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THE OHIO STATE UNIVERSITY BULLETIN

Volume XIX Number 28

OHIO BIOLOGICAL SURVEY

BULLETIN 5

VOLUME II, No. I

THE ASCOMYCETES OF OHIO, 1

Preliminary Consideration of Classification
BRUCE FINK

THE ASCOMYCETES OF OHIO, II

I

The CoUemaceae

BRUCE FINK AND C. AUDREY RICHARDS

June. 1915

PUBLISHED BY THE UNIVERSITY AT COLUMBUS, OHIO
Entered as second-class matter November 17. 1905. at the postoffice at
Columbus. Ohio, under Congress. July 16. 1894

OHIO BIOLOGICAL SURVEY

Herbert Osborn, Director

OHIO STATE UNIVERSITY IN CO-OPERATION WITH
OTHER OHO COLLEGES AND UNIVERSITIES

Administrative Board consisting of Representatives from co-operat-

ing institutions.

E. L. Fullmer,

Baldwin Wallace University,

Berea, O.

E. R. Gregory,

University of Akron,

Akron, O.

M. E. Stickney,

Denison University,

Granville, O.

L. B. Walton,

Kenyon College,

Gambier, O.

Miss M. Getman,

Lake Erie College,

Painesville, O.

Bruce Fink,

Miami University,

Oxford, O.

F. O. Grover,

Oberlin College,

Oberlin, O.

E. L. Rice,

Ohio Wesleyan University,

Delaware, O.

H. M. Benedict,

University of Cincinnati,

Cincinnati, O.

F. H. Herrick,

Western Reserve University,

Cleveland, O.

C. G. Shatzer,

Wittenberg College,

Springfield, O.

ANNOUNCEMENT

The Bulletins of the Ohio Biological Survey will be issued as work
on any special subject is completed, and will form volumes of about
500 pages each.

They will be sent to co-operating institutions and individuals,
libraries and colleges in Ohio and to such surveys, societies and other
organizations as may offer suitable exchange material.

Additional copies of each Bulletin and of completed volumes will be
sold at such price as may cover the cost of publication.

Special rates on quantities to schools for classes.

Subscription for entire volumes . $2.00

Price of this number . 50

All orders should be accompanied by remittance made payable to
Ohio Biological Survey and sent to the Director.

Correspondence concerning the Survey, applications for exchanges
and purchase of copies of Bulletins should be addressed to the Director —
Professor Herbert Osborn, Columbus, Ohio.

tIhiKARY

‘/•^rvErsiTv Of Illinois

urbana

THE UBRARY OF THE

Volume II, No. 1 SEP 3 1943

UNIVERSflY OF ILLINOIS

Ohio Biol ogical Survey

THE ASCOMYCETES OF OHIO, I

Preliminary Consideration of Classification
BRUCE FINK

THE ASCOMYCETES OF OHIO, II

The Collemaceae

BRUCE FINK AND C. AUDREY RICHARDS

Published by

THE OHIO STATE UNIVERSITY

Columbus,

1915

Bulletin 5

PRINTED BY

THE STONEMAN PRESS

Columbus, Ohio

^'cO

I. ■

THE ASCOMYCETES OF OHIO.— I
Preliminary Consideration of Classification*

BRUCE fink

All classifications of ascomycetes hitherto followed have been highly
artificial, nor are the data for a satisfactory disposition at hand.
Hitherto, these fungi, except for lists from two states and certain works
mentioned in this paper, have been placed in two groups. In the work to
follow, all the ascomycetes of Ohio will be treated as a single class of
fungi, and some defense of this method is in order.

The reasons for treating ascomycetous lichens and other ascomycetes
together involve mainly the following three considerations : first, the
nature of the lichen in general ; second, a review of the similarity of
lichens and other ascomycetes ; third, a discussion of the origin of all
ascomycetes, with a view to showing relationships within the group.
These three matters will be treated in order and as briefly as is consistent
with clearness. Frequent reference will be made to other papers where
more lengthy discussions of certain phases of the problem may be found.

Until the time of Schwendener (57), it was generally believed that
the chlorophyllous cells found in lichens were the chloioplasts of these
plants. Bornet (12) grew lichens from spores in cultures and saw them
attack algae. Bonnier (11), by similar methods, obtained mature fruited
plants of Xanthoria parietina and Rinodina sophodes. Moller (47) fol¬
lowed by growing Calicium parietinum in cultures without algae and
obtained plants that produced spermagonia and probably archicarps as
well. Fink (31) has reported on the frequent growing of various algal
hosts in the same habitats with lichens.

Had it not been for clinging to erroneous tradition, the researches
referred to above and others like them would have convinced all botanists
that the lichen is a fungus living in some peculiar relation with an alga.
Unfortunately, Reinke (53), Schneider (55), and others have aided in
retarding progress, and a considerable number of botanists still cling to
some modification of the early view. Peirce (52), Elenkin (25), and
Danilov (20), with their researches which greatly strengthen belief in the
parasitism or the saprophytism of lichens on algae, have succeeded in
turning the attention of botanists toward more plausible conclusions.

We need only to advance definitely to the widely but rather hazily
understood idea that the lichen is a fungus, the alga being its host and

* Contributions from the Botanical Laboratory of Miami University — XI.

3

4

Ohio I Biological Survey

therefore forming no part of it. Some of the arguments for this position
may lie reviewed liriefiy. The algal host is in a disadvantageous position
with respect to light and food materials. The cells of the host become
large and may divide rajiidly because of undue stimulation, showing a
hypertrophied and pathologic condition, resulting in marked vegetative
activity, which balances the rai)id killing of the algal-host individuals.
The bright-green, normally-shaped, algal individuals seen in lichens are
usually those that are not yet parasitized, or those that have been recently
parasitized. The relationship between the lichen and the alga is peculiar
on account of the sim])le structure of the latter ; but the lichen seems to
be a parasite or a saprophyte, or perhaps partly a parasite and partly a
saprophyte, on the alga. The haustoria of lichens are similar to those
of other fungi and in all probability function in like fashion. For an
extended discussion, see the paper hy Fink (31).

One cannot deal with colonies in studies of ontogeny, phylogeny,
mori)hology, or taxonomy, nor does the use of Collcma piilposurn, Plivscia
stcllaris, or other binomial by one group of botanists to designate a plant
and by another group to designate a compound colony conduce to clear¬
ness. In order that one may be clear in his thinking regarding lichens,
he must abandon the historical idea that the chlorophyllous cells found
in these plants, or in which the lichens live in a few instances, are part
of the lichen, or he must hold consistently to the view that the lichen is
not a plant, but a compound colony. Critical examination of statements
about lichens by a large number of botanists shows that it is not possible
to regard the lichen in one of these ways for one purpose and in another
manner for another purpose, or even to consider the compound colonies
for one purpose and the fungi of the colonies for another purpose, with¬
out committing some of the inconsistencies so commonly found in state¬
ments about lichens (see discussion in Fink 31). The conclusion that
the lichen is a fungus is the only one that can be followed equally well
by the physiologist, the ecologist, the morphologist, the cytologist, and the
systematist. In a previous paper (Fink 31), the typical lichen has been
defined as a fungus which lives in parasitic relationship with an alga
during all or part of its life, and also sustains a relationship with an
external substratum, organic or inorganic. The lichens will be dealt
with according to this definition in the treatment of the ascomycetes
throughout this series of papers.

Idchens possess few distinct characters other than physiological, and
these have no weight in the classification of these plants. The dual
hypothesis can scarcely be considered seriously in view of recent studies

Classification of Ohio Ascomycetes 5

(Fink 31). With this hypothesis falls the argument that lichens con¬
stitute a special biological group, nor can classihcation consistently con¬
sider such groups in any event. Some botanists have supposed that we
must know the relation of every lichen or every group of lichens to a
non-lichen ancestor with some degree of certainty before we attempt
to distribute lichens, but this position is manifestly erroneous. Phyl-
ogeny is a science of probabilities, and such exact knowledge of rela¬
tionships has not been attained in any group of plants of considerable
size. Lichens resemble other ascomycetes in their methods of sexual
reproduction, and their special manner of vegetative reproduction by
soredia and soralia can have no considerable weight in favor of treat¬
ment as a distinct group of plants. Higher lichens exhibit certain pecul¬
iarities of vegetative structure, due to their parasitism on algie ; but
their fruits are like those of other ascomycetes, which in turn vary
greatly with respect to somatic structure. In view of the facts regarding
the reproductive tracts and the fruits, the peculiarities in regard to
somatic areas are not sufficient ground for distinct consideration of
lichens, especially since they are connected with other ascomycetes by
hundreds, if not thousands, of intermediate orders, genera, or species.

Present separate incorporation of lichens in herbaria, the large litera¬
ture of lichenology, and the fact that students of ascomycetes, for most
part, may be divided into two groups, those who study lichens and those
who study other ascomycetes, present real obstacles to be overcome ; but
arguments of expediency and temporary convenience are hardly valid
reasons for maintaining a system of classification that is not in accord
with present knowledge regarding the structure, the ontogeny, and the
probable phylogeny of ascomycetes. For this reason, many botanists
who have considered the matter carefully believe that our method of
treatment, which is, in spite of temporary inconvenience, in accord with
knowledge of the ascomycetes, should he followed in any systematic
treatment of the whole group.

There are many genera of ascomycetes which contain some lichen
and some non-lichen members, and, as stated above, the bridges that
connect ascomycetous lichens with other ascomycetes are many. The best
way to facilitate comparison of these forms and thus strengthen our
knowledge of the ascomycetes as a whole is to treat the group as a unit.
This is, without doubt, also the best way to express what is known
regarding these fungi.

Some of the points of nearest approach of lichens to non-lichen
ascomycetes are the following. Endomyccs scytoncmatum is a lichenoid

6

Ohio Biological Survey

plant. The genus Mycocalicium, of the Caliciaceae, has been segregated
from Calicium to receive members which do not live with algae. Sclero-
tinia tubcrosa is known to live rarely in parasitic relation with an alga
(Synes58). Some of the Cclidiaccae are algicolous, others not.
Agyrhim riifum has been considered both lichen and non-lichen. The
species of Lccideopsis seem to be partly lichens and partly non-lichens.
Tobler’s study of Phacopsis vulpina (62) proves that this plant is a lichen,
and the species of Conida and Celidium await similar study. The Patel-
lariaccae present similar conditions in that the genera KarscJiia,Biatorella,
Lcciographa, and Melaspilea all contain some algicolous and some non-
algicolous ascomycetes. Among the Lecanactidaceac are species which
are non-algicolous. Zukal’s investigation (67) has shown that Pariiphrd-
dria heimerlii and Geopcma rehmii of the Cenangiaceae are both
algicolous, though other members of the family are non-algicolous. Some
species of Dichaena are supposed to be lichens, while other members of the
Dichacnaceae are not. The lichen genus Arthonia is well known to con¬
tain non-lichen species and other species that are non-algicolous during
part of their lives. The same condition is found in the lichen genera
Graphis and Opegrapha (Frank 33), and no one knows how many
species are non-algicolous during part or all of their lives, or how many
other genera of the Arthoniaceae and the Graphidaceae contain both lichen
and non-lichen members. Further statements regarding forms transi¬
tional between lichen and non-lichen ascomycetes will be found in a
previous paper (Fink 31), and only extended biological research will
give definite knowledge of hundreds or perhaps thousands of such plants
which are poorly understood.

The research on the sexuality of ascomycetes is all comparatively
recent, nothing of importance dating back of 1863 (De Bary 3). The
whole literature of the subject is readily available and is convincing with
respect to the rather close relationship among all undoubted ascomycetes.
The lichens that have been studied, except those that are degenerate with
respect to sexual reproduction, show the Collema-Mke type of female
reproductive tract, which is very similar to that found in some other
ascomycetes that have been studied. Likewise, most lichens and other
ascomycetes agree in that they develop similar ascocarps, simple or com¬
pound. It is admitted that the topography, the cytolog}-, and the full
development of the sexual reproductive tracts and the ascocarps are not
completely known in any of the species that have been studied, and a
large amount of research is still necessary before we can hope to approxi¬
mate a natural classification of ascomycetes. The nuclear behavior is

Classification of Ohio Ascomycetes

7

known more or less completely in a considerable number of non-algi-
colous species, but in only one algicolous species (Bachman 2). Though
the results are fragmentary and some of them unreliable, one who con¬
siders carefully even the small portion of the research that can be dis¬
cussed in this paper can scarcely doubt that the ascomycetes form a
natural group of fungi.

Through the researches of several botanists, we know something
regarding the sexual reproductive tracts and sexual reproduction in
lichens (Bachman 1 and 2; Baur 4 and 5; Darbishire 2; Funfstiick 37
and 38; Krabbe 42 and 43; Lindau 44 and 45; Nienburg 50; Schulte
55; Stahl 59; Sturgis 60; Wahlberg 63; Wolff 66; and others). The
many-celled archicarp, with its long, many-celled trichogyne and its coiled,
septate ascogonium, is quite uniform and recalls conditions found in
higher non-lichen ascomycetes, especially certain members of the
PezizaleSy the Hypocreales, the Sphaerialcs, and the Lahoulheniales.

One can imagine readily enough how a one-celled ascogonium sur¬
mounted by a one-celled trichogyne, as in Pyronema (Harper 39),
Claussen 16) may be related to the complicated, multicellular, and
imperfectly understood female reproductive area of certain lichens and
other ascomycetes (Fig. 1, et seq.) ; but the simpler female tracts usually
occur with undoubted antheridia, and these cannot be homologized so
easily with the spermagonia usually found in ascomycetes which possess
the more highly differentiated female reproductive tracts. Hence, we
may also suspect, among those non-lichen ascomycetes that have sperma¬
gonia and the more complicated female reproductive tracts, a close rela¬
tionship with certain of the polyphyletic lichens. Rather unexpectedly,
only part of the Pezizales examined have furnished types of reproductive
tracts so nearly like those of the gymnocarpous lichens examined as have
several species of angiocarpous, non-lichen ascomycetes. While the
reproductive tracts of a large majority of the Pezizales and the Lecano-
rales remain tO' be studied, we may still expect to find many closely related
forms connecting these two orders ; but, for the present, usual similarity
in the topography of the reproductive areas has been one factor which
has led to placing the gymnocarpous lichens, for most part, in a con¬
tinuous series. Future studies may require more or less modification of
this method.

The discussions to follow will further elucidate this matter and will
show especially a striking and rather surprising similarity between the
reproductive tracts of the Lecanorales and the pyrenomycetes in general.
Polystigma rtibrum, whose archicarp has been described and figured

8

Ohio Biological Survey

(Fisch 32 and B'rank 34), has spermagonia, sperinatia, ascogonia, and
jierhaps trichogynes especially like those found in many of the Leca~
noralcs. Gnomonia crytJirostoma also has given similar results (Frank
35). Working on the former plant, Blackman and Welsford (10)
confirmed the findings of Fisch and Frank regarding the female tract in
general, but thought that their supposed trichogynes might be vegetative
hyphie which protrude through the stomata, that the spermatia might be
functionless, and that the ascogenous hyphie might arise from somatic
tracts (Fig. 3). However, all of these results on Polystigma are offset to
some extent by Nienburg’s results given below. Brooks (13) found in
Gnomonia- crytJirostoma spermatia attached to the trichogynes, but
thought that both spermatia and trichogynes might be functionless. As
was noted by several workers, these reproductive tracts, however degen¬
erate they may be, show in both spermagonia and archicarps a strong
resemblance to conditions well known in certain lichens which are con¬
sidered below. Higgins (41) has recently found in Sphacrclla very similar
appearances, which need further investigation.

It is convenient to call the coils of an ascocarp an ascogonium and
the remainder of the structure a trichogyne, but such use of terms is often
inexact. Nienburg (51), working on Polystigma rubrum, seems to have
found that one uninucleate cell of the coiled portion of the archicarp
is the ascogonium. A multinucleate cell next the uninucleate cell in the
same coil, he regards an antheridium. He finds what he supposes to be
functionless trichogynes and spermatia, and regards the antheridium and
the method of reproduction respectively a secondarily acquired reproduc¬
tive organ and a secondarily acquired sexual process. He believes that
the ascogenous hyphse do arise from the ascogonium. The question
remains whether he may not have missed a temporary perforation of the
transverse sejita of the trichogynes and the passage of sperm nuclei
through the trichogyne and other cells of the highly differentiated coil
to the ascogonium. His paper is remarkably interesting and suggestive
and will of course result in a re-examination of the much-studied but
still not fully understood Polystigma rubrum and many other ascomycetes
which have similar reproductive tracts. We must pass over Nienburg’s
evidence of delayed nuclear fusion and his discussion of probable rela¬
tionship between the Ascomycctac and the Oomycctac. In this matter,
his views are similar to those of Claussen (16). Recent work by Dodge
also sheds light on the real nature of the ascogonium (23 and Fig. 7).

According to Miss Nichols (49), Ccratostoma brevirostrc has a one-
celled, coiled ascogonium, to whose tip an antheridium becomes attached

Classification of Ohio Ascomycetes

9

directly. The ascogonium septates after fertilization, and the resulting
ascogenous structure (ascogenous hypha?) appears at this stage much
like the septate ascogonium of a lichen. Miss Nichols suspected that
De Bary might have been mistaken about the ascogonia degenerating and
the asci arising independently in Xylaria, and that other earlier workers,
Hartig (4) and Woronin, might have failed to follow the development
carefully in Roscllinia and Sordaria. If her suspicions are correct, and
H. B. Brown (14) has since proved them so for a species of Xylaria,
conditions found in these non-lichen species are somewhat similar to
those known in lichens with respect to the female reproductive tract. If
any of the results which indicate that degeneration of the female repro¬
ductive tract and the independent origin of ascogenous hyphse and asci
are reliable, such degenerate conditions would be similar to certain degen¬
erate states noted below in certain lichens.

Miss Dawson (22) has also found, in Poronia punctata, conditions
much like those reported in lichens by several workers. The female
reproductive tract consists of a multicellular, coiled hypha surmounted by
a more slender, muticellular portion which rises a short distance above
the surface of the stroma. In later stages, the supposed ascogonium is
absent, and ascogenous hyphse appear. Miss Dawson could not connect
the last two structures, nor did she find spermagonia. However, she
considered the female tract similar to that found in lichens, and her
figures indicate one of the nearest approaches hitherto discovered among
non-algicolous ascomycetes to conditions well known among lichens, at
least in regard to appearance of the archicarp.

Ascogonia not accompanied by trichogynes have been reported for
Pcltigera, Pcltidca, and Nephroma (Ffinf stuck 37), and it is supposed
that, in these plants in which spermagonia are absent or rare, the
apothecia arise apogamously (Fig. 8). Baur (5) has reported similar
conditions in Solorina saccata, and several other lichens have given prac¬
tically like results (see especially Sturgis 60). All of these lichens show
degeneration of the sexual tracts comparable to that found in some of
the non-lichen ascomycetes considered above.

Some slightly different conditions among lichens must be noted.
The genus Bacomyces seems to present conditions similar to those already
discussed for other ascomycetes^ from the results of Brooks, those of
Nienburg, and those of Blackman and Welsford. Nienburg (50) has
also worked on lichens and found no trichogynes in Bacomyces, nor was
he sure that he found ascogonia. He thought that the asci might arise
either from ascogenous hyphse or directly from ascogonia. Spermagonia

10

Ohio Biological Survey

are rare in the genus, and he concluded that the apothecia arise apogam-
ously. In his work on Sphyridium byssoides, he found reduced and
probably functionless trichogynes, while here again the spermagonia are
rare. In this species, however, he succeeded in tracing the origin of
ascogenous hyphse from ascogonia. Reduction of the sexual apparatus
does not seem to have proceeded so far here as in Baeomyces ; but the
plant may be apogamous, and, according to Nienburg’s results, it is inter¬
mediate between Baeomyces and those lichens which have normal sexual
organs, both male and female. In the Caliciaceae, sexual reproductive
areas seem to be for most part absent, though Neubner (48) supposed
that functionless spermagonia might exist in certain species. Neubner
thought that the apothecial development must be wholly vegetative.
Mrs. Gertrude Wolff Tobler (66) noted somewhat uncertainly a single
transverse septum in the trichogynes of Graphis clegans, and we seem
to have here a condition intermediate between the unicellular trichogyne
of some non-lichen ascomycetes and the typical lichen trichogyne. It
is not quite certain whether these simpler archicarps are primitive or de¬
generate, and more work on the Hysteriales, whether algicolous or non-
algicolous, would certainly give other helpful results.

It seems probable that in Pyreniila nitida the female sexual tract is
perhaps more like the well-developed conditions found in certain disco-
mycetous lichens that have been studied than those of the non-lichen
pyrenomycetes that have been examined. Baur’s work on Pyreniila
needs confirmation and extension, since there is room to suspect that the
resemblance of the female reproductive tract to the typical, fully devel¬
oped form found in discomycetous lichens may not be so close after all.
Much work is needed on both lichen and non-lichen pyrenomycetes before
we can know certainly whether the pyrenomycetous lichens are related
more closely to non-lichen pyrenomycetes than to discomycetous lichens.

Conditions among the Laboiilbeniales (Thaxter 61 and Faull 26
and 27) are considerably like those found in lichens. A complete series
from the one-celled trichogyne to a multicellular and much-branched
form has been found (Fig. 4). Non-motile male cells become attached
to the trichogynes as in lichens, and later the ascogonium septates or in
some manner gives rise to a septate structure somewhat similar to the
septate ascogonium of lichens. Some of the conditions known in the
Laboulbeniales appear to be intermediate between those found in Cera-
tostoma and those that have been observed so frequently in lichens.

There are also conditions among the PezPalcs which strongly sug¬
gest the female reproductive tract in lichens. In Laehnca stcrcorea,

Classification of Ohio Ascomycetes

11

Fraser (36) found a one-celled ascogonium surmounted by a several-
•celled trichogyne (Fig. 2). The cells of the female reproductive tract
are multinucleate, and the conditions may be intermediate between those
found in Pyronema (Harper 39 and Claussen 16) and those well
known in lichens. In Cutting’s (19) discussion of results obtained in a
study of Ascophanus carneus, it appears that the archicarp consists of a
multicellular ascogonium and a multicellular trichogyne comparable to
those of lichens. Two recent researches show a strong resemblance
between the sex organs of lichens and non-lichen discomycetes. Miss
Bachman’s results (1 and 2) in a member of the Collcmaceae (Fig. 6)
and those of Dodge (23) in Ascobolus carhonarius (Fig. 7) furnish
striking resemblances which, if later found to occur often in both algi-
colous and non-algicolous discomycetes, will demonstrate a much closer
relationship between certain gymnocarpous lichens and the PcAnalcs than
we are at present showing in our classification. There is good reason for
believing that, in many supposed apogamous lichens and in others as well,
internal trichogynes and internal spermatia (antheridia?) may occur,
and that further studies will give results similar to those obtained by
these workers. Among the lichens, some of the promising plants are
•other members of the Collemaceae, some of the lichens supposed to have
degenerate reproductive tracts and apogamy, and members of the Gyalec-
taceae and the Lecideaceae, which are probably close relatives of the
Collemaceae.

The various results concerning sexual reproduction among ascomy¬
cetes recounted above and others not considered herein surely lend color
to the supposition that the ascomycetes are monophyletic ; but the diffi¬
culty of homologizing the various types of male reproductive organs still
remains, somewhat bridged, however, by types of male organs apparently
intermediate between spermatia and antheridia, and suggesting a possible
relationship with conidia as well. It has been suggested by some workers
that all multicellular ascogonia in ascomycetes are post-fertilization con¬
ditions, but the evidence is against this view. The multicellular
structures that arise after fertilization from a one-celled ascogonium in
such ascomycetes as the Erysibaceae, Gymnoascus, and Ceratostoma seem
rather to represent ascogenous hyphse. If this be true and the ascogonia
of lichens and certain other ascomycetes be really many-celled, such post¬
fertilization areas cannot be homologous with these muticellular asco¬
gonia. It will be seen that this discussion is made with allowance for
lack of knowledge, in most instances, whether one cell or more than one
of the coil of the archicarp constitutes the ascogonium.

12

Ohio Biological Survey

Only a very small proportion of the many groups of ascomycetes
has been studied with respect to sexual reproductive areas and life
histories in general, and much of the work accomplished is preliminary,
fragmentary, unconlirmed, and unsatisfactory. In all present arrange¬
ments of ascomycetes, we hnd occasionally in juxtaposition families or
larger groups that have very different reproductive tracts; and further
studies are certain to reveal more of this sort of inconsistency in systems
of classiheation based largely on spore characters and mature conditions
of ascocarps. However, one cannot make very great changes based on
the comparatively few and often fragmentary and uncertain researches
that have been made. For example, the lichens must be interpolated,
for the present, in large groups among other ascomycetes, though they
mav need further distribution after we know more about life histories.
The general similarity of the reproductive tracts in a large majority of
the lichens that have been studied, the resemblance of the somatic tracts
in large groups of lichens, and the rather limited knowledge of the rela¬
tionship between lichens and other ascomycetes justify this conservatism
in distribution.

In previous arrangements of ascomycetes in a single system, no dis¬
cussion of possible or probable homologies and relationships has been
attempted. With our present limited knowledge, such discussion must
serve as an argument for the method rather than a very great help in the
arrangement. Unfortunately, we must continue to base our taxonomic
treatment of ascomycetes quite largely on spore characters and on the
structure of mature fruits until we have sufficient data to enable us to use
more fundamental characters. However, in the plan of classification to
follow, the Lccanoralcs have been kept distinct from the Pczizalcs, partly
because the lichens that have been examined from the former group have,
on the whole, a similar type of reproductive organs, more complicated
than those in the latter group, in which these tracts are as a whole more
degenerate, or possibly more primitive. Which of these two orders
should stand higher is still an open question, regarding which certain
statements are made below. For the present, we have placed the order
which has the more specialized somatic structure higher. However, it is
readily admitted that the higher form of vegetative structure may indi¬
cate either less of degeneration or a better development because of the
advantageous conditions under which the lichens live. Again, if the
Ascomycetac have arisen from the Rhodophyceae, a consideration to be
taken up below, the Lecanorales may yet be proved to have, as a whole,
a shorter line of descent than the Pezizales.

Classification of Ohio Ascomycetes

13

Whatever consideration we may give to somatic characters and
carpologic structure as a basis for classification, it seems certain that
the character of reproductive tracts forms a more valuable criterion.
Yet it needs to be kept in mind that sometimes, as in the Collemaccac,
forms which differ greatly with respect to structure of sexual organs and
sexual behavior are so much alike with respect to vegetative characters
and mature carpologic structure that the latter features have had greater
weight in taxonomic disposition. The plant studied by Miss Bachman
(1 and 2 ) illustrates well the point suggested above. It is probably more
primitive than members of the Collcinaccae whose sexual reproductive
areas, show spermatia in spermagonia, the evolution of the male repro¬
ductive area having probably lagged while carpologic structure and
somatic conditions advanced in the direction followed in the evolution of
the genus Collcma. In spite of the rather striking resemblance of the
reproductive areas of the so-called Collcma piilposum studied by Miss
Bachman and those of Ascobolits carbonariits examined by Dodge (23),
one cannot overlook somatic characters, to the extent of placing the
former plant with the Ascobolaccac rather than with the Collcmaccae.
This is one striking instance in which vegetative characters and carpo¬
logic structure have great value in classification ; but it is scarcely possible
that plants which have such different male reproductive areas as Miss
Bachman’s plant and the one examined by Stahl (59) (Fig. 5) should
both be assigned to the same species, Collcma pitlposum.

Studies of certain ascomycetes show that forms which differ greatly
with respect to somatic and carpologic structure and therefore are far
removed from each other in present systems of classification have very
similar reproductive areas, while others much alike with respect to
somatic and carpologic structure, but very different in regard to male or
female reproductive areas or both, have been placed in the same family,
the same genus, or even in the same species. For instance, the genus
Ascobolits shows a wide range of variation with respect to sexual areas,
while the range is much wider for the Ascobolaccac as a whole. A full
discussion of conditions found in this family may be found in one of the
papers cited from Dodge (23). Wdiile the student who deals with
ascomycetes from the taxonomic point of view must endeavor to famil¬
iarize himself with all studies of the life histories of these fungi, it is by
no means clear at present just what changes should be made in existing
systems of classification on account of the facts more or less clearly,
fully, and certainly ascertained regarding reproductive processes and
ontogeny. At least, a good deal of conservatism may well be observed

14

Ohio Biological Survey

until our information is greatly increased so that many facts which have
a bearing on classification may be known and considered in correlation.

Our knowledge of the development of the ascocarp is perhaps even
more fragmentary than that of the sexual apparatus, toward which a
large portion of the recent work has been directed. It is still an open
question whether the cleistothecium, the perithecium, or the apothecium
is the most primitive form of the ascocarp ; and the whole development
of many carpologic structures in various groups of ascomycetes must be
carefully studied before we may know their worth as criteria for con¬
structing a system of classification.

The Besseys (8 and 9) think that there is good reason to believe that
C ollcma-\\kt plants with apothecia, like the one studied by Miss Bachman,
may be the most primitive of existing Ascomycetae and closely related
to the Rhodophyceae. This suggestion seems reasonable since the species
of Collcma and Miss Bachman’s plant are low forms of lichens vegeta-
tively, the thalli being mycelia devoid of plectenchymatous structure,
while the plants are apparently near relatives of such families as the
Lccideaccac, the Gyalectaceae, and the Ascoholaceac, all of which may be
rather primitive ascomycetes. Of these groups. Miss Bachman’s plant
and Ascobolus carbonarius, studied by Dodge, are nearest the
Rhodophyceae of those that have been examined with respect to repro¬
ductive tracts, and may well be regarded as the most primitive ascomy¬
cetes whose life histories have been investigated. Of the two plants.
Miss Bachman’s, on account of the character of its male reproductive
organs, appears to be nearer the Rhodophyceae. Yet the view that the
first ascomycetes were algicolous is opposed to the one commonly held
by botanists to the effect that lichens may be traced back to non-
algicolous ascomycetes. If the theory that the most primitive ascomy¬
cetes are those having apothecia be true, of course much more investiga¬
tion is needed before we can safely decide whether the primitive forms
were lichen or non-lichen. If lichen, it may be that we have the most
primitive of existing forms among such members of the Collemaceae as
possess the low vegetative structure noticed above and reproductive
tracts and sexual behavior most like those known in the Rhodophyeeae.
The primitive forms are probably to be sought among those that live in
loose relationship with the algal host, but are still largely dependent on
this host. The Collemaceae fulfill these conditions, since they do not,
with one or two exceptions, form haustoria, but simply penetrate through¬
out the gelatinous Nostoc colonies, confining their course to the sheaths
and scarcely injuring the protoplasts, while they are but weakly attached

Classification of Ohio Ascomycetes

15

to the substratum by rhizoids and are almost wholly dependent on the
host for nourishment. From such a primitive form of ascomycete, there
could be two lines of descent, one of which would give rise to higher
lichens with foliose and fruticose thalli, close relationship with the algal
host, and the apothecial form of ascocarp retained, while the other
would produce the lower lichens which are crustose upon the substratum
or sunken into it and which live in increasingly loose relations with the
algal host as they sink into the substratum. Some of these forms could
have lost the algal habit gradually as they came into closer and closer re¬
lationship with the substratum and so have given rise to non-algicolous
ascomycetes. As a rule, lichens which have perithecia or hysterothecia
are those that are sunken into the substratum, while those that are
crustose upon the surface, or are foliose or fruticose, usually have
apothecia. On the theory that the apothecium is the primitive form of
ascocarp, perithecia and hysterothecia would arise as protective struc¬
tures in reaction to xerophytic habit or sinking into the substratum. Of
course we may begin with such a form as Ascobolus carbonarius,
suppose that it evolved a form having the algicolous habit, and then
apply the above reasoning equally well. Therefore, the rather unim¬
portant question whether primitive ascomycetes were lichens or not, is not
so easy to settle, though it is reasonable to suppose that certain
Rhodophyceae which had become parasitic on other marine algge may
have acquired a terrestrial habit, passed into parasitic relationship with
terrestrial algae, and acquired the ascomycetous forms of reproduction
and carpologic structure as a response to changed conditions of environ¬
ment. The relation of the lichen to its algal host and to the substratum
touches this problem and is discussed in a previous paper (Fink 31).

To generalize from finding one or two plants which have apothecia
and at the same time possess reproductive areas and processes much like
those found in the Rhodophyceae would be a very uncertain procedure ;
but the discussion by Dodge (24) shows plainly that the generally
favored theory that the perithecium is the primitive form of the ascocarp
rests on very uncertain evidence at best. While it is certain that many
careful studies of carpologic structure and development will be neces¬
sary before we may know with any reasonable degree of certainty
whether the apothecium or the perithecium is more primitive, it has
seemed reasonable, in view of all the evidence at hand, to adopt a
sequence of orders of ascomycetes which comports in a general way with
the view that the apothecium is the primitive form of ascocarp. In fol¬
lowing this view, it has seemed best to place the algicolous discomycetes

16

Ohio Biological Survey

above the non-algicolous forms because of their higher somatic develop¬
ment and the general resemblance between their sexual reproductive
tracts and those of the pyrenomycetes above them. In so doing, it is
plainly seen that any method that could be followed is open to criticism
and liable to be proved incorrect by further research.

The resemblances between the sex organs of the Ascomycctae and
the Rhodophyccac have been a matter of repeated discussion since the
time of Stahl’s work on the Collemaccae and have been greatly empha¬
sized since by the researches of Thaxter (61), Bachman (1 and 2),
Dodge (24), and others. The ascogenous hyphae of the Ascomycctae like¬
wise have been homologized with the ooblastema filaments of the
Rhodophyccac, while a relationship has been suggested between asci and
the gonidioblasts which arise as the result of secondary fusions among
the red algie. If the evident parallelism in development among the
Ascomycctae and the Rhodophyccac rests on supposed homologies, we
may also conclude that ascocarps and cystocarps are homologous struc¬
tures, and that the former group arose from the latter, or that the two
groups had at least a common origin. One of the two suppositions just
stated must be adopted, or we must follow those who think that these par¬
allelisms do not rest upon homology. This matter is fully treated by
Dodge (24 ) .

In this connection, it might prove worth while to reexamine the
forms of parasitism found among the Rhodophyccac, and reproductive
and somatic tracts in these parasitic algae, with a special view of attempt¬
ing to shed some light on the problem of possible affinities between the
Rhodophyccac and the Ascomycctae. It is even possible that, if asco¬
carps are to be homologized with glomerules and cystocarps, and much
more fundamental and significant homologies exist between structures
which lead up to these mature fruits, such studies would indicate a
multiple origin of the Ascomycctae from the Rhodophyccac and suggest
that more than one type of ascocarp may be primitive. Such studies
might also help to decide whether forms of ascomycetes which have simi¬
lar ascocarps are all closely related, or whether certain groups which have
very different ascocarps are not sometimes nearly related.

^^4latever the future may prove regarding the various possible or
jirobable homologies between the Ascomycctae and the Rhodophyccac, it
is certain that the theory of the origin of the former from the latter is the
most plausible one that has been offered regarding the phylogeny of all
undoubted ascomycetes, algicolous and non-algicolous.

Cytological studies must prove increasingly valuable in our studies

Classification of Ohio Ascomycetes

17

of the relationships among ascomycetes. Especially will knowledge of
nuclear behavior in the reproductive tracts finally have considerable
weight in the arrangement of ascomycetes, since it will enable us to
decide just what portion of an archicarp is the ascogonium. At present
nuclear behavior is not known in the sexual reproductive tracts of any
lichens save one (Bachman 2), nor is it well understood in many other
ascomycetes. According to Claussen (16), delayed nuclear fusions and
homologous alternation in ascomycetes strongly resemble the conditions
found in certain basidiomycetes. This points to the origin of the latter
group from the former, and further studies of nuclear behavior in both
groups may shed some light on the relative rank of the various ascomy¬
cetes. The rather uniform character of ascogenous hyphae and the
peculiar origin of asci from them, together with the characteristic be¬
havior of the nuclei within these structures, is probably the strongest
proof of the common, or at least similar, ancestry of all ascomycetes.

There is another theory of the origin of ascomycetes that must be
noted. Bucholtz (15) has presented a modification of the theory of
origin of the Ascomycetae from the Phycomycetae which is especially
worthy of consideration. He considers Endogone to be a plant whose
fruit results from the conjugation of heterogametes. After cytoplasmic
fusion, the two nuclei pass together from the oogonium into a protrusion,
which might be regarded as a primitive ascus. Here the nuclei fuse.
Not far removed from this is the formation of asci as direct outgrowths
from the single fertilized ascogenic cell described by Faull (26) for
Amoj'phomyces and said to occur in some other genera of the Laboul-
beniales. Nienburg (51), Claussen (16), and others have also' presented
strong arguments for this theory, which would make trichogynes and
ascogenous hyphae structures which arose independently in ascomycetes,
would deny the origin of the Ascomycetae from the Rhodophyceae, and
would account for the resemblances between these two groups by
parallelism alone. However, the evidences of relationship between the
two groups are too strong to be ofifset by such facts as have been ad¬
vanced hitherto for the origin of the Ascomycetae from the
Phycomycetae.

It will be seen from the above presentation that no attempt has been
made to consider the problem of classification of ascomycetes merely as
it applies to our studies in Ohio. The outline of classification that is
presented herewith is the result of a study of the problem concerned
with the phytogeny of all ascomycetes. To base any system of treatment,
even for a small area, on anything less than a consideration of all

18

Ohio Biological Survey

ascomycetes as to their origin and relationships would be to invite
serious failure. Consequently, in preparing the system to follow, all the
orders of ascomycetes have been considered, while many families that
do not occur in our flora, and, consequently, do not appear in the outline,
have been kept in mind, and have been discussed in part. How these
families should be treated will be taken up more fully in the third paper
of the series on “The nature and classification of lichens” (Fink 30 and
31), in which the whole problem of the classification of ascomycetes will
be reconsidered with such additional data as may come to light in the
interval.

As stated in the introductory paragraph of this paper, ascomycetes
have been treated previously as a whole. Lists from Nebraska (Webber
64 and 65) and California (McClatchie 46) have distributed lichens
among other ascomycetes, and certain writers (Bessey 6, 7 and 8;
Clements 17 and 18; Fink 28, 29 and 31, and Schaffner 54) have either
united all ascomycetes into one system or intimated that this method of
treatment is in accord with present knowledge of the group. Certain
European mycologists regard lichens as fungi, and some of them have
either advocated their distribution among other fungi, or have admitted
that they may, without inconsistency, be so treated (Fink 31). The dis¬
position proposed below is based on the belief that we have sufficient
data, in part briefly outlined above, to warrant the distribution of lichens
in the best manner possible in order that we may present relationships
better and have a good basis for comparative studies of all ascomycetes.
The statement in this paragraph expresses views that are held by various
botanists and have been used in certain arguments, keys, catalogues of
plants, and systems of classification cited. However, this series of
papers will be the first to treat lichens as fungi with other ascomycetes in
a descriptive catalogue.

A summary regarding the arrangement of the orders and some of
the families will elucidate some points. The Hemiascales {Hernias cine ae)
of some authors have not been admitted, though Eremascus and some
others of these plants may yet prove to be ascomycetes rather than
phycomycetes. The Lahoulbeniales have been placed first in the outline
of classification, since the work of Thaxter (6) and Faull (26 and 27)
makes them undoubted though aberrant ascomycetes, apparently near to
the Rhodophyceae, which have similar archicarpic characters. With
respect to ascogenous structures, they must be regarded either degenerate
or closely related to those members of the Rhodophyceae that produce
ooblastema filaments, while the closed ascocarp would seem to indicate

Classification of Ohio Ascomycetes

19

higher rank than is given to the order. In any event there seems to be
no good place to interpolate the Laboulbeniales among the other orders,
and first place appears to accord better with present knowledge than the
place at the end of the ascomycete series to which the order has been
assigned more commonly. In placing the order first, one could not afford
to commit himself to the belief that the Laboulbeniales are nearer to
the Rhodophyceae than are the Pezizales and the Lecanorales, which in
turn are probably more plainly related to the orders that follow in the
outline. The two orders just named are placed second and third in
the series on the theory, based on carpologic development and structure,
that they, too, are closely related to the Rhodophyceae. The Lecanorales
have been kept in one large group rather than split up into two or more
orders, partly on account of their close relationship among themselves
and partly because further study may result in a more or less radical
rearrangement, which is likely to involve the Pezizales as well. For
instance, the Collemaceae, the Gyalectaceae, and the Lecanactidaceae seem
to be, partly in light of studies reviewed above, nearer to some of the
Pezizales than we are able to indicate in our scheme of classification..
Some students of ascomycetes favor separating the inoperculate mem¬
bers of the Helvellales from those that are operculate. If this be done,
the group may well be distributed among the Pezizales. The order
Helvellales has been retained for the present and is placed near the
Pezizales, to which it seems to be closely related. The Tuberales, which
have been variously treated by students, seem to be near relatives of both
the Pezizales and the Helvellales, the peridium being probably a reaction
to subterranean position of the ascocarp. Interpolation of the Lecan¬
orales prevents placing the Helvellales and the Tuberales in succession
immediately after the Pezizales, but this is only one of the many un¬
avoidable difficulties met in efforts to formulate schemes of classification
which will bring together forms that seem to be closely related. Proper
sequence of the families of the Pezizales depends on whether the simpler
archicarps are primitive or degenerate, a point not fully determined.
Consequently, the families are arranged in an order which leads up to
the Lecanorales, which are placed next above. This arrangement shows
the relationship between the two orders by passing from strictly non-
algicolous to partially algicolous families of the Pezizales, thence to the
algicolous Lecanorales. Of course this method must eventually give
way, if need be, to better knowledge of the reproductive areas and related
development. Since the Lecanorales in general show great similarity in
sexual reproductive areas, the sequence of families is based on apothecial

20

Ohio Uiological Survey

and vegetative structure, though it is plainly seen that increasing knowl¬
edge of degeneration in the sexual areas and the sexual processes may
soon require some modification of this method. Regarding the Phacidiales
and the Hysterialcs, it is quite possible that further study may result
in placing some of the lichens in the former order, nor is it certain that
these orders are intermediate between discomycetes and pyrenomycetes.
Concerning the angiocarpic groups, it is not probable that the perithecia
and the cleistothecia all arose in similar manner, or that they are so
closely related as the order adopted would indicate. The Pyrcmilales
are placed after the non-algicolous groups for about the same reason that
the Lecanorales are placed after the Pezisales. Here again, further
studies are likely to require some modification of the method used herein.
The Perisporialcs, the Aspergillales, the Exoascalcs, and the SaccJiaromy-
cctales have been placed toward the close of the series since, if the theory
of origin of the Asconiycctae from the Rhodophyceac be true, each of
these orders, as a whole, seems to be composed of either degenerate or
doubtful ascomycetes. Degeneration seems probable, on this basis of
origin, in the simple reproductive tracts tending toward likeness in form
and size of gametes, in reduction and final loss of the trichogynes, and in
undoubted isogamy in a few instances among the saccharomycetes,
though in this order the conjugati.^n, rarely observed, may be a
secondarily acquired character.

In closing, it should be stated that all speculations regarding the
relationships of ascomycetes are rendered more uncertain by the prob¬
ability that carpologic evolution may have proceeded in nearly parallel
lines in various groups so that forms that differ considerably with respect
to archicarps and ascocarps may be more closely related than others that
show considerable similarity with respect to these areas.

In the studies of the orders and families of Ohio ascomycetes to
follow in this series of papers, the better-known groups will be published
while field work and laboratory investigations are increasing our knowl¬
edge of orders at present unknown to our flora, or poorly understood.
The system of classification proposed herein will serve as a basis for the
work and will indicate where each group studied is related in our scheme
of classification adopted for the ascomycetes as a whole. The final work
on the ascomycetes of Ohio should be to bring these studies together
under a system of classification revised sufficiently to show the advance
in knowledge of the group while the preliminary studies are in progress.
The provisional arrangement which is to serve as a basis for the study
of various groups of Ohio ascomycetes is given below.

Classification of Ohio Ascomycetes

21

CLASS ASCOMYCETAE

L LABOULBENIALES (LABOULBENIINEAE)

Many species of this order doubtless are present in Ohio, though
none have been listed.

II. PEZIZALES (PEZIZINEAE)

1. Pyronemaceae. The genus Pyronema is known in Ohio.

2. Pezizaceae. Genera known in Ohio' are Peziza^ Lachnea,
Geopy.vis, Hmnaria, Otidea, and Urmila.

3. Ascobolaceae. Genera known in Ohio are Ascoholus, Lasio-
holns, Ascophaniis, Saccoholus, and Rhyparobiiis.

4. Helotiaceae. Genera known in Ohio are Helotiiim, Sarcos-
cypha, Chlorosplenium, Ciboria, Sclerotinia, Arachnopeziza,
Dasycypha, Stamnaria, Ombrophila, Coryne, and Phialea.
Sclerotinia tiibcrosa sometimes grows with an alga.

5. Mollisiaceae. Genera known in Ohio are Mollisia, Tapcsia,
Orbilia, Belonidium, and Pezizella.

6. Cenangiaceae. Genera known in Ohio are Cenangiiim, Cen-
angclla, Dermatea, Tympanis, Sarcosoma, and Holwaya. There
are some primitive lichens in this family.

7. Patellariaceae. Genera known in Ohio' are Patellaria,
Patellea, Patinella, Karschia, Leciographa, Diirella, and Bia-
torella. Certain species of Patellea, Patinella, Karschia, Lecio¬
grapha, and Biatorella are lichens.

III. LECANORALES (LECANORINEAE)

1. Lecanactidaceae. The genus Lecanactis is known in Ohio.

2. Lecideaceae. Genera known in Ohio' are Lecidea including
Biatora), Bacidia, Lopadinm, Buellia, Psora, Toninia, and
Rhizocarpon.

3. Cladoniaceae. The genus Cladonia is known in Ohio.

4. Gollemaceae. Genera known in Ohio are Collema, Synecho-
blastus, Leptogium, and Mallotium.

22

Ohio Biological Survey

5. Pannariaceae. Genera known in Ohio are Pannaria, Heppia,
and Placynthinm.

6. Stictaceae. Genera found in Ohio are Sticta and Lobaria.

7. Peltigeraceae. Genera known in Ohio are Peltigera, Peltidea,
and Nephroma.

8. Acarosporaceae. The genus Acarospora is known in Ohio.

9. Lecanoraceae. Genera known in Ohio are Lecanora and
0 chr ole cilia .

10. Pertusariaceae. The genus Pertusaria is known in Ohio.

11. Parmeliaceae. Genera known in Ohio are Parmelia, Anzia,
Cetraria, and Nephromopsis.

12. UsNEACEAE. Genera known in Ohio are Usnea, Ramalina,
Alectoria, and Evernia.

13. Teloschistaceae. Genera known in Ohio are Teloschistes,
Placodium (Calopisma) , and Xanthoria.

14. Physciaceae. Genera known in Ohio are Physcia, Pyxine,
Anaptychia, and Rinodina.

15. Gyrophoraceae. Genera known in Ohio are Gyrophora and
Umbilicaria.

IV. CALICIALES (CALICIINEAE)

1. Caliciaceae. Genera known in Ohio are Calcium, Chaenothcca,
Stenocybe, and Cyphelium. These plants are mostly lichens.

V. HELVELLALES (HELVELLINEAE)

1. Geoglossaceae. The genus Leotia is known in Ohio.

2. ELelvellaceae. Genera known in Ohio are Helvella, Morchella,
and Gyromitra.

VI. TUBERAEES (TUBERINEAE)

No members of this order have been reported from Ohio.

VII. PHACIDIALES (PHACIDIINEAE)

1. Stictidiaceae. Genera known in Ohio are Stictis, Propolis,
Mellittiosporium, Schizoxylon, and Lichen opsis.

Classification of Ohio Ascomycetes

23

2. Tryblidiaceae. Genera known in Ohio are Tryblidium and
Scleroderris.

3. Phacidiaceae. Genera known in Ohio are Phacidium, Coc-
comyces, and Rhytisma.

VIII. HYSTERIALES (HYSTERIINEAE)

1. Hypodermataceae. Genera known in Ohio are Angelinia and
Lop hod ermiii m.

2. Dichaenaceae. The genus Dichaena is known in Ohio. Some
of the species may be lichens.

3. Hysteriaceae. Genera known in Ohio are Hysterium, Hystero-
g rap Ilium, and G Ionium.

4. Acrospermaceae. The genus Acrospermum is known in Ohio.

5. Graphidaceae. Genera known in Ohio are Graphis and
Opegrapha.

6. Arthoniaceae. Genera known in Ohio are Arthonia and
Arthothelium.

IX. HYPOCREALES (part of the PRYENOMYCETINEAE)

1. Hypocreaceae. Genera known in Ohio are Hypocrea, Hypo-
crella, Hypocreopsis, Hypomyces, Melanospora, Letendraea,
Nectria, Pleonectria, Ophionectria, Cordyceps, Claviceps, Ty-
phodium, and Gibberella.

X. DOTHIDIALES (part of the PYRENOMYCETINEAE)

1. Dothidiaceae. Genera known in Ohio are Dothidea, Phylla-
chora, and Plowrightia.

XL SPHAERIALES (part of the PYRENOMYCETINEAE)

1. Chaetomiaceae. The genus Chaetomium is known in Ohio.

2. SoRDARiACEAE. Genera known in Ohio are Sordaria and
Sporormia.

3. Sphaeriaceae. Genera known in Ohio are Sphaeria, Lasios-
phaeria, Trichosphaeria, Herpotrichia, Bombardia, Acanthos-
tigma, Rosellinia, Bertia, and Zignoella.

24

Ohio Biological Survey

4. Ceratostomaceae. Genera known in Ohio are Ceratostoma and
Ophioceras.

5. CucuRBiTARiACEAE. Genera known in Ohio are Cucurbitaria,
Nitschkia, and Coelosphacria.

6. Amphispiiaeriaceae. Genera known in Ohio are Ohleria,
Trcmatosphaeria, and Caryospora.

7. Mycospiiaerellaceae. Genera known in Ohio are Mycos-
phaerella and Giiignardia.

8. Pleosporaceae. Genera known in Ohio 3.re, Pleospora, Ventiiria,.
Didymella, Leptospracria, Ophiobohis, and Pyrenophora.

9. Gnomoniaceae. Genera known in Ohio are Gnomonia, Gnomo-
niclla, and Phomatospora.

10. Valsaceae. Genera known in Ohio are Valsa, Anthostoma,.
Kalmusiay Diaporthe, Eiitypa, and Eutypella.

11. Melanchonidaceae. Genera known in Ohio are Valsaria,
Galospora, and Pseudovalsa.

12. Diatrypaceae. Genera known in Ohio are Diatrype, Diatry-
pella, and Qiiaternaria,

13. Melogrammataceae. The genus Botryosphacria is known in
Ohio.

14. Xylariaceae. Genera known in Ohio are Xylaria, Nummidaria,
Ustidina, Hypoxylon, Daldinia, and Poronia.

XII. PYRENULALES (PYRENULINEAE)

1. Verrucariaceae. Genera known in Ohio' are V errucaria and
Staurothele.

2. Pyrenulaceae. Genera known in Ohio are Pyremda, Arthopy-
rcnia, and Leptorhaphis.

3. Dermatocarpaceae. Genera known in Ohio are Dermatocarpon
and Endocai'pon.

4. Trypetheliaceae. The genus Trypethclium is known in Ohio..

5. Mycoporaceae. The genus Mycoporuni is known in Ohio.

Classification of Ohio Ascomycetes

25

XIII. PERISPORIALES (part of the PYRENOMYCETINEAE)

1. Erysibaceae. Genera known in Ohio are Erysibe, Sphaerotheca,
Podosphaera, Microsphaera, Uncinula, and Phyllactinia.

2. Perisporiaceae. Genera known in Ohio are Dimerosporium^
Scorias, Antennaria, and Apiosporium.

3. Microthyriaceae. The genus Asterina is known in Ohio.

XIV. ASPERGILLALES (ASPERGILLINEAE)

1. Gymnoascaceae. The genus Gymnoascus must occur in Ohio.

2. Aspergillaceae. Genera known in Ohio are Aspergillus and
Penicillium.

XV. EXOASCALES(EXOASCINEAE)

1. Exoascaceae. Genera known in Ohio are Exoascus and Taphria.

XVI. SACCHAROMYCETALES (SACCHAROYMCETINEAE)

1. Saccharomycetaceae. The genus Saccharomyces is known in
Ohio.

LIST OP PAPERS CONSULTED

1. Bachman, Freda M. A new type of spermagonium and fertilization in

CoUema- Ann. Bot. 24: 747-760. pi. 49. 1912.

2. Bachman, Freda M. The origin and development of the apothecium in

Collema pulposum (Bern.). Ach. Archiv. Zellforsch. jo: 369-430. pi.
30-36. 1913.

3. Bary, A. de. Recherches sur le developpement de quelques champignons

parasites. Ann. Sci. Nat. IV 20’ 5-148. pi. 1-13. 1863.

4. Baur, Erwin. Zur Frage nach der Sexualitat der Collemaceen. Ber.

Deutsch. Bot. Ges. i6’- 363-367. pi. 23. 1898.

5. Baur, Erwin. Untersuchimgen iiber die Entwickelungsgeschichte der

Flechtenapothecien. Bot. Zeit. 62: 21-44. f. 1. pi. 1-2. 1904.

6. Bessey, C. E. A synopsis of the plant phyla. Univ. Nebraska Stud.

1-99. 1907.

7. Bessey, C. E. Outlines of plant phyla. 1-20. Univ. Nebraska, 1909.
See also the second, third, and fourth editions; 1911, 1912, and 1914.

8. Bessey, C. E. Revisions of some plant phyla. Univ. Nebraska Stud..

14: 37-73. 1914.

26

Ohio Biological Survey

9. Bessey, E. A. Some suggestions as to the phytogeny of ascomycetes.
Myc. Centralb. 5: 149-153. 1913.

10. Blackman, V. H. and Weisford, E. J. The development of the perithe-

cium of Polystigma riibrum DC. Ann. Bot. 26’’ 761-767. pi. 70-71. 1912.

11. Bonnier, G. Recherches sur la synthese des lichens. Ann. Sci. Nat.

VII. 9: 1-32. f. 1-6. pi. 1-5. 1889.

12. Bornet, E. Recherches sur les gonidies des lichens. Ann. Sci. Nat. V.

jy: 45-110. pi. 6-16. 1873.

13. Brooks, F. T. The development of Gnomonia erythrostoma Pers. The

cherry-leaf scorch disease. Ann. Bot. 24: 585-605. pi. 48-49. 1910.

14. Brown, H. B. Studies in the development of Xylaria. Ann. Myc. ii:

1-13. pi. 1-2. 1913.

15. Bucholtz, F. Beitrage zur Kenntnis der Gattung Endogone Link. Beih.

Bot. Centralb. 29^: 147-225. pi. 3-10. 1912. '

16. Claussen, P. Zur Entwicklungsgeschichte der Ascomyceten. Pyronema

conduens- Zeits. Bot. 4'. 1-64. f. 1-13. pi. 1-6. 1912.

17. Clements, F. E. The polyphyletic disposition of lichens. Am. Nat. 51:

277-284. 1897.

18. Clements, F. E. The genera of fungi. 1-227. Minneapolis, The H. W,
Wilson Co., 1909.

19. Cutting, E. M. On the sexuality and development of the ascocarp in
Ascophanus carneus Pers. Ann. Bot. 23' 399-417. pi. 28. 1909.

20. Danilov, A, N. Ueher das gegenseitige Verhaltnis zwischen den Goni-

dien und den Pilzkomponenten der Flechtensymbiose. I. Morphologische
Daten iiber das gegenseitige Verhaltnis der Pilzhyphen und Chlorokok-
ken bei heteromeren Flechten. Bull. Jard. Imp. Bot. St, Petersb. jo:
33-70. f. 1-9. pi. 1-3. 1910.

21. Darbishire, .0. V. Ueher die Apothecienentwickelung der Flechte

Physcia pulverulenta (Schreb.) Nyl. Jahrb. Wiss. Bot. 34: 329-345. pi.
11. 1900.

22. Dawson, Maria. On the biology of Poronia punctata (L.) Ann. Bot. 14:
245-262. pi. 14-15. 1900.

23. Dodge, B. O. Methods of culture and the morphology of the archicarp

in certain species of the Ascoholaceae. Bull. Torr. Bot. Club 39: 139-197.
f. 1-2. pi. 10-15. 1912.

24. Dodge, B. O. The morphological relationships of the Florideae and the

ascomycetes. Bull. Torr. Bot. Club 41: 157-202, f. 1-13. 1914.

Classification of Ohio Ascomycetes

27

25. Elenkin, A. Zur Frage der Theorie des Endosaprophytismus bei Plech-

ten. Bull. Soc. Imp. Nat. Moscou II. i8‘. 164-186. 1904.

26. Faull, J. H. The cytology of the Laboulbeniales. Ann. Bot. ^-5: 649-654.
1911.

27. Faull, J. H. The cytology of Laboulbenia chaefophora and L. gyrinidarum-

Ann. Bot. 26’> 325-355. pi. 37-40. 1912.

28. Fink, Bruce. The lichens of Minnesota. Cont, U. S. Nat. Herb. 14:
I-XVII. 1-269. Frontispiece, f. 1-18. pi. 1-51. 1910.

29. Fink, Bruce. The treatment of lichens in “The genera of fungi.” The

Bryologist 15: 180-183. 1910.

30. Fink, Bruce. The nature and classification of lichens. — I. Views and
arguments of botanists concerning classification. Mycologia 5: 231-269.
1911.

31. Fink, Bruce. The nature and classification of lichens. — II. The lichen

and its algal host. Mycologia 5: 97-166. 1913.

32. Fisch, Carl. Beitrage zur Entwickelungsgeschichte einiger Ascomyce-
ten. Bot. Zeit. 40' 851-870. 875-897. 899-906. pi. 10-11. 1882.

33. Frank, A. B. Ueber die biologischen Verhaltniss des Thallus einiger

Krustenfiechten. Cohn. Beitr. Biol. Pfianz. 2'. 123-200. pi. 7. 1877.

34. Frank, A. B. Ueber einige neue und weniger bekannte Pfianzenkrank-

heiten. Ber. Deutsch. Bot. Ges. j: 58-63. 1883.

35. Frank, A. B. Ueber Gnomonia erythrostoma, die Ursache einer jetzt

herrschenden Blattkrankheit der Susskirschen im Altenlande, nebst
Bemerkungen fiber Infection bei blattbewohnenden Ascomyceten der
Baume fiberhaupt. Ber. Deutsch. Bot. Ges. 4: 200-205. 1886.

36. Fraser, H. C. I. On the sexuality and the development of the ascocarp

in Lachnea stercorea- Ann. Bot. 21’- 349-360. pi. 29-30. 1907.

37. Funfstuck, M. Beitrage zur Entwickelungsgeschichte der Lichenen.

Jahrb. Konigl. Bot. Gard. Berlin 5: 155-174. pi. 3-5. 1884.

38. Funfstuck, M. Thallusbildung an den Apothecien von Peltidea aph-

thosa (L.) Ach. Ber. Deutsch. Bot. Ges. 2: 447-452. pi. 11. 1884.

39. Harper, R. A. Sexual reproduction in Pyronema conduens, and the

morphology of the ascocarp. Ann. Bot. 14: 321-400. pi. 19-21. 1900.

40. Marti g, R. Der Eichenwurzeltodter, Rosellinia iRhizoctonia) quercina.

Unters. Forstbot. Inst. Mfinchen j: 1-32. pi. 1-2. 1880.

41. Higgins, B. B. Life history of a new species of Sphaerella- Myc. Cen-

tralbl, 4'. 187-193. f. 1-2. 1914.

28

Ohio Biological Survey

42. Krabbe, G. Morpliologie imd Entwicklungsgeschichte der Cladoniaceen,

Ber. Deutsch. Bot. Ges. i: 64-77. 1883.

43. Krabbe, G. Entwicklungsgeschichte und Morphologie der polymorphen
Flechtengattimg Cladonia- 1-166. pi. 1-12. Leipzig, 1891.

44. Lindau, G. tjber die Anlage und Entwicklung einiger Flechtenapothe-

cien. Flora yi'. 451-489. pi. 10. 1888.

45. Lindau, G. Beitrage zur Kenntniss der Gattung GyropJiora- Festschrift
Schwendener. 19-36. pi. 2. Berlin, Gebriider Borntraeger. 1899.

46. McClatchie, A. M. Seedless plants of Southern California. Protophy-

tes-Pteridophytes. Proc. Southern Calif. Acad. Sci. j: 336-398. 1897.

47. Mdller, A. tjber die Cultur flechtenbildener Ascomyceten ohne Algen.

Unters. Bot. Ins. Konigl. Akad. Munster. 1887 : 1-52. 1887.

48. Neubner, E. Untersuchungen iiber den Thallus und die Fruchtanfange

der Calycien. Wissenschaft. Beilage Jahresb. Konigl. Gymnasium Paulen
5: 1-12. pi. 1. 1893.

49. Nichols, Mary A. The morphology and development of certain pyreno-

mycetous fungi. Bot. Gaz. 22: 301-328. pi. 14-16. 1896.

50. Nienburg, W. Beitrage zur Entwicklungsgeschichte einiger Flechten-

apothecien. Flora g8‘- 1-40. f. 1-3. pi. 1-7. 1908.

51. Nienburg, W. Zur Entwicklungsgeschichte von Polystigma riibrum DC.

Zeits. Bot. 5: 369-400. f. 1-17. 1914.

52. Peirce, G. J. The nature of the association of alga and fungus in lichens.

Proc. Calif. Acad. Sci. Bot. III. j: 207-240. pi. 41. 1899.

53. Reinke, J. Abhandlungen iiber Flechten. — II. Die Stellung der Flechten

im Pflanzensystem. Jahrb. Wiss. Bott. 26’ 524-542. 1894. — V. Das nat-

iirliche Flechtensystem. 2g: 171-236. f. 196-209. 1896.

54. Schaffner, J. H. The classification of plants, VIII. The Ohio Naturalist,

jj: 70-78. 1913.

55. Schneider, A. Reinke’s discussions of lichenology. Bull. Torr. Bot.

Club 23-. 439-448. 1896. 24'. 32-37. 237-243. 1897.

56. Schulte, F. Zur Anatomie der Flechtengattung Usnca- Beih. Bot. Cent.

13: 2-22. f. 1-8. pi. 1-3. 1905.

57. Schwendener, S. tjber die Beziehungen zwischen Algen und Flechten-

gonidien. Bot. Zeit. 26' 289-292. 1868.

58. Seynes, J. de. Recherches pour servir a 1’ historie naturelle des vege-
taux inferieurs. 1-85. f. 1-3. pi. 1-3. Paris, G. Massou, 1886.

Classification of Ofiio Ascomycetes

29

59. Stahl, E. iBeitrage zur Entwickelungsgeschichte der Flechten. — I, Ueber
die geschlechtliche Fortpflanzimg der Eollemaceen. 1-55. pi. 1-4. Leip¬
zig, Arthur Felix, 1877.

60. Sturgis, W. C. On the carpologic structure and development of the

Collemaceae and allied groups. Proc. Am. Acad, Arts Sci. 25'^ 15-52.
pi. 1-8. 1890.

61. Thaxter, R. Contribution towards a monograph of the Laboulbeniaceae-

Mem. Am. Acad. II. 12: 195-429. pi. 1-26. 1896. Part II. Mem. Am.

Acad. II. ij: 217-469. pi. 28-61. 1908.

62. Tobler, F. Zur Biologie von Flechten und Flechtenpilzen. I. Jahrb.

Wiss. Bot. 4g\ 389-409. f. 1. pi. 3. 1911.

63. Wahiberg, A. tjber die Apothecienentwickelung bei einiger Flechten

der Gattungen Anaptychia und Physcia. Ofvers. Finska Vet. Soc. For-
handl. 44’. 91-120. pi. 1. 1901.

64. Webber, H. J. Catalogue of the flora of Nebraska. Rep. Neb. State

Board Agr. for 1889. i8go'- 37-162. 1890.

65. Webber, H. J. Appendix to the catalogue of the flora of Nebraska.

Trans. Acad. iSci. St. Louis 6: 1-47. 1892.

66. Wolff, Gertrude P. Beitrage zur Entwicklungsgeschichte der Flechten-

apothecien. Flora 95: 31-57. f. 1-22. 1905.

67. Zukal, H. Halbflechten. Flora 74: 92-107. pi. 3. 1891.

30

Ohio Biological Survey

EXPLANATION OF PLATES I AND II

The figures are intended primarily to illustrate some types of female reproduc¬
tive tracts in ascomycetes. The magnification is given wherever this datum was

available.

