsely echinulate, prominently apicul-
ate. Basidia 25-30 ju, long, 5. 3-6.4 /r wide, clavate; sterigmata up to 4.2 \ u long,
no clamps seen at base.

Tramal features: Hyphae in surface layer 4. 2-5. 3 fi, loosely interwoven,
with slightly colored walls; hyphae of inner layer 3. 2-4. 2 ju, incrusted with dark
granules; hyphae of the spines 2. 1-3 .2 ju, some with heavy dark granules,
branched from immediately below the septa.

Habit, Habitat, and Distribution: Solitary to gregarious or finally concres¬
cent, in sandy pine woods, aspen-pine woods, or in mixed hardwoods, through¬
out the state. Collections have been examined from Emmet, Livingston,
Washtenaw, Oakland, Chippewa, Luce, Cheboygan, Jackson, Keweenaw, and
Presque Isle Counties. It occurs generally in eastern North America and is also
known from Europe.

216

THE MICHIGAN BOTANIST

Vol. 7

Material examined: Kauffman Sept. 2, 1905; Oct. 4, 1907. Mains 6584.
Harding 225, 277, 306. Povah 339. Thiers 4120. Shaffer 2789. A. H. Smith
32-315, 1128, 7200, 18494, 18528, 18730, 18744, 20629, 21151, 32650,
32744, 33054, 33083, 35900, 36181, 38077, 38416, 38619, 39319, 39690,
42369, 42407, 42538, 42636, 62846, 63981 , 63994, 64196A, 64197, 67742.

Discussion: The differences between var. niger and var. alboniger are more
quantitative than qualitative. The young basidiocarps of var. alboniger are lighter
in color than those of var. niger and when mature are dark brownish instead of
dark gray. Also in var. alboniger the basidiocarps are usually larger, the surface is
more irregular and is rarely depressed. A study in culture of these two varieties
would be highly desirable.

2. Phellodon melaleucus (Fries) Karst., Rev. Myc. 3: 19. 1881.

Hydnum melaleucum Fries, Syst. Myc. 1: 406. 1821.

Hydnum delicatum Schweinitz, Syn. N. Am. Fung. 161. 1834.

Phellodon ellisianus Banker, Mem. Torrey Bot. Club 12: 174. 1906.

Phellodon delicatus (Schw.) Banker, Mem. Torrey Bot. Club 12: 173. 1906

Phellodon brunneo-olivaceus Coker & Beers, Stip. Hyd. East. U. S. 28. 1951.

Phellodon brunneoroseus Snell, Dick, & Jackson, Lloydia 19: 171. 1956.

Pileus: 1-6 cm broad, or broader when in concrescent groups, plane or disc
depressed and rough, sometimes with pileoli, appressed fibrillose, appearing
zonate when moist but on drying azonate, color pale to dark violaceous brown
fading to very pale violaceous brown in age, margin light grayish drab. Context
thin, soft, fibrillose, zonate, nearly azonate when faded, cinnamon drab, darker
if wet; odor none; taste mild.

Spines: 1-1.5 mm long, irregularly decurrent on stipe, close, grayish drab
becoming dull drab, sordid brown when bruised.

Stipe: 3-6 x 0.2-0. 6 cm, compound and rooting, uneven and rough, dark
violaceous brown, cinnamon-drab (dull violaceous) above, dark vinaceous brown
in the base; pseudorhiza carbonaceous, very uneven and up to 10 cm long.

Chemical reactions: KOH on context dark olivaceous.

Spores and Hymenium: Spores 4-5 \ u, white in mass, globose to sub-
globose, finely echinulate, apiculate. Basidia 25-30 x 4. 5-5. 5 ju.

Tramal features: Hyphae (2) 3-4 (5) n broad, in Melzer’s reagent no
reaction in the spines or flesh, but in the surface hairs and in some of the hyphae
next to the hairs a slight apparent-amyloid reaction present.

Habit, Habitat, and Distribution: Gregarious to concrescent in large
masses, under conifers. I have seen two collections from Michigan, Smith 38618
and 43111. There are scattered records from the Carolinas to Canada but it is
most abundant in the north and particularly in the Pacific Northwest.

Discussion: This is a small species that has the strong fragrant odor of
most Phellodon species when dried. It grows in deep humus and recurs year after
year in the same spot. When wet and growing rapidly it is black with a white
margin. The long black pseudorhiza are not often seen as they are very brittle
and easily lost when the specimens are collected.

1968

THE MICHIGAN BOTANIST

217

The stipes are sometimes branched. Harvey (1958) describes the develop¬
ment of the branched stipes of Auriscalpium vulgare Fries and his observations
on the process in that species are probably the best explanation of what happens
during the growth of P. melaleuciis. He found that stipes may be retarded by
lack of moisture after they start to grow, then new branches may develop later
until finally one produces a pileus. Careful observations on sporophores
developing in a locality over a number of years to verify this supposition are
needed.

3 .Phellodon tomentosus (Fries) Banker. Mem. Torrey Bot. Club 12: 171. 1906.
Figs. 3,4.

Hydnum tomentosum Fries, Syst. Myc. I: 405. 1821.

Hydnum coriaceo-membranaceum Schw., Trans. Am. Philos. Soc. 2. 4:

162. 1834.

Phellodon cyathiformis (Schaeff. ex St.-Amans) Karst., Rev. Mycol. 3:19.

1881.

Calodon cyathiformis (Schaeff. ex St.-Amans) Quel., Ench. Fung. 191.

1886.

Hydnum graveolens subzonatum Peck, Bull. N. Y. State Mus. 75: 24. 1904.

Pileus: 1-6 cm broad, often pitted on disc, typically zonate, plane but
usually depressed to umbilicate at the center; surface radially fibrillose, streaked,
sepia to light brown, pallid to vinaceous buff on margin, when wet dark brown
in center; margin wavy and often elevated, thick, tomentose when actively
growing. Context 1-1.5 mm thick, pliant, leathery, more or less light brown;
odor faint to slightly fragrant of fenugreek as basidiocarps dry; taste mild or
slightly sweetish with a slight biting reaction in the throat.

Spines: Up to 2 mm long, decurrent to a line, crowded, pliant, white,
shaded with buff in age, staining vinaceous-buff when bruised.

Stipe: 2-5 x 0.2-0. 5 cm, often flattened, zonate, irregular in shape,
expanding into the pileus, arising from a felty layer of mycelium in the duff,
concolorous with pileus, unpolished. Context fibrous, zoned, some duff included
in base.

Chemical reactions: When fresh KOH blackens surface and flesh, the
margin darkens slightly; FeS04 grayish on disc then black, flesh grayish. Not
reacting when dried.

Spores and Hymenium: Spores white in mass, 3 A p, globose to sub-
globose, finely echinulate, apiculate; basidia 20-25 x 4-5.3 p, 4-spored, sterig-
mata 3.5-4 p.

Tramal features: Hyphae of pileus about 4 p, thin-walled, septate; in the
stipe up to 6.3 p and with thicker walls, branches rarely seen.

Habit, Habitat, and Distribution: Gregarious to cespitose and often
forming extensive concrescent patches or arcs under conifers. In Michigan it is
known from the Upper Peninsula, and from Emmet County below the Straits of
Mackinac. It occurs generally throughout conifer areas in northern regions
throughout North America and Euope.

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Vol. 7

Material examined: Smith 38617, 43655, 63984, 64017.

Discussion: This species has thin, strongly zonate basidiocarps rather
variable in many of the other characters. Two North American color variations
have been given names but since no correlated microscopic differences have been
found, they are both included here under the one species.

4. Phellodon confluens (Pers.) Pouz., Ceska Mykol. 10: 74. 1956.

Fig. 5.

Hydnum confluens Pers., Mycol. Europ. 2: 165. 1825.

Hydnum amicum Quel., Grevillea 8: 115. 1880.

Calodon amicus (Quel.) Quel., Compt. Rend. Assoc. Fr. Av. Sci. 12: 504.

1884.

Hydnum vellereum Peck, Rep. N. Y. State Mus. 50: 1 10. 1897.

Phellodon vellereus (Peck) Banker, Mem. Torrey Bot. Club 12: 168. 1906.

Phellodon amicus (Quel.) Banker, Mycologia 5: 62. 1913.

Pileus: 4-7 cm broad, concrescent in masses up to 11 cm wide, rarely
plane, usually depressed, but often complicate with ascending pileoli, very
irregular, surface conspicuously velvety, pallid to creamy buff to grayish or
grayish buff, becoming dull brown to dark brown where the pubescence wears
off of the older parts, margin even to lobed, sterile, white becoming fuscous to
dark brown where bruised. Context duplex, with a soft cottony upper layer
concolorous with surface, and a firmer dark lower layer with zones, dark brown
when bruised, taste mild to disagreeable, odor of rotting wood combined with
fenugreek (phellodon odor) when dried.

Spines: 2-3 mm long, decurrent, side view violaceous gray, tips pallid
violaceous gray, dark vinaceous brown in age.

Stipe: Up to 2.5 cm long, usually fused, central or eccentric, very
irregular, with a spongy layer around base, concolorous; context central core
firm, zonate, brown.

Chemical reactions: KOH darkens both layers; FeS04 both layers oliv¬
aceous; dried material inert in KOH.

Spores and Hymenium: Spores 4. 3-4. 7 (-5.3) x 3. 7-4. 2 (^E5) ju, white in
mass, subglobose, echinulate, strongly apiculate. Basidia 25-30 x 4. 2-6.4 /jl ,
4-spored, sterigmata slender, 3.5 ju long.

Tramal features: Hyphae of pileus without clamps, 3. 8-5. 7 /jl broad, thin
walled, in outer layer loosely interwoven, abruptly compacted inward, in stipe
the same as in pileus but tending to be more nearly parallel and with fewer
interspaces.

Habit, Habitat, and Distribution: Gregarious in deciduous or mixed
woods, frequently associated with aspen. It is rare, in part perhaps because it
matures and collapses much more quickly than the other species in this genus
and hence is not as frequently collected. It has been found abundantly in a few
areas however. In Michigan it is known from Cheboygan, Emmet, Gratiot,
Livingston, Oakland, and Washtenaw counties.

1968

THE MICHIGAN BOTANIST

219

Material examined: Potter 3901, 10013, 10036, 11111. Kauffman 1074,
and Aug. 15, 1915. Mains 5308, 6593, 8154, July 15, 1915. Smith 32-323,
1127, 6854, 6874, 6879,6949,7096,7157,7181,7287, 10971, 15207, 18464,
18481, 18504, 18544, 18551, 18742, 18750, 20555, 62816, 62832, 63898,
64108,64196,64238.

Discussion: This species has had a long history involving many names
indicated in the synonymy. It resembles P. niger var. alboniger to some extent
but can be distinguished at once by the lack of a dark inner layer of the context.

BANKERA Coker & Beers ex Pouzar, Ceska! Mykol. 9: 95. 1955.

Bankera Coker & Beers, Stip. Hyd. East. U. S. 33. 1951.

Sporophores terrestrial, pileate, mesopodus, context thick, fleshy, brittle
throughout, nearly white when fresh, not duplex or zonate in texture or color;
spore deposit pure white ; spores subglobose, minutely tuberculate orsubechinulate.
Type species: B. fuligineo-alba (Schmidt ex Fries) Pouz.

This genus, but without a Latin description, was published by Coker &
Beers (1951) for Hydnum fuligineo-album which is distinguished by its brittle
flesh and white subspiny spores. Pouzar validated it by publishing the Latin
description. The genus is small, with two species in North America, and bears the
same relationship to Phellodon in the echinulate white-spored group that
Hydnum does to Hydnellum in the brown-spored.

KEY TO SPECIES OF BANKERA

1. Pileus glabrous but surface layer soft and debris adhering to it conspicuously

. B. fuligineo-alba

1 . Pileus glabrous to scaly and surface not holding debris conspicuously .... B. carnosa

v

1. Bankera fuligineo-alba (Schmidt ex Fries) Pouz., Ceska Mykol. 9: 96. 1955.
Fig. 6.

Hydnum fuligineo-album Schmidt, in Kunze and Schmidt, Mykol., Hefte
I: 88. 1817.

Hydnum fuligineo-album Schmidt ex Fries, Syst. Myc., I: 400. 1821.
Hydnum fragile Fr., Sv. Vet-akad. Forh. 1851: 51. 1852.

Sarcodon fuligineo-albus Schmidt ex Fr.,Quel., Enchir. Fung. 189, 1886;
FI. Myc. 447. 1888.

Sarcodon reticulatus Banker, Mem. Torrey Bot. Club 12: 139-140. 1906.
Hydnum virginianum Murr., Bull. Torrey Bot. Club 67: 276. 1940.
Hydnum reticulatum (Banker) Lloyd, Letter 54, Note 224. 1915.

Pileus: 6-13 cm broad, convex, becoming plane to depressed or in age
umbilicate, surface unpolished, water-soaked in rainy weather, when dry with
adhering debris partly due to growth of surface hyphae binding the debris to the
surface, rarely diffracted scaly, disc dull dark brown progressing through light
brown to fawn and to vinaceous fawn to the margin, much darker when wet;
margin incurved, sterile when young, fertile in age, lobed and wavy. Context

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THE MICHIGAN BOTANIST

Vol. 7

brittle to slightly fibrous, tan to vinaceous buff, azonate, when fresh the odor
and taste not distinctive, when dried the odor distinctive (as in most species of
Phellodon).

Spines: 3-5 (-8) mm long, slightly decurrent, close, white becoming pallid,
pinkish on young specimens when dried.

Stipe: 2-4 x 1-3 cm, smooth, unpolished, concolorous with pileus, lightest
in color next to the spines. Context fibrous, pale vinaceous-buff, fawn or tan.

Chemical reactions: Inert to KOH and Melzer’s reagent when dried, not
tested when fresh.

Spores and Hymenium: Spore deposit white; spores 4-5 (-6) x 4-5 \ u,
subglobose, coarsely echinulate. Basidia about 40 x 5.5 ju; sterigmata 3-4 \ u,
close, no clamps seen.

Tramal features: Hyphae of context 4-6 /!, thin-walled, not clamped,
interwoven; hyphae of pileus cutis stain heavily in phloxine and are strongly
bibulous.

Habit, Habitat, and Distribution: Solitary to scattered or widely

gregarious, usually under pine. In Michigan it is known from Chippewa, Luce,
Marquette, and Midland Counties. It is known in addition from Oregon, Wash¬
ington, North Carolina, Tennessee, Virginia, New Jersey, and New York in the
United States and in Canada from Nova Scotia.

Material examined: Bartelli 252. Smith 35833, 38424, 38496, 67761,
67788.

Discussion: This is usually larger and stouter than B. carnosa and the disc
of the pileus tends to yellow-brown or vinaceous brown rather than the grayish
tones of B. carnosa. The pileus of the latter is more likely to become lacerate or
diffracted scaly. The surface of the pileus in B. fuligineo-alba has much debris
adhering to it and the cutis is very absorbent-taking up water instantly. B.
carnosa has neither of these features.

2. Bankera carnosa (Banker) Snell, Dick, & Taussig, Lloydia 19: 174. 1956.

Figs. 7,8.

Phellodon carnosus Banker, Mycologia 5: 65. 1913.

Hydnum carnosum (Banker) Trotter, Syll. Fung. 23: 472. 1925.

Pileus: 3-10 cm broad, depressed, plane or lobed and wavy; surface at first
smooth, unpolished, matted fibrillose, diffracted or lacerate scaly in age, scales
small and appressed especially on small fruit bodies; color grayish brown, dull
brown to dark brown in age, whitish to pale fawn on the margin. Context soft,
sissile, pallid to light brown, not zonate; odor faintly fragrant, strongly fragrant
when dried, taste mild.

Spines: 5-6 mm long, decurrent, fairly close, white to pallid, becoming
pinkish buff or slightly sordid.

Stipe: 3-6 x 1-2 cm, smooth becoming lacerate scaly, darker than pileus,
base darker or occasionally blackened in old specimens; context at apex as in the
pileus, harder downward, browner to dark umber at base in age.

1968

THE MICHIGAN BOTANIST

221

Chemical reactions: KOH turns dried context pale olivaceous, fresh
material not tested.

Spores and Hymenium: Spores 4-5.5 x 4-5 jit, white in mass, subglob ose,
coarsely echinulate, hyaline in KOH and Melzer’s reagent; basidia 35-40 x 4-4.5

Id.

Tramal features: Hyphae in pileus 4-6 /!, interwoven, not clamped,
phloxine did not differentiate a layer of bibulous tissue in the cuticle.

Habit, Habitat, and Distribution: Solitary, gregarious, to cespitose under
conifers, widely distributed but generally rare. In Michigan it is known mostly
from around Douglas Lake in Cheboygan County. In Canada it is common in
Nova Scotia and also known from New Brunswick, Quebec, and British Colum¬
bia. Kauffman found it in Oregon and the author found it in the Sangre de
Cristo Mountains of New Mexico.

Material examined: Bigelow Aug. 21, 1953. Povah Sept. 4, 1927. Smith
33170, 35946, 37298, 39373, 42514, 63012, 64032, 72165, 75033.

Discussion: The species under discussion here and considered identical
with Phellodon carnosus Banker has never been observed as pure white but only
as pallid to ‘'pale avellaneous.” The type collection of Banker’s at New York
Botanical Garden and Coker’s material at the University of North Carolina
appear identical with the dried young stages of this species from Nova Scotia,
where it has been possible to follow the development of the fruit-body from
small stipes to expanded pilei. The young stages are smooth but may become
fibrillose scaly if conditions are too dry and later become infundibuliform and
lacerate scaly toward the center, a condition most common in western collec¬
tions. Also as they get older they become dark and badly tunnelled by insect
larvae.

This species attains its maximum size in Nova Scotia, Quebec, and north¬
ern Michigan. There are no typical well-grown fruit-bodies of this species in the
herbaria of Coker or Banker. The occurrence of this species in Europe is still
considered doubtful although recently Maas Geesteranus placed it in synonymy
under Bankera mollis (P. Karst.) Maas G., largely because Banker stressed the
white color of B. carnosa when young and Fries speaks of the graying of the
pileus though his H. molle (pi. 2 upper fig. in Icon. Sel. Hym. 1: 4. 1867) is
expanded with a thin margin characteristic of fully grown hydnums. Fortunately
there is in the herbarium at Michigan a collection from Lundell: “Hydnum
violescens A. & S. ex Fr. among mosses in boulder strewn spruce woods. Upland:
Borge parish, ‘Klista skog’ (near Upsala) 8 IX 1941.” This consists of four pilei,
three very light colored depressed and lacerate at the center and one much
darker. All have been lightly pressed when dried but not too much to hide the
characters. The dark specimen is identical with mature specimens of B. carnosa
as known in this country. The light colored ones, however, are too light even for
very young material of B. carnosa. There can be little doubt that three of these
specimens are B. violescens. The color changes to green in KOH, a character
stressed by Maas Geesteranus (1958) and possessed by both. However, it is also
apparent that the darker member of this collection cannot be separated from B.
carnosa of America. The light color of the majority of the specimens and the

222

THE MICHIGAN BOTANIST

Vol. 7

Friesian illustration seem to make it necessary to maintain a white species as H.
molle, and we cannot agree with forcing the wood-brown American species into
synonymy with it on the evidence available. It would appear from the Lundell
collection that there definitely is a light colored form in Sweden and that at
times there is a darker form as well. At this stage they should both be main¬
tained as distinct taxa, at least until the situation as regards the distinctive
characters can be evaluated by more detailed studies on adequate fresh material.

HYDNELLUM Karst., Medd. Soc. Fauna FI. Fenn. 5: 41. 1879.

Calodon Karst, pro. p., Rev. Mycol. 3 (9): 20. Jan. 1, 1881.

Basidiocarps terrestrial, arising out of a mycelial pad, thin to stout,
solitary to gregarious or connate; regular to irregular in outline or compli¬
cate; context tough, fibrous, zonate, often duplex, upper layer soft, lower layer
more compact; spines brown at maturity; spores brown, angular, subglobose,
coarsely or finely tuberculate.

Type species: H. suaveolens (Scop, ex Fries) Karst, (selected by Banker).

The genus Calodon as used by Quelet and Karsten contained the hyaline
spored species treated in this paper under Phellodon and Bankera. Miller and
Boyle (1943) accepted Calodon but had very few species for study. Coker and
Beers (1951) were the first to recognize three genera. The white echinulate
spores of Phellodon and Bankera are easily recognized, and the odor is very
distinct in dried specimens in all members of both genera. It is harder to separate
the slightly fibrous context of some Hydnellum species from the fleshy context
of some Hydnum species. Hydnellum species usually have zonate flesh, a char¬
acter that has been seen in only one Hydnum. The zones are a record of periods
of high and low humidity usually associated with the higher humidity at night
and give a fairly accurate record of daily growth. (52 bands were counted in one
fruit body of Hydnellum suaveolens in 1964.) Hydnellums also have an indeter¬
minate habit of growth and are able to expand and grow around small sticks,
grass blades, ferns, and pine needles. There are also numerous fusions between
adjacent pilei so that mature fruit-bodies may have several stipes and be almost
any shape with quantities of foreign material imbedded in the flesh.

The following key is for the hydnellums found in Michigan, but three
which are extralimital are included as it is highly probable that these species,
commonly occurring in Canada, will be found along the shores of Lake Superior.
These are H. peckii, H. geogenium, and H. mirabile. They are easily recognized.
Several other rare eastern species with bluish spines and apparently restricted
ranges have been omitted although it is possible they will be found in the
northern coniferous forests of the Upper Peninsula.

KEY TO STIRPES OF HYDNELLUM

1. Basidiocarp with bluish zones in stipe or pileus context, or on surface of

pileus or on teeth at least when young . Stirps Caeruleum (p. 224)

1. Not with above colors . . 2

2. Pileus surface strigose; stipe with a watery yellowish juice . . . . Stirps Mirabileip. 242)
2. Pileus and stipe not as above . 3

1968

THE MICHIGAN BOTANIST

223

3. Pileus bright yellow when young; basal mycelium (and that in the

duff) yellow . Stirps Geogenium (p. 241)

3. Colors not as above . 4

4. Longitudinal section of basidiocarp and/or pileus surface showing orange
to orange brown, rusty orange, or pinkish-orange tones .... Stirps Aurantiacum(p. 226)

4. Lacking rusty orange to orange tints . 5

5. Context of fresh basidiocarps brittle to sub fleshy; never with red

droplets on pileus . Stirps Humidum\(y. 223)

5. Context coriaceous to semi-woody; red drops often present when fresh . 6

6. Stipe conspicuously felted by a thick soft layer of mycelium

. Stirps Spongiosipes (p. 232)

6. Stipe not with a conspicuous soft outer layer or zone . 7

7. Taste burning acrid and persistent in the throat; KOH not giving a

blackish reaction on mature tissue . Stirps Peckii (p- 235)

7. Taste mild to farinaceous; KOH blackening mature tissue . . Stirps Scrobiculatum (p. 235)

Stirps Humidum

1. Hydnellum humidum Banker in White, Bull. Torrey Bot. Club 29: 553. 1902.
Figs. 10-12.

Pileus: 3-12 cm broad, broadly infundibuliform to depressed, surface
pubescent becoming appressed fibrillose, zonate, matted fibrillose, with
“pinkish-buff’ zones more or less alternating with “tawny” to “russet” zones,
pileoli sometimes forming in the depressed centers; margin pubescent, obtuse
becoming thin, discoloring to dull brown where handled. Context fleshy, soft,
thin, not duplex, becoming fibrous, zonate with dark brown lines and with areas
between near “wood-brown,” unchanging when cut; odor of freshly cut flesh
faintly pungent, taste becoming acrid and disagreeable.

Spines: Up to 3 mm long, fine, close, decurrent, “Verona-brown” to
“warm sepia” with the lighter tips giving a pallid sheen, frequently fused at the
bases to form ridges or a network.

Stipe: 1^1 x 1-2 cm, solid, very irregular, frequently fused, unpolished and
dull vinaceous-rusty brown where free of decurrent teeth, often enlarged; con¬
text zoned, darker than in the pileus, semi-woody.

Chemical reactions: Hyphae of all parts of the fruiting body reviving
poorly in KOH or Melzer’s; no color reactions in dried material from either.
When fresh, the surface olive-black in KOH, olive grayish in FeS04.

Spores and Hymenium: Spores 4. 5-5. 5 (-6.5) x 4-4.5 (-5.5) p, in mass
wood-brown, subglobose, finely tuberculate with 8 or 10 processes on the cir¬
cumference; basidia could not be revived and sterigmata were not seen.

Tramal characters: Hyphae of the teeth 3-4 p, flexuous, parallel, occasion¬
ally branched; hyphae of the pileus mostly 5-7 p with a few larger cells, Intri¬
cately' interwoven. No clamps were seen and branching was from below the
septa.

Habit, Habitat, and Distribution: Gregarious in low oak woods in Mich¬
igan and the eastern seaboard of North America.

Material examined: Smith 6853, 7097, 7197, 7549, 18751, 64248,
64404,64368.

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THE MICHIGAN BOTANIST

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Discussion: The appearance of these collections fits Coker and Beers’
(1951) description closely but they vary in having a strong odor and acrid taste.
Banker’s original description calls for a tawny to light yellowish brown, scro-
biculate, minutely pubescent species, but Coker felt certain his specimens were
the same as Banker’s. In the herbarium this species is easily confused with
Hydnum stereosarcinon, but the latter has small oblong, nodulose spores which
are distinct from the subglobose finely tuberculate ones of H. humidum.

This is a rare species but in 1961 Dr. A. H. Smith made three collections in
Michigan and obtained detailed field notes on the fresh material. From these
collections it is evident that H. humidum is intermediate between Hydnum and
Hydnellum in all characters except those of determinate and indeterminate
growth. H. humidum has flexuous hyphae forming the thick edge of the pileus
margin and growth will proceed indefinitely when conditions are favorable. For
this reason it has been decided that H. humidum should be left in Hydnellum.
The flesh of the young plants is softer than in H. stereosarcinon but the system
of zonations is often similar. However, the tramal cells are thin-walled, inter¬
woven, and somewhat inflated so that the flesh is similar to that of some species
of Hydnum.

Stirps Caeruleum- Key to Species

1. Odor of fresh specimens sickening-sweetish; stipe context and surface

typically violaceous but surface fades to pallid . 2. H. suaveolens

1. Not as above . 2

2. Freshly growing areas of pileus blackish when rubbed . (4. H. ferrugipes )

2. Freshly growing areas of pileus and of teeth typically bluish and not

turning blackish when rubbed . 3. H. caeruleum

2. Hydnellum suaveolens (Scop, ex Fries) Karst. Medd. Soc. Fauna FI. Fenn.
5:41. 1879.

Hydnum suaveolens Scopoli, FI. Carn. 2: 472. 1772.

Hydnum suaveolens Fries, Syst. Myc. I. 402. 1821.

Hydnum boreale Banker in White, Bull. Torrey Bot. Club 29: 553. 1902.

Hydnellum rickeri Banker, Mycologia 5: 201. 1913.

Pileus: 4-15 cm, convex to plane, surface tomentose, soft, irregular,
uneven, often pestle-like when very young (may develop pileoli with spines at
the center), sordid white changing slowly to snuff-brown at center and gradually
outwards to the white margin, occasionally thin and lobed; margin with broad
sterile band. Context fibrous, thin to thick, zoned with violet — brightest in
stipe; odor strongly fragrant; taste mild or slightly cinnamon-like.

Spines: Up to 5 mm long, crowded, fine, “avellaneous” and becoming
“wood brown” with “army brown” base (pale to dark vinaceous brown).

Stipe: 3-5 x 1-2.5 cm, woody, interior zoned with “dark madder blue”
bands, surface tomentose with a bright violet shade that darkens on being
rubbed.

Chemical reactions: KOH stained the violet tomentum on the stipe a
blue-green and a trace of the same shade appeared when the colored flesh was
tested. Melzer’s reagent has no effect.

1968

THE MICHIGAN BOTANIST

225

Spores and Hymenium: Spores 4. 5-6. 5 x 3-4 ju, vinaceous brown in
deposit, oblong, angular, irregular and unevenly shaped, coarsely tuberculate.
Basidia 16-20 x 5-6 p, clavate, 4-spored.

Tramal characters: Hyphae of spongiose layer 4-5 ju in diameter, clamped,
thin to about 2.5 ju thick, complexly interwoven, ends of hyphae forming a loose
trichoderm (tomentum); hyphae of the more compact inner layer up to 5-7 ju
wide, but often very thin, collapsing but reviving well in water followed by 3%
KOH; in water strands of dark blue hyphae evident, these hyphae turning
greenish in KOH, and then slowly fading; in Melzer’s reagent these hyphae are
seen to be incrusted with very fine granular material that gives them an
“apparent-amyloid” appearance, and in section the granules are coarse as seen
under oil immersion lens; hyphae thin-walled and with frequent clamps at the
septa, often branched near the septa. Hyphae in teeth 3-4 n in diameter, parallel,
clamped and with fine incrusting “apparent-amyloid” granules.

Habit, Habitat, and Distribution: Scattered, gregarious, occasionally in
arcs, under conifers; primarily a northern and montane species. Found from New
Mexico to British Columbia in the west and along the Applachians from North
Carolina into Canada in the East. Reported as common in the Upper Peninsula
but no collections have been deposited in the herbarium.

Discussion: This species is normally thick and stout but in regions of high
rainfall and humidity it is thin, expanded and very rough with some of the blue
of the flesh showing through on the disc. It also may develop projections that
develop as pileoli with spines bearing a hymenium and spores. The youngest
stage of this species is a small cushion of violet colored mycelium that develops a
white colored apex which enlarges and becomes the stipe and fruit body. This is
in contrast to H. caeruleum, whose primary cushion is straw-colored and whose
developing apex is bluish.

3. Hydnellum caeruleum (Hornem. ex Pers.) Karst., Medd. Soc. Fauna FI. Fenn.
5:41. 1879.

Figs. 9 & 13.

Hydnum caeruleum Hornemann, Flora Danica VIII, fasc. 23: pi. 1374.1 808.

Hydnum suaveolens Scop. var. caeruleum Fries, Syst. Myc. I: 403. 1821.

Hydnum caeruleum Pers., Myc. Europ. 162. 1825.

Hydnum cyaneotinctum Peck, Bull. Torrey Bot. Club 30: 98. 1903.

Hydnellum cyaneotinctum (Peck) Banker, Mem. Torrey Bot. Club 12:

164. 1906.

Pileus: 3-11 cm broad, convex, plane, sometimes depressed, surface
smooth, or irregular to colliculose, cottony tomentose, finally matted and
pitted; color light blue when young, becoming almost white, gradually darkening
in age to dark brown, staining ferruginous when bruised. Context duplex; upper
layer spongiose; lower layer tough, fibrous, compact, buff, zonate with bands of
mauve and brown, shading to a reddish brown to ferruginous in the lower part of
the stipe; taste and odor not distinctive.

Spines: 3-5 mm long, decurrent, close, fine, those on the margin whitish
with a shade of blue, finally dark brown with lighter tips.

226

THE MICHIGAN BOTANIST

Vol. 7

Stipe: 2 -4 x 1-2 cm, base bulbous from felty mycelium, buff colored;
context duplex, outer layer thin, inner layer orange or reddish, lighter parts
frequently with bands of blue, tough and fibrous; the thick felty layer around
the base often disappearing from being eaten by slugs.

Chemical reactions: The blue flesh turns blue-green in KOH and the red¬
dish parts a dull “dark olivaceous,” a greenish shade shows in the teeth both
young and old, and a dark olivaceous color diffuses out of teeth and flesh into
the liquid. In Melzer’s reagent reddish incrusting granules are seen adhering to
the hyphae, and “apparent-amyloid” hyphae and particles have been seen.

Spores and Hymenium: 4.5-6 (7) x 4.5-5 p, colored, subglobose, coarsely
tuberculate, vinaceous brown in deposit. Basidia 6-7 ju wide with long sterigmata
(up to 4.5 ju), 4-spored.

Tramal characters: Upper layer of spongiose tissue with hyphae 3-5 p in
diameter, slightly incrusted in places, frequently branched and loosely inter¬
woven, only one clamp seen after lengthy examination, occasional short section
of a hypha showing blackish incrustations in both water and Melzer’s reagent.
Lower layer of compact hyphae 3^1.5 p wide, tending to be parallel, tubular,
rarely with clamps at the septa, heavily incrusted with reddish brown granules
and quantities of fine granular debris present, some sections of hyphae show
blackish granules in both water and Melzer’s; some granules dissolve in KOH and
the hyphae appear greenish. The hyphae in the teeth appear amyloid in Melzer’s
but when crushed out to a thin layer the blackish color of the walls is diluted
and the blackish granules on the walls give the “apparent-amyloid” effect.

Habit, Habitat, and Distribution: Gregarious to concrescent under
conifers, often under Pinus spp., late summer and fall. It is relatively common in
the Upper Peninsula, and in northern conifer areas of North America and
Europe.

Material examined: Bartelli 233. Imshaug 3452. Mains 6527. Povah 1240.
Thiers 4328. Shaffer 2557. Smith 1129. 32895, 33086, 33087, 33088, 38528,
38664, 39368, 42637, 42639, 58017A, 62853, 63985, 64027, 72004, 72319,
73311,74517,74851.

Discussion: The fruit body develops from an ovoid buff colored cushion
of mycelium that becomes the bulbous base of the stipe. At first the stipe is
apparent as a bluish outgrowth that elongates for 1 or 2 cm and then starts to
expand to a convex pileus. It is very compact and solid at this stage. As it
matures it is a squat species that shows bluish shades only on the parts actively
growing. The color is brighter in cool weather. The species is recognized by the
thick, bright colored, blue-zoned flesh of the pileus and the reddish brown color
of the stipe. Old specimens become quite brown, others may be brown at the
center shading to a blue or pallid margin. This is the result of secondary growth.
Fruit-bodies persist for a month or more. Aged specimens are easily confused
with H. pineticola K. Harrison but can be separated by examining the context.

Stirps Aurantiacum — Key to Species

1. Pileus thin and infundibuliform at maturity, not distinctly
zonate .

8. H. conigenum

1968

THE MICHIGAN BOTANIST

227

1. Pileus convex to shallowly depressed; context relatively thick . 2

2. Spines with sulphur yellow tips when young; pileus surface more or

less smooth but often complicate or with only a few pileoli . 7. H. earlianum

2. Not as above . 3

3. Orange tones muted and showing mostly as pale orange-yellow on

margin but this changing to bister or fuscous when growth has ceased . . 5. H. septentrionale

3. Orange to orange-brown tones obvious . 4

4. Context of dried specimens showing dingy slate-bluish zones,

pileus soon dull rusty ferruginous . 4 . H. ferrugipes

4. Pileus whitish on fresh growing surface, becoming orange; context

never with slate-blue zones . 6. H. aurantiacum

4. Hydnellum ferrugipes Coker, Jour. Elisha Mitchell Sci. Soc. 34: 188. 1919.
Fig. 14.

Calodon ferrugipes (Coker) Snell, Mycologia 37: 48. 1945.

Pileus: 5-15 cm broad, plane, disc slightly depressed, colliculose, surface
dry and velvety; color “light ochraceous-buff ’with the margin blue, buff area
staining dingy yellow-brown when injured and finally blackish, turning brown
then blackish in age from the disc outward. Context duplex; inner layer semi-
woody, zoned with blue to pale buff bands; outer layer thin, buff and rather
spongy; with a pungent farinaceous odor when cut and taste pungent-
disagreeable.

Spines: Up to 6 mm long, crowded, sharp, deep vinaceous brown in age.

Stipe: 3-6 x 1-1.5 (2.5) cm, woody, fibrous, the core zoned as in the
pileus and outer layer a soft coat of felted mycelium, orange tawny, slowly
turning dark brown when handled.

Chemical reactions: In fresh material the blue zones of flesh and teeth
turn green in KOH, no reaction to FeS04; when sections are placed in KOH the
blue zones and teeth turn olivaceous green with some muddy brown pigment
leaching out.

Spores and Hymenium: Spores in deposit wood-brown, 5.5-7 x 5. 5-6. 5 p,
semiglobose, tuberculate with 8 or 9 low processes. Basidia 6-7 x 20 p,
sterigmata fine up to 4 p in length. A number of abnormalities were seen in the
hymenium. (It appeared that some of the first basidia had been checked before
normal sterigmata were formed and the ends became filled with a dense globule
of granular material resembling a large spore, resulting in a capitate ‘cystidium’.)

Tramal characters: There seemed to be three layers to the context, a
rather thin upper interwoven spongiose layer zoned with fine dark bands, then a
layer of parallel hyphae that tended to form dark bands and next a layer of very
hard tissue often bluish. The parallel hyphae were in strands, and were 4-6 p in
width, tough, but easily teased apart. Clamps were occasional on the parallel
hyphae. Hyphae are somewhat incrusted and areas that were blue had hyphae
that were “Apparent-amyloid” with “apparent-amyloid” globules free in the
context. Orange incrustations were seen in the tramal hyphae of the teeth but no
clamps were seen there.

228

THE MICHIGAN BOTANIST

Vol. 7

Habit, Habitat, and Distribution: Occurring in deciduous or mixed woods.
Most frequently found in North Carolina. In Michigan one collection from
Cassidy Lake Road near Ann Arbor, and one from the George Reserve, Living¬
ston County.

Michigan material examined: Smith 64102 and 64190.

Discussion: H. fermgipes has been confused with H. aurantiacum in North
America. It is a distinct species as based on the type and original description, but
by 1951 when Coker and Beers published their final description of Hydnellum
species their concept of this one had changed. The original concept had been
altered and the very blue-fleshed collections were identified as H. cyaneotinctum
Peck. Coker also was confused about H. aurantiacum, for he identified a typical
collection of H. aurantiacum from Bar Harbor, Maine as H. ferrugipes for Dr. W.
H. Snell and this was passed on through identifications by Snell to Canadian
mycologists. Thus records of H. ferrugipes in the northern part of the country
prior to 1957 are questionable. Coker and Beers (1951) compared H. ferrugipes
with H. aurantiacum but their description of H. aurantiacum is actually of H.
earlianum Banker. H. aurantiacum and H. caeruleum (=H. cyaneotinctum Peck)
were unknown to Coker in the fresh condition.

5. Hydnellum septentrionale K. Harrison, Canad. Jour. Bot. 42: 1224. 1964.

Fig. 15

Pileus: 4-15 cm broad, convex, soon dividing into sections that develop
and become processes and pileoli, often forming rosettes; margin elevated, thick,
tomentose, spreading and thinner in age and then striate; surface deeply
scrobiculate, sometimes radiately ridged, yellowish pallid to “pale orange-
yellow” on actively growing parts, becoming “cinnamon-drab” to “fuscous” and
finally “blackish-brown” or “bistre” at center; each process and pileolus may
show a range of color so that the color pattern appears very complex. Context
soft, becoming fibrous and hard in age, woody in the stipe, zonate from numer¬
ous dark lines, “clay -color” or near “pinkish-buff ’; odor and taste varying from
faint to strong and disagreeable.

Spines: Up to 5 mm long, decurrent, fine, close, sometimes fused, pale
yellowish, becoming “cinnamon-drab,” almost black in age, tips lighter with a
tint of yellow.

Stipe: 3-5 x 0.6-2 cm, short, stout, subbulbous, tapering to a small root¬
like tip, “salmon buff’ to “apricot buff’ becoming “blackish brown,” with
salmon-colored my celia in duff around base; context duplex: outer layer thin
and spongy, felted; inner layer hard, sordid “clay-color.”

Spores and Basidia: 4.5-5 .5 x 3. 5 A. 5 p, colored, oblong to subglobose,
with low tubercles, appearing to be slightly amyloid in Melzer’s reagent (but
after many tests it was decided that the color was not intensified by the solu¬
tion). Basidia 30 x 6-7 p; sterigmata 3 A p long.

Tramal characters: Hyphae uniform throughout, 4-5 p wide, flexuous,
interwoven and the septa far apart and without clamps, 3-4 p diameter in the
radicating stipe, flexuous, closely interwoven and the outer walls coated with
fine, faintly colored granules.

Chemical reactions: No reaction in KOH or Melzer’s solutions.

1968

THE MICHIGAN BOTANIST

229

Habit, Habitat, and Distribution: Gregarious, concrescent, frequently in
rosettes, under fir and spruce, near the shore of the Straits of Mackinac.

Material examined: Smith 33082, 33121 , 34000A, 361 10, 37138, 38371 ,
38569, 39379, 41971 , 42394, 60795.

Discussion: This is another of the American species in stirps Aurantiacum.
It is not bright colored and is more solid and woody than most of the group.
When old, it resembles//, frondosum K. Harrison, but when young the yellowish
color distinguishes it. The spores are larger.

Collection Smith 34000A, from the Mackinaw City hardwoods, September
1949, consists of a large number of carpophores. They are stout, turbinate, and
the spines are very close. On some of these there are concentric bands of poroid
tissue with normal spines near the stipe and on the pileus margins.

6. Hydnellum aurantiacum (Batsch ex Fries) Karst., Medd. Soc. Fauna FI. Fenn

5: 41. 1879.

Fig. 16.

Hydnum aurantiacum Fries, Syst. Myc. 1 : 403. 1821.

Hydnum floriforme Schaeff., Fung. Bav. 4: 97. 1774, pro parte

Pileus: 6-18 cm broad, convex, plane or depressed, often colliculose,
sometimes quite smooth and then breaking into processes with teeth forming
large rosettes, often extremely rough with projections, surface uneven, tomen-
tose, felted and becoming matted in age, sometimes zonate, margin “orange-
ferruginous,” center dark rusty brown to “bay”; margin whitish to tan in
some northern collections. Context fibrous, tough, zonate, rusty orange in
stipes and buff in pileus; odor and taste pungent, disagreeable.

Spines: 5-7 mm long, decurrent, close, coarse, dark brown with grayish
buff tips. Pileus margin sterile or with tiny white undeveloped spines.

Stipe: 2-7 x 0.5-2 cm, base bulbous, single or several growing from a
cushion of mycelium; context woody at center, orange-red, zonate; surface
matted with orange to brownish tomentum, orange mycelium incorporates the
duff in the bulbous base.

Chemical reactions: Further tests are needed on this and related species.
The surface turns black in KOH and the flesh darkens to a dirty olive.

Spores and Hymenium: Spores 5. 5-7. 5 x 5-6 ju, appearing dark as though
slightly amyloid, subglobose, roughly tuberculate. Basidia about 30 x 6 g,
clavate, 4-spored.

Tramal characters: The outer spongiose layer of rather closely interwoven
hyphae _+ 3 ju wide, branched, septate, clamps not seen; the harder inner layer
with hyphae 3-3.5 ju wide, parallel, compact, tubular, septate, sparsely branched,
no clamps observed. The hyphae in the spines are 2.5-3 \ u wide, septate, and
lacking clamps. Some granules present in the context of the pileus and spines but
most were free of the hyphal walls.

Habit, Habitat, and Distribution: Solitary to gregarious or concrescent,
under conifers in Europe and North America generally.

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THE MICHIGAN BOTANIST

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Material examined: Bartelli Oct. 3, 1965. Mains 6529. Imshaug Sept. 10,
1949. Harding 296. Smith Aug. 13, 1940, Aug. 1949, 42387, 42398, 72005,
72785.

Discussion: This species is one of the striking species in the genus and has
been frequently renamed because of the extreme variability that it displays in
different regions. Usually each region has rather distinct forms that are con¬
sistent within that area and are more or less distinct from those of other regions.
We have limited our species to the group with large spores, slightly darkened in
Melzer’s reagent. The spores are wide (5-6 p) so that the subglobose spores are
easily recognized as large. However, even with this restriction a multiplicity of
forms and variations are encountered. The colors range from dull brown to
brilliant orange; the texture and spongy layer on the surface grades from soft
and thick to firm and thin. The large spores separate this species from H.
earlianum Banker and H. conigenum (Peck) Banker. The brighter colors readily
separate it from H. septentrionale K. Harrison. Any specimens with slaty shades
in the flesh should be compared with H. ferrugipes. One collection from Baraga
County has spores up to 8.5 x 6.3 p.

7. Hydnellum earlianum Banker, Mem. Torrey Bot. Club 12: 161. 1906
Fig. 17.

Hydnum earlianum (Banker) Sacc. et Trott., Syll Fung. 21 : 271. 1912.

Pileus: 3-10 cm, in rosettes up to 15 cm with numerous overlapping pilei,
single pileus depressed but margin lobed and sometimes faintly zoned, usually
somewhat lighter when actively growing, surface smooth, occasionally irregular,
tomentose or as though dusted with orange powder, bright color well retained in
dried specimens, color becoming darker in center ranging from deep ferruginous
to cinnamon-rufous. Context duplex, with a soft upper layer and a somewhat
harder lower layer, soft and brittle, bright colored, usually a slightly lighter tint
than the cap. Coker & Beers (1951) report a faint fragrant odor when dry.

Spines: Up to 3 mm long, decurrent, fine, close, color toward the margin a
peculiar sulphur shade that persists for a long time, eventually becoming dark
brownish shading from the stipe outward, tips lighter, sometimes yellowish.

Stipe: 2-5 x 1-2 cm, expanding into a bulbous base, compound or several
starting from one cushion of mycelium, tapering upwards, expanded near spines;
base 2-8 cm, globular, a mixture of orange-colored mycelium and duff; duplex,
the outer covering smooth, concolorous or brighter than pileus, rusty brown in
age.

Chemical reactions: KOH stains the bright colored tissue a very dull dark
“olivaceous.”

Spores and Hymenium: 4.5-5 .5 x 4^4.5 (many 4.5 x 4) p; oblong- to
subglobose-tuberculate, tubercles rather short, dark under the microscope as
though faintly amyloid. Basidia about 20 x 6 p, clavate, 4-spored; a few reddish
granules present in sections in the subhymenial region.

Tramal characters: Hyphae of the outer (spongiose) layer 3-3.5 p wide,
heavily incrusted with reddish brown granules; in the inner (harder) layer the

1968

231

THE MICHIGAN BOTANIST

hyphae 3. 5-4.2 ju wide, parallel, septate, heavily incrusted with granules and
many granules free in the context of both layers. Hyphae in teeth about 3 /i in
diameter, incrusted with reddish granules. Clamp connections absent.

Habit, Habitat, and Distribution: Gregarious becoming concrescent, in
open oak woods, common in southern Michigan during wet seasons. It occurs
widely scattered in the eastern United States and in Ontario.

Material examined: Kauffman 1069, 1072. Fisher Aug. 27, 1905. Mrs.
Cohn Aug. 5, 1927. Mains 6528. Povah 342. Shaffer 2502. Thiers 3679. Smith
1591, 1723, 6851, 7055, 7184, 18458, 18731, 38377, 62599A, 62851, 63416,
63417, 64126, 64223, 64358, 64845, 72005, 72785.

Discussion: This species is outstanding because of the brightness of the
orange color of the pileus and the sulphur yellow spines when they are actively
growing. The Michigan collections show quite a range in color and toughness but
until single basidiocarps have been watched throughout the growing season it
will be unwise to attempt any further separation of minor taxa. This species
typically occurs under oak whereas H. aurantiacum is usually associated with
conifers.

8. Hydnellum conigenum (Peck) Banker, Mem. Torrey Bot. Club 12: 160. 1906.
Fig. 18.

Hydnum conigenum Peck, Bull. Torrey Bot. Club 30: 97. 1903.

Pileus: 3-7 cm broad, depressed to deeply infundibuliform, complicate,
lobed, flabelliform, concrescent forming rosettes 8 to 10 cm across; surface
irregular, radially ridged, fibrillose, tomentose, striate, or ridged, faintly to
strongly zoned more or less “pinkish cinnamon” to “cinnamon-buff,” “apricot-
buff’ to “mahogany red” (one collection tinted “orange-cinnamon”), darker in
age; margin wavy and lobed. Context thin, tough, pliant “pinkish cinnamon”;
taste strongly farinaceous; odor none to farinaceous.

Spines: Up to 3 mm long, deeply decurrent, close, fine, concolorous with
cap (or a brighter orange).

Stipe: 3-6 x 0. 5-2.0 cm, simple or compound, bulbous at the base, more
or less orange-brown, with a mass of felted tomentum at the base.

Chemical reactions: In fresh condition KOH turns flesh drab and in Mel-
zer’s reagent and FeS04 it becomes dull and water-soaked in appearance. Tramal
tissue after drying turns dull olivaceous in KOH and remains orange in Melzer’s.

Spores: 4-5.5 x 3. 5-4. 5 p subglobose to oblong, angular, with young
spores showing a prominent mucro, dark as though slightly amyloid.

Tramal characters: Hyphae heavily incrusted, _+ 3 ju wide in the teeth and
up to 6 ii in the pileus, frequently swollen at the septa, no clamps seen; numer¬
ous hyphae (laticifers?) seen in the hymenial layer with heavy concentrations of
reddish granules; granules not darkening in Melzer’s.

Habit, Habitat, and Distribution: Gregarious or concrescent in coniferous
forests. Common in the Pacific Northwest. Collections seen from New Mexico to
British Columbia; type locality Idaho; one collection from Isle Royale, Michigan.

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Vol. 7

Discussion: This is a very distinct species, especially bright colored at first
and in age with a dark, very rough radially ridged surface. It is almost always
concrescent and frequently complicate. It is in this condition that it incorporates
cones into the basidiocarp and two of the eight collections examined contained
cones embedded in the base of the compound stipe.

This species is separated from H. aurantiacum by its distinctly narrower
spores; from H. septentrionale by its thin, brightly colored flesh; and from H.
earlianum by the thinner non-duplex flesh.

H. conigenum, from the description, resembles the specimens in Europe
that have been identified by Maas Geesteranus as H. auratile Britzelmeyer. How¬
ever we are retaining the American name for several reasons. First the type of H.
auratile has not been found and in our experience it is unwise to identify any
species in the Hydnellum aurantiacum complex without specimens; second the
western collections are much larger than the Euopean specimens identified as H.
auratile. Also the original description called for “margine albidum” which does
not apply to H. conigenum or the collections identified by Maas Geesteranus.
The long-persistent orange shade of the margin is diagnostic. Many of the
western collections are strongly zonate in age. The Michigan collection is young,
only slightly zonate, and the surface is radially ridged, but otherwise it agrees
well with H. conigenum.

Stirps Spongiosipes
v

9. Hydnellum spongiosipes (Peck) Pouz., Ceska Mykol. 14:130. 1960
Fig. 19.

Hydnum spongiosipes Peck, Rep. N. Y. St. Mus. 50: 111. 1897.

Hydnellum velutinum (Fries) Karst, var. spongiosipes (Peck) Maas G.,

Fungus 27: 62. 1957.

Pileus: 2-10 cm broad, usually convex to irregularly plane, occasionally
depressed, azonate or rarely with a concentric ridge of secondary growth on the
margin, at times misshapen through fusing; surface finely tomentose, uneven or
slightly colliculose, rarely sub striate towards margin if margin thin, darkening
when bruised when young and margin drying black where bruised; color cinna¬
mon-brown, walnut to Vandyke-brown and with a grayish-brown bloom. Con¬
text duplex, upper layer thick, spongy; lower layer dark and rather thin,
connecting with the hard core of the stipe, cinnamon-brown, compact layer in
stipe darker; odor and taste mild.

Spines; Up to 6 mm long, decurrent, close, slender, dark to light brown
with tips slightly lighter when young, darkening when bruised and injuries drying
blackish.

Stipe: 3-10 x 0.5-2 cm, appearing short, narrowest next the spines,
expanding quickly to a very broad bulbous base 4-5 cm wide, wider when several
arise from a tomentose pad of mycelium; context duplex, central core hard,
zonate, dark brown, surrounded by a thick felty layer of spongiose tissue.

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233

Chemical reactions: Sections of dried material in KOH give a violet flash
and then go black, and a dark olivaceous-brown color leaches into the mounting
medium.

Spores and Hymenium: Spore deposit cocoa-brown; spores 5.5-7 x 5-6 p,
many 6.5 x 5.5 /}, walls thick, apiculus prominent, shape subglobose, moderately
to coarsely tuberculate, (7 or 8 tubercles around circumference), tubercles stud¬
like to angular with truncated ends. Basidia 6-7 x 20-25 p, sterigmatajt 4 p long.

Tramal characters: Tramal hyphae of teeth 3-4 p wide, tubular, parallel,
septate but not clamped. In Melzer’s reagent the hyphae 3. 5-6.0 \ u, thin-walled,
end cells at surfaces with russet colored contents; hyphae of interwoven
spongiose layer changing abruptly to the radial tissue of the compact lower
layer, 4. 5-5. 5 j u wide, small pockets of amorphous russet colored material occur¬
ring in the layer, occasional laticiferous hyphae present and filled with russet
contents but these could not be traced to the pockets. The hard radial tissue
supports the teeth and is continuous with the hard tissue in the stipe.

Habit, Habitat, and Distribution: Solitary, gregarious, or connate; on
rather dry hillsides as well as on low ground in deciduous woods, usually oaks
predominate. Common in the deciduous woods of southern Michigan where oaks
are found. We have no collections from the Upper Peninsula. Widely reported
from Iowa east to the Atlantic seaboard. Reports from other areas under pines
should be checked against H. pineticola.

Material examined: Mains July 7, 1915, Aug 7, 1915, 6531, 8153. Shaffer
2750, 2788, 2818. Kauffman Aug 1, 1915; Rea 971. Potter 3805, 11135,
11136, 12764. Smith 32308, 1724, 6784, 6891, 6904, 7070, 6138, 7294,
18529, 18743, 18768, 20871, 62815, 62837, 62852, 63078, 63889, 64118,
64163, 64198, 64250, 64360, 64806, 73288.

Discussion: One collection was found where the margin of the fungus
encountered the margin of a narrow leaf of a deciduous shrub. The point of
contact appeared to be the dividing line between the spongiose tissue and the
hard layer bearing the teeth. The spine-bearing layer produced one or two teeth
below the leaf margin before growth ceased, but the spongiose tissue continued
growing, covering the leaf with a layer of tissue 5-6 mm thick. No layer of hard
spine-bearing tissue developed on the lower surface until after the spongiose
tissue had completely covered the leaf surface and extended over the far side.
Then spines were formed and the darker firm tissue was in evidence again. It is
concluded that the spongiose tissue is primary and produces the compact layer
with spines under suitable stimulus.

The name H. spongiosipes is reserved for the species under oak originally
described by Peck. The presence of a rather similar fungus, H. pineticola, from
under Pinus spp. was not considered until reported by Harrison (1964) although
Maas Geesteranus has pointed out that Coker & Beers’ description of the same
fungus as H. velutinum = H. spongiosipes varied somewhat from the usual con¬
cept of that species.

Maas Geesteranus (1957) placed H. spongiosipes Peck as a variety of H.
velutinum when making his interpretation of the European forms. This dis¬
position was followed by Harrison (1961) for the species from Nova Scotia now

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considered H. pineticola. H. spongiosipes has not been found in Nova Scotia so
now after a study of both species as they occur in Michigan it seems best to treat
all these variations with the violet to black to olive-green KOH reaction under
their original names. The finding of the species H. cumulatum in Nova Scotia in
1962 and by A. H. Smith in Michigan stresses the point that it may take many
years of careful collecting to find the hydnums in even well-collected regions. It
is no solution to the problem to consider the American species as a variety of H.
velutinum. The Friesian description varies considerably from the American
species under oak and there is no certainty that careful collecting in the type
locality may not turn up a form that will fit the Friesian description. Until
lectotypes are established from the type region that conform to the original
description, it will not be possible to have stability in this erratic-fruiting group
of fungi with such long-lived basidiocarps.

10. Hydnellum pineticola K. Harrison, Canad. Jour. Bot. 42: 1226. 1964.

Fig. 20.

Hydnellum velutinum var. spongiosipes (Pk.) Maas G., in Harrison, Stip.

Hyd.N. S.44 and PI. 3, fig. 10. 1961.

Pileus: 3-14 cm broad, convex, plane to slightly or irregularly depressed,
whitish pubescent becoming matted, surface roughened; “pale vinaceous cin¬
namon,” “vinaceous brown,” finally “russet” to “Mars brown,” occasionally
with pink drops in humid weather; when handled staining “vinaceous-cinnamon”
and finally “blackish-brown”; margin thick becoming thinner in age, pallid to
“vinaceous-fawn.” (This is also the color of secondary growth which may cover
the surface or occur only as a broad band around the margin.) Context dark
cinnamon, duplex, moist, tough, zonate, lower layer firm, with bands of crystals
in zones, upper thick and spongy and also showing zonations (daily growth
pattern); odor faintly fragrant to slightly acid; tase slowly disagreeable.

Spines: Up to 9 mm long, decurrent, fine, close, “pinkish-cinnamon”
becoming “Rood’s-brown” or darker (vinaceous brown), paler toward the mar¬
gin.

Stipe: 2 -4 (8) cm long, 1-2 (3) cm thick, solid, duplex, bulbous, zonate,
concolorous with pileus context, surface layer felty from a layer of soft tissue,
spongiose, inner layer hard, showing numerous crystals, base often radicating.

Chemical reactions: In KOH instantly black, in FeS04 olive-black on the
dark bands. KOH on the dried flesh flashes violet, then black and quickly leaches
leaving the tissue dull olivaceous.

Spores and Hymenium: Spores 4-5 x 4.5-6 p, angular, oblong to subglo-
bose, tuberculate, mucro inconspicuous, tubercles usually short. Basidia 5-6 x
20-25 ju, sterigmata 2-3 p long.

Tramal characters: Hyphae of the teeth 2.5-6 p flexuous, parallel, sparsely
branched from close to the septa. Hyphae of the pileus trama parallel and
flexuous in the firm part, loosely interwoven in the spongiose layer; compactly
and complexly interwoven at the base of the teeth, 4-6 p wide. Numerous
oleiferous hyphae 7 p wide with colored cell contents occur in the epicutis of
the pileus; also frequent dark-colored spots and pockets of crystalline material
which dissolve in KOH occur in the context.

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THE MICHIGAN BOTANIST

235

Habit, Habitat, and Distribution: Gregarious, concrescent, usually . under
Pinus spp. on sandy plains, occasionally in mixed conifers. Northern Michigan
and the Annapolis Valley of Nova Scotia.

Material examined: Imshaug 4682; Smith 34036, 38342, 38423, 38495,
38579, 42634, 42635, 42640, 50121, 50135, 50264, 58017,61257, 63420,
64025, 67627, 72006, 72316.

Discussion: This species occurs in great numbers in stands of Pinus
resinosa in Nova Scotia and under Pinus banksiana in northern Michigan. It
grows from deep in the pine straw as a whitish column that starts to expand into
a pileus as soon as it reaches the surface of the duff and in humid weather may
exude red drops of liquid. This is a slow-growing species and if growth is checked
by a change of weather, the manner in which it starts again will affect the shape
of the pileus. If growth recurs over the whole surface, the resulting fruit body
will be thick and turbinate or if only in a narrow band around the margin, it will
be thinner and spreading. Hydnellum spongiosipes (Pk.) Pouz. has not been
found in Nova Scotia, but is common in Michigan and, when the two species
were compared, with adequate field notes, points of difference were found.

The spores are the most useful means of separating this species from H.
spongiosipes (Pk.) Pouz. They average somewhat smaller (are mostly around 5.5
x 4.5 ju), more angular, and with shorter processes. H. spongiosipes has spores
with a large mucro which are mostly around 6.5 x 5.5 ju with processes that may
reach 2 p in height. H. pineticola is very subject to discoloring from bruising and
when growing actively cannot be touched without becoming discolored. The
contents of the hyphae of the pileus are noticeably dark. Coker and Beers’
(1951) description of Hydnellum velutinum and the illustration of the spores
(PI. 60, Fig. 8) are good for H. pineticola.

The latter differs from H. spongiosipes in that it is whitish at first and in
Michigan occurs most abundantly under pines. In the herbarium H. pineticola
has a “pinkish buff’ to “vinaceous cinnamon” pileus, the spongiose tissue is
“russet” to “Verona brown” and the flesh of the stipe is “Mars brown,” while H.
spongiosipes is “ochraceous tawny” to cinnamon” and the hard layer is “bister,”
“warm sepia,” and “burnt umber.” These differences will not separate old or
weathered material.

Stirpes Peckii and Scrobiculatum- Key to Species
1. Spores with fine tubercles (appearing echinulate under ordinary mag¬

nifications) . Y2.H. cumulatum

1. Spores with coarse tubercles . 2

2. Fresh context with a strong peppery to disagreeable after-taste in the

throat . 11. H. peckii

2. Taste not distinctive or merely slightly farinaceous to fungoid . 3

3. Pileus regular, thin and distinctly zonate, usually the disc smooth to

only slightly scrobiculate . 13. H. scrobiculatum var. zonatum

3. Pileus thicker than in above choice and typically scrobiculate and often

complante in addition . 13. H. scrobiculatum var. scrobiculatum

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THE MICHIGAN BOTANIST

Vol. 7

Stirps Peckii

11 . Hydnellum peckii Banker, Mycologia 5: 203. 1913.

Fig. 21.

Calodon peckii (Banker) Snell and Dick, in Snell et al., Lloydia 19: 163.

1956.

Hydnellum diabolus forma reticulatum Coker & Beers, Stip. Hyd. East. U.

S. 72. 1951.

Pileus: 3-14 cm broad, plane to depressed, margin slightly elevated; sur¬
face smooth, then uneven in center, with numerous scrupose projections,
cottony fibrillose toward margin, becoming matted, glabrous in age, white at
first, soon tinged pinkish, becoming buff-pink, in age “dull purplish-black” in
the center and shading to a vinaceous margin, frequently exuding drops of
reddish juice. Context coriaceous, zonate “pecan-brown” or darker, often
exuding red juice when squeezed; odor none or slightly disagreeable, to pungent
when squeezed, taste farinaceous but with a peppery after-taste in the throat,
remaining peppery in dried material; margin and teeth stain reddish brown on
handling.

Spines: 2-6 mm long, crowded, decurrent, “onion skin-pink” becoming
“Pecan-brown” or dark umber at maturity.

Stipe: 2-7 cm long, 1 .3 cm thick, surface irregular and rough expanding to
a bulbous base of mycelium and duff, concolorous with pileus; context solid,
dark “vinaceous-brown,” drying with numerous granular inclusions.

Chemical reactions: The dull purplish black areas on the surface are dark
in water under microscope and are “apparent-amyloid” in Melzer’s reagent. The
“apparent-amyloid” reaction has never been observed in H. diabolus. The same
tissue turns blue-green in KOH and the inclusions dissolve in the solution. The
reaction is usually faint and best observed under the stereoscopic microscope.
The mycelium and granules are not changed when the hyphae are flooded with
alcohol and resoaked in water.

Spores and Hymenium: Spores 4. 5-5. 5 x 3 .5-4.5 ju, “army-brown” in
mass, subglobose, tuberculate, tubercles rather low and some truncated.

Tramal characters: Hyphae loosely interwoven at the surface, with
numerous yellowish laticiferous hyphae 7-8 p in diameter, included in the layer;
context zonate, parallel, clamp connections present regularly on the hyphae,
with numerous amorphous inclusions in tissues of the stipe, these less frequent
in pileus.

Habit, Habitat, and Distribution: Solitary, gregarious or subcespitose
under conifers.

Discussion: This is one of the common species throughout the Pacific
Northwest but has been a poorly understood species in the east. A few collec¬
tions have been made in Nova Scotia but it was not properly understood until
western collections were studied. Coker found it in North Carolina but did not
recognize it and named it H. diabolus forma reticulatum. In the field H. peckii
has a slightly disagreeable odor and is a squat, stout plant but the surface is very

1968

THE MICHIGAN BOTANIST

237

uneven with numerous projections. The covering is cottony fibrillose and this
mats down and takes on a very dark appearance. H. diabolus Banker, which is
closely related, has a strong pleasant odor of bee balm (Monarda didyma) and
the surface appears hirsute (unevenly strigose under the lens) and rarely matted.
The differences in the surfaces of the two are recognizable in the herbarium.

The descriptions of this species and H. diabolus read very much like those
of H. ferrugineum of Europe, and herbaria in North America contain many
collections of these two under the latter name because of the presence of the red
juice. However, the acrid taste persists in the dried material, and both//, peckii
and H. diabolus can be recognized on this character. H. diabolus has now been
recognized in Europe.

Stirps Scrobiculatum

12. Hydnellum cumulatum K. Harrison, Canad. Jour. Bot. 42: 1225. 1964.

Fig. 22.

Pileus: 3-9 cm broad individually, connate, often concrescent, plane, con¬
vex, or concave, imbricate, rarely with concentric ridges near margin, spongiose
to tomentose, “vinaceous-buff’ to avellaneous”, becoming “Hessian-brown” or
“burnt-umber” or darker — almost black when wet; margin thin, elevated, with a
zone-like growth band, “vinaceous-buff’ to “avellaneous” to “brick-red”;
context fibrous, firm, “brick-red” to ““Hessian-brown”; odor slightly aromatic,
taste mild.

Spines: Fine, short, close, “liver-brown,” becoming darker, not changing
color when bruised (but rather mature when collected).

Stipe: 2 -4 cm long, 0.5-1 cm thick, equal to subclavate, very little surface
showing, concolorous with pileus to darker; context hard, fibrous, “brick-red”
to almost black: a small amount of spongiose tissue present around the base.

Chemical reactions: No reaction in Melzer’s solution but the KOH reac¬
tion was typical of stirps Velutinum.

Spores and Hymenium: Spores 4-5.5 x 4-5 p, colored, subglobose, thin-
walled, finely tuberculate (almost appearing echinulate) with prominent mucro.
Basidia 5-7 x 20 p, sterigmata up to 4 p, fine.

Tramal characters: Hyphae in the spines 2.5-3 p, parallel and flexuous;
spongious tissue of interwoven hyphae at the surface, mostly 3-5 p.

Habit, Habitat, and Distribution: Solitary to gregarious in connate masses,
appearing sessile in the duff. On hillsides in mixed stands of mature Pinus
resinosa and Tsuga canadensis, found on the edges of small hollows and in
depressions around exposed roots, in unused rodent burrows and partly hidden
by duff. Previously known only from Kings and Annapolis Counties, Nova
Scotia, Canada.

Material examined: Smith 72082, 72317, 72479, from Luce County,
Upper Peninsula.

Discussion: This species bears a remarkable outward resemblance to
mature carpophores of H. diabolus Banker and this was supported by the slight
aromatic odor. However, there was no trace of an acrid taste. The dried plants

238

THE MICHIGAN BOTANIST

Vol. 7

are near snuff brown in color. The spores have a typical ornamentation that
enables this species to be separated from H. diabolus in the herbarium. It bears
little resemblance in outline to other species in the velutinum-scrobiculatum
group. Apparently it fruits only at long intervals. The area where it was first
found in the ravine on the Research Station, Kentville, Nova Scotia, has been a
favorite collecting area for 35 years, and the species was first found in 1962. It
was also found the same season in quantity in a similar habitat in another
county. The heaped up appearance of the connate masses of pilei gives this
fungus a characteristic appearance.

13. Hydnellum scrobiculatum (Seer, ex Fries) Karst., Medd. Soc. Fauna FI.
Fenn. 5: 41. 1879. var. scrobiculatum,

Fig. 23.

Hydnum cyathiforme a. Bull, ex Fries, Syst. Myc. 1: 405. 1821; non H.
cyathiforme Schaeff. 1774.

Hydnum concrescens Pers., Mycol. Europ. 2: 164. 1825.

Hydnum scrobiculatum Seer, ex Fries, Mycogr. Suisse 2: 522. 1833.

Pileus: 2-7 (15) cm broad, singly to gregarious and concrescent in masses,
convex, to shallowly infundibuliform or broadly depressed; margin thick, some¬
times thin, elevated; surface finely to coarsely scrobiculate especially in the
center or occasionally colliculose, rough, irregular or striately ridged towards
margin, evenly coarsely velvety tomentose on smoother parts, dull onion skin-
pink, becoming Rood’s-brown or dark ferruginous, occasionally seen with red¬
dish brown droplets of juice; margin zonate sometimes with alternating bands
pecan-brown to buff-pink, staining bone-brown to blackish when injured. Con¬
text zonate, tough, fibrous to near woody, cinnamon-rufous to light brown,
typically dry; odor and taste mild, slightly pungent to farinaceous disagreeable.

Spines: 1-2 (3.5) mm long, decurrent, very fine, close chestnut-brown to
chocolate.

Stipe: 1-3.5 cm long, 1-4 cm thick, extremely variable from fusing, arising
from a spongy tomentum, narrowing upward then expanding into pileus, con-
colorous; context tough, fibrous, woody in base.

Chemical reactions: When fresh: KOH dark olive, FeS04 olive black, Mel-
zer’s negative; when dried: KOH giving a violet flash then black and a dark olive
color leaches out.

Spores and Hymenium: Deposit pale vinaceous brown; spores 5. 5-6. 5 x
4. 5-5. 5 ju, subglobose to slightly angular, many tubercles stud-like, blunt, 10 to
12 showing on circumference. Basidia 5. 5-6. 5 x 15-20 n, clavate with broad
ends. Hymenium up to 30 ii thick, no clamps seen.

Tramal characters: Spongiose layer very thin but the hyphae are 4. 2-5. 3 n
wide — mostly tubular but a few irregulary swollen, septate, thin-walled, no
clamps seen. Some hyphal ends are filled with dark contents and dark debris was
present amongst the strands. The harder inner layer of hyphae 3. 8-4. 5 ju wide,
tubular, septate, no clamps seen. Two types of incrusting granules seen: the
usual type was dark reddish brown and rarely pockets of granules were present

1968

THE MICHIGAN BOTANIST

239

amongst the hyphae. Laticiferous hyphae with similarly colored contents were
also present. The second type is present only on short segments of the hyphae.
The hyphae are dark brown and this gives the impression that they are segments
that would correspond to the “apparent-amyloid” hyphae in some other species
of Hydnellum. The intense dark olive reaction in KOH in this species prevents
any observations on whether these same tissues give the blue-green reaction that
is usually associated with these dark granules in the other species. Hyphae of the
spines are undulating, 2.5 A \ u wide, thin-walled, septate, and no clamps were
seen.

Habit, Habitat, and Distribution: Solitary, gregarious, or concrescent, in
deciduous and conifer woods throughout the state. We have collections from 9
counties. It is one of our most common hydnellums. It is widely reported in
North America from the Atlantic to the Pacific Ocean in forested areas and from
Canada to Alabama.

Material examined: Mains 4532, 6590. Harding 326. Shaffer Aug. 6,
1960. Thiers 1096. Smith 1725, 7081, 18500, 18501, 18531, 18532, 32884,
33121, 35839, 50263, 50583, 57835, 62853, 64028, 64189, 64229, 64329,
64835,72464,75034,75120.

Discussion: The species described here forms part of a complex group that
includes “H. zonatum” and in this paper is being restricted to collections with
spore ornamentation in the form of tubercles. The two varieties are quite
uniform in their appearance and in most cases the var. zonatum can be dis¬
tinguished by being thinner, rarely as scrobiculate and typically more conspic¬
uously zonate. When collections of this complex from all over North America
are brought together and studied it is fairly easy to pick out eight or ten distinct
forms associated with the different regions. It is hoped that eventually it will be
possible to arrive at a classification that can be used to differentiate these into
recognizable varieties that will be useful for studying the variation within the
group or to aid in understanding their ecological reactions.

Fries originally described H. scrobiculatum and H. zonatum as distinct but
they have been the subject of confusion ever since. Banker treated the North
American complex under a number of names and attempted to associate them
with European names. He also added the new species H. parvum Banker, a small
southern form growing in pine straw. Miller and Boyle (1943) were doubtful if
two species could be distinguished. Coker and Beers (1951) treated the eastern
North American forms under five species names. Harrison (1961) working in
Nova Scotia considered H. zonatum a variety of H. scrobiculatum while Lundell
in 1959 issued zonatum as a subspecies of Hydnum scrobiculatum. Maas
Geesteranus in Europe considers them all varieties of H. velutinum, which is
another species described by Fries which has to be interpreted from the original
description and tradition as no type exists. The name has been applied to a
number of species none of which fit the original description accurately.

Hydnellum scrobiculatum (Seer, ex Fries) Karst, var zonatum (Batsch ex Fries)
K. Harrison, Stipp. Hyd. N. S. 43. 1961.

Figs. 24 & 25.

240

THE MICHIGAN BOTANIST

Vol. 7

Hydnum zonatum Batsch, Elenchus, Contin. sec. 109, pi. 40, fig. 224.
1789; non//, zonatum Batsch, Elenchus 111. 1783.

Hydnum cyathiforme b. Fries, Syst. Myc. I: 405. 1821; non H.
cyathiforme Schaeff. 1774

Hydnum zonatum Batsch ex Fries, Epicr. Syst. Myc. 509. 1836-38.
Hydnellum zonatum (Fries) Karst., Medd. Soc. Fauna FI. Fenn. 5: 41.
1879.

Hydnellum velutinum var. zonatum (Fries) Maas G., Fungus 27: 64. 1957.

Pileus: 3-6 cm broad, convex, infundibuliform, margin thin, rarely
“thick” unless small, zonate, usually elevated, center often slightly scrobiculate,
surface rough, finely fibrillose on margin, bright to dark rusty cinnamon ; margin
fibrillose, thin, sterile, rarely “thick” unless basidiocarp is young, usually ele¬
vated, near vinaceous pink, staining vinaceous cinnamon when bruised. Context
2. 5-3. 5 mm thick, zonate, soft-fibrous to tough, sometimes soft, light brown to
vinaceous russet; taste mild, odor slight. One collection had a rancid taste.

Spines: Up to 3 mm long, decurrent, very close and slender, dark vin¬
aceous cinnamon.

Stipe: Up to 3 cm long, 0.5-1 (2.5) cm thick, simple or compound, arising
from a tomentose pad of mycelium; context zonate and hard.

Chemical reactions: When fresh: KOH instantly fuscous to olive black;
FeS04 dark olive fuscous next to teeth; dried: KOH gives a violet flash andthe
black dissolves in the liquid as a dark olive pigment.

Spores and Hymenium: Spores 4. 5-5. 5 x 4-4.5 pi, angular, tuberculate, in
Melzer’s reagent as seen in groups showing a yellowish-red tint. Basidia 20 x 5.3
pi, clavate, 4-spored.

Tramal characters: Only a few strands of spongiose tissue present on
surface of pileus. Hyphae 3A pi wide, thin-walled, frequently septate, not
clamped, many ends filled with reddish brown contents and resembling laticifers.
The ends often associated with the band of crystalline material marking the
hyphal zones; some hyphae incrusted but granules usually intercellular, occa¬
sional visible pockets of reddish brown material present, no evidence of
apparent-amyloid hyphae seen; both spongiose and context hyphae radially
oriented; inner layer of hyphae 3 .5-4 pi wide, thin-walled and some slightly
incrusted, tubular, septa numerous. Hyphae of spines 2.5-3 pi, parallel except in
the base where they appear interwoven.

Habit, Habitat, and Distribution: Solitary, gregarious, or in concrescent
masses in conifer and deciduous woods. It is common throughout the state as
well as in forested areas of North America generally. It was described, of course,
from Europe.

Material examined: Kauffman Sept. 24, 1904, Oct. 1907. Mains July 30,
1915, Aug. 1, 1915, Aug. 6, 1915, 1076, 1071. Lowe 124, 172. Harding 250,
293, 301. Smith 32-266, 1739, 6876, 6950, 6955, 7046, 7141, 7155, 7180,
7285, 7288, 18502, 18530, 18543, 18729, 18751, 18752, 18769, 20730,
20781, 33085, 34000, 37139, 37283, 38111, 38487, 38498, 62611, 64059,
64113, 64228, 64356, 64631, 67500, 67793.

1968

THE MICHIGAN BOTANIST

241

Discussion: The spore differences between this and var. scrobiculatum are
slight. In the field the thinner, more zonate, smoother pilei are the major distinc¬
tion. For additional comment see var. scrobiculatum.

Stirps Geogenium

14. Hydnellum geogenium (Fries)Banker, Mycologia 5: 204. 1913.

Hydnum geogenium Fries, Sv. Vet.-akad. Forh., 131. 1852.

Calodon geogenius (Fries) Karst., Medd. Soc. Fauna FI. Fenn 6: 16. 1881.

Pileus: Concrescent clumps up to 25 cm across with individual pilei up to
6.5 cm broad, variable in shape, shelving, substipitate, thin, pliant, subzonate,
surface fibrous with radiating ridges, center scrobiculate; margin variously lobed,
undulating or irregular, color “deep colonial-buff’ to “olive-yellow” when
rubbed buffy-olive. Context thin, pliant, olive beneath the honey-yellow surface
layer; no odor or taste.

Spines: 3-6 mm long, decurrent, close, fine, “olive-ocher” (olive-yellow)
becoming dark “vinaceous-brown” in age with “olive-ocher” tips.

Stipe: None or very short. The soil around the base filled with numerous
strands of greenish-yellow mycelium.

Chemical reactions: In dried material a dark olivaceous pigment dissolved
out into the KOH solution and areas of green hyphae were evident in the tramal
tissue near the surface and some of the zones.

Spores and Hymenium: Spores dark “vinaceous-brown” in mass, 3-4.5 x
34 \i, angular to subglobose, tuberculate, tubercles rather low, prominent
apiculus. Basidia 1040 x 4-5 \ u, clavate, 4-spored.

Tramal characters: In KOH, the surface hyphae 2. 5-5. 5 [i wide, thin-
walled, interwoven, with branches and usually near a branch a septum with a
large medallion clamp ; the loop of the clamp nearly as broad as the hypha. There
is also a system of hyphae with somewhat darker contents (laticifers?) and the
dark contents often end abruptly at a clamped septum. In Melzer’s solution the
laticiferous system shows up plainly and many of the surface hyphae contain
black granules. The main context of the pileus consists of hyphae 3-4 ju wide,
tubular parallel, septate and with large clamps at some of the septa. The latici¬
ferous hyphae are 5.5 fi wide. The hyphae in the spines are 2. 5-3.8 ju wide,
parallel, thin-walled, and with clamps.

Habit, Habitat, and Distribution: Solitary, gregarious, concrescent under
conifers. The only collection known from the west is Smith 52917, San Juan,
Colorado. Common in Nova Scotia. Reported along Atlantic seaboard from
North Carolina to Quebec. Probably will be found in the Upper Peninsula.

Discussion: This is a common species in the northern coniferous forests in
the eastern part of the continent. It occurs on slopes and along stream banks in
large concrescent masses. It is much rarer south but has been found in Georgia.
There it occurs as individual stipitate fruit bodies. When growing luxuriantly it
starts as a group of bright yellow columns on a cushion of yellowish mycelium.
It grows so compact and close to the ground that it often appears to be sessile,

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and it may grow out on the underside of exposed roots and lumps of soil. Its
bright yellow margin and mycelium, along with the dull olivaceous pileus are
characters that make this species conspicuous and easy to recognize.

Stirps Mirabile

15. Hydnellum mirable (Fries) Karst., Medd. Soc. Fauna FI. Fenn. 5: 41. 1879.

Hydnum mirabile Fries, Monogr. Hymen. II: 349. 1863

Calodon mirabilis (Fries) Karst., Medd. Soc. Fauna FI. Fenn. 6: 16. 1881.

Pileus: 4-15 cm broad, broadly convex or flat, surface becoming irregular
and the obtuse margin lobed, densely tomentose to short strigose, plush-like near
margin but tomentum aggregated into coarse upright projections over most of
the surface; color very sordid yellowish buff becoming dirty dark yellowish-
brown. Context thick, coriaceous, woody, softer toward surface, zoned with
dark brown lines, pallid to dull buff otherwise, with a copious dull umber juice,
in base of stipe “pale grayish brown” with lighter areas near surface; odor faint
of raw cucumber or subfarinaceous, taste faint, subfarinaceous but finally slowly
acrid.

Spines: 2-5 mm long, decurrent, distant, crowded, in age frequently flat¬
tened, often with several tips, pallid wood-brown to clay-color.

Stipe: 2-3 x 2-4 cm, enlarged upward, hard and firm but breaks cleanly
and exudes a dingy watery juice, scabrous from undeveloped teeth, yellowish
brown, tomentose in patches; context corky, dingy (pale grayish brown).

Chemical reactions: No reactions obtained with KOH or Melzer’s reagent.

Spores: Spores 4. 5-6. 5 x 4-5.5 ju, colored, subglobose, coarsely tuber-
culate, tubercles usually short, 8 or 9 showing on circumference.

Tramal characters: Context drying very hard and woody and the teeth
hard and brittle, not reviving readily in Melzer’s solution.

Habit, Habitat, and Distribution: Gregarious, subcespitose, in mixed and
coniferous woods. Recorded under pine and hemlock from Idaho, Oregon, and
Washington. Rare in eastern United States, Canada, and Europe. One collection
from Tennessee. Its occurrence in the coniferous forests of Nova Scotia and the
Pacific Northwest leads to its inclusion here as a species that will probably be
found in the Upper Peninsula.

Discussion: This species is rarely collected, possibly because it is so attrac¬
tive to slugs. It starts as straw-colored cushions of mycelium that exude coffee
colored drops of juice and this characteristic is apparent when the flesh and
particularly the stems are broken. The juice dries and darkens the mycelium so
that old specimens become very sordid and dark in appearance. One or more
fruit-bodies may arise from one cushion and then they are very irregular in
shape. There is a strong tendency for regrowth following injury and when this
happens the new mycelium is white at first. The teeth at first are distant and
terete, but in age they are often fused in plates with two to several tips. The
decurrent teeth next to the stipe frequently fuse with it and become partially

1968

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243

buried. This species is very low and squat and when seen actively growing resem¬
bles Polyporus schweinitzii. There is a layer of spongy tissue near the surface
which dries hard especially in young basidiocarps.

ACKNOWLEDGMENTS

The preparation of this paper was possible because of the efforts of many amateur and
professional mycologists who gathered hydums over the years and deposited them in the
University of Michigan Herbarium. My thanks are extended to these many persons past and
present. It is with gratitude that I acknowledge the importance of the contributions made to
this paper by Prof. A. H. Smith, Director of the University of Michigan Herbarium. He not
only has found the majority of the collections recorded here but generously granted per¬
mission to make full use of the numerous field notes and photographs accompanying the
specimens. The microscopic studies were made and the paper prepared during the tenure of
NSF grant GB 4853 to the University of Michigan and the financial support essential for its
completion is gratefully acknowledged.

REFERENCES

Banker, H. J. 1906. A contribution to a revision of the North American Hydnaceae. Mem.
Torrey Bot. Club 12: 99-194.

Bresadola, J. 1932. Iconographia mycologica. 21: pis. 1035-1050; 22: pis. 1051-1060.
Coker, W. C., & A. H. Beers. 1951. The Stipitate Hydnums of the Eastern United States.
Univ. N. C. Press, Chapel Hill.

Cooke, M. C., & L. Quelet. 1878. Clavis Synoptica Hymenomycetum Europaeorum. Lon¬
don.

Harrison, K. A. 1961. The stipitate hydnums of Nova Scotia. Canada Dep. Agr. Publ. 1099:
1-60.

Harrison, K. A. 1964. New or little known stipitate hydnums. Canad. Jour. Bot. 42:
1205-1233.

Harvey, R. 1958. Sporophore development and proliferation in Hydnum auriscalpium Fries.
Trans. Brit. Mycol. Soc. 41: 325-334.

Karsten, P. A. 1879. Symbolae ad Mycologiam Fennicum. Medd. Soc. Fauna FI. Fenn. 5:
40-42.

Maas Geesteranus, R. A. 1957. The stipitate hydnums of the Netherlands. II. Hydnellum P.
Karst. Fungus 27: 50-71.

Maas Geesteranus, R. A. 1958. The stipitate hydnums of the Netherlands. III. Phellodon P.

Karst, and Bankera Coker & Beers ex Pouz. Fungus 28: 48-61.

Miller, L. W., & J. S. Boyle. 1943. The Hydnaceae of Iowa. Univ. Iowa Stud. Nat. Hist. 18:

1-92

V, /V //V f \/ 1 o v f

Pouzar, Zd. 1956. Pnspervek k poznam nasich kloboukatych losaku. Ceska Mykol. 10:

65-76.

Quelet, L. 1886. Enchiridion Fungorum in Europa Media, 1880-93. Lutetia.

244

THE MICHIGAN BOTANIST

Vol. 7

Fig. 1. Phellodon niger var. alboniger xl Smith

Fig. 2. Phellodon niger var. alboniger xl Smith

1968

THE MICHIGAN BOTANIST

245

Fig. 3. Phellodon tomentosus xl Smith 63984

Fig. 4. Phellodon tomentosus xl Smith 70681

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THE MICHIGAN BOTANIST

Vol. 7

Fig. 5. Phellodon confluens xl Smith 64249

1968

THE MICHIGAN BOTANIST

247

Fig. 6. Bankera fuligineo-alba xl Smith 67761

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Vol. 7

Fig. 7. Banker a carnosa xl Smith 72165

1968

THE MICHIGAN BOTANIST

249

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Fig. 10. Hydnellum humidum xl Smith 64248

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251

Fig. 11. Hydnellum humidum xl Smith 64248

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Vol. 7

252

Fig. 12. Hydnellum humidum xl Smith 64404

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253

Fig. 13. Hydnellum caeruleum xl Smith 73311

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Fig. 14. Hydnellum ferrugipes xl Smith 61402

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255

Fig. 15. Hydnellum septentrionale xl Smith 33082

4 A*.

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Fig. 16. Hydnellum aurantiacum xl Smith 66445

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257

Fig. 17. Hydnellum earlianum xl Smith 62851

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Fig. 18. Hydnellum conigenum xl Harrison 6226

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259

Fig. 19. Hydnellum spongiosipes xl Smith 73288

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Fig. 20. Hydnellum pineticola xl Smith 38579

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261

Fig. 21. Hydnellum peckii xl Smith 23699

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Fig. 22. Hydnellum cumulatum xl Smith 72317

Fig. 23. Hydnellum scrobiculatum var. scrobiculatum xl Smith 64028

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Fig. 24. Hydnellum scrobiculatum var. zonatum xl Smith 64059

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265

f lature education feu tare --

CONTACT DERMATITIS CAUSED BY PLANTS1

A. P. Ulbrich, D. 0.

4401 N. V/oodward Ave., Royal Oak, Michigan

The major plants causing contact dermatitis are in the genus Toxi¬
codendron (often included in the more comprehensive sumac genus, Rhus).
These include, in the midwest and eastern United States, poison ivy
(Toxicodendron radicans ) and poison sumac ( Toxicodendron vernix). In the
southeastern United States there is also poison oak ( Toxicodendron
quercifolium), and on the Pacific side of the country there is the western poison
oak ( Toxicodendron diversilobum). While there are species differences, to a
patient these are probably incidental. These plants all presumably contain the
same antigen, namely, pentadecylcatechol.

Other plants contain the same or a closely related toxin, urushiol. The hull
of the cashew nut and, in fact, all of the anacardiaceous family of plants have
this resin. It is also found in the hull or skin of the mango of Florida and the
tropics. The lacquer tree ( Toxicodendron vernicifera), from which the Japanese
have long made beautiful lacquer ware, has the same antigen. This tree was
introduced into Japan from China, where it is native, and where the Chinese in
addition used the fruit of the ginkgo tree as a base for lacquer. The pulp of the
seed of this tree acts as a primary irritant, and epidemics of contact dermatitis
have been traced to areas where students have walked under a ginkgo tree and
stepped on the plumlike seeds.

Physicians are aware that a host of other plants may produce contact
dermatitis, but not by the same antigen, in sensitive persons. The wild carrot or
Queen Anne’s lace ( Daucus carota) is one example. A trip to the country and
back to nature in which children or adults pick buttercups ( Ranunculus species)
may result in a contact dermatitis. These plants contain a substance called
anemonin which may cause blistering. Mayapple ( Podophyllum peltatum),
particularly the root, stem, or rhizome of the plant which contains podophyllin
(a cytotoxin), may cause contact dermatitis. Dermatologists use podophyllin in
the treatment of verruca acuminata and seborrheic keratoses, and some work has
been done using podophyllin resin in the treatment of skin cancer.

The lady-slipper genus ( Cypripedium ) of wild orchids have long been a
much admired group by naturalists. With the increasing use of our state parks
and wilderness areas, these plants are found by persons who feel compelled to
“pick the pretty wild flowers” or attempt to take the plants home. Lady -slippers
can be admired and photographed, but should not be handled. Any of our

* Reprinted by permission, and with slight revisions by the author, from The Journal
of the American Osteopathic Association 64: 1023-1029 (June 1965). Much of this material
was also included in an illustrated talk before the state meeting of the Michigan Botanical
Club at the Cranbrook Institute of Science, March 17, 1968.

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species may produce a toxic reaction. (Furthermore, in Michigan all orchids are
protected and it is unlawful to pick them.)

Some cultivated plants also cause reactions. Often involved is the
Euphorbia group. The juice of one species, white-margined spurge, grown as an
ornamental, “snow-on-the-mountain” ( Euphorbia marginata), “has been used for
branding cattle in preference to a hot iron.” (Kingsbury, 1964) Chrysanthemums
contain pyrethrumlike substances which may be antigenic. Primrose ( Primula ) is
another of the flower garden plants that may produce contact dermatitis; potted
plants have even become an occupational hazard for nurses in hospitals. The
rhizome or rootstock of iris may irritate some people. Lily-of-the-valley
( Convallaria majalis) not only contains a digitalislike substance but also may
produce dermatitis.

However, poison ivy, the poison oaks, and poison sumac produce the
major portion of contact dermatitis from plants. There is much folklore about
poison ivy, and much of it is unreliable. Reported treatments for the dermatitis
range from fuming nitric acid to marshmallows and cream. The supposition that
the serum of the dermatitis carries the antigen is false. The uninjured leaf of
poison ivy, oak, or sumac does not emit the toxin, as the plants do not have
hydathodes (special groups of cells which excrete fluid— which appears like
dewdrops on plants such as tomatoes when humidity is high).

The flowers of poison ivy are small, in clusters, and often unisexual. The
pollen is one part of the plant which does not contain the allergen. The fruit, less
than a quarter-inch in diameter, is a dryish ivory drupe, readily found through¬
out the fall and winter. (A drupe is a fleshy fruit the seeds of which are con¬
tained in a hard “pit,” as in peach and cherry.) The waxy flesh around the pit is
relatively scant but enough to attract and be eaten by over 60 species of
birds, which relish the fruit as winter food. The pits or “seeds” pass through the
bird’s digestive tract with only the flesh being removed; the viable seed remains
intact, germinates, and the bird has helped disseminate poison ivy. Poison ivy is
often profuse along fencerows and near bird feeders or where birds are fed in the
winter.

There is no real substitute for recognition of these plants. This, of course,
leads to avoidance, which is the best prophylaxis. The potential antigenic effect
is phenomenal. Kligman (1958a) had a patient who was not sensitive to the
toxin of poison ivy apply crushed leaf to his thumb. The thumb was applied to a
nonsensitive person in a new area each time for a total of 500 times. The
pressure each 100th time was applied to a highly sensitive individual. The
sensitive skin reacted at all five sites of pressure. Even the 500th pressure caused
a reaction to the infinitesimal quantity transmitted. A second study was done in
which the juice of a crushed leaf was applied to sensitive individuals. It was
allowed to remain in contact varying lengths of time— 5, 10, 30, and 60 min¬
utes— and the skin was then washed with Ivory soap for two minutes. In highly
sensitive persons, after five minutes the fixation was complete and could not be
altered; subsequently, dermatitis developed.

The use of soap and water, therefore, is in a questionable position. Its use
will reduce the incidence of transfer but not remove the antigen in its entirety,

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267

only the excess. Washing with equal parts of acetone and alcohol or U.S.P.
semisolid green soap did not affect the removal of the antigen. The antigens of
other plants have not been as thoroughly studied, but a good cleansing with soap
is a help only to remove excess antigen.

The transfer of antigen should be considered. Cats and dogs are pets often
involved; they may carry the toxin on their fur and transport it to humans.
Persons who have an allergy to pentadecylcatechol may develop dermatitis from
contact with their pet’s fur, without leaving the house. Fireplace wood that has
been contaminated by vines or leaves of poison ivy is another source. Smoke
may be a vector which precipitates either directly on the host or on tools and
clothing. Contaminated wood, golf balls, shoes, car tires, clothing, and lawn
mowers are other means of transmitting the material. The pentadecylcatechol
deteriorates slowly. Thorough laundering will remove it from washable clothing.
On gloves, shoes, and clothing not laundered the resin will undergo oxidation,
polymerization, and deterioration in a moist atmosphere more rapidly than in a
dry one.

If the plant is intact and there is no escape of the resin, the antigen is not
on the surface of the leaf. It is only when the leaf is traumatized or injured that
the sap containing the antigen is present. The trauma may be minimal, from
brushing or bruising the plant or from injuries produced by insects. Perhaps we
do not think of this plant as being attractive to insects, but it is palatable and
many of the leaves will show insect damage. The trauma to the plant as one
walks by and injures the leaves and stems or as one treads on it and it springs
back tends to encourage contact with the resin. There has been speculation that
some fungi may cause enough injury to allow escape of the catechol. A micro¬
scopic examination of poison ivy shows that the resin ducts are large in propor¬
tion to those of many other plants, but no suggestion can be found that poison
ivy, oak, or sumac is any more brittle than other plants. Nor has there been any
suggestion that the resin is under unusual pressure in the ducts.

Protective creams, lotions, etc., are not prophylactically effective to
prevent contact. In fact, oils and creams enhance absorption and help the
dissemination of pentadecylcatechol, which is dispersed in the oil base residues.
There has been much proprietary advertisement about oral desensitization with
poison ivy extract, alum-precipitated. The material is inert, and there is no
clinical or patch test evidence of poison ivy desensitization (Wechsler, 1964).

Once the allergen has joined with the protein of the skin, what is the
therapy of choice? This, of course, depends upon the potential quantity of
allergen on the skin. As mentioned previously, washing with soap removes only
that antigen not already fixed in the tissue (the type of soap is incidental). Also,
some type of compress can be used, the character depending on the experience
of the prescribing doctor. Beyond simple compresses or a mild lotion no therapy
is recommended. A physician should be consulted and his orders followed.

REFERENCES

NOTE: There axe many additional sources of reading available in medical libraries and in
some public libraries.

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Vol. 7

Fisher, A. A. 1967. Contact Dermatitis. Lea & Febiger, Philadelphia. 324 pp.

Kingsbury, J. M. 1964. Poisonous Plants of the United States and Canada. Prentice-Hall,
Englewood Cliffs, N. J. 626 pp.

Kligman, A. M. 1958a. Poison ivy (Rhus) dermatitis: An experimental study. AMA Arch.
Dermat. 77: 149-180.

Kligman, A. M. 1958b. Hyposensitization against Rhus dermatitis. AMA Arch. Dermat. 78:
47-72.

Sowers, W. F., et al. 1965. Ginkgo-tree dermatitis. AMA Arch. Dermat. 91: 452-456.
Wechsler, H. L. 1964. Poison ivy extract, alum precipitated. Jour. AMA 190: 845.

ADDITIONAL WISCONSIN RECORDS FOR THE
EBONY SPLEENWORT, ASPLENIUM PLA TYNEURON

Lytton J. Musselman
Department of Botany and Zoology,

University of Wisconsin, Rock County Campus, Janesville

Asplenium platyneuron (L.) Oakes has previously been known from only
two Wisconsin counties: Dane (Tryon et al., 1953) and LaCrosse (Hartley,
1959). Fell (1955) cites several localities in Winnebago County, Illinois, just
south of the Wisconsin state line. In conjunction with the present study I have
examined specimens in the following herbaria: University of Wisconsin, Mil¬
waukee Public Museum, University of Wisconsin— Milwaukee, Beloit College, and
Milton College. Specimens of the following collections are deposited in the
herbarium of the University of Wisconsin, Rock County Campus, Janesville.

I have collected Asplenium platyneuron at two sites in Rock County,
Wisconsin. In Magnolia Township, T3N, R10E, two individual plants were found
in October 1967 in a white birch— mixed oak woods on a north-facing sandstone
slope in the county park east of Croak Road, SE 1/4 of section 18 (Musselman
1592). Repeated visits to this area have failed to reveal additional plants.

On January 22, 1968, I encountered a population of approximately 22
individuals in a smooth sumac thicket on a well eroded east-facing sandstone
slope in the NW 1/4 of section 23, Plymouth Township, T2N, R11E (Musselman
1659).

LITERATURE CITED

Fell, E. W. 1955. The Flora of Winnebago County, Illinois. Nature Conservancy, Washing¬
ton, D.C. 207 pp.

Hartley, T. G. 1959. Notes on some rare plants in Wisconsin-I. Trans. Wis. Acad. 48: 57-64.
Tryon, R. M. Jr., N. C. Fassett, D. W. Dunlop, and M. E. Diemer. 1953. The Ferns and Fern
Allies of Wisconsin. 2nd ed. Univ. Wis. Press, Madison. 158 pp.

1968

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269

THE MILTON COLLEGE HERBARIUM

Lytton J. Musselman
Department of Botany and Zoology,

University of Wisconsin, Rock County Campus, Janesville

The herbarium of Milton College, Milton, Wisconsin, contains an ex¬
tremely valuable collection of several thousand specimens of vascular plants and
cryptogams which has apparently escaped notice during the past seventy years.
The existence of the collection was recently brought to the author’s attention in
connection with floristic studies in southern Wisconsin. The collection is of more
than passing interest to Wisconsin botanists since it contains many species now
extremely rare or extinct in southern Wisconsin and may provide additional
locality records for persons working on the state flora.

The Milton collection consists largely of specimens either donated by or
purchased from a Mr. S. H. Watson, a little known Wisconsin botanist active
from ca. 1859-1862. All of his collections were made in southern Wisconsin and
are valuable indicators of a bygone vegetation. Several orchids and other plants
now nearly extinct in southern Wisconsin are noted by Watson as being
“common” or “not rare.” Although most of his collecting was done in Dane
County (largely Dunkirk Township but also Madison, Pine Bluff, and Blue
Mounds) it is not mentioned by those who have studied the flora of Dane
County (Cheney and True, 1893; Ellarson, 1949). Occasional specimens are
from Rock County (Janesville, Fulton, Porter, and Union Townships); Iowa
County (vicinity of Arena), and Sauk County (bluffs of the Wisconsin River).
His labels simply read “S. H. Watson, Madison, Wis.” with the name of the plant,
location (usually very general), and year of collection. Few of Watson’s spec¬
imens are in any Wisconsin herbaria other than the herbarium of the University
of Wisconsin (Madison) which received several hundred Watson collections from
the Academy of Natural Sciences of Philadelphia about ten years ago.

Pennell (1935) in his listing of collectors of Scrophulariaceae, includes S.
H. Watson as having specimens deposited in the herbarium of the University of
Michigan. No mention is made of any Wisconsin herbarium.

Mr. Watson apparently collaborated with his better known fellow-botanist,
T. J. Hale. Hale cites several Watson specimens in his papers on the flora of
Wisconsin (Hale 1858, 1860). Several of Watson’s labels read “Ex. Coll. T. J.
Hale.” Hale had several of his specimens identified by D. C. Eaton, a prominent
botanist of the period.

The Milton collection is rich in specimens from the collection of the well
known botanist of Watson’s day, Wm. Canby. Other collectors represented by a
lesser number of specimens than those of Canby include Increase A. Lapham and
T. J. Hale (Wisconsin); George Vasey, Samuel B. Mead, Michael S. Bebb, Elihu
Hall (Illinois); Mary H. Clark (Ann Arbor and Detroit areas of Michigan); C. M.
Short (Kentucky); and H. Jackson (Pennsylvania and Maryland). Worthy of note
are a few specimens of Carex from the collections of Schweinitz and Torrey. In
addition there are a few specimens deposited by staff members and students of

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Milton College about the turn of the century. The source of the older specimens
is a matter of conjecture. They may have been gathered by Watson or by a
member of the Milton College staff. However, evidence seems to point to their
acquisition by S. H. Watson since duplicates of many of the above collectors
were found in Madison newspapers dated 1860-1862.

Despite an absence of care and a century’s deposition of coal dust, the
collection is in excellent condition. Fortunately, many duplicates are available
and are being distributed to Wisconsin herbaria.

It is hoped that Wisconsin botanists will now avail themselves of the use of
this collection. Facilities and staff are not available for the loaning of specimens.
Persons interested in examining the collection should contact the author or
Professor F. M. Van Horn, Chairman, Department of Biology, Milton College,
Milton, Wisconsin 53563. The symbol suggested for the next edition of the Index
Herbariorum is MCW.

ACKNOWLEDGMENTS

I am indebted to L. M. Van Horn, J. L. Skaggs, and A. L. Millard for their assistance.

REFERENCES CITED

Cheney, L. S., & R. H. True. 1893. On the flora of Madison and vicinity, a preliminary
paper on the flora of Dane County, Wisconsin. Trans. Wis. Acad. 9: 45-136.

Ellarson, R. S. 1949. The vegetation of Dane County, Wisconsin in 1835. Trans. Wis. Acad.
39: 2145.

Hale, T. J. 1858. Additions to the flora of Wisconsin. Trans. Wis. State Agr. Soc. 5:
417424.

Hale T. J. 1860. Additions to the flora of Wisconsin. Trans. Wis. State Agr. Soc. 6: 258-263.
Pennell, F. W. 1935. The Scrophulariaceae of Eastern Temperate North America. Acad. Nat.
Sci. Phila. Monogr. 1. 650 p.

Editorial Notes

INDEX: The Michigan Botanist has published a three-year index at the end of Vol. 3 and of
Vol. 6, in the expectation that each three-volume unit would be a better size for binding. No
index is issued now for Vol. 7, but there will be another cumulative one at the end of Vol. 8
or 9.

The May number (Vol. 7, No. 3) was mailed May 25, 1968.

Program Notes

March 16, 1969: State membership meeting, Michigan Botanical Club.

May 23-25: State membership meeting and annual Spring Campout, Michigan Botanical
Club: near Irons, Lake Co., in the Manistee National Forest.

News Notes

PUBLICATIONS: Since it appeared in 1940, Deam’s Flora of Indiana has stood as one of
the finest state floras ever produced. Out of print and extremely scarce on the secondhand
market, it is now being reprinted by J. Cramer, Publisher (P.O. Box 48, D-3301 Lehre,
Germany). Subscription price is $20.00; after publication, the price will be $25.00. . . . The
Michigan Department of Conservation (now Michigan Department of Natural Resources),
due to lack of funds, is unable to reprint its excellent county maps. The complete books
have been out of print for a year; some individual counties are still available.

HONORS TO OUR FRIENDS: Genevieve Gillette, founder and president of the Michigan
Parks Association (and honorary member of the Michigan Botanical Club) was one of five
persons in the nation to receive an award this year for contributions to a “Beautiful Amer¬
ica” from Holiday Magazine, which described her as “conscience for the nation’s land¬
scape.” Michigan Governor George Romney presented the “golden tree” award to Miss
Gillette, who is also a member of President Johnson’s Citizens Advisory Committee on
Recreation and Natural Beauty. In 1966 she was named forest and park conservationist of
the year by the National Wildlife Federation and the Sears Roebuck Foundation in coop¬
eration with the Michigan United Conservation Clubs. . . . Dale J. Hagenah, research
associate with the Cranbrook Institute of Science, now has a fern named for him. Poly-
stichum x hagenahii, the hybrid between P. acrostichoides and P. lonchitis, was published by
W. J. Cody in the American Fern Journal for January-March 1968. . . . Dr. Harold B.
Tukey, Sr., who retired in 1962 as chairman of the Department of Horticulture at Michigan
State University -where he sat at Liberty Hyde Bailey’s old desk-received the Liberty Hyde
Bailey Medal of the American Horticultural Society for 1968. Also honored by the A.H.S.
was former Michigan Stater and Botanical Club member F. L. S. (‘Steve’) O’Rourke, now at
Colorado State University, who received a Citation from the Society. . . . Dr. Stanley A.
Cain returned in August to his duties at the University of Michigan as Professor of

Conservation and of Botany, after serving since February 28, 1965, as Assistant Secretary of
the Interior for fish, wildlife, and parks. . . . And the Michigan Botanical Club received the
award as conservation organization of the year at the annual convention of Michigan United
Conservation Clubs at Sault Ste. Marie in June. Past president Charles Barclay accepted the
award on behalf of the Club.

IN MEMORIAM: Shirley W. Allen, honorary member of the Botanical Club and an early
leader in the Michigan Natural Areas Council, died September 12, 1968, in Ann Arbor, at
the age of 85. Mr. Allen was professor emeritus of forestry at the University of Mich¬
igan. . . . Dr. John H. Ehlers died in Colorado November 17, 1967, two weeks after his
99th birthday. Dr. Ehlers taught systematic botany at the U. of M. Biological Station on
Douglas Lake from 1916 to 1938, in addition to his Ann Arbor teaching and service as
Curator of Phanerogams in the Herbarium, and was one of the most prolific of Michigan
plant collectors.

CONTENTS

Sex Determination in Mosses

Dale H. Vitt . 195

Mosquito Pollination of Habenaria obtusata (Orchidaceae)

Warren P. Stoutamire . 203

Studies on the Hydnums of Michigan. I.

Genera Phellodon, Bankera, Hydnellum

Kenneth A. Harrison . 212

Nature Education Feature-

Contact Dermatitis Caused by Plants

A. P. Ulbrich . 265

Additional Wisconsin Records for the Ebony
Spleen wort , A splenium platyneuron

Lytton J. Musselman . 268

The Milton College Herbarium

Lytton J. Musselman . 269

Editorial Notes . 271

Program Notes . 271

News Notes . 271

(On the cover: Sporophytes of the ‘bug-on-a-stick’ moss,
Buxbaumia aphylla Hedw. , photographed in an aspen
stand at the University of Michigan Biological Station
by Norton G. Miller, July 28, 1 968.)

HEHSHH

Vol. 8, No. 1

THE

MICHIGAN BOTANIST

THE MICHIGAN BOTANIST-Published by the Michigan Botanical Club four times per

year: January, March, May, and October. Second-class postage paid at Ann Arbor,

Michigan. © Michigan Botanical Club, 1969.

Subscriptions: $3.00 per year. Single copies: $.75.

All back issues are available, at the following prices per volume:

Vol. 1 (2 numbers, at $1.00 each): $2.00
Vol. 2-5 (4 numbers, at $.50 each): $2.00
Vol. 6- (4 numbers, at $.75 each): $3.00

Subscriptions (from those not members of the Michigan Botanical Club) and all orders
for back issues should be addressed to the business and circulation manager, who will as¬
sume that subscriptions received prior to the last number of a volume are intended to begin
with the first number of that volume unless specified To the contrary.

Articles and notes dealing with any phase of botany relating to the Upper Great Lakes
Region may be sent to the editor in chief. The attention of authors preparing manuscripts
is called to “Information for Authors” (Vol. 6, p. 202; reprints available from the editor).

Editorial Board

Edward G. Voss, Editor in Chief

Herbarium, The University of Michigan, Ann Arbor, Michigan 48104

Laura T. Roberts, Business and Circulation Manager
1509 Kearney Rd., Ann Arbor, Michigan 48104

Virginia L. Bailey Rogers McVaugh

Burton V. Barnes Alexander H. Smith

Richard Brewer W. H. Wagner, Jr.

THE MICHIGAN BOTANICAL CLUB

Membership in the Michigan Botanical Club is open to anyone interested in its aims:
conservation of all native plants; education of the public to appreciate and preserve plant
life; sponsorship of research and publication on the plant life of the State; sponsorship of
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sanctuaries and natural areas; and cooperation in programs concerned with the wise use and
conservation of all natural resources and scenic features.

Dues are modest, but vary slightly among the chapters and with different classes of
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may send $3.00 dues to the Membership Chairman listed below. In all cases, dues include
subscription to THE MICHIGAN BOTANIST. (Persons and institutions desiring to sub¬
scribe without becoming members should deal with the business and circulation manager as
stated at the top of this page.)

President: W. H. Wagner, Jr., Botanical Gardens, University of Michigan, Ann Arbor,
Michigan 48105

Vice President: William F. Hopkins, 6210 Cowell Rd., Brighton, Michigan 48116
Recording Secretary: Mary Cooley, 703 S. Forest, Ann Arbor, Michigan 48104
Corresponding Sec.: Margaret Buswell, 19204 Plainview, Detroit, Michigan 48219
Treasurer: Charles Buswell, 19204 Plainview, Detroit, Michigan 48219
Membership Chairman: Harriette V. Bartoo, Department of Biology, Western Michigan Uni¬
versity, Kalamazoo, Michigan 49001
Corresponding Secretary, Southeastern Chapter:

Catherine Hamilton, 9591 Grandmont, Detroit, Michigan 48227
Secretary, Huron Valley Chapter:

Ursula R. Freimarck, 704 Madison PI., Ann Arbor, Michigan 48103

1969

3

THE MICHIGAN BOTANIST

POLLEN ANALYSIS AT THE THALLER MASTODON
SITE, GRATIOT COUNTY, MICHIGAN

Edith R. Held and Ronald 0. Kapp
Luther College, Decorah, Iowa, and
Alma College, Alma, Michigan

INTRODUCTION

While digging an irrigation pond on his farm in June of 1965, Louis Thaller
discovered mastodon bones (SE/4, NE1/*, NE14, sec. 18, T 12 N, R 4 W, Gratiot
Co., Michigan). Excavation was undertaken by the landowner and a preliminary
study of the site and bones was made by biologists from Central Michigan
University (Mahan & Whitney, 1966). Samples of peat and wood were collected
by the junior author for radiocarbon dating and pollen analysis. The results of
this investigation permit the reconstruction of the vegetation associated with
Mammut americanum (Kerr), the American Mastodon, and are reported here.

The bones were embedded in gray sand immediately above the glacial till.
The remains include :

20 ribs
1 femur
1 patella
1 fibula

1 set of foot bones
1 humerus

1 ulna
7 vertebrae

1 complete hyoid apparatus

1 right (?) tusk (broken)

2 third molars

1 second molar

The molars were deeply worn with the cusps almost removed on the third
molars; the second molar was highly polished and deeply concave. The size of
the tusk and femur, the deeply worn molars, and the elevated and roughened
muscle and ligament insertions, all indicate that this was an old animal.

SITE DESCRIPTION

The depositional basin is elliptical, about 100 x 200 yards (90 x 180 m) in
extent, and oriented along an east-west axis. Peat occurs to a depth of 80 cm;
beneath a transitional zone, gyttja occurs from 100 to 105 cm; there is silty
gyttja from 105 to 135 cm, and sand from 135 to 150 cm (Fig. 1). Samples were
collected for C-14 dating from the same channel section from which pollen
analytical samples were taken.

The poorly-drained depression apparently contains no more than 2 meters
of accumulated sediments; it is surrounded by low sandy hills. Deposition is no
longer occurring but water stands in the center of the marsh during the spring
and early summer. The vegetation of the marsh is dominated by Carex, Typha,
various grasses, Eupatorium, and Verbena. Alnus rugosa, Spiraea alba, and sever¬
al species of willow ( Salix spp.) form the dominant shrub cover fringing the
marsh; occasional red maple (Acer rubrum ) and trembling aspen (Populus
tremuloides), and a single tamarack (Larix laricina), are present in the basin.

4

THE MICHIGAN BOTANIST

Vol.8

Second-growth and selectively cut woodlands near the excavation site,
and at Lumberjack Park in the same section, provide an indication of the
composition of the upland forests that surrounded the depositional basin before
settlement. Well-drained sandy sites are dominated by white pine ( Firms strobus),
with occasional hemlock ( Tsuga canadensis ), big-toothed aspen ( Populus
grandidentata) , and white oak ( Quercus alba); the understory is dominated by
witch hazel ( Hamamelis virginiana) and juneberry ( Amelanchier sp.). On more
poorly drained sites red maple {Acer rubrum) is dominant, with white pine and
paper birch {Betula papyrifera ) frequent; red oak (Q. borealis ), white oak,
swamp white oak ( Q . bicolor ), wild black cherry {Prunus serotina), sugar maple
(Acer saccharum), and hemlock are occasionally present. These observations and
previous studies of the presettlement forests in nearby Richland Twp., Montcalm
Co. (Hushen, et al., 1966), indicate that the area was originally heavily forested

O cm.

30 cml

60 cml

90 cml

120 cml

150 cml

■peati

gray

sand

AAA

till

7120 ±250 B.R

(wood)

72 20 - 250 B.P-

(wood)

7390± 280 B. P.

(pea))

11 200 - 400 B. P

(gyttja)

9910 ±350 B.P

(bone)

Fig. 1. Sedimentary sequence at the Thaller mastodon site. Left column shows depths of
the samples analyzed; C-14 dates are shown at the right at the appropriate depths.

1969

THE MICHIGAN BOTANIST

5

with white pine; mixed hardwood and hemlock forests occupied moist sites in
presettlement times.

METHODS

Samples were collected from the exposed face of the excavation at 5 cm
intervals to a depth of 150 cm. Samples for pollen analysis were placed in glass
vials in the field; the open vials were pushed into the freshly exposed sediments
to prevent contamination. Peat and wood samples were sealed in polyethylene
bags.

Calcareous samples were flooded with 6% HC1 to remove carbonates. All
samples for pollen analysis were processed according to the standard acetolysis
procedure (heated to boiling in 9:1 mixture of acetic anhydride and concen¬
trated sulfuric acid) and the concentrated residue was suspended in glycerine
jelly for storage and slide preparation.

Pollen counting was performed at 100 diameters magnification due to the
high percentage of pine and spruce; small or difficult entities were studied at 450
or 950 X. Counting was continued along regular traverses of the mechanical
stage until approximately 200 pollen grains (excluding sedges and aquatic plants)
had been tallied. At 140 cm a full count was not tallied because, possibly due to
a rapid rate of sedimentation, pollen grains were very scarce. At 145 and 150 cm
pollen was absent.

RESULTS

The results of pollen analysis are presented in the pollen diagram (Fig. 2),
which includes those entities which are consistently or abundantly represented
in the samples. Cyperaceae (sedge) and aquatic species, which are of restricted
distribution near the depositional site and are frequently overrepresented, are
excluded from the pollen sum. Percentage frequencies are based on the pollen
sum and are plotted for each stratigraphic level and then connected vertically to
give a pollen profile for each of the pollen types. Frequencies of Cyperaceae,
aquatic plants, and ferns are calculated as percentages of the pollen sum, even
though they are excluded from the sum. Percentages of infrequent pollen and
spore types are shown in Table I. Of particular interest in this table is the
occurrence of Tilia and Tsuga throughout the record and the identification of a
single grain of Linum (flax) pollen from the 140 cm level.

Samples of wood collected from the peat zone were identified by micro¬
scopic examination. Three types were recovered: Pinus strobus, Picea sp., and
Fraxinus sp.

Radiocarbon dates on samples of peat, wood, and mastodon bone (inner
tusk) are given in Fig. 1 .

DISCUSSION

There is a general trend from high (greater than 50%) to lower (less than
30%) spruce pollen in the record. This is similar to other sequences from Michi¬
gan which typically have an early TVcetf-dominated zone which began soon after
deglaciation and ended between 8,000 and 10,000 years B.P. (Andersen, 1954;

6

THE MICHIGAN BOTANIST

Vol. 8

Zumberge and Potzger, 1956; Gilliam, et al., 1967; Kapp, ms.).

Characteristic of the Spruce Pollen Zone are high percentages of Picea and
nonarboreal pollen (NAP). Abies is present only in this zone; this is consistent
with other late-glacial pollen records from Michigan. The Thaller diagram has
significant percentages of Ambrosia and Artemisia in the Spruce Pollen Zone;
these herbs, along with some of the composites (Compositae) and grasses
(Gramineae), probably occupied ecologically disturbed and open habitats within

the spruce forest.

The Pine Pollen Zone (Zone II, Fig. 2) is characterized by high percentages
of pine and fern spores and low percentages of spruce and NAP.

Typically, a Hardwood Pollen Zone follows the Pine Pollen Zone in pollen
diagrams from the region. At Vestaburg Bog, only 6 miles from the Thaller farm,
the Hardwood Pollen Zone begins about 8,000 B.P. and is characterized by high
percentages of Quercus and other hardwood pollen types. In the Thaller pollen
diagram the Hardwood Zone is subdued by local overrepresentation of pine and
spruce pollen. Black spruce ( Picea mariana ) apparently persisted after the begin¬
ning of the postglacial period at about 10,000 B.P. and probably increased in
cover; this is reflected in the increase in the spruce pollen percentages (from 45
to 5 cm). The high pine pollen frequency in Zone II is due to the abundance of
white pine on the surrounding hills; a similar forest composition persists to the
present at nearby Lumberjack Park (located 3A mile southwest of the site) and at
nearby Thaller’s woods. The postglacial persistence of spruce at the site, and the
local occurrence of white pine, is confirmed by the discovery of wood of these
trees in the peat zone.

Modern pollen rain studies from moss polsters in Thaller’s woods have
yielded pollen spectra with high pine and low hardwoods (Fig. 1, upper spectra).
Picea is much less prominent in the modern pollen rain when compared with the
period 7,300-7,100 B.P.; spruce trees are now very infrequent in this vicinity.
The stand of spruce in the Thaller peat bog must have disappeared subsequent to
the time that deposition ceased. Comparison of the modern and fossil pollen
spectra substantiates the foregoing interpretation of the Pine Pollen Zone and
explains the paucity of the hardwoods in the record.

Ecological succession gradually changed the site from lake to bog to a
seasonally wet marsh. The presence of Sagittaria and Nuphar in the lowest level
substantiates the existence of a small shallow lake in late-glacial time. The
change from lake to bog is inferred from the changing composition of the
sediments. Gray sand and gyttja immediately above the glacial till indicate the
presence of an alkaline lake; the gradual transition to peaty gyttja and subse¬
quently peat reflects the change to bog, and later marsh, conditions. A substan¬
tial increase in the frequency of fern spores in the upper zone is probably related
to increased dryness associated with this succession. Sediments stopped accumu¬
lating when the bog filled in completely about 7,000 years B.P.

Gilliam, et al. (1967) conclude that Tsuga migrated into Montcalm County
about 8,000 years B.P.; yet Tsuga was present in the Thaller farm about 10,000
B.P. (120 cm). Assuming that Tsuga could not have entered Michigan from the
south because of the presence of the prairie peninsula (Benninghoff, 1964),

1969

THE MICHIGAN BOTANIST

7

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1969

THE MICHIGAN BOTANIST

9

these central Michigan records suggest that hemlock entered the state from the
east or north rather soon after deglaciation. Fagus pollen occurred at 95 cm
(about 8,000 B.P.) and is consistent with other pollen diagrams from central
Michigan. Gilliam also suggests that Tilia is a more recent addition to the flora,
while the Thaller diagram indicates the presence of Tilia as early as 10,000
years B.P. (120 cm).

The radiocarbon dates of the peat, wood, and gyttja (30 cm, 7,120 ± 250
years B.P. on wood, M-1740; 60 cm, 7,220 ± 250 on wood, M-1741; 80 cm,
7,390 ± 280 on peat, M-1742; 130 cm, 11,200 ± 400 on marly gyttja, M-1743)
are in stratigraphic sequence and are accepted as accurate. Different rates of
sedimentation are indicated for the gyttja and peat, however. The gyttja (after
compaction) apparently accumulated at a rate of approximately 1.3 cm/ 100
years while about 18 cm of peat were forming per 100 years (Fig. 1).

The C-14 date on mastodon tusk (9,910 ± 350 B.P., M-1739) is open to
question since it lies stratigraphically beneath the gyttja, but dates younger. While
the possibility that the bones were intruded into the deeper sediments cannot be
conclusively disproved, because of the shallowness of the deposit and method of
excavation, we believe that they were covered in situ. The bones were embedded
in compact sand, lying on glacial till without evidence of peat or gyttja beside or
beneath them. In other instances radiocarbon dates on mastodon bones have been
shown to be too young. Skeels (1962) reported an instance of erroneous dating due
to the penetration of modern plant roots ;Kapp (ms.) concluded that mandible and
tusk dates for the Rappuhn mastodon were too young in an excavation where there
was good stratigraphic control and a consistent series of other radiocarbon dates.

These observations suggest that radiocarbon dates on mastodon bones
need careful scrutiny. Previous conclusions regarding the time of extinction (ca.
6,000 B.P.) are almost certainly too young and will require reinterpretation.
Although corroboration is necessary, we suggest that mastodon extinction oc¬
curred very soon after the beginning of postglacial time, probably in response to
climatic changes and alteration of the favorable late-glacial ecosystems. The role
of man (Martin, 1967) in these ecosystem alterations is uncertain.

This pollen record, and apparently all others in which the vegetation asso¬
ciated with mastodons has been analyzed, indicates that the American mastodon
was a member of glacial and late-glacial ecosystems in eastern North America.
This was a period of cool-moist climate. Pollen records from this period in the
Great Lakes region are dominated by spruce pollen, but the presence of hard¬
wood pollen probably indicates a variable mosaic of conifers and hardwoods. We
tend to accept the hypothesis of Brown and Cleland (1968) that the mastodon
required both a cool-moist climate and a plant community which included
woody angiosperms.

SUMMARY

The vegetational patterns associated with the Thaller Mastodon correspond with
existing Michigan pollen records, but with minor differences. The Spruce Pollen Zone ex¬
tended from deglaciation to about 8,000 years B.P. and the Pine Pollen Zone continued
until deposition ceased at about 7,000 B.P. The typical Hardwood Pollen Period is obscured
by local overrepresentation of pine and spruce pollen. Spruce, white pine, and ash wood

10

THE MICHIGAN BOTANIST

Vol. 8

were embedded in the peat which was deposited during the Pine Pollen Period at the site.
This explanation is confirmed in modem pollen rain studies from moss polsters near the site.

Radiocarbon dates of peat, wood, and gyttja range from 7,120 ± 250 B.P. to
11,200 ± 400 B.P. The date (9,910 ± 350 B.P.) on the bone is open to question because it
was lying in sand stratigraphically lower than the gyttja, yet the bone has a younger date.

ACKNOWLEDGMENTS

These investigations were supported by the National Science Foundation-Under¬
graduate Research Participation program (Grant GY-2871). Appreciation is extended to the
Michigan Memorial Phoenix Project Radiocarbon Laboratory for providing the C-14 dates.
Thanks are extended to Mr. and Mrs. Louis Thaller for their continuous cooperation. The
facilities of the Department of Biology, Alma College, were used throughout.

LITERATURE CITED

Anderson, Sv. Th. 1954. A late-glacial pollen diagram from southern Michigan, U.S.A. Dan-
marks Geol. Underst^gelse II (80): 140-155.

Benninghoff, W. S. 1964. The prairie peninsula as a filter barrier to postglacial plant migra¬
tion. Proc. Indiana Acad. 72: 116-124.

Brown, J., & C. Cleland. 1968. The late glacial and early postglacial faunal resources in
biomes newly opened to human adaptation. In The Quaternary of Illinois, a Sympo¬
sium (Univ. Ill., Coll. Agr. Spec. Publ. 14), pp. 114-122.

Gilliam, Jeanne A., R. O. Kapp, & R. D. Bogue. 1967. A post-Wisconsin pollen sequence
from Vestaburg Bog, Montcalm County, Michigan. Pap. Mich. Acad. 52: 3-17.

Hushen, T. W., R. O. Kapp, R. D. Bogue, & J. T. Worthington. 1966. Presettlement forest
patterns in Montcalm County, Michigan. Mich. Bot. 5: 192-211.

Kapp, R. O. (manuscript). Pollen and macrofossils associated with the Rappuhn Mastodon,
Lapeer County, Michigan.

Mahan, H., & M. Whitney. 1966. Preliminary report on the Thaller mastodon of Gratiot
County. (Unpublished report to Mich. Acad., 70th Annual Meeting.)

Martin, P. S. 1967. Pleistocene overkill, pp. 75-120 in Pleistocene Extinctions, ed. by P. S.

Martin & H. E. Wright, Jr. Yale Univ. Press, New Haven.

Skeels, M. A. 1962. The mastodons and mammoths of Michigan. Pap. Mich. Acad. 47:
101-132.

Zumberge, J. H., & J. F. Potzger. 1956. Late Wisconsin chronology of the Lake Michigan
basin correlated with pollen studies. Bull. Geol. Soc. Am. 67: 271-288.

Review

LIVING WITH YOUR LAND A Guide to Conservation for the City’s Fringe. By John
Vosburgh. Cranbrook Institute of Science (Bulletin 53), Bloomfield Hills, Michigan.
1968. 149 pp. $1.00 (plus tax).

Funds from the Garden Club of Michigan aided preparation of this excellent little
paperback, which no intelligent and concerned citizen should be without. The book is de¬
signed for suburban dwellers in the Northeast, Middle Atlantic, and Midwest states, with the
advice to them “not to dash ahead in a mad effort to conquer nature but to recognize the
character of the natural landscape and to bend your plans to it.” This comes as a refresh¬
ingly helpful and practical, surprisingly complete guide for dealing with native and introduced
plants and animals which may live on or visit one’s suburban premises. It suggests significant
forms of community action to maintain or restore natural beauty and to educate the public.
Appendices on where to find advice, help, and material are good but exceedingly short.
Michigan is occasionally cited as an example and the Huron-Clinton Metropolitan Authority
is specially mentioned.

-E. G. V.

1969

THE MICHIGAN BOTANIST

11

AN ECOLOGICAL ANALYSIS OF
A SOUTHERN MICHIGAN BOG

Garrett E. Crow

Department of Botany and Plant Pathology,

Michigan State University, East Lansing

INTRODUCTION

The term “bog” refers to a soil-vegetation complex made up of specialized
herbs, and trees growing on a wet, acid peat soil.

Gates (1942, p. 217) defines a bog as

... an area of vegetation, developing in undrained or poorly drained situations, which
by the development of a mat invading open water, forms a cover over a body of
water. It consists of a series of successional stages or associations beginning with
mat-forming sedges and passing through shrub (mostly ericad) and Sphagnum stages
to an expected culmination in the development of a characteristic coniferous tree
association.

Rigg (1916, p. 310) says that a Sphagnum bog is

. . . that stage in the physiographic succession of an area during which its surface is
entirely devoid of ordinary “hard” soil, and is composed completely of living
sphagnum moss under which is fibrous brown peat composed mainly or entirely of
partially decayed sphagnum.

A bog generally has a high water table, but usually there is little standing
water other than the associated bog lake. The upper peat and bog waters are
typically strongly acid.

Bogs occur in depressions and lowlands. Many form in kettle holes in
glacial moraines and in low spots in uplands. Bog conditions may develop in
quiet bays of large lakes or stream channels. They are characteristic of cold
regions with long winters and high rainfall. The shallow, little differentiated
podzols of these regions often support a coniferous forest. Dansereau (1957, p.
87) cites these regions as “. . . the zone of bogs, par excellence, with their typical
moss, sedge, and ericad flora: the tendency to peat accumulation is in evidence
everywhere that drainage is impeded.” The term “bog” is often loosely applied
to any peat-covered land of cold, northern regions. However, bogs become scat¬
tered and less numerous as the major vegetational units of North America pass
from the northern conifer forest, toward the glacial boundary, through the
hemlock— white pine— northern hardwood tension zone, to the eastern deciduous
forest.

Transeau (1905) noted that peat deposits of considerable extent occur
occasionally south of the limit of Wisconsin glaciation. Many of these are be¬
lieved to be associated with earlier glacial advances of the Pleistocene.

Bog formations associated with former localized glaciation are also known
in the mountains of the eastern and western United States.

Other situations in which there is considerable peat accumulation are asso¬
ciated with coastal plain phenomena of eustatic movement, irregular depositions

12

THE MICHIGAN BOTANIST

Vol. 8

of alluvial materials in deltas, and reef building causing extension of the land. In
North America the greatest development of the southern bog condition is in
eastern Virginia and North Carolina, Florida, and the Mississippi floodplain. The
vegetation in these habitats is noted for its density and luxuriance and this
combined with stagnant water allows peat formation in spite of high tempera¬
tures (Transeau, 1905).

Although these southern peat deposits are sometimes referred to as
“bogs,” they have a very different appearance from those to the north. In
comparing the vascular plants of two sphagnous wetlands in New Jersey, Mont¬
gomery and Fairbrothers (1963) noted that the northern bog, located within the
glacial boundary, had a floating mat encroaching upon open water and the peat
was deep. The southern bog, on the other hand, located on the unglaciated
coastal plain at the southern tip of New Jersey, had no floating mat and the peat
deposit was shallow. The northern bog reflected a dominance of woody species
in contrast to a chiefly herbaceous flora in the southern bog. The northern bog
had a constant water level and continuous Sphagnum distribution, whereas the
southern bog was characterized by fluctuating seasonal water levels and a scat¬
tered Sphagnum distribution.

Despite their differences, northern and southern bogs have a strong physio¬
logical similarity. Factors such as acidity, peaty soils, high water table, and the
presence of a number of species with otherwise northern distributions suggest
the possibility of southern bogs having served as refugia during Pleistocene times.

DESCRIPTION OF PENNFIELD BOG

The bog which is the subject of this study is located in Calhoun County,
Michigan, 1% miles north of Pennfield and 7 miles northeast of Battle Creek.
Pennfield Bog, as it will be referred to here, is associated with two adjacent
lakes, Goose Lake and Little Goose Lake, each of which is 4 acres in area.
During years of high water table the lakes are connected by a narrow channel of
water filled with Nuphar luteum and Nymphaea tuberosa. They appear as two
distinct lakes during years of low water table (as indicated by an aerial photo¬
graph made in 1961). The bog possesses all the features necessary to fulfill the
bog definitions given by Rigg (1916) and Gates (1942).

Little Goose Lake, now owned and set aside as a plant preserve by the
Eastern Michigan Nature Association, has had the greatest amount of bog devel¬
opment, being completely surrounded by a floating mat. This mat is most exten¬
sive to the northwest, the direction of prevailing winds.

Goose Lake has a floating mat on the northwest side, but although the
lake shore is completely organic, there is virtually no floating mat on the south¬
east side of the lake. The only outlet of the bog is at the east corner of Goose
Lake. It probably is functional only during periods of high water table. The
shallow outlet runs a short distance into Deep Lake, a 7-acre lake also with a
completely organic shore (Humphrys & Green, 1962).

Pennfield Bog, in one of the numerous depressions of the Kalamazoo end
moraine of Wisconsin age, is at an elevation of 860 feet above sea level. The
glacial till of the morainal system is generally sandy and is underlain by a

1969

THE MICHIGAN BOTANIST

13

bedrock of sandstone (Leverett and Taylor, 1915). The glacial ice flow in the
locality was that of the Saginaw ice lobe progressing from a northeast to a
southwesterly direction (Flint, et ah, 1959).

The morainal ridges surrounding the study area support various vegetation
types. The northernmost ridge supports an oak woods and a small reforestation
of red pine; the northwest side is pasture; the southwestern and southern border
is an old orchard returning to an oak woods; the northeast ridge is a black
oak-red maple woods.

CLIMATE

The climate of the region of Pennfield Bog is modified considerably by
Lake Michigan, about 65 miles to the west. When there is little or no wind the
climate is continental, but when there are strong winds from the Lake the
weather becomes semimarine. Winter winds are warmed in transit across Lake
Michigan and summer winds are cooled, thus creating a more moderate climate
than that on the western side of the Lake. Spring temperatures tend to be
depressed while fall temperatures are elevated (Eichmeier, n.d.).

Weather data for the Battle Creek area show the highest temperature re¬
corded was 104°F in July 1936 and the lowest was -24° in February 1899. Days
with 90° temperatures occur on an average of 20 per summer and the mean
temperature of January, the coldest month, is 1 1.4°. The Climatological Survey
(Eichmeier, n.d.) for the period 1926-1955 at the nearest weather station at
Battle Creek shows the mean daily maximum temperature to be 58.6° for the
year, while the mean daily minimum temperature is 38.8°. The warmest month
is July with a mean temperature of 85.1°. The average dates of the last freezing
temperature of spring and the first freezing temperature of fall are May 8 and
October 9, respectively. In the fall of 1967 signs of frost were in evidence at
Pennfield Bog about 20 days earlier than the average date recorded at the nearest
station.

Precipitation averages 46% greater for the months May-October than for
the rest of the year. May and June are the peak months with heaviest amounts of
rainfall due to thundershower activity. Average precipitation ranges from a high
of 4.04 inches in May to 1 .73 inches in February with the annual mean at 33.43
inches.

Snowfall averages 39.8 inches during Battle Creek winters. The first snow¬
fall usually occurs in November and the last in April, with an average of 57 days
with snow on the ground.

Sunshine is only 30% of the possible total during January, while in the
June-August period sunshine reaches about 70% of the possible amount.

Relative humidity averages of the summer season range from 78% in early
morning to 50% in early afternoon while winter averages are 85%-75%.

WATER AND SOIL

The open lake waters of Pennfield Bog are neutral in pH (6.9) and are rich
in planktonic organisms. The water is stagnant and abounds with colloids. The
shore is 100% organic and the substrate of the entire bog is made up of peat. The

14

THE MICHIGAN BOTANIST

Vol. 8

waters of the bog mat are acid (see Table 1).

Transeau (1906) considered putrefaction to be the chief process of reduc¬
tion by which disintegration is carried on by anaerobic organisms and is favored
by low temperatures, presence of acids, and low oxygen availability due to high
water table. Products of putrefaction are of little use as nutrient materials for
higher plants and nitrogen remains unavailable to plants.

METHODS

Field studies were made at Pennfield Bog from mid-April to mid-October,
1967. The bog was visited once each week during spring and fall and two
collection and study trips a week were made during the summer months. Vouch¬
er specimens for all species reported from the bog are deposited in the Michigan
State University Herbarium. Collections were also made of representative species
in the major vegetation types surrounding the bog.

Quantitative data were collected during August to support recognition of
the several vegetational societies. Two transects were run in the bog. The long
axis ran at 16° west of magnetic north and was 660 meters in length. The
shorter, perpendicular axis was 480.5 meters long. Because of the structural
nature of the habitat two kinds of sampling procedures were employed. The
herb and subshrub societies were sampled by quadrats one-half meter square. In

TABLE 1. Average pH values in Pennfield Bog.

Zone

pH value

Open water

6.9

Nuphar-Eleocharis

Surface water

7.15

Submerged mat

6.25

Decodon

Surface water

6.93

Submerged mat

6.5

Carex-Vaccinium macrocarpon

6.03

Depressions

6.8

Sphagnum hummocks

4.7

Cassandra

4.76

Larix

Vaccinium corymbosum understory

4.5

Deep Sphagnum mat

4.18

Larix-Salix subzone

6.07

Acer rubrum

4.81

Openings at outer edge

6.42

1969

THE MICHIGAN BOTANIST

15

the tree communities quadrats of the same size were used to examine the ground
cover, while point intercepts were used to survey canopy and understory shrub
and tree species (nearest species to quadrat recorded). One hundred quadrats
were distributed (as determined by a table of random numbers) along each
transect. Point intercepts recorded in the Larix and Acer mbrum zones were at
the same locations as the quadrats.

In each quadrat the species and their approximate percent cover were
recorded. Species with little significant cover were recorded as having 1% cover.
Frequency, cover, and importance values were then tabulated for each vegetation-
al zone. Determination of importance values follows the method of Beaman and
Andresen (1966) which is a summation of percent total frequency and percent
total cover data, thus giving a summation constant of 200.

A Beckman Pocket pH meter (model 180) was used to obtain pH values of
surface soil and water in the major plant communities. The total of 53 readings
was taken on August 3 1 .

VEGETATION

The bog consists of seven major plant zones: a Nuphar-Eleocharis zone, a
Decodon zone, a Carex-V actinium macrocarpon zone, a Cassandra zone, a
Cassandra-Thelypteris zone, a Larix zone, and an outermost Acer rubrum zone.
The bog community, undergoing the natural processes of succession, is slowly
encroaching upon the open water of Goose and Little Goose Lakes. The general
aspect of the bog can be seen in Figs. 2 and 3. Vegetational zones are shown in
Fig. 1 .

A number of species occur in nearby open water just off the bog mat and
are floating or submerged aquatics. The most conspicuous species include
Ceratophyllum demersum, Myriophyllum verticillatum, Nymphaea tuberosa,
Brasenia schreberi, Utricularia vulgaris, U. purpurea, Potamogeton illinoensis, P.
natans, and P. gramineus.

THE NUPHAR-ELEOCHARIS ZONE (TABLE 2)

The rhizomes and fibrous roots of Eleocharis smallii and the stout
rhizomes of Nuphar luteum provide the major network for the submerged mat
of this zone which is the forerunner of the quaking bog mat.

The dominant species, Eleocharis smallii and Nuphar luteum, are peren¬
nial, rhizomatous aquatics. Other important species which contribute to the
structure of the substrate include Pontederia cordata, Nymphaea tuberosa,
Potentilla palustris, and Carex lasiocarpa. A number of floating and submerged
aquatic species are also present, but have little importance in terms of construc¬
tion of the bog mat.

THE DECODON ZONE (TABLE 3)

Another pioneer zone is the Decodon mat which has developed on the
west shores of Little Goose Lake and Goose Lake. Like the Nuphar-Eleocharis
zone, the Decodon zone is characterized by a submerged mat. The surface water
of the zone is nearly neutral while the submerged mat is slightly acid. The mat is

16

THE MICHIGAN BOTANIST

Vol. 8

Decodon zone

Carex-Voccinium macrocarpon zone

Cassandra zone

Cassandra-Thelvpteris zone

Open Larix subzone

Larix zone

Larix-Salix subzone

Acer rubrum zone

Vaccinium corvmbosum subzone

Transect

Deep Lake

Fig. 1. Vegetation map of Pennfield Bog showing vegetation zones and subzones and tran¬
sect locations. Adapted from an aerial photograph (BDF-5AA-102 by USDA Agricultural
Stabilization and Conservation Service) enlarged with a Saltzman’s projector.

1969

THE MICHIGAN BOTANIST

17

Fig. 2. Oblique aerial view of Little Goose Lake encircled by bog mat (view looking north¬
east). October 22, 1967.

Fig. 3. Decodon mat encroaching upon Little Goose Lake (view looking northwest). July
22, 1967.

18

THE MICHIGAN BOTANIST

Vol.8

TABLE 2. Frequency, cover, and importance values of 21 species encountered in 19 quad¬
rats in the Nuphar-Eleocharis zone.

Species

%

Frequency

%

Cover

Importance

Value

Nuphar luteum

73.68

40.05

47.26

Eleocharis smallii

89.47

28.58

40.73

Potamogeton gramineus

63.15

11.79

21.79

Nymphaea tuberosa

42.10

16.32

21.55

Pontederia cordata

47.36

3.84

12.18

Utricularia vulgaris

36.84

6.31

12.24

Brasenia schreberi

21.05

2.16

5.79

Potentilla palustris

21.05

2.16

5.79

Potamogeton illinoensis

21.05

1.05

5.26

Car ex lasiocarpa

10.52

2.37

3.95

Sparganium minimum

15.78

0.79

3.64

Sparganium chlorocarpum

15.78

0.68

3.55

Lemna minor

15.78

0.16

3.12

Decodon verticillatus

5.26

2.37

2.95

Dulichium arundinaceum

10.52

0.26

2.20

Sagittaria latifolia

10.52

0.16

2.12

Cassandra calyculata

5.26

0.53

1.43

Triadenum virginicum

5.26

0.21

1.16

Lemna trisulca

5.26

0.05

1.03

Sium suave

5.26

0.05

1.03

Utricularia purpurea

5.26

0.05

1.03

a quaking structure and will occasionally break under the weight of a person.
Decodon verticillatus is the dominant species, comprising 23.58% of the total
cover. Other important species include Cassandra calyculata, Typha latifolia,
Nuphar luteum, Eleocharis smallii, and Thelypteris palustris, all of which are
rhizomatous herbs except the shrubby C. calyculata.

The majority of plants are emergent aquatics. They give a hummock-like
aspect to the mat where the dense growth— especially that of Decodon verticil¬
latus, with its thickened, spongy stem base— collects some litter and provides a
microhabitat for bryophytes and such angiosperms as Drosera rotundifolia,
Campanula aparinoides, Ly copus uniflorus, and Viola nephrophylla.

Growth of the mat is enhanced by the ability of Decodon verticillatus to
root freely at the tips when the arching branches touch the water.

THE CAREX- VA CCINIUM MACROCARPON ZONE (TABLE 4)

The establishment of the floating mat requires a dense community of
sedges around the lake margin. The rhizomes of the sedges grow out into the
water behind the Nuphar-Eleocharis zone, forming an interlacing of roots and
rhizomes, buoyed up by the water. As the mat encroaches upon the open water
it loses contact with the bottom.

The sedge mat is dominated by three clonal species: Care x lasiocarpa,
Vaccinium macrocarpon, and Thelypteris palustris. The table indicates that
Vaccinium macrocarpon is the most important species of the zone, but if an

1969

THE MICHIGAN BOTANIST

19

TABLE 3. Frequency, cover, and importance values of 25 species encountered in 9 quadrats
in the Decodon zone.

Species

%%

Frequency

%

Cover

Importance

Value

Decodon verticillatus

88.88

25.00

37.37

Cassandra calyculata

33.33

12.22

16.69

Brasenia schreberi

22.22

13.89

16.54

Typha latifolia

33.33

11.67

16.17

Nuphar luteum

22.22

11.11

13.92

Eleocharis smallii

44.44

6.11

12.65

Thelypteris palustris

44.44

5.67

12.23

Ly copus uniflorus

55.55

0.56

9.14

Osmunda regalis

11.11

6.67

8.01

Utricularia vulgaris

33.33

1.56

6.64

Spiraea to men to sa

22.22

3.11

6.37

Potentilla palustris

22.22

2.56

5.85

Sium suave

33.33

0.33

5.48

Triadenum virginicum

22.22

2.00

5.32

Sagittaria latifolia

22.22

1.56

4.91

Carex comosa

22.22

0.22

3.64

Sparganium minimum

22.22

0.22

3.64

Salix petiolaris

11.11

0.56

2.24

Dulichium arundinaceum

11.11

0.22

1.92

Pontederia cor data

11.11

0.22

1.92

Campanula aparinoides

11.11

0.11

1.82

Drosera rotundifolia

11.11

0.11

1.82

Epilobium strictum

11.11

0.11

1.82

Vaccinium macrocarpon

11.11

0.11

1.82

Viola nephrophylla

11.11

0.11

1.82

importance value could be determined from a summation of relative frequency,
cover, and extensiveness of root system, Carex lasiocarpa would certainly be
primary in importance, with Vaccinium macrocarpon the secondary species. The
importance value of Cassandra calyculata is valid where the zone is relatively
narrow, but at the north end of the bog where the development of the sedge mat
is extensive, the species occurs almost exclusively where the sedge mat borders
on the Cassandra zone.

The mat surface consists of small rises and pocket-like depressions. In early
spring the mat is covered with water to a depth of approximately 30 cm. By
mid-summer the water subsides, leaving only shallow pools in the depressions.
Sphagnum, when it is present, occurs on the rises and the pH is an average 4.7 in
these places. The pH is nearly neutral in the depressions, averaging 6.8, and the
entire zone has an average pH of 6.03. During late summer the aerobic decay
process is enhanced by higher temperatures, near-neutral pH, and lowered water
table. Some decay is apparent in the wet depressions. Only two species, Utri-
cularia intermedia and Drosera intermedia, were observed to inhabit these de¬
pressions, to which, in fact, they were restricted.

The greatest diversity of species apparently occurs in this zone, but most
have very low importance values.

20

THE MICHIGAN BOTANIST

Vol. 8

TABLE 4. Frequency, cover, and importance values of 32 species encountered in 1 8 quadrats
and point intercepts in the Car ex-V actinium macrocarpon zone.

Species

%

Frequency

%

Cover

Importance

Value

Quadrats

V actinium macrocarpon

88.88

47.22

43.64

Carex lasiocarpa

94.44

32.83

34.44

Thelypteris palustris

55.55

13.17

16.04

Cassandra calyculata

55.55

10.22

14.01

Triadenum virginicum

55.55

4.28

9.93

Potentilla palustris

38.88

3.94

8.83

Andromeda glaucophylla

22.22

4.44

5.84

Nuphar luteum

16.66

5.00

5.52

Sium suave

38.88

0.61

5.30

Typha latifolia

22.22

3.61

5.18

Ly copus uniflorus

33.33

1.22

5.02

Salix pedicellaris

27.77

1.39

4.44

Sagittaria latifolia

27.77

0.78

4.02

Sarracenia purpurea

22.22

1.11

3.55

Sphagnum spp.

11.11

2.83

3.33

Eleocharis smallii

11.11

2.50

3.10

Potamogeton gramineus

16.66

1.06

2.81

Onoclea sensibilis

16.66

0.39

2.35

Utricularia intermedia

16.66

0.39

2.35

Hypericum boreale

11.11

1.39

2.34

Carex comosa

11.11

1.22

2.22

Campanula aparinoides

16.66

0.17

2.20

Boehmeria cylindrica

11.11

1.11

2.15

Toxicodendron vernix

5.55

1.67

1.83

Spiraea tomentosa

11.11

0.72

1.81

Bide ns corona ta

11.11

0.55

1.76

Scutellaria galericulata

11.11

0.28

1.58

Drosera rotundi folia

11.11

0.11

1.46

Carex aquatilis

5.55

0.56

1.07

Drosera intermedia

5.55

0.28

0.88

Dulichium arundimceum

5.55

0.06

0.73

Point intercepts

Larix laricina

11.11

TABLE 5. Frequency, cover, and importance values of 6 species encountered in 4 quadrats
in the Cassandra zone on the east side of Little Goose Lake.

Species

%

Frequency

%

Cover

Importance

Value

Sphagnum spp.

100.00

87.50

77.31

Cassandra calyculata

100.00

71.25

67.60

Vaccinium macrocarpon

75.00

5.25

21.88

Drosera rotundifolia

75.00

0.75

19.19

Typha latifolia

25.00

1.25

6.99

Sarracenia purpurea

25.00

1.25

6.99

1969

THE MICHIGAN BOTANIST

21

THE CASSANDRA ZONE (TABLE 5)

The Cassandra zone succeeding the Carex-Vaccinium macrocarpon zone is
characterized by a dense growth of this low shrub, with Sphagnum forming a
cushion about the stems and branches. The substrate of the zone is very acid,
having an average pH value of 4.7.

The portion of the zone on the east side of Little Goose Lake is represent¬
ative of what might be considered a “typical” Cassandra zone. The zone is
dominated by C. calyculata with a deep cushion of Sphagnum which grows up
around the base of the shrubs. The number of species present is small.

THE CASSANDRA-THELYPTERIS ZONE (TABLE 6)

In the northern part of the bog the Casstf^dra-dominated zone takes on a
quite different appearance. Although C. calyculata is the most important species
in this very wet zone, Thelypteris palustris and Typha latifolia nearly equal it in
importance; thus the zone is referred to as the Cassandra-Thelypteris zone. The
zone has become a tension zone, being bordered on the south by the Nuphar-
Eleocharis zone, on the west and north by the Larix zone, on the northeast by
the Larix-Salix subzone, and on the east by the Carex-Vaccinium macrocarpon
zone. Because of the very wet nature of the zone, Sphagnum has a much lower
importance value in this portion of it, making up only 12.83% of the total cover.

This part of the bog is very diverse in number of species, and there are two
distinct areas of subdivision. The column heading “Subzone A” in Table 6 refers
to the area to the immediate north of Little Goose Lake (see Lig. 1). Cassandra
calyculata , Thelypteris palustris, and Typha latifolia are still the dominant
species, but Sphagnum is more prevalent in this area and makes up 24.08% of
the total cover, second only to C. calyculata.

The area farther north along the transect is very wet and scattered clumps
of Larix are numerous. The data for this area in Table 6 are headed “Subzone
B.” In this transitional area the dominant species are again Cassandra calyculata,
Thelypteris palustris, and Typha latifolia. Sphagnum, however, is much less im¬
portant, making up only 4.56% of the total cover. The clumps of Larix provide a
niche for some species that would not otherwise be present in the zone, such as
Menyanthes trifoliata, Calla palustris, Cypripedium acaule, Maianthemum
canadense, Carex aquatilis, and Rosa palustris.

Data for both areas are totaled in a third column in Table 6. Although the
frequency of Larix is nearly the same in both areas, the trees in subzone A are
often single in occurrence while the trees mostly occur in clumps in subzone B.
Lifty percent of the species in the zone are not duplicated in the two areas, and
those of subzone B have affinities with the Larix zone and the Larix-Salix
subzone.

THE LARIX ZONE (TABLES 7 & 8)

Larix forms a very extensive zone encircling the bog and dominates the stage
of succession replacing the Cassandra zone. The zone can be divided into three
subzones: the openL^ra: subzone, the Larix-Salix subzone, and the closed Larix
bog forest.

TABLE 6. Frequency, cover, and importance values of 32 species encountered in 43 quadrats and point intercepts in the Cassandra-Thelypteris
subzone A (15 quadrats) and subzone B (28 quadrats).

22

THE MICHIGAN BOTANIST

Vol. 8

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1969

THE MICHIGAN BOTANIST

23

TABLE 7. Frequency, cover, and importance values of 24 species encountered in 17 quad¬
rats and point intercepts on the deep Sphagnum mat associated with the open Larix subzone
of the west side.

Species

%

Frequency

%

Cover

Importance

Value

Quadrats

Sphagnum magellanicum

88.23

80.59

69.12

Thelypteris palustris

47.05

17.18

20.45

Vaccinium macrocarpon

58.82

8.65

17.40

Cassandra calyculata

47.05

9.52

14.11

Andromeda glaucophylla

29.41

8.53

11.37

Menyanthes trifoliata

23.52

4.76

9.25

Carex lasiocarpa

29.41

4.88

9.05

Osmunda regalis

11.76

6.76

6.68

Typha la ti folia

17.64

3.57

6.37

Eriophorum virginicum

17.64

3.06

5.51

Carex interior

11.76

4.41

5.18

Pogonia ophioglossoides

17.64

0.29

3.75

Potentilla palustris

11.76

0.94

2.97

Vaccinium corymbosum

11.76

0.94

2.97

Acer rubrum

11.76

0.53

2.71

Campanula aparinoides

11.76

0.12

2.45

Sagittaria lati folia

5.88

0.47

1.48

Sarracenia purpurea

5.88

0.47

1.48

Salix petiolaris

5.88

0.24

1.34

Potamogeton gramineus

5.88

0.12

1.26

Triadenum virginicum

5.88

0.12

1.26

Ly copus uniflorus

5.88

0.06

1.22

Scutellaria galericulata

5.88

0.06

1.22

Trientalis borealis

5.88

0.06

1.22

Point intercepts

Larix laricina

Toxicodendron vemix

23.52

17.64

The younger stages of the zone are characterized by an open bog forest.
This has a general appearance somewhat suggestive of savanna, with the fre¬
quency of Larix laricina being 23.52% and Toxicodendron vemix occurring with
a frequency of 17.64%. There is a deep, luxuriant mat of Sphagnum magellani-
cum carpeting the zone.

In Table 7, data are tabulated for the open portion of the Larix zone. The
mat is acid, having an average pH value of 4.18. The most acid reading, pH 3.7,
was taken on this Sphagnum mat.

The Larix -Salix subzone occurs in the northern part of the bog. Since
neither transect passed through this area, no quantitative data were recorded.
The subzone is, however, easily recognized. Larix laricina is the most conspicu¬
ous tree species and Salix bebbiana the most abundant shrub. The area is very
wet and takes on the aspect of a young swamp forest with tall shrubs and small
trees with hummocks formed at the base. The average pH is 6.07. A number of
the species present are common in swampy habitats. These include Cornus

24

THE MICHIGAN BOTANIST

Vol. 8

stolonifera, Betula pumila, Rosa palustris, Ribes americanum, R. hirtellum, Salix
serissima, Bidens coronata, B. cernua, and Scirpus acutus.

A number of years ago a path was constructed by dragging logs through
the Larix-Salix subzone to provide easy access for cranberry pickers. The path
now consists of very loosely consolidated peat and is under water most of the

TABLE 8. Frequency, cover, and importance values of 28 species encountered in 36 quad¬
rats and point intercepts in the mature Larix zone.

Species

%

Frequency

%

Cover

Importance

Value

Quadrats

Sphagnum spp.

72.22

39.67

81.58

Cassandra calyculata

16.66

7.36

16.16

Vaccinium corymbosum

27.77

4.91

16.04

Osmunda regalis

5.55

5.28

9.57

Acer rubrum

27.77

0.44

9.42

Car ex interior

16.66

2.50

8.90

Maianthemum canadense

19.44

0.44

6.79

Trientalis borealis

16.66

0.50

6.00

Carex trisperma

16.66

0.17

5.51

Sarracenia purpurea

11.11

3.51

4.82

Ilex verticillata

8.33

0.83

3.85

Cypripedium acaule

11.11

0.22

3.83

Ly copus uniflorus

8.33

0.36

3.16

Thelypteris palustris

8.33

0.28

3.04

Rubus hispidus

8.33

0.08

2.74

Rosa palustris

2.77

1.11

2.51

Salix pedicillaris

2.77

1.11

2.51

Quercus velutina

5.55

0.17

2.00

Vaccinium macrocarpon

5.55

0.17

2.00

Carex aquatilis

2.77

0.56

1.69

Typha lad folia

2.77

0.14

1.07

Menyanthes trifoliata

2.77

0.06

0.95

Viola affinis

2.77

0.06

0.95

Woodwardia virginica

2.77

0.06

0.95

Drosera rotundifolia

2.77

0.03

0.91

Habenaria ciliaris

2.77

0.03

0.91

Monotropa uniflora

2.77

0.03

0.91

Rhynchospora alba

2.77

0.03

0.91

Point intercepts

Vaccinium corymbosum

86.11

Larix laricina

72.22

Nemopanthus mucronata

13.88

Acer rubrum

8.33

Ilex verticillata

8 33

Quercus velutina

5.55

Toxicodendron vernix

5 55

Nyssa sylvadca

2.77

Salix pedolaris

2.77

Ulmus rubra

2,77

1969

THE MICHIGAN BOTANIST

25

summer. By the end of the summer, when the water table is much lower, the
path takes on a consistency of a very wet muck. A few plants were recorded
from along this path and nowhere else in the bog. Among these are Ranunculus
sceleratus, Rumex orbiculatus, Scirpus acutus, and Boehmeria cylindrica (shade
form).

The mature Larix zone is a closed bog forest where Larix laricina has a
frequency of 72.22%. It has a very dense understory with Vaccinium corymbo-
sum being the dominant shrub. The data for the ground cover are given in Table
8. As the table indicates, there is very little ground cover. Sphagnum is by far the
most frequent plant, but has a cover value of only 39.67%. The remaining
species provide very little cover. Most of the species listed in Table 8 occur near
the margin of the zone.

The zone is acid with an average pH value of 4.5.

THE ACER RUBRUM ZONE (TABLES 9 & 10)

The climax stage of the bog forest encircles the Larix zone and is dom¬
inated by Acer rubrum, which has a frequency of 65.78%. Other tree species are
infrequent, as indicated by the low incidence of point intercepts. The frequen¬
cies of Nyssa sylvatica and Pinus strobus are inflated. Nyssa sylvatica was located
in only two stations, both of which occurred on the transect line. Only two tall
trees and two saplings of Pinus strobus were noted in the bog.

The understory is dominated by Ilex verticillata and Vaccinium corymbo-
sum. Together the two species have a frequency of 73.68%. Nemopanthus
mucronata is also frequent in the zone and should, perhaps, have a higher per¬
cent frequency than is indicated in Table 9. Toxicodendron vernix occurs in this
zone also, but tends to frequent places where the shade is light.

The Acer rubrum zone has a much denser ground cover than the Larix
zone. The predominant species of Sphagnum is S. fimbriatum, which forms
clumps rather than a deep, continuous cushion as does S. magellanicum in the
Larix zone. Maianthemum canadense is the most frequent herb and Ilex verticil¬
lata is the most frequent woody seedling. Osmunda cinnamomea is also an
important species of the zone and is abundant in very wet places.

Along the outer margin of the bog there are numerous openings in the
Acer rubrum zone. The species present in these openings tend to be typical of
swampy sites. These include such species as Viburnum lentago, Cornus stoloni-
fera, Corylus americana, Echinocystis lobata, Vitis vulpina, Leersia oryzoides,
Glyceria striata. Aster umbellatus, Urtica dioica, Bidens cemua, B. coronata,
Impatiens biflora, and Alisma plantago-aquatica. The pH of these openings aver¬
ages 6.42 while the pH of the Acer rubrum zone averages 4.81 .

On the west side of Little Goose Lake there is a large Vaccinium corymbo-
sum subzone associated with the Acer rubrum zone. Vaccinium corymbosum,
with a 100% frequency value, forms a very dense thicket. A ronia melanocarpa is
also frequent, occurring at one-third of the point intercepts. A cer rubrum occurs
with a frequency of 22.72% and the trees are mostly very large and quite
scattered. They cast very little shade on the area. There are very few ground-
cover species and these provide very little cover. Sphagnum attains its highest

26

THE MICHIGAN BOTANIST

Vol. 8

TABLE 9. Frequency, cover and importance values of 23 species encountered in 38 quad¬
rats and point intercepts in the Acer rubrum zone.

Species

%

Frequency

%

Cover

Importance

Value

Quadrats

Sphagnum spp.

42.10

13.95

53.07

Maianthemum canadense

85.71

5.60

40.84

Ilex verticillata

85.71

3.87

35.72

Osmunda cinnamomea

15.78

5.24

19.97

Trientalis borealis

28.94

1.50

12.62

Ly copus uniflorus

26.31

0.47

8.84

R ubus hispidus

18.42

0.58

6.92

Vaccinium corymbosum

7.89

0.92

4.94

Acer rubrum

5.26

0.71

3.59

Thelypteris palustris

7.89

0.18

2.76

Carex trisperma

7.89

0.08

2.46

Galium trifidum

5.26

1.49

1.63

Woodwardia virginica

2.63

0.18

1.27

Cypripedium acaule

2.63

0.13

1.12

Rosa palustris

2.63

0.13

1.12

Toxicodendron vernix

2.63

0.13

1.12

Quercus velutina

2.63

0.05

0.88

Mitchella repens

2.63

0.03

0.82

Point intercepts

Acer rubrum

65.78

Ilex verticillata

39.47

Vaccinium corymbosum

34.21

Nemopanthus mucronata

7.89

Nyssa sylvatica

5.26

Pinus strobus

5.26

Aronia melanocarpa

2.63

Betula lutea

2.63

Quercus velutina

2.63

Toxicodendron vernix

2.63

importance value here since it has high frequency and cover values and since few
other species contribute to the ground cover. It is interesting to note that no
Acer rubrum seedlings were encountered in the quadrats. Data for this subzone
are given in Table 10.

ACKNOWLEDGMENTS

This paper represents the first portion of a thesis submitted to Michigan State Univer¬
sity in partial fulfillment of the requirements for the degree of Master of Science; a phyto¬
geographic analysis will follow.

I wish to express my sincere appreciation to my major professor, Dr. John H.
Beaman, for his guidance and encouragement during this study. I also appreciate the helpful
suggestions concerning the thesis provided by Drs. William B. Drew and Stephen N.
Stephenson and those on phytogeography by Dr. Hugh H. litis. Mr. Norton G. Miller made
determinations of several bryophytes and Mr. Loren Perry flew me over the bog.

1969

THE MICHIGAN BOTANIST

27

TABLE 10. Frequency, cover, and importance values of 10 species encountered in 22 quad¬
rats and point intercepts in the Vaccinium corymbosum subzone associated with the Acer
rubrum zone on the west side.

Species

%

Frequency

%

Cover

Importance

Value

Quadrats

Sphagnum spp.

77.27

52.73

116.51

Rubus hispidus

31.81

14.89

19.38

Maianthemum canademe

36.36

1.45

19.22

Vaccinium corymbosum

18.18

4.77

15.77

Trientalis borealis

22.72

0.32

11.11

Osmunda cinnamomea

18.63

2.82

10.67

Aronia melanocarpa

13.63

0.59

7.27

Point intercepts

Vaccinium corymbosum

100.00

Aronia melanocarpa

36.36

Acer rubrum

22.72

Toxicodendron vernix

9.09

A melanchier arbor ea

4.54

I am also greatly indebted to Miss Bertha A. Daubendiek and the Eastern Michigan
Nature Association for permitting and encouraging this study of the Pennfleld Bog Plant
Preserve and for the generous grant which made the field studies possible.

LITERATURE CITED

Beaman, J. H., & J. W. Andresen. 196 6. The vegetation, floristics, and phytogeography of
the summit of Cerro Potosi, Mexico. Am. Midi. Nat. 75: 1-33.

Dansereau, P. 1957. Biogeography. Ronald Press, New York. 394 pp.

Eichmeier, A. H. [n.d.] Climatography of the United States. 1926-1955. Station 20-20
Battle Creek, Mich.

Flint, R. F., et al. 1959. Glacial Map of the United States East of the Rocky Mountains.
Geol. Soc. Am., New York.

Gates, F. C. 1942. The bogs of northern lower Michigan. Ecol. Monogr. 12: 213-254.
Humphrys, C. R., & R. F. Green. 1962. Calhoun County Lake Inventory. Mich. Lake
Invent. Bull. 13. Dep. Resource Develop., Mich. St. Univ., East Lansing. 4 pp.
Leverett, F, & F. B. Taylor. 1915. The Pleistocene of Indiana and Michigan and the History
of the Great Lakes. U. S. Geol. Surv. Monogr. 53. 529 pp.

Montgomery, J. D., & D. E. Fairbrothers. 1963. A floristic comparison of the vascular plants
of two sphagnous wet lands in New Jersey. Bull. Torrey Bot. Club 90: 87-99.

Rigg, G. B. 1916. A summary of bog theories. PI. World 19: 310-325.

Transeau, E. N. 1905. The bogs and bog flora of the Huron River valley. Bot. Gaz. 40:
351-375; 418-448.

Transeau, E. N. 1906. The bogs and bog flora of the Huron River valley [concl.] . Bot. Gaz.
41: 17-42.

28

THE MICHIGAN BOTANIST

Vol.8

BRYOPHYTES NEW TO MICHIGAN. II1

Howard Crum and Norton G. Miller
Herbarium, The University of Michigan, Ann Arbor,
and Department of Botany and Plant Pathology,

Michigan State University, East Lansing

A continuation of bryological activity in various parts of Michigan during
the past year has resulted in a number of interesting additions to the state flora.
In addition to the four taxa reported here, 14 species new to the state, to be
reported by others but named or confirmed by us, might be given cursory men¬
tion in the interest of unifying records: Lophozia muelleri (Nees) Dum. (Delta
Co.), Sphagnum annulatum var. porosum (Schlieph. & Warnst. ex Warnst.) Maass
& Isov. ex Maass (Cheboygan and Dickinson cos.), S. auriculatum Schimp.
(Antrim and Chippewa cos.), S. contortum Schultz (Emmet, Houghton, and
Mackinac cos.), S. platyphyllum (Lindb. ex Braithw.) Sull. ex Warnst. (Chip¬
pewa, Cheboygan, Kalkaska, Keweenaw, and Schoolcraft cos.), S. pulchrum
(Lindb. ex Braithw.) Warnst. (Chippewa, Mackinac, and Schoolcraft cos.), S.
riparium Angstr. (Emmet and Luce cos.), S. subfulvum Sj^rs (Chippewa and
Luce cos.), S. subnitens Russ. & Warnst. ex Warnst. (Mackinac Co.), Fissidens
exilis Hedw. (Eaton Co.), Eucladium verticillatum (Brid.) BSG (Delta Co.),
Physcomitrella patens (Hedw.) BSG (Wayne Co.), Sematophyllum demissum
(Wils.) Mitt. (Eaton Co.), and Schwetschkeopsis fabronia (Schwaegr.) Broth.
(Berrien Co.).

Specimens cited below are deposited in the herbaria of the University of
Michigan (MICH), Michigan State University (MSC), University of Michigan Bio¬
logical Station (UMBS), or the private collection of Norton G. Miller. Unless
otherwise stated, the distributional ranges given are those which we have con¬
firmed by actual study of specimens.

Pottia davalliana (Sm. ex Drake) C. Jens. Bay Co. On soil in second-year
clover field, Eight Mile Rd. 0.3 mi. N. of Cottage Grove Rd., Kawkawlin Town¬
ship, H. Crum, Oct. 26, 1968 (MICH). Emmet Co. On dry, sandy loam in fallow
field, 1.4 mi. N. of Brutus, N. G. Miller 5005 in part, with Pleuridium subula-
tum, Aug. 16, 1968 (in herb. Miller). Genesee Co. On soil, mowed area at road¬
side 3 mi. W of Clio on M57, H. Crum, Oct. 26, 1968 (MICH). Presque Isle Co.
On calcareous soil in bare spots of alfalfa field, 2.5 mi. E. of Posen, TV. G. Miller
4875, Aug. 3, 1968 (MICH), also Erica M. Frank, Aug. 3, 1968 (MICH); on bare,
calcareous soil, 1 mi. W. of M65 on Metz Rd. near Posen, H. Crum, Oct. 25,
1968 (MICH); on calcareous soil in mowed area, new section of Catholic ceme¬
tery, Posen, H. Crum, Oct. 25, 1968 (MICH). Saginaw Co. On soil in mowed

iThis work was supported in part by a grant from the National Science Foundation
(NSF-GB-6095) for workshops in Bryology at the University of Michigan Biological Station,
Pellston, Michigan.

1969

THE MICHIGAN BOTANIST

29

area, outskirts of Zilwaukee, H. Crum , Oct. 26, 1968 (MICH). — A minute moss
of rare occurrence on limey soils in disturbed places, such as old fields, pastures,
lawns, and roadsides; southern Ontario; throughout Europe.

Ephemerum crassinervium (Schwaegr.) Hampe. Emmet Co. On sandy
loam, moist depression in fallow field, 1.4 mi. N. of Brutus, TV. G. Miller 5004,
Aug. 16, 1968 (in herb. Miller; MICH), also//. Crum, Aug. 16, 1968 (MICH).
Jackson Co. On soil in corn field, on Mt. Hope Rd. near creek ca. 0.8 mi. N. of
Seymour Rd., H. Crum, D. H. Vitt, W. W. Patrick, & Paula Collier, Oct. 9, 1968
(MICH). Presque Isle Co. On soil at edge of stubble field, 1 mi. W. of Posen on
Grand Lake Rd., H. Crum, Oct. 25, 1968 (MICH). Saginaw Co. On soil at edge
of corn field, Birch Run , H. Crum, Oct. 24, 1968 (MICH). Washtenaw Co. On
soil, cornfield, 0.7 mi. N. of Joy Rd. on Webster Church Rd., Webster Township,
H. Crum, Oct. 17, 1968 (MICH); on soil, cornfield, on Walsh Rd. 0.6 mi. E. of
Scully Rd., Webster Township, H. Crum, Oct. 17, 1968 (MICH). — A rare,
ephemeral “pygmy” moss growing as scattered, minute buds on an abundant
protonema, on damp soil in old fields, on river banks, along paths, or at road¬
sides. The species in endemic to North America where it ranges from Quebec and
Ontario southward to Florida and Louisiana.

Pohlia annotina (Hedw.) Lindb. var. annotina. Alger Co. On wet sand,
seepy roadside bank, with Dicranella cerviculata, NW. of Chapel Falls, ca. 7 mi.
W. of Van Meer, N. G. Miller 4999, D. H. Vitt 1160, Aug. 13, 1968 (UMBS). -
A moss best recognized by its dull appearance and regular production of small,
yellowish, ovoid brood-bodies in leaf axils. The var. loeskei Crum, Steere, &
Anders., which has elongate, somewhat twisted, vermicular brood-bodies, has
also been found in a few Michigan localities (Bryologist 70: 118, 1967). The
typical variety has been found in northern and central Europe, British Columbia
to California, and a few localities in eastern North America (Michigan, New
Hampshire, New York, and Ontario).

Campy lium radicale (P.-Beauv.) Grout. Cheboygan Co. On moist humus
and muck, bog forest, Livingston Bog, 2.5 mi. SE. of University of Michigan
Biological Station, Burt Township, TV. G. Miller 3186, June 30, 1966 (MICH).
Ontonagon Co. Porcupine Mts., G. E. Nichols & W. C. Steere, Aug. 1935 (MICH,
as Amblystegium furatzkanum). — A seldom recognized species resembling some¬
what C. polygamum (BSG) C. Jens, but differing in smaller size and shorter leaf
cells which are not or only slightly inflated at the basal angles. It may be recog¬
nized also by its remote, wide-spreading leaves which are abruptly narrowed to a
channeled acumen from an ovate base. Only a few specimens have been seen,
from Florida, Michigan, New Jersey, Pennsylvania, and Virginia; it has also been
reported, by various authors, from Newfoundland and Maine.

30

THE MICHIGAN BOTANIST

Vol. 8

Hutu re education fea ture —

NATURAL AREA PRESERVATION
IN THE AGE OF THE MEGALOPOLIS1

Ronald O. Kapp
Department of Biology,

Alma College, Alma, Michigan

We live in an era in which the most obvious relationship of man to nature
is disturbance and destruction. Bulldozing of hills, damming of streams, levelling
of forests, spraying of roadsides, fields, timberlands— these are the hallmarks of
an advanced technology and vigorous economy. These evidences of man’s prow¬
ess as a manipulator of nature fill the daily press, second in precedence only to
the news of political and social issues.

But the dedication of a tract of land as a natural area marks a different
kind of relationship of man to the natural world. When we set aside an area to
remain undisturbed, allowing native species of plants and animals to survive and
maintain their natural ecological relationships, we underscore the urgency that
man became a respector and protector of nature. The dedication of each natural
area in Michigan, whether large or small, is therefore an important event worthy
of public attention.

Some might question the importance of establishing natural area preserves,
nature study areas, or wilderness tracts. In a world of burgeoning population
growth and a finite supply of resources, should not all areas be exploited to yield
their maximal production? Of what value are the vanishing species— those which
may soon be added to the long list of extinctions caused by man? Of what con¬
sequence are the nature preserves with their confusing array of organisms and
perplexing complex ecological processes? These questions press conservationists
continually ; they are especially insistent if short-term economic gain is an alter¬
native to preservation. Develop the undeveloped; stretch the subdivision; provide
more resorts; increase the tax base— these are the demands heard throughout
Michigan and the entire country. Even the choicest areas, dedicated as state and
national parks by early visionaries, are being threatened. Too frequently “inex¬
pensive” rights-of-way are granted across public land; mineral, water, and timber
rights are sought on these public lands, usually under the guise of saving the
taxpayer money.

In the face of such economic pressures, let us now reassert that nature
preservation is an important, even critical, state and national goal. The reasons
are many, and mostly very practical; let me cite a few:

1. Nature Study Areas provide “living museums” or outdoor classrooms which
allow observation and study of the world of nature in its unmanipulated, nearly

1 Remarks by the Chairman of the Michigan Natural Areas Council, delivered on the
occasion of the dedication of the Nature Study Area at Femwood, 1720 Range Line Road,
Niles, Michigan, on October 6, 1968.

1969

THE MICHIGAN BOTANIST

31

natural state. Students of all ages, those engaged in formal schooling or scouting,
family groups, the casual adult, or the nature enthusiast, may observe firsthand
the native plants, birds, mammals, and invertebrates. Frequently, as at Fernwood
and some of our other outdoor nature centers, interpretive nature trails and dis¬
plays supplement the direct observation of nature. No indoor classroom can pro¬
vide such educational opportunities.

2. Scenic Sites in Michigan provide some of the most exquisite scenery in the
world. Unless dedicated and preserved, they are Fable to despoiling by buddings
or other “human improvements” or they may fall into private ownership and
their enjoyment be denied to the public.

3. Managed Tracts are natural areas dedicated to the preservation of interesting
and rare species or communities and which require artificial management for
their maintenance. In these instances, human management is encouraged, after
appropriate ecological study, to promote preservation. Frequently this may re¬
quire control of water level or periodic burning to maintain an area in an early
stage of ecologic succession. The extinction of these species or demise of these
unique ecological communities would be an irreparable loss; a loss significant
even to Homo sapiens, the super-dominant species. For who can tell when we
may need to turn to some of these native species for germ plasm for developing
new species and varieties of economic importance? Who can assay the economic
value of rich and diversified natural communities to effective tourism? Who can
safely minimize the importance of natural areas as a setting for therapy to cure
mental Flness? Who will assert that humankind wdl be more humane and spiritu¬
ally richer after he has despoiled, destroyed, or exterminated?

4. Natural Area Preserves contain one or more typical or unusual examples of
plant and animal communities; they are dedicated for maximum preservation
and for scientific observation and study of the biota. In some cases, these in¬
clude substantial tracts of wilderness where man is but an occasional visitor.
Preservation of such areas in their pristine state requires limitation of the num¬
ber of visitors to those engaged in the serious study of nature. The traffic of
thousands of human visitors would soon trample and destroy the very features
to be protected. A very practical reason for the preservation is the provision of
areas for scientific research. Aldo Leopold has said ( A Sand County Almanac,
1949):

The science of land-health (i.e. conservation), needs, first of all, a base datum
of normality of how healthy land maintains itself as an organism.

. . . Wilderness, then, assumes unexpected importance as a land laboratory.

. . . each biotic province needs its own wilderness for comparative studies of
used and unused land. It is of course too late to salvage more than a lop-sided system
of wilderness remnants, and most of these are far too small to retain their normalcy.
All wilderness areas, no matter how small or imperfect, have a large value to land
science. The important thing is to remember that recreation is not their only, or even
their principle, utility.

At the present time the need for such areas is being felt keenly by scientists
participating in the International Biological Program. Ecologists studying pro¬
ductivity and pollution need undisturbed areas representative of aU of the major

32

THE MICHIGAN BOTANIST

Vol. 8

biomes of the world (e.g., prairie, deciduous forests, coniferous forests, etc.) to
serve as controls in experimental ecological study. We cannot afford to lose an¬
other acre of wilderness for practical as well as aesthetic reasons!

ACTIVITIES OF THE MICHIGAN NATURAL AREAS COUNCIL

Since its independent organization in 1952, after several years as a com¬
mittee of the Michigan Botanical Club, the Michigan Natural Areas Council has
been instrumental in locating, evaluating and dedicating more than thirty natural
areas in Michigan. As the Michigan representative of the Nature Conservancy, the
Council pioneered in the development of preservation categories and has defined
use patterns and safeguards suitable to each. Nature Study Areas, Scenic Sites,
Managed Tracts, and Natural Area Perserves require different standards for pres-

Fig. 1. Natural Areas in Michigan: Circles represent dedicated areas in public ownership;
stars represent private areas; squares represent some of the tracts now under study which
should eventually be dedicated. Most of the dedications indicated here were accomplished
with cooperation of the Michigan Natural Areas Council or the Michigan Botanical Club; the
several sanctuaries of the Michigan Audubon Society and those dedicated by the Eastern
Michigan Nature Association are not shown. The general location or name of area, acreage,
and ownership are indicated below. Key: MDNR = Michigan Department of Natural Re¬
sources (formerly, Michigan Department of Conservation); USFS = United States Forest
Service; NWR = National Wildlife Refuge; P = private.

1. Porcupine Mountains State Park, 45,210, MDNR

2. Tahquamenon Falls State Park, 4,868, MDNR

3. Betsy Lake area, 14,137, MDNR

4. Wagner Falls Scenic Site, 10, MDNR

5. Laughing Whiteflsh Falls Scenic Site, 160, MDNR

6. Huron Mountain Club, 17,700, P

7. Strangmoor, Seney Wildlife Refuge, 640, NWR

8. Northern hardwoods, Hiawaths National Forest, 200, USFS

9. Virgin cedar swamp, Ottawa National Forest, 233, USFS

10. Craig-Crooked Lake State Park project, MDNR

11. Sylvania, USFS

12. Bois Blanc Island, 696, MDNR

13. Wilderness State Park, 2,390, MDNR

14. South Manitou Island, 1,589, MDNR

15. Besser Natural Area, 135, MDNR

16. Kirtland’s Warbler Management Area, 4,010, USFS

17. Roscommon virgin red pine stand, 160, MDNR

18. Tobico marsh, 1,652, MDNR

19. Newaygo prairie, Manistee National Forest, 80, USFS

20. Woldumar, about 80, P

21. Shadbush tract, Utica Recreation Area, 70, MDNR

22. Haven Hill, Highland Recreation Area, 806, MDNR

23. Proud Lake bog, 105, MDNR

24. Homer Woods, 44, University of Michigan Botanical Gardens

25. Black Spruce bog, Waterloo Recreation Area, 40, MDNR

26. Ann Arbor Outdoor Center, P

27. Ann Arbor wet prairie

28. Middlebelt prairie

29. Warren Dues State Park, 320, MDNR

30. Warren Woods, 312, MDNR

31. Fernwood, 18, P

1969 THE MICHIGAN BOTANIST

33

ervation. In all of these, however, there are two common denominators for pres¬
ervation: maintenance in indefinite perpetuity and minimal alteration or disturb¬
ance.

The Michigan Natural Areas Council does not own property. It identifies
appropriate sites, and arranges for their dedication and preservation under aus¬
pices of an appropriate permanent owner. Most of the dedicated natural areas
are in public ownership. The Council seeks to assure the zoning of state parks so
that recreation and natural area preservation are maintained in appropriate bal¬
ance. About 20% of the lands owned and managed by the Michigan Department
of Natural Resources’ Parks Division have been dedicated in a suitable category.
The Council also cooperates with administrators of State and National Forests in
selecting areas for preservation, recreation, and production. These natural areas
in public ownership range in size from 45,000 acres in the Porcupine Mountains
State Park and nearly 19,000 acres in the Tahquamenon Falls and Betsy Fake
area, to an 80 acre relic-prairie Managed Tract in Newaygo County and a 10 acre

34

THE MICHIGAN BOTANIST

Vol. 8

Scenic Site at Wagner Falls in Alger County (Fig. 1). In recent years private or¬
ganizations, corporations, and individuals have also become interested in natural
area preservation. The Huron Mountain Club pioneered in this effort by dedicat¬
ing 17,700 acres in Marquette County. The Michigan Audubon Society has es¬
tablished sanctuaries, often in connection with nature centers; the Michigan Bo¬
tanical Club has purchased plant sanctuaries for transfer to suitable managing
agencies; the Eastern Michigan Nature Association has purchased several tracts.
Private corporations have established nature centers and nature study areas;
Woldumar, southwest of Fansing, and Fernwood, which we dedicate today, are
examples (Fig. 1). In addition, many universities and colleges have ecological
tracts which may or may not be under dedication.

Calculation of the total acreage preserved by these organizations is diffi¬
cult. I believe, however, that the dedication of Fernwood Nature Study Area is a
particularly auspicious occasion because it pushes that total over the 100,000-
acre mark in Michigan.

Fig. 2. Fernwood and adjacent lands, Berrien Co., Michigan. A: St. Joseph River; B: Nature
Study Area, about 18 acres of woodland dedicated October 6, 1968; C: Nature Center area
and gardens; D: Future arboretum and nursery area; E: Entrance gate at 1720 Range Line
Road, the extreme southeast comer of the property. The road forms the east boundary and
the river the west margin. (Photo by Roger E. Lorensen, March 31, 1965)

1969

THE MICHIGAN BOTANIST

35

FERNWOOD NATURE STUDY AREA

The tract of land we dedicate today, as a Nature Study Area at Fern wood,
is rich in plant species. In spite of its limited area, the ecologic diversity of the
tract also provides nesting sites for many kinds of birds.

The area provides an important addition to the biotic communities which
have been preserved in Michigan. At Warren Dunes and Warren Woods, which are
also in Berrien County, a dunes community, deciduous hardwoods, and rich
mesophytic beech-maple forest have been dedicated earlier as natural areas. Now
this Fernwood tract adds a mosaic of river border habitats (Fig. 2). These three
areas provide a fine nucleus of nature preserves for southwestern Michigan. If as
many outstanding sites could be identified and dedicated in every county, the
state would have an admirable and more nearly adequate system of natural areas.

In vast expanses of the state, particularly in the south-central, central, and
northern Lower Peninsula, there are no dedicated natural areas. It might occur
to someone that prior dedication of 100,000 acres is ample. However, 80% of
this acreage is concentrated in five large tracts in the Upper Peninsula; southern
Michigan has inadequate coverage and the dedicated areas are small. Those of us
who are committed to the cause of nature education might take as a challenge,
and an attainable goal, the establishment of natural area reserves in every county
of the state.

Demographers predict the development of three huge strip cities in Ameri¬
ca by the year 2000. These will be Bos wash on the east coast, and Sansan on the
west, and Chipitts, into which Fernwood will fall. We can already see the devel¬
oping urban sprawl from Chicago to Detroit, Cleveland, and Pittsburgh. It is
most urgent that our work proceed quickly in this belt of high population densi¬
ty in southern Lower Michigan. Nature Study Areas and Nature Centers like
Fernwood will serve increasing numbers of school children, college students, and
researchers; the general public will be increasingly anxious to see, hear, and sense
nature at its best.

SAMUEL H. WATSON, PIONEER BOTANIST
OF SOUTHERN WISCONSIN

Lytton J. Musselman
Department of Botany and Zoology,

University of Wisconsin, Rock County Center, Janesville

In a previous note (Musselman, 1968) the collection of S. H. Watson in the
Milton College herbarium was briefly mentioned. Further investigation has pro¬
vided some information on the life and collection of this early botanist.

Samuel Harding Watson was born in Mildenhall, England, on December 1,
1837, and came to this country with his parents in 1847 when they settled on a

36

THE MICHIGAN BOTANIST

Vol. 8

farm in Dunkirk Township in the southeast part of Dane County, Wisconsin.
Watson entered the University of Wisconsin about 1858 and was most active
from 1858 to 1861, collecting during the school year in Madison and in the vi¬
cinity of his parents’ farm during the summer.

After leaving the university, Watson taught school near Waunakee and
Okee, Wisconsin. In 1876 he began law practice in the office of Richard Lindsay
in Lodi. According to his obituary in the Lodi Enterprise , Watson took an active
part in community affairs and retained his interest in natural history until his
death, caused by complications following pneumonia. He died at the home of his
daughter in Waunakee on December 12, 1928, at the age of 91.

Watson’s interest in botany may be traced to 1854 when at the age of 17
he prepared a list of 303 species entitled “Alphabetical index of plants sent to
the University of Wisconsin,” dated Lebruary, 1854. These specimens were un¬
doubtedly destroyed in the fire which later burned the entire Wisconsin herbari¬
um (Bryan, 1950).

Based on information gleaned from his labels, Watson took botany in 1859
at the University of Wisconsin, a course then taught by Dr. Ezra Carr (Bryan,
1950). Mr. T. J. Hale apparently was a student in the same class. Unlike his lesser
known classmate, Hale published two papers on the Wisconsin flora, in which he
made several references to Watson’s collections (Musselman, 1968).

Many of the specimens from other collectors found in the Watson collec¬
tion may have been obtained by Hale and passed on to Watson. A portion of
Watson’s collection is contained in nine 10” x 12” volumes along with many
specimens obtained by Hale from other botanists. It is unclear as to how Wat¬
son’s collections came to Milton but college records indicate that two of his
younger brothers attended Milton College. Watson’s collection has been undis¬
turbed for nearly a century except for some mounting of specimens done around
the turn of the century and some annotations by Albert Rogers Crandall
(“ARC”), professor of natural history and physiology at Milton from
1902 to 1918.

Mr. Watson was an avid collector of all plants both cultivated and native.
His collection contains such difficult groups as grasses, sedges, and cryptogams.
Most of his collections were made in the Madison vicinity and in Dunkirk Town¬
ship with only an occasional specimen from other localities. I have never en¬
countered any Watson collections from outside Wisconsin although Pennell
(1935) lists Michigan as one of the states in which he collected. Below is a listing
of localities in which he collected:

COLLECTION SITES OF S. H. WATSON IN SOUTHERN WISCONSIN
“Janesville” (T3N, R12E, Rock Co.)

“Fulton” (Section 18, Fulton Township, T4N, R12E, Rock Co.)

“Porter” (Porter Township, T4N, R11E, Rock Co.)

“Union” (Section 10, Union Township, T4N, R10E, Rock Co.)

“Dunkirk,” “D-” (Dunkirk Township, T5N, R11E, Dane Co.)

“Rutland” (Rutland Township, T5N, R10E, Dane Co.)

1969

THE MICHIGAN BOTANIST

37

“Madison,” “Mad.” (T7N, R9E, Dane Co. Many collections were made on or near the uni¬
versity campus or lakes.)

“Pine Bluff” (Section 27, Cross Plains Township, T7N, R7E, Dane Co.)

“Mazo-manie” (Mazomanie Township, T9N, R8E, Dane Co.)

“Middleton” (Middleton Township, T7N, R8E, Dane Co.)

“Blue Mounds” (Section 11, Brigham Township, T6N, R5E, Iowa Co.)

“Arena” (Arena Township, T8N, R5E, Iowa Co. Many specimens labeled “Arena” are from
the sand areas along the Wisconsin River.)

“Bluffs of the Wisconsin River, Sauk Co.”

“Fort Atkinson” (T5N, R14E, Jefferson Co.)

“2nd Lake,” “Dead Lake” (Perhaps in reference to some of the Madison lakes?)

Unfortunately many of Watson’s labels contain little data other than the
name of the plant. Figure 1 illustrates a representative label. Watson’s collec¬
tion is extremely valuable as an indication of the vegetation of southern Wiscon¬
sin as it existed over one hundred years ago. Of particular interest are his collec¬
tions of prairie species which are now rare or uncommon, such as Gerardia auri-
culata, Agoseris cuspidata, and many others.

Fig. 1. Representative label of S. H. Watson.

ACKNOWLEDGMENTS

I wish to thank Mr. J. L. Skaggs (Registrar of Milton College), Mrs. Ruby Murphy
(granddaughter of S. H. Watson), Dr. E. G. Voss, The Lodi Enterprise, and the Wisconsin
State Historical Society for valuable assistance. Much help in the preparation of this paper
was obtained from Dr. R. L. Stuckey’s studies of early botanists. Portions of this work com¬
pleted during the summer of 1968 were incidental to research supported by a University of
Wisconsin Center System Research Grant.

LITERATURE CITED

Bryan, G. S. 1950. A brief history of the development of botany and the department of
botany at the University of Wisconsin to 1900. Trans. Wis. Acad. 40: 1-27.
Musselman, L. J. 1968. The Milton College Herbarium. Mich. Bot. 7: 269-271.

Pennell, F. W. 1935. The Scrophulariaceae of Eastern Temperate North America. Monogr.
Acad. Phila. 1. 650 pp.

38

THE MICHIGAN BOTANIST

Vol. 8

DARK-BARKED BIRCHES OF SOUTHERN MICHIGAN

Bruce P. Dancik

Department of Forestry, School of Natural Resources,

The University of Michigan, Ann Arbor

Yellow birch ( Betula alleghaniensis Britton) is a highly variable member of
the dark-barked birch group (subsection Costatae Regel) in the Great Lakes, St.
Lawrence River, and Appalachian Mountain regions of North America. Original¬
ly named B. lutea Michaux fil., yellow birch is now called B. alleghaniensis be¬
cause Michaux’s name has been interpreted as illegitimate (Little, 1953; Bray-
shaw, 1966). Over much of its range, yellow birch may be quickly and easily
identified by its characteristically yellowish, generously exfoliating bark and
very pubescent and ciliate, palm-like pistillate bracts. Likewise, its relative of the
Appalachian Mountains, the black, sweet, or cherry birch (. B . lenta L.) is distinc¬
tive because of its dark reddish or black, cherry-like, nonexfoliating bark; and
glabrous, somewhat cruciform, pistillate bracts. Upon discovering several dark
and tight-barked birches in southeastern Michigan (Dancik, 1967), I initiated a
search for past reports and interpretations of this taxon.

At various places in the range of yellow birch, particularly in southern
Michigan, Minnesota, and Wisconsin, and northern Indiana and Ohio, a dark and
tight-barked birch has been recognized. It has been called B. lenta L., B. lutea
forma fallax Fassett (. B . alleghaniensis var . fallax Brayshaw), and B. alleghanien¬
sis Britton. Blanchard (191 1) published one of the earliest accounts of this taxon
and probably put it into the proper perspective at that time:

The provincial botanists, having never seen the real black birch think that when
they find a yellow birch ( B . lutea L.) with black bark they have found a black birch,
and lumbermen everywhere from the Adirondacks to New Brunswick are sure that
they have two birches which they are loth to believe can be age variations only. . . .
[The black birch’s] range as well as its abundance in Michigan, Wisconsin, Minneso¬
ta, and Iowa is unknown. In short, the two birches B. lenta and B. lutea have been so
confounded by lumbermen and botanists that no dependence whatever can be placed
on any published statement as to either range or frequency of the black birch in the
north, northeast, or northwest.

Reports of these dark-barked birches led to many exaggerations of the
range of the sweet birch. When Fassett (1932) described the dark-barked birch
taxon as a form of yellow birch, he stated, “This tree occurs in company with
typical B. lutea, and differs from it only in having bark closely simulating that of
B. lenta. Such individuals probably explain numerous sheets in the University
herbarium misidentified as B. lenta, and perhaps form the bases of reports of the
cherry birch from this state [Wisconsin] .” Similarly, in Minnesota, Rosendahl
(1916) recognized that several specimens collected in that state and called B.
lenta were not identified correctly and “the specimens can therefore be referred
to B. lutea without any doubt whatever.” He was working with herbarium ma¬
terial and concluded this without realizing that they were probably misidentified
because of their bark. Similar records in southern Michigan and northern Indiana
of sweet birch localities (Beal, 1905; Bingham, 1945; Cole, 1901; and Pepoon,

1969

THE MICHIGAN BOTANIST

39

1927) probably reflect the occurrence of this form. As recently as 1958, Harlow
and Harrar continued to show the southern third of the Lower Peninsula of
Michigan within the range of sweet birch, though there are no confirmed locali¬
ties for the species there.

At the time when both B. lutea and B. alleghaniensis were accepted as
names for separate taxa, several authors felt that the dark-barked birch was B.
alleghaniensis. According to Dodge (1921, p. 186), “Professor C. S. Sar¬
gent ... is inclined to call this birch Betula allegheniensis Britton, southern yel¬
low birch, and doubts whether we have Betula lenta L., cherry birch, in Michigan
or western Ontario.” In Nova Scotia, Fernald (1921) felt that reported speci¬
mens of sweet birch were actually B. lutea var. alleghaniensis (Britton) Ashe. In
Indiana, Deam (1921) referring to yellow birch, stated:

That there are geographic races or Mendelian segregates of this species is evidenced by
the different interpretations given this species by different authors. Betula alleghani¬
ensis Britton appears to be one of them. The descriptive difference between Betula
lenta and Betula lutea is not clear, which has resulted in many authors crediting
Betula lenta to Indiana and the area west of Indiana.

The morphological characteristics of this confusing, dark and tight-barked
birch, in view of the apparent east of hybridization between the birches (Dancik,
1967), have suggested that this taxon may be a hybrid between sweet birch and
yellow birch. Britton’s original description of B. alleghaniensis (1904) indicated
that his new taxon was easily and often confused with both sweet birch and
what he called B. lutea, suggesting the possibility of hybridization. Others felt
that B. alleghaniensis was in fact this hybrid. Eames (Harger et al., 1917) recog¬
nized this possibility from specimens he examined in Connecticut: “A form
which is intermediate between, and possibly a hybrid of, this species [B. lenta]
and B. lutea has been described as B. alleghaniensis Britton and is rare in the
western part of the state.” Deam (1921) felt that at least some of the peculiar
forms of yellow birch found in Indiana could be accounted for by hybridization
with sweet birch. Braun (1961) suggested that many individuals of B. lutea
forma fallax Fassett in Ohio could be confused with sweet birch and under the
latter observed that “. . . combinations of characters shown by some specimens
from northeastern Ohio suggest hybridization” with yellow birch. This hybrid
interpretation is rather inconsistent in the many areas where the unusual birch
occurs but the sweet birch does not.

Professors Burton V. Barnes and Warren H. Wagner, Jr., and I have found
many of these unusual birches in and around several swamps and lakes in Wash¬
tenaw, Jackson, Kalamazoo, and Allegan counties in Michigan. A detailed study
was made of one population at Walsh Lake, Washtenaw County, Michigan, and
will be reported elsewhere. The distinctive bark of these birches (Fig. 1) is ri¬
valed in interest by their pistillate bracts, many of which are quite glabrous and
so variable in shape that they can be easily confused with those of sweet birch,
paper birch (B. papyrifera Marsh.), or river birch ( B . nigra L.) (Dancik, 1967). At
this time it appears that these forms are not sufficiently genetically distinct to
warrant specific, or for that matter, infraspecific, status. Enough biotypes with
bark types similar or intermediate to those of “normal” yellow birch appear in
these populations to indicate that the dark-barked character is not true breeding

40

THE MICHIGAN BOTANIST

Vol. 8

or particularly unique. Similarly, none of the pistillate bract forms at Walsh Lake
are particularly unique or dominant. This was hinted at by Hanes and Hanes
(1947) when they mentioned that in Michigan “Where the species and the varie¬
ty [B. lutea var. macrolepis Fernald] both occur, . . . trees intermediate between

Fig. 1. Bark of representative yellow birches at Walsh Lake, a) Tree 1, 13.6 in. dbh; b) tree
19, 8.0 in. dbh; c) tree 44, 15.0 in. dbh; and d) tree 64, 8.4 in. dbh.

1969

THE MICHIGAN BOTANIST

41

the two can be found.” Evidently these dark-barked birches (with highly variable
pistillate bracts) are yellow birches with the bark characteristics influenced by
the local environment, genetic differentiation subsequent to isolation from the
more northerly and easterly portions of the gene pool of yellow birch, or intro-
gressed genes from bog birch (. B . pumila L.), a plant that frequently hybridizes
with yellow birch in this area (Dancik, 1967). However, the possibility of intro-
gression is not supported by present evidence. Studies are underway to compare
yellow birches of northern Michigan with those of southern Michigan and fur¬
ther clarify the identity of the dark-barked forms.

Any readers knowing of localities for yellow birches in the southern Lower
Peninsula of Michigan or adjacent parts of Indiana, Ohio, and Illinois are en¬
couraged to contact the author.

ACKNOWLEDGMENTS

Support of investigations on the dark-barked birches by funds provided under the
Mclntire-S tennis Law (P.L. 87-788) is gratefully acknowledged. Acknowledgment is also
made to the Michigan Department of Natural Resources for allowing field research in the
Waterloo Recreation Area.

LITERATURE CITED

Beal, W. J. “1904” [1905] . Michigan flora. Rep. Mich. Acad. 5: 1-147.

Bingham, Marjorie T. 1945. The Flora of Oakland County, Michigan. Cranbrook Inst, of Sci.
Bull. 22. 155 pp.

Blanchard, W. H. 1911. The range of black birch to be restricted. Rhodora 13: 206-207.
Braun, E. Lucy. 1961. The Woody Plants of Ohio. Ohio State Univ. Press, Columbus. 362

pp.

Brayshaw, T. C. 1966. The names of yellow birch and two of its varieties. Canad. Field-Nat.
80: 160-161.

Britton, N. L. 1904. Four new North American birches. Bull. Torrey Bot. Club. 31:
165-166.

Cole, Emma J. 1901. Grand Rapids Flora. A. Van Dort, Grand Rapids. 170 pp.

Dancik, Bruce P. 1967. A Population Study of the Birches , Betula alleghaniensis, B. pumila,
and their Hybrid. Master of Forestry thesis, Univ. Mich. 89 pp.

Deam, Charles C. 1921. Trees of Indiana. Indiana Dep. Cons. Pub. 13. 317 pp.

Dodge, C. K. 1921. Miscellaneous Papers on the Botany of Michigan. Mich. Geol. and Biol.
Survey Publ. 31. 234 pp.

Fassett, Norman C. 1932. Notes from the herbarium of the University of Wisconsin- VIII.
Rhodora 34: 95-96.

Femald, M. L. 1921. The Gray Herbarium expedition to Nova Scotia, 1920. Rhodora 23:
257-278.

Hanes, Clarence R., & Florence N. Hanes. 1947. Flora of Kalamazoo County, Michigan.
Schoolcraft, Mich. 295 pp.

Harger, E. B., C. B. Graves, E. H. Eames, C. H. Bissell, L. Andrews, & C. A. Weatherby.

1917. Additions to the flora of Connecticut. Rhodora 19: 128.

Harlow, William M., & Ellwood S. Harrar. 1958. Textbook of Dendrology. Fourth Edition.
McGraw-Hill Book Co., New York. 561 pp.

Little, Elbert L., Jr. 1953. Check List of Native and Naturalized Trees of the United States
(Including Alaska). U.S. Dep. Agr. Handb. 41. 472 pp.

Pepoon, H. S. 1927. An Annotated Floraof the Chicago Area. Chicago Acad. Sci. 554 pp.
Rosendahl, C. O. 1916. Observations on Betula in Minnesota with special reference to some
natural hybrids. Minn. Bot. Stud. 4: 443-459.

42

THE MICHIGAN BOTANIST

Vol. 8

AN UNUSUAL FRUITING OF

ASTEROPHORA LYCOPERDOIDES

Florence V. Hoseney

Herbarium, The University of Michigan, Ann Arbor

Atmospheric conditions in southeastern Michigan in 1968 were very
favorable for the fruiting of mushrooms. The total rainfall in May, June, and
July was 16 inches. This included a flood in June which resulted in the water in
most ponds and marshes remaining nearly one and a half feet above the normal
level for most of the summer. High humidity and moderate temperatures
contributed to maintain an exceptional fruiting of mushrooms over a much
longer period than in most years.

Not only were common species out in great numbers but also the rarer
ones appeared in surprising quantity. One of these which appears to be of special
interest, Asterophora lycoperdoides (Bull.) Ditmar ex Fr. (Fig. 1), was found in
a pine plantation at the University of Michigan Stinchfield Forest near Dexter,
Washtenaw County, Michigan, on July 25, 1968. This fungus is considered very
rare and usually only one or two specimens are found during a good season; but
upon this occasion Mrs. Ruth Zehner and I in two hours collected more A.
lycoperdoides than we could carry (three market baskets full), from one small
area. In October, it fruited again and an equal amount was gathered.

This species of Asterophora was found parasitizing Russula nigricans. A.
lycoperdoides is a cantharelloid fungus, which means it has thick gills with
obtuse edges, though in many specimens the gills are poorly formed. The young
caps are white and moist, becoming dry and brown at maturity and breaking up
into a mass of powdery chalamydospores. It probably reproduces more readily
by means of the chlamydospores, which form from the breaking up of the
pileus, than from basidiospores. In a second species, A. parasitica, basidiospores
form more regularly.

Further study of A. asterophora might be of interest. For instance, this
parasite preserves the host mushroom for a considerable length of time (two
weeks), suggesting antibiotic properties. Technical information about Astero¬
phora (Nyctalis) can be found in A Monograph of Cantharelloid Fungi by E. J.
H. Corner (Oxford University Press, 1966).

The host, Russula nigricans, seemed to prefer the “trashier” sections of the
pine plantation where trees had been cut down and left to rot or lower branches
had been trimmed but not cleared away, thus creating a debris-laden floor rather
than a clean, needle carpet over the earth.

Uninfested Russulae were few, but time did not permit thorough coverage
of the entire forest. Such an investigation appeared overwhelming because there
were several hundred basidiocarps in one small area alone. A large collection is
preserved in the Herbarium of the University of Michigan as a record of this
fruiting.

*■*

1969

THE MICHIGAN BOTANIST

43

Fig. 1. Asterophora lycoperdoides (Smith 6907)

44

THE MICHIGAN BOTANIST

Vol. 8

MICHIGAN PLANTS IN PRINT
New Literature Relating to Michigan Botany

This section lists new literature relating to Michigan Botany under four categories. A.
Maps, Soils, Geography, Geology (new maps and selected bulletins or articles on soils and
geology as these may be of use to field naturalists and students of plant distribution), B.
Books, Bulletins, etc., and C. Journal Articles (listing, respectively, all separate publications
and articles in other periodicals which cite Michigan specimens or include research based on
plants of wild origin in Michigan ;-not generally including work on cultivated plants nor
strictly economic aspects of forestry, conservation, or agriculture); D. History, Biography,
Exploration (travels and lives of persons with Michigan botanical connections). When the
subject matter or relation to Michigan is not clear from the title, annotations are added in
brackets. Readers are urged to call to the editor’s attention any titles (1960 or later) which
appear to have been overlooked— especially in less well known sources.

A. MAPS, SOILS, GEOGRAPHY, GEOLOGY

The following topographic maps for Michigan have been published by the U. S.
Geological Survey, 1200 S. Eads St., Arlington, Virginia 22202, since the previous listing in
our January, 1968. issue. All are 71/2-minute quadrangles (scale of 1:24,000 or about 2Vz
inches to a mile). Maps are supplied with green overprint showing wooded areas unless
request is made to the contrary. Standard topographic quadrangle maps are $.50 each, and
maps in the 1:250,000 series are $.75 each, with a 20% discount on orders totalling $20.00
or more.

Following the name of the quadrangle, the county or counties in which it primarily
lies are added in brackets in the list below:

Alabaster [Iosco, Arenac]

Au Gres [Arenac]

Au Sable Point [Alger]

Au Sable Point SE [Alger]

Au Sable Point SW [Alger]

Limited revisions of the following sheets in the 1:250,000 series have been issued:
Escanaba (NL 16-8), Flint (NK 17-1), Midland (NK 16-3), and Toledo (NK 17-7).

Chiappetta, Jerry. 1968. Great Lakes, Great Mess. Audubon 70(3): 30-45. [A timely, but
unpleasant, article on pollution, with considerable reference to Michigan and many
data on the Great Lakes.]

(Grand Mere Association) [1968] Unique Grand Mere. [20] pp. [Well illustrated account of
geological and cultural history of this Berrien Co. area, with lists, by common name
only, of trees and some of the other plants which inhabit it. Copies of the booklet are
available from The Grand Mere Association, Box 140, Stevensville, Michigan 49127.]
Kelley, R. W., & W. R. Farrand. 1967. The Glacial Lakes Around Michigan. Dep. Conserva¬
tion, Geol. Surv. Bull. 4. 23 pp. [A concise history, with dates, of the various ice
advances in Michigan, and the related moraines and stages in development of the
Great Lakes.]

Kixkby, E. A. 1967. Index to Michigan Geologic Theses. Dep. Conservation, Geol. Surv.
Circ. 7. 33 pp. [List of 387 master’s and doctor’s theses from 26 Michigan and other
institutions, arranged alphabetically by author, with full titles; two-page subject
index. Most, but not all, were written since 1930. Includes no citation of number of
pages and no indication as to whether thesis has been published. One unexpected
omission is George Babcock Cressey’s classic “The Indiana Sand Dunes and Shore
Lines of the Lake Michigan Basin” (Univ. Chicago, 1923; published 1928).]

Grand Sable Lake [Alger]

National City [Iosco, Arenac]

Point Lookout [Arenac]

Toledo (revised) [Monroe, plus Ohio]

1969

THE MICHIGAN BOTANIST

45

B. BOOKS, BULLETINS, SEPARATE PUBLICATIONS

Chase, Clarence D. 1968. Michigan’s Timber Volume. North Central For. Exp. Sta., U. S.
For. Serv. Res. Note NC-50. 4 pp. [Gives tables on volume of growing stock and
volume of sawtimber on commercial forest land, by counties and species groups.
General comparisons are made between the 1966 data presented and 1955 data.]

Neal, James. 1967. Familiar Trees of Michigan. Mich. St. Univ. Extension Bull. E-616. 22
pp. [Compiled for 4-H Club members and others, with illustrations, brief comments,
and illustrated terminology, for some of the native and planted trees. Sloppy errors
in spelling and otherwise. Drawing of “American Hornbeam ( Ostrya virginiana )” is
actually fruiting Carpinus caroliniana; drawing of eastern cottonwood looks more like
quaking aspen; several of the fruit and leaf types illustrated on the last two pages-
e.g., ginkgo-do not apply to any of the species included in the booklet. The key does
not cover the same species as are illustrated, includes several statements without
alternative contrasts, and hence is often impossible to use. Although free, the booklet
is hardly worth the postage to request it.]

Peterson, Roger Tory, & Margaret McKenny. 1968. A Field Guide to Wildflowers. Boston,
Houghton Mifflin Co., 420 pp. $4.95 [Distribution stated for a number of species
mentions Michigan. See review in Mich. Bot. 7: 190.]

Scora, Rainer W. 1967. Interspecific Relationships in the Genus Monarda (Labiatae). Univ.
Calif. Publ. Bot. 41. 71 pp. $2.50 [Maps in this monograph show M. didyma, M.
punctata, and M. fistulosa in Michigan; “representative specimens” (2 and 1, respec¬
tively) are cited from Michigan only for the latter two species. ]

Thomson, John W. 1967. The Lichen Genus Cladonia in North America. Univ. Toronto
Press. 172 pp. $12.75 [Michigan included in the range stated in the key for several
species, and several other species are illustrated by photos of Michigan specimens.
Although Michigan and Colorado are mentioned in the key to C. carneola, the range
stated for this species in the text would seem to exclude these states. Useful introduc¬
tion and illustrations on such topics as lichen chemistry.]

C. JOURNAL ARTICLES

Brewer, Richard. 1966. Notes on vegetation and birds of a maple forest in southwestern
Michigan. Jack-Pine Warbler 44: 48-49. [Includes some vegetational data on a tract in
Kalamazoo Co.]

Clausen, Knud E. 1968. Variation in height growth and growth cessation of 55 yellow birch
seed sources. North Central For. Exp. Sta., U. S. For. Serv. Res. Pap. NC-23 (Proc.
8th Lake States For. Tree Improvem. Conf.): 1-4. [Five of the sources from which
seed was evaluated were in Michigan.]

Core, Earl L. 1968. The range of Carex trichocarpa Muhl. Castanea 33: 151-152. [Map
includes dots in Ingham and Washtenaw cos.; records from Kent and Genesee have
also been published by Hermann.]

Culberson, William Louis, & Chicita F. Culberson. 1968. The lichen genera Cetrelia and
Platismatia (Parmeliaceae). Contr. U. S. Natl. Herb. 34: 449-558. [C. olivetorum
cited and mapped from Isle Royale; P. glauca and P. tuckermanii also cited from Isle
Roy ale but not mapped there, though other Michigan collections of the latter are
mapped.]

Curtis, E. J. C., & J. E. Cantlon. 1968. Seed dormancy and germination in Melampyrum
lineare. Am. Jour. Bot. 55: 26-32. [Material from Crawford Co.]

Davis, H. A., Albert M. Fuller, & Tyreeca Davis. 1968. Contributions toward the revision of
the Eubati of eastern North America. II. Setosi. Castanea 33: 50-76. [Reports type of
Rubus notatus var. boreus (Kalkaska Co.) is inadequate for evaluation; several species
credited to Michigan: R. dissensus, R. junceus, R. superioris, R. wheeleri, R. specta-
tus, R. regionalis, R. wisconsinensis, R. angustifoliatus (?), R. glandicaulis, R. per-
spicuus, R. missouricus (inch R. mediocris & R. jcjuns); R. conabilis not evaluated.]

Dudley, Theodore R. 1968. Alyssum (Cruciferae) introduced in North America. Rhodora
70: 298-300. [A. alyssoides said to be naturalized in Michigan.]

46

Vol. 8

THE MICHIGAN BOTANIST

Fenwick, Mason G. 1968. Lake Huron distribution of Tabellaria fenestrata var. geniculata A.
Cleve and Coelastrum reticulatum var. polychordon Korshik. Trans. Am. Micr. Soc.
87: 376-383. [Considerable data from Michigan side of the lake.]

Gibbons, J. Whitfield. 1968. Carapacial algae in a population of the painted turtle, Chry-
semys picta. Am. Midi. Nat. 79:517-519. [Work done in “a marsh in southwestern
Michigan.”]

Hale, Mason E., Jr. 1967. New taxa in Cetraria, Parmelia, and Parmeliopsis. Bryologist 70:

414-427. [Distribution map for P. ambigua includes three dots in Michigan.]

King, James P. 1968. Seed source variation in tracheid length and specific gravity of five-
year-old jack pine seedlings. North Central For. Exp. Sta., U. S. For. Serv. Res. Pap.
NC-23 (Proc. 8th Lake States For. Tree Improvem. Conf.): 5-9. [Three of 34 seed
sources used were in Lower Peninsula.]

Lester, Donald T. 1968. Variation in cone morphology of balsam fir, Abies balsamea.
Rhodora 70:83-94. [Although map indicates that collections were made in Michigan,
no Michigan data are included in tables and text.]

Maas, W. S. C. 1967. Studies on the taxonomy and distribution of Sphagnum IV. Nova
Hedwigia 14: 187-214. [Cites Michigan specimens of S. majus and S. annulatum var.
porosum, and the stations are included on maps.]

Mangaly, Jose K. 1968. A cytotaxonomic study of the herbaceous species of Smilax: Sec¬
tion Coprosmanthus. Rhodora 70: 55-82; 247-273. [Michigan material cited and
mapped for S. lasioneuron (sic), S. illinoensis, and S. ecirrhata (sic) -the second of
these a new species intermediate between S. lasioneura and S. ecirrata. Citations are
riddled with careless errors (at least 25 errors in 31 Michigan citations, mostly involv¬
ing spelling of names of collectors or localities). Chromosome numbers for S. lasio¬
neura (2n = 26) and S. illinoensis (n = 13, 2n = 26) are given for Washtenaw Co.
material.]

McGregor, Ronald L. 1968. The taxonomy of the genus Echinacea (Compositae). Univ.
Kan. Sci. Bull. 48: 113-142. [ E. pallida cited from Beaver Island and Baraga Co.,
mapped at latter and two other locations; E. purpurea mapped in Michigan but no
specimens cited.]

Miller, H. E., et al. 1968. Infraspecific variation of sesquiterpene lactones in Ambrosia
psilostachya (Compositae). Am. Jour. Bot. 55: 316-324. [One of the collections
studied was from “Michigan.”]

Nilsson, Siwert. 1967. Pollen morphological studies in the Gentianaceae-Gentianinae.
Grana Palyn. 7: 46-145. [The pollen studied and illustrated of Swertia caroliniensis
was from Michigan (coll. Camps in 1897, presumably Jackson Co.) and that studied
oiHalenia deflexa was also from the state (Dodge in 1894).]

Owston, Peyton W. 1968. Multiple flushing in eastern white pine. For. Sci. 14: 66-67.
[Double branch whorls observed in southeastern Michigan -whether on native or
planted saplings is not stated.]

Pringle, James S. 1967. Royal Botanical Gardens herbarium. Gardens Bull. (Hamilton, On¬
tario) 21(3): 9-24. [One of the representative specimens illustrated is Solidago hough-
tonii from the shore of Douglas Lake, with comment on the unusual inland loca¬
tion.]

Rexrode, Charles O. 1968. Tree-wounding insects as vectors of the oak wilt fungus. For. Sci.
14: 181-189. [Map of reported distribution of oak wilt in U. S. (through 1964), by
counties, shows occurrence in southern Michigan.]

Richards, Edward Leon. 1968. A monograph of the genus Ratibida. Rhodora 70: 348-392.

[Cites R. pinnata from 8 Michigan counties and R. columnifera from one (Wayne).]
Sajdak, Robert L. 1968. Variation in bark characters and wood specific gravity of sugar
maple. North Central For. Exp. Sta., U. S. For. Serv. Res. Pap. NC-23 (Proc. 8th
Lake States For. Tree Improvem. Conf.): 10-14. [Four bark types illustrated by
photos; study on trees in Houghton Co.]

Steiner, Erich E. 1968. Dormant seed environment in relation to natural selection in Oeno¬
thera. Bull. Torrey Bot. Club 95: 140-155. [Two of the races studied represented
Upper Peninsula O. parviflora.]

Program Notes

March 16, 1969: State membership meeting, Michigan Botanical Club.

May 23-25: State membership meeting and annual Spring Campout, Michigan Botanical
Club: near Irons, Lake Co., in the Manistee National Forest.

Editorial Notes

There is still a need for black and white photographs (no smaller than 5 x 7), of artistic
merit, on winter appearance and uses of wild plants, for the Botanical Club’s projected pub¬
lication on “Winter Wildflowers’’ (see Mich. Bot. 7: 95. March, 1968). Chairman of the
committee is Dr. Helen V. Smith, 1766 Glenwood Rd., Ann Arbor, Michigan 48104.

Alert readers may detect that this issue of THE MICHIGAN BOTANIST is
copyrighted. The sole reason for this is to protect ourselves against any unauthorized
reproduction (e.g., by microfilm) and sale of complete sets. The existence of an occasional
copyrighted issue will mean that, so long as they last, complete sets of back issues will be
legally available only from us (and thus remain a valuable source of income). We are not
intending, by this action, to place any limitation on normal quotation and use of material
with credit to the source.

The October number (Vol. 7, No. 4) was mailed October 23, 1968.

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POO font, Mo y 1968

CONTENTS

Pollen Analysis at the Thaller Mastodon Site,

Gratiot County, Michigan

Edith R. Held & Ronald 0. Kapp . 3

Review— Living With Your Land . 10

An Ecological Analysis of a Southern Michigan Bog

Garrett E. Crow . 11

Bryophytes New to Michigan. II

Howard Crum & Norton G. Miller . 28

Nature Education Feature-

Natural Area Preservation in the Age of the Megalopolis

Ronald O. Kapp . 30

Samuel H. Watson, Pioneer Botanist of Southern Wisconsin

Lytton J. Musselman . 35

Dark-Barked Birches of Southern Michigan

Bruce P. Dancik . 38

An Unusual Fruiting of Asterophora lycoperdoides

Florence V. Hoseney . 42

Michigan Plants in Print . 44

Program Notes . 47

Editorial Notes . 47

(On the cover: Tips of willow branches along the
Huron River near Island Park, Ann Arbor,
photographed by Doug Fulton, February 1968.)

■ 7

r'

7 33

THE

Vol. 8, No. 2

MICHIGAN BOTANIST

LIBRARY

■"R 18 1969

NEW YORK
tOTANICAL GARDEN

March, 1969

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THE MICHIGAN BOTANIST-Published by the Michigan Botanical Club four times per

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Editorial Board

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THE MICHIGAN BOTANICAL CLUB

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1969

THE MICHIGAN BOTANIST

51

A PHYTOGEOGRAPHICAL ANALYSIS OF
A SOUTHERN MICHIGAN BOG

Garrett E. Crow

Department of Botany and Plant Pathology,

Michigan State University, East Lansing

Pennfield Bog is located in Calhoun County, Michigan. A description
and an ecological analysis of the bog were recently presented (Crow, 1969).
The present article is concerned with the phytogeography of the floristic com¬
position of the bog.

In order to assess the geographical affinities of a northern bog one must
be greatly concerned with the effect of Pleistocene glaciation on the flora of
North America. Certainly there is no question that as the ice sheets advanced
the vegetation standing in their paths was annihilated. There is much contro¬
versy, however, concerning the effect of the glaciers of Wisconsin age on the
flora south of the glacial boundary.

One school of thought maintains that there was a major displacement of
the boreal forest during Wisconsin glaciation, but “that Full-glacial tundra,
boreal forest, and deciduous forest formations were not identical in species
composition or even in vegetational structure with their present bioclimatic
analogues” (Martin, 1958, p. 378). Whitehead (1965) feels that palynological
evidence from Carolina Bay sediments demonstrates beyond doubt profound
changes in both climate and vegetation in this region during late Pleistocene
time.

On the other hand, some believe that the climate south of the glaciers
did not differ greatly from that of the present. Braun (1951, p. 145) main¬
tains that “. . . the deciduous forest zone, although narrowed, maintained itself
on the Appalachian Plateaus in southern Ohio and Kentucky while glaciers ex¬
tended southward in Ohio.”

Wolfe (1951) raises two pertinent questions concerning some species of
narrow ecological range which are present in microclimates near the glacial
border. If a developing tundra caused these species to retreat to the southern
Appalachians, “. . . how, with their narrow ecological amplitude and lack of
continuous migration routes, did they get back near the glacial border?” And
how did a plant like Sullivantia sullivantii (T. & G.) Britt., which is known
only from limestone and sandstone cliff faces of southern Ohio, southern Indi¬
ana and northern Kentucky, return to its present range and “. . . not persist in
some of the millions of grotto microclimates of the southern mountains”
(Wolfe, 1951, p. 137)?

litis (1965, 1966) has observed that there is a large western floristic ele¬
ment in northeastern North America. In the East these western taxa are re¬
stricted to glaciated areas. He feels that this distribution pattern strongly sug¬
gests that during the Pleistocene the eastern deciduous forests were essentially
closed communities and unchanged from those of today, except where subject
to periglacial phenomena.

52

THE MICHIGAN BOTANIST

Vol. 8

The following phytogeographic analysis takes the latter viewpoint.

The North American ranges of all species in the bog have been deter¬
mined as accurately as possible from the published literature. Five basic distri¬
bution patterns of the Pennfield Bog species are evident: 1) widespread
species, occurring throughout much of the United States and adjacent Canada
and often widespread in other parts of the world; 2) those occurring primarily
in the eastern half of the United States and adjacent Canada; 3) those which
occur in northeastern North America and also on the Coastal Plain; 4) those of
northeastern North America, primarily north of the glacial boundary; and 5)
northern, chiefly circumboreal species.

WIDESPREAD SPECIES

About 10% of the species, of which Typha latifolia (Map 1) is an ex¬
ample, occur widely throughout much of the United States. They are chiefly
plants of swamps and meadows, quiet waters, and shores. The majority of
these plants occur in the pioneer zones and only Typha latifolia is an im¬
portant constituent of the bog. Other species with this distribution include
Agrostis stolonifera, Ceratophyllum demersum, Glyceria striata, Lemna tri-
sulca, Monotropa uniflora, Lemna minor, Potamogeton gramineus, P. illinoen-
sis, Ranunculus sceleratus, Sagittaria latifolia, Scutellaria lateriflora, and Sium
suave. These widespread species probably extended their ranges northward as
the glaciers receded.

EASTERN UNITED STATES SPECIES

Approximately 26% of the species of the bog have distributions through¬
out the eastern half of the United States and adjacent Canada. These are spe¬
cies of shores, thickets, swamps and marshes, and woodlands which have also
extended their ranges northward with the retreat of the glaciers. Nuphar lute-
urn subsp. macrophyllum (Map 2), an important pioneer of the bog, illustrates
this pattern. Other species with this distribution include Acer rubrum, Agalinis
purpurea, Amelanchier arborea, Apios americana, Boehmeria cylindrica,
Brasenia schreberi, Carex comosa, Cephalanthus occidentalis, Cyperus di-
andrus, Decodon verticillatus, Dulichium arundinaceum, Echinocystis lobata,
Eleocharis compressa, Eupatorium perfoliatum, E. purpurea, Impatiens biflora,
Juncus effusus var. solutus, Leersia oryzoides, Mitchella repens, Nyssa sylva-
tica, Onoclea sensibilis, Osmunda regalis, O. cinnamomea, Par them cissus quin-
que folia, Polygonum hydropiperoides, P. sagittatum, Prunus virginiana,
Quercus velutina, Rosa palustris, Sambucus canadensis, Scirpus cyperinus,
Ulmus rubra, Viola affinis, Vitis vulpina, and Woodwardia virginica.

NORTHEAST-COASTAL PLAIN SPECIES

About 8% of the species occur in the Northeast and South along the
Coastal Plain. These are mostly plants of acid or peaty soils. Sarracenia purpu¬
rea (Map 3), a typical bog plant, illustrates the pattern. Most of these plants
have apparently migrated northward from the Coastal Plain into the newly

1969

THE MICHIGAN BOTANIST

53

Map 1. Distribution of Typha latifolia after Hotchkiss and Dozier, 1949.

Map 2. Distribution of Nuphar luteum subsp. macrophyllum after Beal, 1956.

54

THE MICHIGAN BOTANIST

Vol. 8

opened, glaciated northeast. Other species having this pattern are Drosera
intermedia, Habenaria ciliaris, H. clavellata, Hydrocotyle umbellata (chiefly a
Coastal Plain species), Isotria verticillata, Pogonia ophioglossoides, Toxico¬
dendron vernix, Triadenum virginicum, Utricularia gibba (chiefly a Coastal
Plain species), and U. purpurea.

The ranges of Habenaria ciliaris, H. clavellata, and Isotria verticillata also
extend southward in the Appalachian Mountains. These have probably sur¬
vived glaciation in the mountains as well as on the Coastal Plain.

NORTHEASTERN NORTH AMERICA SPECIES

Approximately 28% of the Pennfield Bog species are basically of a
northeastern North American distribution with nearly the entire range occur¬
ring within glaciated territory. The general habitats of this group of plants in¬
clude bogs, peaty soils, swamps and wet meadows. The distributions of these
plants strongly suggest that most of these species survived the Wisconsin glaci¬
ation in the Appalachians or on the Coastal Plain, not far from the ice front.

Nearly half of these species occur chiefly in the mountains south of the
glacial boundary and include such important species of the bog as Vaccinium
macrocarpon, Nemopanthus mucronata, Ilex verticillatus, and Andromeda
glaucophylla. The distribution pattern is represented by Rubus hispidus (Map
4). Other species which were apparently harbored by the mountains are
Aronia melanocarpa, Be tula lutea, Carex trisperma, Corylus americana,
Cypripedium acaule, Eriophorum virginicum, Gaultheria procumbens, Glyceria
canadensis, Iris versicolor, J uncus brevicaudatus, Pinus strobus, Ribes hirtell-
um, Sparganium chlorocarpum, Spiraea tomentosa, Trientalis borealis, and
Viburnum lentago.

Eleocharis smallii, Thelypteris palustris var. palustris, and Vaccinium
corymbosum are important constituents of the Pennfield Bog flora which
apparently persisted chiefly on the coastal plain just south of the glacier.
Rhynchospora alba (Map 5) illustrates the pattern. Others include Aster um-
bellatus, Betula pumila, Bidens coronata, Campanula aparinoides, Cyperus
engelmannii, Eleocharis intermedia, Epilobium strictum, Hypericum boreale,
Liparis loeselii, Nymphaea tuberosa, Ribes americanum, and Rumex orbicu-
latus.

Another possible refugium for a few plants with this basic distribution
pattern was an area to the southwest of the Great Lakes. Asclepias incamata
subsp. incamata. Spiraea alba (Map 6), and Viola nephrophylla appear to have
migrated into glaciated territory from this refugium. Spiraea alba and S. lati-
folia, which apparently survived the Pleistocene in the Appalachians, form a
complex centered in glaciated territory where the two taxa come together (for
a more detailed discussion of this complex, see Kugel, 1958).

NORTHERN SPECIES

About 28% of the species range throughout much of northern North
America; many of these are circumboreal in distribution. Most of them are

1969

THE MICHIGAN BOTANIST

55

Map 3. Distribution of Sarracenia purpurea after McDaniel, 1966.
Map 4. Distribution of Rubus hispidus after Braun, 1937.

56

THE MICHIGAN BOTANIST

Vol. 8

Map 5. Distribution of Rhynchospora alba after Hulte'n, 1958.
Map 6. Distribution of Spirea alba after Kugel, 1958.

1969

THE MICHIGAN BOTANIST

57

plants of bogs and peaty soils, shores, swamps, and marshes. Three of the
most important bog species, Carex lasiocarpa, Cassandra calyculata, and Larix
laricina, are in this category.

Nearly one-third of the northern species (9% of the total) occur south
of the glacial boundary only in the east and thus appear to have survived
glaciation in the Appalachians and the Coastal Plain. Larix laricina (Map 7) is
illustrative. Others include Alisma plantago-aquatica, Aralia nudicaulis, Bidens
cernua, Cassandra calyculata, Coptis trifolia var. groenlandica, Drosera rotundi-
folia, Habenaria obtusata, Ly copus uniflorus, Maianthemum canadense, Meny-
anthes trifoliata, and Rubus pubescens.

About half of the northern species of Pennfield Bog (14% of the total)
constitutive a western element. Most of the western species survived the
Pleistocene in the Rocky Mountains, as is indicated by the distribution of
Cornus stolonifera (Map 8). These taxa are restricted by the glacial boundary
in the East. Other species which probably survived in the Cordillera include
Carex aquatilis, C. interior, Muhlenbergia racemosa (litis, 1965), Myriophyllum
verticillatum, Populus tremuloides (litis, 1965 ), Potamogeton natans, Trigloch-
in maritima, Salix Candida (litis, 1965), S. pedicellaris (Fernald, 1925), S.
petiolaris and S. serissima (litis, 1965), and Urtica dioica var. procera.

The unglaciated interior of Alaska appears to have enabled many species
of the northern forests and of the arctic tundra to survive glaciation (Heusser,
1965). A few species now present in Pennfield Bog have probably migrated
from Alaska after the retreat of the ice. The distributions of Calla palustris
(Map 9), Carex chordorrhiza, and Equisetum fluviatile suggest such a migra¬
tion.

Several species of the bog cannot be accounted for by any of the pre¬
ceding patterns. Some of these are aquatics whose present ranges are entirely
within the glacial boundary and might be considered part of a periglacial ele¬
ment. It is also possible that some of these may have survived on nunataks in
and around Newfoundland, as Fernald (1925) has suggested, or in unglaciated
coastal refugia of the Pacific Northwest (Heusser, 1965). For some there is
insufficient information to allow an assessment of their geographical affinities.

The probable sources of species which now occur in Pennfield Bog are
summarized in Map 10. There appear to be four important sources from
which the bog flora has been derived: 36% of the species are southern and are
chiefly in pioneer zones or at the outer margin of the bog; 42% have an east¬
ern affinity; 10% are Cordilleran; and 2% are of Alaskan affinity.

SUMMARY

Pennfield Bog, Calhoun County, Michigan, lies in one of the many depressions of
the Kalamazoo morainal system. Field work was carried out from April to October,
1967. The bog consists of seven major plant zones: a Nuphar-Eleocharis zone, a
Decodon zone, a Carex- Vaccinium macrocarpon zone, a Cassandra zone, a Cassandra—
Thelypteris zone, a Larix zone, and an outermost Acer rubrum zone. Frequency, cover,
and importance values have been tabulated for each vegetational zone.

58

THE MICHIGAN BOTANIST

Vol. 8

Map 7. Distribution of Larix laricina after Little, 1965.

Map 8. Distribution of Cornus stolonifera after Rickett, 1944.

1969

THE MICHIGAN BOTANIST

59

Map 9. Distribution of Calla palustris after Hulte'n, 1962.

Map 10. Migration streams represented in Pennfield Bog and extent of Wisconsin glaciation (glacial boundary after
Flint, 1957, 1959).

60

THE MICHIGAN BOTANIST

Vol. 8

A total of 144 species of vascular plants was collected. Their North American
ranges have been determined from the literature. Five basic patterns are represented by
the bog species: 1) widespread species; 2) eastern United States species; 3) Northeast-
Coastal Plain species; 4) northeastern North American species; and 5) northern species.
Postglacial migration from Pleistocene refugia may account for the present flora. The bog
flora appears to have been derived from four important sources: 36% of the species have
southern affinities, 42% have eastern affinities, 10% represent a Cordilleran element, and
2% are of Alaskan affinity.

AC KNOWLEDGM ENTS

In addition to the acknowledgments in the previous article, I wish to thank the
Department of Botany and Plant Pathology, Michigan State University, for financial sup¬
port for publication expenses. The maps are based on Goode Base Map No. 102, copy¬
right by The University of Chicago, and are used by permission.

LITERATURE CITED

Beal, E. O. 1956. Taxonomic revision of the genus Nuphar Sm. of North America and
Europe. Jour. Elisha Mitchell Sci. Soc. 72: 317-346.

Braun, E. L. 1937. Some relationships of the flora of the Cumberland Plateau and Cum¬
berland Mountains in Kentucky. Rhodora 39: 193-208.

Braun, E. L. 1951. Plant distribution in relation to the glacial boundary. Ohio Jour. Sci.
51: 139-146.

Crow, G. E. 1969. An ecological analysis of a southern Michigan bog. Mich. Bot. 8:
11-27.

Fernald, M. L. 1925. Persistence of plants in unglaciated areas of boreal America. Mem.

Am. Acad. 15: 239-342 [also Mem. Gray Herb. 2).

Flint, R. F. 1957. Glacial and Pleistocene Geology. John Wiley, New York. 553 pp.

Flint, R. F., et al. 1959. Glacial Map of the United States East of the Rocky Mountains.
Geol. Soc. Am., New York.

Heusser, C. J. 1965. A Pleistocene phytogeographical sketch of the Pacific Northwest and
Alaska, pp. 469-483 in The Quaternary of the United States, ed. by H. E. Wright & D.
Frey. Princeton Univ. Press, Princeton.

Hotchkiss, N., & H. L. Dozier. 1949. Taxonomy and distribution of N. American cat¬
tails. Am. Midi. Nat. 41: 237-254.

Hulte'n, E. 1958. The Amphi-Atlantic Plants and their Phytogeographical Connections. Sv.
Vet-akad. Handl. IV. 7(1). 340 pp.

Hulte'n, E. 1962. The Circumpolar Plants. I. Vascular Cryptogams, Conifers, Monocoty¬
ledons. Sv. Vet.-akad. Handl. IV. 8(5). 275 pp.
litis, H. H. 1965. The genus Gentianopsis (Gentianaceae): transfers and phytogeographic
comments. Sida 2: 129-154.

litis, H. H. 1966. The western element in the eastern North American flora and its phyto¬
geographic implications. (Abstract) Am. Jour. Bot. 53: 634.

Kugel, A. R. 1958. Variation in the Spiraea alba-latifolia Complex. Unpublished Ph.D.
thesis, Univ. Mich. 124 pp.

Little, E. L. 1965. In H. A. Fowells, Silvics of Forest Trees of the United States. U. S.
Dep. Agr. Handb. 271. 762 pp.

Martin, P. S. 1958. Pleistocene ecology and biography of North America, pp. 375-420 in
C. S. Hubbs (ed.), Zoography. Am. Assoc. Advancem. Sci. Publ. 51.

McDaniel, S. 1966. A Taxonomic Revision of Sarracenia (Sarraceniaceae). Unpublished
Ph. D. thesis, Univ. Florida.

Rickett, H. W. 1944. Comus stolonifera and Comus occidentalis. Brittonia 5: 149-159.
Whitehead, D. R. 1965. Palynology and Pleistocene phytogeography of unglaciated east¬
ern North America, pp. 417-432 in The Quaternary of the United States, ed. by H. E.
Wright & D. Frey. Princeton Univ. Press, Princeton.

Wolfe, J. H. 1951. The possible role of microclimate. Ohio Jour. Sci. 51: 134-138.

1969

THE MICHIGAN BOTANIST

61

WHITE PINE AT THE EDGE OF
ITS RANGE, IN OAKLAND COUNTY, MICHIGAN

Sylvia Taylor

Department of Botany, The University of Michigan, Ann Arbor

In eastern Lower Michigan, white pine ( Pinus strobus ) occurs naturally
no farther south than Oakland County. In much of the northeast quarter of
that county it occurs as widely scattered single trees or in small groves among
hardwoods, and is a familiar volunteer tree as well as a planted tree around
lakefront cottages. In other areas of the county, individuals or groups of a few
trees occur locally on hummocks in swamps and these occasionally seed into
bordering sandy uplands.

A different view is expressed in late nineteenth century accounts, which
set the southern boundary in Lapeer County, the next county to the north
(Wheeler, 1898; Spalding, 1899). Those observations, however, were probably
more concerned with merchantable timber than mere occurrence of white
pines, and overlooked branchy, slower growing members of the species in Oak¬
land County. An estimate made just before the turn of the century of 20
acres of standing pine for the whole county (Morse, as quoted by Wheeler,
1898) may reflect similar thinking, for now, even after much clearing of land
and lowered water tables, many times that area supports at least a mixture of
white pine and other trees. The actual range of white pine probably has re¬
mained about the same since presettlement times. Sight records made by the
author (1964-1967), of individual stands appearing to be naturally seeded are
shown in Fig. 1 .

Widely separated outlying stands of white pine occur on the interlobate
morainic area extending west-southwest through southern Oakland County,
where the surface is a mosaic of hills and lakes amid varied glacial and post¬
glacial deposits. Observations of one such stand in Springfield Township pro¬
vide clues to the occurrence and persistance of the species on these sites.

White pines occur in about 15 acres of swamp and adjacent old fields
southeast of Davisburg, between Clark and Andersonville roads, NEJ/4, sec. 21,
T4N, R8E, Springfield Township (Fig. 2). A dozen large pine trees approxi¬
mately 90-100 years old are scattered through a mixed hardwood stand which
rims a shrubby sphagnum swamp.

In the swamp several dozen younger pines up to about 70 years of age
are growing on broad, low hummocks a little higher than the general surface.
These trees are about 50 yards apart and are generally shorter and more wide¬
ly separated near the center of the swamp.

On the east side of the swamp-fringing hardwood stand, a sandy field,
last cultivated twenty-five years ago, has been invaded by a nearly pure stand
of very robust young white pines (Fig. 3). These, in turn, are associated with
equally robust patches of colic-root (Aletris farinosa), trailing arbutus ( Epigaea
repens ), and, in season, the edible mushrooms Suillus brevipes and Suillus
granulatus (which possibly forms mycorrhizae with the pines).

62

THE MICHIGAN BOTANIST

Vol. 8

Local residents claim that a great number of white pines once grew on
the area now occupied by the old field, pasture, and adjacent railroad right-
of-way southeast of Davisburg, but were clear-cut in the mid 1800’s when the
Grand Trunk Railroad was laid. At this time the pine population may have
been nearly wiped out, as there are indications that the present swamp was
then a marsh or bog and has only recently been invaded by trees. The only
surviving pines, after the extensive cutting, would have been seedlings or sap¬
lings restricted to a narrow border between field and water.

As hydrarch succession proceeded in the marsh, seedling pines apparent¬
ly were able to become established on hummocks. More recent abandonment
of the formerly cultivated sandy field has allowed old-field succession of pine
similar to that long familiar in central New England (Raup, 1964). The popu¬
lation continues to expand at present in spite of grass fires, Christmas tree
hunters, and a series of abnormally dry years.

R 7 E R 8 E R9E R 10 E R 1 1 E

Fig. 1. Sight records of white pine stands in Oakland County, Michigan, 1964-1967.

1969

THE MICHIGAN BOTANIST

63

The aggressiveness of these trees in an area detached from the main
range of the species may seem surprising to persons who are inclined to regard
plants of more northern range growing in southern Michigan swamps as glacial
relicts. Local conditions associated with this particular occurrence, however,
provide a favorable habitat in which white pine may not only persist, but
achieve dominance.

Three miles northeast, at Bridge Lake, another group of pines exists on
a wet slope in association with white cedar ( Thuja occidentalis). Only a few
scattered individuals and small groups occur farther south in the country.

It has long been recognized that throughout the Lake States white pine
occurs commonly on sand and near water (Spalding, 1899). Comparison of
the Oakland County distribution of this species with a map of Michigan sur¬
face formations (Martin, 1955) shows a high correlation with parent materials
which are usually sandy in character. Even those pines growing locally on or¬
ganic soil in bogs are near sandy soil, possibly seeded there from upland trees
removed in the last century. Although white pine grows well on a wide variety
of soil types (Frothingham, 1914), competition from hardwoods has tended to
restrict it to well drained, sandy soils where it may persist as a physiographic
climax (Wilson & McQuilkin, 1963). The sandy old field which is presently
being invaded by white pine and which may formerly have supported a ma¬
ture group of these trees appears edaphically suited to maintain such a stand.
Those growing in the richer woods and swamp, however, may be regarded as
successional intermediates in a hardwood forest (Oosting, 1956, p. 251).

In addition to favorable soil, white pines require a cool, humid climate
with a moisture surplus in all seasons (Wilson & McQuilkin, 1963). The nu¬
merous lakes and slight elevation of the land (averaging about 1000 ft. above
sea level) may contribute to this southern extension in the limit of the species
in Oakland County. Predictably, the most southern .outliers are found in
swamps and around lakes.

The pines in the old field are between a marsh and swamp which are
part of the headwaters of the Shiawassee River. The swamp is characterized
by slow drainage and cold water. On a hot summer afternoon one may stand
among the old-field pines in the shade of the woods, and feel a cool breeze
filtering through the trees from across the swamp. A short hike to the west
side of the swamp leads to another sandy old field which has been invaded by
a lone pine and numerous aspen (Populus tremuloides). Here, an equal dis¬
tance from the water, one feels only the hot afternoon sun. In addition to
afternoon shade and a position downwind from the swamp, the field on the
east side is downwind from the large pines near the edge of the woods. Seed
from these trees falls into patches of Polytrichum moss which serve as ideal
seedbeds for germination and development of new plants (McQuilkin, 1959).

In the marsh, bordering hardwoods, and adjacent abandoned field, other
plants which indicate a greater than usual clustering of more northern species
include white birch ( Betula papyrifera ), yellow birch ( Betula lutea ), black
spruce {Picea mariana), club-moss ( Lycopodium dendroideum), low-bush blue¬
berry ( Vaccinium sp.), wintergreen ( Gaultheria procumbens), and

64

THE MICHIGAN BOTANIST

Vol. 8

Fig. 2. Location of Springfield Township white pine stand (dashed line) in relation to
surrounding features.

1969

THE MICHIGAN BOTANIST

65

partridge-berry ( Mitchella repens). They are characteristic of a forest type
which is common farther north but unusual in this region, the white pine-
hemlock— northern hardwoods type.

If left undisturbed this small parcel of vegetation may mature into a lo¬
calized duplication of a forest type well known to early settlers, and produce
rivals of the magnificent white pines of the past which were worthy of being
called our State Tree. It already contains the densest cover of trailing arbutus
and colic-root I have seen in southern Michigan. The future threat to these
white pines is not so much a matter of their position at the southern limit of
their natural range as it is their position on the northern limit of the expand¬
ing Detroit Metropolitan area. A sign on the property reads: “For Sale.”

Fig. 3. Springfield Township
white pine stand, January 2,
1969; facing west on north
side of fence between pasture
and old field.

LITERATURE CITED

Frothingham, E. H. 1914. White Pine under Forest Management. U. S. Dep. Agr. Bull.
13. 70 pp.

Martin, H. M. 1955. Map of the Surface Formations of the Southern Peninsula of Michi¬
gan. Mich. Geol. Surv. Publ. 49.

66

THE MICHIGAN BOTANIST

Vol. 8

McQuilkin, W. E. 1959. Effect of site and stocking on reproduction, pp. 16-19 in What’s
Known About Managing Eastern White Pine. Northeastern For. Exp. Sta., Sta. Pap.
121.

Oosting, H. J. 1956. The Study of Plant Communities, 2nd. ed. Freeman, San Francisco.
440 pp.

Raup, H. M. 1964. Some problems in ecological theory and their relation to conservation.
Jour. Ecol. 52, suppl.: 19-28.

Spalding, V. M., & B. E. Femow. 1899. The White Pine (Pinus Strobus L.) U. S. Dep.
Agr. Forestry Bull. 22. 185 pp.

Wheeler, C. F. 1898. A sketch of the original distribution of white pine in the lower
peninsula of Michigan. Mich. Agr. Exp. Sta. Bull. 162: 4-5.

Wilson, R. W., & W. E. McQuilkin. 1963. Silvical Characteristics of Eastern White Pine. U.
S. For. Serv. Res. Pap. NE-13. 29 pp.

Publications of Interest

From time to time we call attention, under this heading, to new publications
which might be of particular interest to some of our readers but which, because of less
explicit geographic or botanical orientation, do not qualify for regular review or inclusion
in the listings of new literature relating to Michigan botany. All annotations are by the
Editor unless ascribed to another.

MAPS OF MICHIGAN AND THE GREAT LAKES 1545 - 1845 from the private collec¬
tion of Renville Wheat with additions from the Burton Historical Collection. Cata¬
logue of an Exhibition in the Detroit Public Library October 13 to December 3,
1966. Burton Historical Collection, Detroit Public Library, 1967. 44 pp. Twelve of
the 65 maps catalogued are reproduced, much reduced, and show the development of
knowledge about the configuration of the Great Lakes as well as the antiquity of
many place names.

THE QUATERNARY OF ILLINOIS. A Symposium in Observance of the Centennial of
the University of Illinois. Edited by Robert E. Bergstrom. University of Illinois Col¬
lege of Agriculture Special Publ. 14. 1968. $5.00 from College of Agriculture, 123
Mumford Hall, Urbana, Ill. 61801. This volume represents prompt publication of the
papers presented February 12-13, 1968, at a symposium sponsored by the University
of Illinois, Illinois State Geological Survey, and Illinois State Natural History Survey.
Several of the papers are on the ‘Prairie Peninsula’ and other biogeographical phenom¬
ena.

VASCULAR PLANTS OF OHIO. A Manual for Use in Field and Laboratory. By Clara G.
Weishaupt. Revised ed. Wm. C. Brown Book Co., Dubuque, Iowa, 1968. 280 pp.
$7.50. A comprehensive set of field-tested keys to the Ohio flora; no illustrations nor
material on ecology or distribution. Over-priced for a paper-bound, lithoprinted vol¬
ume, but should be useful in southern Michigan.

SCIENCE FOR BETTER LIVING. The Yearbook of Agriculture 1968. 386 pp. A com¬
pendium of recent research relating to agriculture, marketing, and various topics of
interest to the consumer. A chapter of particular local interest on cherry harvesting
has several references to the Michigan industry and cites the first mechanical tart
cherry harvester, introduced in Michigan in 1959. This yearbook is available for $3.00
from the Superintendent of Documents (Washington 20402) or without charge from
one’s congressmen.

[More on page 107]

1969

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67

THE OCCURRENCE OF THE LICHEN COMPLEX,
CETRARIA CILIARIS, IN THE STRAITS REGION OF MICHIGAN

William L. Graham

Department of Botany, The University of Michigan, Ann Arbor

Investigators have long known of morphologically identical but chemical¬
ly distinct strains of lichens. Well-known examples include Cladonia chloro-
phaea, Parmelia cetrarioides, and some groups in which slight differentiation
may be demonstrated, as in the Parmelia ambigua-P. hyperopta complex,
where the cortex color is different but the morphology is similar. Although
Nylander’s use (1866) of potassium hydroxide and calcium hypochlorite to
differentiate taxa helped to strengthen the use of chemistry in the taxonomy
of lichens, many lichenologists have continued to be skeptical of using chemi¬
cal characters to delimit species. In recent years monographic treatments have
stressed chemistry as a valid criterion in delimiting lichen taxa, but workers
often remain ambivalent when chemical variation does not closely coincide
with more standard characters, such as ecology, geography, morphology, etc.
Our lack of understanding completely the bases of chemical variation makes
most investigators very cautious of using chemical differences as the sole crite¬
rion upon which to base a taxon.

Cetraria ciliaris Ach. ( sensu lato )* provides an excellent example of a
species which can be subdivided geographically and chemically. Various lichen
substances may be represented in the medulla in some parts of the range:
protolichesterinic acid, a fatty acid; alectoronic acid, a depsidone; olivetoric
and microphyllinic acids, depsides. (Depsides and depsidones are orcinol deriv¬
atives of phenylcarboxylic acids.) The microphyllinic acid population is found
only on the island of Hokkaido, Japan. The other acid strains are found in
North America, northern Europe, central Asia, Siberia, and Japan (Culberson
and Culberson, 1967). From herbarium specimens, Hale (1963) plotted the
distribution of the acid populations in the United States and Canada. He also
performed field studies in the central Appalachian region, where the popula¬
tions converge. Although protolichesterinic acid is present, the alectoronic and
olivetoric acid populations greatly predominate in that region. In the extreme
western United States the only acid found is protolichesterinic, but all three
are present in the eastern United States and southeastern Canada.

In 1967, William L. and Chicita F. Culberson gave specific rank to the
four chemical elements. Cetraria ciliaris Ach. sensu stricto contains olivetoric
acid as a diagnostic essential; C. orbata (Nyl.) Fink contains protolichesterinic
acid; C. halei Culb. & Culb. and C. microphyllica Culb. & Culb. contain
alectoronic acid and microphyllinic acid, respectively. A complete discussion

lC. ciliaris consists of several elements. These have been variously termed “strains”
and “chemical species.” As will later be shown in this paper, these elements have most
recently been termed species. This work will follow that terminology, except when noted
otherwise.

68

THE MICHIGAN BOTANIST

Vol. 8

of the chemistry and phytogeography of these plants is in the papers of Hale
and the Culbersons.

Richard Harris, of Michigan State University, introduced me to the C.
ciliaris complex, and especially to certain questions surrounding the distribu¬
tion of the acids in the lichen in the region of the University of Michigan
Biological Station near Pellston, Michigan, near the Straits of Mackinac. In
that summer of 1967 I became curious about ecological factors which might
affect the distribution of C. ciliaris.

Many corticolous (bark-inhabiting) lichens have fairly specific substrate
preferences. My observations were that C. ciliaris is essentially limited to
conifers in the area of the Biological Station. Previous work has shown that
the species is a pioneer, growing commonly on younger branches and high in
the trees, but only occasionally on old boles. Studies of vertical distribution
of cryptogams are few. Hale’s earlier study (1952) was performed in a
Thuja- Acer- Be tula association; his later one (1965) in a hardwood swamp
dominated by Acer rubrum, and the studies are not therefore necessarily
comparable. In the region of the Straits of Mackinac the plants grow most
commonly on Abies balsamea in Thuja swamps, on Larix laricina in open
bogs, and on Pinus banksiana in the dry uplands. I have also found occasional
specimens on Acer rubrum, Aims rugosa, Juniperus communis, Picea glauca,
P. mariana, Pinus resinosa, P. strobus, Populus sp., and Thuja occidentalis.
Studies of herbarium labels indicate that the lichen also grows on Betula,
Malus, and Prunus and is occasionally saxicolous (rock-inhabiting).

In northern Michigan, C. ciliaris is common enough to make sampling
for the elements in this complex possible and satistically valid. I chose samples
by walking in a straight to erratic manner through a stand of trees, gathering
thalli at convenient heights. In areas which were difficult to move about in,
fewer trees may have been sampled than in other stands, even though there
may have been a tremendous number of thalli present. The sampling error in
areas the size of which I studied does not exceed 5% (Hale, 1963). No trees
were climbed, but treefalls were sampled when available. Approximately 200
thalli were collected in each stand. A total of 1,896 thalli were gathered in
the study (cf. Table I).

METHODS OF ANALYSIS

In the summer of 1967 I used the potassium hydroxide— sodium hypo¬
chlorite (KC) test on the exposed medulla of the thallus. KC+ orange was as¬
sumed to indicate alectoronic acid, KC+ red was assumed to indicate olive-
toric, and KC- was assumed to indicate protolichesterinic acid. This prelim¬
inary survey led me to believe that there was need for much more work with
larger collections. In 1968 I continued with a more efficient technique. Hale
(1956) had noted that lichen depsides and depsidones fluoresced under ultra¬
violet light (UV) supplied them at wavelengths between 3200 and 4000
angstrom units. Alectoronic acid reacts strongly under these wavelengths,
fluorescing a brilliant bluish-white.

1969

THE MICHIGAN BOTANIST

69

Thalli from a single gathering were spread on a black table in a dark¬
room and exposed to a UV lamp, and their cortices were scratched to expose
the medullae. Those thalli of unquestionable fluorescence in the manner of
alectoronic acid were set aside as C. halei. Those of absent or ambiguous flu¬
orescence were tested microchemically as follows: Those of questionable but
obvious fluorescence were checked for alectoronic acid by extracting the
lichen substances with acetone on a warm microslide. GAW (1:1:1 glycerol,
ethanol, and water) was used as a recrystallizing agent. The slides were
warmed with the reagent and set aside until the diagnostic crystals had
formed; they were then examined microscopically to determine the shape of
the acid crystals which appeared. Those which did not exhibit fluorescence in
UV were tested as above in GE (1:1 glycerol and acetic acid) to determine if
protolichesterinic acid was present and in GAW to look for olivetoric acid.
The latter were also tested with a commercial chlorine bleach (5% sodium
hypochlorite) in the medullae. If they reacted C+ red, olivetoric acid was as¬
sumed to be present.

RESULTS

In the following sections, nomenclature follows that of Culberson and
Culberson (1967). Table I presents an analysis of the distribution of the spe¬
cies on various substrata. Out of a total of 1,896 specimens, all could be re¬
ferred to C. halei with the exception of eight C. orbata thalli and two of C.
ciliaris sensu stricto.

Fig. 1. The distribution of Cetraria halei in Michigan, showing counties from which it has
been collected.

Fig. 2. The distribution of Cetraria ciliaris sensu stricto and C. orbata in Michigan, show¬
ing localities from which they have been collected.

70

THE MICHIGAN BOTANIST

Vol. 8

TABLE I. Occurrence of the Species on Various Substrata.

Substratum

Number and Percentage of Thalli

C. halei

C. orb at a

C. ciliaris

Total

Abies balsamea

319

98%

4

1.2%

2

0.6%

325

Larix laricina

454

100%

0

0%

0

0%

454

Picea mariana

431

100%

0

0%

0

0%

431

Pinus banksiana

509

99.8%

1

0.2%

0

0%

510

Unrecorded: Abies,
Larix, or Picea

173

98%

3

2%

0

0%

176

Total

1,886

99.5%

8

0.4%

2

0.1%

1,896

Collections were made at seven localities given in Table II. The voucher
specimens mentioned in the table are on file in the Lichen Herbarium of the
University of Michigan Biological Station. All collection numbers are mine un¬
less otherwise noted. (Table I does not include data from locality five below,
which was sampled by a class in lichenology, because of data loss. This col¬
lection included an additional thallus of C. ciliaris sensu stricto, not statistical¬
ly considered in Table I.)

The geographic distribution of the Cetraria ciliaris group in Michigan is
shown in figures 1 and 2, based on data presented in this paper and from
herbarium specimens at the University of Michigan (MICH) and Michigan State
University (MSC).

TABLE II. Data for Voucher Collections.

Collection Area

Coll. No.

Species

Substratum

1. Cheboygan Co., Indian River,

Sec. 33, T35N, R2W

L134

C. halei

Pinus banksiana

2. Cheboygan Co., Livingston Bog,
Sec. 2, T36N, R3E.

L139

s.n.

C. halei

C. halei

Larix laricina

Picea mariana

3. Cheboygan Co., Mud Lake Bog,

Sec. 7, T37N, R3W.

L138

L137

C. halei

C. halei

Larix laricina

Picea mariana

4. Cheboygan Co., Reese’s Bog,

Sec. 4, T36N, R3W.

s.n.

s.n.

L140

C. halei

C. orbata

C. ciliaris

Abies balsamea
Abies balsamea
Abies balsamea

5. Cheboygan Co., Douglas Lake,
UMBS, Sec. 33, T37N, R3W.

L132

L131

L130

C. halei

C. orbata

C. ciliaris

Pinus banksiana
Pinus banksiana
Pinus banksiana

6. Montmorency Co., Hillman,

Sec. 10, T31N, R4E.

Hale, s.n.

C. orbata

Pinus banksiana

7. Presque Isle Co., Evergreen

Beach, Sec. 21,T36N, R4E.

L136

L135

C. halei

C. orbata

Unrecorded: Abies,
Larix, ox Picea

1969

THE MICHIGAN BOTANIST

71

DISCUSSION

The species of the C. ciliaris complex are (seemingly) randomly scattered
in all communities which occur within areas in which the species are sym-
patric. No differing ecological requirements have been shown to be attributa¬
ble to one species or the other in this or other works. This type of investiga¬
tion must be undertaken in areas in which the three species occur sympatrical-
ly, such as the region of the central Appalachians investigated by Hale in
1963. However, in his work there, C. orbata was represented by only 0.3% of
the thalli collected (cf. 0.4% in my study near the Straits region of Michigan)
and would prevent any ecological or microenvironmental study of its distribu¬
tion.

In the Appalachian region the percentage of C. halei varied from 0 to
96, and was consistently dominant (96% to 70%) only in the shale barrens in
Maryland and West Virginia (cf. Hale, 1963, esp. o. 130). In the Straits region
of Michigan C. halei is consistently dominant, for C. ciliaris represents no
more than 0.1% of the population. C. orbata is represented by approximately
the same percentage in both regions.

The thalli of these species produce no effective vegetative propagules,
such as isidia or soredia, and the diaspores are the fungal ascospores and
conidia. The predominant means of dispersal of this species is probably by
means of ascospores. If this is so, the distribution of these species may be a
reflection of physiological and genetical differences acting upon or with the
environment. On the other hand, the little investigated, but important effects
of pleiotropy (the control of several paths of development by one gene, or of
several phenotypes by one gene) must receive more consideration in what we
call “chemical taxonomy” in any group of plants.

ACKNOWLEDGMENTS

I wish to thank Howard Crum, Mason E. Hale, Jr., and Richard Harris for their
intense and continued encouragement and assistance throughout this study. I also thank
Barbara Moore and Dale Vitt for their aid, and Melinda Denton and Rudolf Schmid for
their critical reading of the manuscript.

LITERATURE CITED

Culberson, W., & C. Culberson. 1967. A new taxonomy for the Cetraria ciliaris group.
Bryologist 70: 158-166.

Hale, M. 1952. Vertical distribution of cryptogams in a virgin forest in Wisconsin. Ecolo¬
gy 33: 398-406.

Hale, M. 1956. Fluorescence of lichen depsides and depsidones as a taxonomic criterion.
Castanea 21: 30-32.

Hale, M. 1963. Populations of chemical strains in the lichen Cetraria ciliaris. Brittonia 15:
126-133.

Hale, M. 1965. Vertical distributions of cryptogams in a Red Maple swamp in Connecti¬
cut. Bryologist 68: 193-197.

Nylander, W. 1866. Circa novum in studio lichenum criterium chemicum. Flora 66:
198-201.

72

THE MICHIGAN BOTANIST

Vol. 8

SYSTEMATIC AND ECOLOGICAL STUDIES

ON PL ANT AGO CORD AT A

Melvern F. Tessene

Department of Botany and Botanical Gardens,

The University of Michigan, Ann Arbor

The genus Plantago L., consisting of 250-275 species, is represented by
one or more species in most parts of the temperate regions of the world. Al¬
though Plantago is often considered a weedy genus, in reference to such
“camp-followers” as the common plantains, P. major L. and P. lanceolata L.,
in reality over 90% of the species are of restricted distribution. Some species,
such as the four which occur only at the summit of Mt. Waialeale, Hawaii, are
endemic to very small areas and are highly specialized ecologically. Other spe¬
cies have broader geographical ranges but are not common anywhere. Plantago
cordata Lam. is in the latter category and is surely the rarest of the North
American plantagoes.

My study of P. cordata was made in connection with a long-range study
of the broad evolutionary relationships of the genus Plantago. Pilger’s (1937)
monograph of the Plantaginaceae recognized 19 sections in Plantago. The sec¬
tion Paleopsyllium, in which he included P. cordata, was considered a natural
group and the least specialized of the genus. My own studies indicate, how¬
ever, that Paleopsyllium is actually a potpourri of quite unrelated and special¬
ized species. In reviewing this and the other taxonomic sections as used by
Pilger, I have tried to obtain living material of as many species as possible.
The first population of P. cordata was not found until a year after I had be¬
gun an intensive search for the species. The difficulty in locating extant popu¬
lations and the apparently subtle ecological requirements of P. cordata under
natural conditions posed several problems which this paper attempts to re¬
solve. The rarity of the species, and perhaps its imminent extinction, call for a
thorough study of it at this time.

MATERIALS AND METHODS

Living plants were obtained from the following stations: (1) Plum Run
Quarry, 2 mi E of Peebles, Adams Co., Ohio ( Tessene 1881)', (2) Farm of Mr.
Harvey Lockney, ca. 1.5 mi NW of Thedford, Bosanquet twp., Lambton Co.,
Ontario ( Tessene 1941)', (3) Stockport Landing, Columbia Co., N. Y. ( Tessene
2065); (4) Small stream leading into NE side of Lake Murphy sboro, Jackson
Co., Ill. ( Tessene 2066); (5) 3 mi NW of Denton, Flat Swamp Creek, Davidson
Co., N. C. ( Stephenson 618); (6) Chekomeko Creek, Columbia Co., N. Y.
( Hines 6825); (7) Messenger Woods, 2 mi N of New Lenox, Willis Co., Ill.
( Tessene 2176). Seeds were supplied by Dr. I. John Bassett from 3 mi W of
Millcreek, Madison Co., Mo. ( Bassett & Spicer 4501). All specimens were
planted in pots and grown under uniform conditions at the University of
Michigan Botanical Gardens. These plants served as stock cultures for the

1969

THE MICHIGAN BOTANIST

73

ecological and systematic studies. Field observations and ecological data were
recorded in the field at the time of collection. In addition, specimens from a
number of herbaria were examined. Standard techniques were used in the
cytological and histological studies.

Voucher specimens for chromosome counts, morphological variation,
and drawings will be deposited in the Herbarium of the University of Michigan
(MICH).

DESCRIPTION OF THE SPECIES

Plantago cordata was originally described by Lamarck in 1791 from ma¬
terial at the Jardin Botanique in Paris grown from seeds attributed only to
“Canada.” The type specimen (P, photo MO! DAO) is unfortunately a mix¬
ture of the leaves of P. cordata and the spike of P. major. Although the mixed
collection was probably an accident, the two species are often confused espe¬
cially in the vegetative state. The superficial resemblance of the large, expand¬
ed phyllodes of P. cordata, P. major, and P. rugelii (Fig. 1) may lead the casual
observer to believe the three species are closely related. However, the gross
morphology and the anatomy of the flowers and seeds do not support this
hypothesis. The specialized fleshy roots and usually hollow mature peduncle
of P. cordata, as well as its phenology and its ecology, show it to be the most
divergent of the three species and at most, only distantly related. The follow¬
ing key will distinguish P. cordata from other broad-leaved species of the
genus in Eastern North America.

1. Leaves cordate to broadly elliptic, with petiole and blade clearly differenti¬
ated; sepals and bracts glabrous

2. Major roots massive and fleshy, 0.5-1. 3 cm in diameter; peduncle 3-4 mm
thick, with a central cavity usually present and equalling 3/4 or more of
total diameter; seeds 2 (-4) per capsule; major veins of leaf not parallel to

margin, appearing to arise from “mid-rib”; leaves leathery . P. cordata

2. Major roots cord-like to filamentous, 0.1-0. 3 cm in diameter; peduncle 1-2
mm thick, with central cavity, if present, less than 1/3 of total diameter;
seeds 4-12 per capsule; major veins of leaf parallel to margin; leaves thin

3. Capsules oblong, dehiscing near base; seeds black, 4-9 per capsule; bracts
and sepals lanceolate-attenuate; leaves broadly elliptic, the margins bearing

3-7 small teeth, glabrous . P. rugelii

3. Capsules rhombic-ovate, dehiscing medially; seeds brown, 8-12 per cap¬
sule; bracts and sepals ovate-obtuse; leaves cordate-ovate, the margins entire,

glabrate or hirsutulous . P. major

1. Leaves spatulate to linear, with petiole and blade poorly if at all differenti¬
ated; sepals and/or bracts usually pubescent

4. Leaves glaucous, glabrous, erect; roots fleshy; bracts and sepals ovate-obtuse

. P. eriopoda

4. Leaves not glaucous, hirsutulous; roots thin; bracts and sepals lanceolate-

attenuate . P. media

Plantago cordata Lam. Tabl. Encycl. Meth. 1: 338. 1791.

P. kentuckensis Michx. FI. Bor. -Am. 1: 94. 1803.

P. multiplinervia Steud. Flora 32: 410. 1849.

Fig. 1. Habit drawings of Plantago cordata and superficially similar species. A) P. major,
B) P. rugelii ; C) P. cordata ; a,b,c) Leaves of respective species.

1969

THE MICHIGAN BOTANIST

75

Plants herbaceous perennials. Major roots massive and fleshy, branching several
times, 0.5-1 .3 cm thick. Minor roots sparse, fibrous. Leaves (Fig. 2) in winter
rosette broadly spatulate-acuminate, 1-3 cm wide; in late spring and summer
cordate-ovate but never truly cordate, 8-19 cm wide, 12-23 (-50) cm long;

Fig. 2. Silhouettes of Plantago cordata leaves collected Sept. 15, 1967, at Stockport
Landing, N.Y. The largest leaf drawn is 7 inches wide.

76

THE MICHIGAN BOTANIST

Vol. 8

major veins (3) 5,7 (9), not parallel to margin, appearing to arise from a “mid¬
rib” 1/3-1 /2 of the way from the base of the blade; margin entire or undulate;
lamina with a leathery texture, frequently drying black or dark-brown; petiole
base often purple. Inflorescence a spike, its total length including peduncle
12-60 cm, glabrous; a central cavity usually present and equalling 2/3 of total
diameter of peduncle; flowers 2 -4 per cm at center of spike, becoming scat¬
tered at basal and distal ends of inflorescence. Flowers (Fig. 8) perfect, proto-
gynous, obovate in outline; sepals 4, free and distinct, 2-2.3 mm long; bract
solitary, glabrous, resembling sepals, obovate, pigmented central area bordered
by a very narrow hyaline margin, tip truncate; petals 4, connate basally, lobes
2-3 mm long, deltoid, chartaceous, light brown, appressed to sepals at anthe-
sis; stamens 4, epipetalous, filaments 0.9-2. 3 cm long; anthers versatile, pale
yellow and/or purple, horned at basal end, pollen copious; gynoecium superi¬
or, bicarpellate; stigma with two parallel rows of hairs extending from below
corolla to slightly bifid tip. Capsule (Fig. 3) a pyxis, ovate, 9-11 mm long,
circumscissile sub-medially, often retaining old corolla when mature. Seeds
(Fig. 3) 2 (rarely 3-4) per capsule, 3-3.8 mm long, tan-brown; testa smooth,
mucilaginous covering heavy, planar area conspicuous on one end of seed, ob¬
scured on the other; hilum and micropyle separated.

Fig. 3. Reproductive structures of Plantago cordata. A) Seeds; B) Seeds connected to
placental septum as shed from capsule; C) Medial cross-section of seed (drawn to slightly
larger scale).

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THE MICHIGAN BOTANIST

77

LEAVES

Most manuals and floras which include P. cordata (e.g. Small, 1933;
Fernald, 1950; Gleason, 1952; McVaugh, 1958; Fassett, 1959; Gleason &
Cronquist, 1963; Steyermark, 1963) distinguish it from other members of the
genus by the venation of the leaf. In P. cordata, lateral veins supposedly arise
from a midvein or midrib rather than from the base of the leaf as is more
common in the genus. Although the false midrib is present in many mature
specimens of P. cordata (Figs. 1, 4, & 10), it is not invariably present nor, on
any individual plant, is it present on all leaves or at all times of the year. The
emphasis on this character is particularly unfortunate as the “midribbed” leaf
usually does not develop until after the plant has finished flowering and fruit¬
ing. Therefore, at certain times, specimens of P. cordata are identifiable to
genus but the keys will not always clearly distinguish the species. This confu¬
sion is, perhaps, one reason why P. cordata has been questioned as a “good
species” by those who have never seen it and why it may be overlooked or
ignored by collectors in the late spring and summer.

The mature summer leaves of P. cordata, P. macrocarpa of the Pacific
Northwest and the P. princeps complex of the Hawaiian Islands are highly
specialized and are the most extreme examples of a pseudolamina ( sensu
Arber, 1918) in the genus. The mature leaves of P. cordata are very similar to
the basal leaves of Mertensia virginica and sterile specimens of the two species
may be confused.

Fig. 4. Seasonal heteroblastic leaf sequence in Plantago cordata.

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THE MICHIGAN BOTANIST

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The leaves of the winter rosette of P. cordata (Figs. 1 & 4) are lanceo¬
late to spatulate and usually parallel-veined. In the spring, a heteroblastic se¬
quence terminates in a more or less “midribbed” leaf. The sequence may be
interrupted and arrested at any stage. An interruption is readily brought on by
stress conditions, e.g. drought, prolonged periods of direct sunlight, or high
temperatures. If the sequence is not interrupted, three or four summer leaves
will be produced before the reverse sequence in the fall again results in leaves
of the winter rosette type. In the greenhouse, many plants flower, fruit, and
subsequently slough off the peduncles, thus initiating what is, in the field, the
summer stage; but they never develop leaves beyond those of the spring type.
Completion of the cycle is apparently under environmental control as the
same plants will, under other conditions, progress through the complete series.
If the plants are grown continuously, without being subjected to a cold treat¬
ment, and under a 12-hour or longer day, the rosette will consist only of sum¬
mer leaves. I have experimentally held plants at this stage for over a year.
However, either short days (eight hours) or a cold treatment will initiate a
reversal to the winter rosette. Plants in the winter rosette, even when held
under short days and/or cool (50° F) storage, will, if the temperature is above
freezing, spontaneously initiate spring leaves and inflorescences after about a
month. However, they will not produce summer leaf types under these condi¬
tions.

Leaf shape and size vary considerably within any population, in addition
to the variation associated with the heteroblastic development. Fig. 2 shows
the variation in shape and maximum size of summer leaves collected Septem¬
ber 15, 1967, at the Stockport Landing, N. Y., station. In general, the largest
leaves were from plants in more shady habitats; the smallest leaves were from
plants growing on a gravel bar in direct sunlight.

Petiole length and leaf width, at any heteroblastic stage, also vary con¬
siderably and are under direct influence of light intensity, available water, and
temperature. Low light intensity, wet soil, and low temperatures result in the
longest petioles and widest leaves. High light intensity, dry soil, and high
temperatures result in the shortest petioles and narrowest leaves. Various com¬
binations of these factors result in intermediate proportions.

The variation resulting from the interplay of the heteroblastic series and
environmental conditions is expressed in all the populations, both in the field
and in the greenhouse. This plasticity must be kept in mind when looking for
the plant in the field and when identifying herbarium specimens.

Anatomically, the leaves of P. cordata show little variation. The leathery
leaf texture, typical of this species, is the result of a well developed spongy
mesophyll and, in some cases, an aerenchyma-like tissue. This is most highly
developed in plants growing in the open sun in the water and results in a
“flabby” or almost succulent leaf texture. In cross-section, the leaf shows a
palisade layer on both the adaxial and abaxial sides. The epidermal patterns
(Fig. 5) are similar on both leaf surfaces. The scattered stomata are usually
surrounded by four auxiliary cells having more or less sinuate margins.
The simple, 2-celled, capitate hairs have a single stalk cell and are of sporadic

1969

THE MICHIGAN BOTANIST

79

odaxial

abaxia I

submersed emersed

Fig. 5. Epidermal cells on leaves of Plantago cor data.

occurrence on emersed leaves. The plant appears glabrous to the naked eye.

Leaves produced on submersed plants have generally larger epidermal
cells with only slightly sinuate margins (Fig. 5). Stomata are present but they
are not so abundant as on emersed leaves; and abortive stomatal mother cells
which did not undergo further development are common. The latter may be
recognized in Fig. 5 by the radially arranged auxiliary cells. No hairs are pres¬
ent on submersed leaves although a few specialized epidermal cells resembling
trichome bases are found on the adaxial surface.

STEM

The stem of P. cordata is a short fleshy caudex, 2-4 cm in diameter and
1-3 cm high. This caudex consists of the true stem (a corm) and the greatly
swollen, persistent hypocotyl. The upper portion is covered with numerous
adjacent leaf scars. It is glabrous except at the bases of the current leaves and
at the apical region where a silky tomentum of hyaline hairs 1 cm or less
long, covers the meristematic regions. The older portions of the stem become
covered with a spongy endodermis which merges into the fleshy root cortex.

There is no evidence of internodal elongation. The approximate age of a
plant may be estimated by counting the spiralling ridges formed by the leaf
scars. The broader leaf scars formed by the summer leaves alternate with the
narrow ridges formed by the winter leaf bases. Lateral buds, which may occur
sporadically on older specimens, give rise to small rosettes. If these become
detached through mechanical injury or sloughed off from a dying parental
stem, they will reproduce the plant vegetatively . I have not observed the later¬
al shoots producing inflorescences while attached to the main parental stem.

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ROOT

The root is the most characteristic morphological feature of P. cordata.
The genus Plantago has both of the two basic types of root systems: tap and
fibrous. Most sections ( sensu Pilger, 1937) contain species showing both types.
The tap root is usually correlated with annual or biennial life cycles ( P .
macrocarpa which is a perennial is a notable exception). In those species with
a fibrous root system, e.g., P. major and P. mgelii, the roots are relatively
narrow and the primary food storage organ is the caudex. Plantago cordata
and P. eriopoda are unusual in Plantago in having a fleshy fibrous root system
which serves as the primary food storage organ (Figs. 1 & 7). Likewise the
development of the roots is unusual in these species.

In Plantago, as in most dicotyledonous angiosperms, the radicle is the
primary root. In species with a tap root system, the major root develops di¬
rectly from the radicle and can be considered a developmental extension of it.
The fibrous root system differs from this basic plan in that the primary root
soon dies or its development is determinate, whereupon numerous lateral roots
develop. In most Plantago species with a fibrous root system, the lateral roots
either arise from this much shortened primary root (e.g., P. indica ) or, in most
perennial species having a caudex, the lateral roots arise adventitiously from
the caudex (e.g., P. lanceolata ). However, P. cordata and P. eriopoda produce
fibrous roots not as laterals from the primary root nor from the caudex, but
adventitiously from the persistent hypocotyl, as shown in Fig. 6. In these spe¬
cies, the primary root derived from the radicle is short and withers soon after
the adventitious roots are initiated. The lateral roots are also limited in de¬
velopment, averaging 8-10 cm in length. After maximum length is reached,
these become thickened and, in turn, give rise to tertiary roots which likewise
become thickened, and so on. Since the length of each root is finite, the axis
of a lateral root system is usually zigzagged. In cases where a secondary or

Fig. 6. Seedling development in garden-grown Plantago cordata.

1969

THE MICHIGAN BOTANIST

81

tertiary root happens to become aligned, by overtopping, on the same axis as
the root from which it arose, a definite constriction indicates the point of ori¬
gin of the new root. Scattered across the entire root system, but concentrated
at the distal ends, are small thread-like roots bearing very few, if any, root
hairs.

The large fleshy roots of P. cordata and particularly the seemingly con¬
tinuous texture of the roots, hypocotyl, and caudex have led to some confu¬
sion in the literature. Tyler (1901) interpreted the large root as part of a
rhizome which had rotted out in the center and had subsequently broken up.
His photograph of this condition actually shows a lateral shoot which apparent¬
ly was torn from the original plant along with one large fleshy root. (The
herbarium specimen from which the photograph was made is F. J. Tyler,
October 2, 1901, OS). An old plant can have a hollow caudex and, if the
apex dies, it is easy to see how a lateral shoot could become an independent
plant. It is sometimes difficult to distinguish between a fleshy root and a
caudex on herbarium specimens. This lack of differentiation between the
caudex and the large storage roots could easily lead to their misinterpretation
as a rhizome.

Anatomically the roots of P. cordata have several modifications reflect¬
ing its aquatic habitat. The highly developed cortex containing abundant
aerenchyma (Fig. 7), the paucity of root hairs, and the production of adventi¬
tious roots are all characteristic of aquatic plants (Esau, 1953). The root cor¬
tex is the important storage tissue in P. cordata. The cells are frequently filled
with starch grains which may be single or in groups of 2 -4 (Fig. 7). These are
the largest starch grains of the 97 species of Plantago I have observed to date.

REPRODUCTIVE BIOLOGY
PHENOLOGY

Plantago cordata is a short-day plant. Initial blooming may begin in
March in the south (Harper, 1944; Godfrey, 1960) and in mid-April in the
north. Rarely the plant will bloom again in September and October. Although

Fig. 7. Left: Photomicrograph of cross section of a root of Plantago cordata. Right:
Starch grains from cortex of root (drawn from projected image).

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anthesis takes place normally in the spring, the young inflorescences are initi¬
ated in the fall. These young spikes are well protected throughout the winter
by the imbricate petiole bases. Following a cold treatment, which may consist
of simply an early frost, the peduncles elongate and the plant blooms. There
is no evidence that the fall-blooming plants set seed. Even if they did, the
resulting seeds would not have time to germinate and become established be¬
fore winter set in. Thus the reproductive potential of the plant for the follow¬
ing season would be lost (cf. below, “Life History”).

INFLORESCENCES

Under field conditions, inflorescences are subtended only by leaves in
the winter or the spring and fall condition (Figs. 1C & 4) although summer
leaves may be present on the plant. Therefore, the number of inflorescences
(averaging six per plant) is partially determined by the duration of the cool
fall season and/or the first frost. In the greenhouse, individual plants have
sporadically produced inflorescences in the axes of summer leaves when the
plants have been maintained under extended (10 months) summer conditions.

In the spring, the young inflorescences grow up beyond the protecting
petioles and the leaf blades, through the activity of an intercalary meristem.
Under cool, moist conditions, the peduncle grows to a length of 15-30 cm in
6-7 days. Simultaneously, the flowers are maturing and meiosis occurs. Ap¬
proximately one week after growth begins, the peduncle reaches its maximum
length and the stigmas of the lowest flowers are extended. At this time, the
meristematic activity shifts to the rachis. Beginning at the bottom of the spike
and progressing upward, the elongation of the rachis coincides approximately
with the extension of the stigmas of the flowers. Three to four days after the
extension of the stigma, the individual flowers protrude their stamens. This
sequence progresses acropetally. The processes of stigma extension, elongation
of the rachis, protrusion of anthers, and, to a small extent, initiation of addi¬
tional flowers at the tip of the spike, continue (under favorable conditions)
for a period of one and one-half to two weeks, at which time an inflorescence
totalling 40-60 cm in length and bearing 80-130 flowers is produced. If envi¬
ronmental conditions are usually dry, elongation of both the peduncle and the
rachis is inhibited and frequently the terminal portion of the inflorescence
aborts, resulting in a depauperate inflorescence. This condition is frequently
observed on herbarium specimens.

The young inflorescences, up to the time of anther protrusion and the
beginning of seed set, are soft, brittle, and solid. As the capsules start to de¬
velop, the peduncle becomes hard, tough, and hollow. Depauperate inflores¬
cences often retain solid peduncles, so peduncle hollowness should be used
with caution as a diagnostic characteristic of this species.

In contrast to most Plantago species, the peduncle and rachis of P. cor-
data remain alive through the time of capsule dehiscence.

FLOWERS, FRUIT, AND SEEDS

The flowers of P. cordata (Fig. 8) are typical of the genus. They are
well adapted to wind pollination as shown by the reduced, chartaceous

1969

THE MICHIGAN BOTANIST

83

corolla, the long, plumose stigma, and the versatile anthers bearing copious
pollen. For a comparative description of the morphology of the pollen, see
Bassett (1968). Each flower is subtended by a green, oval, glabrous bract. The
bracts beneath the lowest flowers of the spike may be enlarged and leaf-like,

Fig. 8. Flowers of Plantago cordata, P. eriopoda, and hybrid. Abnormal gynoecial parts
of P. cordata x eriopoda.

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THE MICHIGAN BOTANIST

Vol. 8

often obscuring the included flower. No floral abnormalities have been ob¬
served in this species.

The gynoecium, at the time of anthesis, contains four ovules although
usually only two or three seeds are produced per capsule. The capsules of P.
cordata are unique in the genus in that they are alive at the time of de¬
hiscence. The abscission layer of the pyxis is well developed even before the
seeds are mature. The top of the capsule is usually held on only by the con¬
necting septum. When the seeds are ripe, the capsule readily dehisces at the
slightest touch. The seeds and the entire fleshy placenta fall out as a unit, an
unusual character in the genus.

CYTOLOGY AND HYBRIDIZATION EXPERIMENTS

Bassett (1967) reported chromosome counts of P. cordata from a popu¬
lation in Missouri and one in North Carolina as 2n = 24. I have observed
meiotic figures (Fig. 9) from the following populations. Meiosis was normal in
all plants observed.

Collecting Site

No. of plants
observed

Collection

Bivalents at
Meiosis

Ill., Lake Murphysboro

2

Tessene 2066

12

Ill., New Lenox

2

Tessene 21 76

12

Mo., Millcreek

1

Bassett & Spicer 4501

12

N.Y., Stockport Landing

5

Tessene 2065

12

N.C., Denton

1

Stephenson 618

12

Ohio, Peebles

5

Tessene 1881

12

Ontario, Thedford

5

Tessene 1941

12

Plants of P. cordata are self-compatible but some outcrossing is facili¬
tated by the protogynous development of the flowers. Once pollination and

Fig. 9. Camera lucida
drawings of meiotic chro¬
mosomes of Plantago cor¬
data: N = 12 in all cases.
A) Hudson River, New
York ( Tessene 2065)] B)
Peebles, Ohio ( Tessene
1881)] C) Thedford, On¬
tario ( Tessene 1941)] D)
New Lenox, Ill. ( Tessene
2176).

1969

THE MICHIGAN BOTANIST

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TABLE I. Hybridization experiments involving Plantago cordata.

Female Parent

Male Parent

No. of
spikes
pollinated

No. of
seeds
set

No. of
,Fi

plants

F| pollen
staina-
bility

Comments

P. cordata
Peebles, Ohio

P. eriopoda

Wyo. (n=12)

3

10

1

3%

See text

P. eriopoda
Centennial, Wyo.

P. cordata
Thedford, Ont.

4

6

-

-

-

P. cordata
Peebles, Ohio

P. cornuti

Bot. Gard. Coim¬
bra, Portugal.
(n=6)

3

2

P. cordata
Thedford,
Ontario

P. reniformis

Bot. Gard.
Coimbra,
Portugal. (n=6)

2

P. rugelii (n=12)
Ann Arbor,

Mich.

P. cordata
Stockport
Landing, N.Y.

6

4

Seeds

abortive

P. cordata
Peebles, Ohio

P. rugelii

Ann Arbor,

Mich.

2

P. cordata
Stockport
Landing, N.Y.

P. media (n=12)
Berlin, Germany

3

‘

P. cordata
Peebles, Ohio

P. pachyphylla v.

montis-eekis
Maui, Hawaii
(n=12)

3

6

P. pachyphylla
v. montis-
eekis

Maui, Hawaii

P. cordata

Peebles, Ohio

2

P. cordata
Peebles, Ohio

P. princeps

Maui, Hawaii
(n=6)

4

'

'

'

'

P. princeps

Maui, Hawaii

P. cordata
Stockport Land¬
ing, N.Y.

3

3

Seeds

abortive

P. cordata
Peebles, Ohio

P. sparsiflora
Allenhurst, Geo.

6

-

-

-

-

P. spar si flora
Allanhurst, Geo.

P. cordata

Peebles, Ohio

4

2

-

-

No seed
germination

86

THE MICHIGAN BOTANIST

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germination of the pollen grain has occurred, the stigma rapidly withers and
dries— often before the particular flower protrudes its stamens. The lowest
flowers have a high probability of setting seed via outcrossing, but flowers on
the upper portion of the spike may well be fertilized by pollen arising from
the lower flowers. All my greenhouse populations of P. cordata, except Hines
6825, have been intercrossed and are completely interfertile.

The protogynous condition and rapid withering of the pollinated stigma
make Plantago an excellent group with which to perform hybridization stud¬
ies. If the experimental plants are isolated (either mechanically or spatially)
and pollinated as soon as the stigmas appear, self-fertilization is almost com¬
pletely eliminated. Utilizing an aluminum foil jacket to isolate the young
spikes, I made a series of controlled crosses to determine the degree of pro¬
miscuity of P. cordata. The results are presented in Table I.

As can be observed in Table I, P. cordata apparently does not readily
hybridize with the species listed. The one hybrid plant obtained, P. cordata x
eriopoda (Figs. 8, 10, & 11), is intermediate between the parents in most
characters (Figs. 8 & 10). The hybrid is sterile, having abortive, non-emergent
anthers, less than 5% pollen stainability, and, occasionally, extra gynoecial
parts which are abortive (Fig. 8).

Structural abnormalities of the androecium are common in Plantago
hybrids (Hope-Simpson, 1939; Rahn, 1956; Gorenflote, 1959, 1961; Cartier,
1963; Stebbins & Day, 1967); however, the only abnormalities of the gynoeci-
um which have been reported have been induced by herbicides (Hitchcock &
Zimmerman, 1947) or were spontaneous (Gorenflote, 1959). Gorenflote also
reported teratological gynoecia in hybrids of P. coronopus and P. maritima
but these did not include supernumerary parts. What, if any, evolutionary
significance there may be in the occurrence of unusual gynoecia in P. cordata
X eriopoda will not be discussed here.

LIFE HISTORY

Plantago cordata is readily grown under standard greenhouse conditions
in a 2:1 mixture of rich loam and sand and without need for flooding. Al¬
though flooding more closely approximates the aquatic conditions under
which the plants grow in the field, I have found no need to endeavor to simu¬
late natural conditions in the greenhouse. The plants require frequent water¬
ing, and I have grown them in flooded pots.

The seeds and seedlings are apparently the most critical stages in the life
of P. cordata. As indicated above, the seeds remain attached to the spongy
placenta (Fig. 3) when first shed. The resulting unit is highly buoyant and
rapidly disperses the seeds if it happens to fall in the water. When the seeds
come in contact with the water, the hygroscopic mucilaginous coat surround¬
ing them swells, eventually forcing the seed away from the placenta and break¬
ing the funiculus. Once freed, the seeds float individually with their planar
surfaces upward. The mucilaginous coating causes a seed to adhere to any ob¬
ject which it contacts in the water. The seeds will germinate after 6-14 days
even if they are still floating, but will not germinate under water. The

1969

THE MICHIGAN BOTANIST

87

seedlings will continue to float for another week before eventually sinking and
dying. It is during this critical three-week period of seed dispersal and estab¬
lishment that the reproductive potential of P. cordata is realized. If the seed
or a floating seedling does not become established in a favorable location, it is

lost. Longevity experiments show that
they live only about a month in storage
on how the seeds yielding the type of
managed to germinate.)

If a plant becomes established, it

Pcordata x erio

the seeds must germinate rapidly, as
at 40°F. (It is interesting to speculate
P. cordata ever got to Paris and still

will bloom the following year. The

P cordata

o d a

Fig. 10. Leaves of Plantago cordata, P. eriopoda, and hybrid.

88

THE MICHIGAN BOTANIST

Vol. 8

longevity of plants is not known but I have kept mature plants from the field
alive and vigorous in the greenhouse for three years. These specimens were
estimated to be approximately four years old at the time they were collected.

From field observations and herbarium specimens, it appears that P. cor¬
data has few natural enemies. Close observation of most collections will reveal
5-10 small cysts 1-2 mm across on one or more leaves. The cause of these
cysts is not known but the lack of fungal hyphae or insect larvae suggests
perhaps a virus or bacterium. Plants bearing these cysts have been grown in
culture for two years with no apparent deterioration. Similar cysts are found
on many other species of Plantago, e.g. P. lanceolata, P. rugelii.

Dodder, Cuscuta sp., was found on several plants of P. cordata at the
Hudson River station; however, this resulted in no noticeable harm to the
plantago.

In the greenhouse, the inflorescences of P. cordata are subject to infesta¬
tion by a downy mildew. The same mildew (species unknown) attacks most of
the 103 Plantago species in my collection but usually grows on both the
leaves and inflorescences. The mildew apparently does little harm to the
plants.

Insect damage is also rare. Three specimens (all from Illinois) have “nib-
blings” along the margin of the leaf suggesting the activity of leaf-cutting
wasps. In the greenhouse, red spider mites often feed upon the inflorescences

1969

THE MICHIGAN BOTANIST

89

of P. cordata. The silky threads left by the mites may inhibit the full protru¬
sion of the stamens. This infestation has not been observed in the field.

DISTRIBUTION AND ECOLOGY

The geographical distribution of P. cordata (Fig. 12) is correlated with
the occurrence of limestone and dolomite bedrock. The apparent discrepancy
shown by the southern Alabama and central North Carolina localities is ex¬
plained by the presence of limestone erratics and hard-water springs in these
areas. At Peebles, Ohio, P. cordata grows in a shallow stream directly on dolo-
mitic bedrock. Frequently the roots penetrate the joints of the rock for a dis¬
tance of 30 cm or more. At Thedford, Ontario, the limestone is within a few

Fig. 12. Distribution of Plantago cordata.

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THE MICHIGAN BOTANIST

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feet of the surface although the stream in which the plant is found has a mud
bottom. However, the water is highly calcareous as evidenced by marly incrus¬
tations remaining around old pools. The bedrock in the immediate vicinity of
Lake Murphysboro, Illinois, is a calcareous sandstone. Here P. cordata forms
small colonies in a shallow pebble-bottomed stream. Although the New Lenox,
Illinois, station is located on glacial till, the rocks in the stream bed are main¬
ly calcareous. Many of the old localities which I visited in Michigan, Illinois,
Wisconsin, Missouri, and Ohio but where the plantago could no longer be
found, were small streams, frequently flowing on limestone bedrock. Although
the Stockport Landing, New York, station is within the area supposedly hav¬
ing limestone, the plantagos were growing in thick muddy ooze and non-
calcareous alluvial beaches. Bassett (1967) indicates a locality in north-eastern
Florida ( S . R. Buckley, GH) but this is questionable as it is far beyond the
more typical limestone areas, the ecology of the area is completely different,
and it would be the only station on the Coastal Plain.

Associated species occurring at the various stations are listed in Table II.
In general, the physiognomy and composition of the major communities in
which the plantago is found are quite diverse. The Thedford, Ontario, popula¬
tion occurs in and along a small, mud-bottommed, slow-moving stream which,
as shown by the mud bars and their vegetation and as described by Mr. Bill
Henry who guided us to the area, exists as a series of small pools at least part
of the year. The stream meanders through a beach-maple woods. The largest
tree observed was an Acer saccharum with a dbh of approximately 18 inches.
The woods and surrounding fields had been grazed and signs of recent use of
the stream by cattle were evident. It is possible that this population is, in
part, artificially controlled (see below, “Economic Uses”). The population
consisted of about 100 individuals spread out along the stream for a distance
of approximately 350 m. The largest group— about 40 plants— was the last one
seen downstream. (Fig. 13) Dodge, in his flora of the county (1916), implies
that the species was at one time, at least, not rare: “Frequent along streams,
in pools, ditches, wet places and wooded swamps.”

The station at Peebles, Ohio, was visited four times during the course of
this study: November, 1965; May, 1966; April, 1967; and June, 1967. The
general habitat is a small valley with dolomite walls on both sides and a rapid¬
ly flowing stream in the center. Several old stream beds occur scattered in the
ravine, showing the old stream course with several switch-backs. The water in
the stream is murky with dolomite dust from an adjacent quarry. The area is
shady and protected. From the conglomeration of twigs and leaves on many
shrubs and trees, it appears that the entire valley is periodically inundated to a
depth of several feet. In general, the area is an alluvial plain bordered by a
Southern mixed mesophytic forest containing a few trees of Thuja occiden-
talis. Until the summer of 1967, this station was fairly undisturbed; some of
the sycamores on the alluvial plain were over four feet dbh. Unfortunately,
the valley is now being filled with refuse from the quarrying operations, many
trees have been felled, and roads have been built. This situation is most regret¬
table as the Peebles station supported the largest population of P. cordata I

1969

THE MICHIGAN BOTANIST

91

TABLE II. Vascular plants associated with Plantago cordata. Listed are those species
which occurred within about 10 feet of a plant of P. cordata. In some cases, where the to¬
pography of the area was very steep, cliff and upland species are included. Although the
stations were visted at different times of the year, one can approximate the community
composition by the species indicated. The letters following each species give the station(s)
where the species was found: I = Lake Murphysboro, Ill.; M = Messenger Woods, New
Lenox, Ill.; N = Hudson River, Dutchess Co., New York; O = Plum Run Quarry, Peebles,
Ohio; T = Thedford, Ontario. (See text for a general description of each station.)

TREES

Acer negundo L

A. rubrum L M

A. saccharum L NT

Amelanchier sp. T

Carpinus caroliniana T

Carya glabra L

C. ovata L O

Cornus alternifolia N O

C. florida L

Crataegus spp. L M TO

Fagus grandifolia T

Fraxinus americana L M N O

Gleditsia triacanthos L M

Liriodendron tulipifera O

Ostrya virginiana M N T

Platanus occidentalis L N O

Populus deltoides O

P. tremuloides N

Prunus virginiana T

Quercus alba L M

Q. muehlenbergii L O

Q. rubra M O

Thuja occidentalis O

Tilia americana M T

Ulmus americana M N T O

U. rubra M

SHRUBS, SUBSHRUBS, & VINES

Cephalanthus occidentalis T

Cornus racemosa M

Corylus americana M

Euonymus atropupureus L N

E. obovatus N T O

Hamamelis virginiana T O

Lindera benzoin L O

Parthenocissus quinquefolia T

Physocarpus opulifolius N O

Rhamnus cathartica M

Rhus radicans L M N O

Ribes cynosbati L TO

Salix interior N O

Sambucus canadensis T O

Smilax bona-nox L

Staphylea trifolia M

Viburnum lentago M T

Vitis riparia N

Xanthoxylum americanum L N O

HERBS & PTERIDOPHYTES

Allium sp. L T O

Ambrosia trifida M

Anemone quinquefolia T

Arisaema atrorubens M N T O

Asclepias incamata M

Aster novae-angliae M

Bidens sp. L N

Boehmeria sp. L

Campanula americana M

Carex albursina T

Carex spp. L M N T O

Cicuta sp. M T

Circaea canadensis M

Claytonia virginica T

Cuscuta sp. N

Cyperus sp. N

Decodon verticillatus

N

Desmodium illinoense

L

Dryopteris marginalis

T

Equisetum arvense

0

E. hyemale

L

N

Eupatorium rugosum

M

E. serotinum

L

Erythronium americanum

T

Fragaria virginiana

M

T

Galium aparine

L

M

N

T

G. circaezans

L

Glyceria sp.

N

T

Geum rivie

L

M

T

Helianthus sp.

L

Hepatica acutiloba

M

Hystrix patula

M

Impatiens sp.

L

M

N

T

Iris sp.

M

N

Lobelia siphilitica

L

M

Lonicera dioica

M

Luzula campestris

T

Lycopus americanus

N

Lysimachia ciliata

T

L. nummularia

M

T

L. quadrifolia

N

Mertensia virginica

0

Mimulus alatus

L

Mitella diphylla

T

Muhlenbergia sobolifera

L

Nasturtium officinale

M

Onoclea sensibilis

N

T

Osmorhiza longistylis

M

Panax trifolius

T

Panicum boscii

L

Pellaea atropurpurea

0

Phaseolus polystachios

M

Phlox divaricata

M

T

Phryma leptostachya

M

Pilea pumila

M

Plantago lanceolata

0

P. rugelii

L

M

N

T

0

Podophyllum peltatum

T

Polemonium reptans

M

0

Polygonum sp.

N

Polymnia uvedalia

M

Polystichum acrostichoides

T

Pontederia cordata

N

Prunella vulgaris

L

Rumex verticillatus

L

Sagittaria sp.

N

Sanicula marilandica

M

Sanguinaria canadensis

M

Scirpus atrovirens

L

Solanum dulcamara

N

0

Sorghastrum nutans

N

Thalictrum dioicum

T

0

Thaspium barbinode

M

Typha latifolia

N

Tiarella cordifolia

M

Trillium grandiflorum

T

Uniola latifolia

L

Verbena hastata

M

Viola pensylvanica

T

V. renifolia

M

V. rostrata

T

V. sororia

L

M

T

THE MICHIGAN BOTANIST Vol. 8

Fig. 13. Plantago cor data in muddy-bottomed stream near Thedford, Ontario. From a
color transparency by E. G. Voss, May 6, 1967.

Fig. 14. Plantago cordata on gravelly stream bank near Peebles, Ohio, May 8, 1966.

1969

THE MICHIGAN BOTANIST

93

have seen. In May, 1966, I thoroughly explored the area and counted 600
individual mature plants of P. cordata. In addition, many seedlings were scat¬
tered along the shores of the stream and on the various alluvial bars. (Fig. 14)
But here, as at Thedford, the plantago population was not continuous along
the stream even though the habitat appeared to extend considerably beyond
the area where the last plantago was found. Cedar Creek, above its juncture
with Plum Run, did not support any P. cordata, although here again the habi¬
tat appeared to be the same as that along Plum Run and along Cedar Creek
below the juncture of the two streams. The problem of moving upstream is
apparent, but why does the population disappear abruptly downstream? This
phenomenon I have seen repeatedly but can not explain.

The Hudson River, New York, localities were originally discovered in the
early 1800’s (Wiegand & Eames, 1926) and have been revisited by several col¬
lectors since. I visited these areas in September, 1967. Plantago cordata is
known from New York only at the Hudson River although there is a report of
an old inland locality (Wiegand & Eames, 1926). The New York stations are
of particular interest as the plantago grows in slightly brackish water (2-5
ppm, Muenscher, 1937), is subjected to partial inundations by the tide, and
grows in open sunlight. The population studied occupies an area of about two
acres and consists of several hundred plants. The plantagos occur within three
distinct adjacent communities: (1) An association of Peltandra virginica,
Typha latifolia, and Decodon verticillatus. The plantago is common through¬
out the association but most abundant in the area exposed at low tide. The
plants were apparently completely inundated at times as all the leaves and
most of the surrounding vegetation were covered with a fine coating of silt.
(2) High, open alluvial beach along the river. Here P. cordata grows alone or
in groups of a few to over 20 individuals. The plants were in full sunlight at
10:00 a.m. The high-beach plantagos are the smallest, although mature, of the
combined Hudson River population. (3) A Salix interior-Acer negundo
swamp. The plantagos in this association are the largest I have observed in
either the field or the herbarium. The dense overstory shades the ground-level
plants. The resulting shade forms of P. cordata, as with most other members
of the genus, have unusually wide leaf blades with delicate texture, elongation
of the petioles, and a corresponding elongation and narrowing of the spike.

At Lake Murphysboro, Illinois, a man-made lake 47 years old, the plan¬
tago occurred as a small population of 30 individuals along a shallow feeder
stream for a distance of about 50 m. The woods through which the stream
flows is a mixed southern mesophytic association. The dominant trees are
Quercus muhlenbergii, Acer saccharum, and Platanus occidentalis. Plantago
cordata grew in the stream and showed no preference for the edges or the
middle. The deepest part of the stream was only 10 cm. Three plants of P.
cordata were also found growing in an unshaded drainage ditch along the
road. These are the only plants of P. cordata I have seen growing in an obvi¬
ously disturbed and man-made habitat. No yearling plants were found.

A station near New Lenox, Illinois, was visited in October, 1968. Wheth¬
er or not this is the same locality that E. J. Hill visited in 1879 is not known

94

THE MICHIGAN BOTANIST

Vol. 8

as his data are not specific (In rivulet, New Lenox, Ill. May 17, 1879; Hill
15. 1879, ILL). If today’s station were the same as Hill’s locality, this would be
the oldest known extant population (see Fig. 15). However, the ecology of
the area does not suggest 90 years of undisturbed habitat.

Approximately 250 plants of P. cordata grew in the rocky shallows of a
small stream for a distance of approximately 550 ft. at the bottom of a ra¬
vine. Here, as at the other stations, the habitat appeared to continue both
upstream and downstream from the plantago population but the plantago
abruptly stopped. Downstream, the creek “went underground” as the stream-
bed became very rocky. The moist streambed was explored for a distance of
about one-half mile downstream from where the surface water stopped but no
additional plantagos were found. Upstream from the plantago population, the
habitat remained similar for a distance of several hundred yards. Beyond that,
the canopy was open and the stream passed through an old field.

The dominant trees in the ravine were Quercus rubra, Ulmus americana,
Fraxinus americana, and Crataegus sp. Impatiens sp., Rhus radicans, and
Polymnia uvedalia were common in the ravine. The creek itself supported no
other vascular plant except P. cordata. Several stumps three feet or more in
diameter were scattered throughout the area indicating high-cropping in the
past.

A comparison of the species associated with P. cordata in the five sta¬
tions studied is given in Table II. The habitat of P. cordata is usually a ravine
where the immediate vegetation is that of a lowland forest but the nearby
surrounding hills support oak-hickory, beech-maple, or mixed mesophytic for¬
ests.

As is readily ascertained from the above discussion and Table II, the fac¬
tors governing the habitat and occurrence of P. cordata can not be readily
determined from those species and species-associations occurring with it.
Therefore, it becomes necessary to look for other sources of data.

Although the absence of an object can not, logically, be shown to prove
anything positive, a survey of those habitats known to have once supported
P. cordata and those habitats which would appear to be able to support it
today but do not, can, when compared with extant stations, give some insight
in understanding the factors critical to the existence of the species. These
data, in conjunction with what historical information one can obtain from the
literature, herbarium specimens, and word of mouth, can be correlated into a
working hypothesis to explain the ecological behavior and geography of the
plant.

REVIEW OF COLLECTING HISTORY AND COLLECTORS

Roland M. Harper (1944) drew an interesting parallel between P. cordata
and a plant described from Georgia by William Bartram in June, 1773, but whose
report was not published until 1943 (F. Harper, ed., 1943; see p. 139). As
pointed out by R. M. Harper, the plant mentioned fits the description of P.
cordata better than any other Plantago species. Bartram wrote:

1969

THE MICHIGAN BOTANIST

95

... & in & on the Verge of the Rapid rivulets a great Species of Plantago, in every
respect like the common Plantain; except that it is of an incredible magnitude, the
leaves with the footstalk commonly 3 feet in length, & in the Broadest part above
12 inches wide, the number of ribs of the leaf being the same with the common
Plantain, & run in just the same direction, & by the smell & taist of the Plant
seemes to possess the same virtues, but in a greater degree, it grows in the little
coves, of the Rivulets & brooks where the eddie water hath collected & raised a
Bed of slush or in the middle of the Creeks, where it forms a green Island, about
some Rocks or trash.

In interpreting Bartram’s description, Harper assumed that it was not P. corda¬
ta in that (1) the size of the leaf surpassed any specimens known to him; (2)
the “ribs” (= veins) were not the same as the common plantago. Here Harper
assumes P. major but it might just as well have been P. rugelii (Tessene,
1968); (3) Harper did not believe that P. cordata occurred in Georgia. If
Harper had had more specimens available to him, he might have reserved com¬
ment on Bartram’s description. The size of the leaf, although exceptionally
large, is of a magnitude approached by specimens of P. cordata growing in
very shady areas. (A specimen collected in Illinois in 1961 (G. N. Jones
32335, ILL) has a blade 10.2 inches wide.) Georgia in Bartram’s time still
contained large areas of virgin habitat: One would expect a dense overstory.
Plantago cordata does occur in Georgia, as has been reviewed recently (God¬
frey, 1960). I have no doubt that Bartram’s plant was indeed P. cordata.
Lrom this note and Lamarck’s description in 1791, we know that the north¬
ern and southern distributional boundaries of P. cordata had been established
before these and the intervening areas had been markedly influenced by Euro¬
pean man.

After these two early descriptions, I have found no record of the plant
until 1833 when George Engelmann found it in “Rivers and streams, west of
St. Louis” ( Engelmann April 1933, MO). I. A. Lapham collected it in “Wet
places, Columbus O.” in April, 1836 (PH). Prom that time until the 1880’s, P.
cordata was collected sporadically throughout its range. Between 1878 and
1900, many new localities were found (Pig. 15) and most of the specimens
extant today date from this period. The reasons for this sudden wealth of
collections are probably many and complex and an attempt to explain them
would, of necessity, include considerable speculation. Plantago cordata may
have been collected more frequently before 1880 than the graph indicates;
many herbaria of the early botanists have been lost or exist only as remnants.
But this is not as important as the paucity of collections since 1900 when
more collectors have been in the field than ever before. However, I know of
only seven stations for P. cordata today. It is a rare plant, but it may always
have been rare. It is not necessarily becoming extinct.

During the past three years, I have made numerous trips to many areas
and specific localities where P. cordata had been collected in the past. Most of
the searching was in areas where the plant had been collected since 1920, as
these would be the most promising. The seven stations known to be maintain¬
ing a population of P. cordata (Pig. 15) have been discovered only recently.

96

THE MICHIGAN BOTANIST

Vol. 8

Many of the old localities have been obviously destroyed, particularly those
near large cities such as Chicago, St. Louis, etc.; but as recently as 1929 P.
cordata was collected in downtown Milwaukee (stream where North Shore
crosses Howell Ave. Throne May 12, 1929, UWM). In Table III are listed the
old localities which have been searched for and the possible cause for the ap¬
parent local extinction of P. cordata or failure to find it.

TABLE III. Old localities visited during this study but at which Plantago cordata was not
found. Most of the localities were suggested by herbarium labels (see appendix for complete
citations). The “Present Status” column indicates how each locality differs from what ap¬
pears to be the natural habitat of P. cordata as deduced from extant stations. “Suburbia”
covers general destruction of natural habitats in metropolitan areas.

LOCALITY

PRESENT STATUS

MICHIGAN: Brooks and
ditches about Alma. Col¬
lected 1890-95.

No exact location was given. The general area has been
extensively cleared; the remaining woods are widely used
as cattle pastures. Streams carry considerable silt from
erosion of fields.

MICHIGAN: General vicinity
of Michigan State University
campus. Collected 1886-95.

The area is now part of the campus of Michigan State
University.

MICHIGAN: Vicinity of Port
Huron. Collected 1838-1897.

No exact location is given for the Wright or Dodge col¬
lections of P. cordata; however, the Black River is pol¬
luted as is the St. Clair. Many of the small feeder
streams in the area look as though they could support

P. cordata but, to date, it has not been rediscovered.

MICHIGAN: Bank of creek
in hardwoods, 5 mi S of Ann
Arbor. Collected 1924-25

All streams in the area have been exhaustively scoured
but no P. cordata has been found. Most of the streams
have been rerouted into drainage ditches.

MICHIGAN: West branch of
Paint Creek, Ypsilanti town¬
ship. Collected in 1891.

Farwell returned to his locality in 1928 to look for

P. cordata, which he did not find. The area, as men¬
tioned by Farwell (1930), has been partly converted
into a golf course. Today, the course is overgrown and
the area is a gun club. The vegetation along the creek is
all second growth. The creek now flows in a drainage
ditch for about half its length.

MISSOURI: Vicinity of Alien-
ton. Collected 1879-1923.

The area is now part of suburbia.

MISSOURI: West fork of
Fourche a Renault Creek,
Washington Co. Collected in
1955.

The creek has been dammed to make a series of ponds
upstream. The spring still flows but not profusely
enough to fill the creek bed which is now a dry cattle
run.

MISSOURI: 3 mi East of
Millcreek. Collected in

1965.

This locality was discovered in 1965 but apparently has
already been destroyed. The creek is part of a cattle
pasture. When visited in October 1968, the upper, less
disturbed portions of the creek were dry.

1969

THE MICHIGAN BOTANIST

97

LOCALITY

PRESENT STATUS

NORTH CAROLINA: 5V4 mi

SE of bridge on N.C. 49 over
Yankin River. Collected in

1953.

The locality is now under water as the river has been
dammed downstream.

OHIO: Streams west of

Toledo. Collected in 1900.

This and several other localities in the vicinity of

Toledo are now part of suburbia.

OHIO: End of swamp, E of
Indian Lake, Logan Co. Col¬
lected in 1928.

The area has been almost completely drained and is
now occupied by summer homes. Most of the trees
have been removed.

VIRGINIA: Potomac River at
Alexandria Light. Collected
in 1902.

The area is now an industrial site.

WISCONSIN: Bellevue Creek
cascade, Brown Co. Collected
1881-87.

The stream has been rerouted into an irrigation
ditch. The cascade is dry.

WISCONSIN: Stream where
North Shore crosses Howell

Ave., Milwaukee. Collected
in 1929.

The stream no longer exists.

WISCONSIN: Woods NE of

S 35th Street and Morgan Ave.,
Milwaukee. Collected in 1938.

The area is now part of suburbia.

The lack of morphological variability in P. cordata throughout its range,
its early blooming and short-lived seeds, its apparent lack of ecological plastici¬
ty, and its resulting rarity, all suggest a highly specialized plant well adapted
to a particular biological situation. Controlled experiments in both the green¬
house and in the field have not clearly elucidated the factors controlling the
occurrence and maintenance of a population of P. cordata under natural condi¬
tions, as will be discussed below, have only brought the problem into clearer
focus.

Mature plants of P. cordata will grow completely submersed for at least
four months. They will also survive three months without watering, although
under the latter conditions, they will drop their leaves and produce a winter
rosette. Six plants survived watering with a 10% NaCl solution for four
months after which time they were again given standard treatment. At first
the plants showed signs of plasmolysis but after a month and after losing the
original leaves, the plants recovered, produced new leaves (with a distinct salty
taste reminiscent of P. maritima), and appeared quite vigorous. Fifteen adult
plants were placed in a sunny, mesic field plot at the University of Michigan
Botanical Gardens in the fall. They survived the winter, bloomed and set seed
the following spring, survived the summer in a depauperate condition, and
have currently produced a winter rosette initiating their second dormant peri¬
od. The phenological experiments with controlled day-lengths have already

98

THE MICHIGAN BOTANIST

Vol. 8

been discussed above.

It is apparent that the adult plants of P. cordata can withstand consider¬
able “stress” for at least short periods of time. If there were an ecological
catastrophy during a particular growing season, a population would probably
survive. However, under none of the experimental conditions did the seeds
produce additional plants. The hypothesis is thus supported that the seeds and
seedling stages are the critical periods in the life history and biological poten¬
tial of the species. If the stress conditions lasted over several growing sea¬
sons— such as the draining of streams, the opening of the forest canopy, con¬
tinuous water pollution, damming of streams— the original population would
not reproduce and would die either through eventually succumbing to the
stress conditions or “old age.”

The critical data which are unfortunately lacking at this time concern
how P. cordata establishes new populations across an ecologically disharmoni¬
ous geographical area. Although Fig. 15 suggests that P. cordata is as “abund¬
ant” today as in the past, all extant stations are in areas which are relatively
undisturbed. There is little reason to believe that these areas have changed
much in the last 100-150 years. In light of our present knowledge, Fig. 15

Fig. 15. Longevity of known populations of Plantago cordata as indicated by their col¬
lecting history. Each arc represents the period of time (if two or more years) during
which specimens examined (see Appendix) were collected. (Dotted line for Hudson River
represents literature reports which may indicate the same population as more recent
[1933-1968] collections.)

1969

THE MICHIGAN BOTANIST

99

could be read as an extinction chart: All presently known stations may have
supported populations before European man’s disturbance of the area, but
gradually other populations have become extinct. As natural areas become
fewer and farther between, collectors have to botanize new areas and/or more
exhaustively investigate the diminishing older areas. In other case, P. cordata
has been and surely will be discovered in areas where it had previously not
been known. However, this does not mean that the population, per se, is new
but rather that the population only recently became known to science. Con¬
siderable insight on the biology of P. cordata could be obtained if the plant
suddenly appeared in a well known area which is ecologically suitable for the
species but which had not previously supported it. There are many streams in
the vicinity of known stations of P. cordata where this kind of study could be
carried out. Dispersal over larger areas, i.e., ten or more miles, would be inter¬
esting to study but extreme discontinuities between potential habitats prob¬
ably prohibit such dispersal today.