PLATE I

Fig. 1. Three archicarps of Pyronema confluens with spheroidal, one-celled
ascogonia surmounted by one-celled trichog>nes. Each trichogyne tip is in contact
with an antheridium. From Harper.

Fig. 2. A transverse section of an archicarp of Lachnea stercorea. The one-
celled ascogonium is at the left and passes into the several-celled trichogyne, which
extends to an antheridium at the right above, x 600. From Miss Fraser.

Fig. 3. A section of a leaf showing the coiled archicarp of Polystigma rubrum
and the vegetative hyphae which pass through the stoma, x. 900. From Blackman
and Welsford. See also the discussion of Nienburg’s results on this plant.

Fig. 4. A reconstruction from sections of Laboulbenia chaetophora to show
the complete outer wall of the perithecium, one cell of the inner wall, the enclosed
ascogonium surmounted by the trichophoric cell, and the terminal, branched tricho¬
gyne. X 450. From Faull.

Ohio Biological Survey

VoL. II — Plate I

Bruce Fink,

31

32

Ohio Biological Survey

PLATE II.

Fig. 5. The many-celled archicarp of Collema micro pJiyllum. The coiled
ascogonium passes into the erect trichogyne, which in turn extends above the sur¬
face of the algal-host colony, x 300. From Stahl.

Fig. 6. The many-celled archicarp of Miss Bachman’s Collema-Wko. plant.
The coiled ascogonium is at the left, and the trichogyne extends to a group of in¬
ternal spermatia (antheridia ?) at the right, x 520. From Miss Bachman.

Fig. 7. A mature archicarp of Ascoboliis carbonarius : a, a conidium which
gives rise to an archicarp ; b, the conidiophore ; d to g, the stalk of the archicarp ; g
to j, the ascogonial coil; h, probably the ascogonium proper; k to m, the trichogyne;
m, the antheridial conidum ; n, the conidiophore. x 400. From Dodge.

Fig. 8. The archicarp of Peltigera canina. One of four cells of the ascogonium
is shown at A, and a cell of the vegetative hypha from which the archicarp arose is
shown at s. There is supposed to be no trichogyne. x 750. From Funfstuck.

Ohio Biological Survey

VoL. II — Plate II

Bruce Fink.

ilKRARY

UNIVErSITV Of SLUN0I3
. URBANA

THE ASCOMYCETES OF OHIO.— II

I

The Collemaceae*

BRUCE FINK AND C. AUDREY RICHARDS

General Considerations

Hitherto comparatively few descriptions of lichens as fungi have been
undertaken. In many short diagnoses of lichens, mention of the algal
hosts has been omitted, and many lichens have been treated thus in¬
advertently as fungi. But in more lengthy diagnoses or descriptions, the
common practice has been to consider the algal host as part of the lichen.
In order to dispose of typical lichens as fungi, no greater departure from
the ordinary methods is required than to omit from the descriptions all
reference to the algal host, the thing which is now often done through
inadvertency. However, in order to apply to a few families of lichens
the methods commonly employed in the taxonomic disposition of fungi, a
method must be followed which diverges considerably from that which
has been employed by lichenists.

If it should be found impracticable to treat any group of ascomy-
cetous lichens as fungi, the whole plan of treatment of ascomycetes
proposed in the first paper of this series would be impossible of execution.
Consequently, we have considered, in this second contribution toward a
knowledge of the ascomycetes of Ohio, a family of lichens, the treatment
of which as fungi probably involves as wide a departure from the
method commonly employed by students of lichens as any group that
could be chosen. In order to demonstrate as early as possible in this
series of papers how slight are the changes required in order to treat the
large majority of lichens as fungi, the Lecideaceae, another family of
lichens, will receive consideration in the next paper of this series.

The study of the Collemaceae has been made classic through the
contributions of several botanists. De Bary studied this group and
ascertained that the chlorophyllous cells are algal (3). Schwendener
soon extended this view to include the chlorophyllous cells found in all

* Contributions from the Botanical Laboratory of Miami University — XII.

35

36

(3iii() Ijiologjcal Survey

lichens (6). Stahl chose members of the same family for the first im¬
portant work on sexual reproduction in lichens (7). Sturgis (8) and
Baur (4) have since confirmed Stahl's results. Miss Bachman (1 and 2)
has found in a member of the Collcinaccac an interesting condition with
respect to the morphology and the behavior of the sexual reproductive
tracts and has obtained in the same plant the only important cytological
results thus far secured in the study of the sexual organs of lichens. The
results obtained by these workers are known to many botanists, have been
considered in part in the first paper of this series, and need no further
discussion here.

In working out a new taxonomic method, there is a distinct ad¬
vantage in considering a group of plants in which many botanists are
interested. However, our prime motive in selecting the Collemaccac was,
as explained above, to attack first a family of lichens, the treatment of
which as fungi probably presents as many difficulties as any that could
be selected. The species of this family grow mainly within the algal
hosts instead of containing these hosts in a limited area within somatic
tracts of the parasites as do the great majority of lichens; and the de¬
scriptions of gross characters hitherto given as morphological features
of members of the Collemaccac belong largely to the algal hosts rather
than to the lichens themselves. Finding a satisfactory way to obviate
this difficulty and devising a new method for the study of the minute
morphology of these lichens have required spending much time and energy
on details, some of which have proved to be of little if any taxonomic
.value and have not been included in the results presented in this paper.
One year was given to working out a method, which, in the last two years,
has been further elal)orated in the studies of the Collemaccac of Ohio
presented herein.

In connection with many difficulties encountered, it is fortunate that
the tissues of members of the Collemaccac appear more distinctly in
sections than do those of the majority of lichens. This condition has
made it comparatively easy to study most of the details with respect to
structure of cortices, medullse, sexual reproductive tracts, and apothecia.
In a few instances, we have resorted to imbedding and serial sectioning ;
but most of our results have l^een obtained by free-hand sectioning with
the razor. Flemming’s triple stain, Delafield’s haematoxylin, and safranin
..were used with good results; but we depended mainly on the solution
of iodine which is used commonly in the study of lichens. Other
reagents used were water, potassium hydrate, Schulze's macerating fluid,
and alcohol. For macerating, potassium hydrate was found more satis-

The Collemaceae of Ohio 37

factory than Schulze’s solution. The measurements of microscopic
features were taken from specimens treated with water only.

Limiting the area studied to a region near at hand gave the ad¬
vantage of being able to secure material in fresh condition easily. While
our area conforms to the requirements of the Ohio Biological Survey,
the collecting has been conhned to the southwestern portion of the state,
extending roughly about one-fourth of the distance from the southern
boundary to the northern and somewhat more than one-third of the way
from the western boundary to the eastern. The principal collecting
grounds are about Oxford, Eaton, West Alexandria, Mason, Washington
Court House, Greenheld, Paint, Bainbridge, Peebles, and Cincinnati.
Collections made by others in various portions of the state have been
studied through material loaned from the herbarium of the Ohio State
University and have given a fairly satisfactory knowledge of the dis¬
tribution of several of the species within the state as a whole.

Just what changes in limitations of species would result from the
employment of a new taxonomic method could not be known until we
had examined a considerable number of species of the family. While
our paper covers only the sixteen species of the Collcinaccac known in
Ohio, we have made preliminary examinations of about as many more.
This means that we have covered approximately one-fourth of the
species known to occur in North America. Thus far our method has not
required any changes in species-limits. Each species studied has shown
a sufficient number of morphological characters to mark it as distinct,
or has been found to modify the size, form, color, or general appearance
of the algal host in a peculiar manner. In our descriptions, the power
of modifying the algal-host colony has been treated as a specific,
physiological character of each species of lichen. Since the algal host
rather than any portion of the lichen itself, except the apothecia, is con¬
spicuously visible in the field, we have given first under each name of a
species a statement concerning the manner in which the lichen has
modified its host. This brings into prominence that which is seen plainly
on first observation, and at the same time avoids the error of describing
the algal host as a portion of the lichen. Eollowing the statement re¬
garding the transforming action of the lichen on the algal host, such
characters of the lichen itself as have seemed to have considerable
diagnostic value have been recorded.

We have tried to present a method workable for the species found
in the area treated in this paper and also like that which must be em¬
ployed when our method is extended to include the Collcinaccac of a

38

Ohio Biological Survey

larger area, or finally to the family as a whole. It is evident that the
taxonomic value of the diagnostic characters used, both morphological
and physiological, may be lietter known after a large number of species
has been studied carefully, but it is not probable that extension of the
method given herein will involve other than minor modifications.

In deciding on a method of treatment of the algal host, other
ascomycetous lichens which will require treatment similar to that which
is given to the Collemaceac in this paper were kept in mind. Other
nosticolous lichens may be treated in essentially the same manner as the
Collemaceac. Other types are illustrated by Epliebc, the thallus of which
is similar to that of the Collemaccce, but is found penetrating throughout
the tissue of the branched, filamentous alga, Sirosiplion, and by
Coenogonium, which has a mycelial thallus that covers a portion of the
filaments of the algal host, Trentcpohlia. From a purely technical point
of view, it would be better, in treating the Collemaceac and all other
lichens which require special methods for disposition of the modified
algal host, to treat the host in a paragraph following the description and
dealing with habitat and distribution. However, the practical advantage
of placing immediately under the name of the species that which is first
noticed in field or laboratory is so great that all who use our descriptions
will doubtless agree that we are justified in departing from a more exact
scientific procedure sufficiently to place the statement about the trans¬
formation of the algal host before the description of the lichen, which,
excepting its apothecia, is known only through microscopic study.

The penetration of the thallus of members of the Collemaceac
throughout the algal-host colony as a distinct mycelial structure unat¬
tached to the algal cells (Fig. 1) renders the somatic areas so plainly
visible that even the sexual reproductive areas, imbedded in the vegetative
structures, are much more readily visible in section than those of most
other lichens. In beginning our work, it was thought that these re¬
productive structures, so plainly visible, might be available in this family
of lichens for diagnostic purposes. The archicarps (Fig. 18) and the
spermagonia (Fig. 25) were carefully described for each species of
Synechoblastus studied, and later those of a majority of the species of
our other genera were more or less carefully examined, even though the
work on Synechoblastus indicated that these organs do not dififer enough
in the various species examined to make them of diagnostic value, except
in a few instances. Added to this is the difficulty, sometimes very great,
of finding specimens that bear the sex organs, and the areas of the thallus
in which these organs occur.

The Collemaceae of Ohio

39

The sex organs grow, for most part, during two months in the spring
and for the same length of time in autumn. These organs may be found
in specimens not collected during these periods ; but the cells are likely
to be dead, and the parts of the archicarps collected out of season are
not likely to have quite their natural form. The spermagonia likewise
must be studied at the right time, since the spermatia decrease in num¬
bers and finally disappear entirely after the prime condition is past, while
the basidia become considerably enlarged. The spermagonia usually ap¬
pear as minute protuberances, which are commonly some shade of yellow
or brown. They occur most frequently on the lobes and near the
margins, or at least externally to the areas occupied by mature apothecia.
They may be seen easily with a hand lens, especially after the lichen and
the algal host are moistened ; but they are not easily distinguished, in
gross morphology, from young apothecia, which have not yet ruptured
the mass of structures which lie above them. There is no way of de¬
tecting the presence of archicarps through external appearance, but they
usually occur toward the margins from the position occupied by the
scattered apothecia. Sometimes, however, the archicarps are found
within the region of the thallus occupied by the apothecia, and they rarely
occur well toward the center of the thallus. The more central ones are
likely tO' be past prime and more or less distorted.

Owing to the uncertainty of finding the sex organs and their slight
or infrequent taxonomic value, we have, after working them out labor¬
iously, decided not to burden the diagnosis with matter of so little value
in determining species. Instead, we have given the results from studies
of these organs toward the close of the paper where they may be acces¬
sible to those who may wish to consult them. Though the sexual tracts
are not of sufficient diagnostic worth to warrant lengthening the diagnoses
considerably in order to include them, it is true that critical workers on
the Collemaceae must hereafter take these structures into account. Miss
Bachman’s results (1 and 2) and our own findings from several regions
widely separated (Figs. 14 and 20) prove that forms so much alike with
respect to somatic characters and their method of modifying the algal-
host colonies may differ so much with respect to the sexual tracts that
they can scarcely belong to the same genera (Pis. Ill and IV). This
means that critical workers on the Collemaceae must know at least the
general nature of the reproductive tracts in every specimen in which they
can be found in order to be reasonably certain that forms so much alike
otherwise do not differ greatly with respect to the sex organs and their
behavior. Some of the minor differences in size and form of the sexual

40

Ohio Biological Survey

tracts within the family may be seen by consulting our hgures (Pis. Ill
and IV), and the measurements given in the descriptions of the sex
organs. Our work on these areas in Syneclioblastus illustrates the
rather slight dififerences within the species studied from this genus. The
differences in size of ordinary spermatia and the internal male repro¬
ductive cells found in certain of the Collemaccae may be seen by com¬
paring Figs. 14, 20, 21, 22, and 23.

One of the writers of the present paper had Miss Bachman’s material
for examination during the progress of her studies. It seemed certain
enough that she was working on Collema pulposuni until the peculiarities
with respect to the morphology and the behavior of the sex organs were
discovered. It then became apparent that her plants could not be
Collema pulposiim, if those previously studied by Stahl (7) and
Sturgis (8) are this species. Unfortunately, the reproductive areas can
be studied successfully from herbarium material so seldom that it is
doubtful whether microscopic study of the type of the genus Collema
and the type specimen of Collema pulposum would help to decide which
plants are like the types with respect to the sexual tracts. To attempt to
treat in this paper the material with internal sex organs found in Ohio
would be premature. One of the writers will continue with taxonomic
studies of the material already examined by us from Wisconsin, Iowa,
Alinnesota, Ohio, Missouri, and New York, with any other that we may
discover, while the other author pursues further cytologic investigations.

The thalli of members of the Collemaceae are composed entirely of
hyphae. The hyphae are commonly long-celled and loosely interwoven in
such fashion as to form a mycelial structure, which usually composes
the larger portion of the somatic areas. In the hypothecia and the
exciples, the hyphie are closely interwoven. In these areas, the cells of
the hyphae may be long, or they may be approximately isodiametrical and
so closely placed as to exclude interhyphal spaces. In Leptogiuin and
Mallotium there is a distinct cortex, usually one layer of cells in thick¬
ness (Fig, 1 ), composed of a layer of roughly isodiametric end-cells of
hyphae which extend from the dorsal to the ventral surface. Lindau (5)
has proposed the term plectenchyma for the densely interwoven
parenchymatoid hyphal tissues of fungi and has distinguished two kinds,
the paraplectenchyma with cells roughly isodiametrical and the proso-
plectenchyma with the cells more or less elongated and the ends narrowed.
The paraplectenchyma is found in the cortices in certain genera of the
Collemaceae and in the exciples and the hypothecia of certain species.
The prosoplectenchyma is rarely seen in the plants which we have studied.

The Collemaceae of Ohio

41

and we have not attempted to differentiate. Loosely interwoven hyphse
with much elongated cells we have designated mycelial structure. The
mycelial structure is probably present in the medullse of all somatic
tracts within the group, though a plectenchymatoid appearance is some¬
times seen in the medulla in instances in which the hyphse are so densely
interwoven and so interspersed with algal-host cells as to obscure the
structure. The hypothecia and the exciples may be composed of ordinary
long-celled, but closely interwoven hyphse (Fig. 9), or they may be
plectenchymatous (Fig. 10). The hyphse of the medullse extend in various
directions (Fig. 1), and the number that lie perpendicular to the surfaces
of the thallus is usually largest about the apothecia and toward the
margins of the thalli. This distribution gives strength to the thalli.

Perhaps no part of our work presented greater difficulty than de¬
termining whether certain hypothecia, exciples, and medullse, or certain
portions of these structures, are plectenchymatous or whether they are
composed of closely interwoven hyphse which have elongated cells.
Transitional conditions occur, and one may easily mistake interwoven
hyphse with elongated cells for a plectenchyma when the hyphse are
closely packed. We found a plectenchymatoid appearance in the medullse
of certain specimens of Leptogium tremelloides (Fig. 2) and Leptogium
cliloromelum, and even in the same section, certain portions may show a
plainly mycelial structure while others, in which the hyphse are densely
packed, may appear like a plectenchyma. Careful maceration failed to
show a true plectenchyma in any instance in our specimens of these two
species. Other species are said by certain workers to be plectenchy¬
matous throughout, but it may well be doubted whether this is true. We
examined Leptogium tcnuissinium, which is figured in Engler and Prantl
as plectenchymatous throughout, but found a plainly but densely mycelial
structure toward the lower surface. Leptogium plectenchymiim, de¬
scribed in this paper, varies greatly with respect to position and amount
of plectenchyma, but we found no specimens which were wholly
plectenchymatous in the vegetative areas. Certain hypothecia and
exciples present greater difficulty than any of the medullje that we have
examined, and only the use of high powers, the oil-immersion lens, and
careful maceration will solve the problem in many instances. We do
not wish to leave the impression that such obstacles arise in the study
of all the species of the Collemaceae. On the other hand, the typical
plectenchyma (Fig. 3) and the typical mycelial structure (Fig. 1) are
readily recognizable.

42

Ohio Biological Survey

The apothecia of all of the Collemaceae studied possess a more
or less strongly developed exciple. Surrounding the exciple and often
extending above it is a thalloid margin, composed of elements of the
lichen thallus, with which algal-host filaments are intermingled. The
thalloid margin is sometimes absent or overgrown, when the exciple is
naked as in many other ascomycetes. In some of the species, one must
resort to the oil-immersion lens to demonstrate the exciple. This struc¬
ture extends below into the hypothecium, which is much thicker as a
rule and is always easily demonstrable. It needs to be made clear that
the thalloid margin is not a part of the apothecium, but belongs rather
to the somatic tract of the lichen, and is intermingled with elements of
the algal-host colony, which, in this family, obscure the real thalloid
margin.

The Collemaceae form a very distinct group of ascomycetes, especially
peculiar with respect to conditions of parasitism and resulting thallus
disposition and structure. In the lower members of the family, the
thallus is a mycelium pure and simple as found in Syneclioblastiis and
Collema. In certain other genera as Leptogium and Mallotiiini, the
mycelial structure gives way in part to a cortical plectenchyma. In rare
instances, the plectenchyma extends throughout a large portion of the
thallus, which, as noted above, is said by some students to be wholly
plectenchymatous in some species. A plectenchyma sometimes occurs in
the hypothecium and in the exciple, even in the lower members of the
family, in which the vegetative areas are wholly mycelial. Hence, it
seems safe to assume that the most primitive plectenchyma is that found
in the fruit rather than that found in the somatic areas. About and
below the apothecia of Lcptogla, the plectenchyma is usually several
layers of cells in thickness (Fig. 3), and it is similarly thickened below
the apothecia in the material of Mallotium at hand for examination.

An interesting series of transitional conditions which indicates how
a cortical plectenchyma may have arisen came to light in our studies of
Synechohlastus Haccidus (Figs. 6, 7, and 8). Short cells at the upper
and the lower ends of erect hyphse often coalesce over limited areas of
the thallus and form either a typical plectenchyma, or a structure strongly
suggestive of a cortical plectenchyma. In some instances, the cells which
coalesce are elongated and form a palisade which dififers considerably
from a true plectenchyma (Fig. 6). Over other areas the short end-
cells are formed but do not cohere (Fig. 7). On account of these
transitional conditions, this species is often placed in the genus Leptogium
in herbaria. In Leptogium chloromclum, the plectenchymatous cortex is

The Collemaceae of Ohio

43

often poorly developed, and in some instances it is discontinuous and
recalls the conditions just noted for Synechoblastiis Haccidus. These
two species bridge fairly well the gap between members of the
Collemaceae which have no cortex and those which have one. Some of
the material of Mallotiiim also showed a poorly developed and scarcely
continuous cortical plectenchyma.

Sweeping statements to the effect that paraphyses are uniformly
simple in many families of ascomycetes are due to the careless manner
in which the examinations are made. We have not studied a species of
the Collemaceae in which branched paraphyses do not occur (Fig. 11).
In instances in which it is difficult to ascertain the structure, maceration
and examination with an oil-immersion lens will always reveal branched
paraphyses, often in large numbers. Though simple paraphyses occur
in all of the species that we have studied, branched ones sometimes appear
to be more numerous than the simple ones when one makes a careful
examination.

The spores of the Collemaceae are hyaline and vary from simple
ones to muriform conditions (Fig. 13). The occurrence of muriform
spores in Collema and simple ones in genera that have a plectenchymatous
cortex makes it probable that there are at least two lines of descent within
the family, one expressed mainly in the spore evolution, and the other
showing the spore evolution and also the development of a cortical
plectenchyma. In separating Synechoblastus, with several-celled spores,
from Collema, which has muriform spores, we are following the tendency
among students of ascomycetes to give more prominence than formerly
to spore characters, in limiting genera. A gradual evolution from simple
spores to several-celled ones and from these to muriform types is well
known in many groups of ascomycetes ; and transitional conditions in
which a species may have either simple or septate spores, or either
several-celled or obscurely muriform ones are occasionally met. There¬
fore, one sometimes meets difficulties in using spore characters as
taxonomic criteria in the Collemaceae and in some other ascomycetes.
However, we have encountered no difficulty in the species treated in this
paper.

The Collemaceae are related, with respect to apothecial and somatic
structure, to the Pannariaceae and tO' the Lecideaceae. The difference in
structure of the thallus, in the Lecideaceae on one side and the Colle¬
maceae and the Pannariaceae on the other, is mainly explainable as a re¬
action to different hostal environment and to inorganic, substratic con¬
ditions. With respect to the sexual tracts, the Collemaceae are closely

44

Ohio Biological Survey

related to many lichens and other ascoinycetes. These relationships are
considered in the previous jiaper of this series.

Eleven genera of Collcmaccac are recognized in Engler and Plantl’s
“Naturlichen Phanzenfainihen.” Of the genera recognized in Engler
and Prantl, only Physma, Leptogium, and Collema are known in North
America. However, there seems to be no sufficient reason for exclud¬
ing the monotypic North American genus, Hydrothyria, from the
Collemaceae. Seven of the foreign genera are monotypic, and the other
one is ditypic. Of approximately 200 species of the Collemaceae, some¬
what more than half occur in North America.

Thirteen species of the Collemaceae had been listed for Ohio when
we began our work three years since. Of these, Synechoblastiis crystas-
pis, Collema crispiim, Leptogium juniperinum and Mallotium myochroum
do not seem to occur in the state, the determinations being faulty or rest¬
ing on insufficient or sterile material. Our studies have added seven
species to the flora of Ohio, two of which are new. The total number of
species now known for the state is sixteen. Besides this, the species with
internal reproductive organs remains to be described. Sets of the material
on which this paper is based may be found in the state herbarium and
in the herbarium of Bruce Fink.

During most of the time that we have worked on the Collemaceae of
Ohio, Miss Leafy J. Corrington has given her forenoons to the work
under our direction, and we are under obligations to her for assistance.

Systematic Account

COLLEMACEAE

Thallus, with rare exceptions, composed wholly or mainly of a
mycelium which penetrates throughout and usually surrounds the algal-
host colony ; hyphse not attached to the cells of the hosts in any of our
species ; the mycelial structure surrounded, in the higher genera, by a
cortical plectenchyma, usually a single layer of cells, except around and
below the apothecia ; attached to the substratum by hyphal rhizoids,
which are usually seen only in sections; female sex organ an archicarp,
differentiated into an ascogonium and a trichogyne ; male sex organ usually
a spermagonium, which contains spermatia, but sometimes an internal
spermatium (antheridium, or antheridial conidium?) ; external conidia
known in one member of the family; apothecia with an exciple, either
plectenchymatous or of interwoven hyphae, a hypothecium, the structure

The Collemaceae of Ohio

45

of which is usually like that of the exciple into which it is produced, and
a hymenium composed of the asci and the paraphyses, the former blue
and the latter yellow-brown in iodine ; spores simple, several-celled, or
muriform, brown in iodine; apothecia usually surrounded by a thalloid
margin.

The algal host is a form of Nostoc and is strongly modified and
transformed into a more or less plainly foliose structure, which has
peculiar features of form, size, or color dependent on the transforming
efifect of the particular lichen parasite. The algal-host colony becomes
gelatinous when wet, and the gelatinization is more marked when the
alga functions as the host of a species of the CoUcinaceae which is devoid
of a cortical plectenchyma.

KEY TO THE OExXERA

Without a typical plectenchymatous cortex.

Spores 2- to several-celled . Synechoblastus

Spores muriform . Collema

With a typical plectenchymatous cortex.

Rhizoids not plainly noticeable with the hand lens . Leptogium

Rhizoids plainly noticeable with the hand lens . Mallotium

Synechoblastus Trev. Caratt. Gen. Collem. 2. 1853.

Transforming the algal-host colony into a blue-green, olivaceous, or
nearly black, irregular or foliose body, with the lobes often ascending or
erect and covering the horizontal portions, often to the center of the
colony ; thallus not differentiated into a typical plectenchymatous cortex
and a mycelia medulla, but the whole structure commonly mycelial and
the hyphse usually more densely disposed toward the surfaces, especially
the upper surface; hyphse hyaline, perpendicular, horizontal, and extend¬
ing in various directions, straight or curved, monopodially branched ;
rhizoids inconspicuous, but commonly seen in sections of some of the
species ; apothecia scattered ; disk flat to convex, red-brown to black-
brown, surrounded by a thalloid margin which is sometimes overgrown ;
exciple either densely mycelial or plectenchymatous, usually tinged with
brown ; hypothecium similar in structure and color ; hymenium hyaline
below to pale- or darker-brown above ; paraphyses hyaline at the base
and gradually becoming tinged with brown toward the enlarged apex,
simple or once to several times monopodially branched toward the a])ex
or farther back, all species examined showing both simple and liranched

46

Ohio Biological Survey

ones, the number of cells ranging from 5 to 20, the length from 4b to
115 mic., and the width from 1.5 to 5 mic. ; asci clavate, the walls 0.5 to
2 mic. thick laterally and 2 to 9 mic. thick at the apex ; spores hyaline,
2- to several-celled, but not muriform, 8 in each ascus and irregularly
arranged in all of ours.

On account of the flexible nature of the thallus, which is devoid of
a strengthening plectenchymatous cortex, the algal host becomes enlarged
and more or less plainly gelatinous when moist.

KEY TO THE SPECIES OF SYNECHOBLASTUS
Spores 2-celled.

Apothecia marginal on the densely aggre¬
gated, erect lobules . 2. Synechoblastus pycnocarpus

Apothecia scattered over the surface . 1. Synechoblastus ohioensis

Spores more than 2-celled.

A rudimentary plectenchymatous cortex

often seen, usually sterile . 5. Synechoblastus flaccidus

Wholly devoid of cortex, usually fertile.

Spores less than 40 mic. long . 4. Synechoblastus ryssoleus

Spores more than 40 mic. long . 3. Synechoblastus nigrescens

1. Synechoblastus ohioensis Fink sp. nov.

Transforming the algal-host colony into a rather small, suborbicular
or irregular, foliose body, which is 2 to 5 cm. across, 225 to 500 mic.
thick, and rather loosely adnate, with ascending lobes, the margins of
which are sometimes closely rugose lobulate, with the upper surface
dark-olivaceous, varying toward black or green, with the lower surface
lighter colored, and with the algal chains more numerous toward the
surfaces, especially the upper; thallus of rather densely disposed hyphse,
which are somewhat uniformly distributed with their course, mainly
perpendicular to the surfaces, but in part irregular, or roughly horizontal
toward the center, 3 to 8 mic. wide ; rhizoids well developed and com¬
monly seen in sections of thalli ; apothecia common, small to middle-
sized, sessile, scattered, 0.5 to 2.5 mm. in diameter; disk flat or convex,
red- tO' dark-brown, surrounded by an entire thalloid margin, which may
extend above it or may rarely become overgrown ; exciple hyaline, of
interwoven hyphse, but plectenchymatoid above ; hypothecium of inter¬
woven hyphse, 80 to 120 mic. thick; hymenium 65 to 105 mic. thick;
paraphyses frequently and variously branched; asci 55 to 65 mic. long,
and 13 to 16 mic. wide; spores ovoid to ellipsoid-pointed, 2-celled, 12
to 24 mic. long and 5 to 8 mic. wide. See Fig. 25.

The Collemaceae of Ohio

47

Found on limestone rocks near Peebles, Adams County, usually
growing with mosses. The plant was collected for Collcma pulposum^
but differs considerably in its effect on the appearance of the algal-host
colony and much more in its anatomical characters. This new species
resembles Synechoblastus pycnocarpus strongly, but it modifies the algal-
host colony in a different manner and has several distinct features of
thallus and fruit. The plant seems to resemble Synechoblastus texanus
(Tuck.), both in its physiological action on the algal-host colony and in
its anatomical characters. Three collections were made near Peebles.

2. Synechoblastus pycnocarpus (Nyl.) Fink Cont. U. S. Nat. Herb.
14^: 134. 1910.

Collema pycnocarpum Nyl. Syn. Lich. i: 115. 1858.

Transforming the algal-host colony into a rather small, suborbicular,
foliose body, which is 1.5 to 6 cm. across and 150 to 875 mic. thick, the
horizontal portions closely adnate and sometimes sparingly perforate,
with the lobes long and more or less branched, rising toward the margin
into a suberect position, the erect or ascending portions densely rugose
lobulate and obscuring more or less the parts below, especially when
covered with apothecia, with the upper surface olivaceous varying toward
green or black, and sometimes granulate, with the lower surface lighter
colored, and with the algal chains distinctly more numerous toward the
surfaces; thallus of hyphse which are densely disposed toward the sur¬
faces but somewhat scattered toward the center, variously placed and in
part perpendicular or horizontal, the latter especially toward the center,
2 to 4 mic. wide; rhizoids seen in large numbers in certain sections of
the thallus ; apothecia small, numerous, sessile, marginal on erect or
ascending lobules, 0.4 to 0.8 mm. in diameter ; disk flat to convex, brown
or red-brown, surrounded by an entire thalloid margin, which does not
extend above the disk and is sometimes overgrown ; exciple of interwoven
hyphae, tinged with brown ; hypothecium of interwoven hyphse, 33 to 85
mic. thick; hymenium 55 to 85 mic. thick; paraphyses frequently and
variously branched ; asci 45 to 62 mic. long and 10 to 14 mic. wide ; spores
ovoid to ellipsoid-pointed, 2-celled, 11 to 20 mic. long and 3.5 to 6.5 mic.
wide. See Figs. 9, 11, 18, and 24.

Widely distributed in the state, but not common. On tree trunks.
Examined from areas about Peebles, Springfield, Amanda, Cedar
Swamp, Paint, Prospect, and Columbus.

48

Ohio Biological Survey

3. Synechoblastus nigrescens (Muds.) Stizenb. Rer. St. Gall. Ges.

Naturw. 1861-1862: 144. 1862.

Lichen nigrescens Huds. FI. Angl. 450. 1762.

Transforming the algal-host colony into a middle-sized, orbicular or
irregular foliose body, which is 2 to 7 cm. across, 60 to 185 mic. thick,
and closely adnate, with the upper surface closely beset with pustules,
on many of which apothecia are situated, or with the pustules largely
replaced by radiating ridges, with short, rounded, entire, and usually
somewhat raised lobes, with the upper surface olive-green and blacken¬
ing, with the lower surface lighter colored and marked by depressions
which correspond with the pustules or the ridges of the upper surface,
and with the algal chains somewhat more numerous toward the upper
surface; thallus of hyphse which are densely disposed, somewhat uni¬
formly distributed with course largely perpendicular or horizontal, and
1 to 2.5 mic. wide; rhizoids rarely seen in the sections; apothecia rather
small, numerous, sessile, crowded toward the center of the thallus, which
is sometimes obscured by them, 0.4 to 1.2 mm. in diameter; disk flat or
convex, brown or red-brown and surrounded by a thin thalloid margin,
which is sometimes overgrown ; exciple plectenchymatous, tinged with
brown ; hypothecium plectenchymatous below, of interwoven hyphse
above, 40 to 65 mic. thick; hymenium 85 to 115 mic. thick; paraphyses
sometimes anastomizing by a bridge; asci 65 to 85 mic. long and 14 to
22 mic. wide; spores 4- to several-celled, long-fusiform to acicular, 40
to 70 mic. long and 3.5 to 6.5 mic. wide. See Figs. 12 and 13.

Known in Ohio only through specimens collected by T. G. Lea near
Cincinnati, in 1841. However, the plant should occur in other portions
of the state. On the trunks of trees.

4. Synechoblastus ryssoleus (Tuck.) Fink Cont. U. S. Nat. Herb. 14^:

135. 1910.

Collema nigrescens ryssoleum Tuck. Lich. Calif. 34. 1866.

Transforming the algal-host colony into a rather small, suborbi-
cular, foliose, membranaceous body, which is 2 to 5 cm. across, 60 to 200
mic. thick, and rather smooth, with rounded, ascending lobes, with pli¬
cate, undulate and crisped margins, with the upper surface olivaceous to
black-brown and rugose-papulose, with the lower surface lighter colored
and reticulate, and with the algal chains somewhat more numerous
toward the surfaces ; thallus of densely disposed hyphge, which are more

The Collemaceae of Ohio

49

numerous toward the surfaces and largely perpendicular, but in part
horizontally or irregularly placed, 2 to 5 mic. wide ; rhizoids seldom seen
in the sections ; apothecia minute to middle-sized, numerous but scattered,
sessile, 0.2 to 1.5 mm. in diameter; disk brown or red-brown, flat or
convex, surrounded by a thin, entire thalloid margin, which is sometimes
overgrown ; exciple of interwoven hyphae tinged with brown ; hypothe-
cium of interwoven hyphae, 40 to 85 mic. thick ; hymenium 85 to 95 mic.
thick; asci 60 to 85 mic. long and 12 to 20 mic. wide; spores long
ellipsoid-pointed, 6-celled, 22 to 34 mic. long and 4 to 6 mic. wide.

A single specimen was collected by W. A. Kellerman, in Delaware
County. The specimen is sterile and without number. On tree
trunks. The microscopic apothecial characters were taken mainly from
Lichenes B oreali-Americani, No. 199.

5. Synechoblastus flaccidus (Ach.) Trev. Caratt. Gen. Collem. 1853.

Lichen haccidiis Ach. Vet. Akad. Handl. i6\ 14. 1795.

Transforming the algal-host colony into a middle-sized, suborbi-
cular or irregular foliose body, which is 3 to 7 cm. across, 40 tO' 185 mic.
thick, and rather loosely attached to the substratum, with rather wide,
rounded, entire, and often imbricated lobes, with the upper surface
olivaceous, varying toward black or green, usually granulate, with the
lower surface commonly lighter colored, and with the algal cells some¬
what more numerous toward the upper surface ; thallus of densely dis¬
posed hyphae, which extend in various directions and are 2.5 to 4 mic.
wide, those perpendicular to the surface of the thallus often passing at
one end or at both ends into short, enlarged cells, or into branches, the
end cells of which are short and wide and sometimes cohere laterally to
form a true plectenchyma over limited areas ; rhizoids rarely seen in the
sections; apothecia rare, small to middle-sized, sessile, 0.5 to 1.5 mm. in
diameter ; disk concave to slightly convex, brown or red-brown, sur¬
rounded by an entire, often granulate thalloid margin, which may extend
above it ; exciple plectenchymatous, tinged with brown ; hypothecium
plectenchymatous below, of interwoven hyphse above, 34 to 88 mic. thick ;
hymenium 85 to 125 mic. thick; asci rather large, 80 to 110 mic. long
and 14 to 21 mic. wide; spores long ellipsoid-pointed, 4- to 6-celled, 28
to 40 mic. long and 5 to 9 mic. wide. See Figs. 6, 7, 8, 13, and 17.

Widely distributed in the state. On trees and rarely on rocks.
Examined from areas about Amanda, Eaton, Oxford, Peebles, Paint, and
Orwell.

50

Ohio Biological Survey

Collegia (Hill.) Web. in Wig. Prim. FI. Hols. 89. 1780.

Transforming the algal-host colony into a squamulose or an irregu¬
larly foliose body, with the upper surface olivaceous, varying toward
green or black and the lower surface somewhat lighter colored ; thallus
wholly mycelial and the hyphse more densely disposed toward the sur¬
faces ; hyphse hyaline, variously and often densely disposed, straight or
curved, monopodially branched ; rhizoids inconspicuous but commonly
seen in sections; apothecia commonly scattered; disk varying from con¬
cave to convex, from light brown to dark brown, surrounded by a thalloid
margin, which is rarely overgrown ; exciple either densely mycelial or
plectenchymatous, usually hyaline ; hypothecium usually of interwoven
hyphse, hyaline below to pale brown above ; hymenium hyaline below to
brown above ; paraphyses hyaline at the base and tinged with brown
toward the somewhat enlarged apex, simple or sometimes monopodially
branched toward the apex or farther back, all species showing both
simple and branched ones, the number of cells varying from 5 to 9, the
length from 55 to 160 mic., the diameter from 1.5 to 4 mic. ; asci clavate,
the walls 9.5 to 2.5 mic. thick laterally and 5 to 13 mic. thick at the apex;
spores hyaline, ellipsoid, muriform, 8 in each ascus, irregularly arranged.

KEY TO THE SPECIES OF COLLEMA

Attached to calcareous rocks.

Spores usually less than 8 mic. wide . 4. CoHema plicatile

Spores usually more than 8 mic. wide . 5. Collema furvum

Not attached to calcareous rocks.

On calcareous soil and on mosses, usually over calcareous rocks.

Thallus and transformed algahhost colony 200 to

750 mic. thick . 1. Collema pulposum

Thallus and algal-host colony less than 100 mic.

thick . 2. Collema tenax

On tree trunks . 3. Collema microphyllum

1. Collema pulposum (Bernh.) Ach. Lich. Univ. 632, pi. 14. f. 9. 1810.

Lichen pulposus Bernh. Journ. Bot. Schrad. pi. 1. f. 1. 1799.

Transforming the algal-host colony into a middle-sized, orbicular or
more or less irregular body, which is 1.5 to 8 cm. across, 200 to 750 mic.
thick, and closely attached to the substratum, with thick, entire or repand-
crenate and often imbricated, rosulate lobes, which are often reduced in
the central portions, to mere granules, with the upper surface olivaceous,

The Collemaceae of Ohio

51

varyiiij^ toward black, with the lower surface scarcely lighter colored,
and with the algal chains more numerous toward the surfaces ; thallus of
scattered or rather densely disposed hyphse with course largely irregular,
but roughly horizontal toward the center, 1 to 4 mic. wide ; rhizoids
numerous at the scattered points of attachment ; apothecia common, small
to middle-sized, adnate or sessile, 0.6 to 2.5 mm. in diameter; disk concave
to slightly convex, red-brown, surrounded by an entire or rarely crenate
thalloid margin, which extends above it, or is rarely overgrown ; exciple
thin, of interwoven hyphse, tinged with brown ; hypothecium of inter¬
woven hyphse, tinged with brown, 40 to 80 mic. thick ; hymenium 90 to
125 mic. thick; asci 70 to 90 mic. long and 11 to 21 mic. wide; spores
longitudinally 4- to 5-celled and transversely 2-celled, 17 to 32 mic. long
and 6 to 11 mic. wide. See Figs. 9, 13, 15, 16, and 21.

Widely distributed in southern Ohio. On calcareous earth, or on
mosses over calcareous rocks. Examined from areas about Oxford,
Cincinnati, Eaton, Fultonham, and Peebles. Not previously reported for
Ohio, but collected by T. G. Lea near Cincinnati, in 1841.

2. Collema tenax (Sw.) Ach. Lich. Suec. 128, 237. 1798.

Lichen tenax Sw. Nov. Act. Soc. Sci. Ups. 4: 249. 1784.

Transforming the algal-host colony into a small to middle-sized,
orbicular or more or less irregular body, which is 0.5 to 7 cm. across,
65 to 85 mic. thick, and closely attached to the substratum, usually
crenately lobed and imbricated, but sometimes depauperate and reduced
to squamules when attached directly to rocks, with the upper surface
olivaceous, varying toward black, with the lower surface scarcely lighter
colored, and with the algal chains numerous and somewhat evenly dis¬
tributed ; thallus of densely disposed hyphae, which are often perpendicu¬
lar to the surfaces and are 2.5 to 4. mic. wide ; rhizoids numerous at the
points of attachment ; apothecia common, small to middle-sized, sessile
or adnate-immersed, 0.5 to 2 mm. in diameter ; disk concave to slightly
convex, red-brown and darker, surrounded by an entire or a crenate
thalloid margin, which does not extend above it ; exciple hyaline, of
interwoven hyphae ; hypothecium of interwoven hyphae, 80 to 95 mic.
thick; hymenium 130 to 160 mic. thick; asci 65 to 75 mic. long and 16
to 20 mic. wide ; spores longitudinally 4- to 5-celled, transversely 2-celled,
16 to 21 mic. long and 7 to 10 mic. wide.

Collected near Paint, by Bruce Fink, No. 460. Usually on calcar¬
eous soil over rocks, often growing with mosses. Ours a depau])erate

52

Ohio Biological Survey

condition on calcareous rocks. Previously recorded for Ohio, but we
have not seen the specimens.

3. Collema microphyllum Ach. Lich. Univ. 630. 1910.

Transforming the algal-host colony into a minute, suborbicular or
irregular body, which is 0.1 to 1 cm. across, 110 to 160 mic. thick, and
closely adnate or sometimes raised at the margin, with the marginal
lobes rounded and those toward the center reduced, granulate-crenate,
and often imbricate, with the upper surface black-olive, and with the
algal chains somewhat more numerous toward the upper surface; thallus
of densely disposed, much branched hyphse, which are uniformly distri¬
buted, extend in various directions, and are 2.6 to 4 mic. wide; rhizoids
hyaline, numerous at certain points, especially under the apothecia ;
apothecia numerous, minute or small, sessile, or adnate, 0.3 to 0.6 mm. in
diameter ; disk concave to flat, red- to dark-brown, surrounded by an
entire thalloid margin which does not extend above it and often becomes
overgrown ; exciple plectenchymatous, hyaline ; hypothecium plectenchy-
matous below and of interwoven hyphae above, 55 to 80 mic. thick;
hymenium 90 to 120 mic. thick; asci 85 to 95 mic. long and 21 to 26
mic. wide; spores longitudinally 4-celled and transversely 2-celled, 18
to 25 mic. long and 9 to 13 mic. wide. See Figs. 10, 12, and 23.

Collected near Peebles, by Bruce Fink, No. 64. On tree trunks.
Not ])reviously recorded for Ohio.

4. Collema plicatile Ach. Lich. Suec. 129, 237. 1798.

Lichen plicatilis Ach. Vet. Akad. Flandl. i6: 11. pi. 1. f. 2. 1795.

Transforming the algal-host colony into a small, irregular or subor¬
bicular, somewhat coriaceous body, which is 0.4 to 2.5 cm. across, 185
to 370 mic. thick, and loosely attached to the substratum, with more or
less imbricated, ascending or erect lobes with plicate-crisped margins,
with the upper surface black-olive, with the lower surface scarcely lighter
colored, and with the algal chains scattered toward the center but numer¬
ous toward the surfaces ; thallus of densely and variously disposed
hyphie, which are 3 to 5 mic. wide ; rhizoids rare, except at the points of
attachment of the thallus to the substratum; apothecia small, sessile, scat¬
tered, 0.5 to 1 mm. in diameter ; disk concave to flat, red- to dark-brown,
surrounded by an entire thalloid margin, which does not extend above it ;
exciple very thin, of interwoven hyphie, tinged with brown ; hypothecium
of interwoven hyph?e, light brown, 90 to 105 mic. thick; hymenium 130
to 160 mic. thick; asci 80 to 95 mic. long and 14 to 18 mic. wide; spores

The Collemaceae of Ohio

53

longitudinally 4-celled, becoming transversely 2-celled, 24 to 30 mic. long
and 5 to 9 mic. wide.

Collected near Peebles, by Bruce Fink, No. 94. On calcareous
rocks. Not previously recorded for Ohio.

5. Collema furvum Ach. Lich. Suec. 132, 236. 1798.

Lichen fnrvns Ach. Lich. Suec. 132. 1798.

Transforming the algal-host colony into a suborbicular or irregular,
small or middle-sized body, which is 0.7 to 5 cm. across, 145 to 310 mic.
thick, and loosely attached to the substratum, with rounded or somewhat
irregular, entire and ascending lobes, which have undulate or crenulate
and somewhat crisped edges, with the upper surface black-olive and cov¬
ered with granules, with the lower surface lighter colored, and with the
algal chains somewhat more numerous toward the surfaces; thallus of
densely and irregularly disposed hyphse, which are infrequently either
perpendicular or horizontal to the surface, 1.3 to 5 mic. wide; rhizoids
numerous at the points of attachment of the thallus to the substratum;
apothecia small, few and scattered, sessile, 0.3 to 1 mm. in diameter;
disk concave or flat, light brown, becoming dark, surrounded by an entire
thalloid margin, which may extend slightly above it ; exciple plectenchy-
matous, hyaline ; hypothecium plectenchymatous below, of interwoven
hyphse above, 90 to 160 mic. thick; hymenium 105 to 120 mic. thick;
asci 90 to 105 mic. long and 20 to 32 mic. wide; spores longitudinally
4- to 6-celled, transversely 2- to 3-celled, 17 to 26 mic. long and 8 tO' 12
mic. wide. See Fig. 13.

Collected near Peebles, by Bruce Fink, No. 184. On calcareous
rocks. Not previously recorded for Ohio.

Leptogium Ach.; S. F. Gray, Nat. Arr. Brit. PI. i: 400. 1821

Transforming the algal-host colony into a foliose, usually lead-col¬
ored or olivaceous body, which is variously lobed and sometimes granu¬
late or isidioid branched, and in which the algal-host chains are com¬
monly more numerous toward the upper surface of the colony ; thallus
of a plectenchymatous cortex surrounding a medulla which is mainly
or wholly mycelial, or rarely more largely plectenchymatoid ; hyphse of
the medulla hyaline, straight or curved, for most part densely and vari¬
ously disposed, more or less branched monopodially ; plectenchymatous
cortex usually well developed and of a single layer of cells, except around
and below the apothecia, where it is usually several layers of cells in

54

Ohio Hkjlogical Survey

thickness; rhizoids rarely seen; ajjothecia usually scattered; the disk
commonly hrown and surrounded hy an entire thalloid margin; exciple
almost always in ])art or wholly j)lectenchymatous and usually hyaline;
hy])othecium hyaline helow to pale hrown above, more often plectenchy-
matous ; hymenium hyaline helow to pale- or darker-hrown above ; para-
])hyses hyaline toward the base and tinged with hrown toward the
slightly enlarged aj^ex, all s|)ecies examined showing some simple ones
and others one- to several-times mono])odially branched near the apex
or farther hack, ranging from 60 to 160 mic. long, from 2 to 3.5 mic.
wide, and from 6- to 14-celled in the si)ecies examined; asci usually
clavate, 65 to 105 mic. long and 12 to 21 mic. wide, the walls 0.5 to 2
mic. thick laterally, and 2 to 11 mic. thick at the apex (thicker in Lcpto-
giiim laccnim) ; si)ores hyaline, becoming muriform, 8 in an ascus in all
of ours, irregularly arranged.

On account of the rigid nature of the thallus, due to the presence of
a t)lectenchymatous cortex, the algal-host colony does not become so
much enlarged and so plainly gelatinized when moist as does the same
host when parasitized hy Syiicclwblastiis or Collcina, genera which are
less rigid because of absence of a cortex.

KEY TO THE SPECIES OF LEPTOGIUM

Plectenchymatoiis cortex of several layers of
cells around the apothecia.

Spores 4 cells in length.

Exciple plectenchymatoiis . 5. Leptogium plechtenchymum

Exciple of interwoven hyphge, but often

more or less plectenchymatoid . 4. Leptogium chloromelum

Spores usually more than 4 cells in length.

Medulla plainly mycelial . 1. Leptogium pulchellum

Medulla of densely interwoven hyphge, and

the structure often plectenchymatoid ... .3. Leptogium tremelloides

Plectenchymatoiis cortex of one or rarely two

layers of cells around the apothecia . 2. Leptogium lacerum

1. Leptogium pulchellum (Ach.) Nyl. Syn. Licit. /: 123. 1858.

Collcma pulcliclliiin Ach. Syn. Licit. 321. 1814.

Transforming the algal-host colony into a rather large, suhorbicular
body, which is 4 to 12 cm. across, 90 to 265 mic. thick and closely adnate,
with the marginal lobes free and somewhat raised, with the upper surface
lead-colored and thrown into delicate, plicate wrinkles, with the lower

The Collemaceae of Ohio

.■)0

surface lighter colored, similarly wrinkled, and more or less pitted, with
the lobes rounded and sometimes imbricated, with entire or undulate
margins, and with the algal chains more numerous toward the surfaces,
especially the upper; medulla of scattered hyphse, which are in part
irregularly arranged and in part perpendicular to the surfaces, the latter
arrangement more noticeable toward the margins, 1.75 to 2.6 mic, wide,
and slightly more numerous toward the upper surface ; cells of the upper
cortex 4 to 8 mic. long and 4 to 7.5 mic. wide, those of the lower cortex
distinctly smaller, the cortex several cells thick around the apothecia;
rhizoids rarely seen in the sections ; apothecia common, small to middle-
sized, 0.5 to 2 mm. in diameter, sessile or raised on a pseudostipe, and
surrounded by a thalloid margin ; disk flat and light- or darker-brown ;
exciple plectenchymatous ; hypothecium plectenchymatous below, of inter¬
woven hyphse above, 90 to 140 mic. thick ; hymenium 80 to 105 mic.
thick ; spores ellipsoid to ellipsoid-pointed, longitudinally 6-celled, trans¬
versely 2- to 4-celled, 18 to 33 mic. long and 9 to 16 mic. wide. See Figs.

1, 3, 5, and 13.

Generally distributed over the state, but not common. On the
trunks of trees, and rarely on rocks. Examined from areas about
Peebles, Washington C. H., Springfield, Paint, Cedar Swamp, George-
ville. Sugar Grove, Chester Hill, and Columbus.

2. Leptogium lacerum (Retz.) S. F. Gray, Xat. Arr. Brit. PI. i: 401.

1821.

LicJicn laccr Retz. FI. Scand. Prodr. 228. 1774.

Transforming the algal-host colony into a middle-sized, suborbicular
or irregular body, which is 1.5 to 7 cm. across, 60 to 210 mic. thick, and
reticulately wrinkled, with the closely adnate horizontal portions obscured
by ascending and expanding lobes, with the margins sinuate or more often
finely jagged and lacerately fringed, with the upper surface lead-colored
to olive-brown, with the lower surface scarcely lighter colored, and with
the algal chains more numerous toward the upper surface ; medulla of
densely disposed and uniformly distributed hyphae, which are 2.6 to
5 mic. wide and extend in various directions ; cells of the upper cortex
8 to 13 mic. long and 11 to 13 mic. wide, those of the lower cortex
scarcely differing in size ; cortex rarely thickened to two layers of cells
around and below the apothecia ; rhizoids rarely seen in the sections ;
apothecia rare, small to middle-sized, sessile, 0.4 to 1.5 mm. in diameter;
disk flat or concave, red-brown, surrounded by a thalloid margin, which

56

Ohio Biological Survey

does not extend above it and sometimes becomes concolorous, giving a
biatoroid appearance ; exciple and hypothecium plectenchymatous, the
latter 125 to 170 mic. thick; hymenium 140 to 175 mic. thick; asci clavate
or rarely obclavate, 115 to 130 mic. long, and 25 to 35 mic. wide, with
the walls 2 to 3 mic. thick laterally and 20 to 25 mic. thick at the apex ;
spores ellipsoid and sometimes pointed, longitudinally 8- to 12-celled,
transversely 2- to 4-celled, 27 to 46 mic. long and 11 to 18 mic. wide.
See Fig. 13.

^^4dely distributed in the state, but rarely fruited. On rocks
among mosses. Examined from areas about Eaton, Paint, Peebles,
Oxford, Collinsville, Fultonham, Kelley’s Mills, Prospect, and Spring-
field.

3. Leptogium tremelloides (L.) S. F. Gray, Nat. Arr. Brit. PI. 2 \ 400.

1821.

Lichen tremelloides L. Suppl. PI. 450. 1781.

Transforming the algal-host colony into a middle-sized, suborbi-
cular body, which is 3 to 8 cm. across, 65 to 210 mic. thick, and
rather closely adnate, with ascending and often imbricated lobes, with
entire or finely undulate or dentate, crisped and irregular margins, with
the upper surface lead-colored and beset more or less with isidioid
branchlets, which may pass into minute lobules, with the lower surface
somewhat lighter colored, and with the algal chains more numerous
toward the upper surface; medulla of densely interwoven and uniformly
distributed, vertically and variously disposed hyphse, which are 2.4 to

2.7 mic. wide, often in part or wholly plectenchymatoid ; cells of the
upper cortex 5 to 10 mic. long and 5 to 8 mic. wide, those of the lower
cortex scarcely differing in size ; cortex several cells in thickness around
and below the apothecia ; rhizoids rarely seen in the sections ; apothecia
small to middle-sized, scattered, sessile or on a short pseudostipe, 0.5 to

1.7 mm. in diameter; disk from slightly concave to slightly convex,
brown, surrounded by an entire or an irregular thalloid margin, which is
sometimes overgrown ; exciple at least very largely of interwoven hyphse ;
hypothecium of interwoven hyphse, except perhaps at the lower limit,
where it is plectenchymatous or plectenchymatoid, 80 to 100 mic. thick:
hymenium 90 to 115 mic. thick; spores ovoid-ellipsoid to ellipsoid-
pointed, longitudinally 4- to 6-celled, transversely 2- to 3-celled, 18 to
32 mic long and 7 to 12 mic. wide. See Fig. 12.

A rare plant in Ohio, the distribution of which is not well known.

The Collemaceae of Ohio

57

On rocks and on trunks of trees. Examined from areas about Colum¬
bus, Sugar Grove, Peebles, and Eaton.

4. Leptogium chloromelum (Sw.) Nyl. Syn. Lich. j: 128. 1858.

Lichen chloromelos Sw. El. Ind. Occ. y.' 1892. 1808.

Transforming the algal-host colony into a middle-sized, suborbicular
or irregular, thin body, which is 2 to 7 cm. across and 50 to 185 mic.
thick, with sometimes scattered, usually isidioid-granulate, frequently
imbricate, ascending lobes, with crisped, undulate, or isidioid-lobulate
margins, with the upper surface lead-colored varying toward green or
olivaceous, with the lower surface scarcely lighter colored, and with the
algal chains somewhat evenly distributed ; medulla of densely, uniformly,
and variously disposed hyphse, which are 2 to 5 mic. wide, sometimes
plectenchymatoid in part or throughout ; cortex rather poorly developed
and not always continuous, the cells of the upper cortex 7 to 16 mic. long
and 4.5 to 12 mic. wide, those of the lower cortex scarcely differing in
size; cortex several cells in thickness around and usually below the
apothecia ; rhizoids often seen in sections, especially under the apothecia ;
apothecia commonly present and often numerous, small to middle-sized,
sessile, 0.5 to 1.5 mm. in diameter; disk slightly concave to slightly con¬
vex, brown, surrounded by a granulate, entire or irregular thalloid mar¬
gin, which may extend above it ; exciple and hypothecium of interwoven
hyphge, but both structures wholly or in part plectenchymatoid, the latter
80 to 105 mic. thick; hymenium 105 to 155 mic. thick; spores ovoid to
ellipsoid-pointed, longitudinally 4-celled, transversely 2-celled, 15 to 30
mic. long and 8 to 13 mic. wide. See Eigs. 4, 19, and 22.

Widely distributed in the state, but infrequent. On rocks and tree
trunks. Examined from areas about Springfield, Scipio, Oxford, Paint,
Mineral Springs, and Peebles.

5. Leptogium plectenchymum sp. nov.

Transforming the algal-host colony into a small or middle-sized,
suborbicular or irregular body, which is 0.8 to 7 cm. across, 35 to 110
mic. thick, and somewhat loosely adnate, with ascending and often imbri¬
cated lobes, with entire or irregular, often minutely lobulate margins,
with the upper surface lead-colored, with the lower surface somewhat
lighter colored, and with the algal chains somewhat evenly distributed ;
medulla of densely and uniformly distributed, vertically and variously
disposed hyphse, which are 3 to 5 mic. wide; cells of the upper cortex
9 to 19 mic. long and 5 to 14 mic. wide, those of the lower cortex some-

38

Ohio JJiologjcal Survey

what smaller; cortex thickened from 2 to several layers of cells around
and below the apothecia, the whole area below the apothecia sometimes
being- plectenchymatous ; the occurrence of the ])lectenchymatous cortex
varying greatly, this structure being in some specimens thickened only
under the apothecia, while in others it is thickened from 2 to 8 or 9
layers of cells around the apothecia, the thickening sometimes passing
outward 500 mic. or more in all directions from each apothecium ; rhizoids
numerous under the apothecia ; apothecia middle-sized or large, sessile,
scattered, 0.5 to 3 mm. in diameter; disk flat or convex, red-brown, sur¬
rounded by an entire or irregular thalloid margin, which often becomes
concolorous with the disk and resembles in appearance a true exciple ;
exciple plectenchymatous, slightly tinged with brown, often passing im¬
perceptibly into the thickened plectenchymatous cortex about the apothe¬
cium ; hypothecium usually composed wholly of interwoven hyphse, but
sometimes in part plectenchymatous below, 45 to 115 mic. thick; hy-
menium 150 to 170 mic. thick; paraphyses narrow, 1 to 1.5 mic. wide;
spores ovoid-ellipsoid to ellipsoid-pointed, longitudinally 4-celled, becom¬
ing transversely 2-celled, 22 to 30 mic. long and 8 to 14 mic. wide.
See Fig. 13.

Collected three times near Oxford, by Bruce Fink. On damp clay.

Mallotium Ach. ; S. F. Gray, Nat. Arr. Brit. PI. i: 399. 1821.

Transforming the algal-host colony into a middle-sized or large,
foliose, lead-colored to black-green body, which has large, often more or
less granulate, sometimes rugose lobes, in which the algal chains are
commonly more numerous toward the upper surface of the colony ;
thallus of a mycelial medulla and a plectenchymatous cortex ; medulla of
hyaline, usually densely and variously disposed hyphae, which are some¬
what uniformly distributed, straight or curved, more or less branched
monopodially ; cortex of one layer of cells, except around and below the
apothecia where there may be two or more layers of cells, not so well
developed as in Lcptogimn, and sometimes difficult to demonstrate over
certain areas of the thallus ; rhizoids numerous and strong, appearing
as a dense, light-colored nap, which usually covers the whole lower sur¬
face and forms the chief differential character of the genus; apothecia
scattered ; the disk usually brown and commonly surrounded by a thalloid
margin ; exciple and hypothecium wholly or largely of interwoven
hyplue, but both or the latter may be at least partly plectenchymatous ;
hymenium pale below and light-brown above ; paraphyses hyaline toward

The Collemaceae of Ohio

59

the base and tinged with brown toward the more or less enlarged apex ;
asci usually clavate; spores hyaline, becoming muriform.

Thallus somewhat thicker than in Lcptogiiiin and rigid enough to
overcome largely the tendency of the algal-host colony to gelatinize con¬
spicuously when moist.

1. Mallotium saturninum (Dicks.) Mass. ]\Iem. Lich. 95. pi. 18. f. 114.
1853.

Lichen saturniniis Dicks. PL Crypt. Brit, i*.' 21. pi. 6. f. 8. 1790.

Transforming the algal-host colony into a rather large, suborbicular
or irregular body, which is 4 to 15 cm. in diameter, 80 to 250 mic. thick,
and somewhat ascending, rounded, subentire, or undulate, with the upper
surface lead-colored varying toward olivaceous or black, more or less
granulate and sometimes rugose, and the lower surface clothed with light-
colored rhizoids, which form a delicate nap, and with the algal chains
more numerous toward the upper surface of the colony ; medulla of
somewhat densely and uniformly disposed hyphae, which are 2.5 to 5
mic. in diameter, in part irregularly arranged, but many of them roughly
perpendicular while others toward the center of the thallus tend toward a
horizontal position; cortex not always well developed and distinct, cells
of the upper cortex 5 to 8 mic. long and 4.5 to 5 mic. wide, those of the
lower cortex slightly smaller; apothecia rare, middle-sized, subsessile, 1
to 2 mm. in diameter, the disk flat and red-brown, surrounded by a more
or less granulate and irregular thalloid margin ; spores ellipsoid, becom¬
ing muriform, 23 to 30 mic. long and 7 to 9 mic. wide.

Known to us from Ohio only through No. 572, E. E. Bogue, col¬
lected near Columbus. Some of the specimens distributed under this
number are in part or wholly Syneclioblastus flaccid us. The only speci¬
men seen was sterile, and the apothecial characters are taken from Tucker-
man. We have never seen any American specimens in fruit. On tree
trunks, and rarely on rocks.

Descriptions of Sex Organs.

Ascogonia coiled, hyaline, single or in groups ; trichogynes usually
passing directly or indirectly to and above the surface, one from each
ascogonium ; spermagonia marginal or submarginal ; basidia of short
cells, more or less branched ; spermatia hyaline, one-celled, oblong-con¬
stricted, located on the ends and sides of the basidia. Internal male sex
organs present in an Ohio species not yet named.

60

Ohio Biological Survey

Synechoblastiis pycnocarpus. Ascogonia on young lobes, 40 to 215
mic. below the surface, horizontal diameter 20 to 42 mic., vertical diam¬
eter 40 to 215 mic., 10- to 18-or more-celled, cells 5 to 17 mic. long and

3 to 8 mic. wide; trichogynes extending 15 to 30 mic. above the surface,
where branched or unbranched, 85 to 265 mic. long, cells 7 to 59 mic.
long and 3 to 7 mic. wide, the two distal cells much longer than the
others ; spermagonia occurring with the archicarps, horizontal diameter
55 to 200 mic., vertical diameter 55 to 205 mic. ; basidia becoming 20-or
more-celled, cells 3.5 to 8 mic. long and 3 to 6 mic. wide; spermatia 2 to

4 mic. long and 0.5 to 1 mic. wide, some seen as low as the fifth cell from
the tip of the basidium. See Figs. 18 and 24.

Syncchoblasius ryssolctis. Ascogonia 46 to 95 mic. below the sur¬
face, horizontal diameter 19 to 28 mic., vertical diameter 17 to 31 mic.,

7- to 15-celled, cells 5 to 9 mic. long and 4 to 7 mic. wide; trichogynes
not seen; horizontal diameter of spermagonia 55 to 155 mic., vertical
diameter 65 to 125 mic. ; basidia 7- to 12-or more-celled, cells 4 to 6 mic.
long and 2 to 5 mic. wide ; spermatia 2 to 3 mic. long and 1 mic. wide.

Syncchoblastus flaccidus. Ascogonia 50 to 60 mic. below the sur¬
face, horizontal diameter 14 to 22 mic., vertical diameter 12 to 20 mic.,
many-celled, cells 6 to 7 mic. long and 4 to 5 mic. wide ; trichogynes
extending about 8 mic. above the surface, unbranched, 56 to 65 mic. long,
cells 10 to 11 mic. long and 4 to 5 mic. wide; spermagonia submarginal
on young lobes, horizontal diameter 150 to 165 mic., vertical diameter
105 to 150 mic., basidia 6- to 15-celled, cells 2 to 5 mic. long and 3 to 5
mic. wide ; spermatia 2 to 3 mic. long and 0.7 to 1 mic. wide. See Fig.
17.

Synechoblastiis nigrcsccns. Ascogonia 34 to 54 mic. below the sur¬
face, horizontal diameter 15 to 26 mic., vertical diameter 15 to 17 mic.,

8- to 10-or more-celled, cells 6 to 8 mic. long and 3 to 4 mic. wide; trich¬
ogynes extending to and above the surface, 50 to 70 mic. long, cells 8 to
10 mic. long and 3 to 3.5 mic. wide ; horizontal diameter of spermagonia
65 to 230 mic., vertical diameter 56 to 190 mic. ; basidia 4-to 12-or more-
celled, cells 3 to 5 mic. long and the same in width ; spermatia 2 to 4 mic.
long and 1 to 1.5 mic. wide.