In general, when looking for possible stations of P. cordata, one should
search for shaded shallow streams with rocky bottoms in calcareous areas
where there is little evidence of recent disturbance to the canopy trees. Usual¬
ly the plantago occurs in streams at the bottom of ravines where little or no
alluvial deposition occurs. The plants always occur in the shallowest parts of
the stream and usually on gravel bars or rocky areas, but do not occur in
areas subject to periodic drought. Many of the streams are subject to high
flood levels in the spring as evidenced by the accumulation of debris around
trees and in the branches of nearby shrubs.

ETHNOBOTANY AND ECONOMIC USES

The search for P. cordata in the Thedford, Ontario, area resulted in a
first-hand account of the plant’s use as a medicinal herb. Insufficient data on
the exact location were given by Gaiser (1966) or on her labels (data supplied
by I. J. Bassett) for us to find the plant on our own. Consequently we located
Mr. Bill Henry, who is apparently respected among the Potawatomi Indian
community of Kettle Point as a “Root-man,” i.e. an herbalist. At first, he was
unwilling to reveal his locality for the plant, although he immediately recog¬
nized Plantago cordata from a sketch rapidly made by W. H. Wagner. But
after a short discussion which established his fee for services, he led us to the
colony and freely described the regular harvest of the plant, of which the
roots could be used either fresh or dried.

Among the virtues attributed to P. cordata, or “King Root,” at this time
were healing of burns and relieving of symptoms of sore feet when a tea of its
roots is drunk. Mr. Henry’s sister, Mrs. Baxter Shawkence, said it would “Heal
you when you hurt.” Bill Henry mentioned that sometimes P. cordata is mixed
with other herbs in a tea. He frequently mentioned that it was good for “50
diseases” but this may well have been an exaggeration. The plantago was sup¬
posed to be particularly good as a physic. If it worked “too well,” Henry said

100

THE MICHIGAN BOTANIST

Vol. 8

he had another plant that would “shut you off.” He also mentioned that P.
cordata “could kill you.” We were told that King Root was also “good for
claps” but were not given any details. On the way back to our car, he said “I
think it can cure stomach cancer.” I asked “You mean it helps a belly-ache?”
He said “Yes.”

Psyllium seed or Plantago seeds have long been used in many cultures as
a cathartic, and the leaves have been applied to wounds to promote healing
(Shyreu, 1935). H. H. Smith (1933) in discussing the use of Plantago major
[Plantago rugelii?] among the forest Potawatomi, refers to a slippery fluid ob¬
tained by boiling the root. This was then used to lubricate the throat when one
was choking or coughing. I attempted to make such a “tea” out of the roots
of P. major and P. rugelii but did not obtain a “slippery fluid.” However,
when P. cordata roots are treated similarly, a slimy gruel is readily obtained.
“King Root” may therefore have been used in this preparation rather than P.
major or P. rugelii , a theory supported by the anatomical evidence of abund¬
ant starch in the roots of P. cordata and little or no starch in the other spe¬
cies. Smith continues discussing the Indian uses of plantain, listing the use of
a heated leaf to bind on swellings and inflammations, of fresh leaves to dress
wounds and heal sores, and of a tea made from the plant for its supposed
alterative, diuretic, antiseptic, and antisyphilitic properties.

Material of P. cordata was given to Dr. Ara Paul, Department of Pharma¬
cognosy, University of Michigan, for analysis. His screening resulted in discov¬
ery of no particular medicinal properties of the plant, save its well-known
cathartic one.

Regardless of the after-effects, P. cordata can apparently be eaten as
food. Steyermark (1963) in his discussion of the species states that the
“young fleshy leaves and tender petioles may be cooked as a vegetable, and of
all the native plantains is the most tender.”

Plantago cordata is a showy plant, as pointed out by Steyermark (1963)
and has been cultivated in gardens since the time of Lamarck. Even today, it
occasionally appears in commercial plant catalogues specializing in aquatic
plants.

As a rare and fascinating species, hopefully, P. cordata ’s last domain will
not be that from which it was, perhaps ominously, originally named— a garden.

APPENDIX: SPECIMENS EXAMINED

CANADA. ONTARIO: Near Conrad River, Booth, May 17, 1863 (US); Near Amherst-
burg, Macoun, June 10, 1882 (US); Lambton Co., lVi mi N Thedford, Tessene, Hines,
Voss, & Wagner 1941, May 6, 1967 (MICH); Local in small stream and on muddy bank
where slightly marly, low deciduous woods (of Harvey Lockney) W of Hgy. 82, ca. 2 mi.
NW Thedford. Roots medicinal, fide Bill Henry, part-Indian herbalist of Kettle Point,
Voss 12424, May 6, 1967 (MICH). UNITED STATES. ALABAMA: Colbert Co., Shallow
rocky bed of Fox-trap Creek, Harper 3892, July 25, 1942 (MO, US); Wet rocky bank of
Ligon’s Creek about 2 mi W of Lusliville [ Littleville? ] , Harper 3933, April 20, 1943
(MO, US); Wilcox Co., Buckley, April 18, ’10 (NY). DISTRICT OF COLUMBIA: Vicinity
of Washington, Ward, May 2, 1881 (NY); Ward, May 18, 1884 (MSC). GEORGIA:
Catoosa Co., Shallow soil in Cedar Barrens 1.9 mi E, 18 mi N Fort Oglethorpe, Duncan

1969

THE MICHIGAN BOTANIST

101

& McDowell 12236, 31 March 1951 (MO); Frog Co., Banks of Hordey Creek near Rome,
Chapman 4366 (MO, US). ILLINOIS: Champaign Co., In running water, Urbana, Mamis
17606, May 6, 1882 (ILL); Christian Co., Taylorville, Andrews, 5-27. 99 (ILL); Cook
Co., Chicago, Boerde, May 22, 1886 (ISC, US); Glencoe, in sandy mud in small creek at
base of ravine, Gates 1692.1, June 22, 1907 (ILL, US); Johnson, May 8, 1889 (MICH);
In water, ravines at Winnetka, Moffatt, May 10/91 (ILL, MINN, NY); Rivulet, Avondale,
Moffatt 2495, May 14, ’94 (WIS); N of Chicago, Moffatt, June 10/94 (ILL); Ravines,
Glencoe, Pepoon 10187, June 9, 1898 (WIS); North Evanston, Price, 1893 (US); Edge of
streams, Evanston, Shipman, 1897 (MICH, PH); Englewood, Williamson, Aug. 1893 (PH).
Grundy Co., Morris, Slossow (PH). Hancock Co., Augusta, Mead, May, 1842 (NY )\ Mead
6681, May, 1860 (MO). Lake Co., Rivulets in ravine, Highland Park, Hill 16.1880, May
22, 1880 (ILL); Umbach, June 9, 1900 (MINN, PH, WIS). Jackson Co., Stream of NE
end of Lake Murphysboro, Tessene 2066, Oct. 13, 1967 (MICH). LaSalle Co., In water at
bottom of canyon, Matthiessen State Park, near Oglesby, G. Jones 32335, June 30, 1961
(ILL). Marshall Co., Along streams, Heading, May, 1879 (ILL). Menard Co., Revulets
[sic] , Athens, Hall 31313, April, 1863 (ILL). Peoria Co., Peoria, Brendel, May 22, 1859
(ILL); In brooks, Canton, Pekin, abundant, Buckley (NY). Piatt Co., Creeks, Mansville
[Mansfield?], Johnson, May 14, 1900 (US). St. Clair Co., Mascoutah, Welsch, 1862-1871
(ILL). Wabash Co., Chiefly about Mt. Carmel, Schneck (ILL). Will Co., In rivulet, New
Lenox, Hill 15.1879, May 17, 1879 (ILL). INDIANA: Boone Co., Lebanon, Cunky, May,
1861 (PH). Porter Co., Swampy ground by streams, Wheeler,///// 56.1884, July 4, 1889
(ILL). Wells Co., Rock Creek, Deam, 5-27-1900 (MINN). Whitley Co., Boggy stream on
Gradless farm about 7 mi E of Columbia City, Friesner 20424, 6-13-1946 (MICH, MO,
NY). MICHIGAN: Eaton Co., Bellevue, Mr. [D. A.] Shoop [ca. 1868] (MICH). Genesee
Co., Flint, Clark (MO, NY, OS, PH, US). Gratiot Co., Brooks and ditches in mud, vicinity
of Alma, Davis, June 10, 1890 (MSC); Davis, May 22, 1893 (US); Davis, May 26/93
(MINN); Davis, May 29/95 (ILL, MICH, WIS). Ingham Co., Lansing, [Cooley coll.] 5/5/81
(MSC); Dr. Beal’s woods, Agricultural College, Skeels, June 6, 1894 (MSC); Lansing,
Michigan Agricultural College, [Collector unknown], May 22, 1886 (MSC). Ionia Co.,
Wet ground, Hubbardston, Wheeler (MICH, MINN); Macomb Co., Vicinity of Washington,
Cooley, 1843 (MSC). St. Clair Co., Port Huron, Dodge 423 (MO); Dodge 6193 (MINN,
WIS); along shaded streams plentiful, Dodge, 5-22-92 (MICH); Dodge, May 9, 1897 (ILL);
along streams in woods, occasional, often water, Dodge, May 31, 1897 (MICH, MSC);
Wright, June 2, 1838 (MICH). Shiawassee Co., Owosso, in slow streams, Hicks, May 18,
1889 (BLH, MICH). Washtenaw Co., Along bank of creek in hardwoods at “Hickory
Flats” 5 mi S of Ann Arbor, Erlanson 29, May 15, 1924 (MICH); Ditch in low, rich
woods 5 mi S of Ann Arbor, Ehlers, May 16, 1924 (MICH); Ehlers 3154 (MICH); Ehlers
3156 (MICH, WIS); [Hammond woods near Ypsilanti. Frequent, fide Farwell’s field cata¬
log] , Farwell 1087, 5-28-91 (BLH). MISSOURI: Adair Co., Moist woods, bank along
stream and in water of stream in ravine, Tributary of Chariton River between Kirksville
and Moroings, Steyermark 18857, May 4, 1935 (US). Atchison Co., Near Fairfasse [Fair¬
fax?], Eggert, 20 May, 1887 (MO). Boone Co., Webster, Eggert, 24 May, 1887 (ISU,
MO). Dent Co., Growing in a creek, Miss Shoop’s farm, near Sligo, Lodewyks 284, May
12, 1935 (MINN, MO); Margin of stream bed in gravel where seepage accumulates along
north prong of Mehang River T34N, R4W, sec 11, Steyermark 12517, Aug. 4, 1936
(MO). Carter Co., Mann 2064, 5-1-1891 (MO). Howard Co., Along Creek around rocky
exposures, rich woods, slopes and ravine along Maniteau Creek from Blue Bluff south and
SW T51N, R14W, sec 10, 4 mi NE of Bunke Hill, 8 mi NE of Burton, Steyermark 74265
(MO). Jefferson Co., In shallow running water or rocky stream bed near mouth of Buff-
dale Hollow, SW of Selma, Steyermark 8048, April 15, 1934 (MO). Montgomery Co., In
creeks, Montgomery, Engelmann, April, 1863 (MO). Ripley Co., At base of limestone
bluff along spring branch of prong of south fork of Buffalo Creek, T34N, R1E, sec 35, 5
mi E of Bradley, Steyermark 11792, July 14, 1936 (MO). St. Louis Co., St. Louis, Buck-
ley (ISU, MO); Wet places, Eggert, 27 April, 1887 (ILL, MO, NY, US); In streams
and . . . [sic!] etc. west of St. Louis, Engelmann, April, 1833 (MO); Engelmann, April,
1842 (PH, US, WIS); In small rivulets, Geyer, April, 1842 (PH, MO); On bed of small

102

THE MICHIGAN BOTANIST

Vol. 8

stream, Flat Rock, Allen [ton?], Greenman 4214, April 15, (NY); Flat Rock, Hurshel
5378, April 15, 1923 (US); Allenton, Kellogg, June 18, 1886 (MO) ; Kellogg 15302, May
20, 1930 (MO ); Letterman, Dec. 23, 1879 (OS ); Letterman, Mar., 1880 (NY); Letterman,
April 10, 1888 (ILL); Letterman, 1890 (MO, NY, US); Letterman, July 1, 1894 (MO,
PH); Eaux Contantes, St. Louis, Riehl, April 1838 (NY); Sandy beds of streams, Kirks-
ville, Sheldon, 5/31/’83 (OS). Shannon Co., Monteer, Common in streams, Bush 373, May
24, 1900 (NY, US). Washington Co., Sugar Camp Hollow (tributary to Indian Creek)
T39N, R1W, sec 24, 5Vi mi (by air) E of Anthonies Mill, moist limestone rocks along
creek, Steyermark 77806, Sept. 27, 1954 (ILL); Common in spring branch and slough at
base of limestone bluffs along west fork of Fourche a Renault Creek, T37N, R1E, sec 27,
1 mi S of Shirley, Steyermark 78632, June 13, 1955 (ILL). NEW YORK: Columbia Co.,
N side of Chekomeko Creek, “Ralph Jenson Kill,” Livingston tnsp., Hines 6825 (MICH);
Along creek at RR bridge, Stockport Landing, Tessene, S.J. Smith, et al. 2065 (MICH).
Dutchess Co., Gravelly shores of the Stony Creek S of the Madaline Bridge, at Ward
Manor, Elwert, May 14, 1933 (NY); In brackish water on the gravelly shores of Crueger’s
Island in the Hudson, between the stations of Barytown and Tivoli, Elwert, June 8, 1933
(NY); Rocky banks of Imbrocht Bay, Muenscher & Curtis 5810, Sept. 3, 1936 (WIS);
South Bay, Muenscher & Curtis 5811, Aug. 31, 1936 (PH). Green Co., Tidal marshes of
Hudson River, Green Point 4 mi S of Catskill, House 25135, Sept. 2, ’37 (NY). NORTH
CAROLINA: Stanley Co., 5xh mi SE of bridge on N.C. 49 over Yadkin River, Radford
7071, June 4, 1953 (ILL). OHIO: Adams Co., Stream at Beaver Pond, Bartley & Pontius
958, May 27, 1945 (NY, US); Local in bottom of Plum Run Creek, by Plum Run Quar¬
ry, off rt. 31, ca. 2.6 mi E of Peebles, water and soil very limey, Meigs twp., Stuckey
2880, 29 May 1966 (OS); Tessene, W. H. Wagner, et al. 1881, Nov. 13, 1965 (MICH);
Tessene & Stuckey 1918, May 29, 1966 (MICH); Common in a small but fast-flowing
stream through juniper savana, N of Plum Run quarry, with Floyd Bartley, Voss 1531,
April 9, 1953 (MICH). Auglaize Co., St. Marys, Wetzstein, 22 April, 1897 (OS). Erie Co.,
Moseley, May 19, 1895 (BGSU). Champaign Co., From watery pools on Mad River,
Urbana, Samples, May 26th 1838 (MICH). Franklin Co., “Sellsville,” Columbus, Craig
6975, June 1889 (OS); Columbus, Kellerman, 1 June 1895 (OS); Lapham, April, 1836
(PH); Lloyd, April 1891 (ISU, NY); Osborn, July ’97 (OS);Selby, April 25, 1891 (OS);
Tyler, October 2, 1907 (OS); Werner, May, 1891 (OS); Georgesville, Werner, 24 Sept.
1892 (OS); Worthington, [Collector unknown], June (ISC). Logan Co., End of swamp
east of Indian Lake, Richland tnsp., Hine, 16 May, 1928 (OS). Lorain Co., Very wet
woods, infrequent, Oberlin, Cowles, May 21, 1892 (OS); Dick, 12 May, 1894 (OS); Dick,
4-29-95 (MINN); Ricksecher, May 12, 1894 (NY, US); shallow stream, South woods,
Strong, May 21, 1892 (MICH). Lucas Co., Stream 3 miles W of Toledo on Dorr St.,
Burglehaus, 6 May, 1900 (OS). Madison Co., London, Sharp, 1900 (OS). VIRGINIA:
Smyth Co., Vicinity of Marion, Alt. 2100 ft. Ravine of the Holston, N. & E. Britton &
Vail, June 8, 1892 (NY); On bar extending into Potomac River at Alexandria Light,
Chesapeake Bay rgion, Shull 217, Aug. 13, 1902 (MO, NY). WISCONSIN: Brown Co.,
Bellevue Creek Cascade, Schuette, May, 1887 (NY); Ledge near Bellevue Creek, Schuette,
May 15, 1881 (MINN). Kenosha Co., Somers, Wadmond, June 3, 1899 (MINN, WIS).
Milwaukee Co., Creek S.W. of Layton Park, Milwaukee, Finger, June 17,-01 (MIL); Root
River, Milwaukee, just north of Racine Co., line, Elm woods, growing in water (flood
pools), Fuller 4524, May 12, 1932 (MIL); Oak Creek, Goessl, Sept. 21, 1935 (WIS);
Milwaukee, Hassee 13557 (MIL); House, May ’84 (PH); Lapham (NY, WIS); In stream,
woods NE of S 35th and Morgan Ave., Shinners, May 11, 1938 (UWM); New Coeln,
Along ditch in woods, H. Smith, May 24, 1921 (MIL); Slow moving stream where North
Shore crosses Howell Ave., Throne, May 12, 1929 (UWM); Schwiechert’s Woods,
Wauwautosa, Wheeler 7596, June 24-26, 1887 (MIL). Racine Co., Racine, Davis, June 28,
1897 (WIS); Davis, May 21, 1880 (WIS); Root River, “near Milwaukee Co. line-LMS,”
Oppel U-15, Aug. 1935 (WIS). Outagami Co., Kellerman, June 1879 (US).

1969

THE MICHIGAN BOTANIST

103

ACKNOWLEDGMENTS

This research was supported, in part, by grants from the National Science Founda¬
tion, GB-3366 to T. H. Hubbell and GB-6230 to N. G. Hairston, of The University of
Michigan, for research in Systematic and Evolutionary Biology and by the University of
Michigan Graduate Student Research Fund. I wish to thank W. H. Wagner, Jr., Edward G.
Voss, and Ronald L. Stuckey for their interest and encouragement in this project, for
reading the manuscript, and, along with Robert H. Mohlenbrock and Stanley J. Smith,
for being willing to get their feet wet in search of this rare plant. I would also like to
express my gratitude to the directors and curators of the following herbaria who so gen¬
erously lent specimens: BGSU, BLH, ILL, ISC, MICH, MIL, MINN, MO, MSC, NY, OS,
PH, US, UWM, and WIS. The specimens in PH were examined for me by R. L. Stuckey.
Others who have contributed in various ways include Rogers McVaugh, I. John Bassett,
David M. Hines, and John Ludwig. Part of the cost of additional pages in this article has
been defrayed by the Asa Gray Bulletin Fund, originated by the late Prof. H. H. Bartlett.

REFERENCES

Arber, Agnes. 1918. The phyllode theory of the monocotyledonous leaf, with special ref¬
erence to anatomical evidence. Ann. Bot. 32: 465-501.

Bassett, I. J. 1967. Taxonomy of Plantago L. in North America: Sections Holopsylhum
Pilger, Paleopsyllium Pilger, and Lamprosantha Decne. Canad. Jour. Bot. 45:
565-577.

- 1968. Pollen morphology and chromosome numbers of the family Planta-

ginaceae in North America. Canad. Jour. Bot. 46: 349-361.

Cartier, D. 1963. Etude caryologique des diffe'rentes sous-especies du Plantago atrata.

Compt. Rend. Acad. Paris 256: 2900-2902.

Clute, W. N. 1942. Plantago cordata in Indiana. Am. Bot. 48: 95.

Dodge, C. K. 1916 [“1914”] . The flowering plants, ferns and fern allies growing without
cultivation in Lambton County, Ontario. Rep. Mich. Acad. 16: 132-200.

Esau, K. 1953. Plant Anatomy. John Wiley, New York. 735 pp.

Farwell, O. A. 1930. Botanical gleanings in Michigan, VI. Am. Midi. Nat. 12: 113-134.
Fassett, N. C. 1959. Spring Flora of Wisconsin. 3rd ed. University of Wisconsin Press. 189

pp.

Femald, M. L. 1950. Gray’s Manual of Botany. 8th ed. American Book Co., New York.
1632 pp.

Finch, V. C. 1957. Elements of Geography, Physical and Cultural. 4th ed. McGraw-Hill,
New York. 693 pp.

Gaiser, L. O. 1966. A Survey of the Vascular Plants of Lambton County, Ontario. Comp,
by R. J. Moore. Canad. Dep. Agr. 122 pp.

Gleason, H. A. 1952. The New Britton and Brown Illustrated Flora of the Northeastern
United States and Adjacent Canada, vol. 3. New York Botanical Garden, New
York.

- & A. Cronquist, 1963. Manual of Vascular Plants of Northeastern United

States and Adjacent Canada. D. Van Nostrand Co., Princeton, N. J. 810 pp.
Godfrey, R. K. 1960. Plantago cordata still grows in Georgia. Castanea 26: 119-120.
Gorenflot, R. 1959. Le polymorphisme de Plantago coronopus L. ses manifestations et
ses causes. Revue Cytol. Biol. Veg. 20: 237-497.

- 1961. Un exemple d’introgression experimental. Bull. Soc. Bot. Fr. 108:

1-16.

Harper, F., ed. 1943. Travels in Georgia and Florida, 1773-74: A report to Dr. John
Fothergill [by] William Bartram. Trans. Am. Philos. Soc. 33: 121-242.

Harper, R. M. 1944. Notes on Plantago, with special reference to P. cordata. Castanea 9:
121-130.

Hitchcock, A. E., & P. W. Zimmerman. 1947. Response and recovery of Dandelion and
Plantain after treatment with 2,4-D. Contr. Boyce Thompson Inst. 14: 471-492.

104

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Vol. 8

Hope-Simpson, J. F. 1939. Reduction of the androecium in Plantago lanceolata. Phyto¬
pathology 31: 1089-1098.

McVaugh, R. 1958. Flora of the Columbia County Area, New York. N. Y. St. Mus. Bull.
360. 433 pp.

Muenscher, W. C. 1937. Aquatic vegetation of the lower Hudson area. Chapter vii (pp.
231-248) in A Biological Survey of the Lower Hudson Watershed, Rep. St. N. Y.
Conserv. Dept. 26, suppl. (Biol. Surv. 11).

Pilger, R. 1937. Plantaginaceae in Pflanzenreich IV: 269 (Heft 102). 466 pp.

Radford, A. E. 1954. Range extensions in the flora of North Carolina II. Jour. Elisha
Mitchell Sci. Soc. 70:101-104.

Rahn, K. 1957. Chromosome numbers in Plantago. Bot. Tidsskr. 53: 369-378.

Shyreu, E. W. 1935. Drugs derived from the genus Plantago with botanical sources. Jour.
Pharmacology 1935: 1-12.

Small, J. K. 1933. Manual of the Southeastern Flora. Publ. by the Author. 1554 pp.
Smith, H. H. 1933. Ethnobotany of the forest Potawatomi Indians. Bull. Public Mus. Mil¬
waukee. 7: 1-230.

Stebbins, G. L., & Alva Day. 1967. Cytogenetic evidence for long continued stability in
the genus Plantago. Evolution 21: 409428.

Steyermark, J. A. 1963. Flora of Missouri. Iowa State Univ. Press, Ames. 1725 pp.
Svenson, H. K. 1935. Plants from the estuary of the Hudson River. Torreya 35: 117-125.
Tessene, M. F. 1968. Preliminary reports on the flora of Wisconsin no. 59. Planta-
ginaceae-Plantain Family. Trans. Wis. Acad. 56: 281-313.

Tyler, F. J. 1901. Geophilous plants of Ohio, II. Ohio Nat. 2: 132-133.

Wiegand, K. M., & A. J. Eames. 1926. The Flora of the Cayuga Lake Basin, New York,
Vascular Plants. Cornell Univ. Agr. Exp. Sta. Mem. 92. 491 pp.

A NEW LOCATION FOR PORELLA PINNA TA
IN LOWER MICHIGAN1

John S. Russell
School of Natural Resources,

The University of Michigan, Ann Arbor

On August 11, 1965, during a search for aquatic flowering plants, a
large liverwort subsequently identified as Porella pinnata L. was collected
( Russell , UMBS, MICH) near the mouth of Mud Creek at the west side of
Black Lake in Cheboygan County. Specimens (Fig. 1) were found in large,
luxuriant, olive-green or blackish clumps on tree roots submerged by as much
as two feet of murky water. Visual evaluation of the habit of the plant was
not possible owing to the darkness of the water, although it appeared to occur

I Contribution from the University of Michigan Biological Station.

1969

THE MICHIGAN BOTANIST

105

in proximity to, and often mixed with, Riccia fluitans L. Fragments of Porella
were seen floating near the surface of the water, and depauperate clumps were
observed on tree roots above the surface. The creek is cold, slow-moving, and
probably slightly alkaline. The site was visited two days later and additional
material collected by Miller and Crum (1966).

*

Fig. 1 (left). Porella pinnata, about
natural size, from Mud Creek near
Black Lake.

Fig. 2 (below). A portion of the same
collection, enlarged about 8 times.

106

THE MICHIGAN BOTANIST

Vol. 8

The only previous authenticated collections of Porella pinnata from
Michigan are those made by Nichols (1935, 1938) in Marquette County in the
Upper Peninsula and Gilbert (1958) in Iosco County in the Lower Peninsula.
The present find confirms the reported occurence of this species in the Uni¬
versity of Michigan Biological Station region (Gilbert, 1958). Both of Nichols’
specimens were found attached to submerged rocks, while Gilbert listed “sub¬
merged rocks and other materials” as the substrate. An investigation of data
accompanying specimens from North America in the University of Michigan
Herbarium confirms the fact that in northerly locations Porella pinnata is al¬
most always found submerged, while on more southerly sites it frequently
thrives on moist soil, and sometimes in rich woods. Schuster (1953) noted
that in the South it occurs on cypress knees, on the bark of beeches, or as a
xerophyte on rocks far from any source of constant water. He suggested the
existence of a diverse series of genotypes to account for this apparent broad
range of ecological tolerance, to which Conard has also recently called atten¬
tion (1968).

Because of its preference for submerged substrates in the northerly
reaches of its range, Porella pinnata may be more common in this area than
heretofore suspected. A more thorough knowledge of the ecology of its pre¬
ferred habitat may indicate that this liverwort is not rare, but merely local in
occurrence and difficult to find. Plants of this species are characterized by
leaves scarcely touching, narrow non-decurrent underleaves, and similarly
tongue-shaped underlobes (Fig. 2).

According to distribution records given by Frye and Clark (1946),
Porella pinnata L. is widespread in the South and East. I have examined speci¬
mens from Vt., Mass., R.I., N.J., Pa., Va., W.Va., N.C., S.C., Ga., Fla., Tenn.,
La., Tex., Mo., Ark., Ind., Ill., Ohio, Mich., Ont., N.S.

LITERATURE CITED

Conard, H. S. 1968. Porella pinnata in North America. Bryologist 71: 29-30.

Frye, T. C., & Lois Clark. 1946. Hepaticae of North America. Part IV. Univ. Wash. Publ.
Biol. 6: 565-733.

Gilbert, W. J. 1958. Porella pinnata in Michigan. Bryologist 61: 246-7.

Miller, N. G., & Howard Crum. 1966. Bryophytes new to the Douglas Lake area. Mich.
Bot. 5: 57-61.

Nichols, G. E. 1935. The bryophytes of Michigan with special reference to the Huron
Mountain region. Bryologist 38: 11-19.

- . 1938. Bryophytes of the Huron Mountain region, Marquette County, Michi¬
gan. Bryologist 41:25-31.

Schuster, R. M. 1953. Boreal Hepaticae: A Manual of the Liverworts of Minnesota and
Adjacent Regions. Am. Midi. Nat. 49: 257-684.

Editorial Notes

We hope to see all members and friends of the Michigan Botanical Club at the
annual Spring Campout, to be held this year in the Manistee National Forest, near Irons,
in Lake County, Michigan. Full information will be sent to all members in advance.

With some pride, we observe that, entirely by coincidence, all of the excellent arti¬
cles in this issue are by graduate students at Michigan universities.

The January number (Vol. 8, No. 1) was mailed January 3, 1969.

Errata

In addition to a few more obvious transpositions of letters or other spelling altera¬
tions, the following may be noted:

Vol. 7, No. 3, p. 141, line 3 from bottom: for “4183” read 4138
p. 188, line 8 from bottom: for “44:” read 45:

Vol. 8, No. 1, p. 39, line 19: for “east” read ease

Publications of Interest

(Continued from p. 66)

PAPERS ON PLANT SYSTEMATICS. Selected by Robert Omduff. Little, Brown & Co.,
Boston, 1967. 429 pp. $6.00. This is an anthology of 14 biosystematics papers select¬
ed from the massive literature of experimental taxonomy. Included are papers pub¬
lished in 14 different journals, mostly in America and mostly in the last 20 years. In
his preface to the ecology section, Omduff notes that the role of ecological studies in
plant systematics is not as influential as it should be. Systematists are largely ignoring
potentially profitable and simpler techniques of ecology while using the more complex
techniques of numerical and chemical taxonomy. Overall, the book is an excellent in¬
troduction to the literature of biosystematics. The papers illustrate the major areas of
morphological, cytological, biochemical, and ecological approaches to solving system¬
atic problems.

— E. L. McWilliams

WEEDS OF THE WORLD. By Lawrence J. King. Interscience Publ., New York, 1966.
525 pp. $18.00. Although there are numerous books available on the subject of
weeds, this one is unique both in its scope and in its strong emphasis on the biology
of weeds. Taxonomic identification of weeds is not considered. The topics covered
include origins of weeds, ecological classification of weeds, parasitic weeds, germina¬
tion, establishment, growth, dispersal, and evolution of weeds. There is an entire chap¬
ter on weed sociology with a bibliography of over 175 references. The extensive bibli¬
ographies at the end of each chapter are among the most useful aspects of the text.
The book is well illustrated with photographs and diagrams. The last section deals
with weed control, both chemical and non-chemical, with some consideration to the
impact of herbicides on our ecosystem. This volume will be useful to a variety of
biologists, especially students of weed biology, as well as agriculturists.

— E. L. McWilliams

CONTENTS

t

A Phytogeographical Analysis of a Southern
Michigan Bog

Garrett E. Crow . 51

White Pine at the Edge of Its Range, in
Oakland County, Michigan

Sylvia Taylor . 61

Publications of Interest . 66, 107

The Occurrence of the Lichen Complex, Cetraria
ciliaris, in the Straits Region of Michigan

William L. Graham . 67

Systematic and Ecological Studies on
Plantago cor data

Melvern F. Tessene . 72

A New Location for Porella pinnata in Lower Michigan

John S. Russell . 104

Editorial Notes . 107

(On the cover: Spring is coming!

April photo of Hepatica americana taken near
Midland , Michigan, by Harold F. Tweedie.)

33

THE

LIBRARY

MAY 28 1969

NEW YORK
BOTANICAL GARDEN

Vol. 8, No. 3

MICHIGAN BOTANIST

May, 1969

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THE MICHIGAN BOTANICAL CLUB

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1969

THE MICHIGAN BOTANIST

111

THE INTRODUCTION AND SPREAD OF
LYCOPUS ASPER (WESTERN WATER HOREHOUND)

IN THE WESTERN LAKE ERIE AND
LAKE ST. CLAIR REGION1

Ronald L. Stuckey
College of Biological Sciences,

The Ohio State University, Columbus

In many studies of eastern North American plants distinctions are usual¬
ly made between those species which are native (or indigenous) and those
which are introduced (or non-indigenous). This distinction is often clear and
seems to present few problems. The basic idea is that those species which
were here originally (i.e., before the coming of European man) are native, and
those species that have appeared since the arrival of European man and that
are known to be indigenous to another continent have been introduced, have
spread, and in many cases have become established as part of the flora. Little
evidence is ever presented that a native species may be expanding its range,
and that in reality it, too, is a newcomer or non-indigenous member of a
particular floristic region. These non-indigenous members may be considered
truly part of the original flora until their proper status is determined.

Because habitats and the environment of the continent are continually
changing, often as the result of man’s activities, changes in the distribution of
certain species of the native flora are to be expected. It is imperative that
those studying plants understand the changing flora in order to evaluate man’s
impact on the environment. The distributions of individual species give clues
to these changes, which can be recognized only after a comparison is made of
information from past literature and herbarium specimen records with modern
surveys of plants in the field. This paper brings together data from these
sources as evidence to show that Ly copus asper Greene (western water hore-
hound), a species indigenous to western North America, is non-indigenous to
the shore flora of the western Lake Erie and Lake St. Clair region.

Western water horehound (fig. 1), a member of the mint family (Labi-
atae), is a short, 4-8-dm-tall plant with sessile, toothed, oblong-lanceolate,
spreading to ascending leaves. The white-petaled flowers are somewhat incon¬
spicuous in the axils of leaves, where later in the year four nutlets appear.
Vegetative reproduction is by short, thick stolons and runners, which form
long, thick tubers at their tips. Because of the absence of showy parts, the
plants blend together with other members of the community, and therefore
individuals of L. asper may go unnoticed in the field.

Henderson (1962), in his taxonomic revision of the genus Ly copus,
maps the known occurrence of L. asper for North America. The species is
distributed throughout western United States and southern Canada and from
his map appears to be more commonly found in western Minnesota, western

^Contribution from the Botany Program (Paper No. 756), the Herbarium, and The
Franz Theodore Stone Laboratory.

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Fig. 1. Ly copus asper Greene (western water horehound) among cat-tails in wet sand at
the east end of Erie Road along the shore of Lake Erie, Monroe County, Michigan. Photo¬
graphed 14 September 1968 by William H. Anderson.

Iowa, Nebraska, South Dakota, and North Dakota. Henderson points out that
a gap exists in its distribution in eastern Iowa and Minnesota, and throughout
most of Wisconsin. Across northern Illinois and about Lake Michigan, Lake
Huron, and western Lake Erie the species occurs at several stations. Henderson
suggests that this distribution pattern could indicate either an accidental intro¬
duction and establishment eastward or a lack of collecting in the intervening
area, or both. He concludes, however, that this distribution perhaps reflects
the actual occurrence of this species rather than omissions in collecting.
Lernald (1950) points out that L. asper is occasionally adventive along railways
eastward in its range. Gleason (1952) notes that it is rare and probably adventive
east to southwestern Ontario. In Illinois, where it is known from Bureau, White-
side, Cook, Henry, and Lake counties, L. asper is treated as a native species by

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113

Dobbs (1946), Jones et al. (1951), Jones and Fuller (1955), and Jones (1963).
Deam (1940) excludes western water horehound from Indiana because an earlier
report was based on a misidentified specimen. The species was found in 1946 in
Lake County, Indiana, along a weedy shore of Wolf Lake opposite the city park at
Whiting ( Friesner 21121, MICH, US). Henderson (1962), who also observed L.
asper at the same lake shore, reports that this is the only known Indiana locality.
Koeppen (1958) reports L. asper as occasionally adventive in Wisconsin along rail¬
road tracks and Lake Michigan harbors in Sheboygan, Waukesha, and Milwaukee
counties. Because Waterman (1960) does not indicate that L. asper is non-
indigenous in Michigan, her treatment implies that the species is native where
it occurs in the counties of Berrien, Cheboygan, Mason, Monroe, St. Clair, and
Wayne, all of which border the Great Lakes and connecting waters. Waterman
(personal communication) has informed the author that she did not consider
or attempt to evaluate the status of L. asper in the Michigan flora at the time
of her study. Voss (1957) says that Michigan is apparently the eastern limit
of its native range, but believes western water horehound is not indigenous to
Cheboygan county. In papers concerned with introduced plants, Montgomery
(1957) and Rousseau (1968) do not mention L. asper for Ontario and
Quebec, respectively. Isolated localities for western water horehound are
known in Ontario at Hamilton, Wentworth County ( Caesar , 8 Aug 1957,
HAM; Tamsalu 1 Aug 1957 and 22 Jul 1958, HAM), at Keene, Peterborough
County ( Campbell , Aug 1922, DAO), at Rideau River near White’s Bridge,
Carleton County ( Minshall 3895, 16 Aug 1947; Dore & Erskine 14086, 13
Sep 1952; Dore 18507, 4 Sep 1960; Dore, Koyama, Gillett 20007, 22 Jul
1962, all DAO); in Quebec at Saint Nicholas (Marie- Victoria & Rolland-
Germain 56805, 24 Aug 1942, photo DAO) and Pointe Mesakonan (Dutilly &
Lepage 36732, 16 Aug 1958, photo DAO); and in Bergen Swamp, Genesee
County, New York, where Zenkert (1934) reports it as very rare, based on
House’s collection of 1916. Muenscher (1946) does not mention the species
from Bergen Swamp. From the above information it appears that in the Great
Lakes region L. asper is widely scattered and its status is somewhat confused.
Careful examination of the available evidence presented below leads me to
conclude that L. asper is non-indigenous at least to the region about the
western end of Lake Erie and Lake St. Clair.

Because of a long history of floristic study, the western Lake Erie-Lake
St. Clair region is particularly well suited to the study of the changing
aquatic and shore flora. Two of the earliest and very detailed aquatic plant
surveys for the United States were conducted on portions of these two lakes
(Pieters, 1894, 1901). Through the years various investigators have reported
on the flora of the surrounding region as well as on these lakes. Newberry
(1860), Beardslee (1878), Kellerman and Werner (“1893” [1894]), Kellerman
(1899), Schaffner (1914, 1932), and Weishaupt (1960, 1968) do not list L.
asper for Ohio. For Michigan the species is not mentioned by Wheeler and
Smith (1881) or by Beal and Wheeler (1892), but Beal (1905) included it
under the name Lycopus lucidus Turcz., now known to have been misapplied
to American plants. Macoun (1883) did not list it for southwestern Ontario.

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Vol. 8

Several studies on the aquatic and shore plants of more limited portions
of the western Lake Erie-Lake St. Clair region provide documentation for the
presence or absence of L . asper and the approximate time when western water
horehound appeared. The species is not reported for Lake St. Clair by Pieters
(1894), for Detroit (Farwell, 1902), for parts of the Detroit River (Foerste,
1882; Campbell, 1886), for the Canadian shore of western Lake Erie (Burgess,
1888-9; Dodge, 1914; Montgomery, 1957), or for the Ohio portions of western
Lake Erie (Claassen, 1892; Moseley, 1899; Pieters, 1901 ; Burr, 1901 ; Schaff-
ner, 1902; Kellerman, 1904; Kellerman and Jennings, 1904; Fullmer, 1916;
Core, 1948, 1949; Pinkava, 1963; Wood, 1966; Albright, 1966; Lowden, in
press). Specimens of L. asper have not been found, under any name, among
the collections of most of these botanists.

Dodge (1900) is the earliest known writer to mention L. asper in the
western Lake Erie-Lake St. Clair region, as well as for the entire state of
Michigan. Using the name L. lucidus, he reported it as “plentiful about the
elevators in the city of Port Huron. Only locality noticed. An immigrant from
the west.” Dodge’s specimens at MICH were obtained in 1892 and 1896.
Dodge (1910) did not mention western water horehound in a preliminary
report on the flora of Lambton County, Ontario, but he later (1916) wrote
that the plants were “well established around grain elevators and along rail¬
ways” in Lambton County. The species was not known farther south until
Farwell (1918) reported it from the banks of the Detroit River at Wyandotte,
based on a specimen collected 20 August 1916. Billington obtained a speci¬
men from Grosse Isle along the Detroit River on 4 September of the same
year. The earliest specimen known for the shore of western Lake Erie dates
from August 1920 and was collected by Moseley at Toledo Beach in Monroe
County. Based on field work in 1949, McDonald (1951) recorded the species
as rare on the beach at Pointe Mouille'e Marsh.

L. asper was not known in the Lake St. Clair vicinity until 1920 and
1924 when the plants were found on Squirrel Island in the St. Clair River
( Farwell 5680%) and Walpole Island {Farwell 7085), respectively. The species
was again found on Walpole Island in 1944 {Groh 2271). In 1948 plants were
obtained from near waste ground on the bank of the Sydenham River at
Wallaceburg {Bassett & Mulligan 772) and from sandy soil near the Thames
River Light House {Soper & Dale 4077). Although not specifically studying
the shore plants of Harsens Island, Hayes (1964) did report some of the
species of wet habitats, such as Juncus balticus, J. torreyi, Lycopus ameri-
canus, Lythrum alatum, Potentilla anserina, and Verbena hastata, but he did
not mention Lycopus asper. Gaiser and Moore (1966) list L. asper for Walpole
Island only. Western water horehound’s present distribution in the Lake St.
Clair region bears investigation.

In the Ohio portion of western Lake Erie, the flora of the islands and
nearby mainland has been extensively studied by professionals, amateurs, and
students, many of whom are or were associated with the botany program at
The Franz Theodore Stone Laboratory. However, L. asper was not known as
part of the flora until a plant was obtained from Honey Point on North Bass

1969

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115

Island by Doris Albright in 1965 (not reported by Albright, 1966) and from
the shallow water over dolomite rock on the south shore of East Point on
South Bass Island by me in 1967 (Stuckey, 1968a). In August 1968, I ob¬
served four plants on the sandy beach of Lake Erie at Lakeview Park in Port
Clinton. Western water horehound may have first appeared at this site in 1968
because paints were not found there when I extensively surveyed the Park’s
shore in 1967. In 1968 the plants were in places where fresh sand, washed by
the waves earlier in the season, came into contact with the stable sand of the
beach. In September 1968, I saw numerous plants in wet beach sand at the
east end of Erie Road and at Sterling State Park along the western shore of
Lake Erie in Monroe County, Michigan. It appears that western water hore¬
hound is spreading throughout western Lake Erie, particularly at sites where
there is some beach sand, such as at Honey Point, Lakeview Park, Erie Road,
and Sterling State Park. I have searched especially for this species in beach
sand blown in among cat-tails along the lake shore at Magee Marsh (Crane
Creek State Park) in the southeastern corner of Lucas County. Other sites in
western Lake Erie where sand is common and where I have conducted exten¬
sive field work and found no L. asper are Middle and East Harbor (East
Harbor State Park), Sand Point Marsh at the southeastern corner of the Mar¬
blehead Peninsula, the north shore of Kelleys Island at the State Park, and
along the beach and about the ponds at the north and south points of Pelee
Island. Based on its apparent present colonization behavior, L. asper would be
expected to invade these sandy beach sites earlier than other less favorable
habitats such as those along the rocky and rubble shores.

How the plants have spread in western Lake Erie is an interesting ques¬
tion about which we have little information. The location of plants along the
relatively fresh sand at Lakeview Park suggests that the seeds were washed
ashore and then germinated. Ridley (1930, p. 222) points out that Ly copus
europaeus (a species of similar habitats and seed morphology) is specially
adpated for dispersal in rivers. Its seeds apparently remain viable after floating
from 12 to 15 months. Because L. asper is a rather inconspicuous member of
the flora at locations where plants have been found, it is doubtful that man
has deliberately been responsible for its dispersal in our region.

The apparent distributional history of L. asper is remarkably similar to
that of Butomus umbellatus (flowering-rush) (Stuckey, 1968b). Both species
appear to have moved into western Lake Erie from along the Detroit River.
Knowledge of the distributional history of flowering-rush in our area is cer¬
tainly more complete than it is for L. asper, because the former is a much
more conspicuous plant. Its showy pink flowers on tall stalks are easily seen
by both professional and amateur botanists throughout the entire summer and
fall. B. umbellatus is also more likely to have been moved around, planted,
and cultivated by man for its showy flowers and possible food source for
wildlife, than is the western water horehound.

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Vol. 8

Fig. 2. Known distribution of Lycopus asper Greene in the western end of Lake Erie and
Lake St. Clair region based on herbarium specimens cited in this paper. The date(s) indicate
the known collection(s) from each locality represented by a dot. The arrows indicate the
probable pathway by which the species has migrated. This pathway follows the general
flow of water through the St. Clair River, Lake St. Clair, the Detroit River, and south¬
eastward through the western end of Lake Erie. Letters along the south shore and islands
of Lake Erie represent locations of sandy beaches where I have searched unsuccessfully for
L. asper.

A: Crane Creek State Park

B: East Harbor State Park

C: Fox’s Pond and Fishing Point, Pelee Island

D: Kelleys Island State Park

E: North Lagoon and Lighthouse Point, Pelee Island

F: Sand Point Marsh

1969

THE MICHIGAN BOTANIST

117

SUMMARY AND CONCLUSIONS

A review of literature and herbarium records, together with field studies, has made
it possible to work out the probable distributional history of Lycopus asper, a species
native to western North America, in the region about the western end of Lake Erie and
Lake St. Clair. In recent literature it has been treated in our region as an indigenous
species, perhaps because it occurs in naturally disturbed habitats, such as wet, shifting
beach sands. The conclusion that L. asper has been introduced and then spread in our
region is based on

1. The assumption that field studies in this region have adequately sampled the
flora over the past 80 years.

2. Absence of the species from most of the early floristic literature of Michigan,
Ohio, and southwestern Ontario, as well as from studies on small areas within
this region. Even though any one individual studying and reporting on the flora
of a given area might have overlooked L. asper , the fact that a large number of
early investigators of this region did not find and report it is indicative that the
species must have been absent or very local.

3. The earliest known records of Dodge (1900, 1916), who reported that the
species was a western immigrant found about grain elevators and along railways
in Port Huron and the surrounding area. Dodge knew the flora well in the field
and his specimens are labeled with data similar to his published reports.

4. The subsequent dates and locations of plant collections following Dodge, which
reveal the approximate time and the probable pathway that L. asper has used in
spreading southward through the St. Clair River, Lake St. Clair, the Detroit
River, and western Lake Erie (fig. 2). Its direction of spread in western Lake
Erie is consistent with the prevailing direction of surface water flow (see map,
Federal Water Pollution Control Administration, 1968, fig. 25). The conclusion
that L. asper has had this distributional history further assumes that separate
introductions did not occur, as suggested by the absence of the species from
Ohio floristic literature concerning western Lake Erie until 1968.

SPECIMENS EXAMINED FROM THE
WESTERN LAKE ERIE-LAKE ST. CLAIR REGION

MICHIGAN: MONROE CO.: Monroe Piers, 27 May 1924, Farwell 7074 (MICH);
barrier beach, Pointe Mouille'e State Game Area, 10 Aug 1949, McDonald 5386 (MICH);
Toledo Beach, 6 Aug 1920, Moseley s.n. (BGSU); locally common in wet sand along edge
of cat-tail marsh along shore of Lake Erie at the e end of Erie Road, n edge of Sec. 14,
Erie Twp., ca. 3 mi e of the town of Erie, 14 Sep 1968, Stuckey 7757 (OS, PH);
occasional in wet sand on beach along shore of Lake Erie, s w corner of Sec. 35,
Frenchtown Twp., n edge of Sterling Monroe State Park, e edge of the city of Monroe,
14 Sep 1968, Stuckey 7761 (OS); frequent in low hollow back of sandy shore of Lake

Erie, n edge Sec. 14, Erie Tp., ca. 3 mi e of Erie, 14 Sep 1968, Voss 12843 (MICH,

MSC). ST. CLAIR CO.: Plentiful, about grain elevator, city of Port Huron, 25 Aug 1892,
Dodge s.n. (MICH); plentiful, near grain elevator, city of Port Huron, 20 Aug 1896,
Dodge s.n. (MICH); frequent along railroads and grain elevators, near Port Huron, Dodge
s.n., 5 Aug 1904 (US). WAYNE CO.: Low, damp ground, Grosse Isle, 4 Sep 1916,
Billington s.n. (MICH); Wyandotte, 20 Aug 1916, Farwell 4384 (MICH); River Rouge, 9
Sep 1930, Farwell 8767 (MICH).

OHIO: OTTAWA CO.: Honey Point, North Bass Island, Albright 105 (OS); local
(three plants), in shallow water along dolomite shore, s e side of East Point, South Bass
Island, ca. 14 mi s of Buckeye Island, 11 Aug 1967, Stuckey 4950 (MICH, OS); portion

from one of four plants seen on wet sandy beach, NE !4 Sec. 5, T6N, R17E, Portage

Twp., Lakeview Park, e edge of Port Clinton, 19 Aug 1968, Stuckey 7450 (CAN, DAO,
MICH, OS, PH).

ONTARIO: ESSEX CO.: Sandy soil dredged from drain 0.5 mi s of Thames River
Light House, Tilbury Twp., 26 Jul 1948, Soper & Dale 4077 (DAO, US). LAMBTON

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THE MICHIGAN BOTANIST

Vol. 8

CO.: Near waste ground, bank of Sydenham River, Wallaceburg, 24 Jul 1948, Bassett &
Mulligan 772 (DAO); Squirrel Island, 16 Sep 1920, Farwell 568014 (MICH); Walpole
Island, 3 Sep 1924, Farwell 7085 (MICH); Walpole Island, 25 Jul 1944, Groh 2271
(DAO).

ACKNOWLEDGMENTS

The field study was conducted as part of the teaching and research program at The
Franz Theodore Stone Laboratory, Put-in-Bay, Ohio, during the summers of 1967 and
1968. Financial support for field work has been provided by The Ohio Biological Survey.
I am particularly grateful for the field and laboratory assistance of Mr. W. Alan Wentz,
Mr. Ronnie Johnson, and Mr. Harold Harlan. My thanks is also extended to Mr. Wentz
for the preparation of the typewritten manuscript for publication and to Mr. William H.
Anderson for the photograph of Lycopus asper.

I have examined all specimens cited immediately above. All specimens cited else¬
where in the text have been seen except those from the Royal Botanical Gardens Herbari¬
um (HAM), Hamilton, Ontario, for which the data were supplied by Dr. James S. Pringle.
Sincere appreciation is given to the curators of the various herbaria. Specimens or records
of specimens have been seen from the following herbaria: BGSU, CAN, DAO, HAM,
MICH, MO, NY, OS, PH, TRT, US.

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Albright, Doris Marilyn. 1966. Vascular plants of Honey Point, North Bass Island, Ottawa
County, Ohio. M. Sci. Thesis, Ohio State Univ. 47 pp.

Beal, W. J. 1905. Michigan Flora. Rep. Mich. Acad. 5: 1-147.

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(1891) 30: 471-650. (Reprinted and repaged 1-180.)

Beardslee, Henry C., Sr. 1878. Catalogue of the plants of Ohio, including flowering plants,
ferns, mosses and liverworts. Ann. Rep. Ohio State Board Agric. (1877) 32: 335-363.
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Burr, Harriet G. 1901. Plant study at Sandusky Bay. Ohio Nat. 1: 93-94.

Campbell, Douglas H. 1886. Plants of the Detroit River. Bull. Torrey Bot. Club 13: 93-94.
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Core, Earl L. 1948. The Flora of the Erie Islands. An Annotated List of Vascular Plants.
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- . 1949. The plants of western Lake Erie, after fifty years. Unpublished

Manuscript. 28 pp.

Deam, Charles C. 1940. Flora of Indiana. Dep. Conservation, Div. Forestry, Indianapolis.
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Dobbs, Raymond J. 1946. Illustrated notes on the flora of Henry County. Trans. Ill. Acad.
39: 47-49.

Dodge, C. K. 1900. Flora of St. Clair County, Michigan, and the western part of Lambton
County, Ontario. Ann. Rep. State Hort. Soc. Mich. (1899) 29: 231-314.

- . 1910. Plants growing wild and without cultivation in the county of Lambton,

Ontario. Ottawa Nat. 24: 45-52.

- . 1914. Annotated List of Flowering Plants and Ferns of Point Pelee, Ont., and

Neighbouring Districts. Canad. Dep. Mines. Geol. Surv. Mem. 54. No. 2, Biol. Ser.
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- . 1916. The flowering plants, ferns and fern allies growing without cultivation

in Lambton Co., Ontario. Rep. Mich. Acad. 16: 132-200.

Farwell, O. A. 1902. A catalogue of the flora of Detroit. Rep. Mich. Acad. 2: 31-68.
- . 1918. Rare or interesting plants in Michigan. Rep. Mich. Acad. 19: 251-262.

1969

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119

Federal Water Pollution Control Administration. 1968. Lake Erie Environmental Summary
1963-1964. U. S. Dep. Interior, Federal Water Pollution Control Administration,
Great Lakes Region. 170 pp.

Femald, Merritt Lyndon. 1950. Gray’s Manual of Botany. 8th ed. American Book Co., New
York, lxiv + 1632 pp.

Foerste, Aug. F. 1882. Plants of Belle Isle, Michigan. Bot. Gaz. 7: 202-203.

Fullmer, E. L. 1916. The Toledo Cedar Point. Ohio Jour. Sci. 16: 216-218.

Gaiser, Lulu O., & Raymond J. Moore. 1966. A Survey of the Vascular Plants of Lambton
County, Ontario. Plant Research Institute, Canad. Dep. Agr. Ottawa. 122 pp.

Gleason, Henry A. 1952. The New Britton and Brown Illustrated Flora of the Northeastern
United States and Adjacent Canada. Vol. III. N. Y. Bot. Gard., New York. 589 pp.

Hayes, Bruce. 1964. An ecological study of a wet prairie on Harsens Island, Michigan. Mich.
Bot. 3: 71-82.

Henderson, Norlan C. 1962. A taxonomic revision of the genus Lycopus (Labiatae). Am.
Midi. Nat. 68: 95-138.

Jones, George Neville. 1963. Flora of Illinois. 3rd. ed. Am. Midi. Nat. Monogr. 7. 401 pp.

- , & George Damon Fuller. 1955. Vascular Plants of Illinois. Univ. Ill. Press,

Urbana. 593 pp.

■ - - , H. E. Ahles, G. D. Fuller, & G. S. Winterringer. 1951. Additional records of

some Illinois vascular plants. Am. Midi. Nat. 45: 500-503.

Kellerman, W. A. 1899. The fourth state catalogue of Ohio plants, consisting of a serially
numbered systematic check list of the pteridophytes and spermatophytes. Bull. Ohio
State Univ. Ser. 4, 10 (Bot. Ser. 1): 1-65.

- . 1904. Flora of Hen and Chicken Islands, 1903. Ohio Nat. 4: 190-191.

- , & O. E. Jennings. 1904. Flora of Cedar Point. Ohio Nat. 4: 186-190.

- , & W. C. Werner. “1893” [1894]. Catalogue of Ohio plants. Rep. Geol.

Surv. Ohio 7: 56-406.

Koeppen, Robert C. 1958. Preliminary reports on the flora of Wisconsin. No. 41.
Labiatae-Mint Family. Trans. Wis. Acad. 46: 115-140.

Low den, Richard M. [In Press.] A vascular flora of Winous Point, Ottawa and Sandusky
Counties, Ohio. Ohio Jour. Sci.

Macoun, John. 1883. Catalogue of Canadian Plants. Part I.-Polypetalae. Geol. Surv. Canad.
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McDonald, Malcolm E. 1951. The ecology of the Pointe Mouillee Marsh, Michigan, with
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Arbor. 243 pp. + 3 maps. (Univ. Microfilm No. 2421) (Diss. Abstr. 11: 312-314).

Montgomery, F. H. 1957. The introduced plants of Ontario growing outside of cultivation
(Part II). Trans. Royal Canad. Inst. 32: 1-34.

Moseley, E. L. 1899. Sandusky Flora. A catalogue of the flowering plants and ferns growing
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Pieters, A. J. 1894. The plants of Lake St. Clair. Bull. Mich. Fish Comm. 2: 1-12 + map.

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distribution. Bull. U. S. Fish Comm. 21: 57-79.

Pinkava, Donald J. 1963. Vascular flora of the Miller Blue Hole and Stream, Sandusky
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Quebec. Nat. Canad. 95: 49-169.

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- . 1932. Revised Catalog of Ohio Vascular Plants. Bull. Ohio Biol. Surv. 5:

87-215.

Stuckey, Ronald L. 1968a. Aquatic flowering plants new to the Erie Islands. Ohio Jour. Sci.
68: 180-187.

- . 1968b. Distributional history of Butomus umbellatus (flowering-rush) in the

western Lake Erie and Lake St. Clair region. Mich. Bot. 7: 134-142.

Voss, Edward G. 1957. New records of vascular plants from the Douglas Lake region
(Emmet and Cheboygan Counties), Michigan. Pap. Mich. Acad. 42: 3-34.

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Univ. Biol. Ser. 1: 269-302.

Weishaupt, Clara G. 1960. Vascular Plants of Ohio. A Manual for Use in Field and
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Wheeler, Charles F., & Erwin F. Smith. 1881. Michigan Flora. Ann. Rep. State Hort. Soc.
Mich. (1880) 10: 427-529. (Reprinted with title page & repaged 1-105, as Catalogue
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Wood, Duke Willard. 1966. Vascular plant flora of Haunck’s Pond, Middle Bass Island,
Ottawa County, Ohio. M. Sci. Thesis. Ohio State Univ. 59 pp.

Zenkert, Charles A. 1934. The Flora of the Niagara Frontier Region. Buffalo Soc. Nat. Sci.
Bull. 16. 328 pp.

MOSSES FROM SOUTHERN MICHIGAN:

NEW DISTRIBUTIONAL RECORDS1

Paul L. Redfearn, Jr., and Richard L. Halbert^

Life Sciences Department,

Southwest Missouri State College, Springfield, and
Department of Botany and Plant Pathology,

Michigan State University, East Lansing

During the summer of 1967, a course in Bryology was offered by the W.
K. Kellogg Biological Station of Michigan State University. As a result of field
studies connected with this course and our own independent studies, two
species new to the state, Schwetschkeopsis denticulata and Sematophyllum
carolinianum, and 246 new county records were established that were not
listed in Darlington’s recent Mosses of Michigan (1964). These new distribu¬
tional records are cited in the following list. Voucher specimens are deposited
in the herbaria of the W. K. Kellogg Biological Station and/or Southwest

^Contribution No. 165 from the W. K. Kellogg Biological Station of Michigan State
University.

Present address: Department of Botany, University of British Columbia, Vancouver.

1969

THE MICHIGAN BOTANIST

121

Missouri State College. Duplicate collections of the Sphagnaceae and new state
records have also been deposited in the Herbarium of the University of Michi¬
gan. Numbers in parentheses are the collection numbers of the senior author
unless otherwise indicated. Details of the localities cited in this list are as
follows:

Barry County:

Glass Creek. Hardwood forest near junction to tributary of Glass
Creek and Pine Lake Road. NE 14 sect. 7, T. 2 N., R. 9 W.

Otis Lake. East edge of Otis Lake. NE lA sect. 31, T. 3N, R. 9 W.

Sager Road. Hardwood forest near junction of Sager and Shaw
Roads. NE sect. 1, T. 2 N, R. 10 W.

Purdy Lake. Bog and adjacent hardwood forest. NW lA sect. 36, T.
1 N, R. 9 W.

Lawrence Lake. Alkaline Bog. Sect. 27, T. 1 N, R. 9 W.

Hardwood Swamp. East of Hwy. 37, ca. 4 mi. south of Dowling,
SE 14 sect. 16, T. 1 N, R. 8 W.

Berrien County:

Berrien Springs. Steep slopes along small spring-fed gullies. Ca. 1.5
miles NE of Berrien Springs. Sect. 8, T. 6 S., R. 17 W.

Warren Woods. Warren Woods State Park. Sect. 27, T. 7 S., R. 20
W.

Eaton County:

Grand Ledge. Wooded slopes and sandstone bluffs. Fitzgerald Park.
SW !4 sect. 2, T. 4 N, R. 4 W.

Ingham County:

Wooded Deciduous Swamp. Sect. 33, T. 2 N., R. 1 E.

Sanford Woodlot. Michigan State University. East Lansing.

Kalamazoo County:

Kellogg Biological Station. Sect. 6, T. 1 S., R. 9 W.

Kellogg Bird Sanctuary. Sect. 8, T. 1 S., R. 9 W.

Kellogg Forest. Spruce forest and hardwood-pine forest. Sect. 22,
T. 1 S., R. 9 W.

Larix Bog. Ca. 2 miles south of Portage. Sect. 28, T. 3 S., R. 11
W.

Larix-hardwood Bog. Near Gull Lake. Sect. 20, T. 1 S., R. 9 W.
Mason County:

Hardwood Swamp. Ca. 3 miles north of Ludington. Sect. 33, T.
19 N., R. 18 W.

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Ludington State Park. Beech forest and hardwood swampy areas.
Sect. 17, T. 19 N., R. 18 W.

Sand Dunes. Ca. 3 miles north of Ludington. Sect. 33, T. 19 N., R.
18 W.

Shiawassee County:

Rose Lake Wildlife Research Station. Llood plain forest. Sect. 28,
T. 5 N., R. 1 E.

SPHAGNACEAE

Sphagnum capillaceum (Weiss) Schrank var. tenellum (Schimp.) Andr.
Barry Co.: Purdy Lake (22226). Kalamazoo Co.: Larix Bog (21942, 22072,
22075, det. by H. Crum). Mason Co.: Ludington State Park (22011, 22038).

Sphagnum cuspidatum Ehrh. ex Hoffm. Kalamazoo Co.: Larix Bog
(21946, det. by H. Crum).

Sphagnum fimbriatum Wils. ex Hook. f. Barry Co.: Otis Lake (22101,
det. by H. Crum).

Sphagnum girgensohnii Russ. Kalamazoo Co.: Hardwood-Larix Bog
(22085).

Sphagnum papillosum Lindb. Barry Co.: Purdy Lake (22224). Kalama¬
zoo Co.: Larix Bog (21943).

Sphagnum squarrosum Sw. ex Crome. Mason Co.: Hardwood Swamp
(22068, det. by H. Crum).

Sphagnum subsecundum Nees ex Sturm. Mason Co.: Hardwood Swamp
(22050).

TETRAPHIDACEAE

Tetraphis pellucida Hedw. On humus and logs. Barry Co.: Hardwood
Swamp (22242). Mason Co.: Ludington State Park (22004). Shiawassee Co.:
Rose Lake (Halbert 1234).

FISSIDENTACEAE

Fissidens adianthoides Hedw. On humus and decaying logs. Barry Co.:
Lawrence Lake (22215). Mason Co.: Ludington State Park (21989).

Fissidens bryoides Hedw. On soil. Kalamazoo Co.: Kellogg Forest
(21898).

Fissidens cristatus Hedw. ex Mitt. On humus and decaying logs. Barry
Co.: Glass Creek (22127). Mason Co.: Hardwood Swamp (22067).

Fissidens osmundioides Hedw. On log. Mason Co.: Hardwood Swamp
(22046).

Fissidens taxifolius Hedw. On soil. Barrien Co.: Berrien Springs (22305).
Eaton Co.: Grand Ledge (22390).

1969

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123

DITRICHACEAE

Ceratodon purpureus (Hedw.) Brid. On sandy soil and bases of trees.
Barry Co.: Sager Road (22096). Kalamazoo Co.: Kellogg Biological Station
(22188). Mason Co.: Sand Dunes (22042).

Ditrichum pallidum (Hedw.) Hampe. On soil and bases of trees. Barry
Co.: Glass Creek (22131). Kalamazoo Co.: Kellogg Bird Sanctuary (21953).

DICRANACEAE

Dicranella heteromalla (Hedw.) Schimp. On soil. Barry Co.: Purdy Lake
(22236). Shiawassee Co.: Rose Lake (Halbert 784).

Dicranum bonjeanii De Not. ex Lisa. On humus. Mason Co.: Ludington
State Park (22022).

Dicranum flagellare Hedw. On humus and decaying wood. Berrien Co.:
Berrien Springs (22300). Mason Co.: Ludington State Park (21991).

Dicranum fulvum Hook. On vertical sandstone. Eaton Co.: Grand Ledge
(22379).

Dicranum montanum Hedw. On humus and tree bases. Barry Co.: Glass
Creek (22155). Berrien Co.: Warren Woods State Park (22263). Mason Co.:
Ludington State Park (21992).

Dicranum polysetum Sw. On humus. Mason Co.: Ludington State Park
(22027), Hardwood Swamp (22055).

Dicranum scoparium Hedw. On humus and soil. Barry Co.: Otis Lake
(22100), Glass Creek (22150). Berrien Co.: Berrien Springs (22303). Kalama¬
zoo Co.: Kellogg Lorest (21913). Mason Co.: Ludington State Park (22009).
Shiawassee Co.: Rose Lake (Halbert 795).

Dicranum viride (Sulk & Lesq. ex Sulk) Lindb. On tree trunks. Barry
Co.: Glass Creek (22156). Berrien Co.: Warren Woods State Park (22271).

LEUCOBRYACEAE

Leucobryum glaucum (Hedw.) Angstr. ex Fr. On soil, humus, and decay¬
ing logs. Barry Co.: Purdy Lake (22227). Berrien Co.: Berrien Springs
(22321). Kalamazoo Co.: Hardwood-Larix Bog (22079). Mason Co.: Luding¬
ton State Park (21990).

POTTIACEAE

Barbula unguiculata Hedw. On soil. Berrien Co.: Berrien Springs
(22298). Kalamazoo Co.: Kellogg Biological Station (22333).

Desmatodon obtusifolius (Schwaegr.) Schimp. On stone walk Kalamazoo
Co.: Kellogg Biological Station (22194).

Gymnostomum recurvirostrum Hedw. On creek bank. Berrien Co.: Ber¬
rien Springs (Halbert 628).

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THE MICHIGAN BOTANIST

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Phascum cuspidatum Hedw. var. americanum Ren. & Card, ex Card. &
The'r. On old field soil. Kalamazoo Co.: Kellogg Biological Station (Halbert
1-13).

Tortula mucronifolia Schwaegr. On stone wall. Kalamazoo Co.: Kellogg
Biological Station (22221).

Weissia controversa Hedw. On soil. Barry Co.: Glass Creek (22153).
Kalamazoo Co.: Kellogg Biological Station (Halbert 1-13).

GRIMMIACEAE

Grimmia apocarpa Hedw. var. alpicola (Hedw.) Hartm. On stone walls.
Eaton Co.: Grand Ledge (22380). Kalamazoo Co.: Kellogg Biological Station
(22193).

FUNARIACEAE

Funaria hygrometrica Hedw. On soil. Berrien Co.: Berrien Springs
(22297). Mason Co.: Sand Dunes (22044).

Physcomitrium pyriforme (Hedw.) De Not. On old field. Kalamazoo
Co.: Kellogg Biological Station (Halbert 1-26).

BRYACEAE

Bryum angnstirete Kindb. ex Mac. On sandstone ledges. Eaton Co.:
Grand Ledge (22350, 22362).

Bryum argenteum Hedw. On soil. Berrien Co.: Berrien Springs (22316).
Kalamazoo Co.: Kellogg Biological Station (Halbert 8). Mason Co.: Sand
Dunes (22043).

Bryum caespiticium Hedw. On rotting stumps. Shiawassee Co.: Rose
Lake (Halbert 94).

Bryum capillare Hedw. On tree bases. Barry Co.: Otis Lake (22103).
Mason Co.: Ludington State Park (22035).

Bryum pseudotriquetrum (Hedw.) Gaertn., Meyer, & Scherb. On logs.
Barry Co.: Lawrence Lake (22220). Berrien Co.: Berrien Springs (22302).
Kalamazoo Co.: Kellogg Biological Station (Halbert 415). Mason Co.: Hard¬
wood Swamp (22062).

Leptobryum pyriforme (Hedw.) Wils. On vertical sandstone. Eaton Co.:
Grand Ledge (22361).

Pohlia nutans (Hedw.) Lindb. On soil. Berrien Co.: Berrien Springs
(22326). Kalamazoo Co.: Kellogg Forest (Halbert 194). Mason Co.: Ludington
State Park (22013). Shiawassee Co.: Rose Lake (Halbert 629).

Rhodobryum roseum (Hedw.) Limpr. On soil banks. Berrien Co.: Ber¬
rien Springs (22295).

1969

THE MICHIGAN BOTANIST

125

MNIACEAE

Mnium affine Funck. On soil. Barry Co.: Glass Creek (22120, 22124).
Berrien Co.: Berrien Springs (Halbert 594). Kalamazoo Co.: Kellogg Forest
(21888). Eaton Co.: Grand Ledge (22389). Mason Co.: Hardwood Swamp
(22061).

Mnium cuspidatum Hedw. On soil and decaying logs. Barry Co.: Glass
Creek (22139). Berrien Co.: Warren Woods (22265). Kalamazoo Co.: Kellogg
Bird Sanctuary (21958). Shiawassee Co.: Rose Lake (Halbert 1233).

Mnium marginatum (With.) Brid. ex P. Beauv. On soil and sandstone.
Barry Co.: Glass Creek (22162, 22174). Eaton Co.: Grand Ledge (22377).

Mnium medium B. S. G. On soil. Berrien Co.: Berrien Springs (22328).

Mnium punctatum Hedw. On humus and logs. Berrien Co.: Warren
Woods (22288). Eaton Co.: Grand Ledge (22392). Mason Co.: Ludington
State Park (22019).

Mnium punctatum var. elatum Schimp. Base of shrub. Mason Co.: Hard¬
wood Swamp (22057).

Mnium stellar e Hedw. On humus and decaying logs. Barry Co.: Glass
Creek (22154). Berrien Co.: Berrien Springs (22309, 22319).

AULACOMNIACEAE

Aulacomnium androgynum (Hedw.) Schwaegr. On humus. Mason Co.:
Ludington State Park (22017).

Aulacomnium heterostichum (Hedw.) B. S. G. On soil. Barry Co.: Glass
Creek (22134). Berrien Co.: Berrien Springs (22304). Kalamazoo Co.: Kellogg
Forest (21911).

Aulacomnium palustre (Hedw.) Schwaegr. On humus and decaying
wood. Barry Co.: Otis Lake (Halbert 1228). Berrien Co.: Berrien Springs
(22320). Kalamazoo Co.: Larix Bog (22069). Mason Co.: Ludington State
Park (22023). Shiawassee Co.: Rose Lake (Halbert 781).

BARTRAMIACEAE

Bartramia pomiformis Hedw. On soil. Barry Co.: Glass Creek (22136).
Berrien Co.: Berrien Springs (22332).

Philonotis fontana (Hedw.) Brid. var. caespitosa (Jur.) Dix. On log.
Barry Co.: Lawrence Lake (22213, duplicate det. by H. Crum).

TIMMIACEAE

Timmia megapolitana Hedw. On soil. Barry Co.: Glass Creek (22169).

ORTHOTRICHACEAE

Orthotrichum ohioense Sull. & Lesq. ex Aust. On hardwood trees. Barry
Co.: Glass Creek (22165).

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Vol. 8

Orthotrichum pumilum Sw. On hardwood tree. Ingham Co.: Sanford
Woodlot (Halbert 8a).

CLIMACIACEAE

Climacium americanum Brid. On soil, edge of creek. Berrien Co.: Berrien
Springs (22312). Shiawassee Co.: Rose Lake (Halbert 1230).

Climacium dendroides (Hedw.) Web. & Mohr. On log. Mason Co.: Hard¬
wood Swamp (22052).

THELIACEAE

Myurella sibirica (C. Mull.) Reim. On vertical, shaded sandstone. Eaton
Co.: Grand Ledge (22373).

Thelia asprella Sull. Base of oak. Barry Co.: Otis Lake (S. N. Stephen¬
son, Aug. 1967).

FABRONIACEAE

Schwetschkeopsis denticulata (Sull.) Broth. On base of maple. Berrien Co.:
Warren Woods (22276).

LESKEACEAE

Leskea gracilescens Hedw. On tree trunks and logs. Ingham Co.: Sanford
Woodlot (Halbert 20). Kalamazoo Co.: Kellogg Forest (21932).

Leskea obscura Hedw. On base of tree. Berrien Co.: Warren Woods
(22286).

Lindbergia brachyptera (Mitt.) Kindb. On tree trunk. Ingham Co. San¬
ford Woodlot (Halbert 8a).

THUIDIACEAE

Anomodon attenuatus (Hedw.) Hub. On soil, humus, rocks, and tree
bases. Barry Co.: Otis Lake (22110). Berrien Co.: Warren Woods (22270).
Eaton Co. Grand Ledge (22352). Ingham Co.: Sanford Woodlot (Halbert 13).
Kalamazoo Co.: Hardwood-Larix Box (22082). Mason Co.: Hardwood Swamp
(22047).

Anomodon rostratus (Hedw.) Schimp. On humus. Barry Co.: Glass
Creek (22140). Shiawassee Co.: Rose Lake (Halbert 789).

Haplocladium microphyllum (Hedw.) Broth. On log. Berrien Co.: Warren
Woods (22264).

Thuidium delicatulum (Hedw.) B. S. G. On soil, humus, and decaying
logs. Barry Co.: Hardwood Swamp (22248). Berrien Co.: Warren Woods
(22268). Kalamazoo Co.: Kellogg Forest (21900). Mason Co.: Ludington State
Park (22016). Shiawassee Co.: Rose Lake (Halbert 1235).

Thuidium minutulum (Hedw.) B. S. G. On decaying log. Shiawassee Co.:
Rose Lake (Halbert 79).

1969

THE MICHIGAN BOTANIST

127

AMBLYSTEGIACEAE

Amblystegium compactum (C. Mull .) Aust. On sandstone. Eaton Co.:
Grand Ledge (22354).

Amblystegium juratzkanum Schimp. On logs. Barry Co.: Glass Creek
(22138). Mason Co.: Ludington State Park (22031).

Amblystegium serpens (Hedw.) B. S. G. On rocks and trees, Barry Co.:
Purdy Lake (22232). Berrien Co.: Berrien Springs (22324). Ingham Co.: San¬
ford Woodlot (Halbert 7). Kalamazoo Co.: Kellogg Biological Station
(22189).

Amblystegium varium (Hedw.) Lindb. On tree bases and logs. Berrien
Co.: Berrien Springs (22290). Mason Co.: Hardwood Swamp (22065). Shia¬
wassee Co.: Rose Lake (Halbert 51).

Calliergon cor difolium (Hedw.) Lindb. On log. Mason Co.: Hardwood
Swamp (22066).

Calliergonella cuspidata (Hedw.) Loeske. On humus and logs in wet
places. Barry Co.: Lawrence Lake (22218). Mason Co.: Hardwood Swamp
(22054).

Calliergonella schreberi (Brid.) Grout. On soil and humus. Kalamazoo
Co.: Larix Bog (22071).

Campy Hum chry sophy llum (Brid.) J. Lange. On soil. Kalamazoo Co.:
Kellogg Biological Station (22199).

Campy lium hispidulum (Brid.) Mitt. On soil and decaying wood. Barry
Co.: Glass Creek (22122, 22151). Berrien Co.: Berrien Springs (22317).

Campulium stellatum (B. S. G.) C. Jens. On moist soil. Barry Co.:
Lawrence Lake (22214).

Cratoneuron filicinum (Hedw.) Spruce. On humus and soil in wet places.
Barry Co.: Glass Creek (22163). Berrien Co.: Berrien Springs (22310). Kala¬
mazoo Co.: Kellogg Biological Station (22191).

Drepanocladus aduncus (Hedw.) Warnst. var. poly carpus (Bland, ex Voit)
Roth. f. gracilescens (B. S. G.) Moenk. On wet humus. Barry Co.: Glass Creek
(22160, dup. examined by H. Crum).

Drepanocladus uncinatus (Hedw.) Warnst. On wet soil. Kalamazoo Co.:
Kellogg Lorest (21896, dup. det. by H. Crum).

Hygro amblystegium fluviatile (Hedw.) Loeske. On sandstone in creek.
Eaton Co.: Grand Ledge (22383).

Hygroamblystegium tenax (Hedw.) Jenn. On logs in wet places. Barry
Co.: Glass Creek (22173). Berrien Co.: Warren Woods (22266). Kalamazoo
Co.: Kellogg Biological Station (22196).

Leptodictyum kochii (B. S. G.) Warnst. On log. Barry Co.: Sager Road
(22098).

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Vol. 8

Leptodictyum riparium (Hedw.) Warnst. On humus, wood, and rocks in
wet places. Barry Co.: Glass Creek (22142). Berrien Co.: Berrien Springs
(22331).

Leptodictyum trichopodium (Schultz) Warnst. On humus. Barry Co.:
Lawrence Lake (22217). Kalamazoo Co.: Kellogg Biological Station (Halbert
416).

BRACHYTHECIACEAE

Brachythecium acuminatum (Hedw.) Rau & Herv. Base of tree. Berrien
Co.: Warren Woods (22284, examined by H. Robinson).

Brachythecium campestre (Hedw.) B. S. G. On soil. Kalamazoo Co.:
Kellogg Forest (21903, examined by H. Robinson).

Brachythecium oxycladon (Brid.) Jaeg. & Sauerb. On soil and sandstone.
Berrien Co.: Berrien Springs (22318, dup. examined by H. Crum). Eaton Co.:
Grand Ledge (22351). Kalamazoo Co.: Kellogg Biological Station (22195, ex¬
amined by H. Robinson).

Brachythecium plumosum (Hedw.) B. S. G. On soil. Kalamazoo Co.:
Kellogg Forest (21901, examined by H. Robinson).

Brachythecium rivulare (Hedw.) B. S. G. Soil and humus in wet places.
Barry Co.: Glass Creek (22118, examined by H. Robinson). Berrien Co.: Ber¬
rien Springs (22311, examined by H. Robinson). Kalamazoo Co.: Kellogg
Forest (21894, examined by H. Robinson). Shiawassee Co.: Rose Lake (Hal¬
bert 777).

Brachythecium rutabulum (Hedw.) B. S. G. On soil and decaying logs.
Berrien Co.: Warren Woods (22293, examined by H. Robinson). Kalamazoo
Co.: Kellogg Forest (21935, examined by H. Robinson).

Brachythecium salebrosum (Web. and Mohr) B. S. G. On soil and logs.
Barry Co.: Glass Creek (22147, examined by H. Robinson). Shiawassee Co.:
Rose Lake (Halbert 1229).

Brachythecium velutinum (Hedw.) B. S. G. On soil. Barry Co.: Otis Lake
(22106, det. by H. Robinson). Kalamazoo Co.: Kellogg Forest (21914, ex¬
amined by H. Robinson).

Bryhnia graminicolor (Brid.) Grout. On soil. Barry Co.: Glass Creek
(22175).

Bryhnia novae-angliae (Sull. & Lesq. ex Sull.) Grout. On soil. Mason
Co.: Ludington State Park (22001).

Eurhynchium hians (Hedw.) Sande Lac. On log. Berrien Co.: Berrien
Springs (22301).

Eurhynchium pulchellum (Hedw.) Jenn. On soil. Berrien Co.: Berrien
Springs (22322). Kalamazoo Co.: Kellogg Forest (Halbert 208).

Eurhynchium pulchellum var. praecox (Hedw.) Dix. On soil. Kalamazoo
Co.: Kellogg Forest (21905, 21924).

1969

THE MICHIGAN BOTANIST

129

Rhynchostegium serrulatum (Hedw.) Jaeg. and Sauerb. On soil and de¬
caying logs. Barry Co.: Hardwood Swamp (22243). Berrien Co.: Warren
Woods (22287). Kalamazoo Co.: Kellogg Forest (21931).

ENTODONTACEAE

Entodon cladorrhizans (Hedw.) C. Mull. On rocks and decaying wood.
Barry Co.: Glass Creek (22133, 22168). Shiawassee Co.: Rose Lake (Halbert
1231).

Entodon seductrix (Hedw.) C. Mull. On logs and rocks. Barry Co.: Glass
Creek (22144). Berrien Co.: Warren Woods (22262). Eaton Co.: Grand Ledge
(22386). Shiawassee Co.: Rose Lake (Halbert 793).

PLAGIOTHECIACEAE

Isopterygium deplanatum (Sull.) Grout. On log. Berrien Co.: Berrien
Springs (22307).

Plagiothecium denticulatum (Hedw.) B. S. G. On moist soil and logs.
Barry Co.: Otis Lake (22104). Kalamazoo Co.: Kellogg Bird Sanctuary
(21961). Mason Co.: Ludington State Park (22026). Shiawassee Co.: Rose
Lake (Halbert 66).

Plagiothecium laetum B. S. G. On decaying wood and soil. Barry Co.:
Glass Creek (22159). Kalamazoo Co.: Kellogg Bird Sanctuary (21955). Mason
Co.: Ludington State Park (22007).

Plagiothecium roeseanum B. S. G. On soil. Barry Co.: Glass Creek
(22145). Berrien Co.: Warren Woods (22289, 22308). Shiawassee Co.: Rose
Lake (Halbert 794).

SEMATOPHYLLACEAE

Brotherella recurvans (Michx.) Fleisch. On humus and tree bases. Barry
Co.: Otis Lake (22114). Mason Co.: Ludington State Park (22037).

Heterophyllium haldanianum (Grev.) Kindb. On soil, humus, and logs.
Barry Co.: Otis Lake (22105). Berrien Co.: Warren Woods (22275). Eaton
Co.: Grand Ledge (22357). Ingham Co.: Wooded deciduous swamp (Halbert
10). Kalamazoo Co.: Kellogg Forest (21908). Mason Co.: Ludington State
Park (22008).

Sematophyllum carolinianum (C. Mull.) Britt. On sandstone. Eaton Co.:
Grand Ledge (22393).

HYPNACEAE

Ctenidium molluscum (Schimp.) Mitt. On soil. Berrien Co.: Berrien
Springs (22314).

Homomallium adnatum (Hedw.) Broth. On sandstone. Barry Co.: Glass
Creek (22158). Kalamazoo Co.: Kellogg Forest (Halbert 206).

Hypnum curvifolium Hedw. On rocks and logs. Barry Co.: Glass Creek
(22148). Shiawassee Co.: Rose Lake (Halbert 776).

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THE MICHIGAN BOTANIST

Vol. 8

Hypnum imponens Hedw. On log. Mason Co.: Ludington State Park
(22024). Shiawassee Co.: Rose Lake (Halbert 9).

Hypnum lindbergii Mitt. On soil and logs in wet places. Barry Co.: Glass
Creek (22125). Berrien Co.: Warren Woods (22283). Kalamazoo Co.: Kellogg
Forest (21895). Mason Co.: Ludington State Park (22032).

Hypnum pallescens (Hedw.) P. Beauv. On rocks and logs. Barry Co.:
Otis Lake (22115). Kalamazoo Co.: Kellogg Forest (21926). Mason Co.: Lud¬
ington State Park (22033).

Hypnum pratense Koch ex Spruce. On humus. Barry Co.: Hardwood
Swamp (22249). Shiawassee Co.: Rose Lake (Halbert 1232).

Platygyrium repens (Brid.) B. S. G. On tree trunks, logs, and rocks.
Barry Co.: Sager Road (22097). Berrien Co.: Warren Woods (22285). Eaton
Co.: Grand Ledge (22388). Kalamazoo Co.: Kellogg Biological Station
(22200). Shiawassee Co.: Rose Lake (Halbert 4).

HYLOCOMIACEAE

Rhytidiadelphus triquetrus (Hedw.) Warnst. On soil. Kalamazoo Co.:
Kellogg Forest (21910).

BUXBAUMIACEAE

Diphyscium foliosum (Hedw.) Mohr. On soil. Barry Co.: Glass Creek
(22164).

POLYTRICHACEAE

A trichum angustatum (Brid.) B. S. G. On soil. Barry Co.: Glass Creek
(22146). Berrien Co.: Warren Woods (22272). Kalamazoo Co.: Kellogg Forest
(21899).

A trichum undulatum (Hedw.) P. Beauv. On moist soil. Berrien Co.: Ber¬
rien Springs (22296). Kalamazoo Co.: Kellogg Forest (21897).

Poly trichum commune Hedw. On soil and humus. Kalamazoo Co.: Kel¬
logg Forest (21912). Mason Co.: Ludington State Park (22036).

Polytrichum juniperinum Hedw. var. alpestre B. S. G. On humus. Barry
Co.: Purdy Lake (22229). Kalamazoo Co.: Larix Bog (21945).

Poly trichum ohioense Ren. & Card. On soil. Berrien Co.: Warren Woods
(22280). Mason Co.: Ludington State Park (22015).

Poly trichum piliferum Hedw. On sandy soil. Kalamazoo Co.: Larix Bog
(21947). Mason Co.: Ludington State Park (22014).

ACKNOWLEDGMENTS

We wish to thank Dr. Howard A. Crum and Dr. Harold Robinson for their assistance
in the identification of a number of the taxa cited above.

LITERATURE CITED

Darlington, Henry T. 1964. The Mosses of Michigan. Cranbrook Inst. Sci. Bull. 47. 212 pp.
+ 147 figs.

1969

THE MICHIGAN BOTANIST

131

SPECIES OF VASCULAR PLANTS OF PENNFIELD BOG,
CALHOUN COUNTY, MICHIGAN

Garrett E. Crow

Department of Botany and Plant Pathology,

Michigan State University, East Lapsing

Recently a description and ecological analysis (Crow, 1969a) and a
phytogeographical treatment (Crow, 1969b) of Pennfield Bog were presented.
A listing of the species of vascular plants of the bog is now given to complete
the documentation.

A total of 144 species of vascular plants were collected in the bog.
Collections were made during the period of mid- April to mid-October, 1967.
Voucher specimens are deposited in the Michigan State University Herbarium.
Collection numbers are those of Crow.

The families are named and delimited after Gleason and Cronquist
(1963) for pteridophytes and gymnosperms and Takhtajan (1966) for angio-
sperms. They are listed here in alphabetical order.

The occurrence of each species in the vegetation zones of the bog is
indicated by a code in parentheses following the collection numbers. The zone
in which the species is most frequent is listed first. The zones are those
presented in the ecological analysis and vegetation map (Crow, 1969a).

D Decodon zone

C-V Carex-Vaccinium macrocarpon zone

C Cassandra zone

C-T Cassandra-Thelypteris zone

oL open Larix subzone

L Larix zone

L-S Larix-Salix subzone

A Acer mbrum zone

A* outer margin of Acer rubrum zone

V Vaccinium corymbosum subzone

N-E Nuphar-Eleocharis zone

oW open water of lake

LIST OF SPECIES

Aceraceae

Acer rubmm L. 157. (A, L, C-V).

Alismataceae

Alisma plantago-aquatica var. americanum Roem. & Schult. 320, 400,
420. (A*, openings of L).

Sagittaria latifolia Willd. var. latifolia 136, 325, 347, 511. (oL, C-V,
L-S).

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THE MICHIGAN BOTANIST

Vol. 8

Anacardiaceae

Toxicodendron vernix (L.) Ktze. 132, 515. (Throughout the bog).
Apiaceae

Hydrocotyle umbellata L. 191, 350, 507. (Infrequent, on slightly sub¬
merged edges of floating islands associated with D).

Slum suave Walt. 382, 497, 507. (D, C-V, C-T, A*).

Aquifoliaceae

Ilex verticillata (L.) Gray var. verticillata 169, 408. (A).

Nemopanthus mucronata (L.) Trel. 60, 170, 276, 335. (A, L).

Araceae

Calla palustris L. 137, 156, 237, 516. (L, A).

Araliaceae

Aralia nudicaulis L. 483. (A*).

Asclepiadaceae

Asclepias incarnata L. subsp. incamata 203, 523. (D, C-V, C).

Asteraceae

Aster junciformis Rydb. 517. (C-V).

A. umbellatus Mill. 410. (A*).

Bidens cemua L. 513, 518, 555. (A*, light shade of L-S).

B. coronata (L.) Britt. 197. 423, 426, 493, 558. (A*, C-V).

Eupatorium perfoliatum L. var. perfoliatum 389, 409, 536. (C-V, A*).
E. purpureum L. 388, 534. (C-T, C-V).

Balsaminaceae

Impatiens biflora Walt. 387, 399. (A*).

Betulaceae

Betula lutea Michx. f. 77, 90, (A).

B. pumila var. glabra Regel 75, 290. (L-S).

Corylus americana Walt. 542. (A, one station).

Campanulaceae

Campanula aparinoides Pursh var. aparinoides 201, 280. (C-V, D, C-T).
Caprifoliaceae

Sambucus canadensis L. 234. (A*).

Viburnum lent ago L. 492. (A*).

Ceratophyllaceae

Ceratophyllum demersum L. 428. (oW).

Cornaceae

Cornus stolonifera Michx. 72, 192, 288, 345. (L-S, A).

Cucurbitaceae

Echinocystis lobata (Michx.) T. & G. 418. (A*).

Cyperaceae

Carex aquatilis Wahl. 84, 194, 286, 293. (C-V, L-S).

C. canescens var. disjuncta Pern. 152, 195. (C-V, L-S).

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133

C. chordorrhiza L. f. 158, 161. (D).

C. comosa Boott 143, 274, 327, 521. (D, N-E).

C. interior Bailey 159, 221, 369. (L, C-V).

C. lasiocarpa var. americana Fern. 142, 223, 227. (C-V, C-T).

C. trisperma Dew. 226, 367. (L, A).

Cyperus diandrus Torr. 398, 508. (D, one station).

C. engelmanni Steud. 393, 505. (C-V).

Dulichium arundinaceum (L.) Britt. 224, 381. (N-E, C-V, oL).

Eleocharis compressa Sull. 95. (C-V).

E. intermedia (Muhl.) Schultes 396, 509. (D, two stations).

E. smallii Britt. 147, 149, 209, 429. (N-E).

Eriophorum gracile Koch 93, 151, 222. (N-E, C-V).

E. virginicum L. 213, 329, 331. (oL, C).

Rhynchospora alba (L.) Vahl. 331, 535. (C-V, oL).

Scirpus acutus Muhl. 193. (S-L).

S. cyperinus (L.) Kunth 332, 385. (oL).

Droseraceae

Drosera intermedia Hayne 266, 318, 355. (C-V).

D. rotundifolia L. 319, 354. (C, D).

Ericaceae

Andromeda glaucophylla Link. 54, 63, 189, 239. (C-V, oL).

Cassandra calyculata (L.) D. Don 42, 188, 240, 295. (C, C-T, D, L-S,
oL). [Chamaedaphne incorrect name. See Wood (1961).]

Gaultheria procumbens L. 376. (A, one station).

Vaccinium corymbosum L. 69, 92, 153, 363. (L, A, V).

V. macrocarpon Ait. 36, 134, 238. (C-V, oL, C).

Equisetaceae

Equisetum fluviatile L. 51, 199. (L-S, L).

Fabaceae

Apios americana Medic. 365, 531. (L).

Fagaceae

Quercus velutina Lam. 278. (L, A).

Haloragaceae

Myriophyllum verticillatum L. 56, 183, 230, 352, 561. (oW).
Hypericaceae

Hypericum boreale (Britt.) Bickn. 308, 357, 504. (C-V).
Triadenum virginicum (L.) Raf. 307, 390, 406. (C-V, D, L-S).

Iridaceae

Iris versicolor L. 411. (A*).

Juncaceae

Juncus brevicaudatus (Engelm.) Fern. 384, 503. (N-E).

J. effusus var. solutus Fern. & Wieg. 214, 330. (oL).

Juncaginaceae

Triglochin maritima L. 340, 557. (C-V, two stations).

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Lamiaceae

Lycopus unifloms Michx. 341 , 343, 359, 484, 520, 537. (N-E, D, C-V, L).
Scutellaria galericulata L. 133, 202, 281, 404, 539. (C-V, C-T).

S. lateriflora L. 391 , 405, 482. (D, A*).

Lemnaceae

Lemna minor L. 356. (Rare, in pools in C-V).

L. trisulca L. 228. (L, one station).

Lentibulariaceae

Utricularia gibba L. 397, 506. (Infrequent, on slightly submerged edges
of floating islands associated with D).

U. intermedia Hayne 139, 268b. (C-V).

U. purpurea Walt. 233, 269, 525. (oW).

U. vulgaris L. 57, 104, 247, 268a, 533. (oW, N-E).

Lihaceae

Maianthemum canadense Desf. var. canadense 81, 207, 244b, 272. (A).

M. canadense var. interim Fern. 164, 187, 206, 244a, 538. (A).

Lythraceae

Decodon verticillatus (L.) Ell. var. verticillatus 314, 328. (D).
Menyanthaceae

Menyanthes trifoliata L. 71, 163. (oL, C-V, L-S).

Monotropaceae

Monotropa uniflora L. 186, 323, 512. (L).

Nymphaeaceae

Brasenia schreberi Gmel. 184, 527, 559. (oW).

Nuphar luteum subsp. macrophyllum (Small) Beal 140, 528. (C-V).
Nymphaea tuberosa Paine 138. (N-E, oW).

Nyssaceae

Nyssa sylvatica Marsh. 360, 427, 492. (A, L, only two stations noted).
Onagraceae

Epilobium strictum Muhl. 338, 358, 480. (C-V, A*).

Orchidaceae

Cypripedium acaule Ait. 70, 273, 334. (L, oL).

Habenaria ciliaris (L.) R. Br. 312, 530. (oL).

H. clavellata (Michx.) Spreng. 321, 478. (A).

H. obtusata (Banks ex Pursh) Rich. 324. (oL).

Isotria verticillata (Willd.) Raf. 100. (L).

Liparis loeselii (L.) Rich. 166, 348. (C-V, L-S).

Malaxis monophyllos var. brachypoda (Gray) F. Morris 336. (C-V).
Pogonia ophioglossoides (L.) Ker. 141, 337. (C-V).

Osmundaceae

Osmunda cinnamomea L. 146, 235, 368. (A).

O. regalis var. spectabilis (Willd.) Gray 144, 242

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135

Pinaceae

Larix laricina (DuRoi) K. Koch 35, 236. (L, C, C-T, C-V).

Pinus strobus L. 91. (L, A, two large trees, seedlings rare).

Poaceae

Agrostis stolonifera var. major (Gaud.) Farw. 150, 211, 279, 292. (L).
Glyceria canadensis (Michx.) Trin. 210, 419. (L).

G. striata (Lam.) Hitchc. 401. (A*).

Leersia oryzoides (L.) Sw. 501. (A*).

Muhlenbergia racemosa (Michx.) BSP. 342, 414, 522. (C-V).

Polygonaceae

Polygonum hydropiperoides Michx. var. hydropiperoides 313, 353, 383.

(In open water just off edge of Decodon mat).

P. sagittatum L. 514. (A*).

Rumex orbiculatus Gray 519. (L-S, one station).

Polypodiaceae

Onoclea sensibilis L. 386, 488. (A, C-V).

Thelypteris palustris var. pubescens (Laws.) Fern. 339. (C-V, C-T, D, A).
Woodwardia virginica (L.) Smith 271, 315. (oL).

Pontederiaceae

Pontederia cordata L. 145, 168, 241, 407. (N-E, C-V).

Potamogetonaceae

Potamogeton gramineus L. 218, 220, 349, 529. (N-E, C-V).

P. illinoensis Morong 182. (oW).

P. natans L. 181, 185, 246, 526, 560. (oW).

Primulaceae

Lysimachia thyrsiflora L. 101, 554. (D).

Trientalis borealis Raf. 76, 232, 245. (L, A).

Ranunculaceae

Coptis trifolia var. groenlandica (Oeder) Fass. 65, 510. (A).

Ranunculus sceleratus L. 562. (L-S, one station).

Rosaceae

Amelanchier arborea (Michx. f.) Fern. 33, 374. (A).

Aronia melanocarpa (Michx.) Ell. 82, 362, 496. (V, A, L).

Potentilla palustris (L.) Scop. 103, 135, 282. (D, C, oL).

Prunus virginiana L. 88. (A*, one station).

Rosa palustris Marsh. 73, 297. (Margin of L and D, A, L-S).

Rubus hispidus var. obovalis (Michx.) Fern. 167, 208, 277, 364, 417.
(L, A).

R. pubescens Raf. 83. (L-S, one station).

Spiraea alba DuRoi 283, 289, 344, 361 , 489. (C-V, C-T, oL, L-S, A*).

S. tomentosa L. 283, 326, 532. (D, C-V, C-T, L).

Rubiaceae

Cephalanthus occidentalis L. var. occidentals 296, 333, 416. (D).

Galium trifidum L. var. trifidum 231, 242b, 284, 415. (C-V, L-S, A*).
Mitchella repens L. 375. (A, one station).

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Vol. 8

Salicaceae

Populus tremuloides Michx. 79, 285. (C-V, V).

Salix bebbiana Sarg. 52, 291 , 492. (L-S).

S. Candida Fliigge 38, 39, 40, 86. (Margin of C-V and L).

S. pedicellaris var. hypoglauca Fern. 41, 55, 59, 154, 216. (C-V).

S. petiolaris Sm. 37, 370. (C-V, F-S, D).

S. serissima (Bailey) Fern. 87, 89. (C-V, C).

Sarraceniaceae

Sarracenia purpurea F. 64, 131, 366. (C-V, C, oF).

Saxifragaceae

Ribes americanum Mill. 200, 549. (F-S, one station).

R. hirtellum Michx. 541. (F-S, one station).

Scrophulariaceae

Agalinis purpurea (F.) Pennell 329, 486. (C). [Gerardia misapplied. Agalinis
is the conserved name (Fanjouw, et al., 1966, p. 336).]

Solanaceae

Solatium dulcamara F. 180. (C-T, one station).

Sparganiaceae

Sparganium chlorocarpum var. acaule (Beeby) Fern. 229, 380. (N-E).

S. minimum (Hartm.) Fries 351. (oW at edge of D).

Typhaceae

Typha latifolia F. 267. (D, C-T).

Ulmaceae

Ulmus rubra Muhl. 198, 217. (A, F).

Urticaceae

Boehmeria cylindrica (F.) Sw. 205, 287, 346, 540. (C-V, F-S, A).

Urtica dioica var. procera (Muhl.) Wedd. 556. (A*).

Violaceae

Viola af finis Fe Conte 85. (A*, one station).

V. nephrophylla Greene 322, 394, 524. (C-V, C, D).

Vitaceae

Parthenocissus quinquefolia (F.) Planch. 494. (A*).

Vitis vulpina F. 498. (A*, one station).

FITERATURE CITED

Crow, G. E. 1969a. An ecological analysis of a southern Michigan bog. Mich. Bot. 8:
11-27.

Crow, G. E. 1969b. A phytogeographical analysis of a southern Michigan bog. Mich. Bot.
8: 51-60.

Gleason, H. A., & A. Cronquist, 1963. Manual of Vascular Plants of Northeastern United
States and Adjacent Canada. D. Van Nostrand Co., Princeton. 810 pp.

Lanjouw, J., et al. 1966. International Code of Botanical Nomenclature. Regnum Vegeta¬
ble 46. 402 pp.

Takhtajan, A. 1966. Sistema I Filogeniya Tsvetkovjkh Rasteniy. Science Press, Moscow-
Leningrad. 611 pp.

Wood, C. E. 1961. The genera of Ericaceae in the southeastern United States. Jour.
Arnold Arb. 42: 10-80.

1969

THE MICHIGAN BOTANIST

137

THE LOG FERN ( DRYOPTERIS CELS A) AND ITS HYBRIDS
IN MICHIGAN— A PRELIMINARY REPORT

W. H. Wagner, Jr., Florence S. Wagner, & D. J. Hagenah
Botanical Gardens, The University of Michigan, Ann Arbor,
and Cranbrook Institute of Science, Bloomfield Hills, Michigan

BACKGROUND

The woodferns, Dryopteris, are conspicuous plants in the pteridophyte
flora of the eastern United States. Large, robust plants with massive and scaly
rhizomes, many of the woodferns are showy and ideal for cultivation in
shaded gardens. They are notorious for their tendency to cross and form
vigorous hybrids, and indeed woodfern hybrids may become common to
abundant in certain localities. The presence of these plants, more or less
intermediate in appearance between the parent species, has resulted in much
taxonomic confusion. The same entity has been variously interpreted as a
species, as a variety of another species, or as a hybrid.

Fortunately the development of techniques for cytological analysis, and
the duplication of wild hybrids by experimental hybridization, have gradually
brought an understanding of the basic relationships within the genus in the
last two decades. For the benefit of those readers who are not familiar with
this “genome analysis” approach we will briefly review the current knowledge
of the woodferns of eastern North America.

The basic building blocks, as they might be called, are the genomes of
the diploid species, in which there are two duplicate, compatible sets of 41
chromosomes each. For convenience, these basic sets are identified by a kind
of shorthand in which the initial of the specific epithet is used to designate
the genome, as G for goldiana, L for ludoviciana, M for marginalis, etc., with
the letter appearing once for each set of this genome. Thus the giant
woodfern, Dryopteris goldiana, is GG. In a few cases, hybrids of the basic
species have somehow managed to double their chromosome sets, as in D.
celsa, GGLL.

The discovery of natural crosses between these plants with doubled
chromosome complements and other species provides a means for determining
what basic sets or building block genomes are present in such polyploid
species and in what proportions they are present. Thus our continuing search
for natural hybrids of the various woodferns is a search for additional links in
the chain of evidence by which we can analyze the phylogenetic development
of the various species. Of the approximately three dozen distinguishable kinds
of Dryopteris in the eastern United States, in fact, only 12 are fertile, sexual
plants with normal spores and prothallia. All of the rest are sterile, unbalanced
hybrids, with non-matching sets of chromosomes. For example, our common
“Boott’s woodfern,” Dryopteris Xboottii, is the natural hybrid of “crested
fern,” D. cristata, with the cytogenetic formula SSLL, and “fancy fern,” D.
intermedia, with the formula II. The hybrid may be formed from a sperm

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Vol. 8

with the formula SL and an egg with I, so that D. Xboottii will have the
formula SLI. The chromosomes of set S cannot pair with those of L, nor L
with I, nor I with S. The individual chromosomes of one set do not pair
properly in spore formation with those of the other sets, and the
chromosomes are thus distributed with great irregularity and in such
imbalance that all of the spores tend to be abortive and inviable. The plant is
therefore sterile.

On the contrary, the fertile woodferns, as shown in the diagram in
Figure 1 , do have balanced genomes which pair normally and produce living
spores. In D. spinulosa, which has four complements of chromosomes, there
are two sets of I chromosomes and two sets of S. The I chromosomes pair
with each other, and the S chromosomes do likewise, so that all of the spores
receive the IS constitution.

In research on Dryopteris we are thus able to determine relationships to
a large extent by observing how the chromosomes pair at the time of spore
production. We can tell that D. campyloptera, the “mountain woodfern,”
shares a genome with D. intermedia by studying their hybrid. This cross, DDII
X II, produces a triploid DII, in which we find two complements of
chromosomes neatly paired (i.e., two sets of I) and one set completely
unpaired (i.e., D). Since the basic number of chromosomes per set or genome
in Dryopteris is X = 41 , we see 41 pairs and 41 singles in the hybrid of D.
campyloptera and D. intermedia. By combining studies of morphology and
ecology of the plants with observations of their chromosomes, it is possible to
work out an objective scheme of relationships, as shown in Figure 1 .

The summary of fertile kinds of woodferns shown in this figure results
largely from the basic work of Stanley Walker of the Department of Genetics,
University of Liverpool (1955, 1961, 1962), who studied the interrelationships
of eastern American Dryopteris taxa as well as those in Europe. In connection
with our own cytofloristic investigations of ferns, especially in Michigan,
Virginia, and New York, we have supplemented Walker’s work with numerous
additional observations. The summary in Figure 1 shows that D. filix-mas and
D. marginalis are not related to the others, according to chromosome data.
Taxa #3 through #12, however, are connected with each other through shared
genomes. One of the fertile woodferns, which we have designated “D.
semicrista ta is still only a hypothetical plant, not yet discovered. The
postulated plant may be extinct or it may still exist, but it will resemble D.
cristata or the hybrid D. Xuliginosa (= D. cristata X spinulosa) so closely that
it may be overlooked. It probably differs subtly from D. cristata in having
slightly more dentate margins and more divided leaflets.

There are still other problems. For example, we are not certain that D.
campyloptera is really DDII (Wagner, 1963). Our only evidence for the D
genome in this fern is the discovery of D. campy lop tera-like plants in the
Huron Mountain area of Michigan in company with D. intermedia and what
we have called “Lake Superior D. dilatata which we regard as possibly an
undescribed species. The campyloptera- like plants were hybrids of D.
intermedia and Lake Superior dilatata and they were sterile, with little or no

1969

THE MICHIGAN BOTANIST

139

pairing at meiosis. We interpreted their cytogenetic constitution to be Dl,
which, if doubled, would produce the fertile tetraploid DDII (Wagner and
Hagenah, 1962). Widen and Sorsa (1968) have recently contributed new data
from chemical studies of the spreading woodfern of the Lake Superior region.
These show striking differences from the spreading woodfern of the western
United States and Europe,/), assimilis, which is a similar-appearing diploid. In
the words of these authors, “Our results ... fit in with the view of Wagner
and Hagenah (1962) that the ‘western diploid’ and the ‘Lake Superior diploid’
are separate species rather than with the opinion of Britton (1967) that they
are conspecific.”

CLINTON’S WOODLERN AND THE LOG FERN

Two of the swamp-inhabiting taxa, D. clintoniana and D. celsa, are
noted for their close similarities. In general, fortunately, they do not occur
together. Plants from New England to the western Great Lakes area are the
former, known as “Clinton’s woodfern”; and plants from approximately New
Jersey southward are the latter, the “Log Fern,” which was originally

1 .

D. filix-mas

2.

D. marginalis

3.

D. "dilatata"

4.

D. campyloptera .

5.

D. intermedia

6.

D. spinulosa

( 7.

D. semicristata .

8.

D. cristata

9.

D. ludoviciana

10.

D. celsa

1 1 .

D. goldiana

12.

D. clintoniana

p i F 1 p 2p 2

MM

DD

DDII

/

II

IISS

/

SS )

rSSLL

/

LL

I

LLGG

/

GG

\

SSLLGG

Fig. 1. Fertile (sexual) cytomorphotypes in the woodfems, Dryopteris, of eastern United
States, exclusive of D. fragrans. The evidence for the existence of taxon #7 is entirely in¬
ductive; it has not yet been discovered as a living plant.

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THE MICHIGAN BOTANIST

Vol. 8

described from plants growing on logs and at the bases of tree-trunks in
Dismal Swamp, in Coastal Plain Virginia.

Dryopteris clintoniana is one of the most interesting members of the
woodfern complex because it has the highest chromosome number. It is
hexaploid with the genomic constitution SSLLGG, a total of 246
chromosomes. Its hybrids with D. cristata are pentaploid with the formula
SSLLG, thus showing 82 pairs and 41 singles at meiosis. Its hybrids with D.
goldiana are tetraploid, i.e., SLGG, with 41 pairs and 82 univalents at meiosis.
Clinton’s woodfern is very common in appropriate habitats in southern Michi¬
gan. It flourishes in rich, wet, shady swamps, where it is often accompanied
by D. cristata and D. goldiana and readily forms hybrids with them. As a
result of this hybridization we find a gradient series of ferns that run from the
narrow, pale-scaled D. cristata to the broad, black-scaled D. goldiana, i.e.,
SSLL-SSLLG*-SSLLGG-SLGG*-GG. (Sterile hybrids in the series here and be¬
low are marked with an asterisk.) Obviously, considering the natural variability
of the plants involved, such a series can be very confusing to the collector.

We have never expected to find log fern in Michigan. Indeed, in view of
our present difficulties of making identifications of the more critical hybrids
we have always congratulated ourselves that we were so far from its known
range to the south and east. If, in fact, log fern did grow in southern Michigan
in the area of Clinton’s woodfern, we would have the formidable potentiality
of a bewildering and subtle cytomorphological series, i.e.,
SSLL-SSLLG*-SLLG*-SSLLGG-SLLGG*-SLGG*-LLGG-LGG*-GG! And this
does not include all of the D. cristata and D. clintoniana hybrids with D.
marginalis, D. intermedia, and D. spinulosa- hybrids which are already known
from Michigan: MSL*, ISL*, ISSL*, MSLG*, ISLG*, and ISSLG*. These
known hybrids more or less resemble members of the foregoing list in various
respects. If D. celsa should also exist in the state, then three more confusing
hybrids could potentially occur, namely MGL*, IGL*, and ISGL* (Cf. illustra¬
tion in Wagner and Wagner, 1965). Thus if D. celsa were to occur in our area,
the number of possible members of this group would jump from 1 1 to a
disheartening 18.

Dryopteris celsa is very similar to D. clintoniana in gross appearance.
The latter is so like D. cristata that some authors (e.g., Fernald, 1950) have
treated it as a broad-fronded variety, viz. D. cristata var. clintoniana. In gener¬
al D. celsa can be distinguished from D. clintoniana on the following char¬
acters: basal leaflets broader, more ovate-lanceolate (rather than triangular),
midrib slightly scaly (rather than glabrous or nearly so), and scales on the
rhizomes and petiole bases darker and glossier (blackish rather than brown or
tan). Also, the spores of D. celsa average smaller than those of D. clintoniana,
being only 35 to 45 microns in longest diameter rather than between 45 and
50. The chromosomes of D. celsa number 164, but those of D. clintoniana, as
mentioned above, number 246.

The extent of D. celsa northward along the Atlantic coastal region is
still not well established, but there is evidence of its occurrence, at least in the
past, to as far as Staten Island, New York. Plants which have sometimes been

1969

THE MICHIGAN BOTANIST

141

identified as D. clintoniana from eastern Pennsylvania to South Carolina and
Louisiana north to Arkansas and Missouri have all proved to be D. celsa. Log
fern is a rare and sporadic plant, but may be locally abundant in appropriate
habitats. Recently we discovered disjunct populations of it in the vicinity of
Rochester, New York, isolated over 200 miles from the nearest other localities
in the southeast and growing well within the territory of D. clintoniana. With
the single exception of the Rochester occurrences of D. celsa we had absolute¬
ly no reason to anticipate finding it in Michigan.

Readers may well imagine our astonishment, therefore, to discover that
D. celsa and at least a couple of its hybrids do grow in Michigan— over 400 air
miles from the nearest other known localities in Rochester, N. Y., and over
450 miles from its localities in Missouri. One can picture, from what has been
stated above, the multiple problems that this discovery will present, with
respect not only to the origin of the Michigan populations, but also to the
inherent difficulties we expect to face in our future explorations, when we
encounter all of its potential hybrids.

CIRCUMSTANCES OF THE DISCOVERY

On December 1, 1968, W. H. Wagner and D. J. Hagenah explored two
localities in Kalamazoo County, Michigan, each a large swamp dominated by
black ash, Fraxinus nigra, yellow birch, Be tula lutea, and American elm,
Ulmus americana, the latter largely dead or dying from Dutch elm disease.
The first swamp we visited was on the north side of Section 20, along W
Avenue (T4S, R12W— Prairie Ronde Tp.), the second in the northeast quarter
of Section 6 (T4S, R1 1W— Schoolcraft Tp.), near Sugarloaf Lake. The former
of these localities was mentioned earlier by Wagner and Chen (1965) as an
excellent woodfern locality which we strongly recommended to students of
Dryopteris. The second locality, unfortunately, is in the process of being
destroyed. Around the base of the “Island,” a wooded knoll in the middle of
the swamp, the ferns in question were exceedingly abundant, but now the
knoll (as shown on the left in Figure 2) is nearly cleared except for a few
large trees, and the swamp (on the right) is scheduled to be cleared, probably
in spring and summer of 1969. In a short time the whole area will be a
housing development.

On this date, we encountered a large number of luxuriant woodferns
that appeared to be either Dryopteris clintoniana X goldiana (SLGG*) or the
similar D. celsa X goldiana (LGG*). We found plants of this type at both
localities’, and they so suspiciously resembled the plants of D. celsa X goldiana
we had studied from Rochester and from Staten Island, New York, and from
Tennessee, that we took a rather large sample of rhizomes for later cytological
investigation, as well as some plants of different morphology growing near by.

The plants in question were potted and grown in a research greenhouse
at the University of Michigan Botanical Gardens by Donald Trout. By Febru¬
ary, the new fronds had become large enough to begin spore production, at
which time their chromosomes could be studied. F. S. Wagner surveyed the
plants cytologically, discovering that indeed our supposition was correct— the

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plants did conform cytologically to D. celsa X goldiana, with the pairing
behavior of LGG, showing 41 pairs and 41 singles. One of the parents, therefore,
had to be D. celsa. Fortunately among the plants in our sample we had taken one
which superficially resembled D. clintoniana except for its dark scales and some¬
what broader, and less triangular pinnae. This specimen proved to be the predicted
D. celsa, displaying 82 normal pairs at meiosis. Another plant which resembled
D. clintoniana X crista ta in the field proved to be D. celsa X cristata, showing
dark scales and SLLG chromosome pairing behavior, 41 pairs and 82 singles.

Both of these log fern hybrids are remarkably similar to their other
parents, and we feel certain that at least some plants previously identified
from the Kalamazoo area as D. goldiana were in fact D. celsa X goldiana. The
central plant in the profiles of living leaves (Fig. 3) shows the narrow blade

Fig. 2. Winter view of swamp where Dryopteris celsa and its hybrids are abundant. This
locality will soon be destroyed for a housing development. (Photograph by Terry L.
Sharik.)

1969

THE MICHIGAN BOTANIST

143

and short petiole of the hybrid D. celsa X cristata. Note especially the “Vene¬
tian blind” arrangement of the pinnae, those except at the tip twisted into a
plane almost parallel to the ground. Dryopteris celsa X goldiana (profiles to
either side in fig. 3), on the other hand, shows little of this pinna orientation.
Its very broad blades and long petioles are but little different from those of
the giant woodfern, D. goldiana. Both of these hybrids, though sexually ster¬
ile, are extremely attractive and vigorous, and they may give promise horti-
culturally .

Subsequent investigations, still incomplete, in the herbaria of University
of Michigan, Western Michigan University, and Michigan State University, in¬
dicate that there are probably other localities in Kalamazoo County and in
nearby Allegan County. Plants heretofore identified from this area as “D.
clintoniana,” “D. cristata or “D. goldiana ” are likely to be D. celsa or its
hybrids. On the basis of what we know now, we strongly suspect that all of
the hybrids described above will ultimately be found in southwestern

Fig. 3. Profiles of living fronds of log fern hybrids: Dryopteris celsa X goldiana on both
sides. D. celsa X cristata in middle. Note the distinctive pinna orientation, narrow blades,
and short petioles of D. celsa X cristata. (Photograph by James P. Bennett.)

144

THE MICHIGAN BOTANIST

Vol. 8

Michigan. In the following list of potential new taxa in our flora, those which
have already been demonstrated are marked with a dagger:

tDryopteris celsa . LLGG

D. Xleedsii (= D. celsa X marginalis) . MLG*

D. Xseparabilis (= D. celsa X intermedia) . ILG*

D. celsa X spinulosa . ISLG*

fD. celsa X cristata . SLLG*

fD. celsa X goldiana . LGG*

D. celsa X clintoniana . SLLGG*

DISCUSSION

If all of the above woodfern taxa are found in the near future, the total
number of known species and hybrids of pteridophytes in the state of Michi¬
gan will be increased to 122 from Hagenah’s 1968 (unpublished) total of 115.
Three of them were actually collected, as reported here, so the total is
presently 118. For those botanists in Michigan who are appalled at the pros¬
pect of identification of these plants, we hope that as our further investiga¬
tions of the woodfern swamps of Kalamazoo and nearby counties move ahead
we shall finally be able to construct a reliable key to differentiate all of the
taxa discussed here. At the present time no such key exists.

The range extensions represented here are striking. Over four hundred
miles from the nearest known localities to the east and southwest, the dis¬
covery of log ferns in this state is very remarkable. This is the third one of
the pteridophytes now known in Michigan that occur only in the south¬
western portion of the state and which constitute major disjunctions. Many
years ago Liberty Hyde Bailey encountered stands of the narrow chainfern,
Woodwardia areolata, in Van Buren County (cf. Billington, 1952, p. 209).
Richard W. Pippen in 1966 reported that a population of the American climb¬
ing fern, Lygodium palmatum, grows in Kalamazoo County, not far from the
finds described here of the log fern. Is there something distinctive about this
area which permits southern and coastal plain plants to occur at long distances
from their major ranges? (Cf. Hanes, 1942)

In conclusion, we should point out that our “Dryopteris celsa ” could
possibly represent a different cytotype from the typical plants of the south. It
has occurred to us that the northern plants might actually involve a different
genome from true D. celsa, making it possible for it to grow in the main range
of D. cristata, D. spinulosa, and D. clintoniana, all plants of the higher lati¬
tudes. One hypothesis might be that instead of our “D. celsa” being an amphi-
diploid of the southern woodfern, D. ludoviciana (LL) and the giant wood¬
fern, D. goldiana (GG), it could have arisen as the amphidiploid of the un¬
known woodfern, D. semicristata (SS), and giant woodfern. Its formula would
then be SSGG. We must remember, however, that the existence of D. semi¬
cristata is still a purely hypothetical conjecture itself, and as our studies con¬
tinue we may be forced to conclude that there was or is no such plant, past
or present. The postulation of such a plant as D. semicristata is simply the
only means we have at present of explaining the currently available facts.

1969

THE MICHIGAN BOTANIST

145

ACKNOWLEDGMENTS

This research was supported by National Science Foundation Grant GB-8113, the
Evolutionary Characters of Ferns. We wish to acknowledge the aid and advice of Irving
W. Knobloch, E. T. Wherry, Stanley Walker, and Conrad V. Morton.

LITERATURE CITED

Billington, Cecil. 1952. Ferns of Michigan. Cranbrook Inst. Sci. Bull. 32. 240 pp.

Britton, Donald M. 1967. Diploid Dryopteris dilatata from Quebec. Rhodora 69: 1-4.
Femald, M. L. 1950. Gray’s Manual of Botany. 8th ed. American Book Co., New York,
lxiv + 1632 pp.

Hanes, Clarence R. 1942. The Atlantic Coastal Plain element in the flora of Kalamazoo
County, Michigan. Pap. Mich. Acad. 27: 37-43.

Pippen, Richard W. 1966. Lygodium palmatum, the climbing fern, in southwestern Michi¬
gan. Mich. Bot. 5: 64-66.

Wagner, Warren H., Jr. 1963. Pteridophytes of the Mountain Lake area, Giles County,
Virginia, including notes from Whitetop Mountain. Castanea 28: 113-150.

- , & Dale J. Hagenah. 1962. Dryopteris in the Huron Mountain Club area of

Michigan. Brittonia 14: 90-100.

- , & Katherine Lim Chen. 1965. Abortion of spores and sporangia as a tool

in the detection of Dryopteris hybrids. Am. Fern Jour. 55: 9-29.

- , & F. S. Wagner. 1965. Rochester area log ferns (Dryopteris celsa) and their

hybrids. Proc. Rochester Acad. 11: 57-71.

Walker, Stanley. 1955. Cytogenetic studies in the Dryopteris spinulosa complex. I.
Watsonia 3: 193-209.

- r 1961. Cytogenetic studies in the Dryopteris spinulosa complex. II. Am.

Jour. Bot. 48: 607-614.

- . 1962. Further studies in the genus Dryopteris: the origin of D. clintoniana,

D. celsa and related taxa. Am. Jour. Bot. 49: 497-503.

Widen, Carl-Johan, & Veikko Sorsa, 1968. On the intraspecific variability of Dryopteris
assimilis S. Walker and D. spinulosa Watt. Hereditas 62: 1-13.

News of Botanists

Dr. Eric A. Bourdo, Jr., Director of the Ford Forestry Center of Michigan Techno¬
logical University, has been named the first dean of that University’s new School of For¬
estry and Wood Products, which includes the Department of Forestry, the Institute of
Wood Research, and the Ford Forestry Center.

Recent deaths of former University of Michigan botanists include Dr. Edwin B.
Mains (Director of the Herbarium 1931-1960), on December 23, 1968; Dr. Bessie B.
Kanouse (Curator of Fungi in the Herbarium, 1926-1960), on January 28, 1969; and Dr.
Felix G. Gustafson (plant physiologist in the Department of Botany since 1920, retiring
as Professor in 1959), on January 13, 1969.

Dr. Frederick K. Sparrow, appointed Director of the University of Michigan Bio¬
logical Station for the three years until his retirement, was one of two recipients of an¬
nual Merit Awards from the Botanical Society of America at its 1968 annual meeting.

146

THE MICHIGAN BOTANIST

Vol. 8

MARCH ANTIA POLYMORPHA IN NORTHERN MICHIGAN

Ella O. Campbell

Department of Botany, Massey University,

Palmerston North, New Zealand

Certain of the variations of Marchantia polymorpha L. have not been
sufficiently taken into account in northern Michigan or in North America as a
whole. Because of a background of experience with the various expressions of
this species in other parts of the world, I made a special attempt in the
summer of 1968 to study its diversity in the general vicinity of the University
of Michigan Biological Station, in northern Michigan, where colonies re¬
sembling the var. polymorpha and others resembling the var. aquatica Nees are
found.

Marchantia polymorpha is very widely distributed and varies con¬
siderably according to environmental conditions (Burgeff, 1943; Maravolo &
Voth, 1966; Voth, 1941, 1943; Voth & Hamner, 1940). Nees (1838) char¬
acterized the variations known to him according to the left-hand side of the
following outline; for purposes of comparison the subdivision used in this
paper is given on the right-hand side.

M. polymorpha L.

_ var. aquatica Nees

A. communis
a. aquatica

a 1 . nuda a 1 *. brevipedunculata
a 2. denticulata a2*. brevipedunculata
a 3. lobulata
p. riparia

P 1 . dilatata p 1 *. latissima
P 2. angusta

_ var. polymorpha

7 . domestica

7 1 • receptaculis grandiusculis
7 2. receptaculis minutis

_ var. alpestris Nees

B. alpestris
B*. crispa

B 1 3. riparia
B 7. domestica

B 7 1 . macrophylla
B 7 2. microphylla

Muller (1906-11), in dealing with the liverworts of Germany, Austria,
and Switzerland, reduced Nees’ confusing array of taxa to three, namely M.

1969

THE MICHIGAN BOTANIST

147

polymorpha (corresponding to A. communis 7 . domestica Nees) and the
forma aquatica (including A. communis a . aquatica Nees and j3 . riparia Nees)
and alpestris (equating with B. alpestris Nees and its variations). It is this view
of the species as a whole that is presented (at the varietal level) in this paper.
Muller also recognized the forma mamillata Hagen, which has not been gener¬
ally accepted by other authors, for, as Evans (1917) states, “it is based on
exceedingly vague characters.” Muller, in a later publication (1951-58), treated
his forms as varieties.

The essential features of Marchantia polymorpha are well known from
accounts in textbooks of botany. Should goblet-shaped gemma cups be pres¬
ent on the upper surface of a flat, forking thallus, the plant is immediately
recognizable as a Marchantia. Marchantia polymorpha typically has ventral
scales in several rows (at least three) on each side of the midline, with the
outermost projecting beyond the thallus edge. In this respect it differs from
both Preissia quadrata (Scop.) Nees and Conocephalum conicum (L.) Dum.,
which in northern Michigan grow in situations somewhat similar to those
where Marchantia is found but have median scales only. In the upper part of
the thallus are air chambers containing photosynthetic filaments and opening
to the exterior by barrel-shaped pores which contrast with the simple pores of
Conocephalum. When gametophores are present, Marchantia is easily recog¬
nized by the characteristic shapes of the male and female receptacles.

Descriptions of the var. aquatica are given by many authors, including
Nees (1838), Macvicar (1912; 1926), and Burgeff (1943). From their accounts
we can conclude that the characteristic features of var. aquatica are the thin
thallus showing little brandling, the sunken, dark median line where air cham¬
bers are lacking, the poorly developed ventral scales, and in some instances
infrequent occurrence of gemma cups and gametophores. The diagnostic
features of the var. alpestris (not particularly pertinent to the present discus¬
sion) are well set out by Warncke (1968).

There is considerable evidence that var. aquatica and var. alpestris differ
from var. polymorpha in other than morphological features. Burgeff (1943), as
a result of extensive cultural and genetic studies, concluded that the three
taxa warrant specific rank. Cytological work on Burgeff’s material was carried
out by Haupt (1932), who reported that although these taxa all have the same
haploid chromosome number of 9 (8 + m), the m (or sex) chromosome of
female plants of var. polymorpha is four times the mass of that in male
plants; but in var. aquatica and var. alpestris the sex chromosomes are of an
equivalent size in both male and female plants. Material from Michigan, cul¬
tured and treated according to the method described by Haupt, gave compara¬
ble results, although observed differences were slight, owing to the minute
size of the chromosomes. (This included female material of var. polymorpha
collected by M. Frohlich in Benzie County and male material of the same
collected by H. Crum in Presque Isle County, as well as male and female
material of var. aquatica collected by the author in Cheboygan County.) Bio¬
chemically, also, the var. aquatica may be distinct, for Tucker (1963) reported
that the thallus contains a specific antigenic protein in contrast to that found

148

THE MICHIGAN BOTANIST

Vol. 8

in var. polymorpha. In range of distribution, the three varieties often overlap,
although it has been shown that var. aquatica alone has become established in
glass houses and shade houses within recent years in New Zealand (Campbell,
1969) and that var. alpestris is tolerant of high concentrations of copper and
other heavy metals (Warncke, 1968).

Living material of M. polymorpha var. polymorpha was collected in
northern Michigan by D. Vitt in Luce County, from a sandstone bluff near
Upper Tahquamenon Tails; by M. Lrohlich in Benzie County, between Platte
River and Goose Road underneath the steps of a cabin built in 1957 on a
filled area in a Thuja swamp; and by H. Crum in Presque Isle County, 1.5
miles northwest of Mast Point on U. S. Highway 23, on the banks of a
roadside ditch. The thallus is green to yellow-green and has air chambers over
its entire width. On the upper surface, polygonal areas are evident because of
the colorless, or nearly colorless, partitions between air chambers. The pores
are normally open but become closed when the thallus is immersed in glyc¬
erol, indicating that they are functional. On the under surface, the scales are
well developed and in several rows. The median scales carry appendages which
are often made conspicuous by a magenta coloring in their cell walls. This
color may be present in the scales and the flat portion of the thallus, as well.
Rhizoids are most abundant and often brown in color. No gemma cups were
found, but some thalli bore antheridiophores, others archegoniophores with
maturing sporogonia.

Comparable specimens in the Herbarium of the University of Michigan
(MICH) or of the Biological Station (UMBS) include the following:

CHEBOYGAN CO. Bryants Bog, on burnt ground, C. M. Roberts, 1921 (UMBS).
MACKINAC CO. Bois Blanc Island, Frances Wynne, 1942 (UMBS); Mackinac Island, burnt
soil near lime kiln, Lime Kiln Trail, N. G. Miller, 1965 (UMBS, MICH). DELTA CO.
Northeast part of Summer Island, among sedges on damp organic debris over limestone
pavement, N. G. Miller, 1968 (UMBS, MICH).— All these specimens had either gemma
cups or antheridiophores or archegoniophores.

Plants representative of the var. aquatica were found growing in mats
in the seepage pools and streamlets bordering Carp Creek, in the vicinity of
the Iron Bridge. The thallus is submerged or likely to be submerged at times
of high water, is dark-green in color and thin in texture. Air chambers are
normally present except at the very dark midline which shows up with its lack
of air chambers as a prominent feature of the upper surface; only in a few
totally submerged colonies was a suppression of air chambers over the entire
width of the thallus observed. In no plants are polygonal areas corresponding
to the air chambers conspicuous, for the partitions between the air chambers,
like the filaments within the chambers, contain many chloroplasts. The pores
are normally closed but may open in air of high humidity (as in a plastic bag),
only to close again immediately if the thallus is immersed in glycerol, showing
that they are responsive to turgor changes. On the lower surface the scales
are poorly developed. However, more particularly near the growing apices,
median scales with appendiages are usually observable, and in places at the
edge of the thallus small marginal scales generally can be seen, usually

1968

THE MICHIGAN BOTANIST

149

projecting somewhat. Multiplication of thalli is chiefly from fragmentation, for
gemma cups are infrequent or absent, and although in some colonies antheri-
diophores or archegoniophores are found, rarely are any sporogonia present.
Adjustment to drier conditions by submerged forms is too slow for survival,
since plants stranded when the water dries up in summer were seen to die
away. In this way, many colonies are obliterated, but, provided that the dry
period is not too extended, adventitious shoots may develop ventrally from
the midline region and eventually re-establish the colony.

The following specimens are considered to be comparable:

CHEBOYGAN CO. Reeses Bog, Frances Wynne, 1942 (UMBS); Iron Bridge, Carp Creek,
on wet soil, Margaret Fulford, 1951 (UMBS). LUCE CO. Upper Tahquamenon Falls, H.
Crum & N. G. Miller, 1966 (UMBS). ALPENA CO. Growing over Drepanocladus
uncinatus at bottom of a sink hole, A. G. Miller, 1968 (UMBS).

It would seem that M. polymorpha var. polymorpha is the plant which
grew as extensive colonies on burnt-over areas at the time when burning of
bog and forest was a regular practice in northern Michigan (Nichols, 1922;
Gates, 1942). Roadbanks and old campfire sites were also favored locations
for this variety (Fulford, personal communication). Now, with fewer fires and
road construction of a different type, not leaving wet, shady spots favorable
to Marchantia, only a few habitats are left available to it, such as limestone
rocks and some swampy areas.

Although it is true that var. aquatica is not always sharply distinct from
the many forms of var. polymorpha, it is, in its usual form, strikingly differ¬
ent and easily recognized. For this and other reasons mentioned above, it
seems useful and appropriate to recognize it at the varietal level. It grows in
particularly wet situations and is at times completely submerged. Judging from
specimens in the liverwort herbarium of the American Bryological Society at
the University of Cincinnati, the North American distribution of var. aquatica
is predominantly northeastern, in a circle around the Great Fakes including
Ontario, Nova Scotia, Pennsylvania, Wisconsin, and Minnesota, and north¬
western, in Oregon, Washington, Alberta, and Montana; but it is also known
from the High Plains of Nebraska (at 3120 ft.) and in New Mexico (at 8500
ft.).

ACKNOWFEDGMENTS

Acknowledgment is made to the National Science Foundation for funds for partici¬
pation in a workshop in bryology at the University of Michigan Biological Station in the
summer of 1968 (grant number NSF-GB-6095) and also to Massey University for a study
leave grant and to the Department of Biological Sciences of the University of Cincinnati
for the use of facilities.

FITERATURE CITED
Burgeff, H. 1943. Genetische Studien an Marchantia. Jena.

Campbell, Ella O. 1969. Marchantia polymorpha var. aquatica in New Zealand. Tuatara
16: 179-184.

Evans, A. W. 1917. The American species of Marchantia. Trans. Conn. Acad. 21:
201-313.

150

THE MICHIGAN BOTANIST

Vol. 8

Gates, F. C. 1942. The bogs of northern Lower Michigan. Ecol. Monogr. 12: 213-254.
Haupt, Gertraud. 1932. Beitrage zur Zytologie der Gattung Marchantia (L.) I. Zeitschr.
Abst. Vererb. 62: 367-428.

Macvicar, S. M. 1912. The Student’s Handbook of British Hepatics. Ed. 1. London.

- - - . 1926. Ibid. Ed. 2. Eastbourne.

Maravolo, N. C., & P. D. Voth. 1966. Morphogenic effects of three growth substances on
Marchantia gemmalings. Bot. Gaz. 127: 79-86.

Muller, K. 1906-11. Die Lebermoose Deutschlands, Osterreichs, u. d. Schweiz. In Raben-
horst’s Kryptogamen-Flora von Deutschland usw. Band 6. Ed. 2. Leipzig.
- . 1951-58. Ibid. Ed. 3.

Nees von Esenbeck, C. G. 1838. Naturgeschichte der europaischen Lebermoose. IV.
Breslau.

Nichols, G. E. 1922. The bryophytes of Michigan with particular reference to the Douglas
Lake region. Bryologist 25: 41-58.

Tucker, W. G. 1963. Serological relationships with antigens extracted from green tissues.
Nature 200: 597-598.

Voth, P. D. 1941. Gemma-cup production in Marchantia polymorpha and its response to
calcium deficiency and supply of other nutrients. Bot. Gaz. 103: 310-325.
- . 1943. Effect of nutrient solution concentration on the growth of Mar¬
chantia polymorpha. Bot. Gaz. 104: 591-601.

Voth, P. D., & K. C. Hamner. 1940. Responses of Marchantia polymorpha to nutrient
supply and photoperiod. Bot. Gaz. 102: 169-205.

Warncke, E. 1968. Marchantia alpestris in Denmark. Bot. Tidsskr. 63: 358-368.

Review

NATURAL AREAS IN INDIANA AND THEIR PRESERVATION. By Alton A. Lindsey,
Damian V. Schmelz, & Stanley A. Nichols. The Report of the Indiana Natural Areas
Survey. Department of Biological Sciences, Purdue University, Lafayette, Indiana
47907. 1969. 594 pp.

Our neighbor to the south has produced a handsomely printed and bound report,
magnificent in both concept and execution, which can elicit only praise and envy from
other states. Members of the Michigan Natural Areas Council, which is engaged in a much
more modest inventory of natural areas, will applaud the first 67 pages of background
material on principles of preservation, the nature of this compendium, and the natural
vegetation divisions and soils of Indiana. Cataloging of 162 areas follows. In addition to
information on size, exact location (with directions where needed), ownership, priority
for preservation, and general features of each area, there are often data on vegetation, site
and vegetation maps, photographs (including sometimes stereopairs), and literature ref¬
erences. The volume concludes with the text of the 1967 act by the General Assembly of
Indiana establishing a state system of nature preserves. Indiana residents and all others
who share the same concerns will be profoundly grateful to the industrious authors of
this survey and to the Ford Foundation, which financed their work 1967-1969.

-E. G. V.

1969

THE MICHIGAN BOTANIST

151

BIOLOGY OF THE MOUSE-EAR CHICKWEED,

CERASTIUM VULGATUM

Sylvie-Ring Peterson

Department of Botany, The University of Michigan, Ann Arbor

INTRODUCTION

The common Mouse-ear Chickweed, Cerastium vulgatum L. (Caryo-
phyllaceae), has spread widely throughout the temperate and subtropical re¬
gions of the world, evidently as a result of features that make it especially
successful in following human cultivation and disturbance of sites. This investi¬
gation of the plant’s ecological life history endeavored to determine some of
the attributes which have led to its great success as a weed.

The success of this species is due in part to pre-adaptation of its growth
form, germination characteristics, and life cycle to the environmental
conditions of the disturbed or cultivated areas which it colonizes; and in part
to its large production of easily dispersed seeds, to its adventitious growth,
and to the variability of its populations. I attempted to investigate these
various aspects of the species. A few questions were resolved with adequate
sampling and reliable results, but for other questions, only preliminary results
were obtained under the conditions of the study.

The problem of nomenclature and circumscription of the species is
unresolved at the moment due to an essentially worldwide distribution and
possible hybridization. In Europe it is treated as C. fontanum Baumg. ssp.
triviale (Link) Jalas (Tutin, 1964). It is identified here on the basis of char¬
acterization by Gleason (1963).

Plant materials used for the experiments came largely from seeds col¬
lected from lawn and border weed plants on the main campus of the Univer¬
sity of Michigan and from lawn and old-field weed plants at the University
Botanical Gardens, Ann Arbor. Except when stated to the contrary, all obser¬
vations reported below are based on plants grown from these seed sources. In
addition, collections of seed were made at nine localities across the United
States and Canada in order to sample the regional variation of C. vulgatum.
These nine seed sources with the dates of collection are:

1. Ridgewood, New Jersey; July 15, 1967

2. High Point, New Jersey; August 12, 1967

3. Madison, Wisconsin; June 25, 1967

4. Minneapolis, Minnesota; June 25, 1967

5. Iowa City, Iowa; July 12, 1967

6. Pine Bluffs, Wyoming; July 11, 1967

7. Nanaimo, British Columbia, Canada; July 2, 1967

8. Victoria, British Columbia, Canada; July 2, 1967

9. Olympic, Washington; July 4, 1967

152

THE MICHIGAN BOTANIST

Vol. 8

All the plants were grown experimentally under uniform conditions of light
and water on a soil composed of 50% washed sand, 25% sterilized humus-rich
silt loam, and 25% perlite. None received artificial light.

MORPHOLOGY

C. vulgatum is a non-fleshy perennial herb with a prostrate pattern of
growth resulting from weak stems and adventitious rooting at the nodes and
infrequently at the internodes. Larger plants have a stem length of about 0.5
meters; the largest plant grown under glass in this study had a stem length of
0.57 m. The majority of plants studied had stem lengths of 0.1 to 0.2
meters. Some plants flowered at less than 0.18 m, although most plants usual¬
ly had longer stems at time of flowering due to an elongation of internodes
near the apex in preparation for flowering.

Most commonly the basic growth form consists of several long stems
radiating from one central point, but this is often obscured by adventitious
spreading of the plant. All the populations except the one from Pine Bluffs,
Wyoming, exhibited this growth pattern. In contrast, the plants from Pine
Bluffs had masses of tiny leaves growing around the point at which the roots
entered the soil and on the few portions of branched stems. Since only 4% of
the seed from Pine Bluffs germinated, it is not possible to assess the cause of
this unusual form.

The leaves are opposite and have no stipules. They vary from oblong to
ovate, taper at the apex, and are 1-3 cm long and 0.3-1 .5 cm wide. The three
West Coast populations sampled had leaves which were shorter and wider than
the leaves of the other populations. Leaf variation may be divided into three
categories for comparison:

A. Long and narrow (length/width > 2.6/1): High Point, Madison
(mixed with roundish), Minneapolis (many without the characteristic
narrowing of the blade as it joins the stem), Iowa City, and Pine
Bluffs.

B. Intermediate (length/width = 2.5/1): Nanaimo and Olympia.

C. Roundish (length/width < 2.4/1): Victoria

The plant is covered with fine hairs. On the stem these hairs are of
variable lengths and slant downward. The Madison plants had very little hair
on the stems; those from Minneapolis and Victoria had very hairy stems. Most
of the hair on the leaves is on the upper surfaces; the hair on the under
surfaces lies along the veins and margins. The hairs on the leaves are multicel¬
lular and uniseriate. Leaves of plants from all regions were similar in hairiness,
except those from High Point, which had very hairy leaves.

The plant bears its flowers toward shoot apices. The flowers are either
solitary or terminal in dichotomous cymes. Tire mature pedicels are from 0.5
to 1.2 cm long (Gleason, 1963) and elongate as the fruit matures.

In a plant that was sectioned for observations of stem anatomy, there
were nine collateral vascular bundles in a ring surrounding a pith cavity. Out¬
side the vascular bundles were three layers of sclerenchyma cells in the cortex.

1969

THE MICHIGAN BOTANIST

153

The outer cortex was chlorenchymatous and had more air spaces than cells.
Although Korsmo (1954) has reported secondary growth in C. arvense, no
evidence of secondary growth was seen in the plant studied.

Other external morphological characters of C. vulgatum are described in
Gleason (1963). In addition, seedling morphology is treated in detail by Kum-
mer (1951), the morphology of the flower is covered by Thompson (1942),
and the morphology of the seed is discussed by Woodcock (1928).

From experimentation with four light levels— achieved by using 0, 1, 2,
and 3 layers of dark green shade cloth over growing plants— it was found that
C. vulgatum can grow under any of these light conditions but thrives in full
sunlight. Four plants of uniform size, each in a separate 6-inch pot, were
placed under each combination of shade cloth layers. After a period of four¬
teen weeks, the growth of the plants under all four light conditions was
measured by fresh weight, overall length of stem, and number of nodes. Then,
after drying the plants in an oven at about 68 C for one week, their dry
weights were measured.

The results of the shade experiment are shown in the following table,
in which the weights of plants from less shaded conditions are compared with
those from the most shaded:

No. of Layers

Average Fresh

Average Dry

Comparative Weights

of Shade Cloth

Wt./Plant

(grams)

Wt./Plant

(grams)

Fresh

Dry

3

0.32

0.063

2

0.76

0.08

2.4X

1.3X

1

2.78

0.31

8.7X

4.9X

0

19.2

3.05

60. OX

48. 6X

Obviously, growth is by far the greatest in full sunlight, at least under green¬
house conditions November through March. In addition, stem length (the sum
of all the branching of the plant stem) is directly proportional to the weight
before drying.

Individual plants under each shade layer did not necessarily have a
correlation between increased length of stem and an increased number of
nodes. For example, in full sunlight, one plant, which had flowered, had a
greater number of nodes (395) for less stem length (400 cm) and fresh weight
(21 .75 g) than did another plant (with 345 nodes, length of 602 cm, and
weight of 23 g) which had not flowered. Plants which flowered showed, in
general, increase in both stem length and number of nodes. Average number
of nodes is correlated with amount of light received and seems to be a simple
index to amount of growth, as suggested by the following data:

No. of Layers of
Shade Cloth

Average No. of Nodes/Plant

3

2

1

0

33

44

75

315

However, the average length of internode appears to be relatively constant or
variable in response to some factors other than shade and amount of growth:

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Vol. 8

No. of Layers of
Shade Cloth

Average Length of Internode

3

2

1

0

1.33 cm
0.96 cm
1.30 cm
1.40 cm

No firm conclusions may be drawn from these data on growth under different
shade conditions. Under increased shade, elongation of internodes would be
expected, but this is not supported by these experiments. The general obser¬
vation that plants grown in full sun have longer internodes toward the apex
compared to shade-grown plants is also not clearly supported.

The percentage of water in C. vulgatum varied from 80.0% to 90.4%
based on measurements of water loss of each plant from each light environ¬
ment after oven drying for one week. A low, anomalous figure of 60% was
found for one of the plants grown under the three layers of shade cloth, but
this was probably a result of an error in weighing the plant, since the plant
had an extremely low weight. Disregarding this low figure, the average amount
of water in the plants is 86.2% of the fresh weight.

Different shade intensities produced morphologic differences in the
plants. The characteristics of the plants grown in full sun -light were larger
leaves, denser pubescence, denser matting, more branching, and a darker shade
of green compared to plants grown under the shade cloth. In addition to the
above differences, shade plants had leaves with curling margins and roots that
were weak and exposed above the soil surface. Although Kummer (1951) has
described C. vulgatum as a plant primarily of shady areas, these results indi¬
cate that the best growth is made in full sun.

PHYSIOLOGICAL ASPECTS

On a temperature gradient germination plate with a range of 12-35 C,
38 seeds out of 112 seeds of C. vulgatum germinated in the range from
21-24 C. One seed germinated at 25 C.

Germination between wet paper towels at room temperature (24 C) was
quite successful: on the fifth day, 24% had germinated; on the twelfth day,
56% had germinated; beyond this, germination was negligible.

Variation of percent germination of seeds from different regions could
have resulted from any of several factors, including age and maturity of the
seeds at the time of collection. Maturation of seed was undoubtedly under
different environmental influences in the various regions where collections
were made, but these could not be appraised. After cold, dry treatment, the
seeds were planted under glass in November, with the following results:

Site (and No. of Seeds) Germination Percentage

Ridgewood (60) 63%

High Point (85) 84%

Madison (35) 34%

Minneapolis (45) 81%

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155

Site (and No. of Seeds)

Iowa City (100)
Pine Bluffs (50)
Nanaimo (10)
Victoria (6)
Olympia (10)

Germination Percentage

65%

4%

40%

40%

80%

Cold treatment for one week seemed to encourage faster germination
and development, since at the end of three weeks 50% of the cold-treated
seeds had germinated to seedling stage, while only 21% of the non -treated
seeds had produced seedlings. Dry cold treatment, as opposed to wet cold
stratification, produced the most seedlings; at the end of four weeks, 79% of
the planted seed had developed. As more time elapsed, more seedlings ap¬
peared, as shown by the fact that at the end of six weeks, 62% of the planted
non-treated seeds had produced seedlings.

Brett (1955) mentions that the seeds of C. vulgatum are viable after
three or more years when stored in bags at room temperature and states
results similar to those found here, such as the finding that cold treatment
(two to three days at -2 C) hastened germination.

In my experiments, crowding of seeds when planted seemed to make
little difference in the percentage of seedlings that appeared. Depth of plant¬
ing was important, however (50 seeds planted at each depth):

Depth below Surface
0

0.5 cm
1.0 cm
2.0 cm

Percentage Appearance

86%

50%

11%

18%

Robbins, Crafts, and Raynor (1952) state that the seeds of chickweed
germinate throughout the year. Furthermore, the seeds have only a short
dormancy and thus germinate promptly after they are released from the
dehisced capsule in the spring and summer.

Plants grown from seeds collected in Ridgewood, High Point, Minneapo¬
lis, and Iowa City grew more vigorously than plants from the other non-Michi¬
gan collections. The growth of plants from the Pine Bluffs and Olympia seed
collections was distinctly less vigorous than the average growth of other non-
Michigan samples.

A significant East-West gradient may be indicated by the presence of
anthocyanin in the stem. No red color was present in the plants from Ridge¬
wood seed, but the plants from High Point, Madison, and Minneapolis seed
had bright red stems. Seed from the remaining localities (Iowa City and
westward) produced plants with brownish stems. A distinct increase of antho¬
cyanin in the stems of other C. vulgatum plants under greenhouse culture was
noted starting with the week of February 3. This increase may have been due
to increased periods or intensity of daylight.

Flowering appears to be related to the length of daylight received by
the plants, since plants both fourteen weeks old and nine weeks old were

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Vol. 8

found to have opened flowers first on the same date (February 3). Plants that
were germinated subsequently also flowered at nine weeks of age (March 29).
It would seem that nine weeks is the (minimum?) age for flowering if light
conditions are favorable. The fourteen-week-old plants produced more flowers
than did the nine-week-old plants over a period of two months after the onset
of flowering. As much as the apical third of a plant may be reproductive.
After flowering and the formation of capsules are completed, the upper part
of the plant turns yellow and dies. However, the plant is perennial from the
basal parts of aerial shoots.

C. vulgatum reproduces both sexually, by either insect or self-pollina¬
tion (Knuth, 1908), and asexually, by rooting adventitiously at the nodes and
internodes. Adventitious rooting, at least at the nodes, can be induced by
keeping a portion of the plant buried for some time; in this study rooting was
effected by this technique in approximately two weeks. Cutting with a lawn
mower stimulates the plant to a more vigorous prostrate growth (Fogg, 1945),
primarily through increased brandling and adventitious rooting.

C. vulgatum seems to have a basic chromosome number of 9. Chromo¬
some counts of 72, 126, 144, and 180 have been taken (Index to Plant
Chromosome Numbers, 1958).

BIOTIC INTERREFATIONSHIPS

Cerastium vulgatum is found in ecologically disturbed habitats such as
old crop fields, newly abandoned fields, truck and flower gardens, mowed
meadows, lawns, roadsides with fine-grained soils, naturally occurring alluvium,
and slumping, earthen banks. The plants used in this study were found on
silt-loam or clay -loam soils in well-watered lawns and fields, and along river
banks.

Cows are known to eat the weed indiscriminately, encouraging matted
growth; and dispersal of the seed by horses, reindeer, and birds is known.
Furthermore, seeds of C. vulgatum can be dispersed by the wind, if the plants
are in areas open to the wind (Ridley, 1930).

Insect pollination by long-tongued bees, short-tongued bees, other
Hymenoptera, and Diptera has been observed (Robertson, 1929). The pollen,
as reported by Edgeworth (1877), is subspheric with 6-12 pores and is ap¬
proximately 33 microns in diameter.

C. vulgatum is susceptible to fungus diseases induced by species of
Ascomycetes, Fungi Imperfecti, and Phycomycetes (von Tubeuf, 1897). In
addition, it is susceptible to infection by a strain of tobacco-mosaic virus
(Holmes, 1939). However, in my experience, diseased plants are uncommon.

ECONOMIC SIGNIFICANCE

Cerastium vulgatum is of economic significance only as a pest. It is an
unattractive weed in grain fields, gardens, and lawns. Although the seeds are
sometimes found in samples of grain (Harper, 1960), the dry biomass is small,
so the plant is not a serious pest in this respect. The plant has no known
toxic properties.

1969

THE MICHIGAN BOTANIST

157

SUMMARY

In this preliminary description and analysis of C. vulgatum, plants from seeds from
the vicinity of Ann Arbor were used for most observations. In addition, plants from seeds
collected at nine localities on a mid-latitude traverse across North America were studied
under uniform conditions and compared for presence of anthocyanin in the stems, hairi¬
ness, variation in percent germination, leaf variation, growth pattern, and growth vigor.
No anthocyanin was seen in the plants from Ridgewood, whereas bright red pigment was
seen in the plants from High Point, Madison, and Minneapolis, and the stems of the
plants from Iowa westward were brownish. Leaves of the West Coast populations were
shorter proportionately to width than were the leaves of other populations. No other
characters varied consistently on a regional basis.

Using the seeds from Ann Arbor, a second experiment measured growth of plants
under four different intensities of light. The results indicated that C. vulgatum grows best
in full sunlight, at least under greenhouse conditions. Dry cold treatment was most
effective in achieving early seed germination. Nine weeks is probably the minimum age of
a plant before it flowers, given adequate intensities or duration of daylight. Reproduction
is both asexual by adventitious rooting and sexual by insect and self-pollination.

ACKNOWLEDGMENTS

I wish to thank Dr. W. S. Benninghoff for his aid and encouragement and the
University of Michigan Botanical Gardens for its help.

LITERATURE CITED

Brett, Olive E. 1955. Cyto-taxonomy of the genus Cerastium. New Phytol. 14: 138-148.
Edgeworth, M. P. 1877. Pollen. Hardwick & Bogue, London. 92 pp.

Fogg, John M. 1945. Weeds of Lawn and Garden. Univ. Pennsylvania Press, Philadelphia.
215 pp.

Gleason, Henry A. 1963. The New Britton & Brown Illustrated Flora of the Northeastern
United States and Adjacent Canada. Hafner Publishing Co., New York. 3 vol.
Harper, John. 1960. The Biology of Weeds. Blackwell Scientific Publ., Oxford. 256 pp.
Holmes, Francis O. 1939. Handbook of Phytopathogenic Viruses. Burgess Publishing Co.,
Minneapolis. 221 pp.

Index to Plant Chromosome Numbers. 1958 [for 1957] . 61 pp.

Knuth, Paul. 1908. Handbook of Flower Pollination. Clarendon Press, Oxford. 695 pp.
Korsmo, Emil. 1954. Anatomy of Weeds. Kirstes Boktryykkeri, Oslo. 413 pp.

Kummer, Anna P. 1951. Weed Seedlings. Univ. Chicago Press, Chicago. 435 pp.

Ridley, Henry N. 1930. The Dispersal of Plants throughout the World. L. Reeve & Co.,
Ashford, Kent. 744 pp.

Robertson, Charles. 1929. Flowers & Insects. Science Press Printing Co., Lancaster, Pa.

221 pp.

Robbins, Wilfred, Alden S. Crafts, & Richard N. Raynor. 1952. Weed Control. McGraw-
Hill, New York. 503 pp.

Thompson, Betty Flanders. 1942. The Floral Morphology of the Caryophyllaceae. Am.
Jour. Bot. 29: 333-349.

Tubeuf, Karl Freiherr von. 1897. Diseases of Plants Induced by Cryptogamic Parasites.
Longmans, Green & Co., Glasgow. 598 pp.

Tutin, T. G., et al. 1964. Flora Europaea. Vol. 1. Cambridge Univ. Press, Cambridge. 464

pp.

Woodcock, Edward F. 1928. Observations on the morphology of the seed of Cerastium
vulgatum. Pap. Mich. Acad. 8: 233-238.

158

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Vol. 8

AN UNUSUAL FRUITING OF LEOTIA LUBRICA

Florence V. Hoseney

Herbarium, The University of Michigan, Ann Arbor

Reluctance on my part to stop mushroom collecting for the season of
1968 led to the discovery of an unusual fruiting of Leotia lubrica Fr. It was
found on a sandy hill which supported little vegetation except occasional
scrub oaks, a few herbaceous flowering plants, and some mosses and lichens.
Many interesting fungi have been found in this particular part of the Waterloo
Recreation Area, Washtenaw Co., Michigan, and some curious habitat adjust¬
ments were observed. Among the species gathered late in 1968, partially or
entirely covered by sand, were two large collections of Pisolithus tinctorius ,
some Scleroderma geaster, Astreus hygrometricus, Cortinarius sp., and Leotia
lubrica. Some of these fungi are considered rare in Michigan.

Leotia lubrica was abundant during the normal fruiting season in 1968:
so much so that one tended to disregard it when it was growing in its usual
way. However the collection cited here was found late in the season (Novem¬
ber 1) when few fleshy fungi were fruiting. The unusual dull green color and
the fact that it grew buried in the sand were most irregular.

Preliminary microscopic study showed this to be L. lubrica. It possessed
the same characteristics as the common L. lubrica which grows on humus or
sandy soil. In the collection under discussion the fruit-bodies were fully ma¬
tured but were still covered by the moist sand. Their presence was indicated
by cracks in the sand. Only scattered caps were visible on the surface. Most
fungous sporocarps when found below the surface of the soil are pallid and
weakly pigmented. The pigment of these L. lubrica ascocarps was well de¬
veloped even though most of the fruiting bodies were covered by about 2 cm
of sand.

The overall height of the ascocarps (3 .6-8.3 cm) plus the depth of the
sand cover indicate that the point of origin is deep in the soil. Normally,
ascocarps of L. lubrica appear on humus or sandy, barren soil, at the surface.
This collection, therefore, is of considerable ecological interest.

The following is a description by Nancy Jane Smith of Leotia lubrica
Fr. ( F \ Hoseney 1170 , 1 Nov 1968, Waterloo Area, Washtenaw Co., Mich.):

Gregarious to cespitose in sandy area, predominantly hypogeous.

Pile us flattened, cushion-like, 0.6-0. 8 cm high, 0.9-2 .7 cm broad; hymenium
smooth, dull olive green, margin curved under and the area sometimes
exhibiting lobing; gelatinous when fresh.

Stipe 3. 0-7. 5 cm long, 0.4-1 .2 cm in diam., terete to compressed with a
vertical groove, slightly lubricous under the sand, pale olive green to a
dull ochraceous color, green when underground and more yellow in
older and emerged asocarps, macroscopically smooth.