Syncchoblastus ohioensis, Ascogonia 90 to 225 mic. below the sur¬
face, horizontal diameter 16 to 37 mic., vertical diameter 16 to 30 mic.,
7- to many-celled, cells 6 to 8 mic. long and 5 to 6 mic. wide ; trichogynes
passing directly or indirectly to the surface and 20 to 40 mic. above.

The Collemaceae of Ohio

61

where they are sometimes branched, 105 to 150 mic. long, cells 11 to 25
mic. long and 4 to 6 mic. wide ; spermagonia submarginal, horizontal
diameter 190 to 265 mic., vertical diameter 212 to 250 mic.; basidia 14-
to 20-celled, cells 3 to 5 mic. long and 3 to 4 mic. wide; spermatia 3.5
to 5.5 mic. long and 0.8 to 2 mic. wide. See Fig. 25.

Collema pidposum. Ascogonia 155 to 600 mic. below the surface,
horizontal diameter 16 to 39 mic., vertical diameter 13 tO' 57 mic., 5- to
many-celled, cells 9 to 13 mic. long and 4 to 6 mic. wide ; trichogynes
extending 17 to 24 mic. above the surface, where sometimes branched,
155 to 625 mic. long, cells 8 to 54 mic. long and 6 to 8 mic. wide, distal
cells much longer than others ; spermagonia submarginal, horizontal
diameter 65 to 240 mic., vertical diameter 75 to 240 mic.; basidia 7- or
more-celled, cells 3 to 5 mic. long and 3 to 4 mic. wide; spermatia 1.5
to 4 mic. long and 0.5 to 1 mic. wide. See Figs. 15, 16, and 21.

Collema plicatile. Archicarps not seen ; spermagonia numerous,
commonly tinged with red, horizontal diameter 210 to 345 mic., vertical
diameter 210 to 235 mic.; basidia many-celled, cells 5 to 6 mic. long and
2.6 to 5 mic. wide; spermatia 5 to 7 mic. long and 1 to 1.5 mic. wide,
often slightly curved, located on the ends and sides of the basidia.

Collema microphyllum. Archicarps not seen; spermagonia sub¬
marginal, horizontal diameter 135 to 175 mic., vertical diameter 125 to
140 mic. ; basidia several-celled, cells 4 to 8 mic. long and 4 to 5 mic.
wide; spermatia 2 to 3 mic. long and 0.5 to 1 mic. wide.

Leptogiiim chloromelum. Ascogonia 60 to 95 mic. below the sur¬
face, horizontal diameter 18.5 to 24 mic., vertical diameter 26.5 to 29
mic., many-celled, cells 8 to 10 mic. long and 3 to 5 mic. wide ; trichogynes
not seen; spermagonia submarginal, horizontal diameter 60 to 210 mic.,
vertical diameter reaching 250 mic. ; basidia 4-to 8-or more-celled, cells
2 to 6 mic. long and 3 to 5 mic. wide ; spermatia 2 to 5 mic. long and 0.5
to 1.3 mic. wide. See Figs. 19 and 22.

Leptogium pulchellum. Ascogonia about 65 mic. below the surface,
horizontal diameter about 14 mic., vertical diameter about 17 mic., 8-or
more-celled, cells 7 to 8 mic. long and 5 mic. wide ; trichogynes passing
indirectly toward the surface, not fully studied ; spermagonia marginal
or submarginal, horizontal diameter 85 to 135 mic., vertical diameter 90
to 120 mic. ; separate basidia not found ; spermatia 2.5 to 3.5 mic. long
and 1 to 1.5 mic. wide.

Leptogium lacerum. Ascogonia about 80 mic. below the surface,
horizontal diameter about 18.5 mic., vertical diameter about 24 mic..

62

Ohio Biological Survey

many-celled, cells nearly isodiametrical ; trichogynes not seen ; sperma-
gonia submarginal, horizontal diameter 60 to 135 mic., vertical diameter
70 to 120 mic.; spermatia 2.6 to 5 mic. long and 0.75 to 1 mic. wide.

Note. Only spermagonia which reached nearly to the surface and
contained spermatia were measured. However, some of the sections
may not have passed through the center of the spermagonium. This
doubtless makes some of the minimum measurements too small.

LIST OF THE TITLES CITED.

1. Bachman, Freda M. A new type of spermagonium and fertilization in

Collcina- Ann. Bot. 24: 747-760. pi. 49. 1912.

2. Bachman, Freda M. The origin and development of the apothecium in
Collcma piilposiiin (Bern.).Ach. Archiv. Zellforsch. 10: 369-430. pi. 30-36.
1913.

3. Bary, A. de. Morphologie imd Physiologie der Pilze, Flechten, und
Myxomyceten. 1-316. f. 1-100. Leipzig, 1866.

4. Baur, Erwin. Zur Frage nach der Sexualitat der Collemaceen. Ber.

Deutsch. Bot. Ges. i6'- 363-367. pi. 23. 1898.

5. Lindau, G. Beitrage zur Kenntniss der Gattung GyropJiora- Festschrift
Schwendener 19-36. pi. 2. Berlin, Gebriider Borntraeger, 1899.

6. Schwendener, S. Die Algentypen der Flechtengonidien. 1-42. pi. 1-3.
Basel, C. Schultze, 1869.

7. Stahl, C. E. Beitrage zur Entwickelungsgeschichte der Flechten. I.
Ueber die geschlechtliche Fortpflanzung der Collemaceen. 1-55. pi. 1-4,
Leipzig, Arthur Felix, 1877,

8. Sturgis, W. C. On the carpologic structure and development of the

Collcmaccac and allied groups. Proc. Am. Acad, 2^: 15:52, pi. 1-8. 1890.

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Ohio Biological Survey

EXPLANATION OF PLATE III

Fig. 1. A section through the thallus of Leptogiiim piilchellum, showing the plec-
tenchymatous cortex above and below and the mycelial medulla within.
The algal-host chains are shaded, x 435.

Fig. 2. A similar section through the thallus of Leptogium tremelloides, highly
magnified to show the close plectenchymatoid interweaving of the medullary
hyphse. x 1,000.

Fig. 3. A section showing a hypha and a portion of the thickened plectenchyma-
tous cortex below an apothecium of Leptogium pulchellum. x 1,000.

Fig. 4. Four cells of the upper plectenchymatous cortex of Leptogium chloromelum,
isolated by maceration, but still cohering to the hyphse from which they
were cut off. x 1,000.

Fig. 5. A portion of a section of Leptogium pulchellum cut through the thickened
plectenchymatous cortex below an apothecium, macerated, but the isolated
cells still showing somewhat of the usual arrangement in vertical rows.

X 1,000.

Pig. 6. A section through the thallus of Syncchohlastus daccidus to show a plec¬
tenchymatous cortex over a limited area above and closely packed hyphae,
forming a palisade over a limited area below. The horizontal lines above
and below indicate the limits of the algal-host colony, x 875.

Fig. 7. Another section of Syneclioblastus daceidus, showing roughly isodiametrical
cells cut off at the dorsal and the ventral surfaces, but not plainly plecten¬
chymatous. X 875.

P"ig. 8. xVnother section through the thallus of Syueehoblastus daceidus, taken from
a region where there was little of the peculiar modification, but showing
toward the center above two isodiametrical cells cohering in plectenclwma-
like fashion, x 875.

Ohio Biological Survey

VoL. II — Plate III

Fink and Richards.

65

66

Ohio Biological Survey

EXPLANATION OF PLATE IV.

Fig. 0. A section through an apothecium of Collema pulposum, showing a portion
of the hypothecium of interwoven hyphse below and the lower portions of
three asci and several paraphyses above, x 1,000.

h'ig. 10. A portion of a section showing a few cells of a medullary hypha and three
cells of the plectenchyma in the lower portion of the hypothecium of
Collema micro phyllum. x 1,000.

Fig. 11. Three types of branching of paraphyses in Synechoblashis pycnocarpiis.
X 580.

I''ig. 12. Three paraphyses of Synechoblastus mgrescens, showing two of the bridge¬
like anastomoses sometimes seen in this species and a branch arising from
the middle paraphysis. x 580.

Fig. 13. Types of spores found in the Collemaceae ; a, 2-celled spores of Syneclio-
blastus ohioensis; h and c, several-celled spores of Synechoblastus haccidiis
and Synechoblastus nigrescens ; d, e, f, g, h, and i muriform spores of
Collema pulposum (d), Collema microphyllum (e), Collema furvuni (f),
Leptogium plectcnchymum (g), Leptogium lacerum (h), and Leptogium
pulchellum (i). x 1,000.

Ohio Biological Survey

Fink and Richards,

13

68

Ohio Biological Survey

EXPLANATION OF PLATE V.

Fig. 14. Internal reproductive organs in a member of the Colleinaceae collected at
Fayette, Iowa. Parts of two ascogonia at the left and two trichogynes
passing toward a group of male reproductive bodies at the right, x 1,000.

h'ig. 15. Sixteen trichogynes of CoUeina seen projecting above the surface

of the algal-host colony within one field of the microscope, x 580.

Fig. 16. A single archicarp of Collema piilposiini from the same material from
which Fig. 15 was taken, x 1,000.

Fig. 17. The tip of a trichogyne of SynccJioblastus flaccidiis branched more than
usual. X 580.

Fig. 18. The coiled ascogonium of Synechoblastus pycnocarpiis, with a portion of
the trichogyne above and a portion of the supporting hypha below, x 580.
Note how this differs in form of coil and form of cells from the archicarp
of Collema piilposum shown in Fig. 16.

Fig. 19. The coiled ascogonium and portions of the supporting hypha and the
trichogyne of Leptogiiim chloromchim for comparison with similar struc¬
tures from other members of the Colleinaceae shown in Figs. 14, 16 and
18. X 580.

Ohio Biological Survey

VoL. IT — Plate

Fink and Richards.

69

1HE UBRARY OF THE

SEP 3 1943

UNIVERSITY OF ILLINOIS

70 Ohio Biological Survey

EXPLANATION OF PLATE VI.

P'ig. 20. Three groups of internal male reproductive organs from a Collema-Vikt
plant collected near Peebles, Ohio, x 1,000. Note how these male cells
compare in form and size with the spermatia in Figs. 21, 22 and 23, drawn
to the same scale of magnification.

Fig. 21. Portions of three basidia of CoUema pulposiim, showing five spermatia
attached, x 1,000.

Fig. 22. Portions of two basidia of Leptoghim chloromelum, showing four sper¬
matia attached, x 1,000.

Fig. 23. Portions of four basidia of CoUema micro phylhim, showing ten spermatia
attached, x 1,000.

Fig. 24. Portions of three basidia of SynecJwblastiis pycnocarpiis, showing twelve
spermatia attached, x 870.

Fig. 25. A vertical section through a spermagonium of Synechoblastus ohiocnsis,
showing the ostiole, many basidia, numerous spermatia, some attached and
others free in the cavity of the spermagonium, hyphge surrounding the
spermagonium, and a few chains of cells and two free heterocysts of the
algal host x 580.

Ohio Biological Survey

Plate VI

71

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lIBifJARY

iiNIVErSITV Of ILLINOia
URBANA

THE LIBRARY OF THE

SEP 3 1943

Volume II, No. 2 university of Illinois Bulletin No. 6

Ohio B iological Survey

QUALITIES AND USES
OF THE

WOODS OF OHIO

WILLIAM R. LAZENBY, FOR. ENG.

Professor of Forestry

Published by

THE OHIO STATE UNIVERSITY

Columbus,

1915

Introduction.

CONTENTS

PAGE

Economic Importance of Wood . 75

Qualities of Wood . 76

Color, Luster or Gloss, Texture, Grain, Odor, Weight, Moisture,
Volume, Shrinkage, Checking, Toughness, Durability, Fuel Value,
Miscellaneous Qualities.

Table I — Weight and Specific Gravity . 84

Capacity of Wood to Absorb Moisture . 85

Methods of Testing . 87

Explanatory Notes . 89

Summary of Results . 90

Absorption and Volumetric.

Table II — Rate and Percent of Absorption and Expansion. . . 92

Experiments with Coniferous Woods . . 94

Method and Summary.

Table III — Results in Percentages . 96

Table IV — Air Drying Compared with Kiln Drying . 96

Kinds and Quantity of Woods Used in Ohio . 96

Classification of the Uses of Wood . 98

Building and Construction Work, Sashes Doors and Mantles,

Boxes, Barrels, Crates, Cars, Wagons. Boats, Furniture and Cabinet
Work, Veneers, Agricultural Implements, Fencing Material, Rail¬
road Crossties, Telegraph and Telephone Poles, Household Utensils
and Wooden Ware, Mining Industry, Caskets and Coffins, Wood
Pulp and Paper, Street Paving, Wood Distillation, Fuel.

Woods Used for Specific Purposes . 109

Products from Wood and Trees . Ill

f J ^ . h! ' .

QUALITIES AND USES OF THE WOODS OF OHIO.

By William R. Lazenby, For. Eng.

INTRODUCTION.

For some years it has been the duty of the writer to teach the
subject of wood technology. The observations and tests recorded
in this paper are a part of the studies undertaken in preparing for
this instruction. Comparatively little has been published regarding
the structure and physical properties of wood, and many wood users,
as well as students of trees, have felt the need of more knowledge
along this line. The woods used in the tests reported come from
Ohio trees, and it was the aim to secure as far as possible average
samples.

Many publications that treat of wood are too technical for the
average reader. This I have sought to correct by avoiding the use
of terms unfamiliar to the general public. I trust that these notes

Fig. 1. Western Catalpa grown as a roadside tree, about 25 years old. “Useful for
living fence posts and poles, also for supports for telephone wires. Planted
close so that thinnings can be made when the trees are large enough.”

75

76

Ohio Biological Survey

color. Rosewood, mahogany, and ebony are good examples. Some
of our native Ohio woods are well colored. The holly, basswood,
and buckeye are light colored, but the wild cherry, black walnut, and
persimmon are dark. Most woods when exposed to light and air
change their color in course of time. Mahogany may change from
brown to black; ash, from brownish white to light violet; alder, from
a yellowish white to a reddish brown ; and yellow poplar, from a light
to a dark brown. Heartwood is nearly always darker than sapwood
and, in some cases, the dif¥erence is very marked.

Fig. 2. A corner in the Forestry Museum of the Ohio State University.

Luster or Gloss. — Some woods are glossy or lustrous. Maple
and ash when well planed have a shining appearance. This is like^
wise seen in some of the oaks, especially when they are quarter-
sawed. Pine woods can be easily smoothed, but, as a rule, they are
not glossy or shining. The luster or shining quality of wood is due
to the way light is reflected from the fibers that compose it.

Texture. — There is a difference between the texture and the
grain of wood although these qualities are often confused in the public
mind. The texture of wood depends upon the character of the tissues

Qualities and Uses of the Woods of Ohio

77

slight and imperfect as they are, will incite some of our numerous
wood users and students to observe more closely, and take a greater
interest in our rapidly disappearing Ohio woods.

I desire to express my obligation to Mr. O. W. Pflueger, and
Mr. W. L. DuBois for assistance in the preparation of Table II, and
for many of the photographs used.

QUALITIES OF WOOD.

Wood is one of the most abundant products of nature. It is
found wherever trees grow and people live. Excepting food staples
no material is so universally used, and no product contributes more
to the convenience and comfort of mankind. More than one-half of all
the houses of the world are built of wood and most of the other half
use wood for doors, window frames, floors, and other interior parts.
Nearly all barns and out-buildings are made of wood. W^ood warms
the dwellings and cooks the food of more than two-thirds of all the
people on the earth. When we sit down it is on chairs or benches
made of wood ; every day we eat from wooden tables. The news¬
papers, magazines, and books we read are printed on paper made of
wood. Whenever we ride out it is in a wagon, carriage, or car made
largely, or at least partly, of wood. Our furniture, musical instru¬
ments, household utensils, toys, and ornaments are mainly wood.

We ship the products of our farms, orchards, gardens, and fac¬
tories in baskets, crates, barrels, and boxes made of wood. For
transportation by land or water we rely on wood, and for communica¬
tion by telegraph and telephone millions of wooden poles are used.
We cannot work our farms or our mines without wood. So de¬
pendent are we upon wood that our progress and our well-being can
be measured by its use.

Why is wood so useful, and for what special purposes are our
different woods best adapted? Let us briefly consider.

The reason wood can be put to so many uses is the fact that it
has many qualities that make its use desirable. These qualities may
be divided into outer and inner qualities. The outer qualities are
those that can be recognized by the eye, by touch, or by scent.

Color. — Color is one of these qualities and wood that has an
agreeable or attractive color is in demand for the interior of our
houses, for fine furniture, and for many other uses. Wood that
comes from trees that grow in tropical countries is usually rich in

78

Ohio Biological Survey

that compose it. Ironwood, black gum, buckeye, and willow are ex¬
amples of fine texture, while oak chestnut, coffee tree, and elm are
good illustrations of coarse texture.

Grain. — Wood of rapid growth having wide annual rings is coarse
grained, and wood of slow growth and narrow rings is fine grained.
Wood is called rough grained when it does not work easily under
smoothing tools. W^ood that can be split readily is usually straight
grained. When the fibers are more or less twisted, woven or inter¬
locked the wood is said to be cross-grained or spiral-grained. When
the fibers move in and out but do not cross, the grain of the wood
is called wavy. Certain twisted or abnormal growths are highly
prized by cabinet makers and go under the names of birdseye, mot¬
tled, curly, etc. Some wood, like oak or sycamore, when quarter
sawed, show what is called silver grain.

Odor. — AVhile odor is not an important quality of wood, it serves
a purpose in certain receptacles for furs or other clothing liable to
insect injury. It is undesirable in other cases, injuring the flavor
of food-stuffs and liquids. Sweet birch, sassafras, and red cedar have
characteristic and useful odors. In the process of seasoning most
woods lose the odor they have when fresh cut.

Weight . — As is well known, the same volume of wood of different
species, shows a marked variation in weight. This variation depends
mainly upon the amount of pure wood fiber and the amount of mois¬
ture that the wood contains. The weight of pure wood fiber is prac¬
tically the same for al 1 kinds of wood. It is a little more than one
and one-half times the weight of an equal volume of water. As a
rule, the weight of wood is a good indication of hardness and strength,
but it is not a sign of durability.

The weight of wood of the same kind is quite variable and de¬
pends upon the following factors: (1) percentage of moisture; (2)
rate of growth; (3) age of tree; and (4) soundness of fiber. Woods
that weigh less than 25 pounds per cubic foot may be called light,
those that weigh 25 to 35 pounds may be called medium light. When
the weight is 35 to 45 pounds they may be regarded medium heavy,
and when more than 45 pounds they may be considered heavy.

In a series of tests in the laboratory of the Forestry Department
of the Ohio State University, where over 60 species of wood were
used, it was found that among the heaviest were hickory, dogwood,

Qualities and Uses of the Woods of Ohio

79

persimmon, osage orange, ironwood, juneberry, and some of the oaks.
These weighed when well dried, over 45 pounds per cubic foot. In
the medium heavy list — or those that weighed from 35 to 45 pounds
per cubic foot — are found red and yellow oak, hard maple, ash, yellow
locust, beech, birch, and elm. Among the medium light woods may
be named sycamore, soft maple, tulip poplar, chestnut, black walnut,
cedar, and others. These weigh from 25 to 35 pounds per cubic foot.
Among the lighter woods — those weighing less than 25 pounds per
cubic foot — are found arbor-vitae, cottonwood, willow, white pine,
catalpa, buckeye, and basswood.

Fig. 3, A view of the State University woodlot showing fine young growth by
natural regeneration.

Moisture. — Ordinary well seasoned, or air dried, wood contains
from 10 to 15 per cent of water. Lumber seasoned in the usual way
and kept indoors, retains on the average a little more than 10 per
cent of moisture. When exposed to the weather, or in contact with
soil, seasoned wood absorbs moisture quickly or slowly depending
upon the character of the wood tissue. If dry wood is immersed in
water it increases in weight with greater or less regularity and rapid¬
ity. This increase in weight may continue for several months. A
piece of oak that had been kept under water for more than six months
was found to be still gaining in weight. As a rule, however, long
continued immersion causes wood to lose in weight. This is due to

80

Ohio Biological Survey

the washing out of certain soluble material that is in the cells of
wood tissue.

In testing the absorptive capacity of 69 different kinds of Ohio
woods it was found that the soft woods absorbed the most water in
proportion to the weight, and the hard woods the least. The average
total absorption of 27 kinds of the softer and lighter woods was 94
per cent of their original weight. The average total absorption of
ten medium woods was 74 per cent of their original weight. Thirty-
two hard or heavy woods absorbed 64 per cent of their original
weight.

Volume. — It is a well-known fact that dry wood when subjected
to moisture increases in volume. In some cases it assumes a greater
size than it had before it \vas seasoned. The expansion is unsteady
or irregular in most cases. We might think that the woods which
absorb the most moisture and dry most rapidly would expand and
shrink to a greater degree than the woods that take up less moisture
and lose it less rapidly. This is not the case. The soft lighter woods,
such as willow, white pine, and basswood, expand less than the per¬
simmon, oak, and other hard woods. The average increase in volume
for 69 seasoned woods, tested while each was saturated with moisture
was 12^ per cent of the original volume. Where the shrinkage of
wood is irregular the condition known as warping often follows.

Shrinkage. — When wood loses is moisture, it usually shrinks
more or less, and with this often follows warping and checking.
W arping may be caused by irregular shrinkage. If a board dries
more rapidly on one side than on the other, the shrinkage that follows
causes the board to bend at the ends and sides.

Checking depends largely upon the rapidity with which wood
loses its moisture, and upon the variation in the wood tissue. Hard¬
woods are more liable to check than softwoods. Outdoor air drying,
quarter sawing, steaming, followed by slow drying, painting or oiling
are among the means used to prevent checking.

Toughness. — The toughness of wood depends upon various fac¬
tors. Among the more important are weight, soundness, freedom
from knots, soil where grown, rate of growth, and how stored and
seasoned after cutting. Other things being equal the more rapid
wood grows the tougher it is. This is why wagon makers and others
like second growth ash, hickory, and oak.

Qualities and Uses of the Woods of Ohio

81

Fig. 4, A border of young catalpa trees near the University woocllot.

Durability. — The duration or lasting- quality of wood depends
upon the nature of the wood and the conditions surrounding the wood
The character of the wood fiber; the presence of resinous gums, tan¬
nin or other substances, the power to absorb moisture, — the percent¬
age of sap wood, the rate of growth, and the freedom from knots, all
influence the durability of wood. Many woods, if kept dry, are ex¬
tremely durable, while others fall a prey to dry rot fungus or insect
injury. Some woods that are durable when dry, decay rapidly when
subject to repeated moisture and partial drying, as is seen when
woods are used for posts. Locust, red cedar, and cypress are more
subject to fungus diseases when living than when dead.

It has been clearly demonstrated that fence posts from rapid¬
growing trees decay sooner than those from slow-growing trees from
the same species. Of the woods grown in Ohio, osage orange makes
the most durable fence posts. This is followed by yellow locust, red
cedar, red mulberry, chestnut, and white oak. Bur oak and chestnut
oak and a few other varieties are about as durable in the form of
posts as white oak. A white oak or a cedar growing' raj)idly in the
open may be much less durable, however, than a slow-erowine oak
or cedar in the forest, and a locust or catalpa grown in rich soil is
much less durable than those that grow on a thin soil.

82

Ohio Biological Survey

Fig. 5. Section of the trunk of a “birdseye” maple. This tree was found on the
banks of the Olentangy river, and was taken up and planted on the Univer¬
sity Campus by the author in 1884.

The life of the less durable woods, such as cottonwood, basswood,
sycamore, beech, and hickory, may be greatly lengthened by the use
of certain preservatives. Painting, smoking, charring, kiln drying,
and treating with some antiseptic, like creosote or corrosive sublimate,
are means employed to prolong the life of wood.

Qualities and Uses of the Woods of Ohio

83

Fuel Value. — As wood is used so largely for fuel, its value for
this purpose is a matter of great economic importance. The real
heating power of dry wood is directly proportionate to its weight.
Light dry woods are more inflammable and give a quicker heat of
short duration. Five pounds of wood yield as much heat as one
pound of hard coal. A cord of green wood which contains on the
average 45 per cent moisture, has but little more heating value than
V2 cord of well seasoned or air dried wood. The reason is that a
cord of green wood contains about 250 gallons of water, and the heat
required to convert the water into steam is lost to the general heating
purposes. The heating capacity of unsound wood is reduced in pro¬
portion to the amount of wood tissue destroyed. The presence of
gum or pitch increases the capacity to give off heat. The following

Fig. 6. Remnant of a fine beech forest. The beech is rapidly disappearing in
many sections of Ohio where is was formerly abundant. The growth of
weeds and brush with the open conditions prevent further reproduction by
natural means.

woods have high heating capacity : hickory, beech, locust, ironwood,
oak, ash, birch and maple. The following have low heating capacity:
cottonwood, basswood, white pine, hemlock, and chestnut.

Miscellaneous Qualities. — Among other qualities of woods that
make them useful for certain specific purposes are, — resonance,— de¬
manded in wood used for musical instruments, and ‘^nail-holding’’

84

Ohio Biological Survey

power, demanded in building stuff, wood for railroad ties, boxes, fur¬
niture, etc. Among the woods that have excellent nail-holding power
may be named, white oak, hemlock, and yellow pine.

TABLE I

Table showing the weight per cubic foot, in pounds, and the weight in kilograms
per cubic decimeter which is the same as the specific gravity, arranged in order
from lightest to heaviest.

Weight in Specific

Specio. Lbs. Gravity.

Arbor-Vitae, Thuja occidentalis . 20.28 .32482

Western Catalpa, Catalpa speciosa . 21.72 .34788

Balsam Poplar, Populus balsarnifcra . 21.99 .35221

Black Willow, Salix nigra . 22.86 .36614

Ohio Buckeye, Aesculus glabra . 23.43 .37527

Hemlock, Tsiiga canadensis . 23.77 .38072

Basswood, Tilia amcricana . 24.00 .38440

White Pine, Pinus strobus . 24.02 .38546

Box Elder, Acer negundo . 25,15 .40282

Common Catalpa Catalpa catalpa . 25.15 .40282

Cucumber Tree, Magnolia acuminata . 26.63 .41692

Red Mulberry, Morus rubra . 26.57 .42556

Butternut, Juglans cinerea . 26.57 .42556

Largetooth Aspen, Populus grandidentata . 26.57 .42556

Yellow Buckeye, Aesculus octandra . 26.79 .42909

Cottonwood, Populus deltoides . 27.15 .43385

Chestnut, Castanea dentata . 28.28 .45295

Red Cedar, Juniperus virginiana . 28.28 .45295

Tulip, Liriodendron tulipifera . 30.10 .48211

Red Elm, Ulnius fulva . 30.86 .49428

Sassafras, Sassafras sassafras . 31.13 .49860

Sycamore, Platanus occidentalis . 31.13 .49860

Black Ash, Fraxinus nigra . 32.27 .51686

Black Walnut, Juglans nigra . 32.92 .52727

Tulip, Liriodendron tulipifera . 32.96 .52792

Sweet Gum, Liquidamber styracifliia . 33.76 .54073

Blue Ash, Fraxinus quadrangulata . 34.00 .54457

Jersey Pine, Pinus virginiana . 34.29 .54922

Red Maple, Acer rubruni . 35.36 .56636

Wild Black Cherry, Prunus virginiana . 35.82 .57372

Beech, Fagus grandifolia . 35.82 .57372

Pitch Pine, Pinus rigida . 35.82 .57372

Tamarack, Larix laricina . 35.82 .57372

White Basswood, Tilia americana . 35.93 .57549

Red Oak, Quercus rubra . 35.93 .57549

Yellow Birch, Betula lutea . 37.72 .60816

Shingle Oak, Quercus imbricaria . 38.00 .60864

White Elm, Ulmus americana . 38.65 .61905

Qualities and Uses of the Woods of Ohio

85

Hackberry, Celtis occidentalis . 38.65 .61905

Blue Beech, Carpinus caroliniana . 38.65 .61905

Silver Maple, Acer saccharinum . 38.86 .62242

Yellow Locust, Robinia pseudoacacia . 40.00 .64068

Scarlet Oak, Quercus coccinea . 40.00 .64068

Sugar Maple, Acer saccharum . 41.14 .65893

Black Oak, Quercus nigra . 42.06 .67367

Honey Locust, Gleditsia triacanthos . 42.69 .68376

Coffee Tree, Gymnocladus dioica . 42.91 .68728

Swamp White Oak, Quercus bicolor . 43.43 .69561

Bitternut, Hicoria cordiformis . 44.39 .71099

Beech, Fagus grandifolia . 44.39 .71099

Black Maple, Acer nigrum . 44.50 .71275

Pepperidge, Nyssa sylvatica . 44.50 .71275

White Ash, Fraxinus americana . 44.50 .71275

White Oak, Quercus alba . 44.85 .71836

Shellbark Hickory, Hicoria laciniosa . 44.86 .71846

Chestnut Oak, Quercus prinus . 44.86 .71846

Iron Oak, Quercus muhlenbergii . 45.72 .73229

Bur Oak, Quercus niacrocarpa . 46.29 .74142

Sweet Birch, Betula lenta . 46.57 .74591

Persimmon, Diospyros virginiana . 46.58 .74607

Iron Wood, Ostrya virginiana . 46.92 .75151

Juneberry, Amelanchier canadensis . 47.48 .76048

Crab-Apple, Malus coronaria . 47.66 .76337

Mocker Hickory, Hicoria alba . 48.73 .78010

Shagbark Hickory, Hicoria ovata . 49.14 .78707

Pignut, Hicoria glabra . 51.56 .82583

Yellow Oak, Quercus acuminata . 55.72 .89246

Average . 37.00 .59262

THE CAPACITY OF WOOD TO ABSORB MOISTURE AND
ITS RELATION TO WEIGHT AND VOLUME.

As has been stated, the weight and volume of different sorts of
wood are largely affected by the water content. This is not only true
of the wood of different species but is also true of the wood of each
individual tree and of each part of each tree.

The weight of wood is mainly dependent upon two factors : first,
the amount of cellulose and other compounds constituting the wood
substance, and second, the amount of water it contains.

Lumber seasoned in the usual way and used or kept in a dry
room retains on the average a little less than 10 per cent of moisture

86

Ohio Biological Survey

and when well sheltered out of doors retains about 15 per cent. The
amount is by no means constant but varies with the humidity and
temperature of the air.

The real substance of wood is something more than one and
one-half times the weight of water, and the actual wood fiber of the
basswood or poplar is just as heavy as that of the oak or hickory.

Fig. 7. A shellbark hickory that has grown under good forest conditions. The
long unbranched trunk furnishes timber, free from knots, that is always in
demand.

Qualities and Uses of the Woods of Ohio

87

Since it has been clearly demonstrated that both the crushing
and shearing strength of wood are greatly reduced by moisture, the
amount and rapidity of water absorption is a question of no little
importance.

Dry wood when subjected to moisture increases in volume and
in some cases assumes a greater dimension than when green. When
the dry wood is immersed in water it increases in weight with greater
or less regularity and rapidity, and usually the increase in weight
continues for several months or more. From the results of this in¬
vestigation we see that the volumetric increase is not exactly corre¬
lated with the increase in weight.

Prolonged immersion causes some woods to lose in weight in
consequence of the washing out of the constituents soluble in water.

In order to learn something about the relative rapidity of absorp¬
tion and amount of moisture absorbed by air-dried samples of Ohio
woods the following tests n ere made by the author.

Sixty-nine (69) samples representing sixty-two (62) species were
selected for the tests. The samples were from sections of the trunks
of living, thrifty trees about 15 inches in diameter where the sections
were made. This size was selected as representing a fair average of
the timber trees in Ohio, used for ordinary economic purposes. The

Fig. 8. Forestry students making measurements and estimates of the amount of
merchantable wood in trees cut for paper pulp.

88

Ohio Biological Survey

diameter of the section of some of the smaller-growing species was
less than 15 inches and in a few instances in some of the larger-
growing species it was considerably more. In some cases the sec¬
tions were taken close to the stump of the tree and at other times at
some distance up the trunk. As nearly as possible each sample rep¬
resented the wood midway between center and circumference of sec¬
tion of trunk. Sap wood and the oldest heart wood were equally
excluded. Wherever possible sections representing a fair average of
the merchantable timber portion of the tree were taken.

After the sections were selected they were split into small pieces
carefully numbered and allowed to remain in a dry room for more
than a year.

In seven cases a duplicate sample was taken and carefully marked
as such. By a mechanical planer and trimmer each sample was re¬
duced to the exact dimensions, 9 inches in length, IY2 inches wide,
and % inch in thickness, and therefore contained approximately
0.00585 of a cubic foot.

It is well known that the rate of absorption as well as the rapid¬
ity of drying, depends very largely on the size and shape of the piece,
as well as on the structure of the wood and the species. As the sam¬
ples were of equal size and all treated alike, the test, while not con¬
clusive for any one species, are at least comparable in results.

After the samples were prepared each one was carefully weighed
and from this weighing calculations were made of the weight per
cubic foot of each species. (See Table I.) In some cases these
weights agree with those published by other authors but in other
cases they do not. On the average, however, they do agree fairly
well for thoroly air-dried woods of the size and shape named.

Immediately after the weighing the samples were submerged in
water, the water being that of the temperature of the room where the
tests were made.

The samples were first put into water January 17, at ten o’clock
a. m. and were taken out and weighed just 24 hours thereafter. The
samples were removed from the water one at a time and after the ad¬
hering water was wiped off with a cloth, they were weighed, and im¬
mediately placed in fresh clear water. They were weighed the second
time 48 hours after the first. The third weighing was one week after
the second ; the fourth two weeks after the third ; the fifth two weeks

Qualities and Uses of the Woods of Ohio

89

after the fourth and so on until the last interval which was only one
week in duration.

Immediately after the last weighing wdiich determined the water
absorption for the whole period, the samples were placed in a dry¬
ing oven and so arranged that each should get, as nearly as pos¬
sible, the same amount of heat. At the end of 24 hours in this drying

I

oven they were removed and weighed, and this was repeated at in¬
tervals of 48 and 72 hours respectively. This was done in order to
determine whether the species that absorbed water most rapidly
would also give it up most rapidly.

Fig. 9. Students sawing and splitting cottonwood and willow for shipment to
a paper and pulp mill.

After the samples had been allowed to dry in the air of the room
for one month, they were again submerged in water and the results
noted as before.

NOTES.

One (1) liter is equivalent to one (1) cubic decimeter.

One (1) gram is equal in weight to one (1) cubic centimeter of
distilled water.

One (1) pound equals 0.4536 kilograms.

One (1) cubic foot equals 28.32 cubic decimeters.

0.4536 divided by 28.32 equals 0.016017, a factor which multi-

90

Ohio Biological Survey

plied by the number of pounds per cubic foot gives kilograms per
cubic decimeter, which is the same as the specific gravity.

The reciprocal of the weight of a cubic foot of water equals the
approximate weight per cubic decimeter in kilograms.

Each piece contained 10.125 cubic inches.

SUMMARY OF RESULTS.

In all cases except one, where the weight was the same, the
pieces were lighter after soaking and drying than before they were
first placed in water.

In many cases the samples suddenly absorbed more water after
they had once nearly or entirely ceased absorption. This may be
due to a slight increase in temperature or the water penetrating some
place that it had not previously entered.

There was a marked difference in the absorption of the two
pieces of Beech, considerable in the two pieces of Tulip-tree (Yellow
Poplar), less in the two pieces of Jersey Pine and White Hickory,,
while the two pieces of Ohio Buckeye and the two pieces of Cotton¬
wood are in each case, nearly equal.

In general the woods having the least absorptive capacity are
the most durable species. For a given amount of water absorbed the
more porous woods show a less amount of volumetric increase than
the less porous sorts.

The per cent of absorption is correlated with absorptive capacity.

Average absorptive capacity of sixty-nine (69) samples was eigh¬
ty-eight (88) per cent of the original weight of the samples.

In the first soaking the per cent of absorption was least in the
Yellow Locust, forty-eight (48), and greatest in the Ohio Buckeye,
one hundred and eighty-one (181). The average daily absorption
varied from forty-one hundredths (.41) grams in the Yellow Locust
to one (1) gram per day in the Ohio Buckeye. The average of all
was sixty-seven hundredths (.67) grams per day.

In the second test the average daily absorption was the same as
in the first in only one case, that of the Cucumber Tree. In four
cases it was greater, and in all other cases less. The average was
fifty-five hundredths (.55) of a gram each day.

The average total absorption of twenty-seven (27) samples of
the softer woods was ninety-nine (99) grams each, or ninety-four (94)
per cent of their original weight; of ten (10) medium woods the aver-

Qualities and Uses of the Woods of Ohio

91

age total absorption was ninety-four and five tenths (94.5) grams
each, or seventy-four (74) per cent of their original weight; of thirty-
two (32) of the harder woods ninety-five grams each or sixty-four
(64) per cent of their original weight. Of the sixty-nine (69) sam¬
ples the average per sample was ninety-six and seven-tenths (96.7)
grams, or an average of about 70 per cent for all the woods tested.

After the samples were well air-dried after the second test, they
were for the third time placed in water in order to compare the water
absorption with the volumetric increase. (See Table II.) It is seen
that the increase in weight and volume was the same during the first
period in the Swamp White Oak ; the same during the second period
in the June-berry and Black Walnut; the same during the fourth pe¬
riod in the Cucumber Tree. The cases where the volumetric increase
exceeded the weight increase are during the first period. White Pine
(nearly double), Pitch Pine and Cucumber Tree; second period,
AVhite Hickory, Pig-nut; third period. King-nut, Gray Birch, Arbor-
Vitae ; none during the fourth period. The White Elm during the
second period shows decrease of three (3) grams in weight and in¬
crease of three (3) cc. In the Pepperidge, Hackberry and Western
Catalpa the expansion was all during the first period of one day.
There was excessive expansion in the Red Elm during the last period.
The two Catalpas were the same in volume at the beginning and each
increased the same in volume. Eight samples each increased twenty-
five (25) cc. in volume. The White Elm and the Hack-berry showed
the same per cent of volumetric increase.

The average per cent of increase in volume was twelve and five
tenths (12.5) of the original volume. The total amount of increase
in grams was two and ninety-two hundredths (2.92) times as much
as increase in cubic centimeters.

The expansion was unsteady or more or less irreguar in most
cases.

The original volume varied from one hundred and seventy (170)
cubic centimeters in the Papaw to two hundred and two (202) cc. in
the Persimmon. It might be thought that a piece of timber like
Basswood, for example, would shrink to a less volume than an oak,
but by comparison of the table this will be seen not to be the case.

The volume in all cases was determined by placing the sample
in a graduate containing water in sufficient quantity to completely
immerse it, and recording the amount of water displaced.

92

Ohio Biological Survey

Fig. 10. A typical sugar maple in an Ohio farm woodlot. On account of the open
character of the woods it is branched too low to be of much value. It will
furnish either sap for sugar or good fuel wood.

TABLE II.

Table showing the rate in grams of water absorbed in seven days; the or¬
iginal volume in cubic centimeters and the amount and per cent of increase in
weight and volume, in samples of sixty-two (62) species of Ohio timber. The
periods were one, three, five and seven days respectively. Species arranged accord¬
ing to per cent of increase in volume from the least to the greatest.

Qualities and Uses of the Woods of Ohio

93

a

b

c

d

e

f

g

h

i

j

k

1

m

n

o

Orig.

Orig.

Increase

Increase

Increase

Increase

Total

Per cent of

Species

wei'ht

vol.

after

at end of

at end of

at end of

Increase

Increase at

in

in Cc.

soaking

3 days

5 days

7 days

at end of

end of

grams

24 hours

7 days

7 days

In

In

In

In

In

In

In

In

In

In

In

In

gms.

Cc.

gms.

Cc.

gms.

Cc.

gms.

Cc.

gms.

Cc.

wgt.

vol.

Red Mulberry .

74.75

178

13

7

5

0

7

3

11

1

36

11

48.1

6.1

Arborvitae .

57.82

178

8

5

15

7

0

0

14

0

37

12

63.9

6.7

Tamarack .

97.53

187

16

7

9

4

12

2

9

0

46

13

47.1

6.9

Butternut .

76.60

180

16

7

9

3

7

0

16

3

48

13

62.6

7.2

Black Ash . . .

96.65

187

16

10

8

1

8

2

14

1

46

14

47.0

7.5

Pawpaw . . .

92.07

170

5

12

12

0

9

1

7

0

33

13

27.9

7.6

Black or Y e'l ow

Locust... . .

117.89

184

10

6

6

5

7

0

10

3

33

14

18.4

7.6

Ohio Buckeye .

69.42

185

67

3

14

11

11

0

14

0

106

14

152 7

7.6

Boxelder . . ...

74.10

184

48

11

16

4

21

0

17

0

102

15

138.2

1 8.2

Ironwood .

142.79

190

40

8

16

7

21

0

9

0

86

15

60.6

1 8.2

White Pine._ .

98.02

180

4

7

16

6

10

2

11

0

41

15

41.8

; 8.3

Juneberry .

140.68

185

19

9

7

7

11

0

10

0

47

16

31.1

8.6

Blue Ash._ . .

101.83

187

21

11

7

4

5

2

14

0

47

17

46.2

' 9.1

Cucumber

1

Magnolia .

77.13

185

7

11

17

4

9

0

2

2

35

17

45.4

9.2

Chestnut. . .

81.53

180

16

10

10

3

12

4

15

0

53

17

64.9

9.4

Hemlock . . .

70.43

185

20

13

14

5

11

0

9

0

54

18

76.8

9.7

Red Oak .

103.31

179

28

9

26

9

11

0

14

0

79

18

76.3

10.0

Sassafras .

88.25

177

13

9

8

5

7

4

11

0

39

18

44.2

10.2

Black Walnut .

96.49

183

12

10

2

2

10

4

20

3

44

19

45.6

10.4

Cotton Wood . .

80.16

185

21

7

43

12

17

1

14

0

95

20

116.3

10.8

Jersey Pine .

101.61

185

19

11

16

4

11

0

10

5

56

20

55.2

10.8

Silver Maple. .

117.64

189

16

11

13

5

14

5

17

0

60

21

34.0

11.1

Sycamore .

92.23

189

25

13

11

7

6

1

20

0

60

21

65.1

11.1

"Sl^ite Basswood..

50.07

187

64

17

17

4

7

0

13

0

101

21

201.7

11.2

Downy Poplar .

63.40

180

18

8

44

12

18

0

12

1

90

21

142.2

11.7

Western Catalpa..

59.14

170

19

20

9

0

10

0

14

0

52

20

87.9

11.8

Common Catalpa

68.48

170

19

15

7

2

11

1

10

2

47

20

68.7

11.8

Black Willow. .

67.74

185

18

13

15

5

21

3

18

1

72

22

106.2

11.9

Red Maple _ _

101.94

180

20

15

9

2

21

5

21

0

79

22

77.5

12.2

Basswood . . .

71.88

187

45

14

21

5

12

4

21

0

99

23

137.8

12.3

Persimmon .

155.18

202

44

11

18

3

10

6

16

5

88

25

56.7

12.4

White Oak .

131.46

183

14

7

7

5

11

7

16

44

48

23

36.4

12.6

Sugar Maple .

123.33

188

47

20

14

0

7

3

25

1

93

24

75.4

12.8

Black Maple .

135.42

190

37

15

7

2

10

6

21

2

75

25

55.4

13.2

White Ash . . .

135.42

190

25

11

10

8

9

4

6

2

50

25

36.9

13.2

Beech . .

135.08

190

45

14

13

4

14

1

9

6

81

25

59.9

13.2

Blue Beek . .

111.43

180

26

13

29

6

17

4

12

1

84

24

75.8

13.3

Black Oak . .

124.63

185

23

10

21

13

14

2

12

0

70

25

56.2

13.5

Wild Black

Cherry _ _ ..

111.30

194

39

14

19

12

5

0

6

0

69

26

61.9

13.5

Yellow Poplar.. ..

87.74

182

35

21

28

3

9

0

7

1

79

25

90.0

13.7

Red Elm .

88.97

180

21

17

12

0

4

0

10

8

47

25

52.9

13.9

Bur Oak . . .

138.65

187

16

6

10

9

11

6

17

5

54

26

38.9

13.9

Sweet Gum.. .

100.58

186

58

26

21

0

14

0

4

0

107

26

106.2

14.0

Honey Locust .

121.03

177

34

14

16

6

10

3

20

2

80

25

65.9

14.1

Yellow Oak .

172.24

193

24

17

14

11

11

0

14

0

63

28

36.6

14.5

Grey Birch .

111.29

183

20

14

21

8

4

5

10

0

55

27

49.4

14.8

Swamp White

Oak .

127.30

183

8

8

9

8

9

6

10

5

36

27

28.1

14.8

Kentucky Coffee-

tree . .

132.71

180

28

15

11

5

24

4

5

3

68

27

51.3

15.0

Kingnut Hickory.

134.35

187

18

10

9

6

3

7

20

5

50

28

37.2

15.0

Pepperidge or

Tupelo .

135.32

190

56

29

16

0

11

0

3

0

86

29

63.6

15.3

Pitch Pine .

102 . 12

178

14

17

12

6

9

5

9

0

44

28

43.1

15.7

Sweet Birch _ _

141.72

190

27

12

10

8

7

5

20

6

64

31

45.2

16.3

Chestnut Oak . .

131.48

183

23

12

16

12

14

6

10

0

63

30

48.7

16.4

Bitternut

Hickory... .

132.96

187

17

13

13

11

5

4

19

3

54

31

40.7

16.6

Pignut Hickory. ..

158.46

192

16

11

6

9

15

10

22

2

59

32

37.2

16.7

White Hickory. ..

137.81

187

21

18

2

10

19

3

26

1

68

32

49.4

17.1

Shingle Oak .

106.51

175

24

12

32

16

11

2

10

0

77

30

72.3

17.1

White Elm .

112.05

181

58

26

3

3

20

3

12

0

90

32

80.3

17.7

Hackberry .

114.52

185

61

34

21

0

7

0

16

0

105

34

91.8

18.4

Crabapple. . .

145.04

190

58

27

15

8

1

0

11

0

85

35

58.5

18.4

Scarlet Oak .

115.32

180

26

21

25

9

16

4

8

0

75

34

65.0

18.9

Iron Oak .

131.81

180

35

30

18

3

12

4

9

2

74

39

56.2

21.7

Average .

106.17

182.4

27

13

14.3

5.7

11

2.5

13

1.4

64

24.5

63.8

12.5

EXPERIMENTS WITH CONIFEROUS WOODS.

The results of a series of tests to learn something of the weight
and absorptive power of the following eight coniferous species of
Ohio are given in Tables III and IV :

No. 1, — White Pine — Pinus strobus.

No. 2, — Arbor-Vitae — Thuja occidentalis.

No. 3, — Red Cedar — Juniperiis virginiana.

No. 4, — Hemlock — Tsiiga caiiadeiiais.

No. 5, — Norway or red pine — Pinus rcsinosa.

No. 6, — Jersey or scrub pine — Pinus zdrginiana.

No. 7, — Pitch Pine — Pinus rigida.

No. 8, — Tamarack — Larix laricina.

The numbers used in the tables refer to the specimens named
above.

The weight of the kiln-dried wood is represented by 100 per cent.
Using this as a unit the per cent of water in the air-dried wood is
found, also the percentage of the original weight gained in soaking.
The woods were reduced to the same size as that recorded for the
tests already reported, viz.: Nine inches long, three-quarters of an
inch thick and one and one-half inches wide.

The pieces were first weighed, then dried at a temperature of
250 degrees F. until the weight remained constant.

In two instances, viz.: Hemlock (No. 4) and Pitch Pine (No. 7)
the water was all expelled in the first twenty-four hours, while in the
case of Red Cedar (No. 5) and Tamarack (No. 8) the minimum
weight was reached only after ninety-six hours or four days of kiln-
drying. In all specimens the loss in weight was comparatively slight
between the air-dried and kiln-dried woods. In the Hemlock (No. 4)
the water was expelled with the greatest rapidity.

The same pieces after being thoroly kiln-dried were carefully
measured and the shrinkage in width recorded in per cent is shown
in Table III.

The pieces were then placed in water where they remained for
thirty days. They were taken from the water and weighed at the
intervals stated, and the percentage of gain in weight determined as
given in Table III.

Another series of tests (Table I\^) with pieces of the same di-

94

Qualities and Uses of the Woods of Ohio

95

mensions, but not kiln-dried, gave results nearly identical, excepting
that the air-dried pieces absorbed water more rapidly at first. The
results obtained in these tests would not hold true for pieces of dif¬
ferent size and shape.

Fig. 11. A fungus enemy of veteran trees.

96

Ohio Biological Survey

TABLE III.

No. of SPECIES

White Pine

1

i

t* Arbor Vitae

w Red Cedar

Hemlock

O' Norway Pine

0)

c

s

>»

CA

6

Pitch Pine

O

rt

C3

e

rt

H

8

Air-Dried . . .

1.08

1.09

1.09

1.11

1.09

1.08

1.08

1.08

Kiln-Dried . .
In water 1

1.00

1.00

1.00

1.00

1.00

1.00

1.00

1.00

day (24 hrs.) .

1.35

1.34

1.20

1.33

1.25

1.48

1.31

1.29

2 days .

1.47

1.40

1.26

1.44

1.37

1.54

1.49

1.36

3 days .

1.54

1.46

1.32

1.49

1.44

1.56

1.52

1.42

4 days .

1.62

1.49

1.37

1.54

1.49

1.59

1.55

1.47

5 days .

1.65

1.54

1.40

1.58

1.53

1.60

1.55

1.50

7 days .

1.76

1.58

1.43

1.64

1.53

1.63

1.55

1.51

9 days .

1.79

1.64

1.47

1.68

1.54

1.66

1.60

1.57

11 days .

'1.84

1.68

1.49

1.73

1.58

1.67

1.62

1.59

14 days .

1.87

1.73

1.50

1.76

1.60

1.68

1.62

1.61

17^ days . . .

1.92

1.74

1.52

1.76

1.64

1.69

1.63

1.61

days . . .

1.98

1.81

1.55

1.80

1.65

1.70

1.66

1.61

30 days .

2.07

1.89

1.68

1.86

1.68

1.75

1.68

1.66

TABLE IV.

NUMBER AND

COMMON NAME

Weight per cu.
ft. air dried,
lbs.

Weight per cu.
ft. kiln-dried,
lbs.

Loss in weight
per cu. ft., lbs.

Per cent, of
weight lost in
kiln dried

1

j

Weight per cu.
ft. after 30-day
soaking, lbs.

Inc’se in weig’t
by 30 days soak¬
ing, lbs.

Per cent, gain
in 30 days
soaking

Per cent, loss
in 24 hours
drying

Per ct. shrink¬
age in width jn
drying

1.

White Pine

23

21

2

8.7

43/

22/

107

7.2

0.5

2.

Arbor-Vitae

25!4

23/3

2

9.2

43/

20

89

8.5

1.

3.

Red Cedar

31/3

3

8.7

50

IS/

68

7.7

1.5

4.

Hemlock

27^3

24/3

D

O

10.8

45^

21

86

10.8

1.6

5.

Norway or

Red Pine

35

31/

3/3

9.5

5314

21/

68

8.5

1.3

6.

Jersey Pine

43!^

38^3

3/

8.6

67

28/

75

7.8

1.24

7.

Pitch Pine

38

36

0

7.9

58/

22/3

68

7.9

1.12

8.

Tamarack

36Jd

2

7.8

56^

20y3

66

4.3

1.8

KINDS AND QUANTITY OF WOODS USED IN OHIO.

There are more than one hundred kinds of woods used in Ohio
in some way and in some quantity. Sixty of these woods are promi¬
nent in the lumber trade. About one-tenth of the number grow in
foreign countries and another tenth comes from the Pacific Coast
States. The states of the South furnish the larger quantity of our im¬
ported wood. The wood working factories of Ohio use each year more

Qualities and Uses of the Woods of Ohio

97

than two hundred million feet of lumber. The wood consumers of
Ohio pay over $40,000,000 annually for the wood they use and fully
three-fourths of this amount goes out of the state. There can be no

Fig. 12. A fine white ash timber tree badly injured by lightning.

doubt that our indifference to wood production Avill compel future gen¬
erations to pay exhorbitant prices for lumber. It will also have the
effect of driving some of the important wood-working industries from
the state.

98

Ohio Biological Survey

Our farm woodlots should be producing more and better timber.
A good tree grows as thriftily and surely as a poor one and if our
woodlots were given fair treatment the wood grown therein would
soon take the place of much that now comes from outside the state.
W e see today on many farms from ten to forty acres of indifferent
cord wood, where, with slight cost and care, there might be an
equal quantity of choice timber worth many times as much.

CLASSIFICATION OF THE USES OF WOODS.

The demands for wood are so varied that it is difficult to make
any satisfactory classification of its many uses, but the following
general divisions show the principal ways in which wood is used :

1. Construction work, buildings, bridges, trestles, derricks, etc.

2. Special mill work — sashes, doors, mantles, fixtures, frames, mouldings, etc.

3. Boxes, crates, baskets, barrels, tanks, silos, etc.

4. Transportation vehicles, cars, automobiles, wagons, carriages, boats, etc.

5. Furniture, cabinet work, musical instruments, kitchen cabinets, refrigerators,

etc.

6. Agricultural implements, handles, dairy supplies, bee keepers’ supplies, etc.

7. Posts, boards, rails, pickets, and other fencing material.

8. Steam and electric railroad crossties.

9. Poles for support of telegraph, telephone, electric light and power wires.

10. Household utensils, wooden ware, garden tools, matches, hoops, bungs and

faucets, brushes, excelsior, laundry supplies, athletic goods, etc.

11. Mining industry.

12. Caskets and coffins including boxes.

13. Wood pulp, paper, cardboard, etc.

14. Street paving, board and plank walks, conduits, etc.

15. Wood distillation, charcoal, creosote, wood alcohol, etc.

16. Fuel.

BUILDING AND GENERAL CONSTRUCTION WORK—

One of the largest demands for wood is general construction work.
The general “bill stuff” of architects and builders, such as joists,
studding, rafters, stringers, etc., used in framing and structural work,
consists mainly of yellow pine, white pine, hemlock and white oak.

There are four species that go under the common name of yellow
pine, the most important of these are the long leaf pine, the short leaf
pine, and the loblolly pine. The first named being the hardest and the
strongest, is the most valuable for frame work and is what all good

Qualities and Uses of the Woods of Ohio 99

builders try to secure. It has largely taken the place of hemlock and
white oak.

For siding there is no wood in such demand as tulip or yellow

Fig.13. A plantation of common locust completely ruined by the locust borer
(Cyllene robiniae). Fungus growths often follow the ravages of the
borer.

poplar. In fact, nearly all of the wood-using industries call for this
species. The reasons for its popularity are, that it is free from knots.

100

Ohio Biological Survey

seasons without injury, is easy to work, is durable when kept dry, is
a good nail holder, and takes paint and stains well.

Yellow pine is also used as a finishing wood, although white and
red oak are regarded as its superior. The reasons are that these oaks
have a pleasing grain, are durable, and best lend themselves to artistic
treatment.

Maple is used largely for floors and stairways. Whenever floors
are subjected to hard wear hard maple is regarded as the ideal wood.
It stands usage well, wears smoothly, and retains a fine gloss or
polish. For shingles, redwood, cedar and white pine are the most
satisfactory. As lath are usually made from the waste or leavings
of sawed timber, they are often poor in quality. All soft woods
together with yellow poplar, bass wood, and cottonwood are used
in their manufacture.

Boxes, Barrels, Crates, and Retainers Generally — It is estimated

that in this division 20 per cent of all wood cut finds use. In this
division is included all forms of cooperage, veneer wood, fibre boxes,
baskets, trunk frames, and retainers used for storage and transporta¬
tion. Although the amount of wood seems incredible the quality is
of low average value. Over fifty diflferent kinds of wood are used.
Years ago when white pine was plentiful and cheap, it was in de¬
mand for wooden packages. This species has now given way to
cheaper woods, or when used is of a quality unfit for anything else.
For ordinary store boxes and crates, all the softer and lighter woods
are used. Pine of different species probably takes the lead, but cot¬
tonwood, basswood, chestnut, yellow poplar, and many other woods
are in demand.

For large tanks and silos, cypress is used and for casks and bar¬
rels for liquids and meats white oak is most prized. For all rough
cooperage work elm is the wood most largely used. Elm is also
the principal wood used in the manufacture of baskets, and fruit and
vegetable packages. In addition to elm, beech, sycamore, basswood,
and cottonwood are used for fruit and market baskets. Since long
distance shipments are now so common, every effort is made to re¬
duce the weight of packages in order to lessen freight charges. Wood
is cut as thin as possible, provided the required strength is secured.
This has brought about the use of the veneer box and wood fibre
package.

Qualities and Uses of the Woods of Ohio

101

Cars, Automobiles, Carriages, Wagons, Boats, Etc. — Although
the demand may be less in quantity, there are few if any of the divis¬
ions of wood consumption that use a greater number of varieties than

Fig. 14. A portion of a steep hillside held in place by trees. Note how the de¬
forested portion is eroding.

the one just named. In wagons and carriages the following woods
are largely used : White oak, hickory, white ash, yellow poplar, and
yellow pine. To a less extent, locust, elm, gum, cottonwood, black

102

Ohio JIiological Survey

walnut, birch, beech, black cherry, basswood, and catalpa are em¬
ployed. In nearly all heavy wagons the hubs, spokes, felloes, and
all the running gear, except the axles and the tongue, are made of
oak. The axles are made of hickory and the tongue of white ash.
In lighter vehicles elm and yellow locust are used for the hubs, and
hickory for the spokes and felloes. The shafts are generally made
of ash.

The boxes or beds of carriages and wagons are usually made
of yellow pine, ash, and yellow poplar. Yellow poplar is in great
demand by automobile makers because it is strong, not liable to
warp, and has the quality of taking a fine polish without the use of
any previous filler. For this reason its finish is more permanent.
Its size and clear quality permits its use in the form of wide boards,
and this is an especial attraction to the manufacture of fine vehicles.

The building of our modern Pullman and palace cars calls for the
greatest variety and most expensive kinds of wood. Among the
thirty or more different kinds of wood used, vermillion, mahogany,
teak, rosewood, and other tropical species are found.

Furniture and Cabinet Work — The amount and value of furniture
now in use is almost beyond computation, and the demand for wood
to keep up the needed supply of home and public furnishings is
greater than the supply produced. Not long ago the dark woods such
as black walnut, black cherry and ebony were in general use, but that
time has passed. The lighter colored woods are now in vogue, and
maple, birch, and oak have very largely taken the place of walnut
and cherry.

Oak is now in great demand, and the quarter-sawed work of
the cabinet makers of this country is not surpassed in beauty of
design and perfection of finish elsewhere in the world. Next to
oak, birch, ash, and maple are commonly used for the better kinds
of furniture. For paneling and cylinder desks, where lightness, flexi¬
bility and toughness are required yellow poplar is called for. For the
more common furniture, elm, chestnut, beech, gum, and other cheaper
woods are used. Flard maple is in demand for pianos and organs.
Cabinet making is carried on to a greater or less extent in every
important city in the United States. It furnishes thousands of men
employment and consumes millions of feet of lumber every year.

The use of wood in the form of thin strips or veneers enables
many to secure what they desire in wood work, where the price of

Qualities and Uses of the Woods of Ohio

103

the wood in the solid form is prohibitory. Again, the use of veneers
has the merit of making an unusually strong wood basis, preventing
many woods from warping and checking, as they will when the pieces
used are of ordinary thickness. Another merit in the use of veneers
is that it admits of single designs, or efifect by exact duplication that
are practically impossible in the solid wood. Among the native woods

Fig. 15. A fine crowned red oak. Valuable for shade and ornament, but furnish¬
ing little good timber.

that are often used as veneers are oak, maple, black walnut, birch,
black gum, sycamore, butternut, holly, and sweet gum. Woods se¬
lected that contain knots, burls, curls, or some other abnormal growth,
when sawn or cut into thin slices, present surfaces showing beautiful
figures.

Other sections of these woods may show a wavy, mottled ap-

104

Ohio Biological Survey

pearance equally attractive. These abnormalities in structure, when
turned into veneers, furnish an almost endless variety of figures of
pleasing shape and color. In this way what is more or less of a de¬
formity or disease in nature becomes a thing of beauty in economic
art. V eneers vary in thickness according to the value of the wood
from which they are cut and according to the uses to which they are
put. From ten to fifty, or even a hundred veneers may be cut from
a single inch thickness of solid wood.

One form of veneering is known as “pressed work.” It consists
of gluing together several rather thick veneers of a cheaper wood as
lilack walnut, yellow poplar, or ash, and facing them with the more
expensive mahogany, rosewood, English oak, vermillion, etc., in the
form of thin veneers. These thick veneers are then steamed and
shaped in proper moulds to furnish the backs and arms of chairs,
and curved work of all kinds. Treated in this way, the wood has
greater strength, and is less liable to crack or warp than would be
the case if a single, solid piece were used.

Agricultural Implements — The manufacture of farm implements
and labor saving machinery for the farm has been stimulated by the
recent great advance in agricultural education and the growing scar¬
city of farm labor. The woods that are in demand for the implements
and machinery of the farm are oak, yellow pine, maple, hickory, and
ash, but more than thirty other woods are used. The growing scar¬
city of tough, strong woods has led' to a greater use of steel and iron.
For the same output of implements of certain kinds, as drills, harrows
and cultivators, the amount of wood used in their construction is
often 50 per cent less than it was a decade ago.

Durability united with strength, elasticity and comparative
lightness, is found in ash. No wood can properly replace it for
tongues in the larger agricultural implements, and for handles for
forks, spades, shovels and hoes. Hickory is heavy, strong and elastic,
and for the handles of axes, sledges and hammers can not be equaled.
AVhere hardness and the quality to w^ear smooth is desired, maple
and beech are in demand. Elm, red gum, birch, cottonwood and
cypress are quite largely used in special parts of farm machinery.
Catalpa is being used for doubletrees and neck yokes.

Fencing Material — Nearly all kinds of wood have been used in
the construction of fences. As timber becomes scarcer and more ex-

Qualities and Uses of the Woods of Ohio

105

pensive, wire has gradually taken the place of fence rails, fence boards
and pickets. The demand for wooden fence posts is still large, and is
likely to continue, because neither hedge fences nor cement or metal
posts are yet generally popular. The value of wood for posts is based
mainly upon durability when in contact with the soil. The most dur¬
able woods are desired. Men who have used fence posts all their lives
are not agreed as to which are best. Not durability alone but adapta¬
bility to soil and climate, the rate of growth, freedom from insects and
fungous diseases, and other factors must be considered. It is safe to
say that where the following trees grow surely and thriftily no mis¬
take will be made in selecting them for post timber : Osage orange,
yellow locust, red cedar, mulberry, white cedar, hardy catalpa, chest¬
nut, and black ash. Some species of oak and black walnut make good
posts, but they are of slow growth and are too valuable for other pur¬
poses to be recommended.

Crossties — Our railroads are great consumers of wood. Although
the average life of a tie has been increased by the use of wood pre¬
servatives, the annual demand makes one of the heaviest drafts on
our forests. The kinds of woods most largely used are oak, the yellow
pines of the south, the Douglas fir, western pine, cedar, chestnut,
cypress, tamarack, hemlock, and redwood. Less durable woods
usually treated with preservatives are white pine, lodgepole pine,
gum, spruce, and beech.

White ash is considered one of the best, although when untreated it
is less durable than cedar or cypress. Oak can be worked to a more
uniform size and holds the spikes better than most other woods.
Chestnut is slightly more durable than other woods but does not hold
spikes as well. Cedar ties are not well adapted to heavy pressure,
and are not usually used on sharp curves, unless protected by plates.

Telegraph and Telephone Poles — The woods most in demand for
poles are red cedar, white cedar, birch or tamarack and chestnut. Ex¬
cept the last named these species are not found in marketable quan¬
tities in Ohio, so they are brought in from outside the state, and the
transportation must be added to the first cost.

For the shorter poles yellow locust and catalpa are found satis¬
factory. Since there is likely to be a continuously increasing demand
for these poles, with a steadily diminishing supply, plantations o/
these and perhaps other species could scarcely fail to yield a profit to

106

Ohio Biological Survey

the owner. The average life of poles is about ten years. In a little
less than double this time they can be grown.

Woods that answer all the requirements of a good telegraph or

Fig. 16. A typical beech is an open woodlot. It occupies too much ground, and
its low branching makes knotty wood, of little value except for fuel.

telephone pole are few, and will always command good prices. The
demand in Ohio should be met at as early a date as possible by home
grown chestnut, catalpa and locust.