Microscopic features:

Paraphyses filiform, typically slightly enlarged apically, straight, branching in
the lower part.

1969

THE MICHIGAN BOTANIST

159

Asci narrowly clavate, 8 spored, (134) 150-164 (203 )ja X 8-9 (10.6 )/i, not
turning blue at the apex in Melzer’s reagent.

Ascospores straight to slightly curved, narrowly ellipsoid to slightly allantoid
(?) probably multiguttulate when fresh but the oil droplets coagulate
when rehydrated after drying, (16.5) 18-22.5 (25.5)ju X (4) 4.5-5 .2
(5.7)M.

ACKNOWLEDGMENTS

I wish to thank Nancy Jane Smith for the description of the material and Dr.
Alexander H. Smith for his encouragement and the photograph.

Fig. 1. Leotia lubrica, approximately natural size (Hoseney 1170).

160

THE MICHIGAN BOTANIST

Vol. 8

MICHIGAN PLANTS IN PRINT
New Literature Relating to Michigan Botany

This section lists new literature relating to Michigan Botany under four categories: A.
Maps, Soils, Geography, Geology (new maps and selected bulletins or articles on soils and
geology as these may be of use to field naturalists and students of plant distribution); B.
Books, Bulletins, etc., and C. Journal Articles (listing, respectively, all separate publications
and articles in other periodicals which cite Michigan specimens or include research based on
plants of wild origin in Michigan;-not generally including work on cultivated plants nor
strictly economic aspects of forestry, conservation, or agriculture); D. History, Biography,
Exploration (travels and lives of persons with Michigan botanical connections). When the
subject matter or relation to Michigan is not clear from the title, annotations are added in
brackets. Readers are urged to call to the editor’s attention any titles (1960 or later) which
appear to have been overlooked -especially in less well known sources.

A. MAPS, SOILS, GEOGRAPHY, GEOLOGY

Dutton, Carl E. 1968. Summary report on the geology and mineral resources of the
Huron, Seney, Michigan Islands, Green Bay, and Gravel Island National Wildlife
Refuges of Michigan and Wisconsin. U. S. Geol. Surv. Bull. 1260-1. 14 pp. $.20
[General geographical and topographical information for the areas, with conclusion
that the mineral-resource potential is poor.]

Ktichler, A. W., & Jack McCormick. 1967. Bibliography of vegetation maps of North
America. Excerpta Botanica, Sec. B, 8: 145-289. [On pp. 215-224 are listed over 60
vegetation maps for Michigan or portions of the state (even small bogs) -some of them
separately published and some in other publications. Titles are given, together with
scale, but there are no annotations as to area covered (e.g., what is the “North Tip
Block”?) nor kinds of features shown (e.g., that the Land Economic Survey cover
maps show size categories, etc.). Arrangement is chronological by date represented by
the vegetation mapped; there is an index to authors but not to counties or other
geographic subdivisions.]

Ktichler, A. W. 1967. Potential Natural Vegetation. National Atlas [of the U. S.] Sheet
90. U. S. Geol. Surv. $1.50. [Michigan, of course, is included, with 8 vegetation types.
This is a revised edition, based on the author’s 1964 larger scale map published by the
Am. Geogr. Soc.; the small scale of this edition (1:7,500,000) provides a general
picture but cannot be reliable for every tiny detail.]

Pregitzer, Karl E. 1968. Soil Survey of Muskegon County, Michigan. U. S. Dep. Agr. 95
pp. + 90 once-folded map plates + 5 folded tables and indexes. [As in other recent
soil surveys, the map plates include complete aerial photography for the county at a
scale of about 4" = 1 mile, with boundaries of soil types overprinted. An indispensible
field aid for anyone wanting quickly to survey the wooded and wild areas of the
county. $5.50 from Govt. Printing Office, Washington 20402.]

U. S. Forest Service. 1968. Sylvania Recreation Area Management Plan, Ottawa National
Forest. 47 pp. + 2 maps. [Indicates various management zones and the standards and
facilities concerned with management of each. A major portion of this wild tract in
Gogebic Co. is designated as a “Botanical Zone” (i.e., natural area) and much of the
rest is a “Pioneer Zone.”]

Zahner, Robert, & David R. Hedrich. 1966. Moisture release characteristics of forested
sand entisols in northern Lower Michigan. Soil Sci. Am. Proc. 30: 646-649. [Variation
in some moisture properties of Grayling sand under pine stands.]

B. BOOKS, BULLETINS, SEPARATE PUBLICATIONS

Brewer, Richard, Arlo Raim, & Jerome D. Robins. 1969. Vegetation of a Michigan
Grassland and Thicket. Occas. Pap. Adams Ctr. Ecol. Stud. 18. 29 pp. [Map, tables,

1969

THE MICHIGAN BOTANIST

161

photos, discussion reporting on the “Colony Farm Tract” within the city limits of
Kalamazoo.]

Federal Committee on Research Natural Areas. 1968. A Directory of Research Natural
Areas on Federal Lands of the United States of America. 129 pp. $.70 (Govt. Printing
Office, Washington 20402). [Six areas in Michigan are included, with acreages in
primary and other natural areas types. Exact locations are not given, only the county
and Federal project-forest, park, refuge, etc.]

Shaffer, Robert L. 1968. Keys to Genera of Higher Fungi, ed. 2. Univ. Mich. Biol. Sta.,
Ann Arbor. 131 pp. $3.00. [Although the keys themselves include no locality data,
the preface points out that they are designed primarily for Michigan students.]

C. JOURNAL ARTICLES

Barr, Margaret E. 1968. The Venturiaceae in North America. Canad. Jour. Bot. 46:
799-864. [Several species of these fungi are attributed to Michigan, rarely with more
precise locality data. Includes keys, index to host plants.]

Boivin, Barnard. 1966. Etudes Ptdridologiques. III. Variations du Woodsia oregana. Bull.
Soc. Bot. Fr. 113: 407-409. [Includes Keweenaw Peninsula and Porcupine Mts. in stated
range of f. cathcartiana, and cities Porcupine Mt. collection of var. squammosa (W.
abbeae) as an alleged extension of range (though cited in Billington’s “Ferns of Michi¬
gan,” 1952, pp. 145 & 224).]

Cain, Stanley A. 1967. Studies on the stemmed yellow violets of eastern North America
II. Mass-collections of Viola pubescens and V. eriocarpa in the Michigan area. Nat.
Canad. 94: 79-129. [Two of the collections studied were from Bruce Peninsula, Ontar¬
io; 18 were from Michigan, of which 11 were from Oakland Co.]

Crum, Howard. 1966. A taxonomic account of the genus Thelia. Natl. Mus. Canada Bull.
216: 123-127. [Michigan cited in the range of two of the three species of these
mosses.]

Davis, H. A., Albert M. Fuller, & Tyreeca Davis. 1968. Contributions toward the revision
of the Eubati of eastern North America III Flagellares. Castanea 33:206-241. [R.
flagellaris said to range into Michigan. Michigan material is cited of R. baileyanus
(including R. tenuicaulis* & R. uvidus*), R. plicatifolius (including R. exutus*), R.
roribaccus, R. meracus* , R. michiganensis* (including R. complex * & R. florenceae*),
R . vagus *, R. ithacanus, R. schoolcraftianus* , and R. tantalus *, the last a stunted
type perhaps referable to R. meracus. (Types of all names with asterisk are from
Kalamazoo Co.) R. peracer (type locality in Cheboygan Co.) is referred to R. multi¬
formis. ]

Davis, Margaret B. 1968. Pollen grains in lake sediments: Redeposition caused by seasonal
water circulation. Science 162: 796-799. [Studies done at Frains Lake, Washtenaw
Co.]

Gaertner, A. M., & F. K. Sparrow, Jr. 1966. A preliminary study of aquatic Phyco-
mycetes in the lakes of the Huron Mountains, Michigan. Veroff. Inst. Meeresforsch.
Bremerhaven 10: 93-106.

Grund, D. W., & D. E. Stuntz. 1968. Nova Scotian Inocybes. I. Mycologia 60: 406-425.
[Michigan collections, with localities, are cited for several of the species.]

Haard, Karen. “1968” [1969]. Taxonomic studies on the genus Arthrobotrys Corda.
Mycologia 60: 1140-1159. [Much of the material studied was from northern Michigan
localities.]

Harris, Betty D. 1968. Chromosome numbers and evolution in North American species of
Linum. Am. Jour. Bot. 55: 1197-1204. [Includes count of n = 8 for L. catharticum
from Chippewa Co.; generalized distribution maps of the L. cathartica complex and
the virginiana and sulcata subgroups include Michigan.]

Helium, A. K., & R. Zahner. 1966. The frond size of bracken fern on forested outwash
sand in northern Lower Michigan. Soil Sci. Am. Proc. 30: 520-524. [Research done at
Univ. of Mich. Biological Station. Size and density of fronds strongly influenced by
soil profile and overstory forest cover.]

Iwatsuki, Zennoske, & Aaron J. Sharp. 1967. The bryogeographical relationships between
eastern Asia and North America, I. Jour. Hattori Bot. Lab. 30: 152-170. [One

162

THE MICHIGAN BOTANIST

Vol. 8

Michigan specimen (the type of B. subcylindrica, from Kalamazoo Co.) cited for
Buxbaumia minakatae. ]

Johnson, Robert G. 1968. Notes on the distribution of Disporum maculatum (Buckl.)
Britton. Castanea 33: 262-266. [Cites Farwell’s type of D. cahnae and includes dot in
Oakland Co. in distribution map, but offers no explanation of this disjunct occur¬
rence, merely asserting that “Farwell’s meticulous description of the collection docu¬
ments this as the northernmost known station for D. maculatum." This is perhaps the
first time in history that Farwell has been termed “meticulous.’’]

Knight, Allen W., & J. Whitfield Gibbons. 1968. Food of the painted turtle, Chrysemys
picta, in a polluted river. Am. Midi. Nat. 80: 558-562. [Stomach contents of 46
turtles from Kalamazoo River included algae and higher aquatics, not named.]

Lippert, Byron E. 1967. Sexual reproduction in Closterium moniliferum and Closterium
ehrenbergii. Jour. Phycol. 3: 182-198. [One of the strains of C. moniliferum included
in this study of desmids came from near Levering in Emmet Co.]

Mickelson, Carol J., & Hugh H. litis. “1966” [1967] . Preliminary reports on the flora of
Wisconsin No. 55 Compositae IV-Composite family IV (Tribes Helenieae and
Anthemideae). Trans. Wis. Acad. 55: 187-222. [Map and discussion of range of
Tanacetum huronense includes Michigan data; distribution maps for Cirsium pumilum
ssp. hillii and Besseya bullii also include Michigan county distribution.]

Miller, Orson K., Jr. 1968. A revision of the genus Xeromphalina. Mycologia 60:
156-188. [Several of the species are cited from “Michigan,” never with more precise
locality.]

Reeder, John R., & D. N. Singh. 1967. Chromosome number in Calamovilfa. Bull. Torrey
Bot. Club. 94: 199-200. [Count of 2n = 40 from C. longifolia var. magna grown at
Yale from a clone collected at Warren Dunes State Park, Berrien Co.]

Smith, Alexander H., & Harry D. Thiers. 1968. Notes on Boletes— I 1. The generic
position of Boletus subglabripes and Boletus chromapes 2. A comparison of four
species of Tylopilus. Mycologia 60: 943-954. [Results from study of Michigan boletes;
Part 2 includes description of T. cyaneotinctus from “Michigan.”]

Strandhede, Sven-Olov. 1967. Eleocharis, subser. Eleocharis in North America. Taxonomi-
cal comments and chromosome numbers. Bot. Not. 120: 355-368. [Material of Michi¬
gan origin of E. erythropoda gave counts of 2n = 18 (Botanical Gardens, Ann Arbor)
and 2n = 19 (Cheboygan Co.). Material of E. smallii sens. str. from Ingleside, Douglas
Lake, gave count of 2n = 16; but an “unknown taxon” resembling E. smallii morph¬
ologically from Finster Marsh, Cheboygan Co., gave count of 2n = 36.]

Stuckey, Ronald L. 1968. Aquatic flowering plants new to the Erie Islands. Ohio Jour.
Sci. 68: 180-187. [Includes summary of southeastern Michigan records for Ly copus
asper and Epilobium hirsutum. ]

Tessene, Melvem F. 1968. Preliminary reports on the flora of Wisconsin No. 59. Planta-
ginaceae -plantain family. Trans. Wis. Acad. 56: 281-313. [For a few of the species,
brief comments are included on distribution or biology in Michigan.]

Thomas, Paul M., & Dennis D. Bromley. 1968. The establishment of aquatic vegetation in
and around artificial fish shelters in Douglas Lake, Michigan. Am. Midi. Nat. 80:
550-554. [Concerns influence of 10 fish shelters constructed in 1937 on establishment
of aquatic vascular plants and Chara.]

Ulken, Annemarie, & Frederick K. Sparrow, Jr. 1968. Estimation of chytrid propagules in
Douglas Lake by the MPN-pollen grain method. Veroff. Inst. Meeresforsch. Bremer-
haven 11: 83-88.

Wells, Virginia L., & Phyllis E. Kempton. 1968. Studies on the fleshy fungi of Alaska. II.
Mycologia 60: 888-901. [Psathyrella rigidipes mentioned as frequently found in lawns
at Ann Arbor, according to A. H. Smith.]

Wetzel, Robert G., & Don L. McGregor. 1968. Axenic culture and nutritional studies of
aquatic macrophytes. Am. Midi. Nat. 80: 52-64. [Some of the Chara studied came
from Barry Co.]

Yeatman, C. W. 1967. Biogeography of jack pine. Canad. Jour. Bot. 45: 2201-2211.
[Chiefly on postglacial history of the species; includes reference to Michigan pollen
profiles.]

Editorial Notes

The Editorial Board has approved a policy of modest payment for cover pictures.
At least the cost of materials may be covered by the $5.00 now offered photographers.
(Members of the Editorial Board and officers of the Michigan Botanical Club are ineligi¬
ble for payment.) Good black and white prints suitable for cover pictures are earnestly
solicited and may be submitted to the editor at any time.

Several defective copies of the March issue have been discovered, pages 68, 69, 88,
& 89 being blank. Defective copies returned to the business manager will be promptly
replaced.

The March number (Vol. 8, No. 2) was mailed March 7, 1969.

Program Notes

The state membership meeting of the Michigan Botanical Club was held March 16,
1969, at Nazareth College, Kalamazoo, with Dr. Elaine Hurst, head of the Biology De¬
partment, in charge of local arrangements. The new Southwestern Chapter was welcomed
into the Club and was presented with a handsome charter. This chapter now has about
80 members. The officers (as pictured below, left to right) are Mrs. Helen Wiles, Parch¬
ment, Michigan, secretary; Dr. John Beaman, Michigan State University, vice president;
Dr. Richard Pippen, Western Michigan University, president; and Dr. Harriette Bartoo, W.
M. U., treasurer. The program included illustrated talks on Michigan wildflowers and early

spring fungi by Helen V. Smith and Dr. Alexander H. Smith. Several field trips were led
to points of interest: the Nazareth College Ecology Center, by Dr. Hurst; a locality for
the rare climbing fern, Lygodium, by Dr. Pippen; Kellogg Forest for early fungi, by Ellis
Becker; the Audio-Tutorial Center for Biological Science at Western Michigan University,
by Donna Schuman; the M. S. U. Biological Station at Gull Lake, by Peter Rich; and the
Kalamazoo Nature Center, by Dr. Bartoo.

-Helen V. Smith

The Southeastern Chapter will hold its fall campout at Camp Kett (4-H), near
Cadillac, October 3-5, 1969.

CONTENTS

♦

The Introduction and Spread of Lycopus asper
(Western Water Horehound) in the Western Lake
Erie and Lake St. Clair Region

Ronald L. Stuckey . Ill

Mosses from Southern Michigan:

New Distributional Records

Paul L. Redfearn, Jr., & Richard L. Halbert . 120

Species of Vascular Plants of Pennfield Bog,

Calhoun County, Michigan

Garrett E. Crow . 131

The Log Fern (. Dryopteris celsa ) and Its Hybrids
in Michigan— A Preliminary Report

W. H. Wagner, Jr., Florence S. Wagner, & D. J. Hagenah . 137

News of Botanists . 145

Marchantia polymorpha in Northern Michigan

Ella 0. Campbell . 146

Review— Natural Areas in Indiana . 150

Biology of the Mouse-Ear Chickweed,

Cerastium vulgatum

Sylvie-Ring Peterson . 151

An Unusual Fruiting of Leotia lubrica . 158

Michigan Plants in Print . 160

Editorial Notes . 163

Program Notes . 163

(On the cover: Patterns in driftwood
on the shore of Lake Superior at Whitefish Point, Michigan.

Photographed July 20, 1967, by John S. Russell.)

THE

Vol. 8, No. 4

MICHIGAN BOTANIST

LIBRARY

.'40V 13 I%9

NEW YORK
BOTANICAL GARDEN

October, 1969

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V i t \ > 1

Edward G. Voss, Editor in Chief

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1509 Kearney Rd., Ann Arbor, Michigan 48104

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THE MICHIGAN BOTANICAL CLUB

Membership in the Michigan Botanical Club is open to anyone interested in its aims:
conservation of all native plants; education of the public to appreciate and preserve plant
life; sponsorship of research and publication on the plant life of the State; sponsorship of
legislation to promote the preservation of Michigan native flora and to establish suitable
sanctuaries and natural areas; and cooperation in programs concerned with the wise use and
conservation of all natural resources and scenic features.

Dues are modest, but vary slightly among the chapters and with different classes of
membership. Persons desiring to become state members (not affiliated with a local chapter)
may send $3.00 dues to the Membership Chairman listed below. In all cases, dues include
subscription to THE MICHIGAN BOTANIST. (Persons and institutions desiring to sub¬
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stated at the top of this page.)

President: W. H. Wagner, Jr., Botanical Gardens, University of Michigan, Ann Arbor,
Michigan 48105

Vice President: William F. Hopkins, 6210 Cowell Rd., Brighton, Michigan 48116
Recording Secretary: Mary Cooley, 703 S. Forest, Ann Arbor, Michigan 48104
Corresponding Sec.: Thelma Thomson, 15093 Faust, Detroit, Michigan 48223
Treasurer: Charles Buswell, 19204 Plainview, Detroit, Michigan 48219
Membership Chairman: Harriette V. Bartoo, Department of Biology, Western Michigan Uni¬
versity, Kalamazoo, Michigan 49001
Corresponding Secretary, Southeastern Chapter:

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Secretary, Huron Valley Chapter:

Ursula R. Freimarck, 704 Madison PI., Ann Arbor, Michigan 48103
Secretary, Southwestern Chapter:

Helen Wiles, 7113 N 25th St., Rt. 2, Kalamazoo, Michigan 49004

1969

THE MICHIGAN BOTANIST

167

STUDIES OF THE BYRON BOG IN SOUTHWESTERN ONTARIO.
XL. DISTRIBUTION OF SPHAGNUM MOSSES

William W. Judd

University of Western Ontario, London, Ontario

The Byron Bog has been described by Judd (1957). It lies in a hollow at
the southwest corner of Oxford Street and Hyde Park Sideroad in London,
Ontario. There are three vegetation zones, A, B, and C (Pig. 1), in the bog.
The central part is a mat of floating Sphagnum moss (Zone A). Surrounding
this is an expanse of low woods, permanently damp or flooded (Zone B). In
this zone Sphagnum grows beneath an overstory of shrubs. At its inner limits
there are scattered black spruce and larch trees and at its outer limits are vari¬
ous deciduous trees. The outer rim of the bog consists of dry, wooded slopes
(Zone C). In the floating bog is Redmond’s Pond (D). Descriptions of the veg¬
etation in the various zones include accounts of shrubs and vines (Judd,
1966), club-mosses and ferns (Judd, 1967), and orchids (Judd, 1968). In 1968
a study was made of the distribution of Sphagnum mosses in the bog.

METHODS

Sphagnum mosses were collected in the bog on October 6, 1968. Seven
series of samples were taken by starting as close as possible to the edge of
Redmond’s Pond at seven points around the perimeter of the pond and walk¬
ing outward in a straight line from each point to the limits of the Sphagnum
growth. At regular intervals along the sample line, samples of mosses were
taken, consisting of a handful of moss made up of the tips of fronds about
three inches long. Along four of the sample lines five samples were taken and
along three of the sample lines ten samples were taken, thus yielding fifty
samples. In addition, fourteen samples were taken at the outer limits of
Sphagnum growth where Zone B meets Zone C. The location of the sixty-four
sampling points is shown by circles on the map (Pig. 1). The mosses were
pressed lightly in newspaper until dry. They were identified by Dr. Howard
Crum, Department of Botany, University of Michigan. Dr. Crum retained rep¬
resentative specimens for the University of Michigan Herbarium. Other speci¬
mens are deposited in the herbarium of the writer with accession numbers 738
to 750 as in the following account. Nine species and one variety were recog¬
nized and the distribution of them on the bog is shown on the map (Pig. 1).
While most of the samples included only one species, some included two or
three growing closely together.

ACCOUNT OP SPECIES COLLECTED

Sphagnum mosses formed the floating mat of Zone A and extended
throughout Zone B where they grew on solid soil and formed cushions around
the bases of larch and black spruce trees at the inner limits of Zone B. No
Sphagnum grew on the wooded slopes of Zone C. The species collected are

168

THE MICHIGAN BOTANIST

Vol. 8

discussed in order of decreasing numbers of samples, and the accession num¬
bers of specimens in the writer’s collection are included.

S. magellanicum Brid.— 23 samples (744, 745, 747). This species is wide¬
ly distributed in the bog, from the edge of Redmond’s Pond to the outer
limits of Zone B. The purple colour of the fronds, as described by Darlington
(1964), was evident in the specimens. S. magellanicum is widely distributed in
North America in bogs and has been reported from Michigan (Crum, 1964;
Darlington, 1964), Indiana (Welch, 1957), and Pennsylvania (Jennings, 1951).

sphagnum.-

ca - capillaceum
ce - centrale
f - f im briatum
m - magellanicum
p- papillosum

r\

\ \ v

r

\

r'

(

m re

• -i- -

) .f

re -

rec urv um

ri -

r i p a r i u m

rt -

recurvum

var. tenue

rt.

\ 'V

•ca

_y

r

V;-

s

s - subsecundum
t - teres

i j\\

O LA

/

\

/

\

)

\

^ Ve Cm°

rtS

•er.cV n\'*m

.. mrDlf 3

s 3,

i

i/

rh

I

re '•••
( ca* re

V *re
^ *f

•re

f*

• ca

\

B

\

\

(

V

>

/ A

<

- 1

V

/

Fig. 1. Map of Byron Bog showing location of sampling sites for Sphagnum and distribu¬
tion of species of Sphagnum.

A: Floating mat B: Low, damp woods C: Wooded slopes D: Redmond’s Pond

1969

THE MICHIGAN BOTANIST

169

S. recurvum P.-Beauv.— 23 samples (739, 743, 747, 748). This species is
widely distributed in the bog, from the edge of Redmond’s Pond to the outer
limits of Zone B. It is widely distributed in North America in bogs and has
been reported from Michigan (Crum, 1964; Darlington, 1964) and Indiana
(Welch, 1957). Jennings (1951) reports that it is the commonest species in
Pennsylvania.

S. fimbriatum Wils. ex Hook.— 20 samples (738, 750). This species is
widely distributed in the bog, and is the species most commonly piesent in
the shade of trees where Zone B meets Zone C. It is found over much of
North America in bogs. It has been reported from Michigan (Darlington,
1964), Indiana (Welch, 1957), and Pennsylvania (Jennings, 1951).

S. capillaceum (Weiss) Schrank— 16 samples (741, 744, 748). This species
is widely distributed in the bog. It occurs over much of North America and
has been reported from Michigan (Crum, 1964; Darlington, 1964), Indiana
(Welch, 1957), and Pennsylvania (Jennings, 1951).

S. recurvum var. tenue Klinggr.— 3 samples (747, 749, 750). All three
samples of this moss were found in the shade of trees where Zone B and Zone
C meet. This species is recorded by Darlington (1964) as occurring from New¬
foundland to Pennsylvania and across the continent, and it has been recorded
from Michigan (Darlington, 1964), Indiana (Welch, 1957), and Pennsylvania
(Jennings, 1951).

S. centrale C. Jens.— 2 samples (742, 746). These samples were found at
the edge of Redmond’s Pond. This species is widely distributed in North
America. It is reported from Michigan (Crum, 1964; Darlington, 1964), but
not from Indiana (Welch, 1957) or Pennsylvania (Jennings, 1951).

S. subsecundum Nees— 1 sample (742). This moss was collected at the
edge of Redmond’s Pond. It is a species widely distributed in North America
in bogs and has been recorded from Michigan (Crum, 1964; Darlington, 1964),
Indiana (Welch, 1957), and Pennsylvania (Jennings, 1951).

S. teres (Schimp.) Angstr. ex C. Hartm.— 1 sample (740). This moss was
found on the floating mat. It occurs in the northern part of North America
southward to New Jersey and Michigan (Darlington, 1964) and has been re¬
ported from Pennsylvania (Jennings, 1951) but not Indiana (Welch, 1957).

S. papillosum Lindb.— 1 sample (743). This species was collected from
the floating mat. It occurs in the northern part of North America southward
to New Jersey and the Great Lakes (Darlington, 1964) and has been reported
from Michigan (Crum, 1964; Darlington, 1964) and Pennsylvania (Jennings,
1951), but not Indiana (Welch, 1957).

S. riparium Angstr.— 1 sample (746). This was found at the edge of Red¬
mond’s Pond. It is a moss of the far north, i.e. Greenland, Labrador and
Alaska (Andrews, 1915). It is rare in Michigan (personal communication, H.
Crum, October 25, 1968) and has not been reported from Indiana (Welch,
1957) or Pennsylvania (Jennings, 1951).

The four species which are common and generally distributed in the
bog, S. magellanicum, fimbriatum, recurvum, and capillaceum, are widely dis-

170

THE MICHIGAN BOTANIST

Vol. 8

tributed in North America. S. recurvum var. tenue is generally distributed in
North America and was found only in the shaded outer reaches of the bog. S.
papillosum and S. teres , found only on the floating mat of moss, are species that
occur in the northern part of North America. Of the three species found at the
edge of the pond, S. subsecundum and S. centrale are found generally in North
America and S. riparium is a moss of the northern part of the continent.

LITERATURE CITED

Andrews, A. L. 1915. Notes on North American sphagnum VI. Bryologist 18: 1-6.

Crum, H. 1964. Mosses of the Douglas Lake region of Michigan. Mich. Bot. 3: 3-12,
48-63.

Darlington, H. T. 1964. The Mosses of Michigan. Cranbrook Inst. Sci. Bull. 47. 212 pp.
+ 147 figs.

Jennings, O. E. 1951. A Manual of the Mosses of Western Pennsylvania and Adjacent
Regions. Am. Midi. Nat. Monogr. 6. ed. 2. 396 pp.

Judd, W. W. 1957. Studies of the Byron Bog in southwestern Ontario I. Description of
the bog. Canad. Entomol. 89: 235-238.

Judd, W. W. 1966. Studies of the Byron Bog in southwestern Ontario. XXVI. Distribu¬
tion of shrubs and vines. Mich. Bot. 5: 51-56.

Judd, W. W. 1967. Studies of the Byron Bog in southwestern Ontario XXVIII. Distribu¬
tion of club-mosses and ferns. Canad. Field-Nat. 81: 110-113.

Judd, W. W. 1968. Studies of the Byron Bog in southwestern Ontario XXX. Distribution
of orchids in the bog. Rhodora 70: 193-199.

Welch, W. H. 1957. Mosses of Indiana. Indiana Dep. Conservation. 478 pp.

POPULATION STUDIES IN THE CHEMICAL
SPECIES OF THE CLADONIA CHLOROPHAEA GROUP1

Richard Wetherbee

Department of Botany, The University of Michigan, Ann Arbor

It has become increasingly popular in lichenology to stress character¬
istics based solely on chemical constituents rather than the traditional mor¬
phological features typical of plant systematics. The basis for chemical species
formation is not clearly understood. The chemical differences are believed to
be of genetic origin. However, the effects of habitat and substrate on the
chemical nature of the podetia have not been extensively explored. Investiga¬
tions with Parmelia bolliana (Hale, 1967) and Cetraria ciliaris (Graham, 1969)
have indicated that the environment is not related to the cause of chemical

^Contribution from the Department of Botany, University of Michigan, and the
University of Michigan Biological Station.

1969

THE MICHIGAN BOTANIST

171

species formation. Zopf (1903) did find evidence of such a relationship with
the environment in the Pseudevernia complex; however, similar results have
not been extensive enough to justify any reliable, overall conclusions.

The Cladonia chlorophaea group consists of four chemical species each
of which differs essentially by chemical constituents. The acid fumarproto-
cetraric is the only acid present in Cladonia chlorophaea (Florke) Spreng. The
other chemical species, which may or may not include fumarprotocetraric
acid, are C. cryptochlorophaea Asah. containing cryptochlorophaeic acid, C.
grayi Merr. containing grayanic acid, and C. mero chlorophaea Asah. containing
merochlorophaeic acid. C. chlorophaea ranges throughout the United States;
C. cryptochlorophaea and C. grayi are widespread in the northeastern and
midwestern portions of the country; and C. merochlorophaea occurs in the
Great Lakes region and northern New England. These distinct ranges suggest
taxonomic significance.

The purpose of this investigation was fourfold: First, to establish the
percentage of each species within the total population in the vicinity of the
University of Michigan Biological Station (Cheboygan County, Michigan). Sec¬
ond, to determine whether these percentages vary within any of the ecological
communities tested. Third, to test for substrate specificity. Fourth, to deter¬
mine whether there are morphological differences in the podetia of the various
species.

At the beginning of the study I attempted to determine the relative size
of a colony and thereby judge what should constitute a random sample. I
chose a colony at or near the center of a clearing to act as a point of refer¬
ence, and then took one podetia every inch up to one foot, and thereafter
one podetium every foot along a transect of about 1 5 feet. After analysis I con¬
cluded that a population of C. chlorophaea, at least, extends over an area of
almost 10 feet when podetia arise only from the soil. However, podetia taken
from bark (usually but not always at the base of trees) within the same area

MERO- CRYPTO¬

CHLOROPHAEA CHLOROPHAEA

GRAYI

CHLOROPHAEA

Fig. 1. Percentage of each species within the entire population studied.

172

THE MICHIGAN BOTANIST

Vol. 8

typically belong to other members of the complex. Thus it was possible for
two or more species to be within inches of each other. After repeating this
procedure with similar results, I concluded that the remainder of my sampling
could be accomplished by walking at least 10 feet before sampling again in a
similar way, unless there was a suitable change in the substrate.

I sampled stands of bigtooth aspen ( Populus grandidentata), red oak
(< Quercus rubra), jack pine (Pinus banksiana), and paper birch ( Betula papy-
rifera). Twenty samples were taken at one time, half on soil and the other
half on bark. Each sample was analyzed by crystalline tests and, at first, by
thin-plate chromatography as well. (Later, chromatography was used only for
occasional verification.) I then computed the percentages indicated in Figs. 1-3
and made morphological comparisons. The procedure for the crystaline test
may be found in Hale (1961, p. 67).

The procedure for thin-layer chromatography was as follows: (1) Several
closely adjacent podetia were placed on a numbered microscope slide and the
acids extracted in acetone. (2) Numbered chromotographic plates cut to four

Fig. 2. Comparison of percentages of each species on soil and bark.

1969

THE MICHIGAN BOTANIST

173

inches square were spotted with acid residue and transferred from the slide by
micropipettes. (A concentrated spot, often taking several minutes of repeated
application, worked best.) (3) The plates were then lowered into the chamber
making sure that the residue was well above the level of the solvent. The
plates were left in the chromatographic chamber for 30 minutes with a solvent
mixture of benzene, p-dioxane, and acetic acid in the ratio of 4:6:1. (4) The
acids, if present, made spots on the plate corresponding to known RF values.
These were checked first with ultraviolet light, but a good percentage of the
spots would not show up unless sprayed with a 5% normal solution of sulfuric
acid and dried in a oven at 75° centigrade for 30 minutes. The RF values
were then checked and crystalline tests performed when the results were in¬
conclusive.

The percentage of each species within the entire population, as shown in
Fig. 1, is based on 500 samples. Specificity as to substrate is indicated in Fig.
2. Though C. merochlorophaea did not show much preference, C. crypto-
chlorophaea selected soil over bark five to one while C. grayi was four to one
bark over soil. The specificity of C. chlorophaea was less pronounced, occur¬
ring on soil twice as often as on bark. The percentage of each chemical species
in the different stands (Fig. 3), showed variations as well. The pine habitat is
particularly selective for G. grayi while having a low concentration of the
other species. The birch and oak stands noticeably favor C. chlorophaea, while
both C. cryptochlorophaea and C. chlorophaea were prevalent in the aspen
habitat. All of the specimens determined to be the same chemically were ex¬
amined under a dissecting microscope and found to exhibit no clearly discerni¬
ble morphological characteristics which could distinguish them from the other
species similarly studied. There seemed to be no empirical differences; how¬
ever, at the conclusion of the study, I was able to predict quite accurately

80
60
40
20

PINE BIRCH ASPEN OAK

Fig. 3. Percent of each species within individual stands.

Solid black = merochlorophaea
White = cryptochlorophaea
Stipples = grayi
Bars = chlorophaea

174

THE MICHIGAN BOTANIST

Vol. 8

which podetia belonged to which species. This can most likely be accounted
for by a combination of characteristics which I suspect are more ecological
than morphological.

Since this is the first study of its kind, my evaluation of these results
can only be an educated guess. Mason E. Hale, who is familiar with this
group, has told me that the overall percentages within the entire population
are not out of the ordinary and would be expected to vary in different locali¬
ties. However, the fact that individual communities may show percentages dif¬
ferent from those of the population as a whole, strongly indicates an ecologi¬
cal preference of some kind. The results of the substrate specificity test also
lend strong support to this possibility.

Whether the substrate or the microclimate is the major determining fac¬
tor remains to be seen. What can be seen is that the various chemical species
occur in different stands in different percentages, making up a percentage in
the total population dissimilar to that of any individual stand. Two of the
species, C. cryptochlorophaea and C. grayi, clearly show substrate preferences.
Whether the various species result from their habitats or in spite of them is
not for me to say.

ACKNOWLEDGMENTS

I wish to express my appreciation to Mason E. Hale and Howard Crum for their
aid and encouragement throughout the project, and to Gordon McBride for critically re¬
viewing the manuscript. This work was made possible by an NSF grant-in-aid provided by
the University of Michigan Biological Station and is also a result of a workshop in bryolo¬
gy and lichenology financed by NSF grant GB-6095.

LITERATURE CITED

Graham, William L. 1969. The occurrence of the lichen complex, Cetraria ciliaris, in
the Straits region of Michigan. Mich. Bot. 8: 67-71.

Hale, Mason E., Jr. 1961. Lichen Handbook. Smithsonian Inst. Press, Washington. 178 pp.
Hale, Mason E., Jr. 1967. The Biology of Lichens. Edw. Arnold, London. 176 pp.

Zopf, W. 1903. Vergleichende Untersuchungen iiber Flechten in Bezug auf ihre
Stoffwechselprodukte. Beih. Bot. Centralbl. 14: 95-126.

1969

THE MICHIGAN BOTANIST

175

STUDIES IN THE GENUS CORTINARIUS, I: SECTION
DERMOCYBE, CORTINARIUS AUREIFOLIUS COMPLEX

Joseph F. Ammirati, Jr., and Alexander H. Smith
Herbarium and Department of Botany,

The University of Michigan, Ann Arbor

INTRODUCTION

Throughout the genus Cortinarius color has always been important in
the classification of species. The colors of the pileus, stipe, lamellae, and con¬
text of the basidiocarp, as well as color changes in fresh material, are among
the characters used. More recently the type and location of the pigment has
been emphasized.

Taxa in the section Dermocybe are characterized by an innately fibril-
lose to squamulose, usually dry pileus, a dry stipe, and lamellae which are red,
orange, yellow, greenish, olive, or brown when fresh. Some examples are Corti¬
narius sanguineus Fries, C. chrysolitus Kauff., C. cinnamomeus (Fries) S. F.
Gray, and C. raphanoides Fries. The section is divided into smaller groups ac¬
cording to the color of the lamellae. This paper concerns some of the taxa
which have greenish, olivaceous, or yellow lamellae, including C. aureifolius Pk.

Taxa in the section Dermocybe have tramal hyphae and/or basidia
which often contain pigments occurring as granules or amorphous masses or as
diffuse color in or between the cells. These pigments are observed in dry spec¬
imens sections of which are revived and mounted in 2.5% KOH. The following
are a few of the taxa in which these pigments occur.

Cortinarius raphanoides has pale olivaceous-brownish lamellae when
fresh. Hyphae of the lamellar trama contain a diffuse pale pinkish to pale red¬
dish purple pigment and some red pigment-masses. The basidia contain red to
reddish purple pigment and some red pigment-masses. The basidia contain red
to reddish purple granules. In C. chrysolitus the lamellae are chrysolite green.
The tramal hyphae contain red to reddish purple pigment-masses and the
basidia contain a diffuse red to reddish purple pigment and similarly colored
granules. In both of these species the color and nature of the pigment appear
characteristic.

In Cortinarius cinnamomeus sensu lato the pigments are presently being
studied. So far, we have observed two types of pigments. In some collections
the pigment is yellow to yellow-orange in the tramal hyphae and basidia while
in other collections the pigment is red to reddish purple. These different pig¬
ments occur in collections which appear to be otherwise similar, e.g., in habit,
size, shape and roughness of spores, etc., but additional notes on fresh materi¬
al are needed to clarify the features of the basidiocarps.

Two types of pigments similar in color to those in C. cinnamomeus
sensu lato occur in the C. aureifolius complex. The following is a short intro¬
duction to the C. aureifolius complex, descriptions of the taxa, and a compari¬
son of the important characteristics including the pigments.

176

THE MICHIGAN BOTANIST

Vol. 8

Cortinarius aureifolius Complex

Cortinarius aureifolius was described by C. H. Peck in 1885 from ma¬
terial collected in Karner, New York. Since then it has only been collected
occasionally and so far has not been reported from Michigan. Basidiocarps of
this species are 4-5 cm high, grow in sandy soil under pines, and have colors
of the stipe, lamellae, and pileus similar to those of C. cinnamomeus.

During the fall of 1967 and 1968 a Cortinarius resembling C. aurei¬
folius was found at Fralick’s Tree Farm, Hell, Livingston County, Michigan.
Basidiocarps were growing under pines in sandy soil on bare or needle-covered
areas and were often buried in the sand to the pileus margin. In October 1968
at least one hundred were collected on a small sandy slope.

The type collection of C. aureifolius was studied and compared with
the Michigan material. Below are descriptions of the type collection of C.
aureifolius and of the Michigan material.

Cortinarius aureifolius Pk.

Pileus 1.7-3. 3 cm broad, convex, then plane or slightly depressed,
densely fibrillose-tomentose, sometimes slightly squamulose, especially on the
disk, cinnamon brown. Context yellow or pallid, odor like that of radishes.

Lamellae adnexed, yellow becoming yellow-cinnamon, rather broad,
moderately close, subventricose, rounded behind.

Stipe fibrillose, yellow, short, solid, equal, brownish within; plants
2.5-3. 8 cm high.

Spores (Fig. 1) 9.5-13.5(16) X 3.0-4.2/u, in side view obscurely elon¬
gate-inequilateral to subfusiform or fusiform-elliptic, in face view elliptic to
oblong-elliptic, smooth or the surface finely irregular; in KOH, single spores
dull light pale yellowish; in Melzer’s solution, single spores yellowish, groups
of spores light orange-yellow.

Basidia (23)27-40 X 5.5-Ijjl, broadly clavate, irregularly clavate, or elon¬
gate clavate, 4-spored, clamp connections present; in H2O hyaline to yellowish
or some containing yellow or reddish pigment; in KOH nearly hyaline or pale
pinkish to pale purplish, some containing deep red to reddish purple granules;
in Melzer’s pale yellow to golden yellow, some with ± refractive contents and
often containing yellow-orange, orange, or reddish orange granules. Basidioles
20-37 X 5.5-8/a, clamp connections present, similar in shape to basidia, in
H2O, KOH, and Melzer’s similar in color to basidia. No pleurocystidia seen.
Cheilocystidia scattered or in clusters, broadly clavate to vesiculose (difficult
to revive); lamellar edge fertile. Lamellar trama sub-parallel-interwoven; sec¬
tions in KOH pale reddish to pale pinkish purple or nearly hyaline, some
hyphae containing masses of dark red to purple-red pigment; sections in H2O
yellowish, some hyphae containing masses of reddish to reddish orange pig¬
ment; sections in Melzer’s yellowish to yellow-orange, some hyphae containing
masses of reddish orange to red pigment. Hymenium in KOH ± colored similar
to the lamellar trama; in H2O yellowish; in Melzer’s yellowish to yellowish
orange; when sections of lamellar trama are mounted in KOH a reddish pig¬
ment diffuses into the mount.

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177

Pileus cuticle of hyphae ± irregularly interwoven, up to 12-1 5/^ broad,
clamp connections present, hyphal end-cells rounded to clavate or tapered, oc¬
casionally broadly mucronate; sections in KOH pale “ochraceous tawny” to
brownish orange, some individual hyphae nearly hyaline; in Melzer’s yellowish,
yellow-orange to dull orange (“ochraceous orange”). Pileus trama composed of
interwoven hyphae; sections in KOH and Melzer’s similar in color to lamellar
trama, some hyphae containing masses of red to reddish purple pigment.

Hyphae on the surface of the stipe apex subparallel to interwoven,
hyphal end-cells not differentiated (basidia and basidioles from the hymenium
may occur at the immediate apex), clamp connections present; when sections
are mounted in KOH a reddish pigment diffuses into the mount; hyphae in
KOH nearly hyaline to pale pinkish or reddish (some surface hyphae yellow¬
ish), some hyphae containing dark red, (“scarlet red” or purplish red) pig¬
ment; hyphae in Melzer’s “lemon yellow” to “lemon chrome” or orange-
yellow, some hyphae containing golden yellow, orange, or reddish orange pig¬
ment. Surface hyphae from stipe base irregularly interwoven, hyphal end-cells
not differentiated, clamp connections present, some hyphae with slightly yel¬
lowish thickened walls; in KOH and Melzer’s similar in color to the surface
hyphae at the apex but hyphae with reddish contents less common.

Growing in sandy soil under pines. October, 1885, Karner, New York.
Collected by C. H. Peck.

R. Singer (F-1677) collected this species in Gainesville, Florida, Decem¬
ber 30, 1942, in sandy soil under pines. His field notes agree closely with the
description of Cortinarius aureifolius. The pileus was described as “snuff
brown” to “Saccardo’s Umber”; otherwise the coloration is similar to C. aurei¬
folius. Our microscopic examinations revealed scattered to abundant vesiculose
to broadly clavate cheilocystidia (13.5-25 X 8-12 n) which often form a sterile
lamellar edge. Cheilocystidia of this nature were present in the type of C.
aureifolius but did not form sterile lamellar edges as far as could be deter¬
mined (hymenial elements of the type were often difficult to revive). The
abundance of these sterile elements is apparently variable in this species.

Fig. 1 (left). Spores of Cortinarius aureifolius, X1200.

Fig. 2 (right). Spores of Cortinarius psammophilus, XI 200.

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Cortinarius psammophilus sp. nov. (Figs. 3 & 4).

Pileus 1-3.5 cm latus, conicus vel convexus, demum plano-convexus,
margine decurvus, fibrillosus demum squamulosus, brunneus, margine suboli-
vaceus vel luteolus; caro albida demum sulphurea; lamellae stramineae vel
luteae demum brunneae; stipes 1.5-4 cm longus, 3-7 mm crassus, stramineus
vel luteus, deorsum brunnescens; cortina lutea; sporae 10-13 X 3-4. 5ju, sub-
fusiformis, laevis. Typus: J. F. Ammirati, Jr. 2693 (MICH).

Pileus 1-3.5 cm broad, conic to convex becoming plano-convex, margin
incurved to decurved but rarely plane, surface moist to dry, not hygrophan-
ous, densely fibrillose to squamulose, “snuff brown,” “buckthorn brown,” or
“sayal brown” with “Isabella color” or “light brownish olive” tints toward the
margin, frequently tinted “wax yellow” on the outer margin, in some “bister”
to “chestnut brown” on the disc, when rubbed discoloring “dark vinaceous
drab” to “Mars brown,” becoming ± purplish brown when wet. Context 2-3
mm thick on the disc, thinner over the lamellae, solid, firm to spongy, whitish
to “pale yellow-green,” “naphthalene yellow,” or “sulphur yellow”; odor and
taste of radish or slight to none.

Lamellae sinuate to uncinate or subdecurrent to decurrent, close, 2-7
mm broad, at first “straw yellow,” “amber yellow,” or “wax yellow” to
“strontian yellow,” then “yellow ocher” to “Isabella color,” at length “snuff
brown” to “buckthorn brown,” some reddish spotted in age; when fresh red¬
dish with the application of KOH.

Stipe 1.5-4 cm long, apex 3-7 mm thick, base 4-10 mm thick, clavate
to slightly bulbous; surface appressed fibrillose or with ± scattered loose
fibrils, “barium yellow,” “straw yellow,” or “amber yellow” to “wax yellow,”
lower stipe discoloring “snuff brown,” “cinnamon,” “natal brown,” “bister,”
or rusty brown. Context “pale yellow green,” “sulphur yellow,” or colored as
the stipe surface, especially in the cortex, becoming sordid in age, stuffed to
hollow, cortex pliant. Cortina in young basidiocarps “barium yellow,” “amber
yellow” to “wax yellow.”

Spore deposit on white paper deep rusty brown (“amber brown” or
rusty “cinnamon-brown”); spores (Fig. 2) (9)10-13(15) X 3-4.5ju, shape some¬
times ± irregular, in side view obscurely elongate-inequilateral to subfusiform
or broadly subfusiform, in face view narrowly ovate, fusiform-elliptic or ellip¬
tic; in KOH near “ochraceous tawny” in groups, individual spores pale yellow¬
ish; in Melzer’s dull yellowish orange in groups, individual spores light yellow¬
ish.

Basidia 25-30(35-37) X 6-8jU, 4-spored, clavate, narrowly clavate or ir¬
regularly clavate, clamp connections present; in KOH hyaline, greyish or light
brownish, some containing yellow pigment as droplets or diffuse in the cell,
occasionally containing reddish granules; in H2O some containing bright yel¬
low pigment; in Melzer’s nearly hyaline to pale yellowish, greyish, or brown¬
ish. Basidioles 18-30(35) X (3-4)5-7ju, similar in shape to basidia, clamp con¬
nections present; in KOH, H2O, and Melzer’s similar in color to basidia. No
pleurocystidia seen. Cheilocystidia absent or when present scattered or in clus¬
ters, broadly clavate to vesiculose or less commonly cylindrical; lamellar edges

1969

THE MICHIGAN BOTANIST

179

fertile. Lamellar trama subparallel-interwoven; sections in KOH and H2O
hyaline to pale yellowish, some hyphae containing masses of bright yellow pig¬
ment; sections in Melzer’s light yellow to pale yellow, some hyphae containing
yellow pigment; hymenium in KOH and H2O hyaline to yellowish, in Melzer’s
± “ocher yellow”; when sections of lamellar trama are mounted in KOH a
reddish brown to reddish or pinkish pigment diffuses into the mount.

Pileus cuticle hyphae ± radially to irregularly interwoven, up to 12-1 4^
broad, clamp connections present, hyphal end-cells rounded to clavate, less
commonly tapered to broadly mucronate; when mounted in KOH a “tawny”
to “vinaceous brown” or light reddish brown pigment diffuses into the mount,
the hyphae finally light “tawny,” “ochraceous tawny” to light brownish or
some nearly hyaline; in Melzer’s “tawny,” “amber brown” to “hazel”; in KOH

Fig. 3. Cortinarius psammophilus, XL

Fig. 4. Cortinarius psammophilus, XL

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THE MICHIGAN BOTANIST

Vol. 8

some with yellowish thickened walls. Pileus trama composed of interwoven
hyphae; sections in KOH hyaline to pale yellowish, some hyphae containing
masses of pigment as in the lamellar trama; sections in Melzer’s pale yellow.

Hyphae on the surface of the stipe apex subparallel to interwoven, hyphal
end-cells not differentiated, clamp connections present, some hyphae en¬
crusted; when sections are mounted in KOH a reddish to reddish orange pig¬
ment diffuses into the mount; in KOH the hyphae hyaline to yellowish or
golden yellow to dull yellow-orange; in Melzer’s pale yellow to pale rusty yel¬
low. Surface hyphae from stipe base interwoven, hyphal end-cells not differen¬
tiated, clamp connections present, some hyphal walls encrusted; in KOH and
Melzer’s similar in color to surface hyphae at apex.

Gregarious to scattered or solitary under pines in sandy soil, basidio-
carps often partially buried in the sand. October, Fralick’s Tree Farm, Hell, Liv¬
ingston Co., Michigan. J. F. Ammirati, Jr. 1350, 1351, 1378, 1379, 2671,
2693 (type). Under pines, Waterloo Recreation Area, Washtenaw Co., Michi¬
gan, F. Hoseney 1183.

The elongate-subfusiform, ± smooth spores of C. aureifolius and C.
psammophilus are very unusual in the genus Cortinarius and unite these taxa
into a readily definable group. The spores are very similar in shape to those in
the genus Boletus.

The color of the pileus in C. aureifolius is more uniformly brown than
in C. psammophilus in which the margin is often tinted olivaceous and the
outer margin is commonly yellowish when the basidiocarps are fresh. In other
respects the colors of the basidiocarps when fresh are similar.

C. aureifolius and C. psammophilus differ in the color of the pigment
in the tramal hyphae and basidia (mentioned in the introduction). When sec¬
tions of the former are mounted in KOH the pigment masses in the hyphae of
the pileus and lamellar tramae and the pigment and/or granules in the basidia
are red to reddish purple. In C. psammophilus the pigment masses of the
tramal hyphae and the pigment in the basidia are yellow and reddish granules
are only occasionally seen in the basidia. Furthermore, in KOH the hyphae on
the surface of the stipe of C. aureifolius contain red to reddish purple pigment
while in C. psammophilus it is lacking.

The taxa in the Cortinarius aureifolius complex are clearly related by
their spores, habit, and ecology. Differences in pileus color and the color and
location of the pigment as described above form the basis for recognizing the
taxa.

Pigment differences in other taxa of the section Dermocybe when cor¬
related with other characteristics of the basidiocarps may prove important in
defining the taxa, especially in the Cortinarius cinnamomeus group.

ACKNOWLEDGMENTS

The authors wish to thank S. J. Smith, New York State Museum, for providing
material of the type of Cortinarius aureifolius Pk. In addition we wish to thank the Na¬
tional Science Foundation (Grant GB-6876X) for financial support during this study.

1969

THE MICHIGAN BOTANIST

181

THE BOTANICAL ACTIVITIES OF
THOMAS J. HALE, 1858-18621

Lytton J. Musselman
Department of Botany and Zoology,

University of Wisconsin, Rock County Center, Janesville

Thomas J. Hale is best known for his two papers entitled “Additions to
the Flora of Wisconsin” which appeared in the 1858 and 1860 volumes of the
Transactions of the Wisconsin State Agricultural Society (Hale 1858, 1860a).
The 1858 paper was a joint effort with Increase A. Lapham, who did the
mosses, liverworts, and lichens. Very little is known concerning Hale’s other
botanical activities. This paper comments on the activities of this important
plant collector.

Hale lists Racine, Wisconsin, as his address on many of his earlier labels
but I have found no information on where he was born or where he lived.
According to University of Wisconsin records he entered the classical course of
the university in 1857 at the age of 23. He was active in the Hesperian Soci¬
ety, a debating and literary society, and served as president from 1859 to
1860. Hale was apparently an excellent student with grades ranging from 94
to 96. He received his bachelor’s degree in 1860 and his master of arts degree
three years later.

About the time Hale received his B.A. the state geological and agricul¬
tural survey was formed. Hale began work in the survey as a field assistant
under the direction of Dr. Ezra Carr, professor of botany at The University of
Wisconsin, Madison. During the years 1860-1862 he worked for the survey as
a field assistant under Prof. James Hall, a geologist. His assignment was to
collect fossils throughout the state and to map certain sandstones, at the same
time collecting plants. During this period (1860-1862) he spent some of his
winters in Albany, New York, where Hall was working on a geological survey
of that state.

Hale’s letters provide an interesting comment on the activities of a field
botanist a century ago. His request for funds for the summer of 1860 was
$1.50 per day and $170.00 with which he outfitted himself with a horse,
wagon, and harness (Hale, 1860c). On July 15, 1860, he was in Whitewater
when he wrote Prof. Hall regarding his problem in finding lodging because so
many farmers mistook him for a peddler. “I have tried to stay at farm houses
several times but they all seem to think that I want to pay them in pins”
(Hale, 1860d). In several letters to Hall, Hale seems to dwell on his several
aches and pains which periodically plagued him (Hale, 1860e, 1860f). Once
while suffering from the ague he commenced the use of gentian, perhaps the
ague weed, Gentianella quinquefolia (Hale, 1860b).

^Presented at the Botanical Club of Wisconsin section of the 99th meeting of the
Wisconsin Academy of Sciences, Arts, and Letters at Whitewater, May 3, 1969.

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One of Hale’s greatest problems as a field botanist was the preparation
of good specimens. He frequently excused his poor specimens when writing
other botanists. He wrote Daniel Cady Eaton stating that “I have so little time
to spare from my geology that I necessarily do my work in haste” (Hale,
1860j). Later he wrote that he must acknowledge his inability to make fine
specimens while journeying with night air only to dry his papers (Hale,
1861c). These poor specimens irked at least one of his botanical correspond¬
ents, the eminent Kentucky botanist, Dr. Charles Wilkins Short, who wrote
Increase Lapham saying that Mr. Hale had sent him some interesting collec¬
tions from Wisconsin but that Hale did not take the pains which Lapham did
to “make his specimens fine” (Short, 1862).

Hale established correspondence with most of the leading botanists of
his day. In 1859 he wrote Lapham asking for the addresses of botanists in
various parts of the country as he has “no botanical acquaintance” (Hale,
1859). Two years later he wrote D. C. Eaton asking for the addresses of addi¬
tional botanists although he was already corresponding or exchanging speci¬
mens with an illustrious list of botanists including Asa Gray, Samuel Barnum
Mead, [C. J.?] Sprague, George Vasey, Elias Durand, M. S. Bebb, [T. C.?]
Porter, George Engelmann, Moses A. Curtis, [J. W.?] Chickering, Stephen
Thayer Olney, Reuben Nevius, Wm. M. Canby and “many lesser lights” (Hale,
186 Id).

To distribute specimens to so many correspondents required the collec¬
tion of large numbers of plants. By the end of the collecting seasons Hale had
several hundred species, apparently with duplicates, to trade. In 1860 he col¬
lected enough Asclepias lanuginosa , now rare in southern Wisconsin, to “satis¬
fy every American botanist” (Hale, 1860c). (This species was described in his
1858 paper as a new species, Acerates monocephala Lapham “unless it should
be united with A[ cerates] lanuginosa Decaisne”). He mentions collecting such
“rare” plants as Agoseris cuspidata, Castilleja sessiliflora, and some grasses in
quantity for trade (Hale, 1860g). He offered George Engelmann “several rare
species in quantity” (Hale, 1860h). Hale also collected many mosses but felt
constrained to give up their study in order to concentrate on rocks and herb¬
aceous plants. He asked for the address of the moss expert William Starling
Sullivant in order to offer him his collection (Hale, 1860j). He engaged the
services of a “young man at Madison” to collect plants common in that area
for trade (Hale, 1861a). This may be a reference to Samuel H. Watson, then a
student at the University of Wisconsin (Musselman, 1969).

Although best known for his plant collections in Wisconsin, Hale also
collected a few specimens in Minnesota, Iowa, and Illinois. Some representa¬
tive labels of T. J. Hale are shown in Fig. 1. The localities in which he col¬
lected are mapped in Fig. 2. From his letters it is also evident that Hale did
geological work in many areas from which I have seen no plant specimens.

Hale’s collections are of interest because many plants he collected have
not or rarely been seen in these areas since he collected them. For example,
he sent a Senecio to Gray in 1861 which Gray determined to be Senecio
palustris (S. congestus), a northern species which Hale had collected in south-

1969

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183

ern Wisconsin at Madison (Hale, 1860i). Hale’s specimen is not in the Herbari¬
um of The University of Wisconsin, Madison, but the specimen he sent to
Gray may still be in the Gray Herbarium. In 1860 he collected Ruellia humi-
lis (wild petunia) at Beloit where it has only been collected once since.2 Ap¬
parently many of Hale’s specimens survive in herbaria which contain the col¬
lections of Hale’s correspondents. If Hale kept a collection of his own, I have
not been able to locate it.

Hale found plants which he assumed to be new species and sent them to
Gray, Engelmann, and Eaton to be named. He took particular interest in the
genus Euphorbia as he was trying to “work out the genus in the state” (Hale,
1861c). Hale also took an interest in the carices, mosses, and the gerardias. In
1861 he sent Gray a parcel of plants which Hale assumed to have several un¬
named species. He left Gray to describe the new species “trusting that you
will do me such honors as I have deserved, if any of the plants should prove

T. J. HALE,

RACINE, WISCONSIN.

"WMv g" v*,l 1 nr1 I

Fig. 1. Representative labels of T. J. Hale: an original label apparently from Hale’s own
collection and an exchange label.

2“ Ruellia ciliosa Pursh, Beloit, Aug. 30, 1894, G. B. Olds.” In the Beloit College
Herbarium.

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THE MICHIGAN BOTANIST

Vol. 8

to be not described” (Hale, 1861b). Included with this group of plants was a
Gerardia which Gray later named Gerardia tenuifolia var. asperula (Gray,
1879).

Hale’s botanical career ended abruptly in 1862 when the legislature
abolished the state geological and agricultural survey. Hale had fallen out with
James Hall and left Milwaukee hastily while teaching a summer course (Hale,
1862). He apparently died before 1879 because Gray, when describing the
Gerardia which Hale had sent, refers to him as the “late Mr. Hale” (Gray,
1879).

ACKNOWLEDGMENTS

Portions of this work completed during the summer of 1968 were incidental to
research supported by a University of Wisconsin Center System Summer Research grant.
This work would not have been possible without the generous assistance of many librari¬
ans and curators who kindly searched manuscript collections for letters of Hale. Special
credit is due J. E. Boell, director of the archives of the University of Wisconsin, for ob¬
taining information of Hale’s university activities and also to Margaret Gleason of the
State Historical Society of Wisconsin for checking several federal census reports.

REFERENCES CITED

Gray, A. 1879. Gerardia tenuifolia, Vahl, var. asperula. Bot Gaz. 4: 189.

Hale, T. J., & I. A. Lapham. 1858. Additions to the flora of Wisconsin. Wis. St. Agr. Soc.
Trans. 5: 417-424.

Hale, T. J. 1859. Letter to Increase A. Lapham, Sept. 20. The State Historical Society of
Wisconsin, Madison.

- . 1860a. Additions to the flora of Wisconsin. Wis. St. Agr. Soc. Trans.

6: 259-263.

- . 1860b. Letter to James Hall, May 25. New York State Library, Albany.

- . 1860c. Letter to D. C. Eaton, June 4. Yale University Library, New

Haven, Conn.

- . 1860d. Letter to James Hall, July 15. New York State Library, Albany.

- . 1860e. Letter to James Hall, Aug. 10. New York State Library, Albany.

Fig. 2. Map showing collection sites of T. J. Hale in Wisconsin, Minnesota, Iowa, and
Illinois, compiled from labels in the Milton College Herbarium (MCW). Numbers corre¬
spond to localities as listed below:

Wisconsin

1. St. Croix Falls

2. St. Croix County

3. St. Croix River

4. Lake Pepin

5. Black River Falls

6. Sturgeon Bay

7. Green Bay

8. De Pere

9. Kaukana

10. La Crosse

11. Lamartine

12. Kilbourn City

13. Sauk City

14. Hartford

15. Arena

16. Black Earth

17. Madison

18. Lake Mills

19. Waukesha

20. Milwaukee

21. Racine

22. Kenosha

23. Burlington

24. Beloit

25. Fulton

26. Blue Mounds

27. Dodgeville

28. Benton

29. Platte Mounds

Minnesota

30. St. Paul-Minneapolis Area

31. Dakota County

32. Cannon River

33. Fillmore County

Iowa

34. Dubuque

Illinois

35. Freeport

1969

THE MICHIGAN BOTANIST

185

- . 1860f. Letter to James Hall, Oct. 27. New York State Library, Albany.

- . 186Qg. Letter to ?, Nov. 28. Yale University Library, New Haven, Conn.

(In the D. C. Eaton papers.)

- . 1860h. Letter to George Engelmann, Dec. 3. Missouri Botanical Garden

Library, St. Louis.

- . 1860i. Letter to D. C. Eaton, Dec. 13. Yale University Library, New

Haven, Conn.

- . 1860j. Letter to D. C. Eaton, Dec. 17. Yale University Library, New

Haven, Conn.

- . 1861a. Letter to D. C. Eaton, Jan. 5. Yale University Library, New Haven,

Conn.

- . 1861b. Letter to Asa Gray, Nov. 14. Yale University Library, New Haven,

Conn. (In the D. C. Eaton papers.)

- . 1861c. Letter to George Engelmann, Nov. 15. Missouri Botanical Garden,

St. Louis.

- . 186 Id. Letter to D. C. Eaton, Dec. 13. Yale University Library, New

Haven, Conn.

- . 1862. Letter to Increase Lapham, July 3, The State Historical Society of Wis¬
consin, Madison.

Musselman, L. J. 1969. Samuel H. Watson, pioneer botanist of southern Wisconsin. Mich.
Bot. 9: 35-37.

Short, C. W. 1862. Letter to Increase Lapham, March 24. The State Historical Society of
Wisconsin, Madison.

186

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Vol. 8

Bessie Bernice Kanouse (November 21, 1889— January 28, 1969)

(About 1930)

1969

THE MICHIGAN BOTANIST

187

DR. BESSIE BERNICE KANOUSE- 1889- 1969

Alexander H. Smith

Herbarium, The University of Michigan, Ann Arbor

Bessie Bernice Kanouse was born at Quincy, in Branch County, Michi¬
gan, November 21, 1889. She attended the secondary schools of Quincy, and
later taught in the public schools of the state from 1908 to 1912. In 1915 she
received the bachelor’s degree from Eastern Michigan University, which at that
time was known as the State Normal College at Ypsilanti, Michigan. From
1915 to 1917 she taught nature study in the public schools of Gary, Indiana,
and from 1917 to 1921 was supervisor of general science for 33 public
schools at Camden, New Jersey. In 1921 she enrolled at the University of
Michigan, where she received her A.B. in 1922, her M.S. in 1923 and her
Ph.D. in 1926. She was appointed as Curator in the University Herbarium in
1926, and assistant to the Director in 1928, positions she held until her retire¬
ment in 1960. She continued to reside in Ann Arbor for a number of years
following her retirement, and then moved to Coldwater, Michigan, where she
passed away January 28, 1969.

Because of her long association with natural history as a hobby, then as
a teacher, and finally as the one responsible for the care of one of the coun¬
try’s important research collections of fungi, it is not at all surprising that she
was one of the founders of the Michigan Wildflower Association— which even¬
tually became the Michigan Botanical Club. At the organizational meeting May
31, 1941, she was elected as the first secretary-treasurer of the Wildflower As¬
sociation.

Dr. Kanouse’s scientific work was centered in the fungi, where she did
important research both on watermolds and on Discomycetes, on which sub¬
ject she was a frequent contributor to the Papers of the Michigan Academy of
Science, Arts, and Letters. She belonged to the Michigan Academy; Mycologi-
cal Society of America; Michigan Parks Association; Michigan Natural Areas
Council; Women’s Research Club of the University of Michigan; Washtenaw
Historical Society, of which she was a life member; the Midwest Museums So¬
ciety, of which she was a charter member and later an honorary member; the
National Society of the Women of New England and a member of the Ann
Arbor Colony; Sigma Xi; American Association for the Advancement of Sci¬
ence (fellow); and the National Society of the Daughters of the American
Revolution. She also served on the board for the Wesley Foundation, and was
a patroness for Phi Beta Phi.

Dr. Kanouse was deeply interested in philosophy and American litera¬
ture, and when no longer able to engage in the strenuous field work of col¬
lecting fungi turned to reading, and also tried hei hand at writing. She was an
artist at heart, as well as a scientist. At the University she will be remem¬
bered for her efforts to build up the fungous collections, and although she did
not teach classes, will be remembered by many graduate students for the help
and advice she gave them.

188

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BIBLIOGRAPHY OF BESSIE BERNICE KANOUSE

The life-history of a new homothallic Mucor. Pap. Mich. Acad. 3: 123-129. 1924.
Physiology and morphology of Pythiomorpha gonapodioides. Bot. Gaz. 79: 196-206.

1925.

On the distribution of the water molds, with notes on the occurrence in Michigan of
members of the Leptomitaceae and Blastocladiaceae. Pap. Mich. Acad. 5: 105-114.

1926.

A monographic study of special groups of the water molds I. Blastocladiaceae. Am. Jour.
Bot. 14: 287-306. 1927.

A new species of the genus Pythium in the subgenus Aphragium. Pap. Mich. Acad. 7:
129-139. 1927. (with Tryphena Humphry).

A physiological and morphological study of Saprolegnia parasitica. Mycologia 24:
431-452. 1932.

Notes on new or unusual Michigan Discomycetes. I. Pap. Mich. Acad. 19: 93-106. 1934.
Notes on new or unusual Michigan Discomycetes. II. Pap. Mich. Acad. 20: 65-78. 1935.
Notes on new or unusual Michigan Discomycetes. III. Pap. Mich. Acad. 21: 97-104. 1936.
Notes on new or unusual Michigan Discomycetes. IV. Pap. Mich. Acad. 22: 117-122.

1937.

Notes on new or unusual Michigan Discomycetes. V. Pap. Mich. Acad. 23: 149-154.

1938.

Studies of two species of Endogone in culture. Mycologia 28: 47-62. 1936.

Songs to Birchwood. 63 pp. Privately published, Ann Arbor, 1939.

Notes on new or unusual Discomycetes. Pap. Mich. Acad. 24: 25-29. 1939.

Two new genera of Discomycetes from the Olympic National Forest. Mycologia 32:

756-759. 1940. (with Alexander H. Smith).

New and unusual species of Discomycetes. Mycologia 33: 461-467. 1941.

Doctor Howard Atwood Kelly. Mycologia 35: 383-384. 1943.

A new Discomycete from the Olympic National Forest. Mycologia 36: 460-463. 1944.
Some studies in the genus Helvetia. Pap. Mich. Acad. 32: 83-90. 1948.

A survey of the Discomycete flora of the Olympic National Park and adjacent areas.
Mycologia 39: 635-689. 1947.

The genus Plectania and its segregates in North America. Mycologia 40: 482-497. 1948.
Studies in the genus Otidea. Mycologia 41: 660-677. 1949.

A study of Peziza bronca Peck. Mycologia 42: 497-502. 1950.

Some species of the genus Trichophaea. Mycologia 50: 121-140. 1958.

1969

THE MICHIGAN BOTANIST

189

A SEEDLING POPULATION OF ASPENS IN
SOUTHEASTERN MICHIGAN

Gary E. Andrejak and Burton V. Barnes
School of Natural Resources and Botanical Gardens,

The University of Michigan, Ann Arbor

The North American aspens, Populus tremuloides Michaux (trembling
aspen) and P. grandidentata Michaux (bigtooth aspen) typically occur in
clones, aggregations of genetically identical stems (ramets), produced asexually
by root suckering (Barnes, 1966). Disturbances such as fire, logging, and graz¬
ing favor the development of sucker stands (Baker, 1925; Fetherolf, 1917;
Sampson, 1919; Weigle and Frothingham, 1911).

Most of the aspens in existence today were probably derived asexually
and did not arise directly from seedlings as is typical of most forest trees.
Graham et al. (1963) conducted extensive studies in aspen stands in the Lower
Peninsula of Michigan and failed to find a single seedling stand of merchanta¬
ble size. A similar predominance of sucker stands was found in northern
Minnesota (Kittredge & Gevorkiantz, 1929).

Although a clone must originate from a seedling (the ortet), establish¬
ment of seedlings is very difficult and seedling stands are rare (Barnes, 1966).
Seedlings are rare because a favorable combination of biotic and environment¬
al conditions is necessary for establishment. However, during investigations of
hybrids between the two species of aspen in southeastern Michigan (Barnes,
1961) many putative seedling hybrids were observed. They were numerous
along the edges of abandoned fields particularly where the field was adjacent
to a woodlot. The discovery of such a natural population growing in an
abandoned portion of a cultivated field provided the opportunity to study the
structure, origin, and development of a stand markedly different from most
present-day aspen stands in Lower Michigan.

The objectives of the study were to determine (1) the number of seed¬
lings, i.e., different genotypes, in the population, (2) the origin of the popula¬
tion, and (3) the incidence of hybridization.

THE RESEARCH AREA AND METHODS

The population under study, about 0.17 acre in size, is located along the
west and south edges of an abandoned portion of a cultivated field adjacent
to Stinchfield Woods, a forest property of The University of Michigan (Fig.
1). The elevation is approximately 990 feet. The soil type is a Boyer sandy
loam. Second-growth stands of oak and hickory, predominantly Quercus
velutina Lam., Q. rubra L., Q. alba L., Carya ovata Mill., and C. ovalis Wang.,
surround the population to the west, north, and south. Clones of bigtooth
aspen were found in the stands north and east of the population.

The history of the establishment of the population was determined with
the help of the original General Land Office Surveys, various settlement his¬
tories of Washtenaw County, aerial photographs, and personal communication

190

THE MICHIGAN BOTANIST

Vol. 8

OAK -HICKORY

no

Oil I

113*2

I

I

I

I

l

l

Fig. 1. Map of the research area showing the location of P. tremuloides (#), P. grandi-
dentata (O), and hybrid individuals (©). The road forms the northeastern border of lot
59 of Stinchfield Woods, forest property of the University of Michigan, Washtenaw Co.,
Michigan.

1969

THE MICHIGAN BOTANIST

191

with Mr. Frank Murray, former manager of the forest properties of the Uni¬
versity of Michigan.

All aspen stems 2 inches in diameter at breast height (dbh) and larger
were numbered and mapped. The dbh of each stem was measured with a di¬
ameter tape and the height was measured with an Abney level. Increment bor¬
ings of several of the larger individuals were taken to determine their age.

Vertification of the seedling origin of each stem would have necessitated
observing seed fall, germination, and subsequent development of the plant.
Seedlings might also be identified by root excavations since the root system of
a seedling is distinguishable from that of a sucker, at least initially. Although
we were unable to use these methods, we adopted a method which approxi¬
mates the number of seedlings by determining the number of different geno¬
types (clones) in a given stand. Of the total number of stems, the greater the
number of different clones represented, the greater the number of seedlings
established. For example, if a population of 100 stems were all similar in
morphology and phenology, we would consider them as 100 ramets of 1 clone
or possibly 1 seedling ortet and 99 ramets. If 80 of the 100 stems were differ¬
ent in their characters, i.e., represented different genotypes, a predominantly
seedling stand would exist. Obviously, various combinations of seedlings and
multi-stemmed clones may exist in a given stand due to establishment condi¬
tions and subsequent disturbance. This method tends to underestimate the
number of seedlings, particularly if there are many stems involved. Two or
more genotypes may have some similar characters and time is usually not suf¬
ficient to study all possible differentiating characters.

Following the guidelines formulated by Barnes (1969), the characters
used to distinguish different clones were as follows:

1. Taxon: P. grandidentata, P. tremuloides, or P. grandidentata X P.
tremuloides.

2. Sex: 6, 9, or

3. Leaf morphology: Overall shape and size, blade base shape, blade
tip shape, petiole length, number of teeth per side of blade.

4. Phenology: time and stage of flowering, time and stage of leaf
flushing, time of leaf drop, autumnal leaf color.

From all numbered stems, 4 to 6 shoots bearing “early” leaves were col¬
lected in June, 1967, after the foliage had expanded fully. The shoots, 1.5 to
3 in. long, were taken from inside lateral branches in the middle and lower
portions of the crown. Five undamaged leaves from the middle portion of the
shoots of each tree verified to be a different genotype were selected for meas¬
urement. Two leaves at most were taken from any 1 shoot. The following
measurements were made on each leaf:

1. Total blade length, cm.

2. Maximum blade width, cm.

3. Petiole length, cm.

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THE MICHIGAN BOTANIST

Vol. 8

4. Number of teeth per leaf side.

5. Total length of shoot from which the leaf was taken, cm.

6. Distance from the bud scar at base of shoot to the point of leaf
attachment, cm.

Leaves of selected individuals were mounted and photographed.

In March, 1967, before buds began to swell, 10 to 20 terminal buds
were taken from each tree. These were then examined for amount of pubes¬
cence on the bud scales. Stage of flowering was observed on 3 different dates
in 1967 and periodically in 1968.

Before pollen discharge began, branches of male individuals bearing flow¬
er buds were taken to the greenhouse of The University of Michigan Botanical
Gardens and placed in containers of water. Pollen was collected 4 to 8 days
later. The pollen was stored in a refrigerator and later examined to determine
percentage of abortion and diameter of grains. The pollen of each tree was
mounted on a glass slide and stained with aceto-carmine. At each of 10 ran¬
dom positions on the slide, the diameters of 10 grains were measured. At each
position all grains in the field of view were counted and the percentage of
aborted grains, those collapsed and lacking cytoplasm, was computed. The
average number of grains observed per clone was 275.

After pollination had taken place in the spring of 1967, a branch from
each female individual bearing fruit was taken to the greenhouse, and placed
in a container of water. The seeds were subsequently collected. After about 2
weeks’ storage in a refrigerator, seeds of each female tree, with 4 exceptions,
were counted into 3, 50-seed lots and placed on moist filter paper in petri
dishes. Germination was observed daily and, at the end of 5 days, total germi¬
nation was recorded.

To facilitate comparison of the 2 aspen species with their putative hy¬
brids, a hybrid index was devised using 6 leaf and bud characters. Coefficients
of variation were computed to compare the variability of leaf traits and pollen
diameter of the taxa.

RESULTS AND DISCUSSION
HISTORY OF ESTABLISHMENT

At the time of settlement by white men, Washtenaw County was prima¬
rily covered by a hardwood forest. General Land Survey notes of 1819 indi¬
cate that the area where the research site is now situated was “very hilly oak
land.” In pre -settlement forests, aspens were probably uncommon and incon¬
spicuous trees, as they apparently were in other parts of the Lake States
(Barnes, 1966). However, on sites disturbed by fire or Indian cultivation, they
probably were more frequent in occurrence. The Land Survey notes show that
aspens were occasionally used as witness trees in swamps and along the edges
of lakes, rivers, and streams. The surveyors used “aspen” in their notes and
distinguished it from cottonwood or “poplar.” Aspens made up 2.2% of the
total number of species mentioned in Washtenaw County. The closest aspen

1969

THE MICHIGAN BOTANIST

193

cited was located approximately 1 mile east of the research area, where the
section line crossed the Huron River.

The land on which the stand is located was probably cleared some time
before 1880. The area that is now Stinchfield Woods was grazed by sheep
until about 1935, when the University of Michigan began purchasing the sec¬
ond growth oak-hickory stands. In 1953, the University purchased a corner of
the field, 0.57 acres, and during the summer of that year constructed a road
giving better access to hardwood stands east and north of the field. Aerial
photographs taken in 1940 and information provided by Mr. Frank Murray
indicate that, prior to the time of road construction, no aspens existed where
the population grows today; that area was then part of the cultivated field.

Evidence from increment borings and aerial photographs indicates that
aspens probably became established during the 1953 or 1954 growing season.
Seeds probably came from female clones growing along the perimeter of the
field to the north. When the road was constructed across the corner of the
field, the end of the field was narrowed so that it became difficult to maneu¬
ver equipment for plowing. Thus, in 1954, a strip of ground at the end of the
field was not planted although it may have been plowed. The abandonment of
this portion of the field and the process of road building provided exposed
mineral soil, a highly desirable seedbed for aspen seedlings. The weather rec¬
ords of 1954 show that June was above average in temperature and was the
second wettest June in Michigan on record up to that time (U. S. Dep. Com¬
merce, 1954). Therefore, the seedbed and moisture conditions were very fa¬
vorable for the establishment of aspen seedlings.

THE SEEDLING STAND

Of the 78 stems numbered and mapped, 54 (or 69% of the total) were
conclusively verified as different genotypes: 35 of P. grandidentata, 14 of P.
tremuloides, and 5 of putative Fj hybrids. The remaining 24 stems could not
be positively distinguished from the other clones. However, none of these in¬
dividuals could be definitely identified as a ramet of one of the 54 clones. It
is probable that more study would reveal that many of these also represent
different clones and thus would increase the percentage of sexually produced
individuals.

Generally, each of the 54 stems constituted what might be called “single¬
tree clones,” presumably the ortet only— a seedling plant. Along the edges of
the field adjacent to the population, suckers from one or two of the ortets
have begun to appear where cultivation has severed or injured roots. Apparent¬
ly, only in this area are the typical multi-stemmed clones currently being initi¬
ated. The finding of no signs of other disturbance and that the small stand is
even-aged, support the conclusion that the remaining 24 stems also may be
seedlings.

Reports of seedling aspens are infrequent. In the Great Basin region of
Utah, Baker (1925) suggested that conditions for seedling establishment last
occurred in the late Pleistocene. Similarly, Cottam (1954) stated that P.
tremuloides ceased reproduction by seed in Utah at the onset of the Post-

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Vol. 8

pluvial climate, approximately 8,000 years ago. In the eastern United States, a
preponderance of sucker stands has been noted (Graham et ah, 1963; Kit-
tridge & Gevorkiantz, 1929). Although it is obvious that seedlings must have
been established before multi-stemmed clones could originate, reports of recent
seedling establishment are also rare. Larson (1944) reported several hundred
established seedlings along the shoreline of a receded reservoir in Utah. Barnes
(1966) discovered numerous aspen seedlings on recently burned sites in north¬
ern Idaho. Thus, seedlings can become established in certain areas of the West
where both climatic and soil conditions are suitable.

In the East, Faust (1936) discovered seedlings established along a re¬
ceded reservoir; and many regeneration surveys after fire have indicated that
aspen “seedlings” were among the invading species (Ahlgren, 1959; Gauvin,
1964; MacArthur, 1964; Mills, 1961; Vogl, 1969). In Michigan, Graham et al.
(1963) reported several seedling populations that arose after fire or logging, and
in one instance in the seedbeds of a forest nursery.

Evidence of seedling establishment in northern Lower Michigan was indi¬
rectly provided by Bertenshaw (1965) who found that on a given acre the
number of clones ranged from 26 (average of 15 ramets per clone) to 174
(average of 5 ramets per clone). Because of periodic fires in the area, it is
unlikely that any of the trees on these acres are the original seedlings. How¬
ever, the evidence does indicate that many seedlings were established some¬
time in the recent past.

It is not surprising that seedling aspens are uncommon. First, seedling
establishment is difficult and only occurs through the favorable combination
of many environmental and biotic factors. Second, aspens sucker profusely
following disturbance, and a seedling stand may be transformed quickly into a
sucker stand. Suckers may arise as soon as the second year on the root system
of a seedling (Day, 1944). Sucker-promoting disturbances such as fire, logging,
and grazing are very common throughout the ranges of the aspens.

Although establishment of seedlings is difficult, they are probably much
more frequent than published reports indicate. Reports of seedlings are infre¬
quent because of the above reasons and also because it is very difficult to
establish that a given stem is a seedling unless the tree is young. Root connec¬
tions decay and suckers establish independent root systems (Barnes, 1966;
DeByle, 1964). For these reasons determining the proportions of different
genotypes to the total number of stems in a population is a more feasible
method to resolve the number of seedlings established than root excavations.
However, it cannot pinpoint the date of origin of the clones without addi¬
tional information of seedfall, seedbed, and other environmental conditions.

CHARACTERISTICS OF THE POPULATION
Height , Diameter, and Growth

Considerable variation among the 54 verified seedlings was found in the
growth attributes of each taxon (Table I). The hybrids slightly exceeded the
parent species in dbh, total height, and annual growth. Although the differ-

1969

THE MICHIGAN BOTANIST

195

Table I. Mean dbh, height, and growth rate for P. tremuloides, P. grandidentata, and their
hybrid in the population studied. Age approximately 13 years.

No. of Mean Annual

Taxon

Individuals

DBH

Mean Range

Total Height
Mean Range

Growth

Diameter

Height

-in-

-ft-

in

ft

P. tremuloides

14

3.5 2.0-4. 7

29.5

24.0-37.0

0.24

2.4

P. grandidentata

35

4.5 2.2 -6.4

34.3

17.0-42.0

0.32

2.6

Hybrid

5

4.6 3. 7-5. 5

35.3

30.0-41.5

0.33

2.7

ences were negligible and only a few hybrids were present, it was apparent
that on this site hybrids could become established and compete successfully
with both parents to an age of approximately 13 years.

Sex and Hermaphroditism

The ratio of males to females for trembling aspen was 7:5; 2 additional
clones were hermaphrodites. The corresponding ratio for bigtooth aspen was
17:14. Four clones did not flower. Three of the 5 hybrids were female and 2
were hermaphroditic. Of the latter 1 was predominantly female and 1 was pre¬
dominantly male. Although aspens are reported to be dioecious, it is well
known that many clones are bisexual (Erlanson & Hermann, 1928; Lester,
1961; Pauley & Mennel, 1957; Santamour, 1956). Einspahr (1960) reported
that the incidence of the bisexual condition in wild populations of trembling
aspen may range from 4% to 20%. Following the terminology of Westergaard
(1958), both parents and the hybrid should be considered “subdioecious,” i.e.,
some bisexual plants occur regularly in nature besides exclusively male and
female plants.

Leaf Flushing

The average date of leaf flushing (bursting of buds and expansion of
leaves in the spring) of trembling aspen was approximately 8 days earlier than
that of bigtooth aspen. In northern Lower Michigan, Barnes (1969) found the
difference between the 2 species to range from 2 to 4 weeks depending on
site conditions. Differences in the initiation and progress of flushing for differ¬
ent stems of trembling aspen were conspicuous. In 1967, the earliest trembling
aspen clone began flushing on April 16 and the latest on May 6, a difference
of approximately 3 weeks. In northern Lower Michigan a similar observation
was made for trembling aspen (Barnes, 1969). Although bigtooth aspens were
more uniform in the initiation of flushing, notable differences appeared as
flushing progressed. The flushing of hybrid individuals most nearly resembled
trembling aspen in initiation and sequence.

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Leaf and Bud Characteristics

As expected, marked variations in leaf shape, size, margin, base, and tip
were observed in both species and the hybrids. The leaf margin is a very good
trait by which the parents may be differentiated and hybrids distinguished
from the parents (Fig. 2). Terminal buds of trembling aspen were glabrous
except for minute hairs on the basal scales of some individuals. Bigtooth aspen
was characterized by dense pubescence on the central portion of each scale;
pubescence was sparse along the margins of the scales. The hybrids lacked the
extremely dense pubescence of bigtooth aspen, but had instead only relatively
dense pubescence on the lower scales and numerous scattered hairs on the up¬
per scales.

Flowering

Trembling aspen began shedding pollen before bigtooth aspen. The mean
first flowering date separating trembling and bigtooth aspen in Ontario is ap¬
proximately 8 days (Bassett et al., 1961). No comparable data are available
from this study. Although trembling aspen flowered first, at least 4 days be¬
fore the earliest bigtooth aspen, considerable overlap was observed. For ex¬
ample, receptive female flowers of trembling aspen, as judged by fully ex¬
tended and brilliant red stigmas, were present as late as April 8, 1967, well
into the an thesis period of bigtooth aspen (March 26 to April 16).

HYBRIDIZATION

To determine the relationship of the putative hybrids to the parent spe¬
cies, 5 leaf characters (blade width, blade length, ratio of blade width to blade
length, petiole length, number of teeth per leaf side) and the pubescence rat¬
ing of terminal buds were selected and the mean, standard deviation, and coef¬
ficient of variation determined (Table II). The mean values of the hybrids
were approximately mid-way between the means of the parents except for
blade width and petiole length, in which they were closer to trembling aspen.
There was considerable overlap between the hybrids and trembling aspen in

Table II. Variation of 5 leaf characters of P. tremuloides, P. grandidentata, and their hybrid
in the population studied. Means (m), standard deviation (s), and coefficient of variation (cv).

Blade

Petiole

No. of Teeth

Taxon Width

Length

Width/Length

Length

Per Leaf Side

m s cv

m s cv

m s cv

m s cv

m s cv

-cm- %

-cm- %

%

-cm- %

%

P. tremu-

loides 6.3 0.88 14.0

6.0 0.72 11.9

1.0 0.09 8.6

4.8 1.14 23.8

31.4 4.52 14.4

Hybrid 6.7 0.86 12.8

8.0 0.45 5.6

0.8 0.08 9.5

5.4 0.60 11.2

20.2 2.07 10.2

P. grandi-

dentata 8.5 0.72 8.5

9.3 0.72 7.8

0.9 0.07 7.6

6.9 0.94 13.6

9.9 1.30 13.1

1969

THE MICHIGAN BOTANIST

197

Fig. 2. Silhouettes showing leaf size and margin differences of representative individuals
of P. tremuloides, P. grandidentata, and their hybrid in the population studied. Scale:

X0.6.

198

THE MICHIGAN BOTANIST

Vol. 8

these characters (Andrejak, 1968). Similar findings were reported by Barnes
(1961). Trembling aspen was the most variable species in all characters. Big-
tooth aspen was the next most variable with the exception of blade width.

Two hybrid indices were computed to examine the relationship of the
hybrids to the parents; one utilized 5 leaf characters and the pubescence rat¬
ing; the other used the 3 best distinguishing characters between the parents:
blade length, number of teeth per leaf side, and pubescence (Fig. 3). Both
indices showed the hybrids approximately mid-way between the parents. From
this and observation of other leaf traits, we conclude the intermediate plants
are Fj hybrids. Although the hybrids were similar to trembling aspen in peti¬
ole length and blade width, we believe it is unlikely that these individuals are
backcrosses or introgressants.

HYBRID INDEX

Fig. 3. Graph of frequency distribution of hybrid index using 3 characters. P. tremu-
loides (+) 14 plants, P. grandidentata (x) 35 plants, and hybrid (*), 5 plants. (Graph was
prepared by a CalComp Plotter from a Fortran IVG program in the Michigan Terminal
System.)

1969

THE MICHIGAN BOTANIST

199

Hybrids between the species may be produced by controlled pollination
with ease (Heimburger, 1936; Pauley, 1963). Natural hybrids have been re¬
ported from Massachusetts (Pauley, 1956), Quebec, Canada (Victorin, 1930),
and the Lake States (Barnes, 1961; Einspahr, 1960). Barnes (1961) found
numerous hybrids in southeastern Michigan in contrast to their rarity in north¬
ern Lower Michigan. He estimated they might represent from 1 to 5% of the
clones in the areas studied. The reasons postulated for the occurrence of
hybrids were an overlap in flowering time, and favorable climatic and seedbed
conditions.

Although P. tremuloides usually flowers in advance of P. grandidentata,
this isolating mechanism is not complete. Barnes (1961) and Einspahr and
Joranson (1960) reported that intraclonal variation in both species may some¬
times be great enough to cause the flowering times to overlap substantially.
Also, Pauley (1956) suggested that temperature inversions in some areas may
retard the flowering of female P. tremuloides to coincide with the pollen dis¬
charge of P. grandidentata .

Observations of the Stinchfleld Woods population show that considera¬
ble overlap in flowering exists. In 1967, there was a 14-day period when P.
tremuloides flowers (late flowering females) were receptive and P. grandiden¬
tata males (early flowering clones) were discharging pollen. On April 8, 1967,
P. tremuloides stem #60 was observed to have receptive female flowers on
some catkins at a time when P. grandidentata clones were shedding pollen.
Similarly on April 5, 1968, P. grandidentata #12 was at the peak of pollen
discharge while P. tremuloides #35, 25 ft away, had many receptive female
flowers.

Excluding temperature inversions and other environmental factors, over¬
lap in flowering of female trembling aspen and pollen shedding of male big-
tooth aspen may occur in the following ways:

1. All of the flowers of inherently late-flowering female trembling
aspen clones may be receptive and receive pollen from early flow¬
ering clones of bigtooth aspen.

2. Some female flowers on the distal portion of some catkins of
trembling aspen are apparently receptive after flowers on the rest
of the catkin have been pollinated. We have observed such flowers
to be receptive during the normal pollen shedding period of big¬
tooth aspen.

3. Late-flowering whole catkins of certain clones produce receptive
flowers after most of the catkins on the same ramet are pollinat¬
ed. Einspahr and Joranson (1960) observed this pehnomenon on 5
female clones and 1 male clone of trembling aspen in the green¬
house and on 1 female trembling aspen in nature. We also ob¬
served this feature in the greenhouse and in nature and found that
flower buds at the base and tip of the shoot are most likely to
produce late-flowering catkins.

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Vol. 8

Although trembling aspen seems more likely to be the female parent of
hybrids, male trembling aspens also may have the opportunity to pollinate big-
tooth aspens. For example, in 1967 and 1968, P. tremuloides #1 was observed
shedding pollen when P. grandidentata #2, 10 ft away, had receptive female
flowers. On April 1, 1968, during the anthesis of P. tremuloides, a branch
from P. grandidentata #18, containing receptive female flowers, was removed
and taken indoors, eliminating the possibility of further pollination. This was
accomplished 4 days before any P. grandidentata males were observed to begin
shedding pollen. Pollen grains were observed on the stigmas.

In summary, flowering time is clearly not a complete isolating mechan¬
ism in southeastern Michigan. If the population described here is any indica¬
tion, many hybrid seeds must be formed. Although the cross P. tremuloides 9
X P. grandidentata 6 is most likely, the reciprocal is also possible. The 5
hybrids, 9% of the verified clones and 6% of all individuals, were undoubtedly
established as a result of the cultivation and the favorable soil moisture condi¬
tions. Because of the relative incompleteness of the flowering time barrier in
southeastern Michigan where the parent species occur together, lack of ade¬
quate seedbed conditions may be a more important factor in restricting the
occurrence of hybrids than time of flowering.

Pollen

The mean pollen abortion values for trembling and bigtooth aspen, 2.1%
and 4.9% respectively, were relatively low, and in this study were not signifi¬
cantly different (P > .05). Clonal differences in pollen diameter, coefficient of
variation for pollen diameter, and abortion percentage were not great for the
5 trembling aspens from which pollen was available. However, the pollen of
bigtooth clones showed marked variations in diameter and abortion rate.
Abortion percentage varied from 0.4% to 12.2%. Six of the 16 clones had
abortion percentages above 5%. Within-clone variation in pollen grain diameter
was significantly correlated (P < .01) with abortion percentage (r = 0.77). The
one male hybrid had a comparatively high abortion percentage, 18.7%, and
showed more variability in size of pollen grains than either parent (standard
deviation about twice that of either parent).

Seed Germination

Seed germination was excellent for the 1 trembling aspen, 93%, and the
13 bigtooth aspens, mean = 95%. The 2 hybrids tested on filter paper both
had a germination of 89%. Seeds from 4 clones, 2 bigtooth and 2 hybrids,
were tested only on a sand substrate. The percentages for the former were
96% and 43%, and for the latter, 87% and 4%. The low germination of the 1
hybrid may have been due to self-pollination of this hermaphroditic individual
and the possibility that most of the seeds used in the test were not sound
seeds. Maini (1964) found that bisexual flowers of P. tremuloides had a large
percentage of aborted seeds. We conclude that the germination of open pol¬
linated seeds from hybrids is only very slightly reduced from that of the par¬
ents. This agrees with the experience of the junior author who has repeatedly

1969

THE MICHIGAN BOTANIST

201

found high seed set and germination from open pollinated flowers of hybrids.
Heimburger (1936) reported that artificially produced hybrids between these
species “yielded good seed.”

Although several individuals suspected of being backcrosses or introgres-
sants were reported by Barnes (1961), there is as yet no compelling evidence
of these phenomena in Michigan aspens. However, this study has shown that
hybrids do have a high germination capacity and can compete with the parent
species well into flowering and fruiting age. Given suitable seedbed conditions,
backcrosses may be established in areas where hybrids coexist with the parent
species.

SUMMARY

A population of 78 aspens, Populus tremuloides, P. grandidentata, and their hybrid,
growing along the edge of a cultivated field adjoining the University of Michigan’s Stinch-
field Woods, Washtenaw Co., Michigan, was investigated to determine the number of seed¬
ling individuals, to account for their establishment, and to determine the incidence of
natural hybridization. The stand was established about 1954. Fifty-four of the stems were
verified as different genotypes through studies of phenology and leaf morphology. They
were presumed to be seedlings. Due to the even-aged character of the stand and lack of
disturbance the remaining individuals probably are also seedling plants.

Considerable overlap was observed in the flowering of trembling and bigtooth
aspen. There was approximately a 2-week period when late flowers of trembling aspen
individuals were receptive to pollen from bigtooth aspens. Less frequently, female flowers
of bigtooth aspen were receptive when pollen from trembling aspen was being discharged.
Five individuals were determined to be Fj hybrids between trembling and bigtooth aspen.
Seeds from 3 of the 4 female hybrids were highly germinable. Except for the relative
scarcity of hybrids there seems to be no stronger barrier to the production and establish¬
ment of successive backcross generations than there is for the production and establish¬
ment of seedlings of the parent species.

LITERATURE CITED

Ahlgren, C. E. 1959. Some effects of fire on forest reproduction in northeastern Minne¬
sota. Jour. Forestry 57: 194-200.

Andrejak, Gary E. 1968. A study of a seedling population of Populus tremuloides, P.

grandidentata, and their hybrid. Master’s Thesis, University of Michigan. 62 pp.

Baker, Frederic S. 1925. Aspen in the central Rocky Mountain region. U. S. Dep. Agr.
Bull. 1291. 47 pp.

Barnes, Burton V. 1961. Hybrid aspens in the Lower Peninsula of Michigan. Rhodora 63:
311-324.

Barnes, Burton V. 1966. The clonal growth habit of American aspens. Ecology 47:
439-447.

Barnes, Burton V. 1969. Natural variation and delineation of clones of Populus tremul¬
oides and P. grandidentata in northern Lower Michigan. Silvae Genetica [in press]
Bassett, I. J., R. M. Holmes, & K. H. MacKay. 1961. Phenology of several plant species at
Ottawa, Ontario, and an examination of the influence of air temperature. Canad. Jour.
PI. Sci. 41: 643-652.

Bertenshaw, James L. 1965. The clonal structure of selected aspen stands in northern
Lower Michigan. Master’s Thesis, University of Michgian. 56 pp.

Cottam, Walter P. 1954. Prevernal leafing of aspen in Utah mountains. Jour. Arnold Arb.
35: 239-250.

Day, Maurice W. 1944. The root system of the aspen. Amer. Midi. Nat. 32: 502-509.
DeByle, Norbert V. 1964. Detection of functional intraclonal aspen root connections by
tracers and excavation. Forest Sci. 10: 386-396.

202

THE MICHIGAN BOTANIST

Vol. 8

Einspahr, D. W. 1960. Abnormal flowering behavior in aspen. Iowa St. Jour. Sci. 34:
623-630.

Einspahr, Dean W., & Philip N. Joranson. 1960. Late flowering in aspen and its relation
to naturally occurring hybrids. Forest Sci. 6: 221-224.

Erlanson, E. W., & F. J. Hermann. 1928. The morphology and cytology of perfect flow¬
ers in Populus tremuloides Michx. Pap. Mich. Acad. 8: 97-110.

Faust, M. E. 1936. Germination of Populus grandidentata and P. tremuloides with partic¬
ular reference to oxygen consumption. Bot. Gaz. 97: 808-821.

Fetherolf, James M. 1917. Aspen as a permanent forest type. Jour. Forestry 15: 757-760.
Gauvin, G. 1964. Regeneration study in 1955 burn on the Q. N. S. Paper Company
Limits. Pulp Paper Mag. Canad. 65: WR452-453.

Graham, Samuel A., Robert P. Harrison, Jr., & Casey E. Westell, Jr. 1963. Aspens:

Phoenix Trees of the Great Lakes Region. Univ. Mich. Press, Ann Arbor. 272 pp.
Heimburger, Carl C. 1936. Report on poplar hybridization. Forestry Chronicle 12:
285-290.

Kittredge, Joseph, & S. R. Gevorkiantz. 1929. Forest Possibilities of Aspen Lands in
the Lake States. Univ. Minn. Agr. Exp. Sta. Tech. Bull. 60. 84 pp.

Larson, George C. 1944. More on seedlings of western aspen. Jour. Forestry 42: 452.
Lester, D. T. 1961. Observations on flowering in the aspens. Proc. Eighth N. E. For. Tree
Imp. Conf., pp. 35-38.

MacArthur, J. D. 1964. A Study of Regeneration after Fire in the Gaspe Region. Publ.
Dep. For. Canada 1074. 20 pp.

Maini, J. S. 1964. Anomalous floral organization in Populus tremuloides. Canad. Jour.
Bot. 42: 835-839.

Mills, B. B. 1961. The Beddington Bum Stages a Comeback. U. S. For. Serv. N. E. For.
Exp. Sta. Res. Note 117. 6 pp.

Pauley, S. S. 1956. Natural Hybridization of the Aspens. Minn. For. Note 47. 2 pp.
Pauley, S. S., & G. F. Mennel. 1957. Sex Ratio and Hermaphroditism in a Natural Popu¬
lation of Quaking Aspen. Minn. For. Note 55. 2 pp.

Pauley, S. S. 1963. Performance of some aspen seed sources and hybrids in eastern Massa¬
chusetts. F.A.O. World Con. For. Gen. Tree Imp. 63/-2b/2. pp. iii-17.

Sampson, A. W. 1919. Effect of Grazing upon Aspen Reproduction. U. S. Dep. Agr. Bull.
741. 29 pp.

Santamour, F. S. 1956. Hermaphroditism in Populus. Proc. Third N. E. For. Tree Impt.
Conf. pp. 28-30.

U. S. Dep. Commerce Weather Bureau. 1954. Climatological data. State of Michigan. 69:
81-98.

Victorin, Fr6re Marie. 1930. Les variations Laurentiennes du Populus tremuloides et du P.

grandidentata. Contrib. Inst. Bot. Univ. Montreal 16. 16 pp.

Vogl, Richard J. 1969. One hundred and thirty years of plant succession in a southeastern
Wisconsin lowland. Ecology 50: 248-255.

Weigle, W. G., & E. H. Frothingham. 1911. The Aspens: Their Growth and Management.
U. S. Dep. Agr. For. Serv. Bull. 93. 35 pp.

Westergaard, M. 1958. The mechanism of sex determination in dioecious flowering plants.
Advances in Genetics 9: 217-281.

Editorial Notes

INDEX: Notice to Librarians, Subscribers, Binders. No index appears in this
final number of Vol. 8. A three-year index, covering Volumes 7, 8, & 9, is
planned at the end of Vol. 9, following the pattern established in Vol. 3 and
Vol. 6. Thus far, three volumes have made a unit of practical size for binding.
In the future, if we are able to continue an increased size of volumes, an an¬
nual index may be prepared. Expressions of opinion from readers would be
welcome, regarding a useful frequency of indexing.

The May number (Vol. 8, No. 3) was mailed May 16, 1969.

Errata

Vol. 8, No. 1 (Jan. 1969, p. 16. Scale of map should read: 1 cm = approx.
63 m.

Vol. 8, No. 2 (March 1969), p. 74. Right half of scale should be headed cm.

p. 92. Date of Fig. 14 should read May 29, 1966.
p. 96. Right column of Table III, line 17, for “has
been” read had been.

p. 97. line 15 from bottom, for “as will be” read
and as will be.

Vol. 8, No. 3 (May 1969), p. 164 (rear cover). Add name of author after
fourth line from end of Contents: Florence V. Floseney.

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CONTENTS

Studies of the Byron Bog in Southwestern Ontario.

XL. Distribution of Sphagnum Mosses

William W. Judd . ’ . 167

Population Studies in the Chemical Species of the
Cladonia chlorophaea Group

Richard Wetherbee . 170

Studies in the Genus Cortinarius, I:

Section Dermocybe, Cortinarius aureifolius Complex

Joseph F. Ammirati, Jr., & Alexander H. Smith . 175

The Botanical Activities of Thomas J. Hale, 1858-1862

Lytton J. Musselman . 181

Dr. Bessie Bernice Kanouse— 1889-1969

Alexander H. Smith . 187

A Seedling Population of Aspens in Southeastern Michigan

Gary E. Andrejak & Burton V. Barnes . 189

Editorial Notes . 203

Errata . 203

(On the cover: Jordan River below Chestonia Bridge,

Antrim County, Michigan. Photo by John P. Schnell, July 26, 1969; »
reprinted by permission from newsletter of the
Mackinac Chapter, Sierra Club.)

LIBRARY

THE

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BOTANICAL GARDEN

MICHIGAN BOTANIST

January, 1970

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Editorial Board

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1970

THE MICHIGAN BOTANIST

3

POST-SETTLEMENT INFLUENCES UPON A
SOUTHERN MICHIGAN MARL LAKE1

Peter H. Rich

W. K. Kellogg Biological Station, Michigan State University,

Hickory Corners, Michigan

The origins of several excavations in the marl basin of Lawrence Lake
were investigated during an intensive limnological investigation of the lake. In¬
terviews with local residents revealed a long history of marl removal and much
additional information concerning post-settlement land and water usage in the
area. Patterns in the cultural history of Lawrence Lake are discernible, and
may be common to other marl lakes in southern Michigan.

Lawrence Lake, also known as Willison’s and East Lake, is 2.1 km (1.3
mi) east of the village of Hickory Corners in southern Barry County (Pig. 1).
Hickory Corners enjoyed prosperity in the 19th century out of proportion to
its present size (population ± 220). In 1880, prior to the establishment of a
railroad on the west side of Gull Lake, Hickory Corners was an important
trading center for the area, and had a population of 150 people. Hickory
Corners Road, immediately north of Lawrence Lake, was built in 1837. The
road has since been regraded but has not been significantly relocated in the
Lawrence Lake area.

Lawrence Lake, Little Lawrence Lake, and the marsh associated with
them are contained in a basin lying along the southern outwash apron of the
Kalamazoo morainic system (Leverett & Taylor, 1915). The soil surrounding
the basin is -generally a sandy loam of moderate fertility with good to rapid
drainage (Deeter & Trull, 1928; Veatch, 1953). The marsh bordering Lawrence
Lake is extensively underlain by marl. Casual observation suggests that peat is
a major component of the marsh south of the lake.

Lawrence Lake has an area of 4.9 ha (12.1 a) and a maximum depth of
12.6 m (41.3 ft). It receives two small streams from a drainage basin roughly
10 times the surface area of the lake (Fig. 1). Lawrence Lake drains into a
small unnamed lake, then south through an extensive marsh to Augusta Creek.
The lake has a distinct marl bench at its border, unless removed by dredging.
Dredged areas are discernible on the morphometric map (Fig. 2) by the de¬
parture from simple concentric contour lines on the north side, the west end,
and a small area in the southeast corner of the lake. The bench occurs from
waters-edge to a depth of 1 m and may extend up to 20 m from shore. Exca¬
vations have shown that the marl underlies a much broader zone around the
lake. For instance, marl occurs, and is reputed to be 1 1 m deep, at the ex¬
treme western end of the lake which is now 50 m back from the original
shoreline. Recent cores by Dr. R. O. Kapp of Alma College indicate that marl
is at least 5 m thick beneath the deepest point in the lake.

Contribution No. 181, W. K. Kellogg Biological Station of Michigan State Uni¬
versity.

4

THE MICHIGAN BOTANIST

Vol. 9

AGRICULTURE

The first house in the Hickory Corners area was built near a flowing
spring just west of Little Lawrence Lake in 1834. Watercress (. Nasturtium
officinale ) was immediately introduced into this and nearby springs. Pepper¬
mint {Mentha piperita) was added about 1890. Both species are present today.
Two bogs containing native cranberries {Vaccinium macrocarpon and V.
oxycoccos) surrounded springs near Little Lawrence Lake and were exploited.
Furthermore, the land surrounding many springs was actually plowed and cul¬
tivated for various garden crops. “Swamp shoes” on the hooves of horses pre¬
vented miring in the treacherous ground. The shoes were hardwood planks,
about 2 decimeters wide and 3 dm long, with an iron harness for attachment.
Despite the danger of miring and snakebite, cattle and sheep were also allowed
on the marshes to graze. Although Deeter and Trull (1928) state that prairie
sites and well drained “oak openings” were preferred agricultural land in the
county, my information suggests that marshes also received significant atten¬
tion.

The earliest lumbering in this region may well have been concentrated
on wet areas. The original tamarack {Larix laricina) was tall and straight

HICKORY

CORNERS

0

2 KM.

Fig. 1. Map of Lawrence Lake and vicinity, Barry County, Michigan, delineating marshy
area.

B.M •
281.6m

1970

THE MICHIGAN BOTANIST

5

6

THE MICHIGAN BOTANIST

Vol. 9

enough to be preferred for rafters in houses and barns, with the result that no
large tamarack have existed near Lawrence Lake in the memory of those now
living in the area. White cedar ( Thuja occidentalis) was also removed early. A
narrow border of hardwoods along the southern shore of Lawrence Lake is
largely virgin, but has suffered from soil compaction by cattle, selective
cutting, and Dutch elm disease. All other trees around the lake are recent
second growth.

In the early 20th century, from about 1918 to 1930, the marsh west of
Lawrence Lake was ditched, irrigated, and intensively cultivated for commer¬
cial crops of onions, celery, mint, etc. A smaller area of marsh northeast of
the lake was planted to rye for one season, 1891, and has remained fallow
since. Today, both areas are dominated by shrubby cinquefoil ( Potentilla
fruticosa). The presence of blazing-star ( Liatris spicata) in the older, less dis¬
turbed area suggests that the original community may have been a wet prairie.
Since 1930, agricultural use of land near Lawrence Lake has declined to occa¬
sional pasturing and watering of stock at the extreme eastern corner.

LAND RECLAMATION

Attempts to drain large areas of the marsh near Lawrence Lake began
about 1870 and were related to the expansion of agricultural enterprise. The
very large number of deaths attributed to the “summer fever” in the Law¬
rence family records suggests that mosquito control must have motivated later
reclamation efforts. Natural drainage from the west side of the marsh south of
Lawrence Lake was first diverted to Little Lawrence Lake. This modification
is known to have made Little Lawrence appear muddier than it had previously.

Lawrence Lake and the small, unnamed lake to the south were allowed
to continue draining to Augusta Creek through the east side of the marsh.
Later, water from Little Lawrence Lake was channeled east to the unnamed
lake. The unnamed lake, originally having a barren marl shelf, was almost
divided in two by the spoil from the ditching operation. The lake now has the
appearance of two connected ponds almost overgrown by aquatic plants (Lig.

i).

Attempts to straighten and deepen the drainage from Lawrence Lake to
Augusta Creek were hindered by an abandoned beaver dam encountered
50-100 m south of the unnamed lake. Dynamite was used at the site in 1930
to open the channel further. The dynamiting, which coincided with several
drought years, was followed by reduced water levels in Lawrence Lake for
two years. Beaver still exist on the upper parts of Augusta Creek.

MARL REMOVAL

Marl has been removed from the Lawrence Lake area since settlement.
Lrom 1840 to the middle 1890’s a lime kiln operated just west of Lawrence
Lake. A narrow-gauge tramway ran south from the kiln to Little Lawrence
Lake where the marl was mined. A dirt road now covers the tramway bed.
Quicklime produced from the kiln was used for plaster in nearby homes. The
demand for firewood required by the kiln may have accelerated tree cutting

1970

THE MICHIGAN BOTANIST

7

nearby, but no marl was removed from Lawrence Lake itself during this
period.

Marl was first removed from the western end of Lawrence Lake about
1918. Demand may have resulted from the introduction of alfalfa, which was
thought to require calcium, into the agriculture of the period. Marl was re¬
moved during the summer, and transportation was accomplished by horse-
drawn sledges during the winter.

Shortly after the publication of the county soil survey (Deeter & Trull,
1928), which recommended liming in both light and heavy soils, increased
demand and the introduction of mechanized draglines brought about large-
scale marl removal at Lawrence Lake. The western end of the lake was
dredged from 1930 to about 1960, and spoil banks of marl are still in evi¬
dence. Marl removal at this site since 1918 has produced a bay roughly 2500
square meters in area and 3 m deep. The northern edge of the lake was
dredged for 3 years, ending in the early 1950’s. The resulting excavation is in
the form of an elongated trough about 4 m deep which is parallel to the main
basin (Fig. 2). The trough is isolated from the main basin by a vestige of the
marl shelf associated with the original shoreline. Marl at this site was found to
be only 4 m deep.

Changes during the period of active marl removal at both the eastern
and northern sites are partly documented by a series of aerial photographs
obtained from the National Archives and Records Service. Approximately 11%
of the present surface area of Lawrence Lake and about 6% of its volume
have resulted from marl removal since 1918. Marl dredging still continues at
Little Lawrence Lake.

An examination of the current aquatic macrophyte assemblage in Law¬
rence Lake has given evidence that marl removal has had a drastic effect upon
the aquatic flora. The broad marl shelf existing along undisturbed shorelines
supports very sparse and stunted patches of Chara vulgaris, C. globularis, Najas
flexilis, and Potamogeton gramineus. Much of the natural marl shelf is entirely
barren.

Beyond the 1 m contour, the bottom slopes very sharply downward, and
maintains a high angle of descent well beyond the limit of macrophyte
growth. From the 1 m contour to 6 m a submerged sedge, the water bulrush
(Scirpus subterminalis ), grows very densely (up to 200 g/m2ash-free dry
weight).2 The nearly pure stand of water bulrush is significantly invaded by
another species ( Potamogeton praelongus) only at a depth of 5 m and below.
The P. praelongus zone seldom extends below 7 m.

In contrast, dredged areas are generally 3-4 m deep, and support dense
stands of Potamogeton praelongus, P. illinoensis, and P. pectinatus. The
absence of the sterile marl shelf also allows aquatics to grow to the shoreline.

The aquatic vegetation was sampled each month throughout a year by means of a
core sampler specially designed for the types of vegetation and bottom material found in
Lawrence Lake. Four replicate samples were taken at each 1 m depth interval over four
transects in the lake. Biomass for each species was determined on a dry weight and ash¬
free dry weight basis.

8

THE MICHIGAN BOTANIST

Vol. 9

As a consequence of the barren marl shelf and steep sides of Lawrence
Lake, the total crop of macrophytes is relatively small. The dominance of the
water bulrush limits the invasion and growth of pondweed species still further.
However, marl dredging drastically disrupts the typical lake morphology. Thus,
the relatively small addition of 11% to the total lake area by marl removal
probably represents a significant increase in substrate suitable for aquatic
macrophytes and allows a major increase in Potamogeton growth.

MISCELLANEOUS ACTIVITIES

Ice was cut from both Lawrence and Little Lawrence Lake from settle¬
ment until complete electrification of the Hickory Corners area in 1940.

A sugar bush and sugaring equipment existed south of the lake outlet
until 1930.

Intensive muskrat trapping in the area is known during all periods of
past habitation. Muskrat activity persists in Lawrence Lake at the present
time.

The lake enjoyed a local reputation as a good fishing spot in the past.
This reputation may be due to knowledge of some stocking done by a past
owner between 1910 and 1925. Species mentioned include warmouth, crappie,
yellow perch, bluegills, black bass, and pike. Present fishing effort is minor.

To date, Lawrence Lake has escaped major pollution. Inevitably, the
next step in the evolution of human influence upon the lake appears to be
nearby residential development.

SUMMARY

Three major patterns of human activity have influenced Lawrence Lake since set¬
tlement.

Pioneer agriculture utilized the marshy border of the lake, and evolved into special¬
ized, irrigated crops in the early 20th century. The original tamarack and white cedar
were removed during the early stages of settlement.

Large-scale efforts to drain the marsh south of Lawrence Lake were initiated short¬
ly after settlement to extend agricultural sites and to reduce insect vectors of human
disease.

Marl removal occurred in three phases. In the 19th century marl was removed and
dehydrated for plaster. Early in the 20th century agricultural marl was removed by primi¬
tive means, probably in response to the introduction of alfalfa as a field crop. Following
the publication of the county soil survey in 1928, marl was removed to a much greater
extent and by mechanical means. There is evidence that marl removal has allowed a sig¬
nificant increase in the quantity and diversity of aquatic macrophyte species in Lawrence
Lake.

Several human activities having less drastic impact upon the lake are cited.

ACKNOWLEDGMENTS

Much of this account was gathered from personal and telephone interviews either
with people directly involved with marl dredging at Lawrence Lake and with shorefront
owners, or their relatives. Mr. Curtis Lawrence of Hickory Corners, who supplied most of
the information from the 19th century and a great deal from the 20th, is a direct de¬
scended of the first settler, a life-long resident, and a conscientious local historian. His
statements were supported in part by photographs, records, and old maps (1860, 1873,
1895) in his possession.

1970

THE MICHIGAN BOTANIST

9

Miss Jenny Greer, who has resided within sight of Lawrence Lake during most of
her life and who permits Station personnel free access to the lake over her property,
convincingly corroborated Mr. Lawrence on all points. Miss Greer, scarcely more than a
generation removed from the hardy people who settled Hickory Corners, exemplifies their
incredible health and spirit.

General information concerning Hickory Corners was gleaned from compilations
distributed by the Hickory Corners Home-Coming Committee.

Information concerning the morphology and aquatic macrophytes of Lawrence
Lake became available in the course of joint efforts under Dr. R. C. Wetzel (Atomic
Energy Commission Contract AT(1 1-1)-1 599, COO-1599-23) and from the author’s thesis
research.

LITERATURE CITED

Deeter, E. B., & F. W. Trull. 1928. Soil Survey of Barry County, Michigan. U. S. Dep.

Agr. Bur. Chem. Soils, Ser. 1924, No. 14. 22 pp. + map.

Leverett, F., & F. B. Taylor. 1915. The Pleistocene of Indiana and Michigan and the

History of the Great Lakes. U. S. Geol. Surv. Monogr. 53. 529 pp. + maps.

Veatch, J. O. 1953. Soils and Land of Michigan. Mich. State Coll. Press, East Lansing.
241 pp. + map.

VASCULAR PLANTS OF THE BRUCE PENINSULA:

A REVIEW, WITH COMMENTS AND ADDITIONS

George Wm. Thomson

Michigan Botanical Club, Southeastern Chapter,

15093 Faust Avenue, Detroit, Michigan

For the last few weeks I have been reliving over 22 years of enjoying
the wild flowers of the Bruce Peninsula of Ontario during many holidays long
and short. The Bruce Peninsula lies between Lake Huron and Georgian Bay
and has been a mecca for botanists and plant lovers for a century. Being dedi¬
cated botanophiles, Mrs. Thomson and I were usually replete with hand lens,
camera, and just about every field manual which might help. Above all, our
constant companion was “Krotkov,” Professor P. V. Krotkov’ s fine paper,
Botanical Explorations in the Bruce Peninsula, Ontario (Trans. Roy. Canad.
Inst. 23: 3-65. 1940), published seven months after his death. This paper has
an excellent description of the ecology of the Bruce and the important plant
associations. It includes an annotated list of 916 different plant taxa repre¬
senting his collections and a few by others.

It was inevitable that Professor Krotkov would miss some plants, since
he made his major trips to the northern and central portions of the Peninsula
and actually excluded one of the richest areas, Keppel Township of Grey
County, from his region of interest. Since then most workers have considered
the Bruce to be more naturally bounded by the highway between the towns

10

THE MICHIGAN BOTANIST

Vol. 9

of Southampton and Owen Sound, with minor extensions just east of Owen
Sound. This boundary, without the Owen Sound extension, is used in the new
Check-List of Vascular Plants of the Bruce Peninsula prepared by the Wild-
flower Committee of the Federation of Ontario Naturalists.1 There is a short,
good introduction by Peter A. Peach, a geologist, on the botany of the Bruce
Peninsula. The new List is stated to be based on Krotkov, without any at¬
tempt to check herbarium specimens.

The scientific names in the Check-List are based on M. L. Fernald’s
eighth edition of Gray's Manual, designated “Gray” below. We must differenti¬
ate carefully between name equivalencing and Bruce plants. The first is a
necessary exercise in botanical semantics— the relationship between the names
in Krotkov and in Gray. The authors emphasize that they did not attempt to
verify the actual identity of Krotkov’s specimens. They give the closest Gray
name to the Krotkov name. I think that this was wise since Krotkov relied
heavily on Fernald’s advice and the identity of Krotkov’s plants would be ex¬
pected to be close to the eighth edition of the Manual which was nearing
completion at that time. On the other hand about one hundred new plants are
in the Check-List, swelling Krotkov’s total of 916 (not 915 as he says) to a
total of over 1000. For these new plants the scientific names should have
been based on the best taxonomic opinion of today, not that of over twenty
years ago. Alternative names would have been very useful. Unfortunately no
areas or collection citations are given for new entities in the List.

I estimate that at least another thirty entities should be added, so that
the Bruce flora includes at least 1040 different kinds of plants. There is good
internal evidence that the leading authority on the southeastern Ontario flora,
Professor James H. Soper, was not consulted— doubtless he could have added
many more.

It would require too much space to list here the 107 plant entities in
the Check-List not in Krotkov. (A list of these and of the excluded species
would have been very useful.) One at least is really new, Listera ovata (L.) R.
Br., the Common Twayblade of Europe and Asia, first discovered in North
America on the Bruce near the Lake Huron Shore in July 1968. (See Elliott,
Ontario Nat. 1/69: 15. 1969).) I can vouch personally for its identity since I
photographed it (Figs. 1 & 2) in full blossom on July 9, 1969, and had seen it
growing in the Edinburgh Botanic Garden in 1962. There were about twenty
plants nearby in various stages of development. On that trip I had taken with
me two British floras to check some of the Bruce waifs. The twayblade
matched exactly the fine line illustration by John Hutchinson in his British
Wild Flowers (Vol. II, p. 875. 1955). The plant is enormous compared with
our tiny native twayblades, being two to three feet tall with the single pair of
leaves about four inches across.

1 Check- List of Vascular Plants of the Bruce Peninsula, by Marion S. Shivas, M. D.
Kirk, W. K. Kirkwood, & Carol Rolfe. Federation of Ontario Naturalists, 1262 Don Mills
Road, Don Mills, Ontario, 1969. 62 pp. $1.50.

1970

THE MICHIGAN BOTANIST

Fig. 1. (left). Common Twayblade, Listera ovata, near Lake Huron, Bruce Peninsula,
Ontario.

Fig. 2. (right). Inflorescence of the same plant. Both photos by George W. Thomson, July
9, 1969.

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Seventeen ferns are new, although most have already been noted from
the Bruce in the botanical literature. Of note are Asplenium platyneuron (L.)
Oakes, the Ebony Spleenwort, and Woodsia Cathcartiana Robins., Cathcart’s
Woodsia. Fern students would have appreciated approximate localities or col¬
lection records of these rarities. There will probably be more records for the
Moonworts, Botrychium spp., and the Wood Ferns, Dryopteris spp., when
Dale J. Hagenah completes his studies, since he has more field knowledge of
the Bruce ferns than anyone else. The Dryopteris hybrids have been extensive¬
ly studied on the Bruce by Mr. Hagenah and by Dr. D. M. Britton, who
“checked” the ferns of the Check-List. However, these hybrids are not even
hinted at. Britton even gives the old Gray name for Dryopteris spinulosa var.
fructuosa although he has discussed it as the hybrid D. intermedia X spinulosa
in other publications. There are a number of other slips and technical errors in
the fern section reflecting the dependence on only translating the names to
Gray and the difficulty of relating the results of recent research to plants in
the field. The distinctive hybrid between the Northern Holly Fern and the
Christmas Fern, Polystichum XHagenahii W. J. Cody (Am. Fern Jour. 58: 30.
1968) was named from material collected on the Bruce by Dale Hagenah and
W. H. Wagner, Jr. Note that the name of the Lady Fern, according to Gray,
should be var. Michauxii (Spreng.) Farwell and not the species itself as shown
on p. 10 of the List.

Before leaving the ferns I must point out the most astonishing omission
of them all. Schizaea pusilla Pursh, the Curly-Grass Fern, was collected in
July, 1928, by a keen-eyed moss expert, E. A. Moxley, in an area now built
up with cottages at Sauble Beach. This record is noted in Gleason’s edition of
Britton and Brown’s Illustrated Flora but not in Gray. The information was
not known to Krotkov since the collection was not published until 1945 (Am.
Fern Jour. 35: 40). Perhaps this tiny fern will be found again since there is no
doubt either of its identity or of the exact location where it was collected.

The Bruce has been famous for a century as a rich orchid area. Krotkov
has 28, the Check-List has 38, and Fred Case’s recent book, Orchids of the
Western Great Lakes Region (Cranbrook, 1964) shows 33 from the Bruce. The
new items in the List not shown on Case’s maps are Spiranthes gracilis (Bigel.)
Beck, Slender Ladies Tresses; Malaxis unifolia Michx., Green Adder’s Mouth
Orchid; Listera ovata, discussed above; a color form of Spotted Coral-root;
and the doubtful Habenaria media which is often said to be the cross between
H. dilatata, Tall White Bog Orchid, and H. hyperborea, Tall Northern Green
Orchid. These intergrade so much in the field that H. media is usually rejected
as a worthless name. The Check-List also accepts H. macrophylla, as did
Krotkov, which many orchid specialists including Case and Correll believe to
be a larger form of H. orbiculata, Large Roundleaved Orchid or Dinner Plate
Orchid, as we call it on our Michigan Botanical Club field trips. A strange
omission is Spiranthes cernua (L.) Richard, Nodding Ladies Tresses, which
many of us have seen and which is mapped from the Bruce in Case. If the
boundaries of the Bruce are stretched to a few miles east of Owen Sound, yet
another orchid can be added. The Hagenahs have found a good stand of

1970

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13

Orchis spectabilis L., Showy Orchis, there and we would expect to find it in
similar habitats to the west.

The following are among the other entities in the Check-List which are
not in Krotkov: Trillium cernuum, Nodding Trillium; Ranunculus flabellaris,
Yellow Water Crowfoot; Menispermum canadense, Moonseed; Corydalis
sempervirens, Pale Corydalis; Saxifraga Aizoon var. neogaea, called the “Yel¬
low Mountain Saxifrage” in the List although this name is generally applied to
S. aizoides, while “Encrusted Saxifrage” has been used for S. Aizoon; Sorbaria
sorbifolia, False Spiraea; Pterospora andromedea, Pine Drops; Centaurium
umbellatum, Centaury; Myosotis verna. White Forget-me-not; Hedeoma
pulegioides , American Pennyroyal; Conopholis americana, Squaw Root;
Veronica longifolia, Longleaved Speedwell; Veronica officinalis, Common
Speedwell; Penstemon Digitalis, Foxglove Beardtongue; Campanula rapun-
culoides, Creeping Bellflower; Actinea herbacea, Stemless Rubber Weed;
Grindelia squarrosa, Gumweed; Tragopogon porrifolius, Common Salisify or
Purple Goatsbeard; Tussilago Farfara, Coltsfoot. These are splendid additions
to the flora. I have seen only a few of them and am looking forward to find¬
ing the others.

About ten entities have been dropped from Krotkov; in other words, I
could not find a corresponding item in the Check-List. Potamogeton longi-
ligulatus Fern., a Pondweed; Oryzopsis pungens (Torr.) Hitchcock, a Rice
Grass; Smilax rotundifolia L., Common Greenbrier; Populus tremuloides var.
intermedia Viet., a variety of the Quaking Aspen not maintained in Gray and
var. magnifica Viet., not only given in Gray (p. 522) but mentioned as from
the Bruce (!); Betula microphylla Bunge, an arctic and western Birch not in
Gray but collected in northern Canada by Hugh Raup; Apocynum sibiricum
var. salignum (Greene) Fern., the western extreme of Clasping-leaved Dogbane,
not in Gray; Chaenorrhinum minus (L.) Lange, which I have found in
Wiarton; Inula Helenium, Elecampane, common along roadsides south of
Wiarton; Solidago altissima, Tall Goldenrod. I believe that these items ap¬
parently omitted in transcribing the Krotkov list should be included in the
Check-List when a new edition appears.

Listed below are some plants which are included neither in Krotkov nor
in the Check-List, but which I have seen. I have also included some shrubs
recorded as growing on the Bruce on the maps by Soper and Heimburger (100
Shrubs of Ontario, 1961); these are marked “S,H” below. Many of the plants
have been noted by our friends and associates in the Michigan Botanical Club,
not only on innumerable short trips, but also during our expeditions over the
entire peninsula in 1957, 1958, and 1959 (with 40 to 50 each time) and on
our memorable State meeting in 1966 attended by 199 members.

Lycopodium complanatum var. flabelliforme Fern., Running Ground-
Pine. The common form in the rich dune forests of Sauble Beach, etc.

Claytonia virginica L., Common Spring Beauty. This narrow-leaved
species is often seen with the commoner C. caroliniana Michx. in rich hard¬
woods.

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THE MICHIGAN BOTANIST

Vol. 9

Ranunculus bulbosus L., Bulbous Buttercup, is widespread both along
roadsides and in open woods.

Ranunculus trichophyllus Chaix, Hairleaved Water Crowfoot, was seen
once in a muddy stream near Lindenwood. These were not R. longirostris
Godr., which is listed in Krotkov.

Clematis verticillaris DC., Purple Clematis. (S,H)

Lindera Benzoin (L.) Blume, Spice Bush. (S,H) We have seen it many
times in rich mixed woods.

Berberis vulgaris L., Common Barberry. (S,H) Only found in the wild
occasionally. A sturdy plant with very large leaves and spines was seen in a
rocky pasture in the shelter of large dolomite blocks, not near a farm house,
east of Clavering.

Berberis Aquifolium Pursh, Mahonia; Oregon Holly Grape. A very large
bush about 15 feet in width is flourishing and reseeding on an abandoned
farm site southwest of Lindenwood.

Erucastrum gallicum (Willd.) O. E. Schulz, Dog Mustard. A tumbleweed
mustard pointed out by Dr. E. G. Voss of the University of Michigan on our
1966 expedition. He said it was becoming widespread around the Great Lakes.
Since then we have seen it at many places along the Huron shore.

Hesperis matronalis L., Dame’s Rocket. An abundant plant throughout
on shores, talus, and roadsides.

Dentaria laciniata Muhl., Cutleaved Toothwort. Plentiful in rich woods
north of Kemble.

Barbarea vulgaris R. Br., Yellow Rocket. Plants along the roadsides near
Johnson Harbour were not B. orthoceras.

Ribes Cynosbati var. atrox Fern., Bristly Gooseberry. (S,H) We have
seen it at many places in the northern part of the Bruce. Type locality is
Manitoulin Island.

Ribes triste var. albinervium (Michx.) Fern., glabrous form of Wild Cur¬
rant. Much commoner than the hairier form.

Ribes odoratum Wendland f., Golden Currant, and R. glandulosum
Grauer, Skunk Currant. Both S,H.

Trifolium procumbens L., Low Hop Clover. Many Bruce specimens
checked were not T. agrarium L. Both species are common along roadsides in
gravel on both sides of the Peninsula.

Gleditsia triacanthos L., Honey Locust. There are old planted trees on
the Big Bay Road near Owen Sound.

Euphorbia marginata Pursh, Snow-on-the-mountain. Well established
escape, Oliphant Cemetery and elsewhere.

(Two spurges noted in the Check-List, E. Cyparissias L., Cypress Spurge,
and E. Esula L., Leafy Spurge, are dominating large areas of the sandy fields
north of Sauble Beach and are appearing elsewhere.)

Euonymus atropurpureus Jacq., Burning Bush. Rich woods on lower
talus slopes, Sydney Bay.

We have all sorts of other escapes, so let us include Althaea rosea Cav.,
Hollyhock.

1970

THE MICHIGAN BOTANIST

15

Viola renifolia Gray, Hairy Kidneyleaved Violet. The densely pubescent
form is common in similar wet wood habitats with var. Brainerdii (Greene)
Fern., the almost glabrous form mentioned in the Check-List.

Viola rostrata Pursh, Longspurred Violet, in rich wet woods with V.
conspersa Reichenb., Dog Violet.

Aegopodium Podagraria L., Bishop’s Goutweed. A bad garden escape,
long persistent.

Levisticum officinale W. D. J. Koch, Lovage. An astonishing omission
from the Check-List. The plants at the Big Bay Dock and nearby at North
Keppel have been flourishing since the village was founded about a hundred
years ago. They are known to all the members of the Michigan Botanical Club
who have visited the Bruce.

Pyrola rotundifolia L., Roundleaf Pyrola. Noted on Flowerpot Island.
Not var. americana (Sweet) Fern.

Kalmia polifolia Wang., Bog Laurel. (S,H)

Vinca minor L., Myrtle or Common Periwinkle, is well established as a
ground cover in Wiarton and other built-up areas.

Myosotis scorpioides L., Forget-me-not. Wet sand and beaches with M.
laxa Lehm.

Hackelia virginiana (L.) Johnst., Beggar Lice. Dock at Little Port Elgin,
Cape Croker.

Ajuga reptans L., Bugle. A well-established garden weed and escape,
Colpoy’s Bay.

Thymus serpyllum L. (sensu Amer. authors), Wild Thyme. A garden
escape noted at Red Bay, Cabot Head, and in other locations where it is well
established often far from habitations.

Orobanche fasciculata Nutt., Bunched Cancer Root. Photographed at
Dorcas Bay. Also found on Manitoulin Island.

Veronica peregrina L., Neckweed. A white speedwell common in the
grass near the shore at Hope Bay.

Viburnum cassinoides L., Witherod, and Lonicera villosa (Michx.) R. &
S., Mountain Fly-Honeysuckle. (Both S,H.)

This brings us to the Compositae. The most striking omission is the low
yellow matformer, Hieracium Pilosella L., Mouse-ear Hawkweed, which grows
by the millions, covering entire fields, in Keppel Township. Several of the rare
Moonworts {Botrychium spp.) listed in the Check-List, perhaps from Dr. D. M.
Britton’s records, grow hidden among this hawkweed!

Another bad pest of this marginal soil is Carduus acanthoides L., a spiny
thistle with a characteristic “skinny” look. A good common name might be
Acanthus Thistle; the Keppel farmers have a better one, I’m sure.

Cirsium muticum Michx., Swamp Thistle, is in the Sauble Beach area.

Lapsana communis L., Nipplewort, is becoming an active invader of
roadsides and disturbed places. I have seen plants six feet high in Owen
Sound.

Another hawkweed which is said to be having a population explosion
around the Great Lakes is Hieracium florentinum All., King Devil. Only a few

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THE MICHIGAN BOTANIST

Vol. 9

of the big yellow hawkweeds turn out to be this species, but we did see it at
Berford Lake. Much more worthy of the name King Devil is the other big
yellow one, H. pratense Tausch, Field Hawkweed (in the Check-List as a new
item), which often grows with the ubiquitous Mouse-ear Hawkweed.

Everybody notices Beilis perennis L., English Daisy, in the Wiarton
lawns but somehow or other it got left out of the Check-List.

Tragopogon major Jacq., Pale Yellow Goatsbeard, is the commonest of
the three salsifies found on the Bruce. Also known as T. dubius Scop., it can
be easily distinguished from T. pratensis L., which we call the Orange Goats¬
beard. All three species are common roadside plants. None of the hybrids de¬
scribed by Hall, Persell, and Thompson (Mich. Bot. 5: 79-91. 1966) have been
noted.

Solidago hispida var. tonsa Fern. (Gray, p. 1393), the almost completely
glabrous form of the Hairy Goldenrod, has apparently been included in var.
amoglossa Fern, in Gleason’s Illustrated Flora (III, p. 420). It is a very north¬
ern form which is the principal goldenrod in the dolomite crevices in exposed
wet places like the rocks near the Tobermory lighthouse.

Although we have seen and photographed it in northern Michigan, we
have looked for and not yet found in the Bruce the famous Upper Great
Lakes disjunct, Adenocaulon bicolor Hook., Trail Plant. It was collected by A.
Y. Massey in 1895 on the Cape Croker Indian Reserve. Gray (p. 1465) cites it
from the Bruce. Let us hope that the next edition of the Check-List will in¬
clude Adenocaulon as well as Schizaea.

In closing, I believe that the Check-List will be of great value to both
botanists and botanophiles. Its worth will be enhanced when the authors are
able to add the many other records of the numerous people who have studied
and enjoyed the plants of the Bruce for over a century.

Publications of Interest

[See also page 63]

NATURAL DYES IN THE UNITED STATES. By Rita J. Adrosko. U. S. National Muse¬
um Bulletin 281, 1968. 160 pp. $3.25, clothbound (Supt. of Documents, Govt. Print¬
ing Office, Washington 20402). Part 1 (53 pp.) is on dyestuffs used in America during
the 18th and 19th centuries; part 3 is on home dyeing with natural dyes. The ap¬
pendixes are chiefly historical, presenting accounts and recipes from early American
dye literature.

NIGHTSHADES The Paradoxical Plants. By Charles B. Heiser, Jr. W. H. freeman & Co.,
San Francisco, 1969. $5.95. A popular account, for the general reader, of the night¬
shade family (Solanaceae), including potatoes, tomatoes, hot peppers, tobacco, Luther
Burbank’s “Wonderberry,” mandrake, belladonna, and other edible, poisonous, or
ornamental members.

1970

THE MICHIGAN BOTANIST

17

THE GENUS PANELLUS IN NORTH AMERICA

Orson K. Miller, Jr.

Forest Disease Laboratory, Forest Service,

U. S. Department of Agriculture, Laurel, Maryland

The genus Panellus has been thoroughly studied in Europe and the
species clearly delimited by Kiihner and Romagnesi (1953), Moser (1967),
Singer (1962), and others. The species have been individually studied in North
America and variously reported by Kauffman (1918), Groves (1962), Krieger
(1947), and Murrill (1915) under Pleurotus, Panus, and other taxa. However,
the use of the amyloid reaction in connection with a systematic restudy of
the morphology of the basidiospores and the sporocarp of the North Ameri¬
can species has not been undertaken. The object of this study is to examine
the North American taxa which might be included in Panellus, and to report
on the number, distribution, and ecology of the North American species. Keys
are provided which utilize among others many new or previously unused char¬
acters to aid in the identification of fresh and dried specimens.

Photomicrographs were taken with a Leitz Orthomat camera mounted
on a Leitz-Ortholux research microscope. The line drawings were made with
the aid of camera lucida. Ridgway (1912) colors are indicated in quotation
marks, e.g., “snuff brown.” The location of each collection studied is indi¬
cated by the standard herbarium designation, e.g., (MICH), following the col¬
lection number.

PANELLUS P. Karst. Bidr. Kami. Finl. Nat. Folk 32: XIV. 96. 1879.

Pileus 8-45 (-100) mm broad, 5-35 mm wide, viscid to dry, glabrous to
woolly-pubescent, smooth, white, green, brown, or violaceous. Lamellae close
to subdistant, even, edges entire, white, pinkish-buff, ochraceous, olivaceous,
to vinaceous or brown. Veil present on young sporophores of one species.
Stipe present or absent, if present very short (1-12 mm) and eccentric or later¬
ally attached, dry, minutely pubescent or fibrillose, concolorous with the
pileus. Sporocarps revive when moistened. Taste mild to slightly acrid. Odor
not distinctive.

Spores 3.0-7 .0(- 1 1 .0) X (0.5-)l .0-2.0(3.5) /i, allantoid, oblong, cylindric
to narrowly elliptical, smooth, thin-walled, amyloid (blue to purple in Melzer’s
solution); spore print white to yellow. Basidia small, 11-28 X 2.5-4.2(-6.3) m,
clavate, thin-walled, 4-spored, hyaline in KOH and Melzer’s solution. Cheilo-
cystidia cylindric, fusiform, clavate to bifurcate or irregularly shaped, thin-
walled, hyaline, embedded to protruding, abundant to infrequent. Pleuro-
cystidia present or absent, when present similar to cheilocystidia. Pileocystidia
usually present, typically long, narrow, thin- or thick -walled, and single or in
fascicles. Cuticle either a gelatinous pellicle (Fig. 27) or of interwoven thick-
and thin-walled hyphae (Figs. 28, 29). The latter usually with hairs (pileo¬
cystidia) but sometimes aggregated into fascicles appearing hispid. Trama of
pileus of two types: 1. duplex with an upper gelatinous zone and a lower

18

Vol. 9

THE MICHIGAN BOTANIST

zone of interwoven, non gelatinous, thick- and thin-walled hyphae; 2. homo¬
geneous, of interwoven thick- and thin-walled hyphae. Trama of the lamellae
of interwoven usually thick- and sometimes thin-walled hyphae. Clamp con¬
nections are present in all species.

Habit, habitat, and distribution.— Imbricate, often very crowded (e.g.
Fig. 34), rarely single, always on twigs, branches, logs, and stumps of woody
plants. Most commonly found on hardwoods but occasionally on conifers,
noted especially on Betula, Salt c, Populus, Alnus, Quercus, Tsuga, Picea, and
Pinus. Fruiting from midsummer in the boreal forests to fall in the temperate
forests. Known from Europe, North America, South America, Asia, Australia,
and New Zealand.

Observations.— The genus Panellus is a lignicolous group of white- to yel-
low-spored species in-the Tricholomataceae. It differs from all other genera by
having a combination of amyloid, smooth, allantoid to narrowly elliptical
spores; thick-walled hyphae in the trama of the lamellae (see Figs. 4, 10); even
gill edges; a sessile to short-stipitate fruiting body; and thin-walled cheilo-
cystidia. Specimens revive when moistened and could be confused with
Lentinellus, Phyllotopsis, and Plicaturopsis. Lentinellus has short elliptical to sub-
globose amyloid spores but the serrate gill edges, cellular cutis, amyloid pileus
trama in some species, and lack of gelatinous tissue separate it from Panellus.
The nonamyloid, allantoid spores of Phyllotopsis, which are pink in deposit, are
distinctive. Plicaturopsis appears closely related to Panellus but its allantoid
spores are nonamyloid, the lamellae are often discontinuous or reduced to
veins, and it has distinctive basidia constricted near the apex (Reid, 1964).
Type: Panellus stipticus (Bull, ex Fr.) Karst.

KEY TO THE SPECIES OF PANELLUS

1. Pileus viscid, with a thin pellicle (Fig. 27), glabrous or with pileocystidia (Fig.

28), brown to green; upper trama a thick refractive, gelatinous layer; with or

without a veil when young; usually on hardwoods . 2

1. Pileus dry or moist, white, violaceous to brown; if brown distinctly stipitate;
cuticle with pileocystidia and/or loosely interwoven hyphae; lacking a veil; on

hardwoods or conifers . 3

2. Pileus small (8-15 mm broad), brown, villose to fibrillose in age; a
thin-membranous veil (Fig. 32) present on young sporophores but absent at

maturity; cheilocystidia (3. 5-5.0 ju wide, Fig. 3), pleurocystidia absent .

. 1 . P. patellaris

2. Pileus large (25-100 mm broad), slimy viscid when moist, green to dark
olive-green (drying green); veil absent; cheilocystidia and pleurocystidia

abundant (6.5-12 n wide, Figs. 7, 8) . 2. P. serotinus

3. Pileus small (5-10 mm), moist, white (drying white), with a thick gelatinous
upper trama; gelatinous gill edge (Fig. 31) furnished with small contorted

cheilocystidia 12-19 X 3. 2-7. 6 p (Fig. 13); always on conifer wood . 3. P. mitis

3. Pileus larger (10-35 mm), violaceous to brown; pileus trama and gill edge not

gelatinous; on hardwoods and conifers . 4

4. Pileus tan, distinctly laterally stipitate; cheilocystidia abundant, protruding,
narrow, 18-45 X 2.5-6 .0 p (Fig. 19); spores elliptical to weakly allantoid in

profile (Fig. 17); abundant and common on many hardwoods . 4. P. stipticus

4. Pileus violaceous, sessile; cheilocystidia infrequent, embedded; spores

allantoid to oblong; often on birch, willow, alder, and conifers (usually spruce) ... .5

1970

THE MICHIGAN BOTANIST

19

5. Spores (4.5-)5.0-6.5(-7.5) x 1. 2-2.0 yu; lamellae not distinctly intervenose;

usually on hardwoods; common in Europe and North America . 5. P. ringens

5. Spores 6-11 X 2-3.5 ju; lamellae conspicuously intervenose; usually on conifers;

known from eastern Canada and Europe . 6. P. violaceofulvus

1. Panellus patellaris (Fr.) Konr. & Maubl. Figs. 1-4, 27, 32.

Icon. sel. fung. VI. p. 379. 1924.

Panus patellaris Fr. Epicr. p. 400. 1838.

Pileus 8-15 mm broad, sessile to short-stipitate, convex, viscid to gluti¬
nous at first, soon becoming fibrillose to villose and moist to dry, brown,
“snuff brown.” Context a narrow water-soaked brown (“sayal brown”) layer
just beneath the gelatinous cuticle, remainder of context of pileus and stipe of
firm pale cinnamon (“pale pinkish cinnamon”) tissue. Stipe when present 1-3
mm long, 0.5-1. 5 mm wide, eccentric, equal, dry, usually somewhat curved.
Lamellae subdistant, narrow, “cinnamon-buff,” covered at first by a thin
membranous, dry “pinkish buff’ veil which soon disappears leaving no traces
at maturity.

Spores 4.5-5 .5 X 0.5-1. 5 p, allantoid, thin-walled, amyloid (light blue in
Melzer’s solution); spore print white. Basidia 17-28 X 3. 5-4.2 p, narrowly
clavate, 4-spored. Cheilocystidia 22-35 X 3.5-5 .0 p, clavate to fusiform, thin-
walled, hyaline with some yellowish contents in KOH and Melzer’s solution,
protruding 1/3 to 1/2 of total length. Pleurocystidia none seen.

Cuticle of pileus a thick gelatinous layer of interwoven hyphae 1.5-3. 5
p diam., thin-walled, clamps frequent, hyaline except for a thin pigmented
zone near the surface; trama of pileus of interwoven hyphae 2.5-3. 5 (-4.5) p
diam., thin-walled, clamps frequent, yellow to brown near cuticle, otherwise
yellowish to hyaline in KOH or Melzer’s solution. Trama of lamellae of inter¬
woven hyphae 2. 5-4 .5 p diam., thin- and thick-walled (walls up to 1.5 p
thick), hyaline to yellowish in KOH and Melzer’s solution.

Habit, habitat, and distribution.— Solitary, imbricate, or even in caespi-
tose clusters on dead branches or on fallen twigs. Branches usually not
decorticated. Noted on species of Alnus, Betula, Salix, and Fagus. Widespread
in Europe and North America but encountered infrequently. Fruiting from
July to October.

Observations.— P. patellaris is the only member of the genus with a veil.
At maturity the veil has disappeared and one must then look for the combina¬
tion of a viscid cuticle, gelatinous upper trama, narrow cheilocystidia, small
size, and brown coloration. P. serotinus is the other pigmented species with a
viscid cuticle, but it is much larger, lacks a veil, has wide cheilocystidia and
pleurocystidia, and the pileus is green to olivaceous when fresh. Fresh spore
prints (Miller, 1968) were used to demonstrate the amyloid spore wall.

Material examined. Canada: Ontario; R. I*’. Cain & A. H. Smith 4118 (MICH), L.
Overholts 18603 (MICH). Yukon Terr.; O. K. Miller 5547 (BEDL). U.S.A.: Maine; F. L.
Harvey 2010 (MICH). Miehigan; D. V. Baxter & C. H. Kauffman Aug. 18, 1922 (MICH),
A. H. Smith 22308 (MICH), D. E. Stuntz 5508 (MICH). New York; Man. 1884 (NY).

20

THE MICHIGAN BOTANIST

Vol. 9

Figs. 1-4. Panellus patellaris. 1. Basidia. 2. Basidiospores. 3. Cheilocystidia. 4. Hyphae in
the trama of the lamellae.

Figs. 5-10. P. serotinus. 5. Basidia. 6. Basidiospores. 7. Pleurocystidia. 8. Cheilocystidia.
9. Pileocystidia. 10. Hyphae in the trama of the lamellae.

Figs. 11-15. P. mitis. 11. Basidia. 12. Basidiospores. 13. Cheilocystidia. 14. Pileocystidia.
15. Hyphae in the trama of the lamellae.

Figs. 16-21. P. stipticus. 16. Basidia. 17. Basidiospores. 18. Pleurocystidia. 19. Cheilo¬
cystidia. 20. Pileocystidia. 21. Hyphae in the trama of the lamellae.

Figs. 22-26. P. ringens. 22. Basidium. 23. Basidiospores. 24. Cheilocystidia. 25. Pileo¬
cystidia. 26. Hyphae in the trama of the lamellae.

(Figs. 1-26 x750)

1970

THE MICHIGAN BOTANIST

21

2. Panellus serotinus (Pers. in Hofmann ex Fr.) Kiihner Figs. 5-10, 33.

FI. Analytique Champ, sup. p. 67. 1953.

Agaricus serotinus Fries, Syst. Mycol. 1: 187. 1821.

Pileus 2. 5 -6 .0(- 10.0) cm broad, 2.0-3. 5 cm wide, convex, fan-shaped,
conchate; margin often lobed or wavy, viscid, “vetiver green,” “olive ochre”
to “deep colonial buff’ rarely ochraceous in center to “dark greenish olive,”
“dark ivy green,” “citrine drab” to “dark olive buff’ occasionally toned
purple over the margin, always sessile. Lamellae close to subdistant, narrow,
even, clearly marginate, pale orange to “honey yellow” with “yellowish olive”
to “brownish olive” rarely “cinnamon buff’ on the margin.

Spores 4.0-5 .5 X 1.0-1. 5 p, allantoid to cylindric, with amyloid (blue)
walls and yellow contents in Melzer’s solution; spore print yellow. Basidia
17-21 X 3.0-4 .0 p, narrowly clavate, 4-spored. Pleurocystidia 35-64 X 6.5-11
p, fusiform, broadly fusiform, to clavate, thin-walled, sometimes with incrusta¬
tions over the apex, hyaline in KOH, yellowish in Melzer’s solution, frequent,
projecting up to 1/2 of total length. Cheilocystidia 40-60 X 6.5-12 p, nearly
cylindric, fusiform to narrowly clavate, thin-walled, hyaline in KOH, yellowish
in Melzer’s solution, abundant, projecting 1/2 to 3/4 of total length. Cuticle
with numerous tangled or decumbent, nearly cylindric, fusiform to long-
clavate, thin-walled pileocystidia 50-90 X 6-9 p with yellow contents in
Melzer’s solution, hyaline in KOH, olive in H2O, and arising from a thin cutis
composed of interwoven thin- and thick-walled hyphae 2. 5-8. 5 p diam., the
hyphae yellow in Melzer’s solution, hyaline in KOH. Trama of pileus duplex
with an upper gelatinous layer containing interwoven, thin-walled hyphae
1.5-3. 5 p diam. and is often folded or partially gelatinized. Lower trama of
non gelatinous, interwoven, thin- and thick-walled hyphae 3.5-18.0 p diam.,
bearing clamp connections, dingy yellow-brown in Melzer’s solution, light yel¬
lowish in KOH. Trama of lamellae of loosely interwoven, mostly thick-walled
hyphae 2.5-11.0 p diam., yellowish in Melzer’s solution, light yellowish in
KOH.

Habit, habitat, and distribution.— Occasionally solitary but usually imbri¬
cate on the sides of logs and sticks of various hardwoods and conifers, usually
before they become decorticated. Noted on Alnus, Populus, Ulmus, Fagus,
Betula, and Quercus, but also on Tsuga heterophylla in the western United
States. Fruiting in the fall especially during late September and October. Wide¬
ly distributed throughout North America, Europe, and Asia.

Observations.— This robust species has a characteristic greenish to olivace¬
ous color of the pileus, is viscid when fresh, and has numerous tangled pileo¬
cystidia (Fig. 28). This combination of characters is distinctive in Panellus.
Herbarium specimens without notes could be recognized by the combination
of long, broad, pleurocystidia and cheilocystidia; the distinctive pileocystidia;
and the thick gelatinous cuticle.

Material examined. -U.S. A.: California; A. H. Smith 56323 (MICH), 56773 (MICH).
Idaho; O. K. Miller 1472 (BFDL), 2222 (BFDL), 2248 (BFDL), 2081 (BFDL), 2851
(BFDL), 2890 (BFDL), 2917 (BFDL). A. H. Smith 53899 (MICH), 54585 (MICH),
54601 (MICH), 55247 (MICH), 55299 (MICH), 67702 (MICH), 70927 (MICH), 71033

22

THE MICHIGAN BOTANIST

Vol. 9

Fig. 27. Viscid cuticle of P. patellaris. Fig. 28. Viscid cuticle of P. serotinus with tangled
pileocystidia. Fig. 29. Dry cuticle of P. stipticus, a dense tangle of thick- and thin-walled
hyphae. Fig. 30. Elongate cheilocystidia in P. stipticus. Fig. 31. Gelatinous edge of
lamella on P. mitis.

1970

THE MICHIGAN BOTANIST

23

(MICH), 76559 (MICH), 76946 (MICH). J. R. Weir and A. S. Rhoads F.P. 38593
(BFDL). Michigan; O. K. Miller 131 (BFDL); A. H. Smith 43864 (MICH), 44063 (MICH),
50995 (MICH), 58322 (MICH), 58360 (MICH), 60809 (MICH), 62042 (MICH), 62043
(MICH), 62063 (MICH), 62210 (MICH), 62359 (MICH), 62394 (MICH), 64754 (MICH),
75227 (MICH). Montana; J. R. Weir F. P. 11348 (BFDL), F. P. 11411 (BFDL). New
Hampshire; P. Spaulding F.P. 2318 (BFDL). North Carolina; G. C. Hedgecock F.P. 55086
(BFDL). Canada: British Columbia; O. K. Miller 4856 (BFDL).

3. Panellus mitis (Pers. ex Fr.) Sing. Figs. 11-15, 31, 35.

Ann. Mycol. 34: 334. 1936.

Agaricus mitis Pers. ex Fr., Syst. Mycol. 1: 188. 1821.

Pileus 5-8 mm long, 5-6 mm wide, spathulate to conchate, flat to broad¬
ly convex, a dull white ground color, covered with a fine white canescence
from appressed fibrils (best seen with a handlens). Lamellae subdistant, even,
narrow, white to pale pinkish cinnamon. Stipe 1-2 mm long, 1 mm or less
wide, laterally attached, covered with minute white fibrils over a pallid ground
color.

Spores 3.5-6.0 X 0.9-1. 2 (-1.5) p, allantoid to cylindric, smooth, thin-
walled, amyloid (purple in Melzer’s solution). Basidia 11-22 X 2.5-3.5 p, nar¬
rowly clavate, 4-spored. Cheilocystidia 12-19 X 3.0-7 .6 p, fusiform, clavate to
contorted and irregular, thin-walled, hyaline in KOH and Melzer’s solution,
projecting somewhat or embedded in the sterile gelatinous layer which covers
the edges of the lamellae. Cuticle with long, thin-walled hairs (1.5-3. 5 p wide)
solitary or in loose often tangled fascicles, hyaline, often with a basal clamp
connection. Trama of the pileus duplex with a wide (300-400 \ u) gelatinous
upper layer, composed of gelatinous hyphae (1.5 -3. 5 p wide) embedded in a
refractive, hyaline matrix; lower layer non gelatinous, of interwoven, thin-
walled hyphae (2.5-7 .0 p, in diam.), ochraceous in Melzer’s solution, dingy
yellowish brown in KOH. Trama of lamellae of interwoven hyphae similar to
the lower trama of the pileus, with the exception of the edge which is com¬
posed of a broad gelatinous layer containing gelatinous hyphae.

Habit and habitat.— Usually scattered or in clusters along the sides of
conifer logs and sticks; particularly abundant on western larch ( Larix occi-
dentalis). Fruiting in the fall especially during September and October in
Europe and North America.

Observations.— The small white sporocarps containing a gelatinous upper
layer in the pileus trama and at the gill edges (Fig. 31) combined with the
unusual cheilocystidia (Fig. 13) are a combination of characters which is
possessed by no other species of Panellus.

Material examined. -Canada: B.C.; A. Foster 44413 (DAOM), 44474 (DAOM). N.
B.; Stillwell 87394 (DAOM). Nfld.; Stillwell 46768 (DAOM). Ont.; M. Nobles 46790
(DAOM), H. S. Jackson 50124 (DAOM), 8060 (DAOM). U.S.A.: Idaho; O. K. Miller
2850 (BFDL), 2856 (BFDL), 2901 (BFDL). A. H. Smith 54578 (MICH), 54842 (MICH),
55296 (MICH). Oregon; A. H. Smith 18133 (MICH), 55403 (MICH). Europe: Sweden; S.
Lundell, Fungi Exsiccati Suecici 9. (BPI). Norway; J. Stordal 65000 (DAOM).

4. Panellus stipticus (Bull, ex Fr.) Karst. Figs. 16-21, 29, 30, 34.

Bidr. Kann. Finl. Nat. Folk. 32: XIV. 96. 1879.