Qualities and Uses of the Woods of Ohio

107

Household Utensils and Wooden Ware — The lighter, softer and
cheaper woods are generally used for utensils and wooden ware. Cot¬
tonwood, basswood, white pine, buckeye, yellow poplar, black gum,
and other species are in demand. Buckeye, on account of its uniform
texture and its creamy color, is in great demand for wooden ware. It
is also used for artificial arms and legs. Yellow poplar and cucumber
are much used by the toy manufacturer and novelty maker. Wooden
wire used for mattings and screens, is made from willow, basswood,
and cottonwood. Wooden shoe pegs come from birch and hard
maple ; and matches from white pine, cottonwood, spruce, and bass¬
wood. Basswood is in demand for fine excelsior. About ten million
feet board measure of yellow poplar is used in Ohio annually for the
manufacture of bungs and faucets.

Mining Industry — This industry consumes vast quantities of
wood of various kinds. Oak, ash, hickory, yellow pine, and other
tough strong woods are used for supporting roofs and slopes and
for the construction of the machinery and appliances incident to
mining. Thousands of acres are annually stripped of all valuable
timber trees for this purpose alone.

Caskets and Coffins — Receptacles for the dead including the
burial or outer cases are made largely of chestnut and white pine.
The latter is used for the outer cases and the former for caskets,
especially those that are covered with cloth.

These woods are comparatively cheap, light in weight and easy
to work. The chestnut has the desirable quality of being durable
underground. It also holds glue remarkably well, which is a de
mand in cloth covered caskets.

Other woods that are largely used are yellow poplar, hemlock,
white and red oak. These woods take a fine polish, stain ,well, and
are strong and durable. The higher priced caskets are not cloth
covered, and the woods used are often finished so as to resemble
mahogany, walnut or cherry. Occasionally these more expensive
woods are used and are finished naturally, sometimes being elab¬
orately carved.

Wood Pulp and Paper — The use of wood for the manufacture
of paper has greatly increased during the last 25 years. In 1900 about
2,000,000 cords of wood were consumed for this purpose. In 1914
over 4,000,000 cords were used. One cord of wood yields on the

108

Ohio Biological Survey

average 1 ton of ground pulp or ^2 ton of chemical fiber. The prin¬
cipal woods used for paper are spruce, hemlock, and poplar. In
smaller quantities balsam, pine, white fir, cottonwood, willow, beech,
gum, birch, chestnut, maple, basswood, buckeye, and a few others
have been tested. In some places mill and factory waste constitutes
a large part of the wood used. The states of New York and Maine
are the largest consumers of pulp wood. In Ohio the pulp mills
consume about 40,000 cords of wood a vear.

Street Paving — With the improved and more economical meth¬
ods of preservation, more wood is being used now for street paving,

Fig. 17. A woodlot showing some timber trees that are over mature, with few
standards and practically no trees of younger growth.

conduits, and similar purposes. Yellow pine and cedar are among
the woods commonly used as street paving blocks. Some of the
cheaper forms of wood are still used for sidewalks, but this use of
wood is growing less.

Wood Distillation — One of the modern chemical industries is
the distillation of wood. This results in the formation of various
gases, vapors, tarry liquids, and certain solid residues, of which the
chief one is charcoal. The condensible vapors give us wood alcohol
and various acids. The tarry liquids yield wood tar, creosote, paraf¬
fine, naphthalene, and other allied products.

Qualities and Uses of the Woods of Ohio

109

About one and one-half million cords of wood are now distilled
annually. Both hardwoods and softwoods are used. Beech, birch,
and maple are the principal hardwoods consumed with much smaller
quantities of oak, chestnut, hickory, elm, and ash. The yellow pine of
the south is the principal softwood used. In the charcoal kiln process
the gases, vapors, and tar products of wood are usually lost. Charcoal
is the product saved. Two cords of hardwood yield 1 ton of charcoal.
In this rather wasteful form of making charcoal the wood used loses
about 75 per cent in weight and 50 per cent in volume.

Fuel — Inasmuch as nearly all the woods are used to a greater or
less extent as fuel, it is unnecessary to specify them. The abundance
and cheapness of coal, oil, and gas have largely taken the place of
firewood except as a waste product. Farmers who own woodlots are
the principal users of wood for heating and cooking purposes. In
some sections wood is still largely used as fuel in the manufacture
of tile, brick, lime, etc. Slabs, sawdust, and other forms of wood
waste are quite generally used as a fuel to furnish power for sawmills
and woodworking factories.

WOODS USED FOR SPECIFIC BUT MINOR PURPOSES.

Alder — Cigar boxes, wooden bowls, toys.

Ash — Barrel hoops, tool handles, ball bats, oars, tennis racquets.

Beech — Shoe lasts, plane stocks, clothes pins, hames, paving blocks.

Birch — Toothpicks, clothes pins, spools, bobbins, spindles, skewers, brushes, but¬
tons, match boxes, imitation mahogany.

Black gum — Mangles, rollers in glass factories, wooden shoes.

Black cherry — Surveyors’ instruments, backing for metal engraving plates.

Buckeye — Artificial limbs, wooden hats.

Catalpa — Single trees, neckyokes, boys’ ball bats.

Cedar — Buckets, pencils, chests, racing boats.

Chestnut — Coffins, Jacob staff for compasses, backing for veneers.

Common locust — Police clubs, tree nails, insulator pins, cross bars.

Cotton woods — Wooden ware, excelsior, matches.

Crabapple — Pipes, canes, mallets.

Cucumber — Ox yokes, pump logs.

Cypress — Tanks, porch floors, window blinds, molasses barrels.

Dogwood — Spools, bobbins, skewers, engraving blocks.

Elm — Carriage hubs, saddle trees, bicycle rims, wheelbarrows.

Fir — Masts, flag poles, packing cases.

Hackberry — Hames, ax handles.

110

Ohio Biological Survey

Fig. 18. Well wooded ravine where some cleaning up and improvement cutting
should be made.

Hickory — Handles, barrel hoops, screws, spokes, felloes, bows of ox yokes.

Holly — Mallets, engraving blocks, mathematical instruments.

Ironwood — Levers, wagon brakes, shoes, wedges.

Linden or basswood — Parts of musical instruments, wooden shoes, butter churns,
papier-mache.

Maple — Violins, shoe pegs and lasts, pulley blocks, band wheels, gun stocks
wooden type.

Mulberry — Small boats, ax handles.

Oak — Axle trees, brake bars, paving blocks, baskets, plow beams.

Osage orange — Paving blocks, mallets, canes, tooth picks.

Persimmon — Golf club heads, shoe lasts.

Pine — Matches, excelsior, baled shavings, ship masts, patterns for foundry work.
Sassafras — Light skiffs, clothes chests.

Spruce — Oars, paddles, step ladders, wood type, wooden thread.

IHE LIBRARY OF THE

SEP 3 1943

UNWERSnV OF ILLINOIS

Qualities and Uses of the Woods of Ohio 111

Sweet gum — Coiled hoops, tobacco boxes, pipes.

Sycamore — Butchers’ blocks, plug tobacco boxes, butter trays.

Tupelo gum — Root is used for bicycle handles and buoys for fish nets.

Tulip or yellow poplar — Bungs, biscuit boxes, tobacco hogsheads, cigar boxes.
Walnut — Gun stocks, carpenter tool handles.

White ash — Oars, tennis racquets, snow shoes, ball bats, clothes pins, tooth picks.
Willow — Baby carriages, boys’ ball bats, artificial limbs, charcoal for smokeless
powder.

USEFUL PRODUCTS FROM WOOD AND PARTS OF TREES.

Chewing gum from pitch of spruce, fir and pine.

Cordage from inner bark of basswood and pawpaw.

Dyes from sumac and oak bark.

Flavoring substances from hickory bark, birch bark, fruit of cucumber tree.
Healing extract from witch hazel and balsam fir.

Lampblack from white cedar.

Maple syrup and sugar from sap of sugar maple.

Oil of wintergreen from sweet birch bark.

Prussic acid from wild cherry bark.

Sarsaparilla from sassafras roots.

Spruce beer from young branches of hemlock.

Substitute for tea from flowers of basswood, sassafras root bark and buds.
Tanning substances from bark of hemlock, oak, chestnut, spruce, etc., extract
of chestnut wood.

Vanillin from spruce wood.

Wood vinegar, distilled from birch wood.

volume 11. No. 3

Bulletin No. 7

Oh io Biol ogical Survey

THE PHYSIOGRAPHIC ECOLOGY

OF THE

CINCINNATI REGION

E. LUCY BRAUN

Published by

THE OHIO STATE UNIVERSITY
Columbus.

1916

PRINTED BY

THE STONEMAN PRESS

Columbus. Ohio

THE PHYSIOGRAPHIC ECOLOGY OF THE CINCINNATI REGION.

BY E. LUCY BRAUN.

Page

CONTENTS.

Page

I. Introduction .

H. Upland Series .

A. Forest Associations . 121

1. Depressions . 121

2. Undrained flats . . 128

3. Stages of the succession in depressions and on undrained

flats . 13^

4. Drained and gently rolling uplands . 135

5. The pre-erosion climax forest . 137

6. Morainal belt . . . . 137

7. Margins of uplands; narrow divides . 138

B. Clearing Associations . 139

1. Meadows . 139

2. Reforesting areas . 142

HI. Slope Series . 144

A. Ravines . 14®

1. The typical ravine . 146

2. Modifications of the typical ravine . 157

B. River Bluffs . 1^2

1. Rock bluffs . 162

2. Clay bluffs . 169

3. Gravel bluffs . I’i'l

C. Clearing Associations . 173

D. The Erosion Climax Forest . 176

IV. Terraces and Filled Valleys . 177

A. Gravel Terraces . . . . . 177

1. Forest associations . 177

2. Clearing associations . 178

B. Sand and Silt Terraces . 179

C. Filled Valleys . 180

1. The Norwood trough . 180

2. Upper Mill creek valley area . 182

3. The Miami-Whitewater valley . 186

V. Flood Plain Series . 187

A. The Beach-like Shores . . . 187

1. Gently sloping beach-like shores . 188

2. Steep river banks . 189

B. Islands . 102

C. The Flood Plain . . . 194

1. Pond-swamp associations . 194

2. Meadow associations; clearings . 198

3. Forest associations . 198

VI. General Conclusions and Summary . 207

I. INTRODUCTION

Dynamic plant geography or physiographic ecology, is intimately
related to topography and to soil, and consequently, to physiographic
history and to geologic formations. It is therefore necessary to know the
essential features of the geologic and physiographic history of a region
before its plant associations can be understood.

In this paper, the area considered is that generally known as the Cin¬
cinnati region. It includes, in Ohio, Hamilton and Clermont counties,
and the southern parts of Butler and Warren counties ; in Kentucky, the
northern parts of Boone, Kenton, and Campbell counties (see map, fig.

59)-

Considerable has been published on the geology and on the topo¬
graphy of the Cincinnati region ; and a recent contribution by Fuller and
Clapp (1912) gives a brief treatment of both the geology and topography,
preliminary to a discussion of the underground waters. But believing
that the reader of plant geography does not wish to search geological lit¬
erature, especially when it is as scattered as it here is, a brief summary
of the features most essential to the ecologist is here given.

The flat hill-tops or uplands are but remnants of the Tertiary pene¬
plain, uplifted and partially dissected. Near Cincinnati, these uplands arc
merely flat-topped divides, a half mile or a mile in width, for near the
Ohio and its larger tributaries — the Miami, Little Miami, Licking, and
Mill creek — dissection of the peneplain has progressed far enough to
leave few level areas. Farther from drainage lines, in Butler, Warren,
and Clermont counties, the upraised peneplain becomes the general level,
over which sluggish creeks meander in valleys little below this level, and
into which occasional streams are cutting toward the temporary base level
of the Ohio, nearly 500 feet below.

Almost the whole of the Cincinnati region has been glaciated. Drifts
of two epochs, the Illinoian and the Earlier Wisconsin, are represented
within the area. Over the uplands is spread a covering of drift from five
to twenty-five feet thick (fig. 2). Little of the Illinoian till now remains
on hillsides or in ravines, but some of the older valleys are partially filled
with it. It is compact and impervious, for it is composed largely of
clay, with relatively few and usually small pebbles (Leverett, 1902).

116

Ecology of the Cincinnati Region

117

This older drift affects but little the topography of the region, but does
help to produce poor surface drainage on the flat uplands.

North of Cincinnati, and extending into Hamilton county in the
vicinity of Mill creek and the Miami, is the covering of the younger
drift — the Earlier Wisconsin. This, both in the morainal tract and inner
border district, commonly contains numerous and often large pebbles
held in a clayey matrix (Leverett, 1902).

In many places within the area of Wisconsin glaciation, a distinctly
glacial topography may be found. Low gently sloping hills, and valleys
not undrained but still not stream valleys, are common features of the
upland topography within the area which was covered by the Earlier
Wisconsin ice. Some areas on the uplands are flat, and except for soil
differences, are much like those covered only by the Illinoian. As a rule,
it is the uplands covered with the younger drift which are the better
drained. Glacial hills occur outside the upland districts, along the west
side of Mill creek valley from Lockland almost to Hamilton.

The loess, or white clay, which is associated with the Illinoian
deposits, is not found north of the Wisconsin glacial boundary. It
covers the highest terraces and filled valley flats (at an altitude of 600
feet or more) and is spread out in varying thickness over the upland.
It is a very fine-grained material which holds water for a long time.

Outwash gravels from the Wisconsin ice sheet partially filled some
of the valleys, and now remain as terraces in many places along the Ohio,
the 'two Miamis and Mill creek (Hg. 2). These terraces are composed
of coarse gravels in the northern parts of the area, along the Adiami and
Little Miami ; and of finer material, farther south along the Ohio. The
gravel, because of its large pore content, has a high water capacity, but
drains rapidly at the surface, and where exposed on river bluffs. It is
for this reason that terrace edges are among the most xerophytic habitats
of the region.

In most parts of the area, stream valleys cut through the glacial
deposits, and expose the bed-rock beneath. However, along the small
sluggish streams of the upland, and in the deeply filled valleys, bed-rock
is seldom seen.

The bed-rock is of Upper Ordovician age, and five formations are
represented in the region (iig. 2). Three of these, the Pt. Pleasant and
the Utica at the bottom, and the Richmond at the top, seldom outcrop
within the area, and have little or no effect upon the kind of vegetation.
The other two, the Eden shale and the Maysville (Fairview and McMil-

118

Ohio Biological Survey

Ian), it will be shown, affect the character of vegetation, because of their
different physical natures.

The Eden shale is made up of compact thinly bedded shales inter¬
rupted at intervals of about ten feet by thin limestone layers. This
material is easily eroded and weathers rapidly. Consequently slopes on
it are gentle except in the immediate proximity of the stream, where
active erosion produces a gorge. The altitude of its upper surface varies
from over 700 to less than 600 feet within the area considered, owing to
the dip of the strata. This variation in altitude of outcrop has consid¬
erable effect upon stream profile and ravine slopes, and through these,
upon the vegetation.

The Maysville is made up of alternating thin-bedded shale and lime¬
stone layers. The frequent occurrence of limestone layers makes this
formation much more resistant than the underlying Eden shale. Only
one division of it (the Bellevue) outcrops except where relatively rapid
erosion is in process. The Maysville, however, weathers so much more
slowly than the Eden shale, that it produces noticeably steeper hillsides
where removed from the influence of actively eroding streams. And
it is resistant enough to withstand extremely rapid stream cutting, so
that valleys in it are less gorge-like than those in the Eden shale.

The lower formations, Utica and Pt. Pleasant, are usually covered

by the thick alluvium which fills the valleys to a depth of 100 to 200 feet
(fig. 2). Along the larger streams — especially, the Little Miami and

Miami above New Baltimore — the broad flood plains are a conspicuous
feature of the topography. The flood plains along the Ohio and lower

Fig. 1. View from edge of upland above South Milford, showing the flood
plain of East Fork and Little Miami rivers, the terrace upon which Terrace
Park is situated, and the level sky line of the uplands beyond.

Ecology of the Cincinnati Region

119

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120

Ohio Biological Survey

courses of its tributaries reach an altitude of 500 feet, rising upstream
to 600 feet in the vicinity of Hamilton and Loveland. This large area
is occasionally flooded, and supports the characteristic flood plain vege¬
tation of the region.

Thus it may be seen, that the topography of the region presents a
complex of forms — some erosional and some constructional. Uplands,
flat or rolling, broad or much dissected ; ravines and valley slopes ; broad
flood plains ; glacial terraces and filled valleys, are striking features of
the topography. Fig. 7 is a view showing parts of each of the principal
topographic areas.

Soil water is a controlling factor in a consideration of the vegetation
of these four areas. While the water table in general conforms to sur¬
face contour, it is nevertheless in a measure dependent upon the character
of the soil. The approximate position of the water table in the four
areas is shown in the accompanying diagram (£g. 2).

The compact clays of the upland retain the ground-water, so that
in wet seasons, the water table beneath the broad uplands may rise to
within five feet of the surface, while in spring it so nearly coincides with
the surface, that depressions but a few inches 'deep are filled with water
for several months. Cultivated fields in such areas are always trenched
at ten or twelve foot intervals, and it is no uncommon sight to see the
trenches filled with water, and the fields dotted over with crayfish holes.

The different rock formations differ in their ability to retain water.
Both the Richmond and the Maysville, because of their relatively large
percentage of limestone, are traversed by numerous solution channels,
through which ground-water rapidly drains away. The Eden shale,
which is very impervious, nevertheless has little effect upon the water
table, for it is never the surface rock over any considerable area.

Terrace gravels and the deeper valley fillings are much alike in their
high water capacity (porosity 25 to 40%)^; but the loose gravels are un¬
able to hold the ground-water much above neighboring drainage lines.
For this reason, the water table is relatively high in the pre-glacial valleys
no longer occupied by large streams, and low in terraces, which usually
stand from 40 to 70 feet above the flood plain or stream which they border.

In the flood plain area, the height of the water table necessarily
varies with stream height, rising with the spring floods, and sinking during
the dry summers. Here flood water and not ground-water, is the impor-

^ U. S. Geol. Surv., Water-supply paper 259, p. 32.

Ecology of the Cincinnati Region

121

tant factor, as the vegetation must be able to withstand prolonged periods
of complete submergence.

The four topographic forms previously mentioned present different
conditions of soil and ground-water. Each possesses its own general
vegetational types, dependent for their existence upon the cumulative
effects of past and present conditions ; but constantly changing in accord
with the constructional and destructional processes tending toward the
ultimate mesophytic base level.

My thanks are due to Prof. H. M. Benedict of the University of
Cincinnati and to Prof. Henry C. Cowles of the University of Chicago
for their suggestions during the course of my work.

To Prof. N. M. Eenneman, I am under obligations for the use of the
surficial geology maps of the Cincinnati, Hamilton, and Mason quad¬
rangles.

^ 11. UPLAND SERIES.

The upland is the oldest habitat presented, as well as temporarily the
most stable. On many parts, no indications of erosive work are seen.
Wherever undissected, the till plains and loess-covered areas are still
essentially as they were at the close of the glacial epoch. Since that time,
a forest cover has appeared — by what steps it is now impossible to say.
Conditions somewhat similar have been noted by Cowles (1901) in the
Chicago region.

A. EOREST ASSOCIATIONS

1. Depressions

The till plains are not everywhere even, but sometimes have shallow
depressions upon them a few feet in depth, and a hundred or several
hundred feet in diameter. Depressions in the Illinoian drift sheet are
almost imperceptible, and would be passed by unnoticed, were it not for
the character of their vegetation. In central and northern Ohio, and
elsewhere, in regions having a distinctly glacial topography, depressions
of a more pronounced character and containing a very decided swamp
or bog vegetation, are not uncommon. These have been described or
mentioned by a number of authors, — those in Ohio by Bonser (1903),
Dachnowski (1910, 1912, 1912a), Detmers (1912), and Schaffner, Jen¬
nings, and Tyler (1904) ; at Woods Hole by Shaw (1902) ; and in north¬
ern Michigan by Whitford (1901).

One of the most pronounced of the depressions in the Illinoian till
plain of this region, and one which may be considered as typical, is located

122

Ohio Biological Survey

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Ecology of the Cincinnati Region

123

on the Mill creek-Miami divide a mile north of College Hill. The lowest
part of the elongate depression, as is shown on the map, fig. j, lies seven
feet below the general level of the flat upland, and two feet lower than
the bordering tract toward the ravine. Most of the depression is but
slightly more than a foot below this ravineward margin. Yet this slight
depression is sufficient to produce a hydrophytic^ forest. The associa¬
tions are of a character commonly found in shallow morainic depressions
elsewhere. A very similar association is noted by Cowles (1901) in the
morainic areas of the Chicago region.

Fig. 4. Red maples (Acer rubrum) in the lowest part of the depression.

Facies of depression. — The lowest part of the depression is occupied
exclusively by swamp trees, red maple (Acer rubrum)^ and some shell-
bark hickory (Cary a ovata), and swamp herbs, Cardamine bulbosa, Bidens
frondosa, Eupatorium perfoliatum, and species of Carex (dg. 4).

^ The term hydrophytic is used in connection with a type of forest of this region, in which
the soil is saturated, with puddles standing in all slight depressions, during the spring and
early summer — the growing season — and often dry at the surface during the late summer and
fall.

^ Names used are those of Gray’s New Manual of Botany, seventh edition.

124

Ohio Biological Survey

Up to an elevation of five feet above the center of the depression,
swamp trees — red maple, shell-bark hickory, sour gum (Nyssa sylvatica),
pin oak (Qiiercus palustris), and white oak (Qiiercus alba) — are com¬
mon (^g. 5).

Fig. 5. Shell-bark hickory (Carya ovata) and sour gum (Nyssa sylvatica) in zone
around center of depression.

Higher than this, more mesophytic trees appear, advancing into the
depression, and finally becoming the facies of the forest.

Along the top of the ravine slopes, drainage conditions are better
and such trees as mulberry (Monts rubra), hackberry (Celtis occiden-
talis), honey locust (Gleditsia triacantJios) , and pignut (Carya glabra),
are found. The close proximity of hydrophytic trees of the depression
and xero-mesophytic trees of the ravine is one of the striking features
of this bit of woodland, and illustrates plainly the impervious nature of
the till.

Ecology of the Cincinnati Region

125

It seems evident that the distribution of larger trees was determined
by the extent and depth (or wetness) of the depression. The horizontal
succession of large trees accords with the contours. The young ravine
which now cuts across this depression drains its highest parts. In the
next generation, mesophytic, and even xero-mesophytic trees will advance
far into the depression, for the saplings now growing ^beneath the hydro-
phytic trees are largely mesophytes and xero-mesophytes. Of course,
saplings of the swamp trees are scattered over the whole depression, but
the facies of the undergrowth everywhere but in the center is of meso¬
phytic nature as evidenced by the presence of such trees as sugar maple
(Acer saccharum) , red elm (Ulmns ftilva), wild cherry (Primus sero-
tina), and white oak (Querciis alba). The center of the depression
maintains its swamp character, and saplings of red maple, sour gum,
and shell-bark hickory far out-number those of the other trees. The
hydrophytic herbs still persist in every local depression, and commonly
near the center of the major depression.

Fig. 6, Diagram representing the two successions in progress in the depression
shown in Figs. 3, 4, and 5. Horizontal distances represent time interval ;
vertical, progress of succession, toward or away from mesophytism.

Elimination of depression. — Two agencies in particular are at work
in the elimination of the depression : one biotic, the gradual accumulation
of humus, which is slowly building it up ; the other topographic, running
water, which is cutting a ravine across it, and so draining it. The first
is progressive, tending directly toward mesophytism. The latter is retro-

126

Ohio Biological Survey

gressive, for it initiates a long and varied xerarch* succession, which
must intervene between the early hydrophytic stages, and ultimate
mesophytism (iig. 6). The end of this succession is much more distant
than that of the hydrarch succession, which has already advanced so near
to mesophytic conditions. And given time enough, this ravine is destined
to destroy the mesophytic forest already established around the depres¬
sion.

Fig. 7. An upland hydrophytic forest in spring (Terra Alta).

Abundance of depressions. — Many other such depressions have been
found upon the uplands near Cincinnati, some larger and some smaller,
but all similar to the one here described. Few, however, show so com¬
pletely the remarkable succession above described ; for they are often more
irregular, and have no definite center around which the zones are arranged.
The amount and character of the undergrowth in the scattered depression

* The terms xerarch and hydrarch were first used by Cooper in 1913. The following is his
footnote defining them: “The terms xerarch and hydrarch are here used for the first time, for
the purpose of indicating a natural and important classification of plant successions. The
former is applied to those successions which, having their origin in xerophytic habitats, such
as rock shores, beaches, and cliffs, become more and more mesophytic in their successive stages;
the latter to those which, originating in hydrophytic habitats such as lakes and ponds, also
progress toward mesophytism.”

Ecology of the Cincinnati Region

127

forests varies considerably. In some, a carpet of grass, sedge, and swamp
herbs covers the wetter parts (fig. 4), and a dense undergrowth of sap¬
lings occupies the other zones (fig. 5J. In others, the ground is almost
bare, except for scattered plants or patches of Ranunculus abortivus, Car-

Fig. 8. A small depression, overgrown with poison ivy (Rhus Toxicodendron) ,
southwest of Mount Carmel.

damine bulbosa, Impatiens pallida, Impatiens biflora, and Carex typhi-
noides, mixed with a few saplings. It is in the latter type of woods that
standing water in spring is most noticeable (fg. y).

In some woods, there are occasional groups of hydrophytic trees and
herbs, and the intervening spaces are occupied by a mixed forest of a

128

Ohio Biological Survey

common upland type. This will be described later in connection with
the forests of imdrained flats. Often the depression, though small, is
decided enough to produce a pond. Such ponds, however, are temporary,
and dry up during the summer. They are bordered by sedges and but-
tonbush (Cephalanthiis occidentalis) , or overgrown by a rank growth of
poison ivy (fig. 8).

Depressions of all sizes occur on the till plain of the upland, from the
small ones just described, to those so large that they are scarcely distin¬
guishable from the undrained flats.

Fig. 9. Forest of pin oak, near Bethel.

2. UXDRAINED FLATS

Wherever the uplands are broad and undrained, a hydrophytic forest
similar to that of depressions is found. In the immediate vicinity of
Cincinnati, such areas rarely occur, as stream dissection has progressed
too far. The hydrophytic forest of the undrained flat is never extensive
here. But eastward, in Clermont and Brown counties, where the old
peneplain is uncut for many miles, except for shallow valleys, the swamp

Ecology of the Cincinnati Region

129

forest is the prevailing type. Were it not for cultivated fields, this type
of forest would be unbroken except along shallow ravines, or deeper
valleys, where it gives way to mesophytic or xerophytic woodlands.

The dominant trees of the upland hydrophytic forest are not every¬
where the same, nor is the general appearance of the woods necessarily

Fig. 10. A beech tree in the pin oak forest; notice that the roots diverge above
the ground so that the trunk appears to be raised on three props.

the same. Some woodlands are open and grassy^ and others are dense
with scant herbaceous vegetation. But they are alike in that- the trees
of which they are composed are hydrophytes, or largely so. Pin oak
(Quercus palnstris) is often the dominant tree, and may make almost
pure stands for several miles, a condition not reported, to my knowledge.

130

Ohio Biological Survey

anywhere else. In such cases, the trunks usually stand about 8 to 10
feet apart, and very little herbaceous vegetation is found in the dense
shade beneath them (iig. p). Again pin oak is mixed with other trees,
especially white oak (Quercus alba), sour gum (Nyssa sylvatica), sweet
gum (Liquidambar Styracidua) , swamp white oak (Quercus bicolor), red
maple (Acer rubrum), shell-bark hickory (Carya ovata), and sometimes
beech (Fagus grandifolia) . Where beech occurs in the swamp forest, its
larger roots may lie on the surface of the ground, or may be raised above
the surface, appearing as prop roots at the base of the trunk, as illustrated
in dg. 10.

With the exception of swamp white oak and red maple, any tree
which commonly occurs in the pin oak woods, may become dominant.
Where one tree is dominant, it is usually dominant over large areas, or
it may give way locally to a primary species.

Thus, in the northern part of Clermont county (Milford to Blan-
chester), pin oak is usually dominant, but often mixed with white oak,
which locally is more abundant than the pin oak. Farther south, in
Clermont county and in Brown county, in the vicinity of Bethel and
Hamersville, pin oak, though still common, does not produce the pure
stands before noted, but is mixed with sweet gum, and indeed, is often
only one of the primary species in a sweet gum forest. White oak and
red maple are common here also, but seldom occupy any considerable
area exclusively. As the sweet gum forest is so similar to the pin oak
forest in density and in moisture conditions and therefore in under¬
growth, it may be considered as belonging to the pin oak type, which in
this region is the more common.

The extensive forests of the undissected plateau can not of course
occur in the immediate vicinity of Cincinnati. But smaller areas of the
same type are found, wherever similar conditions are imposed. Although
rather exceptional here, they may be considered as outliers of a prevailing
forest type of southwestern Ohio, bearing the same relations to their
surroundings as do the more extensive tracts.

Much that has been said concerning depression forests, applies
equally well to those of the flat uplands. All zonation and horizontal
succession are absent here, except where ponds are present, introducing
the earlier hydrophytic stages, or ravines are working headward into the
plateau, introducing drained conditions.

Pin oak forest. — The pin oak type of forest is well represented by
small areas. The undergrowth is either very sparce, consisting of Car-

Ecology of the Cincinnati Region

131

damine bulbosa, Lilium philadelphicum, Ranunculus abortivus, Onoclea
sensibilis, Aspidium Thelypteris, Scirpus atrovirens, several species of
Carex, and a number of other plants, usually less common ; or it may be
dense, consisting largely of poison ivy, and saplings of the same trees
which make up the forest (dg. ii). In either case, fleshy fungi are

Fig. 11. An upland forest near Mount Washington in which pin oak (Quercus
palustris) is dominant.

usually abundant, and such mosses as Thuidium, Climacium, Polytrichum,
Catharinea, Leucobryum, and occasionally Sphagnum, form extensive
mats.

Forest openings. — White oak is a minor constituent of almost all
these upland woods, where individual trees or groups of trees occur scat¬
tered through the denser pin oak forest. Such spots are more open than
the rest of the woods, and are grassy (dg. 12). Here are usually found
a number of herbs characteristic of sunny swamps and meadows, Scirpus
cyperinus, Ludvigia alternif olia, Bidens frondosa, Lobelia siphilitica, Soli-
dago gramini folia, Eupatorium perfoliatum, Ilysanthcs dubia, Steironema
lanceolatum, and a number of Carices.

132

Ohio Biological Survey

Where white oak becomes the dominant tree, the ground is usually
less swampy, though still very wet ; the herbaceous vegetation is more
abundant, but made up largely of the same species as are found in the
other parts of the forest. Kentucky blue grass (Poa pratensis) often
forms a close sod in the more open parts, and other grasses, Poa sylvestris,
Cinna anindinacea, and Agrostis perennans, are not infrequent.

Fig. 12. Opening in a hydrophytic forest.

White oak. — The general occurrence of white oak (Qucrcus alba),
which is so well known everywhere as a xerophyte or xero-mesophyte,
in the hydrophytic forests of the uplands, is of special interest. Mixed
with pin oak, sweet gum, and red maple, and surrounded by hydrophytic
herbs and mosses, its hydrophytic propensities can not be doubted
(hg. ig). Neither does it seem to mark any decided advance in the
hydrarch succession in which it is here found. Where mixed with pin
oak, it is subjected to the same conditions as the latter — an undrained
clay soil where water may stand in puddles for several months. Where
white oak becomes the dominant tree, the ground is not so wet, and water

Ecology of the Cincinnati Region

133

is found on the surface in but few places. Such a forest does not, how¬
ever, seem to succeed the pin oak forest, but rather to occupy areas which
always have been different. Either a pin oak or a white oak forest may
pass directly into a beech forest.

In the advancing mesophytic forests, white oak rapidly drops out
with the other hydrophytic trees. In the ultimate mesophytic forest of
this succession, it forms no part. It will be shown later, that white oak
is a character tree in the xero-mesophytic associations of slopes. It is

Fig. 13. White oak tree (Quercus alba) in the upland hydrophytic forest near
Bethel ; a hummock of mosses built up around it. The lighter patches
are Sphagnum.

equally characteristic of these two opposing types of forest. Possibly
it may be considered as the ecological equivalent of some of the conifers;
for example, white pine, and to take the place of such trees in this region.

Mixed forest. — Contrasted with the almost monotonous forest of the
pin oak type, are the mixed woods found upon many parts of our up¬
lands. What evidences of succession there are indicate that these mixed
hydrophytic forests probably were once similar to the more extensive
forests previously discussed, and that they are rapidly becoming more
mesophytic.

134

Ohio Biological Survey

Upon the older drift, a representative area of mixed forest about 400
by 1000 feet is found a little less than a mile west of Hazelwood. It is
estimated, that here 84 per cent of the large trees belong to species com¬
monly found in wet soil in this region; 31 per cent are shell-bark hickory
(Carya ovata), and the remaining 53 per cent include in the order of their
abundance, red maple (Acer ruhrum), pin oak (Querciis palustris), white
oak (Querciis alba), white-heart hickory (Carya alba), white ash (Fraxi-
nns americana), shingle oak (Quercus imbricaria), swamp white oak
(Querciis bicolor), sour gum (Nyssa sylvatica), and white elm (Ulmus
americana) . The red maples usually occur in groups, in the wettest part
of the woods. No decided mesophytes are found among the larger trees,
though white ash and shingle oak (18 per cent) often occur in mesophytic
woods also, and might be listed here as mesophytes. Xero-mesophytes
are, however, well represented (16 per cent), and include redbud (Cercis
canadensis) (8 per cent), black oak (Quercus velutina), black walnut
(Juglans nigra), and locust (Robinia Pseudo-Acacia).

Xero-mesophytes make up a much larger proportion of the saplings
than of the large trees. Here red elm, walnut, redbud, hackberry, mul¬
berry, and Sassafras are common, and compose probably 50 per cent of
the young tree growth. Wild black cherry (Primus serotina), a meso-
phyte, is found, and white ash has increased in importance.

The large trees stand rather far apart, so that the ground in many
places is not well shaded. This may in part account for the scarcity of
saplings of mesophytic trees, which are appearing only in the more shaded
portions.

In the wetter parts of the woods, are saplings of the hydrophytic
trees, and many herbaceous plants, among which are Scirpus atrovirens,
Scirpus lineatus, Juncus tenuis, Potentilla canadensis, Epilobium color-
aturn, Ludvigia alternifolia, Pycnanthemum dexuosum, Bidens frondosa,
and several species of Carex. Carex squarrosa frequently forms large
patches.

Few such areas are found upon the Illinoian till plain; but upon the
gently rolling surface of the younger drift, and even on its flattest parts,
the mixed swamp forest is more common than that in which pin oak pre¬
vails. Extensive flats of Wisconsin age are absent in this region, and
for this reason the swamp forest is limited in extent. The forest suc¬
cession seems to have advanced more rapidly here, probably because of
the more porous nature of the sub-stratum, so that the areas of the two
drifts may often be distinguished by the character of their plant cover.

Ecology of the Cincinnati Region

135

3. Stages of the succession in depressions and on undrained flats.

The earliest steps in the hydrarch succession in progress in the de¬
pressions and on the undrained flats are problematic. At first, of course,
the succession was climatic, and the earliest stages were not such as could
exist in these places today. If we may compare the sequence of events
here after the retreat of the glaciers, with what we find in the north today,
we would expect tundra and meadow vegetation, slowly encroached upon
and restricted by the advancing forests. Today, small natural openings
are found in the upland forests. Some are still ponds, with their typical
marginal vegetation of sedges and buttonbush ( Cephalanthus occidenta-
lis). In others (dg. 4), are a few red maples and a number of sun plants
instead of the shade plants of the rest of the forest. Still others are the
openings about the white oak trees (dg. 12), with their more meadow-like
vegetation. Are these perhaps, relicts of a former and more extensive
formation, in time doomed to destruction by the encroachment of the
forests? At least, they give some clue to possible earlier stages leading
up to the present hydrophytic forests. The succession indicated here is
also further substantiated by what is now going on in many clearings.

Upon the broadest uplands, there are no indications of what will
follow the swamp forest. In the next generation, at least, it will succeed
itself. But on the smaller uplands, and wherever streams are draining
the flats, or around the margins of depressions, is developed a mesophytic
forest.

4. Drained and gently rolling uplands

It was shown in discussing the depression forest, that it is being suc¬
ceeded by a more mesophytic vegetation, because of the draining and
gradual filling of the depression. Small streams are working headward
into the broad flats, carrying off the surface water, and leaving areas
drained but still moist. Few such upland valleys are cut through the till
mantle ; often indeed, their bottoms are but five or ten feet below the gen¬
eral level of the upland. This slight drainage has served to change a
hydrophytic into a mesophytic habitat.

Succession. — Occasionally a horizontal succession from the hydro¬
phytic to the mesophytic forest is seen. More rarely, the succeeding meso¬
phytic vegetation is foretold by the appearance of mesophytic trees in the
undergrowth of a hydrophytic forest. The succession consists of but few
stages. Sometimes a mixed forest intervenes between the pure hydro¬
phytic and the ultimate association of the succession. Often, however,
the jump is sudden. If horizontal succession, upon a very gradually

136

Ohio Biological Survey

changing topography, may be used as an indication of vertical succession
in a place bound to pass through those same topographic stages shown in
the area of the horizontal succession, a beech forest will follow the hvdro-
phytic association. A beech forest is the climax of the hydrarch succes-
sion of the upland.

The beech forest. — A beech woods is sometimes found upon flats
which are still so wet that swamp herbs are not uncommon. Sometimes
even, water stands between the trees. But a beech tree in such a situa¬
tion, occupies a mound one or two feet above the wetter places (hg. g).

Fig. 14. An upland beech forest, the pre-erosion climax, near Madeira.

Thus, its habitat is essentially different from that of the swamp herbs.
The same feature was noticed by the writer along the C. E. and W. Elec¬
tric R. R., in Lorain Co., Ohio, where, in a forest composed chiefly of
beech and red maple, beech occupies distinctly higher ground, being either
on low ridges or on individual mounds.

The mesophytic forest of the upland is composed almost entirely of
beech (hg. 14). Occasionally sugar maple (Acer sacchai urn) and white

Ecology of the Cincinnati Region

137

ash are found, but beech forms about ninety-five per cent of the tree
growth. Of the saplings, sometimes fifty per cent are sugar maple ; beech
is next in abundance, and white ash, hickory, and wild cherry are found.
The large percentage of sugar maple among the saplings does not neces¬
sarily indicate that it will become an important species in the succeeding
forest, as the mortality among the smaller trees is higher in this species
than in beech.

Herbaceous plants are not common in these beech forests. The
vernal vegetation is composed chiefly of spring beauty ( Claytonia virgin-
ica), toothwort (Dentaria laciniata) , waterleaf ( Hydro phylliim appendicu-
latuni), and May apple (Podophyllum peltatum). The aestival vegeta¬
tion is very scanty, and the most common plant is beech-drops (Epifagus
virginiana). Spikenard (Aralia raccmosa) , white snakeroot (Eupatori-
iim urticaefolium) , spleenwort (Asplenium angustifolium) , and young
plants of Hydrophyllum make occasional patches of green.

Wet places still remain in most of the upland mesophytic forests.
Here are found a few swamp trees, usually sour gum, but sometimes pin
oak or red maple ; and the herbaceous vegetation is similar to that of the
hydrophytic forests.

5. The pre-erosion climax forest.

This climax forest is but temporary. As long as the topography
changes but slightly, the beech forest remains unchanged. But it can be
no more lasting than the topographic form upon which it is found. This
climax forest is one developed on a topography little modified by erosional
changes. It is found in but one other situation — filled valleys — and
there also upon a youthful topography. In contrast to the truer -climax
of the region, it will be termed the pre-erosion climax. It is best devel¬
oped on the youthful topography of the Illinoian drift area, but it is also
found upon the pre-erosion topography of the uplands covered with Wis¬
consin drift.

6. ^lORAINAL belt.

Although no distinct terminal moraine marks the most southerly
position reached by the Earlier Wisconsin ice sheet, the topography in
this morainal belt is decidedly more varied than that to the south. The
character of the vegetation of the two drifts is often different, even on
the flatter parts, where both successions are progressing from. the hydro¬
phytic to the mesophytic. In portions of the morainal belt, a xerarch
succession is in progress. The conditions imposed here were never ex-

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Ohio Biological Survey

treme. The slopes are gentle (about 10 degrees) ; the soil a stifif bowlder-
clay.

The forest association today is mesophytic, but scarcely climax.
Beech and sugar maple are dominant trees, but with them occur black oak
(Quercus velutina), white oak (Qiiercus alba), the small-fruited hickory
(Carya microcarpa), and Sassafras. Only a few remnants of this forest
association remain upon the uplands. The character of the undergrowth
is nowhere displayed.

The contrast between this morainal forest, and the hydrophytic
forest of the flat upland to which it is adjacent, is striking. The sudden
change in vegetation, which is so generally found along the geologic boun¬
dary between the two drift sheets in this region, is due chiefly to the
change in topography. The vegetational line is almost as marked as the
topographic, and is closely related to it ; for wherever the topographic line
is indistinct, the plant associations grade into one another. In places, the
morainal forest is almost lacking, and the mesophytic or hydrophytic
forest of the flat uplands to the south (area of Illinoian till) passes indis¬
tinctly into the mixed or hydrophytic forest of the rolling Wisconsin till
plain.

This distinction between the vegetation of the two drifts is only pos¬
sible upon the uplands, whose topography has been changed by the till
cover. Soil fertility in the two areas is very different, and farms within
the area of Wisconsin drift are worth about one and one-half times as
much per acre as those south of its margin. But similar topographic
situations upon the two areas support similar types of natural vegetation,
and the differences which can be noticed are due chiefly to the differences
in the topography.

7. Margins of uplands; narrow divides.

In some places, the remnants of the Tertiary peneplain are so small
or so well drained, that none of the characteristic upland forest associa¬
tions are found. Such conditions are to be seen in the vicinity of the
major streams, especially near the Ohio and along such narrower divides,
as the Ohio-Licking, upon which Fort Thomas is situated. In these
places, the multitude of ravines afford rapid surface drainage, and their
deep valleys have so lowered the water table beneath the upland, that a
xero-mesophytic habitat is presented.

The forest association is essentially that of the dryer ravine slopes.
It is the association of the tops of ravine bluffs, where xerophytic con¬
ditions remain the longest. The association is a retrogressive step in the

Ecology of the Cincinnati Region

139

succession, and with the continued dissection of uplands, and the head-
ward working of ravines, will spread back farther and farther into the
uplands. The hydrophytic forests will give place to mesophytic ones,
which mark the temporary climax of many upland successions. But this
temporary climax forest will in time give way to the xerophytic associa¬
tion, which initiates the xerarch succession characteristic of slopes.

A forest of oaks (Quercus Muhlenhergii, Q. rubra), sugar maple
(Acer saccharum), red elm (Ulmus fulva), basswood (Tilia americana) ,
hop hornbeam (Ostrya virginiana) , and thorns (Crataegus) , replaces the
mesophytic beech forest of the upland. The undergrowth is usually of
saplings of the same kinds of trees, although the proportions vary con¬
siderably, depending on the stage in the succession. In the more meso¬
phytic places, sugar maple is dominant. Near the margins of steep
bluffs, it may be entirely absent. Even where it is an important constitu¬
ent of the forest, it may be almost lacking from the undergrowth, if the
trend of the succession is toward a less mesophytic forest.

B. CLEARING ASSOCIATIONS

The forest cover has been removed from a large area of the uplands,
and in many parts of this area, a clearing succession is in progress.

1. Meadows.

It will be remembered that natural openings in the upland forests
contained a characteristic herbaceous vegetation. Wherever a few trees
are removed, this herbaceous vegetation appears, so that in a short time
such a clearing has much the appearance of a natural opening. More
extensive clearing is attended by similar results. If the cleared upland
is wet, a meadow association resembling that of wet prairies appears ;
if dry, it is soon occupied by a number of xerophytic annuals and per¬
ennials.

The clearing successions are probably similar to, but more rapid than
the original upland successions. Upon the wetter uplands, the earlier
stages of a hydrarch succession are shown.

Wet meadows. — Depressions are filled with water for at least a part
of the year. Larger ones exhibit a typical pond-swamp succession (dg,
13). The number of aquatics is determined by the permanency of the
pond. Submerged aquatics, other than algae, have not been found, and
pondweed (Potamogeton dimorphus) was observed in only one instance.
The rush (Juncus effusiis) frequently forms a distinct marginal zone (dg.

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Ohio Biological Survey

-i^) and with it may occur the bulrush (Scirpus validus) and species of
Eleocharis. Encroaching upon these, is a zone made up of a number of
Carices and Scirpus. In places, cat-tails (Typha latifolia) may nearly
fill the pond; or such species of broad-leaved amphibious plants as Alisma
Plantago-aqiiatica, Litdvigia pahistris, and Penthonnn sedoidcs may com¬
pose the marginal zone.

Very shallow depressions on the wet uplands are filled with water
for only a short time. Their position is indicated throughout the year by

Fig. 15. Juncus effusns forms the marginal zone of the pond in this clearing south¬
west of Mount Carmel.

the patches of Scirpus atrovirens and Carex typhinoides, which usually
surround or even fill them (fig. i6).

The sedge zone is followed by a meadow association in which grasses,
sedges and rushes often predominate. Other plants which commonly
occur in the wet grassy meadows are Arisaema Dracoiitium, Cardamine
hulhosa, Polygala sangiiinea, Viola papilionacea, Lythrum alatiun, Ludvi-
gia altcrmfolia, llysanthcs dubia, Gerardia tenuifolia, Diodia teres, Hons-

Ecology of the Cincinnati Region

141

tonia coerulea, and Solidago graminifolia. A number of mosses (Hyp-
num, Mnium, Polytrichum, Climacium, and Dicranum), and occasional
hepatics and lichens are found. In a few places, Bidens trichosperma is
the dominant plant, and grows so abundantly, that the meadows are a
mass of golden-yellow blossoms early in September.

Fig. 16. A grassy meadow at Terra Alta. The darker patches are Scirpus atro-
virens and Carex typhinoides.

Dry Meadozvs. — The dryer parts of the uplands, when cleared, are
soon occupied by a number of the more xerophytic grasses (Poa com-
prcssa, Aristida gracilis, Panicuni capillare, Andropogon virginicuni) .
As was the case in the wet meadows, other plants also occur, among which
are Pentstemon glaber, Pycnanthenium flexuosum, Rudbeckia hirta, Soli¬
dago nemoralis, and A. canadensis, Anaphalis margaritacea, and Aster
ericoides var. villosiis. In some places, this dry meadow is a later stage
in the pond-swamp-meadow succession.

The meadow may be only a rapidly passing stage in the forest suc¬
cession, or it may be relatively permanent. The first condition is ex¬
pected ; the second is difficult to explain.

142

Ohio Biological Survey

Certain areas of the upland which are known to have been cleared of
their forest cover at least twenty-five years ago, are still meadow. Such
an area is shown in fig. i6. It is true, a few scattered trees ( Quercus bi¬
color, Q. paliistris, and Acer rttbnnn) occur, but not enough to be con¬
sidered as the beginnings of a forest cover. That conditions are still suit¬
able to the growth of hydrophytic trees, even though the water table may
have been somewhat depressed and conditions of light and exposure made
severe by the removal of the forest cover, is proven by the red maples
which have been planted along the roadsides. It is possible, that during
the interval, the succession may have been set back by burning, or by
cutting. Grassy meadows, however, are common upon the Little Miami-
Mill creek divide. Few of the flattest parts of this upland show any ten¬
dency toward reforestization. Woodlots retain their straight edges, and
do not encroach upon the meadows. Cleared areas reach, in a few years,
a meadow stage which is much more persistent than elsewhere in the
region.

The causes which produce this condition are not understood. Cer¬
tainly the meadow can be but temporary, and must in time give way to
forest. Meadows, not now reforesting, are in all cases covered with a
thick and dense turf of grass and a few other low herbaceous plants.
Where this is occasionally broken, as at the edges of plowed fields or
along roadsides, a few trees are found. It may be that as the turf be¬
comes thinner in the shade of the few trees which do appear, seedlings
will be able to get a foothold, and a new forest will start. Conditions
which appear to be similar to these of our cleared uplands, have been
noted by Shimek (1912) as occurring near Amana, Iowa, in the heart of
a forest region.

2. Reforesting areas.

The usual clearing succession reaches the meadow stage either direct¬
ly, by the rapid spreading of meadow plants growing in natural openings,
or through the earlier hydrophytic stages. Early in the succession, sap¬
lings usually begin to appear: xerophytes (Sassafras, red elm, and locust)
in the dry meadows, and mostly hydrophytes (pin oak, red maple, shell-
bark hickory, and white ash) in the wet meadows.

If the clearing is large, or the meadow distant from areas of original
forest cover, the saplings are apt to be somewhat scattered. The suc¬
cession, however, definitely shows its forest trend ; and this meadow differs
from the more permanent ones in that grass is a less important constitu-

Ecology of the Cincinnati Region

143

ent. The aspect of such a meadow is therefore much more varied (fig,
ly). A large number of species are found, among which are a few
grasses (Poa pratensis and P. compressa, Aristida gracillis), J uncus
tenuis, and /. marginatus, Scirpus atrovirens, Habenaria peramoena, Hy¬
pericum punctatum, Ludvigia alternifolia, Steironema lanceolatum, Ly th¬
rum alatum, Poly gala sanguinea, Verbena angustifolia, Pycnathemum
flexuosum Pentstemon glaber, Eupatorium coelestinum and E. perfoliat-
um, Solidago nemoralis, and vS. graminifolia.

Fig. 17. A clearing meadow near Mount Washington returning to forest.

The species of saplings appearing are controlled to a large extent
by the kind of trees in the vicinity. In the region of sweet gum forests,
this species is an important constituent of the new growth. Elsewhere it
is entirely absent. Pin oak is the most common species, and with it oc¬
cur red maple, swamp white oak and white ash. In almost every case, a
considerable number of red elms (Ulmus fulva) appear.

144

Ohio Biological Survey

Forest openings indieated in the elearing successions. — Very near to
a tract of forest — within about fifty feet (probably the effective limit of
seed distribution) — saplings are much more numerous than at greater
distances. This growth is composed of the same species as are found
in the neighboring woodland, and is often so close, as to almost exclude
meadow vegetation. But openings are found in this, and each contains a
rich meadow flora. These will grow smaller as the trees increase in size,
and the quantity of meadow vegetation, and probably also the number
of species, will decrease. It is such spots as these which probably form
the openings in the upland forests.

It is likely that the original upland forests grew up in a manner veri¬
similar to that now shown in clearings. Natural meadows do not occur
in this region, and the only meadow plants, outside of clearings, are found
in the small openings in the forests. The history of such spots is proba¬
bly that outlined above. Conditions are still favorable to the development
of extensive meadows, if the forest is removed. Then the plants, which
still maintain a foothold in the openings, will spread rapidly and cover the
cleared area, only to be slowly restricted again by the growth of new
forests.

III. SLOPE SERIES

The slopes of the region naturally fall into two principal divisions,
ravine slopes, and major valley slopes or river bluffs. That such a sepa¬
ration of slopes should be made, may at first sight seem strange. In fol¬
lowing a growing ravine from its source to its entrance into one of the
larger streams crossing the region, a connected history is involved. Such
a stream is, in physiography, termed susequent. But here the sequence
may stop. The large valley is not always older than the small one. The
Ohio and lower Miami rivers did not in preglacial times follow their
present course. Because of the drainage changes shown in the accom¬
panying map, hg. i8, parts of the Ohio and Miami valleys exhibit marks
of extreme youth. And youth in a large valley does not impose the same
conditions upon vegetation, that youth in a ravine does. The narrow and
deep ravine is quite different from the xerophytic bluff (Cowles, 1901 ,a).
The history of the river valley is not the history of its tributary valley.

The river series of Cowles (1901), including the ravine, the river
bluff, and the flood plain, is somewhat comparable to the slope series and
the flood plain series of the present paper. But here, the river bluff and
the flood plain are such prominent and important features of the tbpo-

Ecology of the Cincinnati Region

145

Fig. 18. Pre-glacial and present drainage in the Cincinnati region. (After Fenneman.)

146

Ohio Biological Survey

graphy, that they are deserving of separate treatment. The ravine, as
here described, includes blufifs and flood plains of small size only.

A. RAVINES

The smaller streams of the region cut through a number of different
materials, which to some extent, modify the character of the valley, and
with it, the plant successions. Usually, these are met with, in definite parts
of the course. A typical valley (Hg. 2j) with its attendant plant associa¬
tions is here traced.

Fig. 19. Patches of Acorus calamus, and scattered willow trees (Salix nigra)
mark the source of a stream on the cleared uplands.

1. The TYPICAL RAVINE

The ravine begins almost imperceptibly in the beech woods of the
upland. Its head is broad and rounded, serving to collect the surface
waters from different directions. Upon a cleared upland, a stream often
commences in an Acorns Calamus swamp, around which are a few wil¬
lows (Salix nigra). Such a stream source is shown in fig. zp.

Ecology of the Cincinnati Region

147

First stage. — The shallow valley in the till of the upland contains a
stream, for it will be remembered, the water table is high. This upland
stream is sluggish and meandering. It has many of the marks of old
age. The flood plain which it is building supports a hydrophytic vegeta¬
tion wherever the forest is not very dense. Such plants as Cardamine
pennsylvanica, Asclcpias incarnata, Mimulus ringens, Lobelia siphilitica,
Eiipatorium perfoliatum, Helianthus strumosus, and Bidens laevis, are
found scattered along the stream. The aspect of the upland beech woods
is, however, little modified by the valley, for its slopes are not yet steep
enough or high enough to greatly influence the vegetation.

Fig. 20. The broad open valley of Gunpowder creek.

Across the cleared uplands, the stream is bordered by hydrophytic
herbs, and occasional willows. The plants found are the same as those
along the stream in the woodland, but here they are much more numerous.

Second stage. — Gradually the ravine increases in size. Its banks
become higher, and steeper (5 to 10 degrees). Its flood plain becomes
broader, but less hydrophytic, for the stream is lowering its channel below

148

Ohio Biological Survey

this flood plain, and leaving it as a terrace. The bed of the stream is
limestone. Beech is still dominant on the ravine slopes, and extends down
nearly to the stream (fig. 20). But other trees are beginning to appear,
sugar maple (Acer saccharum) first, and later, wild black cherry (Primus
serotina), tulip tree (Liriodendron Tidipifera) , sweet buckeye (Aesculus
octaiidra), and sometimes white oak ( Qucrcus alba) and red oak ( Quercus
rubra). Occasionally sugar maple or tulip almost entirely takes the place
of beech. If the dominant tree is tulip, the lower zones of the forest are
well developed, because tulip admits a greater amount of light than either

Fig. 21. The tulip tree (Liriodendron Tnlipifera) is the facies in this woods.

beech or sugar maple (fig. 21). This is a mesophytic forest, very differ¬
ent in appearance from the mesophytic forest of the upland. The under¬
growth is more varied and denser, being made up of a large variety of
saplings, among which beech and maple are most numerous ; dogwood
(Cornus dorida) and papaw (Asimina triloba) are abundant. Instead of
the scant herbaceous vegetation of the upland beech woods, a varied
spring flora is found in which Claytonia virginica (spring beauty), Phlox
divaricata, Hydrophyllum appendiculatum, Trillium erectum, Dentaria
laciniata (toothwort), and violets (Viola papilionacea, V. canadensis, V.
striata, and V. pubescens) are perhaps the most characteristic plants.

Ecology of the Cincinnati Region

149

Thus, with the young ravine, there seems to have come an increase
in mesophytism of the forest, because of the protection afforded by the
slopes, which are still too gentle, however, to cause xerophytic conditions.
But this is only temporary ; for, on the steeper parts of the hills, oaks are
appearing. These indicate that as the stream cuts deeper, and its slopes
become steeper, the mesophytic forest must change.

Occasionally on the remnants of the old bottoms, are large sycamores
(Platanus occidentalis) and bur oaks (Quercus macrocarpa) , relicts of a
former hydrophytic association ; but most of the newer vegetation is xero¬
phytic. Red elm (Ulmiis fulva), red-fruited thorn (Crataegus mollis),
locust (Robinia Pseudo-Acacia) , and hop hornbeam (Ostrya virginiana)
grow along the top of the low bluff. The stream is not yet actively cut¬
ting, and has some flood plain. Its bed is only full of water after heavy
rains. Pools remain upon the unequal surfaces of the limestone floor.
Here, and along the margin of the stream, are found a few hydrophytes — •
Eupatorium perfoliatuui, Bidens laevis, several species of Polygonum,
Carex, and Cyperus.

Third stage. — The stream is now flowing about one hundred feet
below the level of the upland, and has traversed about one-third of its
course. Its gradient has materially increased, but is very irregular. For
the heavier limestone layers of the Maysville are fairly resistant, and so
the stream must flow for sometimes a hundred feet over the horizontal
surface of a stratum, before it drops several feet over shale and thin
limestone layers, to another level stretch. The stream is actively cutting,
and it is only on the inside of a sharp curve, that any flood plain is being
built. The higher valley floor has entirely disappeared, and the banks
rise steeply (10 to 20 degrees) from the water’s edge.

A xero-mesophytic association whose coming was indicated higher
up the stream, now occupies the ravine slopes. The valley is open, and
fairly steep sided ; a combination of factors which has produced the xero¬
phytic conditions. In sheltered parts of the slopes, as along tributary
ravines, or on north exposures, a mesophytic association is found. The
soil is thin, so that occasionally rock ledges, or more often loose slabs of
limestone are found. On these a few species of lichens occur. But here
they have little importance in the succession, for soil formation by weath¬
ering of the shales is so rapid that even on fresh exposures, herbaceous
plants and seedlings of trees appear the first year.

On undercut banks, are shown the earliest stages of the xerophytic
succession. Carrot (Daucus Carota), parsnip (Pastinaca sativa), sweet

150

Ohio Biological Survey

clover (Melilotus alba), and wire grass (Poa compressa) are scattered
over the almost bare bank. At the top of the bluff are red elm (Ulmus
fidva), hop hornbeam (Ostrya virginiana) , locust (Robinia Pseudo-Acac¬
ia), red-fruited thorn (Crataegus mollis), blue ash (Fraxinus quadrangu-
lata), and chestnut oak (Quercus Muhlcnbergii) . Toward the upper end
of the bluff, these are seen advancing down the slopes. Fig. 22 shows, in
the left foreground, the upper end of an undercut bank, and in the dis¬
tance, a higher and barer bluff.

Fig. 22. Muddy creek under-cutting its banks ; scattered herbaceous plants on the
bluffs, trees advancing onto the upper end.

As conditions become less severe, and a little soil is formed, a number
of other trees appear, among which are white oak ( Quercus alba), hack-
berry (Celtis occidentalis), redbud (Cercis canadensis), basswood (Tilia
amcricana), and sometimes red cedar (Juniperus virginiana) , red oak
( Quercus rubra), bur oak ( Quercus macrocarpa) , and sugar maple (Acer
saccharum) . Many of the trees which have been mentioned are of low
growth, and it is not long before the oaks reach above them.

Ecology of the Cincinnati Region

151

Then the slope appears to be covered with an oak forest (Hg. 2^), but
beneath the oaks, are all the other trees which were pioneers with them.
Locust does not retain its early importance after the other trees are started,
and is seldom found within the oak forest. The herbaceous vegetation.

Fig. 23. Slopes forested chiefly with oak, Sharon creek.

in spring, is made up chiefly of groundsel (Scnecio obovatus) in large
patches, coral root (Corollorrhiza Wisteriana) , wake robin (Trillum ses¬
sile), larkspur (Delphinium trieorne), rue anemone (Anemonella thalie-
troides), buttercup (Ranunenliis hispidus), sickle-pod (Arabis eanaden-

152

Ohio Biological Survey

sis), and a number of the more xerophytic sedges (Carex laxiiiora, C.
pnhescens, and C. varia). Later, are found species of Desmodium and
Lespedeza, Thaspitim pinnatifidum, Cynoglossiim virginianum, Scutellaria
canescens, Seymcria macrophylla, Ruellia strepens, Solidago ulmifolia,
Aster cordifolius, and Aster Shortii.

This type of xero-mesophytic forest is found only on the steeper
south exposures. Hillsides sloping in other directions and gentler slopes

Fig. 24. Patches of hydrophytic herbs occur along the swampy margins and on
small islands, MuHy creek.

are more mesophytic, and on these the kinds of trees are more numerous.
Sugar maple is an important species, sometimes becoming more abundant
than the chestnut oak. Other trees which may occur are wild black
cherry, sweet buckeye, tulip, basswood, Carya microcarpa, shell-bark hick¬
ory, black oak and red oak. In such a forest, sugar maple is dominant
in the undergrowth. The vernal vegetation is richer than on the more
xerophytic hillsides, and a number of more mesophytic plants are ap¬
pearing.

Ecology of the Cincinnati Region

153

North slopes are always more mesophytic than south slopes of equal
steepness. In some places, the north side of the stream may be bordered
by the xero-mesophytic association above described, and the south side
by a more mesophytic association, in which beech is a secondary species.

The stream bed is broad, and level reaches of limestone are frequent.
The margins of the stream are swampy, and in places on the limestone
floor, are swampy patches where small islands have been built (fig. 24).

Fig. 25. The stream drops suddenly into a narrow gorge of shale. West Fork creek.

Such places carry a hydrophytic vegetation, which is usually varied.
Among the plants which may be found here are Echinochloa Criisgalli,
Leersia oryzoides, Cyperns diandrus, C. escidentus, Eleocharis ovata,
Typha latifolia, Polygonum hydropiper, P. persicaria, P. pennsylvanica,
Mentha spieata, M. piperita, and Bidens laevis.

Fourth stage. — After flowing through this open valley, with its broad
flat bottom, for a considerable distance (as a rule about two miles), the
stream drops suddenly into a narrow gorge of shale (hg. 2y). This is

154

Ohio Biological Survey

never more than ten feet deep at its beginning, but its walls are almost
vertical. The stream has reached the Eden shale, and the character of
the valley has changed. The fall is rapidly receding, carrying the gorge
back into the open valley through the Maysville limestone. The walls
of the gorge are dripping with moisture in many places. But the shale
erodes so rapidly that almost no vegetation is found upon it. Here and
there are patches of algae, and occasionally some moss (Hypnum sp.).
Near the top of the gorge, or wherever the banks are not rapidly cutting,

Fig. 26. Lower course of Indian creek, bordered by a typical marginal vegetation.

are patches of Impatiens and Hydrangea. Instead of the bluff xerophytes
found higher up the stream, there are saplings of sugar maple, ash and
white elm (see fig. ^o). This narrow gorge is mesophytic. But the
mesophytic forest gives way higher up the slope to the xero-mesophytic
forest described above.

The length of this gorge is largely dependent on the altitude of the
Eden-Maysville contact, and of the flood plain of the river, into which
this stream is flov/ing.

Ecology of the Cincinnati Region

155

Fifth stage. — Soon the valley widens out and the slopes become gen¬
tle. Its bed is full of broken pieces of limestone ; its valley floor built up
with rubble, into which the stream cuts during freshets. The flat lime¬
stone reaches, with their groups of hydrophytic plants, are gone. Scat¬
tered sycamores and willows mark the course of the stream.

The valley sides are gently sloping, except near the top, where the
limestone of the Maysville still retains much of the steepness character¬
istic of a younger part of the ravine.

Sixth stage. — The stream emerges into the flood plain of the river.
Its valley, now but a few feet deep, is cut in silt. The stream is sluggish,
and meanders across the flood plain. It is bordered by a typical marginal
vegetation composed of willows (Salix nigra and S. alba vitellina), syca-

B

Fig. 27. Profiles of streams: — A, a normal valley; B, the typical ravine of the
region.

more (Platanus occidentalis) , silver maple (Acer saccharinum) , white elm
(Ulmus americana) , and, in sunny spots, Helianthus strumosus, Ambrosia
triiida, and Silphium perfoliatum (hg. 26). Nothing but the marginal
vegetation belongs to the small stream. The rest is a part of the flood
plain of the river. Here upon the flood plain, the vegetation of the small
stream merges with that of the river.

Because many of our valleys present the features above described,
they are considered as typical of the region. The ^profile of the stream of
this typical ravine (hg. 2y, B) \s not the normal stream profile (hg. 2^, A).
Streams commencing near the margin of the upland, and lacking the
sluggish upland course, have profiles more nearly resembling the normal.

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156

Ohio Biological Survey

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Ecology of the Cincinnati Region

157

2. Modifications of the typical ravine

Many of the ravines of the region do not exhibit all of the features
above outlined ; few have all well developed. Some exhibit peculiar fea¬
tures which are dependent upon the particular physiographic and soil
conditions of the area through which they flow.

Fig. 29. A climax forest in August. Spleenworth (Aspleniiuii angiisnfoiiuui)
forms large patches.

A number of the smaller streams begin near the margin of the up¬
land. In these, the sluggish upland course is lacking. The fall of the

158

Ohio Biological Survey

stream is rapid, and its valley sides much steeper than is the case with
longer streams. Along such streams are found many of our steepest
slopes. The valleys are so narrow everywhere that the contrast between
the parts in the Maysville limestone and in the Eden shale is not often
well marked.

N orthward flowing streams. — North flowing and south flowing
streams of this sort show much the same differences that were noted be¬
tween north and south slopes. Along the northward flowing streams, are
the most mesophytic forests of the region. The tree growth is more
varied than elsewhere, and almost every mesophytic species of the region
may occur. In such a forest, beech forms about fifty per cent of the
tree growth. The remaining fifty per cent is made up chiefly of sugar
maple (Acer sac chariim) , tulip tree (Liriodendron Tulipifera), wild black
cherry (Primus serotina), rock elm (Ulmus racemosa) , big shell-bark
hickory (Carya laciniosa), mulberry (Morns rubra), and basswood (Tilia
americana). On some slopes beech predominates, but rarely forms over
eighty per cent of the forest. The line transect, fig. 28, shows the compo¬
sition of a representative mesophytic forest of this type.

The undergrowth is dense, consisting of saplings of the trees named
above, with a large amount of dogwood (Cornus florida), ironwood (Car-
pinus caroliniana) , papaw (Asimina triloba), and spice bush (Benzoin
aestivalc), and sometimes moosewood (Dirca palustris) and elder (Sambu-
cus canadensis) . Lianas are represented by wild grape (Vitis aestivalis) ,
Virginia creeper (Psedera quinquefolia) and green brier (Sniilax his-
pida). Because of the very dense shade of this forest, summer herba¬
ceous vegetation is scarce. It consists only of a few thin-leaved shade
plants, as Pilea pumila, Impatiens pallida and I. biflora, Actaea alba, Des-
modiuni pauciflorum and D. gran dido rum, Aralia racemosa, and a number
of ferns, often in large patches (fig. 2q). The vernal vegetation is ex¬
tremely luxuriant. The ground is carpeted with dense mats of squirrel
corn and Dutchman’s breeches (Dicentra canadensis and D. Cucullaria),
dog’s tooth violet (Erythronium americanum and E. albidum), twinleaf
(Jeffersonia diphylla), and many other species less abundant.

This mesophytic forest prevails on all slopes up to about 25 degrees.
Steeper slopes have some oaks upon them ( Quercus Muhlenbergii, Q.
alba, and Q. rubra), and the herbaceous growth is less mesophytic. Here
are usually found buttercup (Ranunculus hispidus), Greek valerian
monium re plans), rue anemone (Anemonella thalictroides) , meadow rue

Ecology of the Cincinnati Region

159

(Thalictmm dioicuni), Hepatica (Hepatica acntiloba) and Christmas fern
(Polystichum acrostichoides ) .

Southzvard flowing streams. — The forests developed along the steep
southward flowing streams are always less mesophytic than those of the
northward slopes. It is only in very sheltered spots that a truly meso¬
phytic association is found. Elsewhere, the associations are xero-meso-
phytic or even xerophytic on the steepest slopes. Beech is almost always
absent, and sugar maple is not everywhere abundant. Instead of Dicentra
and Erythronium, are larkspur (Delphinium trieorne), false rue anemone
(Isopyrum biteniatum) , and wild hyacinth (Camassia eseulenta) .

The short steep ravines seldom carry permanent streams. The slug¬
gish lower course is always short. The marginal vegetation, if present
at all, is hardly more than an inward lobe of the marginal flood plain
association of the larger stream.

Absenee of gorge. — In a number of streams of the region, the gorge¬
like valley through the Eden shale is absent. This is due either to the
relatively high level of the stream into which the tributary flows or to the
deep and broad valley filling through which the little stream must cut
during the latter part of its course. The altitude of the Eden-Maysville
contact is a contributing factor in the elimination of the gorge-like valley.

Elongated sluggish eourse. — When the stream must cross a broad
flood plain, or flow through a filled valley, the sluggish part of its course
is often long. This is true of many streams flowing into the Little Miami
below East Eork, and of some flowing into the Ohio. But the greatest
development of this sluggish portion of the stream course is found along
Mill creek valley. In fact, Mill creek flows for most of its length through
the wide pre-glacial valleys of the Ohio and Licking rivers.

Streams at edge of Wiseonsin drift. — Eour streams in the region,
Simpson creek, Sharon creek. West Eork of Mill creek, and Banklick
creek, flow for part of their courses, at or very near the margin of the
Wisconsin drift (Cincinnati Eolio). These streams were in places pushed
southward by the advancing ice, and so forced to cut new channels
through rock. In this part of the course, whether the stream is cutting
through Maysville or through Eden shale, the valley is very narrow and
deep, and the sides steeper than found along the typical ravine. Fig. go
shows the gorge in the Eden shale and the narrow valley of Banklick
creek. The south slope is quite xerophytic, except near the base. The
north slope is damp, and even on the open parts, a number of shade plants
are found.

160

Ohio Biological Survey

The deep lacustrine deposits through which Sharon creek and
Banklick creek cut before entering the gorge, present a habitat different
from any found elsewhere. Here clay slopes of 60 degrees are not un¬
common (fig. ^i). On the newest slopes, nothing but herbaceous vege¬
tation is found. This presents a strange medley of xerophytes and hy-

Fig. 30. A steep sided, but mesophytic valley in the Eden shale, Banklick creek.

drophytes. The ridges are extremely dry, and on them are Desmodium
canescens, Erechtites hieracifolia, Euphorbia maculata, Oenothera biennis,
Aster Shortii, and Equisetiim arvense. Here and there are wet pockets,
in which may be found Lobelia siphilitica, Amphicarpa monoica, and Hy-

Ecology of the Cincinnati Region

161

drangea arborescens. Where shaded, the moist clay banks are carpeted
with mosses, Dicranella and Physcomitrium. On older parts of the
banks, are red elm, sumach, Ostrya, and sugar maple, with sycamore
and willow in the wetter parts. The succession which is here indicated
is very rapid. The most trying conditions are light and instability of
the material. Slides are frequent (dg. gi). Wherever a few trees have
gained a foothold and the ground is shaded, mesophytes appear, such as
beech, sugar maple, and ironwood. The tops of these bluffs retain their

Fig. 31. Steep clay banks support little vegetation ; Sharon creek.

xerophytic character longer than the slopes. They are bordered with
locust, walnut, Ostrya, and oaks ( Qiierciis Miihlenbergii, Q. rubra, Q.
coccinea, and Q. velutina). In many respects this succession resembles
that found on clay river bluffs. It differs only in that it is much more
rapid, because better sheltered.

162

Ohio Biological Survey

B. RIVER BLUFFS

1. Rock bluffs

The development of rock bluffs along streams was indicated in the
ravine, where such bluffs are small and relatively short lived. The suc¬
cession there is rapid, as it is everywhere along a ravine. The rock
bluffs along the rivers (the Ohio especially) are not changing so rapidly.
Their plant succession is slower and exhibits more stages. These bluffs
are not shaded or protected from wind by the opposite bank of the
stream, for the valley is too wide. River erosion is the primary cause

Fig. 32. Under-cut rock bluff on the Miami river at New Baltimore.

of the existence of the bluffs, but their steepness is renewed in two
ways — by under-cutting at the foot of the bluff ( iig. g2 ), and by slump¬
ing high up on the bluff.

Almost everywhere a flood plain of greater or less width has been
built at the foot of the bluff, so that the earliest stages in the develop¬
ment of rock bluffs by under-cutting are seldom seen. Such rock bluffs
are similar to, but much larger than under-cut banks along ravines. The

Ecology of the Cincinnati Region

163

rock exposed is always largely shale, and therefore the earliest stages
of rock bluff vegetation are not shown.

The bluffs of the Ohio rise from 300 to 400 feet above the river,
gently at first but precipitous above. The steepest bluffs and those
showing the youngest stages in the succession, are found in the position
of pre-glacial cols and on the inside of meander curves (Hg. i8). It is
here that slumping is most effective in producing the bare rock bluffs
upon which the pioneer associations start. The Bellevue division of the
Maysville, which is by far the most resistant rock of the formation,
stands out prominently. It outcrops near the tops of the higher bluffs,
there forming almost vertical cliffs, fifteen feet high. The slope above
this is gentle ; below, steep as far down as the Eden-Maysville contact
about 100 feet below, where it changes to a gentler slope on all but the
youngest bluffs.

Along the Miami and Little Miami valleys. Mill creek valley, and
the pre-glacial valleys, are lower rock bluffs, which are seldom as steep
as the bluffs of the Ohio valley.

Pioneer herbaceous association. — Except on the most resistant rock
cliffs, the pioneer vegetation is herbaceous. The only plants are a few
annuals. The earliest associations in the bluff succession may be com¬
pared to those found on abandoned quarry faces. The first plants to
appear are black mustard (Brassica nigra), sweet clover (Melilotus alba
and M. officinalis) , parsnip (Pastinaca sativa), and Aster eric aides. A
few years suffice to cover all but the steepest parts with this herbaceous
growth. The shale, which is everywhere interbedded with the limestone,
weathers rapidly and soon supplies enough soil for woody plants.

The first tree stage. — Ability to withstand strong light is apparently
a factor in determining what trees shall appear. The first are locust
(Robinia Pseudo-Acacia) , honey locust (Gleditsia triacanthos) , red elm
(Ulnius fulva), and blue ash (Fraxinus quadrangulata) . With or soon
after these, are redbud (Cercis canadensis) , and red-fruited thorn (Crn^
taegns mollis). Sycamores are not uncommon as pioneers, appearing
in gullies or along horizons of abundant seepage of ground-water. The
herbaceous plants of the open slopes begin to disappear, and in their

places come the purple aster (Aster Shortii), cleavers (Galium Aparine),
motherwort (Leonurus Cardiaca), catnip (Nepeta Cataria), and wild
onion (Allium cernuum).

All parts of the slope do not reach this stage with equal rapidity.
The succession progresses most rapidly upon the gentler slopes below.

164

Ohio Biological Survey

Here, and on the steeper slopes beneath the cliffs the succession is very
similar. After the first tree stages are passed, mesophytism increases
more rapidly on the shale area than above, and the undergrowth becomes
denser than that found on steeper slopes.

Bellevue limestone cliffs. — In every situation thus far mentioned, the
pioneer stages are passed rapidly. This is not the case on the cliffs of
the Bellevue limestone. No soil is found here except in crevices. The
rock surfaces are covered with crustose lichens (Endocarpon pusilhim
Hedw., V errucaria nigrescens Pers., and Placodhim sideritcs (Tuck.)),
and patches of xerophytic mosses. In the crevices are a few herbaceous
plants, Saxifraga virginiensis, Aster Shortii, Solidago canadensis, and
Poa compressa, with here and there a small red elm, blue ash, or hop
hornbeam. The cliffs have still but few trees upon them, when the suc¬
cession on the slopes below has reached the oak forest stage. Where
the Bellevue does not cap the bluffs, the slopes are often less steep, and
the summits rounded instead of cliff-like. In either case the tops of the
bluffs support a similar vegetation, which is always in a more advanced
stage than that of the steepest slopes. Thus the cliffs may be bordered
above by large oaks ( Quercus Mulilenbergii, Q. rubra, and Q. coccinea)
and sometimes sugar maples a short distance back from the edge.

The succession on the Bellevue limestone cliffs is typical of resistant
rock bluffs. It is similar to that outlined by Cowles (1901, a) as occur¬
ring on limestones and dolomites of the Illinois region, except that the
conifer stage is here entirely absent. The first tree stage is its ecological
equivalent, being followed here, as in the Illinois region, by an oak forest.
The river bluffs nowhere show a later stage in the succession than this
oak forest.

The oak forest. — The oak forest varies somewhat on different expo¬
sures. On south slopes, oak predominates, and with it are some honey
locust, hackberry, blue ash, and red elm. The undergrowth is made up
largely of redbud and Crataegus with saplings of the trees named above.
Sugar maple is occasionally an important constituent of the younger
growth. On north slopes, sugar maple is sometimes a secondary species
of the forest. Sweet buckeye (Aesculus octandra), basswood (Tilia
americana), wild black cherry (Primus serotina), and walnut (Juglans
nigra) are also mixed with oak, which is still the most abundant tree.

Considerable difference is noted in the spring herbaceous growth of
north and south bluffs. On the north slope, the ground cover is dense,
and the plants are those commonly found in mesophytic forests — pale

Ecology of the Cincinnati Region

165

corydalis (Corydalis flavula), waterleaf (Hydrophyllum appeiidiculatum) ,
squirrel corn and Dutchman’s breeches (Dicentra canadensis and D.
cneullaria), celandine poppy (Stylophoruni diphyllum), spring beauty
f Claytonia virginica) , and yellow adder’s tongue ( Erythroniiiin anieri-
canum), — and a few plants of less mesophytic species — larkspur (Delphi¬
nium tricorne), wake robin (Trillium sessile), and bloodroot (Sanguinaria
canadensis) .

Few of these mesophytes are found on the southward facing bluff;
instead are early meadow rue (Thalictrum dioicum), larkspur (Deh
phinium tricorne), twinleaf (Jeffersonia diphylla), wild hyacinth (Camas-
sia esculenta) , shooting star (Dodecatheon Meadia), and Virginia cow¬
slip (Mertensia virginica) .

Logs and outcropping ledges of rock on the north slope are covered
with mesophytic mosses — Mnium affine, and species of Hypnum ; on the
south slope, they are bare except for scattered patches of Grimmia.

Rate of progress in bhiif successions. — Some idea of the rate of
progress of the bluff succession may be gained from clearing successions
of known age. These are of course more rapid than the original succes¬
sions. Many of the river bluffs were cleared by the early settlers and
vineyards planted upon them. In 1864 and 1865, many of these vine¬
yards were cut down, and some of the hillsides left uncultivated.

The hillside above Anderson’s Ferry is one of these. Here the
south slope toward the river varies from 15 degrees to 40 degrees, but
the west slope toward a deep ravine presents most of the features of
steeper bluffs. The river and ravine slopes are forested now with second
growth, and a few old oaks remain near the top. Fig. gg is a view from
the opposite side of the ravine, showing the face of the steep ravine bluff
bare near the top of the hill, and the gentler slope toward the river. The
bare bluff was produced by slumping.

The succession on this south slope has reached the thicket stage.
Redbud (Cercis canadensis) and red-fruited thorn (Crataegus mollis) are
dominant, and with them is a low growth of hackberry, blue ash, honey
locust, red elm, locust, Ohio buckeye, and hop hornbeam mixed with sap¬
lings of chestnut oak (Quercus Muhlenbergii) , the beginning of the oak
stage. The vernal vegetation is very characteristic of warm south slopes.
The dwarf larkspur (Delphinium tricorne) is by far the most abundant
plant. With it are found wake robin (Trillium sessile), wild hyacinth
( Camassia esculenta) , meadow rue (Thalictrum dioicum), toothwort (Dcn-
taria ladniata), Solomon’s seal (Polygonatum bidorum), sickle-pod

166

Ohio Biological Survey

(Arabis canadensis) , wild onion (Allium cerniinm), and blue violet (Viola
sororia).

In large part, the vegetation of the west slope is similar, with this
difiference, however, that the ground layer is more open. The larkspur
is much less abundant, and in some of the steepest places, Viola triloba is
almost the only plant growing under the few scattered trees. Rock
ledges and scattered limestone slabs are lichen-covered, and patches of
xerophytic mosses occur here and there.

Fig. 33. Reforested slopes of a hill at Anderson’s Ferry. The right hand slope
is toward the Ohio river.

At the edge of the bare clifif, are red oak and Ostrya with spring
beauty (Claytonia virginica), meadow rue (Thalictrnm dioicum), alum
root (Heuchera americana) , and early saxifrage (Saxifraga virginiensis)
beneath them (dg. ^4). As the slope becomes gentle above the top of
the cliff, this association gives way to a more mesophytic one. On the
flat hilltop, basswood and ash are found, together with numerous small
sugar maples. Instead of the xerophytic herbs of the steep hillsides, are

Ecology of the Cincinnati Region

167

spring beauty, Dutchman’s breeches (Dicentra CucuUaria) , and patches
of the bladder fern (Cystopteris fragilis).

The flat does not continue long, for the ravine heads to the east.
The north slope toward this ravine, is as steep as the south slope toward
the river. It is somewhat comparable to north river bluffs, though more
protected by the other ravine slopes, and therefore more mesophytic.
The differences in vegetation on the two slopes of this hill are due

Fig. 34. Red oak and hop hornbeam at the edge of the cliff ; early saxifrage and
spring beauty beneath them.

directly and indirectly, to differences of exposure, and the contrast be¬
tween them is marked. Mesophytes form a large proportion of the
forest. Redbud is entirely absent ; the more mesophytic sweet buckeye
(Aesciilns octandra) replaces the Ohio buckeye (A. glabra) ; sugar maple
and wild black cherry are common. Almost no xerophytes are found
among the saplings. The herbaceous vegetation is mesophytic, consisting
of Dice^itra canandensis, Corydalis davula, Cystopteris fragilis, Hydro-
phyllum appendiculatum, and Stylophorum diphyllmn (Celandine poppy).

168

Ohio Biological Survey

The vegetation of the lower part of the slope is even more mesophytic.
But it belongs essentially to the ravine.

Opposite the large ravine, a clearing succession of the same age is
found on east and southeast slopes. The associations here are very
similar to those of south and west slopes, but somewhat less xerophytic.
An east exposure has a richer spring flora than the opposite west slope
of equal steepness. The prevailing winds of this region are west-south¬
west, and their drying ef¥ect is evidenced in this difference in the vegeta¬
tion of slopes sheltered from and exposed to these winds.

Fig. 35. A young bluff of clay and sand on the Little Aliami ; annuals along the
wet (dark) bands, older bluff association in center background, and flood
plain associations to left.

Thus, all the different exposures are exhibited in this limited area.
The succession on the south slope is the slowest ; the associations here
the most xerophytic. While the associations of south slopes are still
pioneers those of north exposures have progressed almost to a climax
forest.

Ecology of the Cincinnati Region

169

If the associations now existing on river bluffs of different exposures
are compared with one another, it is seen that the relations are similar
to those found between the associations of the cleared slopes.

2. Clay bluffs

Rarely one of the rivers of the region may be seen cutting bluffs in
deep clay or clay and sand deposits. These are not the low steep banks
formed where the stream is under-cutting its own flood plan, but high
bluffs cut in till deposits, or in sandy terraces.

Eroding bluffs. — The newer bluffs support little vegetation of any
sort. On banks of stratified sand or clay, some of the bands are usually
wet, and along these, are a few annuals and young perennials (fig.
Occasionally, a row of young willows, cotton-woods, and sycamores is
seen along a wet band, if erosion has not been excessive for a few years.
The vegetation of the till banks is more scattered than that of the stratified
deposits. Each year the banks are undermined by floods, and the plants
that have started are destroyed. At the up-stream part of the cut, where
active river erosion has ceased, perennials and woody plants advance
rapidly. Locust is a common pioneer. Occasionally red cedar is the
first tree to advance, and sometimes these may be seen dotted over the
face of the bluff (dg. g6). The herbaceous vegetation is composed of
the common perennials of dry habitats, and a few more characteristic
herbs — Lepachys pinnata, Siliphium terehinthinaceuni, Lithospermum
canescens, Euphorbia corollata, and Aster laevis — which in this region are
confined to very steep dry banks.

The succession resembles that of the less resistant rock bluffs, but is
more rapid. The greatest vicissitudes are encountered in the early part
of the succession because of the instability 'of the material. Parts of these
clay bluffs usually slump each spring, but during the remainder of the
year, these banks are firm unless interbedded with sand. Slumping plays
an iniportant part in the clay succession, by bringing down plants from
above. Thus woody plants may advance not only from the up-stream
margin of the bluff, but also from slump centers.

A clay or sand bluff rapidly loses its original steepness, and after the
ground is shaded, the succession progresses rapidly. Oaks appear on
relatively young bluffs, and the first forest cover is composed largely of
oaks (Quercus Mtihlenbergii, Q. alba, and Q. rubra).

170

Ohio Biological Survey

Old bluffs. — The old clay bluff is clothed with a forest much resemb¬
ling the mixed oak forest of the typical ravine. The vernal vegetation
usually consists of a large variety of plants, among which may be men¬
tioned Hepatica acutiloba, Ranunculus hispidus, Polemonium rcptans
(Greek valerian), Uvularia grandiflora (bellwort), Ancmonella thalic-
troidcs (rue anemone), Thalictrum dioicuni, Cynoglossmn virginianum,
and Erigeron pulchcllus. Beech and sugar maple are abundant among

Fig. 36. Young clay (till) bluff of the Little Miami river near Miamiville; red
cedar a pioneer.

the saplings, as are also the hickories ( Carya laciniosa and C. glabra viE
losa), basswood (Tilia americana), dogwood (Cornus dorida), and the
oaks which are the facies of the upper layer of the forest. The red
cedar has almost entirely disappeared. In sheltered ravines in the old
bluff, is found a mixed mesophytic forest similar to that found on north¬
ward flowing streams — a forest of the erosion climax type.

Ecology of the Cincinnati Region

171

3. Gravel bluffs

The Little Miami and Miami rivers are, in many places, cutting bluffs
in the gravels with which these valleys were partly filled during the
Wisconsin glacial epoch. The gravel bluffs vary from forty to seventy
feet in height. The under-cut banks stand at high angles, owing to a
partial cementation of the gravel at some horizons.

The new bluffs are devoid of vegetation, except that brought down
from the top by slides, as the loose gravel is constantly slipping (fig. ^y).
At the top of the bluffs, and in those parts where erosion is not very
active, a few xerophytic herbs appear, among which are Verbena stricta,
Kiihnia eupatorioides, Arabis Drummondi, and Poa conipressa. These
are all sun plants, and persist but a short time in the life of the succes¬
sion.

Fig. 37, Gravel bluff of the Little Miami river near Milford.

Toward the upper end of the under-cut bluff, a few xerophytic trees
and shrubs are seen advancing upon the open herbaceous association
37)- Foremost among these are red elm, chestnut oak, blue ash,
hop hornbeam, red cedar, and Rhus canadensis.

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Ohio Biological Survey

Up-stream, the bluff decreases in steepness as it becomes older. At
the same time, the vegetation becomes denser. A young forest made
up of oaks (Qiierciis Muhlenbergii, Q. rubra, and Q. alba), hackberry,
redbud, basswood, thorn (Crataegus mollis), hop hornbeam, and red
cedar, shades the banks. The undergrowth is made up of a few smaller
saplings, and the shrubs, Rhus canadensis, Physocarpus opidifolius (nine-
bark), and Cornus asperifolia. The Rhus is by far the most important
and characteristic of these shrubs, as in this locality it is entirely re¬
stricted to such habitats. It often forms large patches to the exclusion
of other vegetation. The original xerophytic herbs are largely driven
out, and in their places are Aquilegia canadensis (wild columbine), Pha-
celia Purshii (Miami mist). Phlox divaricata, Sanguinaria canadensis
(bloodroot), Arabis canadensis, Silcnc virginica (fire pink), and Aster
Shortii. The numerous bowlders which lie on the surface are covered
with lichens and moss.

Soon a number of marginal trees appear at the foot of the bluff, but
these belong to the life history of the flood plain which is now being
built. The bluff succession in progress is entirely independent of this.
The demarcation between flood plain and bluff is sharp. The original
xerophytes retain their position at the top of the bluffs long after they
have disappeared from the slopes. Above the marginal trees at the
foot of the bluffs, a few mesophytes appear, sugar maple first and then
beech.

The terrace retains its steep and bluff-like margin long after the
food plain has grown wide at its foot, and all stream erosion has ceased.
The mesophytic tendency displayed on the lower slopes is an indication
of what the future forest will be. But even the oldest bluff*s show little
more than a small percentage of mesophytic trees. Oaks predominate,
and in some places along the Little Miami, red cedar is important. Sugar
maple saplings are not uncommon, and sometimes flowering dogwood'
(Cornus dorida) is seen. The herbaceous flora retains much the same
aspect as on the younger bluffs, but is a little more varied and denser.
The summer blooming plants are not abundant, for the steep gravel
slopes are very dry except during the spring when rains are frequent.

The gravel bluff exhibits many peculiarities not seen on bluffs of
rock or clay. Under-cutting banks in gravel are even more unstable
than those in clay ; at least as far as the surface layer is concerned, and
more xerophytic than most rock bluffs. A number of the plants of these
bluffs fRhus canadensis, Physocarpus opidifolius, Verbena stricta, Kuhnia

Ecology of the Cincinnati Region

173

eupatorioides, and Arabis Drummondi) are seldom found elsewhere.
Gravel bluffs and hillsides may be said lo have the most characteristic
vegetation of any habitat of the region.

C. CLEARING ASSOCIATIONS

The use to which a cleared slope has been put influences the char¬
acter of the earlier clearing associations. Pastured land is occupied by a

Fig. 38. Abandoned pasture land in Ault Park; honey locust (Gleditsia triacanthos)
dominant on this 25 degree south slope. The large trees are oaks (Quercus
velutina and Q. Muhlenbergii) and beeches.

turf composed chiefly of Kentucky blue grass (Poa pratensis), over
which are scattered a few scrubby trees and herbaceous plants, among
which ironweed (V ernonia fasciculata) is always prominent. Long after
the pasture land is abandoned, the sod retains its hold (dg. j8). Slowly
the number of broad-leaved herbs increases, and more trees, sometimes
in great variety, appear.

174

Ohio Biological Survey

Cultivated lands, lying waste, grow up in tall weeds, among which
may be a few of the taller grasses, but never blue grass, at least in the
earlier stages. Saplings are more abundant on such areas than on pas¬
ture land.

Land which has been cleared, or partially cleared and not cultivated,
returns to forest much more rapidly than is the case in the two former
instances, doubtless because of the presence of living seeds and roots in
the soil.

The stage in the original succession influences the character of the
new growth in two very different ways. First, the former association
depended for its being upon the nature of the soil, its physical state and
water content, as influenced by the plant cover and by the stage in the
physiographic cycle. The former plant cover affects the new association
because of its modifying influence on the soil. Second, if any of the
original trees remain in the clearing, they will be represented in the new
growth, and often in larger numbers than would otherwise be possible.
For this reason, a few mesophytes are sometimes represented in the first
clearing association.

Clearing always has a retarding effect on the succession. An oak
forest is commonly replaced by one composed almost entirely of the pio¬
neers of new slopes. In many places groves of locust (Rohinia -Pseudo-
Acacia) occupy the cleared slopes. Though not an indigenous tree, this
is probably the most common pioneer of clearings. In bluff successions,
it does not long retain the precedence which it may have originally had,
but is soon replaced by native trees. In clearings there is, however,
often no indication of what will succeed it — the groves are pure stands
of locust. The predominance of locust, when once it is introduced into .
a clearing, may be partially accounted for by the rapid vegetative propaga¬
tion of this tree by root shoots,^ which gives it an advantage over other
bluff xerophytes.

Other common clearing pioneers are honey locust ( Gleditsia triacan-
thos), thorns (Crataegus mollis, C. Crus-galli, and C. punctata), Sassafras
and red elm. These may be found together composing a mixed copse,
or separately forming almost pure stands (dg. g8).

In a few localities, especially in the Little Miami drainage basin, red
cedar (Juniperus virginiama) is the facies of the early stages of slope
successions, in clearings as well as on young bluffs. Locust may be

® The terms water shoot, water sprout, shoot, sprout, and sucker, are largely used indis¬
criminately for shoots from roots, both at and far from the main stem. It is here sug¬
gested, that root shoot be used to designate those shoots arising from roots, at a distance
from the original plant.

Ecology of the Cincinnati Region

175

mixed with it in greater or less abundance, but in older associations, the
red cedar stands almost alone (fig. jpj. In other places it is mixed with
white and chestnut oaks, in which case it forms a very prominent and
important part of the young forest. In most of the area under consid¬
eration, however, red cedar is not even a secondary species in any of the
pioneer associations, and never becomes the facies. Its limited distribu¬
tion within the region is not understood.

Fig. 39. Red cedar (Jtinipcrus virginiana) is the facies in this clearing association
(northeast of Milford).

The direction and steepness of the slope, together with the nature of
the soil, influence the rapidity of the clearing succession, as they do the
original succession. Slopes of from 20 to 30 degrees usually retain their
xerophytic flora long after gentler slopes have become mesophytic.
Steeper slopes seldom ever become mesophytic. It is in this tension zone
of intermediate slopes that some of the most interesting clearing succes¬
sions are seen. It is here that the influence of soil and exposure is most
strongly felt.

176

Ohio Biological Survey

A slope of 30 or 40 degrees may be covered with a second growth
mesophytic forest. The direction of slope must be northerly — northeast,
north, northwest, or rarely east or west. The soil must be deep, and of
such a nature that it retains water. These requirements are met along
a number of ravines, which cut in deep glacial deposits composed largely
of fine sand and silt. A single clearing formation changes from a xero-
mesophytic forest association on the south slope to a mesophytic one on
west and northwest exposures of equal steepness. The succession passes
from an association in which locust and sumach (Rhus typhina) are domi¬
nant among the trees, to one in which beech, sugar maple, and tulip are
most abundant. Very few species are common to the two associations.
With the locust and sumach of the south slope are red elm, hackberry,
and mulberry. Blackberry thickets form a lower layer in this open forest,
and on the ground are a few herbs among which are V enus’s looking-glass
(Specularia perfoliata) , wild pansy (Viola RahnesquU) , the ebony spleen-
wort (Asplenium platyneuron) , and some Cystopteris fragilis. On the
west and northwest slopes, with the beech, maple, and tulip, are dogwood,
sweet buckeye, iron wood, a few large chestnut and red oaks, and bass¬
wood. The herbaceous flora is not that of a climax forest, but is remark¬
ably different from that of the south slope. Here Cystopteris fragilis
is common, and with it are the maidenhair fern (Adiantum pedatum),
Jack-in-the-pulpit (Arisacma triphyllum) , bloodroot (Sanguinaria cana¬
densis), May apple (Podophyllum peltatum), black snakeroot (Cimicifuga
racemosa) , and Solomon’s seal (Polygonatum hidorum).

Later stages in the clearing succession are very similar to corre¬
sponding associations of the original succession.

D. THE EROSION CLIMAX EOREST

The mesophytic forest which is found upon a number of the gentler
or more protected slopes, it has been shown, differs from that of the
uplands. It contains a larger variety of trees, and its herbaceous growth
is extremely rich and varied (dgs. 28, 2^). The relatively simple life
history of the upland forest is not found here. Topographic changes
have played an important part in the succession. Retrogressive periods
must have had some place in it. This is the mesophytic forest which
develops at the end of a long and varied succession. It is the climax
of every erosion succession now in progress, and as the forces of erosion
must sooner or later modify all areas, it may be considered as the prob¬
able climax forest of the region. In contrast to the pre-erosion climax

Ecology of the Cincinnati Region

177

forest of the upland, is this forest, which will be termed the erosion
climax.

IV. TERRACES AND FILLED VALLEYS

Combining, in a measure, the features of uplands and of slopes, of
pre-erosion and erosion vegetational types, is a topographic division of
great prominence, and a vegetation series of somewhat intermediate
character. The flats of abandoned river valleys and the younger glacial
terraces of present streams have not been greatly modifled by the forces
of erosion. Older terrace remnants are much dissected, and their vegeta¬
tion belongs more properly to the slope series.

The terraces and pre-glacial valleys of the region show less of their
original vegetation than any other equal area in the region under con¬
sideration, for they are very favorably situated for both agricultural and
commercial pursuits. The discussion is necessarily based on isolated
and sometimes poorly preserved remnants.

Terrace associations may be grouped into two classes, the associa¬
tions of gravel terraces, and the associations of sand and silt terraces,
each dependent on the physical character of the terrace material. Between
these two classes, there are a number of intermediate forms for the ter¬
race materials intergrade.

A. GRAVEL TERRACES

Gravel terraces are limited to a few small areas. Most of these lie
along the Miami and Little Miami rivers. Because of their very favor¬
able situation above the reach of floods, and their nearly level surface,
the terraces have been cleared for many years, and it is only in a few
spots that any of their original vegetation remains.

1. Forest associations

A few large trees remain upon the terraces. These are usually
beech, but here and there is an occasional wild black cherry or sugar
maple. From these few remnants it would appear, that the original
forest of the gravel terrace was dominated by beech, and probably resem¬
bled the beech forest of the upland. Near the top of river bluffs or of
ravines in the terrace, the mesophytic forest is replaced by one resembling
the forests of the margins of uplands!' Red oak, white oak, chestnut
oak, and sometimes red cedar replace the mesophytes of the level terrace.

178

Ohio Biological Survey

2. Clearing associations

Most of the clearings are occupied by a dry meadow association
different from that found in any other location. Wire grass fPoa com-
pressa ) is the most common grass, except in some of the older clearings,
where Kentucky blue grass fPoa pratensis) almost supersedes it. Sap¬
lings are usually scarce, and represented only by a few red elms, locusts,
and red cedars. It is among the herbaceous vegetation that the charac¬
teristic plants are found. Kuhnia eupatorioides, Verbena stricta, Arabia
drummondi, Pentstemon hirsutus are the most characteristic. With these,
some other widespread herbs occur.

These cleared fields are covered with the layer of soil which has
accumulated since the deposition of the gravels. Although xerophytic,
they do not present the extreme condition of dryness of exposed gravels.

Gravel pits are very artificial habitats, but possibly they present con¬
ditions somewhat parallel to those existing before the accumulation of
the soil cover. The material is so coarse and loose, that almost no water
is retained near the surface. In no place, except on under-cut gravel
bluffs or on bare rock, are conditions of ecpial dryness found. Only the
most extreme xerophytes can exist. Even these are not abundant. The
association is an open one. The ordinary xerophytes of the region, which
usually occur in dry situations are stunted. Verbena stricta, occasional
plants of Pentstemon hirsutiis, Oenothera laciniata, Croton monanthogy-
nus, and stunted Oenothera biennis make up the open gravel association.
In a few places, the prickly pear cactus, Opnntia Rahnesquii, is the facies
of the association, and Yucca iilamentosa or Andropogon virginicus, pri¬
mary species (hg. 40).

On long-abandoned gravel pits, the association is less open, and
comes to resemble the dry meadow association of clearings.

The facies of sun plant formations of gravel terraces is not the same
everywhere. In some localities, a usually common species may be absent,
and others so restricted in range that they are not even mentioned here,
may be dominant. Of all the herbaceous plants, Verbena stricta is by
far the most widespread and characteristic plant of gravel terraces, dr
other gravel deposits. It is, as far as known, restricted to such habitats.
JArbena angnstifolia, which occurs frequently in upland meadows, never
occurs in the fields on the terraces. The two plants are entirelv distinct
in range, the former species (V. stricta) never occurring on uplands,
except on the gravel roadbeds of some of the railroads. Furthermore,
the species are sharply defined, and intermediate forms, such as are found

Ecology of the Cincinnati Region

179

at Stony Island, Chicago, do not occur between the plants of the two
habitats.

B. SAND AND SILT TERRACES

The forest cover of the sand and silt terraces is similar to that sup¬
posed to have been the original cover of gravel terraces. It is a climax
mesophytic forest somewhat intermediate between the pre-erosion and
erosion types. Beech is usually dominant, growing with sugar maple,
tulip, sweet buckeye, and walnut. Tulip, in one instance, becomes the
facies, constituting nearly 80 percent of the tree growth.

Fig. 40. A clearing association on the gravel terrace; Opuiitia rafiiiesquii the facies.

The undergrowth of this mesophytic forest of the terrace is rela¬
tively dense, consisting of saplings of the trees noted, with summer grape
(Vitis aestivalis) , poison ivy (Rims Toxicodendron) , spice bush (Benzoin
aestivate), and mesophytic herbs, among which blue violet (Viola papilio-
nacea), waterleaf (Hydrophyllnni appendiculatiini) , black snakeroot
(Cimicifnga racemosa), richweed (Pilca pumila), Joe-pye weed (Eupa-

180

Ohio Biological Survey

toriinn pnrpurcni ) , and white snakeroot (Eiipatorium ^irticacfoliiim) , are
abundant.

The sand and silt terraces differ from those of gravel, in that they
lack the usual xerophytic margin of the latter. The trees of the flood
plain are followed immediately by the mesophytes of the terrace, even
where there is considerable difference in the altitude of the flood plain
and terraces.

Clearings on these terraces have nothing characteristic about them.
The associations are not extremely xerophytic, as are those of gravel ter¬
races, and the characteristic herbs of the latter are not found on these
sand and silt terraces.

C. FILLED VALLEYS

The pre-glacial valley of the Ohio, which is partially filled with deep
glacial deposits, is divisible into three parts, differing somewhat in soil
features and plant associations. One of these commonly known as the
Norwood trough, extends from the Little Miami in the vicinity of Plain-
ville and Red Bank to Alill creek. This part of the valley is filled with
deposits of Illinoian age, covered with a thick mantle of loess.

A second portion of this valley, includes the upper part of Mill creek
valley from about Hartwell northward to Hamilton, and the belt of low
hills on the west side of the valley. The hills are of Wisconsin till, the
valley filled with undifferentiated glacial deposits.

A part of the former valley of the Ohio, which is not included in the
present discussion, is now occupied by the Miami river, from Hamilton
almost to New Baltimore. Nothing but recent alluvium is exposed in
this valley, and its associations belong almost entirely to the flood plain
series. The remainder of the valley, extending from the Miami in the
vicinity of New Baltimore westward to the Whitewater river at Harrison
is filled with till of Illinoian and Wisconsin age.

Other valleys of the region were filled in the same way. Much of
the glacial filling has since been removed from those now occupied by
large streams, but in many places along these valleys, small areas of deep
drift deposits remain. The vegetation of these areas is considered under
clay bluff's.

In general features, the vegetation of all filled valleys is similar.

1. The Norwood trough

The plant associations of the Norwood trough are most closely re¬
lated to those of the uplands covered with Illinoian till. Undissected

Ecology of the Cincinnati Region

181

flats (here of limited extent) and small depressions occur, as well as
gentle ravine slopes.

Poorly drained and drained areas of slight relief. — The few depres¬
sions which are found in the Norwood trough are usually smaller, but
more decided than those of the upland. These not uncommonly contain
shallow ponds of small size in which the only aquatic plants are algae.
They are bordered by the usual marginal plants — buttonbush ( Cephalan-
thiis oeeidentalis) , bitter cress (Cardamine pennsylvaniea), poison ivy,
and sedges. There is no gradation between depression vegetation and
that of the surrounding area. A swamp may succeed the pond, by the
closing in of the marginal vegetation, but this is not succeeded by meadow.
The area is so small, and usually so well shaded, that the plants of the
surrounding forest can occupy the area as soon as it is dry enough.
Occasionally, there are a few hydrophytic trees around the depression.
These are the same as found on the uplands — Nyssa sylvatica, Quercus
hieolor, and Q. palustris.

On the flat and undissected portions, drainage conditions are poor,
and a wet meadow association much like that of the uplands prevails in
the cleared areas. The environs of Norwood — the divide between east
and west drainage — probably exhibited in the largest degree, the features
common to flat uplands, but here little of the original vegetation remains.
Other flat areas of the valley exhibit the upland forest types. Both a
mixed hydrophytic forest similar to that of uplands, and a beech forest —
the pre-erosion climax — occur here.

Disseeted areas. — The marginal parts of the Norwood trough are
so well dissected that they present none of the features of uplands. The
relief is never great, however, and ravine slopes in the clays are gentle
except on under-cut banks. The forests of these areas are therefore
prevailingly mesophytic. Beech is often, if not usually, the dominant
tree, but with it are many of the other most pronounced mesophytes.
This forest extends a short distance up the slopes of surrounding hill¬
sides, usually to the top of the gentle slopes of the till. It is in strong
contrast to the oak forest of the rest of the hillside. The view across the
Norwood trough (fig. 41) shows a number of forested areas in the valley,
in all of which, beech is dominant. The beech forest is also seen on
the lower slopes of the hill in the foreground. The oak tree in the left
foreground higher up this hill, is a remnant of the oak forest characteristic
of rock slopes.

182

Ohio Biological Survey

It is only along the crests of narrow divides, or upon unusually steep
slopes, that the beech forest is superseded by a xero-mesophytic or mixed
forest. Such areas are, however, always small.

The prevalent forest of dissected areas is a mesophytic forest inter-

Fig. 41. View over the Norwood trough from top of valley slopes. Beech is dom¬
inant in all the wooded areas ; the oak ( Quercus muhlenbergii) in the fore¬
ground belongs to the slopes.

mediate in character between the pre-erosion and erosion types, but most
closely approaching the latter. Its development has been more rapid,
with consequent greater simplicity of the forest, than has that of the
erosion climax forest previously discussed, but it nevertheless represents
an erosion type.

2. Upper Mill creek valley area

The valley. — A part of the broad flat valley of upper Mill creek is
covered with recent alluvium, and belongs to the flood plain area. In
the rest of the valley, glacial deposits are at the surface. Nothing of the
original vegetation of this part remains. It is all farm land.

Ecology of the Cincinnati Region

183

Another pre-glacial valley, filled with deposits similar to those of
upper Mill creek, lies for the most part, north of the area considered.
It runs from the Little Miami above Kings Mills, past Glenwood, to the
Miami at Middletown. This valley, whose greatest relief is scarcely
more than twenty feet (similar in this respect to upper Mill creek valley),
contains a few forest remnants which indicate the probable previous vege¬
tation of Mill creek valley.

On regions of low relief, is a mixed mesophytic forest composed of
various oaks ( Quercus ruhra, Q. Muhlenhergii, Q. macrocarpa, and
Q. alba), Kentucky coffee-tree (Gymnocladus dioica), white ash (Fraxi-
nus americana) , and hickory (Carya alba), but containing no beech, and
only an occasional sugar maple. Although the relief is very slight, the
forest is less mesophytic than that of steeper slopes elsewhere, for the
soil is gravelly. Verbena stricta, which is so characteristic of gravel ter¬
races, is not uncommon.

Hydrophytic associations of this valley belong to two opposing types,
those of half-drained and those of undrained situations — depressions and
flats removed from streams (Cowles, 1901).

Depressions contain swamp white oak (Quereus bieolor), white oak
(Quercus alba), pin oak (Quercus palustris), and red maple (Acer
rubrum).

Half-drained swampy areas are not met with elsewhere in the region
except along the margins of some streams, and in very limited areas
around springs or on springy slopes. In the vicinity of Glenwood, how¬
ever, are extensive flats through which small sluggish streams wander in
beds from one to three feet below the general level (dg. 42). The area
occupied by the swamp is not a flood plain, yet the plant associations are
much like some of those of flood plains. The forests are dominated by
silver maple (Acer saccharinum), with an occasional sycamore (Platanus
occidentalis) near the stream, and a few trees of black ash (Fraxinus
nigra), and swamp white oak (Quercus bicolor).

Clearings, and openings in this forest, support a luxuriant herbaceous
vegetation, different, however, from that of undrained depressions and
wet upland flats. There are but few plants of button-bush ( Cephalanthus
occidentalis) — a character plant of undrained situations. On the other
hand, Bidens laevis, Eupatorium perfoliatum and Verbena hastata — all
typical brookside plants — Iris versicolor. Hibiscus Moscheutos, Physos-

184

Ohio Biological Survey

tcgia virginiaiia, Sparganiiim curycarpiim, Typha latifolia and many
sedges, are abundant (^ig. 4g).

This area is swampy, not because there are no streams to carry off
the surface waters, as on uplands, but because the water table is always
near the surface in this low-lying area of slight relief. Instead of the
gradual sinking, and spreading of ground water, as under the flat uplands,
there is a constant flow from the hills bordering the lowland area under
the swamp lands to the shallow drainage lines.

Fig. 42. The swamp at Glenwood extends to the foot of the distant hills; Hibiscus
in bloom ; shrubs are Salix nigra.

In all probability, upper iMill creek valley, and the Little Miami-
Miami pre-glacial valley supported similar types of vegetation. Leverett
(1902) speaks of a “marshy tract” in Mill creek valley above Sharonville,
which was still undrained and not fit for cultivation in 1889. Water in
wells in IMill creek valley stands practically at the surface (Leverett,
1902), and the same condition is found near Glenwood.

Glacial hills. — The low glacial hills along the western side of the
valley are a continuation of the morainal hills of the upland. In origin

Ecology of the Cincinnati Region

185

and topography, they are related to these. In situation within an old
filled valley, they are a part of the latter. Their vegetational features are
to some extent a combination of those of both areas. Their slopes are
always very gentle (3 to 8 degrees), their tops broad and rounded.

Fig. 43. Near view in the same swamp, showing Hibiscus Moschcufos and Scirpus
atrovirens.

The forests of this area are mesophytic, and for the area as a whole,
beech is the dominant tree. There is, however a larger admixture of oaks
than in the forests of the Norwood trough; indeed, the forests are less
mesophytic than those of the latter area. Beech occupies the tops and
the longest and gentlest slopes of the hills and much of the space between
the hills. With the beech on the tops of the hills, are other trees, the
most important of which are sugar maple (Acer sacchanim) , big shell-
bark hickory (Carya laciniosa) , white oak (Qucrciis alba), and red oak
( Querciis rubra). On steeper slopes, beech is absent, and then the above-
mentioned trees are relatively more abundant. On the flats between the
hills, the hickories (Carya ovata, C. alba, and C. cordiformis) , bur oak

186

Ohio Biological Survey

(Quercus macrocarpa) , and white ash (Fraxiniis americana) occur with
beech.

In the upper Mill creek valley area, low hills are built up above the
general level ; in the Norwood trough, valleys are cut beneath it. The
soil of these hills — a compact but gravelly till without a covering of
loess — is more porous than the Illinoian till and the loess of the Norwood
trough. These two factors — relatively small area of hill top, and porous
soil — tend to make the water table lower than in areas of equal or even
greater relief in the Norwood trough, and therefore, the forests less
mesophytic.

Many of the streams in this area wander among the hills, finding
rather than making their valleys. The larger streams which cross the
area have, however, cut fairly deep valleys in the till. These valley slopes,
though gentle (6-10 degrees) are not clothed with beech forest, as was
the case on ravine slopes in the Norwood trough. In fact, beech is
always entirely absent, for the porous soil of this gravelly till drains
readily. The forest here is composed of oaks ( Quercus Muhlenhergii,
Q. velutina, Q. texana, and Q. alba), sugar maple, walnut, and white ash.
A short distance back from the edge of the ravine slopes, beech is again
dominant.

3. The Miami-Whitewater valley

Much of that part of the Miami-Whitewater valley covered with till
of Wisconsin age, is flat. In places there is a poorly-developed sag and
swell topography. The valley filling is partly water-laid ; gravel is an
important component in much of the area.

Streams start in indefinable depressions much as do those of the flat
uplands. Their margins are swampy and they are bordered by willows
(Salix nigra and N. longifolia) , cat-tail (Typha latifolia), ditch stonecrop
(Penthorum sedoides), swamp milkweed (Asclepias incarnata), monkey
flower (Mimuhis ringens), and boneset (Eupatorium perfoliatum) .

The area is little dissected, except near the margins adjacent to the
low-lying flood plains. These slopes are terrace edges, and as the till
contains a relatively high percentage of gravel, the vegetation is that of
gravel slopes.

The forests of the Miami-Whitewater pre-glacial valley are essentially
like the mixed forests of undrained uplands. Here, as on the uplands,
is a pre-erosion forest type. Groups of red maples (Acer rubrum) occur
in a mixed forest of oaks ( Quercus alba, Q. velutina), sugar maple (Acer

Ecology of the Cincinnati Region

187

saccharumj, sour gum (Nyssa sylvatica), shell-bark hickory (Cary a
ovata) , and white-heart hickory (Carya alba).

The most striking feature of this area, as well as of the valley of
upper Mill creek, and the Little Miami-Miami valley, is the absence of
beech. It is found neither on the flats nor on the slopes. The forests,
though mesophytic, are far from either climax type.

V. THE ELOOD PLAIN SERIES

In places along the ravine course, small areas of flood plain with
their attendant vegetation were noted. Along the rivers of the region,
flood plains are sometimes several miles wide. The vertical range is
greatest along the Ohio, which has an annual variation in height of about
sixty feet. It is the flood plains of the larger streams — the Ohio, Miami,
Little Miami, and Licking rivers, and Mill creek — which constitute one
of the four great topographic and vegetational areas of the region. Along
these rivers, flood plain vegetation reaches its greatest development.

Although the beach-like shore and the islands are genetically a part
of the flood plain, it is convenient here to separate the frequently sub¬
merged and open shore associations from the closed associations of the
broad and higher flood plain.

A. THE BEACH-LIKE SHORES

The river shores here discussed are comparable to lake beaches
(Cowles, 1899). Their vegetation is controlled by a similar set of fac¬
tors, most important of which are topography, moisture, and varying
river stages. The latter aids in producing zones comparable to the
wave zones of lake shores, but as the motion of the water is slower, the
corresponding zones usually have more plants.

The lower part of the shore — the first vegetation zone — is submerged
many times each year and not infrequently during the growing season.
It may be compared with the lower beach of lakes, although it usually
supports a scant growth of annuals, most of which are hydrophytes.

A higher part — the second vegetation zone — corresponds to the mid¬
dle beach, with its xerophytic annuals. This is not often submerged
except during the winter and early spring.

That part of the Ohio river affected by the Fernbank dam (Dam 37)
has a nearly constant summer height. Here the lower shore is con¬
stantly submerged, and consequently the first vegetation zone is almost

188

Ohio Biological Survey

absent, the upper shore association approaching close to the water’s edge.
Vegetation is more abundant and more luxuriant than on the upper
shore elsewhere, and some hydrophytes are included among the annuals.

1. Gently sloping beach-like shores

In many places the river shore is broad and rises gently from the
water’s edge. Along the Ohio, such shores are of sand or silt. On the
two Miamis, they are usually of gravel or cobblestones with more or less
mud. There is not a great deal of difference in the vegetation of these
two kinds of shores.

Sandy shores. — On sandy shores, the density of the vegetation in¬
creases up-shore. The demarkation between lower and upper shore is
indistinct, so that there sometimes appear to be no definite zones below
the first tree zone. The wettest places are occupied by Eleocharis oliva-
cea. On other parts of the shore, chiefly the second zone, may be found
a variety of plants, among which are a number of grasses fEchinochloa
Crusgalli, Panicunt dichotomiflonim, Cenchrus tribuloides, and Eragrostis
hypnoides) , knotweeds, (Polygonum pennsylvanicum and P. lapathifo-
lium), Amaranthus hyhridus, Strophostyles helvola, Ambrosia trihda, and
Bidens laevis. Here and there are seen a few small willows ( Salix nigra
and S. longifolia), and cotton- wood (Populus deltoides) advancing into
the zone of annuals. These presage the advance of the marginal tree zone.
The width of the zone of annuals is determined by the slope of the shore
and the average summer fluctuation of the river. The plants of the
shore are mainly transients. They are plants which come late, after the
rivers have gone down to low summer heights.

Cobblestone shores. — The shores of gravel or cobblestones are occu¬
pied by an association of herbaceous plants very similar to that found on
sandy shores. The principal differences are that there is no definite
advance of woody plants from a distinct tree zone, and that the upper
shore association is more xerophytic. The willows are few and scattered.

Along very flat bowlder shores, there is sometimes a marginal zone
composed of Dianthera americana, advancing out into the river (hg. 44).
This is usually found where the shore is narrow, and bordered by bluff¬
like banks of the flood plain. In such places the shore is shaded by the
tall trees along the banks, and much of the vegetation characteristic of
sunny shores is absent.

The second zone of cobble shores is occupied by an open association,
in which herbaceous plants have a prominent place. In contrast to the

Ecology of the Cincinnati Region

189

lower shore, which is occupied by hydrophytic herbs, is the rough upper
shore, with its xerophytic annuals and perennials. With the scattered
willows, sycamores, and poplars of this upper zone, are clammyweed
(Polanisia graveolens) , blue false indigo (Baptisia australis), crab grass
(Digitaria sanguinalis), black mustard (Brassica nigra), peppergrass
(Lepidium virginicum) , sweet clover (Mclilotus alba), and evening prim¬
rose (Oenothera biennis). Fig. 4j is a view of a wide cobble shore of
the Little Miami; fig. 4p, AA' , a belt transect of the same.

Fig. 44. Narrow flat shore at the foot of steep wooded banks of the flood plain.
Water willow (Dianther a americana) advancing out into the river.

The third zone is a closed association similar to the corresponding
zone of sandy shores. This belongs more properly to the flood plain
forest than to the beach-like shore.

2. Steep river banks

The open shore association extends higher up on steep river banks
than on gently sloping ones. But this shore association is not the same

190

Ohio Biological Survey

as that previously described. In vertical range it corresponds approxi¬
mately to the first three zones of the flood plain forest to be described
later. It is itself divided into distinct belts.

Bed-rock banks. — These steep shores are of several kinds. In rare
instances they are of bed-rock (Pt. Pleasant) in position, with large
pieces broken off and scattered over the surface. Such banks have very
little vegetation. A few annuals in the crevices of the rock, and scattered
willow bushes are all that is found.

Fig. 45, Beach-like shore of cobblestones on the Little Miami river near Gravel Pit.

Rocky banks. — A later development of this type of river bank is
sometimes found. Instead of ledges of rock, and large blocks lying
around, the banks are covered with small angular fragments (dg. 46).
The slope of this kind of shore averages about 12 degrees. There are
more plants here than on the bed-rock banks, but the association is often
open. For a few feet back from the water’s edge, the banks are almost
bare. This zone corresponds approximately to the first zone of gently
sloping shores.

Ecology of the Cincinnati Region

191

Above this are a number of belts composed largely of xerophytic per¬
ennials, among which Desmanthus illinocnsis is the most abundant. Other
plants commonly found on such banks are Baptisia australis, Apocynum
cannabium, Tecoma radicans, Salix longifolia (low bushes), Echinochloa
Crusgalli, Panicum dichotomidorum, Aster ericoides, Xanthium canadense,
and Ipomoea hederacea. These belts, with increasing density, and larger
plants, extend upward about forty feet, or to the foot of the sudden rise
bordering the flood plain. They correspond to the second and third zones
of other shores.

Fig. 46. A steep rocky river bank near North Bend on which Desmanthus illinoensis
and Baptisia australis are common.

On the low steep banks bordering the flood plain is a zone of trees,
not marginal trees, but the bluff trees — red elm, hackberry, black locust,
scarlet thorn (Crataegus coccinea), and honey locust.

Although submerged at some periods of the year, the plants of the
rocky shores are not hydrophytes. During the summer months, these
banks are dry. They are in some ways similar to the high cobblestone
shores. The perennials of such localities are deeply and strongly rooted.

192

Ohio Biological Survey

Such banks as these are not depositional. They are found only along
eroding parts of a river’s course. These herbaceous plants, which present
no obstruction to flood waters, retain their foothold, even when the willows
of the same association, or the trees higher up, are torn out.

Steep river banks are found at or near the foot of b’ufifs. They are
always narrow and often short-lived, as a flat beach-like shore may be
built out in front of them.

Under-cut banks. — All the streams except the Ohio are under-cutting
their flood plains. These under-cut banks are comparable to bluffs in
steepness, but they belong essentially to the flood plain area. Whatever
vegetation may appear upon them during one year is swept away the next
spring. It is only after under-cutting ceases, and the slopes become more
gentle, that the flood plain vegetation gets a foothold.

The plant associations of these steep river banks in silt or sand re¬
semble most closely those of the gently sloping shores in the same mate¬
rial. On the narrow flat at the base of such banks are the hydrophytic
herbs of the broader shores. Above this are the other associations of
gently sloping banks. These are often so intermingled that they can not
be separated into zones (fig. 44). In other places the zones are very nar¬
row and at times one or more are eliminated entirely.

P>. ISLANDS

The islands of the Miami and Little Miami rivers are residual — rem¬
nants of a former valley Ailing, partially reworked, but produced by ero¬
sive rather than by depositional forces. They vary in size from a few hun¬
dred to several thousand feet in length. It is only the smaller ones which
can properly be called islands. The larger ones are areas of the flood
plain separated from the mainland by cut-offs (sloughs). They can be
considered as islands only in that they are bounded on all sides by parts
of the river and that their origin is the same as that of the smaller islands.
It is the smaller islands which will be considered here.

All gradations between the island and the cobble shore are found.
Many projecting stretches of beach-like shore are separated from the
higher flood plain by a depression, which during the summer is either
swampy or partly filled with water (fg. 4/). Some stretches of shore
connected with the mainland in but one or two places become islands dur¬
ing high water. The islands may be well in the center of the river so that
the waters divide nearly equally in passing them (fg. 42), or so much to
one side that the river is very unequally divided. Sometimes the shallow

S' - Zo2'l7. Tiig'ra, Black VJiUow

longzfol'idjSoirLd 3ur WiUour
Tda f a. n u s occide nlahs. Sgc a mort .

M-Acer saccJiari num,Si h-er Ma pie.

L -GhdiBia IriacanlhoSjHoney Locust
H -Celii 3 occidenlahSyHacklerry

VJ-Juglaris nigra^ Black Walnut Bk-iree , size indicated }y circles.
C- Prunus seroiina, Wild Black Cherry. 0 - Lush , or shrulrly tree
E- Ulmus amencana, While Elm. d - herlaceous plants.

J' Uj. 4 < .

Map showing general dislriljution of trees and shrubs on island and neighboring shores; Little Miami liver near Gravel "The

river banks, is taken as base, and the fall of the river, 1 to 3 feet at each rapids, disregarded. Contours therefore also indicate vegetation nones.

Ecology of the Cincinnati Region

193

channel at one side of an island is filled up during a flood, and the island
becomes a part of the shore. And, vice versa, a projecting portion of the
shore may be suddenly transformed into an island by the gouging out of
a channel across it. All parts of an island formed in this way are of
very nearly the same age. For this reason, the normal asymmetry of
vegetation of river islands as noted by Cowles ( 1901 ) , is not seen here.

Fig. 48. The steep lower end of one of the residual islands of the Little Miami
river ; near Gravel Pit.

Instead, the zonation of associations is radial. The number of zones is
dependent on the height and size of the island ; their width upon the
steepness of the island’s shores. Fig. 48 shows the lower end of the
island in the map (fig. 4/). In this case, the lower end of the island is
steep, and resembles the eroding portion of depositional islands. The
upper end of this same island has very gently sloping shores, occupied
by the first two vegetation zones, both open associations. Tree zones
occupy the higher parts of the island — zone III that part between the 15
and 20 foot contours, and zone IV, the area above 20 feet (fgs. 47 and
49, BB' and CC).

194

Ohio Biological Survey

Factors limiting associations of cobble shores. — The plant associations
of the cobble or gravel shore, whether of mainland or island, are not con¬
trolled by newness of land, but by water height and topography. The
lower shore association occupies the area which is exposed during low
water. The second zone, or upper shore, is submerged in the summer
only for very short periods, if at all, during unusually high water. The
lower shore is not being built up or extended outward by deposition ;
neither are the islands increasing in size by gradual accession of deposited
material at their lower ends. The succession is not related to slow con¬
structional changes, as is the succession on depositing shores or deposi-
tional islands ; nor is it accompanied by gradual encroachment of one
association upon the next younger. The streams along which the cobble
shores, and in which the residual islands are found, are essentially eroding
streams. The lower shore is cut below the upper shore ; the upper shore,
cut below the general level of the flood plain. The materials of which
the islands are composed are too coarse to be moved during low water,
even by the swift current assumed by these rivers in passing through the
narrow channels around the islands. Changes are effected only during
high water, when the cutting and carrying power of the stream is enor¬
mously increased. Nevertheless the beach-like shores and the residual
islands of the two Miamis are remarkably stable. Changes of sufficient
magnitude to destroy the upper shore vegetation rarely occur.

C. THE FLOOD PLAIN
1. Pond-swamp associations.

Swampy conditions or ponded waters are found in a number of places
on the flood plain. Here are included natural and artificial depressions,
and sloughs.

Low places which are filled with water for at least a part of the year,
are very commonly found at the foot of a pronounced rise in the flood
plain. They owe their existence to the levee nature of the alluvial de¬
posits. Other depressions are found whose origin is not as easily ex¬
plained. Artificial conditions also have contributed more or less in the
formation of a number of flood plain ponds. And a number also have
been destroyed by draining or filling. Most of those that remain are
small, and do not contain the wealth of aquatic vegetation which local
check lists credit to larger ponds of the region which were drained
years ago.

I'k;. -10. licit transects along- the lines AA', BB', and CC', in map, Fig. 47; symbols as in Fig. 47.

Ecology of the Cincinnati Region

195

True aquatics are scarce or entirely absent, but the amphibious plants
are numerous. These are seldom arranged in distinct zones, as these
flood plain ponds are often very irregular in outline. Cat-tail (Typha
latifolia), arrow-head (Sagittaria latifolia), and bulrush (Scirpus validus)
usually occupy the margin of the pond, or are scattered through it in
patches. Each wetter spot in the general swampy area is occupied by

Fig. 50. Ponds on the Ohio-Little Miami flood plain.

these plants, and the intervening spaces are filled with other amphibious
plants, among which are Alisma Plantago-aquatica, Cicuta maculata, Pen-
thorum sedoides, Ludvigia alternifolia, L. palustris, Rumex altissimus,
and Asclepias incarnata. Here and there are patches of the buttonbush
(Cephalanthus occidentalis) , and advancing upon the herbaceous plants
of the pond and its swampy margins are willows and cottonwoods, or
sedges and grasses. Fig. 50 is a view of a flood plain swamp, in which
are a number of small stretches of open water.

196

Ohio Biological Survey

The sloughs of the Little Miami and Miami rivers are similar in
many respects to ox-bow cut-offs. They differ from them, however, in
mode of origin, and in their almost universal connection with the river
at both ends, at least during high water. Between the swampy depres¬
sion which may cross the landward side of a wide shore (fig. 4/), and
the slough of almost river dimensions, there are all gradations..

The water of the slough flows very sluggishly, if at all, during dry
seasons, but in times of flood is swift. In very quiet places, the water is
covered with duckweed (Lemna minor) and floating species of Riccia are

Fig. 51. The shallow parts of sloughs are occupied by swamp plants; duckweed on
the surface of the water.

occasionally found. The shallower parts are occupied by Sagittaria lati-
folia, Eleocharis, Typha latifolia, Leersia oryzoides, and Echinochloa Crus-
gain (fig. 51).

Whether the slough is full of water, or merely swampy, and occu¬
pied by swamp herbs, the margins are bordered by the typical marginal
vegetation of rivers. In this respect, the sloughs resemble rivers with
their well drained margins, much more than they resemble ox-bow lakes,

Ecology of the Cincinnati Region

197

around which are always found plants characteristic of undrained swamps
(Cowles, 1901). Fig. 52 shows one of the sloughs of the Miami, in which
the channel is full of quiet water. It is bordered by willows ( Salix nigra,
and .S', cordata). .

Fig. 52. A slough of the Miami bordered by willows.

This marginal zone of bushy willows is a continuation of the first
tree zone of the rivers. It completely surrounds the “island’’ which is
enclosed between the slough and the river. Commencing with this zone
of willows, the slough succession and the river bank succession are
identical.

198

Ohio Biological Survey

2. Meadow associations ; clearings.

In its natural condition, almost the whole of the large flood plain was
forested. Now most of it is cleared and cultivated, and only a few nat¬
ural woodlands and uncultivated clearings remain.

The pond-swamp associations are usually followed by forest asso¬
ciations. In places, however, an herbaceous association composed of
sedges, grasses, and field plants (H elianthus strumosus, V ernonia fascicu-
lata. Ambrosia trifida, and Solidago canadensis) follows the amphibious
herbs. This condition is only temporary, and if undisturbed, trees begin
to appear in the meadow — willows, poplar, honey locust, and elm.