24

THE MICHIGAN BOTANIST

Vol. 9

Agaricus stipticus Bull, ex Fr., Syst. Mycol. 1: 188. 1821.

Panus stipticus (Bull, ex Fr.) Fr., Epicr. Mycol. 399. 1838.

Pileus (5-) 12-32 mm long, 12-1 5(-25) mm wide, convex to plano-convex,
crenulate margin in age, dry areolate, woolly-pubescent, sometimes with sev¬
eral concentric ridges or zones, ochraceous buff, “pale ochraceous buff’ to
“light ochraceous buff,” various shades of tan or brown when dried. Lamellae
close, narrow, often forked, “ochraceous-buff” to “ochraceous salmon” even
“tawny olive” in age. Stipe 6-12 mm long, 3-8 mm wide, usually eccentric,
often lateral, minutely fibrillose, dull white, constricted somewhat just at base.
Taste slightly acrid, perhaps somewhat astringent. Odor not distinctive.

Spores (3.0-)3.5-4.6 X 1. 2-2.2 p, elliptical to somewhat allantoid, with
yellow contents and amyloid (blue) walls in Melzer’s solution. Basidia 15-20 X
2. 5-3. 5 p, narrowly clavate, 4-spored. Cheilocystidia 17-45 X 3. 5-6.0 p, nar¬
rowly clavate, cylindric, clavate-rostrate, fusiform to bifurcate, thin-walled,
hyaline, abundant, protruding 1/2 total length or more (Fig. 30). Pleuro-
cystidia 19-40 X 3.0-4.5 p , fusiform, narrowly clavate, to bifurcate at apex,
thin-walled, hyaline, scattered or in a dense cluster, occasionally protruding up
to 1/2 total length. Cuticle of pileus a tangle of thick- and thin-walled,
clamped hyphae 1. 2-6.0 p diam. (Fig. 29) with scattered inconspicuous pileo-
cystidia 40-55 X 3.5-5 .5 p, cylindric to cylindric-capitate, thin-walled, often
with a basal clamp, yellowish in Melzer’s solution, hyaline in KOH and with
scattered amorphous, embedded, dingy brown, incrusted material on the wall.
Trama of pileus obscurely duplex with a layer of upright hyphae blending into
a lower layer of interwoven hyphae (2.5-8 .0 p diam.), thin- and thick-walled,
with clamp connections, light ochraceous to dingy yellowish in Melzer’s solu¬
tion, light yellowish to nearly hyaline in KOH. Trama of the lamellae of inter¬
woven hyphae, mostly thick-walled, similar in all aspects to the lower pileus
trama.

Habit and habitat.— Usually in tightly packed imbricate clusters as a
saprophyte on the sides of logs, limbs, and stumps of hardwoods. Noted on
Quercus, Acer, Alnus, Betula and on wood in service, particularly structural
timbers. Fruiting in the fall in September, October, and November in Europe
and North America.

Observations.— The brown pileus, luminescent lamellae, and constant
presence of a short lateral stipe, combined with abundant, protruding cheilo¬
cystidia (Fig. 30) and the unique woolly-pubescent cuticle (Fig. 29) clearly
separate P. stipticus from all other species in the genus. The amyloid (blue)
spore, thick -walled hyphae of the trama of the lamellae and pileus, ability to
revive when moistened, and lignicolous habitat, combined with the smooth
(non-serrate) edges of the lamellae were used as central characters in erecting
Panellus as a genus in the Tricholomataceae. It is spelled “P. stypticus ” by
many authors and is named for its styptic or astringent properties.

Buller (1924) investigated the luminescence of P. stipticus and found
that the lamellae of the North American fruiting bodies are luminescent but
the phenomenon cannot be observed in European specimens. Twelve hours ex¬
posure of the gills to film in complete darkness yielded negative results with

1970

THE MICHIGAN BOTANIST

25

Fig. 32. P. patellaris (x 2) Photo by J. Lindsay.
Fig. 33. P. serotinus (x Vi) Photo by A. H. Smith.
Fig. 34. P. stipticus (X Vi) Photo by A. H. Smith.

26

THE MICHIGAN BOTANIST

Vol. 9

European material eliminating the possibility of very low levels of lumines¬
cence. Mycelium was examined by Buller (1924) and again luminescence of
the vegetative cells could only be demonstrated in North American material.
He compared spores and other morphological features of the fruiting bodies
on both continents but no additional constant differences could be found. He
concluded that two physiological races could be discerned and that both were
variants of P. stipticus. The luminescence is usually visible after several min¬
utes in a dark room and best observed in temperatures from 10 to 25°C.

Material examined. -U.S.A.: Idaho; O. K. Miller 2937 (BFDL). Louisiana; A. Laska
2639 (BFDL). Maryland; O. K. Miller 3668 (BFDL), 5100 (BFDL), 5102 (BFDL), 7119
(BFDL). Michigan; O. K. Miller 43 (BFDL), 1308 (BFDL), A. H. Smith 75145 (MICH).
New York; O. K. Miller 3550 (BFDL), R. L. Gilbertson 6828 (BFDL). Washington D. C.;
O. K. Miller 4949 (BFDL).

5. Panellus ringens (Fr.) Romagnesi. Figs. 22-26, 36.

Bull. Soc. Mycol. Fr. 61: 38. 1945.

Lentinus ringens Fr. Epicr. Syst. Mycol. 396. 1836-38.

Panus salicinus Peck. N. Y. State Mus. Rep. 24: 77. 1872.

Panus ringens Fr. Hymen. Europe. 490. 1874.

Pileus (5-) 10-30 mm in diam., sessile, margin even to somewhat crenate,
sometimes irregularly striate (see Fig. 36), light purple to purple drab or lilac
colored with vinaceous tints (“purple-drab,” “dark purple drab”) fading in age
sometimes to “vinaceous fawn” with a conspicuous pallid pubescence which is
dense over the lateral attachment of the cap (Fig. 36) to the substrate; dried
material is “vinaceous gray” to “lilac gray” at point of attachment. Lamellae
radiating from the point of attachment, fawn to pink (“fawn color,” “vinace-
ous-fawn” to “vinaceous-pink”) often fading in age, reddish-brown when dried
(“mikado brown” to “verona brown”).

Spores (4.0-) 5. 0-7.0 X 1. 2-2.0 p, oblong to allantoid, amyloid (blue in
Melzer’s solution), hyaline in KOH, often hard to find. Basidia 17-27 X
(2.5-)3.5-6.3 p, narrowly clavate, 4-spored. Pleurocystidia not found. Cheilo-
cystidia 24-38 X 4.2-5 .5 p, clavate, cylindric, fusiform, thin-walled, hyaline in
Melzer’s solution and KOH, infrequent. Cuticle of pileus of single, tangled, or
fasciculate thin- and thick-walled clamped hyphae 2.5-5 .0 p diam., hyaline to
light yellowish in Melzer’s solution and KOH. Trama of pileus and lamellae of
interwoven, thick-walled and infrequent thin-walled hyphae 3.5-7 .0 p diam.,
light yellowish in Melzer’s, hyaline in KOH. Subhymenium a thin layer of
tightly interwoven hyphae 1.0-2. 5 p diam.

Habit, habitat, and distribution.— Several to gregarious, often lining the
sides of sticks and limbs on the ground, sometimes nearly imbricate. Recorded
on species of Alnus, Betula, and Salix in Europe and North America but in
Europe it is also occasionally found on conifers. It is widespread in North
America, from Maine to Idaho and north into Canada. Fruiting in late summer
and fall but the persistent sporophores have been collected throughout the
winter.

Observations.— P. ringens is closely related to P. violaceofulvus but has
smaller spores, smaller basidia, lacks a cellular hymenopodium, and does not

1970

THE MICHIGAN BOTANIST

27

Fig. 35. P. mitis (X 1) Photo by A. H. Smith.
Fig. 36. P. ringens (x 6) Photo by O. K. Miller.

mMtmliim

28

THE MICHIGAN BOTANIST

Vol. 9

develop a venose hymenium between the lamellae. However, both species are
purple to violet or violet tinted. P. ringens has been mistakenly identified as P.
violaceofulvus (Batsch ex Fr.) Sing, by American authors and is discussed fur¬
ther by Miller (1967). The two collections cited by Coker (1944) were ex¬
amined and both have the short spores and other characters of P. ringens.

Material examined.— Europe: Finland; Karsten Jan. 2, 1866 (NY). Italy; G.
Bresadola, March 13, 1896 (BPI). Sweden; Bruun and Lundell, Fungi Exsiccati Suecici,
231 (BPI), A. Melderis 66411 (DAOM). Switzerland; Quelet 1890 (NY). North America:
Canada. Ontario; J. Dearness (Peck 203) (MICH), J. W. Groves 17446 (MICH), J. Dear¬
ness 203 (NY), Feb. 1890 (NY), Elies & Everhart 2503 (NY), R. F. Cain 21551 (NY),
80577 (DAOM), A. H. Smith 80576 (DAOM). North America: U.S.A.: Colorado; Seaver
& Bethel Aug. 30, 1910 (NY). Idaho; O. K. Miller 2908 (BFDL), 2933 (BFDL), 2937
(BFDL), 4893 (BFDL), J. R. Weir 5541 (NY), J. R. Weir & A. S. Rhodes 16145 (BPI),
16146 (BPI). Maine; Murrill 2017 (NY), 2026 (NY). Maryland; O. K. Miller & J. Lindsay
OKM 5026 (BFDL). New York. C. H. Peck, Center, New York (type of Panus salicinus
Pk.) (NYS), O. K. Miller 3594 (BFDL), C. L. Shear 108 (NY). Pennsylvania; Herbst
(NY).

6. Panellus violaceofulvus (Batsch ex Fr.) Sing.

Ann. Mycol. 34: 335. 1936.

Pileus 3-15 mm diam., sessile or with a short rudimentary stipe; margin
even to slightly wavy, inrolled at first, dry, brown tinted violet, or darker
violet-brown with a conspicuous white, woolly pubescence. Lamellae sub-
distant, sometimes venose, pale violet to brownish in age.

Spores 6.5-10.0 X 2.2-3.5 /a, cylindric to somewhat allantoid, amyloid.
Basidia 25-35 X 4.5-6.0 p, clavate, 4-spored. Cystidia, none seen. Cuticle of
pileus of interwoven or loosely fasciculate, thick -walled, clamped hyphae
3.5-6.0 diam., light yellow brown in Melzer’s, hyaline in KOH. Trama of
pileus and lamellae of interwoven thick -walled, clamped hyphae 3.5-10.0 p
diam., light yellowish in Melzer’s, hyaline in KOH. Subhymenium a cellular
hymenopodium, particularly evident near gill attachment to pileus.

Habit, habitat, and distribution.— Several to many, often somewhat im¬
bricate, on small limbs and logs. The only collection examined from North
America was on balsam fir ( Abies balsamea) from Anticosti Island in eastern
Quebec, Canada. In Europe it has been reported on silver fir ( Abies alba or A.
pectinata of some authors) and other conifers, usually at high elevations in the
Alps.

Observations.— Collections identified as Panus, Resupinatus, and Panellus
violaceofulvus have been examined from a number of herbaria (MICH, QFB,
BPI, BFDL, DAOM and NY). All North American collections which I have
studied with one exception discussed above are Panellus ringens (Fr.) Sing,
and are easily identified by the short spores, absence of a cellular hymenopod¬
ium, short basidia, and their occurrence on species of hardwood, particularly
Alnus, Salix, and Betula. It is quite likely that P. violaceofulvus will be en¬
countered frequently in the northern boreal forests.

Material examined. -Canada: Quebec; G. B. Ouellette 5083 (QFB). Europe: Aus¬
tria; Magmus (NY), Italy; Bresadola 1896 (NY). Switzerland; 405. Les Champ, du Jura et
des Vosges (NY).

1970

THE MICHIGAN BOTANIST

29

EXCLUDED SPECIES

The following list deals only with species found in North America which have been
referred to Panellus.

Panellus dealbatus (Berk.) Murrill. N. Am. Flora 9: 245-246. 1915.

^Asterotus dealbatus (Berk.) Sing. Mycologia 35: 161. 1943.

Panellus eugrammus (Mont.) Murrill. N. Am. Flora 9: 245. 1915.

-Nothopanus eugrammus (Mont.) Sing. Mycologia 36: 364. 1944.

Panellus haematopus (Berk.) Murrill. N. Am. Flora 9: 246. 1915.

-Panus suavissimus (Fr.) Sing.

Panellus mirabilis Singer. Mycologia 47: 770-771. 1955.

=Mycena mirabilis (Sing.) O. K. Miller comb. nov.

The type specimens of this minute fungus have central or slightly eccentric stipes
which recurve very sharply from the host. It does not revive as the species of Panellus do,
and the hyphae of the trama of the lamellae are thin-walled. In addition, the spores are
very weakly amyloid, elliptical, pointed at one end, 6.0-9. 5 x 2. 5-4.0 p, and not the
usual allantoid to narrowly elliptical spores of Panellus. The cheilocystidia are of two
types: (A) Clavate end-cells with dingy yellow contents 20-30 X 8-10 p, thin-walled,
often embedded or protruding slightly; (B) Narrowly clavate to cylindric cells with
branched projections 18-35 X 2-4 p, thin-walled, hyaline. Pleurocystidia 21-35 X 5-7 p,
clavate to fusiform, with a short blunt tapered apex, thin-walled, and hyaline. In addi¬
tion, the subcutis and trama of the pileus is deep red in Melzer’s solution. The above
combination of characters is found only in the genus Mycena, Eumycena in the section
Corticolae. The cheilocystidia resemble those of Mycena corticalis A. H. Smith, while the
spores are very much like those of Mycena madronicola A. H. Smith (Smith, 1947, Fig. 3).

Panellus vulpinus (Sow.) Murrill. N. Am. Flora 9: 246. 1915.

=Lentinellus vulpinus (Sow. ex Fr.) Kuhn et R. Maire, Bull. Soc. Myc. Fr. 61: 44. 1946.

SUMMARY

Six species of Panellus are described from North America. A number of new char¬
acters have been found or used to clearly delimit species using material in either the fresh
or dried condition. New ecological data, hosts, and wider distributions have been pre¬
sented for many species. A key to the species is provided along with camera lucida draw¬
ings and photomicrographs to illustrate the important characters. Macrophotographs of
five species are included.

ACKNOWLEDGMENTS

I would like to thank Dr. Alexander H. Smith for notes, photographs, and helpful
criticism during the preparation of this paper. In addition, I would like to acknowledge
the technical help of Miss Linnea Stewart and Mr. John Lindsay. The curators of the
following herbaria very kindly loaned material during the course of the study: The Na¬
tional Fungus Collections; The University of Michigan Herbarium; The Mycological Her¬
barium, Ottawa, Canada; and the Laboratoire de Biologie Forestiere, Quebec, Canada.

LITERATURE CITED

Buller, A. H. R. 1924. Researches on Fungi. Vol. III. Longmans, Green, and Co., Lon¬
don. 611 pp.

Coker, W. C. 1944. The smaller species of Pleurotus in North Carolina. Jour. Elisha
Mitchell Sci. Soc. 60: 71-95.

Groves, J. W. 1962. Edible and Poisonous Mushrooms of Canada. Res. Branch Canada
Dep. Agr. Publ. 1112. 298 pp.

30

THE MICHIGAN BOTANIST

Vol. 9

Kauffman, C. H. 1918. The Agaricaceae of Michigan. Vol. I. Mich. Geol. & Biol. Surv.
Publ. 26 (Biol. Ser. 5). 924 pp.

Krieger, L. C. C. 1947. The Mushroom Handbook. Macmillan, New York. 5 38 pp. + 32 pi.
Kiihner, R., & H. Romagnesi. 1953. Flore Analytique des Champignons Superieurs.
Masson et Cie, Paris. 556 pp.

Miller, O. K. 1967. Notes on western fungi. I. Mycologia 59: 504-512.

- . 1968. Interesting fungi of the St. Elias Mountains, Yukon Territory, and

adjacent Alaska. Mycologia 60: 1190-1203.

Moser, M. 1967. Kleine Kryptogamenflora. Basidiomyceten II. Die Rohrlinge und Blatter-
pilze. Gustav Fischer Verlag, Stuttgart. 443 pp.

Murrill, W. A. 1915. Panellus, N. Am. Flora 9: 244-247.

Reid, D. A. 1964 [“1963”]. Notes on some fungi of Michigan-I. Persoonia 3: 97-154.
Ridgway, R. 1912. Color Standards and Color Nomenclature. Washington, D. C. 43 pp. +
53 pi.

Singer, Rolf. 1962. The Agaricales in Modern Taxonomy. 2nd ed. J. Cramer, Weinheim.
915 pp. + 73 pi.

Smith, A. H. 1947. North American Species of Mycena. Univ. Mich. Press, Ann Arbor.
521 pp. + 99 pi.

A NEW CLITOCYBE FROM MICHIGAN

Howard E. Bigelow and Alexander H. Smith
Department of Botany, University of Massachusetts, Amherst,
and Herbarium, The University of Michigan, Ann Arbor

One studying the fleshy fungi soon becomes almost immune to surprises,
but we must confess that a fungus collected several times in the spring of
1969 in a rose house in Mt. Clemens, Michigan, shocked us both. The basidio-
carps of the species were very much like those of the Clitocybe irina group
(Bigelow & Smith, Brittonia 21: 144-177. 1969), but the color of the spore
deposit was dark vinaceous brown. A survey of the literature did not reveal
the presence of such a species in any region of the world. Hence we describe
the fungus here as new and take pleasure in dedicating it to Prof. E. S.
Beneke of Michigan State University, who first recognized it as a mushroom
extraordinary.

Clitocybe benekei sp. nov. Figs. 1-3.

Pileus 8-15 cm latus, convexus, margine incurvatus ad crenatus, super¬
ficies glaber, humidus, hygrophanus, pallide incarnato-bubalinus turn sordido-
pallidus. Caro albidus. Odor et sapor ingratus. Lamellae adnatae vel breviter
decurrentes, confertae, latae, pallide turn vinaceo-brunneae. Stipes 6-12 cm
longus, apice 1-3 cm crassus, aequalis vel basi bulbosus, solidus, fibrosus,
superficies asper, sordido-pallidus. Sporae 5.5-7 X 4.5-5 /i, verrucosae, in
cumulo vinaceo-brunneae. Typum legit E. S. Beneke (A. H. Smith no. 77176),
Mt. Clemens, Mich. (Holotypus MICH; isotypus MSC).

1970

THE MICHIGAN BOTANIST

31

Fig. 1. Clitocybe benekei, slightly reduced. Type collection.

32

THE MICHIGAN BOTANIST

Vol. 9

Pileus 8-15 cm broad, broadly convex with an incurved margin, becom¬
ing wavy to crenate or scalloped in age, surface glabrous, moist, hygrophan-
ous, color a watery pale pinkish buff to cartridge buff when moist, fading to
dingy pallid, in age often brownish pallid from spores. Context white, not
changing overnight after being broken or bruised, floccose; odor and taste dis¬
agreeable. FeS04— no reaction.

Lamellae adnate to short decurrent, seceding but tearing the context
when separating, close, broad, pallid but finally becoming “wood brown,” not
discoloring overnight where bruised, edges even, thin.

Stipe 6-12 cm long, 1-3 cm thick at apex, equal or the base bulbous,
solid, fibrous, whitish within, unchanging or merely slightly dingy from bruis¬
ing; surface uneven, pallid like the faded pileus. Veil absent.

Spores 5.5-7 X 4.5-5 /u, elliptic, distinctly verrucose (warts up to 0.5 ii
high), hyaline in KOH, in Melzer’s solution fresh spores showing rare amyloid
particles within, particles not evident in dried spores revived in Melzer’s but
the ornamentation appearing weakly amyloid in some, deposit “wood brown”
to “army brown.”

Basidia 18-33 X 6.5-9.5 /i, 4-spored. Cystidia not differentiated. Pileus
with a surface layer of somewhat gelatinous hyphae in KOH as viewed under
the microscope, the hyphae cylindric and 2-5 ju wide; pileus tramal hyphae
cylindric to somewhat inflated, 6.5-13 ju wide, cells often short. Hymeno-
phoral trama of hyphae parallel to subparallel, cylindric to somewhat inflated,
2-14 n wide. Clamp connections present.

Fig. 2. Clitocybe benekei, x 3/4. Type collection.

1970

THE MICHIGAN BOTANIST

33

Cespitose on mulch in rose beds in greenhouses near Mt. Clemens,
Macomb County, Michigan, March and April, 1965; type Smith 77176
(MICH), comm, by Dr. E. S. Beneke.

The type collection consisted of a market basket full of basidiocarps and
it is from these that our description is drawn and the photographs taken. In
general aspect Clitocybe benekei appears to belong to the group of species
centered around Clitocybe irina, but the dark-colored spore deposit and rela¬
tively coarse spore ornamentation at once distinguish it. The habitat and man¬
ner of fruiting are typical for members of section Verruculosae of Clitocybe
such as C. subconnexa, the type of the section. Members of this section are
often found in quantity fruiting on mulch or on compost heaps out of doors.

The unusual features are the dark vinaceous-brown spore deposit which
becomes gray-brown as it air dries, and the slight amyloid reaction of the
spore ornamentation. When fresh the spore deposit has the color of that of
some species of Agaricus. The rare amyloid particles found within freshly de¬
posited spores when mounted in Melzer’s solution have not been reported
previously for any taxon in Clitocybe, but we do not regard them as an aid in
identifying the species because they are so rare and easily overlooked. The
obscure amyloid reaciton of the ornamentation on the spore wall appears to
be more noteworthy at least as far as postulating relationships of the section
to other groups in the Tricholomataceae is concerned. It may very well be
true that the difference between amyloid and non-amyloid spores is not of
major significance in establishing phylogenetic lines.

Fig. 3. Clitocybe benekei, x 3/8. Type Collection.

34

THE MICHIGAN BOTANIST

Vol. 9

CHARLES HENRY SWIFT-1881-19661

Walter L. Palmer

School of Medicine, The University of Chicago,

Chicago, Illinois

With the death of Charles H. Swift on November 17, 1966, from a
ruptured abdominal aneurysm, the world lost one of its last naturalists and
scholars reared in the tradition of the nineteenth century; his many friends
lost one of their most charming companions.

Charles Swift was born in the bluegrass country of Kentucky, August
17, 1881, the son of extraordinary parents. His father’s family was of English
(Lincolnshire) stock; his grandfather was a teacher and city librarian of Lex¬
ington, Kentucky, renowned for his encyclopedic knowledge. Charlie’s father,
William Swift (1834-1914), was a perpetual student and teacher with degrees
in law (LL.B., Lexington) and medicine (M.D., Lexington); he become profes¬
sor of all the sciences in Christian College at Millersburg, Kentucky, occupy¬
ing, as someone has said, not a chair but a settee! Charlie’s instruction in
science began as a small child when his father taught him the lore of trees,
flowers, and birds and inspired his son with a love of knowledge. The Louis¬
ville public schools provided the formal education culminating in a sort of
collegiate high school affiliation with the old University of Louisville from
which, on graduation, Charlie was awarded the A.B. degree.

The Swift family had the Chautauqua habit in the nineties, from which
followed at least two important consequences. The first of these was acquaint¬
ance with William Rainey Harper and a decision to move to Chicago so that
the children could attend the new University of Chicago. The idea of moving
to Chicago and the decision to do so is ascribed to Charlie’s mother, Helen
Mar Kelly Swift (1840-1929), a strong-minded woman of Scotch-Irish ancestry
who was determined to obtain the best possible educational opportunities for her
son Charlie and her daughter Verger Rebecca Swift. The mother exerted a
profound influence on both of her children until the time of her death at the
age of 88 years.

Charlie matriculated at the University of Chicago in 1899 and received
an A.B. in History with the minor in Latin and Greek in 1903. His early inter¬
est in nature then led him to the sciences; he qualified for and in 1906 re¬
ceived a third bachelor’s degree, a B.S. in Botany and in Anatomy. The inter¬
est in anatomy led him to the broad field of medicine and resulted in 1910 in
an M.D. degree from Rush Medical College. Dr. Swift never practised medicine
but he derived great satisfaction from the fact that he was a qualified physi¬
cian and proudly renewed his license annually in order to be able to prescribe
for his friends. Upon his graduation from medical school Dr. Swift became an

Expanded version of a notice which appeared in the Anatomical Record 165:
268-269 (1969); used by permission of the Wistar Institute Press.

1970

THE MICHIGAN BOTANIST

35

Instructor in Anatomy at the University of Chicago; in 1913 he was awarded
the Ph.D. degree with anatomy as the principal subject and botany secondary.
In 1918 he was promoted to Assistant Professor. Advancement to Associate
Professorship came in 1929 and retirement as Associate Professor Emeritus in
1946.

Dr. Swift began his research during his Fellowship in Anatomy in
1908-1909 and continued it after his appointment as Instructor. His papers on
Primordial Germ Cells in the Chick and the Yolk Nucleus of Balbiani were
published in the American Journal of Anatomy in 1914, 1915, and 1916.
Appreciation was immediate and general; within a year this work was famous
in Europe and America! Recognized at the time as a classic study, it is still so
rated more than 50 years after its publication.2’3

During World War I, Dr. Swift’s heavy teaching schedule interrupted his
research. He abhorred the lecture system and even with the relatively large
classes, preferred, as I remember him in 1917, to spend his time at the dissect¬
ing tables, sitting on a stool, discoursing on anatomy. His photographic mem¬
ory enabled him to quote at length from Gray’s Anatomy, Cunningham,
Spalteholtz, or any other treatise of importance. In the early 1920’s Dr. Swift
organized the first course in the University on the History of Medicine. It was
an immediate success and was continued until his retirement in 1946.

Dr. Swift’s knowledge encompassed almost all significant subjects and
some not so significant, such as the railroad timetables of the United States
and Canada. He is known to have worn out one Encyclopedia Britannica and
started on a second! His memory for faces and names was excellent; he
possessed the gift of instant recall so that, meeting a former student any¬
where, he would recognize him immediately and give his name and class. The
mutual affection between students and teacher was evidenced by the appreci¬
ative remarks made by his former students at his retirement in 1946 and again
after his death in 1966.

Two fascinating questions arise. Why did Dr. Swift fail to continue his
investigative career and why did he fail advancement to full professorship?
Some of his friends have thought the explanation to be that the teaching and
administrative responsibilities of the war years drew him away from research;
this is not an adequate answer. Charlie possessed abundant intellectual curiosi¬
ty, vigor, and ability. His wide-ranging mind was filled with his studies of his¬
tory, botany, ornithology, and science; everything interested him. Pressure
for research surrounded him. In the history of the world, at least in modern
times, there has never been a department of anatomy with so many giants as
at Chicago in the first third or so of this century:2 Robert Russell Bensley
(1867-1956), Chairman of the Department from 1905 to 1933, has been
ranked as the greatest of the American anatomists.4 George W. Bartelmez

William Bloom, personal communication.

3Lincoln V. Domm, personal communication; Basil C. H. Harvey, “After Forty-
Seven Years: Indian Summer,” Univ. Chicago Mag. December 1946, pp. 5-6.

4Basil C. H. Harvey, remarks at the memorial service for R. R. Bensley, November
13, 1956; Howard Wakefield, personal communication.

36

THE MICHIGAN BOTANIST

Vol. 9

l ig. 1. Charles Henry Swift (1881-1966). Photograph by Joseph Merante.

1970

THE MICHIGAN BOTANIST

37

(1885-1967) was one of the world’s leading investigators of human embryolo¬
gy;5 C. Judson Herrick (1868-1960), a recognized genius, devoted his life to
the study of the brain and nervous system, published at least 13 books and
200 scientific scholarly papers, and remained active until his death at the age
of 92 years; A. A. Maximow (1874-1928), even before his migration from
Russia to Chicago in 1922, was described by Florence Sabin as the greatest
histologist in the world.2 Basil C. H. Harvey (1875-1958) in the years from
1901 to 1917 rose rapidly through the five faculty ranks from Assistant to
Professor before being called into military service. Upon his return Dr. Harvey
resumed his research with Dr. Bensley and his beloved teaching. The Universi¬
ty soon induced him to accept the position of Dean of Medical Students, an
assignment he filled with distinction and yet found time to write on topics in
anatomy and medical education.

Dr. Swift devoted himself to the teaching of gross anatomy and yet
balked at investigation and writing. Why? It is tempting to suggest that he
reacted against the tremendous pressure for research, as did most of his stu¬
dents. Charlie himself alleged in later years that he had failed to continue his
early superb studies because of his clumsiness with the microtome and the
punishment he inflicted on his own fingers and thumbs. Neither he nor the
department could afford the luxury of a technician and so the investigation
ceased. This explanation does not account for his failure to carry out other
types of creative work or to write on the history of medicine he knew so
well. Charlie was not a man to yield to pressure from any source. He knew
nothing of grantmanship, of the “publish or perish” philosophy of the modern
university, although in a sense he was an early victim. He loved knowledge for
its own sake, as something to be relished and savored, as some people prize a
fine wine; he would not seek knowledge for money or promotion, but only
for enjoyment. He delighted in every new fact he learned. I can picture in my
mind the merry way he would have ridiculed and rejected any suggestion that
he prostitute his talents for forty pieces of silver! And yet to many of the
academic idealists in science in both the past and present generations, no man
merits full professorship without continued activity in research. Dr. Swift
never complained, even to his closest associates. Nevertheless, it is distressing
to think of the number of lesser men to whom the accolade of full professor¬
ship has been given!

Reference was made earlier to the Chautauqua habit of the Swift family
in the nineties. This led to participation in a similar colony at Bay View,
Michigan, established under the auspices of a group of Methodists. The Swift
family bought a cottage; Charlie took an active role in the Bay View enter¬
prise and returned every summer for the rest of his life. It was there that he
met Juliana Streid of Washington, Illinois, the sister of Charlie’s close friend
and classmate, Joseph Streid, who had died in his senior year at Rush. Charlie
and Juliana were married in 1920; Juliana survives him.

5George W. Corner, “George W. Bartelmez,” Anat. Record 161: 533-534. 1968.

38

THE MICHIGAN BOTANIST

Vol. 9

Charlie’s father had been an artillery officer in the Confederate Army
serving with Bragg, Morgan the Raider, and Breckenridge. As a boy and as a
youth, Charlie attended the annual reunions with his father and listened with
rapt attention to the tales of the old soldiers. From this grew an avid interest
in the history of the conflict. He read every book he could find on the War
and its leaders; with his marvelous memory he could discourse at length. I
think his favorite work was Douglas Southall Freeman’s three-volume Lee's
Lieutenants - A Study in Command (Scribners, 1943), given him by Juliana in
1942 and graciously passed on by her to me after his death. Charlie is said to
have been an especial authority on the Battle of Gettysburg, but he knew the
disposition of troops in every battle and the comments of the generals after¬
wards. His appraisals of the War, the armies, the leaders, and the issues at
stake, were all remarkably objective. Dr. Swift was almost equally familiar
with the other great wars of history. One day I casually asked him what had
happened to Alexander the Great in India; a look of incredulity came over his
face and he proceeded to give me a detailed account of Alexander’s last cam¬
paign.

Dr. Swift’s keen interest in ornithology continued throughout his life.
He kept a diary dating the first appearance of the various birds each spring as
they came through Chicago. Frequent expeditions were made almost until the
last to the sand dunes and other areas favored by the bird watchers. The num¬
bers of birds he would identify from his apartment window amazed his
friends. Whether he talked of a bird, a tree, a bush, or a flower, Charlie could
expound on its common and scientific names, its habits and characteristics.

Economics and politics were of much less interest to him than were his¬
tory and science, but at times in a discussion he would begin with Adam Bede
or before, and end by giving his views of contemporary problems. A Democrat
born and raised, he learned to vote the Republican ticket and laughingly re¬
marked that, were his father to know that he had done so, he would turn
over in his grave!

Active, alert, energetic until the day of his death at the age of 85,
Charlie Swift will be remembered by his friends, colleagues, and students for
his warm friendship, his encyclopedic knowledge, his gay wit, his keen sense
of humor, and his rare charm. He was large of heart, mind, stature, and talent.
We shall not see his like again.

CHARLES H. SWIFT’S BOTANICAL WORK IN MICHIGAN6

As soon as he came to the University of Chicago in 1899, Charles Swift
associated in the botany department with John Merle Coulter and Henry
Chandler Cowles. He immediately started learning the Chicago area plants.
Whether Coulter, who had been director of the Bay View Summer School
from 1893 to 1896, helped to influence the Swifts in their decision to sum¬
mer at Bay View, Michigan, I do not know. At any rate, from his first sum¬
mer at Bay View, in 1899, until the end, Charles Swift was devoted to the

6Appended by Edward G. Voss.

1970

THE MICHIGAN BOTANIST

39

s?

I)

Vi

~*N

v

€
c k

V *

^ c,
Sft’
<*

? \

,5v

u<$

5k

(N

£
ox) <y~>

r-^

t C/5

Two pages from the original F allass and Swift manuscript list of the Emmet County flora. The larger writing
:’s. (Reprinted from Asa Gray Bull. N. S. 3: 91. 1955)

40

THE MICHIGAN BOTANIST

Vol. 9

plants of that area. On July 1, 1903, he met Charles W. Fallass (1854-1 942), 7
an ardent botanist by avocation, who had moved to Petoskey in 1898 and who
had summered at Bay View since 1887. Both Fallass and Swift had been
botanizing that day west of Harbor Springs, and when they met in the field,
each with a vasculum over his shoulder, sharing their joy at having found the
rare Orchis rotundifolia, there began an acquaintance which lasted for forty
years.

The Fallass and Swift team explored together, at first by train or on
foot and later by auto, many parts of Emmet County and the Jack pine plains
of Cheboygan County. About 1917 Swift suggested that a list of the vascular
plants of this region be prepared. He drew up an annotated list of about 600
species and Fallass added another 200. Two pages from the original list (pre¬
sented to me in 1965 by Dr. Swift) are reproduced in Fig. 2. A few copies of
the list were made and distributed, and although it was never published it
became cited in various works on the flora of Michigan. Dr. Swift’s prime
interest was in identification; once he learned a plant and noted it in his
manual, he generally preserved no specimen. Mr. Fallass was the collector, and
his herbarium, now at Albion College (Albion, Michigan), includes almost all
of the few specimens actually collected by Swift. Two exceptions are the
types of some unusual forms of the purple-fringed orchid, Habenaria psycodes,
found by Dr. Swift at Round Fake near Bay View in July of 1913. These
were described, with full credit to Swift, by Mary M. Bryan (Ann. Missouri
Bot. Gard. 4: 37-42. 1917) as var. varians (lacking the central lobe of the lip)
and var. ecalcarata (lacking both spur and fringe on the lip); these specimens
are in the herbarium of the Missouri Botanical Garden.

From the time of my first correspondence with him in 1954 until my
last visit with him and Mrs. Swift at Bay View in 1965, it was clear to me not
only that Dr. Swift was a botanist intensely interested in the same local flora
as I— and one who had been actively studying it years before the nearby Uni¬
versity of Michigan Biological Station was founded in 1909!— but also that he
possessed those qualities of scholarship and charm which are so well portrayed
in the preceding sketch.

7See Edward G. Voss, “Charles W. Fallass (1854-1942), a Pioneer Michigan Botan¬
ist,” Asa Gray Bull. N. S. 3: 77-96. 1955.

1970

THE MICHIGAN BOTANIST

41

TYPE SPECIMENS OF FLOWERING PLANTS DESCRIBED
FROM LONG’S EXPEDITION TO THE SOURCE
OF THE ST. PETER’S RIVER IN 18231

Ronald L. Stuckey
College of Biological Sciences,

The Ohio State University, Columbus

In the herbarium of the Academy of Natural Sciences of Philadelphia is
a small collection of plants that once was a part of the large herbarium of the
Rev. Lewis David von Schweinitz. These specimens are the basis for Schwein-
itz’s catalogue of plants collected in the northwestern territory by Mr. Thomas
Say on Major Stephen H. Long’s Expedition in 1823 (Schweinitz, 1824).
Schweinitz enumerated 130 taxa including 11 new species and five new
varieties of flowering plants. The present paper summarizes the historical back¬
ground for these collections, and enumerates the existing type specimens with
their limited data.

THE EXPEDITION

As recorded by Keating (1824), the Honorable J. C. Calhoun, Secretary
of War, ordered an expedition to the source of the St. Peter’s River (Minne¬
sota River) in 1823 under the command of Stephen H. Long, Major, United
States Topographical Engineers. The United States government wished to as¬
semble more information for a topographical description of the region, to
ascertain the latitude and longitude of all the remarkable points, and to ex¬
amine and describe its animal, vegetable, and mineral productions. The coun¬
try to be explored was that bounded by the Missouri and Mississippi Rivers
and the northern boundary of the United States. The exploring party left Phil¬
adelphia on 30 April 1823, and traveled westward across Pennsylvania and
Maryland to Wheeling, then through central Ohio to Fort Wayne and Chicago.
From Chicago the expedition proceeded across Wisconsin Territory to Prairie
du Chien, up the Mississippi River to Fort St. Anthony, then to the source of
the St. Peter’s River, down the Red River past the International Boundary and
on to Winnepeek Lake [Lake Winnipeg, Manitoba] . Turning eastward the ex¬
ploring party went via Lake of the Woods, Rainy Lake, Falls of Kakabeka, the
north shore of Lake Superior, down through Lake Huron, across Lake Erie,
and homeward— reaching Philadelphia 26 October 1823. A detailed map of the
route traveled appears in Keating (1824), and a more general map is in Barber
(1928) which was later reprinted in Weiss and Ziegler (1931, p. 189).

Dr. Edwin James, botanist of a previous expedition under the command
of Major Long to the Rocky Mountains in 1819-20, was appointed botanist,
geologist, and physician to the 1823 expedition. Because he failed to receive
his instructions for making connections with the party at Wheeling or Colum-

^Contribution from the Botany Program (Paper No. 764) and the Herbarium of
The Ohio State University.

42

THE MICHIGAN BOTANIST

Vol. 9

bus, he waited for them at Pittsburgh while the party was passing through
Wheeling. By the time James received the communication he could not catch
the party. Accordingly, Thomas Say, zoologist and antiquary on the expedi¬
tion, undertook to collect such plants as might appear to him interesting. Ac¬
cording to Brendel (1879) several boxes containing botanical collections dis¬
patched during the expedition were lost. Brendel’s undocumented statement
has not been commented upon or documented by later botanical historians.
However, the plants that were brought back were available for study. Thomas
Nuttall, perhaps the most noted naturalist, explorer, and botanist of the time,
applied successfully to Major Long for the opportunity of preparing the
botanical report from the dried specimens. As discussed by Graustein (1967,
p. 195), when it came time for Nuttall to sail for England, in about mid-
November of 1823, he had merely begun the work on the plants. He returned
the materials, with the intent of working on them upon his return in the
spring. When no word had been heard from Nuttall by the first of the follow¬
ing July, Keating arranged for Schweinitz to prepare the plant list and descrip¬
tions. Within about four months, Schweinitz organized the plants, and the
complete report was published before the end of 1824 (Keating, 1824).

SCHWEINITZ’S CATALOGUE

Two published versions of Schweinitz’s Catalogue appeared, the first
printed in Philadelphia in 1824 and the second printed in London in 1825.
Both editions contain exactly the same botanical information. However, the
type was reset and the pagination changed in the second printing. The names
of the new taxa date from the 1824 edition published as Section II in the
Appendix with pages numbered consecutively from Keating’s Narrative. In the
1825 printing, the Appendix begins with page one.

In the introduction to the Catalogue, Schweinitz points out that Nuttall,
who had begun the identifications and the writing of the text, had not re¬
turned from Europe in time to complete the work. Schweinitz reveals that he
undertook the task with “great diffidence and sincere regret.” Although the
published introduction differs from that in Schweinitz’s original manuscript
copy in the library of the Academy of Natural Sciences, both versions convey
essentially the same information.

One-hundred thirty plant taxa are listed. Schweinitz relied on Pursh’s
Flora (“1814” [1813]) and Nuttall’s Genera (1818), as his primary sources
for the identification of the plants. References and page numbers to these
floras usually accompany the names of the plants. Locality information is
lacking for many of the species. However, in certain cases, if Schweinitz knew
the species, he commented on its occurrence or abundance elsewhere, such as
in the eastern or southeastern United States or in Europe— geographical areas
where he had had previous experience with the flora. The localities mentioned
especially for the northwest were places where plants were obtained along the
route between Fort Wayne and the north shore of Lake Superior. Those local¬
ities most often cited were Prairies of the St.- Peter and Red Rivers, Lake of
the Woods, Falls of Kakabeka, and the north shore of Lake Superior.

1970

THE MICHIGAN BOTANIST

43

According to the introduction in Schweinitz’s Catalogue (1824), and as
mentioned by Upham (1884) and Graustein (1967), the first five plants were
named by Nuttall. Four of the first five entries bear names given to them by
Nuttall. According to Schweinitz number four, “Cyperus *alterni floats,” was
“labelled Cyperus by Mr. Nuttall,” but Schweinitz states that he himself pre¬
pared the description and I assume he also provided the epithet. Rodgers
(1942, footnote, p. 53) believed that Torrey aided Schweinitz in the prepara¬
tion of the Catalogue, but there is no indication in the Catalogue itself or in
the published correspondence between Schweinitz and Torrey (Shear &
Stevens, 1921), that Torrey helped Schweinitz in organizing or naming the
plants from Long’s 1823 expedition.

In the published version of the Catalogue, each plant (taxon) bears a
number. Some of the type specimens and other specimens in the collection
also have numbers on their labels. Each plant specimen number is one digit
lower than the published number in the Catalogue, with one type, specimen
number 5, Eriophorum angustifolium ft *megastachyon, being an exception.
This discrepancy in numbering can be explained after comparing the manu¬
script version of the Catalogue with the published version. The former does
not list Limnetis glabra, but it was inserted in the latter as number 6. Those
taxa listed after number 6, therefore, have numbers one digit higher than the
numbers on the specimen labels. Nuttall’s new Eriophorum, specimen number
5, was unaffected by this insertion. In the manuscript version of Schweinitz’s
Catalogue the taxa are not numbered and Nuttall’s description of Eriophorum
angustifolium 0 *megastachyon is missing, both items evidently having been
added at the time of printing.

THE TYPE SPECIMENS

After examining some of the type specimens, I realized that certain past
and recent monographers were either unaware of these species and their type
specimens at the Academy or had overlooked these specimens in their studies
even after annotating them. To cite one example, M. L. Fernald annotated the
type specimen of Eriophorum angustifolium (3 *megastachyon as Eriophorum
viridi-carinatum (Engelm.) Fernald, but did not mark it as a type, did not
treat it in synonymy, or even mention it in his treatment of the genus for
North America (Rhodora 7: 81-92. 1905).

In January of 1969 while at the Academy Herbarium, I undertook to
locate the type specimens collected on Long’s 1823 expedition, and to corre¬
late any information on their labels with Schweinitz’s published Catalogue,
Keating’s Narrative, Schweinitz’s original handwritten manuscript of the Cata¬
logue, and the Systematically Arranged Catalogue of the Schweinitz Herbari¬
um (Schweinitz, unpublished and undated), the latter two being in the manu¬
script collection in the Academy’s library. Thirteen type specimens were
found, representing all the new taxa except Crataegus flexuosus, Vicia triden-
tata, and Hieracium canadense f3 scabrum.

In the annotated list below, the data on the labels of these type speci¬
mens are quoted. As described by Pennell (1935) and Stuckey (1967), the

44

THE MICHIGAN BOTANIST

Vol. 9

data with the specimens in Schweinitz’s herbarium are meager and cryptic.
The specimens taken on Long’s Expedition also have cryptic data, consisting
of only the name of the species, a notation of the fact that they were taken
on the Long Expedition, and sometimes a number and general location. The
handwriting, unless otherwise noted, is that of Schweinitz. Data following
“Hab.” are from Schweinitz’s Catalogue. Where localities are given in the Cata¬
logue or on the specimen label, I have added the approximate date or dates
when the plant was collected, based on the expedition’s itinerary given in the
Narrative by Keating. All of the taxa are listed in Schweinitz’s Herbarium
Catalogue (unpublished) except Apocynum androsaemifolium (3 pubescens,
Ranunculus filiformis (3 hispidus, and Vernonia corymbosa, but unfortunately
no additional data can be supplied from this Catalogue, as most of the identi¬
fying notes accompanying the entries read simply, “Long’s Exped.”

ANNOTATED LIST OF NEW TAXA2 DESCRIBED BY
SCHWEINITZ FROM LONG’S 1823 EXPEDITION

4. Cyperus * alterniflorus Schwein. in Keat., Narrative ... p. 381. 1824. p.

106-107. 1825. = C. schweinitzii Torrey
“Cyperus alterniflorus LvS Longs Exped. Winnipeg Lake” 18-19 Aug.

Hab. No information.

Because of the existence of Cyperus alterniflorus R. Br. (Prodr. 216.
1810), Torrey renamed C. alterniflorus Schwein. in Keat. as C. schwein¬
itzii based on the Schweinitz type (Ann. Lyc. Nat. Hist. N. Y. 3: 276.
1836).

5. Eriophorum angustifolium [Honckeny] (3 *megastachyon Nutt, in

Schwein. in Keat., Narrative . . . p. 381-382. 1824. p. 107-108. 1825.
= E. viridi-carinatum (Engelm.) Fernald (det. M. L. F[ernald], with¬
out date).

“ ‘5’ Eriophorum angustifolium (3 *megastachyon [in Nuttall’s writing]
Long’s 2d Exped. [in Schweinitz’s writing]” 29 May-4 Jun.

Hab. Prairies between Fort Wayne and Lake Michigan.

Although the specimen was annotated by Fernald, he did not include
it in his list of specimen citations and omitted the name from the list of
synonyms (Rhodora 7: 89-91. 1905).

13. Triticum *pauciflorum Schwein. in Keat., Narrative ... p. 383. 1824. p.
108. 1825. = Agropyron trachycaulum (Link.) Make. (det. M. L.
F[ernald], 1932); fide Fernald, Rhodora 36: 417-420. 1934; Hitch¬
cock & Chase, Man. Grasses. 2nd. ed., p. 801. 1950.

“Triticum pauciflorum LvS [two illegible words]” 9-22 Jul.

Hab. Prairies of the St. Peter.

26. Apocynum androsaemifolium [L.] [3 pubescens Schwein. in Keat., Narra¬
tive . . . p. 384. 1824. p. 109-1 10. 1825.

9

One new combination is also included.

1970

THE MICHIGAN BOTANIST

45

“25 Apocynum androsaemifol[ium] (Long’s exp)”

Hab. No information.

This variety has not been listed in the Gray Card Index and was not
considered by Woodson (Ann. Missouri Bot. Gard. 17: 1-212. 1930; N.
Am. FI. 29: 103-192. 1938).

28. Gentiana *rubricaulis Schwein. in Keat., Narrative ... p. 384-385. 1824.
p. 110. 1825. IJames S. Pringle, 21 Jun 1965, but not cited by him
(Brittonia 19: 23-24. 1967).

“27 Gentiana *rubricaulis Long’s 2d Expd.” 9-22 Jul.

Hab. Prairies of St. Peter’s River.

43. Prunus *incana Schwein. in Keat., Narrative . . . p. 387. 1824. p. 111-112.
1825. = P. pumila Michaux; fide Wight, U. S. Dep. Agr. Bull. 179:
65. 1915.

“Prunus incana LvS Lake of the Woods Longs Exped.” 25-27 Aug.

Hab. Islands in the Lake of the Woods.

46. Crataegus *flexuosus Schwein. in Keat., Narrative . . . p. 387-388. 1824. p.
112. 1825.

Specimen not found. 31 Aug.-3 Sept.

Hab. Near Rainy Lake.

50. Rosa *sayi Schwein. in Keat., Narrative ... p. 388-389. 1824. p. 113.

1825. = R. acicularis Lindl. subsp. sayi (Schwein. in Keat.) W. H.
Lewis. (The second specimen det. W. H. Lewis, 1956); fide Lewis,
Brittonia 11: 19. 1959.

“49 Rosa *Sayi Long’s 2d Exped”

“ ‘Rosa Sayi LvS Long’s Exp’ (herb Schw) [Charles Pickering’s writing] ”
Hab. No information but Lewis (Brittonia 11: 20. 1959) quotes the loca¬
tion as “Mouth of the St. Peter River: 44° 53' 49” N; 93° 8' l" W.”; this
location has nothing to do with the type locality (Lewis, in litt., 24 Oct.
1969). In the Narrative, Keating (1825, Vol. 1, p. 255) wrote: “The eye
is charmed by the abundance of wild roses which are strewed over the
country, . . .” This location was between Prairie du Chien and Fort St.
Anthony.

51. Potentilla fruticosa [L.] var. floribunda (Pursh) Schwein. in Keat., Narra¬

tive . . .p. 389. 1824. p. 113. 1825.

This new combination is based on Potentilla floribunda Pursh, FI.
Am. Sept. 1: 355. “1814” [1813]. It has not been listed in the Gray
Card Index and was not considered by Rydberg (Mem. Bot. Columbia
Univ. 2: 1-233. 1898; N. Am. FI. 22: 239-533. 1908) or by Wolf (Bibl.
Bot. 71: 1-714. 1908).

57. Ranunculus filiformis [Michx.] (3 *hispidus Schwein. in Keat., Narra¬
tive . . . p. 389-390. 1824. p. 114. 1825. = R. reptans L.

“56 Ranunculus filiformis (3 *hispidus [Schweinitz’s writing] (Long’s
exp.) [Pickering’s writing] .”

46

THE MICHIGAN BOTANIST

Vol. 9

Hab. No information.

This variety is not considered by Benson (Am. Midi. Nat. 40: 1-261.
1948).

60. Stachys *velutina Schwein. in Keat., Narrative ... p. 390. 1824. p. 114.
1825. = Stachys ambigua Smith in Engl. (det. C. Epling, 1932); fide
Epling, Fedde repert. Sp. Nov. Beih. 80: 64. 1934.

“Stachys velutina Longs Exp.”

Hab. No information.

63. Melampyrum *brachiatum Schwein. in Keat., Narrative . . . p. 391. 1824.
p. 115. 1825. = M. lineare Desv. var. typicum [lineare] (Both det. M.
lineare Desv., F. W. P[ennell], 1933); fide Pennell, Monogr. Acad.
Nat. Sci. Phila. 1: 509. 1935.

“62 Melampyrum *brachiatum Long’s 2d Exped”

“Say Head of Miss [source of St. Peter’s River] Melampyrum aut nov
Gen” 22-25 Jul.

Hab. No information.

73. Vicia *tridentata Schwein. in Keat., Narrative ... p. 392-393. 1824. p.
116. 1825. = V. americana Muhl. ex Willd. var. minor Hook, fide
Hermann, U. S. Dep. Agr. Handb. 168: 84. 1960.

Specimen not found. 31 Aug.-3 Sept.

Hab. Rainy Lake.

80. Hieracium canadense [Michx.] j3 var. scabrum Schwein. in Keat., Narra¬

tive . . .p. 394. 1824. p. 117. 1825. = H. scabriusculum Schwein. in
Keat. var. scabrum (Schwein. in Keat.) LePage; fide LePage, Nat.
Canad. 87: 67. 1960.

Specimen not found.

Hab. No information.

LePage does not cite Schweinitz’s specimen.

81. Hieracium * scabriusculum Schwein. in Keat., Narrative . . . p. 394. 1824.

p. 117-118. 1825.

“80 Hieracium ^scabriusculum Long’s Exped” A second sheet bears simi¬
lar data but lacks the number 80 and the asterisk.

Hab. No information.

Although LePage annotated these specimens in 1959 as H. scabriu¬
sculum, he did not mark either one as a type and did not cite them in his
publication (Nat. Canad. 87: 62-71. 1960).

83. Vernonia *corymbosa Schwein. in Keat., Narrative . . . p. 394-395. p. 118.
1825. = V. corymbosa Schwein. in Keat.; fide Gleason, Bull. N. Y.
Bot. Gard. 4: 209. 1906.

“82 Vernonia *corymbosa Long’s 2d Expedit.”

Hab. No information.

86. Erigeron canadense [L.] grandifiorum Schwein. in Keat., Narra¬
tive ... p. 395. 1824. p. 118-119. 1825. = Conyza canadensis (L.)

1970

THE MICHIGAN BOTANIST

47

Cronquist; in accordance with the viewpoint of Cronquist, Bull. Tor-
rey Bot. Club 70: 632. 1943.

“85 Erigeron canadense grandiflorum (Long’s exp.)” A second sheet
bears similar data, but lacks the number 85 and the specific and varietal
epithets.

Hab. No information.

ACKNOWLEDGMENTS

My thanks are extended to Miss Ruth E. Brown and Dr. Alfred E. Schuyler, librar¬
ian and curator of the herbarium, respectively, of the Academy of Natural Sciences of

Philadelphia, for providing me the access and use of the materials for this research.

LITERATURE CITED

Barber, H. S. 1928. Thomas Say’s unrecorded journey in Mexico. Ent. News 39: 15-20.

Brendel, Frederick. 1879. Historical sketch of the science of botany in North America
from 1635 to 1840. Am. Nat. 13: 754-771.

Graustein, Jeannette E. 1967. Thomas Nuttall Naturalist Explorations in America
1808-1841. Harvard Univ. Press, Cambridge. 481 pp.

Keating, William H. 1824. Narrative of an Expedition to the Source of St. Peter’s River,
Lake Winnepeek, Lake of the Woods, &c. Performed in the Year 1823, by order of
the Hon. J. C. Calhoun, Secretary of War, under the Command of Stephen H. Long,
U. S. T. E. 2 Vols. H. C. Carey and I. Lea, Philadelphia. 439 pp. and 459 pp. Reprint,
Geo. B. Whittaker, London. 458 pp. and 248 pp. + 156 pp. 1825.

Nuttall, Thomas. 1818. The Genera of North American Plants and a Catalogue of the
Species, to the Year 1817. 2 Vols. Printed for the author by D. Heartt, Philadelphia.
312 pp. and 254 pp. + index, eratum [sic] , and additions.

Pennell, Francis W. 1935. The botanist Schweinitz and his herbarium. Bartonia 16: 1-8.

Pursh, Frederick. “1814” [1813]. Flora Americae Septentrionalis; or, a Systematic Ar¬
rangement and Description of the Plants of North America. 2 Vols. White, Cochrane,
and Co., London. 751 pp. + 24 pi.

Rodgers, Andrew Denny, III. 1942. John Torrey A Story of North American Botany.
Princeton Univ. Press, Princeton. 352 pp. Reprint, Hafner Publishing Co., New York.
1965.

Schweinitz, Lewis David von. [1824] . A Catalogue of Plants Collected in the North-West¬
ern Territory by Mr. Thomas Say in 1823. 11 pp. Manuscript in the library of the
Academy of Natural Sciences of Philadelphia, coll. no. 4, item 16.

- . 1824. Appendix. Section II. Botany. A Catalogue of Plants Collected in

the North-Western Territory by Mr. Thomas Say, in the Year 1823, Vol. 2, pp.
379-400. In William H. Keating, Narrative of an Expedition to the Source of St.
Peter’s River, . . . H. C. Carey and I. Lea, Philadelphia. Reprint, Geo. B. Whittaker,
London, Vol. 2, pp. 105-123. 1825.

- . [Undated] . [Systematically Arranged Catalogue of the Schweinitz Herbar¬
ium]. Unpublished manuscript in the library of the Academy of Natural Sciences of
Philadelphia, coll. no. 137, item 12.

Shear, C. L., & Neil E. Stevens, [ed.] 1921. The correspondence of Schweinitz and Tor¬
rey. Mem. Torrey Bot. Club 16: 119-300.

Stuckey, Ronald L. 1967. Daniel Steinhauer, early Ohio plant collector, and his cor¬
respondence with the botanist Schweinitz. Bartonia 36: 1-24.

Upham, Warren. 1884. Catalogue of the flora of Minnesota, including its Phaenogamous
and vascular Cryptogamous plants, indigenous, naturalized, and adventive. 12th Ann.
Rep. Geol. Nat. Hist. Surv. Minn. 1883(6): 1-193.

Weiss, Harry B., & Grace M. Ziegler. 1931. Thomas Say Early American Naturalist.
Charles C Thomas, Springfield, Ill. 260 pp.

48

THE MICHIGAN BOTANIST

Vol. 9

A PRELIMINARY REPORT ON THE VARIETIES OF

MAIANTHEMUM CANADENSE IN NORTHERN MICHIGAN1

Stephen G. Weller

Department of Botany, The University of Michigan, Ann Arbor

Maianthemum, a small circumboreal genus, is noted among the Liliaceae
for having its flower parts arranged in multiples of two. Differences between
the species are slight, and have caused some argument (Farwell, 1915). Never¬
theless, three species are generally recognized. M. bifolium occurs in Europe
and Asia, M. dilatatum occurs along the Pacific coasts of both Asia and North
America, and M. canadense extends from Labrador south to upland Georgia
and westward to northern British Columbia, south to southern Alberta. On
the basis of pubescence found on the undersurface of the leaves of M. cana¬
dense growing in the western part of its range, Fernald (1914) described var.
interim. Butters (1927) listed many other characters separating the two vari¬
eties. He found no evidence of ecological preferences, and when the two vari¬
eties were occasionally found growing together in Wisconsin and Minnesota,
few intermediate forms were noted.

The purpose of this investigation was to study the occurrence and pos¬
sible intergradation of the varieties in the vicinity of the University of Michi¬
gan Biological Station. Previous investigations (Butters, 1926) and herbarium
records indicated the presence of both varieties throughout the state, and
specifically in Emmet and Cheboygan counties, the area surrounding the Bio¬
logical Station.

A wide range of habitats was sampled in 1969 in both counties. Mai¬
anthemum is nearly ubiquitous, occurring commonly in all shaded habitats
except very dry aspen woods and jack pine forests. At each study area, large
numbers of specimens were collected for the purpose of analyzing the varia¬
tion occurring in that area. Representative samples of these collections have
been deposited in the University of Michigan Biological Station Herbarium.
Because many of the previously described differences between the varieties
were based on flower and inflorescence characters, collections intended for
verification of these differences consisted primarily of blooming individuals.
No differences between the varieties were restricted to the leaves of sterile
shoots. Uprooted rhizomes were extensive, and invariably broke before the
entire length of the rhizome could be exhumed. These rhizomes produced
both fertile shoots and sterile shoots with a single leaf. Because of the clonal
nature of the plant, and the difficulty of distinguishing between the clones, no
attempt to apply precise sampling methods was made. Only in those areas
where both varieties were found growing together was the presence of two
clones definitely indicated. In these areas bias was minimized because of the
superficial similarity of the two varieties, in spite of Butters’ remark,
“. . . these two plants are so distinct even though they are occasionally found
growing side by side, as to give one the feeling that they are to be regarded as
species rather than varieties.”

^Contribution from the Biological Station of The University of Michigan.

1970

THE MICHIGAN BOTANIST

49

Failure to detect ecological preference on the part of either variety was
the most immediate result of this study. Although Baldwin (1958) stated that
in the clay belt of northern Ontario and Quebec var. interim preferred drier
woodland habitats than var. canadense, no such preference was found in the
area studied. In northern Michigan both varieties occur in aspen-pine wood¬
lands, coniferous forest bordering the Straits of Mackinac, northern hardwood
associations, and also in bog forests consisting of Thuja, Abies, and Picea. In
the forest surrounding Galloway Lake bog north of Levering, Emmet Co., var.
interim was growing on a rotting stump almost completely surrounded by
open water. In each of these major habitats where M. canadense was found,
both varieties grew together in at least one area, in some cases to the extent
that the rhizomes were densely intermingled.

Butters (1927) cited numerous differences between the two varieties. He
stated that generally var. interim was taller than var. canadense, and produced
more flowers in a denser inflorescence, although the length of the inflores¬
cence of var. interim was usually greater. In addition, the leaves of var.
interim were supposedly more cordate than the typical form, with the second¬
ary veins and commissures of var. interim more obscure than those of var.
canadense. Compared to var. canadense, Butters found the leaves of var.
interim to be more plicate. He also stated that var. canadense typically
bloomed somewhat earlier than var. interim. Minute differences in flowers and
rhizome anatomy were also cited. Fernald (1950) stated that the leaves of var.
interim were usually a darker blue-green compared to those of var. canadense.

In order to determine the validity of these claims, measurements of four
characters were made from specimens in the University of Michigan Herbari¬
um, and the Michigan State University Herbarium. Specimens measured were
primarily from Michigan, but collections from other parts of the United States
and Canada were also used. The characters recorded are shown in Table I,
where the results are summarized. Statistically significant differences were
obtained in all cases, and the measurements agreed with Butters’ results. How-

TABLE I. Character values based on specimens of Maianthemum canadense from the
herbaria of the University of Michigan and Michigan State University.

Variety

1. Overall

2. No. of

3. Inflorescence length

4. Angle of the

height

flowers per

versus no. of flowers

apex of the

inflorescence

lowest leaf

No. of

Length

Average

plants

of

no. of

raceme

flowers

M. canadense

13 cm

20

2

4-5 cm

23

52°

var. canadense

36

3-4 cm

24

(Values averaged

131

2-3 cm

20

on the basis of

41

1-2 cm

17

210 specimens.)

M. canadense

17 cm

27

1

5-6 cm

32

65°

var. interius

19

4-5 cm

38

(Values averaged

46

3-4 cm

29

on the basis of

48

2-3 cm

24

126 specimens.)

12

1-2 cm

22

50

THE MICHIGAN BOTANIST

Vol. 9

ever, the leaf angle character was very unsatisfactory. The average difference was
so slight, and the individual variation so great, that no general impression of con-
sistant differences could be gained by looking at the leaves of the two varieties.

In addition to these characters, other differences mentioned by Butters
were verified by specimens collected in Emmet and Cheboygan counties. The
intermediate veins and commisures of var. interim were distinctly more ob¬
scure than those of var. canadense. However, the leaves of var. interim did not
appear to be more plicate than leaves of the typical form. Butters’ statement
that the anthers of var. interim are clavate, compared to the subulate form of
anthers of var. canadense , did not appear to be borne out. Neither was any
consistent difference in the pistils of the two varieties evident, although
Butters claimed that the pistil of var. interim is more prominent and deeply
two-lobed. Other characters mentioned by Butters and Fernald appear much
more variable. This is especially true in the case of leaf shape. In some popula¬
tions of var interim, the leaves are very acute, and in other populations they
are blunt and more cordate. The same range of variation also occurs with var.
canadense. In some areas, acute-leaved var. interim occurs together with
blunt-leaved var. canadense, a situation exactly opposite from that described
by Butters. Similarly, although color differences between the leaves of the
varieties are encountered, the differences are not stable and may be totally
reversed from one situation to the next.

The most interesting variation is in the range of flowering periods. On

Fig. 1. Maianthemum canadense var. interius (three plants at left) and var. canadense
collected at Readmond Township Park, Emmet County, Michigan, June 13, 1969. Nearly all
the flowers of var. interius are open. Flowers of var. canadense are still in young bud. The
more broadly cordate shape of the leaves of var. interius is also evident. Photographed by
Gary R. Williams.

1970

THE MICHIGAN BOTANIST

51

several occasions, var. interim has been reported to flower as much as two
weeks later than var. canademe (Butters, 1927; Ingram, 1966; Morley, 1969).
In this study a fixed relation between the blooming periods of the two varie¬
ties was not found. In some situations where they were found growing to¬
gether, both varieties flowered at the same time; in others, var interim
bloomed later than or before the typical variety (Fig. 1). In populations where
both of the varieties occur together, and the blooming periods overlapped,
intermediate plants were found. These are assumed to be hybrids because they
are intermediate in amount of pubescence, height, and number of flowers. The
number of intermediates varied according to the degree of overlap in flower¬
ing. In areas where little overlap occurred, few putative hybrids were present.
Conversely, where overlap was complete, large numbers of intermediates were
found. Measurements similar to those made on herbarium specimens from
throughout the range of the two taxa were made on the plants collected in
Emmet and Cheboygan counties. Differences between the varieties have ap¬
parently not been affected by extensive hybridization. Statistics based on the
plants collected during this study compare closely with measurements of speci¬
mens from throughout the range of the taxa (see Table II). This fact, and the
large numbers of putative hybrids found in Emmet and Cheboygan counties,
indicate the possible occurrence of hybrids wherever the varieties grow to¬
gether. However, this is not suggested from herbarium records. In the Universi¬
ty of Michigan Herbarium only 1 1 intermediate specimens were found among
317 specimens examined of both varieties of M. canadense. Lack of extensive
collecting in a particular area and the clonal nature of the species probably
contribute to this situation.

Butters states that in Minnesota intermediate forms “are very rare and
can easily be explained as hybrids.” He makes this claim on the basis of in¬
fertile pollen taken from intermediate forms. Pollen was not examined in this
study, but an attempt was made to show the occurrence of introgressive
hybridization, which was suggested by the widely differing degrees of pubes¬
cence. Four groups of putative hybrids were recognized, as listed in Table III.
A certain amount of bias in recording the data was unavoidable, because of
the difficulty of distinguishing between group 4 and var. interim. To some
extent, this probably accounts for the greater numbers of individuals in groups
1 and 2, where the degree of pubescence is much easier to categorize. The
heights and number of flowers of individuals in each group were measured to
determine possible correlations between variation of these characters with vari-

TABLE II. Character values based on specimens of Maianthemum canadense collected in
Emmet and Cheboygan counties. (Because the overall height and number of flowers per
inflorescence probably reflect the length of the inflorescence, this character was not meas¬
ured for these plants. The leaf angle character was eliminated because of its unreliability.)

Variety Overall height Number of flowers

per inflorescence

var. canadense

(Values averaged on the basis of 203 specimens.) 13 cm. 21

var. interius

(Values averaged on the basis of 122 specimens.)

17 cm.

25

52

THE MICHIGAN BOTANIST

Vol. 9

TABLE III. Average height and number of flowers for groups of putative hybrids be¬
tween Maianthemum canadense var. canadense and M. canadense var. interius, separated
by differing degrees of pubescence.

Amount of pubescence

Height

No. of flowers
per inflorescence

1. Leaf margins and petioles minutely pubes¬
cent (based on 77 specimens)

16

23.6

2. Pubescence longer and thicker, but still con¬
fined to the leaf margins and petioles (based on

70 specimens)

15.5

23.4

3. Pubescence same length as in group 2, but
covering the basal portion of the undersurface
of the leaves as well as the leaf margins and
petioles (based on 37 specimens)

15.9

25.9

4. Entire undersurface of leaf pubescent in ad¬
dition to leaf margins and petioles. Pubescence
on basal portion of leaves similar in length to
basal leaf pubescence in group 3, but more
abundant. Pubescence on distal portion of
leaves shorter and more thinly distributed than

16.2

25.0

in var. interius (based on 49 specimens)

ation of pubescence (see Table III). It was expected that correlations between
these characters would appear, but none were found.

SUMMARY

In Emmet and Cheboygan counties, Michigan, differing habitat preferences be¬
tween the two varieties of Maianthemum canadense do not seem to exist. Some morpho¬
logical differences cited by former workers were verified. Blooming time varied widely in
both varieties, and this variation directly influenced the formation of intermediates. These
putative hybrids were found when overlap in blooming periods occurred. When the varie¬
ties bloomed at distinctly different times, very few intermediates were present. Because
of the widely differing degrees of pubescence shown by the putative hybrids, introgressive
hybridization was suggested, but not adequately demonstrated.

ACKNOWLEDGMENTS

I wish to thank Edward G. Voss for his considerable aid throughout the course of
this research and also Gary R. Williams for the photograph. The work was supported in
part by NSF grant GB-8440 to the University of Michigan Biological Station.

LITERATURE CITED

Baldwin, W. K. W. 1958. Plants of the Clay Belt of Northern Ontario and Quebec. Natl.
Mus. Can. Bull. 156. 324 pp.

Butters, F. K. 1926. Notes on the range of Maianthemum canadense and its variety
interius. Rhodora 28: 9-11.

Butters, F. K. 1927. Taxonomic studies in the genus Maianthemum. Minn. Stud. PI. Sci.
1: 429-444.

Farwell, O. A. 1915. Notes on the Michigan Liliaceae. Bull. Torr. Bot. Club 42: 351-358.
Fernald, M. L. 1914. The western variety of Maianthemum canadense. Rhodora 16:
210-211.

Fernald, M. L. 1950. Gray’s Manual of Botany, ed. 8. Am. Book Co., New York, lxiv +
1632 pp.

Ingram, J. 1966. Notes on the cultivated Liliaceae 3. Maianthemum. Baileya 14: 50-59.
Morley, T. 1969. Spring Flora of Minnesota. Univ. Minn. Press, Minneapolis. 283 pp.

1970

THE MICHIGAN BOTANIST

53

THE BARNES HYBRID ASPEN,

POPULUS XB ARNE SIP HYBR. NOV. -
A NOMENCLATURAL CASE IN POINT

W. H. Wagner, Jr.

Matthaei Botanical Gardens,

The University of Michigan, Ann Arbor

Natural hybrids are much more common in plants than animals. In some
groups like the ferns as many as 20 per cent of the “kinds” in a flora may be
hybrids. Among trees of the Great Lakes area, such hybrids as black times
sugar maple, Acer nigrum X saccharum, produce such a confusing array of
intermediates that some collectors doubt whether the parents are true species.
Others of our native hybrids, such as Bebb’s oak, Quercus Xbebbiana ( Q . alba
X macrocarpa ) and the Purpus birch, Betula Xpurpusii (B. alleghaniensis X
pumila ) are so well marked that the competent dendrologist recognizes them
immediately. I wish to describe another well defined hybrid tree of this region
here.

First it should be stated that from a number of standpoints, hybrids are
often inconvenient and troublesome to field workers, taxonomists, and evolu¬
tionists. For example, crosses are hard to key out in many cases, and they
tend to blur the distinctions of normal species; for in practically all respects,
hybrids tend to be intermediate between their parents. This fact has caused
wide variation in nomenclatural usage (Wagner, 1968, 1969). The hybrid aspen
to be designated below might be named (1) as a variety of Populus grandiden-
tata; (2) as a variety of P. tremuloides, its other parent; or (3) as one of three
varieties, the parents and the hybrid, of a single species; (4) by its formula, P.
grandidentata X tremuloides; (5) by a straight binomial, as if it were a normal
species; or (6) by a hybrid binomial, i.e., interpolating the multiplication sign
to indicate its origin. It seems hard to believe that authors find arguments to
support each of these different methods. I personally believe that a hybrid
binomial should be used for all hybrids except those so rare that they are not
likely to be encountered. Whatever we do, designation of hybrids should be
convenient; the name should be a useful handle for denoting the plant. It
should be stable, so that the name will not have to be changed in future. And
it should be meaningful, i.e., we should be able to tell right away whether the
plant named is a species or hybrid.

The plant we have been referring to clumsily as Populus grandidentata X
tremuloides is now known to be widespread and locally common in the Upper
Great Lakes and elsewhere in the eastern United States. Dr. Burton V. Barnes,
who has been studying it intensively for over a decade in connection with his
broader researches on biology and classification of poplars and aspens, reports
numerous localities in Michigan (cf. Barnes, 1961, 1967; Andrejak and Barnes,
1969). The hybrid aspen ranges from Minnesota to New England (Pauley,
1956; Barnes, pers. comm.). In my own field work in Michigan I have ob¬
served it so many times I no longer bother to collect it. At least in southern

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THE MICHIGAN BOTANIST

Vol. 9

Michigan, I do not doubt that it will be found in every county. It occurs
especially in second-growth communities, especially woods along railroads,
edges of fields, and along streams and rivers where there has been much past
disturbance. One professor of my acquaintance, unaware of the widespread
presence of this plant, thought it was a rarity. Indeed, when he began taking
classes to one of his favorite localities, he taught the hybrid as “P. grandiden-
tata ” because trees of that species were not present nearby, only P. tremul-
oides. Compared to the latter, the misidentified trees did have leaves with
“fewer and larger teeth.”

It must not be overlooked that all of our aspens form large stands by
root suckering (Barnes, 1966). A single aspen hybrid in this way can theoreti¬
cally produce dozens of trees by vegetative propagation underground. An
aspen woods is not made of individual trees that came from separate seeds;
only one or a few original plants can produce a large stand.

I have argued elsewhere (Wagner, op. cit.) that the hybrid binomial has
certain advantages over other methods of designating hybrids. One can quickly
refer to a specimen in teaching and discussion, using a stable name. And if we
install the X sign, it immediately tells that the plant is a “between species,” a
special kind of taxon differing in various respects from normal species. I be¬
lieve that Barnes’ hybrid aspen is a good case in point and accordingly pro¬
pose that this plant be given a hybrid binomial. The choice of epithet is ap¬
propriate because Dr. Barnes has made the bulk of contributions to our
knowledge of this plant, and most of the literature is to be found under his
name.

Populus xbarnesii W. H. Wagner, hybr. nov. Arbor clones faciens, inter P. grandidentatum
et P. tremuloidem media, gemmis toinentellis, foliorum petiolis 3-6 cm longis, laminis
ovatis vel subdeltoideis vel suborbicularibus, acuminatis, 4-9 cm longis, 3. 5-7. 5 cm latis,
dentibus marginalibus, 12-25 in quoque margine, aliquantuin irregularibus et sinubus
rotundatis 0. 5-3.0 mm. profundis separatis.

Type: Michigan, Washtenaw County, Waterloo Recreation Area (NE!4SE!4 sec. 9,
T2S, R3E). Barnes no. 65W88. (Holotype MICH; isotypes NY, UC, US, WTU).

LITERATURE CITED

Andrejak, Gary E., & Burton V. Barnes. 1969. A seedling population of aspens in south¬
eastern Michigan. Mich. Bot. 8: 189-202.

Barnes, Burton V. 1961. Hybrid aspens in the Lower Peninsula of Michigan. Rhodora 63:
311-324.

- . 1966. The clonal growth habit of American aspens. Ecology 47: 439-

447.

- . 1967. Indications of possible mid-Cenozoic hybridization in the aspens of

the Columbia Plateau. Rhodora 69: 70-81.

Pauley, Scott S. 1956. Natural Hybridization of the Aspens. Univ. Minn. for. Note 47. 2

pp.

Wagner, W. H., Jr. 1968. Hybridization, Taxonomy, and Evolution. In V. H. Heywood
(ed.), Modern Methods in Plant Taxonomy, Chapt. 9, pp. 113-138. Academic Press,
London and New York.

- . 1969. The role and taxonomic treatment of hybrids. BioScience 19:

785-789, 795.

1970

THE MICHIGAN BOTANIST

55

Hatu re education feature —

WINTER FIELD KEY TO SOME TREES COMMON IN MICHIGAN

James R. Wells

Cranbrook Institute of Science
Bloomfield Hills, Michigan

The increasing interest in environmental education brings to mind that
the winter season is too often neglected for its potential value in nature edu¬
cation. Our various parks and recreation areas are open in the winter and their
trails provide easy access to an outdoor laboratory for winter botany.

Perhaps trees in the winter condition are a most suitable point of de¬
parture for the student of plant science during this season. In winter, as in
any other season, trees constitute the most conspicuous part of many plant
communities. It should be borne in mind that one does not really “know” a
tree when a scientific or common name can be associated with certain leaf
characters. To really “know” a tree one should be able to recognize and
identify it during all seasonal aspects as well as in youth, maturity, and old
age. I wonder how many of us are able to recognize genera of trees on the
basis of flower characters alone, for instance.

In addition to the well-known Michigan Trees by C. H. Otis (9th ed.,
University of Michigan Press, 1931), the selected references listed and annotat¬
ed below are suitable for use in the western Great Lakes area, as all apply to
the eastern or northeastern States. All are illustrated. It is intended that at
least one of these references be available as a field manual and thus a supple¬
ment to this “first key” to trees in the winter condition in this area. I there¬
fore have omitted a glossary, drawings, and photographs, as well as citation of
authorities following scientific names.

Core, Pari L., and Nellie P. Ammons. 1958. Woody Plants in Winter. 218 pp. The Box¬
wood Press, P.O. Box 1717, Pittsburgh, Pa. 15213. Clothbound $4.00; paper $2.75.

This handy volume is, in my opinion, the best single “field manual” reference for
native woody plants in the winter condition in the western Great Lakes area. As its title
implies, shrubs are treated, as well as certain vines and trees. Some plants which have
been introduced are included. It is profusely illustrated by line drawings.

Trelease, William. 1931. Winter Botany. 3rd rev. ed. 396 pp. Reprinted 1967 by Dover
Publications, Inc., 180 Varick St., New York, N. Y. 10014. $2.00 (paper bound).

This volume lists over 1000 woody plants, including some escaped species and
vines. Conifers are excluded. Perhaps the strongest point of this volume is the line draw¬
ings, which are particularly diagnostic at the generic level.

Harlow, William M. 1959. Fruit and Twig Key. 50 + 56 pp. Dover Publications, Inc.
(address above). $1.35 (paper bound).

This small volume is a reprint of two earlier publications: 1) a fruit key (1946)
which treats both deciduous and evergreen tree species in its 50 pages; 2) keys (1954) to
deciduous woody plants (both trees and shrubs) based on twig characters. In all there are
over 350 photographs, some of which are of questionable value. This book might well be
used as an additional reference with either of the preceding books.

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THE MICHIGAN BOTANIST

Vol. 9

Symonds, George W. D. 1958. The Tree Identification Book. 272 pp. M. Barrows and
Co., 425 Park Ave. S., New York, N. Y. 10016. $10.00 (cloth bound).

This book, along with its companion volume, The Shrub Identification Book (379
pp.) by the same author and publisher but $15.00, clearly provides the practical approach
to woody plant identification. While keys in the traditional couplet sense are not in¬
cluded there are in each several “keys” based on such characters as thorns, leaves, twigs
and buds, bark, and flowers. These separate keys are essentially photographic treatments
(in black and white) and the photographs in the shrub book are of somewhat better
quality than those in the book on trees. In both volumes species that are strictly south¬
ern or subtropical are omitted, as are hybrids in general and certain genera whose species
are very numerous, e.g., Crataegus (hawthorn). Although several dozens of species of oaks
occur in eastern North America and adjacent Canada, only nine species are included. In
each volume both popular and scientific names are given.

KEY

1.

1.

3.

3.

5.

5.

7.

7.

9.

9.

11.

11.

13.

13.

Leaves evergreen, persisting through winter . 2

Leaves deciduous, not persisting through winter . 8

2. Leaves scale-like or awl-shaped, opposite or whorled . 3

2. Leaves linear, needle-like, alternate or in bunches of 2-5 . 4

Leaves all scale-like, branchlets flattened . Thuja occidentalis (arbor vitae)

Leaves all or in part awl-shaped; branchlets not strongly flattened .

. Juniperus virginiana (red cedar)

4. Leaves in bunches of 2-5 on dwarf branches . Pinus spp. (pine)

4. Leaves alternate, not in bunches on dwarf branches . 5

Leaves 4-sided in cross section . Picea spp. (spruce)

Leaves in cross section flattened from top to bottom . 6

6. Leaves of 2 lengths, those on the upper side of twig shorter than those on

lower side of twig . Tsuga canadensis (hemlock)

6. Leaves of about equal length . 7

Branchlets flexible; cones hanging, 3-pronged bracts conspicuous beyond cone

scales; leaf stalks remaining on twigs after leaves have fallen .

. *Pseudotsuga menziesii (Douglas fir)

Branchlets stiff; cones erect, 3-pronged bracts absent; leaf stalks absent .

. Abies balsamea (balsam fir)

8. Dwarf branches bearing crowded leaf scars; gymnosperms . 9

8. Dwarf branches, if present, lacking crowded leaf scars; angiosperms . 10

Bundle scars 2; leaf bases not persisting as ridges down the stem .

. *Ginkgo biloba (ginkgo)

Bundle scar 1; leaf bases persisting as ridges down the stem . . . .Larix laricina (larch)

10. Leaf scars whorled . Catalpa spp. (catalpa)

10. Leaf scars opposite or alternate . 11

Leaf scars opposite . N .... 12

Leaf scars alternate . . . . 15

12. Twigs stout; terminal bud about 1 cm long or longer, its 10-12 scales

sometimes sticky . Aesculus spp. (*Horse chestnut, buckeye)

12. Twigs not stout; terminal bud shorter, its scales fewer, not sticky . 13

Bud scales hairy or rough, brownish-black; bundle scars more than 3 .

. . . . Fraxinus spp. (ash)

Bud scales not hairy nor rough, reddish or gray; bundle scars 3 . 14

*Tree species not native in Michigan.

1970

THE MICHIGAN BOTANIST

57

15.

15.

17.

17.

19.

19.

21.

21.

23.

23.

25.

25.

27.

27.

29.

29.

31.

31.

14. Visible bud scales 2; leaf scars slightly raised, the 2 opposing scars joined

by a line but not meeting around the stem . Cornus spp. (dogwood)

14. Visible bud scales usually more than 2; leaf scars not raised; often meeting

around the stem . Acer spp. (maple)

Twigs with thorns . 16

Twigs without thorns . 21

16. Sap milky; thorns usually 1 per axil . *Maclura pomifera (osage orange)

16. Sap not milky; thorns not single at each axil . 17

Axillary buds within leaf scar . 18

Axillary buds not within leaf scar . 19

18. Thorns usually branched . ! . Gleditsia triacanthos (honey locust)

18. Thorns unbranched, in pairs opposite leaf scars .

. Robinia pseudoacacia (black locust)

Thorns not formed from modified branch ends . Crataegus spp. (hawthorn)

Thorns formed from ends of modified branches . 20

20. Stipule scars present; the terminal bud usually absent .

. Prunus spp. (plum, cherry)

20. Stipule scars absent, the terminal bud present . *Pyrus communis (pear)

Twigs with stipular rings completely encircling the stem . 22

Twigs lacking stipular rings which completely encircle the stem . 24

22. Leaf scar encircling the lateral bud; terminal bud absent .

. Platanus occidentalis (sycamore)

22. Leaf scar not encircling the lateral bud . 23

Leaf scar round; terminal bud scales 2 . Liriodendron tulipifera (tulip tree)

Leaf scar not round, often U-shaped; terminal bud scale 1 .

. *Magnolia spp. (magnolia)

24. Buds at least 1 cm long, covered by 20 or more spirally arranged scales;

stipular scars almost meeting around the twig . Fagus grandifolia (beech)

24. Buds shorter, scales fewer, and stipular scars not as big . 25

Leaf scars in 2 ranks . 26

Leaf scars not in 2 ranks . 38

26. Bundle scar 1 or several appearing as 1 . *Diospyros virginiana (persimmon)

26. Bundle scars 3 or more, some of them sometimes composed of more than

1 in a close group . 27

Terminal bud naked . 28

Terminal bud absent or if present then covered by scales . 29

28. Buds yellowish; stipular scars present; pith without diaphragms .

. Hamamelis virginiana (witch hazel)

28. Buds covered with reddish brown hairs; stipular scars absent; pith some¬
times with greenish plates or chambered . Asimina triloba (pawpaw)

Stipular scars absent . 30

Stipular scars present . 31

30. Three vertical ridges below leaf scar; terminal bud absent .

. Cercis canadensis (redbud)

30. Vertical ridges absent; terminal bud present .... Amelanchier spp. (serviceberry)

Bud scales 2; buds slanting sideways; pith not 3-sided . . . Tilia americana (basswood)

Bud scales more than 2 or if 2 then pith 3-sided; buds sometimes slanting . 32

32. Bundle scars 3 or in 3 distinct groups . 33

32. Bundle scars more than 3 and not in groups of 3 . 37

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THE MICHIGAN BOTANIST

Vol. 9

33.

33.

35.

35.

37.

37.

39.

39.

41.

41.

43.

43.

45.

45.

47.

47.

49.

49.

51.

51.

Bud scales in 4 ranks; trunk with longitudinal ridges .

. Carpinus caroliniana (blue-beech, ironwood)

Bud scales not 4-ranked; trunk lacking longitudinal ridges . 34

34. Bud scales longitudinally striped with fine lines .

. . Ostrya virginiana (hop hornbeam, ironwood)

34. Bud scales without longitudinal stripes . 35

Terminal bud present; lenticels horizontally elongated; pith 3-sided .

. Be tula spp. (birch)

Terminal bud absent; lenticels not horizontally elongated; pith not 3-sided . 36

36. Lateral buds appressed; pith chambered .

36. Lateral buds not appressed; pith not chambered

Leaf scar triangular; sap not milky; pith 3-sided . . .
Leaf scar rounded; sap milky; pith rounded .

38. Lateral buds superposed .

38. Lateral buds not superposed .

Pith diaphragmed .

Pith not diaphragmed .

Celtis occidental is (hackberry)
. Ulmus spp. (elm)

. Corylus spp. (hazelnut)

. Moms spp. (mulberry)

. 39

. 41

. Juglans spp. (walnut)

. 40

40. Axillary bud contained within the leaf scar; pith not 5-angled nor star¬
shaped . Gleditsia triacanthos (honey locust)

40. Axillary bud not surrounded by the leaf scar; pith 5-angled or star-shaped

. Cary a spp. (hickory)

Bundle scar 1 or several appearing as one . 42

Bundle scars more than 1 and separated . 43

42. Stipular scars absent; buds not superposed; twigs green and aromatic .

. Sassafras albidum (sassafras)

42. Stipular scars present; buds often superposed; twigs not green and

aromatic . Ilex verticillata (winterberry)

Bundle scars 3 or in groups appearing as 3 scars . 44

Bundle scars more than 3 and separate . 49

44. Bud scale 1 . Salix spp. (willow)

44. Bud scales more than 1 . 45

Pith (and sometimes twigs) 3-sided; buds borne on short stalks . . Alnus spp. (alder)
Pith not 3-sided; buds not stalked . 46

46. Bud scales 2 . Cornus alternifolia (alternate-leaved dogwood)

46. Bud scales more than 2 . 47

Pith 5-6-angled and star-shaped; deciduous branch scars present .

. Populus spp. (aspen, poplar, cottonwood)

Pith not 5-6-angled; deciduous branch scars absent . . 48

48. Stipule scars absent . *Pyrus spp. (apple, pear)

48. Stipule scars present . Prunus spp. (cherry, plum)

Buds clustered near tip of twig . . Quercus spp. (oak)

Buds not clustered near tip of twig . 50

50. Terminal bud present and large; pith 5-angled or star-shaped .

. Cary a spp. (hickory)

50. Terminal bud usually absent; pith rounded . 51

Leaf scars U-shaped or round; bundle traces often less than 9; sap resinous,

sometimes milky; some species poisonous to the touch . Rhus spp. (sumac)

Leaf scars shaped like a shield, notched at top; bundle traces usually more
than 9; sap not resinous nor milky . *Ailanthus altissima (tree-of-heaven)

1970

THE MICHIGAN BOTANIST

59

MICHIGAN PLANTS IN PRINT
New Literature Relating to Michigan Botany

This section lists new literature relating to Michigan Botany under four categories: A.
Maps, Soils, Geography, Geology (new maps and selected bulletins or articles on soils and
geology as these may be of use to field naturalists and students of plant distribution); B.
Books, Bulletins, etc., and C. Journal Articles (listing, respectively, all separate publications
and articles in other periodicals which cite Michigan specimens or include research based on
plants of wild origin in Michigan ;-not generally including work on cultivated plants nor
strictly economic aspects of forestry, conservation, or agriculture); D. History, Biography,
Exploration (travels and lives of persons with Michigan botanical connections). When the
subject matter or relation to Michigan is not clear from the title, annotations are added in
brackets. Readers are urged to call to the editor’s attention any titles (1960 or later) which
appear to have been overlooked -especially in less well known sources.

A. MAPS, SOILS, GEOGRAPHY, GEOLOGY

Farrand, William R., Robert Zahner, & William S. Benninghoff. 1969. Cary-Port Huron
Interstade: Evidence from a buried bryophyte bed, Cheboygan County, Michigan.
Geol. Soc. Am. Spec. Pap. 123: 249-262. [Contains important data for interpreting
vegetational history in northern Michigan; implies that the Straits of Mackinac were
not deglaciated during the Two Creeks Interstade but that drainage then occurred
across the Petoskey-Cheboygan lowland. Cf. also paper listed below by Miller & Ben¬
ninghoff.]

Gair, Jacob E., & Robert E. Thaden. 1968. Geology of the Marquette and Sands Quad¬
rangles, Marquette County, Michigan. U. S. Geol. Surv. Prof. Pap. 397. 77 pp. + pi. in
pocket. $3.00 (Govt. Printing Office). [Entirely on bedrock geology, but illustrates algal
structures in an ancient chert-dolomite and some maps show topography as well as
subsurface features and outcrops.]

Mettert, Wesley. 1969. Soil Survey of Osceola County, Michigan. U. S. Dep. Agr. 149 pp.
+ 96 once-folded map plates + 6 folded tables and indexes. $6.00 (Govt. Printing
Office). [Another of the new soil surveys including complete aerial photographic cov¬
erage for the county, at a scale of about 4" = 1 mile, with boundaries of soil types
overprinted. Will be indispensable guide to locating the numerous fine woodlots in this
county.]

Whiteside, E. P., I. F. Schneider, & R. L. Cook. 1968. Soils of Michigan. Mich. Agr. Exp.
Sta. Ext. Bull. E-630. 52 pp. + folded map. [An almost unaltered new edition of
former Spec. Bull. 402, with the same colored map of major Michigan soil associa¬
tions. A good general introduction to Michigan soils, available at county offices of the
Michigan State Cooperative Extension Service or from MSU Bulletin Office, Box 231,
East Lansing 48823.]

B. BOOKS, BULLETINS, SEPARATE PUBLICATIONS

Edman, F. Robert. 1969. A Study of Wild Rice in Minnesota; Minnesota Resources Com¬
mission, State Capitol, St. Paul. 110 + xii pp. (processed). [A compilation which in¬
cludes 6 pages of somewhat garbled data on distribution of wild rice in Michigan.]
Gillett, John M. 1968. The Milkworts of Canada. Canada Dep. Agr. Monogr. 5. 24 pp. [A
handy treatment of Polygala, including keys, drawings, and distribution maps. The
latter indicate a few Michigan locations for five of the species. The only Michigan
species in the genus which apparently is not found in Canada is P. cruciata, which
ranges as close as Algonac (St. Clair Co.) and Monroe Co. Available from Information
Division, Canada Department of Agriculture, Ottawa.]

Heiser, Charles B., Jr., with Dale M. Smith, Sarah B. Clevenger, & William C. Martin, Jr.
1969. The North American Sunflowers (Helianthus). Mem. Torrey Bot. Club 22(3).
218 pp. $5.00. [A thorough summary of present knowledge of this difficult genus,

60

THE MICHIGAN BOTANIST

Vol. 9

with keys and distribution maps, the latter showing Michigan stations for many of the
species.]

Schuyler, Alfred E. 1969. Three New Species of Scirpus (Cyperaceae) in the Southern
United States. Not. Nat. 423. 12 pp. [Cites Muskegon County records for related S.
hallii and illustrates achenes of one of them.]

Smith, Alexander H., & L. R. Hesler. 1969. The North American Species of Pholiota.
Hafner Publ. Co., New York. 402 pp. + 90 pi. $22.50. [A thorough monograph, with
many taxa including numerous new ones cited from Michigan-the new ones from
definite localities. The plates all consist of photographs of these mushrooms.]

Tubbs, Carl H. 1969. The Influence of Light, Moisture, and Seedbed on Yellow Birch
Regeneration. North Central For. Exp. Sta., U. S. For. Serv. Res. Pap. NC-27. 12 pp.
[Study made in a forest opening on the Upper Peninsula Experimental Forest near
Skandia (Marquette Co.).]

C. JOURNAL ARTICLES

Erbisch, Frederic H. 1969. Ascus and ascospore development of five species of the
lichen-forming genus Pertusaria. Bryologist 72: 178-199. [“All species examined were
collected in Michigan, in both upper and lower peninsulas. ”-no further locality data.]
Genys, John B. 1968. Geographic variation in eastern white pine. Silvae Genet. 17: 6-12.
[Three of the 99 sources of seed grown in Maryland were in Michigan; data on 9
characters given in a table.]

Gibbons, J. Whitfield. 1968. Reproductive potential, activity, and cycles in the painted
turtle, Chrysemys picta. Ecology 49: 399-409. [Includes mention of common plants
of Sherriff’s Marsh and Wintergreen Lake, T. 1 S., R. 9 W., Kalamazoo Co.]

Hagenah, Dale J. 1969. Polypodium virginianum not restricted to rocky habitats. Am.
Fern Jour. 59: 80-81. [Documents with citation of several Michigan collections the
occurrence of this species on sand, gravel, and rotten logs.]

Hennen, Joe F., & George B. Cummins. 1969. The autoecious species of Puccinia and
Uromyces on North American Senecioneae. Mycologia 61: 340-356. [Cites and illus¬
trates teliospores of type material of P. recedens from Senecio aureus at Ann Arbor.]
Herbst, Richard P. 1969. Ecological factors and the distribution of Cladophora glomerata
in the Great Lakes. Am. Midi. Nat. 82: 90-98. [Includes brief report on “massive
growths” of this alga at cities along eastern shore of Lake Michigan.]

Ireland, Robert R. 1969. Taxonomic studies on the genus Atrichum in North America.
Canad. Jour. Bot. 47: 353-368. [One Michigan collection is cited for each of three of
the species in this moss genus.]

Kimbrough, James W. 1969. North American species of Thecotheus (Pezizeae, Peziza-
ceae). Mycologia 61: 99-114. [T. apiculatus, a new species from Ontario, cited also
from Ann Arbor.]

Koponen, Timo. 1968. The moss genus Rhizoinnium (Broth.) Kop., with description of
R. perssonii, species nova. Mem. Soc. Fauna IT. Fenn. 44: 33-50. [Specimens of the
new species are said to have been seen from several North American states and prov¬
inces, including Michigan, but Eurasian material is cited.]

Miller, Norton G., & William S. Benninghoff. 1969. Plant fossils from a Cary-Port Huron
Interstade deposit and their paleoecological interpretation. Geol. Soc. Am. Spec. Pap.
123: 225-248. [Site was not far north of Douglas Lake, Cheboygan County; species
reported from deposit 12,500-13,300 years old which are now characteristics of north¬
ern boreal coniferous forest, forest-tundra ecotone, and adjacent tundra of the far
north. Cf. paper cited under “A” above by Farrand et al.]

Miller, O. K., & Roy Watling. 1968. The status of Boletus calopus Fr. in North America.
Notes Roy. Bot. Card. Edinburgh 28: 317-325. [This species not known in eastern
North America; material identified as it or the synonymous B. pachypus (including
collection cited from Sault Ste. Marie, Michigan) is different.]

Miller, Philip C., & David M. Gates. 1967. Transpiration resistance of plants. Am. Midi.
Nat. 77: 77-85. [field studies of energy budget of several species in Emmet and
Cheboygan counties.]

1970

THE MICHIGAN BOTANIST

61

Pringle, James S. 1969. Documented plant chromosome numbers 1969:1. Sida 3:
350-351. [Includes count of n = 26 for Potamogeton gramineus from Cheboygan
County, with tracing from photomicrograph.]

Smithberg, Margaret H., & Conrad J. Weiser. 1968. Patterns of variation among climatic
races of red-osier dogwood. Ecology 49: 495-505. [One of 21 clones studied under
uniform conditions in Minnesota was from Cadillac, Michigan.]

Thiers, Harry D., & James M. Trappe. 1969. Studies in the genus Gastroboletus. Brittonia
21: 244-254. [G. scabrosus noted from Michigan.]

Thompson, Paul W. 1969. A unique American chestnut grove. Mich. Acad. 1(3-4):
175-178. [Several thousand offspring established from an old planting near Frankfort
in Benzie Co.]

Vuilleumier, Beryl Simpson. 1969. The genera of Senecioneae in the southeastern United
States. Jour. Arnold Arb. 50: 104-123. [Mentions occurrence of Arnica cordifolia on
the Keweenaw Peninsula.]

Watling, Roy. 1969. New fungi from Michigan. Notes Roy. Bot. Gard. Edinburgh 29:
59-66.

Watling, Roy. 1969. The genus Paragyrodon. Notes Roy. Bot. Gard. Edinburgh. 29:
67-73. [Many southern Michigan localities, chiefly in Washtenaw County, cited for P.
sphaerosporus. ]

Wells, James R. 1969. A review of the varieties of Polymnia uvedalia. Rhodora 71:

204-211. [Map shows two varieties in southern Michigan.]

Wells, James R. 1969. Specific relationships between Polymnia canadensis and P. laevigata
(Compositae). Castanea 34: 179-184. [Distribution map includes 15 Michigan localities
for P. canadensis. ]

Wetzel, Robert G. 1969. Excretion of dissolved organic compounds by aquatic macro¬
phytes. BioScience 19: 539-540. [Laboratory data on Najas flexilis from Lawrence
Lake, Barry County (see pp. 3-9 in this issue.)]

Wright, H. E., Jr. 1968. The roles of pine and spruce in the forest history of Minnesota
and adjacent areas. Ecology 49: 937-955. [Includes consideration of percentages of
pine and spruce pollen in Michigan palynological studies.]

Zander, Richard. 1967. The New World distribution of Scopelophila (= Merceya). Bryolo-
gist 70: 405-413. [S. ligulata cited from Alger Co.]

D. HISTORY, BIOGRAPHY, EXPLORATION

Bliss, Lawrence C. 1969. Henry J. Oosting March 12, 1903 - October 31, 1968. Bull.
Torrey Bot. Club 96: 370-372. [Biographical sketch with picture and selected bibliog¬
raphy of the noted ecologist born in Holland, Michigan and a graduate of Hope Col¬
lege (A.B. 1925) and Michigan State (M.S. 1927); his 1932 paper on Potamogeton in
Michigan is not in the selected bibliography.]

Brockhoff, Dorothy A. 1969. Edgar Anderson 1897-1969. [An appreciative sketch with
portrait of one of Michigan State University’s best known graduates (B.S. 1918),
whose Michigan collections, while locally important, are doubtless one of his least
claims to fame; his family ties to Kalamazoo County are not mentioned.]

Degener, Otto. 1968. Earl Edward Sherff (1886-1966). Taxon 17: 189-198. [Biographical
sketch, with bibliography and portrait, of Michigan-born botanist who retired to Hast¬
ings, Michigan, after a professional career largely in Illinois.]

Fulford, Margaret, & Philip B. Whitford. 1968. In Memoriam Raymond Edward Hatcher
(1930-1967). Bryologist 71: 58-59. [Includes photo and bibliography. Hatcher col¬
lected bryophytes extensively in Michigan and elsewhere.]

Smith, Alexander H. 1969. Edwin Butterworth Mains 1890-1968. Mycologia 61: 449-451.
[Brief biography, with photo, of late director emeritus of the University of Michigan
Herbarium.]

Stuckey, Ronald L. 1969. An overlooked plant name (Aesculus maxima) of Daniel Drake
and his lost herbarium. Castanea 34: 185-192. [Mentions that the herbarium of the
Academy of Natural Sciences of Philadelphia includes some specimens collected by
Drake [ca. 1842] at Mackinac Island and Lake Superior, Michigan.]

62

THE MICHIGAN BOTANIST

Vol. 9

Information for Authors The Michigan Botanist

THE MICHIGAN BOTANIST is distributed not only to a considerable number of
individual and institutional subscribers domestically and abroad, but also to all members
of the Michigan Botanical Club, an organization composed in large part of non-profes¬
sionals with concerns for conservation and nature study as well as the local flora. (Total
paid circulation is about 800.) Considering the range of interests among members and
subscribers, we seek to include (in each year if not each issue) the widest possible range
of subject matter in regard both to the entire plant kingdom and to branches of the field
of botany.

Articles should, in general, include some new botanical information, and should
pertain to the Upper Great Lakes region. Not acceptable are manuscripts of primarily
literary nature (including all poetry and articles of an “editorial” orientation). Notes on
techniques useful to the teacher and hobbyist are welcomed, as is material of biographical
and historical nature.

To promote clarity to non-technical readers (without sacrifice of accuracy and
precision), such means are encouraged as illustrations, inclusion of a generally accepted
“common name” (when such exists) at least once in the title or text of any article
devoted to consideration of one or a few species, and introductory material placing the
author’s studies in perspective and briefly indicating their significance. Summaries,
especially of long or complex papers, are desirable.

Manuscripts should be submitted to the editor in chief and are considered on their
botanical merit and clarity of presentation; their acceptability is judged without special
regard for repetition of authors, length, or prospect of subsidies. All manuscripts are
subject to advisory review by botanists prepared to evaluate not only their content but
also their suitability for this journal. Manuscripts are ordinarily published in the order of
acceptance, with allowance as necessary for maintenance of diversity within our space
limitations. In the interests of diversity, acceptable articles occupying more than 10
printed pages can be published only if divided into two or more issues or if the necessary
additional pages can be subsidized (present charge is $12.50 per page). Subsidy of the
entire cost of an acceptable article will permit its publication without delay, and authors
with access to funds for this purpose are encouraged to allow complete subsidy, regard¬
less of length of article, to avoid contributing toward a backlog of papers awaiting
publication.

NOTE: Manuscripts should be neatly typed, double-spaced (especially bibliog¬
raphies, figure legends, and other technical matter), and submitted in duplicate to simpli¬
fy review. Use 8V2 X 11 inch paper and avoid “corrasable” kinds! Follow carefully the
style of current issues in regard to headings, identifications of authors, citation of litera¬
ture, abbreviations, and other matters of form. Complete pagination should be indicated
for all titles in “Literature Cited.” Illustrations and maps are encouraged, and there are no
arbitrary limits on the amount of such materials so long as they .can be reduced to page
size (4(4 X 7 inches); all are numbered consecutively as “Figures.” Page proof is sent to
authors (at least in North America) and must be checked for errors and returned prompt¬
ly; major revisions cannot be made at this time.

An order form for reprints will be supplied when a manuscript is accepted. Re¬
prints (with extraneous matter removed) are supplied at standard prices established by
the printer (rate schedule available from the editor).

-The Editorial Board

Program Notes

April 3-4: Michigan Academy of Science, Arts, & Letters. 74th Annual Meeting, at Wayne
State University, Detroit. The speaker at the Biologists Luneheon Friday, April 3, will be
David Jenkins, Chief, Research and Development Division, Michigan Department of
Natural Resources.

May 29-31: Michigan Botanical Club. Annual Spring Campout, at the University of Michigan
Biological Station on Douglas Lake.

The October number (Vol. 8, No. 4) was mailed October 29, 1969.

News of Botanists

DALE HAGENAH has retired from the Chrysler Corporation and is now Research
Associate and Curator of the Herbarium at the Cranbrook Institute of Science. . . . Two
botanists, both ecologists, have been named provost at their institutions: JOHN E.
CANTLON at Michigan State University and RONALD O. KAPP at Alma Col¬
lege. . . . ALEXANDER H. SMITH, Director of the University of Michigan Herbarium,
received one of the two Certificates of Merit awarded by the Botanical Society of America
at its 1969 annual meeting. . . . WALTER P. NICKELL, who retired in 1968 as natural¬
ist at the Cranbrook Institute of Science, where he was associated for 33 years, has been
awarded an honorary membership in the Michigan Botanical Club. . . . GENEVIEVE
GILLETTE, President of the Michigan Parks Association and honorary member of the
Michigan Botanical Club, was one of four persons (the others were college presidents)
recognized as Distinguished Alumni by Michigan State University last June. . . . STAN-
LP1Y A. CAIN, University of Michigan professor of conservation and former U. S. Assistant
Secretary of the Interior, was presented May 23 with the Mary Soper Pope Medal of the
Cranbrook Institute of Science for “noteworthy and distinguished accomplishment in the
field of plant sciences”; the presentation was made by W. H. Wagner, Jr., a trustee of the
Institute, president of the Botanical Club, and Director of the University of Michigan
Botanical Gardens. Dr. Cain was the Cranbrook botanist 1946-1950, played an important
role in the formation of the Michigan Natural Areas Council, and has served as chairman of
the Michigan Conservation Commission.

Publications of Interest

AN INVENTORY OF MINNESOTA LAKES. Bull. 25, Division of Waters, Soils, and
Minerals, Minnesota Conservation Department. 1968. 498 pp., plastic binding. This
tremendous compilation does for Minnesota much of what “Michigan Lakes and
Ponds,” reviewed at length in this journal in March, 1966, does for Michigan. I;or
each county, all lakes of 10 acres or more (whether named or not) are listed, assigned
a number, located by section(s) and township, the acreage stated. There are additional
notes, summaries, and classifications, and a master alphabetical index to all named
lakes. A map for each county locates its lakes (but almost nothing else except survey
township lines and major towns). In all, 15,291 lakes are tabulated. Unlike the highly
restricted Michigan volume, this one is readily available for $4.64 postpaid from Docu¬
ments Section, 140 Centennial Bldg., St. Paul, Minnesota 55101.

FLOWERS OF EUROPE. By Oleg Polunin. Oxford University Press, London, 1969. 622
pp. + 192 color plates. £4.4s [= $10.08]. Those who are tired of high prices for well
illustrated wildflower books will welcome this exceptional bargain which can be ob¬
tained for about $10.00 plus postage from any British bookseller (considerably more
locally). The plates include over 1000 good color photographs, and in addition 280
species are shown in line drawings. The text keys and describes in detail over 1900
species of the commoner and most attractive seed-bearing plants found in Europe, and
almost another thousand are described less fully. Many of them are also found in
North America.

I More on page 16]

CONTENTS

Post-Settlement Influences Upon a Southern Michigan Marl Lake

Peter H. Rich . 3

Vascular Plants of the Bruce Peninsula:

A Review, with Comments and Additions

George Wm. Thomson . 9

Publications of Interest . 16, 63

The Genus Panellus in North America

Orson K. Miller, Jr . 17

A New Clitocybe from Michigan

Howard E. Bigelow & Alexander H. Smith . 31

Charles Henry Swift— 1 881 -1966

Walter L. Palmer . 34

Type Specimens of Flowering Plants Described from

Long’s Expedition to the Source of the St. Peter’s River in 1823

Ronald L. Stuckey . 41

A Preliminary Report on the Varieties of

Maianthemum canadense in Northern Michigan

Stephen G. Weller . 48

The Barnes Hybrid Aspen, Populus Xbamesii, hybr. nov.—

A Nomenclatural Case in Point

W. H. Wagner, Jr . 53

Nature Education Feature-

Winter Field Key to Some Trees Common in Michigan

James R. Wells . 55

Michigan Plants in Print . 59

Information for Authors . 62

Program Notes . 63

News of Botanists . 63

(On the cover: Longitudinal section of a twig of Black Walnut,
Juglans nigra, showing characteristic chambered pith.
Photograph by James R. Wells from material gathered July 1968
along Ford Road, Washtenaw County, Michigan.)

Vol. 9, No. 2

33

<1*1

THE

MICHIGAN BOTANIST

LIBRARY

MAR 31 1370

MEW yo&k
aOTANICAL. 9AROEN

MARCH, 1970

THE MICHIGAN BOTANIST-Published by the Michigan Botanical Club four times per

year: January, March, May, and October. Second-class postage paid at Ann Arbor,

Michigan.

Subscriptions: $3.00 per year. Single copies: $.75.

All back issues are available, at the following prices:

Vol. 1 (2 numbers, at $1.00 each): $2.00

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Subscriptions (from those not members of the Michigan Botanical Club) and all orders
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with the first number of that volume unless specified to the contrary.

Articles and notes dealing with any phase of botany relating to the Upper Great Lakes
Region may be sent to the editor in chief. The attention of authors preparing manuscripts
is called to “Information for Authors” (Vol. 9, p. 62; copies available from the editor).

Editorial Board

Edward G. Voss, Editor in Chief

Herbarium, The University of Michigan, Ann Arbor, Michigan 48104

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1509 Kearney Rd., Ann Arbor, Michigan 48104

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Richard Brewer W. H. Wagner, Jr.

THE MICHIGAN BOTANICAL CLUB

Membership in the Michigan Botanical Club is open to anyone interested in its aims:
conservation of all native plants; education of the public to appreciate and preserve plant
life; sponsorship of research and publication on the plant life of the State; sponsorship of
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sanctuaries and natural areas; and cooperation in programs concerned with the wise use and
conservation of all natural resources and scenic features.

Dues are modest, but vary slightly among the chapters and with different classes of
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Membership Chairman: Harriette V. Bartoo, Department of Biology, Western Michigan Uni¬
versity, Kalamazoo, Michigan 49001
Corresponding Secretary, Southeastern Chapter:

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Secretary, Huron Valley Chapter:

Ursula R. Freimarck, 704 Madison PI., Ann Arbor, Michigan 48103
Secretary, Southwestern Chapter:

Helen Wiles, 7113 N 25th St., Rt. 2, Kalamazoo, Michigan 49004

1970

THE MICHIGAN BOTANIST

67

CHROMOSOME STUDIES OF SOME MOSSES
OF THE DOUGLAS LAKE REGION

Jerry A. Snider

Department of Botany, Duke University,

Durham, North Carolina

The moss flora of the Douglas Lake region of northern Michigan has
been thoroughly studied during the past 50 years. From the time the first
bryology course was taught at the University of Michigan Biological Station
by George E. Nichols in 1920 through to the present course taught by
Howard A. Crum, a large collection of the mosses occurring in this region has
been made. A detailed list of this flora has been published by Crum (1964;
1965), with additions by Miller and Crum (1966) and Crum and Miller
(1967). No publications are known which deal with the chromosome numbers
of mosses of the area. The present paper reports counts on six species col¬
lected in the Douglas Lake region and adjacent counties.

An outline of the techniques used is given to aid those interested in
doing further study. For additional accounts of methods and techniques the
student is referred to Steere, Anderson, and Bryan (1954) and Anderson and
Crum (1958). The method used in this paper is that adopted by Anderson in
his numerous investigations of moss cytology.

All cytological studies were made on spore mother cells (SMC) at
meiosis. Living plants were collected during July and August, 1969. Immedi¬
ately upon collection the plants were placed in polyethylene bags to keep
them fresh and moist. Plants having well -formed, healthy capsules with a faint
yellowish color in the annulus generally proved to be in the proper stages of
meiosis. Although this color character did vary from species to species and
sometimes even within the same species, it was, nevertheless, an excellent aid
to obtaining sporophytes in meiotic condition. In some instances, such as in
Pylaisiella, the annulus exhibited a translucent to a very faint yellowish color
during meiosis, whereas in Dicranum the annulus had a definite light yellow
aspect to it. The other extreme was observed in species of Brachythecium in
which a very definite yellow-brown color in the annulus was indicative of
meiosis.

The selected capsule was placed on a clean microscope slide, flooded
with fixative (3 parts absolute alcohol : 1 part glacial acetic acid) and cleansed
thoroughly of dirt, sand, and other debris. The calyptra and operculum were
removed by cutting below the annulus. It is important that the capsule and its
contents not be allowed to dry out. The spore mother cells were removed by
squeezing the columella out or by dissection if the columella refused to slip.
When the SMC were freed from the columella, or the sporogenous tissue was
dissected out in small pieces, some of the fixative was allowed to evaporate.
While the SMC were still slightly moist a drop of aceto-orcein was applied.
The cover glass was then gently applied and a slight tapping motion on the
surface of the cover slip was initiated to spread out the SMC. Pressure can

68

THE MICHIGAN BOTANIST

Vol. 9

then be applied to the cover glass in a variety of ways, including continuous
pressure with the slide inverted on absorbent paper, by tapping with various
blunt tools, or by a combination of these procedures. Some species require
only the lightest pressure while others require very rigorous pounding in order
to flatten the SMC. When the SMC were sufficiently flattened the cover glass
was sealed with vaseline or other suitable material and the SMC studied im¬
mediately. In some instances the chromosomes were slow in absorbing the
stain and it was necessary to store the preparation overnight. Finished prepara¬
tions may be stored from several days to a few weeks by placing the slide in a
refrigerator.

Voucher specimens were deposited in herbaria at Duke University, the
University of Michigan, and the author’s personal herbarium.

Figures 1-9. ca. x2300. Figs. 1-2. Bryum argenteum, n = 11. Fig. 3. Dicranum flagellare, n
= 12. Figs. 4-6. Dicranum scoparium, n = 12. Figs. 7-9. Pylaisiella selwynii, n = 10 + m.

1970

THE MICHIGAN BOTANIST

69

Bryum argenteum Hedw. n = 11 (figs. 1,2,10,11). On soil in pasture, Magee
Rd., Charlevoix Co., Michigan, 31 July 1969, Snider 894. Steere, Anderson,
and Bryan (1954) reported the number n = 12 for a California population
identified as var. lanatum BSG. Anderson and Bryan (1958) and Al-Aish and
Anderson (1961) reported the number n = 10 for populations of B. argenteum
from North Carolina. Steere (1954), however, reported the number n = 11
from an arctic Alaskan population of this species, and recognized a large
bivalent dissociating early. Although in my counts the early dissociation of a
single bivalent was recognized (figs. 2,10), no size difference was noted among
the eleven bivalents.

Dicranum flagellare Hedw. n = 12 (figs. 3,12,13). On soil bank of Five-Mile
Creek near Lake Michigan, Emmet Co., Michigan, 2 August 1969, Snider 871 .
Five counts were obtained from this population with the number n = 12

13

14

*

0

18

Figures 10-18. ca. x2300. Drawings made from photographic negatives. Figs. 10-11. Bryum
argenteum, n = 11. Figs. 12-13. Dicranum flagellare, n = 12. Fig. 14. Fissidens cristatus, n =
12. Figs. 15-16. Fissidens osmundoides, n= 12. Figs. 17-18. Pylaisiella selwynii, n = 10 +

70

THE MICHIGAN BOTANIST

Vol. 9

observed in each count. Meiotic behavior was regular and one large bivalent
was observed during metaphase I. This count differs from the number n = 23
on Canadian material by Anderson and Bryan (1958), thus contributing evi¬
dence that chromosome races exist in this species.

Dicranum scoparium Hedw. n = 12 (figs. 4,5,6). On soil near the mouth of
Five-Mile Creek, Emmet Co., Michigan, 2 August 1969, Snider 889. Due to
the ease of obtaining good meiotic metaphase stages numerous counts were
determined from the Michigan population. The number n = 12 agrees with
counts reported by Anderson and Bryan (1958) from populations in North
Carolina.

Fissidens cristatus Wils. n = 1'2 (fig. 14). On moist soil along shore of
Wycamp Lake, Emmet Co., Michigan, 9 August 1969, Snider 898. Anderson
and Bryan (1956) reported the number n = 12 + 1 for this species in Nova
Scotia, and Al-Aish and Anderson (1960) published the number n = 13 +■ 2 as
occurring in a population from Florida. Thus, the n = 12 count in this report
adds still another for this species in North America. One of the larger biva¬
lents disjoins early during metaphase I and could be recognized in all prepara¬
tions by its distinctive shape. Anderson and Bryan (1956) also recognized this
feature in the Nova Scotia population. The presence of an m-chromosome was
not observed in any of the Michigan counts.

Fissidens osmundoides Hedw. n = 12 (figs. 15,16). On soil in a burned-over
Larix bog, Sec. 25, T33N, R6W, west of M75, Charlevoix Co., Michigan, 24
July 1969, Snider 884. Chopra (1961) published the number n = 16 from a
population in the Great Smoky Mountain National Park in which he recog¬
nized four V-shaped chromosomes. In counts from the Michigan population
twelve large bivalents were easily recognized, all of a uniform size. Thus, this
report adds yet another count to the many differences existing in chromo¬
some numbers within the Section Serridium. The presence of intraspecific
chromosome races within this group will have to be studied thoroughly before
any conclusions as to relationships can be made.

Pylaisiella selwynii Kindb. n = 10 + m (figs. 7,8,9,17,18). On trunk of maple,
Five-Mile Creek near Lake Michigan, Emmet Co., Michigan, 2 August 1969,
Snider 863. Reports by Steere (1954) on arctic Alaskan populations and
Anderson and Bryan (1958) on North Carolina specimens agree on the num¬
ber n = 11, including the presence of an m-chromosome Numerous counts
from the Michigan population agree with these, including the precociously
disjoining m-chromosome (figs. 7,8,17,18).

ACKNOWLEDGMENTS

I wish to thank Dr. F. K. Sparrow, director, University of Michigan Biological
Station and the National Science Foundation (NSF GB-8440) for partial support while
conducting this study. Gratitude is also expressed to Dr. Lewis E. Anderson of Duke
University and to Dr. Howard A. Crum of the University of Michigan for their critical
reading of the manuscript, and to Dr. Crum for his aid in obtaining specimens and
verifying the identifications.

1970

THE MICHIGAN BOTANIST

71

LITERATURE CITED

Al-Aish, M., & L. E. Anderson. 1960. Chromosome numbers of some mosses of Florida.
Jour. Elisha Mitchell Sci. Soc. 76: 112-120.

Al-Aish, M., & L. E. Anderson. 1961. Chromosome studies on some mosses of the
southeastern United States. Bryologist 64: 289-314.

Anderson, L. E., & Virginia S. Bryan. 1956. A cytotaxonomic investigation of Fissidens
cristatus Wils. and F. adiantoides Hedw. in North America. Revue Bryol. Lichenol. 25:
254-267.

Anderson, L. E., & Virginia S. Bryan. 1958. Chromosome numbers in mosses of eastern
North America. Jour. Elisha Mitchell Sci. Soc. 74: 173-199.

Anderson, L. E., & H. Crum. 1958. Cytotaxonomic studies on mosses of the Canadian
Rocky Mountains. Natl. Mus. Canada Bull. 160: 1-89.

Chopra, Narinder. 1961. Chromosome numbers of some species of Fissidens Hedw.
Castanea 26: 88-93.

Crum, H. 1964. Mosses of the Douglas Lake region of Michigan. Mich. Bot. 3: 3-12,
48-63.

Crum, H. 1965. Additions to the Douglas Lake Bryoflora. Mich. Bot. 4: 26-30.

Crum, H., & N. G. Miller. 1967. Bryophytes from the Upper Falls of the Tahquamenon.
Mich. Bot. 6: 54-57.

Miller, N. G., & H. Crum. 1966. Bryophytes new to the Douglas Lake area. Mich. Bot. 5:
57-61.

Steere, W. C., L. E. Anderson, & Virginia S. Bryan. 1954. Chromosome studies on
Californian mosses. Mem. Torrey Bot. Club 20: 1-75.

Steere, W. C. 1954. Chromosome number and behavior in Arctic mosses. Bot. Gaz. 116:
93-133.

Review

HOW TO KNOW THE LICHENS. By Mason E. Hale. Wm. C. Brown Co., Dubuque. 1969.

226 pp. $3.00 (spiral); $3.75 (cloth).

Here is a lichen key that really works, and it’s a bargain too. The elegant illustra¬
tions, together with pertinent comments on specific characteristics, comparisons with
related species, and distribution maps, make determinations virtually certain. Though
exact enough to satisfy even professional lichenologists, the keys can be understood and
easily used by the less informed, even to the technical level of a high school student. The
introductory pages give a good background of information on morphology and methods
of lichen taxonomy, including color and crystalline tests (with valuable photographs of
crystals) and chromatography. Students and those who direct the unlimited energies of
students will find use for information on such projects of interest as studies of growth
rates and lichen synthesis and techniques of dyeing.

It is interesting to note that the author never uses a handlens or dissecting micro¬
scope himself, and the characters on which he bases his key are accordingly quite easily
observed. The fact that the author knows his field so well adds authority to a seemingly
effortless and simplified approach to an esoteric study. The key has been tested by three
lichen classes, under the author’s careful supervision, including one of my own at the
University of Michigan Biological Station. This book provides names for nearly all foliose
and fruticose lichens of North America, from the Arctic to the northern boundary of
Mexico, excepting a few poorly understood genera, such as Usnea, Ramalina, and
Collema, which are “summarized rather broadly.”

-Howard Crum

72

THE MICHIGAN BOTANIST

Vol. 9

INVESTIGATIONS IN THE

WHITE WATERLILIES (NYMPH AEA) OF MICHIGAN1

Gary R. Williams

Department of Science, Glenbard North High School,

Carol Stream, Illinois

INTRODUCTION

Distinguishing Nymphaea odorata Ait. from N. tuberosa Paine has been
a problem for botanists for many years. Fassett (1940) recognized that some
profess difficulty in differentiating these species. Sculthorpe (1967) refers to
the Nymphaea odorata complex and to the fact that in Nymphaea there are
abundant natural hybrids, most of which are sterile. Conard (1916) states

It seems highly desirable, therefore, that critical studies should be made of
the white waterlilies of all the Great Lake region, and the Central States. Every
detail mentioned in the table given above should be carefully examined into. Only
thus can the taxonomic value and the range of these plants be determined. At
present I do not place entire confidence in any of the published names.

A search of the literature reveals that many authors have worked with
Nymphaea in some detail, particularly in the late 1800’s and early 1900’s. The
most complete summary exists in H. S. Conard’s The Waterlilies , published in
1905. One must keep in mind that the older literature uses the name Castalia
for our Nymphaea, and Nymphaea for today’s Nuphar.

The diagnostic characters generally used in Nymphaea, although many,
are sometimes qualitative and thus indistinct. Waterlilies having characteristics
of both N. odorata and N. tuberosa are found in many areas. Because of the
difficulty indicated by the above statements— and experienced by me in recog¬
nizing and separating the species in the University of Michigan Biological
Station area— a survey of the full range of variations in Nymphaea in this
region was begun in 1964. The initial attempt was simply to analyze the N.
odorata-N. tuberosa complex as to morphological characters that could be
used to separate the species. During the summers of 1965 through 1969, more
detailed studies were undertaken, with financial assistance from The University
of Michigan Biological Station and the National Science Foundation.

The taxa in the Nymphaea odorata-N. tuberosa complex have been
differentiated by various characters, many of which are summarized in Table
1. All of these characters were evaluated in addition to the number of
peduncle coils when in fruit; presence or absence of petiole or peduncle hairs;
aril size; leaf shape; number of air tubes in petiole; acid-base affinity; water
depth preference; temperature and/or weather conditions required for opening
and closing; and shelter or protection requirements. A checklist was designed
to standardize observations.

iContribution from the Biological Station of The University of Michigan.

1970 THE MICHIGAN BOTANIST 73

TABLE 1. Traditional

characters used in differentiating Nymphaea odorata and N.

tuberosa. All characters from Femald (1950) except those marked with an asterisk (*),

from Fassett (1940).

Character

Nymphaea odorata

Nymphaea tuberosa

1. Number of stamens

36-100

35-106

2. Opening time

Early morning

8:00 a.m.

*7:00 a.m.

*8:00 a.m.

3. Closing time

None

Early afternoon

*1:00 p.m.

*3:00 p.m.

4. Petiole

Purple-green to red

Green striped with brown
* Usually 4-5 purple streaks

5. Sepal length and/or

width

2.8-8 cm x 1-2.5 cm

6. Sepal shape

Ovate-lanceolate

7. Sepal color

Purple or green

Green

*Often purple

*Green

8. Number of petals

17-32

17-32

9. Shape of petals

Tapering ovate

Broadly rounded at summit

*Blunt or pointed

*Often broader than odorata, round

10. Stamens

Inner filaments narrower

Filaments broader than anthers

than anthers

11. Tubers

Occasionally present, but

Present, constricted at base,

not strongly constricted
at base.

readily detachable.

12. Leaf size

0.5-2.5 dm across

0.5-6 dm wide

*as much as 4 dm wide

*as much as 4 dm wide

13. Underside of leaf

Usually purple beneath

Green, rarely purple

*Often purple

*Green

14. Fragrance

Strong

None or faint

* Present

*No

15. Diameter of open

5-15 cm

1-2.5 cm [error for dm]

flowers

*Seldoin more than 12 cm

* Usually larger than odorata

METHODS AND OBSERVATIONS

Observations were made at the
and Ontario, Canada:

ALGER COUNTY
Deer Lake

CHEBOYGAN COUNTY
Vincent Lake
Little Lake 16
East Lake
Duck Lake
Duck Marsh
Upper Black River
Black River

Fontinalis Run at Burt Lake

following locations in northern Michigan

Douglas Lake
Munro Lake
Mullett Lake

Mud Lake (Inverness Tp.)

Black Lake

EMMET COUNTY

Carp (Paradise) Lake
French Lake
Wycamp Lake
Crooked Lake
Hay Lake

74

THE MICHIGAN BOTANIST

Vol. 9

MARQUETTE COUNTY (Huron
Mountain Club)

Mountain Lake
Howe Lake
Rush Lake
Pine Lake
Ives Lake
Lily Lake

OTSEGO COUNTY
Lake Manuka

PRESQUE ISLE COUNTY
Ocqueoc Lake

SCHOOLCRAFT COUNTY
Cusino (Round) Lake
Gemini Lake
Ross Lake

ALGOMA DISTRICT, ONTARIO
Black Trout Lake

At most locations, 35 mm color reference slides were taken and material
collected for herbarium specimens and for analysis in the laboratory while
fresh. Care was taken to collect leaves, flowers, and rhizomes from each loca¬
tion. It is almost useless to collect only leaves and flowers without being sure
they are arising from the same rhizome. It is also much more significant to
look for tubers along the rhizome. Although this is not an easy task, it is not
impossible and can usually be accomplished by wading rather than having to
dive. The rhizome is usually covered by only 1-2 inches of mud and silt, and
can be taken by following a petiole to its base, grasping the rhizome, and

Fig. 1. Nymphaea flower photographed August 12, 1969, at Hook Point, Douglas Lake,
Michigan.

1970

THE MICHIGAN BOTANIST

75

breaking off a section about 1-2 feet in length. Care should be taken to bring
up tubers when present since they break off easily.

Material for chromosome study from root-tips and pollen grains was
collected and fixed in Newcomer’s solution or 3 : 1 ethanol : acetic acid.
Chromosomes were stained with aceto-carmine or aceto-orcein and the squash
technique was used. Efforts met with little success due to the very small size
of the chromosomes and their high number (2n = 84, Langlet & Soderberg,
1929; 2n = 56, Sculthorpe, 1967). On the average, only about 10-30% of the
pollen grains from a single anther were deeply stained. The chromosomes are
very difficult to get into a sihgle plane in order to count them. In the few
slides which were more or less acceptable, it could only be determined that
the chromosome numbers were in the range of 50-70. Not enough data have
been collected for this to be helpful information.

The first clone of white waterlilies examined by me was one at Hook
Point in Douglas Lake (T37N, R3W, Sec. 22). The flowers (Fig. 1) are fra¬
grant, with narrow inner filaments, and in a size range indicating Nymphaea
odorata. The leaves are red to purple underneath, with faintly striped petioles,
also indicating N. odorata . Petal shape, sepal color, presence of distinctly

Fig. 2. Rhizome portion collected and photographed August 12, 1969, at Hook Point,
Douglas Lake, Michigan. Note coiled peduncle, tubers, and bicolored leaf (actually pink
underside with green margin where not in contact with water); cf. Fig. 11.

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THE MICHIGAN BOTANIST

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constricted tubers, petiole and peduncle striping (although faint), and leaves
that are green beneath when not in contact with the water all point to N.
tuberosa (Fig. 2). In retrospect, these plants are nearly perfect intermediates.

The time of opening and closing is widely considered to be diagnostic,
with flowers of Nymphaea odorata opening about one hour earlier (7:00 a.m.)
and closing about two hours earlier (noon or 1:00 p.m.) than those of N.
tuberosa. After tagging plants at Hook Point in Douglas Lake and at Fonti-
nalis Run, and observing plants in these several lakes, I feel that these char¬
acters are not valid. At Hook Point, for example, flowers began to open at
8:30 a.m. and to close at 3:45 p.m. EST. Flowers continued to open through¬
out the morning and some remained open at 5:05 p.m. Although no tagging
was done in the Upper Black River, flowers have been found closed at 11:00
a.m., opening shortly thereafter. Most flowers are still open at 4:30 p.m., but
the time of closing has not been observed. At Carp Lake, Mr. Ed Nantais
observed a clone of Nymphaea at the spillway near his motel during a two-
week period in late July and early August, 1964, and reported that the
flowers open by 9:00 a.m. and close by 4:00 p.m.

Seeds were collected for study from French Lake and the Upper Black
River. These areas were selected because their plants produce abundant seeds
and are easily accessible. The plants from French Lake are similar to the

Fig. 3. Nymphaea flower photographed July 24, 1969, in French Lake, Michigan.

1970

THE MICHIGAN BOTANIST

77

Nymphaea odorata phenotype (flower in Fig. 3) while those from the Upper
Black River are most like N. tuberosa (flower in Fig. 4). Seeds were separated
by making a cross section through the ovary and squeezing the two halves.
The seeds were rinsed several times in tap water, measured, labelled, bottled,
and stored in tap water. Approximately half of the seed collected was kept in
a four-inch finger bowl at room temperature. The water was changed twice
per week. The other half of the seed was kept in a refrigerator at approxi¬
mately 11° C. The refrigerated seeds were further separated after three months
with half of them being put in an aquarium at room temperature and the
other half remaining in a refrigerator. Only among the latter did any germina¬
tion occur. Germination results are summarized in Table 2. Approximately
half of the seeds which germinated were placed in dark jars and half of them
in an aquarium which was illuminated from 18 inches above by a “Gro-Lux”
light. No further growth was noted in any of the seedlings. Figure 5 shows a
Nymphaea seedling— quite unlike the typical monocot or dicot seedling.

Tetrazoleum, a chemical used extensively to estimate percentage of
germination of hybrid seed corn, was tested on Nymphaea seeds in the sum¬
mer of 1967. The powder was mixed with distilled water to make a 1%
solution and seeds from French Lake were soaked in it. Ten seeds were cut
open each day for eight days. No staining was observed until the third day,

Fig. 4. Upper Black River Nymphaea. Photographed August 14, 1969.

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THE MICHIGAN BOTANIST

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TABLE 2. Results of germination tests on Nymphaea seeds.

Seed Source

Condition

Upper Black River

French Lake

Douglas Lake

Carp Lake

Room temperature

0

0

0

0

1 1°C for six months

14.4%

31/216

21.4%

48/224

0

0/180

7.4%

11/149

11°C for three months

0

0

0

0

- to room temperature

when three seeds had a slightly pink cast in longitudinal section. After eight
days, of the ten seeds cut, four displayed some pink to red staining and six
showed no staining at all. Further work is being done on cold-treated seeds.

Tubers were collected in all areas studied and some were allowed to
sprout in finger bowls in the laboratory (Fig. 6). These tubers sent up leaves
for the entire summer, and when they were transferred in September to an
aquarium four inches deep, 18 inches from a “Gro-Lux” light, continued to
send up leaves until the following July, when they were discarded. This gives
some idea of the potential efficiency of tubers as propagules. Sculthorpe
(1967) recounts the plowing of a field near Buffalo, New York, that uncov¬
ered many Nymphaea tubers which when planted in tubs grew into a deep
pink variety of N. odorata. The land had been drained for many years and a
pond had existed there, possibly a century before.

Fig. 5. Germinating Nymphaea seedling.

ASSOCIATED ANIMALS

The following macroscopic animals have been observed and/or collected
in or on the plants.

BRYOZOA:

Plumatella, Fredericella, and Hyalinella on stalks and undersides of leaves.

1970

THE MICHIGAN BOTANIST

79

Fig. 6. Tuber, with leaves and roots, collected from Upper Black River, Michigan, August
14, 1969. Actual length was approximately 5 cm.

HIRUDINEA:

Various, particularly Helobdella, Placobdella, very common under leaves.
INSECTS:

Odonata larva on stalks.

Homoptera— Aphidae kre very common in flowers.

Coleoptera-Chrysomelidae, Curculionidae, Haliplidae, Dytiscidae, and Gyrinidae
larvae on stalks and under leaves. Adults in flowers or on leaf tops.
Donaciinae adults are the commonest beetles on flowers and leaves. The
eggs are apparently laid on the underside of leaves through holes cut in the
leaves.

Diptera-many egg rafts are in association with leaves. Some are stuck on the
underside of the leaves. Chironomidae larvae are common in the petiole and
peduncle air tubes. They are so common in Black Lake and the Upper Black
River plants that the appearance of a red “worm” can almost be considered
a diagnostic character. Drosophila are common in the flower.

HYDRACARINA:

Swimming in near vicinity, sometimes more or less attached to the stalks.

MOLLUSCA:

Various Gastropods quite common on undersides of leaves and on the stalks.

Physa, Bulimnaea, Ferrissia, Stagnicola, Lymnaea, and Goniobasis have been

frequently collected.

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THE MICHIGAN BOTANIST

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The above list deserves mention, because some of these organisms prob¬
ably utilize the leaves and stalks (peduncles and/or petioles) of waterlilies and
may cause many of the problems in maintaining transplanted clones. Some of
the insects listed above probably aid in the pollination of the flowers. Scul-
thorpe (1967) indicates that waterlilies are pollinated by beetles, sweat bees,
and flies that crawl indiscriminately over the flower. The principal attraction
is probably the warmth and smell of the opening flowers and food value of
the stamens, since the flowers are nectarless.

TRANSPLANT STUDIES

In the summer of 1965, initial transplants were made in an attempt to
discover the effect of the environment on the phenotype of the complex.
Additional transplants were made in 1966, 1967, and 1968. Reciprocal trans¬
plants were made whenever possible. In a specified pair of lakes, the same
location in each lake was used both for collecting and transplanting rhizomes.
Two basic methods were used: 1. A rhizome section and soil were put into a
2X3 foot box, six inches deep, and the box was weighted with rocks (Fig.
7). 2. A rhizome section was put into a 4 X 12 inch clay drain tile which was
partially buried in the substrate (Fig. 8). Any attached leaves were adjusted to
allow them to float if possible. Fertilizer when used was a mixture of about Vi
dry cow manure and Vi soil, which was moistened, allowed to sit for a few
days, and then added to the box or tile near the growing rhizome. The tile
method has several advantages: 1. The rhizome is better protected and more
stable. 2. It is much easier— no boxes to build. 3. No size-limiting factors are
presented as the growing tip of the rhizome can penetrate surrounding sub¬
strate. 4. The possibilities of other “unwanted” tubers, seeds, or other propa-
gules washing in is nearly precluded. 5. The tile is more permanent and easier
to relocate than the box.

Table 3 summarizes the transplant results. The most interesting trans¬
plants are those that “perform” by growing and producing vegetative and
reproductive structures. The transplants from Douglas Lake to the Upper
Black River, Fontinalis Run to Neu’s pond in Glen Ellyn, Illinois, and from
the Upper Black River to Cusino Lake are thriving. The July 16, 1966, trans¬
plant from Fontinalis Run (apparently a cultivated strain) to Neu’s pond
produced its characteristic pink-magenta flowers in 1968.

The August 3, 1965, transplant from the Upper Black River to Cusino
Lake is unquestionably the healthiest and thus the most valuable for purposes
of observation even though the comparatively remote location of Cusino Lake
practically precludes frequent observation of the transplant. A brief descrip¬
tion of Cusino Lake is given by Voss (1965). On the cover is a photograph of
a Cusino Lake flower. As Table 4 indicates, the transplant has each year
produced leaves, buds, and presumably flowers (except in 1969), although
fully open flowers have not been observed. One can see that a rhizome from a
plant with good characters of Nymphaea tuberosa produced new leaves each

1970

THE MICHIGAN BOTANIST

81

■ ■ Ww* ’ ' . t;'' "■ '

Fig. 7. Rhizome portion of Nymphaea, demonstrating the box method of transplanting.

Fig. 8. The tile method of transplanting Nymphaea rhizome

82 THE MICHIGAN BOTANIST Vol. 9

TABLE 3. Summary of Nymphaea transplant

Rhizome Source Transplanted to

studies.

Box or Tile
Date F = + fer¬
tilizer

1966

Observations*

1967 1968

1969

1965

Upper Black River

Cusino Lake

8/3

B -F

1

1

l(Bud) 2

Cusino Lake

Upper Black River

8/5

B -F

3

6

6

6

Upper Black River

Douglas Lake

8/5

B -F

6

6

6

6

Fontinalis Run

Douglas Lake

8/5

B -F

6

6

6

6

Upper Black River

Duck Lake

8/6

B -F

6

6

6

6

Mud Lake Bog

French Lake

8/6

B -F

3

4

6

6

Upper Black River

Lancaster Lake

8/9

B

6

6

6

6

Upper Black River

French Lake

8/10

B

4

6

6

6

Duck Marsh

French Lake

8/10

B

3

6

6

6

French Lake

Upper Black River

8/13

B

2

2

2

2

Duck Lake

Upper Black River

8/13

B

5

6

6

6

Douglas Lake

Upper Black River

8/13

B

5

6

6

6

1966

French Lake

Upper Black River

7/1

T -F

3

6

6

Upper Black Lake

French Lake

7/1

T

4

6

6

Lancaster Lake

French Lake

7/8

T

4

6

6

East Lake

Upper Black River

7/8

T

6

6

6

Carp & Black Lakes,

Pool at Ed Neu’s

Fontinalis Run

Glen Ellyn, Illinois

7/16

B

2

1

2

Carp Lake

Upper Black River

7/27

T-F

4

6

6

Upper Black River

Cusino Lake

7/29

T

4

6

6

Cusino Lake

Upper Black River

8/2

T-F

4

6

6

Duck Marsh

Upper Black River

8/9

T

3

6

6

Duck Marsh

Black Lake Bay

8/9

T

5-2

5-2

5-2

Black Lake

Duck Marsh

8/9

T

2

2

2

Upper Black River

Duck Marsh

8/9

T

6

2

2

Canada & Duck Marsh

St. John’s College

Santa Fe, New Mexico

8/16

B

2

2

4

Duck Marsh &

Black Lake

Lake Ellyn, Illinois

8/24

T

2

2

2

1967

Douglas Lake

Upper Black River

6/28

T-F

1

2

Upper Black River

Carp Lake

7/1

T

2

2

Carp Lake

French Lake

7/7

T

6

6

Duck Marsh

Upper Black River

7/13

T-F

4

6

Upper Black River

Douglas Lake

7/19

T

6

6

1968

Carp Lake

French Lake

7/19

T-F

2

Douglas Lake

Upper Black River

7/26

T-F

2

*KEY: 1 = Survived and flowered; 2 = Survived, produced vegetative parts; 3 = Survived,
produced small purple juvenile floating leaves, but died during the summer; 4 = Died
during the winter; 5 = Washed out of position; 6 = Transplant could not be found.

1970

THE MICHIGAN BOTANIST

83

TABLE 4. Summary of Upper Black River-Cusino Lake Nymphaea transplant.

Character Native Cusino Transplant from Upper Black River

Lake Plants

8/3/65

7/29/66

8/19/66 7/21/67

7/31/68

7/30/69

Leaf diameter (mm)

145

180

135

130

115-120

110-135

115-120

Leaf shape*

2 & 3

1

1

1

1

1 (2)

1

Petiole diameter (mm)

4

6

4

3-4

3

3-4

3

Petiole stripes

0

+

0

0

0

+

0

Leaf color, upper

Green

G

G

G

G

G

G

Leaf color, underside

Red

G

R-Pink

R-Pink

R-Pink

G(Pink) R-Purple

Leaves, number of

4

12

12

7

5

22

Buds

2

1

3

3

2

0

:Leaf shapes: 1 =

year which have been smaller and more purple on the average than those of
the original plant. The petioles have tended to be smaller, less distinctly
striped, and pinker. In other words, they tend to be more and more like the
Cusino phenotype (N. odoratal) and less like that of the Upper Black River.
The color change in the undersurface of the leaves is most dramatic (Fig. 9).
This would lead one to the conclusion that the phenotypic differences in the
Nymphaea of the Upper Black River and Cusino Lake are dictated largely by
the environmental factors and less by genotypic factors. In 1967, the color of
the underside of the leaves of the transplant was green with only a slight
touch of pink on some of them. This would seem to be a reversion to the
“parental” phenotype. I interpret this condition to be a reaction to increased
water depth with assumed attendant change in average temperature, pH, light
penetration, etc. Unfortunately, the reciprocal transplant from Cusino Lake to
the Upper Black River did not survive; the leaves were riddled with holes by
an unknown predator and it did not produce leaves after 1966.

DISCUSSION AND CONCLUSIONS

Most of the characters listed earlier in this report— such as leaf color,
flower odor, and tuber production— are not sufficient to differentiate two
species of waterlilies in the Douglas Lake area, although there are certainly
distinct differences in the plants from the various lakes studied. Pink-tinted
outer petals in the East Lake clone; a distinctive green wedge on the underside
of the Cusino Lake leaves (Fig. 9), and leaves with upturned edges from the
Black River make these clones identifiable to most observers. Duck Marsh
plants have leaves with three and five petiole tubes as well as the usual four
(Fig. 10).

The color of the underside of the leaf, long considered to be a good
diagnostic character, is apparently not valid— at least in this area. The dramatic

84

THE MICHIGAN BOTANIST

Vol. 9

color change in the transplanted material in Cusino Lake attests to this fact.
The somewhat less striking changes in the transplants from Douglas Lake to
the Upper Black River and others also bear this out. The transplant in each
case develops red or pink undersides of the leaves if green initially, or lighter
red undersides, if red or purple originally. Juvenile leaves tend to be red or
brown and much smaller than the mature leaves. Another interesting fact is
that in cases where leaves are held up out of the water, the underside will be
green, even though floating leaves on the same plant are red underneath. One
leaf collected ( Williams 76) at Hook Point was floating and in contact with
the water over 80 per cent of its undersurface and about 20 per cent of the
edge was held out of the water by adjacent vegetation. This leaf was red
where in contact with water and green where exposed (Fig. 11). The implica¬
tion here is that red pigment is produced in response to water contact or its
production is inhibited by exposure to air and/or sunlight.

Odor is not a good character to distinguish our taxa, since nearly all
flowers tested were fragrant on the first day of opening. Admittedly, however,
some were more fragrant than others.

Tubers were found in all areas where collections were made. Some
rhizomes had more tubers per foot than others and some were more con-

Fig. 9. Undersides of 3 Nymphaea leaves. At the left is a 25 cm diameter leaf from
Upper Black River, August 14, 1969. At the upper right is an 11 cm leaf from Cusino
Lake. The one on the lower right is a 10 cm leaf taken from the transplant from Upper
Black River to Cusino. Both Cusino specimens were collected August 15, 1969. Note the
lighter (green) color of the Upper Black River leaf and the darker (red-purple) color of
the other two. The Cusino leaf displays a lighter (green) wedge running along the midrib.
The distinctive shape of the leaf apexes is also shown.

1970

THE MICHIGAN BOTANIST

85

Fig. 10. Cross-sections of Nymphaea petioles from Upper Black River (left) and Cusino
Lake (right). The larger petiole measured 6 mm in diameter. Note that both have 4 major
petiole tubes although the former has 2 large and 2 small tubes while all 4 tubes are
more or less equal in the latter.

Fig. 11. Bicolored leaf collected August 12, 1969 at Hook Point, Douglas Lake, Michi¬
gan. The lighter (green) margin at the apex and sinus was not in contact with the water
while the darker (red-purple) portion was floating on the water’s surface. Actual diameter
was 17 cm.

86

THE MICHIGAN BOTANIST

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stricted at the base than others, but tubers were always present (Fig. 2).

The peduncles coil and pull the fruit under the surface of the water
after the flower is fertilized. This characteristic is well displayed in French and
Douglas Lakes, but the number of coils is not constant even in flowers grow¬
ing from the same rhizome (Fig. 2).

I do not believe that hybridization is an important factor because of a
lack of evidence of reproduction by seeds. In the six-summer period, only one
seedling was discovered that could definitely be identified as Nymphaea.
Nuphar seedlings are quite common and could be confused with Nymphaea at
first glance. The lack of good staining points to abortive or sterile pollen. The
size and color difference in seeds and ovules indicates that many, perhaps
most, of the ovules are not fertilized. In many mature ovaries are found
mostly yellow seeds 1 mm long and only a few red to brown seeds 3-4 mm
long. From tetrazoleum and germination tests, apparently only the larger ones
are viable seeds. Most reproduction is by tubers breaking off and floating
away to become lodged and rooted elsewhere (Fig. 6). Germinating tubers can
be readily found in nearly any of the lakes examined. The leaves from tubers
can be identified by their small size (about 15 X 20 mm) and uniform
brown-red color throughout.

The white waterlilies in the Douglas Lake area are a highly variable
taxon and there seems to be no good reason for calling them distinct species.
They are most certainly a single species or very closely related species that are
showing variable responses to their environments.

ACKNOWLEDGMENTS

This work was sponsored in part by a National Science Foundation Academic Year
Institute, the University of Michigan Biological Station with National Science Foundation
support, and National Science Foundation Summer Fellowship Grant #75210. Repre¬
sentative herbarium material is deposited at the University of Michigan Biological Station
and in my own collection at Glenbard North High School, Carol Stream, Illinois. I
especially want to thank Dr. E. G. Voss (The University of Michigan) for his guidance,
the administration of Glenbard Township High School District 87 for their encourage¬
ment, and my wife for deciphering my notes and typing the manuscript. The cost of
printing additional pages in this article has been defrayed by the Asa Gray Bulletin Fund,
originated by the late Professor H. H. Bartlett of The University of Michigan.

LITERATURE CITED

Conard, H. S. 1905. The Waterlilies A Monograph of the Genus Nymphaea. Carnegie
Inst. Publ. 4. 279 pp.

Conard, H. S. 1916. The white waterlily of Iowa. Proc. Iowa Acad. 23: 621-623.

Fassett, N. C. 1940. A Manual of Aquatic Plants. McGraw-Hill, New York. 382 pp.
Fernald, M. L. 1950. Gray’s Manual of Botany, ed. 8. Am. Book Co., New York. 1632
pp.

Langlet, O., & E. Soderberg. 1929. Uber die chromosomenzahlen einiger Nymphaeaceen.
Acta Horti Berg. 9: 95-104.

Sculthorpe, C. D. 1967. The Biology of Aquatic Vascular Plants. Edward Arnold, Lon¬
don. 610 pp.

Voss, E. G. 1965. Some rare and interesting aquatic vascular plants of northern Michigan,
with special reference to Cusino Lake (Schoolcraft Co.). Mich. Bot. 4: 11-25.

1970

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87

ADDITIONAL BRYOPHYTES FROM SINKHOLES
IN ALPENA COUNTY, MICHIGAN,

INCLUDING ORTHOTRICHUM PALLENS NEW TO
EASTERN NORTH AMERICA1

Norton G. Miller2 and Dale H. Vitt
Department of Botany and Plant Pathology,

Michigan State University, East Lansing,
and Herbarium, The University of Michigan, Ann Arbor

One of the results of our field work in northern Michigan during
the summer of 1968 was extensive collecting in several limestone sinks lo¬
cated in northern Alpena County. Many interesting and rare ferns were re¬
ported from these sinks by Hagenah (1955). He listed Matteuccia pensyl-
vanica (Willd.) Raymond, Cystopteris fragilis (L.) Bernh., C. bulbifera (L.)
Bernh., Dryopteris spinulosa (O.F. Mull.) Watt, D. filix-mas (L.) Schott,
Athyrium pycnocarpon (Spreng.) Tidestr., A. thelypterioides (Michx.) Desv.,
Asplenium trichomanes L., and Polypodium virginianum L. Wagner and Lord
(1956) reported Botrychium lunaria forma onondagense (Underw.) Butters &
Abbe, along with Cystopteris dickieana Sim, from a sink near Bolton, noting
that this was the only locality for this form of Botrychium in the Biological
Station region. Robinson and Wells (1956) studied the bryophyte flora of the
sinks and published a list of 110 species including three new to Michigan:
Seligeria calcarea (Hedw.) B.S .G.,Mannia siberica (K. Mull.) Frye & Clark, and
Tritomaria scitula (Tayl.) Joerg.

After several collecting trips to the sinkholes, we can add the moss
Orthotrichum pallens as new to eastern North America and several noteworthy
records of rare or infrequently collected hepatics.

THE SINKHOLE AREA

The sinks at the four sites studied by Robinson and Wells (1956) occur
across a distance of 10 miles in north-central Alpena County from just north
of Long Rapids east to Cathro. All have developed in the Middle Devonian
Alpena limestone and judging from their small size and relatively uneroded
sidewalls, which show no evidence of glacial abrasion, the sinkholes appear to
be postglacial in origin. During the summer of 1968, either jointly or inde¬
pendently, we revisited all of the sinks originally studied by these workers.
However, because our collections from “Bolton Sink” and the “Four Sinks”
are most noteworthy, this report will treat only these two localities.

1 Contribution from the University of Michigan Biological Station. Research sup¬
ported in part by National Science Foundation grant GB-8212 to N. G. Hairston,
University of Michigan, for research in Systematic and Evolutionary Biology and in part
by grant GB-6095 to H. A. Crum for Workshops in Bryology at the University of
Michigan Biological Station.

2Present address: Arnold Arboretum, Harvard University, Cambridge, Massa¬
chusetts.

88

THE MICHIGAN BOTANIST

Vol. 9

Bolton Sink is unquestionably the most interesting sink in the region. It
is located about 2 miles southeast of Bolton near the center of the northern
half of Section 14 (T. 32 N., R. 7 E.)3 and is the only one we visited with an
extensive talus slope and persistent ice deposits. The sink is best reached by
walking south from Daggett Road. Open fields, thin secondary forests and low
limestone outcrops occur around the sink and the surrounding land is decided¬
ly dry. At most places flat-lying limestone bedrock appears to be less than a
meter below the surface. The sink itself is small, about 200 meters in diameter
and 25 meters deep. The east and part of the north side are bare and mostly
covered by a talus of loose boulders. A dense growth of Thuja occidentals oc¬
curs on other slopes and limited exposures of bedrock are found just below the
rim at various locations but particularly on the south side. The more or less flat
bottom, scarcely 10 m across, is mostly shaded and covered by a jumble of
large and small boulders. The talus slopes above the bottom are fairly steep
but stable where tree-covered and in total aspect the sinkhole is funnel-shaped.

It is evident that the talus is largely man-made, for it is a mixture of
native limestone rock and glacier transported igneous and metamorphic
boulders. Local farmers undoubtedly found the sink a convenient dump for
the numerous bothersome rocks that cluttered their fields. Through the years
the talus slopes were gradually built, first slowly by limestone blocks that
broke off the sidewalls and fell into the hole and later, after the settlement of
the region, more rapidly by European man who added many tons of field
stones. The massive rock piles seen frequently in northern Alpena County
attest the former abundance of rock in the soil. In contrast to several nearby
sinks very little trash has been thrown into Bolton Sink. However, charred
stumps and logs are an indication that fire has swept the site at least once.

Ice is found well into the summer near the bottom in spaces between
boulders and may persist year round if the summer months are especially cool
and dry. Robinson and Wells report seeing ice as late as June 26 during 1955
and we observed a large mass of ice near the bottom at the north side on 4
August 1968. However, somewhat later in August of the following year none
was visible even though the air temperature was much lower at the bottom of
the sink than at the rim (M. Frohlich, pers. comm.). The luxuriant growth of
Drepanocladus uncinatus, Hylocomium splendens, Marchantia polymorpha var.
aquatica, and Tritomaria scitula is evidence that the cool humid environment
at the bottom of Bolton Sink, in contrast to the more xeric surrounding
region, is especially favorable to the growth of bryophytes.

The Four Sinks are located 2 miles southeast of Leer in the NW % of
Section 16 (T. 32 N., R. 6 E.). They occur in a woodlot and for the most
part are wooded themselves. Three of the sinks have extensive areas of ex¬
posed limestone sidewalls and are considerably larger than the sink near
Bolton. Talus is mostly absent and the best bryophyte habitats are on ledges

3Robinson and Wells located Bolton Sink in the NW % of Section 12 (T. 32 N., R.
7 E.), but after first searching this quarter-section and later writing to them, we are
certain that the correct location is Section 14.

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89

and in crevices of the limestone, as well as on moist decorticated logs found
at the bottom of the sinks. The collections we record here were made at the
third and fourth sinks in from Leer Road.

ORTHOTRICHUM PALLENS

Orthotrichum pallens Bruch ex Brid., Bryol. Univ. 1: 788. 1827.

Orthotrichum pallens var. typicum Vent., Hedwigia 12: 20. 1873. nom.
illeg.

Dorcadion pallens (Bruch ex Brid.) Lindb., Musci Scand. 28. 1879.

Orthotrichum pallens var. commune Vent, in Husn., Muse. Gall. 189.
1887.

Orthotrichum pallens var. saxicola Burch., Hedwigia 31: 30. 1892.

Orthotrichum pallens ssp. eu-pallens Giac., Atti 1st. Bot. Pavia V. 4:
248. 1947. nom. illeg.

Plants 3-10 mm high, in yellow-green, olive-green, or dark brown, dense
to loose tufts. Leaves loosely erect-appressed and slightly incurved when dry,
erect-spreading to spreading when moist, 1. 6-3.0 mm long, oblong-lanceolate
to lanceolate, narrowly obtuse to broadly acute; margins entire, recurved-
revolute to near apex; costa slender, ending near the apex; upper cells 9-14 p
wide, irregularly rounded to rounded-hexagonal, thick-walled, with 2(3) low,
conical papillae; basal cells rectangular, hyaline, grading to quadrate at
margins; broad bodies sometimes present on leaves. Cladautoecious. Setae
0.5-1. 5 mm long; capsules 1.3-2.25 mm long, emergent to barely exserted,
oblong-cylindric, strongly 8-ribbed and becoming narrowly cylindric and con¬
tracted along the entire length when old, contracted to the setae through a ±
short neck, exothecial cells differentiated into bands 4-6 cells wide; stomata
immersed in 1 or 2 rows at the middle of the capsule, scarcely or not covered
by the poorly differentiated subsidiary cells, the inner walls not or scarcely
thickened; peristome double; exostome teeth 8, evenly and densely papillose,
erect until old, then reflexed; endostome segments 16, rarely 8 of these rudi¬
mentary or lacking, about as long as exostome, incurved and conspicuous,
smooth or finely papillose, margins sometimes crenulate, of one or two rows
of cells. Spores 11-17/1, coarsely papillose. Calyptrae conic-mitrate, plicate,
naked, smooth or slightly papillose on plications. Figures 1-8.