Clearings on the flood plain, which are uncultivated for a number
of years, are usually occupied by a rank growth of tall weeds, only a few
of which are in any way characteristic of flood plains. Most abundant
among these, is the giant ragweed (Ambrosia trifida) which, though com¬
mon in moist soil in clearings everywhere in the region, reaches its highest
development in these situations. Where this is not abundant, the vege¬
tation is sometimes quite varied, consisting of such plants as Helianthus
strumosus, Silphium perfoliatuni, Echinochloa Crusgalli, Lycopus ameri-
caniis, Sicyos angulatus, Lippia lanceolata. Polygonum pennsylvani-
cum, etc.

In some places the fields are grassy instead of being filled with tall
weeds. The aspect of these clearings changes with the season, for they
always contain a number of plants other than grasses and sedges. Miami
mist (Phacelia Purshii), lamb’s lettuce (V alcrianella radiata), field sorrel
(Rum ex acetosella) , cinquefoil (Potentilla monspeliensis) and mist-flower
(Eupatorium coelestinum) are perhaps the most noticeable.

3. Forest associations.

The forests of the flood plain are of two types, belonging to different
successions. One of these, the depression forest, is uncommon, occur¬
ring in undrained situations on the flood plain. The other, which is best
known as the flood plain forest, is almost universal. It occupies the wet
but well drained margins of streams, and the lower flood plains.

Depression forest. — The depression forest of the flood plain is very
similar to the depression forest of the upland. The trees are those most
typical of undrained situations, namely, pin oak, swamp white oak, and
red maple. Only three such areas are known, two on the flood plain of
the Ohio, and one in Mill creek bottoms. They lie on broad plains, and
as on the uplands, the depressions are so slight, that they are not easily

Ecology of the Cincinnati Region

199

noticed. All three are on high flood plains, only reached by exceptional
floods, as that of March, 1913.

It is probable that this type of forest arises late in the pond-swamp
succession. At least the ponds of the flood plain occupy the only other
similar positions. None of these show any tendency toward the develop¬
ment of a swamp forest, but this may be due to the artificial conditions
which surround them.

Little can be learned as to the probable trend of the succession from
the depression forests themselves, for they are completely cleared of their
undergrowth. A comparison of the three depressions shows that their
forests are not in the same stage of development. The largest one (on the
flood plain of the Ohio river at Melbourne, Ky.) is undissected by small
ravines. Its trees are of different ages and range from several inches to
a few feet in diameter, some of the pin oaks being exceptionally large.
On the margin of the depression, and not far from the willows and syca¬
mores of the flood plain forest is a single beech tree. This gives the only
clue as to the possible future of the depression forest of flood plains, if un¬
dissected — a mesophytic forest, probably similar to that which follows the
depression forest of the upland.

The other two depressions are broken by ravines. One, on the Ohio
river flood plain at Finney, Ohio, is trenched by a deep gully, along
which are a few ravine trees. The third depression, north of Lockland,
is traversed by several shallow valleys along which are sycamores. As
on the uplands, the conditions which produced the depression forest are
being destroyed by dissection.

The depression forests of uplands and flood plains are alike in that
they contain the same kinds of trees, and their successions are proceed¬
ing in the same general course, either toward mesophytism by gradual
filling, or toward xerophytism, by dissection. The depressions of the
upland depend upon rainfall, ground-water and impervious soil for their
supply of water. Those of the flood plain depend upon rainfall and
ground-water during only a portion of the year, for at times they are
filled by the river. The alluvial soil is light and porous, very different
from the till and white clay of the uplands. Topographic situation is
a big factor in the production of swamp forests in the flood plain de¬
pressions.

Flood plain forest. — The flood plain forest is closely related, in its
development, to the constructional phases of river activity. It exhibits
a number of zones whose extent and degree of development are depen-

200

Ohio Biological Survey

dent on the slope and breadth of the flood plain (sec fig. 55J. The lower
shore association (zone I) and zone I of the flood plain forest are
identical. But in areas of steadily advancing flood plain forest, the
upper shore is usually occupied by an association composed largely of
young trees. A general view of a forested area on the flood plain of
the Little ]\Iiami river near Gwendolyn is shown in fig. jj.

The succession commences with the scattered willows or poplars on
the margin of depositing shores. The number of tree seedlings in the

Fig. 53. General view of an area of flood plain forest on the Little Miami river
at Gwendolyn.

herbaceous zones of the shore is usually few, indicating a very gradual
advance of trees. In rare instances, however, seedlings almost complete¬
ly take possession of the beach producing a distinct zone. When this
is the case, the seedlings are all of the same age and height, and if more
than one species are represented in the zone, they often produce distinct
bands. Fig. 5^, which is a near view of the flood plain forest seen on
the right hand side of /ig. 55, shows in the foreground a banded zone of

Ecology of the Cincinnati Region

201

first year seedlings of Salix nigra and Popuhis deltoides. Here, although
zone II (dg. 55^ contains many tree seedlings, the plants are all of the
current season’s growth (1913). If these endure, and the banks are built
out in front of them, a new and younger zone is initiated.® The causes

Fig. 54. The same flood plain forest viewed from the shore.

which permit the sudden appearance of this new zone have not yet been
determined. It is probable, as willow and cotton-wood seeds will germi¬
nate only on moist ground, that a great deal depends on the stage of the
river at the time of seed ripening.

The second association of the flood plain formation consists most
commonly of small willows and cotton-woods, but in many places other
trees may enter in, among which are sycamore, white elm, box elder
(Acer Negnndo), and silver maple (Acer saccharinuni) . In the succes¬
sion shown in dg. 5^, this zone is made up of willows and poplars showing

®yhe belt transect, fip. 55, shows the conditions late in June, 1914, one year later. The new
zone in this is called zone O, and the other zones numbered in accordance with the photographs

202

Ohio Biological Survey

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Ecology of the Cincinnati Region

203

the same banding that was noted in the first zone. The trees here are
from eight to ten feet tall. Herbaceous plants are not abundant, but
are represented by a mixture of beech herbs with those of later asso¬
ciations.

Competition becomes keener with the increase in size of individuals.
A large number of the trees which started die out; and those remaining
are standing farther apart. Banding, even if it once existed, is less ap¬
parent in later stages of the succession.

Fig. 56. Within the third zone of this flood plain formation.

In the next association, the third flood plain zone, fig. ^6, willow
and poplar do not everywhere hold the ascendency in numbers that was
once theirs. Silver maple is now proportionately more abundant than
in the two previous zones. This is an association of small flood plain
trees. The compact growth of the second zone, which was due to the
bushy character of its trees is now gone. Here there is very little her¬
baceous or low woody growth. A few plants of wood nettle (Laportea.
canadensis) and a little poison ivy are all that is seen. Of all the asso-

204

Ohio Biological Survey

ciations of the flood plain formation, this has the most open undergrowth.
Each year floods add their deposit of mud and debris, seemingly in great¬
est amount at this horizon.

On very broad flood plains this association is usually wide. Else¬
where, it is narrow. Traced up or down stream, this is found to be the

Fig. 57. Within the fourth zone of this flood plain formation.

marginal zone. It is the most persistent flood plain association, border¬
ing all rivers, except along under-cut banks, and extending up smaller
streams to their upper limit of flood plain formation (ci. fig. 26). It
occupies the area reached by all high water. This zone is flooded several
times during the late winter and spring, but rarely at other seasons.

The flood plain does not rise evenly from the river to its highest
parts, but consists of a number of levels separated by short abrupt slopes.
The rise is usually gradual through the first three zones. Then there is
an abrupt rise of five or ten feet to a higher flood plain. This, though
not often flooded during the growing season, is always covered by all
ordinarily high water.

Ecology of the Cincinnati Region

205

On this level is the fourth zone of the flood plain formation (Hg.

It contains the same hydrophytic trees as the previous associations.
Those commonly occurring here are the willows ( Salix nigra and S', alba
vitellina), cotton-wood (Populus deltoides) , white elm (Ulmiis ameri-
cana), silver maple (Acer saccharinnm) , box elder (Acer Negundo) ,
and sycamore (Platanus occidentalis) . These vary in abundance in dif¬
ferent parts of the association. The fourth zone is in striking contrast
to the previous association with its scant undergrowth. The ground is
covered with herbaceous plants, growing in dense patches of single
species. The most abundant of these are the wood nettle (Laportea
canadensis) , touch-me-not (Inipatiens pallida), skullcap (Scutellaria la-
terifolia) and in sunny spots, the giant ragweed (Ambrosia tridda).
Saplings are not uncommon in this association, but because of the dense
shade, willow, poplar, and sycamore are almost excluded. White elm
and box elder are frequent and sugar maple (Aeer saccharum) occa¬
sionally occurs. This last is the first indication of an advance toward
mesophytic conditions.

This association, in various aspects, is the most wide-spread of any
of the flood plain formation. It often occupies a number of successive
levels, very slowly changing toward a more mesophytic forest, as the
flood plain is built up. On some of the broadest flood plains, it is capable
of further subdivision on a basis of age, and of the character of its
herbaceous growth.

The next zone of the flood plain formation is not of the same char¬
acter along all the streams. Much depends on the history of the valley
and on its topography.

In the area shown in dg. 5y, the fourth zone is the last that is typical
of flood plains (cf., dg. 55j. The next zone occupies the lower slopes
of the hills. It is a zone only reached by exceptional floods, as that of
March, 1913. Its trees are a medley of flood plain types, slope xero-
phytes, and mesophytes. Here are found hackberry ( Celtis occidentalis) ,
walnut (Juglans nigra), honey locust (Gleditsia triacanthos) , red elm
(Ulmns fulva), red-fruited thorn (Crataegus mollis), and buckeye
(Aesculus octandra), with the last remaining trees of the flood plain,
white elm and box elder. Among the saplings, sugar maple (Acer sac¬
charum) and wild cherry (Primus serotina) are abundant, with a few
of the other trees. Herbaceous plants are very scarce in this zone, pos¬
sibly because of the erosive work of the floods. Succeeding associations
do not belong to the flood plain formation. This mixed zone is the last

206

Ohio Biological Survey

which shows any indication of connection with the depositional phases
of river activity.

With slight modifications, the succession just outlined is charac¬
teristic of most of the flood plains of the region. Along the Miami,
however, is found a different and later stage in the flood plain succes¬
sion. Much of the flood plain area of that river lies at a level which is
very rarely flooded; some of it above the level of the highest floods of
historic times. The forest found here is not that typical of the lower

Fig. 58. Destruction of a mature flood plain forest by a stream under-cutting its
banks.

flood plains. On this high-level flood plain, is a mixed mesophytic forest
in which are found white elm (Ulmus americana), bitter nut (Cary a
cordiformis) , sugar maple (Acer saccharum) , wild black cherry (Prunus
serotina), walnut (Juglans nigra), black ash (Fraxinus nigra), red ash
(Fraxinus pennsylvanica) , and oaks (Quercus macrocarpa, Q. imbricaria,
Q. alba, and Q. rubra). The white elms are large and as before stated,
are among the last of the typical flood plain trees to persist. Of the
other trees, Quercus macrocarpa (bur oakj and Carya cordiformis are

Ecology of the Cincinnati Region

207

particularly characteristic of high-level flood plains, both of rivers and
small streams. Although this mixed mesophytic forest of the high-level
flood plain differs from the erosion climax forest in many respects, it re¬
sembles it more closely than it resembles any other forest in the region.
It is probably a fore-runner of that forest.

The most complete successions are found on the inside of broad
curves of the rivers. In such places, the rivers are essentially depositing
streams, and their flood plains are constantly being built upward and
outward. It is this constant addition to the flood plain, which permits
of the gradual advance of flood plain vegetation.

Even the mature flood plain forest is not always permanent. In
the course of its wanderings on the flood plain, the river may destroy
the land it has built up, and the trees that have grown upon it. On the
Miami and Little Miami rivers this process is not slow. On under-cutting
parts of the flood plain, many trees are destroyed each year. Across the
river from the very complete flood plain formation shown in Hgs. S3~S7’
destruction of a mature stage of the fourth flood plain association is in
progress. The trees lying in the water at the foot of the bank (iig. §8)
illustrate the destructive work of this river. Erosion will very likely
cease in this location before the whole of the flood plain has been de¬
stroyed. In this case, a new flood plain succession will commence on
the new shore. Very often, parts of different successions are found
upon the same flood plain. It is by this partial destruction of flood plains,
that this condition is brought about.

Beginning with the shore, the succession is a gradual and progres¬
sive one, culminating in the mixed forests of the high-level flood plains.

VI. GENERAL CONCLUSIONS AND SUMMARY

It has been shown that the plant formations of the Cincinnati region
are closely related to its topography. The four principal topographic
areas — uplands, slopes, flood ‘plains, and terraces — are also, to a large
extent, soil areas. Each exhibits its own plant associations, dependent
as has been shown, primarily upon topography and its changes, and
only secondarily, upon the soil.

Soil influences may be overcome by topographic conditions. Dis¬
similar topographic forms, in the same soil are occupied by different
plant associations. Similar topographic forms, in different soils, support

208

Ohio Biological Survey

plant associations which are related to one another. Although the vegeta-
tional features of young blufifs in gravel, rock, and clay differ from one
another in many respects, they are more nearly related to each other, than
are the associations of a gravel bluff and a gravel shore. Numerous
examples point to the conclusion that topographic influences are usually
more powerful than soil influences.

In most of the region, there is some relation between vegetational
and geological areas due to the influence of soil. Fresh exposures of
gravel, rock, and clay are more different from one another, than are weath¬
ered ones. For this reason, the differences between the vegetation of dif¬
ferent soil areas are more pronounced early than late in the succession.

Soil has an indirect influence on vegetation through its effects on
topographic forms. Resistance or ease of erosion of rock material re¬
tards or accelerates the progress of the erosion cycle, and with that, the
vegetative cycle dependent upon it. The effects of difference in resist¬
ance are best disclosed where two materials, in contact, are acted upon
by the same agent. These effects may be seen along a stream which
cuts through two or more unlike materials. The differences in the asso¬
ciations, along such a stream are partly due to differences in topography
arising as a result of differences in the ease of erosion of the soil. The
associations, as existing along that stream, do not illustrate exactly the
succession of associations at any one place, for this is modified by the
soil of the area in which the succession occurs.

In some instances, topography and the physical character of the soil,
together produce the plant association. An undissected flat is usually
a poorly drained area. If, added to the absence of drainage lines, the
soil is of such a nature that it drains slowly the features of bad drainage
will be more pronounced, and the plant associations which depend upon
poor drainage for their existence, will be better developed.

Roth xerarch and hydrarch successions are illustrated in the Cin¬
cinnati region.

The xerarch succession embraces those of dry flats, whether on up¬
land or terrace, and of slopes, whether of rock, gravel or clay. The suc¬
cessions on dry flats are relatively rapid. They are dependent for their
advance upon biotic agencies. Topography is as yet little changing.
Erosional features are almost absent. Biotic cycles are not complicated
by the progress of a topographic cycle.

The successions on slopes are slow and varied. They are produced
by a combination of biotic and topographic agencies. Of these succes-

Ecology of the Cincinnati Region

209

sions, that on rock bluffs is the slowest and includes the greatest number
of stages. Soil or rock is here influencing the succession, because of
its resistance to erosion, and its effect on soil water.

Hydrarch successions are of two kinds, those of undrained and those
of drained or half-drained situations. The former are exhibited in three
widely different topographic situations, namely, on uplands, in fijled
valleys, and on flood plains ; the latter, along the margins of streams,
where the typical flood plain succession is in progress, and occasionally,
in the pre-glacial filled valleys. During the hydrophytic stages, these
two kinds of successions are characterized by different types of vege¬
tation.

The hydrarch succession of the upland depends upon topography
and soil. An undissected flat, combined with an impervious soil pro¬
duces the moist conditions necessary to the growth of hydrophytes.
That soil is an important factor in producing these conditions, may be
learned by comparing flats having different soils. Hydrophytic associa¬
tions are best developed on uplands covered with a relatively thick
deposit of Illinoian drift. Where this deposit is very thin or absent, and
only the loess (white clay) reaches its usual thickness, as in the southern
part of the region, decidedly hydrophytic forests are not found. Only a
few hydrophytic trees occur in an otherwise mesophytic forest. South
of the limits of glaciation and the extent of loess, wet upland flats appear
to be uncommon. Owen (1857) mentioned the “soggy beech flats” in
Fayette County, Kentucky. There, as in this region, they seem to be inti¬
mately related to soil conditions, i. e., the sub-stratum is composed of
fine-textured mudstones and shales, which produce a poorly drained soil.
On comparing an area covered with till of Illinoian age with one cov¬
ered with the Earlier Wisconsin, a similar, but less pronounced difference
is seen. Thus the imperviousness of surface material on the uplands
modifies the character of the vegetation through its effect on soil water.

The hydrophytic associations of undrained situations in the filled
valleys are similar in character and causal relations to those of the
uplands.

The depression hydrophytic forests of the flood plain occur in habi¬
tats formed by streams, but now removed from the influence of their
activity. Soil is not the important factor here. Low situation in combi¬
nation with undrained conditions determines their existence.

210

Ohio Biological Survey

The typical flood plain succession is a direct result of stream action.
Changing topography is the chief cause of the progress of the suc¬
cession.

All successions in the region, whether xerarch or hydrarch, in
whatever topographic situation they may be, are progressing toward a
mesophytic forest. The relatively short successions on a topography
as yet unchanging, and the long and varied successions on ever chang¬
ing topographic forms, all lead to a mesophytic forest — a climax forest.
The climax mesophytic forest exists in two distinct forms. One is the
beech forest of the uplands and filled valleys, the forest developed on an
unchanged or very youthful topography. The climax of the vegetative
cycle precedes, or accompanies the initiation of the erosion cycle. This
beech forest is the pre-erosion climax.

The second form of the climax forest is the mixed mesophytic forest
of gentle slopes. All long successions dependent on topographic changes
lead to this forest. The succession in ravines, the successions in progress
on flood plains, both end in a mixed mesophytic forest. Both depend on
topographic change, the first on stream degradation, the second on stream
aggradation. The climax of these vegetative cycles follows a period of
erosive activity. It comes within an erosion cycle. This mixed meso¬
phytic forest is the erosion climax.

The pre-erosion climax forest of the Cincinnati region resembles
the beech forests of northern Ohio and of Michigan. Its closest affinities
are northward. The erosion climax forest resembles the deciduous forest
of the Southern Appalachians. It is poorer in variety of trees, but it has
much in common with the southern forest. Northern and southern for¬
est types here occur side by side.

The pre-erosion climax forest is temporary. It can endure only as
long as the pre-erosion topography. The erosion climax forest, a mixed
mesophytic forest, which results after, though not at the end of topo¬
graphic changes, represents more truly the climax forest of the region.

Ecology of the Cincinnati Region

211

LITERATURE CITED

Bonsep, Thomas A.

1903. Ecological study of Big Spring Prairie. Proc. Ohio State Acad. Sci.,
Spec, paper No. 7.

Cincinnati Folio

- . In preparation.

Cooper, William S.

1913. The climax forest of Isle Royale, Lake Superior, and its develop¬
ment. Bot. Gaz., 55:1-44, 115-140, 189-235.

Cowles, H. C.

1899. The ecological relations of the vegetation on the sand dunes of
Lake Michigan. Bot. Gaz., 27:95-117, 167-202, 281-308, 361-391.

1901. Physiographic ecology of Chicago and vicinity. Bot. Gaz., 31:73-108,
145-182.

1901, a. The influence of underlying rocks on the character of the vegeta¬

tion. Bull. Amer. Bur. Geogr., 2:163-176.

1911. The causes of vegetative cycles. Bot. Gaz., 51:161-183.

Dachnowski, Alfred.

1910. A cedar bog in central Ohio. Ohio Nat., 11:193-199.

1912. The succession of vegetation in Ohio lakes and peat deposits. Plant
World, 15:87-103.

1912, a. The peat deposits of Ohio. Geol. Surv. Ohio, Bull. 16.

Detmers, Frederica.

1912. An ecological study of Buckeye Lake. Proc. Ohio State Acad. Sci.,
Spec, paper No. 19.

Fuller, M. L. and Clapp, F. G.

1912. The underground waters of southwestern Ohio. U. S. Geol. Surv.,
Water-supply paper 259.

Leverett, Frank.

1902. Glacial formations and drainage features o the Erie and Ohio ba¬

sins U. S. Geol. Surv., Mon. 41.

Owen, David Dale.

1857. Third report of the Geol. Surv. Kentucky (Fayette Co.).

Schaffner, John H.,, Jennings, O. E., and Tyler, F. J.

1904. Ecological study of Brush Lake. Proc. Ohio State Acad. Sci., Spec,
paper No. 10.

Shaw, C. H.

1902. The development of vegetation in the morainal depressions of the
vicinity of Woods Hole. Bot. Gaz., 33:437-450.

Shimek, B.

1912. Prairie openings. Paper delivered at the Cleveland meeting of the
A. A. A. S.

Whitford, H. N.

1901. The genetic development of the forests of northern Michigan: a
study in physiographic ecology. Bot. Gaz., 31:289-325.

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region, and localities mentioned in the text. (Boundary of Wisconsin drift, as far east as Kings Mills, after I’cnneman; remainder, and bound

nig general topographic features of the Cincinnati
inoian drift, after Leverett).

Volume II, No. 4

Bulletin 8

Ohio Biological Survey

THE TINGITOIDEA OF OHIO

HERBERT OSBORN

and

CARL J. DRAKE

Published by

THE OHIO STATE UNIVERSITY

Columbus.

1916

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CONTENTS

Page

Superfamily Tingitoidea . 217

Family Piesmidae . 218

Genus Piesma . 218

Family Tingitidae . 219

Genus Acalypta . 220

Genus Fenestrella . 222

Genus Corythucha . 224

Genus Gargaphia . 233

Genus Galeatus . 236

Genus Leptostyla . 237

Genus Leptobyrsa . 240

Genus Leptophya . 241

Genus Physatochila . 242

Genus Melanorhopala . 243

Genus Alveotingis . 245

PRINTED BT

THE STONEMAN PRESS

Columbus, Ohio

THE TINGITOIDEA OR “LACE-BUGS” OF OHIO.
Herbert Osborn and Carl J. Drake.

The insects belonging to the superfamily Tingitoidea are commonly
known as “lace-bugs.” They are readily diflerentiated from all other
Hemipterous-Heteropterous insects by the net-, gauze-, or lace-like ap¬
pearance of their dorsal surface. This peculiar appearance is due to the
more or less closely reticulate relief of the membranous pronotal mar¬
gins and the elytra or hemielytra. The lace-bugs are all rather small
insects and plant feeders. They are usually found on the underside of
the leaves of trees and shrubs, also a few herbaceous plants. When
present in sufficient numbers they may do a considerable amount of
damage to cultivated shrubs and plants.

We have included several species not yet collected within the borders
of the state, but which probably belong to our fauna.

In the descriptions of the genera and species we have followed Dr.
Champion (Biologia Centrali- Americana, Rhynch., Vol. II, p. 1), and
the following terms are used for the different portions of the elytra :
(1) costal area (the outer membrane of Stal), (2) subcostal area (the
costal area of Stal), (3) discoidal area, (4) sutural area.

Acknozvledgment. — The authors desire to express their sincere thanks
to those who have very kindly assisted in the preparation of this paper
as follows : Dr. L. O. Howard, Bureau of Entomology, Washington,
D. C., for the loan of specimens from the National Museum and for
plates from the United States Department of Agriculture ; Mr. Otto
Heidemann, Washington, D. C., for the use of specimens from his
private collection ; Dr. E. H. Ewing, Ames, Iowa, for the use of material
from the collection of Iowa State College ; Mr. H. M. Parshley, Harvard
University, for specimens from his private collection ; The Entomo¬
logical Society of Washington for the loan of plates made from draw¬
ings by Mr. Heidemann ; Dr. A. W. Morrill, State Entomologist,
Phoenix, Arizona, for the privilege of using his plate showing life history;
Mr. James Hambleton who has made the original drawings; Mr. Erank
H. Smith for the photographs.

KEY TO THE FAMILIES OF THE TINGITOIDEA.

Pronotum produced posteriorly, forming a triangular process; ocelli wanting;
elytra of a uniform texture throughout, densely areolated, membranous be¬
tween the areolae . Family Tingitidae

217

218

Ohio Biological Survey

Pronotum not produced posteriorly; ocelli present; hemielytra with a distinct

membrane, the rest reticulately punctate . Family Piesmidae

Family Piesmidae.

Head short and broad; juga free, longer than the tylus, produced
before the apex of the head. Rostrum consisting of four segments.
Eyes small, prominent; ocelli present, inserted close to the margins of
the prothorax. Legs short ; tarsi consisting of two segments. Hemi¬
elytra with a distinct non-reticulate membrane.

This family contains only one Nearctic genus.

Genus Piesma Lepelletier et Serville.

Piesma Lepelletier et Serville, Encycl, Method. X, p. 653, 1825.

Zosmenus Laporte, Essai class, syst. des Hemipt, p. 49, 1832.

Aspidotoma Curtis, Entomol. Magaz., I, p. 196, 1833.

Zosmerus Douglas et Scott, Brit. Hem.-Het., p. 237, 1865.

Piesma Stal, Enum. Hemipt., Ill, p. 115, 1873.

Head short, broad, deflected. Antennae short ; first segment strongly
swollen, with a short petiole ; second segment shorter than the first, less
swollen; third segment longer than the first and second conjoined;
fourth segment stout, longer than the first, fusiform. Ocelli small, in¬
serted close to the posterior margin of the head. Legs short ; tarsi
composed of two segments ; claws short, curved. Scutellum small,
triangular. Hemielytra as long as the abdomen, with a distinct non-
reticulate membrane at the apex, the rest reticulately punctate.

The genus contains only one described American species.

Piesma cinerea Say (Plate X, fig. a).

Tingis cinerea Say, Descr. Heteropt. Hemipt., p. 26, 1832; Compl. Writ., Vol. I,
p. 349, 1859.

Piesma cinerea Stal,, Enum. Hemipt., Band. HI, p. 116, 1873.

Piesma cinerea Comstock, Intro. Entom., p. 204, 1888.

Piesma cinerea Summers, Bull. Agr. Exp. Stat. Tenn., Vol. IV, No. 3, p. 90,
1891.

This species is commonly known as the “ash-gray’’ leaf bug. Ac¬
cording to Prof. Summers it often does a considerable amount of damage
to the leaves and flowers of grapes.

Head with a short, stout spine between the antennae and eyes. Antennae
short; first segment strongly swollen, abruptly smaller at the base, longer than the
second; second segment short, less swollen; fourth segment swollen towards the
apex, beset with a few short hairs. Rostrum reaching slightly beyond the first
pair of coxae. Pronotum with four elevated lines, obsolete behind. Length, 2.9
mm; width, about 1.2 mm.

The Tingitoidea or ‘‘Lace-Bugs'' of Ohio

219

Color: Beneath grayish, more or less varied with brown; venter brownish,
with spots or lines of grayish. Prothorax and elytra grayish, more or less spotted
with dark-brown or black. The color varies in different specimens. In some
forms the spots are almost entirely wanting.

Common thruout the state. It may be taken in the winter hiber¬
nating beneath the bark of sycamore and horse chestnut. Columbus,
Berea, Cedar Point, Jefferson, Buckeye Lake, Chillicothe, and Perry Co.

Family Tingitidae.

Head sometimes with dorsal porrect spines ; juga not prominent, not
longer than tylus. Antennae consisting of four segments ; third segment
longest. Rostral sulcus rather deep, sides raised ; rostrum consisting
of four segments. Ocelli wanting. Tarsi consisting of two segments.
Pronotum, in front, with a hood or only tumid; lateral margins folia-
ceous, reticulately projecting or closely lapped over on the pronotum ;
disc with one or three carinae; posterior triangular process usually long.
Elytra of a uniform thickness, more or less closely reticulated thruout ;
membranous between the reticulations.

This family contains several genera that present wide differences.

KEY TO THE GENERA OF OHIO TINGITIDAE.

1. Membranous pronotal margins not expanded; or reflected vertically; or

reflected more or less against the dorsal surface of the pronotum . 8

Membranous pronotal margins more or less reflected, but at an obtuse

angle . 2

8. Pronotum with a pronotal and discal hood, the latter divided behind ;

areolae very large . Genus Galeatus Curtis

Pronotum with only a pronotal hood; areolae very much smaller . 3

3. Rostral sulcus interrupted between the meso- and metasternum by a

prominent, simiose, transverse carina . Genus Gargaphia Stal

Rostral sulcus uninterrupted by a transverse carina . 4

4. Outer edge of membranous pronotal margins and elytra, except distal

third, armed with short, acute spines ; elytra with a tumid elevation

near the antero-inner margin . Genus Corythucha Stal

Outer edge of membranous pronotal margins and elytra unarmed (in the

genus Leptobyrsa closely beset with fine hairs) . 5

5. Pronotum with a single median carina . Genus Fenestrella n. gen.

Pronotum tricarinate . 6

G. Discoidal area reaching beyond the middle of the elytra .

Genus Acalypta Westwood
Discoidal area not reaching the middle of the elytra . 7

7. Elytra very broad, extending about one-half beyond the apex of the abdo¬
men, the outer margin broadly rounded and beset with numerous, fine
hairs . Genus Leptobyrsa Stal

220

Ohio Biological Survey

Slender insects; elytra narrow, shorter, the outer margin more or less

constricted about the middle, hairs on outer margin wanting .

Genus Leptostyla Stal

8. Membranous pronotal margins not expanded . Genus Leptophya Stal

IMenibranous pronotal margins reflected vertically; or reflected more or less

against the dorsal surface of the pronotum . 9

9. Reflected pronotal margins very broad, broadly rounded, touching or

nearly touching the outer carinae . Genus Physatochila Fieber

Reflected pronotal margins very narrow, subquadrate . 10

10. Subcostal, discoidal, and sutural areas undifferentiated; body very thick;

antenniferous tubercles very large . Genus Alveotingis n. gen.

Subcostal, discoidal, and sutural areas distinct; antenniferous tubercles

much smaller . 11

11. Antennae contiguous at the base; elytra broadly rounded at the apex .

Genus Teleonemia Costa.

Antennae not contiguous at the base; elytra greatly narrowed or obliquely
rounded at the apex . Genus M elanorhopala Stal.

Genus Acalypta Westwood.

Acalypta Westwood, Mod. Class. Ins., Vol. II, Syn. Gen., p. 121, 1840.
Acalypta Stal, Enum. Hemipt., Band. II, p. 118, 1873.

Antennae moderately long, rather stout ; first segment swollen,
longer than the second ; second segment less swollen, short ; third seg¬
ment very long, slender ; fourth segment fusiform. Pronotum tricari-
nate ; rather closely reticulated ; hood small, covering the base of the
head ; lateral margins expanded, subquadrate. Costal area narrow,
coarsely reticulated, with one complete or with one complete and a par¬
tial series of areolae ; subcostal area closely reticulated, much narrowed
behind ; discoidal area rather wide, extending beyond the middle of the

elytra; sutural area (hrachypterous form) with the inner margin straight,
narrow. In the macropterous form the sutural area is broad with the in¬
ner margin rounded.

KEY TO THE SPECIES OF ACALYPTA.

Costal area of the elytra with one complete and the greater part of a second

series of areolae . A. thoinsonii Stal

Costal area with a single series of areolae, or uniseriate in greater part

(Figs. 1 and 2) . A. UUianis Bueno

Acalypta thomsonii Stal.

Acalypta thomsonii Stal, Enum. Hemipt., Band. HI, p. 122, 1873.

Acalypta thomsonii Bueno, Bull. Brook. Ent. Soc., Vol. XI, p. 39.

Head armed with two rather stout, blunt porrect spines — one on each side
of the median line just above the antennae. Rostrum reaching almost to the end of
the first abdominal segment. Antennae rather stout, beset with a few short hairs ;
first segment considerably swollen, about twice the length of the second ; second

The Tingitoidea or ‘‘Lace-Bugs"' of Ohio

221

segment short, less swollen; third segment nearly twice the length of the fourth,
slenderest; fourth segment fusiform. Pronotal hood narrow, extending between
the eyes. Pronotum with the discal carina diverging posteriorly. Elytra with the
outer margin rounded ; costal area coarsely reticulated, with one complete and
nearly a second series of areolae; discoidal area extending beyond the middle of
the elytra. Length, 2.5 to 3 mm. ; width, 1.5 to 1.75 mm.

Color: General color griseus. Body beneath and legs ferrugineous. Antennae
ferrugineous ; the basal and apical segments blackish. Thorax and elytra griseus.
Habitat: South Carolina ( Stal ) and Virginia (Barber).

Acalypta lillianis Bueno.

Acalypta lillianis Bueno, Bull. Brook. Ent. Soc., Vol. XI, p. 39, 1916.

Eig. 1. Acalypta lillianis Bueno, brachypterous form (length, 2.3 mm.).

“Black, posterior part of pronotum and hemelytra dark gray; third joint of
antennae of equal thickness throughout, four times as long as the fourth, longest,
second joint shortest, first stoutest and shorter than fourth, which is not quite so
stout as second ; pronotum tricarinate, keels nearly parallel, or at most slightly
diverging posteriorly, the middle keel highest; hood small, not projecting over
head, slightly angulated in front, somewhat higher than the median keel; sides

222

Ohio Biological Survey

foliaceous, more or less biseriately reticulate, anteriorly rounded; costal membrane
uniseriate in greater part. Abdominal segments broadly banded with rufous.”

Fig. 2. Acalypta lillianis Bueno, macropterous form (length, 3 mm.).

“The macropterous form has the membrane complete, reticulately veined ; in
the brachypterous, it is much reduced and is biseriately reticulate. The pronotum
in the former is quite high, but in the latter much reduced in area and nearly flat.
The hemelytra are one-third longer than the body in the macropterous form, and
in the brachypterous exceeds it but very little. The brachypterous form is nearly
oval, while the macropterous is elongate.”

Length (macropterous), 3 mm.; width. 1.6 mm.

Length (brachypterous), 2.3 mm.; width, 1.2 mm.

Habitat: Michigan, New York, Maine, New Hampshire, and Canada
(Ottawa).

Genus Fenestrella gen. nov.

Head with at least three spines, almost entirely covered with the hood.
Antenniferous tubercles small. Antennae rather short; first segment swollen
almost twice the length of the second; second segment short, less swollen; third
segment long cylindrical, slender ; fourth segment inflated, fusiform. Eyes
strongly granulate; a short blunt spine or process between the eyes and antennae.

The Tingitoidea or “Lace-Bugs'" of Ohio

223

Rostral groove uninterrupted. Pronotum with a single median carina; hood broad,
reaching to the vertex of the head; lateral membranous margins broad, coarsely
reticulated; posterior process very obtusely angled. Legs short and slender.
Elytra broad oval ; outer margin rounded ; costal area widely reticulated, with one
complete and a partial series of areolae at base and apex; subcostal area very
broad and long, reaching the apex of the elytra ; discoidal area broad, reaching
far beyond the middle of the elytra; sutural area very narrow at the middle,
widened at the base and apex, inner margin straight, except at the apex slightly
overlapping.

The genus contains a single species, Fenestrella ovata n. sp., the type
of the genus. It is close to Acalypta, but can be readily separated by the
single median carina, the trispinous head, and the larger hood.

Fenestrella ovata spec. nov. (Fig. 2).

Head with three short, blunt spines — a short median spine just above the
antennae and the two lateral converging spines inserted just above the median

Fig. 3. Fenestrella ovata n. sp.

spine. Antennae considerably less than one-half the length of the body; beset
with a few short hairs; first segment nearly twice the length of the second; third

224

Ohio Biological Survey

segment slightly more than twice the length of the other three conjoined; fourth
segment hairy, very slightly longer than the first and second segments. Rostrum
reaching the end of the rostral groove. Eyes very coarsely granulate. Pronotum
reticulated, except near the base of the hood; median carina raised, with two or
three areolae at the middle. Membranous pronotal margins very wide and broad,
subquadrate, widely reticulated ; hood rather coarsely reticulated. Legs short,
beset with a few short hairs. Costal area very coarsely reticulated, with one com¬
plete and a partial series of areolae; subcostal area with five series of areolae;
discoidal area broad, with five or six rows of areolae; sutural area with one com¬
plete and one or two partial rows of areolae at the base and apex. Length, 2 mm.;
width, 1.35 mm.

Color: Body dark gray-brownish. Legs and antennae brownish, more or less
tinged with yellow. Eyes brownish. Pronotum yellowish, tinged with brown.
Elytra brown, slightly tinged with yellow. Areolae hyaline or whitish.

Described from a female specimen, taken at Cincinnati, Ohio, by Mr.
Chas. Dury.

Genus Corythucha Stal.

Corythucha Stal, Enum. Hem., Band. Ill, p. 119, 1873.

Head entirely concealed by the hood, the apex of hood reaching a lit¬
tle in front of the head. Antennae slender, beset with long, bristly hairs ;
first segment more or less swollen, from two to three times the length
of the second ; second segment less swollen, short ; third segment very
long, cylindrical ; fourth segment moderately swollen towards the apex.
Membranous pronotal margins broad, armed on the outer margins with
short spines. Elytra narrowed at the base with outer margin strongly
reflexed, with a tumid elevation near the antero-inner margin ; costal
margin including anterior reflected margin armed with spines, except
about the distal third. The nervures of hood, membranous pronotal
margins, and the elytra irregularly armed with a few erect spines.

KEY TO THE OHIO SPECIES OF CORYTHUCHA.

1. First segment of the antennae twice or slightly more than twice the length

of the second . 2

First segment of the antennae three or almost three times the length of

of the second . 7

2. Hood not very abruptly constricted about the middle; color entirely
white, except elytra usually with a brown spot on the tumid elevation. .

C. ciliata Say

Hood abruptly constricted about the middle; hood and membranous
pronotal margins pallid or hyaline, usually with a few brown markings ;
elytra pallid or hyaline, with one or two transverse brown or fuscous
bands . 3

The Tingitoidea or “Lace-Bugs'" of Ohio

225

3. Color pallid; elytra with a more or less indistinct band near the base and

the apical band very indistinct or entirely wanting . C. pallida n. sp.

Color hyaline or brownish ; elytra with distinct brown or fuscous bands

near the base and apex, the latter band sometimes wanting . 4

4. Globose portion of the hood narrow ; body black, except the margins of

the prosternum, the humeral area, and the last abdominal segment

brownish or fulvous . 5

Globose portion of the hood quite broad; body entirely black . 6

5. Size small (about 3.2 mm.) ; hood sinuate on dorsal surface at the con¬

striction . C. pergandei Heid.

Size larger (4 mm.) ; hood not sinuate on the dorsal surface at the con¬
striction . C. pruni n. sp.

6. Hood moderately deflected; fourth segment of the antennae thick and

brown . C. juglandis Fitch

Hood more strongly deflected ; fourth segment of the antennae less

swollen and fuscous . C. celtidis n. sp.

7. Globose portion of the hood twice or more than twice as broad as triangu¬

lar portion at constriction . 8

Globose portion of hood less than twice as broad as triangular portion at

constriction; elytra marmorate . C. marmorata Uhler

8. Size large (4 mm. or longer); globose portion of the hood very large;

anterior portion directed almost vertically downward . 11

Size smaller (less than 4 mm.) ; globose portion of hood smaller; anterior

portion of the hood directed obliquely forward . 9

9. Elytra narrowed posteriorly, with two black-fuscous bands.. C. contracta n. sp.

Elytra not narrowed posteriorly, with one or two brown hands . 10

10. Hood moderately raised ; nervures of hood and membranous pronotal
margins almost entirely yellowish ; elytra with one or two brown

bands . C. arcuata Say

Hood considerably raised ; nervures of hood and membranous pronotal
margins mostly embrowned ; elytra with two transverse brown bands

C. crataegi n. sp.

11. Globose portion of the hood very large; membranous pronotal margins

and elytra without a continuous brown margin . C. aesculi n. sp.

Globose portion of the hood extremely large; membranous pronotal mar¬
gins and elytra broadly margined with dark-fuscous . C. bulbosa n. sp.

Corythucha ciliata Say (Plate VIII, fig. h).

Tingis ciliata Say, Hemip. New Harm., p. 27, 1832; Compl. Writ., Vol. I, p. 348,
1859.

Corythucha ciliata Morrill, Psyche, Vol. X, p. 133, 1903.

This is the common buttonwood or sycamore lace-bug. It is readily
separated from the other Ohio forms in the genus by its whitish color.

Antennae beset with numerous bristly hairs of various lengths; first segment
twice as long as the second ; fourth segment slightly swollen towards the apex.
Rostral sulcus very broad ; rostrum reaching the meso-metasternal suture. Mem-

226

Ohio Biological Survey

branous pronotal margins broad, biillate about the middle, slightly reflexed at the
anterior and posterior margins. Hood moderately raised, not very abruptly con¬
stricted about the middle; posterior portion subglobose. Apex of triangular pro¬
cess faintly reticulated; the sides feebly raised anteriorly, with a few distinct
areolae. Costal area of the elytra unevenly reticulated, with three complete rows
of areolae. Length, 3.75 mm.; width, 1.6 mm.

Color: General color whitish. Body above and beneath black. Antennae
yellowish ; the apical segment light brown. Legs yellowish ; tarsi brownish. Hood
and membranous margins of pronotum whitish. Elytra whitish, usually with a
brown spot on the tumid elevation, sometimes faint or obsolete.'

We have a form that seems to be a teneral specimen in which the
areolae are opaque and of a creamy-white color.

Common thruout-the state. The leaves of sycamore trees are often
much whitened and wilted from their presence. We have specimens
from Columbus, Cedar Point, Castalia, Put-in-Bay, Berea, Buckeye Lake,
Zanesville, and Malta.

Corythucha marmorata Ilhler.

Tingis marmorata Uhler, Proc. Bost. Soc. Nat. Hist., Vol. XIX, p. 415, 1878.

Antennae slender, beset with a few moderately long bristly hairs ; first segment
three times as long as the second; fourth segment swollen towards the apex.
Hood strongly raised; constricted about the middle; posterior portion globose.
Membranous pronotal margins bullate about the middle, slightly reflected at the
anterior and posterior margins. Pronotum and base of triangular portion dis¬
tinctly punctured. Costal area of elytra unevenly reticulated, with two complete
and a partial third series of areolae. Length, 3.4 mm. ; width, 1.8 mm.

Color: Body black ; the humeral region and pleural margins sometimes paler.
Antennae yellowish. Legs yellowish ; tarsi brownish. Hood and membranous
pronotal margins lacteous ; the nervures more or less embrowned. Elytra lacteous,
with the centers of areolae hyaline, with four irregular, transverse, smoky-brown
bands — the two terminal bands separated by a transverse row of hyaline areolae.

Berea, Cedar Point, Put-in-Bay, Bay Point, Castalia, Columbus,
Zanesville, ]\Ialta, Buckeye Lake, Marietta, and North Kenova.

Corythucha juglandis Fitch.

Corythucha juglandis Fitch, Third Report, Ins. N. Y., p. 466, 1866.

Antennae beset with a few long bristly hairs; first segment slightly more than
twice the length of the second. Rostral groove rounded at the apex; rostrum
reaching the middle of the metasternum. Hood considerably raised, quite evenly
reticulated, abruptly constricted about the middle ; posterior portion globose.
Membranous pronotal margins evenly reticulated, bullate about the middle. Pro¬
notum and base of triangular process distinctly punctured ; the apex of the latter
reticulated. Median carina foliaceous, with a double series of areolae at the
middle. Lateral margins of triangular process feebly raised anteriorly, with two
or three areolae. Length 3.3 to 3.7 mm. ; width 2 to 2.3 mm.

The Tingitoidea or “Lace-Bugs'" of Ohio

227

Color: Antennae yellowish; apical segment sometimes brownish. Legs
brownish; tips of tarsi brownish. Hood and membranous pronotal margins hyaline,
the nervures yellowish with parts more or less embrowned. Pronotum yellowish.
Elytra hyaline, with a transverse band on the costal area near the base, a spot
on the tumid elevation, and more or less of a band on apical third brown.

A common species found on walnut and (sometimes) basswood.
Malta (Drake), Jef¥erson (Sim), Columbus (Hine).

Corythucha celtidis nov. sp.

This species is very close to C. juglandis Fitch, but differs from the
specimens that we have assigned to that species by the less elevated
hood, the more numerous bristly hairs on the antennae, and the some¬
what different markings at the tip of the elytra.

Antennae slender, beset with numerous, moderately long bristly hairs ; first
segment twice as long as the second ; fourth segment swollen towards the apex.
Rostral groove quite wide; rostrum reaching the meso-metasternal suture. Hood
moderately raised, abruptly constricted about the middle ; posterior portion globose,
rather evenly reticulated ; anterior portion triangular, more closely reticulated.
Membranous pronotal margins broad, kidney-shaped, bullate about the middle,
slightly raised at the anterior and posterior margins. Pronotum and base of
triangular process coarsely and regularly punctured, the latter reticulated at the
apex; sides of triangular process feebly raised anteriorly, with a few distinct
areolae. Median carinae foliaceous, with a double row of areolae at the middle;
costal area unevenly reticulated, with thrjse complete rows of areolae. Length,
3.6 mm.; width, 1.8 mm. ’ *

Color: General color hyaline. Body above and beneath black. Legs yellow¬
ish; tips of tarsi embrowned. Antennae yellowish; terminal segment fuscous.
Hood, membranous pronotal margins, and apex of triangular process hyaline,
except nervures around two or three areolae brown. Pronotum and base of tri¬
angular process brownish. Elytra hyaline, a small vitta near anterior margin,
posterior portion of tumid elevation, and more or less of a narrow indistinct band
on distal third fuscous.

Described from a long series of specimens, taken on hackberry in the
various stages of development at Columbus during Sept., 1903, by the
senior author. We have two specimens taken by Mr. Kostir at Cedar
Point.

Corythucha arcuata Say (Plate IX).

Tingis arcuata Say, Hem. Xew Harm., p. 27, 1832; Compl. Writ., Vol. I, p. 350,
1859.

Corythucha arcuata Morrill, Psyche, Vol. X. p. 127-132.

Antennae beset with a few long bristly hairs ; first segment almost three times
as long as the second ; apical segment slightly swollen towards the apex. Hood
moderately raised, rather evenly reticulated, abruptly constricted about the middle ;

228

Ohio Biological Survey

posterior portion globose. Membranous pronotal margins broad, kidney-shaped,
bullate about the middle, slightly raised at the anterior and posterior margins,
Pronotum and base of triangular process distinctly and quite regularly punctured.
Apex of triangular process reticulated; the sides feebly raised anteriorly, with a
few areolae. Length, 3.1 mm. ; width, 1.6 mm.

Color: Antennae yellowish; fourth segment brownish. Legs yellowish; tips
of tarsi brownish. Hood hyaline, with one or two dorsal nervures more or less
embrowned. Membranous pronotal margins hyaline, sometimes one or two areolae
brown. Pronotum yellowish-brown. Elytra hyaline, with a band near the base,
tumid elevation, and more or less of a band near posterior end fuscous.

A common species found feeding on oak and the leaves show whitish
or yellowish-brown patches from their attacks. This species is quite
variable in color. We have specimens with the hood, membranous pro¬
notal margins, and the posterior portion of the elytra almost entirely
hyaline.

No doubt common thruout the state. We have numerous examples
from Columbus and Athens.

Corythucha pergandei Heidemann (Figs. 4 and 5).

Corythucha pergandei Heid., Proc. Ent. Soc., Wash., Vol. VIII, p. 10.

Antennae slender, the basal segment twice as long as the second. Rostrum
reaching the meso-metasternal suture. Pronotal hood considerably raised, ab¬
ruptly constricted about the middle; posterior portion globose, widely reticulated;
anterior portion triangular, more closely reticulated. Membranous pronotal mar¬
gins evenly reticulated, reniform, bullate about the middle, slightly raised at the
anterior and posterior margins. Pronotum and base of triangular process deeply

Pig. 4. Corythucha pergandei Heidemann, adult (after Heidemann).

punctured, the latter with the sides feebly raised anteriorly (a few distinct
areolae) and with the apex irregularly reticulated. Median carina moderately
raised, with five or six areolae and sometimes two additional ones formed by a

The Tingitoidea or “Lace-Bugs'" of Ohio

229

little cross-vein dividing the middle cell. Elytra with the costal area very un¬
evenly and widely reticulated. Length, 3.2 mm. ; width, 1.6 mm.

Color: General color brownish. Body black, except the margins of the
prosternum, the humeral area, and the last two abdominal segments fulvous.
Antennae yellowish ; the apical segment brownish. Legs yellow ; the tips of tarsi
brownish. Areolae of hood and membranous pronotal margins hyaline, a few
slightly embrowned; nervures yellowish or brownish. Elytra hyaline; humeral
area and a band across the apical third dark-fuscous with the centers of areolae
hyaline.

One specimen, taken at Sugar Grove by the junior author. It is
recorded as occurring on Almis.

Corythucha crataegi spec. nov. (Plate X, fig. f).

This species was listed by Morrill in Psyche, Vol. X, p. 132, as a new
subspecies of C. arcuata under the name of Corythucha arciiata crataegi.
Morrill finds the eggs of the oak- and hawthorn-inhabitating .species to
be distinct and gives a brief description of the ova, but no description
of the adult. We have a long series of both forms and find them to be
distinct species.

Antennae slender, beset with numerous bristly hairs of various lengths; first
segment about three times as long as the second ; fourth segment swollen on the
distal half. Rostrum reaching the meso-metasternal suture. Hood considerably
raised, abruptly constricted about the middle; posterior portion globose, widely
reticulated; anterior portion triangular, more closely reticulated. Membranous

230

Ohio Biological Survey

pronotal margins evenly reticulated, reniform, bullate about the middle, slightly
reflected at the anterior and posterior margins. Pronotum coarsely and regularly
punctured ; triangular process reticulated, feebly raised on the sides anteriorly, with
two or three areolae; median carina foliaceous, with one row of areolae and two
additional ones formed by a small nervure dividing two middle cells. Elytra
with the outer margin sinuate, very broadly rounded at the tip ; costal area with
three rows of areolae and a partial fourth at anterior end, widely reticulated;
sutural area not very widely and unevenly reticulated. Length, 3.5 mm.; width,
1.7 mm.

Color: Body black, the apical segment of the abdomen and more or less of
the sides of the abdomen flavous. Antennae yellow ; distal half of apical segment
brownish. Legs yellowish; tips of tarsi brownish. Hood hyaline; parts of the
nervures and areolae brownish. Membranous pronotal margins yellowish, with a
brown spot near the outer margin. Pronotum yellowish. Elytra hyaline; a band
at anterior and one near posterior margin fuscous ; sutural area slightly em¬
browned.

A common insect upon hawthorn. We have a long series, taken
at Athens, Sept. 12th, 1905, by the senior author.

Corythucha contracta n. sp.

Readily distinguishable by the transverse black-fuscous bands near
the anterior and posterior margins of the elytra and the elytra are nar¬
rowed posteriorly.

Antennae slender, beset with a few moderately long bristly hairs; first seg¬
ment three times as long as the second; fourth segment moderately swollen
towards the apex. Hood considerably raised, abruptly constricted about the mid¬
dle; posterior portion globose, evenly reticulated; anterior portion triangular, more
closely reticulated. Membranous pronotal margins broad, reniform, bullate about
the middle, slightly reflected at the posterior margin. Pronotum and base of tri¬
angular process distinctly punctured ; the sides of the latter feebly raised anteriorly,
with a few distinct areolae. Elytra narrowed posteriorly; costal area very broad,
with three complete and one partial series of areolae, very unevenly reticulated.
Length, 3.25 mm.; width, 1.5 mm.

Color: General color griseus. Antennae yellowish; terminal segment brown¬
ish. Legs yellowish; tarsi brown. Pronotum and base of triangular process
brownish. Elytra griseus, with transverse black-fuscous bands near the anterior
and posterior margins.

One specimen, taken at Jefferson, Ashtabula Co., Aug. 8th, 1915
(Sim) .

Corythucha pallida spec. nov.

Easily distinguishable from the other Ohio species by its pallid color.

Antennae beset with long bristly hairs of various lengths; fourth segment
swollen towards the apex. Rostrum very long, almost reaching the end of the
mesosternum. Membranous pronotal margins very broad, evenly reticulated, not

The Tingitoidea or “Lace-Bugs'" of Ohio

231

very strongly bullate about the middle, slightly reflected at the posterior margins.
Hood moderately raised, not very abruptly constricted about the middle. Median
Carina with one complete series of areolae and two cells divided near the middle
forming double cells. Pronotum regularly and coarsely punctured. Apex of tri¬
angular process reticulated; the sides feebly raised anteriorly, with two or three
small areolae. Costal area with three complete and one partial series of areolae,
very unevenly reticulated. Length, 4.1 mm. ; width, 2 mm.

Color: Body brown-black. Lateral portion of prothrax and terminal segment
of abdomen ochraceous. Antennae yellowish ; apical segment embrowned. Legs
yellowish; tarsi slightly embrowned and usually darker at tips. Hood, mem¬
branous pronotal margins, and triangular process pallid. Elytra pallid, with an
indistinct very light brownish band near the base. Length, 4 mm. ; width, 2 mm.

Five specimens, taken on linden at Sugar Grove (Dr. Kellicott), and
at Columbus (Prof. Hine).

Variety idniif We have numerous specimens taken on Ulnnis arneri-
canum (Sanders) at Lisbon, Sept. 4th, 1903. They are slightly smaller
and the transverse band near the base of the elytra is a little more dis¬
tinct. There are also a few more erect spines on the membranous prono¬
tal margins and elytra. The specimens seem to be slightly discolored
and with the material in hand we do not feel warranted in separating it as
a distinct species.

Corythucha pruni spec. nov.

Close to C. aesctila, but readily separated from it by the narrower and
much less elevated hood, the shorter first segment of the antennae, and
the slightly raised median carina at the middle.

Antennae beset with numerous bristly hairs ; first segment slightly more than
twice the length of the second ; fourth segment swollen on the distal half.
Rostrum reaching the meso-metasternal groove. Hood, narrow, moderately ele¬
vated, rather widely reticulated, not very abruptly constricted about the middle;
posterior portion subglobose ; anterior portion triangular. Membranous pronotal
margins very broad, evenly reticulated, bullate about the middle, slightly raised
at the anterior and posterior margins. Pronotum and base of triangular process
coarsely punctured, the sides of the latter feebly raised anteriorly with three or
four areolae. Median carina foliaceous, not strongly elevated about* the middle.
Elytra broadly rounded at the tip, the outer margin sinuate; costal area widely
and unevenly reticulated; with three complete and a partial row of areolae.
Length, 4.2 mm. ; width, 2.3 mm.

Color: Body black; apical abdominal segment fulvous. Antennae yellowish;
the apical segment embrowned. Legs yellowish ; tarsi embrowned. Hood hyaline ;
the nervures, especially on dorsal surface, more or less embrowned and a few of
the areolae somewhat clouded. Membranous pronotal margins hyaline, with one
or two small spots near the outer margin brown. Pronotum yellowish. Elytra
hyaline ; a band near the anterior and one near the posterior margins, and more

232

Ohio Biological Survey

or less of sutural area fuscous; the apical band with one or two large areolae
hyaline.

Described from numerous specimens, taken on the wild cherry, Pru-
nus serotina, at Washington, D. C., by Prof. Hine.

Corythucha aesculi n. sp. (Plate X, fig. g).

Moderately large and broad. Antennae slender, sparsely beset with a few
long bristly hairs ; first segment three times as long as the second ; second very
short; fourth segment moderately swollen towards the apex. Rostral groove
rather broad; rostrum reaching between the intermediate coxae. Membranous
pronotal margins broad; reniform, unevenly reticulated, bullate about the middle,
slightly reflected at the anterior and posterior margins. Hood strongly raised,
abruptly constricted about the middle; posterior portion very large, globose, quite
widely reticulated ; anterior portion triangular, more closely reticulated on the
sides, directed almost vertically downward. Median carina foliaceous, with one
complete and a partial second (about the middle) series of areolae. Pronotum
coarsely punctured. Sides of triangular process feebly raised anteriorly, with a
few distinct areolae. Costal area of the elytra very broad, unevenly reticulated,
with three complete and a partial fourth series of areolae. Length, 4 mm.; width,
3 mm.

Color: General color brownish. Body above and beneath black. Antennae
yellowish; terminal segment light fuscous. Legs yellowish; tips of tarsi brownish.
Pronotum embrowned. Membranous pronotal margins hyaline, with one or two
small areas about the middle near the outer margin brown. Hood more or less
embrowned, with the center of some of the areolae hyaline; nervures yellowish or
brownish. Elytra hyaline, with a broad transverse band near the base and apex
fuscous; in the latter band a few of the areolae hyaline.

Found in great numbers and in the various stages of metamorphosis
on the under sides of buckeye leaves, Aescula glabra, during the month
of April, May, and June, 1915 and 16, on the university campus and at
IMinerva Park, north of Columbus. This species hibernates in the adult
stage; it can be found feeding on the leaves in early spring just as soon
as they have partially opened.

Corythucha bulbosa n. sp. (Plate X, fig. e).

From any member of the genus known to us it is easily dififerentiated
by its larger size, the much larger pronotal hood, and dark-fuscous color.
According to Mr. H. M. Parshley (Harvard University) this species
was given the manuscript name, C. carbonafa, by Uhler, but no descrip¬
tion was ever published.

Antennae slender, beset with a few long bristly hairs ; first segment three
times as long as the second; fourth segment slightly incrassated, the bristly hairs
becoming shorter and more numerous towards the apex. Rostral sulcus wide ;
rostrum reaching beyond the meso-metasternal suture. Hood very large, very

The Tingitoidea or “Lace-Bugs'" of Ohio

233

strongly raised, very abruptly constricted about the middle; posterior portion very
large and wide, globose, very unevenly reticulated ; anterior portion triangular,
quite evenly reticulated, directed nearly vertically downwards. Membranous pro-
notal margins very broad, bullate about the middle, slightly reflected at the an¬
terior and posterior margins. Pronotum and base of triangular process deeply
and coarsely punctured ; the sides of the latter slightly raised anteriorly, with a
single areola at the base. Elytra very broad; costal area with three complete and
a partial series of areolae. Length, 4.6 mm. ; width, 2.6 mm.

Color: General color dark-fuscous. Body above and beneath black. An¬
tennae yellowish; apical segment brownish. Legs yellowish; tips of tarsi brownish.
Hood embrowned; the center of a few areolae hyaline. Membranous pronotal
margins dark-brown; one or two rows of areolae on the inner margin hyaline.
Pronotum and base of triangular process black-brown; apex of the latter yel¬
lowish. Elytra dark-brown ; the centers of two large areolae near the apex, a few
areolae on inner margin of costal area, dark-brown.

Described from numerous specimens, taken at Sugar Grove, May
15th, 1915, by Mr. Delong. We have specimens from Sugar Grove,
1902 (Osborn), and Plummer’s Island, Maryland (taken July 31st,
1906, by D. H. Clemmons).

Genus Gargaphia Stal.

Gargaphia Stal, Stett. Ent. Zeit., p. 324, 1862; Enum. Hem., Band. HI, p. 119
et 124, 1873.

The genus Gargaphia can be separated from the rest of the Tingitidae
by the interrupted rostral sulcus between the meso- and metasternum
by a prominent transverse, sinuous carina.

KEY TO THE OHIO SPECIES OF GARGAPHIA.

1. Size small (length, 3.4 mm.; width, 1.6 mm.); head with short blunt

spines . . . G. angiilata Heidemann

Size larger (4 mm. or longer); head with long, acute spines . 2

2. Membranous pronotal margins broadly rounded; antennae testaceous, ex¬

cept terminal segment black . G. tiliae^ Walsh

Membranous pronotal margins more narrowly rounded; first segment of
antennae black, second testaceous, third yellowish-white, fourth black..

G. solani Heidemann

Gargaphia angulata Heidemann (Fig. 6).

Gargaphia angulata Heidemann, Can. Entom., Vol. XXXI, p. 301; Chittenden,
Bull. 23, XTw Series, U. S. Div. Entom. (fig. 9), p. 32-33.

Head with three, short, rather thick, obtuse spines in front between the
antennae; antennae slender, beset with fine hairs; first segment nearly twice as
long as the second; third segment about four times as long as the fourth. Pro-

*As we have no specimens of G. amorpha Walsh, we do not try to sei)arate it from G.
tiliae in this key.

234

Ohio Biological Survey

notum finely punctate, tricarinate ; the three carinae foliaceous. Membranous
pronotal margins, moderately wide, angularly dilated behind the middle, with two
rows of areolae at the narrowest portion and with three series at the widest part.
Elytra oblong-oval, the sides abruptly converging to the base, broadly rounded at
the apex. Length, 3.4 mm. ; width, 1.6 mm.

Fig. 6. Gargaphia angulata Heidemann (after Chittenden).

Color: Body black beneath. Head black; spines white. Antennae testa¬
ceous; apical segment blackish. Legs testaceous; tips of tarsi blackish. Pro-
notum blackish ; carinae testaceous, the median carina posteriorly whitish. Nerv-
ures of elytra, membranous pronotal margins, and hood testaceous; three or four
transverse nervures of elytra blackish; areolae quite hyaline.

No specimens have been taken in the state. It has been taken in the
eastern and southern part of the United States on bean. We have
specimens from Clarksville, Tennessee, (DeLong), Beach Bluf¥, Mass.,
(Parshley), Washington, D. C., (Heidemann), Bedford, Penn., and
Balsam, N. C.

Gargaphia tiliae Walsh.

Tingis tiliae Walsh, Proc. Entom. Soc. Phil., Vol. Ill, p. 408, 1864.

Gargaphia fasciata Stal, Enum. Hem., Band. HI, p. 125, 1873.

We have found fasciata Stal to be a synonym of tiliae Walsh; this
has been confirmed by Mr. Heidemann in a letter to the senior author.

Head with five long, acute spines. Antennae long, slender, hairy; first seg¬
ment greatly swollen, about three times the length of the second ; second segment
short, less swollen ; third segment very long, slender ; fourth segment slightly
swollen towards the tip. Rostrum reaching a little beyond the middle of the
metasternum. Pronotum tricarinate, the outer carinae slightly sinuate. Mem¬
branous pronotal margins foliaceous, broadly reflexed, composed of four irregular
series of areolae. Elytra broadly rounded at the tip ; costal area broad, unevenly
reticulated; discoidal area not quite reaching the middle of the elytra. Length,
4.2 to 4.6 mm. ; width, 2.1 to 2.2 mm.

Color: General color pale yellowish-brown. Head more or less blackish. Eyes
black. Body beneath blackish or brownish-black. Antennae pale yellowish-brown ;

The Tingitoidea or “Lace-Bugs'" of Ohio

235

fourth segment blackish, Pronotum blackish ; membranous margins, hood, and
apex pale yellowish-brown. Legs pale yellowish-brown ; tips of tarsi blackish.
Nervures of elytra pale yellowish-brown; more or less of three or four transverse
nervures near basal third dark-brown or black ; areolae hyaline.

Common on Basswood; Cedar Point (Kostir), Berea (Drake), Jeffer¬
son (Sim), Columbus (Drake), Oxford (Shideler), Milan and San¬
dusky (Osborn), and Vinton (Hine).

Gargaphia amorphae Walsh.

Tingis amorphae Walsh, Proc. Entom. Soc. Phil., Vol. Ill, p. 408, 1864.

This species is unknown to us and we include the original description
of Dr. Walsh.

“Differs from the above {tiliae'\ only in the large, basal, carinate cell of the
elytra, terminating behind nearly in an angle of 60-80°, and in the veins of the
wings, both those on the basal side of the middle and those at the tip, being on
the average of specimens much more deeply stained with black, though individuals
of the two species occur which are identical in this character. Length about .15
inch. Eighteen specimens on Amorpha fructicosa."

Gargaphia solani Heidemann (Fig. 7).

Gargaphia solani Heidemann, Proc. Entom. Soc. Wash., Vol. 15, p. 136; Fink
(Life Hitory), Bull. 239, U. S. Dept. Agr.

Fig. 7. Gargaphia solani Heidemann (after Fink),

236

Ohio Biological Survey

Head deeply punctured, with five long acute spines. Antennae long, slender,
hairy; first segment greatly swollen, about three times the length of the second;
second segment short, much less swollen ; third segment more than four times the
length of the fourth; fourth segment fusiform. Pronotum feebly convex, with
three longitudinal carinae. Lateral membranous pronotal margins angularly ex¬
panded, somewhat broadly reflexed, with two to five series of irregular areolae.
Pronotal hood rather large, covering the posterior part of the head, with fine,
minute areolae. Head, pronotum, and membranous pronotal margins more o’
less densely covered with fine, soft hairs. Elytra broadly rounded at the tip, feebly
sinuate near the base; discoidal area pyriform, almost reaching the middle of the
elytra. Length, 4mm. ; width, 2 mm.

Color: Body dark brown. Antennae — first segment black; second segment
testaceous; third segment yellowish-white; fourth segment black at the apex.
Pronotum black ; carinae yellowish. Legs pale yellow. Lateral membranous mar¬
gins of the pronotum and nervures of the elytra yellowish, some of them more
or less embrowned or blackish ; areolae somewhat hyaline.

This species is commonly known as the eggplant lace-bug. Accord¬
ing to Fink, this lace-bug destroys from 10 to 15 per cent of the eggplant
crop in the vicinity of Norfolk, Va.

Very common in the southern part of the state; taken on the horse
nettle (Solaniim carolinensc) at Malta, Apple Grove, North Kenova,
Hanging Rock, Crown City by the junior author; and in Clermont Co.
by Mr. Jesse Robinson.

Genus Galeatus Curtis.

Galeatus Curtis, Entom. Mag., p. 196, 1833.

Galeatus Stal, Enum. Hem., Band. HI, p. 118, 1873.

Head armed with long, acute spines. Rostral sulcus rather wide,
uninterrupted; rostrum long and slender. Antennae and legs long and
slender. Lateral margins of the pronotum foliaceous, with a few large
areolae forming a single series. Pronotal hood extending over the me¬
dian anterior and posterior portions of the pronotum ; discal hood large,
globose, divided behind. Median carina foliaceous. Elytra extending
nearly one-half beyond the apex of the abdomen, outer margin near the
base abruptly narrowed, broadly rounded at the apex, composed of very
large areolae ; costal area wide, composed of a single series of areolae ;
inner margin of the subcostal area jointly raised with the margins of
discoidal and sutural areas.

This genus can be readily differentiated from the allied genera by
the very large areolae. Only one Nearctic species has been described.

Galeatus peckhami Ashmead (Plate X, fig. c).

Sphaerocisfa peckhauii A.shmead, Entom. xA.mer., Vol. HI, p. 156, 1888.

The Tingitoidea or “Lace-Bugs'" of Ohio

237

Galeatus peckhami Van Duzee, Canad. Entom., Vol. XXI, p. 5., 1889; Uhler,
Proc. U. S. Nat. Mus., Vol. XXVII, p. 364, 1904; Parshley, Entom. News,
Vol. XXVII, p. 105, 1916

Head with five very long acute spines. Antennae slender, beset with nu¬
merous short hairs; first segment swollen, twice the length of the second; second
short, slightly less swollen ; third segment very long, slender ; fourth segment
rather long, apical third slightly swollen. Rostrum long, almost reaching the
end of the rostral groove. Membranous pronotal margins greatly dilated, composed
of five areolae (anterior and posterior areolae very small, the other three large
and transverse). Pronotal hood extending slightly over the base of the head.
Discal hood very large, globose, divided in the middle behind. Median carina
foliaceous, composed of a single series of areolae, extending from the posterior
end of the pronotum forward between the divided discal hood. Elytra with the
basal angle deeply sinuate, composed of large areolae; costal area wide, composed
of a single row of mostly large quadrate areolae. Length, 4.2 mm.; width, 2.3 mm.

Color: Head and body black. Antennae yellowish ; apical segment em¬
browned. Legs yellowish; tarsi fuscous. Nervures of hood, pronotal margins,
and elytra more or less embrowned or fuscous ; areolae hyaline, except a few
clouded ones on the discal vesicle.

This curious insect has not been taken in the state. It is recorded for
Massachusetts (Parshley), Canada (Van Duzee), Wisconsin (Ash-
mead), and New Mexico (Uhler).

Genus Leptostyla Stal.

Leptostyla Stal, Enum. Hemip., Band. Ill ; p. 120, 1873.

Leptostyla Champion, Biol., Centr.-Amer., Rhynch., Vol. II, p. 11.

The genus includes three Nearctic species. Antennae long, slender;
first segment from two to five times as long as the second ; second seg¬
ment short ; third segment very long, cylindrical ; fourth segment more
or less swollen towards the apex, longer than the first. Rostral sulcus
uninterrupted by a transverse ridge between the meso- and metasternum ;
rostrum reaching the meso-metasternal suture, sometimes shorter or
longer. Pronotum tricarinate ; membranous margins moderately wide.
The hood varies in size. Elytra without a tumid elevation, constricted
about the middle, gradually widening towards the apex, extending far
beyond the tip of the abdomen ; costal area more or less widely reticu¬
lated, with one, two, three, or more irregular rows of areolae ; subcostal
area sometimes very narrow, with one, two, or three rows of areolae ;
discoidal area not reaching the middle of the elytra, with three or four
rows of small areolae ; sutural area more or less widelv reticulated.
Wings not reaching or extending slightly beyond the apex of the abdo¬
men.

238

Ohio Biological Survey

KEY TO THE NEARCTIC SPECIES OF LEPTOSTYLA.

1, Costal area of the elytra with a transverse, fuscous fascia just before the

middle; first segment of the antennae twice as long as the second .

L. clitoriae Heidemann

Costal area of the elytra without a transverse fascia; first segment of the

antennae three or slightly more than three times as long as the second. . 2

2. Costal area of the elytra white, size small (length, 2.75 mm.; width, 1 mm.)

L. oblonga Say

Veins of costal area more or less embrowned; size larger (length, 3 mm.;

width, 1 mm.) . L. heidemanni n. sp.

Leptostyla oblonga Say.

Tingis oblonga Say, Journ. Acad. Sci. Phil., Vol. IV, p. 325; Compl. Writ.,
Vol. II, p. 248, 1859.

This species can be distinguished from L. clitoriae Heid. by its larger
size, the long acute spines on the head, the much longer first segment of
the antennae, and the white costal area of the elytra. From L. heide¬
manni n. sp., it differs in its smaller size, white spines, and white costal
area of the elytra ; the costal area has one complete and a partial series of
areolae about the middle. Length, 2.75 mm. ; width, 1 mm.

No specimens have been taken in the state; we have included it as a
form that probably occurs here.

Leptostyla heidemanni n. sp.

Close to L. oblonga Say, but differing in the more swollen first seg¬
ment of the antennae, the four long acute spines on the head with black
tips, and the costal area of the elytra has two almost complete series of
areolae.

Head short, with four long acute spines. Antennae rather long; first segment
three times as long as the second; second segment very short; third very long;
fourth segment swollen on the distal two-thirds. Rostrum reaching the middle
of the mesosternum. Membranous pronotal margins moderately wide, recurved,
converging anteriorly, with two series of areolae. Hood oval, considerably raised.
Pronotal disc coarsely and regularly punctured. The three longitudinal carinae
raised, each with a complete series of areolae. Elytra with three large areolae
near the apex ; costal area with one complete and another nearly complete series
of areolae, very unevenly reticulated; subcostal and discoidal area closely and
evenly reticulated ; sutural area rather broad, unevenly reticulated. Length, 3 mm. ;
width, 1 mm.

Color: Body black. First segment of the antennae fulvous; second and
third segment yellowish ; fourth segment fuscous, except proximal third yellowish.
Legs yellowish ; tarsi more or less embrowned. Disc of pronotum black. Mem¬
branous pronotal margins hyaline, a few nervures on inner humeral area brown.

The Tingitoidea or “Lace-Bugs'" of Ohio

239

Hood hyaline; the nervures brown. Elytra hyaline, a spot on the discoidal area
(usually extending over in the subcostal area), transverse nervures of costal area,
and nervures of sutural area mostly fuscous.

Described from two specimens, one from Arkansas (Osborn collec¬
tion) and one from Washington, D. C. (Heidemann). We have just
received a specimen from Mr. Parshley that was collected at Newton,
Mass.

Leptostyla clitoriae Heid. (Fig. 8).

Leptostyla clitoriae Heidemann, Proc. Ent. Soc. Wash., Vol. XHI, p. 180.

Leptostyla costofasciata Drake, Ohio Journ. Sci., Vol. XVI, p. 326, 1916.

Head with three short, rather blunt spines — the two smaller, slightly converging
spines situated just above the antennae (one on the inner side of each) ; the larger
frontal spine is just above the other two and extends a little farther forward.
Antennae slender, about one-half the length of the body ; first segment swollen.

twice as long as the second : fourth segment fusiform. Rostrum almost reaching the
meso-metasternal suture. Pronotum norrowed anteriorly, with the membranous mar¬
gins moderately wide, recurved, converging anteriorly, with two rows of areolae ; hood
oval, considerably raised, with about eight areolae on each side ; the three longitud¬
inal carinae raised, composed of a single row of areolae. Costal area of the elytra

240

Ohio Biological Survey

moderately wide, rather widely reticulated, with two rows of areolae; subcostal
area rather closely reticulated ; discoidal area more closely reticulated ; sutural area
unevenly reticulated, with two large areolae near the posterior end. Wings not
longer than the abdomen. Length, 2.2 mm. ; width, 9 mm_.

Color: Body black. Legs yellowish; tips of tarsi fuscous. Antennae — first and
second segments testaceous; third segment yellowish; fourth segment fuscous.
Hood whitish, nervures more or less testaceous ; membranous pronotal margins
whitish, the nervures of four or five areolae on humeri brownish. Elytra whitish,
with a rather broad, transverse, dark-fuscous fascia just before the middle on the
costal area; nervures of subcostal, discoidal, and sutural area dark-fuscous. The
apex of the elytra dark fuscous with the center of the areolae whitish.

We have specimens from Knoxville, Tennessee and Calhoun, South
Carolina.

Genus Leptobyrsa Stal.

Leptobyrsa Stal. Enum. Hemip., Band. Ill, p. 119, 1873.

Leptobyrsa Champion, Biol. Centr.-Amer., Rhynch., Vol. II, p. 25.

Head with, at most, five rather short spines. Antennae slender ; the
basal segment long; third segment very long. Rostral groove uninter¬
rupted, broad on the meso- and metasternum and closed behind ; rostrum
rather short. Pronotal hood large, extending prominently above the
head. Membranous pronotal margins foliaceous, broadly rounded. Pos¬
terior portion of the pronotum abbreviated, obtuse at the tip. Elytra
twice or nearly twice as long as the abdomen, broad, more or less widely
reticulated, strongly rounded at the base and apex. Wings short or
obsolete.

The genus contains only one described Nearctic species.

Leptobyrsa explanata Heidemann (Plate VIII)

Leptobyrsa explanata Heidemann, Proc. Entom. Soc. Wash., Vol. II, p. 105.

Body short, oval in the female and more elongate in the female. Head rather
short, with three frontal spines and two other more slender ones extending from be¬
hind the eyes towards the front. Antennae long, slender, finely pilose, the first two
segments swollen; first segment twice the length of the second; third segment a
little more than three times the length of the fourth. Pronotum feebly convex,
coarsely punctured; membranous margins strongly rounded behind, reflexed,
widening moderately at the sides, with two or three rows of areolae : hood not
much inflated, covering the head (except eyes), with quite large areolae at sides
near the top and smaller ones near the base, cristate and slightly tapering towards
the front. The three pronotal carinae foliaceous; the median carina strongly
raised, as high as the crest of the hood, rounded on top and slowly declining
towards the apex of triangular portion of pronotum; outer carinae feebly raised,
one-half as long as the median carinae. Elytra ovate, strongly rounded from base
to apex, broadest behind the^ middle. Crest of hood, lateral margins of pronotum,

The Tingitoidea or “Lace-Bugs'" of Ohio

241

outer margins of elytra, and most of the nervures beset closely with very fine, long
hairs. Length, 3.6 mm.; width of each elytron across widest part, 1.4 mm.

Plate VII. — Leptobyrsa explanata Heid. Fig. a, adult; fig. b, nymph (after Heide-

mann).

Color: Body shining black. Bucculae yellow. Antennae yellowish; apex in-
fuscated. Pronotum .shining black ; carinae yellowish. Hood, membranous pro-
notal margins, and elytra yellowish ; areolae semitranslucent, pale yellow.

Warren, Ohio, July 15th, 1897 (H. B. Perkins).

Genus Leptophya Stal.

Leptophya Stal, Enum. Hem., Band. Ill, p. 121, 1873.

Antennae rather stout ; first segment less than twice the length of the
second ; second segment short ; third segment not at all slender ; fourth
segment fusiform. Rostral sulcus uninterrupted by a transverse ridge,
open at the tip. Pronotum closely reticulated, with a single, median
Carina ; lateral margins not foliaceous. Pronotal hood very broad, ex¬
tending over the base of the head. Elytra narrowed at the base, broadly
rounded at the tip, a little longer than the abdomen. Wings extending
beyond the apex of the abdomen.

The genus contains only one described Nearctic species.

Leptophya mutica Say (Plate X, fig. b).

Leptophya mutica Say, Hem. New Harm., p. 26, 1832; Compl. Writ., Vol. I,
p. 349, 1859.

Head with two spines deflected against its surface. Antennae rather stout,
about of a uniform thickness; first and second segments about equal in length;
third segment long, cylindrical ; fourth segment fusiform. Rostrum reaching the

242

Ohio Biological Survey

middle of the mesosternum. Pronotum very closely reticulated; apex of the elytra
more widely reticulated. Length, 3 mm. ; width, 1.1 mm.

Color: General color grayish-brown. Body beneath dusky. Elytra grayish-
brown, with a spot or band near the middle and the nervures at tip dark-brown.

No specimens have been collected in the state, altho the species prob¬
ably occurs as it has been taken in Indiana.

Genus Physatochila Fieber.

PJiysatochila Stal, Enum. Hem., Band. Ill, p. 120, 1873.

Head with the spines bent down against its surface. Antennae mod¬
erately slender ; first segment swollen,- less than twice the length of the
second ; second segment short, less swollen ; third segment very long,
slender ; fourth segment fusiform. Rostral groove uninterrupted, open
at the tip. Hood moderately large, extending over a small portion of
the head. Pronotum closely reticulated, with three longitudinal carinae ;
membranous margins foliaceous, obtusely rounded, reflexed back against
the surface of the pronotum. Elytra narrowed at the base, broadly
rounded at the tip, extending beyond the apex of the abdomen ; subcostal
area very long, narrow ; discoidal area extending beyond the middle of
the elytra. Wings longer than the abdomen.

KEY TO THE SPECIES OF PHYSATOCHILA.

Rostrum reaching beyond the first segment of the abdomen; elytra gray-brown,

the nervures more or less fuscous . P. plexa Say

Rostrum reaching between the intermediate coxae ; elytra dull-brown, con-

colorous . P. hrevirosfris n. sp.

Physatochila plexa Say.

Physatochila plexa Say, Hem. New Harm., p. 26, 1832; Compl. Writ., Vol. I,
p. 349.

Physatochila plexa Stal, Enum. Hem., Band. HI, p. 129, 1873.

Head with five spines bent down against its surface. Antennae moderately
long; first segment less than twice the length of the second. Rostrum very long,
reaching the middle of the second segment of the abdomen. Reflexed pronotal
margins extending to or slightly over the outer longitudinal carina. Elytra reach¬
ing a little beyond the apex of the abdomen ; costal area with three complete or
two and a partial third irregular series of areolae. Wings extending a little be¬
yond the apex of the abdomen. Length, 3.4 mm. ; width, 1.5 mm.

Color: General color grayish-brown and more or less marked with fuscous.
Body beneath brownish. Antennae brownish ; apex of terminal segment fuscous.
Legs brownish ; tarsi darker. Pronotum and elytra grayish-brown ; nervures
marked with fuscous.

This species differs from P. brcvirosfris by its more oval shape, the
longer rostrum, and the shorter elytra and wings.

The Tingitoidea or “Lace-Bugs’" of Ohio

243

We have no Ohio specimens, but as it was described from Indiana
we have included it as a form that probably occurs in the state. We have
specimens from Fairfax, Iowa; Agricultural College, Mich.; Ottawa,
Canada; and Berkley Springs, W. Va.

Physatochila brevirostris spec. nov.

From its only Nearctic congener, P. plexa Say, it is readily sepa¬
rated by its broader and less oval shape, the much shorter rostrum, the
three quite regular series of costal areolae, and the concolorus elytra.

Antennae moderately slender ; first segment swollen, slightly longer than
the second; second segment short, less swollen; third segment very long, cylin¬
drical; fourth segment fusiform. Rostral sulcus moderately wide, open at the
apex; rostrum reaching between the intermediate coxae. Pronotum closely reticu¬
lated ; the three longitudinal carinae slightly raised, composed of a single series of
small areolae. Elytra narrowed at the base, very broadly rounded at the tip , con¬
siderably longer than the abdomen ; costal area composed of three quite regular
series of areolae; subcostal area very long, narrow, composed of three incom¬
plete series of areolae; discoidal area reaching beyond the middle of the elytra.
Wings considerably longer than the abdomen. Length, 3.2 mm.; width, 1.6 mm.

Color: General color dull brown. Body beneath brownish. Legs dull brown;
tarsi fuscous. Antennae dull brown; apex of fourth segment black. Hood, re¬
flexed pronotal margins, and pronotum dull brown, a few small areas slightly
darker. Elytra dull brown.

Described from two specimens, one taken at Chillicothe (Ross Co.,
O.), May 21st, 1904, by Mr. O. H. Swezey, and the other at New
York City, N. Y., May 20th, 1889, by Mr. E. B. Southwick.

Genus Melanorhopala Stal.

MelanorJiopala Stal, Enum. Hemip., Band. Ill, p. 130, 1873,

Head armed with five deflected spines. Rostral groove uninterrupted.
Antennae stout, long, clavate or subclavate ; third segment very long,
slightly curved, more or less swollen at the apex ; fourth segment conical
or fusiform. Pronotum closely reticulated, with three longitudinal
carinae ; sides foliaceous, reflected back vertically, or against the sides of
the pronotum. Elytra narrowed at the base, apex narrowed or obliquely
rounded ; costal area composed of a single series of areolae ; subcostal area
narrow, with two series of areolae ; discoidal area reaching beyond the
middle of the elvtra.

KEY TO THE SPECIES OF MELANORHOPALA.

1. Antennae clavate ; apex of third segment greatly swollen, the fourth seg¬
ment conical; membranous pronotal margins reflected, vertical (Plate
VHI, fig. A) . M. clavata Stal

244

Ohio Biological Survey

Antennae subclavate; apex of third segment slightly swollen, the fourth
segment fusiform; membranous pronotal margins reflected back against

or nearly against the surface of the pronotum . 2

2. Antennae very long (Plate VIII, fig. B) . M. lurida Stal

Antennae much shorter (Plate VIII, fig. C) . M. uniformis Stal

Melanorhopala clavata Stal (Plate VIII, fig. A; Plate X, fig. d).

Tingis (Melanorhopala) clavata Stal, Enum. Hemipt., Band. Ill, p. 130, 1873.

Antennae long clavate; first segment strongly swollen one and a half times
the length of the second; second segment less swollen, short; third segment long,
slender, greatly swollen at the apex ; fourth segment conical. Membranous pro¬
notal margins reflected back, vertical. Length, 5 to 5.3 mm. ; width, 1.9 to 2.1 mm.

Color: Dull yellow-brown; the swollen portion of third segment and apical
segment black.

We have specimens from Point Pleasant and Westfield, N. J., and
Newton, Mass. No specimens have been taken in the state.

Melanorhopala lurida Stal (Plate VIII, fig. B).

Tingis (Melanorhopala) lurida Stal, Enum. Hemipt., Band. Ill, p. 131, 1873.

Plate VIII. — Antennae. A, Melanorhophora clavata Stal ; B, M. lurida Stal ; C, M.
uniformis Stal.

Tpie Tingitoidea or “Lace-Bugs'" of Ohio

245

Antennae very long; second segment small at the base and swollen at the
apex; third segment very long, very slightly swollen at the apex; fourth segment
fusiform, longer than in clavata or uniformis. Membranous pronotal margins
reflected back against or nearly against the surface of the pronotum. Length,
male 5.5 mm., female 5.8 mm. ; with, male 1.9 to 2 mm., female 2.1 mm.

Color: Dull yellow-brown, slightly darker than clavata or uniformis; the
apex of the third segment and fourth segment black.

Two specimens, male and female taken at Castalia, O., by Mr. DeLong.
This species was listed for the state as clavata. We have two specimens
from New York City, N. Y. (E. B. Southwick).

Melanorhopala uniformis Stal (Plate VIII, fig. C).

Tingis (Melanorhopala) uniformis Stal, Enum. Hemipt., Band. Ill, p. 131, 1873.

Antennae considerably shorter than in clavata or lurida; second segment two-
thirds the length of the first, less swollen ; third segment much shorter than
clavata or lurida, slightly swollen at the tip; fourth segment fusiform. Mem¬
branous pronotal margins reflected back against or nearly against the surface of
the pronotum. Length, 5.5 mm. ; width, 2 mm.

Color: Dull yellow-brown; the apex of the third segment and fourth segment
black-brown.

This species was described from specimens taken in Illinois. We
have one specimen from South Dakota.

Genus Alveotingis gen. nov.

Head with, at least, five spines; eyes large, strongly granulate. An-
tenniferous tubercles very large and long, resembling the basal segment
of the antennae. Antennae strongly swollen, more than one-half the
length of the body ; first segment about twice the length of the second ;
second segment very short ; third segment very long, more strongly
swollen, largest ; fourth segment very short, conical. Rostral sulcus
rather broad, uninterrupted, open behind. Pronotum narrowed in front,
with three longitudinal carinae, reticulated, except the antero-lateral
areas) ; anterior margin with a single series of areolae; lateral margins
foliaceous, composed of a single series of areolae, reflexed back against
or nearly against the sides of the pronotum ; posterior portion very broad,
triangular. Legs long and robust. Elytra strongly convex; nervures
strongly raised ; costal area composed of a single series of areolae ; dis-
coidal, sutural, and subcostal areas not dififerentiated.

Type of genus, Alveotingis grossocerata n. sp.

Alveotingins grossocerata spec. nov. (Fig. 9).

Head with five spines — a single median spine between the eyes, two short
converging spines between the antenniferous tubercles, and two longer deflected

246

Ohio Biological Survey

spines inserted close to the pronotum and extending along the inner side of the
eyes. Rostral sulcus rather wide, uninterrupted. Eyes large, subglobose, rather
coarsely granulate. Antenniferous tubercles very large and long, appearing some¬
what like the first antennal segment. Antennae very stout, more than one-half the
length of the body, densely beset with short decumbent hairs ; first segment almost
twice the length of the second, slightly more swollen than the second ; second
segment very short; third segment very long, about one and two-thirds the length
of the other three conjoined; fourth segment very short, conical. Legs very long
and robust. Pronotum tricarinate, rather coarsely reticulated ; median carina
raised, composed of a single series of small areolae and becoming obsolete pos-

Fig. 9. Alveotingis grossocerata n. sp.

teriorly, reaching from the base to the apex of triangular process; discal carinae
sinuate, extending from near the base to sides of triangular process, composed
of a single series of small areolae; two series of areolae between the median and
lateral carinae. Membranous pronotal margins reflected back against or nearly
against the pronotum. Elytra broad, very strongly convex, outer margin rounded ;
costal area composed of a single series of large areolae; the rest of the elytra
rather coarsely reticulated. Length, 2.8 mm. ; width, 1.2 mm.

The Tingitoidea or “Lace-Bugs'" of Ohio

247

Color: General color grayish-fuscous. Body dull-black. Nervures of prono-
tum and elytra sooty-black; areolae whitish. Antennae and legs grayish-fuscous.

Described from one specimen, taken at Orono, Maine, Aug. 5th, 1913,
by the senior author. The body of this species is much thicker than that
of any other American Tingitid.

Date of publication, June 28th, 1916.

248

Ohio Biological Survey

EXPLANATION OF PLATE IX.

Corythucha arcuata Say (after Morrill).

Fig. 1. Laterval view of egg.

Fig. 2. Apical end of egg showing cap.

Fig. 3. Nymph, first instar.

Fig. 4. Trumpet-shaped spine, arising from a subconical base.

Fig. 5. Nymph, second instar.

Fig. 6. Nymph, third instar.

Fig. 7. Nymph, fourth instar.

Fig. 8. Trumpet-shaped spine arising from a long thick protuberence.
Fig. 9. Nymph, fifth instar.

The Tingitoidea or “Lace-Bugs'"

OF Ohio

249

Ohio Biological Survey

Vol. II, Plate IX.

I

Cl w dj..

250

Ohio Biological Survey

EXPLANATION OF PLATE X.

Fig. a. Piesma cinerea Say.

Fig. b. Leptophya mutica Say.

Fig. c. Galeatus peckhami Ashmead.
Fig. d. Melanorhopala clavata Stal.
Fig. e. Corythucha biilbosa spec nov.
Fig. /. Corythucha crataegi spec. nov.
Fig. g. Corythucha aesculi spec. nov.
Fig. h. Corythucha ciliata Say.

The Tingitoidea or “Lace-Bugs'^ of Ohio

251

Ohio Biological Survey

Vol. II, Plate X.

Osborn and Drake.

Volume IL No. 5

Bulletin No. 9

OHIO BIOLOGICAL SURVEY

THE GRASSES OF OHIO

John H. Schaffner

THE LIBRARY OF THE

NOV 2 2 1938

UNIVERSITY OF ILLINOIS

Published by

THE OHIO STATE UNIVERSITY
Columbus
1917

Contribution from the Department of Botany,
Ohio State University 98.

ilD.S

OH

0 .

CONTENTS

Introduction . 256

Synopsis of the Tribes . 261

Key to the Genera. . 267

Festuceae . 272

Aveneae . 287

Hordeae . 291

Chlorideae . 296

Agrostideae . 299

Phalarideae . — . 309

Paniceae . 311

Oryzeae . 323

Andropogoneae . 325

Maydeae . 328

INTRODUCTION

The grasses of our state, including both wild and cultivated species,
may properly be regarded as our most important vegetation ; for no other
plants have such a wide and fundamental bearing on the stability and
progress of our prosperity and civilization. There are about 4500 species
of true grass plants known in the world. Of these about 180 species
occur as native, introduced, or commonly cultivated in Ohio.

The grasses, outside of the bamboo tribe, are all herbacous, peren¬
nials, biennals or annuals. The stem, sometimes called the culm, is of
a very extreme type having exceedingly long internodes. It is either
hollow or solid, but if hollow it is always closed at the nodes. The leaves
are two-ranked and alternate. The leaf is also a highly specialized
structure usually having a linear blade with a more or less prominent
midrib and a large sheath which is open on the side opposite the blade.
On the upper side of the leaf between the sheath and the blade there is
often a permanent fringe of hairs or one or more scarious appendages
called ligules. The roots of grasses are fibrous and together with the
creeping rootstocks form the sod so characteristic of many species.

Grasses are very generally distributed throughout the world, often
forming extensive prairies, meadows, or plains, and furnishing great
quantities of food for grazing animals. They are of importance in
making an effective covering for large areas of the earth’s surface and
in protecting the land from erosion. The grasses yielding food grains
are called cereals and constitute the most substantial food plants for maM.
The most important of these grasses are Indian corn, rice, wheat, rye,
barley, oats, sorghum, millet and sugar cane.

Aside from the general uses mentioned above certain species of
grasses are employed in many other ways. Various species furnish
material for basketry and for hats, mats, and braidwork ; many species
are used in paper making ; some furnish various commercial starches, oils
and cellulose used in the arts ; some are used for fuel as for instance the
cobs of Indian corn ; certain species are used as soil binders on levees, in
river bottoms, and on sand dunes ; species belonging to the bamboo tribe
are used extensively for building houses and fences, for furniture, fishing
poles, and an endless assortment of small articles ; various species like rye
are used for thatching roofs ; broom corn, a variety of sorghum, is used
extensively in the manufacture of brooms and brushes ; the peculiar
fruits of job’s-tears are strung as beads; corn cobs are manufactured into
tobacco pipes ; some species are used in medicine ; various species are
cultivated as ornamental grasses in gardens and the panicles of some are
employed for dry winter boquets. Many other uses might be mentioned.

Grasses, of course, belong to the flowering plants, or Anthophyta.
They are classified botanically as follows :

256

GRASSES OF OHIO

257

Phylum, Anthophyta, flowering plants. Seed plants of diverse habit and com¬
monly with showy flowers on the sporophyte, with closed carpels or carpel sets,
with female gametophytes of eight or rarely sixteen, or a smaller number of cells,
usually containing two, or rarely more, polar nuclei which conjugate with each
other and with one of the two nonmotile sperms from the male gametophyte or
pollengrain, which is deposited on a stigma and develops a long pollenttibe. As the
result of this triple fusion a new tissue is produced of greater or less extent sur¬
rounding the developing embryo in the ovule. This peculiar endosperm is called
the xeniophyte generation and it is the presence of this tissue, especially, in many
grains of grasses that gives them their great food value.

The grasses belong to the class Monocqtylae, or monocotyls, which
may be defined as having the following characteristics :

Sporophytes developing as herbs or sometimes as woody plants of large
dimensions ; embryo usually with one terminal cotyledon and a lateral plumule ;
stem with closed, usually scattered vascular bundles, without typical bark and
annual rings of growth, rarely with secondary thickening ; leaves mostly parallel-
veined, sometimes netted-veined ; flowers more commonly trimerous, the ideal
being a trimerous, pentacyclic flower with united carpels.

The Monocotyls are divided into four subclasses, the grasses falling
into the subclass, Glumiflorae :

Glumiflorae (Glume-flowered Monocotyls). Usually grass-like herbs or some¬
times woody plants with hypogynous, inconspicuous flowers. Carpels united ;
stigmas 3-1 ; perianth usually of 6-2 minute segments or bristles or entirely want¬
ing; inflorescence usually consisting of spikes or spikelets variously clustered;
endosperm mealy, starchy or rarely sugary.

The order of the Glumiflorae to which the grasses belong is com¬
monly called- Graminales and is characterized by its unilocular ovulary
containing one anatropous, erect or ascending ovule.

There are two families of Graminales, Cyperaceae or Sedges and
Graminaccae or Grasses. In general the sedges constitute the lower and
less specialized group.

Graminaccae. Grass Family

Highly specialized, perennial, geophilous herbs or annuals, or sometimes woody
plants, with hollow or occasionally solid stems, having prominent internodes, the
nodes closed. Leaves 2-ranked, with sheaths, the sheaths usually split to the base;
upper end of the sheath usually with one or two ligules ; sometimes with a short
petiole between the sheath and the blade, in which a cleavage plane is developed.
Inflorescence a panicle, raceme, or spike, composed of spikelets. Spikelets and
flowers with 2-ranked glumes, each spikelet normally with two empty glumes at
its base and each flower normally with two flowering glumes, the outer one called
the lemma, the inner the palet. Flowers bisporangiate or monosporangiate, mone-
cious or diecious with all intermediate gradations, often completely vestigial.
Perianth of 3 usually 2 small bracts called lodicules which probably represent a
corolla or inner perianth cycle; stamens of the andrecium 6, or usually 3, some¬
times reduced to 2 or 1 ; anthers versatile, with 4 microsporangia ; gynecium highly
specialized, consisting of 3 united carpels forming a unilocular ovulary with one
ovule; stigmas 3, 2 or 1, usually 2, hairy or plumose. Fruit a dry seed-like grain
(caryopsis), or in the lower forms sometimes fleshy; endosperm .starchy or rarely
sugary.

258

GRASSES OF OHIO

TERMINOLOGY OF THE GRASS INFLORESCENCE

The inflorescence of a grass is made up of compact flower-bearing
branchlets known as spikelets. In general, the spikelet of a grass is of
the same importance in identification as the flower in most other groups.
The spikelet usually has two bracts at the base which are called the empty
glumes. These may be distinguished as the outer and inner empty
glumes, or the lower and upper empty glumes. Each flower is also nor¬
mally inclosed in two bracts, called the flowering glumes. The outer of
these glumes is called the lemma, the inner the palet. All of these bracts
can thus be called glumes collectively. Through reduction of the spikelet
and degeneration of the flower, part of the glumes may be absent or
vestigial, or extra glumes may be present. Usually there are 2 (some¬
times 3) minute bracts or scales at the base of the flower, within the
flowering glumes. These are called lodicules and are supposed to repre¬
sent a vestigial perianth. The axis of the spikelet is called a rachilla and
if the inflorescence is a spike, its main axis is called a rachis.

Fig. 1.
Fig. 2.

Fig. 3.

Fig. 4.

Fig. 5.

Fig. 6.

Fig. 7.
Fig. 8.
Fig. 9.
Fig. 10.
Fig. 11.

EXPLANATION OF PLATE XI
Diagram of spikelet of Bromus sp.

Diagram of spikelet of Panicum virgatum. The staminate flower has a
vestigial gynecium.

Diagram of spikelet of Chaetochloa viridis. The vestigial flower con¬
sists only of a lemma. There are bristles at the base of the spikelet.

Diagram of staminate spikelet of Coix lacryma-jobi. Each of the two
flowers has a vestigial gynecium.

Diagram of carpellate spikelet of Coix lacryma-jobi, enclosed in a modi¬
fied bony leaf sheath. The spikelet contains a vestigial lemma and a
carpellate flower with three vestigial stamens ; also two vestigial struc¬
tures marked as oval black spots.

Spikelet of Bromus racemosus showing the two empty glumes and nine
flowers.

Lower or outer empty glume of Panicum virgatum.

Inner or upper empty glume of Panicum virgatum.

Lemma of Eragrostis major.

Palet of Eragrostis major.

The two lodicules (vestigial perianth segments) at the base of the grain
of Triticum aestivum.

OHIO BIOLOGICAL SURVEY

VOL. 11. PLATE XI.

Schaffner

Schaffner

PLATE XTI

Representative stages in the evolution of the grass inflorescence.
Fig. 1. Inflorescence of Arundinaria macrosperma Mx.

Fig. 2. Panicle of Uniola paniculata L.

Fig. 3. Spike-like panicle of Pennisetum glancum (L.) R.Br.

Fig. 4. Carpellate inflorescence (ear with husks) of Zea mays L.

GRASSES OF OHIO

261

SYNOPSIS OF THE TRIBES OF GRASSES

I, Spikelets many-l-flowered ; rachilla usually articulated above the empty glumes

which are persistent after the fall of the flowers ; spikelets usually more or
less laterally compressed.

1. Aerial stems entirely woody or at least woody at the base, perennial ;

lodicules (perianth segments) often 3; leaf-blade with a short petiole
articulated with the sheath.

Subfamily, Bambusatae.

a. Tribe, Bamhuseae.

2. Aerial stems herbaceous and annual; lodicules usually 2; leaf-blades sessile,

without a joint.

Subfamily, Poacatae.

(1) . Spikelets 2-many-flowered, rarely with 1 perfect and 1 staminate

flower; in panicles, spike-like panicles or racemes.

a. Flowering glumes as long or longer than the empty glumes,

unawned or with a straight awn from the apex.

Tribe, Festuceae.

b. Flowering glumes generally shorter than the empty glumes, usually

with a bent awn on the back ; callus and usually the rachilla-
joints hairy. Tribe, Aveneae.

(2) . Spikelets 1-several-flowered, in rows, forming an equilateral or 1-

sided spike or raceme; sometimes monosporangiate.

a. Spikelets sessile in 2 opposite rows^ forming an equilateral spike ;

leaf-blades bearing at base a more or less well-marked pair of
auriculate appendages. Tribe, Hordeae.

b. Spikelets sessile in 2 rows on one side of a flattened axis, forming

1-sided spikes which are digitate or paniculate, or sometimes
solitary. Tribe, Chlorideae.

(3) . Spikelets with but one perfect flower, sometimes with additional

monosporangiate flowers ; always in panicles or racemes, not
in rows.

a. Empty glumes 2, the third glume (lemma) enclosing a perfect

flower just above the empty glumes; palet of the perfect flower
usually 2-nerved. Tribe, Agrostideae.

b. With 2 minute glumes (lemmas) above the 2 normal empty

glumes or sometimes with 2 staminate flowers, the perfect flower
on the top of the spikelet ; palet of the perfect flower usually
1-nerved or nerveless. Tribe, Phalarideae.

II. Spikelets usually 2-flowered or by degeneration 1-flowered ; rachilla articulated

below the empty glumes which are thus deciduous with the flowers ; spike¬

lets usually more or less dorsally compressed ; aerial stems annual.

Subfamily, Panicatae.

1. Flowering glumes, at least of the perfect flowers, similar in texture to the
empty glumes, or frequently coriaceous or chartaceous (indurated),
never thin and hyaline.

(1). Spikelets with 4 or more glumes, more or less dorsally compressed,
a. Flowering glumes of the perfect flowers chartaceous or coriaceous,
very dift'erent from the empty glumes. Tribe, Paniceae.

262

OHIO BIOLOGICAL SURVEY

b. Flowering glumes membranous.

(a) . Inflorescence paniculate, spikelets deciduous singly from the

ultimate branches ; first empty glume usually smaller and
narrower than the rest. Tribe, Tristegineae.

(b) . Inflorescence spikate; spikelets deciduous singly or in groups:

first empty glume usually larger than the rest, the second
one often spiny. Tribe, Zoysieae.

(2). Spikelets laterally compressed, with but 2 normal glumes, the empty
glumes reduced or wanting; spikelets often monosporangiate.

Tribe, Oryzeae.

2. Flowering glumes thin and hyaline, much more delicate in structure than the
thick-membranous or coriaceous empty glumes.

a. Spikelets in pairs, one usually sessile the other pedicellate, the sessile

spikelet with a perfect flower, the pedicellate one with a perfect,
staminate, or sterile (vestigial) flower; lemmas of the perfect flowers
usually awned. Tribe, Andropogoneae.

b, Spikelets monosporangiate, in separate inflorescences or in different

parts of the same inflorescence, the carpellate portion or inflores¬
cence below, the staminate above ; lemmas awnless.

Tribe, Maydeae.

SYNOPSIS OF THE GENERA IN EACH TRIBE

The number after the generic name refers to the list number.

Festuceae.

I. Rachilla and flowering glumes glabrous or with hairs shorter than the glumes.
A. Spikelets all alike.

1. Lemmas 5-many-nerved.

(1) . Spikelets not in dense 1-sided clusters.

((1)). Stigmas arising below a cushion-like appendage at the tip
of the ovtilary. Bronius. (1)

((2)). Stigmas arising at or near the tip of the ovulary.

a. Spikelets with 2 or more of the upper glumes empty, broad,

and enfolding each other. Melica. (2)

b. Spikelets with the upper glumes flower-bearing, or if

empty similar in shape to the other glumes.

(a) . Lemmas rounded on the back; at least below.

((a)). Lemma acute, pointed or awned at apex, its
nerves not prominent. Festuca. (3)

((b)). Lemma obtuse or subacute at the apex, usually
toothed, its nerves prominent.

Paniciilaria. (4)

(b) . Lemmas more or less strongly compressed and

keeled. Poa. (5)

(2) . Spikelets nearly sessile in dense l-sided clusters. Dactylis. (6)

2. Lemmas 1-3-nerved, rarely with faint additional nerves.

(1). Callus and nerves of the lemma glabrous or cobwebby, or the

callus sparingly bearded.

GRASSES OF OHIO

263

((!)). Lemma not coriaceous in fruit; grain beakless, not ex-
serted.

a, Rachilla not prolonged behind the palet of the uppermost

sterile flower; spikelets in an open panicle.

' Eragrostis. (7)

b. Rachilla prolonged behind the palet of the uppermost

flower ; spikelets 2-5-flowered, usually in a more or less
contracted panicle.

(a) . Inner empty glume broad and rounded at the tip;

very dissimilar from the outer glume.

Sphenopholis. (8)

(b) . Inner empty glume similar to the outer.

Koeleria. (9)

((2)). Lemma coriaceous in fruit, grain beaked and exserted.

Korycarpus. (10)

(2). Callus and nerves of the lemma densely bearded, not cobwebby.

((1)). Palet not ciliate-fringed ; internodes of the rachilla short.

Tridens. (11)

((2)). Palet conspicuously ciliate-fringed; internodes of the
rachilla long. Triplasis. (12)

B. Spikelets of 2 kinds, in small clusters, the upper perfect ones surrounded
by sterile spikelets below. Cynosurus. (13)

II. Rachilla with hairs longer than the flowering glumes. Phragmites. (14)

Aveneae.

I. Spikelets not deciduous, the empty glumes persistent.

A. Spikelets 2-many-flowered ; rachilla prolonged beyond the upper glume.

1. Awn projecting from between the lobes of the lemma, flattened and

twisted. Danthonia. (15)

2. Awn of the lemma dorsal, not flattened.

(1) . Spikelets 2-flowered, the upper bisporangiate, the lower staminate

and its glumes strongly awned. Arrhenatherum. (16)

(2) . Spikelets 2-several-flowered ; flowers all bisporangiate or the

uppermost imperfect..

a. Lemma keeled ; awn rising from above the middle ; spikelets

less than Yi inch long. Trisetum. (17)

b. Lemma rounded on the back.

(a) . Spikelets over ^2 in. long; grain furrowed, usually ad¬

herent to the flowering glumes. Avena. (18)

(b) . Spikelets less than ^ in. long; grain not furrowed, free.

Deschampsia. (19)

B. Spikelets with 2 bisporangiate flowers; rachilla not prolonged beyond the

upper flower. Aspris. (20)

II. Spikelets deciduous ; lower flower bisporangiate, upper staminate with a hook¬
like awn; plant velvety. Nothoholcns. (21)

264

OHIO BIOLOGICAL SURVEY

Hordeae.

I, Spikelets solitary at each joint of the rachis.

1. Spikelets with their backs or edges to the rachis. Lolium. (22)

2. Spikelets with their sides to the rachis,

a. Empty glumes 3-many-nerved.

(a) . Empty glumes lanceolate or linear; spikelets 3-many-flowered.

Agropyron. (23)

(b) . Empty glumes ovate; spikelets 2-6-flowered. Triticum. (24)

b. Empty glumes 1-nerved, subulate ; spikelets with 2 perfect flowers.

Secale. (25)

II. Spikelets 2-6 at each joint of the rachis, some of them may be reduced.

1. Spikelets 2-many-flowered.

a. Empty glumes but little smaller than the flowering glumes.

Elymtis. (26)

b. Empty glumes very small or none; spikelets in pairs. Hystrix. (27)

2. Spikelets 1-flowered or with a vestigial second flower; in threes at each joint

of the rachis. Hordenm. (28)

Chlorideae.

I. Spikelets 1-flowered, with no vestigial lemma.

1. Spikelets articulated below the empty glumes.

a. Rachis extending beyond the upper spikelet ; spikelets lanceolate.

Spartina. (29)

b. Rachis of the spikes not produced; spikelets orbicular

Beckmannia. (30)

2. Spikelets articulated above the empty glumes ; spikes digitate.

Capriola. (31)

II. Spikelets with two or more flowers.

1. With 2-several perfect flowers in each spikelet ; spikes usually digitate.

Eleusine. (32)

2. With 1 perfect flower in each spikelet; spikes solitary or racemose.

a. Spikes numerous, each with 12 or less spikelets. Atheropogon. (33)

b. Spikes 4 or less, each with 25 or more spikelets. Boutelona. (34)

Agrostideae.

I. Flowering glumes membranous, not firmer than the empty glumes.

A. Lemma awnless or with a dorsal awn ; flowering glumes loosely enclosing
the grain.

1. Empty glumes not conspicuously compressed; spikelets usually in open
or narrow panicles,

(1) . Lemma 1-nerved, awnless. S porobolus. (35)

(2) . Lemma 3-5-nerved.

GRASSES OF OHIO

265

((!)). Panicle open; lemma usually awned.

a. Flowers not stalked ; stamens 3 ; palet 2-nerved.

(a) . Lemma entire; rachilla not prolonged, or if so with

a tuft of long hairs.

((a)). Callus below the flowering glumes and the
prolonged rachilla with a tuft of long

hairs. Calamagrostis. (36)

((b)). Callus naked or with short hairs; rachilla
not prolonged. Agrostis. (37)

(b) . Lemma long-awed, with a bifid apex; rachilla pro¬

longed into a short bristle. Apera. (38)

b. Flowers plainly stalked ; stamen 1, palet usually 1-nerved.

Cinna. (39)

((2)). Panicle narrow and contracted; lemma awnless.

Ammophila. (40)

2. Empty glumes conspicuously compressed-carinate; spikelets in dense
spike-like panicles.

(1) . Empty glumes awnless.

((!)). Lemma awned below the middle; panicle exserted, cylindri¬
cal. Alopecurus. (41)

((2)). Lemma awnless; panicle partly included, ovoid.

Heleochloa. (42)

(2) . Empty glumes awned; lemma awnless. Phleum. (43)

%

B. Lemma with a terminal awn or awn-pointed ; flowering glumes closely en¬
closing the grain.

1. Rachilla not prolonged behind the palet; empty glumes usually evident.

Muhlenbergia. (44)

2. Rachilla extending beyond the base of the palet into a bristle ; empty

glumes minute. Brachyelytrum. (45)

II. Flowering glumes indurated at maturity, firmer than the empty glumes.

A. Lemmas awnless, their margins inrolled ; spikelets without a basal callus.

Milium. (46)

B. Lemmas awned, their margins flat ; spikelets with a basal callus.

1. Awn simple.

(1) . Lemma broad, its awn straight and deciduous. Oryzopsis (47)

(2) . Lemma narrow, its awn twisted and persistent. Stipa. (48)

2. Awn 3-branched ; callus acute. Aristida. (49)

Phalarideae.

1. Empty glumes equal or nearly so.

a. Third and fourth glumes (lemmas) subtending staminate flowers.

Savastana. (50)

b. Third and fourth glumes small and empty, or vestigial. Phalaris. (51)

2. Empty glumes very unequal; third and fourth glumes dorsally awned.

Anthoxanthum. (52)

266

OHIO BIOLOGICAL SURVEY

Paniceae.

I. Spikelets without a subtending involucre.

1. Spikelets arranged in true panicles. Panicum. (53)

2. Spikelets arranged in 1-sided recemes, sometimes paniculate.

(1) . Fruiting flowering glumes chartaceous, with flat hyaline margins.

a. Spikelets long-pediceled, in a diffuse pinacle. Leptoloma. (54)

b. Spikelets on slender spike-like, often digitate branches of the in¬

florescence. Syntherisma. (55)

(2) . Fruiting flowering glumes indurated, rigid, the margins inrolled, not

hyaline.

a. Some of the glumes awned; spikelets crowded in 1-sided branches

of the panicle. Echinochloa. (56)

b. Glumes awnless ; spikelets plano-convex, on secund branches of the

inflorescence. Paspalum. (57)

II. Spikelets with an involucre of bristles or a prickly bur.

1. Spikelets subtended by bristles. Chaetochloa. (58)

2. Spikelets enclosed in a globular, prickly, bur-like involucre. Cenchrus. (59)

Oryzeae.

1. Spikelets much flattened laterally, with bisporangiate flowers.

Homalocenchrus. (60)

2. Spikelets monosporangiate.

a. Spikelets intermixed in the panicle. Zizaniopsis.

b. With carpellate spikelets in the upper part of the panicle and staminate

spikelets below. Zizania. (61)

Andropogoneae.

1. Inflorescence paniculate, decompound.

a. Pedicellate spikelet present; stem solid. Holcus. (62)

b. Pedicellate spikelet wanting; stem hollow. Sorghastriim. (63)

2. Inflorescence simple or compound made up of 1 or more spike-like racemose

branches.

a. Spikelets alike, all with bisporangiate flowers ; rachis of the inflorescence

branches continuous. MiscantJnis. (64)

b. Sessile spikelets bisporangiate, with awns, the pedicellate spikelets staminate

or empty; inflorescence simple or with 6 or less digitate branches.

Andropogon. (65)

Maydeae.

1. Carpellate spikelets inclosed in pockets of the internodes of the enlarged and

flattened rachis; rachis breaking into joints at maturity; inflorescence stam¬
inate above, carpellate below.

a. Carpellate inflorescence not covered with husks. Tripsacum. (66)

b. Carpellate inflorescence covered with husks. Euchlaena.

2. Carpellate spikelets covered by a bony leaf-sheath or collected in an ear covered

with husks.

a. With a bony leaf-sheath covering the single large grain. Coix. (67)

b. With numerous grains in an ear covered with husks, or with husk-like

glumes around the grains. Zea. (68)

GRASSES OF OHIO

267

KEY TO THE GENERA OF OHIO GRASSES.

The number after the generic name refers to the list number.

1. Spikelets in panicles or racemes, or in spike-like racemes, not in definite
rows on the branches of the inflorescence, usually on distinct pedicles of
varying lengths. 2

1. Spikelets borne in definite rows (usually 2 rows) on a simple inflorescence
or on the branches of a compound inflorescence. 51

2. Spikelets not enclosed in a prickly bur. 3

2. Spikelets enclosed in a prickly bur; an annual grass. Ccnchriis. (59)

3. Spikelets articulated above the empty glumes. 4

3. Spikelets articulated below the empty glumes, which are occasionally minute. 38

4. Spikelets with 2 to many perfect flowers. 5

4. Spikelets with but 1 perfect flower, rarely with 1 or 2 additional staminate
flowers. 23

5. Flowering glumes as long or longer than the empty glumes, unawned or with
a straight awn from the apex. 6

5. Flowering glumes generally shorter than the empty glumes, usually with a
bent or twisted awn on the back below the apex of the lemma. 18.

6. Rachilla of the spikelet naked or with hairs shorter than the flowering glumes. 7

6. Rachilla with hairs longer than the flowering glumes; very tall stout grasses;

lowest flower staminate, the others perfect. Phragmites. (14)

7. Spikelets all alike. 8

7. Spikelets of 2 kinds in a dense spike-like inflorescence, the fertile 1-3-flowered,
the lower spikelets of the inflorescence clusters consisting of empty glumes.

Cynosurus. (13)

8. Lemma 5-many-nerved. 9

8. Lemma 3-1-nerved. 14

9. Spikelets not in dense 1-sided clusters on the inflorescence. 10

9. Spikelets nearly sessile in dense 1-sided clusters at the ends of the panicle
branches. Dactylis (6)

10. Spikelets with the upper flowering glumes like the lower fertile ones, or if
empty not enfolding each other. 11

10. Spikelets with the upper flowering glumes empty and enfolding each other

in a club-shaped mass. Melica (2)

11. Lemma keeled; base of the flowers often with webby hairs. Poa. (5)

11. Lemma rounded cn the back or keeled only at the summit. 12

12. Nerves of the lemma 5-7, prominent, parallel; style terminal; spikelets com¬
pressed cylindric, or little flattened. Panicularia. (4)

12. Nerves of the lemma not prominent. 13

13. Stigmas arising below the apex of the ovulary ; lemma 2-toothed.

Brornns. (1)

13. Stigmas at or near the apex of the ovulary; lemma entire. Festuca. (3)

—14—

14. Flowering glumes not coriaceous when mature, grain beakless and not ex-
serted. 15

14. Flowering glumes coriaceous in fruit, grain beaked and exserted ; upper 2-4

lemmas empty. Korycarpiis. (10)

15. Lateral nerves of the lemma glabrous. 16.

15. Lateral nerves of the lemma pilose. 17

16. Panicle open; spikelets 3-many-flowered. Eragrostis. (7)

16. Panicle dense and spike-like, spikelets 2-4-flowered. Kodcria. (9)

268

OHIO BIOLOGICAL SURVEY

17. Palet not ciliate-fringed ; internodes of the rachilla short; middle nerve
or all three, exciirrent between the acute lobes of the lemma. Tridens. (H)

17. Palet conspicuously ciliate-fringed; internodes of the rachilla long; middle
nerve excurrent between the truncate lobes of the lemma. Triplasis. (12)

18. Rachilla prolonged beyond the upper glumes or flowers. 19

18. Rachilla not prolonged beyond the upper glumes ; spikelets 2-flowered.

Aspris. (20)

19. Awn projecting from between the lobes of the lemma, flattened and twisted.

Danthonia. (15)

19. Awn of the lemma dorsal, not much flattened. 20

20. Spikelets 2-flowered, the upper bisporangiate, the lower staminate and its

glumes strongly awned. Arrhenatherum. (16)

20. Spikelets 2-several-flowered ; flowers all bisporangiate or the uppermost one
imperfect. 21

21. Lemma keeled, awn arising from above the middle; spikelets less than in.

long. Trisetum. (17)

21. Lemma rounded on the back. 22

22. Spikelet over Yz in. long; grain furrowed, usually adherent to the palet.

Avena. (18)

22. Spikelet less than ^2 in. long; grain unfurrowed, free from the palet.

Deschampsia. (19)

23. Spikelets with a single perfect flower often at the side of the rachilla, not
with vestigial flowers or glumes above the empty glumes. 24

23. Spikelets with a perfect flower at the top, sometimes with staminate or
vestigial flowers below. 36

24. Lemma membranous, not indurated, not firmer than the empty glumes. 25.

24. Lemma indurated at maturity, firmer than the empty glumes. 33

25. Lemma awnless or with a dorsal awn, loosely enclosing the grain. 26

25. Lemma with a terminal awn or awn-pointed, closely enfolding the grain. 32

26. Empty glumes awned; inflorescence dense; lemma awnless. Phleum. (43)

26. Empty glumes awnless. 27.

27. Lemma 1-nerved. 28

27. Lemma 3-5-nerved. 29

28. Inflorescence lax, somewhat contracted ; glumes not conspicuously com¬
pressed. Sporobolus. (35)

28. Inflorescence dense, spike-like ; glumes conspicuously compressed-carinate.

Heleochloa. (42)

29. Rachilla prolonged beyond the palet. 30

29. Rachilla not prolonged beyond the palet. Agrostis. (37)

30. Lemma long- or short-awned, sometimes on the back ; panicle lax or con¬

tracted. 31

30. Lemma awnless ; panicle dense and spike-like ; leaves with deep grooves on

the upper surface. Ammophila. (40)

31. Prolongation of the rachilla and callus with long hairs ; lemma short-awned

on the back; perennial. Calaniagrosfis. (36)

31. Prolongation of the rachilla naked; lemma long-awned ; annual. Apera. (38)

32. Rachilla not prolonged beyond the base of the palet ; lemma pointed or

awned. Miihlenbergia. (44)

32. Rachilla prolonged into a bristle behind the palet ; lemma long-awned.

Brachyelytrum. (45)

33. Lemma awnless, its margins inrolled, spikelets without a basal callus.

Milium. (46)

33. Lemma awned, its margins flat ; spikelets with a basal callus. 34

34. Awn simple. 35

34. Awn 3-parted, the branches divaricate. Aristida. (49)

GRASSES OF OHIO

269

35. Lemma broad, glabrous, its awn deciduous ; callus short, obtuse.

Oryzopsis (47)

35. Lemma narrow; its awn persistent; callus usually acute. Stipa. (48)

36. Spikelets with a terminal perfect flower and 2 staminate lateral flowers,

Savastana. (50)

36. Spikelets with but 1 flower and sometimes with mere vestiges of lateral
flowers. 37

37. Empty glumes very unequal; stamens 2; third and fourth glumes (above the

two empty glumes) awmed upon the back. Anthoxanthum. (52)

37. Empty glumes equal; stamens 3; third and fourth glumes small and ves¬
tigial not awned. Phalaris. (51)

—38—

38. Spikelets monecious, the staminate and carpellate spikelets in different parts
of the inflorescence. 39

38. Spikelets not monecious; staminate spikelets if present distributed among the
perfect ones. 40

39. Staminate and carpellate spikelets much alike in appearance, intermixed in

the panicle, but the staminate at the top of the branches. Zizaniopsis.

39. Staminate spikelets in the lower part of the panicle ; the carpellate above,

unlike in appearance. Zizania. (61)

40. Inflorescence a compact spike-like panicle. 41

40. Inflorescence a loose panicle or with slender branches. 42

41. With an involucre of bristles about the base of the spikelets.

Chaetochloa (58)

41. Spikelets without a subtending involucre of bristles. Alopecurns. (41)

42. Spikelets laterally compressed. 43

42. Spikelets round, or dorsally compressed. 45

43. Spikelets 2-3-flowered ; empty glumes dissimilar. Sphenopholis. (8)

43. Spikelets 1-flowered. 44

44. Glumes 4; lemma with a short awn between the bifid apex. China. (39)

44. Glumes only 2, without awns. Homalocenchnts. (60)

45. Flowering glumes chartaceous or coriaceous, differing from the membranous
empty glumes. 46

45. Flowering glumes thin and hyaline, usually much more delicate than the
empty glumes. 48

46. Margins of the fruiting flowering glumes hyaline and flat. Leptoloma. (54)

46. Margin of the fruiting lemma inrolled and not hyaline. 47

47. Sterile lemma awned or pointed; spikelets crowded in 1-sided racemes which

are arranged in a panicle. Echinochloa. (56)

47. Empty glumes and lemmas awnless; spikelets usually in panicles. Panicum (53)

48. Stem solid. 49

48. Stem hollow. 50

49. Sessile spikelets perfect the stalked spikelet vestigial, spikelets on the ends

of slender branches of a spreading panicle. Holciis. (62)

49. Spikelets all perfect; arranged in pairs along the continuous branches of the

panicle. Miscanthus. (64)

50. Lemma with a hooked dorsal awn, spikelets in a loose panicle ; plant softly

pubescent.^ _ Nothoholcus. (21)

50. Lemma with a prominent terminal awn ; spikelets in a slender panicle ; leaves

scabrous and glaucous, the sheaths smooth. Sorghastrum. (63)

—51—

51. Spikelets not covered with a prickly bur. 52

51. Spikelets covered with a prickly bur.

Cenchrus. (59)

270

OHIO BIOLOGICAL SURVEY

52. Inflorescence not moneciotis, altho the plants may be diecious. 52,

52. Monecions, the staminate and carpellate spikelets in dififerent parts of the
same inflorescence or on dififerent inflorescences ; stem solid. 72

52. Spikelets in a simple terminal spike. 54

53. Spikelets usually on one side of digitate or racemose branches of the in¬
florescence. 61

54. Spikelets solitary at each node of the rachis. 55.

54. Spikelets 2-several at each node of the rachis. 58.

55. Spikelets with their backs turned to the rachis. Lolium. (22)

55. Spikelets with their sides turned to the rachis. 56

56. Palet adherent to the grain; wild perennial grasses. Agropyron. (23)

56. Palet free from the grain ; ours cultivated or escaped annuals. 57

57. Spikelets usually 3-several-flowered. Triticum. (24)

57. Spikelets 2-flowered. Secalc. (25)

58. Spikelets 2-several-flowered. 59

58. Spikelets 1-flowered, sometimes with the vestige of a second flower, usually
in threes at the joints of the rachis, some of the spikelets often vestigial.

Hordeuin. (28)

59. Rachis of the spike jointed and readily breaking up at maturity. Sitanion.

59. Rachis not breaking up at maturity. 60

60. Spikes mostly dense; empty glumes a little smaller than the flowering glumes.

Elymus. (26)

60. Spikes very loose; empty glumes very small or none. Hystrix. (27)

61. Joints of the rachis or rachilla with prominent hairs; stem solid; spikelets
in pairs, 1 stalked and 1 sessile. 62

61. Joints of the rachis or rachilla not with hairs, at most only short pubescent. 63

62. Spikelets all perfect. Miscanthus. (64)

62. Sessile spikelets perfect, the pedicellate staminate or sterile and vestigial.

Andropogon. (65)

63. Spikelets oval, articulated below the empty glumes, compact in form. 64

63. Spikelets lanceolate, articulated above the empty glumes, except in Spartina ;
flowering glumes with awns or points. 68.

64. Inflorescence with prominently digitate branches or simple. 65

64. Inflorescence branches racemose. 66

65. Lemma with inrolled margins, not hyaline ; spikelets oblong to orbicular.

Paspalum. (57)

65. Lemma with hyaline margins, not inrolled ; spikelets lanceolate.

Syntherisma. (55)

66. Spikelets dorsally compressed; stems solid or with delicate pith.

Echinochloa. (56)

66. Spikelets laterally compressed. 67

67. Spikelets near the ends of long stalked branches of a spreading panicle.

Hornalocenchrus. (60)

67. Spikelets in two rows on the erect branches of the panicle, the branches

sessile. Beckmannia. (30)

68. Inflorescence with numerous short reflexed branches. Atheropogon. (33)

68. Inflorescence with few spreading or ascending branches or with numerous
digitate branches. 69

69. Inflorescence branches digitate. 70

69. Inflorescence branches racemose. 71.

70. Spikelets 1-flowered; plants extensively creeping. Capriola. (31)

70. Soikelets with 2-several perfect flowers, stems tufted. Eleusine. (32)

GRASSES OF OHIO

271

71. Spikelets strictly 1-flowered ; branches of the inflorescence in our species
usually numerous. Spartina. (29)

71. Spikelets with 1 perfect flower and 1 or more vestigial ones ; branches of the

inflorescence 4 or less. Bouteloua. (34)

72. Carpellate spikelets in pockets of the rachis ; inflorescence without husks or

hardened leaf-sheaths. Tripsacum. (66)

72. Carpellate spikelets covered with husks or with the thickened subtending
leaf sheath. 73

73, Carpellate spikelets covered by a hardened leaf sheath, forming a bead-like

fruit. ■ Coix. (67)

73. Carpellate spikelets in rows on a cob, usually covered with papery husks.

Zea. (68)

272

OHIO BIOLOGICAL SURVEY

Subfamily, Poacatae.

Tribe, Festuceae. Fescue-grass Tribe.

Grasses with laterally compressed, 2-several-floioered spikelets
jointed above the empty glumes and arranged in panicles; empty glumes
usually shorter than the dowering glumes; palets not owned or the awn
terminal and straight, rarely born just below the apex.

1. Bromus L. Chess, Brome-grass.

Annual or perennial grasses with terminal panicles. Spikelets
rather large, few-many-flowered; empty glumes unequal, 1-5-nerved,
acute ; flowering glumes longer than the empty glumes ; lemmas
rounded on the back or sometimes compressed-keeled, 5-9-nerved,
usually 2-toothed at the apex, generally awned ; palet 2-keeled ; stig¬
mas sessile, plumose, inserted below a hairy cushion-like appendage
at the top of the ovulary ; grain adherent to the palet.

1.

1.

2.

2.

3.

3.

4.

4.

5.

5.

6.
6.
7.

7.

8.
8.

9.

9.

10.

10.

11.

11.

12.

12.

Outer or lower empty glume 3-nerved, the inner or upper one 5-9-nerved. 2
Outer empty glume 1-nerved, the inner one 3-nerved. 8
Lemma distinctly awned. 3

Lemma awnless or only awn-pointed, nearly as broad as long; annuals.

B. brizaeformis.

Lemma pubescent on the back, with well developed hairs. 4
Lemma glabrous or minutely roughened on the back ; annuals. 5
Hairs of the lemma long and silky; perennials. B. kalmii.

Hairs of the lemma short; annuals. B. hordeaceus

Leaf sheaths glabrous or only slightly pubescent ; lemmas with strongly in-
rolled margins, the nerves not prominent. B. secalinus.

Leaf sheaths pubescent, often velvety; lemmas not with inrolled margins, the
nerves prominent. 6

Panicle contracted, usually not over 3 in. long, its branches erect or ascend¬
ing, ^2-1/4 in. long. B. hordeaceus leptostachys.

Panicle loose and spreading, usually 6-12 in. long, its lower branches usually
2-6 in. long. 7

Sheaths pubescent; panicle 8 in. or less long; spikelets broadly lanceolate,
usually 1 or 2 on the longer branchlets. B. racemosus.

Sheaths densely velvety-pubescent with reflexed hairs; panicle 8-12 in. long;
spikelets lanceolate, several on the longer branchlets. B. arvensis.

Awn shorter than the body of the lemma, or wanting ; rather tall perennials. 9
Awn longer than the body of the lemma ; low annuals with drooping pan¬
icles. 12

Lemma awnless or merely awn-pointed.

Lemma awned. 10

Leaf sheaths glabrous or softly pubescent. 11 ■

Leaf sheaths strongly retrorse-hirsute.

Lemma pubescent on the margins only.

Lemma evenly pubescent all over the back.

Awn less than ^ in. long; body of the lemma strigose; not over ^4 in. long.

B. tectorum.

Awn about 1 in. long or more ; body of the lamma sparsely hispidulous, Yi
in. long or more. B. sterilis.

B. inermis.

B. asper.
B. ciliatus.
B. purgans.

GRASSES OF OHIO

273

1. Bromus brizaeformis Fisch. & Mey. Awnless Chess. An
annual grass with stems 1-2 ft. high, erect, simple, the sheaths and
blades pubescent. Panicle open and drooping; spikelets broadly
ovate F2-I in. long, awnless; lemma very broad, obtuse, 9-nerved,
shining.

In fields and waste places. July, August. Introduced from
Europe. Cuyahoga County.