Habitat: On the trunks of trees or on tree branches and on dry rocks,
usually calcareous or granitic.

Distribution: Europe— Austria, Germany, Switzerland, Scandinavia, Italy,
France, Russia; North America— see Figure 9.

Specimens Examined from Eastern North America: CANADA— Quebec.
Gaspe Peninsula, Perce, Stifler, July 1938, ex herb. Grout (DUKE). U.S.A.—
Michigan. Alpena Co., Bolton Sink, on Thuja branches, Vitt 886, 887 (MICH),
Miller 4913, 4914; on limestone boulder, Vitt 888, Robinson & Wells, June
26, 1955 (MICH). Keweenaw Co., Fort Wilkins, Sharp, Sept. 1, 1937 (TENN).

Discussion: Orthotrichum pallens is distinguished by having immersed
stomata which are scarcely covered by thin-walled subsidiary cells over the
spore-bearing portion of the capsule. The peristome is double with a well

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THE MICHIGAN BOTANIST

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Fig. 1-8. Orthotrichum pallens Bruch ex Brid. : 1. Leaves, x 33. 2. Peristome from inside,
x 108. 3. Mature capsules, x 19. 4. Old and dry capsule, x 19. 5. Calyptra, x 19. 6.
Stoma, x 177. 7. Basal leaf cells, x 306. 8. Upper leaf cells x 306.

1970

THE MICHIGAN BOTANIST

91

developed endostome of 16 segments which remain intact until the capsule is
quite old. Gametophytically, the species is distinguished by having narrowly
obtuse, oblong-lanceolate leaves. The basal leaf cells are thin-walled and rec¬
tangular. The plants are cladautoecious, that is, the perigonia are at the ends
of separate branches instead of in the usual axillary position. No other North
American species in the Orthotrichaceae has an endostome of 16 segments and
stomata which are scarcely covered by the subsidiary cells.

The specimens examined from the sink area as well as those from the
Gaspe Peninsula have capsules which are almost exserted. These specimens are
identical with the European Orthotrichum pallens var. saxicola Burch. How¬
ever, after studying both European and North American specimens, we have
decided that this and other characters are variable and do not correlate with
each other. Thus the var. saxicola has been relegated to synonomy.

In western North America there exist two forms closely allied to typical
Ortho trichum pallens. These western plants are characterized by being smaller
and having shorter, oblong capsules with a very short seta. These forms have
been referred to variously— and apparently incorrectly— as O. pallens, O. pal¬
lens var. parvum Vent., and O. microcarpum De Not. This problem will be
dealt with in detail in a separate paper in preparation by the junior author.

Orthotrichum pallens, in a broad sense, is distributed in a discontinuous
pattern in boreal and alpine regions over much of the Northern Hemisphere.

Fig. 9. Distribution of Orthotri¬
chum pallens Bruch ex Brid. in
North America, based on speci¬
mens examined. (Goode Base
Map No. 102, copyright by the
University of Chicago, used by
permission.)

Orthotrichum polUnt

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THE MICHIGAN BOTANIST

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In North America the geographic range of this species is broken up into the
following disjunct areas: (1) scattered in the western Cordillera, (2) northern
Michigan including the Keweenaw Peninsula, (3) the Gaspe Peninsula in
Quebec, and (4) Cerro Potosi in northeastern Mexico. The latter is the
southernmost station for the species and represents the only report from
Mexico (Delgadillo & Vitt, 1970).

Fernald (1935) pointed out that the Keweenaw Peninsula as well as the
Gaspe Peninsula contains a well-marked Cordilleran element in the vascular
plant flora. Similarly, Steere (1937) found among the bryophytes of the
Keweenaw Peninsula a distinct Rocky Mountain element. He mentioned
among others Timmia austriaca Hedw., Frullania bolanderi Aust., and
Jungermannia schiffneri (Loitlesb.) Evs. The distribution of Ortho trichum pal-
lens parallels that of these species (see Fig. 9).

It should be noted that the entire sinkhole region was glaciated in Wis¬
consin times and the sinkholes under discussion here presumably originated
after glaciation. Therefore, the occurrence of O. pallens in these sinks seems
clearly to be a result of postglacial migration. If the Cordillera served as a
plant refuge during Pleistocene glaciation, then O. pallens is an example of a
species which has migrated eastward following disappearance of glacial ice.

ADDITIONAL SPECIES NEW TO THE SINKS

To the species listed by Robinson and Wells (1956), we can add the
following as being found at Bolton or Four Sinks. Specimens cited below are
deposited in the University of Michigan Herbarium (MICH), the University of
Michigan Biological Station Herbarium (UMBS), or the private collection of
Norton G. Miller. A total of 101 species of mosses and 24 species of hepatics
are now known from the entire sinkhole region.

Musci

Fissidens minutulus Sull. Four Sinks: On dry limy lithosol, limestone
ledge, third sink in from road, Miller 4602 (UMBS).

Ditrichum flexicaule (Schwaegr.) Hampe Bolton Sink: On moist side of
boulder near sinkhole floor, Miller 4595 (UMBS).

Seligeria campylopoda Kindb. ex Mac. & Kindb. Bolton Sink: On lime¬
stone boulder beneath Thuja, S side, Miller 4920 (with S. doniana ), 4921.

S. doniana (Sm.) C. Miill. Four Sinks: On moist shaded limestone ledge,
third sink in from road, Miller 4605 (UMBS), Vitt 892 (MICH).

Dicranum montanum Hedw. Bolton Sink: On rotting log beneath Thuja,
S side, Miller 4901 (with D. viride).

D. viride (Sull. & Lesq. ex Sull.) Lindb. Bolton Sink: On rotting log
beneath Thuja, S side. Miller 4901 (with D. montanum ), 4902.

Gymnostomum aeruginosum Sm. Bolton Sink: In crevice of dry lime¬
stone cliff face, S side of sinkhole near lip, Miller 4907; Four Sinks: On dry
limestone ledge, sinkhole wall, third sink in from road, Miller 4603 (UMBS).

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93

Didymodon rigidulus Hedw. Bolton Sink: In crevice of dry limestone
cliff face, S side of sinkhole near lip, Miller 4903 ; Four Sinks: On dry limy
lithosol over limestone ledge, third sink in from road, Miller 4601 (UMBS).

Bryum argenteum Hedw. Bolton Sink: On organic turf over exposed
limestone boulder on floor of sink, Miller 4593 (UMBS, with Tortella fragilis).

Rhodobryum roseum (Hedw.) Limpr. Bolton Sink: On rock and organic
debris, shaded slope in sinkhole, Miller 4600 (UMBS, with Pohlia cruda).

Mnium marginatum (With.) Brid. ex P. Beauv. Bolton Sink: On damp
humus beneath Thuja, just above floor of sink, Miller 4598.

M. spinulosum B.S.G. Bolton Sink: On side of boulder near bottom of
sink, Miller 4599 ; on organic debris over log, Thuja thicket, S side, Miller
4911.

Plagiopus oederiana (Sw.) Limpr. Bolton Sink: On ledge, dry cliff face,
S side of sink near lip, Miller 4917, Vitt 875 (MICH).

Orthotrichum obtusifolium Brid. Bolton Sink: On Thuja branch, S side,
Miller 4916 (with O. speciosum var. elegans).

O. speciosum var. elegans (Schwaegr. ex Hook. & Grev.) Warnst. Bolton
Sink: On Thuja branch with O. pallens, S side, Vitt 886 (MICH), Miller 4913,
4914; on Thuja branch, S side, Miller 4916 (with O. obtusifolium ); on Thuja
branches, sinkhole slopes, Vitt 880, 881 (MICH).

Thuidium recognitum (Hedw.) Lindb. Bolton Sink: On partially exposed
limestone rocks, S side, Miller 4922.

Campylium polygamum (B.S.G.) C. Jens. Bolton Sink: On humus, floor
of Thuja thicket, S side, Miller 4899.

Platydictya jungermannioides (Brid.) Crum Bolton Sink: On limestone
cliff, Vitt 878 (MICH).

Eurhynchium pulchellum var. praecox (Hedw.) Dix. Four Sinks: On dry,
disturbed soil bank, third sink in from road, Vitt 856 (MICH).

Plagiothecium roeseanum B.S.G. Bolton Sink: On humus, floor of
shaded Thuja thicket, S side, Miller 4918.

Heterophyllium haldanianum (Grev.) Kindb. Bolton Sink: On humus,
floor of Thuja thicket, S side, Miller 4905.

Diphysium foliosum (Hedw.) Mohr Four Sinks: On damp sand, shaded
crest of fourth sinkhole in from road, A/z7/er 4604 (UMBS), Vitt 893 (MICH).

Hepaticae

Cephalozia media Lindb. Bolton Sink: On moist decorticated log in
shade of Thuja trees just above floor of sinkhole, Miller 4586 (UMBS, with
Tritomaria exsectiformis and Dicranum flagellare); on rotting log beneath
Thuja, S side. Miller 4893 (UMBS, with Tritomaria exsectiformis) .—We have
examined specimens of Cephalozia catenulata collected by Robinson and Wells
(1956) at Bolton and Four Sinks and find them to represent C. media.

Geocalyx graveolens (Schrad.) Nees. Bolton Sink: On charred log be¬
neath Thuja near bottom of sink , Miller 4574 (UMBS).

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Lophozia incisa (Schrad.) Dumort. Bolton Sink: On rotting, shaded,
decorticated log near bottom of sinkhole, Miller 4577 (UMBS).

Leiocolea heterocolpos (Thed.) Buch Bolton Sink: On root of living
Thuja just above ice accumulation near bottom of sink, Miller 4576 (UMBS);
beneath Thuja on organic debris over rock, S sid q. Miller 4887.— Known from
a number of localities across the Upper Peninsula of Michigan but apparently
the only other station in this state south of the Straits of Mackinac is at the
tip of the thumb near Port Austin in Huron County (Nichols, 1933).

Tritomaria exsectiformis (Breidl.) Schiffn. Bolton Sink: On moist de¬
corticated log in shade of Thuja trees just above floor of sinkhole, Miller 4586
(UMBS, with Cephalozia media and Dicranum flagellare); on shaded decorti¬
cated log near bottom of sink, Miller 4587 ; on rock above ice accumulation
near bottom of sink, Miller 4588; on rotting log beneath Thuja, S side, Miller
4893 (with Cephalozia media).

Plagiochila asplenioides (L.) Dumort. Bolton Sink: On humus beneath
Thuja, S side, Miller 4888.

Scapania saxicola Schust. Bolton Sink: On moist side of shaded lime¬
stone boulder near bottom of sink, S side, Miller 4890 (with S. mucronata ),
4892— Known otherwise in Michigan only from the Pictured Rocks near
Munising in the Upper Peninsula (Crum & Miller, 1968).

Frullania eboracensis Gottsche Bolton Sink: On bark of living Thuja
twig, S side, Miller 4895.

Marchantia polymorpha L. Bolton Sink: Over Drepanocladus uncinatus
at mouth of ‘ice cave’ near bottom of sink, Miller 4591.— Miss Ella Campbell
has assigned this specimen to the var. aquatica Nees (see Campbell, 1969).

Conocephalum conicum (L.) Dumort. Bolton Sink: On thin organic turf,
cliff face, S side near lip of sink , Miller 4886.

REFERENCES

Campbell, Ella O. 1969. Marchantia polymorpha in northern Michigan. Mich. Bot. 8:
146-150.

Crum, H., & N. G. Miller. 1968. Bryophytes new to Michigan. Ibid. 7: 132-134.

Delgadillo, C., & D. H. Vitt. 1970. New moss records from Mexico. Madrono (In press).
Fernald, M. L. 1935. Critical plants of the upper Great Lakes region of Ontario and
Michigan. Rhodora 37: 197-222, 238-262, 272-301, 324-341.

Hagenah, D. J. 1955. Notes on Michigan pteridophytes, I. New county records in
Osmundaceae and Polypodiaceae. Am. Fern Jour. 45: 65-80.

Nichols, G. E. 1933. Notes on Michigan bryophytes. II. Bryologist 36: 69-78.

Robinson, H., & J. Wells. 1956. The bryophytes of certain limestone sinks in Alpena
County, Michigan. Ibid. 59: 12-17.

Steere, W. C. 1937. Critical bryophytes from the Keweenaw Peninsula, Michigan.
Rhodora 39: 1-14, 33-46.

Wagner, W. H., Jr., & Lois P. Lord. 1956. The morphological and cytological distinctness
of Botrychium minganense and B. lunaria in Michigan. Bull. Torrey Bot. Club 83:
261-280.

1970

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95

ALGAE IN PARK LAKE,

CLINTON COUNTY, MICHIGAN

Norman A. Andresen

Department of Biology, Taylor University,

Upland, Indiana

INTRODUCTION

Park Lake represents a rapidly aging, eutrophic lake in southern Michi¬
gan, where time and the disturbance of man are causing a profound and
destructive influence. As in many Michigan lakes, pollution of various kinds is
rapidly ruining Park Lake as a source of water and for recreation and home
sites. In the spring of 1966 Dr. G. W. Prescott suggested Park Lake to me as a
possible area for investigation, since a record of its present state would be
desirable for reference in future studies of the changing water quality.

Park Lake is of interest because it has been neglected biologically and
chemically even though it is very close to the Michigan State University
campus and phycology classes have made incidental collections there for many
years, especially during the spring. Knowledge of the biota and the lake chem¬
istry are pertinent both for the history of this lake and for comparison with
other lakes in southern Michigan. Furthermore, the lake is of interest because
the physiography and the hydrography together with pollution factors present
a unique combination of characters. The individuality of Park Lake results in
part from its extreme shallowness; high percentage of floating and emergent
vascular aquatic plants; and its pollution, organic and inorganic, by the resi¬
dents of the area.

THE LAKE

Park Lake is located in the southeast corner of Clinton County, Michi¬
gan (T 5 N, R 1 W, Sec. 28,29), six miles to the northeast of the campus of
Michigan State University. The general outline is approximately pyriform
(pear-shaped) with its long axis lying in a northeast-southwest direction. It is a
very shallow, productive lake of glacial origin. In 1931 it was described in the
Michigan Lakes and Streams Directory (East Lansing) as follows:

Park Lake, Clinton County; near M-78, south of Bath, near Lansing, 120 acres. A
road touches lake. Extensive resort development, boat livery, good swimming, sand
beach, woods on one side, flat land, lake much frequented. Large- and small-mouth
bass, bluegills, perch, pike.

The situation encountered by the author is quite different from that in the
above description. The community has deteriorated and the resort develop¬
ment is non-existent. There is little use of the lake for swimming and the
dirty sand beaches are cluttered with refuse and discarded items of every
variety. It was reported by Bratt (1957) that the Michigan Health Department
had closed the beaches in years past because of what was referred to as
chemical toilet pollution.

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Along the northwest and part of the west side of the lake there is a
swampy zone with marginal vegetation including a floating mat of Decodon
verticillatus bordered shoreward by a zone of trees. The land beyond this is
occupied by a muck farm. Toward the south end, the Decodon mat merges
with a Larix laricina bog mat and beyond this, toward the south, there is a
swampy lowland area. At the northeast end of the lake there is a public park
with a boat landing. The complete shoreline of the public park is overgrown
with Typha lati folia. Along the northern shore there is a fish bait shop and
boat livery near an abandoned ‘beach house’ which at one time apparently
offered some of the same services as the present establishment. Most of the
lake supports dense growths of partly emergent vascular aquatic plants so that
during the summer there is n6 large area of open water.

The residential community almost completely encircles the lake leaving
only the margin along the Larix bog mat and about one-half the Decodon mat
free from residential use.

The lake is eutrophic according to Thienemann’s classificaiton system as
given in Odum (1959). Bratt (1957) mentions Park Lake as a temporate lake
of the third order according to the Forel and Whippel classification system,
with a temporary thermocline being formed when wind and wave action are
limited. Bratt did not indicate at what time of year the thermocline occurs.
Observations bear out Bratt’s statement that the lake is an eutrophic hard-
water type with some marl precipitation.

The lake is surrounded by two major soil associations according to the
map in Veatch (1953), in the legend of which the land characters and geolog-
ic-pedologic relations are indicated as follows:

(1) The Fox-Plainfield-Hillsdale-Bellefontaine Association (sandy loams, plains
sands, or sand-gravel substratum):

Undulating plains, broken by swampy depressions; in part a complex of flats
of upland and swampy lowland irregular in shape and size. Relief 10 to 50 feet.
Textures mostly sandy loams, but various textures included. Substrata locally
gravelly, but also sands, silt and clay. Stone-free or not excessively bouldery. Acid
soils; medium fertility. . . . Till plains and outwash. Gray drift weathering to red¬
dish and yellow colors. Medium influence from limestone; variably weak to
strong influence from local sandstone and shale formations. Profiles on upland
mostly Gray -Brown Podzolic. Forest-on upland mostly oaks-hickory type.

(2) The Carlisle-Rifle Association (peats, mucks):

Mostly swamp but includes some marsh. A few low mounds and ridges of upland.
Mostly blackish muck on surface, but in part brown and yellow strongly acid peat.
Locally very thin deposits over variable substratum clay, sands, marl. . . . Post-
Glacial accumulations. Soil characteristics autogenous, but in local instances plant
matter highly decomposed. Strongly developed glei horizons; locally mineral sub¬
stratum strongly mottled with iron oxide; local bog iron ore pans. Forest-elm,
ash, soft maples, swamp white oak; in part tamarack.

Park Lake is, in my opinion, a highly productive lake. No direct studies
were conducted during my survey to evaluate the productiveness or to give
experimental data to classify the lake as a productive lake. The lake is classi¬
fied as a productive lake on the basis of the following inferential evidence:

1970

THE MICHIGAN BOTANIST

97

1) the observation of the lake and its abundance of vascular aquatic
plants;

2) the large number of algal taxa recorded for the lake;

3) the algal blooms recorded for the lake (1938 “very abundant”; 1967,
present study); (on all other occasions of water bloom reports, the
lake in which the bloom occurred was reported to be productive and
no unproductive lake has had a water bloom recorded for it.);

4) the high alkalinity with an abundance of bicarbonate which tends to
indicate production because of increased photosynthesis.

PREVIOUS SURVEY OF THE LAKE

In 1938 the Institute for Fisheries Research, Michigan Department of
Conservation, conducted a survey of the lake. This was made by a party
headed by Dr. Robert C. Ball and was mostly concerned with fish and the
ecological factors affecting their biology. A list of aquatic vascular plants was
compiled along with some morphometric observations resulting in the hydro-
graphic map from which the map below was redrawn (depths in 5-ft contours).

The hydrographic map of 1938 shows the area to be 185 acres with a
maximum depth of 25 feet. Bratt (1957) estimated the area to be 120 acres
with a maximum depth of twenty feet. Bratt also calculated the surface areas
over given depths from the 1938 map as follows:

58% of surface over a depth of less than five feet

33% of surface over a depth of between five and ten feet
9% of surface over a depth of ten feet or greater.

PARK LAKE

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THE MICHIGAN BOTANIST

Vol. 9

With 91% of the lake’s surface over a depth of ten feet or less, the shallow¬
ness of the lake is emphasized. Although no more recent data are available,
my observations confirm the general extensiveness of shallow water.

The 1938 survey party noted that the vegetation was dense in most
parts of the lake and that on two sides of the lake there were extensive
encroaching bogs. It also reported most of the bottom to be covered by a
pulpy peat of from four to twenty feet thick.

Bratt’s 1957 master’s thesis provides some limited field data which, as
Bratt stated, . . can be used for comparison in future investigations of Park
lake or other aquatic habitats . . as well as a list of the invertebrates found
during his investigation. Some of his data have been utilized in this work.

Since 1938 the Michigan Department of Conservation has maintained
records of the lake concerned mostly with the recreational use of the lake,
with specific reference to fishing. A winter fish-kill was recorded for the year
1939. An interesting letter dated September 10, 1943, from M. J. DeBoer to
R. S. Kocher dealt with the possibility of using the lake for testing chemical
controls on the aquatic plants and possible algicide testing. Apparently no
further consideration was given to the proposal.

Since the first records on Park Lake were kept by the Institute for
Fisheries Research in 1938, reports of pollution from chemical toilets have
been entered periodically. Bratt (1957) was the most recent to record infor¬
mation concerning this pollution. During my collecting trips physical evidence
of pollution was clear. The shorelines were used as a dumping ground for all
types of refuse. Residential units had direct drains into the lake, as observed
during the summer of 1967. Waste water from a household sink was observed
flowing into the lake proper. Outdoor sanitary units were located within
twenty feet of the lake edge where the shoreline was composed mainly of
sand. The garbage dump for the community is located near the edge of the
lake at the south end.

METHODS

For the purpose of the algal survey, permanent and incidental collecting
stations were established. The three permanent stations were chosen for their
biotic differences and to record the change in the flora at these stations
throughout the survey. The incidental stations were mainly samples collected
at random along the shoreline and as far out into the lake as the investigator
could wade. When a boat was available, samples were collected from the lake
area inaccessible from the shore.

The collecting period began on 13 April 1966 and was terminated on 22
November 1967, with occasional collections made up to 5 June 1968. The
occasional samples were made for the purpose of augmenting the qualitative
samples. The collections were made weekly during June through October;
biweekly during November, December, March through May; and monthly in
January and February.

During the period of study 197 samples were collected. The samples
were taken from floating masses, both filamentous and non-filamentous; scrap-

1970

THE MICHIGAN BOTANIST

99

ings of any non-living object in the water having a film or growth appearing
on its surface; scrapings of submerged parts of aquatic vascular plants; squeez¬
ings of bottom debris and vascular aquatic plants; the first centimeter of
bottom substrate; growths on animal life of the lake; masses chopped out of
the ice cover; and plankton samples.

The samples were examined in the living condition and preserved in
6:3:1 solution (water— 95% alcohol— formaldehyde) or FAA solution (formalde¬
hyde-acetic acid— alcohol) (50% preservative, 50% sample) for subsequent study.

WATER BLOOM

A water bloom occurred in the lake for a period of approximately ten
weeks (July 9-Sept. 16) during the summer of 1967. The major components
of the bloom were Microcystis aeruginosa , M. flos-aquae, Anabaena flos-aquae,
Coelosphaerium naegelianum, and C. kuetzingianum.

Jackson (1963) reported that on several occasions during his investiga¬
tion of Lake Lansing, water blooms were observed. The blooms in the fall of
1959 and 1962 in Lake Lansing turned the surface waters of the shallow
littoral region a pea-soup color and were composed primarily of Microcystis
aeruginosa, M. flos-aquae, Coelosphaerium naegelianum, and Anabaena sp.

Jackson reported other blooms occurring with the major components
being Ceratium hirundinella on one occasion and Tolypothrix tenuis on
another. Park Lake had an abundance of C. hirundinella in the spring of 1967
but it was never in bloom proportions. Tolypothrix tenuis was recorded for
Park Lake but it was never in abundance.

Wade (1949) conducted a study on Jordan Lake in southern Michigan
that supported a water bloom during the summer months. The major com¬
ponents of that bloom flora were Anabaena limnetica, A. spiroides var. crassa,
A. circinalis var. macrospora, Aphanizomenon flos-aquae, Microcystis
aeruginosa, and Coelosphaerium naegelianum.

Edelstein (1966), in her paper on the phytoplankton of Spring Lake,
Michigan, mentioned that this lake in 1957 also supported a water bloom. The
major components of that bloom were Aphanizomenon flos-aquae, Anabaena
spiroides var. crassa, A. circinalis, Microcystis aeruginosa, and Aphanocapsa
pulchra.

Lakes Lansing, Jordan, Spring, and Park are all located in southern
Michigan and have been recorded to support water blooms. The major con¬
stituents of the blooms have been the Cyanophyta and some of the same
organisms appear in the blooms recorded for the different lakes. The Cyano¬
phyta are the rrajor elements of the blooms that are the producers of ob¬
noxious conditions while the Chry sophy ta and the Pyrrophyta, though noted
for their bloom ability, are not the most obnoxious water bloomers. It should
be noted that water blooms can occur in most highly productive lakes when
the physical and chemical elements are in the proper balance and proportions.
Odum (1959) states that blooms are characteristic of eutrophic lakes. Water
blooms are of interest, among other aspects, because of their economic im¬
portance. Blooms cause economic loss to a recreational area by their objec-
tional odors and by ruining the water for fishing and swimming.

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SYSTEMATIC LIST OF THE ALGAE IN PARK LAKE

The names and sequence of taxa in the following systematic list are
chiefly those of Prescott (1962), which also served as the basic reference for
identification. New records for North America1 are marked with +; new rec¬
ords for Michigan are marked with *; all are illustrated in figures 1-26.

CYANOPHYTA

Aphanocapsa elachista West & West
A. pulchra (Kiitz.) Rabenhorst
Aphanothece gelatinosa (Henn.) Lemmer-
mann

A. stagnina (Spreng.) A. Braun
Chroococcus dispersus (Keissl.) Lemmer-
mann

C. giganteus W. West
C. limneticus Lemmermann
C. minimus (Keissl.) Lemmermann
C. minor (Kiitz.) Nageli
C. minutus (Kiitz.) Nageli
C. prescottii Drouet & Dailey in Drouet
C. turgidus (Kiitz.) Nageli
C. varius A. Braun in Rabenhorst
Coelosphaerium dubium Grunow in Raben¬
horst

C. kuetzingianum Nageli
C. naegelianum Unger

Dactylococcopsis fascicularis Lemmermann
Gloeothece rupestris (Lyngb.) Bornet in
Wittrock & Nordstedt
Gomphosphaeria aponina Kiitzing
G. aponina var. delicatula Virieux
G. aponina var. multiplex Nygaard
G. lacustris var. compacta Lemmermann
Merismopedia convoluta de Brdbisson in
Kiitzing

M. glauca (Ehrenb.) Nageli
M. punctata Meyen
M. tenuissima Lemmermann
Microcystis aeruginosa Kiitz. emend. Elenkin
M. flos-aquae (Wittrock) Kirchner
M. incerta Lemmermann
Chamaesiphon incrustans Grunow in Raben¬
horst

Pleurocapsa minor Hansg. emend. Geitler
Arthrospira jenneri (Kiitz.) Stizenberger
*Borzia triocularis Cohn (Fig. 3)

Lyngbya aestuarii (Mert.) Liebermann
L. birgei G. M. Smith
L. diguetii Gomont in Hariot
L. epiphytica Hieronymus in Engler &

Prantl

L. hieronymi Lemmermann

L. major Meneghini

L. spirulinoides Gomont

L. taylorii Drouet & Strickland

L. versicolor (Wartmann) Gomont

Oscillator ia agardhii Gomont

O. articulata Gardner

O. bornetii Zukal

O. chalybea Mertens in Jurgens

O. curviceps C. A. Agardh

O. limnetica Lemmermann

O. limosa (Roth) C. A. Agardh

O. nigra Vaucher

O. ornata Kiitzing

O. princeps Vaucher

O. rubescens DeCandolle

O. subbrevis Schmidle

O. tenuis C. A. Agardh

Phormidium tenue (Menegh.) Gomont

Schizothrix tinctoria Gomont

Spirulina princeps (West & West) G. S. West

S. subsalsa Oersted
Trichodesmium lacustre Klebahn
Anabaena flos-aquae (Lyngb.) deBrdbisson
A. oscillarioides Bory

Nostoc comminutum Kiitzing
Microchaete goeppertiana Kirchner in Engler
& Prantl

+Scytonema bohneri Schmidle (Figs. 1,2)
Tolypothrix lanata Wartmann in Rabenhorst

T. tenuis Kiitz. emend. J. Schmidt
Calothrix atricha Frdmy

C. breviarticulata West & West
C. epiphytica West & West
C. stellaris Bornet & Flahault
Gloeotrichia echinulata (J. E. Smith) P.
Richter

G. pisum (C. A. Ag.) Thuret

CHLOROPHYTA

Chlamydomonas globosa Snow
C. polypyrenoideum Prescott
C. pseudopertyi Pascher
Eudorina elegans Ehrenberg
Gonium pectorale Miiller
Pandorina morum (Miill.) Bory
Ploeodorina calif or nica Shaw

1 Scenedesmus armatus var. boglariensis f. semicostatus and S. tibiscensis were previ¬
ously recorded for North America in an unpublished thesis by Ronald W. Hoham (1966).

1970

THE MICHIGAN BOTANIST

101

Volvox aureus Ehrenberg
V. tertius A. Meyer

Aster ococcus superbus (Cienk.) Scherffel
Gloeocystis ampla (Kiitz.) Lagerheim
G. gigas (Kiitz.) Lagerheim
G. planctonica (West & West) Lemmermann
Palmella mucosa Kiitzing
Sphaerocystis schroeteri Chodat
Apiocystis brauniana Nageli
Tetraspora gelatinosa (Vauch.) Desvaux
Dispora crucigenioides Printz
Elakatothrix viridis (Snow) Printz
Binuclearia tatrana Wittrock
Geminella interrupta (Turp.) Lagerheim
G. minor (Nag.) Heering
Hormidium klebsii G. M. Smith
Radiofilum flavescens G. S. West
Ulothrix tenerrima Kiitzing
Uronema elongatum Hodgetts
Microspora elegans Hansgirg

M. loefgrenii (Nordst.) Lagerheim
M. pachyderma (Wille) Lagerheim
M. stagnorum (Kiitz.) Lagerheim
Cylindrocapsa conferta W. West
Aphanochaete polychaete (Hansg.) Fritsch

A. repans A. Braun
Chaetophora attenuata Hazen
C. incrassata (Huds.) Hazen
Pseudulvella americana (Snow) Wille
Stigeoclonium lubricum (Dillw.) Kiitzing
S. subsecundum Kiitzing

S. tenue (C. A. Ag.) Kiitzing
Protococcus viridis C. A. Agardh
Chaetosphaeridium ovalis G. M. Smith
Coleochaete irregularis Pringsheim
C. scutata de Brdbisson
Gongrosira debaryana Rabenhorst
Basicladia chelonum (Collins) Hoffman &
Tilden

B. crassa Hoffman & Tilden
Cladophora crispata (Roth) Kiitzing

C. insignis (C. A. Ag.) Kiitzing
Rhizoclonium crassipellitum West & West
R. hieroglyphicum (C. A. Ag.) Kiitzing

*Oedogonium capitellatum Wittrock (Fig. 5)
O. howardii G. S. West
O. poecilosporum Nordstedt & Hirn in Hirn
O. pratense Transeau

O. spirostriatum Tiffany
Botryococcus braunii Kiitzing

B. sudeticus Lemmermann
Characium ambiguum Hermann

C. dabary anum (Reinsch) DeToni
C. falcatum Schroeder

Pediastrum araneosum (Racib.) G. M. Smith

P. boryanum (Turp.) Meneghini

P. boryanum var. longicorne Raciborski

P. duplex var. clathratum (A. Braun)
Lagerheim

P. duplex var. rugulosum Raciborski
P. integrum Nageli
P. obtusum Lucke
P. tetras (Ehrenb.) Ralfs
P. tetras var. tetraodon (Corda) Rabenhorst
Sorastrum spinulosum Nageli
Coelastrum microporum Nageli in A. Braun
Ankistrodesmus falcatus (Corda) Ralfs
A. falcatus var. tumidus (West & West) G. S.
West

Chlorella vulgaris Beyerinck
Closteriopsis longissima var. tropica West &
West

Gloeotaenium loitelsbergerianum Hansgirg
Nephrocytium agardhianum Nageli

N. limneticum (G. M. Smith) G. M. Smith
Oocystis borgei Snow

O. elliptica W. West
O. gigas Archer

O. parva West & West

O. solitaria Wittrock in Wittrock & Nordstedt
Planktosphaeria gelatinosa G. M. Smith
Quadrigula lacustris (Chod.) G. M. Smith
Rayssiella hemisphaerica Edelstein & Prescott
Tetraedron minimum (A. Braun) Hansgirg
T. muticum (A. Braun) Hansgirg
Trochiscia reticularis (Reinsch) Hansgirg
Zoochlorella parasitica Brandt
Crucigenia rectangularis (A. Braun) Gay
*Pectodictyon cubicum Taft (Fig. 26)
Scenedesmus acutus Meyen
+S. acutus f. alternans Hortob. (Fig. 8)

+S. acutus f. tetradesmiformis (Wolosz.)
Uherkovich (Fig. 9)

S. arcuatus Lemmermann
S. arcuatus var. platydiscus G. M. Smith
*S. armatus var. boglariensis f. semicostatus
Hortob. (Fig. 11)

+S. bicaudatus (Hansg.) Chodat (Fig. 6)

S. brevispinus (G. M. Smith) Chodat
S. ecornis (Ralfs) Chodat
S. ecornis var. discidormis Chodat
S. longus var. naegelii de Brdb.) G. M. Smith
S. ovalternus Chodat
S. quadricauda (Turp.) de Brdbisson in de
Brdbisson & Gody

S. quadricauda var. longispinus (Chod.) G.M.
Smith

S. quadricauda var. maximus West & West

S. quadricauda var. quadrispinus (Chod.) G.

M. Smith

S. spinosus Chodat
*S. tibiscensis Uherkovich (Fig. 7)

Tetradesmus wisconsinensis G. M. Smith
Sirogonium sticticum (Engler) Kiitzing

102

THE MICHIGAN BOTANIST

Vol. 9

Explanation of Figures 1-25 (taxa previously unreported from Michigan).

I. & 2. Scytonema bohneri; plant occurring in small tufts on substrate; trichome branches
frequently exhibiting both Scytonema- type branches and Tolypothrix- type branches;
sheath thin and not lamellated, cells appearing quadrate; veg. cell diam. 8-10m-

3. Borzia trilocularis ; trichome of three to five barrel-shaped cells, terminal cells hemi¬
spherical; trichome diam. 14m, L. 9p.

4. Cosmarium garrolense var. dimido-minus ; variety distinguished from typical by its
smaller size; diam. 14;u, L. 23m, Ism. 3m-

5. Oedogonium capitellatum; oospore spore wall smooth, operculum median, macran-
drous and monoecious; veg. cell diam. 7m, L. 31m; oogonia diam. 24m, L. 22m; oospore
diam. 15m, L. 20m.

6. Scenedesmus bicaudatus', oval cells tightly appressed with the terminal cells of the
coenobium each bearing one spine on opposite poles; diam. 5-6j u, L. 13-15/u, spine L.
10/x.

7. Scenedesmus tibiscensis; coenobium of alternating angular cells; diam. 3m, L. 12-14m.

8. Scenedesmus acutus f. alternans; alternating coenobium, inner cells angular-clavate,
terminal cells lunate with swelling on inner curve; diam. 5m, L. 17.5m.

9. Scenedesmus acutus f. tetradesmiformis', coenobium composed of fusiform cells; diam.
7m, L. 25m.

10. Cosmarium conspersum f. minus', variety distinguished from typical by smaller size;
diam. 42m, L. 54m, Ism. 18m.

II. Scenedesmus armatus var. boglariensis f. semicostatus ; terminal cells of coenobium
bearing a long spine at each pole; median cells having a prominent longitudinal median
rib; diam. 4-4.5m, L. 16m.

12. Cosmarium minutissimum', diminutive size characteristic; diam. 13m, L. 15m, Ism. 2m-

13. Cosmarium humile var. danicum\ apical granules and smaller size distinguish it from
typical; diam. 15m, L. 17m, Ism. 6m-

14. Cosmarium humile var. glabrum; lack of apical granules characteristic, diam. 13.5m,
L. 15m, Ism. 4m.

15. Cosmarium regnellii ; concave apex, open notch at side of apex and lateral depression
characteristic; diam. 14m, L. 17m, Ism. 3p.

16. Euastrum turneri var. strictum\ deep apical notch and the smoothly undulating
margin with projecting spines and granules are characteristic; diam. 27m, L. 41m, Ism. 6p.

17. Staurastrum disputatum f. minus', size characteristic; diam. 13m, L. 20m, Ism. 9p.

18. Cosmarium depressum\ subspecific taxa previously reported for Michigan, first record
of typical; wall is punctate, oval in apical view; diam. 52 m, L. 54m, Ism. 14m-

19. Staurastrum cosmarioides; large size, punctate wall, and triradiate arrangement are
characteristic; diam. 60m, L. 95m, Ism. 30m.

20. Euglena fusca; subcylindrical form, size, and pellicle (smooth in this specimen) are
characteristic; diam. 20m, L. 155m-

21. Staurastrum cyclacanthum var. subacanthum; considered to be the var. subacanthum
on the basis of size and thickness of the central portion of the cell when viewed apically;
diam. 45m L- 40m, Ism. 10m.

22. Phacus pseudonordstedtir, the two large paramylon bodies at the lateral margins, the
straight caudus, and the left to right spiral striations in the pellicle are characteristic;
diam. 15m, L. 33m.

23. Phacus obolus; size, the large central paramylon body, and the straight to slightly
bent caudus are characteristic; diam. 15m, L. 30m.

24. Cosmarium sportella', granular cell wall with central roset of granules are character¬
istic; oval in apical view; diam. 50m, L. 58m, Ism. 17m.

25. Staurastrum punctulatum var. minus', size characteristic; diam. 18m, L. 20m, Ism. 8m-

1970

THE MICHIGAN BOTANIST

103

104

THE MICHIGAN BOTANIST

Vol. 9

160p

Fig. 26. Pectodictyon cubicum (after
Taft, 1945); the unusual coenobium is
very characteristic; veg. cell diam. 7ju-

Spirogyra crassa Kiitzing
S. longata (Vauch.) Kiitzing
Zygnema sterile Transeau in Transeau,
Tiffany, Taft, & Li
Closterium dianae Ehrenberg

C. ehrenbergii Meneghini
C. jenneri Ralfs
C. leiblenii Kiitzing
C. libellula Focke
C. monoliferum (Bory) Ehrenberg
C. parvulum Nageli
C. venus var. westii (West) Krieger
Cosmarium angulare Johnson
C. angulosum de Brdbisson
C. botrytis Meneghini
*C. conspersum f. minus Borge (Fig. 10)

C. denticulatum var. borgei Irende-Marie
*C. depressum (Nag.) Lundell (Fig. 18)

C. depressum var. schroederi Krieger &
Gerloff

+C. garrolense var. dimido-minus (Groblad)
Krieger & Gerloff (Fig. 4)

C. humile (Gay) Nordstedt in DeToni
*C. humile var. danicum (Borge) Schmidle
(Fig. 13)

*C. humile var. glabrum Gutwinski (Fig. 14)
*C. minutissimum Archer (Fig. 12)

C. obtusatum Schmidle
C. portianum Archer
C. pscudopyramidatum Lundell
*C. regnelli Wille (Fig. 15)

C. reniforme (Ralfs) Archer
C. sexangulare f. minimum Nordstedt
*C. sport ella de Brdbisson (Fig. 24)

C. subcostatum Nordstedt
C. subtumidum Nordstedt
C. trilobulatum Reinsch
C. tumidum Lund
C. turpinii de Brdbisson

C. undulatum var. minutum Wittrock
Desmidium aptogonum de Brdbisson

D. baileyi (Ralfs) Nordstedt

Euastrum hypochondrum f. decoratum Scott
& Prescott

*E. turneri var. strictum (Borges.) Scott &
Prescott (Fig. 16)

E. verrucosum Ehrenberg

Hyalotheca dissiliens (Smith) de Brdbisson
H. mucosa (Dillw.) Ehrenberg
Micrasterias truncata var. semiradiata (Nag.)
Cleve

Penium margaritaceum (Ehrenb.) de
Brdbisson in Ralfs

Pleurotaenium ehrenbergii (de Brdb.) DeBary
P. trabecula (Ehrenb.) Nageli
Staurastrum cornutum Archer
*S. cosmarioides Nordstedt (Fig. 19)

S. crenulatum (Nag.) Delpont
+S. cyclacanthum var. subacanthum
Gronblad (Fig. 21)

+S. disputatum f. minus West & West
(Fig. 17)

S. furcigerum de Brdbisson
S. gracile Ralfs var. nanum Wille
S. hexacerum (Ehrenb.) Wittrock
S. johnsonii West & West
S. longiradiatum West & West
S. manfeldtii Delpont
+S. punctulatum var. minus West & West
(Fig. 25)

S. sebaldi Reinsch var. ornatum Nordstedt
S. vestitum Ralfs

Xanthidium antilopaeum (de Brdb.) Kiitzing
Chara globularis Thill.

C. vulgaris Linnaeus

EUGLENOPHYTA
Euglena elastica Prescott

1970

THE MICHIGAN BOTANIST

105

*E. fusca (Klebs) Lemmermann (Fig. 20)

E. gracilis Klebs

E. proximo Dangeard

Lepocinclis acuta Prescott in Prescott,

Silva, & Wade
Phacus caudatus Hiibner
P. contortus var. complicatus Bourrelly
P. curvicauda Swirenko
P. longi cauda (Ehrenb.) Dujardin
P. longicauda var. insectus Kozw.

+P. o bolus Pochmann (Fig. 23)

P. orbicularis Hiibner
+P. pseudonordstedtii Pochmann (Fig. 22)

P. pseudo swirenkoi Prescott
P. suecicus Lemmermann in Pascher &
Lemmermann

P. undulatus (Skv.) Pochmann
Trachelomonas hispida (Perty) Stein
T. volvocina Ehrenberg

PYRROPHYTA

Glenodinium pulvisculus (Ehrenb.) Stein
Peridinium gatunense Nygaard in Ostenfeld
& Nygaard

P. inconspicuum Lemmermann
P. pusillum (Penard)Lemmermann
P. umbonatum Stein
P. volzii Lemmermann
P. willei Huitfeld-Kaas
Ceratium hirundinella (O. F. Mull.)
Dujardin

Cystodinium cornifax (Schill.) Klebs

CHRYSOPHYTA

Ophiocytium cochleare (Eichw.) A. Braun
O. parvulum (Perty) A. Braun
Synura uvella Ehrenberg
Dinobryon cylindricum Imhof ex Ahlstrom
D. sertularia Ehrenberg
D. tabellariae (Lemm.) Pascher in Pascher
& Lemmermann

Uroglenopsis americana (Calkins) Lemmer¬
mann

Cyclotella sp.

Melosira sp.

Tabellaria sp.

Asterionella sp.

Fragilaria spp.

Synedra spp.

Synedra delicatissima W. Smith (Det. Begres)
Eunotia sp.

Cocconeis sp.

Amphipleura sp.

Gyro sigma sp.

Navicula spp.

Neidium sp.

Pinnularia spp.

Stauroneis sp.

Gomphonema spp.

Amphora spp.

Cymbella spp.

Epithemia sp.

Rhopalodia sp.

Nitzschia spp.

Cymatopleura sp.

Surirella sp.

Because of the problems involved in diatom taxonomy, the Bacillario-
phyceae were mostly not identified to species, although those genera are indi¬
cated which were represented by more than one species. In addition, spec¬
imens referable to the following genera were collected but were either im¬
mature or lacked the necessary features for specific identification and hence it
could not be determined whether they represented taxa otherwise unknown
from the lake: Stigeoclonium, Gongrosira, Pithophora, Bulbochaete, Oedogoni-
um, Mougeotia, Spirogyra, Zygnema, Euglena, Lepocinclis, Phacus, and
Vaucheria.

SUMMARY AND CONCLUSIONS

Park Lake is a shallow, eutrophic lake of glacial origin, located in the
southeast corner of Clinton County, six miles from the campus of Michigan
State University. In recent years, the eutrophication process has been en¬
hanced by increased pollution. It is a rapidly aging lake with a high density of
vascular aquatic plants and an abundant algal flora.

The algae can be assigned to the following divisions from the collections

made:

106

THE MICHIGAN BOTANIST

Vol. 9

DIVISION

GENERA

TOTAL TAXA

% OF TOTAL

Cyanophyta

26

74

22.1

Chlorophyta

71

200

59.7

Euglenophyta

4

21

6.3

Pyrrophyta

4

9

2.7

Chrysophyta

27

31

9.2

Of the 335 taxa recorded from the 197 samples collected, 25 are newly
recorded for Michigan, of which 10 are new for North America.

The flora is co-dominated in number of species by the Chlorophyta and
the Cyanophyta. The Euglenophyta, Pyrrophyta, and Chrysophyta comprise
only 18.2% of the total algal flora of the lake. The importance of the Chryso¬
phyta must not be disregarded or minimized by the above percentage as they
were not all identified to species, the Bacillariophyceae (diatoms) being identi¬
fied to the generic level only.

The littoral community includes more than twice the number of taxa as
the limnetic community.

The lake supported a continuous water bloom for a period of approxi¬
mately ten weeks during the summer of 1967. The bloom flora of Park Lake
was similar to that of Lake Lansing, but differed from the bloom flora in
Jordan Lake and Spring Lake in respect to their dominants.

ACKNOWLEDGMENTS

This paper is revised from a thesis submitted to Michigan State University in
partial fulfillment of the requirements for the degree of Master of Science. I wish to
express my sincere appreciation to my major professor, Dr. G. W. Prescott, for his
guidance and encouragement during this study, Drs. R. C. Ball, E. S. Beneke, J. C. Elliott,
I. W. Knobloch, and W. E. Wade, members of my committee, for their helpfulness and
critical advice. I also wish to thank Garrett E. Crow, who generously offered his help at
many stages of this work, and Darryl D. Fritze who assisted in the typing of the manu¬
script. Special thanks must be given to Forrest M. Begres, who helped in and confirmed
the author’s identification of diatoms.

REFERENCES

Bratt, A. D. 1957. A Quantitative Study of the Invertebrate Population of Park Lake,
Clinton County, Michigan. M. S. thesis, Mich. State Univ.

Edelstein, T. 1966. The phytoplankton of Spring Lake, Michigan. Hydrobiologia 27:
137-145.

Edelstein, T., & G. W. Prescott. 1964. Rayssiella, a new genus of Oocystaceae (Chloro¬
phyta) from Spring Lake, Michigan. Phycologia 4: 121-125.

Frdmy, P. 1930, Les Myxophycdes de l’Afrique dquatoriale franqaise. Thesis. Caen. 507

pp.

Hoham, R. W. 1966. The Fresh Water Algae of Mission Well and Tykeson Pond, Mon¬
tana. M. S. thesis, Mich. State Univ.

Irdnde-Marie, Fr. 1939. Flore Desmidiale de la Rdgion Montreal. Laprairie, Canada. 547

pp.

Jackson, D. C. 1963. A Taxonomic and Limnological Study of the Algae in Lake
Lansing, M. S. thesis, Mich. State Univ.

Krieger, W. 1933-1937; Die Desmidiaceen in Rabenhorst, Kryptogamen-Flora 13, Abt. 1.
712 pp. + 96 pi.

Odum, E. 1959. Fundamentals of Ecology. Saunders, Philadelphia. 546 pp.

1970

THE MICHIGAN BOTANIST

107

Patrick, Ruth, & C. W. Reimer. 1966. The Diatoms of the United States. Vol. 1. Acad.
Nat. Sci. Phila. Monogr. 13. 688 pp.

Pochmann, A. 1942. Synopsis der Gattung Phacus. Arch. Protistenk. 95: 81-252.

Prescott, G. W. 1962. Algae of the Western Great Lakes Area. rev. ed. Brown, Dubuque.
977 pp.

Prescott, G. W. 1964. How to Know the Fresh-Water Algae, rev. ed. Brown, Dubuque. 272

pp.

Prescott, G. W., & A. M. Scott. 1945. The fresh-water algae of southern United States III.
The desmid genus Euastrum, with descriptions of some new varieties. Am. Midi. Nat.
34: 231-257.

Scott, A. M., & G. W. Prescott. 1952. The algal flora of southeastern United States VI.
Additions to our knowledge of the desmid genus Euastrum. 2. Hydrobiologia 4:
377-398.

Smith, G. M. 1916. A monograph of the algal genus Scenedesmus based upon pure
culture studies. Trans. Wis. Acad. 18: 422-530.

Smith, G. M. 1924. Phytoplankton of the inland lakes of Wisconsin, Part II. Bull. Wis.
Geol. Nat. Hist. Surv. 57(2): 1-227.

Smith, G. M. 1950. The Fresh-Water Algae of the United States, ed. 2. McGraw-Hill, New
York. 719 pp.

Taft, C. E. 1945. Pectodictyon, a new genus in the family Scenedesmaceae. Trans. Am.
Micr. Soc. 64: 25-28.

Tiffany, L. H. 1937. Oedogoniales. N. Am. FI. 11: 1-102.

Uherkovich, G. 1966. Die Scenedesmus-Arten Ungarns. Akademiai Kiado, Budapest. 173

pp.

Veatch, J. O. 1953. Soils and Land of Michigan. Mich. State Coll. Press, East Lansing.
241 pp. + map.

Wade, W. E. 1949. Some notes on the algal ecology of a Michigan lake. Hydrobiologia 2:
109-117.

Publications of Interest

A PHOTO GUIDE TO THE PATTERNS OF DISCOLORATION AND DECAY IN
LIVING NORTHERN HARDWOOD TREES. By Alex L. Shigo & Edwin vH. Larsen.
Northeastern Forest Experiment Station, Upper Darby, Pennsylvania (U.S.D.A. Forest
Service Research Paper NE-127), 1969. 100 pp. One hundred color photos illustrate
how discoloration and decay form in northern hardwood trees and show how interior
defects can be estimated from external indications on the living tree. Damage from
fungi and animals are both included.

DEFICIENCY SYMPTOMS OF SOME FOREST TREES. By John Hacskaylo, R. F. Finn,
& J. P. Vimmerstedt. Ohio Agricultural Research and Development Center, Wooster,
Ohio (Research Bulletin 1015), 1969. 68 pp. Another bulletin copiously illustrated
with color photos showing results of various nutrient deficiencies on the color and
development of leaves and roots of black walnut, black locust, cottonwood, sweet-
gum, and Scots pine.

108

THE MICHIGAN BOTANIST

Vol. 9

MORPHOLOGY OF THE FUNGAL ASSOCIATION IN
THREE SPECIES OF CORALLORHIZA IN MICHIGAN

Ella 0. Campbell

Department of Botany and Zoology, Massey University,

Palmerston North, New Zealand

In the summer of 1968 I investigated the fungal associations of three
species of Corallorhiza which grow in northern Michigan, namely C. trifida
Chatelain, C. maculata Raf., and C. striata Lindl.

Material of the orchid rhizomes and adjacent tree roots was collected in
July and August. After the material had been washed in water, promising
pieces were killed and fixed in formalin-acetic-alcohol, infiltrated with paraf¬
fin, and sectioned by microtome. In many cases fine tree-roots were lying
close to the orchid rhizomes and held attached in this position by fungal
hyphae or by rhizomorphs. In serial sections cut at 10 n it was possible to
follow hyphae from the roots into the orchid rhizomes. Staining of the sec¬
tions with safranin and fast green gave satisfactory contrast.

Corollorhiza trifida

C. trifida is found in Europe and Asia as well as in North America. Its
mycorrhiza has been studied previously. Jennings and Hanna (1898) using the
name C. innata R.Br. for their specimens, which came from the eastern
European Alps, mention that the plants are not entirely colorless, for, al¬
though the reduced leaves lack chlorophyll, the flowers and the upper part of
the scape may be pale yellow-green. They draw attention to the tufts of
mykokleptic hairs, which Thomas (1893) had earlier observed projecting from
papillae on the rhizome, and describe the passage of septate hyphae through
the hairs and the outer layers of the rhizome to a zone of cells occupied by
fungal coils and eventually to a deeper zone where collapse of the coils oc¬
curs. In one instance they found sporophores of Clitocybe infundibuliformis
Sch. growing from mycelium around the rhizome, but on other occasions the
sporophores found in close proximity to the orchid belonged to other Basidio-
mycetes.

In northern Michigan the plants of C. trifida show slight green colora¬
tion in the flowering stem, the basal leaf, and the ovary. On the surface of the
coralloid rhizomes are numerous papillae-bearing hairs (Fig. 1). Also present
on the rhizomes are fungal hyphae which can be followed either as single
threads or as rhizomorphs to the fine roots of Thuja occidentalis L. or of
Picea spp. In some cases the fungal threads are yellowish in color, in other
cases white.

The yellowish fungus corresponds with Mycena thujina Smith. In living
Thuja roots it is present in the cortical cells and in dead' roots it is in the
vascular cylinder as well. It enters the orchid by way of the hairs (Fig. 1),
spreads in a cylinder around the rhizome at a depth of 3 to 6 cells from the
surface, and then enters deeper cells where the fungal coils, which form at

1970

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Fig. 1. Part of the rhizome of C. trifida in transverse section, x 185. d. zone of fungal
digestion; h. hair; p. papilla.

Fig. 2. Part of the rhizome of C. maculata in transverse section, x 185.

110

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first, eventually collapse. The hyphae are septate with clamp connections and
are of diameter 3/i. In the rhizomorphs the central hyphae lie parallel longi¬
tudinally, while interwoven peripheral hyphae lie horizontally in a cylinder.

The white fungus occurs either as single hyphae or as rhizomorphs, up
to 0.16 mm in diameter, in which the hyphae lie somewhat interwoven in a
lengthwise direction. Most of the hyphae of the rhizomorph are of diameter 3
to 6jU and show clamp connections, but 6 to 8 central hyphae are much
wider, with a diameter up to 47/Lt. The outermost hyphae tend to have thick¬
ened walls and to lack clamp connections. Sometimes the rhizomorphs lie in
the axils of the scale leaves on the orchid rhizomes. They can be followed to
adjacent roots, where, as hyphae, they enter the cells of the cortex and in
dead roots the vascular cylinder also. Hyphae departing from the rhizomorphs
infect the orchid plants through hairs on the papillae and appear within the
rhizomes as does the yellowish fungus.

Corallorhiza maculata

C. maculata has no chlorophyll in any part of the plant. It is endemic to
North America. The 20 plants selected for investigation were from flourishing
colonies, as indicated by the number of new flowering spikes present and of
old fruiting spikes of the previous year. They were collected from a variety of
situations, as from beech-oak forest under trees of Fagus grandifolia Ehrh. and
Quercus rubra L., from beech -maple forest under Fagus grandifolia and Acer
saccharum Marsh., from oak -maple forest under Quercus rubra and Acer
saccharum , and elsewhere under pure stands of Betula papyrifera Marsh., of
Thuja occidentalism of Pinus strobus L., of Pinus resinosa Ait., and of Populus
grandidentata Michx.

The underground portion of the orchid consists of pale, coralloid, fleshy
rhizomes, up to 4 mm in diameter, lying horizontally in the litter layer
amongst decaying twigs, fallen leaves, and the roots of trees. The orchid itself
has no roots. On the surface of the rhizome, besides ring-like depressions at
the nodes and the tattered remnants of vestigial scale leaves, there are num¬
erous conical or hemispherical papillae bearing projecting hairs or appearing as
brown lesions after the hairs have withered off. Against the rhizomes lie the
fine rootlets of trees, many of which are already rotted internally, leaving
only an outer shell to maintain the form of the rootlet. Associated with 17 of
the specimens that were examined, rhizomorphs of Armillaria mellea (Vahl. ex
Fr.) Kummer were found nearby in the litter. These rhizomorphs, as well as
penetrating the roots of adjacent trees, also break up into white strands and
into hyphae which lie along the orchid rhizomes and are anchored at intervals
to the hairs. In two specimens, the fungus was a different one with yellow
strands, and in a further specimen the strands had a few wide, centrally placed
hyphae surrounded by narrower ones. In all the fungi involved, the hyphae
were septate without clamp connections.

The structure of the orchid rhizome was studied microscopically in
transverse and longitudinal sections. There is a central vascular cylinder with 4
to 6 vascular bundles enclosed by an endodermis with Casparian bands. The

1970

THE MICHIGAN BOTANIST

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cortex, 16 to 20 cells in width, consists of starch-storing parenchyma. At the
surface is an epidermis with a cuticle. The papillae scattered over the rhizome,
and sometimes also on the bases of the scale leaves, are 6 to 10 cells wide at
the base and 5 to 6 cells high; they carry rather stiff unicellular hairs up to
0.5 mm in length (Fig. 2).

The fungal infection of the rhizome follows a characteristic course.
Fungal strands and individual hyphae often lie appressed to the epidermis in
the depressions at the nodes, but penetration occurs only through the tips of
hairs on those papillae lying a short distance back from the growing-point.
The entering hyphae grow down the hair and at first spread through the cells
of the papilla in a loose branching meshwork. The nucleus of infected cells
enlarges and becomes lobed as some of the hyphae lose their content and
collapse, but other hyphae remain intact. The basal wall of the hair cell and
sometimes the innermost wall of deeper cells of the papilla becomes thickened
and brown but gives no reaction for lignin or suberin. Later, when the hairs
have withered, the thickened walls persist as a scar over the papilla. Hyphae
from the papilla region enter the cortex and spread laterally and inwards in a
definite pattern (Fig. 2). The epidermis and outer 2 to 3 rows of the cortex
remain free of fungus. There is next a belt, 2 rows of cells in width, which
harbours the fungus without apparently being affected by it. But in deeper
cells the position is different. In a belt some 5 to 9 rows in depth the fungus
first occupies the cells with branching coiling hyphae. The nucleus of infected
cells enlarges and tends to become lobed while the hyphae, which at first
contain much protoplasm, lose their content and collapse, leaving only a
densely staining clump of unabsorbed material at the centre of the cell. A
belt, 3 to 6 rows wide, outside the endodermis is not penetrated by the
fungus and the cells here retain their starchy contents and accumulate fatty
oils as well.

3 ! d

Fig. 3. Part of the rhizome of C. striata in transverse section, x 185. d. zone of fungal
digestion, p. papilla.

112

THE MICHIGAN BOTANIST

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Corallorhiza striata

C. striata is another non-green orchid which is endemic to North Ameri¬
ca. In northern Michigan it grows amongst the roots of a variety of trees. The
10 specimens selected for investigation were collected from under Thuja occi-
dentalis and Picea sp., from under Thuja occidentalis and Betula lutea Michx.
f., from under Quercus rubra and Acer saccharum , and from pure stands of
Pinus strobus, of Pinus resinosa, of Fagus grandi folia, and of Betula
papyrifera.

The underground portion of the orchid consists of a branching system
of rhizomes, each 2 - 5 mm in diameter and lying more or less horizontally in
the litter layer amongst fallen leaves and the fine roots of forest trees. Some¬
times the rhizomes resemble in appearance those of C. maculata; at other
times they are brown in colour with short, white, fungal-free tips. They have
no roots.

Transverse sections of the rhizome show an internal structure resembling
that of C. maculata. The central vascular cylinder has 3 to 12 small vascular
bundles enclosed by an endodermis with Casparian bands. There is a paren¬
chymatous cortex, 16 to 17 cells in width, and at the surface there is an
epidermis with a cuticle. The papillae are 4 to 14 cells in diameter and 3 to 8
cells high. However, their surface cells, unlike those of the two previous
species, have convex outer walls but no projecting hairs (Fig. 3).

Two different fungi were found associated with the rhizomes. In one of
these the hyphae, which are of a diameter of 3 - 4.5/i, have a brown, external
wall except near the growing tips and are septate without clamp connections.
The brown walls give the brown colour to the rhizomes in which they become
established. This fungus in one area is also mycorrhizal on the roots of the
Pinus strobus alongside the orchid and grows as a fungal mantle and Hartig
net on the pine roots. In another area the fungus is mycorrhizal as a fungal
mantle and Hartig net on the roots of either Acer saccharum or Quercus
rubra. In Thuja roots there is no superficial mantle but the fungus is in the
cortex, penetrating first along the walls and later lying in the cell cavities, and
penetrating into the vascular cylinder of dead roots. In the case of the other
fungus the hyphae within the orchid rhizome are colourless and of diameter
1.5 - 3/i, but outside the rhizome they may have brown walls; they are septate
without clamp connections. They are also to be found in the cortical cells of
the fine roots of Betula papyrifera and in the cortex and vascular cylinder of
fine roots of the Fagus grandifolia. Most of the fine roots of Fagus near the
orchid were found to be dead.

The fungus enters the orchid rhizome through surface cells of papillae
near the growing tips. These surface cells appear to be a special point of entry
for the fungus. Their basal walls later become thickened and suberized. The
hyphae grow through the papilla into the cortex where they spread laterally
(Fig. 3). The epidermis and outer 2 to 4 rows of cortical cells remain free of
fungus. There is then a belt, 3 to 5 cells wide, occupied by fungal coils, or, in
the case of the colourless fungus, by a fungal meshwork, followed by a zone 4
to 7 cells wide in which the coils eventually collapse leaving only a darkly

1970

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staining clump of unabsorbed material. The innermost cortical cells remain
free of fungus and contain starch.

DISCUSSION

All three species of Corallorhiza were found to be able to form an
association with more than one species of fungus. This supports the findings of
earlier workers for C. innata (Jennings & Hanna, 1898). The condition, how¬
ever, contrasts with that in such examples as Gastrodia cunninghamii where
the orchid is associated with one particular fungus (Campbell, 1962).

The surface cells of the papillae in C. striata and the tips of the hairs
projecting from the papillae of both C. striata and C. maculata are the point
of entry to the rhizome for certain fungi. However, for each species of
Corallorhiza there is a particular range of fungi, as indicated by the fact that
only in C. trifida do the fungi show clamp connections and only in C. striata
is the brown-walled fungus found. Once the fungus has entered the rhizome,
its advance into the cortex and later its digestion by tolypophagy follows a
similar pattern in all three species.

Some of the fungi concerned are inhabitors of the roots of trees where
they behave as weak parasites. But they also flourish in the soil amongst the
trash of wood and bark left after felling operations or left after snow and
wind damage. Others are mycorrhizal on roots of adjacent trees.

The orchid, particularly in the case of C. trifida , although obviously
obtaining nourishment from the fungus, forms a relatively harmonious associa¬
tion with it, for the infected cells suffer no obvious damage. This situation
contrasts with that in some other orchids, such as Gastrodia minor, where the
infected region of the tuber ultimately collapses (Campbell, 1963).

ACKNOWLEDGMENTS

Acknowledgment is made to the National Science Foundation for funds for re¬
search at the University of Michigan Biological Station in 1968 (grant number
NSF-GB-6095) and also to Massey University for a study leave grant.

LITERATURE CITED

Campbell, Ella O. 1962. The mycorrhiza of Gastrodia cunninghamii Hook.f. Trans. Roy.

Soc. N.Z. 1: 289-296.

Campbell, Ella O. 1963. Gastrodia minor Petrie, an epiparasite of manuka. Trans. Roy.

Soc. N.Z. 2: 73-81.

Jennings, A. V., & H. Hanna. 1898. Corallorhiza innata R.Br. and its mycorhiza. Scient.

Proc. Roy. Dubl. Soc. 9:1-11.

Thomas, M. B. 1893. The genus Corallorhiza. Bot. Gaz. 18: 166-170.

114

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Vol. 9

SPHAGNUM IN CHARLEVOIX COUNTY, MICHIGAN

William M. Zales and Howard Crum
Joliet Junior College, Joliet, Illinois, and
Herbarium, University of Michigan, Ann Arbor

As a result of its history of glaciation, Michigan abounds in poorly
drained habitats favorable to the growth of Sphagnum. In recent years con¬
siderable interest in the taxonomy, distribution, and ecology of the genus, so
poorly understood by American bryologists, has centered around the Univer¬
sity of Michigan Biological Station in northern Michigan. Access to the
abundance of collections made by numerous bryologists visiting the area or by
students in bryology over a period of 50 years has enabled the junior author
to confirm records of 25 species from the Straits of Mackinac area (Cheboy¬
gan, Emmet, Mackinac, and Presque Isle counties). Detailed descriptions of
those species, as well as a key for their determination, will be provided in a
manual of the bryoflora of the Straits region now in manuscript. Charlevoix
County, lying immediately southwest of this group of counties, on the shore
of Lake Michigan, was selected for special study in the summer of 1969
because of ease of access from the Biological Station and because not a single
record of Sphagnum is noted from that county in Darlington’s recent Mosses
of Michigan. We made several trips to the county and found 15 species in the
localities listed below (and subsequently referred to by number only). All of
these are also known frofn the Straits counties, which have a Sphagnum flora
of 25 species, as compared with a total of 27 known from the entire state and
about 45 from all of North America.

1. SE 34 sec. 31, T33N, R7W, Monroe Creek, about 2 miles west of the
south arm of Lake Charlevoix, in a dense, ecologically immature
Thuja-Larix swamp.

2. SW 34 sec. 10, T32N, R4W, south of county road 626, in an open,
sedgy Larix swamp with many trees dead because of alteration of the
water table by road building.

3. SW 34 sec. 31, T33N, R7W, Monroe Creek, about 3 miles west of the
south arm of Lake Charlevoix, in a seepage area in a Fraxinus
swamp.

4. NE 34 sec. 25, T33N, R6W, about 1.5 miles northeast of Boyne City
on Highway M-75, in an open, disturbed Thuja-Larix swamp (with
many trees dead owing to disturbance of the water table).

5. NW 34 sec. 29, T33N, R4W, south branch of Spring Brook, on Major
Road, in a mature Thuja swamp.

6. SE 34 sec. 35, T33N, R4W, 1 mile north of county road 626, on
Magee Road, in a partially drained Larix-Betula-Vaccinium swamp.

S. capillaceum (Weiss) Schrank var. capillaceum 1, 2, 4, 5, 6. Var.
tenellum (Schimp.) Andr. 4.

S. centrale C. Jens. 1, 2, 5, 6.

1970

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115

S. cuspidatum Ehrh. ex Hoffm. 6.

S. fimbriatum Wils. ex J. Hook. 1, 6.

S. fuscum (Schimp.) Klinggr. 2, 4.

S. girgensohnii Russ. 1 , 4.

S. magellanicumBrid. 6.

S. majus (Russ.) C. Jens. 6.

S. recurvum P.-Beauv. var. recurvum 6. Var. ten ue Klinggr. 1, 2, 4.

S. riparium Angstr. 6.

S. russowii Warnst. 1,5,6.

S. squarrosum Sw. ex Crome. 6.

S. teres (Schimp.) Angstr. ex C. Hartm. 1, 6.

S. warnstorfii Russ. 1, 2, 3, 4, 5.

S. wulfianum Girg. 1.

A full set of specimens is deposited in the herbarium of the University
of Michigan Biological Station.

All the species listed here are common and widely distributed, except
for Sphagnum riparium, which is rare and local over a broad range including
northern Europe and Asia, Greenland, Labrador to British Columbia, south¬
ward to New Hampshire, Quebec, Ontario, and Michigan. Although it has not
been reported previously from Michigan, it has been collected in the past few
years in several Michigan localities: Charlevoix Co. (on Magee Road, 1 mi.
north of county road 626, W. M. Zales 892, UMBS, MICH); Emmet Co.
(Stutsmanville Bog, N. G. Miller 4547, UMBS, and W. S. G. Maass, July 27,
1967, MICH); Iron Co. (Teepee Lake, R. Harris 725, MSC); Luce Co. (Rat
Lake, H. Crum, June 27, 1968, UMBS); Otsego Co. (junction of Gibbs and
Marquardt Roads, H. Crum, Aug. 7, 1969, MICH); Schoolcraft Co. (0.3 mi.
west of Luce County line on Michigan Highway 28, W. S. G. Maass, Aug. 3,
1967, MICH).

S. riparium occurs in moderately acid or somewhat basic situations, in
relatively wet boggy habitats, especially at the margins of bog ponds, in black
spruce bogs and Thuja swamps. This is a distinctive species, easy to recognize.
The plants are large and green or whitish-green and resemble S. recurvum in
many ways but may be distinguished by a large terminal bud and stout, turgid
branches with broad, imbricated leaves narrowed to a small, recurved tip and
not undulate. The leaf tip consists of uniformly narrow, green cells, rather
than the usual network of green and hyaline cells. The most striking feature,
however, is the large stem leaves (about 1 .6 mm long as compared with 1 mm
in S. recurvum) which are conspicuously perforated or deeply split at the
middle owing to localized resorption of the walls of hyaline cells.

We are indebted to the National Science Foundation for funds from
Grant GB 8440, made available to the senior author through the Biological
Station, and for Grant GB 6095, provided to the junior author for Workshops
in Bryology.

116

THE MICHIGAN BOTANIST

Vol. 9

A BOTANICAL INVENTORY OF SANFORD NATURAL AREA.

I. THE ENVIRONMENT

John H. Beaman

Beal-Darlington Herbarium, Department of Botany and Plant Pathology

Michigan State University, East Lansing

“We can never have enough of Nature.”

Thoreau in Walden

Sanford Natural Area is one of the few local remnants of the great
forests which once covered most of southern Michigan. For over a century it
has served as an outdoor laboratory for faculty and students of Michigan State
University. It has been the subject of forestry research in timber, firewood,
and maple syrup production, fire protection, and recreation management.
Urbanization of the campus necessitated transfer of this research to less popu¬
lated areas, but at the same time vast increase in University size has intensified
the need for natural areas convenient for teaching. Thus Sanford Natural Area
now serves as a major instructional resource for introductory courses in bio¬
logical science and such upper level courses as plant taxonomy, ecology, in¬
vertebrate zoology, ornithology, entomology, and biogeography.

Among the major universities Michigan State is perhaps unique in having
a highly diversified woodland conveniently located for efficient use by classes.
This very convenience poses the greatest threat to Sanford as a natural area,
for in recent years its periphery has been a choice residence hall building site.
With the greatly increased student population have come uses and abuses which
may ultimately destroy it unless effective protective measures are instituted.

Ten other areas on the University campus, amounting to some 324
acres, are wooded and in varying states of naturalness. None of the others,
however, is as conveniently located for most instructional programs as is San¬
ford. It is also doubtful that any of the other woodlots have a comparably
rich flora. Nor are they yet so seriously endangered as Sanford, although
encroachment on Baker Woodlot has just begun with construction of the first
part of a new medical school complex along its east side.

A principal aim of this account is to record some of the values of
Sanford Natural Area. Any observant person might perceive it as an unusual
feature of the campus landscape. But its scientific legacy, utility, and potential
are not immediately evident to the layman. Hopefully, the information sum¬
marized here will provide a basis for greater appreciation of this remarkable
resource by faculty, students, and the public. Its preservation will require the
concern and cooperation of the entire community. If indeed it can be saved,
it could become an outstanding case history on the conservation of a natural
area in the face of steadily increasing population pressures.

Another aim of the study is to provide a thorough documentation of
one aspect of the Sanford environment— its vascular plant flora. A detailed
understanding of the composition of natural environments is fundamental to

1970

THE MICHIGAN BOTANIST

117

any consideration of environmental quality control and rehabilitation. The
enormous species diversity of Sanford Natural Area and the unusual nature of
many of its species can provide challenging lessons to the student of environ¬
mental quality.

My interest in the woodlot began when I first saw it in 1956. At that
time Claytonia virginica was in full bloom, a beautiful pink carpet of delicate
spring wild flowers. I shortly came to depend on Sanford for three courses,
but not until I began teaching a large new course in introductory plant tax¬
onomy in 1964 did its instructional value become most evident to me. All of
the nearly 250 students annually enrolled in this course have an opportunity
(in small groups of about 16) to study the Sanford flora. The checklist of
vascular plants (to be published in the next issue of The Michigan Botanist)
started as a limited list for use in the course. By 1968 so many species had
been added that it was evident an intensive and systematic search ought to be
made. Field trips were conducted that summer at two- to three-week intervals,
each adding 30 or more new records for the Area. Although the present list of
358 species should be nearly complete, we have continued to find additional
species up to the time of this writing (November, 1969). There may also be
species added, chiefly weedy ones, which are not now in Sanford. Likewise
there may be subtractions, as there have already been, but I fervently hope
the many uncommon native plants can be preserved.

PHYSICAL SETTING

Sanford Natural Area (Fig. 1, Fig. 2) is located in the S Vi of the SEx/4 of
Sec. 18, T 4 N, R 1 W of the Michigan Meridian, Ingham County, on the
Michigan State University campus, immediately south of the Red Cedar River,
between Hagadorn Road on the east and Bogue Street on the west. It is
bordered by dormitories on the west and south. It formerly included 55.5
acres, but was reduced to 35 by building and road construction between 1956
and 1965.

The land surface slopes generally toward the river, but two levels of
benches are apparent on the topographic map (Fig. 1). The soils on these
benches (Fig. 3) commonly differ from those on the lower lying alluvial areas
which are subject to periodic flooding by the Red Cedar River. Natural soil
drainage is good on the highest bench, above elevations of 840 feet, where
Oshtemo loamy sand predominates. The lower bench of Metamora soils is less
well drained, and drainage is poorest on the alluvial Ceresco soils. The natural
levees of Landes soils along the river are better drained. Elevation of normal
water level at Bogue Street Bridge is 829 feet, and extreme high water level at
that location is 836.5 feet. Maximum elevation in the woodlot is 852 feet.

The well drained or zonal soils of Ingham County belong to the sub¬
order of Udalfs in the New Comprehensive Soil Classification in the United
States (Soil Survey Staff, U.S.D.A., 1960 et seq.) and are represented in
Sanford Natural Area by the Oshtemo and Lapeer series. These were formerly
the Gray-Brown Podzolic soils of Central and East-Central parts of the United
States (Veatch et al., 1941; NCR. 3 Committee, 1960).

118

THE MICHIGAN BOTANIST

Vol. 9

Fig. 1. Topographic map of Sanford Natural Area, based on 1965 revision of blueprint by Division of Physical Plant Planning and
Development, Michigan State University. (The narrower trails have developed from recent usage.)

1970

THE MICHIGAN BOTANIST

119

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120

THE MICHIGAN BOTANIST

Vol. 9

Average annual precipitation in the East Lansing-Lansing area for the
period 1864-1968 was 30.84 inches, with a maximum of 48.44 inches in 1883
and a minimum of 18.50 in 1930. For the 30-year period from 1931-1960 the
greatest monthly rainfall was in May (3.73 inches), June (3.34 inches), and
August (3.05 inches); the least in December (1.99 inches), January (1.96
inches), and February (1.95 inches). For the same period the average daily
maximum temperature was 57.1° F and the average daily minimum 38.2°,
with an average annual temperature of 47.6°. The average daily maximum for
this period was highest in July with 83.4° and the average daily minimum was
lowest in February with 16.5°. (Data from the Environmental Science Services
Administration, State Climatologist, East Lansing.)

According to Westveld (1931) and an unsigned, undated report in the
Forestry Department, records (not seen) made by Dr. W. J. Beal show that
about 150 trees, especially beech, died as a result of the drought of 1894 and
1895, with destruction greatest on the west side of the woods. Similarly, the
death of 48 trees, about half of which were beech, was attributed to the
drought of 1934 (unsigned report of January, 1936, in the Forestry Depart¬
ment). These appear to be the only written records which suggest that recent
climatic fluctuations have adversely affected the Sanford vegetation.

Apparently climatically related effects are evident, however, on radial
growth as shown by increment cores from trees in the Area (11 species, 18
trees, listed in Table 3). The earliest records obtained from the cores indicate
an adverse period in the early 1870’s. This was followed by a period of
mostly normal growth until the late 1880’s when another adverse period be¬
gan which lasted to the mid-1 890’s. In a subsequent period through 1923
optimum conditions for growth prevailed. Normal growth for some trees and
slow growth for others followed until the early 1930’s. An especially adverse
period from this time until the late 1940’s is indicated by most trees in the
sample. Relatively normal growth rates have prevailed in about half the sample
from that time to the present, although the early 1960’s was a time of slower
growth for several species. These times of slow, normal, and rapid growth
correspond for the most part with cumulative precipitation for the same
periods.

There was an average precipitation deficit of two inches per year from
1867 through 1875, a deficit of nearly four inches per year from 1888
through 1895, a deficit of two inches per year from 1930 through 1939, and
a deficit of nearly five inches per year from 1960 through 1966. The principal
failure in correspondence of growth rates and precipitation is from 1940-1949
when there was a three-inch average annual precipitation excess for the decade
but the majority of the trees in the sample grew more slowly than normal.
This might possibly reflect a slow recovery from the drought of the 1930’s,
but other factors could be involved. The limited data from the increment
cores and the observations of trees dying during droughts suggest that, even in
this area of relatively high rainfall, precipitation may often be a major factor
in growth rates or survival of trees.

1970

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121

Fig. 3. Map of Sanford Natural Area showing wards, firebreak, and trails from map of 5 November 1895; soil boundaries from map by
A. J. Quinkert with the advice of J. O. Veatch, c. 1931, revised in 1969 with tentative correlations by E. P. Whiteside; Ca, Carlisle
muck; Ce, Ceresco fine sandy loam; Gr, Granby sand; Lan, Landes fine sandy loam; Lap, Lapeer sandy loam; Me, Metamora loam; Os,
Oshtemo loamy sand.

122

THE MICHIGAN BOTANIST

Vol. 9

ADMINISTRATIVE HISTORY

Sanford Natural Area is part of 676.57 acres of forested land purchased in
1855 from A. R. Burr of Lansing as the site for the Agricultural College of the
State of Michigan. Within 40 years all but 150 acres had been cleared for build¬
ings and fields. On December 11, 1894, the State Board of Agriculture
passed a resolution introduced by Charles Garfield, Chairman of the Commit¬
tee on Botany and Horticulture, to the effect that College policy would be to
preserve and place all remaining forest areas on the College farm in such a
condition as to illustrate good woodlot management for continuous and last¬
ing crops. The woodlots were placed under the supervision of the professor of
agriculture who managed them under the guidance of the professor of botany
and forestry. On August 28, 1901, the Forestry Department was established
and charged with management of the woodlots.

One of these was originally called North Woodlot and Woodlot No. 1. It
was later named River Woodlot, the name in general use in the 1930’s. In
1941 Professor Paul A. Herbert, then Head of the Forestry Department, re¬
named it Sanford Woodlot, after Frank Hobart Sanford (1880-1938; B.S.,
1904; M.S., 1913; Forestry faculty, 1906-1920; the second full-time forester
appointed by the College).

On December 11, 1959, President John A. Hannah transferred jurisdic¬
tion of Sanford Woodlot from the Forestry Department to Campus Planning
and Grounds Maintenance (now Division of Campus Park and Planning). In
1964 the name was changed to Sanford Natural Area, to better reflect the
purposes to which it is currently dedicated.

Over the protests of numerous Michigan biologists and conservationists a
new era in its history began in 1956 when about two acres on the west side
were taken over for construction of a women’s cooperative residence hall (Van
Hoosen). Since that time three other residence units (Owen Hall, 1960-65;
McDonel Hall, 1963; Holmes Hall, 1965) have been built into the west and
south sides. Bogue Street Bridge (west side) was built in 1964, and Hagadorn
Road (east side) was widened in 1965. This construction reduced the size of
the Area by about 20 acres, obliterated at least one major habitat, and
eliminated several species from the flora.

In 1895 the woodlot was subdivided into six wards (Fig. 3), each con¬
taining approximately 10 acres (Smith, 1896). The purpose of the subdivision
was for designating and recording conditions of the different portions of the
forest and work performed upon them. At this time a general clean-up was
made to remove debris and fallen trees. A firebreak four rods wide north and
south through the center of the woodlot was cleared of all underbrush, old
logs, and leaves. This strip was not kept up very long, however, since there
was no evidence of its existence in 1931 (Westveld, 1931). While brush from
the clean-up was being burned in the spring of 1895 fire escaped three times
and burned approximately 15 acres, mostly in Wards 1 and 2 but also near
the north end of Wards 4 and 5. There apparently are no more recent records
of fire in the woods.

1970

THE MICHIGAN BOTANIST

123

Cattle have always been restricted from the woodlot. The basis for this
was stated by Crozier (1896):

Two principal enemies of forest property are live stock and fire. Where stock
is confined to a limited section of woodland all underbrush is soon destroyed and
the woods became open and park like, the surface more or less covered with grass,
attractive places for picnic parties, etc., but of little further use as a forest. The
duration of such a forest is necessarily limited to the life of the large timber, and
even this often prematurely dies from the unwonted exposure.

To the principal enemies of forest property noted by Crozier we must
now add the excessive human population.

HARVESTING OPERATIONS

The forest is of mostly natural origin and uneven age, but it is by no
means virgin as some people seem to think. It was managed for 65 years to
demonstrate sound forestry practices for farm woodlots. Prior to that time a
considerable number of trees in certain areas had been cut for fuel. During the
management period harvest cuttings were both selective and conservative. A
major objective was maple syrup production, so virtually all trees of Acer
saccharum were spared. According to Crozier (1896) the large timber was cut
from several acres in one ward (Ward 1) a few years prior to 1896, and a
thrifty second growth took its place. In the fall and winter of 1896-97 about
150 dead or dying trees were converted into fuel wood. In 1912 one acre in
the northeast corner was selected as a practice field for silviculture students
doing thinning operations (Walline, 1940). During the winter of 1931-32 all
dead timber was cut, amounting to 100 standard cords of fuel wood. A
salvage cutting was made in 1936 to remove 48 trees which had been killed
by the drought of 1934. Westveld’s (1931) management plan indicated that
almost the entire woodlot would be given a selection cutting in 15 years, but
the cutting was to be restricted principally to trees 12 inches d.b.h. and over.
It is not clear from the records that this policy was implemented, but a
portable sawmill was located in the woodlot from 1934 to 1948, so it may be
assumed that considerable cutting was done. The lumber manufactured was
used by various departments, partly for building construction, and for the
annual Winter Carnival. The firewood produced was used for evaporating
maple sap, with the excess being sold to local residents and commercial com¬
panies. The last harvest by the Forestry Department was in 1959.

MAPLE SYRUP PRODUCTION

The woodlot was first used for syrup production in 1915, at which time
an evaporator house (referred to subsequently as “sugar house”) was erected
and equipped with a Leader Evaporator. The trees were tapped for the last
time in 1960, and the sugar house shortly removed, but its site remains a
distinctive part of the woodlot (see p. 134). Early research efforts in syrup
production concerned the effect of varying numbers of buckets on sap flow.
This research was gradually broadened to include other problems related to
the operation of a sugar bush, such as fertilizer effect on the yield of syrup,

124

THE MICHIGAN BOTANIST

Vol. 9

the effect of cultural operations on sap yields, and financial aspects of operat¬
ing a sugar bush (Anonymous, 1953). Walline (1940) reported that there were
ca. 748 sugar maple trees large enough for tapping (12 inches or greater
d.b.h.) in the entire woodlot, but only 600 were conveniently situated for
tapping. The average yield per tree based on a nine-year record between 1915
and 1925 was 0.2 gallons of syrup. The highest yield secured was 0.34 gallons
per tree. During this period 1,050 gallons of syrup were produced with an
average annual yield of 116.6 gallons.

RECREATION MANAGEMENT

The recreational values of Sanford Natural Area have been recognized
since management procedures were first instituted. Crozier in 1896 noted that
the roads and other improvements were making it a forest park which was
becoming a popular resort for students and others. The management plans
prepared by Westveld (1931) and Walline (1940) indicate that the area was to
be maintained in a condition suitable for recreation. As early as 1931 keeping
cars off the trails was a problem, a circumstance now paralleled by the pro¬
hibited but all too frequent motorcycles. For a number of years certain trails
were part of the bridle trail system, but not all riders managed to keep their
horses on the trails. Partly because of this abuse the woodlot was closed to
equestrian travel in 1949. Certain trails have been in use for many years by
the cross-country track team. About 20 years ago the road near the river was
made into a nature trail, with labels identifying the plants (Wilcox, 1949).
This was not maintained very long, however, and by 1969 only one label was
still evident. In spite of the recent unsightly apartments on the north side of
the river, Sanford contributes much to the attractiveness of canoeing, a cur¬
rently popular pastime on the Red Cedar. It also serves as a pleasant buffer
between the campus and the apartment and business district to the north.

The most important recreational use of Sanford Natural Area is also the
most difficult to document. Thousands of people live in its vicinity. Some
probably care nothing for it and may even resent its presence. Many others,
however, do appreciate it as a place they can go on a pleasant day and enjoy
nature. Persons who have grown up in the East Lansing area have often
remarked to me of fond memories of childhood outings in the woodlot.

The recreational and cultural values of natural areas have been force¬
fully expressed by litis (1968): “We need nature and its diversity in order to
survive as human beings, as well as a technological and agricultural civilization.
We must, therefore, accept that nature is the most precious of all human
possessions. Our educational system’s main responsibility, therefore, is to train
people, not only in the exploitation, but now also in the protection of na¬
ture— the only nature to which man is adapted, in which man can exist.”

SANFORD AS AN INSTRUCTIONAL LABORATORY

A summary of current use of the Area for formal instruction in 21
courses is presented in Table 1. Obviously not all class time for a particular
course is spent there, but it is nevertheless a basic resource for these classes.

1970

THE MICHIGAN BOTANIST

125

TABLE 1. Courses currently using Sanford Natural Area as a laboratory, grouped by depart¬
ments as follows: Biological Science, Botany and Plant Pathology, Entomology, Fisheries
and Wildlife, Forestry, Geography, Lyman Briggs College, Zoology.

Course

Approximate no. of students per year

B S

202,

Foundations of Biological Science

450

BOT

318,

Introductory Plant Taxonomy

250

BOT

450,

Ecology

5

BOT

823,

Plant Taxonomy

10

ENT

302,

General Entomology

30

ENT

420,

Aquatic Insects

40

ENT

438,

Taxonomy of Immature Insects

7

ENT

820,

Insect Ecology

7

F W

402,

Conservation Education

100

F W

403,

Conservation Education

100

F W

483,

Advanced Conservation Education

15

FOR

302,

Forest Biometrics

36

GEO

411,

Problems in Geography

10

GEO

431,

Advanced Physical Geography

10

GEO

432,

Biogeography

20

LBC

140,

Biology

250

ZOL

204,

Natural History of Birds

17

ZOL

461,

Ornithology

50

ZOL

481,

Invertebrate Zoology

60

ZOL

486,

Mammalogy

10

ZOL

881,

Biology of the Arthropoda

12

TOTAL

1,489

Other important instructional values of the woodlot are less tangible. For
example, many of the more advanced students in plant taxonomy have done
their first fieldwork there, as an examination of collection numbers in the
checklist will demonstrate. The amount of instructional use the Area can
sustain is naturally somewhat limited; a class of 450 students cannot be
turned loose to forage at will. But divided into small groups of 10 to 20,
which stay on the ample trails, a large class can use the area without undue
damage. All classes are expected to avoid picking flowers, breaking shrubs,
trampling the soil, or otherwise altering the natural environment.

The opportunity for special training and research which the woodlot
offers is illustrated by several undergraduate and graduate students currently
working in the Beal-Darlington Herbarium who are developing procedures for
computerizing floristic data. They have already prepared preliminary taxonom¬
ic data matrices for the woody species, herbaceous spring dicots, and spring
monocots of Sanford. These data matrices will be used in connection with the
teaching program in introductory plant taxonomy.

Although University students are the principal users of the Area, it is
noteworthy that secondary schools in East Lansing are also taking classes
there.

126

THE MICHIGAN BOTANIST

Vol. 9

PLANTINGS

On several occasions native and exotic species have been planted in the
woodlot. The most ambitious attempt was in 1896, a description of which is
provided below (from an unsigned, undated report in the Forestry Depart¬
ment; cf. also Crozier, 1897; cf. Fig. 3 for trail locations and ward bound¬
aries).

In this woodlot were planted in the spring of 1896 the following seeds and
young trees. The nuts and acorns were furnished by Dr. W. J. Beal, also the young
redbuds and black locust sprouts. The locusts, about fifty in number, were set in
Ward 1 between the crossroad and the ravine. The redbuds, a dozen or more, were
mostly set on the river bank in the lane from two to ten rods west of the woods.
These were one year old from seed. A few chinquapins were planted in the south
half of Ward 1, north of the south drive. In the same locality a smaller number of
chestnuts were planted, also a few of the same all along the south drive and a
group of the same on newly plowed land in the extreme southeast corner of the
woodlot where they were protected and cared for during the season and made a
good start. Some of the chinquapins were in poor condition when they were
planted here. In Ward 1 in the same general locality as the chinquapins, north of
the south drive, were planted about one fourth of a bushel of shucked walnuts.
About the same amount of hickory nuts were planted in a thin spot in the south
end of Ward 3 where there stands a tree of the same species about ten inches in
diameter. About half a peck of butternuts was planted in the same general locality
but mostly further south and more scattered. A few red oak acorns were planted
in Ward 1 along the sides of the crossroad and near its north end. Also a large
number of red oak acorns were planted along both sides of the drive across the
north end of Ward 4. In the north end of Ward 3, near a large tree of the same
species were planted about two quarts of bur oak acorns. About the same quantity
of swamp white oak acorns were planted along the bayous in the north end of
Wards 3, 4 and 5. About a dozen nuts of Carya porcina [= C. glabra ] were planted
in Ward 2 south of the south drive about four rods west of the point of its
greatest northerly bend. They were planted in open places in a belt about eight
rods long north and south.

At the right of the drive just after entering the woods from the river lane
were set three pawpaws taken from the opposite bank of the river. Along the
south side of the lane were set four groups of trees native to Michigan, deciduous
trees all from the neighboring river banks, woods or elsewhere on the farm. The
conifers were furnished by the Horticulture Department and were from where they
had been grown a few years in the nursery row. At the right of the entrance
leading from the main farm lane were set a mixed group, including hard and red
maples, basswood, tulip, red alder, Juneberry, hazel, red cedar, arbor vitae, and
white pine. All made a good start during the season except the Juneberry. The
white pine was a large spreading tree about five feet high, the larger of the two red
cedars was four to six feet high and the two arbor vitae each about two feet high,
though one of them was partly broken down during the summer. On the steep side
of the river bank opposite this group, were set two red cedars of about the same
size as those set in the group. Further east on the next rise was set a large group
of deciduous trees including three species of elm, beech, hawthorn, Juneberry, and
ninebark. On the next rise of land to the east was set a group of prickly ash, and a
little further east on swampy ground which is now drained where the line fence
begins to run straight were set four thrifty young cedars of the same size and from
the same source as the two set in the first group at the west end of the lane.

Along the river bank just west of the bridge to the east side of the little
stream coming down out of Ward 1, were set, chiefly in the woods, ten red cedars

1970

THE MICHIGAN BOTANIST

127

12 to 18 inches high. By the lagoon, just east of the firebreak, were set ten arbor
vitae about 12 inches high. On the south river bank just west of the bridge, were
set two white spruce, eight or more of the same being set just north of the south
drive in Ward 1. They were about two feet in height. Those in the woods were
badly eaten by squirrels soon after being set and made a poor start. At the angle
between the north and the cross drive were set eleven white pines, mostly mis¬
shapen or injured in some way but vigorous and four to six feet high. These were
obtained from the Horticulture Department.

In the east end of the same woodlot in the vicinity of the gates into the
highway were set ten red pines about three feet high. In Ward 5 on the margin of
an old gravel pit were set about ten hemlocks and in Ward 2 on the north side of
the north drive about twenty more of the same species were planted.

Of the species planted in 1896, the only ones known with certainty to
have survived to 1969 are about a dozen black locusts (the largest now 21
inches d.b.h., but most of them 9-12 inches) and four hemlocks (5-8 inches
d.b.h.). A chestnut persisted until Van Hoosen Hall pre-empted its space.

In addition to the planting program of 1896, four bushels of black
walnuts were planted in the southeast corner running over into Ward 3 in
November, 1904, and in December of that year a part of four bushels of
butternuts were planted in Ward 2 and also running north and south across
Ward 1. Much more recently, subsequent to the construction of Holmes Hall,
shrubs or small trees of Halesia Carolina, Tsuga canadensis, Thuja occidentalis,
and Cornus controversa have been set out on the southeast edge of the woods.

THE FOREST IN 1895

The following notes (reproduced in an unsigned, undated report in the
Forestry Department) were recorded in October, 1895 (cf. Fig. 3 for trail
locations and ward boundaries).

Ward 1. Original timber was mainly beech and maple. The larger trees have
more than half of them been cut from time to time and along the west side the
wind has blown down most of them that were left. On the west side next to Field
No. 7 a strip about ten rods wide is covered with a continuous thicket of young
trees ten to fifteen feet high. Further in, the second growth is smaller and occurs
more in clumps with comparatively open spaces between. The larger timber con¬
sists of the following species: At the north end along the river are a few bass¬
woods, four at least over one foot in diameter; four tall bittemuts of about the
same size as the basswood; three white ash one foot or more in diameter; three red
elm respectively of six inches, one foot, and one and a half feet in diameter. On
the line in the northeast corner of the ward and one rod from the river bank is a
fine large maple six feet, ten inches in circumference at four and a half feet from
the ground and several others nearly as large, consisting of maple and beeches. One
rod from the bank and about one third of the way to the west side of the ward is
a tall thrifty butternut 45 inches in circumference, four feet from the ground.

Further south in the ward the following other trees were noted: Several
bittemuts, one red elm, one rock elm, one white ash, one butternut, one black
cherry, one scarlet oak [in this account scarlet oak probably is Quercus rubra var.
borealis ] , and about a dozen basswoods. Several of the beeches in the southwest
corner of the ward are dead and others dying, with frequently the upper half only
of the trees being affected. A large share of this injury has occurred from drought
this season.

128

THE MICHIGAN BOTANIST

Vol. 9

Near the river is a mulberry stump from which a sprout is growing fifteen
inches in circumference. This stump is 44 inches in circumference. The under¬
growth of this ward consists of the following species grouped as shrubs and trees
and arranged in about the order of abundance.

Trees. Sugar maple-the most abundant. American elm, white ash, mainly at
the south end. Beech, ironwood, in the center. Blue ash in the southwest quarter.
Red elm in the north half. Rock elm in the south half. Bitternut, red maple,
mulberry, basswood, black walnut, all small, butternut sapling, and white oak.

Shrubs. Spice bush, blue beech, red elder, American ivy, choke cherry,
bladdernut, moosewood, green osier, hawthorn, wahoo, gooseberry, black rasp¬
berry, blackberry, witch hazel, poison ivy, prickly ash, greenbrier, red osier.

Ward 2. Similar to Ward 1 with the following exceptions: Beginning at the
south end was noticed a few clumps of sumac, some small trees of black cherry, a
few saplings of white, scarlet and black oak, also younger oaks eight to twelve feet
high, chiefly white oak, several sassafras ten to fifteen feet. The undergrowth is
chiefly sugar maple. On the north half of the Ward are several tall ironwoods. Near
the northwest corner immediately on the river bank and leaning a little toward the
river are two scarlet oaks about three and a half feet in diameter. A smaller tree of
the same species stands about four rods further east. The undergrowth here for
about six rods back from the river is about eight feet high and chiefly beech.

In the northeast corner is about half an acre of bottomland in which are a
white ash 49 inches in circumference, a butternut 75 inches, two American elms
respectively 97 and 84 inches, two scarlet oaks respectively 45 and 71 inches,
these all being on the immediate river bank. Then on the lower land a little further
back are white ash 86 inches and a red elm 46 inches in circumference.

Ward 3. At the north end the bottomland widens eastward and the ridge
along the river broadens and becomes more pronounced in the same direction. In
the northwest corner of the ward, beginning at the corner stakes, are first a small,
diseased and crooked butternut, a second thrifty leaning sycamore about two feet
in circumference with a large dead spot at the base on the northeast side, caused
apparently by fire about four years ago. Other trees are a double sapling butternut,
four thrifty bur oaks about three and a half feet in diameter. The open ground on
the bottomland is covered with poison ivy.

About midway east and west in this ward the road crosses the lower land to
the bluff along the river. In the center of the low land at the edge of the road on
the west side is a hickory about ten inches in diameter, and on the opposite side a
thicket of young black ash six to ten feet high.

On the east side of the road just as it strikes the bluff are two Kentucky
coffee trees now seven feet high which have grown as sprouts from small trees that
had been cut down. These two trees were cut down again by hunters in 1896. The
timber on the river ridge on the east half of this ward consists mainly of half-
grown maples six to ten inches in diameter. Down on the river edge is a fringe of
larger white maple, while scattered over the ridge are beech, white ash, red and
rock elm, ironwood, basswood, and sassafras. Two beeches and one scarlet oak are
of large size.

In the bottomland and extending east across the firebreak into Ward 4 is a
lagoon or series of lagoons now dry bordered by a mixed undergrowth chiefly of
black ash and white maple with a few tall thrifty trees of the same species.

South of this bottomland the remainder of Ward 3 consists of a better class
of timber than Ward 2, it being taller and more thrifty, the maples rather exceed¬
ing the beech in number except at the south end where the large timber is mainly
beech. The timber, especially through the center of the ward is quite open, having
been much of it cut for fuel and the ward as a whole possibly containing no more
large timbers than in Ward 2. The entire upland of this ward is covered with young
growth five to ten feet high and becoming at places very thick and some of it is

1970

THE MICHIGAN BOTANIST

129

from ten to fifteen feet high. Hard maple prevails except on a strip about five rods
wide along the top of the bluff where it is nearly all beech. A tall thrifty butter¬
nut about one and a half feet in diameter was noted near the bluff and a solid top
of the same species over two feet in diameter south in the edge of the firebreak.
Several young butternuts are scattered through the ward and at the south end are
young trees of sassafras, poplar, sumac and white oak, mixed with the sugar maple
and white oak.

Ward 4. Timber on this ward is more uneven and at the south end the oldest
timber consists still more largely of beech than Ward 3. Much of the small growth
at the south end of the ward is red maple and dogwood. The southeast corner of
this ward is the head of a swamp, which opens toward the east. In this swamp is a
thicket consisting of red raspberry, red osier, black alder, poison sumac, and
several species of willow. In the open ground on the margin of the swamp
Osmunda regalis is abundant. On the bluff north of the swamp on a line between
Wards 3 and 4 is a continuous thicket of beech, maple, etc., four to five feet high
which becomes thicker and taller further north. In this undergrowth are also sassa¬
fras, maple-leaved viburnum, dogwood, witch hazel, red maple and several species
of oak.

The central portion of this ward contains but little large timber, most of it
having been cut for fuel. What is left is mainly beech. The surface here is entirely
covered with second growth, thicker than in any previous ward. On the bluff on a
line between this ward and Ward 5, is an especially fine mixture of maple.

Ward 5. The swamp which began on Ward 4 extends across the entire south
end of this ward and a short distance into Ward 6. It also extends a little south
over the line fence. In this swamp are about two dozen tamaracks ten to fifty feet
high and with several still smaller types. The swamp is mostly filled with willows,
ash, poison sumac, etc., but at the outlet, which is just across the line in Ward 6,
there is south of the road an open marsh of about one fourth of an acre and
another of smaller size in the southwest corner of Ward 6. The water from this
swamp discharges across the road above mentioned and loses itself in the woods to
the north. North of the swamp in Ward 5 just north of the road which is the
south drive is a gravel pit from which a large amount of excellent red gravel has
been taken for the public highway.

Ward 6. The south end of this ward is thinly timbered, chiefly with low¬
headed thrifty beeches and a few tall large maples. The surface is mostly grassed
over and bears thin, scattered clumps of beech four to six feet high. At the north
end river bottom occupies about one fourth of the entire ward and most of this
bottomland consists of a broad swell or ridge near the river. This ridge bears a
mixed growth of tall young timber, much of it about one foot in diameter,
consisting of three 'kinds of elm, butternut, black cherry, ironwood, beech, maple,
basswood, hickory of the bitternut variety, rock and red elm prevailing. On the
low land south of the ridge is a miscellaneous thicket of undergrowth. The rock
and red elm seems to prevail.

PRESENT FOREST STRUCTURE

In October, 1969, the occurrence and distribution in Sanford Natural
Area of woody species of tree growth form were analyzed using the point-
quarter method (Cox, 1967). Data for individuals one inch or greater
d.b.h. at 100 random points distributed along four transects (cf. fig. 1) were
recorded and are summarized in Table 2. The importance values were obtained
by summation of relative density, relative dominance (basal area), and relative
frequency values. From these data it is evident that Acer saccharum is by far

130

THE MICHIGAN BOTANIST

Vol. 9

TABLE 2. Density, basal area, relative dominance, frequency, and importance values of
27 woody species one inch or greater d.b.h. encountered at 100 point-quarters in Sanford
Natural Area.

Species

Density
(individuals
per acre)

Basal area
per acre
(sq. ft.)

Relative

dominance

(%)

Frequency

(%)

Importance

value

Acer saccharum

436.9

81.03

62.9

74

137.4

Fagus grandifolia

67.2

20.46

15.8

24

32.2

Acer saccharinum

62.4

14.86

11.5

14

23.4

Fraxinus americana

72.0

3.28

2.5

25

19.8

Tilia americana

43.2

3.08

2.3

25

16.6

Acer nigrum

52.8

2.54

1.9

18

14.4

Carpinus caroliniana

43.2

0.26

0.2

15

10.5

Ostrya virginiana

38.4

0.17

0.1

14

9.5

Ulmus americana

30.7

0.91

0.7

12

8.6

Quercus rubra var. borealis

24.0

1.58

1.2

9

7.2

Fraxinus nigra

7.7

0.00

0.0

3

1.9

Quercus muehlenbergii

7.7

0.18

0.1

2

1.6

Quercus macrocarpa

5.8

0.10

0.1

2

1.3

Prunus serotina

4.8

0.15

0.1

2

1.3

Linder a benzoin

4.8

0.00

0.0

2

1.2

Sassafras albidum

4.8

0.02

0.0

2

1.2

Ulmus rubra

4.8

0.00

0.0

2

1.2

Crataegus mollis

4.8

0.00

0.0

1

0.8

Acer negundo

2.9

0.00

0.0

1

0.6

Acer rubrum

2.9

0.08

0.0

1

0.6

Carya cordiformis

2.9

0.00

0.0

1

0.6

Celtis occidentalis

2.9

0.01

0.0

1

0.6

Crataegus punctata

2.9

0.00

0.0

1

0.6

Hamamelis virginiana

2.9

0.00

0.0

1

0.6

Populus deltoides

2.9

0.00

0.0

1

0.6

Robinia pseudo-acacia

2.9

0.00

0.0

1

0.6

Staphylea trifolia

2.9

0.00

0.0

1

0.6

the most important tree, but nine other species are of considerable signifi¬
cance.

The preponderance of Acer saccharum may have resulted in part from
its being left for syrup production, while other species were more heavily cut.
Sugar maple is a very shade-tolerant species, however, and is now present in
large numbers in the small diameter classes. Although it may have been fa¬
vored by selective cutting, the 1895 account indicates that it was very abun¬
dant at that time. The data for sugar maple and beech in Sanford Natural Area
are in close agreement with those for Tourney Woodlot, a 15-acre virgin tract
on the campus about two miles south of Sanford. For this stand Schneider
(1966) reported (on the basis of a 100% inventory in 1960) that there were
400.3 sugar maple trees per acre one inch or greater d.b.h. and 54.2 of beech.
All other species he noted were lower in density than the major species in
Sanford, except Ulmus rubra for which he reported 38.2 trees per acre (his
study was prior to the epidemic of Dutch elm disease). It appears therefore

1970

THE MICHIGAN BOTANIST

131

that the trees in Sanford are as densely stocked, if not more so, than in a
neighboring woods which is virtually undisturbed.

Distributional patterns of the important trees are not readily apparent
from the data in Table 2. The high frequency value of Acer saccharum sug¬
gests that it is common throughout most of the woodlot. The only place it
does not occur is around the intermittent lagoons near the river, where it is
replaced by Acer saccharinum. The latter is also common on the river margin
and is restricted to very wet habitats. The other most important trees, com¬
mon on both the floodplain and upland, are Fagus grandifolia, Fraxinus ameri-
cana, Quercus rubra var. borealis , and Tilia americana. Each of them, however, is
more abundant in some areas than in others. The ielatively slight dif¬
ferentiation of tree distribution in the woodlot makes a small-scale vegetation
map impractical. I have been unable to clearly distinguish five forest types
which Westveld mapped in 1931.

A recent drastic change in the relative abundance of trees in Sanford (as
elsewhere in south-central Michigan) has been caused by Dutch elm disease. In
1961 Dr. G. W. Parmelee examined all the larger elms in the woodlot and
found in healthy condition 395 Ulmus americana (including a few U.
thomasii) and 220 U. rubra of eight -inch or greater d.b.h. An additional 73
trees were noted which were suspected of harboring the disease. By 1969
almost every elm greater than six inches d.b.h. was dead; only one large tree
(24 inches d.b.h.) was found alive. Many of the dead trees are still standing,
but others were cut or have fallen and are rapidly decaying.

In addition to the dead elms there is some mortality among other im¬
portant species. Much of this is the result of wind damage. At present there
are 66 trees four inches or greater d.b.h. which have recently fallen or are
severely bent or broken. These are as follows:

Species

Number of trees

Basal area
(sq. ft.)

Acer saccharum

33

30.55

Fagus grandifolia

4

7.17

Fraxinus americana

8

14.73

Quercus rubra var. borealis

3

8.75

Tilia americana

11

14.78

Miscellaneous (6 species)

7

10.17

Total

66

86.15

Much of the damage is in the upland portion of the woods, especially near the
south edge. Holes from the root systems of the larger windfalls range from 2-4
feet deep and are generally 8-15 feet across, although one hole 20 feet wide
was observed. “Tip-up” mounds from old windfalls are abundant on the up¬
land.

The dimensions, estimated age, and radial growth rate for 18 trees rep¬
resenting 11 species are indicated in Table 3. It is evident that the trees attain
most of their height long before they become exceptionally large in diameter.
The forest canopy generally ranges from about 85 to 115 feet in total height.
One of the tallest and fastest growing trees is Acer saccharinum. Several trees

132

THE MICHIGAN BOTANIST

Vol. 9

TABLE 3. Dimensions, estimated age, and radial growth of selected trees in Sanford Natural
Area.

Average

Species

d.b.h.

(inches)

Total

height

(feet)

No. of
annual
rings
observed

Estimated

aged

annual

diameter

increment

(inches)

A cer saccharin u m

34a

125

73c

130

0.25

A. saccharinum

18

100

86

90

0.22

A. saccharum

37a

110

64c

210

0.17

A. saccharum

15

70

lioc

135

0.10

A. saccharum

16

95

no

115

0.14

Fagus grandifolia

29b

90

84c

150

0.20

F. grandifolia

14

85

77

85

0.16

Fraxinus americana

33a

115

90c

150

0.21

F. americana

18

95

90

95

0.20

Juglans cinerea

17a

85

64

70

0.24

J. nigra

23a

75

72c

80

0.28

Quercus rubra var. borealis

33a

95

1 15c

190

0.17

Q. rubra var. borealis

16

95

72

80

0.24

Sassafras albidum

18a

80

68

75

0.22

Tilia americana

28a

110

98c

125

0.23

T. americana

19

110

89

105

0.17

Ulmus americana

24a

85

258

280

0.08

U. rubra

13a

85

67

75

0.17

aLargest tree in diameter of this species in the Area.
t»The largest Fagus grandifolia is 30 inches d.b.h. and 125 feet high,
increment core substantially incomplete (11 inches is the maximum length with the
equipment available); all cores taken at 4.5 feet above ground level.

^Based on the number of annual rings observed, the completeness of the increment
core, and in the case of incomplete cores the d.b.h. (excluding bark) and average annual di¬
ameter increment. These estimates do not take into account the possibility that the tree
may have existed as a seedling for an extended period.

of this species around the lagoons are between 115 and 125 feet tall. In the
central upland 95 to 110 feet is a common height for the larger trees. At the
northwest corner the trees are of lower stature, ranging from about 70 to 90
feet, possibly because this is a second-growth stand.

The faster-growing species, in addition to Acer saccharinum, include
Juglans cinerea, J. nigra , Quercus rubra var. borealis, Sassafras albidum, and
Tilia americana. Species with slower growth include Acer saccharum (one tree
of this species with 37-inch d.b.h. is the largest in the woodlot), Fagus grandi¬
folia, and Ulmus americana. None of the trees examined had a uniform rate of
radial growth throughout the period represented by the increment cores (see
p. 120). The most obvious long-term variations involve an extended period of
rapid growth from about 1896 to 1923 and a corresponding period of slow
growth from about 1930 to 1949. From the estimated ages it would appear
that most of the larger trees are at least 150 years old. Rather surprisingly,

1970

THE MICHIGAN BOTANIST

133

the oldest tree in the sample is a 24-inch d.b.h. specimen of Ulmus americana,
which would not be considered especially large, except that all the larger elms
are dead.

The diversity of woody species in Sanford parallels the total diversity of
its flora. Counting a few species no longer present, 81 spontaneous, native
woody species occur there; 56 of these could be regarded as trees (or at least
with the potential for attaining that growth form). This number amounts to
about half the native trees of Michigan. Except for the conifers, of which only
Larix laricina and Thuja occidentalis are spontaneous, the economically im¬
portant tree species of Michigan are well represented.

HABITATS

The 1895 description of the forest provides a detailed view of many
aspects of the woodland habitats. I shall therefore supplement this with a
more generalized account. Two broadly defined situations can be recognized,
the floodplain and the upland. The former has the richer flora, which prob¬
ably reflects both the importance of floodwaters in seed dispersal and the
greater diversity of environments near the river. The large number of species
in the flora is of course directly related to the broad spectrum of habitats the
Area affords. The following ten major habitat types are briefly described, with
distinctive species indicated, proceeding insofar as possible from the wetter to
the drier sites.

1. River margin. This habitat is a distinctive but not homogeneous en¬
vironmental complex. In some places there is a nearly vertical bank up to four
or five feet high; at other locations the bank is gradually sloping, and else¬
where are mud flats. Acer saccharinum is the most common tree on the bank.
Among the more conspicuous herbaceous plants is Saururus cernuus. Many of
the herbs, such as Equisetum arvense, Rumex verticillatus, and Samolus parvi-
florus, are more sporadic. Aquatics in the river include Ceratophyllum
demersum , Elodea nuttallii, and Lemna minor.

2. Intermittent lagoons. A swampy area parallels the river along all but
the west end of the woods. This is narrow to the west but becomes wider
eastward. Characteristic woody plants are Acer saccharinum, Carpinus caro-
liniana, Lindera benzoin, Quercus macrocarpa, and elms (the latter mostly
dead). Herbaceous species include Arisaema dracontium, Boehmeria cylindrica,
Chelone glabra, Dioscorea villosa, Eleocharis acicularis, Erythronium albidum,
Iris virginica var. shrevei, Lobelia cardinalis, and Mimulus ringens.

3. River ridge. A slightly higher ridge (natural levee) generally parallels
the river and separates it from the lagoons. The ridge supports an abundant
population of Toxicodendron radicans and a large assemblage of other spe¬
cies including various grasses, Acer saccharum, A. nigrum, Allium tricoccum,
Celtis occidentalis, Fagus grandifolia, Menispermum canadense, Sassafras
albidum, and Zanthoxylum americanum.

4. Carlisle muck. The muck patch near the southeast corner is of his¬
torical interest only, what with the part left after construction of Holmes Hall

134

THE MICHIGAN BOTANIST

Vol. 9

being so thoroughly drained that it supports few of the species formerly there.
A dense patch of Laportea canadensis is the principal survivor in the area that
once had Caltha palustris, Cornus stolonifera, Ilex verticillata , Larix laricina,
and Toxicodendron vernix. A small vernal pool in the extreme southeast cor¬
ner of the woodlot contained a number of interesting species, but the widen¬
ing of Hagadorn Road drained it.

5. East-central lowland. An extensive low but fairly well-drained area
with several wet depressions occurs in the east-central portion of the woods.
Common trees are Acer saccharum, A. nigrum, Fraxinus americana, and Tilia
americana. A dense undergrowth of Lindera benzoin is frequently present. In
the spring Ranunculus septentrionalis is conspicuous. Less abundant species
include Cardamine bulbosa, C. douglassii, and Cinna arundinacea.

6. Central north slope. The slope north of the sugar house site is a
springy area with a concentration of ferns including Athyrium thelypterioides,
Dryopteris goldiana, and D. spinulosa. Hydrastis canadensis has also been
found in this area, and in the late summer and fall Collinsonia canadensis is
conspicuous.

7. Northwest woods. The lowland wooded portion of about two acres
immediately north of Van Hoosen Hall has perhaps a greater number of native
species than any other small area. Two large Juglans cinerea trees are present,
a small tree of Morus rubra, abundant Fagus grandifolia, Acer nigrum, the
only colony of Hybanthus concolor, a few Dirca palustris, abundant patches
of Asarum canadense, and a number of plants of Caulophyllum thalictroides
and Actaea pachypoda. Species of the edge and riverbank also contribute to
the abundant flora of this locality.

8. Southern upland. The higher area to the south is the most homo¬
geneous portion of the woods, but even it is hardly uniform. It is dominated
by Acer saccharum with smaller amounts of Fagus grandifolia, Tilia americana,
elms (mostly dead), Fraxinus americana, and Quercus rubra var. borealis. In
late April the upland is abundantly covered with Gaytonia virginica. Dentaria
laciniata, Erythronium americanum, and Hepatica acutiloba are also common,
while Dicentra canadensis and Sanguinaria canadensis occur in scattered
colonies. Dicentra cucullaria and Dirca palustris are present but scarce. Slightly
later flowering species include an abundance of Trillium grandiflorum. Com¬
mon later in the spring and summer are Geranium maculatum, Osmorhiza longi-
stylis, Parthenocissus quinquefolia, Phlox divaricata, Viola canadensis, and
V. pensylvanica.

9. Sugar house site. The sugar house site is a more or less unique
habitat. Although it is an opening not much more than 100 feet in diameter,
a number of species occur there but hardly if at all elsewhere in the Area.
These mostly successional species include Aralia racemosa, Dipsacus laciniatus,
Populus deltoides, P. grandidentata, P. tremuloides, Rhus typhina, Rubus
occidentalis, and Scrophularia marilandica.

10. Edges. Almost as many species are found around the edge of the
Area as in its interior. Conspicuous among them are a large number of intro-

1970

THE MICHIGAN BOTANIST

135

duced weeds. The west edge is especially rich in species both native and
introduced. The Gramineae and Compositae are very well represented; one
need only scan the checklist under these two families for an indication of
their diversity. The west edge has been much disturbed by construction of
Bogue Street Bridge, which required a large amount of fill. A number of
species may have arrived by this means, while seeds of others may have been
washed in by the river. The east edge is also an area of major disturbance
from the widening of Hagadorn Road, likewise involving fill. It too has a rich
weed flora, but the number of species is not so great as on the west side. The
south edge of the woods also has its distinctive weed flora, differing consid¬
erably from the east and west edges. Major habitat disturbance has been less
on the south side because the forest was simply chopped down to make room
for dormitories and little if any fill was added. Some of the more character¬
istic species of the south edge are Conyza canadensis, Helianthus decapetalus,
Phytolacca americana, and Polygonum convolvulus .

The permanency of many of the edge species is doubtful, since they will
probably be replaced by other species, but many of them are hardy enough
to persist somewhere around the woods. A few introduced species have even
managed to gain roothold inside the woods, as for example Arctium minus,
Asparagus officinalis, Berberis thunbergii, Hemerocallis fulva, Hesperis
matronalis, Lonicera tatarica, Lysimachia nummularia, and Myosotis scorpio-
ides. If such species become more abundant they will signal decline in the
quality of Sanford as a natural area.

RARE SPECIES

Among the rare native species of south-central Michigan which now
occur or once occurred in Sanford Natural Area, the following 30 are enumer¬
ated. Destruction of natural habitats through urbanization is the major reason
these plants are becoming scarce. Some of the species listed are less common
than others, and from the checklist still others could be added. Those cited
below are at least illustrative of the richness and diversity in Sanford of
relatively uncommon native woodland plants.

Aplectrum hyemale
Arisaema dracontium
Athyrium pycnocarpon
Carex molesta
Corallorhiza maculata
Cy stop tens bulbifera
Dirca palustris
Dryopteris goldiana
Erythronium albidum
Euonymus atropurpureus
Festuca obtusa
Fraxinus quadrangulata
Gymnocladus dioica
Hybanthus concolor
Hydrastis canadensis

Juglans cinerea
Lilium michiganense
Liriodendron tulipifera
Matteucia struthiopteris
Morns rubra
Oryzopsis racemosa
Panax quinquefolius
Poa alsodes
P. nemoralis
P. palustris
Polymnia canadensis
Saururus cernuus
Stachys tenuifolia
Ulmus thomasii
Viburnum recognitum

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Vol. 9

SPECIES LOST FROM THE FLORA

The following nine species formerly occurred in Sanford Natural Area,
but apparently are not there at the present time.

Caltha palustris

Cornus stolonifera

Euonymus atropurpureus

Gymnocladus dioica

Ilex verticil lata

Five of these, Caltha palustris, Cornus stolonifera, Ilex verticillata, Larix
laricina, and Toxicodendron vernix, grew in the swampy area where Holmes
Hall was constructed. The only Gymnocladus dioica had already been cut
down by 1896, and timber-cutting activities of the Forestry Department may
have taken all Liriodendron tulipifera. The basis for the demise of the two
other species is less certain. If early records of the flora were more complete
it might be found that still other species have been lost, but the percentage
probably would be low.

One of the reasons for the very large number of species now in Sanford
is that many introduced species have been added to its flora, and most of the
natives have persisted. We are currently faced with the problem of preventing
further environmental deterioration which would increase the number of
weeds and eliminate the precariously small populations of native species of
limited ecological tolerance.

FUTURE PROSPECTS

In 1965 a group of biologists and campus planners met to discuss the
biological values of Sanford Natural Area and to set up guidelines for its
utilization. Among matters considered were relocation of walks and trails,
fencing, signs and maps, disposal of dead timber, pesticides, a planting pro¬
gram, a formal dedication, and an inventory of the biota. Some recommenda¬
tions of this group have already been implemented. One noteworthy guideline
is that pesticides be completely banned on the grounds that such chemicals
are incompatible with the concept of a natural area. The suggested planting
program has been followed to some extent, although this also seems incom¬
patible with the purposes of the Area. The present study provides an in¬
ventory of one aspect of the biota, but the other major taxonomic groups
remain to be treated.

Maintaining Sanford as a natural area is the principal challenge facing
those concerned with its future. It is not now entirely “natural” and can
never be completely so. Yet it is an irreplaceable resource worth every effort
and expense to preserve. An imminent danger is the foot traffic. More than
7,000 students live in the neighboring dormitories, not to mention all those in
the mass of apartments on the north side of the river. In 1957 President John
A. Hannah noted (letter to Dr. W. B. Drew, Chairman, Department of Botany
and Plant Pathology) that “in the not too distant future ... it will be so
tramped over as to make it fail completely to serve your students as a typical
wood lot situation.” This grim prediction may soon be fulfilled, for even

Larix laricina
Liriodendron tulipifera
Quercus velutina
Toxicodendron vernix

1970

THE MICHIGAN BOTANIST

137

though ample trails are provided new ones are constantly being formed. Dr. G.
W. Parmelee (letter to State News ) describes this process as pounding “the
spongy forest floor with its wealth of delicate forest herbs and tree seedlings
into barren ‘earth-crete’.”

The concentration of people has already made it impossible for certain
studies to be effectively carried out in the Area. For example, disturbance of
stones and fallen trees makes it unsatisfactory for courses in cold-blooded
vertebrates. Experimental ecological studies are impossible because of distur¬
bance and vandalism. According to Dr. G. J. Wallace it is no longer very good
for ornithological field trips because such species as grackles and starlings have
replaced nesting Louisiana water thrushes and oven birds. One indication of
wanton disregard for the Area is the numerous beer bottles, cans, and food
wrappers scattered about.

Several natural factors, however, contribute to the preservation of the
woodlot. During much of the winter the soil is frozen, thus helping prevent
soil compaction. In the spring, when the wild flowers are most attractive,
many of the days are cold, windy, or wet, discouraging the casual wanderer.
By late spring mosquitos are out in full force and they remain a potent
defensive army for much of the summer. In spring, summer, and fall there is
enough poison ivy near the river to limit trampling by the less naive, and
during the summer wood-nettle stands guard at many moister sites.

A very substantial fence, with an entrance only on the west side, would
seem the best solution for reducing unnecessary traffic. The present low fence
with six gates (four on the south side) is entirely ineffective. Some of the new
trails have actually developed from the gate locations. On the north side the
river is an excellent barrier, but frozen it becomes a thoroughfare between the
campus and the apartments and Grand River Avenue shopping district. In
1895 the College invested in a fence on private property along the north side
of the river to prevent cattle from entering the woodlot across the frozen
river. Such a fence for people now seems impossible, but one along the south
and east boundaries would at least discourage use of the woodlot as a short¬
cut and sidewalk.

It is also urgent that there be a vigorous educational program for the
public, especially the academic community, concerning the values of Sanford
Natural Area and the reasons it is seriously endangered. A fence would help in
the educational process, but also needed are frequent press reports, more
intensive supervision of the Area, adequate signs, and an attractive descriptive
brochure. If these measures fail to halt deterioration, closing the woodlot to
all unauthorized persons could be justified on grounds of its formal instruc¬
tional and research values.

Whatever the future holds for Sanford Natural Area, it will be instruc¬
tive as to the possibilities for coexistence of nature’s handiwork and thousands
of university students. It has already served the community well for over a
century. If proper protective measures are taken, I believe it can continue to
benefit future generations of students and faculty, illustrating the biological
principles of a natural area.

138

THE MICHIGAN BOTANIST

Vol. 9

SUMMARY

Sanford Natural Area is a 35-acre woodlot on the Michigan State University
campus which has been used for teaching and research for over a century. For 65 years
it was managed as a demonstration of good woodlot practices for continuous and lasting
crops; maple syrup production was a major research subject. In 1959 administrative
responsibility for the Area was shifted from the Forestry Department to Campus Park
and Planning. With the change in administration came a redefinition of purpose: It now
serves to illustrate the biological principles of a natural area. Nearly 1500 students in 21
University courses in eight departments use it each year as an outdoor laboratory. It is
most conveniently located for teaching of all wooded areas on campus, and is perhaps the
botanically most varied of all woodlots in the vicinity of East Lansing. The large flora of
358 species of vascular plants reflects the diversity of habitats in the Area. Ten major
habitat types are recognized. The dominant tree is Acer saccharum, but nine other tree
species are important components of the forest. Among the 81 native woody species
present 56 are trees. The larger trees are more than 30 inches d.b.h., over 100 feet high,
and many of them are probably more than 200 years old. At least 30 species in the flora
are rare in central Michigan. Nine species formerly present have been lost, in large part
because a major habitat was destroyed by building construction. Expansion of the Uni¬
versity in the past 15 years has resulted in 20 acres formerly occupied by the woodlot
being utilized for four large dormitories. Increased foot traffic from more than 7,000
students in these and other nearby residents units and from the rest of the enlarging
community threatens to obliterate many natural features of the Area. Adequate fencing
and a vigorous educational program appear to offer the most effective means for preserv¬
ing this endangered natural environment.

ACKNOWLEDGMENTS

Students of Michigan State University have made this study possible, and it is to
them that I am particularly indebted. Their role is somewhat evident from the specimens
cited in the checklist to be published as part II. Those who have made major contribu¬
tions include Dr. Norton G. Miller of Harvard University, Dr. William T. Gillis of Fair-
child Tropical Garden, Jerold L. Grashoff of the University of Texas, Larry E. Morse,
Joan M. Hubbard, Martin A. Bordner, Richard R. Nelson, Thomas J. Popma, and Basil G.
Stergios. Miss Darlene M. Valasek has been most helpful in attending to many details in
the final stages of the project. A fine series of collections was made by Dr. L. C.
Anderson of Kansas State University when he was on our faculty in 1962-63. Drs. G. W.
Parmelee and G. Schneider have loaned records of the Division of Campus Park and
Planning and the Department of Forestry, respectively, and have contributed otherwise to
the study, especially by critically reading the manuscript. Dr. E. P. Whiteside examined
the soils in the field, revised the nomenclature on the original soils map of the Area, and
provided other information on the soils. Dr. S. N. Stephenson has provided advice on
ecological procedures and helped in the identification of the Gramineae. Dr. E. G. Voss
identified several species, especially Car ex, and Dr. W. H. Wagner, Jr., checked the fern
identifications. The cost of additional pages and other expenses of the study have been
borne by the Department of Botany and Plant Pathology.

LITERATURE CITED

Anonymous. 1953. Fifty Years of Forestry at Michigan State College. 32 pp.

Cox, G. W. 1967. Laboratory Manual of General Ecology. Wm. G. Brown, Inc. Dubuque,
Iowa. 165 pp.

Crozier, A. A. 1896. The College wood lots. The M.A.C. Record 1 (no. 8, March 3): 1.

- . 1897. Pp. 32-33 in C. D. Smith. Report of the Department of Practical

Agriculture, and Superintendent of the Farm. 35th Ann. Rep. Sec. State Board Agr.
Mich.

1970

THE MICHIGAN BOTANIST

139

litis, H. H. 1968. The optimum human environment and its relation to modern agri¬
cultural preoccupations. Biologist 50: 114-125.

NCR. 3 Committee. 1960. (North Central Regional Technical Committee on Soil Survey
(NCR-3), June, 1960.) Soils of the North Central Region of the United States (North
Central Regional Publication 76). Bull. 544. Agr. Exp. Sta. Univ. Wisconsin, Madison,
Wis.

Schneider, G. 1966. A Twenty -year Ecological Investigation in a Relatively Undisturbed
Sugar Maple-Beech Stand in Southern Michigan. Mich. Agr. Exp. Sta. Res. Bull. 15. 61

pp.

Smith, C. D. 1896. Pp. 35-36 in Report of the Department of Practical Agriculture. 34th
Ann. Rep. Sec. State Board Agr. Mich.

Soil Survey Staff, U.S.D.A. 1960. Soil Classification. A comprehensive system. 7th ap¬
proximation, as amended March, 1967, July, 1968, and April, 1969.

Veatch, J. O., et al. 1941. Soil Survey of Ingham County, Michigan. U.S.D.A. Bur. PI.
Ind. Ser. 1933, no. 36. 43 pp. + map.

Walline, E. 1940. Management plan for the River Woodlot. Manuscript, Mich. State Univ.
Forestry Department. 24 pp.

Westveld, R. H. 1931. Management plan for River Woodlot. Manuscript, Mich. State Univ.
Forestry Department. 17 pp.

Wilcox, A. T. 1949. Recreation management in the Sanford Woodlot. Manuscript, Mich.
State Univ. Forestry Department. 2 pp.

MOSSES NEW TO THE FLORA OF MICHIGAN1

Howard Crum

Herbarium, University of Michigan, Ann Arbor

As a result of continuing field and herbarium studies, seven species of
mosses can be recorded as new to the known flora of Michigan, as follows:

Sphagnum tenerum Sull. & Lesq. ex Sull. Cheboygan Co.: In a cedar
swamp, in the open, 3A mile northeast of Pine Grove State Forest Camp¬
ground, T33N, R1W, sect. 1 6, Richard C. Harris 788 (MICH, MSC). Chippewa
Co.: In a spruce-jack pine area, Dick State Forest Campground, T44N, R5W,
sect. 2, Harris 1153 (MSC), Clare Co.: In a Thuja swamp west of Farwell,
T17N, R5W, sect. 28, Harris 992B (MICH, MSC); in a Thuja swamp, Silver
Lake, T18N, R5W, sect. 30, Harris 1011 (MICH, MSC). Eaton Co.: In a
Vaccinium bog, Kikendall farm, near Dimondale, TIN, R3W, Harris 893
(MSC). Ingham Co.: In an old tamarack-leatherleaf bog, Holt Bog, T3N, R2W,
sect. 21, Harris 874 (MICH, MSC), 882 (MSC). Mackinac Co.: On a log, near a
Thuja , Lake Huron shore west of Hessel, T42N, R2W, sect. 22, Harris 1095

^Contribution from the Herbarium of the University of Michigan and also
from the University of Michigan Biological Station. Grateful acknowledgment is made for
the National Science Foundation for research funds provided under grant GB6095.

140

THE MICHIGAN BOTANIST

Vol. 9

(MSC); Prentiss Bay, J. H. Ehlers, July 30, 1920 (MSC). Otsego Co.: In a
Thuja- spruce-alder swamp on Whitehouse Road, 0.7 miles north of Lake Shore
Drive, H. Crum, Aug. 7, 1969 (MICH). Roscommon Co.: At edge of muck,
without locality, J. Caution & W. Gillis, July 1956 (MSC). — S. tenerum is
quite variable in its overall appearance but most commonly resembles S.
capillaceum (Weiss) Schrank, to which it is obviously related. It differs from
that species essentially in the “hemi-isophyllous” nature of the stem leaves,
which approach the branch leaves to some extent in shape (being somewhat
pointed and concave at the tip and not very strongly bordered at the base)
and in structure (having hyaline cells fibrillose nearly throughout the leaf and
porose on both surfaces). The stem leaves of S. capillaceum also show some
degree of hemi-isophylly, but the fibrils are not so well or so extensively
developed, and the pores, if present at all, are few, large, rounded, and re¬
stricted to the inner surface. In S. tenerum, by contrast, there are three to
five large, rounded pores on the inner surface of each cell and more numerous
elliptic pores along the commissures on the other surface.

The species occurs in bog forests and also in more open situations, such
as blueberry or tamarack -leatherleaf bogs from Newfoundland to Florida and
less commonly inland in scattered localities in Quebec, Ontario, Michigan,
Missouri, and, very disjunctively, in British Columbia.

Fissidens subbasilaris Hedw. Washtenaw Co.: On an old, rotten tree
trunk, woods near Ypsilanti, H. T. Darlington, June 9, 1928 (MSC,
MICH). — This species is included in Darlington’s Mosses of Michigan, but inso¬
far as I have been able to ascertain, on the basis of misidentifications. It is
easily recognized with a microscope or even with a lens because the upper
portion of the costa is obscured by a covering layer of short, green cells.
Though scattered and rare in the North, it is a widely distributed species
throughout eastern North America from Maine and southern Ontario west to
Kansas and south to northeastern Mexico and Florida. It grows on the bark of
trees and sometimes on rotten wood or rock.

Grimmia plagiopodia Hedw. Cheboygan Co.: On a calcareous sandstone
boulder in the railroad right-of-way, in the complete shade of a young spruce
(and covered by its lower boughs), Mill Creek, ca. 3 miles southeast of Mack¬
inaw City on U. S. Highway 23, H. Crum, June 25, 1969 (MICH, UMBS). — A
rare and interesting species in eastern North America, G. plagiopodia greatly
resembles the ubiquitous G. apocarpa because of its brownish leaves ending in
hyaline hair-points, but it is actually very distinctive because of its wide¬
mouthed capsules bulging on one side and thus causing the short, curved or
even curled setae to seem excentric in attachment. The presence of a
peristome is an important distinction between G. plagiopodia and G. anodon
BSG. The plants are typically found on sandstone but sometimes occur on
limestone or concrete. The species is distributed in Europe, the Caucasus, and
central Asia, Alaska to California and New Mexico, and in a few eastern
localities from Ontario and the Dakotas south to Wisconsin, Ohio, and Mary¬
land.

1970

THE MICHIGAN BOTANIST

141

Philonotis capillaris Lindb ex C. J. Hartm. Chippewa Co.: On soil of a
roadside ditch bank, about 3 miles northwest of Homestead, Sugar Island, H.
Crum , July 1969 (MICH). — A species of circumpolar distribution, reputedly
widespread in the East from Maine to Virginia and also in the West from
Alaska to Arizona and New Mexico. Reports in the literature are to be viewed
with skepticism, however, as this and other species of Philonotis are more
often misidentified than not. From specimens readily available to me, I can
confirm the American distribution as including Arkansas, Michigan, New York,
British Columbia, Washington, and California. I have also seen material from
Germany.

Orthotrichum alpestre Hornsch. ex BSG. Keweenaw Co.: Mt. Bohemia,
A. J. Sharp, Sept. 1, 1937 (TENN). — This species, common on the calcareous
rocks so prevalent in the western mountains from Yukon to Arizona, is not
otherwise known from eastern North America. The substrate is not given for
the Michigan collection, but it was probably some acid type of rock. The
species also occurs in northern and central portions of Eurasia. I am indebted
to Dale H. Vitt for calling my attention to this significant range extension,
which we have both confirmed.

Brachythecium digastrum C. M. & Kindb. ex Mac. & Kindb. Ingham
Co.: Woodlot 17, Second College Woods, East Lansing, H. T. Darlington, May
1, 1934 (MSC, MICH). — This species was reported from Clinton and Gratiot
Counties in Darlington’s Mosses of Michigan, but I have not seen any correctly
named material to validate the records. Brachythecium digastrum is somewhat
similar to the common B. oxycladon (Brid.) Jaeg. & Sauerb., as evidenced by
a similar areolation at the basal angles of the leaves, but the stem leaves are
rather broadly decurrent, especially in forms with widely spaced leaves; the
branches are more terete-foliate; and the branch leaves are tapered to short,
bluntly acute, serrate points, with apical cells noticeably shorter than the
median and the costa somewhat serrulate at back near the tip. I have seen
specimens from a broad range in eastern North America (Arkansas, Michigan,
North Carolina, Ontario, Quebec, and Virginia). The species was described on
the basis of collections from Ontario and New Brunswick.

Isopterygium subfalcatum (Aust.) Jaeg. & Sauerb. Eaton Co.: On moist
earth in open woods, west of Fitzgerald Park, south shore of Grand River just
northwest of Grand Ledge,//. T. Darlington, May 2, 1951 (CAN). Chippewa
Co.: On rock in woods, Lower Falls of the Tahquamenon River, R. R. Ireland
4346 (CAN). — A rare species growing on moist siliceous cliffs or sometimes
on the peaty soil of banks, /. subfalcatum occurs disjunctively over a broad
range from Maine to Michigan and south to Arkansas and North Carolina.
Taxiphyllum howellianum Crum & Anders., described not too long ago from
North Carolina collections, belongs in the synonymy of this species. I am
grateful to Robert R. Ireland for supplying the determinations for both speci¬
mens cited here, as well as for clarifying the nomenclature.

142

THE MICHIGAN BOTANIST

Vol. 9

Review

HOW TO KNOW THE AQUATIC PLANTS. By G. W. Prescott. Wm. C. Brown Co.,
Dubuque. 1969. 171 pp. $2.50 (spiral); $3.25 (cloth).

The chief value of this book is its particular scope. The geographic area covered is
the entire U. S. and the definition of aquatic is broad (unlike Muenscher’s manual for the
U. S.), even broader than in Fassett’s northeastern manual, for some marine angiosperms
and bog plants are included. The format is that of the familiar Pictured Key Nature
Series, but the key is only to the genera- about 165 of them-although some species are
mentioned. Beginners who for some reason want to know only the generic name of an
aquatic angiosperm or conspicuous cryptogam and who have a good deal of patience and
intuition with which to approach the rather casual key of 301 couplets may find the
volume helpful.

The key is commendable in including many genera at more than one place, so that
variability of either Nature or interpretation need not lead to error in identification. But
such error is still likely to result from frequent failure of the key to use the most obvious
distinguishing characters or to contrast clearly the characters used. There is a great excess
of the utterly uninformative “Plant otherwise”; after sailing through five consecutive
leads like this and failing to arrive at Elatine, I had to conclude that this genus (strictly
submersed in Michigan) was to be reached only by lead 62b in this couplet, most of
which does not contrast, leaving one helpless with a rooted submersed plant:

62a. Plants submersed and essentially rooted to the bottom, sometimes at 8 M (often
becoming free-floating); with or without floating leaves.

62b. Plants rooted on shore or in shallow water, emergent, sprawling or growing erect
at margins of aquatic habitats.

Couplet 242 will not distinguish Eleocharis (which has a sheath at the base of each
slender stem) from Scirpus. The flowers of Ruppia are not “in a simple umbel” nor are
the leaves “mostly basal” (except perhaps when stranded); this normally submersed
species with extremely long alternate leaves is said to grow “on saline or alkaline shores.”
Littorella, in contrast, is said to grow “submersed,” although the figure shows it flower¬
ing, a condition occurring only on shores. I cannot imagine anyone finding Littorella
“difficult to differentiate” from Ruppia, the former having basal rosettes of short
leaves-which, in Michigan at least, are not “dark green” or “with a broad sheathing
base,” but a distinctive light green, terete, and no broader at base than the middle. The
whole group of small rosette-formers of our softwater lakes (e.g., Littorella, Eriocaulon,
Lobelia dortmanna, Isoetes, Subularia) fares very badly in this key, which makes these
distinctive plants excessively difficult to compare.

The main discussion of Potamogeton, with figures of several species, follows coup¬
lets 58 and 59 which both call for opposite leaves. At couplet 284, under alternate
leaves, there is merely a reference back to this point. Couplet 116 distinguishes Rorippa
(“Flowers greenish white”) from Neobeckia, Cardamine, and Nasturtium (“Flowers white,
yellow or lavender”). In fact, the latter three are not yellow-flowered, while Rorippa
always is. In referring to R. aquatica “with whitish flowers,” the author apparently is
unaware that this name is synomymous with his Neobeckia (Armoracia) aquatica, intend¬
ed to run under the alternate lead! Submersed plants of Megalodonta “may be confused
with some species of Ranunculus or Neobeckia” —and indeed they may when one fails to
mention that the latter two differ most obviously in having alternate leaves.

The glossary combined with the index is frequently weak as well as incomplete.
Alternate is an arrangement, not “leaves . . .”; phyllodia is a plural word; “cilliate” is not
the correct spelling. The Introduction concludes with a list of 15 divisions of the plant
kingdom (9 of which are algae), immediately after which is the statement: “When all the
aquatic plant species are placed in review it is noteworthy that a majority are in the
Monocotyledonae of the seed plants.” That the algologist author could let such an aston¬
ishing assertion slip by is perhaps indicative of the care with which this volume seems to
have been written.

-E. G. V.

Program Notes

Michigan Botanical Club and chapter members will be receiving further information
about these major events:

April 13: Huron Valley Chapter, Annual Meeting at the University of Michigan Botanical
Gardens. Dr. Rogers McVaugh will talk on “Botanical Activities of the Geological
Survey of Michigan, 1837-1840,” relating the story of the early collections made
under the supervision of Douglass Houghton.

May 29-31: Michigan Botanical Club, Annual Spring Campout, at the University of Michi¬
gan Biological Station on Douglas Lake. A full program of field trips is expected,
including all-day ones to the Upper Peninsula limestone country for spring flora and
rare ferns, led by W. H. Wagner; to Trail’s End Bay and Wilderness State Park, led by
Ed Voss; and to Bois Blanc Island, led by Morgan Poole.

June 28: Southeastern Chapter, Annual Meeting, at the Drayton Plains Nature Center.

September 25-27: Southeastern Chapter, Fall Campout, at Yankee Springs.

Editorial Notes

Errata in Vol. 9, No. 1 (January 1970):

p. 16, line 4 from bottom, insert “200 pp.”

line 8 from bottom, for “3” read “2”
p. 61, line 17 from bottom, insert reference: “Brittonia 21: 196-200”

Authors of articles in recent issues who wish to have manuscripts and/or art work
returned should advise the editor immediately, before another “housecleaning” of his files.

The January number (Vol. 9, No. 1) was mailed January 22, 1970.

Publications of Interest

From time to time we call attention, under this heading, to new publications
which might be of particular interest to some of our readers but which, because of less
explicit geographic or botanical orientation, do not qualify for regular review or inclusion
in the listings of new literature relating to Michigan botany. All annotations are by the
Editor unless ascribed to another.

A SHORT HISTORY OF BOTANY IN THE UNITED STATES. Edited by Joseph Ewan.
Hafner Publishing Co., New York, 1969. 174 pp. $7.50. Distributed to all registrants
at the XI International Botanical Congress in Seattle last August, this volume offers a
general survey of American botanical history, with chapters by 13 authors on specific
areas of botany. One of the most interesting contributions is Ewan’s “Calendar of
Events,” 300 B.C. to 1968, wherein one may learn a fascinating series of historical
tidbits and highlights. Ewan also contributed a 23-page chapter on “Early History”
and a 9-page one on “Plant Geography.”

SPRING FLORA OF MINNESOTA Including Common Cultivated Plants. By Thomas
Morley. University of Minnesota Press, Minneapolis, 1969. 283 pp. $7.75. This is a
hardcover edition of the work published in 1966 by the Department of Botany and
enthusiastically noted in this journal in January, 1968. The same type has been used,
except for various corrections and minor revisions. One useful innovation is marking
with an asterisk the names of those genera for which all species native or naturalized
in Minnesota are included; thus, one immediately knows whether a genus includes
additional species which bloom after June 7. Perhaps the most conspicuous change is
the price, which has more than doubled, making this a very expensive binding!

MICHIGAN SCIENCE IN ACTION is the title of a new series of publications which
relates research developments at the Michigan Agricultural Experiment Station. Nine
numbers have been issued during the past year, and copies may be obtained by
writing to the MSU Bulletin Office, Box 231, East Lansing, Michigan 48823: 1. Focus
on Pests and Pesticides, 2. Sod and Turf- Michigan’s $350 Million Carpet, 3. Focus on
Food Science, 4. Focus on the Family, 5. Reshaping the Soil, 6. Focus on Beef
Cattle, 7. Bedding Plants: Michigan’s Blooming Industry, 8. Scientific Achievements:
1888-1969, 9. Focus on Dairy Nutrition.

CONTENTS

Chromosome Studies of Some Mosses of the Douglas Lake Region

Jerry A. Snider . 67

Review— How to Know the Lichens . 71

Investigations in the White Waterlilies ( Nymphaea ) of Michigan

Gary R. Williams . 72

Additional Bryophytes from Sinkholes in Alpena County, Michigan,

Including Orthotrichum pallens New to Eastern North America

Norton G. Miller & Dale H. Vitt . 87

Algae in Park Lake, Clinton County, Michigan

Norman A. Andresen . 95

Publications of Interest . 107, 143

Morphology of the Fungal Association in Three Species of
Corallorhiza in Michigan

Ella 0. Campbell . 108

Sphagnum in Charlevoix County, Michigan

William M. Zales & Howard Crum . 114

A Botanical Inventory of Sanford Natural Area.

I. The Environment

John H. Beaman . 116

Mosses New to the Flora of Michigan

Howard Crum . 139

Review— How to Know the Aquatic Plants . 142

Program Notes . 143

Editorial Notes . 143

(On the cover: A white waterlily, Nymphaea,
blooming at Cusino Lake, Schoolcraft County, Michigan.
Photograph by Gary R. Williams, August 15, 1969.

See p. 80.)

*vM

LIBRARY

h1 I

7 3

THE

MAY 20 1970

H«W YORK
BOTANICAL BANDEW

Vol. 9, No. 3

MICHIGAN BOTANIST

May, 1970

THE MICHIGAN BOTANIST-Published by the Michigan Botanical Club four times per

year: January, March, May, and October. Second-class postage paid at Ann Arbor,

Michigan.

Subscriptions: $3.00 per year. Single copies: $.75.

All back issues are available, at the following prices:

Vol. 1 (2 numbers, at $1.00 each): $2.00

Vol. 2-5 (4 numbers, at $.50 each): $2.00 (per volume)

Vol. 6- (4 numbers, at $.75 each): $3.00 (per volume)

Subscriptions (from those not members of the Michigan Botanical Club) and all orders
for back issues should be addressed to the business and circulation manager, who will as¬
sume that subscriptions received prior to the last number of a volume are intended to begin
with the first number of that volume unless specified to the contrary.

Articles and notes dealing with any phase of botany relating to the Upper Great Lakes
Region may be sent to the editor in chief. The attention of authors preparing manuscripts
is called to “Information for Authors” (Vol. 9, p. 62; copies available from the editor).

Editorial Board

Edward G. Voss, Editor in Chief

Herbarium, The University of Michigan, Ann Arbor, Michigan 48104

Laura T. Roberts, Business and Circulation Manager
1509 Kearney Rd., Ann Arbor, Michigan 48104

Virginia L. Bailey Rogers McVaugh

Burton V. Barnes Alexander H. Smith

Richard Brewer W. H. Wagner, Jr.

THE MICHIGAN BOTANICAL CLUB

Membership in the Michigan Botanical Club is open to anyone interested in its aims:
conservation of all native plants; education of the public to appreciate and preserve plant
life; sponsorship of research and publication on the plant life of the State; sponsorship of
legislation to promote the preservation of Michigan native flora and to establish suitable
sanctuaries and natural areas; and cooperation in programs concerned with the wise use and
conservation of all natural resources and scenic features.

Dues are modest, but vary slightly among the chapters and with different classes of
membership. Persons desiring to become state members (not affiliated with a local chapter)
may send $3.00 dues to the Membership Chairman listed below. In all cases, dues include
subscription to THE MICHIGAN BOTANIST. (Persons and institutions desiring to sub¬
scribe without becoming members should deal with the business and circulation manager as
stated at the top of this page.)

President: W. H. Wagner, Jr., Botanical Gardens, University of Michigan, Ann Arbor,
Michigan 48105

Vice President: William F. Hopkins, 6210 Cowell Rd., Brighton, Michigan 48116
Recording Secretary: Mary Cooley, 703 S. forest. Ann Arbor, Michigan 48104
Corresponding Sec.: Thelma Thomson, 15093 Faust, Detroit, Michigan 48223
Treasurer: Charles Buswell, 19204 Plainview, Detroit, Michigan 48219
Membership Chairman: Harriette V. Bartoo, Department of Biology, Western Michigan Uni¬
versity, Kalamazoo, Michigan 49001
Corresponding Secretary, Southeastern Chapter:

Catherine Hamilton, 9591 Grandmont, Detroit, Michigan 48227
Secretary, Huron Valley Chapter:

Ursula R. Freimarck, 704 Madison PI., Ann Arbor, Michigan 48103
Secretary, Southwestern Chapter:

Helen Wiles, 7113 N 25th St., Rt. 2, Kalamazoo, Michigan 49004

1970

THE MICHIGAN BOTANIST

147

A BOTANICAL INVENTORY OF SANFORD NATURAL AREA.
II. CHECKLIST OF VASCULAR PLANTS

John H. Beaman

Beal-Darlington Herbarium, Department of Botany and Plant Pathology

Michigan State University, East Lansing

A general account of the environment of Sanford Natural Area was
published previously (Beaman, 1970). The following checklist includes vascular
plants which now occur in the woodlot and a few others which are no longer
present. Only spontaneous plants are included, with the exception of three
species which have persisted from planting in 1896. Collections cited are de¬
posited in the Beal-Darlington Herbarium of Michigan State University. A large
number of specimens collected in the 1890’s labeled “College Woods” are
most likely from Sanford, but are not cited because of the possibility that
they are from a different woodlot. All collections cited have been obtained
within the past 30 years, most since 1960.

Flowering dates of the angiosperms are based on specimens in the Beal-
Darlington Herbarium from the southern half of the Lower Peninsula of Mich¬
igan. Many species, especially those of the spring flora, are evident for a very
short period. The dates indicated should help suggest to the student what
species to expect at a particular time of year. Phenological records also have
other uses as in studies of productivity and ecosystem analysis. Optimally they
would include aspects of plant development throughout the year, but a com¬
monly recorded feature is date of first flowering (e.g. Taylor, 1969). Such
data are not available for the species of Sanford Natural Area nor for most
plants in Michigan.

The systematic arrangement is as follows: Horsetail, fern, and gymno-
sperm higher taxa, Cronquist et al. (1966); fern and gymnosperm families,
Melchior & Werdermann (1954); angiosperms, Cronquist (1968); genera and
species of each family, alphabetical.

The checklist includes 90 families, 230 genera, and 355 1 species. Of this
number, 279 species are native, 73 introduced, and three planted. It is note¬
worthy that 60% of the 151 vascular plant families in Michigan occur in
Sanford Natural Area. Well over 10% of the species of vascular plants in the
Michigan flora are present. The largest families are the Gramineae with 21
genera and 33 species and the Compositae with 19 genera and 33 species. The
largest genus is Carex with 16 species. Eight other genera, Quercus, Viola ,
Salix, Cornus, Acer, Viburnum, Solidago, and Poa, each have five or more
species.

^In part I the number of species was reported as 358, but revised determinations
reduce the total by three.

148

THE MICHIGAN BOTANIST

Vol. 9

EQUISETOPHYTA (Horsetails)

EQUISETACEAE (Horsetail Family)

Equisetum arvense L. Horsetail. Lauff 9. Along the river.

E. hyemale L. Scouring rush. Grashoff 307. On riverbank at NW corner.

POLYPODIOPHYT A (Ferns)

OPHIOGLOSSACEAE (Adder’s-tongue Family)

Botrychium virginianum (L.) Sw. Rattlesnake fern. Hutnphrey 204; Throop 6. Scat¬
tered NE of sugar house site. Rare.

OSMUNDACEAE (Royal Fern Family)

Osmunda regalis L. Royal fern. Nelson 153. Middle portion near river.

POLYPODIACEAE (Polypody Family)

Adiantum pedatum L. Maidenhair fern. Humphrey 201; Nelson 188. Scattered NW of
sugar house site in moist areas.

Athyrium asplenioides (Michx.) Desv. Lady fern. Nelson 152. Middle portion near
river.

A. pycnocarpon (Spreng.) Tidest. Narrow-leaved spleenwort. Ciarkowski 56; Miller
4525. E-central part in rich moist soil.

A. thelypterioides (Michx.) Desv. Silvery spleenwort. Nelson 199. In wet soil in cen¬
tral part.

Cystopteris bulbifera (L.) Bernh. Bulblet fern. Hymphrey 203; Nelson 107. On river-
bank in NW section.

C. fragilis (L.) Bernh. Fragile fern. Bordner 171. S bank of river.

Drvopteris goldiana (Hook.) Gray. Goldie’s fern. Humphrey 205; Miller 4524. Central
and E part in moist soil.

D. spinulosa (O. F. Muell.) Watt. Spinulose shield-fern. Bordner 174; Nelson 163. SE
and middle part. Bordner 174 appears to be of hybrid origin.

Matteuccia struthiopteris (L.) Todaro. Ostrich fern. Nelson 154, 199, 251. Middle
portion near river.

Onoclea sensibilis L. Sensitive fern. Marshall 284; Nelson 200, 202. Central and E part
in wet soil.

Polystichum acrostichoides (Michx.) Schott. Christmas fern. Humphrey 200; Nelson
160. N-central area.

Thelypteris hexagonoptera (Michx.) Weatherby. Broad beech-fern. Nelson 103. near
middle of woods. Rare.

T. palustris var. pubescens (Lawson) Fern. Marsh fern. Valasek 439. By intermittent
lagoon in N-central part.

PINOPHYTA (Gymnosperms)

PINACEAE (Pine Family)

Larix laricina (DuRoi) K. Koch. Tamarack. Noted in 1895 in marshy area on SE edge,
the present site of Holmes Hall.

Tsuga canadensis (Endl.) Carr. Hemlock. Nelson 109. Planted specimens in NW quarter
on high ground above riverbank.

CUPRESSACEAE (Cypress Family)

Thuja occidentalis L. White cedar or arbor vitae. Nelson 304. NW portion on river¬
bank.

1970

THE MICHIGAN BOTANIST

149

MAGNOLIOPHYTA (Angiosperms)

MAGNOLIATAE (Dicotyledons)

MAGNOLIIDAE

MAGNOLIACEAE (Magnolia Family)

Liriodendron tulipifera L. Tulip-poplar. Management records indicate that this species
was once present, but specimens have not been located. Flowering in June.

ANNONACEAE (Custard-apple Family)

Asimina triloba (L.) Dunal. Pawpaw. Grashoff 298. Known only from one small tree
on the N bank of the river, which succumbed to apartment construction in the fall
of 1968. Flowering in May.

LAURACEAE (Laurel Family)

Lindera benzoin (L.) Blume. Spicebush. Hubbard 7; Marshall in 1949. Common in wet
areas. Flowering in April.

Sassafras albidum (Nutt.) Nees. Sassafras. Grashoff 320. Infrequent in N-central por¬
tion near the river. Flowering in May.

SAURURACEAE (Lizard’s-tail Family)

Saururus cernuus L. Lizard’s-tail. Anderson 2162; Lauff in 1950. Abundant on river
margin. Flowering June - August.

ARISTOLOCHIACEAE (Birthwort Family)

Asarum canadense L. Wild ginger. Anderson 2253 ; Ellsworth in 1949; Hubbard 25.
Patches scattered in low areas. Flowering April - June.

CERATOPHYLLACEAE (Hornwort Family)

Ceratophyllum demersum L. Hornwort. Nelson 299. Submerged aquatic collected in
the river only near Bogue Street bridge, but probably more abundant. Flowering
July - August.

RANUNCULACEAE (Crowfoot Family)

Actaea pachypoda Ell. White baneberry. Halicki in 1957; Hubbard 44. Scattered
through the woods. Flowering May - June.

Anemone quinquefolia var. interior Fern. Wood anemone. Anderson 2257; Grashoff
283; Hubbard 17. Uncommon. Flowering April - May.

Caltha palustris L. Marsh-marigold. Anderson 2230. The species was once abundant in
the marshy area on the SE side. It was exterminated by construction of Holmes
Hall. Flowering April - June.

Hepatica acutiloba DC. Liverleaf. Anderson 2217; Hubbard 10; Marshall 1517. Fre¬
quent throughout much of the woods. Flowering April - May.

Hydrastis canadensis L. Golden-seal. Grashoff 300. In central part, rare. Flowering in
May.

Isopyrum biternatum (Raf.) T. & G. False rue-anemone. Hubbard 35; Stergios 17.
Infrequent. Flowering April - May.

Ranunculus abortivus L. Small-flowered crowfoot. Grashoff 279; Hubbard 34. Occa¬
sional in disturbed situations. Flowering May - June.

R. pensylvanicus LT. Bristly crowfoot. Grashoff 31. On riverbank, apparently rare.
Flowering in July.

R. recurvatus Poir. Grashoff 315. Apparently rare. Flowering May - June.

R. septentrionalis Poir. Swamp buttercup. Hubbard 24. Common by vernal pools in
E-central portion. Flowering April - June.

Thalictrum dasycarpum Fisch. & Lall. Purple meadow-rue. Nelson 108. On riverbank;
rare. Flowering June - July.

150

THE MICHIGAN BOTANIST

Vol. 9

T. dioicum L. Early meadow-rue. Grashoff 278; Stergios 8, 14. Scattered in low areas.
Flowering April - May.

BERBERIDACEAE (Barberry Family)

Berberis thunbergii DC. Japanese barberry. Grashoff 296; Stergios 7. Introduced; scat¬
tered through the woods. Flowering April - May.

Caulophyllum thalictroides (L.) Michx. Blue cohosh. Anderson 2256; Hubbard 27.
Common. Flowering April - May. (See cover photo.)

Podophyllum peltatum L. May-apple. Hubbard 45. Several scattered patches. Flower¬
ing in May.

MENISPERMACEAE (Moonseed Family)

Menispermum canadense L. Moonseed. Grashoff 311. Common near the river. Flower¬
ing June - July.

PAPAVERACEAE (Poppy Family)

Sanguinaria canadensis L. Bloodroot. Gillis 2886; Hubbard 16. Scattered through the
woods. Flowering April - May.

FUMARIACEAE (Fumatory Family)

Dicentra canadensis (Goldie) Walp. Squirrel-corn. Anderson 2239; Hubbard 30;
Marshall 293. Frequent in patches. Flowering April - May.

D. cucullaria (L.) Bernh. Dutchman’s breeches. Anderson 2240; Grashoff 281; Stergios
25. Infrequent in a few patches. Flowering April - May.

HAMAMELIDAE

PLATANACEAE (Plane-tree Family)

Platanus occidentalis L. Sycamore. Valasek 430. Floodplain in N-central part. Flower¬
ing in May.

HAMAMELIDACEAE (Witch-hazel Family)

Hamamelis virginiana L. Witch-hazel. Anderson 2286; Hubbard 4; Popma 332. Fre¬
quent near the river. Flowering September - October.

ULMACEAE (Elm Family)

Celtis occidentalis L. Hackberry. Nelson 161. Rare tree near the river. Flowering in
May.

Ulmus americana L. American elm. Horning 16. Formerly one of the dominants, but
most large trees killed by Dutch elm disease during the 1960’s. Flowering in April.

U. rubra Muhl. Slippery or red elm. Nelson 309. Near the river. Flowering April -
May.

U. thomasii Sarg. Rock elm. Apparently frequent in 1895, but only one tree (13
inches d.b.h.) on the W side just east of the ravine has been located recently. A
specimen has not yet been prepared. Flowering April - May.

MORACEAE (Mulberry Family)

Morus alba L. White mulberry. Nelson 102. Introduced. Infrequent. Flowering in May.

M. rubra L. Red mulberry. Valasek 427. NW and N-central portion, rare. Flowering
May - June.

URTICACEAE (Nettle Family)

Boehmeria cylindrica (L.) Sw. Bog-hemp. Anderson 2165; Nelson 215, 252, 301. In
low, wet areas near the river. Flowering June - August.

Laportea canadensis (L.) Wedd. Wood-nettle. Anderson 2164; Nelson 165. Forming
dense colonies in moist areas. Flowering June - August.

Parietaria pensylvanica Muhl. Pellitory. Nelson 149. Abundant in patches in disturbed
areas mostly near trails. Flowering June - July.

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THE MICHIGAN BOTANIST

151

Pilea pumila (L.) Gray. Clearweed. Anderson 2166; Nelson 270. Abundant in very wet
soil. Flowering August - September.

Urtica gracilis Ait. Nettle. Nelson 237, 327. In open, wet, disturbed situations.
Flowering July - August.

JUGLANDACEAE (Walnut Family)

Carya cordiformis (Wang.) K. Koch. Bitternut hickory. Horning in 1960 \ Nelson 164.
Scattered through the woods. Flowering in May.

C. ovata (Mill.) K. Koch. Shagbark hickory. Nelson 201. Rare; in central portion.
Flowering in May.

Juglans cinerea L. Butternut. Nelson 116. Near river in NW section. Flowering in May.

J. nigra L. Black walnut. Valasek 446. One large tree between Bogue Street and Van
Hoosen parking lot. Flowering in May.

FAGACEAE (Beech Family)

Castanea dentata (Marsh.) Borkh. Chestnut. Planted in 1896; the last tree was ap¬
parently removed in the construction of Van Hoosen Hall. Flowering June - July.

Fagus grandifolia Ehrh. Beech. Anderson 2278; Horning 10; Hubbard 6. One of the
most important dominant trees. Flowering in May.

Quercus alba L. White oak. Horning 34. Scattered tnroughout. Flowering in May.

Q. bicolor Willd. Swamp white oak. Morse 7004. Rare; on floodplain in N-central
part. Flowering in May.

Q. macrocarpa Michx. Mossy-cup or bur oak. Nelson 203, 308. On the floodplain.
Flowering in May.

Q. muehlenbergii Engelm. Chestnut oak. Nelson 191. Rare; in NW part. Flowering
May - June.

Q. rubra var. borealis (Michx. f.) Farw. Northern red oak. Nelson 119. Frequent.
Flowering in May.

Q. velutina Lam. Black oak. Noted in 1895 but not located in recent years. Flowering
in May.

BETULACEAE (Birch Family)

Carpinus caroliniana Walt. Ironwood or blue-beech. Anderson 2158; 2259. Frequent
near the river. Flowering April - May.

Ostrya virginiana (Mill.) K. Koch. Ironwood or hop hornbeam. Horning in 1960;
Hubbard 8. Scattered through the woods. Flowering April - May.

CAR YOPHYLLIDAE

PHYTOLACCACEAE (Pokeweed Family)

Phytolacca americana L. Pokeweed. Nelson 213. Frequent weed in open area on S.
edge. Flowering June - October.

CARYOPHYLLACEAE (Pink Family)

Lychnis alba Mill. White cockle. Hubbard 60. Introduced weed in open areas. Flower¬
ing May - October.

Stellaria media (L.) Cyrill. Common chickweed. Nelson 196. Introduced weed in open,
disturbed areas. Flowering May - July.

PORTULACACEAE (Purslane Family)

Claytonia virginica L. Spring beauty. Hubbard 12; Shipman 6. Abundant throughout
upland portion. Flowering March - May.

CHENOPODIACEAE (Goosefoot Family)

Chenopodium album L. Pigweed or lamb’s-quarters. Nelson 226. Introduced weed in
open area at edge. Flowering July - September.

152

THE MICHIGAN BOTANIST

Vol. 9

AMARANTHACEAE (Amaranth Family)

Amaranthus tuberculatus (Moq.) J. D. Sauer. Water-hemp. Nelson 295. In open sandy
soil on riverbank; rare. Flowering July - September.

POLYGONACEAE (Buckwheat Family)

Polygonum aviculare L. Knotweed. Nelson 144. Introduced; infrequent weed on open
edges. Flowering July - October.

P. convolvulus L. Black bindweed. Nelson 518. Introduced weed along open edge.
Flowering July - August.

P. lapathifolium L. Smartweed. Nelson 262, 298. Infrequent in open situations near
the river. Flowering July - August.

P. virginianum L. Jumpseed. Grashoff 16. Common throughout the woods. Flowering
July - August.

Rumex crispus F. Yellow dock. Grashoff 3 38A ; Nelson 175. Introduced weed in open
area at edge. Flowering in June.

R. verticillatus L. Water dock. Valasek 434. N-central part along river. Flowering June
- August.

DILLENIIDAE

GUTTIFERAE (St. John’s-wort Family)

Hypericum perforatum L. St. John’s-wort. Nelson 183. Introduced; infrequent weed at
edge. Flowering June - July.

TIFIACEAE (Finden Family)

Tilia americana L. Linden or basswood. Anderson 2279; Horning 5; Hubbard 2. A
common large tree. Flowering May - June.

VIOLACEAE (Violet Family)

Hybanthus concolor (Forster) Spreng. Green violet. Gilly & Parmelee 237; Grashoff
337. One large patch in NW part. Flowering May - June.

Viola canadensis L. Canada violet. Anderson 2238; Gillis 2655; Popma 331. Abundant
throughout the woods. Flowering mostly in May and June but sporadically from
April to October.

V. pubescens var. eriocarpa (Schwein.) Russell. Yellow violet. Anderson 2231; Hub¬
bard 32. Scattered through the woods. Flowering April - June.

V. rostrata Pursh. Long-spurred violet. Anderson 2237. Infrequent. Flowering April -
May.

V. sororia Willd. Blue violet. Anderson 2236; Hubbard 23, 26; Stergios 19. Scattered
through the woods. Flowering April - June.

V. striata Ait. Cream violet. Stergios 20. Infrequent. Flowering April - July, most
abundantly in May.

CUCURBITACEAE (Gourd Family)

Echinocystis lobata (Michx.) T. & G. Prickly cucumber. Nelson 245. Near the river in
NW portion; rare. Flowering in August.

SALIC ACE AE (Willow Family)

Populus deltoides Marsh. Cottonwood. Nelson 169. By the river, on the E edge and
at sugar house site. Flowering April - May.

P. grandidentata Michx. Large-tooth aspen. Nelson 256. Small tree at sugar house site.
Flowering in May.

P. tremuloides Michx. Quaking aspen. Nelson 190. Small tree at sugar house site.
Flowering in April.

Salix amygdaloides Anderss. Peach-leaved willow. Valasek 429. On road bank at E
edge. Flowering May - June.

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153

S. discolor Muhl. Pussy willow. Morse 69 76. On E edge. Flowering April - May.

S. fragilis L. Crack willow. Morse 6968. Introduced; on small mud flat, S bank of
river. Flowering April - May.

S. interior Rowlee. Sandbar willow. Nelson 211. On road bank at E edge. Flowering
May - June.

S. rigida Muhl. Heart-leaved willow. Valasek 428. Swampy open area near E edge.
Flowering April - May.

CRUCIFERAE (Mustard Family)

Arabis laevigata (Muhl.) Poir. Rock cress. Grashoff 331; Hubbard 52. Scattered
through the woods on sandy soil. Flowering May - June.

Barbarea vulgaris R. Br. Winter cress or yellow rocket. Grashoff 324; Hubbard 57.
Introduced weed, infrequent at edge. Flowering May - August.

Berteroa incana (L.) DC. Hoary alyssum. Nelson 212. Introduced weed, infrequent at
edge. Flowering June - early November.

Brassica campestris L. Field mustard. Nelson 208. Introduced weed, infrequent in
open, disturbed areas. Flowering May - August.

Capsella bursa-pastoris (L.) Medic. Shepherd’s purse. Grashoff 340. Introduced weed,
infrequent in disturbed areas. Flowering May - October.

Cardamine bulbosa (Schreb.) BSP. Spring cress. Grashoff 284, 335; Hubbard 51. In
low wet areas near vernal pools. Flowering April - early June.

C. douglassii (Torr.) Britton. Bitter cress. Hubbard 13. In low wet areas near vernal
pools. Flowering April - May.

Dentaria laciniata Muhl. Toothwort. Hubbard 14. Common throughout the upland
portion. Flowering April - May.

Erysimum cheiranthoides L. Wormseed-mustard. Nelson 157. Introduced; near the
river. Flowering July - October.

Hesperis matronalis L. Dame’s-violet. Grashoff 325; Hubbard 61; Nelson 214. Intro¬
duced; occasional near the river. Flowering May - July.

Lepidium campestre (L.) R. Br. Peppergrass. Nelson 126. Introduced weed at edge.
Flowering May - June.

L. virginicum L. Peppergrass. Nelson 224. In open area at edge. Flowering May - July.

Sisymbrium officinale (L.) Scop. Hedge-mustard. Nelson 222. Introduced; in open area
at edge. Flowering May - July.

PRIMULACEAE (Primrose Family)

Lysimachia ciliata L. Loosestrife. Grashoff 2. In low wet areas near the river. Flower¬
ing June - August.

L. nummularia L. Moneywort. Malcolm in 1957. Introduced; in low wet areas near
the river. Flowering June - August.

Samolus parviflorus Raf. Water-pimpernel. Anderson 2160. On riverbank with
Saururus cernuus. Flowering July - September.

ROS1DAE

GROSSULARIACEAE (Gooseberry Family)

Ribes cynosbati L. Prickly gooseberry. Anderson 2285; Grashoff 305. Scattered
through the wood. Flowering in May.

R. sativum Syme. Red currant. Grashoff 304; Hubbard 37; Stergios 15. Introduced;
near the river. Flowering in May. Stergios 15 and Grashoff 304 appear intermedi¬
ate between this species and the native R. triste Pall.

SAXIFRAGACEAE (Saxifrage Family)

Mitella diphylla L. Bishop’s cap. Anderson 2284; Hubbard 39. Scattered through the
woods. Flowering late April - early June.

154

THE MICHIGAN BOTANIST

Vol. 9

ROSACEAE (Rose Family)

Amelanchier laevis Wieg. Serviceberry. Stergios 1. On the riverbank; infrequent. Flow¬
ering late April - early June.

Crataegus macrosperma Ashe. Hawthorn. Morse 6990, 6995. N of Van Hoosen Hall
and by fence NE of Holmes Hall. Flowering May - June.

C. mollis (T. & G.) Scheele. Hawthorn. Valasek 436. NE part. Flowering March - early
June.

C. punctata Jacq. Hawthorn. Nelson 104. Infrequent; near the river. Flowering May -
early June.

C. succulenta Link. Hawthorn. Morse 6991; Nelson 115. Infrequent; near the river.
Flowering May - June.

Geum canadense Jacq. Avens. Marshall 228; Nelson 171. Scattered through the woods.
Flowering June - August.

Potentilla recta L. Five-finger. Nelson 133. Introduced weed in open areas at edge.
Flowering June - August.

Prunus serotina Ehrh. Black cherry. Horning 19. Scattered, mostly large trees. Flower¬
ing in May.

P. virginiana L. Choke cherry. Grashoff 321; Nelson 221, 323. Forming small thickets,
especially near the W edge. Flowering May - early June.

Rubus allegheniensis Porter. Common blackberry. Grashoff 333. Infrequent; at edge.
Flowering May - June.

R. occidentalis L. Black raspberry. Grashoff 343; Hubbard 58. In open areas. Flower¬
ing May - June.

R. strigosus Michx. Red raspberry. Morse 6996. Near the river at NE corner. Flower¬
ing June - July.

LEGUMINOSAE (Pea Family)

Amphicarpa bracteata (L.) Fern. Hog-peanut. This species has not been collected in
Sanford Natural Area, but immature plants were observed there in June, 1968.
They could not be relocated after the flood in late June. Flowering July -
September.

Desmodium nudiflorum (L.) DC. Tick-trefoil. Grashoff 28. Scattered through the
woods. Flowering July - August.

Gleditsia triacanthos L. Honey -locust. Nelson 244. Seedlings along the riverbank, prob¬
ably from trees planted upriver. Flowering in June.

Gymnocladus dioica (L.) K. Koch. Kentucky coffee-tree. Noted in N-central part in
1895, cut down in 1896 and apparently not present since then. Flowering May -
June.

Medicago lupulina L. Black medick. Nelson 138. Introduced weed at edge. Flowering
June - September.

M. sativa L. Alfalfa. Nelson 313. Introduced weed on E side along road bank. Flower¬
ing June - September.

Melilotus alba Desr. White sweet-clover. Nelson 120. Introduced weed at edge. Flower¬
ing June - October.

M. officinalis (L.) Lam. Yellow sweet-clover. Nelson 130. Introduced weed at edge.
Flowering June - September.

Robinia pseudo-acacia L. Black locust. Grashoff 336. Planted in 1896; several large
trees in W-central portion and some smaller spontaneous individuals. Flowering
May - June.

Trifolium hybridum L. Alsike clover. Nelson 128, 180. Introduced weed at edge.
Flowering June - October.

1970

THE MICHIGAN BOTANIST

155

THYMELAEACEAE (Mezereum Family)

Dirca palustris L. Leatherwood or moosewood. Anderson 2228; Marshall 291. Rare; a
few shrubs scattered through the woods. Flowering late April - May.

ONAGRACEAE (Evening-primrose Family)

Circaea quadrisulcata var. canadensis (L.) Hara. Enchanter’s nightshade. Nelson 187.
Scattered throughout and very abundant in local areas. Flowering June - early
August.

Oenothera biennis L. Evening primrose. Nelson 241. Weed in open area on the river-
bank. Flowering July - September.

CORNACEAE (Dogwood Familv)

Cornus alternifolia L. f. Alternate-leaved dogwood or green osier. Grashoff 303. Scat¬
tered through the woods. Flowering May - June.

C. amomum subsp. obliqua (Raf.) J. S. Wilson. Silky cornel or kinnikinnik. Morse
6962, 6989, 6993; Nelson 167; Valasek 435. On E side and NW corner. Flowering
in June. Morse 6993, Nelson 167, and Valasek 435 are from an apparently hybrid
clone (x C. foemina subsp. racemosa) in the SE corner.

C. florida L. Flowering dogwood. Nelson 147. Infrequent in NW part. Flowering in
May.

C. foemina subsp. racemosa (Lam.) J. S. Wilson. Morse 6973; Valasek 431. Near river
in N-central and NE part. Flowering May - June.

C. stolonifera Michx. Red osier. Noted in 1895 in marshy area on SE edge, the
present site of Holmes Hall, and on W wide. Flowering May - June, infrequently in
July and August.

CELASTRACEAE (Staff Tree Family)

Celastrus scandens L. Climbing bittersweet. Nelson 117. In partially open disturbed
areas near riverbank. Flowering May - June.

Euonymus atropurpureus Jacq. Burning bush or wahoo. Noted in 1895 but not lo¬
cated in recent years. Flowering June - July.

E. obovatus Nutt. Running strawberry-bush. Anderson 2167; Grashoff 36, 332; Hub¬
bard 48. Scattered through the woods. Flowering May - June.

AQUIFOLIACEAE (Holly Family)

Ilex verticillata (L.) Gray. Black alder or winterberry. Noted in 1895 in marshy area
on SE edge, the present site of Holmes Hall. Flowering June - July.

EUPHORBIACEAE (Spurge Family)

Acalypha rhomboidea Raf. Three-seeded mercury. Anderson 2168. On roadside at NE
corner. Flowering August - September.

RHAMNACEAE (Buckthorn Family)

Rhamnus cathartica L. Common buckthorn. Nelson 312. Introduced; one small tree in
NE corner. Flowering in May.

R. frangula L. Alder-buckthorn. Nelson 140. Introduced; a few shrubs on W side.
Flowering May - June.

VITACEAE (Grape Family)

Parthenocissus quinquefolia (L.) Planch. Virginia creeper or American ivy. Nelson 210.
Common throughout the woods. Flowering in June.

Vitis riparia Michx. Frost grape. Nelson 170. Near the river and at E edge. Flowering
May - June.

STAPHYLEACEAE (Bladdernut Family)

Staphylea trifolia L. Bladdernut. Hubbard 43; Janson in 1950. Occasional on W side.
Flowering May - June.

156

THE MICHIGAN BOTANIST

Vol. 9

HIPPOCASTANACEAE (Horse-chestnut Family)

Aesculus hippocastanum L. Horse-chestnut. Morse 6984. Introduced; one shrubby
small tree near W edge. Flowering in May.

ACERACEAE (Maple Family)

Acer negundo L. Box-elder. Nelson 168. Infrequent; at E edge. Flowering April - May.

A. nigrum Michx. f. Black maple. Nelson 158. Scattered through the woods but
mostly near the river. Flowering April - early May. In this area the species shows
much evidence of intergradation with ,4. saccharum.

A. platanoides L. Norway maple. Morse 6431. Introduced; seedling by path near S
edge. Flowering April - May.

A. rubrum L. Red maple. Nelson 146. Infrequent, near the river. Flowering April-early
May.

A. saccharinum F. Silver or white maple. Nelson 106. Frequent in wet areas near
river. Flowering April - early May.

A. saccharum Marsh. Sugar or hard maple. Nelson 159. The dominant tree throughout
most of the woods. Flowering April - May. Intergrading with A. nigrum.

ANACARDIACEAE (Cashew Family)

Rhus typhina F. Staghorn sumac. Nelson 194. In open area at the sugar house site.
Flowering in June.

Toxicodendron radicans (F.) Ktze. Poison-ivy. Gillis 6930. Very abundant, especially
near the river. Flowering in June.

T. vernix (F.) Ktze. Poison sumac. No specimens collected from Sanford Natural Area,
but plants occurred on the site where Holmes Hall was constructed. Flowering
June - July.

RUTACEAE (Rue Family)

Ptelea trifoliata F. Wafer-ash or hop tree. Horning in 1960 \Janson 69. Occasional on
W side in partially open areas. Flowering April - early June.

Zanthoxylum americanum Mill. Prickly-ash. Grashoff 301. Occasional near the river in
NE portion. Flowering in May.

OXAFIDACEAE (Wood-sorrel Family)

Oxalis stricta F. Wood-sorrel. Nelson 143. In open area at W edge; weedy. Flowering
late May - August.

GERANIACEAE (Geranium Family)

Geranium maculatum F. Cranesbill. Hubbard 38; Stergios 26. Common throughout the
woods. Flowering May - early June,

BAFSAMINACEAE (Touch-me-not Family)

Impatiens noli-tangere subsp. biflora (Walt.) Hultdn. Spotted touch-me-not. Grashoff
40. Occasional, mostly in areas where the soil is very wet. Flowering July -
September.

ARAFIACEAE (Ginseng Family)

Aralia racemosa L. Spikenard. Gillis 2654; Nelson 192. In open area at the sugar
house site. Flowering in July.

Panax quinquefolius L. Ginseng. Grashoff 44. Wooded slope; extremely rare. Flower¬
ing May - early June.

P. trifolius F. Dwarf ginseng. Ellsworth 85; Suttkus 31; Shipman 30. In thickly
wooded areas, rare. Flowering April - May.

UMBEFFIFERAE (Parsley Family)

Cryptotaenia canadensis (Torr.) DC. Hone wort. Nelson 197. Scattered through the
woods. Flowering June - July.

1970

THE MICHIGAN BOTANIST

157

Daucus carota L. Wild carrot or Queen Anne’s lace. Nelson 179. Introduced weed at
edge. Flowering June - October.

Erigenia bulbosa (Michx.) Nutt. Harbinger-of-spring. Anderson 2216; Hubbard 3;
Marshall 1521. Infrequent; scattered through the woods. Flowering in April, the
earliest species.

Osmorhiza claytonii (Michx.) C. B. Clark. Sweet cicely. Hubbard 46; Nelson 205.
Scattered through the woods. Flowering May - June.

Sanicula gregaria Bickn. Black snakeroot. Grashoff 327; Nelson 150, 320. Scattered
through the woods. Flowering May - early July.

S. trifoliata Bickn. Black snakeroot. Nelson 321. Infrequent in S-central part. Flower¬
ing late May - early July.

Slum suave Walt. Water-parsnip. Gillis 2415; Grashoff 30. Rare; on swampy riverbank.
Flowering July - October.

Taenidia integerrima (L.) Drude. Yellow pimpernel. Gillis 2447. Collected from N
bank of river adjacent to Sanford Natural Area, but apparently not obtained from
within the area. Flowering in June.

Torilis japonica (Houtt.) DC. Hedge-parsley . Nelson 272. Introduced weed; on S edge.
Flowering July - August.

ASTERIDAE

APOCYNACEAE (Dogbane Family)

Apocynum cannabinum L. Indian-hemp. Nelson 239. In open area on the riverbank at
W edge. Flowering May - September.

ASCLEPIADACEAE (Milkweed Family)

Asclepias incarnata L. Swamp milkweed. Grashoff 27. In mucky soil near the river.
Flowering June - August.

A. syriaca L. Common milkweed. Nelson 278. Weed at W edge. Flowering June -
August.

Cynanchum nigrum (L.) Pers. Nelson 118. Introduced; in somewhat disturbed areas
near W edge. Flowering May - July.

SO LAN ACE AE (Nightshade Family)

Physalis subglabrata Mackenz. & Bush. Ground-cherry. Nelson 248. In small ravine
near W edge, rare. Flowering July - August.

Solanum dulcamara L. Red nightshade. Nelson 315. Introduced weed at E edge.
Flowering June - mid August.

S. nigrum L. Black nightshade. Nelson 225, 267. Introduced weed in disturbed areas.
Flowering July - early October.

POLEMONIACEAE (Phlox Family)

Phlox divaricata L. Phlox. Hubbard 33; Stergios 27. Common throughout the woods.
Flowering April - June.

HYDROPHYLLACEAE (Waterleaf Family)

Hydrophyllum appendiculatum Michx. Waterleaf. Grashoff 319. Infrequent in the
woods. Flowering May - July.

H. virginianum L. Waterleaf. Grashoff 323; Hubbard 53. Scattered through the woods.
Flowering May - early June.

BORAGINACEAE (Borage Family)

Hackelia virginiana (L.) I. M. Johnst. Stickseed. Gillis 2656; Nelson 247. In semi-open,
somewhat disturbed area. Flowering July - August.

Myosotis scorpioides L. Forget-me-not. Grashoff 42; Hubbard 54; Nelson 173. Intro¬
duced; in wet mucky soil near the river. Flowering May - October.

158

THE MICHIGAN BOTANIST

Vol. 9

PHRYMACEAE (Lopseed Family)

Phryma leptostachya L. Lopseed. Anderson 2157; Nelson 275. Rare; in upland woods.
Flowering June - August.

VERBENACEAE (Vervain Family)

Verbena urticifolia L. White vervain. Grashoff 13. Scattered through the woods, in
some areas very abundant. Flowering July - early August.

LABIATAE (Mint Family)

Collinsonia canadensis L. Horse-balm. Grashoff 39. Scattered through the woods.
Flowering August - early October.

Leonurus marrubiastrum L. Motherwort. Valasek 403. Introduced weed; on open
riverbank at NW corner. Flowering August - September.

Lycopus virginicus L. Water-horehound. Anderson 2202; Grashoff 29; Nelson 324. In
wet soil near the river. Flowering July - September.

Prunella vulgaris L. Heal-all. Nelson 255. Weedy, in open area at sugar house site.
Flowering June - October.

Scutellaria lateriflora L. Mad-dog skullcap. Grashoff 12; Nelson 250, 296. In wet soil
near the river. Flowering May - August.

Stachys tenuifolia Hedge-nettle. Grashoff 14; Nelson 261, 283, 307. In wet soil near
the river. Flowering July - August.

Teucrium canadense L. Wood-sage. Grashoff 9: In road ditch at E edge. Flowering
July - September.

PLANTAGINACEAE (Plantain Family)

Plantago lanceolata L. Ribgrass or buckhorn. Nelson 135. Introduced weed at edge.
Flowering May - June.

P. major L. Common plantain. Nelson 276. Introduced weed in open disturbed area at
E edge. Flowering July - October.

P. rugelii Dene. Plantain. Kaiser 238. Infrequent. Flowering July - September.

OLEACEAE (Olive Family)

Fraxinus americana L. White ash. Horning 15; Nelson 198. One of the most common
trees on the moister sites. Flowering in May.

F. nigra Marsh. Black ash. Nelson 260, 311. Near the river in NE portion. Flowering
May - June.

F. quadrangulata Michx. Blue ash. Morse 7000. On W side, rare. Flowering in May.

Ligustrum vulgare L. Privet. Valasek 426. Introduced; on riverbank in N-central part.
Flowering May - July.

SCROPHULARIACEAE (Figwort Family)

Chelone glabra L. Turtlehead. Nelson 305. In wet soil near the river. Flowering July -
September.

Linaria vulgaris Hill. Toadflax. Nelson 316. Introduced weed, in open, disturbed soil
at E edge. Flowering June - October.

Mimulus ringens L. Monkey-flower. Grashoff 25. In very wet soil near the river.
Flowering late June - September.

Scrophularia marilandica L. Carpenter’s-square. Nelson 254. In somewhat disturbed,
shaded area at edge of sugar house site. Flowering June - July.

Verbascum thapsus L. Common mullein. Nelson 218. Introduced weed, in disturbed
area at E edge. Flowering June - August.

V er onicastrum virginicum (L.) Farw. Culver ’s-root. Grashoff 15. In wet soil near the
river. Flowering June - August.

1970

THE MICHIGAN BOTANIST

159

OROBANCHACEAE (Broom-rape Family)

Epifagus virginiana (L.) Bart. Beech-drops. Anderson 2204; Nelson 288. Parasitic on
beech, scattered through the woods. Flowering July - October.

CAMPANULACEAE (Bluebell Family)

Campanula americana L. Tall bellflower. Grashoff 1; Nelson 228. Scattered through
the woods, very abundant in some areas. Flowering June - August.

Lobelia cardinalis L. Cardinal-flower. Grashoff 11; Nelson 282. In pools near the river.
Flowering July - October.

L. syphilitica L. Blue cardinal-flower. Grashoff 37, 38. In very wet soil near the river.
Flowering August - early October.

RUBIACEAE (Madder Family)

Cephalanthus occidentalis L. Buttonbush. Nelson 253. By intermittent lagoon in
N-central portion. Flowering June - August.

Galium aparine L. Cleavers. Hubbard 49. Scattered through the woods. Flowering May

- June.

G. circaezans Michx. Wild-licorice. Nelson 162. Infrequent. Flowering May - July.

G. obtusum Bigel. Bedstraw. Nelson 172. Infrequent; in open area at W edge. Flower¬
ing in June.

G. triflorum Michx. Sweet-scented bedstraw. Nelson 148, 189. Scattered through the
woods. Flowering May - July.

Mitchella repens L. Partridge-berry. Valasek 445. Rare; one colony in NW portion near
the river. Flowering June - July.

CAPRIFOLIACEAE (Honeysuckle Family)

Lonicera maackii Maxim. Morse 6975. Introduced; occasional throughout the woods.
Flowering material of Michigan specimens lacking in the Beal-Darlington Herbarium.

L. tatarica L. Tatarian honeysuckle. Grashoff 302. Introduced; scattered through the
woods. Flowering May - June.

L. xylosteum L. Fly honeysuckle. Morse 6980. Introduced; in W-central part. Flower¬
ing in May.

Sambucus canadensis L. Common elder. Nelson 209. At SE edge near vernal pool
drained by road construction. Flowering June - early August.

S. pubens Michx. Red-berried elder. Grashoff 295; Hubbard 41. Scattered through the
woods. Flowering late April - June.

Viburnum acerifolium L. Arrow-wood or maple-leaved viburnum. Morse 6985;
Parmelee 516. Rare; near W edge. Flowering May - June.

V. lantana L. Wayfaring tree. Grashoff 297. Introduced; at W edge. Flowering May -
June.

V. lentago L. Sheepberry or nanny berry. Morse 7012. Rare; one small shrub SW of
sugar house site. Flowering May - June.

V. recognitum Fern. Arrow-wood. Morse 6977; Nelson 156. Rare in this woods and in
Michigan. Near the river in N-central and E part. Flowering material of Michigan
specimens lacking in the Beal-Darlington Herbarium.

V. trilobum Marsh. Highbush-cranberry. Anderson 2368; Donahue 88; Grashoff 314,
Hubbard 56. Scattered through the woods, especially near the river. Flowering May

- July.

DIPSACACEAE (Teasel Family)

Dipsacus laciniatus L. Teasel. Nelson 193. Introduced weed in open area at the sugar
house site. Flowering July - August.

COMPOSITAE (Composite or Aster Family)

Achillea millefolium L. Common yarrow. Nelson 136. Introduced weed at W edge.
Flowering May - October.

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Ambrosia artemisiifolia L. Common ragweed. Nelson 280. Weed in disturbed areas
around the edge. Flowering August - September.

A. trifida L. Great ragweed. Nelson 238. Common in some open areas, especially near
riverbank. Flowering July - August.

Anthemis arvensis L. Corn-chamomile. Nelson 223. Introduced weed in open dis¬
turbed area at W edge. Flowering May - June.

Arctium minus (Hill) Bernh. Common burdock. Nelson 195. Introduced weed; scat¬
tered through the woods. Flowering July - August.

Aster cordifolius L. Nelson 285, 325. In partly open margin on W side. Flowering
August - October.

A. lateriflorus (L.) Britt. Calico aster. Nelson 291. In partly open margin on W side.
Flowering July - early October.

A. macrophyllus L. Large-leaved aster. Miller 3015; Nelson 246. Scattered through the
woods. Flowering August - October.

A. pilosus Willd. Nelson 290. In partly open margin on W side. Flowering August -
October.

Bidens cernua L. Stick -tight. Nelson 297. On riverbank in NW portion. Flowering
August - October.

B. frondosa L. Beggar-ticks. Nelson 294. In wet soil on riverbank. Flowering July -
September.

Cichorium intybus L. Common chicory. Nelson 178. Introduced weed; common
around the edge. Flowering June - October.

Cirsium arvense (L.) Scop. Canada thistle. Nelson 181. Introduced weed in open area
at W edge. Flowering July - October.

C. vulgare (Savi) Tenore. Bull thistle. Nelson 268. Introduced weed in open area along
road bank at E edge. Flowering July - October.

Conyza canadensis (L.) Cronq. Horseweed. Nelson 273, 310. Weed at edge of woods
and along riverbank. Flowering August - September.

Erigeron annuus (L.) Pers. Daisy-fleabane. Nelson 101, 236. Weed at edge of woods.
Flowering June - early October.

E. philadelphicus L. Fleabane. Grashoff 339; Hubbard 63. In open areas at edge.
Flowering May - June.

Eupatorium maculatum L. Joe-Pye-weed. Valasek 402. Infrequent on semi-open river¬
bank in N-central part. Flowering July - September.

E. purpureum L. Joe-Pye-weed. Nelson 232. Scattered through the woods. Flowering
July - September.

E. rugosum Houtt. White snakeroot. Nelson 231. Scattered through the woods. Flow¬
ering July - early October.

Helianthus decapetalus L. Sunflower. Nelson 219. Abundant along open S edge. Flow¬
ering late July - early September.

Matricaria matricarioides (Less.) Porter. Pineapple-weed. Nelson 227. Introduced weed
in open disturbed areas. Flowering June - September.

Polymnia canadensis L. Leafcup. Grashoff 43; Miller 301 7. Infrequent in NE part.
Flowering late May - October.

Prenanthes altissima L. Rattlesnake-root. Nelson 322. Scattered through the woods.
Flowering July - October.

Rudbeckia laciniata L. Coneflower. Nelson 281. 306. In low very wet areas near the
river. Flowering July - early October.

Solidago caesia L. Blue-stem goldenrod. Nelson 286. In partly open W edge. Flowering
late August - early October.

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161

S. canadensis L. Goldenrod. Nelson 284, 289. In open area at W edge. Flowering late
July - October.

S. flexicaulis L. Goldenrod. Anderson 2156; Nelson 230, 279, 287. Along partly open
W edge. Flowering July - September.

S. gigantea Ait. Goldenrod. Nelson 326. In wet area in N-central portion. Flowering
August - September.

S. graminifolia (L.) Salisb. Goldenrod. Nelson 292. In open area at W edge. Flowering
late July - September.

Sonchus arvensis L. Sow-thistle. Nelson 265. Introduced weed; on open road bank at
E edge. Flowering July - October.

S. oleraceus L. Sow-thistle. Nelson 274. Introduced weed on S edge. Flowering June -
August.

Taraxacum officinale Weber. Dandelion. Grashoff 299; Stergios 10. Introduced weed in
open, disturbed areas. Flowering April - October.

LILIATAE (Monocotyledons)

ALISMATIDAE

ALISMATACEAE (Water-plantain Family)

Alisma plantago-aquatica var. americanum Roem. & Schult. Water-plantain. Grashoff
26; Nelson 314. In pools near the river in NE portion. Flowering July - August.

Sagittaria latifolia Willd. Arrowhead. Grashoff 24. Swampy ground near the river in
NW portion. Flowering July - August.

HYDROCHARITACEAE (Frog’s-bit Family)

Elodea nuttallii (Planch.) St. John. Waterweed. Nelson 177. In the river near Bogue
Street Bridge and in vernal pools on the floodplain. Flowering July - August.

COMMELINIDAE

JUNCACEAE (Rush Family)

Juncus tenuis Willd. Path rush. Nelson 145. Near trail along river in N-central portion.
Flowering June - August.

Luzula acuminata Raf. Wood rush. Hubbard 21. N side near river. Flowering April -
May.

CYPERACEAE (Sedge Family)

Carex albursina Sheldon. A nderson 2288. Along paths at E end. Flowering May - July.

C. alopecoidea Tuckerm. Nelson 182. NW edge. Flowering June - July.

C. blanda Dewey. Nelson 122. NW corner by Bogue Street Bridge. Flowering May -
June.

C. careyana Torr. Grashoff 287; Marshall 296; Stergios 3. Along path near river.
Flowering April - June.

C. cephaloidea Dewey. Grashoff 316. Flowering May - early June.

C. convoluta Mackenzie. Nelson 142. NW edge. Flowering late May - July.

C. grayii Carey. Anderson 2203; Hubbard 62. N-central part in moist areas. Flowering
June - August.

C. intumescens Rudge. Hubbard 55. Low area. Flowering May - July.

C. lupulina Muhl. Nelson 216. SE corner by former vernal pond. Flowering July -
August.

C. molesta Mackenzie. Nelson 123. NW corner by Bogue Street Bridge. Flowering June
- July.

C. muskingumensis Schwein. Valasek 443. Rare; edge of floodplain in N-central part.
Flowering May - July.

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THE MICHIGAN BOTANIST

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C. pedunculata Muhl. Valasek 441. NE part near N trail. Flowering May - June.

C. plantaginea Lam. Anderson 2218, 2287; Hubbard 15. In E portion. Flowering April

- May.

C. sprengelii Dewey. Grashoff 310; Marshall 295. Moist shady woods. Flowering April

- June.

C. stricta var. strictior (Dewey) Carey. Marshall 294. Moist shady area. Flowering
April - June.

C. woodii Dewey. Matthews & Matthews 96; Stergios 4. Along trail by river. Flower¬
ing April - May.

Eleocharis acicularis (L.) R. & S. Spike rush. Valasek 438. By intermittent lagoons in
N-central part. Flowering June - September.

GRAMINEAE (Grass Family)

Agropyron repens (L.) Beauv. Quack grass. Nelson 134, 229. Introduced weed; along
W edge N of Van Hoosen Hall. Flowering June - August.

Brachyelytrum erectum (Schreb.) Beauv. var. erectum. Anderson 2159; Nelson 113,
243. Scattered through the woods, mostly near the river. Flowering June - July.

Bromus inermis Leyss. Brome grass. Grashoff 344. Introduced weed on NW edge.
Flowering June - July.

B. latiglumis (Shear) Hitchc. Brome grass. Anderson 2163; Nelson 242. NW corner.
Flowering June - August.

B. purgans L. Brome grass. Parmelee 289. In the woods. Flowering July - August.

B. tectorum L. Brome grass. Nelson 125. Introduced weed near the river at NW
corner. Flowering May - July.

Calamagrostis canadensis (Michx.) Nutt. Parmelee 284. Floodplain. Flowering July -
August.

Cinna arundinacea L. Wood reed. Anderson 2161 ; Nelson 258, 271. By vernal ponds
in central and SE portion. Flowering in August.

Dactylis glomerata L. Orchard grass. Grashoff 338; Nelson 127. Introduced weed at
NW corner. Flowering June - August.

Echinochloa muricata (Beauv.) Fern. Barnyard grass. Nelson 300. On riverbank at NW
corner. Flowering August - September.

Elymus riparius Wieg. Wild rye. Nelson 235. In open weedy area at edge. Flowering in
August.

E. villosus Muhl. Wild rye. Nelson 204. Near trail along river in NW portion. Flower¬
ing July - August.

E. virginicus L. Wild rye. Nelson 234, 263. NE portion of floodplain and in weedy
open edge. Flowering July - August.

Eragrostis pilosa (L.) Beauv. Love grass. Popma 329. Introduced weed on riverbank at
NW corner. Flowering July - October.

Festuca obtusa Biehler. Fescue. Nelson 220. At sugar house site. Flowering June -
July.

F. pratensis Huds. Meadow fescue. Grashoff 334. Introduced weed. Flowering June -
August.

F. ovina L. Sheep fescue. Grashoff 342; Nelson 124, 137. Introduced weed in NW
portion by Bogue Street Bridge and at sugar house site. Flowering May - July.

Glyceria striata (Lam.) Hitchc. Manna grass. Nelson 151, 217. Scattered through the
woods. Flowering June - July.

Hystrix patula Moench. Bottle-brush grass. Marshall 227; Nelson 132. In NW portion.
Flowering June - July.

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163

Leersia virginica Willd. White grass. Nelson 269. E-central edge. Flowering August -
September.

Lolium perenne L. English rye grass. Nelson 129, 277. Introduced weed, at NW and
SW edges. Flowering June - August.

Muhlenbergia frondosa (Poir.) Fern. f. commutata (Scribn.) Fern. Muhly. Nelson 303.
On riverbank at NW corner. Flowering August - September.

M. schreberi Gmel. Muhly. Nelson 266. E-central edge. Flowering August - September.

Oryzopsis racemosa (Sm.) Ricker. Rice grass. Valasek 440. N-central part near trail.
Flowering in July.

Panicum capillare L. Witch grass. Nelson 302. NW portion on riverbank. Flowering
July - September.

Phleum pratense L. Timothy. Nelson 121. Introduced weed at NW corner. Flowering
June - August.

Poa alsodes Gray. Grashoff 329; Hubbard 50. N side along riverbank. Flowering May -
June.

P. annua L. Spear grass. Nelson 257. Introduced weed at sugar house site. Flowering
June - October.

P. nemoralis L. Nelson 110. On riverbank in NW section. Flowering June - August.

P. palustris L. Fowl meadow grass. Nelson 264. On riverbank in NE portion.
Flowering July - August.

P. pratensis L. Kentucky blue grass. Nelson 185, 233. Introduced weed at NW corner.
Flowering June - August.

Setaria glauca (L.) Beauv. Foxtail grass. Nelson 293. Introduced weed on riverbank
near Bogue Street Bridge. Flowering July - September.

Sphenopholis intermedia (Rydb.) Rydb. Grashoff 322; Nelson 111. On riverbank in
NW portion. Flowering in June.

ARECIDAE

ARACEAE (Arum Family)

Arisaema triphyllum (L.) Schott. Jack-in-the-pulpit or Indian turnip. Anderson 2280,
2282; Hubbard 28. Scattered through the woods. Flowering late April - June.

A. dracontium (L.) Schott. Green dragon. Nelson 155. N-central portion near the river
in low, wet areas. Flowering May - June.

Symplocarpus foetidus (L.) Nutt. Skunk cabbage. Anderson 2229; Hubbard 11. Near
river in NW portion and formerly abundant in marshy area on SE edge where
Holmes Hall was constructed. Flowering in April.

LEMNACEAE (Duckweed Family)

Lemna minor L. Duckweed. Nelson 176. In river near Bogue Street Bridge, and in
vernal pools on the floodplain. Flowering June - August.

LILIIDAE

LILIACEAE (Lily Family)

Allium tricoccum Ait. Wild leek. Grashoff 41. In W-central area and near river in
N-central portion. Flowering July - August.

Asparagus officinalis L. Asparagus. Nelson 317. Introduced weed in E section on
floodplain. Flowering May - June.

Erythronium albidum Nutt. White dog’s-tooth-violet. Hubbard 22. N-central portion in
wet area; rare. Flowering April - May.

E. americanum Ker. Yellow adder’s tongue. Anderson 2281; Grashoff 289; Hubbard
18. Frequent in the woods. Flowering April - May.

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THE MICHIGAN BOTANIST

Vol. 9

Hemerocallis fulva L. Day lily. Nelson 166. Introduced weed in SE part. Flowering
July - August.

Lilium michiganense Farw. Michigan lily. Nelson 259. N-central and NE floodplain;
rare. Flowering June - July.

Maianthemum canadense Desf. Canada mayflower. Nelson 186 (var. interius Fern.);
Suttkus 63 (var. canadense ). Near the river. Flowering May - June.

Polygonatum pubescens (Willd.) Pursh. Small Solomon’s seal. Anderson 2283. Scat¬
tered through the woods. Flowering May - June.

Smilacina racemosa (L.) Desf. False Solomon’s seal. Hubbard 47. Scattered through
the woods. Flowering May - June.

S. stellata (L.) Desf. False Solomon’s seal. Hubbard 40. Scattered through the woods.
Flowering May - June.

Trillium grandiflorum (Michx.) Salisb. Trillium. Hubbard 31; Matthews 101. Frequent
throughout the woods. Flowering April - May.

Uvularia grandiflora Sm. Bellwort. Anderson 2258; Grashoff 290; Hubbard 29. Scat¬
tered through the woods. Flowering April - May.

IRIDACEAE (Iris Family)

Iris virginica var. shrevei (Small) Anderson. Iris. Grashoff 312. By intermittent lagoons
in N-central portion. Flowering in June.

SMILACACEAE (Greenbrier Family)

Smilax ecirrata (Engelm. ex Kunth) S. Wats. Hubbard 36. Near the river. Flowering in
May.

S. illinoensis Mangaly. Gillett 606; Grashoff 326. NW part. Flowering May - June.

S. tamnoides var. hispida (Muhl.) Fern. Bristly greenbrier. Grashoff 313; Nelson 114.
NW portion near the river. Flowering in June.

DIOSCOREACEAE (Yam Family)

Dioscorea villosa L. Wild yam. Nelson 174, 202. By intermittent lagoons in N-central
portion. Flowering June - July.

ORCHIDACEAE (Orchid Family)

Aplectrum hyemale (Muhl.) Torr. Adam-and-Eve. GiUy & Parmelee 236. Rare.
Flowering May - June.

Corallorhiza maculata Raf. Coral-root. H. Miller in 1967. In NW portion near trail;
rare. Flowering July - August.

Epipactis helleborine (L.) Crantz. Helleborine. Marshall 27. Introduced; near the river.
Flowering July - August.

LITERATURE CITED

Beaman, J. H. 1970. A botanical inventory of Sanford Natural Area. I. The environment.
Mich. Bot. 9: 116-139.

Cronquist, A. 1968. The Evolution and Classification of Flowering Plants. Houghton
Mifflin Co., Boston. 396 pp.

- , A. Takhtajan, & W. Zimmermann. 1966. On the higher taxa of Embryo-

bionta. Taxon 15: 129-134.

Melchior, H., & E. Werdermann (editors). 1954. A. Engler’s Syllabus der Pflanzenfamilien.

I. Band. Gebriider Borntraeger, Berlin-Nikolassee. 367 pp.

Taylor, F. G. 1969. Phenological Records of Vascular Plants at Oak Ridge, Tennessee.
ORNL- IBP- 69-1, Oak Ridge. 46 pp.

1970

THE MICHIGAN BOTANIST

165

Reviews

HOW TO KNOW POLLEN AND SPORES. By Ronald O. Kapp. Wm. C. Brown Co.,
Dubuque. 1969. 249 pp. $3.25 (spiral); $4.00 (cloth).

This is in some ways a frustrating book, but one which will be useful to most
American palynologists. It could have been very much better and more useful if the
micro-morphology of the pollen were described more carefully, if the drawings were more
detailed, and if greater care had been taken in editing. A serious mistake in editing is the
reproduction of the numerous sketches in the key at different magnifications. This gives a
misleading impression of the relative sizes of different pollen and spores. Pollen from four
species of oak, for example, that differs in size by 30% at the most, is illustrated on the
same page by drawings which differ in size by a factor of two.

The book begins with a general discussion of the morphology and function of
pollen grains and spores, and with a very brief description of sampling and preparation
techniques. The description of exine morphology is quite clear, but is incomplete in
failing to describe “LO analysis.” This technique, involving study of the exine under high
magnification at different levels of focus to interpret 3-dimensional structures, is given
only a brief definition in the glossary and index that appears at the end of the book. Yet
it is essential for distinguishing pitted structures in the exine from raised structures. These
very different kinds of structure are frequently confused at low magnification by inex¬
perienced palynologists. The same difficulty arises with the illustrations, which are incon¬
sistent; they should show structure with stippling and spaces unstippled (e.g., both Salix
(fig. 191) and Rorippa (fig. 194) have a raised netlike structure, but this is stippled in
Salix and light in Rorippa). Although the key is easy to follow, since variable types or
types that are difficult to interpret are repeated in appropriate sections, the author is
unclear in his written descriptions of the fine structure of pollen exines. I would argue,
to give a few examples, with his description of Potamogeton as tectate, Dryas as psilate,
Acer pensylvanicum as intectate.

The main usefulness of the book will be to direct American palynologists to the
family or genus in which to look for analogs to unknown fossil grains. The drawings and
descriptions will indicate possible identifications, although they should not be used them¬
selves for identifications. Kapp cautions that palynologists must have available reference
collections prepared from authoritatively identified plants. He has done us all a service
by including in this treatment spores from mosses and fungi, as well as algae and proto¬
zoans that are frequently confused with pollen grains.

-M. B. Davis

PLANTS OF THE CHICAGO REGION A Check List of the Vascular Flora of the
Chicago Region with Notes on Local Distribution and Ecology. By Floyd Swink.
Morton Arboretum, Lisle, Illinois. 1969. 445 pp. $8.50.

The distribution of each species is indicated, largely by maps, for 22 counties
around the head of Lake Michigan, from three in southeastern Wisconsin to Berrien
County, Michigan. The introduction makes clear exactly what the book intends to do
(and not to do) and lucidly explains the style and conventions employed. The work is
based on critical evaluation of both specimens and literature reports. Perhaps its major
original contribution is extensive listing of associated species and habitat preferences.
The species are in one alphabetical list, with cross-references by synonyms and common
names; all included genera, however, are listed under family names on pp. xvii-xxii.
Nomenclaturists will regret the use of undesignated trinomials, although the introduction
states that all are varieties. The volume is well bound and is reproduced very clearly from
typed copy in a format easy to consult. I can recall examining no other book so closely
without finding a single typographical error! The Morton Arboretum and its industrious
taxonomist can justly be proud of this fine contribution to the local flora literature.

-E. G. V.

166

THE MICHIGAN BOTANIST

Vol. 9

NATURAL PROLIFERATION OF FLOATING STEMS
OF SCOURING-RUSH, EQUISETUM HYEMALE

Warren H. Wagner, Jr., and William E. Hammitt1
Matthaei Botanical Gardens,

The University of Michigan, Ann Arbor

Scouring-rush, Equisetum hyemale L., is one of the most familiar plants
of the northern hemisphere. Its colonies of rigid, upright, cone-tipped stems
are scattered across an enormous range in northern Europe, Asia, and North
America, in the New World running southward to Guatemala. The stems, with
their gritty epidermis encrusted with silica tubercles, were used by our fore¬
bears for scouring pots and pans, and even today this or related species are
used by poor people in other nations for this purpose. To the botanist, the
scouring-rushes and their relatives, the horsetails, have special interest, for they
represent the last survivors of a once prominent element of the world’s flora.
In the Coal Age, some of these plants grew as tall as modern trees, and there
were a number of distinctive genera. There is today but a single genus with
only 15 species.

Because they are unique among plants on the earth today, scouring-
rushes and horsetails have received much attention from morphologists. They
are completely isolated taxonomically, and any common ancestry with other
living plants occurred at least several hundred million years ago.

Although we usually assume that scouring-rushes and horsetails repro¬
duce by spores and gametophytes in a manner similar to the ferns, more and
more evidence suggests that at least under certain circumstances, vegetative
propagation may play an important role in forming new colonies. As anyone
who has ever attempted to dig up a colony of scouring-rushes knows, the
underground stems or rhizomes are rigid and fibrous, thicker than ordinary
clothes rope. These rhizomes form extensive subterranean systems, often ex¬
tending hundreds of feet along railroad embankments, lake shores, and the
banks of streams. Thus, when one discovers a number of seemingly discrete
colonies, say, along a highway shoulder, it is not necessarily true that they are
separate colonies which originated from separate spores. All of the plants and
all of the clumps may represent a single stock from a single fertilization.

An even more interesting mode of vegetative reproduction in scouring-
rushes involves fragmentation of stems. Theoretically, if pieces of stem should
be broken off, for one reason or another, and if they should float across a
lake or down a river, it should be possible for them to found a new colony by
settling on some damp shore and forming new roots and shoots. Contained
within the nodes and inside the leaf sheaths are tissues which are still
embryonic and capable of proliferating. It is these soft tissues which enable us

*We are indebted to the following persons for help in various ways: Peter Fajans,
Richard Hauke, John Ludwig, Robert Sabo, Terry Shank, Sylvia Taylor, and W. C.
Wagner.

1970

THE MICHIGAN BOTANIST

167

to pull the stems of scouring-rushes into separate units or internodes so easily.
Each internode is attached to the node below it by a ring of delicate tissue,
the intercalary meristem, which separates readily when pulled. This meristem
is composed of more or less undifferentiated cells which are capable of divid¬
ing when properly stimulated. The occasional branched specimens of scouring-
rush that we encounter in the field are caused by some injury to the apex or
death of the apex; new side branches are then stimulated to form and these
break through the sheaths. Such specimens have been given formal names (“f.
proliferum ” and “f. polystachyum ”) but they do not deserve such recognition,
because they result from growth conditions, not truly genetic differences.

Hauke (1963) has theorized that the spread of hybrid scouring-rushes
over long distances may be due to vegetative propagation by stem fragments.
All known hybrids between species of scouring-rushes are sterile, and their
spores are abortive, incapable of normal germination and growth; thus they
cannot spread by spores. Hauke states that vegetative propagation via stem
fragments may explain the existence of certain hybrids in places where one or
both of the parents may be absent. For example, the sterile scouring-rush
cross, E. Xferrissii Clute, is an intermediate between common scouring-rush, E.
hyemale, and the smooth scouring-rush, E. laevigatum A. Br. Its spores are
abortive, and the cones fail to open. In the geographical regions where the
two parental species grow together, the hybrid scouring-rush, as would be
expected, is common, as in the Upper Great Lakes region. However, as Hauke
has demonstrated, this hybrid is also present in regions where E. laevigatum is
absent, indeed often at long distances from the range of that parent. There are
records of E. Xferrissii from Quebec and Connecticut south to Virginia, far
east of the Ohio Valley localities at the easternmost edge of the range of E.
laevigatum. Hauke hypothesizes that E. Xferrissii exists in its eastern range
because fragments of its stems were carried by rivers, by boats, or by becom¬
ing lodged in wheels of passing trains. If the fragments happened to fall in
appropriate sites, they proliferated new plants and formed colonies. He per¬
formed experiments in which he showed that portions of the hybrid stem can
be cut and planted in wet soil and will, in fact, produce new plants.

In this article we wish to report studies made during the summer and
fall of 1969 of the occurrence of natural proliferation of stems of the com¬
mon scouring-rush while floating in water. One of the consequences of our
study is the conclusion that it is not necessary for a given segment of stem to
be “planted,” i.e., placed in soil, before it forms a new plant. If the means of
dispersal of the stem fragment is via water— across a lake or down a stream—
the fragment may well have proliferated and produced a plant before it is cast
upon the appropriate shore situation for establishment. In other words, a new
plant will be formed and ready for immediate rooting and growth.

FIELD OBSERVATIONS

In July, 1969, we encountered what appeared to be small plants of
Equisetum growing directly out of the water of Dix Pond (Fig. 1) on the
grounds of the Matthaei Botanical Gardens, due south of Dixboro, Washtenaw

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THE MICHIGAN BOTANIST

Vol. 9

County, Michigan. Closer examination revealed that the small plants were
actually growing from the nodes of floating, detached stems of E. hyemale
(Fig. 2). Further observations in Dix Pond revealed dozens of floating stem
fragments of this species with all stages of development of new plants in
evidence.

Scattered at different localities in the Matthaei Gardens natural areas are
four taxa of Equisetum: Common horsetail, E. arvense L., is abundant in
fields and woods; the water horsetail (or “pipes”), E. fluviatile L., is limited
to wet, marshy spots in the woods along Fleming Creek. The only other
scouring-rush beside E. hyemale that we have found thus far on the some 350
acres of the Gardens is a hybrid, E. Xnelsonii (A. A. Eat.) Schaffn. (= E.
laevigatum X variegatum), known from a couple of localities along pond
shores. (It is interesting to note that neither of the parents of the sterile E.
Xnelsonii have been discovered in the Matthaei Gardens area.) Common scour-
ing-rush, E. hyemale , is abundant in several areas, including the north end of
Dix Pond.

Dix Pond is an artificial pond that resulted from the natural flooding of
an old gravel pit. It is rectangular in outline, approximately 395 feet long, 120
feet wide, and has an average depth of 5 to 6 feet. The pond has occasionally
been used for research purposes by University of Michigan biologists since the
time the gravel pit operation was discontinued around 1950. Pine trees were
planted during the period 1950-55 in the surrounding John Dix Field, but the
vegetation bordering the pond is from natural seeding and has developed
spontaneously. The trees and shrubs along the borders are shrubby St. John’s-
wort, Hypericum prolificum, the green ash, Fraxinus pennsylvanica, cotton¬
wood, Populus deltoides , and several willows, Salix spp. Although the dry,
open fields surrounding the pond contain mainly weedy and introduced
species such as sweet-clovers, Melilotus officinalis and M. alba, star thistle,
Centaurea maculosa, Queen Anne’s lace, Daucus carota, and common milk¬
weed, Asclepias syriaca, the banks of Dix Pond support mainly native aquatic
or semi-aquatic plants— cattail, Typha latifolia, sedges, Carex spp., bulrush,
Scripus validus , spike-rush, Eleocharis sp., and water-plantain, Alisma triviale.
The scouring-rushes along the bank are accompanied by the more abundant
and widespread field horsetail, £. arvense.

As our study of the natural proliferation of floating stems of the scour¬
ing-rush proceeded, it became evident that there were dozens of them, in
some places forming such masses as to produce the appearance of a whole
colony of small scouring-rushes growing out of the water. For the most part
the stems which rose above the surface were not much more than 1 or 1.5
mm in thickness (Fig. 2). Below most of them, clusters of roots hung down
into the water. Most remarkable was the fact that a few of the aerial stems, in
spite of their size and slender habit, bore small strobili at their tips. Thus
essentially complete, although miniature, plants could be formed from floating
stem fragments, entirely without connection to soil. Specimens of various sizes
and showing different forms of proliferations are illustrated in Figure 3.

1970

THE MICHIGAN BOTANIST

169

Fig. 1. Border of Dix Pond, Matthaei Botanical Gardens, showing clones of scouring-rush
along the shore.

Fig. 2. Floating culms of scouring-rush with upright shoots indicated by arrows. Roots
hang into the water below. (Strip = 6 inches).

170

THE MICHIGAN BOTANIST

Vol. 9

Our interest was further stimulated by the fact that there were so many
stem fragments present on the pond surface. This suggested that some agent
besides mere casual breaking off of stems was operating, and the clue to the
cause was found by examining the broken tips. The majority of the fragments
showed that the ends had obviously been chewed off— not merely separated at
the nodes as one might expect. This led us to suspect that some mammal was
involved, and it soon became evident that the animal in question was the
muskrat. Eighteen muskrat tunnel entrances were subsequently located in Dix
Pond, and the animal is obviously common here. Whether or not the muskrats
here actually feed upon the scouring-rush fragments that they chew off is not
known. The cutting of the plants may be merely a by-product of their clear¬
ing of paths to the Typha and Scirpus, which the muskrats do feed on, and of
which practically every stem shows evidence of their chewing.

In order to gain some idea of the role of stem length and stem position
in the origin of new plants from the floating fragments, we collected approxi¬
mately 90 of them, and tabulated the number of nodes, the number and
length of shoots and roots per node, and the positions of the nodes. All of
the proliferations were found to arise from the nodes, i.e., from the inter¬
calary meristems, and none occurred at the broken surfaces of internodes. All
sizes of fragments were found to proliferate, even those with only two nodes
(cf. Fig. 3, fragments on left). However, the larger fragments, those with 10 or
more nodes, tended to show two phenomena: terminalization of the prolifera¬
tions and polarity. The greatest amount of proliferation of any kind was
toward the ends; the least amount in the middle (cf. Fig. 3, fragment on
right). Also, the upper end of the fragment (as can be readily determined
from the orientation of the sheaths) tends to produce mainly shoots, usually
without any evident roots at all, while the lower end forms abundant roots as
well as shoots (data tabulated in Tables 1 & 2). This polarity is extremely
interesting in view of the fact that the fragments float in a horizontal plane. It
suggests therefore that there is an inherent polarity independent of the
orientation of the fragment: the morphologically basal part of the stem is
more prone to proliferate complete plants with roots and shoots, and the
morphologically apical part to proliferate only shoots.

EXPERIMENTAL OBSERVATIONS

To find how long it takes for the scouring-rushes to sprout while float¬
ing, as well as what conditions are needed and how much proliferation will
occur under more or less controlled conditions, we placed fragments in dif¬
ferent situations. These fragments were loosely tied with a string to a weight
or naturally fixed object and left to float freely on the water surface. We
selected sites that showed different combinations of slow and rapid current
and shade and sun. In each site we used a sample of 14 stems, each consisting
of the terminal internode plus 10 or 11 nodes. In each sample, 7 stems were
cut at the node, and 7 between the nodes. The sites selected and the results
of the experiments are shown in Table 3. To summarize the results, the
following points should be made: (1) Within 6 days after fragments were

1970

THE MICHIGAN BOTANIST

171

floated, they began to sprout. (2) Those in full sunlight sprouted first, but
sprouting eventually occurred even in strong shade. (3) The sprouts were
formed at the nodes, beneath the sheaths, but not in the internodes. (4) Still
water is not necessary for proliferation, nor is flotation at the surface. Sprouts
occurred even in rapid current, where the fragments became submerged. (5)
Shoots up to 2 or more inches in length can be formed in only three weeks in
the water. (6) Such rapidity of growth and differentiation of plants by natural
proliferation suggest that many of them would be well prepared for establish¬
ment once they drifted to an appropriate land surface along the shore.

In another experiment we compared rates of production of prolifera¬
tions in water with those on wet sand. On October 3, 1969, we collected
approximately 40 culms and divided them into two groups. One group was
floated in water, the other placed in wet sand on a misting bench in a
research greenhouse. Those floated in water were adjacent to the culms on the
mist bench. No changes were noticed until October 13, when 4 of the stems
in the water, but only 1 on sand, showed primordial bulges. By October 21,
all except one of those on water had formed distinct culms, but less than half
of the stems on sand had evidence at all of proliferation, mainly as mere
primordia. On October 24, all of the stems on water had definite shoots as
well as roots; 75% of those on sand, in contrast, showed evidence of beginning
root growth, but only a few specimens had shoots. These observations suggest
that proliferation occurs more rapidly on water than on wet sand, at least
under the conditions of our experiment.

DISCUSSION

We have described here the consequences of fragmentation by natural
causes (cutting of culms by muskrats) of the common scouring-rush, E.
hyemale, which gave rise to the flotation of numerous pieces on a small lake.
These fragments proliferated shoots and/or roots at the nodes, some of these
shoots even forming strobili at their apices. A tabulation of nearly 90 stem
fragments revealed that all, even those with only two nodes, gave rise to
proliferations. Large fragments with up to 10 or more nodes showed a tend¬
ency toward terminalization of the proliferations, as well as a distinctive
polarity. Proliferation was greatest near the ends and least in the middle. The
morphologically apical parts produced shoots but very few roots, but the basal
parts produced both shoots and roots, indicating that an innate morphogenetic
polarity is present that is unaffected by the horizontal orientation of the
floating stem fragments. Experimental tests in selected aquatic sites revealed
that well formed proliferations could arise in three weeks, that proliferation
could occur in shade, in rapidly moving water, and even when the stem
fragment was submerged. The muskrat has evidently not previously been re¬
ported as an agent in natural fragmentation in Equisetum. The ability of stem
fragments to proliferate new plants entirely without connection to soil has
apparently been previously overlooked.

Of what significance are these observations? It seems to us that the
ability of the scouring-rush (and very likely related hybrids and species) to

172

THE MICHIGAN BOTANIST

Vol. 9

form fairly well differentiated plants during flotation may have significance in
the efficiency of asexual spread and establishment. Hauke (1963) has written
(in regard to E. Xnelsonii ) that

In northern Michigan . . . where there are many lakes and rivers and hundreds of
miles of sandy shores around the Great Lakes, [E. xnelsonii ] is perhaps the most
widespread of all [scouring rushes], and seems to reproduce readily by frag¬
mentation and water transport of the rhizomes. This mechanism was seen in action
in Douglas Lake, Cheboygan Co., Michigan. At a place called “Pine Point,” a sandy
projection into the lake from its east shore, E. xnelsonii was first collected in 1921.
... In 1957 I went to the same locality and found it in abundance. Wave action
was washing the sand away and rhizomes, attached to the shore, had sections up to
one meter in length floating in the water. ... It was apparent that a violent storm
could tear off and scatter some of these.

Fig. 3. Dried specimens of proliferating culms of the scouring-rush . (See text). 1/5 X
Natural Size.

1970

THE MICHIGAN BOTANIST

173

TABLE 1. Data on 30 floating culms of Equisetum Leftmost column, specimen no.;
next column, length in cm. Node number base to apex along top. SHOOT classes: 1 =
less than Vz cm; 2 = Vz - 3; 3 = 3 - 12; 4 = 12 or greater; * = strobili present.
(Superscripts represent no. of shoot proliferations at each node.) ROOT Classes: 1 =

under 1 cm; 2 = less than 10 roots up to 4 cm; 3 = 10 or more roots up to 4 cm or less
than 5 roots over 4 cm; 4 = 10 roots, more than 5 over 5 cm; 5 = 10 or more roots over
4 cm.

SHOOTS ROOTS

TABLE 2. Summary of shoot and root production at basal 4, apical 4, and medial nodes.
Indices computed for shoots by multiplying classes by superscripts and summing prod¬
ucts, for roots by summing classes.

ROOTS

SHOOTS

Nodes with proliferations
Nodes without proliferations

64

120

53.3

41

120 '

34.2

BASAL 4 NODES

M

39

189

20.6

35

189 ‘

18.5

MEDIAL NODES

II

_ 3

120

2.5

II

H °

1 ^

34.2

APICAL 4 NODES

TOTAL OF

PROLIFERATION INDICES

140

=

191

BASAL 4 NODES

=

68

=

75

MEDIAL NODES

=

7

=

102

APICAL 4 NODES

174

THE MICHIGAN BOTANIST

Vol. 9

TABLE 3. Summary of field experiments. Culms floated on August 14, 1969. N = no
proliferations. P = initial protrusions. S = definite shoots.

Locality

Flow

Light

Aug . 1 8

Aug. 20

Sept. 3

Parker

Brook

Pond

Slow

Moving

Full

Sun

14 N

13 S, 1 P

(All lost)

Fleming

Creek

(Pool)

Slow

Moving

Shade

14 N

10 P, 4 N

14 S

(1 cm-2 cm)

Fleming

Creek

(Pool)

Rapid

Current

Full

Sun

14 N

11 N
(3 lost)

11 S

(1 cm-2 cm)

(All submerged)

Fleming

Creek

(Pool)

Rapid

Current

Shade

14 N

10 P, 4 N

14 S

(3 cm-5 cm;
one 1 3 cm)

Dix Pond

Still

Full

Sun

14 N

12 S, 1 P, 1 N

14 S

(3 cm-5 cm)

Thus Hauke suggests that rhizome fragments, torn off of wave-lashed
shores, could serve as propagules. His experiments concerned with aerial stem
sections demonstrated the potentiality of dispersal without rhizomes. How¬
ever, he planted the artificially cut stems directly on or in more or less moist
soil. Is such layering necessary for proliferation to be initiated? Our observa¬
tions here suggest that whole plants, with shoot and root systems, can be
formed during flotation— across a lake, down a canal or stream— and thus be
immediately prepared for establishment once cast upon an appropriate shore
site.

LITERATURE CITED

Hauke, Richard. 1963. A Taxonomic Monograph of the Genus Equisetum Subgenus
Hippochaete. Nova Hedwigia Beih. 8. 123 pp.

1970

THE MICHIGAN BOTANIST

175

CHANGES IN THE VASCULAR FLORA OF SEVEN
SMALL ISLANDS IN WESTERN LAKE ERIE1

Thomas Duncan and Ronald L. Stuckey
College of Biological Sciences,

The Ohio State University, Columbus

Floristic studies of the herbaceous and woody plants growing without
cultivation have been conducted for several or all of the islands in western
Lake Erie by various authors during the past seventy years. Moseley (1899)
published a flora of Erie County, Ohio, and included the peninsula and islands
in Ottawa County, Ohio. A list of plants for each island cannot be prepared
from Moseley’s flora because he did not always cite the islands where each
species was seen. Schaffner (1902) reported 15 species for Little Chicken
Island. Kellerman (1904) listed 39 species for Hen Island, six species for Big
Chicken Island, and 16 species for Little Chicken Island. For the latter two
islands, the number of plants for each species was listed. Dodge (1914) pre¬
pared a flora of Point Pelee, Canada, and adjacent islands, taking records for
the Ohio islands from Moseley (1899). Malcolm E. Stickney presented two
papers entitled “The census of flowering plants on certain small islands of
Lake Erie” and “The distribution of flowering plants on the smaller islands of
Lake Erie” at meetings of the Ohio Academy of Science in 1921 and 1922,
respectively (titles from Ohio Jour. Sci. 22: 19. 1921; 23: 23. 1923). Al¬
though Core (1948) used lists compiled by Stickney, neither a published nor a
manuscript version of either of these papers has been located. Core listed 261
species from the seven small islands that are considered in this study. We have
seen 231 species, 35 of which were not seen by Core.

Floras are continually in a state of flux and any contemporary floristic
analysis should take into account those changes that have occurred through
time, and the possible factors affecting those changes. In this paper, we com¬
pare the lists of plants prepared by Kellerman in 1903 (published in 1904)
and by Core in 1939, 1940, and 1941 (published in 1948) with the records
compiled by us during 1967, 1968, and 1969, in order to understand and
possibly explain the floristic changes that have occurred on some of the small
islands in western Lake Erie. These small islands are particularly desirable for
a study in the dynamics of a flora, because the number of species on each
island is few, and relatively complete lists for each island can be made.

During the summers of 1967, 1968, and 1969 inventories were prepared
and collections were made of the vascular plants of Hen, Big Chicken, and
Little Chicken Islands in Essex County, Ontario, Canada; and Lost Ballast,

Contribution from the Botany Program (Paper No. 768), the Herbarium, and the
Franz Theodore Stone Laboratory. Presented by the senior author at the 73rd annual
meeting of the Michigan Academy of Science, Arts, and Letters, Wayne State University,
Detroit, 3 April 1970, and at the 79th annual meeting of the Ohio Academy of Science,
Wittenberg University, Springfield, 17 April 1970.

176

THE MICHIGAN BOTANIST

Vol. 9

Fig. 1. Map of the islands in western Lake Erie. The small islands whose flora is discussed
are labeled on the map. All of the other islands are designated according to the following
numbers.

1 Ballast Island

2 Chick Island

3 East Sister Island

4 Kelleys Island

5 Marblehead Peninsula

6 Middle Island

7 Middle Bass Island

8 Middle Sister Island

9 Mouse Island

10 North Bass Island

11 North Harbor Island

12 Rattlesnake Island

13 South Bass Island

14 Sugar Island

15 Pelee Island

16 West Sister Island

1970

THE MICHIGAN BOTANIST

177

Starve, Gibraltar, and Green Islands in Put-in-Bay Township, Ottawa County,
Ohio (Fig. 1). During 1967 and 1968, Green Island was visited twice and
Starve Island once. Additional records and specimens were obtained in 1967
from Starve Island by Billy Isom, Rachel Cox Downing, and Richard Deweese,
and from Big Chicken Island by Jane L. Forsyth. In 1969, Lost Ballast Island
was visited once; Hen, Big Chicken, Starve, and Little Chicken Islands were
surveyed twice; and Green Island studied four times. The flora of Gibraltar
Island was investigated frequently during all three summers because the Franz
Theodore Stone Laboratory is located on this island. All specimens collected
are in the herbarium of either The Ohio State University or the Franz
Theodore Stone Laboratory. From the lists of plants, numbers and/or per¬
centages for each island were calculated for (1) all species seen in our surveys,
(2) the species reported by Kellerman (1904) and Core (1948) but not seen
during our surveys, (3) the species not reported in any previous survey, (4)
the non-indigenous species seen, and (5) the annuals, biennials, and perennials
seen (tables 2-6).

DESCRIPTION OF THE ISLANDS

The islands in western Lake Erie were formed by erosion of preglacial
streams from solid limestone [dolomite] bedrock (Carman, 1946). The bed¬
rock of Starve, Gibraltar, and Green Islands is Put-in-Bay dolomite, whereas
that of Lost Ballast, Little Chicken, Big Chicken, and Hen Islands is Raisin
River dolomite (Carman, 1946). The islands are composed of gravel bars or
beaches, rocky -shore cliffs, thin soil over dolomite bedrock, or a combination
of these (Table 1). Lost Ballast, Little Chicken, Big Chicken, and Starve
Islands are small, low, entirely (or partially) dolomitic gravel bar islands,
whereas Hen, Gibraltar, and Green Islands are larger islands covered primarily
with thin soil over dolomitic bedrock. These three islands have high steep
rocky shore cliffs.

LOST BALLAST ISLAND

Lost Ballast Island is almost circular in outline, with a diameter of
approximately 75 feet. The bedrock is covered with coarse gravel. In 1969,
the island extended no more than one and a half feet above the surface of the

TABLE 1. Approximate Percentage of the Physiographic Areas on Each Island.

Physiographic Area

Island*

Lost

Ballast

Little

Chicken

Big

Chicken

(1)

Starve

(2)

Hen

(53/4>

Gibraltar

(6)

Green

(20)

Gravel bar or beach

100

100

100

50

10

10

—

Rocky-shore cliff

—

—

—

50

15

15

15

Thin soil over dolomite bedrock

—

—

—

—

75

75

85

♦Numbers in parentheses indicate the approximate size of each island in acres.

178

THE MICHIGAN BOTANIST

Vol. 9

Fig. 2. Starve Island; gravel bar in the foreground and shrubs of hackberry, willow, and
dogwood in the background. Photographed on 13 July 1969 by Thomas Duncan.

Fig. 3. Gibraltar Island with high exposed north facing cliffs; surface of the island
covered with woody vegetation. Photographed on 28 June 1969 by Thomas Duncan.

1970

THE MICHIGAN BOTANIST

179

lake. Although Core (1948) reported several herbaceous species from this
island, in 1969 only woody species were seen by us. At one time Lost Ballast
Island was part of Ballast Island, but it was separated by wave action and heavy
storms to the extent that now they are connected only by a submerged gravel
bar (Ryall, 1913). The island has never been inhabited by man, but is the site
of many herring gull nests.

LITTLE CHICKEN ISLAND

In 1969, Little Chicken Island was a bar of coarse gravel about 100 feet
long extending one to two feet above the surface of the lake. The bedrock is
covered primarily with angular rock fragments one to three inches in diameter
(Kindle, 1937). Several herbaceous and a few woody species were reported by
Jones (1902, 1912), Schaffner (1902), Kellerman (1904), and Core (1948),
although in 1969 no species of vascular plants were present. In 1969, many
herring gulls were observed on the island. Little Chicken Island has never been
inhabited by man.

BIG CHICKEN ISLAND

In 1969, Big Chicken Island was approximately 300 feet long and ex¬
tended about nine feet above the surface of the lake at the highest point. The
bedrock of the island is covered primarily with slabby limestone [dolomite]
pebbles from one to six inches in diameter (Kindle, 1937). Vegetation in the
summer of 1969 was entirely herbaceous, although woody species were re¬
ported by Jones (1902, 1912), Kellerman (1904), and Core (1948). An eight-
foot depression which is centrally located on the island contains a collapsed
fishing building. Approximately one foot of water was in the depression in the
summer of 1969, but no species of aquatic vascular plants were observed.
Many pairs of cormorants and hundreds of herring gulls were nesting on the
island. The island has been sporadically inhabited by man.

STARVE ISLAND

Starve Island is approximately two acres in area (Hatcher, 1945). In
1969, the island extended two feet above the surface of the lake at its highest
point. The island is divided into two distinct parts by a gravel bar, which was
submerged in July of 1969, but exposed in August (Fig. 2). Vascular plants
were not seen on the bar at the latter time. The larger part of the island is
approximately one and a third acres in area. About half of the bedrock is
covered with coarse gravel. Glacial grooves are prominent on the south side of
the island. The vegetation is composed of herbaceous and woody plants, with
the former mostly on the gravel portion of the island. The island has never
been inhabited by man. Many herring gull nests were observed in 1969.

HEN ISLAND

Hen Island contains an area of five and three-quarters acres (Kindle,
1937). It is surrounded by low rocky cliffs, at the base of which are five- to
ten-foot shelves extending out and along the surface of the lake. Most of the

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THE MICHIGAN BOTANIST

Vol. 9

dolomite bedrock underlying the island is covered by a thin layer of soil, al¬
though outcrops were observed in several places. The Quinnebog Fishing Club,
which purchased the island in 1888 (Ontario Department of Lands and Forests,
pers. comm., 1969) has erected buildings and planted several herbaceous and
woody species, some of which have escaped cultivation. Some areas of the island
have remained forested, although these areas are now crossed with winding paths.

GIBRALTAR ISLAND

Gibraltar Island covers an area of six acres (Core, 1948). Rocky cliffs
occur on the west, northwest, and north sides of the island (Fig. 3). On the
southwest corner is Alligator Bar. This bar, approximately 100 feet of which
is usually above the surface of the lake, extends from Gibraltar Island to
South Bass Island and separates Fishery Bay and Squaw Harbor. The interior
of the island is a lawn which is continually mowed and where many non-
indigenous species occur. Small stands of trees are located on the northeast
and northwest corners of the island. Trees also occur at various places
throughout the lawn area. The island was purchased from J. D. Rivera St.
Jago in 1864 by Jay Cooke who then constructed a mansion (Cooke Castle),
which is still standing. He, his family, and guests used the island and mansion
as a summer home until 1905 (Pollard, 1935). In 1925, the island was pur¬
chased by Julius F. Stone, who donated it to The Ohio State University
(Langlois, 1949). Since 1925, the island has been the headquarters of the
Franz Theodore Stone Laboratory.

GREEN ISLAND

Green Island is approximately 20 acres in area (Ryall, 1913). The island
is surrounded by cliffs and rocky shores similar to those on Hen Island. The
soil is very black, generally six to eight inches deep (Core, 1948). Although
the island is covered primarily by woody vegetation of secondary successional
growth, leaf litter is practically destroyed by the large snail population and
heavy winter storms (Stansbery, personal communcation, 1969). According to
a notation on an 1835 map (of which a copy has been supplied by Jack
McCormick), the island was purchased in 1853 by the United States govern¬
ment. Green Island was occupied by a lighthouse keeper and his family from
about 1860 (Ryall, 1913) until about 1919 (Nate Ladd, pers. comm., 1969).
The house and remnants of the barn and boathouse used by these people still
remain. During the summers prior to World War II, the island was occupied by
a doctor and his family (Robert Dodge, pers. comm., 1969).

CHANGES IN THE TOTAL FLORA2

The vascular flora on each one of four smal1 islands (Starve, Hen,
Gibraltar, and Green) is undergoing change to the extent that 30-40% of the

9

In the discussion below we have chosen 1903, 1939, and 1969 as the reference
years (the years of the most extensive work) when making comparisons. However, it
should be understood that Core’s records and our records are for three-year periods as
mentioned in the introduction.

1970

THE MICHIGAN BOTANIST

181

TABLE 2. Changes in the Total Flora.

Island

Lost

Little

Big

Ballast

Chicken

Chicken

Starve

Hen

Gibraltar

Green

Number of species seen in:

1903

—

16

6

—

39

—

—

1939

38

18

12

38

68

163

185

1969

14

0

15

63

80

144

138

Percentage of the flora of:*

1903 not seen in 1939

—

62

50

—

30

_

_

GO)

(3)

(12)

1903 not seen in 1969

—

100

83

—

44

—

—

(16)

(5)

(17)

1939 not seen in 1969

68

100

67

30

40

30

34

(26)

(18)

(8)

(ID

(26)

(50)

(64)

Percentage of the flora which

were new records in:

1939

—

75

(12)

75

(9)

—

60

(41)

—

—

1969

13

0

73

57

44

23

13

(2)

(0)

(ID

(36)

(35)

(31)

(17)

*The actual number of species is given in parentheses beneath each percentage.

species seen in 1939 were not seen in 1969 (Table 2). More drastic floristic
changes have occurred on Lost Ballast and the Chicken Islands, where 68% of
the species seen in 1939 on Lost Ballast Island and 67% of the species in
1939 on Big Chicken Island were not seen in 1969. All of the vascular flora
has disappeared from Little Chicken Island. Considering the flora as a whole
for these seven islands, 261 species were noted in 1939, of which only 196
were found in 1969, for a 25% reduction in the total flora. This percentage is
lower than that for any one island, because some species which have disap¬
peared from all of the islands where they were seen in 1939 were seen in
1969 on one or more of the other islands.

Lost Ballast and Little Chicken Islands have undergone a similar change
in that between 1939 and 1969 all of the herbaceous species have disap¬
peared. The disappearance of these herbaceous species probably occurred dur¬
ing a much shorter period of time. This change could have happened during
any one year in which there was strong wave erosion, high water level, or
destructive ice action. Winds and wave action during storms, which can change
the water level of the lake as much as four feet (Langlois, 1965a), would
inundate these islands and destroy the plants. The short period of time in
which the amount of vegetation can change on a gravel bar in the Lake Erie
island area has been documented (Langlois, 1965b). For example, during
periods of high water, the gravel bar between South Bass Island and its north¬
east point, Buckeye Island, or Buckeye Point, is inundated and its vegetation
eradicated. After two or three subsequent years of low water level, the bar
becomes revegetated (Langlois, 1965b). Chick Island, or Chicken Reef, located

182

THE MICHIGAN BOTANIST

Vol. 9

northeast of Big Chicken Island, was covered with a dense stand of Poly¬
gonum sp. in 1901 (Jones, 1902, 1912), but these plants had disappeared by
1903 (Kellerman, 1904). During this two-year period, wave erosion and ice
action, accompanied by a one-foot rise in mean water level (U. S. Lake Sur¬
vey, 1969), was probably sufficient to eradicate the vegetation. In 1969, a
year of high water level, Chick Island was no more than a few boulders
protruding above the surface of the water.

The flora of both Little Chicken and Big Chicken Islands has undergone
considerable flux over the past 70 years. For example, on Little Chicken
Island, 62% of the species seen in 1903 by Kellerman were not seen in 1939
by Core, and none of these species were seen by us. On Big Chicken Island,
50% of the species seen in 1903 by Kellerman were not seen by Core in 1939,
and 67% of the species noted in 1939 by Core and 83% of the species noted
in 1903 by Kellerman were not seen by us. In 1903, Big Chicken Island had
six species and in 1969, 15 species were present. The survival of a flora on Big
Chicken Island, especially during a year of high water level as occurred in
1969, and not on Little Chicken Island is apparently related to certain physio¬
graphic features of Big Chicken Island that are not present on Little Chicken
Island. Big Chicken Island is partially protected from wave erosion and ice
action by a gravel storm beach extending approximately nine feet above the
surface of the lake on the northeast side of the island. Behind the storm
beach the surface slopes to the southwest where plants are in a habitat which
is not directly exposed to severe northeast storms. Little Chicken Island, how¬
ever, is but a ridge of gravel lying in an east-west direction which extends
approximately two feet (in 1969) above the surface of the water. Its position
is particularly vulnerable to the destructive action of storms.

A history of some of the species on Big Chicken and Little Chicken
Islands can be reconstructed for the years between 1901 and 1969. The bur
cucumber, Sicyos angulatus, one of the most abundant plants on Big Chicken
Island in 1969, was seen there by Jones in 1901. Kellerman (1904) reported
75 small plants of this species. Since then, Kennedy (1922), Core (1948), and
Forsyth (specimen label, 1967, OS) have stated that the bur cucumber is one
of the predominant species on Big Chicken Island. One box-elder maple, Acer
negundo, 12 inches high was reported by Kellerman on Big Chicken Island.
Core also listed this species for Big Chicken Island. Forsyth (specimen label,
1967, OS) noted a small bent over, beaten tree. In 1969, Acer negundo was
seen by members of the ornithology class from the Franz Theodore Stone
Laboratory during the latter part of June, but the tree was not seen by us in
the later part of July. The tree probably was blown away during the severe
storm of 4 July 1969. The most abundant species on Big Chicken Island in
1969 was Lepidium virginicum , pepper-grass, an annual weed. It was not seen
on the island in 1903 by Kellerman, but was reported by Core (1948). On
Little Chicken Island, Salix amygdaloides, peach-leaved willow, was seen by
Jones in 1901, and Kellerman reported 50 trees of varying sizes. Kindle
(1937) and Core also reported this species, but the peach-leaved willow was
not present in 1969.

1970

THE MICHIGAN BOTANIST

183

The small gravel bar islands, Lost Ballast, Little Chicken, Big Chicken,
and Starve, are not only disturbed by the waves and ice, but they are also
inhabited by large concentrations of herring gulls and other shore birds during
the summer. The large number of birds probably provides a significant means
for the dissemination of plant propagules to the islands. The large amount of
guano left on the islands could be significant for the survival of many of the
plant species and account for some of the changes noted in species composi¬
tion.

From 1939 to 1969, the number of species has increased from 38 to 63
on Starve Island and from 68 to 80 on Hen Island. Conversely, on Gibraltar
Island and Green Island, the number of species has decreased from 163 to 144
and from 185 to 138, respectively. However, if a comparison is made between
the number of species present in 1939 with the number of those same species
present now, we find that 3040% of these species on each one of these four
islands have disappeared (Table 2). The reasons for these changes are not
evident when viewing the total flora, but apparent reasons can be suggested by
a study of the relationships between the longevity of the plants, their status in
the flora, and the disturbance that occurs on these islands.

RELATIONSHIPS BETWEEN LONGEVITY OF PLANTS,

THEIR STATUS IN THE FLORA, AND DISTURBANCE

The longevity (annual, biennial, or perennial) and apparent status (in¬
digenous or non-indigenous) of each species in the flora are included in Table
6. The percentage of annuals has increased substantially between 1939 and
1969 on Big Chicken, Starve, and Hen Islands; has remained the same on
Gibraltar Island; and has decreased slightly on Green Island (Table 3). From
1939 to 1969, the percentage of non-indigenous species has increased
considerably on three of the islands, Big Chicken, Starve, and Hen (Table 3).
The percentage of non-indigenous species has increased slightly on Gibraltar
Island, and has remained the same on Green Island (Table 3).

On two of the small gravel bar islands, Big Chicken and Starve, the
non-indigenous species are primarily annuals, whereas the indigenous species
are mostly perennials (Table 4). One would expect a short-lived annual to
survive and reproduce in a disturbed habitat more favorably than a perennial.
Based on the percentage of species that have apparently invaded (numbers of
new records) since 1939, a trend exists toward the elimination of the in¬
digenous perennials and the invasion and establishment of non-indigenous an¬
nuals. For example, 55% of the new records for Big Chicken Island are non-
indigenous annuals and none are indigenous perennials (Table 5). Indigenous
perennials such as Acer negundo, Celtis occidentalis, Sambucus canadensis,
Scirpus fluviatilis, Urtica dioica, and Verbena hastata have disappeared.
Non-indigenous annuals such as Amaranthus retro flexus, Capsella bursa-
pastoris, Chenopodium album, Digitaria sanguinalis, Lactuca scariola, Portulaca
oleracea, Sonchus arvensis, and S. asper constitute the majority of the present
flora. Starve Island, whose area is about half gravel bar, is showing a trend
toward an increase in both indigenous and non-indigenous annuals and a de-

184

THE MICHIGAN BOTANIST

Vol. 9

TABLE 3. Changes Related to the Longevity of the Plants and their Status in the Flora.

Island

Lost

Ballast

Little

Chicken

Big

Chicken

Starve

Hen

Gibraltar

Green

Percentage of the flora which

was: *

a) non-indigenous species in:

1903

—

12

17

—

29

—

—

(2)

0)

(ID

1939

24

39

50

26

30

34

26

(9)

(7)

(6)

(10)

(22)

(55)

(48)

1969

22

0

73

40

48

37

26

(3)

(0)

(11)

(25)

(38)

(54)

(36)

b) annuals in:

1903

—

62

17

—

23

—

—

(10)

0)

(9)

1939

32

44

50

24

16

26

18

(12)

(8)

(6)

(9)

(ID

(42)

(34)

1969

0

0

80

44

34

26

14

(0)

(0)

(12)

(28)

(27)

(37)

(19)

c) perennials in:

1903

—

38

83

—

72

—

—

(6)

(5)

(28)

1939

68

56

42

74

81

71

78

(26)

(10)

(5)

(28)

(55)

(116)

(144)

1969

100

0

13

52

62

70

80

(14)

(0)

(2)

(33)

(50)

(101)

(111)

d) biennials in:

1903

—

0

0

—

5

—

—

(0)

(0)

(2)

1939

0

0

8

3

3

3

4

(0)

(0)

0)

0)

(2)

(5)

(7)

1969

0

0

7

3

4

4

6

(0)

(0)

0)

(2)

(3)

(6)

(8)

*The actual number of species is given in parentheses beneath each percentage.

crease in both indigenous and non-indigenous perennials. Of the species that
are listed as new records for Starve Island in 1969, 53% are annuals and 45%
are perennials (Table 5). The gravel bar habitat of Starve Island, located most¬
ly in the center of the island, is apparently less disturbed than Big Chicken
Island. Another part of Starve Island, namely the bedrock shore, is consider¬
ably disturbed. Compared to Big Chicken Island, the percentage of invading
non-indigenous annuals has been lower on Starve Island (Table 5).

On the three larger islands, Hen, Gibraltar, and Green, the non-indig¬
enous species are both annuals and perennials and the indigenous species are
mostly perennials (Table 4). Perennials, once established, would have a better

1970

THE MICHIGAN BOTANIST

185

TABLE 4. Longevity of Non-Indigenous and Indigenous Species

Island

Lost

Ballast

Little

Chicken

Big

Chicken

Starve

Hen

Gibraltar

Green

Percentage of non-in¬
digenous species which
were annuals in:*

1903

—

50

(1-2)

0

(0-1)

—

55

(6-11)

—

—

1939

33

(3-9)

43

(3-7)

50

(3-6)

50

(5-10)

27

(6-22)

43

(23-55)

28

(17-48)

1969

0

(0-3)

0

(0-0)

73

(8-11)

60

(15-25)

42

(16-38)

43

(23-54)

33

(11-36)

Percentage of indig¬
enous species which
were perennials in:**

1903

—

36

(5-14)

80

(4-5)

—

86

(24-28)

—

—

1939

79

(22-28)

45

(5-11)

50

(3-6)

86

(24-28)

80

(37-46)

83

(90-108)

85

(117-137)

1969

100

(11-11)

0

(0-0)

0

(0-5)

68

(26-38)

80

(33-41)

73

(69-94)

89

(91-102)

*The first number in parentheses is the number of non-indigenous annuals and the second
is the total number of non-indigenous species.

**The first number in parentheses is the number of indigenous perennials and the second
is the total number of indigenous species.

chance of surviving on these larger islands because the surface is little dis¬
turbed by wave erosion and ice action. Although man has had considerable
influence on these larger islands, his actions have probably not disturbed the
entire islands to the extent that the small gravel bar islands have been dis¬
turbed by environmental influences. The overall decrease in the number of
species for Gibraltar Island is probably a result of continued cutting, maintain¬
ing, and opening of the habitat by the staff of the Franz Theodore Stone
Laboratory. The non-indigenous component of the flora has been rather stable
over the past 30 years (Table 3), with the greatest fluctuation probably occur¬
ring with the occupancy of the island by The Ohio State University personnel
during the years immediately following the establishment of the laboratory.
The species that have moved in over the past 30 years are mostly perennials
(20 of 31 new records, or 64%, Table 5).

Although Core (1948) stated that Green Island has the closest approxi¬
mation to the natural vegetation of any island, the 1885 map shows that 7.21
acres (42% of the island) were occupied by pasture, lawn, or garden. There¬
fore, a larger non-indigenous species component probably existed at the turn
of the century or before, than has occurred on the island since man left in
1919. With increased secondary successional growth and a trend toward more
stable habitats, the once man-disturbed habitats on Green Island are becoming

186

THE MICHIGAN BOTANIST

Vol. 9

TABLE 5. Longevity and Status of the Plants that are New Records for Each of the Islands
in 1969

Island

Lost

Ballast

Little

Chicken

Big

Chicken

Starve

Hen

Gibraltar

Green

Percentage of the new
records which are:*

a) non-indigenous
annuals

0

(0-2)

0

(0-0)

55

(6-11)

31

(11-36)

40

(14-35)

16

(5-31)

18

(3-17)

b) non-indigenous
perennials

50

(1-2)

0

(0-0)

9

(Ml)

14

(5-36)

20

(7-35)

29

(9-31)

12

(2-17)

c) indigenous annuals

0

(0-2)

0

(0-0)

27

(3-11)

22

(8-36)

17

(6-35)

16

(5-31)

12

(2-17)

d) indigenous peren¬
nials

50

(1-2)

0

(0-0)

0

(0-11)

31

(11-36)

17

(6-35)

35

(11-31)

53

(9-17)

*The first number in parentheses is the number of new records in this category and the
second is the total number of new records.

fewer, such that the establishment of non-indigenous annuals is apparently
more difficult, but indigenous perennials are becoming re-established. Of the
new records for Green Island since 1939, 3 of 17 species (18%) are non-
indigenous annuals and 9 of 17 (53%) are indigenous perennials (Table 5).

The trends seen from the data for Gibraltar and Green Islands differ
from those shown in the data for Hen Island. A gradual increase in the
numbers of species, from 39 in 1903 to 68 in 1939 and to 80 in 1969 has
occurred (Table 2). Since 1939, this increase is accounted for mostly by
non-indigenous annuals (14 of 35 new records or 40%, Table 5). The changes
in the flora on Hen Island are probably related to man’s increased use of this
rather remote island in the past thirty years.

CHANGES RELATED TO THE SIZE AND
PHYSIOGRAPHY OF THE ISLANDS

The percentage of species new to any island (new records) is higher on
the small gravel bar islands and lower on the larger islands. Big Chicken Island,
one of the small gravel bar islands, has the highest percentage of new records
(73%). Green Island, the largest island studied, has the lowest percentage of
new records (13%). A comparison of island size and percentage of new records
can be made from the information in Tables 1 and 2. This trend also correlates
with the amount of disturbance on the islands. The habitats on the gravel bar
islands are continually undergoing disturbance (particularly from wave erosion
and ice action), whereas the habitats on the large islands are more stable.
Because of the severe environmental influences and physiographic features
(extreme small island size and slight elevation above the surface of the water),
these correlations are not shown for Lost Ballast and Little Chicken Islands.

1970

THE MICHIGAN BOTANIST

187

SUMMARY

Comparisons of the floras of seven small islands in western Lake Erie have shown
that (1) 30-40% of the species on Starve, Hen, Gibraltar, and Green Islands; 68% of the
species on Lost Ballast; 67% of the species on Big Chicken; and all of the species on
Little Chicken Island have disappeared since 1939; (2) the percentage of annuals and
non-indigenous species has increased and the percentage of perennials and indigenous
species has decreased for most of the islands; (3) of the species that have invaded in the
past 30 years, annuals have apparently been more successful in becoming established on
the disturbed gravel bar islands, whereas perennials have been most successful in becom¬
ing established on the larger islands; and (4) the percentage of species new to any island
is higher on the small gravel bar islands and lower on the larger islands. Some of the
apparent reasons for these floristic changes are the influences of wave erosion, ice action,
fluctuating water level, and bird inhabitation on the small gravel bar islands. Man’s in¬
fluence and general vegetation succession have been important factors in the floristic
changes on the larger islands.

ACKNOWLEDGMENTS

We are grateful to the Ohio Biological Survey for financial support of field work
and partial publication costs. We also thank the Director of the Franz Theodore Stone
Laboratory for providing transportation. Several students, Miss J. Wynne Jones, Mr. W.
Louis Phillips, and Mr. Robert Haynes, assisted in the field work.

LITERATURE CITED

Braun, E. Lucy. 1967. The Monocotyledoneae [of Ohio]. Cat-tails to Orchids. With
Grainineae by Clara G. Weishaupt. Ohio State Univ. Press, Columbus. 464 pp.

Carman, J. Ernest. 1946. The geologic interpretation of scenic features in Ohio. Ohio
Jour. Sci. 46: 241-283.

Core, Earl L. 1948. The Flora of the Erie Islands An Annotated List of Vascular Plants.

Ohio State Univ., Franz Theodore Stone Lab. Contr. 9. 106 pp.

Dodge, C. K. 1914. Annotated List of Flowering Plants and Ferns of Point Pelee, Ont.,
and Neighbouring Districts. Canada Dep. Mines. Geol. Surv. Mem. 54, No. 2 Biol. Ser.
131 pp.

Fernald, Merritt Lyndon. 1950. Gray’s Manual of Botany. 8th ed. American Book Co.,
New York, lxiv + 1632 pp.

Gleason, Henry A. 1952. The New Britton and Brown Illustrated Flora of the North¬
eastern United States and Adjacent Canada. N. Y. Bot. Gard., New York. 3 vols.
Hatcher, Harlan. 1945. Lake Erie. Bobbs-Merrill Co., New York. 416 pp.

Jones, Lynds. 1902. The summer birds of Lake Erie’s Islands. Ohio Nat. 2: 281-284.

- . 1912. A study of the avifauna of the Lake Erie Islands. Wilson Bull. 14:

6-18.

Kellerman, W. A. 1904. Flora of Hen and Chicken Islands, 1903. Ohio Nat. 4: 190-191.
Kellerman, W. A., and Mrs. 1900. The non-indigenous flora of Ohio. Ohio State Univ.
Bull. Ser. 4(27): 1-28.

Kennedy, C. H. 1922. The ecological relationships of the dragonflies of the Bass Islands
of Lake Erie. Ecology 3: 323-336.

Kindle, K. M. 1937. Geology of Pelee and adjacent islands, Ann. Rep. Ontario Dep.
Mines 45(7): 75-116.

Langlois, Thomas H. 1949. The Biological Station of the Ohio State University. Ohio
State Univ., Franz Theodore Stone Lab. Contr. 11. 64 pp.

- . 1965a. The waves of Lake Erie at South Bass Island. Ohio Jour. Sci. 65:

335-342.

- . 1965b. Ecological processes at a section of shoreline of South Bass Island,

Lake Erie. Ohio Jour. Sci. 65: 343-352.

188

THE MICHIGAN BOTANIST

Vol. 9

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. pc Lost Ballast Little Chicken Big Chicken Starve Hen Gibraltar Green

sPecies Longevity & Status 39 69 03 39 69 Q3 39 69 39 69 Q3 39 69 39 69 39 69

196

THE MICHIGAN BOTANIST

Vol. 9

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_ TABLE 6 (Continued) _ _ _ _ _

Lost Ballast Little Chicken Big Chicken Starve Hen Gibraltar Green

Species Longevity & Status 39 69 03 39 69 Q3 39 69 39 69 03 39 69 39 69 39 69

198

THE MICHIGAN BOTANIST

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THE MICHIGAN BOTANIST

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200

THE MICHIGAN BOTANIST

Vol. 9

McDonald, Sr. Elizabeth Seton. 1937. The life-forms of the flowering plants of Indiana.
Am. Midi. Nat. 18: 687-773.

Moseley, E. L. 1899. Sandusky Flora. A Catalogue of the Flowering Plants and Ferns
Growing without Cultivation, in Erie County, Ohio, and the Peninsula and Islands of
Ottawa County. Ohio Acad. Sci. Spec. Pap. 1. 167 pp.

Muenscher, Walter Conrad. 1949. Weeds. Macmillan Co., New York. 579 pp.

Pollard, James E. 1935. The Journal of Jay Cooke or the Gibraltar Records 1865-1905.
Ohio State Univ. Press, Columbus. 359 pp.

Ryall, Lydia J. 1913. Sketches and Stories of the Lake Erie Islands. American Publishing
Co., Norwalk, Ohio. 546 pp.

Schaffner, J. H. 1902. The flora of Little Chicken Island. Ohio Nat. 3: 331-332.

U. S. Lake Survey, Corps of Engineers, U. S. Army. 1969. Hydrograph of monthly mean
levels of the Great Lakes. U. S. Army Engineer District Lake Survey, Detroit,
Michigan.

A REPORT ON THE 1969 MOREL SEASON IN MICHIGAN

Ingrid Bartelli

Upper Peninsula Extension Center, Michigan State University,

Marquette, Michigan

Approximately 250 people from scattered areas of the State of Michigan
reported on their success (or lack of it) gathering morel mushrooms in the
spring of 1969. A questionnaire was mailed to people who had at one time or
another attended one of our mushroom workshops. Their reports and com¬
ment should help us prepare for morel seasons to come.

Respondents replied to the following questions:

1. Approximately how many morels did you harvest this year? 2. Which
kinds were they? 3. Would you say this was a very poor, poor, average, or
better than average year? 4. What were the approximate dates when picking
was the best? 5. In what kind of territory do you find your best picking (tree
cover, slope, exposure, etc.)?

The manner in which the volume of harvest was reported varied. Some
people counted each fruiting body with the numbers varying from “one, only
one” to over 2,000. Amounts were also indicated in meals, pounds, quarts,
gallons, pecks, and bushels.

It was interesting to note that even though people in some areas re¬
ported a harvest of several bushels of morels they rated the season as very
poor. This was particularly apparent in reports coming from the northern part
of Lower Michigan. By comparison, a harvest of several bushels would be
considered a very good season in most parts of the Upper Peninsula.

To the question, “What kinds were they?” answers varied: Such termi¬
nology as black morels, white morels, brown morels, closed-caps, open-caps,

1970

THE MICHIGAN BOTANIST

201

loose-caps, beefsteaks, gray morels, early-capped, late gray, tan morels, open-
stem or woods mushroom, sponge mushroom, early morel, late morel, light
beige morel, large light morel, sandy sponge morel, cone-type morel, grayish
brown morel, large cinnamon morel, pine tree morel, and elephant ears. One
could only guess at the species when tabulating results.

Some respondents used scientific names, particularly in the Traverse City
and Sault Ste. Marie areas. They listed Morchella angusticeps, Morchella
esculenta, Morchella crassipes, Morchella hybrida, Verpa bo he mica, Helvella
esculenta, Helvella gigas, and Morchella deliciosa. This added materially to the
value of their information.

The species distribution in the state made an interesting study. In the
Upper Peninsula the predominant species harvested is Morchella angusticeps
(black morel), but a substantial volume of Helvella esculenta is harvested and
eaten. Verpa bohemica is prolific in the U. P. and Helvella gigas is reportedly
collected there in volume. Very few reported harvest of Morchella esculenta
(white morel) in the U. P., though we know they do grow in the area because
of a few reported large-volume harvests.

In the northern half of the Lower Peninsula, Morchella esculenta (white
morel) is the predominant species. Morchella angusticeps contributes some
volume to the harvest. Verpa bohemica (often reported as the loose or open-
cap morel) is harvested in abundance as is Helvella esculenta. There are some
reports of Morchella crassipes being harvested also.

In the southern half of the state, Morchella esculenta and Morchella
crassipes were the predominant species in the reported harvest. Very few
reported any volume of Morchella angusticeps.

When asked to evaluate the season, the respondents were practically
unanimous in the rating for their area.

In the U. P., the season for Morchella species was rated poor to very,
very poor. The season for Verpa bohemica was better than average. I saw one
four-bushel harvest of Verpa bohemica picked near Marquette in a two-day
period and which was being transported to Ohio for consumption. Three
people indicated that Verpa bohemica made them ill. Those who pick Helvella
esculenta reported an average to good season.

In the northern half of the Lower Peninsula, respondents rated the
season as average and above average for Morchella esculenta and very poor for
Morchella angusticeps. A heavy fruiting of Verpa bohemica was reported also.

In the southern half of the Lower Peninsula, respondents to this survey
reported a very poor to average season. Morchella esculenta and Morchella
crassipes were the dominant species. The poor season seemed to be attributed
to competition among collectors as much as to the actual fruiting.

Most interesting was the reported time of optimum harvest for the 1969
season. In the entire state, best picking was between the second week and the
end of May. Apparently the species most abundantly harvested ( Morchella
crassipes in the southern part of the state, Morchella esculenta in the northern
half of the Lower Peninsula, and Morchella angusticeps in the Upper Penin¬
sula) all fruit at approximately the same time. We know if we were limited to

202

THE MICHIGAN BOTANIST

Vol. 9

any one area we would look first for Helvetia gigas, Verpa bohemica, and
Helvetia esculenta, followed by Morchella angusticeps, then Morchella
esculenta, and last Morchella crassipes.

As would be expected in a survey of the entire state, there were re¬
ported collections in late April to mid June. Generally, the weather was
blamed for the poor season. “Too cold” in all areas and “too dry and too
cold” in some southern and most northern areas. Answers to “In what kind of
territory did you find your best picking?” were naturally general in nature.
No one will lead you to his patch.

In the southern half of the Lower Peninsula, harvesting was best in rich
river-bottom land, ash stands, old orchards, hedge rows, and on wooded slopes
and ridges. Dead or dying elms were indicated present in almost all of the best
areas. {Morchella esculenta and Morchella crassipes were the main species
harvested.)

The terrain varied in the northern half of the Lower Peninsula where the
harvest included Verpa bohemica , Helvetia esculenta , Morchella esculenta, and
a few Morchella crassipes. Best collecting areas indicated for Morchella
angusticeps were wooded slopes, black cherry stands, borders of marshes and
orchards. Morchella esculenta and Morchella crassipes were harvested from elm
slashings, grassy fields, high areas and ridges of wooded slopes with sandy soil,
north facing slopes covered with mixed, sparse, young tree cover and orchards.
Helvetia esculenta was harvested from conifer plantations.

In the U. P., where most of the harvest is Morchella angusticeps, areas
for best picking included wooded slopes and ridges of young mixed tree
growth, aspen slashings (1-, 2-, and 3-year old), berry bramble, fern-filled
fields, old orchards, slashings next to swamps, old pastures, aspen groves, and
along logging roads. Several collectors reported good harvest in Norway Pine
plantations as late as June 15. Mushrooms are easily spotted in a bed of pine
needles. Perhaps we stop looking too soon.

Some scattered reports of good collecting for Morchella esculenta (white
morel) came from areas in the U. P. adjacent to Lakes Michigan and Huron. A
few came from inland areas and an occasional one from areas near the Lake
Superior shore. The types of areas included elm thinnings, along ditches and
right-of-way of gravel roads, and in stands of elm and ash. Verpa bohemica
was collected in quantity on aspen and hardwood covered slopes and along
borders of swamps. Helvetia esculenta was collected mostly in evergreen tree
cover, predominantly pines.

CONDENSATION OF MOREL HARVEST REPORTS, 1969 SEASON, BY AREAS
Species interpretation used by tabulator:

Morchella angusticeps - included those termed blacks, pine-cones, cone-mushroom.

Morchella esculenta - whites.

Verpa bohemica - loose-caps, open-caps.

Helvetia esculenta - beefsteaks.

Southwest Lower Michigan (reports from Niles and Sherwood vicinities):

Very poor season— dry cold weather— best picking between May 11 and May

30 -Morchella angusticeps, Morchella esculenta, and Morchella crassipes found near

cherry and elms, on partially wooded slopes and river bottoms.

1970

THE MICHIGAN BOTANIST

203

Southeast Lower Michigan (reports from Ann Arbor, Fenton, Flint, Holly, Lambertville,
and Davison):

Season rated very poor to good -best picking during last two weeks of May-
Morchella esculenta and Morchella crassipes collected near elms, in hedge rows,
along river banks, on wooded slopes, among landscape plantings- some Morchella
angusticeps found on ash and aspen covered slopes.

Central West Side of Lower Michigan (reports from Baldwin, Bitely, Onekama, Irons,
Manistee, Reed City, Scottsville, Branch, Evart, Luther, Tustin, Ludington, and
Cadillac):

Season reported to be “worst in 20 years”— poor to very poor by two-thirds of
respondents, average by one-third— too cold and dry -better for whites than
blacks -Morchella angusticeps, Morchella esculenta, and Helvella esculenta collected
from last week in April through May with best picking last two weeks of May-
collections made on wooded slopes (hardwood and aspen cover), near dead or
dying elms (for Morchella esculenta >, orchards, and open woods.

Northwest Lower Lower Michigan (reports from Frankfort, Northport, Maple City,
Traverse City, Gaylord, Williamsburg, Omena, Empire, and Mayfield):

Season rated as very poor to poor for Morchella angusticeps (unanimous) and
average to better than average for Morchella esculen ta -harvest began as early as
May 1 and extended to June 9 with best picking May 10-25, peaking on May
16 -Morchella angusticeps collected on wooded slopes, near base of hills, orchards,
marsh borders, in black cherry -Morchella esculenta (as much as 5 bushels) col¬
lected from elm slashings, grassy fields, high areas on wooded sandy slopes, and
near basswood.

Eastern Upper Peninsula, etc. (reports from Goetzville, Paradise, Cedarville, Pickford,
Hessel, Hill Island, Pickford, St. Ignace, Sault Ste. Marie (Michigan & Ontario), Echo
Bay (Ontario), Engadine, Naubinway, Newberry, and McMillan):

Season rated as very poor for all species except Verpa bohemica and Helvella
esculenta -good to better than average for Verpa bohemica, Helvella gigas, and
Helvella esculenta -Morchella angusticeps collected among tree cover (mixed aspen,
beech and maple), old orchards, along edges of woods roads, open areas on high
ground (those on lower ground froze), along ridges of small second growth hard¬
wood and among adder’s tongue and violets. A few Morchella esculenta were
harvested in ditch and along right-of-way of gravel roads -Helvella esculenta col¬
lected in pines and under evergreens along Lakes Huron and Michigan shores.
Collecting extended from last of April till June 3 with second week in May
reported best.

Central and Southern Upper Peninsula (reports from Manistique, Escanaba, Caspian, Iron
Mountain, Gladstone, Cornell, Bark River, Rapid River, Schaffer, Ishpeming, Mar¬
quette, Negaunee, North Lake, Gwinn, Republic, Munising, Wetmore, and AuTrain):
The season was unanimously rated very, very poor to poor because of the cold dry
weather -Morchella angusticeps predominant species collected-average to better
than average year for Verpa bohemica-xaie reports of Morchella esculenta -collect¬
ing best during second and third week of May— found in aspen slashings, mixed
tree cover (aspen, pine, balsam, and maple), slashings next to swamps, sunny wooded
slopes, under small evergreens, aspen groves (frozen in all the exposed areas). Two
unusual dates were reported: On June 15, substantial quantities of Morchella angus¬
ticeps were collected from a 25-year old Norway Pine plantation. At the end of
October 1968, Morchella angusticeps was gathered from along a river bank.

North and Western Upper Peninsula (reports from L’Anse, Skanee, Chassell, Atlantic
Mine, Houghton, Winona, Baraga, Ontonagon, Bruce Crossing, Ironwood, and White
Pine):

A very poor season for Morchella angusticeps- better for Helvella gigas, Helvella

204

THE MICHIGAN BOTANIST

Vol. 9

esculenta, and Verpa bohemica- best collecting during last two weeks of May with

15 and 16 rated best days-found in old pastures, along fence rows next to woods,

among ferns in old fields, aspen groves, ridges, and slopes in young mixed hard¬
wood, aspen, conifer stands.

SELECTED COMMENTS FROM RESPONDENTS

Almost everyone wanted more help in learning how to identify, collect,
prepare, and preserve other mushrooms, edible greens, berries, cherries, nuts,
roots, etc. Respondents wanted to learn about poisonous as well as edible plants.
Many shared experiences, advice, and concerns.

“The whole mushroom family interests me, yet I am frightened each year at the
number of people who pick them without really knowing much about them.”

“Why not encourage more people to harvest and preserve Armillaria mellea and
Naematolotna in autumn? We have about three years out of five when these ‘stump
mushroom’ species could be harvested by the ton. They are good-flavored, easily clearned
in large quantities if picked clean with a knife, easily canned. They are picked in large
quantities by a few people; more should know about them. Couldn’t these mushrooms be
glamorized some way? Many of my ‘shaggy-mane’ picking friends don’t know about these
other species.” Many asked “when to collect” with clues given in the news and on the
radio.

Several resented spraying. “Stop spraying with DDT. Wild land does not need it.”

“No doubt, it’s beyond your scope, but the indiscriminate spraying with herbicide
by the . . . Electric Association hurts any conservation efforts. I stress the word indiscrim¬
inate-like spraying by our gate of an area where no tree branches touched the high line,
but it was there that the rare and protected Nodding Trillium grew. I haven’t had the
heart to look because I know the spray got to the trilliums. So needless-such spraying.
In heavy growth of seedling maples I can understand the spraying, but not in open areas.
I haven’t looked in another open area where arbutus thrives, but I’ll wager it was
sprayed. Yes, we need electricity and help for ‘brushing out’ is scarce these years, but
spraying an open area seems senseless.”

A need for maps and rules was expressed. “I have been wondering if there are
maps of the woods. There are many trails which twist and turn.” “Rules of harvesting,
such as what can be dug up and transplanted, are needed.”

From Iowa: “There is active mushrooming in Iowa. Many, many people collect
here; morels are available in the supermarkets from local collectors at 80 cents per half
pound.”

“I have broken morels and scattered them where I think they would grow (for 33
years) and they do grow there where I planted them in approximately 80 per cent of the
places. I usually pick an area where the humus is rich and rotting and rotted wood is
about, plus rotted poplar leaves.”

“I pick mushrooms more for the enjoyment of being in the woods and out to find
them. Last fall a friend and I picked eight bushels of the stump mushrooms. We did this
in about two and one-half hours. We left many behind in the small area we found them
in; it was great picking.”

“Morels appear about the time the ground brush is starting to leaf out and popple
trees are beginning to green, providing proper amount of rain makes soggy walking on
dead leaves. When ferns and ground cover start vigorous growth, morels are finished.”

“Have the mushroom sellers pay a tax as the byuers. It’s getting to be quite a
racket. Seems to be less mushrooms every year and more pickers. It seems the people
who take them out of state should have to pay a license. Every time anything is free
they come up.”

1970

THE MICHIGAN BOTANIST

205

MICHIGAN PLANTS IN PRINT
New Literature Relating to Michigan Botany

A. MAPS, SOILS, GEOGRAPHY, GEOLOGY

The following topographic maps for Michigan have been published by the U. S.
Geological Survey, 1200 S. Eads St., Arlington, Virginia 22202, since the previous listing
in our January 1969 issue. All are IVi-minute quadrangles (scale of 1:24,000 or about Vh
inches to a mile). Maps are supplied with green overprint showing wooded areas unless
request is made to the contrary, and are $.50 each, with a 20% discount on orders
totalling $20.00 or more. Revised maps are marked with an asterisk in the list below;
others are new or resurveyed. Following the name of the quadrangle, the county or
counties in which it primarily lies are added.

Bay City [Bay, Saginaw]

Bay City NE [Bay]

Berkey* [Lenawee, plus Ohio]

Brich Run N. [Saginaw]

Bridgewater [Washtenaw]

Butternut [Montcalm]

Caledonia [Kent]

Carleton* [Monroe, Wayne]

Carson City [Gratiot, Montcalm]

Cedar Springs [Kent]

Cedar Springs SW [Kent]

Crump [Bay]

Cutlerville [Kent]

Erie* [Monroe]

Essexville [Bay]

Estral Beach* [Monroe]

Flat Rock* [Monroe, Wayne]

Flat Rock NE* [Wayne; formerly
Detroit -Wayne Airport]

Grand Rapids E [Kent]

Grand Rapids W [Kent]

Hubbardston [Clinton, Ionia]

Kalamazoo [Kalamazoo]

Kalamazoo NE [Kalamazoo,

Allegan]

B. BOOKS, BULLETINS, SEPARATE PUBLICATIONS

Arend, John L., & Harold F. Scholz. 1969. Oak Forests of the Lake States and their
Management. North Central For. Exp. Sta., U. S. For. Serv. Res. Pap. NC-31. 36 pp.
[Includes data on oak stands in Michigan].

Kapp, Ronald O. 1969. How to Know Pollen and Spores. Wm. C. Brown, Dubuque. 249
pp. $3.25 (spiral), $4.00 (cloth). [Many of the 499 figures are drawn from Michigan
material. See review on p. 165 of this issue.]

Shetron, Stephen G. 1969. Variation in Jack Pine Growth by Individual Soil Taxonomic
Units in Baraga County, Michigan. Res. Notes Mich. Tech. Univ. Ford Forestry Center
5. 25 pp. (processed).

Smith, Bruce D., ed. 1969. Summer Science Journal. Vol. 1. Summer Science, Inc., Ann
Arbor. 68 pp. [Contains reports on the geology, botany, aquatic biology, and archae¬
ology projects of the 32 students (13-18 age group) enrolled during the summer of
1968 at Summer Science Camp, on 2200-acre Summer Island, off the tip of Michi¬
gan’s Garden Peninsula (Delta Co.). While the reports, especially the orthography of
scientific names, reveal the inexperience of their youthful authors, valuable data on
the island are here recorded.]

Kalamazoo SW [Kalamazoo]
Kawkawlin [Bay]

Maybee [Monroe, Washtenaw,
Wayne]

Monroe* [Monroe]

Mt. Forest [Bay, Arenac]

Otsego [Kalamazoo, Allegan]

Palo [Ionia]

Pinconning [Bay]

Portage [Kalamazoo]

Rockford [Kent]

Rockwood* [Wayne, Monroe]
Saginaw [Saginaw]

Saginaw NE [Saginaw]

Schoolcraft [Kalamazoo, St. Joseph]
Schoolcraft NW [Kalamazoo]

Sparta [Kent]

Standish [Arenac, Bay]

Standish NE [Arenac]

Tecumseh N [Lenawee, Washtenaw]
Vicksburg [Kalamazoo, St.

Joseph]

YpsilantiW [Washtenaw]

206

THE MICHIGAN BOTANIST

Vol. 9

Swink, Floyd. 1969. Plants of the Chicago Region. Morton Arboretum, Lisle, Illinois. 445
pp. $8.50 [Berrien Co., Michigan, is one of the 22 counties included in this checklist
with extensive ecological notes. See review on p. 165 of this issue.]

C. JOURNAL ARTICLES

Bigelow, Howard E., & Margaret E. Barr. 1969. Contribution to the fungus flora of
northeastern North America. V. Rhodora 71: 177-203. [Cites type of Tricholomopsis
bella from Emmet Co.]

Bigelow, Howard E., & Alexander H. Smith. 1969. The status of Lepista— a new section
of Clitocybe. Brittonia 21: 144-177. [Material of 11 species is cited from Michigan
(usually with no further locality data) and a paratype of C. irina var. luteospora. ]

Bonnot, E.-J. 1967. Ambrosia artemisiifolia L. Bull. Mens. Soc. Linn. Lyon 36: 348-359.
[A general survey (in French) of knowledge about common ragweed, with references
to some of the work done in Michigan, including a long footnote summarizing the
thesis of Gebben on ecology of the species in southeastern Michigan.]

Byer, Michael D. 1969. The role of physical environment in some tracheophyte distribu¬
tions along a soil moisture gradient. Bull. Torrey Bot. Club 96: 191-201. [Study area
was in Crawford Co.]

Canham, Susan Carey. 1969. Taxonomy and morphology of Hypocrea citrina. Mycologia
61: 315-331. [Var. citrina cited from Tahquamenon Falls.]

Culberson, Chicita F. 1969. Chemical studies in the genus Lobaria and the occurrence of
a new tridepside, 4-0 methylgyrophoric acid. Bryologist 72: 19-27. [Some of the
herbarium material analyzed for certain of the taxa in this lichen genus was from
Michigan.]

Review

GEOLOGY OF MICHIGAN. By John A. Dorr, Jr., & Donald F. Eschman. University of
Michigan Press, 1970. 476 pp. $15.00.

Get this book! Any further review is only commentary on that three-word summary.
Get it for its abundant handsome illustrations, well displayed on a large page size. (Figure
legends not only cite localities where photographs were taken but also indicate them on
an outline map of Michigan.) Get it for its charts and maps, its lists of collecting sites for
minerals and fossils. Get it for information on Michigan’s economic resources: iron,
copper, salt, limestone, gas, petroleum, etc. Get it for fundamental explanations of the
State’s scenic features: sand dunes and waterfalls, the picturesque harbors of Keweenaw
County, the Pictured Rocks, Mackinac Island’s terraces and fort sites. Get it for the
amply illustrated accounts of Michigan fossils: invertebrates, vertebrates (including man),
and plants, the latter including billion-year-old algae in Ontonagon County-the oldest
evidence of photosynthesis. Get it for the chapter on “Water and Wind in Michigan,”
which will be of particular interest to botanists (although the brief comments on the role
of vegetation in filling of lakes fail to distinguish the unique features of bogs and
swamps). Get it for the history of the Great Lakes, with maps and summary chart of
outlets and dates.

Satisfyingly broad and complete in its scope, this volume will be a joy to anyone
in this region who is curious about the land he lives on and the lakes which surround him
and how they came into their present form. The text is written at a level from which an
intelligent layman can learn much, and there are references to key literature. The only
complaint about this oft-delayed production is the paper-covered boards used in binding;
they do not look resistant to abrasion from the sands of Michigan or the dirty hands of
field-trippers!

-E. G. V.

Program Notes

May 29-31: Michigan Botanical Club, Annual Spring Campout, at the University of Michi¬
gan Biological Station on Douglas Lake. A full program of field trips is planned;
members are receiving complete information on reservations.

June 28: Southeastern Chapter, Annual Meeting, at the Drayton Plains Nature Center.

September 25-27: Southeastern Chapter, Fall Campout, at Yankee Springs.

Editorial Notes

Authors of articles in recent issues who wish to have manuscripts and/or art work
returned should advise the eidtor immediately, before another “housecleaning” of his
files.

The October issue will include a cumulative three-year index to volumes 7-9. It is
contemplated that future volumes will include an annual index.

Prospective authors are urged to read “Information for Authors” on p. 62 of the
January number.

The March number (Vol. 9, No. 2) was mailed March 17, 1970.

Publications of Interest

A GUIDE TO THE FLORA OF ELGIN COUNTY ONTARIO. By William G. Stewart
and Lome E. James. Catfish Creek Conservation Authority [1129 Talbot St., St.
Thomas, Ontario], 1970 [“1969”]. 118 pp. $1.50 + $.25 postage. This is an an¬
notated list of vascular plants and some bryophytes of a 720-square-mile county on
the north shore of Lake Erie, with maps and background information. The list is not
based on published reports, but only on species actually seen by the authors; voucher
specimens for almost all are at the University of Western Ontario.

FOOD FOR US ALL. The Yearbook of Agriculture 1969. 360 pp. Another volume in
the annual series, this one devoted to food: meat cuts, grading of foods, packaging,
recipes, nutritive values, storage, shopping economy, etc. Available for $3.50 from the
Superintendent of Documents (Washington 20402) or without charge fron one’s con¬
gressmen.

CHROMOSOME NUMBERS OF FLOWERING PLANTS. By Z. Bolkhovskikh, V. Grif, T.
Matvejeva, & O. Zakharyeva. An A. Federov (editor). V. L. Komarov Botanical Insti¬
tute, Academy of Sciences of the USSR, Leningrad, 1969. 926 pp. 6R.92K. A vast
index to 80 years’ investigations throughout the world on plant chromosomes, indi¬
cating the numbers reported for 35,000 names of species, including many here newly
listed. Since the preface appears in both Russian and English and the rest of the tome
consists of scientific names and citations to the bibliography of 7,000 titles, this
impressive compilation presents no language barrier. Families, genera, and species are
arranged alphabetically; for genera containing over 10 species, tables summarize the
distribution of chromosome numbers. Soviet botanists have rendered an invaluable
service to Angiosperm systematists!

CONTENTS

A Botanical Inventory of Sanford Natural Area.

II. Checklist of Vascular Plants

John H. Beaman . 147

Reviews— How to Know Pollen and Spores

Plants of the Chicago Region . 165

Natural Proliferation of Floating Stems of
Scouring-Rush, Equisetum hyemale

Warren H. Wagner, Jr., & William E. Hammitt . 166

Changes in the Vascular Flora of Seven Small Islands in
Western Lake Erie

Thomas Duncan & Ronald L. Stuckey . 175

A Report on the 1969 Morel Season In Michigan

Ingrid Bartelli . 200

Michigan Plants in Print . 205

Review— Geology of Michigan . 206

Program Notes . 207

Editorial Notes . 207

Publications of Interest . 207

(On the cover: Flower of blue cohosh, Caulophyllum thalictroides,
a common spring wildflower in Sanford Natural Area,
photographed by John H. Beaman, April 1968. See p. 150.)

r

Vol. 9, No. 4

THE

MICHIGAN BOTANIST

LIBRARY

OCT ly '970

October, 1970

NJIW YOfl*

BOTANICAL (SAROBN

THE MICHIGAN BOTANIST-Published by the Michigan Botanical Club four times per

year: January, March, May, and October. Second-class postage paid at Ann Arbor,

Michigan. ©Michigan Botanical Club, 1970.

Subscriptions: $3.00 per year. Single copies: $.75.

All back issues are available, at the following prices:

Vol. 1 (2 numbers, at $1.00 each): $2.00

Vol. 2-5 (4 numbers, at $.50 each): $2.00 (per volume)

Vol. 6- (4 numbers, at $.75 each): $3.00 (per volume)

Subscriptions (from those not members of the Michigan Botanical Club) and all orders
for back issues should be addressed to the business and circulation manager, who will as¬
sume that subscriptions received prior to the last number of a volume are intended to begin
with the first number of that volume unless specified to the contrary.

Articles and notes dealing with any phase of botany relating to the Upper Great Lakes
Region may be sent to the editor in chief. The attention of authors preparing manuscripts
is called to “Information for Authors” (Vol. 9, p. 62; copies available from the editor).

Editorial Board

Edward G. Voss, Editor in Chief

Herbarium, The University of Michigan, Ann Arbor, Michigan 48104

Laura T. Roberts, Business and Circulation Manager
1509 Kearney Rd., Ann Arbor, Michigan 48104

Virginia L. Bailey Rogers McVaugh

Burton V. Barnes Alexander H. Smith

Richard Brewer W. H. Wagner, Jr.

THE MICHIGAN BOTANICAL CLUB

Membership in the Michigan Botanical Club is open to anyone interested in its aims:
conservation of all native plants; education of the public to appreciate and preserve plant
life; sponsorship of research and publication on the plant life of the State; sponsorship of
legislation to promote the preservation of Michigan native flora and to establish suitable
sanctuaries and natural areas; and cooperation in programs concerned with the wise use and
conservation of all natural resources and scenic features.

Dues are modest, but vary slightly among the chapters and with different classes of
membership. Persons desiring to become state members (not affiliated with a local chapter)
may send $3.00 dues to the Membership Chairman listed below. In all cases, dues include
subscription to THE MICHIGAN BOTANIST. (Persons and institutions desiring to sub¬
scribe without becoming members should deal with the business and circulation manager as
stated at the top of this page.)

President: W. H. Wagner, Jr., Botanical Gardens, University of Michigan, Ann Arbor,
Michigan 48105

Vice President: William F. Hopkins, 6210 Cowell Rd., Brighton, Michigan 48116
Recording Secretary: Mary Cooley, 703 S. Forest, Ann Arbor, Michigan 48104
Corresponding Sec.: Thelma Thomson, 15093 Faust, Detroit, Michigan 48223
Treasurer: Charles Buswell, 19204 Plainview, Detroit, Michigan 48219
Membership Chairman: Harriette V. Bartoo, Department of Biology, Western Michigan Uni¬
versity, Kalamazoo, Michigan 49001
Corresponding Secretary, Southeastern Chapter:

Catherine Hamilton, 9591 Grandmont, Detroit, Michigan 48227
Secretary, Huron Valley Chapter:

Anne Benninghoff, 3315 Alton Ct., Ann Arbor, Michigan 48105
Secretary, Southwestern Chapter:

Helen Wiles, 7113 N 25th St., Rt. 2, Kalamazoo, Michigan 49004

1970

THE MICHIGAN BOTANIST

211

MYXOMYCETES FROM CHEBOYGAN AND
EMMET COUNTIES, MICHIGAN

Laura S. Bradford
Department of Biology,

Washington University, St. Louis, Missouri

The Myxomycetes listed below were collected by the author during
early July 1969 in the vicinity of the University of Michigan Biological
Station. The first weeks of July had favorable conditions for plasmodial for¬
mation. June had been cold and very wet. During the first part of July, the
weather became warmer and drier but due to the heavy rains in June the
ground remained moist for several weeks. During these several weeks there
were plentiful plasmodia and also freshly formed fruit-bodies of Myxomycetes.

Although this was not an intensive collecting program, two of the
species listed below (Lindbladia tubulina and Physarum confertum) were not
included in the preliminary survey for this area done by Brooks and Kowalski
(1963) although they have been reported from Michigan elsewhere (Kauffman,
1910 & 1929; Gilman, 1943). Brooks and Kowalski (1963) had records of
Arcyria denudata, Reticularia lycoperdon, and Lycogala epidendmm from
Emmet County 7but not from Cheboygan County. All 12 genera and 14
species reported here were from Cheboygan County except the single collec¬
tion of Diachea leucopodia which came from a hardwood forest in Emmet
County. The two collections of Stemonitis axifera and the one collection of
Reticularia lycoperdon came from a hardwood forest area near Mackinaw
City. The other Cheboygan County collections came from forested areas be¬
tween and adjacent to Douglas Lake and Burt Lake. The classification used is
that of Martin and Alexopoulos (1969).

CERATIOMYXALES

Ceratiomyxaceae

1. Ceratiomyxa fructiculosa (Mull.) Macbr. — two collections, one on
moss and one on bark.

LICEALES

Reticulariaceae

2. Lycogala epidendrum (L.) Fries — two collections, one on mossy log
and the other on rotting wood.

3. Reticularia lycoperdon Bull, — one collection, on fallen log. Although
many of the spores are free as in the description given in Martin and
Alexopoulos (1969), spore clusters do exist mostly around the
pseudocapillitium.

4. Tubifera ferruginosa (Batsch) Gmel. — one collection, on mossy log.

Cribariaceae

5. Lindbladia tubulina Fries — one collection, on fallen leaves and other
forest floor debris.

212

THE MICHIGAN BOTANIST

Vol. 9

TRICHIALES

Trichiaceae

6. Arcyria denudata (L.) Wettst. — two collections, both on old logs.

7. Arcyria nutans (Bull.) Grev. — one collection, on birch bark.

STEMONITALES

Stemonitaceae

8. Stemonitis axifera (Bull.) Macbr. — two collections, both on old logs.

9. Diachea leucopodia (Bull.) Rost. — one collection, on twig.

PHYSARALES

Physaraceae

10. Fuligo septica (L.) Weber — four collections, on rotting log, a leaf, a
twig, and on bark.

1 1 . Physarum confertum Macbr. — one collection, on moss, which Martin
and Alexopoulos (1969) report as its favorite substrate.

12. Physarum psittacinum Ditm. — one collection, on dead wood. Martin
and Alexopoulos (1969) report this as an “uncommon” species but
Brooks and Kowalski (1963) have also reported two additional speci¬
mens of this particular species from this same area.

Didymiaceae

13. Diderma spumarioides (Fries) Fries — five collections, four of which
were on leaves, the other on a twig.

14. Mucilago Crustacea Wiggers — two collections, one on a dead leaf,
the other on a twig.

ACKNOWLEDGMENTS

I wish to thank Dr. James Maniotis for his assistance in confirming the identifica¬
tion of the Myxomycetes listed in the study. Grateful acknowledgment is made to the
National Science Foundation and the University of Michigan Biological Station for re¬
search funds provided under grant GB-8440.

LITERATURE CITED

Brooks, T. E., & D. T. Kowalski. 1963. A preliminary survey of the Myxomycetes of
Emmet and Cheboygan Counties. Pap. Mich. Acad. 48: 131-136.

Gilman, R. 1943. Slime molds of Benzie County, Michigan. Proc. Iowa Acad. 49:
189-190.

Kauffman, C. H. 1910. Unreported Michigan fungi for 1909. Rep. Mich. Acad. 12:
99-103.

- . 1929. A study of the fungus flora of the Lake Superior region of Michi¬
gan, with some new species. Pap. Mich. Acad. 9: 169-218.

Martin, G. W., & C. J. Alexopoulos. 1969. The Myxomycetes. Univ. Iowa Press, Iowa
City. 561 pp.

1970

THE MICHIGAN BOTANIST

213

BOTANICAL RESULTS OF THE MICHIGAN
GEOLOGICAL SURVEY UNDER THE DIRECTION OF
DOUGLASS HOUGHTON, 1837-1840

Rogers McVaugh
University Herbarium,

The University of Michigan, Ann Arbor

INTRODUCTION

The authors of the pioneer floras of North America, writing in the early
19th Century, knew little of Michigan, and their works contain few references
to the plants of that state. Although such botanical celebrities as Thomas
Nuttall (in 1810), David B. Douglass (in 1820), Edwin James (from 1827 to
1832), and Douglass Houghton (in 1831 and 1832), had visited Michigan and
collected plants there, their collections were made under difficulties (and as a
result were never very large), and the publications based on the collections
were scattered and often brief. For the most part these early explorers
traveled by water, and their botanizing was almost entirely restricted to the
shores of the upper Great Lakes.

Even the first volume of the authoritative and comprehensive Flora of
North America by John Torrey and Asa Gray, which was published in parts in
1838 and 1840, contains very few references to Michigan plants, although it
was intended to cover the continent north of Mexico. It is thus of consider¬
able interest to find that in the North American Botany by Eaton and
Wright,1 published in Troy, New York, in 1840, more than 1000 species of
vascular plants were reported from Michigan, including many woodland and
prairie species in addition to those from the shores of the Great Lakes. The
North American Botany was essentially an eighth and revised edition of Amos
Eaton’s Manual of Botany for North America, of which the seventh edition
had appeared in 1836. As the Michigan records were not included in the 1836
edition, it may be guessed that there is an interesting story behind their
insertion en masse in the 1840 edition. A clue to the story may be found in
the advertisement for the latter work, as quoted by Eaton in the preface (p.
vi):

“Age, and declining vision, have admonished the author that he must call in
the aid of youthful zeal, with talented energies, to supply his loss by the waning
of years. He has been so fortunate as to succeed in associating Dr. JOHN WRIGHT
with him in this edition. He is a very zealous and accurate young naturalist. After
some years of ardent devotedness to the Natural Sciences at this institution [the
Rensselaer Institute at Troy], he had the benefit of Professors Ives’ and Silliman’s
instructions at Yale College. He has since been united with Prof. Hall in the
descriptive catalogue of Troy Plants. In the summer of 1838 he collected and

lAmos Eaton and John Wright. North American Botany; comprising the native and
common cultivated plants, north of Mexico: Genera arranged according to the artificial
and natural methods. 8th edition [of Eaton’s Manual of Botany ]. viii + 625 pp. Elias
Gates, Troy, N. Y. 1840.

214

THE MICHIGAN BOTANIST

Vol. 9

reported, the plants of Michigan, as official Botanist to the State Geological Com¬
mission.”

“Dr. Wright is now going through all the generic and specific descriptions;
correcting and modernizing the language, as far as may be done, without departing
from the original simplicity of manner, peculiar to this work. New, and well
authenticated discoveries, are also added by him. He has before him the works of
Lindley, Torrey, Darlington, Hooker, Beck, &c. The contributions of many friends,
in catalogues, letters, and rare plants, afford him many facilities.”

A little further on (p. 16) Eaton added what he called a “remark”;
“The districts about our N.W. Lakes . . . have been in a great measure deficient
in recorded localities of plants . . . the botanical surveys of Dr. Houghton, Dr.
Wright, and his diligent assistant Mr. G. Bull, have supplied these deficiencies.”

It seems clear from the above that the addition of the Michigan records
(“new, and well authenticated discoveries”) to the 1840 edition was primarily
the work of Wright, and it may be surmised that the original source of many
of the records is to be sought in Wright’s work as “Botanist to the State
Geological Commission.” This is a little-known chapter in botanical history
that deserves a fuller treatment.

In 1840 Michigan was a new and relatively poor State, having been
admitted to the Union only 3 years before. It is remarkable that such a
pioneer community was able to initiate a program which by 1840 made Michi¬
gan one of the better known states, botanically speaking, in the whole area
west of the Alleghenies. In fact the botanical program comprised only a minor
part of the Geological Survey of Michigan.

One of the early acts of the new State Legislature was to provide
(February 23, 1837) for “an accurate and complete geological survey of the
state.” The survey was intended to result in a “full and scientific description
of its rocks, soils & minerals & of its botanical and geological productions, to¬
gether with specimens of the same.” The governor was authorized to appoint a
geologist to take charge of the Survey, and also to appoint some competent
persons to act as assistants under the direction of the geologist.2

The moving spirit in the Geological Survey was a remarkable young
man, Douglass Houghton (1809-1845), a native of New York who had studied
with Professor Amos Eaton at the Rensselaer Institute in Troy. Houghton
came to Detroit in 1830, upon Eaton’s recommendation, to give a series of
lectures on chemistry and other natural sciences. He spent the winter of
1830-31 in the city, before accepting the summer post of surgeon and
botanist to the Schoolcraft Expedition to the sources of the Mississippi River.
He returned from the expedition in October. The following summer he took a
similar part in the second Schoolcraft Expedition, which in fact reached Lake
Itasca. An account of Houghton’s botanical work on the 1832 expedition
appeared in 1962 in this journal.3

2Acts of the Legislature of the State of Michigan 1837: 14. 1837.

^Janice L. Rittenhouse and Edward G. Voss. Douglass Houghton’s botanical collec¬
tions in Michigan, Wisconsin, and Minnesota on the Schoolcraft Expedition of 1832.
Mich. Bot. 1: 61-70. 1962.

1970

THE MICHIGAN BOTANIST

215

After Houghton’s return from the expedition of 1832, he settled
permanently in Detroit. He went into the practice of medicine, and by all
accounts was successful. He speculated in Detroit real estate so profitably that
all his biographers agree he became financially independent. He maintained at
the same time his interest in the sciences, and he seems to have been almost
solely responsible for the planning of the Michigan Geological Survey, and for
the adoption of the project by the Legislature. The first Governor of Michi¬
gan, Stevens T. Mason, close upon the legislative action authorizing the
Survey, appointed Houghton to the post of State Geologist.4

In order to understand and appreciate the botanical work of the Geolog¬
ical Survey, we should digress briefly to describe the conditions in Detroit and
in Michigan at that time. Although forts and settlements on the shores of the
upper Great Lakes had been in existence for almost 200 years, the interior of
northern Michigan was a wilderness; in fact there were few settlements north
of Saginaw Bay except on the lake shores. The population of Detroit in 1830
was 2222; by 1840 it had risen to 9102. Increase Lapham, who later became
known as a competent amateur botanist in Wisconsin, visited Houghton in
June, 1836, when a geological survey “was being talked of.” Lapham de¬
scribed Detroit as “a fine city one hundred and thirty-five years old; it con¬
tains about 8,000 inhabitants, several fine four-story brick buildings, numerous
fine churches and a market-house as large as an ordinary Presbyterian church
and built much in the same style!”

At this same time, according to Lapham, Grand Haven, Michigan, was a
new town of ten houses and one steam mill; Milwaukee, Wisconsin, consisted
of 50 houses, and claimed a thousand inhabitants including the suburbs.5

Travel by road in Michigan was limited to the southern tiers of counties.
When Houghton and his assistant, Bela Hubbard, set out on the first geological
excursion in September, 1837, they travelled by wagon the first day from
Detroit to Royal Oak, where they camped by the roadside. The following day
they continued by wagon as far as Byron (then consisting of a mill and two
houses) in southeastern Shiawassee County, whence they were obliged to con¬
tinue their journey to Saginaw Bay by canoe. Midland was a “village without
inhabitants”; in the Saginaw Valley the travelers found they had “penetrated
into the wilderness, many miles beyond the most remote of the settlements of
the Anglo-Saxon. Wild game was very abundant. . . . Besides deer, we had often
seen along the shore, tracks of the elk, and sometimes of the moose. . . .”6

Earlier the same year, when Houghton was exploring farther west in the
Lower Peninsula, he went from Detroit to Ann Arbor and Jackson by road,

4 Bela Hubbard. A memoir of Dr. Douglass Houghton. Am. Jour. Sci. II. 5:
217-227. 1848. Alvah Bradish. Memoir of Douglass Houghton, xii + 302 pp. + frontisp.
Raynor & Taylor, Detroit, 1889. Edsel K. Rintala. Douglass Houghton, vii + 119 pp. +
frontisp. Wayne Univ. Press, 1954.

^Letters of I. A. Lapham, pp. 324-337, copies in the manuscript collection of the
Ohio Historical Society Library, Columbus.

^George N. Fuller, ed. Geological Reports of Douglass Houghton, pp. 33-34.
Lansing, 1928.

216

THE MICHIGAN BOTANIST

Vol. 9

thence by canoe down the Grand River past the future site of Lansing; he
explored the Maple River as far north as Gratiot County, then descended to
Grand Rapids, all without benefit of highways.

Or consider the following, from Bela Hubbard’s report on Wayne
County in 1838: “Marshes or wet prairies comprise extensive tracts. . . . Indi¬
cations are apparent which prove that very many, at least, of these peat
marshes had their origin in the labors of the beaver, aided by the natural
conformation of the surface. . . . About fourteen sections, two-thirds of which
are in the town of Hamtramck ... are of the character above described; but
not more than one-half this extent is actual prairie. This yields an abundant
growth of wild hay. The marsh is now in the progress of successful drain¬
age. . . . According to reports of Indians, beavers disappeared from this region
thirty years ago. Their numbers previous are said to have been incredible.”

Because of the unbroken and largely uninhabited wilderness that occu¬
pied all but the southern part of the Lower Peninsula, detailed botanical and
zoological surveys were possible only along the major roads that crossed the
State from east to west. The “Chicago road,” following the route of an old
Indian trail, ran from Detroit to Clinton, Jonesville, Coldwater, Sturgis, and
on westward. Another possible road led to Ann Arbor, Jackson, Marshall,
Kalamazoo, and St. Joseph. We shall see how the actual surveys were in¬
fluenced by the existence of these roads, and the towns along the roads.

BOTANICAL ACTIVITIES OF THE SURVEY

Most of what is known, or surmised, about the botanical work of the
Geological Survey is derived from the work by Eaton and Wright mentioned
above; from the geological reports made by Houghton to the Michigan Legisla¬
ture (including the report by John Wright on the field season of 1838); and
from the plant specimens collected by the members of the Survey from 1837
to 1840, and now preserved in the herbarium of The University of Michigan
at Ann Arbor.

The specimens are approximately 800 in number, representing fewer
than 600 species. They provide much information about the routes of the
field parties in each year, but the data accompanying the plants are often
incomplete or misleading, and must be interpreted with caution. It appears
that the collections came to the University Herbarium from several sources.
Whether or not any of them have been continuously in University hands since
1840 or before is not known. Some of them bear the original field labels
written by Abram Sager (Fig. 1), Bela Hubbard (Fig. 2), John Wright (Figs.
8,9), or George Bull (Figs. 10,11), or by Houghton himself (a few only). More
than half the specimens in the herbarium seem to have been duplicates, the
labels for which were prepared 30 years or more after the plants were col¬
lected.

It is surmised that most of the existing collections, including duplicates,
remained in the possession of the University until about 1868, when Mark W.
Harrington (1848-1926), later a distinguished educator, became an assistant in
the University Museum. As part of his duties he is said to have catalogued the

1970

THE MICHIGAN BOTANIST

217

herbarium, collected and identified specimens, and prepared collections for
exchange. Harrington was later (1873-1876) primarily responsible for the de¬
velopment of the collections in the Museum.7 Whatever his duties may have
been, a very large proportion of the labels on the First Survey specimens are
in his handwriting (Fig. 4). Many others are in the hand (Fig. 3) of John F.
Eastwood ( -1911), an assistant in the Museum from about 1871 to 1875

and later a college teacher of chemistry.

Another handwriting (Fig. 5) that appears on miscellaneous old collec¬
tions in the Michigan herbarium, and on many of the specimens collected by
the botanists of the First Survey, although bold and distinctive, is thus far
unidentified. It is presumed to be that of another museum assistant. One
interesting label in this hand, on a specimen of Erythronium albidum from
Ann Arbor, probably was written between 1875 and 1880, as it includes the
words “Frederick & Ferris (’77).” Because this particular label was written on
the back of one of the original labels prepared by John Wright for a plant
collected in 1838, I suppose the Erythronium was collected about the time
the First Survey plants were being mounted for the University Museum, and
that some of the original labels were discarded and used for scrap-paper. We
have no direct evidence as to the date of mounting of any of the specimens
collected between 1837 and 1840, except that on one sheet of Chenopodium
ambrosioides there is a note by V. M. Spalding, stating that the plant came
from the “Houghton collection,” and that when it was mounted, on January
26, 1880, the original odor of the plant was still strong!

Some of the duplicates from the First Survey collections seem to have
found their way into the private herbarium of Abram Sager, which was pre¬
sented to the University in 1886; at least a few specimens from Sager’s herbar-

Fig. 1. Handwriting of Abram Sager, slightly
lected in 1837.

less than natural size, from specimens col-

7e. B. Mains. Univ. Mich. Encyc. Surv. 1444-1446. 1956.

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Fig 2. Above, handwriting of Bela Hubbard, about natural size, from specimens collected in 1837. The date “1838” is a later
addition. Below, handwriting of Elizabeth C. Allmendinger, about natural size, probably 1870-1880. The specimen was
presumably collected by Wright and Bull, not by Houghton.

1970

THE MICHIGAN BOTANIST

219

ium, though cryptically labelled, bear dates and localities indicating their con¬
nection with the First Survey. An even larger number of duplicates apparently
came into the possession of Miss Elizabeth C. Allmendinger (1837-1909), a
gifted amateur botanist who lived in Ann Arbor, and whose herbarium was
eventually given to the University. The duplicates from the Allmendinger col¬
lection often bear labels (Fig. 2) indicating Houghton as the collector, even
though from known dates and localities it is clear that the plants actually
were collected by others.

In addition to all the above, there is a surprisingly large number of
specimens that bear no labels at all except comparatively recent ones prepared
with a typewriter, or written by herbarium assistants of the period 1920-1930.

Thus the specimens now in the Michigan set have been treated in a
variety of ways, and not always for their good. The plants for the most part
were carefully collected and pressed; many of the specimens are of excellent
quality, and many of them are still in an excellent state of preservation, but
as noted above many are inadequately documented. More of this below, in the
discussion of the collections made in the different years of the Survey.

Fig. 3. Handwriting of John F. Eastwood, about natural size, probably 1871-1875.

220

THE MICHIGAN BOTANIST

Vol. 9

THE SURVEY IN 1837

The Survey proceeded slowly during the first year. The State appropri¬
ated $3000.00, mostly to be spent on field expenses. Houghton and his
assistants, with one exception, served without salary. The Botanical and
Zoological departments were united under the direction of the principal
assistant, Dr. Abram Sager (1810-1877), who was later Professor of Botany
and Zoology, and ultimately Dean of the Medical School, at the University of
Michigan.8 Sager was primarily interested in zoology, but apparently devoted
some effort to botanical collecting. Houghton’s report dated December 1,
1837, says “The Botanical and Zoological departments have received due atten¬
tion and larger collections of specimens have been made than could have been
anticipated.”

fcZ Vi *

/7

/v

Fig. 4. Handwriting of Mark W. Harrington, about natural size, probably 1870-1880.

8For a brief notice of Sager, see E. B. Mains, in The University of Michigan [,] an
encyclopedic survey, p. 1444. 1956.

1970

THE MICHIGAN BOTANIST

221

In a report dated January 22, 1838, he said further:9 “The departments
of Botany and Zoology, under the more immediate direction of Dr. Abraham
[sic] Sager, principal assistant, have received due attention. Extensive collec¬
tions of plants have been made, and the preparatory steps taken towards
arranging a Flora of the state.”

There are about 160 herbarium specimens that we suppose were col¬
lected in 1837. These are primarily from a series of localities extending from
Monroe County westward to St. Joseph County and north to Kalamazoo (Fig.
6). Somewhat fewer than half the sheets bear contemporary labels in the hand

Fig. 5. Handwriting presumed to be that of a museum assistant at the University of
Michigan, 1870-1880, about natural size. Many of the duplicate labels, especially of the
1838 collections, are in this hand.

^Fuller, op. cit., p. 137.

222

THE MICHIGAN BOTANIST

Vol. 9

of Bela Hubbard.10 This is especially interesting because the labels for
1838-1840, almost without exception, are written by other persons. It is
surmised that the specimens labelled by Hubbard may have once formed a
part of Houghton’s private herbarium, i.e. duplicates that were especially

Fig. 6. Localities where plants were collected in 1837 by the Botanical Department of
the Geological Survey (dots), and by Douglass Houghton or his assistant (triangles).
Houghton’s travels, as outlined in his Field diary, are indicated by heavy lines.

10Bela Hubbard (1814-1896) was Houghton’s principal geological assistant, who
worked with him in the Geological Survey from 1837 to 1845, and accompanied him on
many of his travels in Michigan. One of his duties seems to have been that of a secretarial
assistant, to assist Houghton in keeping records and field notes. A transcript of Hough¬
ton’s field notes for 1837 and 1838, for example, is in Hubbard’s hand in the original
volume of such notes, between pages copied earlier by Houghton himself, and the field
record for 1839 and succeeding years, also in Houghton’s hand. Hubbard’s writing was
distinctive, easy to recognize and easy to read (Fig. 2). Its presence on the labels of
1837, and on no others, is alluded to in the text. For a short biographical sketch of
Hubbard, who later became a well-known figure in Detroit, see Rep. Mich. Acad. 4:
163-165. 1904.

1970

THE MICHIGAN BOTANIST

223

ticketed by Hubbard for his superior.11 The month and day of collection
usually appear on Hubbard’s labels, but the date 1837 appears on one sheet
only (Dulichium arundinaceum, Branch Co., Aug. 1, 1837).

About 20 sheets not labelled by Hubbard, but also apparently collected
in 1837, bear tickets in what seems to be the writing of Abram Sager; again
but a single sheet ( Heuchera americana, Dundee, July 3, 1837), is dated as to
year.

Most of the rest of the collections in this series (about 60 sheets) seem
to have been labelled long after they were collected, as described above. They
have labels by Miss Allmendinger, Harrington, Eastwood, or the unknown
person who seems to have been a museum assistant of the Harrington period.
Of all the labels written by these copyists, the year is given as 1838 in about
25 instances, and 1837 in 2 or 3 instances only. It is surmised that the copies
were made from the original 1837 labels, which ordinarily did not record the
year of collection.

Even in the face of incomplete and sometimes conflicting data, we can
say with some assurance that these collections were made by the field party
of 1837, and not by Houghton himself as some have supposed. Houghton left
Detroit for Ann Arbor, Jackson, and the upper valley of the Grand River
about the same time that plant collections began in Monroe, and his field
diaries show that his routes for the rest of the summer never coincided with
the road that was travelled by the botanical collectors.12 It appears from an
existing copy of Abram Sager’s report for the summer, moreover, that the
latter travelled the very route indicated by the sequence of plant collections,
namely from Monroe to Dundee, Adrian, Jonesville, and Coldwater.13 We
know pretty certainly, also, that the “1837” collections could not have been
made in 1838, for the routes of the 1838 collectors are well documented, and
it is clear upon comparison of the two series of dates and localities that the
two sets of plant collections could not have been made during the same
season by a single party. After the 1838 season there was no formal plant
collecting, and there were but few specimens from 1839 and 1840, these
usually clearly attributable to Houghton himself.

It seems probable that most of the specimens from along the route from
Monroe to White Pigeon and Kalamazoo were actually collected by Dr. Sager,
who was after all in charge of the botanical and zoological aspects of the
Survey. Bela Hubbard’s duties were primarily geological in nature, and it

llAt least a part of Houghton’s herbarium was given to the University in 1864 by
his widow, Mrs. R. R. Richards; cf. Regents’ Proceedings 1864-1870, p. 39. June Meeting,
1864. Mains (op. cit. p. 1445) reports that the herbarium consisted of 9,000 specimens.
Some of Houghton’s own specimens are identifiable as such because the name
DOUGLASS HOUGHTON is stamped on them in small black capital letters. The collec¬
tions from the 1831 and 1832 expeditions, almost all of which are also labelled in the
hand of Bela Hubbard, are thus stamped.

1 ^Douglass Houghton. Transcript of field journal, pp. 53ff. Manuscript, Michigan
Historical Collections.

^Douglass Houghton. Transcript of field journal. Abstracts from Dr. Sager’s notes
[for part of the summer of 1837]. p. 95. Manuscript, Michigan Historical Collections.

224

THE MICHIGAN BOTANIST

Vol. 9

seems likely that he was not with Sager’s party for any length of time, even
though as noted above he wrote the labels for many of the existing specimens.

In addition to the specimens listed in Table 1, there are a few that
almost certainly were collected by Houghton himself, or by Hubbard, in 1837.
The dates and localities on these specimens correspond precisely to those of
Houghton’s known itineraries, and some of the species (from the vicinity of
salt springs) are mentioned in his diaries. The species represented by specimens
include the following: Scirpus olneyi and Samolus parviflorus, from Gratiot
County; Triglochin palustre, from Ionia County; Triglochin palustre and
Samolus parviflorus, from Kent County; and Goody era pubescens , from White
Rock, southeastern Huron County.

TABLE 1. Collections made in 1837, presumably by Abram Sager or those in his party, in
the southern counties of Michigan. For each locality the total number of collections is
entered in one column, and the number of collections bearing the original label in Hubbard’s
hand in a second column. Hubbard seems often to have recorded merely the name of the
county in which the collection was made; see county names in the Gazetteer.

Labelled by

County

Locality

Date

Total

Hubbard

Monroe

Monroe

June, or undated

6

3

June 20-July 1

10

5

Dundee

July 3

4

1

Lenawee

Adrian

undated

2

1

July 6-8

7

2

Lenawee

July 14

1

1

Hillsdale

Hillsdale

Hillsdale, or

July, or undated

12

4

Hillsdale Co.

July 1-24

6

1

Jonesville

July, or undated

2

2

July 12-25

17

12

Branch

Branch, or

Branch Co.

Mason, or

July 17-August 1

8

4

“Masonville”

July, or undated

2

July 19-30

7

4

St. Joseph

St. Joseph, or

St. Joseph Co.

August 2-8

8

5

Sherman

undated

1

August 1-2

2

2

Sherman prairie

August 2

1

1

Sturges prairie

White Pigeon, or

August 2

3

1

(August 8),
“Pigeon”

August, or undated

5

August 4-23

13

6

Pigeon prairie

August 11, 24

2

1

Kalamazoo

Kalamazoo

August

2

1

August 23-September 5

8

5

1970

THE MICHIGAN BOTANIST

225

THE SURVEY IN 1838

During the winter of 1837-38 the Survey was reorganized on a larger
basis. The legislature passed, on March 22, 1838, an expanded act providing
for a Survey of four departments, namely geological and mineralogical, zoolog¬
ical, botanical, and topographical. A condition of the act, which ultimately
proved detrimental to the progress of the survey, provided that 17 specimens
of each collection should be taken whenever practicable, for distribution by
the regents “amongst the university and its branches.”l * * * * * * * * * * * * 14

As soon as the act of 1838 was passed, Houghton sought and found a
competent person to take charge of the Botanical Department of the Survey.
This was Dr. John Wright (1811-1845), who like Houghton had been associ¬
ated with Amos Eaton, and who was the co-author of a creditable local flora
published two years earlier.15 Wright gave up what was said to be a “lucrative
practice” of medicine in Troy to accept the position in Michigan. As Botanist
of the Geological Survey, he received $1500.00 for the season. This seems a
fairly good salary when measured against those of the Director of the Survey
and the Governor of Michigan, each of whom received $2000.00 for his
services during the same period.16 At the time of Wright’s appointment a Mr.
George H. Bull was appointed as “sub-assistant to the Botanical Department,”
at a salary of $800.00. 17 Dr. Abram Sager remained in charge of the Zoologi¬
cal Department, with the services of two assistants.

A report of the botanical work of 1838 was published by Wright.18
This consisted of a brief introductory statement, followed by an alphabetical
list of names of “between eight and nine hundred native or naturalized
species” of flowering plants and ferns. Because the report is a list of names
and no more, without any supporting information about localities, dates,
habitats or collectors, it is not very useful, but it is interesting as providing
records of certain weeds that had already been introduced into Michigan, and

l4Acts of the Legislature of the State of Michigan 1838: 119-121. 1838.

15john Wright & James Hall. A catalogue of plants growing without cultivation in

the vicinity of Troy. 42 pp. Troy, [New York], 1836.

16Michigan Senate Documents 1839: 109, 117. 1839.

17I have not been able to learn anything about George Bull’s life outside of his

connection with the Survey. He seems to have been an efficient assistant, with some

degree of literacy and some interest in botany. Amos Eaton named a new species,

Gymnandra bullii, and dedicated it to “Mr. Geo. Bull, Ast. Bot. to Mich. Commission.”
When Wright resigned after the 1838 season. Bull continued as an assistant to Houghton.

He accompanied Houghton on a long trip along the shores of the upper Great Lakes

during the summer months of 1839, and apparently was responsible for most of the 1839

plant collections that are still extant. At the conclusion of that season’s field work he

travelled to New York at Houghton’s request “with a suite of the plants collected this

year, for the purpose of submitting them to the examination of Dr. Wright . . .” (Hough¬

ton, letter to John Torrey, 1 Nov. 1839), but whether he came originally from New York

or from Michigan I do not know. He is referred to at least once by Houghton as “Dr.

Bull,” but this seems to have been a slip of the pen.

2nd Ann. Report of the State Geologist (Michigan House Doc. 1839:
421-442. 1839; Mich. Senate Doc. 1839: 305-326. 1839; also in Fuller, op. cit., pp.

223-251.

226

THE MICHIGAN BOTANIST

Vol. 9

records of native plants, particularly prairie species, that are now very rare in
Michigan, or quite unknown except from the collections of the First Survey.
In the words of the report, the collections of 1838 were made “in the two
most southern ranges of counties of the state, from the Detroit river to Lake
Michigan, excepting the county of Monroe, and in St. Clair county.” Actually
the data from existing specimens show that the work was concentrated in the
following counties (see Fig. 7): St. Clair (May 25-June 11), Washtenaw (June
19-July 2), Jackson (July 5-18), Kalamazoo (July 23-Aug. 7), Van Buren
(Aug. 9), Berrien (Aug. 10-20), Cass (Aug. 20-28), St. Joseph (Sep. 8-12), and
Lenawee (Sep. 14-24). Wright, in his report, explained why these particular
localities were chosen, alluding to and at the same time perhaps understating
the difficulties of making extensive field collections in the wilderness:

Fig. 7. Localities where plants were collected in 1838 by the Botanical Department of
the Geological Survey (dots), and in 1839 and 1840 by Douglass Houghton or George
Bull (half-dots).

1970

THE MICHIGAN BOTANIST

227

The extensive requisitions of the bill, making it obligatory ... to collect and
preserve . . . seventeen specimens of each kind . . ., has been the principal induce¬
ment for confining the sphere of action to the above mentioned portions of the
state. The bulky apparatus necessary to be conveyed from place to place, during
the excursions, for the preservation of the plants in such extensive collections, and
the requisite conveniences for drying and protecting them, render it impracticable
to examine a very great extent of country, and particularly such portions of it as
are unsettled, during a single season. . . .

Wright went on to say that with Mr. Bull’s assistance he had examined
the species enumerated in the report, and collected specimens of each,
“amounting, in all, to about nine thousand, which are now in an excellent
state of preservation. More than this number of species were observed growing
in the counties examined, but they were not in a proper condition for the
selection of specimens. ... A considerable number of cryptogamous or flower¬
less plants were also noticed and secured.”

We know very little about the fate of the 9000 and more specimens that
were so laboriously collected in 1838, but we must admire the diligence and
determination displayed by Messrs. Wright and Bull in gathering them. The
field season lasted only about 100 days, which meant the field party had to
dry 90 or 100 specimens every day, at the same time collecting as many as
they could in sets of 17. Any field botanist who has tried to dry that many
specimens, day in and day out, through a long summer, knows how much
work is involved. The mere changing and drying of blotters in such quantities
must have presented some problems, to say nothing of the careful selection of
duplicates in large quantities, the labelling, packing, and otherwise safeguard¬
ing the specimens.

The largest existing set, as far as I know, is that at the University of
Michigan. It consists of about 500 sheets representing more than 400 species.
Like the collections of 1837, these probably came to the University from
several sources. It may be that the Michigan set once included most of the
duplicates; it is possible that many of these were destroyed by the disastrous
fire in the herbarium in 1913, but this is not certainly known.19 A consider¬
able set of duplicates seems to have gone into Miss Elizabeth Allmendinger’s
herbarium, and perhaps an equal or even greater number into Houghton’s
private herbarium. Partial sets were sent by Wright or by Houghton to John
Torrey and to Asa Gray, and these specimens may now be found at the New
York Botanical Garden and at the Gray Herbarium of Harvard University.
Other private collectors who received sets were George Thurber (1821-1890),
who was professor of botany and horticulture at Michigan Agricultural College
from 1859 to 1863, and Zina Pitcher (1797-1872), an early associate of
Houghton in Detroit, and a Regent of the University of Michigan. Thurber’s
specimens may be found today both at Harvard and at New York, and
Pitcher’s at the U. S. National Arboretum. There is also a small set of grasses
and sedges from the First Survey at the Tracy Herbarium of Texas A. & M.
University, and additional specimens from other sets may yet be discovered.

19Mains, op. cit., p. 1447.

228

THE MICHIGAN BOTANIST

Vol. 9

v

y>c>. — / t-yx-^ /l /

/C&A: <

Fig. 8. Handwriting of John Wright, about natural size, from specimens collected in
1838. The words “Platanthera dilatata” appear to be in the hand of Asa Gray. Gray
visited Detroit and Ann Arbor in August, 1838, and may have seen this specimen at that
time. The words “Glyceria aquatica Smith” on the lower label are a later addition.

1970

THE MICHIGAN BOTANIST

229

TABLE 2. Dates and localities of botanical collections in Michigan, May 25-September 24,
1838. The more general localities are those at which the majority of collections were made;
other localities near these much-collected places are listed secondarily.

County

Locality

Inclusive dates

Approximate
number of
collections

St. Clair

Port Huron

Ft. Gratiot

Black River

Clyde

Lake Huron Shore above St.

Clair River

Head of St. Clair River

6 miles below Port Huron

May 25 -June 11

60

Washtenaw

Ann Arbor

Marsh near Whitmore Lake
Northfield, sphagnous swamp
Banks of Huron River

Sphagnous swamp around lakes

2 miles west of Ann Arbor
Pleasant Lake

June 19-July 2

110

Jackson

Jackson

Grand River

July 5-18

75

Calhoun

Marshall

July 20

6-10

Kalamazoo

Kalamazoo

Sand Lake

Austin’s Lake

Grand Prairie

Along Kalamazoo River

Lake 3 miles east of the village
Gull Prairie

Gull Lake

July 23-August 7

75

Van Buren

None stated

August 9

6-10

Berrien

St. Joseph

Lake [Michigan] shore

Mouth of the Paw Paw River

St. Joseph River at Berrien

August 10-20

35

Niles

August 20-21

5

Cass

Edwardsburgh

Edwards Prairie

August 20-28

40

St. Joseph

Centreville

Hog Creek

September 8-12

15

Lenawee

Clinton

Wolf Lake

September [7?] -24

10-15

In the Michigan collection the specimens collected in 1838 are labelled
in a variety of ways, like those from other years of the First Survey. Perhaps
a third of them are provided with the original labels in the hand of John
Wright (Figs. 8,9), in that of George Bull (Figs. 10,1 1), or, in a few instances,
in another hand. The rest of the sheets lack original labels; such information

THE MICHIGAN BOTANIST

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230

Fig. 9. Handwriting of John Wright, about natural size, from specimens collected in
1838. The words “Hemicarpha subsquarrosa, Nees,” “v. setifolia,” and “C. eburnea
Boott,” are later additions.

1970

THE MICHIGAN BOTANIST

231

as they carry has been provided by Miss Allmendinger, by Harrington or one
of his contemporaries, or even by others at much later dates. Bela Hubbard’s
hand, for what reason I do not know, does not appear on any of the collec¬
tions of 1838. Because the specimens of 1838 are so much more numerous
than those of 1837, and because the year of collection was almost invariably
written on the original labels, there is little doubt as to the exact localities
and dates of collections made in this second season of the Survey.

THE SURVEY IN 1839 AND 1840

Botanical work of the Survey, as such, was not continued after the 1838
season. The assistants in the Botanical and Zoological Departments, with the
exception of George Bull, resigned, presumably because they were not expect¬
ing support for the field season of 1839. Some years after this event
Alexander Winchell, the geologist, inquired of the widow of Abram Sager, and
was informed that the “resignations were provoked by ignorant criticisms and
caricatures indulged in by members of the legislature in public debates on
questions connected with the survey, especially in the departments.”20 Read¬
ing between the lines, one may suppose that as the State of Michigan was not
in a good financial condition, the legislators could see little practical value in
paying for more collections of stuffed birds and dried flowers.

In any event, the State Senate inquired formally of the Director of the
Geological Survey, “What loss, injury or detriment might result from a tempo¬
rary suspension of said geological survey?” and “What part of such survey, in
particular, would be prejudiced by such suspension?”

Houghton, in his reply to the Senate on March 7, 1839, emphasized
especially the harm that would result if the work of the Botanical and Zoolog¬
ical Departments were to be discontinued. Of John Wright he said “it was
only after the most urgent solicitations, and at much pecuniary sacrifice upon
his part, that he was induced to accept the appointment; relying upon the
faith of the state to complete the work which he had been induced to under¬
take. . . . Even the most temporary suspension of the work would eventuate in
the loss of the scientific and practical matter which has been collected in this
department of the survey, while it certainly would be acting in no good faith
to the head of that department.”21

Wright seems to have returned to Troy early in October, 1 838 ,2 2 and
perhaps never returned to Detroit for any long period. His report on the 1838
field season, mentioned above, was dated at Detroit, January 1, 1839.

In Houghton’s 3rd Annual Report, dated February 3, 1840,23 he
alluded to the resignation of the botanical and zoological assistants, and con-

^OQuoted in George P. Merrill, ed. Contributions to a history of American state
geological and natural history surveys. Bull. U. S. Natl. Mus. 109: 175. 1920.

^Michigan Senate Doc. 1839: 463^t66. 1839; Fuller, op. cit., pp. 340-345.
22Letter from A. Sager to Houghton, 28 Sep. 1838, in Michigan Historical Collec-

23Michigan Senate Doc. 1840, vol. 2: 87. 1840; Fuller, op. cit., pp. 395-396.

tions.

THE MICHIGAN BOTANIST

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232

Fig. 10. Handwriting of George H. Bull, about natural size, from specimens collected in
1838 and 1839.

*»’ # V

iiIHliill'iiilWi

1970

THE MICHIGAN BOTANIST

233

tinued: “In order to preserve from absolute loss, what had already been done,
and to make such advance as the circumstances might admit, a single sub¬
assistant, Mr. Geo. H. Bull, retains his situation, and by his assistance I have
been enabled to place the botanical portion of the work in such condition
that it may be of use in the final and connected reports which are required to
be made.”

Unfortunately for Houghton’s expectations of a “final and connected
report,” the legislature by an act approved March 28, 1840, abolished the

Fig. 11. Handwriting of George H. Bull, about natural size, from specimens collected in
1839.

234

THE MICHIGAN BOTANIST

Vol. 9

zoological and botanical departments, and the survey became officially re¬
stricted to its strictly geological and topographical aspects.24

The University Herbarium contains about 60 specimens that were ap¬
parently collected in 1839. Most of them were probably collected by George
Bull, and about one-quarter of these bear a contemporary label in what I
suppose is his handwriting (Figs. 10,11). There are also in the herbarium at
least 4 specimens dated 1840, one with an original label by Hubbard. The rest
of the 1839 and 1840 collections, aside from those labelled by Bull or
Hubbard, evidently were eventually handled like those of 1837 and 1838, and
by the same people; their labels reflect the work of Miss Allmendinger, Har¬
rington, and the rest.

According to a statement by Houghton, Bull spent a part of the early
field season of 1839 on an independent venture into the south-central part of
the state.25 It is presumed that the following were collected by Bull; speci¬
mens marked with an asterisk (*) are thought to have been labelled by him:

Larix laricina

Royal Oak

April 26

Salix petiolaris

Royal Oak

April 30

Chamaedaphne calyculata

Royal Oak

May 2

Menyanthes trifoliata

Royal Oak

May 2*

Ribes americanum

Royal Oak

May 2

Aquilegia canadensis

Pontiac

May 24

Fraxinus quadrangulata

Washington, Macomb Co.

May 28

Ranunculus ?aquatilis

Pontiac

June 8

Arethusa bulbosa

Pontiac

June 8

Carex lanuginosa

Oakland Co.

June 11*

Carex buxbaumii

Oakland Co.

June 11*

Carex flava

Oakland Co.

June 11*

Collinsia vema

Saginaw Co.

June 21*

The rest of the collections of 1839 are from the northern part of Michi¬
gan. Houghton left Detroit on July 26, by steamer for Mackinac [i.e.
Mackinac Island] , where he arrived on May 28. Apparently Bull had preceded
him there, for there are at least four collections (two of them labelled by
Bull), dated at Mackinac, July 22-24. Four additional collections from the
same place are dated July 28-30.

On August 2, as recorded in Houghton’s diary, he left Mackinac in a
small boat with Bull and three “day laborers.” The party passed along the
southern shore of what is now Mackinac County to Drummond Island,
circumnavigated that island, ascended the St. Mary’s river to Sault Ste. Marie,
returned to Lake Huron and passed westward along the north shore of Lake
Michigan as far as the mouth of the Menominee River. They visited Green Bay
village and several of the islands in the bay, then left by steamboat for
Mackinac on August 27.

The specimens in the herbarium may be summarized as follows:

24 Acts of the Legislature of the State of Michigan 1840: 126-127. 1840; Merrill,
op. cit.. p. 175.

25“Dr. Bull, the Sub. Asst, occupied a portion of the season in the interior of the
state & a portion, in company with myself, on the coast. . . .” Letter, Houghton to John
Torrey. 1 Nov. 1839. See Fig. 12.

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235

Fig. 12. Handwriting of Douglass Houghton, in a letter to John Torrey, 1 November 1839.

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Vol. 9

County

Locality

Date

Number of
specimens

Mackinac

Mackinac [Island]

July 22-30

8

Chippewa

Drummond Island (south or southeast side).

August 2-3

10

Drummond Island (north side).

August 4

2

Islands of St. Mary’s River (below Sugar

Island)

August 6

4

St. Mary’s River (Lake George to the Sault)

August 7

1

Drummond Island (in the Detour Passage)

August 10

1

Mackinac

Goose Island

August 11

1

North Shore (i.e. between St. Ignace and 9
miles west of Point aux Chenes)

August 14

2

North Shore (from camp to 3 miles west of
Menococien River)

August 15

1

Mackinac or
Schoolcraft

North shore (from camp to east side of Point
Seul Choix)

August 16

1

Schoolcraft

“Point Souchoix”

August 17

1

Delta

North shore (Little Bay de Noc)

August 20

1

?Menominee

North shore, Green Bay

August 21

1

Menominee

[Near mouth of Menominee River]

August 22

3

“Louse Island, Mich.” (=Rock Island, Wis¬
consin, at the tip of the Door Peninsula).

August 26

1

The collections of 1840 are but four in number. The dates and locali¬
ties, when definitely stated, agree with those of Houghton’s known itineraries
as recorded in his field diaries. The specimens are as follows:

Kalmia polifolia Whitefish Point, [Chippewa County] . June, the label

partly in the hand of Bela Hubbard

Polystichum lonchitis Lac Bay de Res, Lake Superior [Bete Gris Bay, Kewee¬
naw County] , June 30, 1840

Ranunculus reptans Lake Superior, June 30, 1840

Listera cordata Isle Roy ale, August 3, 1840

SPRING COLLECTIONS, 1837-1839

Because so many of the specimens are dated to month and day only,
the year of collection must be inferred in many instances. This can be done
with a high degree of confidence for specimens collected along the routes of
the field parties at some distance from Detroit. A specimen labelled merely
“Sherman, Aug. 1” can be assigned the date 1837 because the field party of
that year is known to have been in the southern part of St. Joseph County
for about 10 days in the first part of August.

In the vicinity of Detroit, however, it is quite another matter. The
members of the Geological Survey were based in the city, and presumably had
some opportunities to collect plants, if they wished, as soon as the spring
flora began to appear and before the regular field parties began their work.
Houghton himself was a competent botanist and may well have picked up an
occasional specimen, as may Bela Hubbard, whose handwriting appears on so
many of the 1837 labels. Sager was in the city in 1837 before leaving for the
beginning of the work in Monroe County in mid-June. Wright and Bull must
have been in Detroit at least for a short time in the spring of 1838 before
they began work in St. Clair County on May 25. As the name of the collector

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237

was never written on any of these early specimens, the copyists who labelled
the specimens many years later had no idea who collected them, and it is now
often impossible to guess who was responsible for them, and equally impos¬
sible to be sure of the year of collection.

Listed below are the spring collections in question. It seems likely that
those labelled by Bela Hubbard were collected in 1837, as his writing appears
on many other specimens that are confidently dated from that year, and on
very few specimens except those collected in that year. The date of 1838 is
presumably correct for the specimen of Viola rostrata. The other specimens in
the list may have been taken in 1837, 1838, or 1839. For additional spring
collections attributed to George Bull, see above under the account for 1839.

Populus tremuloides
Ulmus americana
Lonicera canadensis
Senecio aureus
Viola rostrata

Ranunculus flabellaris
Uvularia grandiflora
Sarracenia purpurea
Mitella nuda
Cypripedium reginae
Juniperus communis
Silene virginica
Potamogeton berchtoldii

Lysimachia thyrsiflora

Vicia americana
Lathyrus ochroleucus

Detroit, April 3, 1839
Detroit, April 4, 1839
Detroit, April 23, 1839
Ann Arbor, May 15, 1839
Detroit, May 22, 1838

Washtenaw Co., May 27
Wayne Co., May
Oakland, June 8
Oakland, June 8
Wayne Co.

Grosse Isle, June
Grosse Isle, June 12, 1838
Grosse Isle, June 16

Squaw Island, Detroit River,
June 16

Gibraltar, June 15, 1838
Gibraltar, June 16, 1839

(label by Harrington)

(label about 1870)

(label about 1870)

(label by Allmendinger)

(found 1952 in Houghton’s copy
of Beck’s Botany )

(label by Harrington)

(label by Hubbard)

(label by Hubbard)

(label by Hubbard)

(label about 1870)

(label about 1870)

(label by M. V. Cannon, ca. 1920)
(contemporary label in unknown
hand)

(label by Hubbard)

(label by Harrington)
(label by Harrington)

BOTANICAL ACCOMPLISHMENTS OF THE SURVEY

The activities of the Botanical Department of the Michigan survey had a
modest effect on contemporary botanical thought and literature. It is unfortu¬
nate that with the effective cessation of botanical work after the season of
1838, the complete withdrawal of support by the Michigan legislature in
1840, the death of Wright in April, 1845, and that of Houghton in October of
the same year, the botanical part of the survey not only came to an end but
also lost its principal advocates, and at the same time lost those who might
have been able to prepare a useful and comprehensive report on the collec¬
tions already made.

Wright continued his interest in the botany of Michigan until the end of
his life.2® After he returned to Troy he continued to correspond with Torrey

26Among the Sager documents in the Michigan Historical Collections is a letter
from Thomas C. Brinsmade, Wright’s medical associate in Troy. The letter is clearly dated
April 25, 1845, and describes in painful detail the progress of Wright’s slow decline and
death from “pulmonary phthisis.” Until the very last days he is said to have enjoyed
talking and reminiscing about his experiences with George Bull in the field in Michigan.
The standard biographical reference to Wright, that by S. E. Jelliffe in Howard A. Kelly’s
Cyclopedia of American Medical Biography (vol. 2, pp. 532-533. 1912) gives the date of
his death as April 11, 1846, but apparently it took place a year earlier.

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and consult him about matters of plant identification, but he noted in one
letter (March 21, 1840) that he had brought very few plants with him from
Michigan, and “those mostly single specimens.” He went on to say “Mr. Bull
my Assistant in Mich, has put up a bundle, & I have added a few in a separate
parcel, for you. I have not strength to overhaul my plants to send you all I
wish to.”

Torrey and Gray, in the foreword (June 1, 1840) to their Flora of
North America, state that “From Michigan, Dr. Houghton and Dr. Wright have
furnished us with numerous plants.” In the pages of the Flora these authors
cited a few collections by Houghton and a few by Wright, but it seems
unlikely that they received any very large number of plants of more than
ordinary interest to them.

Wright’s positive contributions after the work in Michigan were his list
of plants published in the Michigan Legislative Documents for 1839, and his
additions of Michigan records to Eaton and Wright’s North American Botany
in 1840. Both of these were in effect mere lists of names. Neither included
information on distributions or habitats of the plants as they occurred in
Michigan. The work of Eaton and Wright presumably became accessible to
most practicing botanists of the time, but the 1839 list must never have
enjoyed very wide circulation. It is interesting to speculate on the means
employed by Wright in preparing this list for publication between the end of
the field season in September, 1838, and the presentation of the report, which
is dated January 1, 1839. In Michigan at that time there was no herbarium of
any size that he could consult, and for his identifications he presumably
depended upon his prior knowledge of the plants of the northeastern United
States, and upon such works as the 7th edition of Eaton’s Manual (1836), or
L. C. Beck’s Botany of the Northern and Midland States (1833). Perhaps the
very need for haste in the preparation of the list prevented the inclusion of
certain species that Wright did not immediately recognize, and did not have
time or opportunity to study. For example the plant called Gymnandra (or
Besseya) bullii seems to have been collected in 1838, but no notice was taken
of it until it was described by Eaton in 1840.

With few exceptions the names in the 1839 list are those of plants that
might be expected in southern Michigan today. The more difficult groups are
not slighted; for example, there are 54 species of Carex in the list. On the
other hand there are only 6 species of Potamogeton, this reflecting perhaps a
lack of interest in aquatic vegetation on the part of the collectors. Many
introduced weeds are recorded, some certainly for the first time from Michi¬
gan. These alien species included Agrostemma githago (called “Cockle” by
Wright), Cannabis sativa (“Hemp”), Datura tatula (“Purple thorn-apple”),
Eleusine indica, Galeopsis tetrahit (“Flowering nettle”). Hibiscus trionum,
Malva neglecta (“Low Mallows”), Nepeta cataria (“Catnip”), Leonurus
cardiaca (“Motherwort”), Saponaria officinalis (“Bouncing Bet”), Trifolium
pratense and T. repens (“Red Clover” and “White Clover,” respectively).

The list as a whole is of interest because of what it suggests with respect
to the vegetation of southern Michigan before 1840, and the changes that

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have taken place since that time. For example, as might be expected, one
finds many characteristic plants of rich woodlands across southern Michigan
(e.g. Verbesina alterni folia, Aesculus glabra, Hydrastis canadensis, Agastache
nepetoides, Hydrophyllum appendiculatum, Panax quinquefolius, Polymnia
canadensis ), but some of these, like Hydrastis and Panax, are rare enough now
to be exciting finds for a botanist. If they had been as rare in 1838, the
chances would have been against their discovery by the Survey. Similarly with
species of primarily northern distribution in the State; plants like Arcto-
staphylos uva-ursi, Scheuchzeria palustris, Clintonia borealis, Coptis trifolia,
Waldsteinia fragarioides, and Gaultheria hispidula are by no means unknown in
the southern part of the Lower Peninsula, but habitats suitable for them are
now much restricted in area, and the fact that so many of them were found
by the Survey parties in 1838 indicates either that they were more common
at that time, or the botanists searched assiduously for them!

Perhaps most interesting from our point of view are the prairie plants.
Undisturbed prairies have now disappeared from Michigan except for a few
small areas along railroads, in cemeteries, and the like, but formerly they were
dotted across the southern end of the State, increasing in size and abundance
toward the southwest. The early settlers applied the term prairie not only to
upland grass-covered areas of deep soil, but also to open flat and mostly
inundated areas that would now be called marshes. These were at one time
very abundant in southeastern Michigan, but were beginning to be drained at
the time of the First Survey. The true prairies, with vegetation related to that
of the Great Plains of central United States, were found primarily from
Jackson County southwestward.

Often associated with the prairie plants, especially around lakes and
other water-filled depressions, were species having their principal areas of dis¬
tribution on the Atlantic Coastal Plain, or at least in the southeastern United
States. The botanists of the Survey discovered not only some of the so-called
“Coastal Plain disjuncts” (e.g. Fuirena pumila ), and some of the maritime
species that occur also around the Great Lakes (e.g. Cakile edentula.
Euphorbia poly goni folia), but also a number of southeastern species nearing
their northern limits in woodlands or in these relatively open prairie margins
and wet meadows, in extreme southwestern Michigan. Notable among these
are such grasses and sedges as Andropogon virginicus, Digitaria filiformis, Stipa
avenacea, Hemicarpha micrantha, Eleocharis equisetoides, and Fimbristylis
drummondif such monocots as Commelina erecta, Smilax rotundifolia, and
the rare orchid Triphora trianthophora; composites including Cacalia tuberosa,
Kuhnia eupatorioides, and Polymnia uvedalia ; and a few other species of
dicots as e.g. Buchnera americana, Eryngium yuccifolium, Swertia carolini-
ensis, and Sabatia angularis. Some of these, including Buchnera americana,
Commelina erecta, Digitaria filiformis, and Polymnia uvedalia, seem never to
have been found in Michigan since the First Survey, and none of the others is
common in the State at the present time.

As for the prairie species, Wright’s list included not only the more
common and adaptable species that are still common or even weedy in many

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places in southern Michigan (e.g. Andropogon scoparius, A. gerardii, Bouteloua
curtipendula, Sorghastrum nutans, Ratibida pinnata, Silphium perfoliatum, S.
terebinthinaceum, and Coreopsis tripteris ), but also some of the rarer species
that have since disappeared (or nearly so) from the Michigan scene (e.g.
Amorpha canescens, Aster sericeus, Baptisia leucophoea, Coreopsis palmata,
and Echinacea purpurea).

For what reason I cannot say, a rather high percentage of the rare and
unusual species reported by Wright are represented by specimens in the Michi¬
gan herbarium. There are no specimens, however, for many of the common
woodland species of southern Michigan. Perhaps some past curator of the
herbarium discarded duplicate sheets of the common species because the col¬
lection at that time contained better and more modern specimens of the same
species. Some few of the most interesting of Wright’s reports cannot be docu¬
mented, but most of the doubtful ones refer to distinctive species, and I am
inclined to credit the reports. Among those species for which there are no
supporting specimens at the University of Michigan are Andropogon virginicus,
Diarrhena americana, Stipa avenacea, Ludwigia altemifolia, Swertia carolini-
ensis, Onosmodium hispidum, and Eupatorium sessili folium. Apparently no
species of Onosmodium is known from Michigan; the others are all known,
but rare, in the southern part of the State.

No discussion of the work of the First Survey would be complete with¬
out some mention of their discoveries of rare species and species new to
science. That there were relatively few of the latter is an indication of how
well the flora of the Midwest was known by 1840, primarily through the
pioneer work of Nuttall, Michaux, Rafinesque, and a few others. Most of the
new species collected at the time of the Survey were not recognized as such
until somewhat later. The most interesting species are as follows:

Asplenium ruta-muraria L. Sp. PI. 1081. 1753. Collected on Drummond Island by
Houghton and Bull on August 3, 1839, on the limestone headland commanding the
passage between Drummond and Cockburn Islands. This, the American representative of
the common Wall-rue of Europe, is apparently unknown from Michigan except at this
locality, where it was re-discovered in 1961 by Jarl K. Hiltunen; see Brittonia 14: 120.
1962. At the Drummond Island locality the plant grows with another and almost equally
rare Michigan fern, Pellaea glabella Mett., which was also collected by Houghton.

Gymnandra bullii Eaton, in Eaton & Wright, N. Am. Bot. 259. 1840. ( Wulfenia
bullii of Gray’s Manual, 8th ed.; Besseya bullii as treated by Pennell, Acad. Nat. Sci.
Philadelphia Monog. 1: 378. 1935). The type locality was stated by Eaton as “Prairies of
Mch.” Pennell, although he examined most of the specimens of Scrophulariaceae in the
United States at the time of his revision, was unable to locate any of the original
material, and there is none in the University of Michigan herbarium. Perhaps the amount
collected in the first place was small, as Wright wrote to Torrey (who apparently had
asked to see a specimen) and informed him he was sending “all the remaining material.”
Perhaps a part of the type material, not recognized as such by Pennell or not seen by
him, may yet be found among the Torrey plants at New York.

Solidago houghtonii Torr. & Gray ex A. Gray, Man. 211. 1848. The type locality
was given as “North shore of L. Michigan; collected in the Michigan State Survey.” An
isotype at Michigan is labelled “Aug. 15th 1839 North shore of Lake Michigan.” As
Voss 27 has shown, the type material, undoubtedly collected by George Bull, came from

27jour. Sci. Lab. Denison Univ. 44: 28. 1956.

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THE MICHIGAN BOTANIST

241

somewhere between a point about four miles southeast of the present site of the village
of Epoufette and four miles west of the present village of Naubinway, in Mackinac
County.

Echinodorus parvulus Engelm. in Gray, Man. ed. 2. 438. 1856. Recorded by Engel-
mann, at the time of the original description, from Michigan, but not known from that
State except through one of the collections made by the First Survey (from White
Pigeon, St. Joseph County, August 11, 1837). Voss28 has discussed this in detail and
concludes that the specimen in the University of Michigan herbarium may be part of the
type-material. A specimen at the Gray Herbarium is labelled by Gray himself, “Michigan
State Coll[ection] .” This is probably the basis for Engelmann’s report of the species from
Michigan, as according to Voss there is no specimen from this state among Engelmann’s
own plants at the Missouri Botanical Garden.

Lysimachia stricta var. producta A. Gray, Man. ed. 2. 272. 1856. Gray’s specimen
was from Michigan, without statement of definite locality or collector, but labelled
“Mich. State collection] .” Voss29 has documented this case and concludes that the
specimen at the University of Michigan, labelled as from Branch County, Aug. 1 [1837],
is an isotype. The plant is now regarded as an interspecific hybrid, Lysimachia Xproducta
(A. Gray) Fern.

Cirsium pitcheri Eaton, Man. Bot. N. Am. ed. 5: 180. 1829. Pitcher’s thistle,
although neither a new species at the time of the First Survey nor a particularly rare one
in its own specialized Jiabitat on the dunes of the upper Great Lakes, was collected by
Wright near St. Joseph, Berrien County, and included by him in his 1839 report. It is
mentioned here because his was presumably the first report from so far south. Earlier
records were from Mackinac Island and from Lake Superior.

Scirpus olneyi A. Gray, Boston Jour. Nat. Hist. 5: 238. 1845. This is a widely
distributed but rather rare sedge, known in Michigan only from a few localities in saline
marshes in the Grand River valley. Douglass Houghton, traveling by canoe in 1837,
ascended the Maple River to a “Salt Works” in southern Gratiot County. On July 23 he
collected this species, or what may be a hybrid between it and S. americanus, in the
saline marshes near the “works.” Exactly 123 years later, July 23, 1960, Dr. Voss found
the very marsh that had been the source of the specimen and, a little further down the
river in Clinton County, by following the directions in Houghton’s original field diary, he
found another locality where Scirpus olneyi was still growing in a similar marsh.

Plantago cordata Lam. Tabl. Encyc. Mdth. 1: 338. 1791. This interesting and
relatively distinctive species of plantain was recently the subject of an article in this
journal. 30 It was formerly rather widespread in moist calcareous soils in northeastern
United States, but is apparently sensitive to pollution and other changes in the environ¬
ment, and has become very rare. It was collected near Port Huron by Wright in 1838,
and at a few other localities in southeastern Michigan by later collectors, but Mr. Tessene
has been unable to find any localities where the plant is now growing in the State.

EPILOGUE

The First Geological Survey of Michigan was conceived and nurtured by
a group of young men. Houghton was not yet 28 years of age when he
became State Geologist; Sager was a year younger, and Wright a year younger
still. The Survey in many ways was ahead of its time; perhaps it asked too
much of a State that desperately needed practical results and ready cash, and
could not afford to wait for the deliberate gathering of data that scientific

28Mich. Bot. 6: 16-17. 1967.

29Mich. Bot. 6: 20-21. 1967.

3hMelvern F. Tessene. Systematic and ecological studies on Plantago cordata. Mich.
Bot. 8: 72-104. 1969.

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methods dictated. All the young scientists that took part in the Survey must
have been disappointed to see their work come to naught. The geological
aspects of the Survey were, it is true, supported by the State until some results
could be seen. The botanists, on the other hand, have had to wait more than
a century and a quarter for any sort of recognition, and it is only to be
regretted that the traces of their work have been so meagerly preserved.

GAZETTEER

The following list does not include names of well-known application
that have been continuously in use for the same place since the period of the
First Survey, except for the names of certain counties that have been (or may
be) confused with specific localities in the same or other counties.

Ann Arbor, Washtenaw County. Collections were made June 23, 1838, in a “sphagnous
swamp” around a small lake 2 miles west of Ann Arbor. This was probably Second
Sister Lake, which until recently supported a sphagnous mat with many characteristic
associated species.

Austin Lake, Kalamazoo Co. See Sand Lake.

Berrien, Berrien County. Collections were made on “banks of the St. Joseph River,
Berrien,” August 17, 1838. As late as 1850 the name Berrien was applied to present
Berrien Springs.

Bete Gris Bay, Keweenaw County. Houghton worked in the vicinity of what he called
Bay de Gres, June 27-July 1, 1840. Specimens labelled “Lac Bay de Res of Lake
Superior,” June 30, 1840, are presumably from the shore of Lake Superior or from
Lac La Belle, almost south of Copper Harbor on Keweenaw Bay.

Black River, St. Clair County. See Clyde.

Branch [County]. On Hubbard’s labels of 1837, “Branch” is written sometimes as if it
were the name of a specific locality, but probably “Branch County” is meant; cf.
Lenawee, Hillsdale, St. Joseph. There was also a settlement called Branch (named, like
the county, for John Branch, Secretary of the Navy under President Jackson), situated
on the Coldwater River, a few miles south of Mason (or Masonville), which was also
on the river. The town of Coldwater was about a mile northeast of Branch, at the
edge of the “Coldwater Prairie,” and east of the river. The modern city of Coldwater
has spread away from the river onto the prairie, and Mason and Branch have long
since disappeared.

Clyde, St. Clair County. At one time a settlement called Clyde or Clyde Mills, now a
township traversed by the Black River, some 8-10 miles northwest of Port Huron.

Edwards Prairie, [Cass County] . Near Edwardsburgh, where collections were made August
20-28, 1838.

Goose Island, Mackinac County. In northern Lake Huron, T 41 N, R 1 W, about 6 miles
south of Hessel.

Grand Prairie, Kalamazoo County. Formerly an extensive prairie area west and northwest
of Kalamazoo.

Gull Prairie, Kalamazoo County. Formerly an extensive prairie area, near present Rich¬
land, west of Gull Lake.

Hillsdale [County]. On Hubbard’s labels of 1837, and on some labels copied in 1870 or
later, “Hillsdale” is written as if it were the name of a specific locality, but “Hillsdale
County” is meant. The present town of Hillsdale was not laid out until about 1839.
The older town, Jonesville, was on the “Chicago road” west of Detroit.

Hog Creek, [St. Joseph County]. Presumably near Centerville, where Hog Creek joins the
Prairie River. Collections were made near Centerville, September 8-12, 1838.

Lac Bay de Res, Lake Superior. See Bete Gris Bay.

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THE MICHIGAN BOTANIST

243

Lenawee [County] . On Hubbard’s labels of 1837, “Lenawee” is written as if it were the
name of a specific locality, but “Lenawee County” is meant.

Louse Island, “Michigan.” Houghton’s party visited here August 26, 1839. Its modern
name is Rock Island; it is the northernmost island in Wisconsin on the east side of
Green Bay.

Mackinac [Island], Mackinac County. At the time of the First Survey, all references to
“Mackinac” refer to the settlement on the island, or to the island itself, which is
about 3 miles east of St. Ignace, Mackinac County. The settlement at the present site
of Mackinaw City, on the mainland south of the Straits, had been abandoned for
more than 50 years.

Mason [or Masonville], Branch County. See Branch.

Menococien River, [Mackinac County]. This is the stream now called Millecoquins River;
it enters Lake Michigan about 1 mile west of Naubinway.

Northfield, Washtenaw County. See Whitmore Lake.

Oakland [County]. On Hubbard’s labels of 1837, “Oakland” is written as if it were the
name of a specific locality, but probably “Oakland County” is meant.

Paw Paw River, mouth of, [Berrien County] . The Paw Paw River empties into the St.
Joseph River at St. Joseph-Benton Harbor.

Pigeon [or Pigeon Prairie] . Evidently both these names refer to places near White Pigeon,
St. Joseph County, where the botanists worked August 4-24, 1837.

Pleasant Lake, [Washtenaw County]. Collections were made here June 28, 1838, while
the Survey party was working in Washtenaw County. This is presumably the Pleasant
Lake in Freedom Township, ca. 12 miles southwest of Ann Arbor.

St. Joseph [County]. On Hubbard’s labels of 1837, “St. Joseph” is written as if it were
the name of a specific locality, but probably “St. Joseph County” is meant. The
collectors passed through the county, from Sturgis to White Pigeon, August 2-24,
1837. Not to be confused with St. Joseph, Berrien County, which was the site of
botanical collections August 10-20, 1838.

Sand Lake, Kalamazoo County. Not definitely located. Austin Lake, and the smaller lakes
surrounding it, are indicated collectively as “Sandy Lakes” on maps of the period
1835-1840. Wright, in a note to Torrey, said Sand Lake was a small lake 8 or 10
miles from Kalamazoo, which is about the right distance for Austin Lake.

Sherman, St. Joseph County. Old maps show Sherman approximately at the present site
of Sturgis, and Sturgis (or usually Sturges) Prairie a little west of Sherman.

Squaw Island, Detroit River. This locality appears on a specimen labelled by Hubbard,
with date of June 16, [1837] . I cannot find any island so called; it may have been an
early name for one of the islands in the river on the Michigan side, or one of the
Canadian islands.

Sturges Prairie, St. Joseph County. See Sherman.

Thunder Bay, [Alpena County] . Specimens of Pinguicula vulgaris and Iris lacustris are
labelled “Thunder Bay.” Houghton was in this vicinity, June 13-17, 1838, and visited
Thunder Bay Island, June 15-17. The island, which is about 4 miles offshore and 12
or more miles east of Alpena, is the southernmost known Michigan locality for
Pinguicula vulgaris.

White Rock, Huron County. A specimen is labelled “White Rock, Oct. 8, 1837.” Hough¬
ton and Hubbard passed here on this date, but did not land. Houghton’s map shows
the rock 1900 feet off shore, in Sect. 29, R 16 E, T 15 N.

Whitmore Lake. Visited by a botanical party, June 21, 1838. The lake is about 10 miles
north of Ann Arbor; its southern half is in Northfield Township, Washtenaw County,
and its northern half in Livingston County.

Wolf Lake, Lenawee County. There is a lake of this name, near the highway, about 12
miles west of Clinton. A single collection is labelled as from Wolf Lake, September
14, 1838.

244

THE MICHIGAN BOTANIST

Vol. 9

GRIMMIA AGASSIZII, A MOSS NEW TO MICHIGAN

F. J. Hermann
Forest Service Herbarium,

Washington, D.C.

Five species of Grimmia (one of them, G. apocarpa, with six recognized
varieties) are ascribed to the state in Darlington’s The Mosses of Michigan
(1964) and a sixth (G. plagiopoda ) has recently been reported (Mich. Bot. 9:
140) by H. A. Crum. Still another species, G. agassizii (Sull. & Lesq.) Lesq. &
James was encountered in Baraga County by the writer on June 27, 1969 (F.
J. Hermann 22646, US, DUKE, NY, MICH). There it occurred as an extensive
colony on a granite ledge at water’s edge along the Sturgeon River, 1 1 miles
SSE. of L’Anse. An earlier collection from Keweenaw County had not at first
been recognized (F. J. Hermann 16370, MICH, CAN): on periodically
submerged rock in Gratiot River, 2 miles SW. of The Cliff, August 31, 1960.

The distribution of this species was given by G. N. Jones, in Grout’s
Moss Flora of North America (2: 11-12. 1933) as “on dry rocks, Quebec and
Ontario; British Columbia; Utah, Europe.” But the plant occurs over a much
wider range than was then supposed, the writer having collected it in Maine,
New York, Alaska, Oregon, Idaho, Montana, Wyoming, and Colorado in addi¬
tion to Michigan. Citations for five of these nine localities were given in The
Bryologist 67: 173. 1964. In every instance, except for the granite ledge along
the Sturgeon River, the moss was growing on boulders, usually granite or
gneiss, in or on the banks of streams or rivers, often blackening them in
colonies of considerable size.

A member of the subgenus Schistidium, the species is nearest allied to
the strictly maritime G. maritima Turn., but it bears a superficial resemblance
to G. alpicola var. rivularis. From this it differs markedly in its funnel-form
(rather than urceolate with round base) capsule with widely flaring mouth.
Vegetatively it differs from G. alpicola and its var. rivularis in having at least
the lower leaves narrowly lanceolate, not carinate, nor repand or dentate at
the apex and the margins not at all consistently recurved, whereas in G.
alpicola the margins remain conspicuously recurved and the leaves are so
carinate that a good proportion of them remain folded when mounted on a
slide. A form of G. agassizii with dimorphic leaves, however, is of frequent
occurrence and may be confusing. In this the upper leaves, especially those of
the sterile stems, are ovate, definitely keeled, at least toward the apex, and
often with the margins in part revolute, so that they are very suggestive of G.
alpicola; but the lower leaves are always distinctive. A further distinction
between the two species lies in the color of the plant, the lower leaves in G.
agassizii being black (never so in G. alpicola) and even the upper leaves much
darker green than those of G. alpicola. The walls of the upper leaf cells in G.
agassizii are greatly thickened. The leaf margins are thickened and bistratose,
contrary to the statement by Jones.

1970

THE MICHIGAN BOTANIST

245

MICHIGAN PLANTS IN PRINT
New Literature Relating to Michigan Botany

A. MAPS, SOILS, GEOGRAPHY, GEOLOGY

Dorr, John A., Jr., & Donald F. Eschman. 1970. Geology of Michigan. Univ. Mich. Press,
Ann Arbor. 476 pp. $15.00 [A magnificent volume which should be invaluable for
anyone interested in the natural history of the state. See review on p. 206 of the May
issue.]

Haines, Donald A., & Rodney W. Sando. 1969. Climatic Conditions Preceding Historically
Great Fires in the North Central Region. North Central For. Exp. Sta., U.S. For. Serv.
Res. Pap. NC-34. 19 pp. [Includes data on the 1871 and the 1881 (Thumb) fires in
Michigan.]

B. BOOKS, BULLETINS, SEPARATE PUBLICATIONS

Chase, Clarence D., Ray E. Pfeifer, & John S. Spencer, Jr. 1970. The Growing Timber
Resource of Michigan 1966. North Central For. Exp. Sta., U.S. For. Serv. Resource
Bull. NC-9. 62 pp. [Presents statistics from the third forest survey of Michigan,
showing sizable gains in growing-stock and sawtimber volumes since 1955. A rough
map of major forest types in Michigan is on the inside rear cover.]

Smith, Bruce D., ed. 1970. Summer Science Journal. Vol. 2, No. 1 & No. 2. Summer
Science, Inc., Ann Arbor. 105 pp. & 87 pp. $1.00 ea. [No. 1 includes student reports
on historical archaeology and independent research during 1969. The latter includes a
study on the diatoms of Summer Island and a hardwood-softwood survey of the
island. No. 2 includes reports on aquatic biology, meteorology, plant ecology (a fern
report here), shoreline geology, and wildlife ecology of Summer Island. As mentioned
in noting Vol. 1 in our May listings, the reports reveal the orthographic and other
inexperience of their youthful authors, but valuable data on this island off the Garden
Peninsula are recorded.]

C. JOURNAL ARTICLES

Bigelow, Howard E. 1970. Omphalina in North America. Mycologia 62: 1-32. [Four of
the species are cited for Michigan, without further locality.]

Byer, Michael D. 1970. Interspecific plant association as influenced by three variables.
Ecology 51: 103-112. [Changes in interspecific association related to variation in
quadrat size, position on a gradient, and abundance measure, studied in southern
Crawford Co.]

Cooperrider, T. S., & G. A. McCready. 1970. Chromosome numbers in Chelone
(Scrophulariaceae). Brittonia 22: 175-183. [2n = 28 or n = 14 reported for C. glabra
var. linifolia from Berrien and Washtenaw counties.]

Davis, H. A., Albert M. Fuller, & Tyreeca Davis. 1969. Contributions toward the revision
of the Eubati of eastern North America. IV. Castanea 34: 157-179. [This revision of
the blackberries, more correctly called subgenus Rubus, continues with the sections
Verotriviales (no species in our region), Canadenses, and Alleghenienses. In the
synonymy of R. canadensis are included R. besseyi (type from Ontonagon Co.) and
R. laetabilis (type from Kalamazoo Co.); the types of these and of R. darlingtonii
(Ontonagon Co.), which cannot be placed, are discussed. In the synonymy of R.
allegheniensis are included var. plausus (type from Lapeer Co.) and R. rappii (type
from Kalamazoo Co.), and the species is also cited from Alger, Clare, and Ontonagon
counties. R. attractus (type from Allegen Co.) is not placed. R. rosa is cited from
Gogebic Co.]

246

THE MICHIGAN BOTANIST

Vol. 9

Davis, H. A., Albert M. Fuller, Tyreeca Davis. 1969. Contribution toward the revision of
the Eubati of eastern North America. V. Arguti. Castanea 34: 235-266. [This portion
of a continuing monograph reduces Bailey’s 120 species of native glandless highbush
blackberries to 50. Kalamazoo Co. types of R. localis, R. associus, and R. licens are
reduced to R. uvidus, for which other collections are cited from Allegan, Kalamazoo,
Muskegon, and Van Buren counties; R. cauliflorus and R. limulus, both also from
Kalamazoo Co., are reduced to R. recurvans, which is also cited from Menominee Co.;
R. hanesii is an “odd plant’’ not definitely placed; R. frondosus is cited from
Kalamazoo Co.]

Dissing, Henry. 1966. A revision of collections of the genus Helvella L. ex St-Amans
emend. Nannf. in the Boudier Herbarium. Rev. Mycol. 31: 189-224 [Cites H. stevensii
from Detroit.]

Dogma, Irineo J., Jr. 1969. Additions to the phycomycete flors of the Douglas Lake
region. VIII. Chytriomyces annulatus sp. nov. and notes on other zoosporic fungi.
Nova Hedw. 18: 349-365. [Observations at various places in Michigan, chiefly the
Douglas Lake region.]

Dogma, Irineo J., Jr. 1969. Observations on some ceilulosic chytridiaceous fungi. Archiv.
Mikrobiol. 66: 203-219. [Notes on several species, mostly at various Michigan
localities.]

Giannasi, David E., & C. Marvin Rogers. 1970. Taxonomic significance of floral pigments
in Linum (Linaceae). Brittonia 22: 163-174. [The material studied of L. medium var.
texanum and L. sulcatum came from Oakland and Newaygo counties, respectively.]

Krai, R., & P. E. Bostick. 1969. The genus Rhexia (Melastomataceae). Sida 3: 387-440.
[Stress in this treatment is on cytological-anatomical data obtained since the 1956
monograph by James. Includes key to species and distribution maps; map for R.
virginica shows only Van Buren and Kalamazoo counties in Michigan, although the
species ranges north to Oceana Co.]

McKnight, Kent H. 1969. A note on Discina. Mycologia 61: 614-630. [Three species
cited from Michigan.]

Montgomery, James D., & David E. Fairbrothers. 1970. A biosystematic study of the
Eupatorium rotundifolium complex (Compositae). Brittonia 22: 134-150. [E. sessili-
folium mapped in southeastern Michigan.]

Stuckey, Ronald L. 1970. Distributional history of Epilobium hirsutum (Great Hairy
Willow-herb) in North America. Rhodora 72: 164-181. [Data, with map, include
Michigan localities with stress on sequence of collections.]

Thieret, John W. 1969. Notes on Epifagus. Castanea 34: 297-402. [Distribution map
includes many Michigan locations for beechdrops.]

Wetzel, Robert G. 1969. Factors influencing photosynthesis and excretion of dissolved
organic matter by aquatic macrophytes in hard-water lakes. Verh. Internatl. Ver.
Limnol. 17: 72-85. [Study based on growth of Naias flexilis from Lawrence and Gull
lakes, Barry and Kalamazoo cos., in axenic culture.]

1970

THE MICHIGAN BOTANIST

247

News of Botanists

STANLEY A. CAIN has been appointed the first director of the University of
Michigan’s new Institute for Environmental Quality. Professor of conservation and of
botany at the University, he was recently designated “Eminent Ecologist,” the highest
honor bestowed by the Ecological Society of America, and he has also been honored by
election to the National Academy of Sciences. . . . .DOUG FULTON, outdoor editor
for the Ann Arbor News, received the Conservation Communications Award of the year
at the Michigan United Conservation Clubs annual convention in Traverse City last
summer. . . . WILLIAM F. HOPKINS, long active in the Botanical Club, has been
elected president of the Michigan Parks Association.

MICHIGAN FLORA PROGRESS

Part I of a flora of the vascular plants of Michigan is nearing completion. It will
include keys to gymnosperms and monocots, together with distribution maps showing the
counties from which each is known. The maps will be based only on specimens actually
examined, not on literature or manuscript reports, as reliable as these may be. Therefore,
all existing collections should be examined. Many Michigan herbaria, and a few outside
the state, have been collaborating for several years, but some significant Michigan
collections have doubtless been overlooked. Persons in charge of private or institutional
herbaria containing Michigan specimens which might be made available and with which
arrangements have not yet been made for checking, are encouraged to notify Edward G.
Voss, Herbarium, North University Bldg., University of Michigan, Ann Arbor, Michigan
48104.

Publications of Interest

LICHENS OF OHIO Part 2. Fruticose and Cladoniform Lichens. By Conan J. Taylor.
Biol. Notes 4, Ohio Biological Survey, Columbus, 1968. pp. 153-227 + 22 pp., plastic
bound. $3.00 from Publications Office, Ohio State University, 2500 Kenny Rd.,
Columbus, Ohio 43210. This work continues with the same kind of excellent
photographs, distribution maps for Ohio, keys, and descriptions as in Part 1, noted
here earlier (Vol. 7, p. 189. 1968). Present plans call for Part 3, on the crustose
lichens, to complete the group with a year or two.

THE OXFORD BOOK OF FOOD PLANTS. By S. G. Harrison, G. B. Masefield, &
Michael Wallis. Oxford University Press, London, 1969. 106 pp. 55s [=$6.60].
Handsomely and thoroughly illustrated in color by B. E. Nicholson, this volume is a
popular guide to food plants of all kinds (including herbs and beverages) from all
parts of the world. There are general notes on culture, origin, distribution, parts used,
and such matters. Scientific names are given but without authorities, the families to
which the plants belong are seldom mentioned, and there are no references to other
literature. The botany student, therefore, may be somewhat frustrated, but the book
is sufficiently complete, attractive, and apparently authoritative to appeal to the
gourmet or housewife. An omission which may be missed is the recently promoted
“kiwi fruit” ( Ziziphus jujuba Mill., in the Rhamnaceae).

NOTES: A reprint edition of Natural Areas in Indiana and Their Preservation, reviewed
over a year ago (Vol. 8, p. 150, May 1969), is now available from the American Midland
Naturalist, Department of Biology, University of Notre Dame, Notre Dame, Indiana
46556. The price is $5.50 and orders placed by individuals must be prepaid. ... A list
of revisions and additions to Check-List of the Vascular Plants of the Bruce Peninsula
appears on three pages (unnumbered) in the Federation of Ontario Naturalists Newsletter,
Vol. 11, No. 3 (May-June-July, 1970). Included are some of the records presented by
George Thomson in the extended review in our January issue (Vol. 9, pp. 9-16). The
discovery of Listera ovata in the Bruce is written up in the April-June number ot
Rhodora (Vol. 72, pp. 274-275).

248

THE MICHIGAN BOTANIST

Vol. 9

MICHIGAN BASE MAP

Because a number of articles in The Michigan Botanist refer to Michigan localities
by county, we respond here to suggestions by readers, and print a base map of the State as
a preface to the Index.

1970

THE MICHIGAN BOTANIST

249

INDEX TO VOLUMES 7-9

With the exceptions here noted, this index is intended to be complete for the
three volumes, including all scientific names of genera and species, titles (by important
words), authors, and subjects. NOT indexed, in general, are the following: news items,
program notes, announcements, etc.; items in literature lists, bibliographies, and sum¬
maries; common names or names of taxa higher than genus or lower than species, except
when there is a major discussion or use of them. Reviews are ordinarily listed only (by
title) under “Reviews”— not under the author or title of the works reviewed, nor under
the reviewer’s name; “Publications of Interest” are not indexed although that heading is.
Pictures are indexed under their explanations (whether on the same page, a facing page,
or in the case of cover pictures, the rear cover). If an article has three or more joint
authors, the others are cross-referenced to the senior author without repetition of title; if
there are only two authors, both are fully indexed with title. Names of persons not
followed by a title or cross-reference refer to information about the person. Most other
references to persons are not indexed.

Some inconsistencies are inevitable as to whether or not species mentioned in
passing or in habitat descriptions are indexed. Casual mention of a species (or a genus) is
usually not indexed, especially if the point is not to give a record or to note taxonomic
comparisons. Species (or genera) in pollen diagrams are not indexed unless there is
further discussion in associated text. Also not in this index, as an economy measure, are
(1) the species cited in tables and descriptions of Pennfield Bog (8: 11-27; 51-60) since
all are included in the annotated list (8: 131-136), which is indexed; (2) the species in
the description of Sanford Natural Area (9: 130-136) as all are indexed from the
annotated list (9: 147-164); (3) the list of associates of Plantago cordata (8: 91). When
an article is devoted to a single genus and there are many species names (including
synonyms), these names are not separately indexed, as the title of the article (at least
when inverted and indexed under the generic name) should indicate its contents.
Similarly, if in an article devoted to a family there are one or more genera with several
species, the index entry will only read “[genus] spp.” to avoid listing a large number of
names in an obvious context.

The attention of phytogeographers is called to the special entry, “Distribution
Maps,” where all species are listed for which partial or complete maps have been
published.

Abies (pollen), 8: 5-7

Abies balsamea, 7: 125; 9: 56

Acalypha rhomboidea, 9: 155, 197

^nelundL, 9: 156, 182, 192
nigrum, 9: 156, 192
platanoides, 9: 156
rubrum, 8: 131; 9: 156
saccharinum, 9: 156, 199
saccharum, 9: 112, 156, 196
A cerates

lanuginosa, 8: 182
monocephala, 8: 182
Achillea millefolium, 9: 159, 198
Actaea pachypoda, 9: 149
Actinea herbacea, 9: 13
Adenocaulon bicolor, 9:16
Additional Bryophytes from Sinkholes in
Alpena County, Michigan, 9: 87-94
Additional Wisconsin Records for the Ebony
Spleenwort, Asplenium platyneuron,

7: 268

Adiantum pedatum, 9: 148

Aedes

canadensis, 7: 208
diantaeus, 7: 208
fitchii, 7: 208
intrudens, 7: 208
vexans, 7: 208, 210
Aegopodium podagraria, 9:15
Aesculus, 9: 56
glabra, 9: 239

hippocastanum, 9: 155, 196
Agalinis purpurea, 8: 136
Agaricus

mitis, 9: 23
serotinus, 9: 21
stipticus, 9: 24
Agastache nepetoides, 9: 239
Agoseris cuspidata, 8: 182
Agropyron

repens, 9: 162
trachycaulum, 9: 44
Agrostemma githago, 9: 238
Agrostis

alba, 9: 1 88

250

THE MICHIGAN BOTANIST

Vol. 9

hyemalis, 9: 197
stolonifera, 8: 135
Ailanthus altissima, 9: 58, 195
Ajuga reptans, 9:15
Albion College, herbarium, 9: 40
Aletris farinosa, 8: 61

Algae in Park Lake, Clinton County, Michi¬
gan, 9: 95-107

Alisma plantago-aquatica, 8: 131; 9: 161
Allen, Shirley W., 7: 271
Alliaria officinalis, 9: 198
Allium

canadense, 9: 195
cernuum, 9: 195
tricoccum, 9: 163, 195
Allmendinger, Elizabeth C., 9: 218-219,

227, 231, 234
Alnus, 9: 58

Alpena County, Michigan, Additional Bry-
ophytes from Sinkholes in, 9: 87-94
Althaea rosea, 9: 14, 192
Amaranthus

retroflexus, 9: 197
tuberculatus, 9: 152
Amblystegium

compactum, 8: 127
juratzkanum, 8: 127
serpens, 8: 127
varium, 8: 127
Ambrosia (pollen), 8: 5-7
Ambrosia

artemisiifolia, 9: 160, 194
trifida, 9: 160, 199
Amelanchier, 9: 57
arborea, 8: 135
laevis, 9: 154
sanguinea, 9: 195

Ammirati, Joseph F., Jr., & Alexander H.
Smith, Studies in the Genus Cortinarius,
I: Section Dermocybe, Cortinarius aurei-
folius Complex, 8: 175-180
Amorpha canescens, 9: 240
Amphicarpa bracteata, 9: 154
Amphipleura, 9: 105
Amphora, 9: 105
Anabaena

circinalis, 9: 99
flos-aquae, 9: 99-100
limnetica, 9: 99
oscillarioides, 9: 100
spiroides, 9: 99

Andrejak, Gary E., & Burton V. Barnes, A
Seedling Population of Aspens in South¬
eastern Michigan, 8: 189-202
Andresen, Norman A., Algae in Park Lake,
Clinton County, Michigan, 9: 95-107
Andromeda glaucophy 11a, 8: 133
Andropogon

gerardii, 7: 63-65; 9: 193, 240
scoparius, 7: 63-65; 9: 240
virginicus, 9: 239-240
Anemone

quinquefolia, 9: 149

virginiana, 9: 197

Animals associated with Nymphaea, 9: 78-80
Ankistrodesmus falcatus, 9: 101
Anomodon

attenuatus, 8: 126
rostratus, 8: 126

Antennaria plantaginifolia, 9: 196
Anthelia juratzkana, 7: 134
Anthemis

arvensis, 9: 160
cotula, 9: 190
Antirrhinum majus, 9: 199
Aphanizomenon flos-aquae, 9: 99
Aphanocapsa

elachista, 9: 100
pulchra, 9: 99-100
Aphanochaete

polychaete, 9: 101
repans, 9: 101
Aphanothece

gelatinosa, 9: 100
stagnina, 9: 100
Apiocystis brauniana, 9: 101
Apios americana, 8: 133
Aplectrum hyemale, 9: 164
Apocynum

androsaemifolium var. pubescens, 9: 44-
45

cannabinum, 9: 157, 190
sibiricum var. salignum, 9: 13
Aquilegia canadensis, 9: 197
Arabis

drummondii, 9: 192
laevigata, 9: 153, 189
shortii, 9: 197
Aralia

nudicaulis, 8: 132; 9: 196
racemosa, 9: 156
Arceuthobium pusillum, 7: 126
Arctium minus, 9: 160, 198
Arctostaphylos uva-ursi, 9: 239
Arcyria

denudata, 9: 211-212
nutans, 9: 212
Arenaria

serpyllifolia, 9: 197
stricta, 9: 199
Arisaema

atrorubens, 9: 199
dracontium, 9: 163
triphyllum, 9: 163
Armillaria mellea, 9: 110
Aronia melanocarpa, 8: 135
Artemisia (pollen), 8: 5-7
Artemisia caudata, 7: 64
Arthrospira jenneri, 9: 100
Asarum canadense, 9: 149
Asclepias

incarnata, 8: 132; 9: 157
lanuginosa, 8: 182
syriaca, 9: 157, 192

Asexual Reproduction in the Burning Bush,
Euonymus atropurpureus, 7: 60-61

1970

THE MICHIGAN BOTANIST

251

Asimina triloba, 9: 57, 149
Asparagus

officinalis, 9: 163, 191
Aspen, The Barnes Hybrid, Populus

xbarnesii, hybr. nov.-A Nomenclatural
Case in Point, 9: 53-54
Aspens, A Seedling Population of, in South¬
eastern Michigan, 8: 189-202
Asplenium

platyneuron, 9: 12
ruta-muraria, 9: 240
trichomanes, 9: 87

Asplenium platyneuron, Additional Wiscon¬
sin Records for, 7: 268
Aster

azureus, 7: 64-65
cordifolius, 9: 160
junciformis, 8: 132
lateriflorus, 9: 160
macrophyllus, 9: 160
pilosus, 9: 160, 193
sericeus, 9: 240
shortii, 9: 193
umbellatus, 8: 132
Asterionella, 9: 105
Asterococcus superbus, 9: 101
Asterophora lycoperdoides, An Unusual
Fruiting of, 8: 42-43
Asterotus dealbatus, 9: 29
Astreus hygrometricus, 8: 158
Athyrium

asplenioides, 9: 148
pycnocarpon, 9: 87, 148
thelypteroides, 9: 87, 148
Atrichum

angustatum, 8: 130
undulatum, 7: 198; 8: 130
Atriplex patula, 9: 193
Aulacomnium

androgynum, 8: 125
heterostichum, 8: 125
palustre, 8: 125
Auriscalpium, 7: 213
Authors, Information for, 9: 62

Bankera, 7: 213, 219
carnosa, 7: 219-222
fuligineo-alba, 7: 219-220
violescens, 7: 221
Baptisia leucophoea, 9: 240
Barbarea vulgaris, 9: 14, 153, 192
Barbula unguiculata, 8: 123
Barnes, Burton V., & Gary E. Andrejak, A
Seedling Population of Aspens in South¬
eastern Michigan, 8: 189-202
Barnes Hybrid Aspen, Populus xbarnesii,
hybr. nov.-A Nomenclatural Case in
Point, The, 9: 53-54

Bartelli, Ingrid, A Report on the 1969 Morel
Season in Michigan, 9: 200-204
Bartelmez, George W., 9:35
Bartramia pomiformis, 8: 125

Basicladia

chelonum, 9: 101
crassa, 9: 101

Bay View, Michigan, 9: 37-40
Beaman, John H., A Botanical Inventory of
Sanford Natural Area. I. The Environ¬
ment, 9: 116-139. II. Checklist of Vas¬
cular Plants, 9: 147-164
Bebb, Michael S., 7: 269; 8: 182
Beilis perennis, 9:16
Bensley, Robert Russell, 9: 35
Berberis

aquifolium, 9:14
thunbergii, 9: 150
vulgaris, 9: 14
Berteroa incana, 9: 153
Besseya bullii, 9: 238, 240
Betula, 9: 58

alleghaniensis, 8: 38-41
lenta, 8: 38-39
lutea, 8: 39, 63, 132
microphylla, 9: 13
nigra, 8: 39

papyrifera, 8: 39, 63; 9: 112
pumila, 8: 41, 132
Bidens

cernua, 8: 132; 9: 160
connata, 9: 194
coronata, 8: 132
frondosa, 9: 160, 194
Big Chicken Island, 9: 179
Bigelow, Howard E., & Alexander H. Smith,
A New Clitocybe from Michigan, 9: 30-
33

Binuclearia tatrana, 9: 101
Biology of the Mouse-ear Chickweed,
Cerastium vulgatum, 8: 151-157
Birches, Dark-Barked, of Southern Michigan,
8: 38-41

Boehmeria cylindrica, 8: 136; 9: 150, 199
Bog, An Ecological Analysis of a Southern
Michigan, 8: 11-27; A Phytogeographical
Analysis of a Southern Michigan, 8: 51-60
Bog, Pennfield, 8: 11-27,51-60, 131-136
Bolbitius, 7: 19, 21, 23
Borzia triocularis, 9: 100, 102
Botanical Activities of Thomas J. Hale,
1858-1862, The, 8: 181-185
Botanical Inventory of Sanford Natural
Area, A. I. The Environment, 9: 116-
139. II. Checklist of Vascular Plants, 9:
147-164

Botanical Results of the Michigan Geologi¬
cal Survey Under the Direction of Doug¬
lass Houghton, 1837-1840, 9: 213-243
Botham, Wilfred, & Howard Crum, Funaria
flavicans, a Moss New to Canada, 7: 24
Botrychium

lunaria f. onondagense, 9: 87
virginianum, 9: 148
Botryococcus
braunii, 9: 101
sudeticus, 9: 101

252

Vol. 9

THE MICHIGAN BOTANIST

Bourdo, Eric A., Jr., 8: 145
Bouteloua curtipendula, 9: 240
Bowers, Maynard C., New Localities for
Mosses Rarely Found in Michigan’s
Upper Peninsula, 7: 93-94
Brachyelytrum erectum, 9: 162
Brachymenium mexicanum, 7: 201
Brachythecium

acuminatum, 8: 128
campestre, 8: 128
curtum, 7: 132
digastrum, 9: 141
oxycladon, 8: 128; 9: 141
plumosum, 8: 128
rivulare, 8: 128
rutabulum, 8: 128
salebrosum, 8: 128
starkei, 7: 132
velutinum, 8: 128

Bradford, Laura S., Myxomycetes from
Cheboygan and Emmet Counties, Mich¬
igan, 9: 211-212
Brasenia schreberi, 8: 134
Brassica

campestris, 9: 153
kaber, 9: 189

Brinsmade, Thomas C., 9: 237
Bromus

inermis, 9: 162
latiglumis, 9: 162
purgans, 9: 162
racemosus, 9: 188
secalinus, 9: 188
tectorum, 9: 162
Brotherella recurvans, 8: 129
Bruce Peninsula, Vascular Plants of the: A
Review, with Comments and Additions,

9: 9-16
Bryhnia

graminicolor, 8: 128
novae-angliae, 8: 128

Bryophytes, Additional, from Sinkholes in
Alpena County, Michigan, 9: 87-94
Bryophytes New to Michigan, 7: 132-134;

II, 8: 28-29
Bryum

angustirete, 8: 1 24
argenteum, 8: 124; 9: 68-69, 93
caespiticium, 8: 124
capillare, 8: 124
pseudotriquetrum, 8: 124
Buchnera americana, 9: 239
Bull, George H., 9: 214, 216, 225-227, 229,
231-234, 236-238, 240
Burning Bush, Euonymus atropurpureus,
Asexual Reproduction in, 7: 60-61
Butomus umbellatus (Flowering-rush), Distri¬
butional History in the Western Lake Erie
and Lake St. Clair Region, 7: 134-142
Buxbaumia aphylla, 7: 272
Byron Bog in Southwestern Ontario, Studies
of the, XL. Distribution of Sphagnum
Mosses, 8: 167-170

Cacalia tuberosa, 9: 239
Cain, Stanley A., 7: 271 ; 9: 63, 247
Cakile edentula, 9: 199, 239
Calamagrostis

canadensis, 9: 162
inexpansa, 7: 5-6, 9

Calhoun County, Michigan, Species of Vascu¬
lar Plants of Pennfield Bog, 8: 131-136
Calla palustris, 8: 132
Calliergidium pseudostramineum, 7: 132
Calliergon cordifolium, 8: 127
Calliergonella

cuspidata, 8: 127
schreberi, 8: 127

Calodon, 7: 213-218, 222, 227, 236, 241-242
Calothrix

atricha, 9: 100
breviarticulata, 9: 100
epiphytica, 9: 100
stellaris, 9: 100
Caltha palustris, 9: 149
Camassia scilloides, 9: 195
Campanula

americana, 9: 159, 193, 198
aparinoides, 8: 132
rapunculoides, 9: 13

Campbell, Ella O., Marchantia polymorpha in
Northern Michigan, 8: 146-150; Morpholo-
ogy of Fungal Association in Three Species
of Corallorhiza in Michigan, 9: 108-113
Campsis radicans, 9: 190
Campylium

chrysophyllum, 8: 127
hispidulum, 8: 127
polygamum, 9: 93
radicale, 8: 29
stellatum, 8: 127
Cannabis sativa, 9: 238
Canby, Wm. M., 8: 182
Cantharellaceae of Michigan, The, 7: 143-183
Cantharellus spp., 7: 143-183
Cantlon, John E., 9: 63
Capsella bursa-pastoris, 9: 153, 194
Cardamine

bulbosa, 9: 153
douglassii, 9: 153
Carduus acanthoides, 9:15
Carex

albursina, 9: 161
alopecoidea, 9: 161
aquatilis, 8: 132
blanda, 9: 161, 188
brevior, 9: 191
canescens, 8: 132
carey ana, 9: 161
cephaloidea, 9: 161
cephalophora, 9: 195
chordorrhiza, 8: 133
comosa, 8: 133
convoluta, 9: 161
eburnea, 9: 195
frankii, 9: 188
grayii, 9: 161

1970

THE MICHIGAN BOTANIST

253

interior, 8: 133
intumescens, 9: 161
lasiocarpa, 8: 133
lupulina, 9; 161
molesta, 9: 161
muskingumensis, 9: 161
pedunculata, 9: 162
plantaginea, 9: 162
rosea, 9: 199
sprengelii, 9: 162, 188
stricta, 9: 162
trisperma, 8: 133
woodii, 9: 162

Carpinus caroliniana, 9: 58, 151
Carya, 9: 58

cordiformis, 9: 151
ovata, 9: 151, 195
Cassandra calyculata, 8: 133
Castalia, 9: 72
Castanea dentata, 9: 151
Castelleja sessiliflora, 8: 182
Catalpa, 9: 56

Caulophyllum thalictroides, 9: 150, 208
Celastrus scandens, 9: 155, 192
Celtis occidentalis, 9: 58, 150, 191
Centaurium umbellatum, 9: 13
Cephalanthus occidentalis, 8: 135; 9: 159, 190
Cephalozia

catenulata, 9: 93
media, 9: 93-94
Cerastium

arvense, 9: 195
fontanum, 8: 151
nutans, 9: 189

vulgatum, 8: 151-157; 9: 197
Cerastium vulgatum, Biology of, 8: 151-157
Ceratiomyxa fructiculosa, 9: 211
Ceratium hirundinella, 9: 99, 105
Ceratodon purpureus, 8: 123
Ceratophyllum demersum, 8: 132; 9: 149
Cercis canadensis, 9: 57
Cetraria

ciliaris, 8: 67-71
halei, 8: 67-71
microphyllica, 8: 67
orbata, 8: 67-71

Cetraria ciliaris, in the Straits Region of
Michigan, The Occurrence of the Lichen
Complex, 8: 67-71
Chaenorrhinum minus, 9: 13
Chaetophora

attenuata, 9: 101
incrassata, 9: 101
Chaetosphaeridium ovalis, 9: 101
Chamaesiphon incrustans, 9: 100
Changes in the Vascular Flora of Seven
Small Islands in Western Lake Erie,

9: 175-200
Chara

globularis, 9: 7, 104
vulgaris, 9: 7, 104
Characium

ambiguum, 9: 101

dabaryanum, 9: 101
falcatum, 9: 101

Charles Henry Swift -188 1-1 966, 9: 34-40
Charlevoix County, Michigan, Sphagnum in,
9: 114-115

Cheboygan and Emmet Counties, Michigan,
Myxomycetes from, 9: 211-212
Chelone glabra, 9:158
Chenopodium

album, 9: 151, 193
hybridum, 9: 189
Chickering, J. W., 8: 182
Chickweed, Mouse-ear, Biology of the, 8:
151-157

Chlamydomonas
globosa, 9: 100
polypyrenoideum, 9: 100
pseudopertyi, 9: 100
Chlorella vulgaris, 9: 101
Chromosome Studies of Some Mosses of the
Douglas Lake Region, 9: 67-71
Chroococcus

dispersus, 9: 100
giganteus, 9: 100
limneticus, 9: 100
minimus, 9: 100
minor, 9: 100
minutus, 9: 100
prescottii, 9: 100
turgidus, 9: 100
varius, 9: 100

Chrysanthemum leucanthemum, 9: 196
Cichorium intybus, 9: 160
Cinna arundinacea, 9: 162
Circaea quadrisulcata, 9:155
Cirsium

arvense, 9: 160
muticum, 9: 15
pitcheri, 9: 241
vulgare, 9: 160, 194
Cladonia

chlorophaea, 8: 170-174
cryptochlorophaea, 8: 171-174
grayi, 8: 171-174
merochlorophaea, 8: 171-173
Cladonia chlorophaea group, Population
Studies of the, 8: 170-174
Cladophora

crispata, 9: 101
insignis, 9: 101

Clark, Howard L., & John W. Thieret, The
Duckweeds of Minnesota, 7: 67-76
Clark, Mary H., 7: 269
Clavaria spp. (see Clavariadelphus)
Clavariadelphus, 7: 143-144
Clavariadelphus, A Preliminary Study of, in
North America, 7: 35-57
Claytonia

caroliniana, 7: 77-93; 9: 13
virginica, 7: 77-93, 96; 9: 13, 151, 189
Claytonia in Michigan, 7: 77-93
Clinton County, Michigan, Algae in Park
Lake, 9: 95, 107

254

THE MICHIGAN BOTANIST

Vol. 9

Clintonia borealis, 9: 239
Clematis verticillaris, 9: 14
Climacium

americanum, 8: 126
dendroides, 8: 126
Clitocybe

benekei, 9: 30-33
irina, 9: 30, 33
subconnexa, 9: 33

Clitocybe, A New, From Michigan, 9: 30-33
Closteriopsis longissima, 9: 101
Closterium

dianae, 9: 104
ehrenbergii, 9: 104
jenneri, 9: 104
leiblenii, 9: 104
libellula, 9: 104
monoliferum, 9: 104
parvulum, 9: 104
venus, 9: 104
Cocconeis, 9: 105
Coelastrum microporum, 9: 101
Coelastrum, Review of the Status of Some
Green Algae in the Genus, 7: 129-131
Coelosphaerium
dubium, 9^ 100
kuetzingianum, 9: 99, 100
naegelianum, 9: 99, 100
Coleochaete

irregularis, 9: 101
scutata, 9: 101

Collinsonia canadensis, 9: 158
Comandra umbellata, 9: 188
Commelina

communis, 9: 198
erecta, 9: 239

Conocephalum conicum, 8: 147; 9: 94
Conocybe, 7: 19, 23
Conopholis americana, 9: 13
Contact Dermatitis Caused by Plants, 7:
265-268

Convallaria majalis, 7: 266
Convolvulus

arvensis, 9: 191
sepium, 9: 192

Conyza canadensis, 9: 46, 160
Cooley, Dennis, 7: 6
Coptis trifolia, 8: 135; 9: 239
Corollorhiza

innata, 9: 108, 113
maculata, 9: 108-113, 164
striata, 9: 111-113
trifida, 9: 108-110, 113
Corollorhiza in Michigan, Morphology of the
Fungal Association in Three Species of,

9: 108-113
Coreopsis

palmata, 9: 240
tripteris, 9: 240
Cornus, 9: 57

alternifolia, 9: 58, 155
amomum, 9: 155
drummondii, 9: 197

florida, 9: 155
foemina, 9: 155
obliqua, 9: 196
stolonifera, 8: 132; 9: 155
Cortinarius

aureifolius, 8: 175-177, 180
chrysolitus, 8: 175
cinnamomeus, 8: 175-176
psammophilus, 8: 177-179
raphanoides, 8: 175
sanguineus, 8: 175

Cortinarius, Studies in the Genus, I: Section
Dermocybe, Cortinarius aureifolius Com¬
plex, 8: 175-180
Corydalis sempervirens, 9: 13
Corylus, 9: 58

americana, 8: 132
Cosmarium

angulare, 9: 104
angulosum, 9: 104
botrytis, 9: 104
conspersum, 9: 102, 104
denticulatum, 9: 104
depressum, 9: 102, 104
garrolense, 9: 102, 104
humile, 9: 102, 104
minutissimum, 9: 102, 104
obtusatum, 9: 104
portianum, 9: 104
pseudopyramidatum, 9: 104
regnellii, 9: 102, 104
reniforme, 9: 104
sexangulare, 9: 104
sportella, 9: 102, 104
subcostatum, 9: 104
subtumidum, 9: 104
trilobulatum, 9: 104
tumidum, 9: 104
turpinii, 9: 104
undulatum, 9: 104
Coulter, John Merle, 9: 38
Crataegus, 9:57

flexuosus, 9: 43, 46
macrosperma, 9: 154
mollis, 9: 154
punctata, 9: 154
succulenta, 9: 154
Craterellus

caeruleofuscus, 7: 149-151, 172

calyculus, 7: 149-150

cinereus var. multiplex, 7: 149, 151, 173

cornucopioides, 7: 149, 152-154

corrugis, 7: 47

fallax, 7: 149, 153-154, 175

foetidus, 7: 149, 152, 174

ochrosporus, 7: 153

pistillaris, 7: 42

sinuosus, 7:151

unicolor, 7: 47

Cratoneuron filicinum, 8: 127
Crow, Garrett E., An Ecological Analysis of
a Southern Michigan Bog, 8: 1 1-27; A
Phytogeographical Analysis of a South-

1970

THE MICHIGAN BOTANIST

255

ern Michigan Bog, 8: 51-60; Species of
Vascular Plants of Pennfield Bog, Cal¬
houn County, Michigan, 8: 131-136
Crucigenia rectangularis, 9: 101
Crum, Howard, Mosses New to the Flora of
Michigan, 9: 139-141

Crum, Howard, & Wilfred Botham, Funaria
flavicans, a Moss New to Canada, 7: 24
Crum, Howard, & Norton G. Miller, Bry-
ophytes New to Michigan, 7: 132-134;
BryophytesNew to Michigan. II, 8: 28-29
Crum, Howard, & William M. Zales, Sphag¬
num in Charlevoix County, Michigan,

9: 114-115

Cryptotaenia canadensis, 9: 156
Ctenidium molluscum, 8: 129
Curtis, Moses A., 8: 182
Cyclotella, 9: 105
Cylindrocapsa conferta, 9: 101
Cymatopleura, 9: 105
Cymbella, 9: 105
Cynanchum nigrum, 9: 157
Cyperus

alterniflorus, 9: 43-44
diandrus, 8: 133
engelmannii, 8: 133
erythrorhizos, 9: 198
ferruginescens, 9: 198
schweinitzii, 7: 64; 9: 44
Cypripedium, 7: 265
acaule, 8: 134

Cystodinium cornifax, 9: 105
Cystopteris

bulbifera, 9: 87, 148
dickieana, 9: 87
fragilis, 9: 87, 148

Dactylis glomerata, 9, 162, 193
Dactylococcopsis fascicularis, 9: 100
Dancik, Bruce P., Dark-Barked Birches of
Southern Michigan, 8: 38-41
Dark -Barked Birches of Southern Michigan,
8: 38-41

Datura tatula, 9: 238
Daucus carota, 7: 265; 9: 157, 194
Decodon verticillatus, 8: 134
Dentaria laciniata, 9: 14, 153, 189
Dentinum, 7: 213

Dermatitis, Contact, Caused by Plants, 7:
265-268

Descurainia pinnata, 9: 189
Desmatodon obtusifolius, 8: 123
Desmidium

aptogonum, 9: 104
bailey i, 9: 104

Desmodium nudiflorum, 9: 154
Diachea leucopodia, 9: 211-212
Diarrhena americana, 9: 240
Dicentra

canadensis, 9: 150
cucullaria, 9: 150, 195
Dicranella

cerviculata, 7: 132; 8: 29
heteromalla, 8: 123

schreberiana, 7: 132-133
Dicranum

bonjeanii, 8: 123
flagellare, 8: 123; 9: 68-70, 93-94
fulvum, 8: 123
montanum, 8: 123; 9: 92
polysetum, 8: 123
scoparium, 8: 123; 9: 68-70
viride, 8: 123; 9: 92
Diderma spumarioides, 9: 212
Didymodon rigidulus, 9: 93
Digitaria

filiformis, 9: 239
sanguinalis, 9: 198
Dinobryon

cylindricum, 9: 105
sertularia, 9: 105
tabellariae, 9: 105
Dioscorea villosa, 9: 164, 188
Diospyros virginiana, 9: 57
Diphyscium foliosum, 8: 130; 9: 93
Dipsacus laciniatus, 9: 159
Dirca palustris, 9: 155

Discomycetes, Odd and Unusual, from Mich¬
igan, I, 7: 58-59
Dispora crucigenioides, 9: 101
Distribution maps:

Abies balsamea, 7: 125

Butomus umbellatus, 7: 137, 139

Calla palustris, 8: 59

Cetraria ciliaris, 8: 69

Cetraria halei, 8: 69

Cetraria orbata, 8: 69

Claytonia caroliniana, 7: 85

Clay tonia virginica, 7 : 84

Cornus stolonifera, 8:58

Gentiana linearis, 7: 101, 108-109

Gentiana rubricaulis, 7: 101, 108-109

Juniperus communis, 7: 128

Juniperus horizontalis, 7: 128

Juniperus virginiana, 7: 128

Larix laricina, 7: 124; 8: 58

Lemna minor, 7: 74

Lemna trisulca, 7: 74

Ly copus asper, 8: 116

Nuphar luteum subsp. macrophyllum, 8:

53

Picea glauca, 7: 126
Picea mariana, 7: 126
Pinus banksiana, 7: 125
Pinus resinosa, 7: 125
Pinus strobus, 7: 124
Plantago cordata, 8: 89
Rhynchospora alba, 8:56
Rubus hispidus, 8: 55
Sarracenia purpurea, 8:55
Spiraea alba, 8: 56
Spirodela polyrhiza, 7: 74
Taxus canadensis, 7: 124
Thuja occidentalis, 7: 127
Tsuga canadensis, 7: 126
Typha latifolia, 8: 53
Wolffia columbiana, 7: 74
Wolffia punctata, 7: 74

256

THE MICHIGAN BOTANIST

Vol. 9

Distributional History of Butomus umbellatus
(Flowering-rush) in the Western Lake
Erie and Lake St. Clair Region, 7: 134-142
Ditrichum

flexicaule, 9: 92
pallidum, 8: 123

Dr. Bessie Bernice Kanouse-1 889-1969, 8:
186-188

Dorcadion pallens, 9: 89
Douglas Lake Region, Chromosome Studies
of Some Mosses of the, 9: 67-71
Douglas, David B., 9: 213
Drepanocladus
aduncus, 8: 127
uncinatus, 8: 127
Drosera

intermedia, 8: 133
rotundifolia, 8: 133
Dryopteris spp., 8: 137-145
Dryopteris

filix-mas, 9: 87
goldiana, 9: 148
spinulosa, 9: 87, 148
spinulosa var. fructuosa, 9: 12
Dryopteris celsa and Its Hybrids in Michi-
gan-A Preliminary Report, 8: 137-145
Duckweeds of Minnesota, The, 7: 67-76
Dulichium arundinaceum, 8: 133
Duncan, Thomas, & Ronald L. Stuckey,
Changes in the Vascular Flora of Seven
Small Islands in Western Lake Erie, 9:
175-200

Durand, Elias, 8: 182

Eaton, Amos, 9: 213-214, 238
Eaton, Daniel Cady, 8: 182
Eastwood, John F. 9: 217, 219
Echinacea purpurea, 9: 240
Echinochloa

crusgalli, 9: 199
muricata, 9: 162

Echinocystis lobata, 8: 132; 9: 152, 191
Echinodontium, 7: 213
Echinodorus parvulus, 9: 241
Eclipta alba, 9: 191

Ecological Analysis of a Southern Michigan
Bog, 8: 11-27
Ehlers, John H., 7: 271
Elakatothrix viridis, 9: 101
Eleocharis

acicularis, 9: 162
compressa, 8: 133
equisetoides, 9: 239
erythropoda, 9: 198
intermedia, 8: 133
smallii, 8: 133
Eleusine indica, 9: 238
Elodea nuttallii, 9: 161
Elymus

canadensis, 9: 194
mollis, 7: 192
riparius, 9: 162
villosus, 9: 162, 197

virginicus, 9: 162, 191
Emmet and Cheboygan Counties, Michigan,
Myxomycetes from, 9: 211-212
Engelmann, George, 8: 182
Entodon

cladorrhizans, 8: 129
seductrix, 8: 129
Ephemerum crassinervium, 8: 29
Epifagus virginiana, 9: 159
Epigaea repens, 8:61
Epilobium strictum, 8: 134
Epipactis helleborine, 9: 164
Epithemia, 9: 105
Equastrum

hypochondrum, 9: 104
turneri, 9: 102, 104
verrucosum, 9: 104
Equisetum

arvense, 9: 148, 168
xferrissii, 9: 167
fluviatile, 8: 133; 9: 168
hyemale, 9: 148, 166-174
laevigatum, 9: 167
xlitorale, 7:115
xnelsonii, 9: 168, 172
variegatum, 9: 168

Equisetum hyemale, Natural Proliferation of
Floating Stems of, 9: 166-174
Eragrostis

megastachya, 9: 198
pectinacea, 9: 198
pilosa, 9: 162

Erechtites hieracifolia, 9: 194
Erigenia bulbosa, 9: 157
Erigeron

annuus, 9: 160, 193
canadensis, 9: 194

var. grandiflorum, 9: 46
philadelphicus, 9: 160, 193
strigosus, 7: 64; 9: 191
Eriophorum

angustifolium var. megastachyon, 9: 43-
44

gracile, 8: 133
virginicum, 8: 133
viridi-carinatum, 9: 43-44
Erucastrum gallicum, 9: 14
Eryngium yuccifolium, 9: 239
Erysimum cheiranthoides, 9: 153
Erythronium

albidum, 9: 163
americanum, 9: 163, 188
Ethnobotany of Plantago cordata, 8: 99-100
Eucladium verticillatum, 8: 28
Eudorina elegans, 9: 100
Euglena

elastica, 9: 104
fusca, 9: 102, 105
gracilis, 9: 105
proxima, 9: 105
Eunotia, 9: 105
Euonymus

atropurpureus, 9: 14, 155, 190

1970

THE MICHIGAN BOTANIST

257

obovatus, 9: 155, 192
Euonymus atropurpureus, Asexual Repro¬
duction in, 7: 60-61
Eupatorium

maculatum, 9: 160
perfoliatum, 8: 132
purpureum, 8: 132, 160
rugosum, 9: 160
sessilifolium, 9: 240
Euphorbia, 7: 266
corollata, 7: 64-65
cyparissias, 9: 14, 195
esula, 9: 14
maculata, 9: 190
marginata, 7: 266; 9: 14
polygonifolia, 9: 239
supina, 9: 197
Eurhynchium
hians, 8: 128

pulchellum, 7: 199-200; 8: 128; 9: 93

Fagus (pollen), 8: 8-9
Fagus grandifolia, 9: 57, 112, 151
Fallass, Charles W., 9: 40
Fenwick, Mason G., Review of the Status of
Some Green Algae in the Genus Coel-
astrum, 7: 129-131

Fern, Log, (Dryopteris celsa) and Its Hybrids
in Michigan- A Preliminary Report, 8:
137-145

Fernwood Nature Study Area, 8: 34-35
Festuca

obtusa, 9: 162
ovina, 9: 162
pratensis, 9: 162
Fimbristylis drummondii, 9: 239
Fissidens

adianthoides, 8: 122
bryoides, 8: 122
bushii, 7: 133
cristatus, 8: 122; 9: 69-70
minutulus, 9: 92
osmundoides, 8: 122; 9: 69-70
subbasilaris, 9: 140
taxifolius, 8: 122

Flowering of Duckweeds, 7: 67-68
Folwell, N.W., 7: 9
Fontinalis antipyretica, 7: 201
Fragaria

vesca, 9: 192
virginiana, 7: 63-64
Fragilaria, 9: 105
Fraxinus, 9: 56

americana, 9: 158, 192
nigra, 9: 158, 190
pennsylvanica, 9: 194, 196
quadrangulata, 9: 158, 197
Frullania

bolanderi, 9: 92
eboracensis, 9: 94
Fuirena pumila, 9: 239
Fuligo septica, 9: 212
Fulton, Doug, 9: 247

Fumaria officinalis, 9: 189
Funaria

flavicans, 7: 24
hygrometrica, 8: 124

Funaria flavicans, a Moss New to Canada, 7:
24

Fungal Association in Three Species of
Corallorhiza in Michigan, Morphology
of the, 9: 108-113

Galeopsis tetrahit, 9: 238
Galeropsis, 7: 19, 21
Galinsoga parviflora, 9: 199
Galium

aparine, 9: 159, 193
circaezans, 9: 159
mollugo, 9: 190
obtusum, 9: 159
trifidum, 8: 135
triflorum, 9: 159, 199
Gastrocybe, 7: 19-20
lateritia, 7 : 20-23
Gastrodia

cunninghamii, 9: 113
minor, 9:113
Gaultheria

hispidula, 9: 239
procumbens, 8: 63, 133
Gazetteer (of First Survey collecting sites in
Michigan), 9: 242-243
Geminella

interrupta, 9: 101
minor, 9: 101
Gentiana

affinis, 7:110
alba, 7: 105, 110
andrewsii, 7: 105, 110
austromontana, 7: 110
autumnalis, 7: 105, 110
catesbaei, 7: 105, 110
clausa, 7: 105, 110
decora, 7:110
linearis, 7: 99-112
pennelliana, 7: 1 10
puberulenta, 7: 105, 110
rubricaulis, 7: 99-112; 9: 45
saponaria, 7: 105, 110
villosa, 7: 105, 110

Gentiana linearis and G. rubricaulis in the
Upper Great Lakes Region, The Status
and Distribution of, 7: 99-112
Gentianella quinquefolia, 8: 181
Genus Panellus in North America, The, 9:
17-30

Geocalyx graveolens, 9: 93
Geological Survey, Michigan, Botanical Re¬
sults of the, Under the Direction of
Douglass Houghton, 1837-1840, 9: 213-
243

Geranium

carolinianum, 9: 192
maculatum, 9: 156
pusillum, 9: 197

258

THE MICHIGAN BOTANIST

Vol. 9

robertianum, 9: 192
Gerardia tenuifolia var. asperula, 8: 184
Geum canadense, 9: 154, 195
Gilbraltar Island, 9: 178, 180
Gillette, Genevieve, 7: 271; 9: 63
Ginkgo, 7: 265
biloba, 9: 56

Gleditsia triacanthos, 9: 14, 57-58, 154, 189
Glenodinium pulvisculus, 9: 105
Gloeocystis
ampla, 9: 101
gigas, 9: 101
planctonica, 9: 101

Gloeotaenium loitelsbergerianum, 9: 101
Gloeothece rupestris, 9: 100
Gloeotrichia

echinulata, 9: 100
pisum, 9: 100
Gloiodon, 7: 213
Glyceria

canadensis, 8: 135
striata, 8: 135; 9: 162
Gomphonema, 9: 105
Gomphosphaeria
aponina, 9: 100
lacustris, 9: 100
Gomphus

bonari, 7: 145, 148
clavatus, 7: 145, 168-170
floccosus, 7: 145, 147-148, 171
pseudoclavatus, 7: 145-147
Gongrosira debaryana, 9: 101
Gonium pectorale, 9: 100
Graham, William L., The Occurrence of the
Lichen Complex, Cetraria ciliaris, in the
Straits Region of Michigan, 8: 67-71
Gratiot County, Michigan, Pollen Analysis
at the Thaller Mastodon Site, 8: 3-10
Gray, Asa, 7: 77; 8: 182; 9: 227-228, 241;
North American Gramineae and
Cyperaceae, 7: 5-9
Green Island, 9: 180
Grimmia

agassizii, 9: 244
alpicola, 9: 244
anodon, 9: 140
apocarpa, 8: 124; 9: 140, 244
maritima, 9: 244
pilifera, 7: 133
plagiopodia, 9: 140, 244
Grimmia agassizii, a Moss New to Michigan,
9: 244

Grindelia squarrosa, 9: 13
Gustafson, Felix G., 8: 145
Gymnandra bullii, 9: 238, 240
Gymnocladus dioica, 9: 154, 195
Gymnosperms in Michigan, A Preliminary
Report on Distribution of, 7: 121-128
Gymnostomum

aeruginosum, 9: 92
recurvirostrum, 8: 123
Gyrosigma, 9: 105

Habenaria

ciliaris, 8: 134
clavellata, 8: 134
dilatata, 7: 210; 9: 12
hyperborea, 7: 210; 9: 12
macrophylla, 9: 12
media, 9: 12
obtusata, 8: 134
psycodes var. ecalcarata, 9: 40
psy codes var. varians, 9: 40
Habenaria obtusata (Orchidaceae), Mosquito
Pollination of, 7: 203-212
Hackelia virginiana, 9: 15, 157
Hadley, James, 7: 8

Hagenah, Dale J., 7: 271 ; 9: 63; (see W. H.
Wagner, Jr., et al.)

Halbert, Richard L., & Paul L. Redfeam, Jr.,
Mosses from Southern Michigan: New
Distribution Records, 8: 120-130
Hale, T. J., 7: 269; 8: 36; The Botanical Ac¬
tivities of, 1858-1862, 8: 181-185
Hall, Elihu, 7: 269
Hamamelis virginiana, 9: 57, 150
Hammitt, William E., & Warren H. Wagner,
Jr., Natural Proliferation of Floating
Stems of Scouring-rush, Equisetum
hyemale, 9: 166-174
Haplocladium microphyllum, 8: 126
Harrington, Mark W., 9: 216-217, 220, 231,
234

Harrison, Kenneth A., Studies on the Hyd-
nums of Michigan. I. Genera Phellodon,
Bankera, Hydnellum, 7: 212-264
Harvey, Basil C. H., 9: 37
Hedeoma pulegioides, 9: 13, 190
Held, Edith R., & Ronald O. Kapp, Pollen
Analysis at the Thaller Mastodon Site,
Gratiot County, Michigan, 8: 3-10
Helianthus

decapetalus, 9: 160
occidental^, 7: 63-64
Helvetia

esculenta, 9: 201-204
gigas, 9: 201-203
Hemerocallis fulva, 9: 164, 195
Hemicarpha micrantha, 9: 239
Hen Island, 9: 179-180
Hepatica

acutiloba, 9: 149, 192
americana, 9: 189

Herbaria, Rare Old Publications in Michigan,
7: 3-13
Herbarium,

Albion College, 9: 40
Allmendinger, Elizabeth C., 9: 217, 227
Gray, 9: 227

Houghton, Douglass, 7: 3; 9: 222-223,
227

Milton College, 7: 269-270 (see also 8:
35-37)

N. Y. Botanical Garden, 9: 227
Pitcher, Zina, 9: 227

1970

THE MICHIGAN BOTANIST

259

Sager, Abram, 9: 217
Thurber, George, 9: 227
Tracy, 9: 227

University of Michigan, 7: 1012; 8: 187;

9: 216-217, 223, 227, 234, 240-241
Watson, Samuel H., 8: 35-37
Wayne State University, 7: 5-9
Hericium, 7: 213
erinaceus, 7: 32

Hermann, F. J., Grimmia agassizii, a Moss
New to Michigan, 9: 244
Herrick, C. Judson, 9: 37
Hesperis matronalis, 9: 14, 153
Heterophyllium haldanianum, 8: 129; 9: 93
Heuchera americana, 9: 192
Hibiscus trionum, 9: 238
Hickory Corners, 9: 3
Hieracium

canadense var. scabrum, 9: 43, 46
florentinum, 9: 15
pilosella, 9:15
pratense, 9: 16
scabriusculum, 9: 46
Holomitrium, 7: 198-199
Homomallium adnatum, 8: 129
Hopkins, William F., 9: 247
Hormidium klebsii, 9^ 101
Hortus Siccus Londinensis, by Mariano
Lagasca, 7: 11-13

Hoseney, Florence V., An Unusual Fruiting
of Asterophora lycoperdoides, 8: 42-43;
An Unusual Fruiting of Leotia lubrica,

8: 158-159

Houghton, D., herbarium, 7: 3; 9: 222-223,227
Houghton, Douglass, Botanical Results of
the Michigan Geological Survey Under
the Direction of, 1837-1840, 9: 213-243
Hubbard, Bela, 9: 215-216, 218, 222-224,
231, 234, 236-237
Humulus lupulus, 9: 195
Hyalotheca

dissiliens, 9: 104
mucosa, 9: 104
Hybanthus concolor, 9: 152
Hybrid Aspen, The Barnes, Populus

xbarnesii, hybr. nov.-A Nomenclatural
Case in Point, 9: 53-54
Hybrids, The Log Fern (Dryopteris celsa)
and Its, in Michigan -A Preliminary Re¬
port, 8: 137-145

Hydnellum spp., 7: 213-214, 222-264
Hydnum spp., 7: 214-242
Hydnums of Michigan, Studies on the, I.
Genera Phellodon, Bankera, Hydnellum,

7: 212-264

Hydrastis canadensis, 9: 149, 239
Hydrocotyle umbellata, 8: 132
Hydrophyllum

appendiculatum, 9: 157, 198, 239
virginianum, 9: 157, 196
Hygroamblystegium
fluviatile, 8: 127
tenax, 8: 127

Hypericum

boreale, 8: 133
perforatum, 7: 64-65; 9:152
Hypnum

curvifolium, 8: 129
imponens, 8: 130
lindbergii, 8: 130
pallescens, 8: 130
pratense, 8: 130
Hystrix patula, 9: 162, 194

Ilex verticillata, 8: 132; 9: 58, 155, 190
Impatiens

biflora, 8: 132
capensis, 9: 194
noli-tangere, 9: 156
pallida, 9: 192

Information for Authors, 9:62
Introduction and Spread of Ly copus asper
(Western Water Horehound) in the West¬
ern Lake Erie and Lake St. Clair Region,
The, 8: 111-120
Inula helenium, 9: 13
Investigations in the White Waterlilies
(Nymphaea) of Michigan, 9: 72-86
Iris

lacustris, 9: 243
versicolor, 8: 133
virginica, 9: 164
Islands (see name of Island)

Islands in Western Lake Erie, Changes in the
Vascular Flora of Seven Small, 9: 175-
200

Isopterygium

deplanatum, 8: 129
distichaeceum, 7: 133
subfalcatum, 9: 141
Isopyrum bitematum, 9: 149
Isotria verticillata, 8: 134

Jackson, H., 7: 269
James, Edwin, 9: 41-42; 213
Judd, William W., Studies of the Byron Bog
in Southwestern Ontario. XL. Distribu¬
tion of Sphagnum Mosses, 8: 167-170
Juglans, 9:58
cinerea, 9: 151
nigra, 9: 64, 151
Juncus

brevicaudatus, 8: 133
dudleyi, 9: 191
effusus, 8: 133
tenuis, 9: 161, 198
Jungermannia schiffneri, 9: 92
Juniperus

communis, 7: 127-128
horizontalis, 7: 128
virginiana, 7: 128; 9: 56, 197

Kalmia polifolia, 9:15
Kanouse, Dr. Bessie Bernice, 1889-1969, 8:
186-188; 145

260

THE MICHIGAN BOTANIST

Vol. 9

Kapp, Ronald O., Natural Area Preservation
in the Age of the Megalopolis, 8: 30-35
Kapp, Ronald O., & Edith R. Held, Pollen
Analysis at the Thaller Mastodon Site,
Gratiot County, Michigan, 8: 3-10
Kempton, Phyllis E., & Virginia L. Wells, A
Preliminary Study of Clavariadelphus in
North America, 7: 35-57
Key, Winter Field, to Some Trees Common
in Michigan, 9: 55-58
Koehleria cristata, 7: 9
Kuhnia eupatorioides, 9: 239

Lactuca

canadensis, 9: 197
floridana, 9: 191
scariola, 9: 194

Lagasca, Mariano, Hortus Siccus Londinen-
sis, 7: 11-13

Lake Erie, Changes in the Vascular Flora of
Seven Small Islands in Western, 9: 175-
200

Lamium amplexicaule, 9: 194

Lapham, Increase A., 7: 269; 8: 182; 9: 215

Laportea canadensis, 9: 150, 188

Lapsana communis, 9: 15, 197

Larix

decidua, 7: 123

laricina, 7: 123-124; 8: 135; 9: 4, 56,
148

Lathyrus palustris, 9: 190
Lawrence Lake, 9: 3-9
Leccinum lactifluum, 7:16
Leersia

oryzoides, 8: 135
virginica, 9: 163
Leiocolea heterocolpos, 9: 94
Lemna

gibba, 7: 71

minor, 7: 68-73, 75; 8: 134; 9: 163
obscura, 7: 69, 71
perpusilla, 7: 69, 71, 73
trisulca, 7: 68, 70, 73, 75; 8: 134
Lemnaceae, of Minnesota, 7: 67-76
Lentinellus vulpinus, 9: 29
Lentinus ringens, 9: 26
Leonurus

cardiaca, 9: 198, 238
marrubiastrum, 9: 158
Leotia lubrica, An Unusual Fruiting of, 8:

158-159

Lepidium

campestre, 9: 153, 197
virginicum, 9: 153, 182, 193
Lepocinclis acuta, 9: 105
Leptobryum pyriforme, 8: 124
Leptodictyum
kochii, 8: 127
riparium, 8: 128
trichopodium, 8: 128
Leskea

gracilescens, 8: 126
obscura, 8: 126

Leucobryum glaucum, 8: 123
Levisticum officinale, 9:15
Liatris spicata, 9: 6

Lichen Complex, Cetraria ciliaris, in the
Straits Region of Michigan, The Oc¬
currence of the, 8: 67-71
Ligustrum vulgare, 9: 158, 190
Lilium michiganense, 9: 164
Linaria vulgaris, 9: 158
Lindbergia brachyptera, 8: 126
Lindbladia tubulina, 9: 21 1
Lindera benzoin, 9: 14, 149
Linum (pollen), 8: 5, 8
Liparis loeselii, 8: 134
Liriodendron tulipifera, 9: 57, 149
Listera ovata, 9: 10-11
Lithospermum arvense, 9: 190
Little Chicken Island, 9: 179
Lobelia

cardinalis, 9: 159
kalmii, 9: 196
syphilitica, 9: 159

Log Fern (Dryopteris celsa) and Its Hybrids
in Michigan-A Preliminary Report, 8:
137-145

Lolium perenne, 9: 163
Long, Stephen H., 9: 41-47
Long’s Expedition to the Source of the St.
Peter’s River in 1823, Type Specimens
of Flowering Plants Described from, 9:
4147
Lonicera

dioica, 9: 196
maackii, 9: 159
tatarica, 9: 159
villosa, 9: 15
xylosteum, 9: 159
Lophozia
incisa, 9: 94
muelleri, 8: 28
Lost Ballast Island, 9: 177
Ludwigia alternifolia, 9: 240
Luzula acuminata, 9: 161
Lychnis alba, 9: 151, 199
Ly cogala epidendrum, 9: 211
Lycopodium

complanatum var. flabelliforme, 9: 13
dendroideum, 8: 63
Lycopus

americana, 9: 192
asper, 8: 111-120
lucidus, 8: 113-1 14
uniflorus, 8: 134; 9: 199
virginicus, 9: 158, 190
Lycopus asper (Western Water Horehound)
in the Western Lake Erie and Lake St.
Clair Region, The Introduction and
Spread of, 8: 111-120
Lyngbya

aestuarii, 9: 100
birgei, 9: 100
diguetii, 9: 100
epiphytica, 9: 100

1970

THE MICHIGAN BOTANIST

261

hieronymi, 9: 100
major, 9: 100
spirulinoides, 9: 100
taylorii, 9: 100
versicolor, 9: 100
Lysimachia,
ciliata, 9: 153
nummularia, 9:153
xproducta, 9: 241
quadriflora, 9: 196
stricta var. producta, 9: 241
thyrsiflora, 8: 135

Maclura pomifera, 9: 57
Macromitrium, 7: 198-200, 202
comatum, 7: 201
Magnolia, 9: 57
Maianthemum
bifolium, 9: 48

canadense, 8: 134; 9: 48-52, 164
dilatatum, 9: 48

Maianthemum canadense in Northern Michi¬
gan, A Preliminary Report on the Vari¬
eties of, 9: 48-52
Mains, Edwin B., 8: 145
Making Rubber Duplicates of Mushrooms,

7: 184-187
Malaxis

monophyllos, 8: 134
unifolia, 9: 12
Malva neglecta, 9: 192, 238
Mammut americanum, 8: 3
Managed Tracts, 8: 31
Mannia siberica, 9: 87
Marchantia polymorpha, 9: 94
Marchantia polymorpha in Northern Michi¬
gan, 8: 146-150

Marl Lake, Post-Settlement Influences upon
a Southern Michigan, 9: 3-9
Marrubium vulgare, 9: 190
Mastodon Site, Thaller, Pollen Analysis at,

8: 3-10

Matricaria matricarioides, 9: 160
Matteuccia

pensylvanica, 9: 87
struthiopteris, 9: 148
Maximow, A. A., 9: 37
McVaugh, Rogers, Botanical Results of the
Michigan Geological Survey Under the
Direction of Douglass Houghton, 1837-
1840, 9: 213-243; Rare Old Publications
in Michigan Herbaria, 7: 3-13
Medicago

lupulina, 9: 154, 197
sativa, 9: 154, 190
Mead, Samuel B., 7: 269; 8: 182
Melampyrum

brachiatum, 9: 46
lineare, 9: 46
Melilotus

alba, 9: 154, 195
officinalis, 7: 64; 9: 154, 190
Melosira, 9: 105

Menispermum canadense, 9: 13, 150, 192
Mentha piperita, 9: 4
Menyanthes trifoliata, 8: 134
Merismopedia

convoluta, 9: 100
glauca, 9: 100
punctata, 9: 100
tenuissima, 9: 100
Michigan

Bryophytes New to, 7: 132-134; II, 8:
28-29

The Cantharellaceae of, 7: 143-183
Map of counties, 7: 79; 9: 248
Mosses New to the Flora of, 9: 139-141
A Preliminary Report on the Distribution
of Gymnosperms in, 7: 121-128
The Spring Beauties (Claytonia) in, 7:
77-93

Studies on the Hydnums of, I. Genera
Phellodon, Bankera, Hydnellum, 7:
212-264

Michigan Botanical Club

Conservation Organization of the Year
award, 7: 271

Organizational meeting, 8: 187
Southwestern Chapter, 8: 163
“Winter Wildflower” project, 7: 95
Michigan Flora Progress, 9: 247
Michigan Geological Survey, Botanical Re¬
sults of the, Under the Direction of
Douglass Houghton, 1837-1840, 9:
213-243

Michigan Natural Areas Council, 8: 32-34
Michigan Plants in Print (see New Literature
Relating to Michigan Botany)

Michigan State University, Sanford Natural
Area, 9: 116-139, 147-164
Micrasterias truncata, 9: 104
Microchaete goeppertiana, 9: 100
Microcystis

aeruginosa, 9: 99, 100
flos-aquae, 9: 99, 100
incerta, 9: 100
Microspora

elegans, 9: 101
loefgrenii, 9: 101
pachy derma, 9: 101
stagnorum, 9: 101
Mielichhoferia meilichhoferi, 7: 134
Miller, Norton G., & Howard Crum, Bryo¬
phytes New to Michigan, 7: 132-134;
Bryophytes New to Michigan. II, 8: 28-
29

Miller, Norton G., & Dale H. Vitt, Addition¬
al Bryophytes from Sinkholes in Alpena
County, Michigan, Including Orthotri-
chum pallens New to Eastern North
America, 9: 87-94

Miller, Orson K., Jr., The Genus Panellus in
North America, 9: 17-30
Milton College Herbarium, The, 7: 269-270;

(see also 8: 35-37)

Mimulus ringens, 9: 158

262

THE MICHIGAN BOTANIST

Vol. 9

Minnesota, The Duckweeds of, 7: 67-76
Mitchella repens, 8: 63, 135; 9: 159
Mitella diphylla, 9: 153
Mnium

affine, 7: 198; 8: 125
andrewsianum, 7: 93
cuspidatum, 8: 125
marginatum, 8: 125; 9: 93
medium, 7: 197, 199; 8: 125
pseudopunctatum, 7: 93
punctatum, 8: 125
spinulosum, 9: 93
stellare, 8: 125
Monarda fistulosa, 9: 190
Monotropa uniflora, 8: 134
Morchella

augusticeps, 9: 201-203
crassipes, 9: 201-202
esculenta, 9: 201-203

Morel Season in Michigan, A Report on the
1969, 9: 200-204

Morphology of the Fungal Association in
Three Species of Corallorhiza in Michi¬
gan, 9: 108-113

Mortemore, Richard A., Making Rubber
Duplicates of Mushrooms, 7: 184-187
Morus, 9: 58

alba, 9: 150, 198
rubra, 9: 150, 191

Mosquito Pollination of Habenaria obtusata
(Orchidaceae), 7: 203-212
Mosses from Southern Michigan: New Dis¬
tributional Records, 8: 120-130
Mosses New to the Flora of Michigan, 9:
139-141

Mosses of the Douglas Lake Region, Chrom¬
osome Studies of Some, 9: 67-71
Mosses Rarely Found in Michigan’s Upper
Peninsula, New Localities for, 7: 93-94
Mosses, Sex Determination in, 7: 195-203
Mucilago Crustacea, 9: 212
Muhlenbergia

frondosa, 9: 163
mexicana, 9: 188
racemosa, 8: 135
schreberi, 9: 163

Musselman, Lytton J., Additional Wisconsin
Records for the Ebony Spleenwort,
Asplenium platy neuron, 7: 268; The
Milton College Herbarium, 7: 269-270;
Asexual Reproduction in the Burning
Bush, Euonymus atropurpureus, 7: 60-
61 ; The Botanical Activities of Thomas
J. Hale, 1858-1862, 8: 181-185; Samuel
H. Watson, Pioneer Botanist of Southern
Wisconsin, 8: 35-37
Mycena

mirabilis, 9: 29
thujina, 9: 108
Myosotis

laxa, 9: 15

scorpioides, 9: 15, 157
verna, 9: 13

Myriophyllum verticillatum, 8: 133
Myurella sibirica, 8: 126
Myxomycetes from Cheboygan and Emmet
Counties, Michigan, 9: 211-212

Najas flexilis, 9: 7
Nasturtium officinale, 9: 4
Natural Area Preserves, 8: 31-32
Natural Areas in Michigan, 8: 32-34
Natural Proliferation of Floating Stems of
Scouring-Rush, Equisetum hyemale, 9:
166-174

Nature Education Feature, 7: 184-187, 265-
268; 8: 30-35; 9: 55-58
Nature Study Areas, 8: 30-31
Navicula, 9: 105
Neidium, 9: 105

Nemopanthus mucronata, 8: 132
Nepeta cataria, 9: 194, 238
Nephrocytium

agardhianum, 9: 101
limneticum, 9: 101
Nevius, Reuben, 8: 182
New Clitocybe from Michigan, A, 9: 30-33
New Literature Relating to Michigan Botany,
7: 25-30, 94, 188-189; 8: 44-46, 160-
162; 9: 59-61, 205-206, 245-246
New Localities for Mosses Rarely Found in
Michigan’s Upper Peninsula, 7: 93-94
New Location for Porella pinnata in Lower
Michigan, A, 8: 104-106
News of Botanists, 7: 271; 8: 145; 9: 63, 247
Nickell, Walter P., 9: 63
Nitzschia, 9: 105

Nomenclature, International Code of Botan¬
ical, 7: 4-5

North American Gramineae and Cyperaceae,
by Asa Gray, 7: 5-9
Nostoc comminutum, 9: 100
Notes on the Genus Suillus (Boletaceae), 7:
14-18

Nothopanus eugrammus, 9: 29
Nuphar, 9: 72, 86
luteum, 8: 134
Nuttall, Thomas, 9: 42, 213
Nymphaea, 9: 72-86, 144
odorata, 9: 72-86
tuberosa, 8: 134; 9: 72-86
Nymphaea of Michigan, Investigations in
the, 9: 72-86
Nyssa sylvatica, 8: 134

Oakland County, Michigan, White Pine at
the Edge of Its Range in, 8: 61-66
Observations on the Bolbitiaceae. IV. A New
Genus of Gastromycetoid Fungi, 7: 19-24
Occurrence of the Lichen Complex, Cetraria
ciliaris, in the Straits Region of Michigan,
The, 8: 67-71

Odd and Unusual Discomycetes from Mich¬
igan, 1.7: 58-59
Oedogonium

capitellatum, 9: 101-102

1970

THE MICHIGAN BOTANIST

263

howardii, 9: 101
poecilosporum, 9: 101
pra tense, 9: 101
spirostriatum, 9:101
Oenothera

biennis, 9: 155, 190
rhombipetala, 7: 64-65
Olney, Stephen Thayer, 8: 182
Onoclea sensibilis, 8: 135; 9: 148, 198
Onosmodium hispidum, 9: 240
Oocystis

borgei, 9: 101
elliptica, 9: 101
gigas, 9: 101
parva, 9: 101
solitaria, 9: 101
Ophiocytium

cochleare, 9: 105
parvulum, 9: 105
Orchis spectabilis, 9:13
Orobanche fasciculata, 9: 15
O’Rourke, F. L. S„ 7: 271
Orthotrichum
alpestre, 9: 141
lyellii, 7: 198-201
microcarpum, 9: 91
obtusifolium, 7: 199-201; 9: 93
ohioense, 8: 125
pallens, 9: 89-92
pumilum, 8: 126
speciosum, 7: 197; 9: 93
Orthotrichum pallens New to Eastern North
America, 9: 87-92
Oryzopsis

pungens, 9: 13
racemosa, 9: 163
Oscillatoria

agardhii, 9: 100
articulata, 9: 100
bornetii, 9: 100
chalybea, 9: 100
curviceps, 9: 100
hmnetica 9: 100
limosa, 9: 100
nigra, 9: 100
ornata, 9: 100
princeps, 9: 100
rubescens, 9: 100
subbrevis, 9: 100
tenuis, 9: 100
Osmorhiza

claytonii, 9: 157
longistylis, 9: 192
Osmunda

cinnamomea, 8: 134
regalis, 8: 134; 9: 148
Ostrya virginiana, 9: 58, 151, 195
Oxalis

europaea, 9: 197
stricta, 9: 156

Palmella mucosa, 9: 101
Palmer, Walter L., Charles Henry Swift-
1881-1966, 9: 34-40

Panax

quinquefolius, 9: 156, 239
trifolius, 9: 156
Pandorina morum, 9: 100
Panellus, The Genus, in North America, 9:

17-30

Panicum

capillare, 9: 163, 193
lanuginosum, 9: 191
perlongum, 7: 63-64
oligosanthes, 7: 63-65
xanthophysum, 7: 5-6, 9
Panus

ringens, 9: 26
salicinus, 9: 26
stipticus, 9: 24
suavissimus, 9: 29

Parietaria pensylvanica, 9: 150, 198
Park Lake, Clinton County, Michigan, algae
in, 9: 95-107
Parthenocissus
inserta, 9: 199

quinquefolia, 8: 1 36 ; 9: 155, 192
Pectodictyon cubicum, 9: 101, 104
Pediastrum

araneosum, 9:101
boryanum, 9: 101
duplex, 9: 101
integrum, 9: 101
obtusum, 9: 101
tetras, 9: 101

Pellaea glabella, 9: 197, 240
Penium margaritaceum, 9: 104
Pennfield Bog, 8: 11-27, 51-60, 131-136
Penstemon

digitalis, 9:13
hirsutus, 9: 196
Peridinium

gatunense, 9: 105
inconspicuum, 9: 105
pusillum, 9: 105
umbonatum, 9: 105
volzii, 9: 105
willei, 9: 105

Peterson, Sylvie-Ring, Biology of the Mouse-
ear Chickweed, Cerastium vulgatum, 8:
151-157

Peziza proteana, f. proteana & f. sparas-
soides, 7: 58-59
Phacus

caudatus, 9: 105
contortus, 9: 105
curvicauda, 9: 105
longicauda, 9: 105
obolus, 9: 102, 105
orbicularis, 9: 105
pseudonordstedtii, 9: 102, 105
pseudoswirenkoi, 9: 105
suecicus, 9: 105
undulatus, 9: 105
Phascum cuspidatum, 8: 124
Phellodon spp., 7: 213-220
Philonotis

capillaris, 9: 141

264

THE MICHIGAN BOTANIST

Vol. 9

fontana, 8: 125
Phleum pratense, 9: 163, 194
Phlox

divaricata, 9: 157, 196
pilosa, 7: 63-64
Phormidium tenue, 9: 100
Phryma leptostachya, 9: 158, 196
Physalis

heterophylla, 9: 194
subglabrata, 9:157
Physarum

confertum, 9: 211-212
psittacinum, 9: 212
Physcomitrella patens, 8: 28
Physcomitrium pyriforme, 8: 124
Physocarpus opulifolius, 9: 198
Phytogeographical Analysis of a Southern
Michigan Bog, A, 8: 51-60
Phytolacca americana, 9: 151, 191
Picea (pollen), 8: 5-7
Picea, 9: 56
glauca, 7: 126
mariana, 7: 126; 8: 63
Pilea pumila, 9: 151, 188
Pine, White, at the Edge of Its Range, in
Oakland County, Michigan, 8: 61-66
Pinguicula vulgaris, 9: 243
Pinnularia, 9: 105
Pinus, 9: 56

banksiana, 7: 123-125

nigra, 7: 124

resinosa, 7: 123-125

strobus, 7: 123-124; 8: 61-66, 135; 9:

112

sylvestris, 7: 124
Pisolithus tinctorius, 8:158
Pitcher, Zina, 9: 227
Plagiochila asplenioid&s, 9: 94
Plagiopus oederiana, 9: 93
Plagiothecium

denticulatum, 8: 129
laetum, 8: 129
latebricola, 7: 133
roeseanum, 8: 129; 9: 93
Planktosphaeria gelatinosa, 9: 101
Plantago

cordata, 8: 72-104; 9: 241
cornuti, 8: 85

eriopoda, 8: 73, 80, 83, 85-87
kentuckensis, 8: 73
lanceolata, 9: 158, 196
major, 8: 73-74, 100; 9: 158
media, 8: 73, 85
multiplinervia, 8: 73
pachyphylla var. montis-eekis, 8: 85
princeps, 8: 85
reniformis, 8: 85

rugelii, 8: 73-74, 85, 100; 9: 158, 198
sparsiflora, 8: 85

Plantago cordata, Systematic and Ecological
Studies on, 8: 72-104
Platanus occidentalis, 9: 57, 150, 195
Platydictya jungermannioides, 9: 93

Platygyrium repens, 8: 130
Pleurocapsa minor, 9: 100
Pleurotaenium

ehrenbergii, 9: 104
trabecula, 9: 104
Ploeodorina califomica, 9: 100
Poa

alsodes, 9: 163
annua, 9: 163

compressa, 7: 63-64; 9: 197
nemoralis, 9: 163
palustris, 9: 163
pratensis, 7: 63-64; 9: 163, 195
Podophyllum peltatum, 7: 265; 9: 150, 189
Pogonatum capillare, 7: 132
Pogonia ophioglossoides, 8: 134
Pohlia

annotina var. annotina, 8: 29
annotina var. loeskei, 8: 29
nutans, 8: 124
Poison ivy, 7: 265-266
Poison oak, 7: 265-266
Poison sumac, 7: 265-266
Pokora, Dan, Seasonal Change in a Sand
Prairie in Van Buren County, Michigan,

7: 62-66

Polanisia graveolens, 9: 199
Pollination, Mosquito, of Habenaria obtusata
(Orchidaceae), 7: 203-212
Polygonatum

canaliculatum, 9: 191
pubescens, 9: 164, 197
Polygonum

aviculare, 9: 152, 198
convolvulus, 9: 152, 191
hydropiperoides, 8: 135
lapathifolium, 9: 152, 193
pensylvanicum, 9: 188
persicaria, 9: 194
punctatum, 9: 188
sagittatum, 8: 135
scandens, 9: 197
virginianum, 9: 152
Polymnia

canadensis, 9: 160, 197, 239
uvedalia, 9: 239

Polypodium virginianum, 9: 87, 188
Polystichum

acrostichoides, 9: 148
xhagenahii, 9:12
Polytrichum

commune, 8: 130
juniperinum, 8: 130
ohioense, 8: 130
piliferum, 8: 130
Pontederia cordata, 8: 135
Population Studies in the Chemical Species
of the Cladonia chlorophaea Group, 8:
170-174
Populus, 9:58

xbarnesii, 9: 53-54
deltoides, 9: 152, 191, 272
grandidentata, 8: 189-201; 9: 152

1970

THE MICHIGAN BOTANIST

265

grandidentata x tremuloides, 8: 189-201;
9: 53-54

tremuloides, 8: 63, 136, 189-201; 9: 53-
54, 152, 191

tremuloides var. intermedia, 9:13
tremuloides var. magnifica, 9:13
Porella pinnata in Lower Michigan, A New
Location for, 8: 104-106
Porter, T. C., 8: 182
Portulaca oleracea, 9: 197
Post-Settlement Influences upon a Southern
Michigan Marl Lake, 9: 3-9
Potamogeton

gramineus, 8: 135; 9: 7
illinoensis, 8: 135; 9: 7
longiligulatus, 9: 13
natans, 8: 135
pectinatus, 9: 7
praelongus, 9: 7
Potentilla

floribunda, 9: 45
fruticosa, 9: 6, 45
norvegica, 9: 189
palustris, 8: 135
recta, 9: 154, 193
Pottia davalliana, 8: 28
Prairie, Sand, in Van Buren County, Michi¬
gan, Seasonal Change in a, 7: 62-66
Preissia quadrata, 8: 147
Preliminary Report on the Distribution of
Gymnosperms in Michigan, A, 7: 121-
128

Preliminary Report on the Varieties of
Maianthemum canadense in Northern
Michigan, A, 9: 48-52
Preliminary Study of Clavariadelphus in
North America, A, 7: 35-57
Prenanthes
alba, 9: 193
altissima, 9: 160
Primula, 7: 266

mistassinica, 7: 134

Pringle, James S., The Status and Distribu¬
tion of Gentiana linearis and G. rubri-
caulis in the Upper Great Lakes Region,
7: 99-112

Proliferation, Natural, of Floating Stems of
Scouring-Rush, Equisetum hyemale, 9:
166-174

Protococcus viridis, 9: 101
Prunella vulgaris, 9: 158, 196
Prunus, 9: 57-58
incana, 9: 45
persica, 9: 189
pumila, 9: 45
serotina, 9: 154, 192
virginiana, 8: 135; 9: 154, 195
Pseudocraterellus calyculus, 7: 149-150
Pseudotsuga menziesii, 9: 56
Pseudulvella americana, 9: 101
Ptelea trifoliata, 9: 156, 195
Pteridium aquilinum, 7: 64
Pterospora andromedea, 9: 13

Publications of Interest, 7: 13, 31, 76, 189,
191; 8: 66, 107; 9: 16,63, 107, 143,

207, 247

Pylaisiella selwynii, 9: 68-70
Pyrola rotundifolia, 9:15
Pyrus, 9: 58
Pyrus communis, 9: 57

Quadrigula lacustris, 9: 101
Quercus (pollen), 8: 5-7
Quercus, 9:58

acuminata, 9: 188
alba, 9: 151, 199
bicolor, 9: 151
macrocarpa, 9: 151
muehlenbergii, 9: 151, 191
rubra, 9: 112, 151, 191
velutina, 8: 133; 9: 151

Radiocarbon dates, Thaller mastodon site,

8: 4,9

Radiofilum flavescens, 9: 101
Ranunculus, 7: 265
abortivus, 9: 149, 191
bulbosus, 9: 14
filiformis var. hispidus, 9: 45
flabellaris, 9: 13
pensylvanicus, 9: 149
recurvatus, 9: 149
reptans, 9: 45
sceleratus, 8: 135; 9: 198
septentrionalis, 9: 149
trichophyllus, 9: 14

Rare Old Publications in Michigan Herbaria,
7: 3-13

Ratibida pinnata, 9: 240
Rayssiella hemisphaerica, 9: 101
Redfearn, Paul L., Jr., & Richard L. Hal¬
bert, Mosses from Southern Michigan:
New Distributional Records, 8: 120-130
Report on the 1969 Morel Season in Michi¬
gan, A, 9: 200-204
Reticularia lycoperdon, 9: 211
Review of the Status of Some Green Algae
in the Genus Coelastrum, 7: 129-131
Reviews:

Disease in Forest Plantations: Thief of
Time, 7: 30

A Field Guide to Wildflowers, 7: 190
Geology of Michigan, 9: 206
How to Know the Aquatic Plants, 9: 142
How to Know the Lichens, 9:71
How to Know Pollen and Spores, 9: 165
Living with Your Land, 8: 10
Natural Areas in Indiana and their Preser¬
vation, 8: 150

Plants of the Chicago Region, 9: 165
Rhamnus

cathartica, 9: 155
frangula, 9: 155
Rhizoclonium

crassipellitum, 9: 101
hieroglyphicum, 9: 101

266

THE MICHIGAN BOTANIST

Vol. 9

Rhodobryum roseum, 8: 124; 9: 93
Rhopalodia, 9: 105
Rhus, 9: 58

aromatica, 9: 190
glabra, 9: 199
radicans, 9: 194
typhina, 9: 156, 192
Rhynchospora alba, 8: 133
Rhynchostegium serrulatum, 8: 129
Rhytidiadelphus triquetrus, 8: 130
Ribes

americanum, 8: 136
cynosbati, 9: 14, 153, 192
hirtellum, 8: 136
odoratum, 9:14
sativum, 9: 153
triste, 9:14

Rich, Peter H., Post-Settlement Influences
upon a Southern Michigan Marl Lake, 9:
3-9

Richards, Mrs. R. R., 9: 223
Robinia pseudoacacia, 9: 57, 154, 195
Rorippa islandica, 9: 189, 193
Rosa

acicularis, 9: 45
Carolina, 7: 63-64; 9: 195
multiflora, 9: 199
palustris, 8: 135
sayi, 9: 45

Rubber Duplicates of Mushrooms, Making,

7: 184-187
Rubus

allegheniensis, 9: 154, 189
hispidus, 8: 135
occidentals, 9: 154, 197
pensilvanicus, 9: 189
pubescens, 8: 135
strigosus, 9: 154
Rudbeckia laciniata, 9: 160
Ruellia

humilis, 8: 183
ciliosa, 8: 183
Rumex

acetosella, 7: 63-64
altissimus, 9: 198
crispus, 9: 152, 193
orbiculatus, 8: 135; 9: 188
obtusifolius, 9: 188
verticillatus, 9: 152
Russell, John S., A New Location for
Porella pinnata in Lower Michigan,

8: 104-106

Russula nigricans, 8: 42

Sabatia angularis, 9: 239
Sager, Abram, 9: 216-217, 220-221, 223-
224, 231, 236, 241
Sagittaria latifolia, 8: 131; 9: 161
St. Peter’s River in 1823, Type Specimens of
Flowering Plants Described from Long’s
Expedition to the Source of the, 9: 41-
47

Salix, 9: 58
alba, 9: 191

amygdaloides, 9: 152, 182, 191
bebbiana, 8: 136
bebbiana, teratology, 7: 115-118
Candida, 8: 136
cordata, 9: 188
discolor, 9: 153, 191
fragilis, 9: 153
glaucophylloides, 9: 188
interior, 9: 153, 193
pedicellaris, 8: 136
petiolaris, 8: 136
rigida, 9: 153
serissima, 8: 136
Sambucus

canadensis, 8: 132; 9: 159, 193
pubens, 9: 159
Samolus parviflorus, 9: 153
Samuel H. Watson, Pioneer Botanist of
Southern Wisconsin, 8: 35-37
Sand Prairie in Van Buren County, Michi¬
gan, Seasonal Change in a, 7: 62-66
Sanford Natural Area, A Botanical Inventory
of, 9: 116-139, 147-164
Sanguinaria canadensis, 9: 150
Sanicula

canadensis, 9: 196
gregaria, 9: 157
trifoliata, 9: 157

Saponaria officinalis, 9: 198, 238

Sarcodon, 7: 219

Sarracenia purpurea, 8: 136

Sartwell, H. P., 7: 5-9

Sassafras albidum, 9: 58, 149

Satureja vulgaris, 9: 193

Saules de la Suisse, by N.-C. Seringe, 7: 9-11

Saururus cernuus, 9: 149

Saxifraga aizodn, 9: 13

Say, Thomas, 9: 42

Scapania saxicola, 7: 134; 9: 94

Scenedesmus

acutus, 9: 101-102
arcuatus, 9: 101-102
bicaudatus, 9: 101-102
brevispinus, 9: 101
ecomis, 9: 101
longus, 9: 101
ovaltemus, 9: 101
quadricauda, 9: 101
spinosus, 9: 101
tibiscensis, 9: 101-102
Scenic Sites, 8: 31
Scheuchzeria palustris, 9: 239
Schistostega pennata, 7: 93-94
Schizaea pusilla, 9:12
Schizothrix tinctoria, 9: 100
Schoolcraft Expeditions, 9: 214
Schweinitz, Lewis David von, 7: 269; 9: 41-
47

Schwetschkeopsis
denticulata, 8: 126
fabronia, 7: 200; 8: 28

1970

THE MICHIGAN BOTANIST

267

Scirpus

acutus, 8: 133
americanus, 9: 193, 241
cyperinus, 8: 133
fluviatilis, 9: 188
olneyi, 9: 241
subterminalis, 9: 7
Scleroderma geaster, 8: 158
Scopelophila ligulata, 7:133
Scouring-Rush, Equisetum hyemale, Natural
Proliferation of Floating Stems of, 9:
166-174

Scrophularia marilandica, 9: 158, 190
Scutellaria

galericulata, 8: 134
lateriflora, 8: 134; 9: 158
Scytonema bohneri, 9: 100, 102
Seasonal Change in a Sand Prairie in Van
Buren County, Michigan, 7: 62-66
Sedum

acre, 9: 195
sarmentosum, 9: 199
ternatum, 9: 189

Seedling Population of Aspens in South¬
eastern Michigan, A, 8: 189-202
Seligeria

calcarea, 9: 87
campylopoda, 9: 92
doniana, 9: 92
Sematophyllum

carolinianum, 8: 129
demissum, 8: 28
Senecio

congestus, 8: 182
palustris, 8: 182

Seringe, N.-C, Saules de la Suisse, 7: 9-11

Setaria glauca, 9: 163, 197

Sex Determination in Mosses, 7: 195-203

Short, Charles W., 7: 269; 8: 182

Sicyos angulatus, 9: 182, 193

Silene antirrhina, 9: 189

Silphium

perfoliatum, 9: 240
terebinthinaceum, 9: 240
Sinkholes in Alpena County, Michigan,
Additional Bryophytes from, 9: 87-94
Sirogonium sticticum, 9:101
Sistotrema, 7: 213
Sisymbrium

altissimum, 9: 199
officinale, 9: 153, 193
Sium suave, 8: 132; 9: 157
Smilacina

racemosa, 9: 164, 193
stellata, 9: 164, 193
Smilax

ecirrata, 9: 164, 188
herbacea, 9: 188
hispida, 9: 188
illinoensis, 9: 164
rotundifolia, 9: 13, 239
tamnoides, 9: 164
Smith, Alexander H., 9: 63

Smith, Alexander H., The Cantharellaceae
of Michigan, 7: 143-183; Dr. Bessie
Bernice Kanouse-1889-1969, 8: 186-188
Smith, Alexander H., & Joseph F. Ammirati,
Jr., Studies in the Genus Cortinarius, I:
Section Dermocybe, Cortinarius aurei-
folius Complex, 8: 175-180
Smith, Alexander H., & Howard E. Bigelow,
A New Clitocybe from Michigan, 9:30-
33

Smith, Alexander H., & Harry D. Thiers,
Notes on the Genus Suillus (Boletaceae),
7: 14-18

Smith, Nancy Jane, Odd and Unusual Dis-
comycetes from Michigan, I, 7: 58-59
Snider, Jerry A., Chromosome Studies of
Some Mosses of the Douglas Lake Re¬
gion, 9: 67-71
Solanum

dulcamara, 8: 136; 9: 157, 198
nigrum, 9: 157, 196
Solenostoma sphaerocarpum, 7: 134
Soli dago

altissima, 9: 13
caesia, 9: 160
canadensis, 9: 161, 191
flexicaulis, 9: 161, 196
gigantea, 9:161
graminifolia, 9: 161, 197
hispida, 9: 16, 191
houghtonii, 9: 240-241
speciosa, 7 : 64
Sonchus

asper, 9: 194
arvensis, 9: 161, 199
oleraceus, 9: 161, 194
Sorastrum spinulosum, 9: 101
Sorbaria sorbifolia, 9: 13
Sorghastrum nutans, 9: 240
Sphagnum

annulatum var. porosum, 8: 28
auriculatum, 8: 28

capillaceum, 8: 122, 168-169; 9: 114,

140

centrale, 8: 168-170; 9: 114
contortum, 8: 28
cuspidatum, 8: 122; 9: 115
fimbriatum, 8: 122, 168-169; 9: 115
fuscum, 9:115
girgensohnii, 8: 122; 9: 115
magellanicum, 8: 168-169; 9: 115
majus, 9:115

papillosum, 8: 122, 168-170
platyphyllum, 8: 28
pulchrum, 8: 28
recurvum, 8: 168-169; 9: 115
var. tenue, 8: 168-170
riparium, 8: 28, 168-170; 9: 115
russowii, 9:115
squarrosum, 8: 112; 9: 115
subfulvum, 8: 28
subnitens, 8: 28
subsecundum, 8: 122, 168-170

268

THE MICHIGAN BOTANIST

Vol. 9

tenerum, 9: 139
teres, 8: 168-170; 9: 1 15
warnstorfii, 9: 115
wulfianum, 9:115

Sphagnum in Charlevoix County, Michigan,
9: 114-115

Sphagnum Mosses, Distribution of, Studies
of the Byron Bog in Southwestern On¬
tario, XL, 8: 167-170
Sparganium chlorocarpum, 8: 136
minimum, 8: 136
Sparrow, Frederick K., 8: 145
Species of Vascular Plants of Pennfield
Bog, Calhoun County, Michigan, 8:
131-136

Specularia perfoliata, 7: 64-65
Sphaerocarpus donnellii, 7: 202
Sphaerocystis schroeteri, 9: 101
Sphenopholis intermedia, 9: 163
obtusata, 9: 195
Spiraea alba, 8: 135
tomentosa, 8: 135
Spiranthes cernua, 9: 12
gracilis, 9: 12

Spirodela polyrhiza, 7: 68-70, 75
punctata, 7: 69
Spirogyra crassa, 9: 104
longata, 9: 104
Spirulina princeps, 9: 100
subssalsa, 9: 100
Sprague, C. J., 8: 182
Spring Beauties (Claytonia) in Michigan,
The, 7: 77-93
Stachys ambigua, 9: 46
tenuifolia, 9: 158
velutina, 9: 46

Staphylea trifolia, 9: 155, 196
Starve Island, 9: 178-179
Status and Distribution of Gentiana linearis
and G. rubricaulis in the Upper Great
Lakes Region, The, 7: 99-112
Staurastrum

cornutum, 9: 104
cosmarioides, 9: 102, 104
crenulatum, 9: 104
cyclacanthum, 9: 102, 104
disputatum, 9: 102, 104
furcigerum, 9: 104
gracile, 9: 104
hexacerum, 9: 104
johnsonii, 9: 104
longiradiatum, 9: 104
manfeldtii, 9: 104
punctulatum, 9: 102, 104
sebaldi, 9: 104
vestitum, 9: 104
Stauroneis, 9: 105
Steccherinum, 7: 213
Stellaria media, 9: 151, 191
Stemonitis axifera, 9: 211-212
Stereum calyculus, 7: 149
Stigeoclonium lubricum, 9: 101
subsecundum, 9: 101

tenue, 9: 101

Stipa avenacea, 9: 239-240
Stone, Franz Theodore, Laboratory, 9: 180,
185

Stoutamire, Warren P., Mosquito Pollination
of Habenaria obtusata (Orchidaceae), 7:
203-212

Strophostyles helvola, 9: 194
Stuckey, Ronald L., Distributional History
of Butomus umbellatus (Flowering-rush)
in the Western Lake Erie and Lake St.
Clair Region, 7: 134-142; The Introduc¬
tion and Spread of Lycopus asper (West¬
ern Water Horehound) in the Western
Lake Erie and Lake St. Clair Region, 8:
111-120; Type Specimens of Flowering
Plants Described from Long’s Expedition
to the Source of the St. Peter’s River in
1823,9:4147

Stuckey, Ronald L., & Thomas Duncan,
Changes in the Vascular Flora of Seven
Small Islands in Western Lake Erie, 9:
175-200

Studies of the Byron Bog in Southwestern
Ontario. XL. Distribution of Sphagnum
Mosses, 8: 167-170

Studies in the Genus Cortinarius, I: Section
Dermocybe, Cortinarius aureifolius Com¬
plex, 8: 175-180

Studies on the Hydnums of Michigan. I.
Genera Phellodon, Bankera, Hydnellum,
7: 212-264
Suillus

albidipes, 7: 14-15
brevipes, 8: 61
granulatus, 7: 15, 18; 8: 61
lactifluus, 7: 16, 18
unicolor, 7: 15-17
Sullivan tia sullivantii, 8: 51
Surirella, 9: 105

Swertia caroliniensis, 9: 239-240
Symplocarpus foetidus, 9: 163
Synedra, 9: 105
Synura uvella, 9: 105
Syringa vulgaris, 9: 196
Systematic and Ecological Studies on
Plantago cordata, 8: 72-104

Tabellaria, 9: 105
Taenidia integerrima, 9: 157
Taraxacum

erythrospermum, 9: 190
officinale, 9: 161, 193
Taxiphyllum howellianum, 9: 141
Taxus canadensis, 7: 122, 124; 9: 194
Taylor, Sylvia, White Pine at the Edge of
Its Range, in Oakland County, Michigan,
8: 61-66

Teratological Stamens and Carpels of a Wil¬
low from Northern Michigan, 7: 113-120
Tessene, Melvem F., Systematic and Ecolog¬
ical Studies on Plantago cordata, 8: 72-
104

1970

THE MICHIGAN BOTANIST

269

Tetradesmus wisconsinensis, 9: 101
Tetraedron

minimum, 9: 101
muticum, 9: 101

Teucrium canadense, 9: 158, 196
Tetraphis pellucida, 8: 122
Tetraspora gelatinosa, 9: 101
Teucrium occidentale, 9: 190
Thalictrum

dasycarpum, 9: 149
dioicum, 9: 150, 189
Thaller Mastodon Site, Gratiot County,
Michigan, Pollen Analysis at the, 8:

3-10

Thelia asprella, 8: 126
Thelypteris

hexagonoptera, 9: 148
palustris, 8: 135; 9: 148
Thlaspi arvense, 9: 195
Thieret, John W., & Howard L. Clark, The
Duckweeds of Minnesota, 7: 67-76
Thiers, Harry D., & Alexander H. Smith,
Notes on the Genus Suillus (Boletaceae),
7: 14-18

Thomson, George Wm., Vascular Plants of
the Bruce Peninsula: A Review, with
Comments and Additions, 9: 9-16
Thuidium

delicatulum, 8: 126
minutulum, 8: 126
recognitum, 9: 93

Thuja occidentalis, 7: 127; 8: 63; 9: 6, 56,
108, 148

Thurber, George, 9: 227
Thymus serpyllum, 9: 15
Tilia (pollen), 8: 8-9
Tilia americana, 9: 57, 152, 192
Timmia

austriaca, 9: 92
megapolitana, 8: 125
Tolypothrix
lanata, 9: 100
tenuis, 9: 99-100
Torilis japonica, 9: 157
Torrey, John, 7: 269; 9: 225, 227, 234-235,
237-238, 240

Tortula mucronifolia, 8: 124
Toxicodendron

quercifolium 7: 265
radicans, 7: 265; 9: 156
vernicifera, 7: 265
vernix, 7: 265; 8: 132; 9: 156
Trachelomonas
hispida, 9: 105
volvocina, 9: 105
Tradescantia ohiensis, 7: 63-64
Transplant studies in Nymphaea, 9: 80-83
Tragopogon
dubius, 9: 16
major, 9: 16
pratensis, 9: 16
porrifolius, 9: 13

Trees Common in Michigan, Winter Field
Key to Some, 9: 55-58
Triadenum virginicum, 8: 133
Trichodesmium lacustre, 9: 100
Trientalis borealis, 8: 135
Trifolium

hybridum, 9: 154
pratense, 9: 195, 238
procumbens, 9: 14, 199
repens, 9: 194, 238
Triglochin maritima, 8: 133
Trillium

cernuum, 9: 13
flexipes, 9: 195
grandiflorum, 9: 164
Triphora trianthophora, 9: 239
Triticum pauciflorum, 9: 44
Tritomaria

exsectiformis, 9: 93-94
scitula, 9: 87

Trochiscia reticularis, 9: 101
Tsuga (pollen), 8: 5-7,9
Tsuga canadensis, 7: 125-126; 9: 56, 148
Tubifera ferruginosa, 9: 211
Tukey, Harold B., Sr., 7: 271
Tussilago farfara, 9: 13
Type Specimens of Flowering Plants De¬
scribed from Long’s Expedition to the
Source of the St. Peter’s River in
1823, 9: 41-47
Typha latifolia, 8: 136
Typhulopsis, 7: 35

Ulbrich, A. P., Contact Dermatitis Caused by
Plants, 7: 265-268
Ulmus, 9: 58

americana, 9: 150, 191
rubra, 8: 136; 9: 150, 191
thomasii, 9: 150, 188
Ulothrix tenerrima, 9: 101
University of Chicago, 9: 34-38
University of Michigan Herbarium, 8: 187;

9: 216-217, 223, 227, 234, 240-241
Unusual Fruiting of Asterophora lyco-
perdoides, An, 8: 42-43
Unusual Fruiting of Leotia lubrica, An, 8:
158-159

Uroglenopsis americana, 9: 105
Uronema elongatum, 9:101
Urtica

dioica, 8: 136; 9: 191
gracilis, 9: 151
Utricularia
gibba, 8: 134
intermedia, 8: 134
purpurea, 8: 134
vulgaris, 8: 134

Uvularia grandiflora, 9: 164, 195

270

THE MICHIGAN BOTANIST

Vol. 9

Vaccinium

corymbosum, 8: 133; 9: 4
macrocarpon, 8: 133
oxycoccos, 9: 4

Vascular Plants of the Bruce Peninsula: A
Review, with Comments and Additions,
9: 9-16

Vasey, George, 7: 269; 8: 182
Verbascum thapsus, 9: 158, 193
Verbena hastata, 9: 190
urticifolia, 9: 158
Verbesina alternifolia, 9: 239
Vernonia corymbosa, 9: 46
Veronica arvensis, 9: 193
longifolia, 9: 13
officinalis, 9: 13
peregrina, 9: 15, 199
Veronicastrum virginicum, 9: 158
Verpa bohemica, 9: 201-204
Viburnum acerifolium, 9: 159
cassinoides, 9: 15
lantana, 9: 159
lentago, 8: 132; 9: 159
recognitum, 9: 159
trilobum, 9: 159
Vicia americana, 9: 46, 190
caroliniana, 9: 199
tridentata, 9: 43, 46
Vinca minor, 9: 15, 196
Viola affinis, 8: 136
canadensis, 9: 152
cucullata, 9: 190
nephrophylla, 8: 136; 9: 196
pubescens, 9: 152, 192
renifolia, 9:15
rostrata, 9: 15, 152
sororia, 9: 152
striata, 9: 152
Vitis riparia, 9: 155, 196
vulpina, 8: 136

Vitt, Dale H., Sex Determination in Mosses,
7: 195-203

Vitt, Dale H., & Norton G. Miller, Addi¬
tional Bryophytes from Sinkholes in
Alpena County, Michigan, Including
Orthotrichum pallens New to Eastern
North America, 9: 87-94
Volvox aureus, 9: 101
tertius, 9: 101

Voss, Edward G., Charles H. Swift’s Botani¬
cal Work in Michigan, 9: 38-40; A Pre¬
liminary Report on the Distribution of
Gymnosperms in Michigan, 7: 121-128;
The Spring Beauties (Claytonia) in Mich¬
igan, 7: 77-93

Wagner, Florence S. (see W. H. Wagner, Jr.,
et al.)

Wagner, W. H., Jr., The Barnes Hybrid
Aspen, Populus xbarnesii, hybr. nov.-A
Nomenclatural Case in Point, 9: 53-54;
Teratological Stamens and Carpels of a

Willow from Northern Michigan, 7: 113-
120

Wagner, Warren H., & William E. Hammitt,
Natural Proliferation of Floating Stems
of Scouring-Rush, Equisetum hyemale,

9: 166-174

Wagner, W. H., Jr., Florence S. Wagner, &

D. J. Hagenah, The Log Fern (Dryopteris
celsa) and Its Hybrids in Michigan-A Pre¬
liminary Report, 8: 137-145
Waldsteinia fragarioides, 9: 239
Waterlilies (Nymphaea) of Michigan, Investi¬
gations in the White, 9: 72-86
Watling, Roy, Observations on the Bolbitia-
ceae. IV. A New Genus of Gastromy-
cetoid Fungi, 7: 19-24

Watson, Samuel H., 7: 269-270; 8: 35-37, 182
Wayne State University Herbarium, 7: 5-9
Weissia controversa, 8: 124
Weller, Stephen G., A Preliminary Report
on the Varieties of Maianthemum cana-
dense in Northern Michigan, 9: 48-52
Wells, James R., Winter Field Key to Some
Trees Common in Michigan, 9: 55-58
Wells, Virginia L., & Phyllis E. Kempton, A
Preliminary Study of Clavariadelphus in
North America, 7: 35-37
Wetherbee, Richard, Population Studies in
the Chemical Species of the Cladonia
chlorophaea Group, 8: 170-174
White Pine at the Edge of Its Range, in Oak¬
land County, Michigan, 8: 61-66
Williams, Gary R., Investigation in the
White Waterlilies (Nymphaea) of Michi¬
gan, 9: 72-86

Willow, Teratological Stamens and Carpels,

7: 113-120

Winter Field Key to Some Trees Common in
Michigan, 9: 55-58
Winter Wildflowers, 7: 95
Wisconsin, Additional Records for the Ebony
Spleen wort, Asplenium platyneuron, 7:
268

Wisconsin, Samuel H. Watson, Pioneer Bot¬
anist of Southern, 8: 35-37
Wolffia

columbiana, 7: 69, 72-75
punctata, 7: 69, 72-75
Woodsia cathcartiana, 9: 12
Woodwardia virginica, 8: 135
Wright, John, 9: 213-214, 216-217, 225-231,
237-241

Wulfenia bullii, 9: 240

Xanthidium antilopaeum, 9: 104
Xanthorhoe munitata, 7: 211

Zales, William M.,& Howard Crum, Sphagnum
inCharlevoixCounty, Michigan, 9: 114-115
Zanthoxylum americanum, 9: 156
Zoochlorella parasitica, 9:101
Zygnema sterile, 9: 104
Zygodon viridissimus, 7: 133

INDEX NOTICE

To Librarians, Subscribers, and Binders

In this issue is a single cumulative index to Volumes 7, 8, and 9 of THE
MICHIGAN BOTANIST— our third triennial index. These three volumes may thus be
bound together as a single unit with a single index. Note that all covers are included in
the pagination and should be bound in. There are no special title pages.

Due to the increasing size of volumes, this is doubtless the last triennial index and
future ones will be prepared annually or biennially.

Editorial Notes

We are deeply indebted to Miss Mary E. Cooley for the considerable labor of
alphabetizing and filing the entires for the index in this issue.

As noted earlier (Vol. 8, p. 47) the only reason for copyrighting an occasional
issue is to protect ourselves against unauthorized reproduction of complete sets (e.g., by
microfilm). No limitation is intended on normal quotation and use or on reprinting by
authors of their own material, so long as the source is credited.

The May number (Vol. 9, No. 3) was mailed May 9, 1970.

BACK ISSUES NEEDED

Our supply of complete sets of back issues is dwindling and there is still a demand
for them from new institutional (and other) subscribers. The two numbers of Vol. 1
(1962) are especially short. The following numbers would be particularly welcome if
subscribers have copies for which they have no further use; send them to the business
and circulation manager (postage will be refunded):

Vol. 1, No. 1 & No. 2 Vol. 5, No. 1

Vol. 2, No. 1 Vol. 5, No. 3 & 3A

Vol. 4, No. 3 Vol. 9, No. 3

The above numbers are no longer available for sale as single copies, as the entire stock is
reserved to make up complete sets.

THANKS TO REVIEWERS

One of the chief functions of the members of the Editorial Board is to review
manuscripts submitted for publication, considering botanical content, clarity of presenta¬
tion, and suitability for this journal. All manuscripts are reviewed by at least one person.
Reviewers are by tradition anonymous, but the editor takes this opportunity to express
his appreciation not only to members of the Editorial Board (names marked with an
asterisk below) but also to all others in the following list, who have served as advisory
reviewers for one or more articles submitted for Volumes 7-9:

William H. Anderson
Virginia L. Bailey*
Burton V. Barnes*
John H. Beaman
W. S. Benninghoff
Richard Brewer*
Garrett E. Crow
Howard A. Crum
Francis C. Evans
William T. Gillis
Dale J. Hagenah
William E. Hammitt
Richard C. Harris
Claude W. Hibbard
Hugh H. litis

Henry Imshaug
Peter B. Kaufman
Gordon McBride
E. L. McWilliams
Rogers McVaugh*
Martin A. Piehl

E. D. Rudolph
Alexander H. Smith*
Helen V. Smith

F. K. Sparrow, Jr.
Eugene F. Stoermer
Ronald L. Stuckey
Floyd Swink

W. H. Wagner, Jr.*
Grady L. Webster

CONTENTS

Myxomycetes from Cheboygan and Emmet
Counties, Michigan

Laura S. Bradford . 211

Botanical Results of the Michigan Geological

Survey Under the Direction of Douglass
Houghton, 1837-1840

Rogers McVaugh . 213

Grimmia agassizii, a Moss New to Michigan

F. J. Hermann . 244

Michigan Plants in Print . 245

News of Botanists . 247

Publications of Interest .... .\ . . 247

„ **■

Index to Volumes 7-9 . . . . 249

* " j ■

••• v

Editorial Notes . 271

P I

(On the cover: Cottonwoods, Populus deltoides,
on the Sleeping Bear Dunes.
Photograph by Rudolf Schmid, October 5, 1969.)

New York Botanical Garden Library

3 5

85 00289 5793

N. MANCHESTER,
INDIANA