2. Bromus kalmii Gr. Kalm’s Chess. A perennial grass with
a slender stem, 1-3 ft. high, the sheaths and blades pubescent.
Panicle open 2-6 inches long, its branches usually flexuous ; spikelets
drooping on capillary peduncles, densely silky pubescent all over;
lemma 7-nerved, with a straight awn.

In dry ground, woods and thickets. June, July. Franklin, Lucas.

3. Bromus hordeaceus L. Soft Chess. An annual grass, the
whole plant more or less pubescent, 1-3 ft. high, erect, with a rather
contracted panicle with erect or ascending branches. Spikelets ap-
pressed-pubescent, on short pedicels ; lemma awned, 7-9 nerved,
awned between the obtuse or acute teeth. The var. leptostachys
(Perr.) Beck, has glabrous or slightly scabrous spikelets.

A weed in fields and waste places. July, August. Introduced,
Wayne, Lorain.

4. Bromus secalinus L. Common Chess. An annual grass
with an erect stem 1-3 ft. high, its sheaths smooth and strongly
nerved but the blades glabrous, rough, or sometimes hairy. Spikelets
glabrous, turgid on somewhat drooping branches of the open panicle;
lemma short-awned, becoming convex, thick and inrolled at the mar¬
gins.

Common in fields and waste places and often a pernicious weed
in grain fields. According to an old superstition wheat changes to
this chess which is, of course, absolutely impossible. Also called
cheat. Naturalized from Europe. General and abundant.

5. Bromus racemosus L. Upright Chess. An annual erect
grass, 1-3 ft. high, with pubescent sheaths and short erect panicles.
Spikelets glabrous ; lemma obtuse, awned, smooth and shining, the
nerves prominent.

In fields and waste places. June-August. Naturalized from
Europe. General and abundant.

6. Bromus arvensis L. Field Chess. An annual grass with
erect stems, 1-3 ft. high, pubescent sheaths, and large open panicles
with long drooping branches. Spikelets, lanceolate, somewhat shin¬
ing; lemma smooth or minutely scabrous, 5-nerved, broadest at the
middle, bearing an erect awn.

In fields and waste places. Summer. Introduced from Europe.
Franklin County.

274

OHIO BIOLOGICAL SURVEY

7. Bromus inermis Leyss. Hungarian Brome-grass. A peren¬
nial grass with tufted stems, 2-3^ ft. high, with smooth and glabrous
leaves and an oblong panicle of erect oblong spikelets. Lemma 5-7-
nerved, awnless or sometimes awn-pointed. An important grass for
pasture and hay.

In fields and waste places. June, July. Introduced from Europe.
Wayne County.

8. Bromus ciliatus L. Fringed Brome-grass. An erect peren¬
nial, 2-4 ft. high, with retrorsely pubescent or nearly smooth sheaths
and a broad, lax, drooping panicle whose branches are spikelet-bear-
ing near the ends. Spikelets 5-10-flowered ; lemma smooth on the
back, pubescent along the margins, 5-7-nerved, obtuse and slightly
bifid at the apex which ends in a straight awn. Very variable.

In moist woods and thickets. July, August. Erie, Wayne,
Eranklin, Champaign, Hocking.

9. Bromus purgans L. Hairy Brome-grass. A perennial grass
with erect rather stout stems, 2-5 ft. high, sparsely retrorse-pilose
sheaths, and large lax nodding panicles. Spikelets, 7-12-flowered ;
lemma lanceolate, acute, densely pubescent, 5-7-nerved, emarginate
and with a short straight awn.

In moist rocky woodlands. June- August. General.

10. Bromus asper Murr. Rough Brome-grass. A perennial
grass with simple erect rough stems, 2-6 ft. high, retrorse-hirsute
leaf-sheaths, and an open panicle with drooping branches. Spikelets
5-10-flowered; lemma acute, hispid near the margins and on the
lower part of the keel, short awned.

In waste places. July, August. Naturalized from Europe. No
specimens.

11. Bromus tectorum L. Downy Brome-grass. A short-lived

annual, ft. high, with slender tufted stems, pubescent sheaths

and leaf-blades, and broad, rather dense, one-sided drooping panicles.
Spikelets nodding, 5-8-flowered ; lemma long-awned, 7-nerved, usually
hirsute-pubescent. A weed.

In waste places and roadsides. May-July. Introduced from
Europe. General and abundant.

12. Bromus sterilis L. Barren Brome-grass. An erect annual
grass, 1-3 ft. high, with a broad lax drooping panicle, the slender
branches usually with but 1 spikelet, and with smooth or sometimes
pubescent sheaths. Spikelets spreading or pendulous; lemma
acuminate, long-awned, 7-nerved, scabrous on the nerves.

In waste places. June, July. Introduced from Europe. Licking,
Sandusky, Cuyahoga.

GRASSES OF OHIO

275

2. Melica L. Melic-grass.

Perennial grasses with simple stems, and rather large spikelets
in narrow panicles. Spikelets 2-several-fIowered, often secund ;
rachilla extended beyond the normal flowers and usually bearing
2-3 club-shaped or hooded glumes, convolute around each other ;
empty glumes large, unequal, membranous, 3-5-nerved ; lemmas
7-13-nerved, sometimes awned, the margins more or less scarious ;
palets broad, 2-keeled ; grain free.

1. Melica nitens Nutt. Tall Melic-grass. An erect simple
smooth and glabrous grass with hollow stems, 2-4 ft. high, and an
elongated open panicle with short spreading or ascending branches.
Spikelets numerous, usually more or less secund, nodding, on flex-
uous pedicels, usually with 3 perfect flowers ; lemmas scabrous, acute.

In rocky woods and open places. May, June. Erie County.

3. Festuca L. Fescue-grass.

Perennials or annuals with flat or convolute leaf-blades and ter¬
minal panicles. Spikelets 2-many-flowered ; empty glumes unequal,
narrow, acute, keeled, the outer 1 -nerved, the inner 3-nerved ; lemma
narrow, rounded on the back, 5-neryed, usually awned at the apex ;
palet usually about as long as the lemma ; styles very short, stigmas
plumose.

1. Leaves flat_ over in. wide; lemma awnless; perennial. 2

1. Leaves setaceous, usually involute or folded, less than in. wide. 4

2. Panicle narrow, with short erect branches; spikelets 5-10-flowered ; lemma

usually in. or more long. F. elatior.

2. Panicle with long, spreading or ascending branches ; spikelets 3-6-flowered ;
lemma less than y in. long. 3

3. Spikelets broadly obovate; panicle branches spikelet-bearing from or below

the middle. F. shortii.

3. Spikelets lanceolate; branches of the panicle elongated, the spikelets loosely

scattered at the ends. F. nutans.

4. Lemma awnless or short-awned ; perennial ; stamens 3. 5

4. Lemma long-awned ; annual ; stamen usually 1. 6

5. Lemma short-awned ; leaves setaceous. F. ovina.

5. Lemma awnless; leaves capillary. F. capillata.

6. Outer empty glume more than Yi as long as the inner ; awn not longer than

the body of the lemma. F . octoHora.

6. Outer empty glume less than as long as the inner; awn of the lemma longer
than its body. F. myuros.

1. Festuca elatior L. Tall Fescue-grass. A loosely tufted per¬
ennial, 2-5 ft. high, often with short creeping rhizomes. Spikelets
broadly obovate when mature, 3-6-flowered, on erect contracted
panicles whose branches are spikelet-bearing nearly to the base ;
lemma oblong-lanceolate, scabrous at the summit, faintly 5-nerved,
acute or rarely short-awned.

In meadows and waste places. Cultivated for hay. Also called

276

OHIO BIOLOGICAL SURVEY

meadow fescue-grass. June-August. Introduced from Europe.
General.

2. Festuca nutans Willd. Nodding Fescue-grass. A perennial
grass with simple, erect, slender stems, 2-3 ft. high, with glabrous
or pubescent leaf-sheaths and a very loose panicle whose branches
are secund and spikelet-bearing near the ends, at first erect and
finally spreading. Spikelets lanceolate, 3-5-flowered ; lemma oblong-
ovate, acute, very faintly nerved.

In moist woods and copses. June, July. General.

3. Festuca ovina L. Sheep Fescue-grass. A densely tufted
perennial 34-2 ft. high, with crowded sheaths at the base and pale
green filiform or setaceous blades. Panicle contracted after bloom¬
ing; spikelets with rather loose flowers; lemma smooth or slightly
scabrous, acute and short awned. Many varieties.

A good pasture grass. In fields and waste places. June, July.
From Europe. Erie, Franklin, Wayne.

4. Festuca capillata Lam. Filiform Fescue-grass. A densely
tufted perennial with slender, smooth and glaucous stems, p2-l34
ft. high, and with filiform leaf blades. Panicle contracted with erect
branches; spikelets 4-5-flowered ; lemma acute, unawned.

In fields and along roadsides. June, July. From Europe. Cuya¬
hoga County.

5. Festuca octoflora Walt. Slender Fescue-grass. A slender,
small, erect annual, often tufted, 34-1/4 ft. high, with sheaths shorter
than the internodes and narrowly linear usually involute blades.
Panicle narrow, erect, usually reduced to a more or less secund
raceme; spikelets 6-13-flowered ; lemma usually very scabrous,
acuminate into a straight awn ; stamens 2.

In dry sterile or sandy soil. June-August. Ashtabula, Erie,
Lucas, Ashland, Delaware, Licking, Lawrence.

6. Festuca myuros L. Rat-tail Fescue-grass. A smooth, glab¬
rous annual, 1-2 ft. high, with solitary or small tufted stems and
overlapping sheaths. Leaf-blades smooth, linear, involute, erect.
Panicle usually 1-sided, contracted, its branches appressed ; spikelets
3-6-flowered ; lemma linear-lanceolate, scabrous above, attenuate into
a slender awn; stamen 1.

In dry fields and waste places. June, July. From Europe. Lake
County.

4. Panicularia Fabr. Manna-grass.

Mostly tall, perennial, hydrophytic grasses with simple stems
and terminal panicles. Spikelets few-many-flowered, terete or some¬
what flattened; empty glumes unequal, 1-3-nerved; lemmas rounded

GRASSES OF OHIO

277

on the back with scarious margins or apex, 5-9-nerved ; palets with
the 2 keels near the margin ; stamens 2 or 3 ; stigmas plumose ;
grain smooth.

1. Spikelets linear or compressed cylindric, in. long or more. 2

1. Spikelets ovate or oblong, in. long or less. 3

2. Lemma acute, much shorter than the palet, about Yz in. long. P. aciitiUora.

2. Lemma obtuse, nearly as long as the palet, about Y in. long. P. septentrionalis.

3. Lemma very broad, obscurely nerved ; panicle open ; spikelets 5-12-flowered.

P. canadensis.

3. Lemma rather narrow, sharply and distinctly 7-nerved. 4

4, Panicle contracted, elongated and narrow; spikelets 3-4-flowered. P. torreyana.

4. Panicle open and lax, not elongated, its branches usually spreading or droop¬
ing. 5

5. Empty glumes obtuse or rounded at the apex. 6

5. Empty glumes truncate and denticulate at the apex ; inner empty glumes nearly

Y in. long; panicle rather narrow, with few spikelets, usually less than 6
in. long. P. pallida.

6. Spikelet Yz in, long or less ; branches of the panicle often drooping ; inner

empty glume about ^ in. long. P. nervata.

6. Spikelet nearly Y in. long; branches of the panicle ascending or spreading;
inner empty glume nearly in. long. P. grandis.

1. Panicularia acutiflora (Torr.) Ktz. Sharp-glumed Manna-
grass. A grass 1-2 ft. high, with flattened weak and slender stems,
and with overlapping sheaths, the uppermost inclosing the base of
the slender panicle. Spikelets linear, 5-12-flowered, erect on the
panicle, lemma acute, scabrous, much exceeded by the long-acuminate
palet.

In wet soil and shallow water. June-August. No specimens.

2. Panicularia septentrionMis (Hlitch.) Bickn. Floating Manna-
grass. An erect grass with somewhat flattened stems, 2-3 ft. high,
from a creeping base and with a very slender panicle of a few re¬
mote branches which are at first appressed or finally horizontal.
Spikelets 7-12-flowered, compressed-cylindric, subsessile ; lemma
hispidulous, 7-nerved, obtuse, somewhat exceeded by the acuminate
palet.

In shallow water. June, July. General as far south as Har¬
rison, Perry, Franklin and Auglaize Counties.

3. Panicularia canadensis (Mx.) Ktz. Rattlesnake Manna-
grass. A grass with erect simple stems, 2-3 ft. high, with com¬
pressed sheaths, and with very loose open panicles, the remote ca¬
pillary branches drooping. Spikelets 5-12-flowered, ovate, tumid;
lemma broad, obtuse or abruptly acute, obscurely 7-nerved.

In swamps and wet places. July, August. Ashtabula, Geauga,
Portage, Cuyahoga, Lorain, Summit, Stark, Wayne.

4. Panicularia torreyana fSpreng.) Merr. Long Manna-grass.
A grass with solitary or few erect stems, 1-3 ft. high, from a run-

278

OHIO BIOLOGICAL SURVEY

nin,^- rhizome. Panicle dense, contracted, with erect branches;
s])ikelets v3-7-rtowered, appressed ; lemma narrow, acute, 7-nerved,

In wet woods. August, September. Ashtabula, Cuyahoga, Sum¬
mit, Erie, Fairfield.

5. Panicularia nervata (Willd.) Ktz. Nerved Manna-grass. A
grass with erect, sim])le, slender stems, l-v3 ft. high, often in large
clumi)s, with an expanded nodding panicle, the capillary branches
drooping. Spikelets 3-7-tiowered, purplish; lemma obtuse or rounded,
with 7 shar]), distinct nerves.

In wet places. June-August. General.

6. Panicularia grandis (Wats.) Nash. Tall Manna-grass. A
grass with stout erect clustered stems, 3-5 ft. high, and a very com¬
pound loose and open ])anicle, nodding at the summit. Spikelets
4-7-dowered with purple flowering glumes and whitish empty glumes ;
lemma 7-nerved, obtuse or rounded at the apex.

In wet soil and ditches. June-August. Stark, Wayne.

7. Panicularia pallida (Torr.j Ktz. Pale Manna-grass. A grass
with pale green slender stems, 1-3 ft. high, ascending from a creeping
base. Panicle lax, few-flowered ; spikelets 4-8-flowered, pale green ;
lemma 7-nerved, truncate and denticulate at the apex.

In shallow water. Jnly, August. Trumbull County. (Ottawa
County — Moseley Herbarium)

5. Poa L. Blue-grass, Meadow-grass, Spear-grass.

Perennial or annual grasses with flat or convolute leaves ending
in a cucullate tip, and with terminal panicles. Spikelets 2-6-flowered,
compressed; empty glumes keeled, 1-3-nerved; flowering glumes
longer than the empty ones, commonly with a tuft of cobwebby
hairs at the base, 5-nerved ; palets 2-nerved or 2-keeled, 2-toothed
at the apex; styles short, stigmas plumose; grain free or sometimes
adherent to the palet.

1. Flowering glumes not with webby hairs at the base, naked or with short
hairs. 2

1. Flowering glumes with webby hairs at the base; stems tufted. 4

2. Stems much flattened, not tufted; rhizome long and creeping; lemma ob¬
scurely 3-nerved. P. cornpressa.

2. Stems little or not at all flattened, tufted; lemma 5-nerved. 3

3. Lemma with the midnerve silky-pubescent for ^ its length ; perennials

P. antumnaUs.

3. Lemma with the midnerve pilose below ; annuals. P. annua.

4. Lemma glabrous; stems distinctly compressed; perennial. P. debilis.

4. Lemma somewhat pubescent above the basal webby hairs; stems round or
only slightly compressed. 5

5. Lateral nerves of the lemma glabrous. 0

5. Lateral nerves of the lemma pubescent. 7

6. Plant yellowish green ; lemma less than % in. long.

6. Plant green ; lemma more than in. long.

P. trivia Its.
P. alsodes.

GRASSES OF OHIO

279

7. Panicle-branches dividing and spikelet-bearing at or below the middle ; basal
leaves much shorter than the stem. 8

7. Panicle-branches usually dividing and spikelet-bearing only at the ends ; basal
leaves very long, the early ones often as long as the stem. P. hrachyphylla.

8. Intermediate nerves of the lemma obscure. 9

8. Intermediate nerves of the lemma prominent. 10

9. Panicle erect, rarely over 5 in. long, its branches ascending; empty glumes

narrow. P. nernoralis.

9. Panicle drooping, often 1 foot or more long, its branches spreading.

P. triflora.

10. Midnerve of the lemma pubescent only below ; spikelets crowded on the
branches. _ P. prat crisis.

10. Midnerve of the lemma pubescent its whole length; spikelets scattered on
the spreading, often reflexed branches. P. sylvestris.

1. Poa compressa L. Flat-stemmed Blue-grass. A perennial
grass of a pale bluish green color with wiry flattened stems, de¬
cumbent at the base, ^-2 ft. high, from long horizontal rhizomes.
Panicles narrow with erect or ascending branches, spikelet-bearing
to the base. Spikelets 3-9-flowered ; lemma obscurely 3-nerved, the
nerves sparingly pubescent toward the base, more or less bronzed
at the summit.

A' grass of some importance in dry mostly sterile soil, in culti¬
vated fields and in woods. Also called English blue-grass. J\Iay-
September. From Europe. General and abundant.

2. Poa trivialis L. Rough-stalked Meadow-grass. A perennial
grass with erect stems from a somewhat decumbent base, 1-3 ft.
high, scabrous below the panicle. Panicle open, its branches usually
spreading or ascending; leaf-sheaths and blades retrorsely scabrous;
spikelets 2-3-flowered ; lemma webby at the base, strongly 5-nerved,
the midnerve silky-pubescent below, the lateral nerves naked.

An important grass in meadows, roadsides, and waste places.
May-August. From Europe. Crawford County.

3. Poa debilis Torr. Weak Spear-grass. A perennial grass
with erect, weak, slender, nearly terete stems, 1-2^ ft. high, with
compressed sheaths much shorter than the internodes, and with an
open nodding panicle, the few, long, capillary branches ascending
or spreading at the ends. Spikelets 2-4-flowered ; lemma obtuse,
glabrous except the webby base.

In rocky woodlands. May-July. Trumbull County. (Erie
County — Moseley Herbarium.)

4. Poa triflora Gilib. Fowl Meadow-grass. A perennial grass
with erect stems, 1-5 ft. high, and with a pyramidal or oblong panicle,
often purplish, its filiform branches spreading. Spikelets 3-4-flowered ;
lemma obtuse with copious webby hairs, its intermediate nerves ob¬
scure, midnerve and marginal nerves pubescent on the lower half.

In wet meadows and swampy places. Also called false red-top.
July-August. Fairfield, Geauga, Lawrence.

280

OHIO BIOLOGICAL SURVEY

5. Poa nemoralis L. Wood Meadow-grass. A perennial grass
with slender leafy stems, 1-2 ft. high, and an open spreading panicle.
Spikelets 2-5-flowered ; lemma with obscure intermediate nerves,
with a few webby hairs at the base.

^Meadows and open woods. June-September. Introduced from
Europe. Lake County.

6. Poa pratensis L. Kentucky Blue-grass. A perennial grass,
sending out numerous running rhizomes from the base, with simple
erect stems, 1-4 ft. high, with compressed sheaths, and with a pyr¬
amidal panicle, the spreading or ascending slender branches divided
and spikelet-bearing above the middle. Spikelets 3-5-flowered,
crowded ; lemma conspicuously webbed at the base, 5-nerved, the
marginal nerve and midnerve pubescent below, the intermediate ones
naked.

A very important grass extensively used for pastures and lawns
and to some extent for hay. In fields, meadows, and woods. May-
August. General and abundant.

7. Poa autumnalis ]Muhl. Flexuous Spear-grass. A perennial
grass with erect slender stems, 1-3 ft. high, and a panicle with long,
capillary, flexuous, spreading branches bearing a few spikelets near
the ends. Spikelets 3-6-flowered, lemma not webby at the base but
pubescent below between the strong nerves, the midnerve silky
pubescent for three-fourths its length.

In woods. March-May. Hocking County.

8. Poa sylvestris Gr. Sylvan Spear-grass. A perennial grass
with simple, slender, erect, slightly flattened stems, 1-3 ft. high, and
oblong-pyramidal panicles with spreading ascending or reflexed
branches spikelet-bearing at the extremities. Spikelets 2-4-flowered ;
lemma webbed at the base, pubescent below, 5-nerved, the midnerve
pubescent nearly its entire length, the marginal nerves pubescent
below the middle.

In meadows, woods, and thickets. May-July. Rather general;
no specimens from the northwestern counties.

9. Poa alsodes Gr. Grove ^leadow-grass. A perennial grass
with simple, erect, slender stems, ^-2^ ft. high, with long sheaths,
the uppermost often sheathing the base of the panicle, and with a
panicle of spreading or ascending branches, spikelet-bearing at the
ends. Spikelets 2-3-flowered ; lemma webbed at the base, faintly
nerved, the midnerve pubescent below.

Wooded hillsides and thickets. ^lay, June. Seneca, Franklin,
Summit, Cuyahoga, Trumbull, Knox.

10. Poa brachyphylla Schult. Short-leaf Spear-grass. A per¬
ennial grass with stems 1-3 ft. high from running rhizomes, with

GRASSES OF OHIO

281

basal leaves often about equalling- the stems and abruptly cuspidate-
tipped, and with an open panicle whose ascending, spreading or
reflexed branches are spikelet-bearing at the ends. Spikelets 3-5-
flowered ; lemma webby at the base, the keel and marginal nerves
sparingly pubescent, the intermediate nerves prominent and naked.

In rocky woodlands. April-June. Lawrence, Perry, Medina,
Cuyahoga, Trumbull.

11. Poa annua L. Annual Meadow-grass. An annual grass
with flattened stems, ft. high, decumbent at the base, with

loose sheaths, very soft leaves, and a pyramidal panicle with open
spreading branches. Spikelets 3-6-flowered, crowded ; lemma dis¬
tinctly 5-nerved, the nerves hairy below.

Cultivated and waste ground. April-October. From Europe.
General.

6. Dactylis L. Orchard-grass.

Tall perennial grasses with glomerate panicles. Spikelets 2-5-
flowered, compressed, nearly sessile, in dense capitate clusters ; empty
glumes thin-membranous, hispid-ciliate on the keel, acute or mucron-
ate ; lemmas 5-nerved, keeled, the midnerve extended into a point or
short awn; palets 2-keeled ; grain free, enclosed in the lemma and
palet.

1. Dactylis glomerata L. Orchard-grass. A coarse tufted
glaucous grass with simple erect stems, 2-4 ft. high, and a panicle
with a few stiff contracted branches, naked below and bearing dense
one-sided clusters of spikelets at the ends. Spikelets 3-5-flowered ;
lemma 5-nerved, short-awned, rough, ciliate on the keel.

An important hay and pasture grass cultivated to some extent.
In fields and waste places. June, July. Naturalized from Europe.
General ; no specimens from the southeastern counties.

7. Eragrostis Beauv. Love-grass.

Annual or perennial grasses with the spikelets in loose or dense
terminal panicles. Spikelets strongly flattened ; 3-many-flowered ;
empty glumes unequal, shorter than the flowering ones, keeled, 1-
nerved or the inner 3-nerved ; lemma membranous, keeled, 3-nerved ;
palet shorter, prominently 2-nerved or 2-keeled ; grain free, loosely
enclosed in the flowering glumes.

1. Spikelets 5-35-flowered ; in. long. 2

1. Spikelets 2-5-flowered ; in. or less long; annuals. 5

2. Stems erect or ascending, simple, rigid. 1-214 ft. tall ; flowers bisporangiate ;

perennials. E. pectinacca.

282

OHIO BIOLOGICAL SURVEY

2. Stems extensively creeping; plants diecious, annual. E. hypnoides.

2, Stems usually decumbent at the base and generally much branched ; flowers
bisporangiate ; annuals. 3

3. Spikelets nearly or quite ]4, in. wide; plants with very strong odor. E. major.

3. Spikelets tV in. wide or less. 4

4. Lemmas usually dull-purple or green, the lateral nerves very prominent;

spikelets about tV in. wide; outer empty glume about ^ as long as the
inner. E. purshii

4. Lemmas usually bright purplish, the lateral nerves faint or wanting; spikelets
less than tV in. wide; outer empty glume ^2 as long as the inner. E.pilosa.

5. Pedicels and branches of the panicle short; stems branched above the base.

E. frankii.

5. Pedicels and branches of the panicle long and capillary; stems branched only
at the base. E. capillaris.

1. Eragrostis pectinacea (jMx.) Steud. Purple Love-grass. An
erect or ascending, perennial grass with a rigid stem, 1-2^ ft. high,
from a short stout rhizome, with over-lapping sheaths, sparingly
villous and densely bearded at the throat. Panicle purple or purplish,
included at the base at least at hrst, its widely spreading or redexed
branches strongly bearded at the base; spikelets 5-12-tIowered, on
stiff pedicels; lemma acute, minutely scabrous, its lateral nerves
prominent.

In dry sandy soil. July-September. Lake, Cuyahoga, Erie,
Auglaize.

2. Eragrostis hypnoides (Lam.) B. S. P. Creeping Love-grass.

An annual grass with an extensively creeping stem sending up
panicle-bearing branches, ft. high. Panicles nearly simple,

sometimes nearly capitate. Spikelets 10-35-flowered,. 1:he flowers
bisporangiate, staminate, or carpellate ; lemma acuminate its lateral
nerves prominent.

In sandy or gravelly soil along shores and ditches. August, Sep¬
tember. Rather general ; no specimens from the central eastern
counties.

3. Eragrostis major Host. Strong-scented Love-grass. A beau¬
tiful erect or ascending or occasionally prostrate grass with rather
flaccid, freely branching stems, ^-4 ft. high or long, with a strong
scent, and with greenish-lead-colored, rather densely flowered
panicles. Spikelets 10-40-flowered, the flowers closely imbricated;
pedicels and keels of the empty glumes sparingly glandular; lemma
thin, obtuse, scabrous, the lateral nerves prominent.

A common weed in cultivated fields and waste places. Also
called stink-grass. July-September. Naturalized from Europe.
General.

4. Eragrostis purshii Schrad. Pursh’s Love-grass. An annual

tufted grass, usually decumbent at the base and much branched,
the stems ft. long. Panicle open, its branches spreading.

GRASSES OF OHIO

28

>

naked in the axils. Spikelets 5-1 S-dowered, dull purple or green;
lemma firm, acute, the lateral nerves prominent.

In dry soil. August, September. General.

5. Eragrostis pilosa (L.) Beauv. Tufted Love-gras^. An an¬
nual grass with slender, tufted, diffusely branching stems, ^4-1^2 ft-
high, with a decumbent or spreading base, and with a diffuse panicle
whose lower axils are usually sparingly bearded. Spikelets 5-18-
fiowered, becoming linear; lemma subacute, usually purplish, the
lateral nerves faint.

In waste places or cultivated ground. August, September.
Rather general ; no specimens from the northwestern counties.
Naturalized.

6. Eragrostis frankii Steud. Frank’s Love-grass. An annual,
diffusely branched grass with tufted erect or decumbent stems, 14-134
ft. high and an oblong open panicle with short spreading branches.
Spikelets ovate, 3-5-fiowered, on more or less appressed pedicels;
lemma very acute ; faintly 3-nerved.

In low or sandy places. August-October. Rather general ; no
specimens from the south-eastern nor from the north-western
counties.

7. Eragrostis capillaris (L.) Nees. Capillary Love-grass. A
slender, erect, sparingly branched annual grass, 34-1/4 ft. high with
panicles more than half the entire length of the plant, oblong-ovoid
in shape, the capillary branches spreading or ascending. Spikelets
ovate, 2-4-fiowered, on long, divergent pedicels ; lemma acute, faintly
3-nerved. Often lemon-scented.

Considered a weed. In dry, sandy soil. August, September.
Ottawa, Madison, Clinton.

8. Sphenopholis Scrib. Eaton-grass.

Slender tufted perennials, usually with narrow terminal panicles.
Spikelets 2-3-fiowered, the rachilla articulated between and extended
beyond the fiowers, the pedicels jointed just below the glumes; outer
empty glume linear, acute, 1 -nerved, the inner much broader, 3-
nerved, usually obtuse or rounded at the apex, the margins scarious ;
lemma chartaceous, with obscure nerves ; palet narrow, 2-nerved ;
grain free, enclosed in the rigid lemma.

1. Empty glumes unequal, the outer shorter and about 34 as wide as the inner. 2

1. Empty glumes nearly equal the outer not less than 34 as wide as the inner;

Branches of the lax panicle more or less spreading. y. nitida.

2. Inner empty glume obovate ; panicle narrow, densely flowered. y. obtusota.

2. Inner empty glume narrowly obovate to oblanceolate ; branches of the lax

panicle more or less spreading. S. pollens.

284

OHIO BIOLOGICAL SURVEY

1. Sphenopholis obtusata (Mx.) Scrib. Blunt-glumed Eaton-
grass. An erect rather stout grass, 1-2^2 ft. high with a dense gen¬
erally spike-like, strict panicle with erect branches. Spikelets 2-3-
flowered ; lemma narrow, obtuse ; empty glumes unequal, often
purplish, the first narrow, shorter than the inner which is sub-
cucullate and about six times as wide.

In dry soil. June-August. No specimens.

2. Sphenopholis pallens (Spreng.) Scrib. Tall Eaton-grass. An
erect, usually slender grass with simple stems, 1-3 ft. high, and
lax, nodding panicles. Spikelets oblong-lanceolate, usually numerous
and somewhat crowded and appressed to the branches ; lemma
lanceolate, acute, rarely awned ; empty glumes unequal, the outer
narrow and about % as broad as the obtuse or abruptly acute inner
one, which is smooth or somewhat rough on the keel.

In woods or moist soil. June, July. General.

3. Sphenopholis mtida (Spreng.) Scrib. Slender Eaton-grass.
A grass with very slender erect stems, 1-2 ft. high, with pubescent
sheaths and leaves, and with a loosely flowered panicle, the branches
spreading at flowering time and later becoming erect. Spikelets
cuneiform, not crowded ; empty glumes smooth, the outer about
one-third as wide as the inner; lemma narrow, smooth, obtuse, rarely
short-awned.

In dry woods. May, June. Cuyahoga, Knox, Licking, Fairfield,
Hocking, Lawrence, Adams.

9. Koeleria Pers. Koeler-grass.

Tufted perennials with narrow leaves and mostly densely
flowered terminal spike-like panicles. Spikelets 2-4-flowered ; rachilla
prolonged into a naked pedicel behind the upper palet; empty glumes
narrow, acute, unequal, keeled, with scarious margins ; lemma faintly
3-5-nerved ; palet 2-keeled ; grain free, loosely inclosed within the
subrigid lemma.

1. Koeleria cristata (L.) Pers. Crested Koeler-grass. A per¬
ennial grass with simple rigid erect stems, l-2j^ ft. high, with
retrorsely pubescent sheaths, and with pale green, spike-like, cylin¬
drical panicles. Spikelet 2-5-flowered ; lemma scabrous^ shining.

A valuable pasture grass in the west. In dry sandy soil, es¬
pecially on prairies. July-September. Ottawa County.

10. Korycarpus Zea. Korycarpus.

Erect, nearly smooth parennials with narrow paniculate or
racemose inflorescence. Spikelets 3-5-flowered, the rachilla articu¬
lated between the flowers ; empty glumes much shorter than the

GRASSES OF OHIO

285

flowering glumes ; lemma broad, coriaceous and shining, 3-nerved ;
palet firm, 2-keeled ; stamens 2, rarely 1 ; grain beaked, free.

1. Korycarpus arundinaceus Zea. American Korycarpus. A
grass with simple erect stems, very rough below the panicle, 1^-4
ft. high, and with very simple panicles. Spikelets short-pediceled,

3- 5-flowered, erect ; lemma somewhat abruptly acuminate.

In rich woods and shaded river banks. July-September. Ross,
Franklin, Auglaize, Highland.

11. Tridens R. & S. Purple-top.

Perennial grasses with long leaves and terminal open or con¬
tracted panicles. Spikelets 3-many-flowered ; empty glumes un¬
equal, keeled, shorter than the spikelet; lemma 3-nerved, bidentate,
the nerves silky-villous below ; palet broad, compressed, 2-keeled.

1. Tridens flava (L.) Hitch. Tall Purple-top. A grass with
erect stems, viscid on the axis of the panicle and below it, 2-5 ft.
high, with the leaf-sheaths bearded at the summit, and with showy
loose and open panicles with slender spreading branches. Spikelets

4- 8-flowered, purple, on long pedicels; lemma oval, the three nerves
pilose and excurrent as short points. Many small insects are caught
by the vicid panicle.

In dry and especially sandy fields and waste places. July-Sep¬
tember. Rather general ; no specimens from the northwestern
counties nor the extreme eastern part.

12. Tnplasis Beauv. Sand-grass.

Perennials with small, nearly simple, contracted or open panicles.
Spikelets 3-6-flowered, rachilla articulated between the remote
flowers ; empty glumes keeled ; lemma short-awned, 2-cleft, with
3 strongly ciliate nerves ; palet 2-keeled, the keels long ciliate ; grain
free.

1. Triplasis purpurea (Walt.) Chapm. Purple Sand-grass. A
tufted grass with widely spreading or ascending stems, 1-3 ft. long,
with short internodes, short convolute leaf blades, the uppermost
minute, and with a small terminal panicle composed of a few rigid
branches which are finally divergent. Smaller panicles produced
later in the season at the nodes of the stem. Spikelets 2-5-flowered,
usually rose-purple ; lemma with a short awn scarcely exceeding its
truncate lobes ; palet with abundant long hairs on the two nerves.
Plant acid to the taste.

In sandy soil, especially on sea beaches. August, September.
Ashtabula, Cuyahoga, Erie.

286

OHIO BIOLOGICAL SURVEY

13. Cynosurus L. Dogtail-grass.

Annual or perennial tufted grasses with dense, erect, spike-like
panicles. Spikelets of 2 kinds, in small clusters ; the lower spikelets
of the clusters consisting of narrow glumes and continuous rachilla,
the terminal spikelets of broader glumes and articulated rachilla and
containing perfect flowers ; empty glumes of the fertile spikelets
1-nerved; lemma 1-3-nerved, pointed or short-awned ; glumes of the
sterile spikelets spreading, 1-nerved; grain finally adherent to the
palet.

1. Cynosurus cristatus L. Dogtail-grass. An erect slender
perennial grass with simple stems, 1-2^ ft. high, with narrow leaves,
and dense spike-like long exserted panicles. Spikelets of two kinds,
in small clusters, the upper with bisporangiate flowers the lower
with sterile vestigial flowers ; lemma of the perfect flower pointed
or short-awned ; lemma of the sterile flowers very narrow, pointed,
and strongly scabrous on the keel.

In fields and waste places. June-August. From Europe. Ma¬
honing County.

14. Phragmites Trin. Reed-grass.

Tall reed-like perennial grasses with stout, hollow, leafy stems,
broad leaf blades, and large terminal panicles. Spikelets 3-7-flowered,
rachilla articulated, with long silky hairs; empty glumes unequal,
lanceolate, acute ; lowest lemma empty or subtending a staminate
flower; lemmas of the perfect flowers glabrous, narrow, long-acumin¬
ate, much exceeding the short palet ; grain free.

1. Phragmites phragmites (L.) Karst. Reed-grass. A tall
reed-like perennial grass with stout, hollow, leafy stems, 5-15 ft. high,
from long creeping rhizomes, with overlapping sheaths, and with a
large, tawney, rather densely flowered panicle whose branches are
ascending. Spikelets 3-7-flowered, the flowers shorter than the long
hairs of the rachilla ; palet 3-nerved, long-acuminate. Can be used
for thatching.

In swamps and wet places. August-October. Ashtabula, Cuya¬
hoga, Erie, Lucas, Huron, Wayne, Franklin.

Pampas-grass (Cortaderia argentea) is a related perennial grass, 6-10 ft.
high with a very large, dense, silky panicle, cultivated for ornament. It might
perhaps be successfully planted in protected places in southern Ohio.

GRASSES OF OHIO

287

Tribe, Aveneae. Oats Tribe.

Grasses zvith laterally eompressed, 2-several-flowered spikelets,
usually jointed above the empty glumes and arranged in panieles ; empty
glumes usually longer than the dovuering glumes; one or more of the
palets usually azvned on the baek or from betzveen the 2-toothed apex.

15. Danthonia DC. Wild-oat-grass.

Mostly erect perennials with narrow leaves and small terminal
contracted or open panicles. Spikelets 3-many-tlowered, the upper¬
most staminate or vestigial ; rachilla pubescent, extending beyond
the dowers ; empty glumes keeled, acute, subequal, much longer than
the lemmas, usually exceeding the uppermost dower ; lemmas
rounded on the back, 2-toothed, with a dat, twisted and bent awn
between the teeth ; palet hyaline, 2-keeled ; grain free.

1. Teeth of the lemma triangular, not awn-pointed; leaves of the stem short.

D. spicata.

1. Teeth of the lemma aristate; leaves of the stem elongated. D. compressa.

1. Danthonia spicata (L.) Beauv. Common Wild-oat-grass. A
grass with terete stems, 1-2^ ft. high, with involute leaves, and with
a few-dowered panicle, often reduced to a raceme, the few short
branches erect or ascending. Spikelets 5-8-dowered, on short stiff
pedicels ; lemma sparsely clothed with stiff hairs, broadly oblong,
its teeth triangular, the bent awn longer than the body of the lemma.

In dry or rocky soil. June- August. General.

2. Danthonia compressa Aust. Flattened Wild-oat-grass. A
grass with simple, erect, slender, dattened stems, often decumbent
at the base, lJ^-3 ft. high, and an open panicle the lower branches
generally spreading. Spikelets 5-10-dowered ; lemma oblong, with
aristate teeth, a ring of short hairs at the base, and an erect or some¬
what bent and twisted awn.

In dry woods. July-September. Portage County.

16. Arrhenatherum Beauv. Oat-grass.

Tall perennials with dat leaf-blades and long narrow panicles.
Spikelets 2-dowered, the lower staminate, the upper bisporangiate,
rachilla extending beyond the dowers ; empty glumes thin-mem¬
branous, keeled, acute, unequal ; lemmas rigid, 5-7-nerved, the first
bearing a long bent, twisted, dorsal awn arising below the middle,
the second unawned ; palet hyaline, 2-keeled ; grain free, ovoid.

1. Arrhenatherum elatius (L.) Beauv. Oat-grass. A grass
with a simple erect stem, 2-4 ft. high, and a narrow, shining, pale or

288

OHIO BIOLOGICAL SURVEY

purplish panicle, the short branches verticillate and usually spikelet-
bearing from the base. Lemma scabrous, the awn of the staminate
flower about twice the length of its body.

A grass cultivated to some extent for hay. In fields and waste
places. June, July. From Europe. Hamilton County.

17. Trisetum Pers. False-oats.

Tufted perennial grasses with spike-like or open panicles.
Spikelets 2-5-flowered, rachilla prolonged beyond the flowers ; empty
glumes unequal, keeled; lemma 2-toothed, keeled, bearing a dorsal
awn; palet narrow, 2-toothed; grain free, enclosed in the flowering
glumes.

1. Trisetum pennsylvanicum (L.) Beauv. Marsh False-oats.
A grass with an erect, slender, often weak stem, 1-3 ft. high, and
with a narrow or sometimes loose and nodding yellowish panicle,
the branches ascending. Spikelets 2-flowered ; lemma of the lower
flower with a small awn; lemma of the upper flower with a long,
bent and twisted awn.

In swamps and wet meadows. June, July. No specimens.

18. Avena L. Oats.

Annual or perennial grasses with terminal panicles of large
spikelets. Spikelets 2-many-flowered, rachilla bearded below the
flowers ; empty glumes subequal, membranous, many-nerved ; lem¬
mas indurated toward the base, generally bearing a dorsal awn ;
palet narrow, 2-toothed ; grain often adherent to the flowering
glumes.

1. Empty glumes much shorter than the spikelets ; flowers with a circle of short

bristles around the base; perennials. A. torreyi.

1. Empty glumes as long or longer than the spikelets exclusive of the awns;

annuals. 2

2. Lemmas pubescent with long bristles; awns long and stiff. A. fatiia.

2. Lemmas glabrous; awn comparatively short and weak, or absent. A. sativa.

1. Avena torreyi Nash. Purple Oats. A perennial grass with
an erect simple slender stem, 1-2 ft. high, and a lax panicle, the
branches erect or ascending, naked below. Spikelets 3-6-flowered ;
lemma scabrous, strongly nerved, with a ring of short hairs at the
base, and with an awn as long as its body or longer.

In woods. July, August. Franklin County.

2. Avena sativa L. Common Oats. An annual glabrous grass
with a hollow stem, 1-4 ft. high, and a loose ample panicle with as¬
cending branches. Spikelets about 1 inch long; lemma glabrous,
awnless or with a small straight awn slightly twisted at the base.

GRASSES OF OHIO

289

Extensively cultivated for its grain and straw. Persisting in
fields and along roads and railways. June, July. Rather general.
Escaped from cultivation.

3. Avena fatua L. Wild Oats. An annual grass with a stout
simple erect stem, 1-4 ft. high, and an open panicle with ascending
branches. Spikelets pendulous ; empty glumes smooth, lemma
pubescent with long rigid brown hairs and a ring of hairs at the
base, its long stiff awn inserted about the middle and bent and
twisted. A weed.

In fields and waste places. July, xAugust. From Europe. No
specimens.

19. Deschampsia Beauv. Hair-grass. |

Tufted perennial grasses with flat or involute leaves and con¬
tracted or open panicles of shining spikelets. Spikelets 2-flowered,
the hairy rachilla extended beyond the flowers or rarely ending in
a staminate one ; empty glumes keeled, acute, membraneous ; lemmas
thin, 4-nerved, the midnerve becoming an awn, toothed at the apex ;
palet narrow, 2-nerved ; grain free enclosed in the flowering glumes,

1. Deschampsia flexuosa (L.) Trin. Wavy Hair-grass. A
glabrous grass with an erect slender simple stem, 1-234 ft. high^
sheaths much shorter than the internodes, involute setaceous leaves,
and an open panicle with erect, ascending or widely spreading, flex-
uous branches naked at the base. Lemma acutely toothed at the
apex and with a twisted bent awn inserted near its base.

In dry places. July, xAugust. Portage County.

20. Aspris xAdans. Hair-grass.

Small, delicate annuals with narrow leaf-blades and contracted
or open panicles. Spikelets small, 2-fIowered, both flowers perfect;
empty glumes thin-membranous, subequal, acute ; lemmas hyaline,
2-toothed, bearing a delicate dorsal awn arising below the middle;
palet hyaline, 2-nerved ; grain enclosed in the flowering glumes and
usually adherent to them.

1. Aspris caryophyllea (L.) Nash. Silvery Hair-grass. A
delicate glabrous annual grass with erect stems, ^4-1 ft. high, in¬
volute setaceous leaves, and an open, silvery, shining panicle.
Spikelets clustered toward the ends of the capillary branches ; lem¬
ma with a bent awn, very acute, 2-toothed.

In fields and waste places. May-July. From Europe. Lake
County.

290

OHIO BIOLOGICAL SURVEY

21. Nothoholcus Nash. Velvet-grass.

Perennial grasses with densely flowered terminal panicles.
Spikelets 2-flowered articulated below the empty glumes, the lower
flower perfect, the upper staminate ; empty glumes membranous,
keeled, the outer 1-nerved, the inner 3-nerved and often short-awned ;
lemmas chartaceous, that of the upper flower bearing a bent awn ;
palet narrow, 2-keeled, grain free, enclosed in the glumes.

1. Nothoholcus lanatus (L.) Nash. Velvet-grass. A light
green, perennial, densely and softly pubescent grass with erect sim¬
ple stems, often decumbent at the base, 1-3 ft. high, and a narrow
purplish panicle. Spikelets nearly as broad as long; lemmas ciliate
at the apex, that of the second flower with a hook-like awn.

In flelds, meadows and waste places. June- August. Lake,

Trumbull, Cuyahoga, Lorain, Erie, V ayne, Fairfield. From Europe.

GRASSES OF OHIO

291

Tribe, Hordeae. Barley Tribe.

Grasses ivitli laterally eoinpressed, 2-several-floiaered spikelefs
jointed above the empty glumes and arranged on opposite sides of a
jointed raehis forming equilateral spikes ; spikelets sometimes zvitli but i
perfeet Hozver and a mere vestige of the second ; leaf -blades zvitli a pair
of more or less zvell-developed appendages at the base.

22. Lolium L. Darnel.

Annual or perennial grasses with simple erect stems, flat leaves,
and terminal spikes. Spikelets several-flowered, solitary, with the
edge turned toward the raehis ; empty glumes rigid ; lemmas rounded
on the back, 5-7-nerved, awned or awnless; palet 2-keeled ; grain
adherent to the palet.

1. Outer empty glume shorter than the spikelet ; short-lived perennials. 2

1. Outer empty glume as long or longer than the spikelet ; lemmas awned or

awnless; annuals. L. teniulentum.

2. Lemmas awnless. L. perenne.

2. Lemmas, at least the upper ones, awned. L. miiltiflorum.

1. Lolium perenne L. Red Darnel. A short-lived perennial
grass with an erect, simple stem, ^-2;^ ft. high, and a flat spike.
Spikelets 5-10-flowered ; lemma obscurely nerved, acuminate, awn¬
less. Cultivated as a pasture grass and also used for hay.

In cultivated fields and roadsides. Also called ray-grass and
rye-grass. June-August. Rather general. From Europe.

2. Lolium multiflorum Lam. Awned Darnel. A short-lived
perennial grass with tufted stems, roughened below and in the in¬
florescence, 2-3 ft. high, and with a loose spike of 20-30 spikelets.
Spikelets 10-20-flowered ; lemma awned. Cultivated for hay.

In fields and roadsides. Also called Italian rye-grass. June,
July. From Europe. Flamilton, Madison.

3. Lolium temulentum L. Poison Darnel. An annual glabrous
grass with simple, erect stems, 2-4 ft. high, and a flat, coarse spike.
Spikelets 5-7-flowered ; outer empty glume equalling or exceeding
the spikelet ; lemma obscurely nerved, awned or awnless. The grain
of this grass is said to be poisonous.

In waste and cultivated fields; often a troublesome weed. June-
August. No specimens. From Europe.

23. Agropyron J. Gaertn. Couch-grass, Wheat-grass.

Perennial grasses with simple stems and terminal spikes. Spike¬
lets 3-many-flowered, solitary in the notches of the raehis. the side

292

OHIO BIOLOGICAL SURVEY

of the spikelet turned toward the rachis ; empty glumes equal, usually
rigid, several-nerved, acute or awned ; lemma rounded on the back,
rigid, 5-7-nerved, usually acute or awned ; palet 2-keeled, the keels
bristly ciliate; grain usually adherent to the palet.

1. Lemma awn-pointed or with an awn usually not more than as long as the

body; stems from creeping rhizomes. A. repens.

1. Lemma wth an awn about twice as long as its body; stems tufted, without

creeping rhizomes. A. caninum.

1. Agropyron repens (L.) Beauv. Couch-grass. A perennial,
usually glaucous grass with stem 1-4^ ft. high, from a long, jointed,
running rhizome, and with a strict spike. Spikelets 3-7-flowered ;
lemma glabrous or more or less scabrous, strongly nerved, acute or
short awned at the apex. Often a bad weed. Frequently infested
with ergot. The rhizome is used in medicine.

In fields, roadsides, and waste places. Also called Quitch-grass.
July, August. Rather general; no specimens from the southeastern
part of the state. From Europe.

2. Agropyron caninum (L.) R. & S. Awned Wheat-grass. A
perennial grass with simple, erect stems, 1-3 ft. high, and more or
less nodding spikes. Spikelets 3-6-flowered ; empty glumes awn-
pointed or bearing an awn ; lemma 3-5-nerved, usually scabrous
toward the apex, bearing an awn sometimes twice the length of its
body.

In cultivated grounds and meadows. July-September. Portage
County. From Europe.

24. Triticum L. Wheat.

Tall annual grasses with flat leaves, with spikelets single at
the nodes of the spike, and with their sides turned to the rachis.
Spikelets 2-6-flowered ; empty glumes rather broad ; lemma rounded
on the back, many-nerved ending in one or more points or awns ;
palet 2-keeled ; grain with a deep groove, hairy at the apex, free.

1. Triticum aestivum L. Wheat. An annual grass with erect
hollow stems, 1-5 ft. high, and compact terminal spikes. Spikelets
3-6-flowered ; lemma awned or awnless. Cultivated in numerous
spring and winter A^arieties for its grain and straw.

In fields and waste places, especially along railways. June, July.
Erie, Belmont, Harrison, Tuscarawas, Morrow, Eayette, Madison,
Preble, Eranklin. Escaped from cultivation.

25. SecMe L. Rye.

Tall annual grasses with flat leaves and simple stems, with the
spikelets single at the nodes of the spike, and with their sides to the
rachis. Spikelets usually with 2 perfect flowers ; empty glumes

GRASSES OF OHIO

293

lanceolate, awn-pointed; lemma keeled, the keel hispid-ciliate,
awned ; palet 2-keeled; grain with a deep groove, hairy at the
apex, free.

1. Secale cereale L. Rye. A tall, light green, annual grass with
slender, erect, hollow stems, 2-6 ft. high, and more or less nodding,
terminal spikes. Spikelets 2-3-flowered ; lemma sharply keeled,
awned. Cultivated as a winter annual for its grain and straw, and
for temporary pasturage. The grain is sometimes infested and dis¬
torted by ergot.

In cultivated fields and waste places, especially along railways.
May-July. Erie, Morrow, Franklin, Scioto. Escaped.

26. Elymus L. Wild-rye.

Tall, erect tufted perennials with flat leaves and dense terminal
spikes. Spikelets 2-several-flowered, sessile, usually in pairs, some¬
times in threes; empty glumes equal, rigid, narrow, 1-3-nerved, acute
or awned ; lemma convex, obscurely 5-nerved, usually with an awn ;
palet 2-keeled, a little shorter than the lemma ; grain adherent to
the palet.

1. Empty glumes narrowly lanceolate ; spike stout. 2

1. Empty glumes narrowly awl-shaped ; spike and stem slender. E. striatiis.

2. Lemma glabrous ; spike erect, empty glumes indurated. E. virginicus.

2. Lemma more or less pubescent. 3

3. Spike erect; empty glumes indurated below. E. hirsutiglumis.

3. Spike nodding, empty glumes not indurated below. E. canadensis.

1. Elymus virginicus L. Virginia Wild-rye. A dark yellow-
green grass with stout, erect, simple stems, 2-5 ft. high, and thick,
rigidly upright spikes, often included at the base in the uppermost
inflated sheath. Spikelets 2-3-flowered ; lemma smooth, bearing a
rough awn. Extensively infested with ergot.

In moist soil, especially in ravines and along streams. July,
August. General.

2. Elymus hirsutiglumis Scrib. Strict Wild-rye. A grass with
an erect stem, 2-3 ft. high, and a stout, erect spike, the rachis
pubescent. Spikelets crowded, in pairs, 2'-5-flowered ; lemma 5>-
nerved, lanceolate, appressed-hirsute, long acuminate into a rough
awn.

On river banks and moist places. July, August. Ottawa, Huron.

3. Elymus canadensis L. Nodding Wild-rye. A light green or
usually glaucous grass with simple erect stems, 2-7yi ft. high, and a
broad, stout, compact, nodding spike interrupted below, its peduncle
much exserted. Spikelets divergent from the rachis, 3-5-
flowered ; lemma hirsute with a long, spreading awn. Often infested
with ergot.

In sandy soil and moist ground. July, August. Rather general.

294

OHIO BIOLOGICAL SURVEY

4. Elymus striatus \\411d. Slender Wild-rye. A more or less
pubescent grass with simple, slender, erect stems, 2-3 ft. high, and
slender, dense, sometimes nodding spikes. Spikelets 1-3-flowered,
diverg'ent from the rachis ; lemma hirsute, bearing a slender, rough
awn. Sometimes infested with ergot.

In woods and on banks. June, July. Wayne, Erie, Auglaize.

27. Hystrix Moench. Bottle-brush-grass.

Usually tall perennials with simple stems, flat leaves, and loosely
flowered terminal spikes. Spikelets 2-several-flowered, in pairs, rarely
in 3’s at each node of the rachis ; empty glumes vestigial ; lemmas
narrow, rigid, rounded on the back, 5-nerved, awned ; palet 2-keeled ;
grain adherent to the palet when dry.

1. Hystrix hystrix (L.) Millsp. Bottle-brush-grass A grass
with simple erect stems, 2-4 ft. high, and short exserted or partially
included lax spikes. Spikelets usually distant, at length widely di¬
vergent ; lemma more or less pubescent, acuminate into an awn
about 1 in. in length.

In rocky woods. June, July. General.

28. Hordeum (Tourn.) L. Barley.

Tufted annual or perennial grasses with terminal spikes, which
disarticulate at maturity. Spikelets with one perfect flower, rarely
2-flowered, usually in 3’s at each joint of the rachis, the middle one
sessile and perfect, the lateral ones pediceled and usually reduced ;
empty glumes equal, rigid, narrow or setaceous; lemma obscurely
5-nerved, awned ; palet slightly shorter, the two keels near the
margin ; grain usually adherent to the palet at maturity.

1. With 3 fertile spikelets at each joint of the rachis; lemma with an awn 4-6
in. long. H. vulgare.

1. With 1 fertile and 2 lateral, more or less vestigial spikelets at each joint of
the rachis. 2

2. Spike flat; awn of the lemma enclosing the fertile flower 4-6 in. long.

H. distichum.

2. Spike not decidedly flattened, awn 3 in. or much less in length. 3

3. Awns 1/2 in. or less in length. H. nodosum.

3. Awns 1-3 in. long, the whole spike appearing like a brush. H. jubatum.

1. Hordeum vulgare L. Common Barley. An annual grass
with simple erect hollow stems, 1-3 ft. high, and dense terminal
spikes. Spikelets 3 at a joint of the rachis each with a perfect flower;
lemma usually with a coarse long awn which breaks readily. Cul¬
tivated for its grain.

In fields and waste places, especially along railways. June,
July. Franklin, Tuscarawas, Portage. Escaped from cultivation.

GRASSES OF OHIO

295

2. Hordeum distichum L. Two-rowed Barley. An annual
grass like Hordeum vulgare but with the 2 lateral spikelets at each
node of the rachis vestigial and containing vestigial flowers.

Cultivated for its grain, usually mixed with H. vulgare. In fields
and waste places. June, July. Escaped in Lake and Erie Counties.

3. Hordeum nodosum L. Meadow Barley. An annual grass
with erect or sometimes decumbent stems, 3^-2 ft. high, and erect
spikes. Spikelets 3 at a node of the rachis, the central one with a
perfect flower, the lateral ones with staminate or vestigial flowers;
lemmas awned.

In thin dry soils and waste places. May, June. Hamilton
County.

4. Hordeum jubMum L. Squirrel-tail Barley. A biennial grass
with simple, erect, usually slender stems, ^4-2^2 ft. high, and a
nodding, densely awned spike. Spikelets in 3’s, the central one con¬
taining a perfect flower, the lateral ones vestigial ; empty glumes
awn-like ; lemma with an awn as long as the empty glumes ; all
the awns scabrous and very slender.

A weed. In dry soil and waste ground. June-August. From
Lake to Lucas County; also in Franklin, Madison, Greene, Allen,
Defiance and Williams. Naturalized from the West.

296

OHIO BIOLOGICAL SURVEY

Tribe, Chlorideae. Chloris Tribe.

Grasses ivith i-several-flozvered, laterally compressed spikelets,
jointed above the empty glumes and borne in 2 rows on one side of a
continuous axis, forming spikes which are digitately or paniculately
arranged, or sometimes solitary.

29. Spartina Schreb. Slough-grass.

Coarse perennial glabrous grasses with strong horizontal
rhizomes, rigid stems, long tough leaves, and an inflorescence of
one-sided spreading or erect alternate spikes. Spikelets 1-flowered,
flattened laterally, borne in two rows on the rachis ; empty glumes
unequal, keeled, acute or bristle-pointed; lemma obtuse, 1-nerved;
palet equaling or exceeding the lemma, 2-nerved ; grain free.

1. Spartina michauxiana Hitch. Tall Slough-grass. A coarse
grass with a simple erect stem, 2-10 ft. high, with an inflorescence
of 5-20 spike-like branches. Leaves with a scabrous cutting edge.
Spikelets much imbricated ; lemma glabrous except the serrulate-
scabrous midnerve which terminates abruptly below the emarginate
or 2-toothed apex.

Used for thatching and in making coarse mats. In swamps and
streams of fresh or brackish water, in wet ravines, and even sand
dunes. August-October. Rather general ; no specimens from the
central, eastern nor from the southwestern counties.

30. Beckmannia Host. Beckmannia.

A tall, erect perennial grass with flat leaf-blades and erect spikes
borne in a terminal panicle. Spikelets 1-2-flowered, globose, lat¬
erally compressed, closely imbricated in two rows along one side
of a continuous rachis ; empty glumes subequal, inflated, boat-shaped,
chartaceous, with scarious margins ; lemma narrow, lanceolate, thin-
membraneous ; palet hyaline, 2-keeled ; grain free, enclosed in the
lemma and palet.

1. Beckmannia erucaeformis fL.) Host. Beckmannia. A glab¬
rous grass with flattened wiry stems, 1^-3 ft. high, and a panicle of
appressed spikes. Spikelets nearly circular in outline, closely imbri¬
cated in 2 rows on the rachis ; empty glumes transversely wrinkled ;
lemma with an acuminate apex.

In wet ground. July-September. Cuyahoga County.

GRASSES OF OHIO

297

31. Capriola Adans. Bermuda-grass.

Low diffusely branched, extensively creeping perennials, with
flat leaf blades and slender spikes digitately arranged at the apex
of upright branches. Spikelets 1-flowered, laterally compressed, ses¬
sile, in two rows along one side of a slender continuous axis ; empty
glumes unequal, narrow, keeled ; lemma broad, boat-shaped, ciliate
on the keel ; palet 2-keeled, hyaline ; grain free.

1. Capriola dactylon (L.) Ktz. Bermuda-grass. A glabrous
grass with flattened, wiry, erect stems, ^-1 ft. high, from long creep¬
ing branched stolons, and a digitate inflorescence of 4-5 spikes.
Spikelets imbricated ; lemma broad and boat-shaped, longer than the
empty glumes.

In flelds and waste places. Cultivated in the south for pasture.
July-Sept. No specimens. From Europe.

32. Eleusine Gaertn. Yard-grass

Coarse tufted annuals or perennials with stout unilaterial spikes,
digitate or approximate at the summit of the stem. Spikelets sev¬
eral-flowered, sessile, closely imbricated in two rows along one side
of a continuous rachis which is not extended beyond them; empty
glumes unequal, shorter than the flowers, scabrous on the keel ; lemma
with a thickened 5-ribbed keel ; palet shorter, acute, the narrowly
winged keels distant ; grain black, loosely enclosed in the lemma
and palet.

1. Eleusine mdica (L.) Gaert. Yard-grass. A glabrous tufted
grass with flattened erect or decumbent stems, ^-2 ft. high, and an
inflorescence of 2-10, more or less digitately arranged spikes. Spike¬
lets appressed, 3-6-flbwered ; lemma with a thickened 5-ribbed keel.
A weed.

In yards, sidewalks, and waste places. June-September. Gen¬
eral. Naturalized.

33. Atheropogon Muhl. Grama-grass.

Perennial grasses with solid stems, narrow flat leaf-blades, and
an inflorescence composed of numerous, short, scattered, one-sided,
spreading or reflex spikes. Spikelets 1-flowered, crowded in two
rows ; empty glumes unequal, acute, narrow, keeled ; lemma 3-
toothed ; palet narrow, hyaline, 2-toothed ; grain free, enclosed in
the flowering glumes.

1. Atheropogon curtipendulus (Mx.) Fourn. Tall Grama-
grass. A gray-green grass with simple, solid, erect stems, 1-4^^ ft.
high and a long slender raceme of 15-20 spreading or reflexed spikes

298

OHIO BIOLOGICAL SURVEY

whose rachis is bifid at the apex. Spikelets 4-12 on each branch ;
lemma scabrous, ending in 3 short awns ; sterile lemma with 2 acute
lobes and 3 straight awns.

A valuable hay and pasture grass. In dry soil. July-Sept. Lake,
Erie, Ottawa, Franklin, Adams.

34. Bouteloua Lag. Mesquite-grass.

Annual or perennial usually gray-green grasses with narrow, flat,
or convolute leaves and numerous spikelets in 1-sided spikes. Spike-
lets 1-2-flowered, arranged in two rows on one side of a flat rachis
and bearing 1-3 awns and 1-3 rudimentary glumes; empty glumes
unequal, keeled ; lem^ma 3-5-nerved, 3-toothed, the teeth awn-pointed
or awned ; palet hyaline, 2-keeled, 2-toothed ; grain free.

1. Rachilla glabrous ; inner empty glume strongly papillose-hispid on the keel.

B. hirsuta.

1. Rachilla with a tuft of hairs; inner empty glume scabrous and sometimes

sparingly long-ciliate on the keel. B. oligostachya.

1. Bouteloua hirsuta Lag. Hairy Mesquite-grass. A gray-
green grass with erect stems l4-l^ ft. high, leafy at the base, and
with an inflorescence of 1-4 erect or ascending spikes whose rachis
extends beyond the spikelets as a conspicuous point. Spikelets
numerous ; outer empty glume setaceous, the inner conspicuously
tuberculate-hirsute on the keel ; fertile lemma pubescent, 3-cleft,
the divisions awn-pointed ; rachilla without a tuft of hairs under the
vestigial glumes and awns.

A valuable pasture grass in the West. In dry soil, especially on
prairies. July-Sept. Waifs in Franklin County.

2. Bouteloua oligostachya (Nutt.) Torr. Smooth Alesquite-
grass. A gray-green grass with erect, simple stems, j4-2^ ft. high
with involute curly leaves, and with an inflorescence of 1-3 spikes
often strongly curved, the rachis terminating in a short inconspicuous
point. Spikelets numerous, outer empty glume hyaline, shorter than
the membranous inner one which is scabrous and sometimes long-
ciliate on the keel or with a few papillae ; fertile, lemma pilose,
3-cleft, the divisions short-awned ; sterile lemma consisting of two
truncate lobes and three equal awns with a tuft of long hairs at the
base.

A valuable pasture grass in the West. On dry prairies. July-
Sept. Waifs in Franklin County.

GRASSES OF OHIO

299

Tribe, Agrostideae. Bent-grass Tribe.

Grasses zvith more or less laterally compressed spikelets jointed
above the empty glumes and arranged in panicles or racemes; spikelets

1- flozvcred, the rachilla sometimes prolonged behind the palet into a
naked or plumose bristle.

35. Sporobolus R. Br. Rush-grass, Dropseed.

Perennial or rarely annual grasses with flat or convolute leaves
and open or contracted panicles, our species with a peculiar constric¬
tion about the middle of the leaf blade. Spikelets usually small, 1-
flowered, awnless ; empty glumes ^usually unequal, membranous ;
lemma 1-nerved, equaling or longer than the empty ones; palet

2- nerved, often splitting at maturity ; grain free, readily falling from
the spikelet.

1. Panicle contracted. 2

1. Panicle open, its branches spreading; stems tufted; perennials. 5

2. Lemma pubescent. 3

2. Lemma glabrous. 4

3. Panicle 2-5 in. long, its branches 1-2 in. long ; palet long-acuminate ; perennial.

5. clandestinus.

3. Panicle 1-2 in. long, its branches in. long or less; palet long-acute; annual.

S. vaginaedoTus.

4. Panicle 3-10 in. long ; sheaths cylindric ; palet obtuse ; perennial. S. asper.

4. Panicle 1-214 in. long, sheaths inflated ; palet acute ; annual. S. neglectus.

5. Spikelets in. long or less ; empty glumes ovate to lanceolate, leaf sheaths

densely pilose at the throat. 5. cryptandrus.

5. Spikelets nearly in. long; outer empty glumes subulate, leaf sheaths glabrous
or sparingly pilose at the summit. S. heterolepis.

1. Sporobolus asper (Mx.) Kunth. Longleaf Rush-grass. A
perennial grass with stout, erect, simple or occasionally branched
stems, 1^-3^ ft. high and terminal linear panicles, partly included
in the inflated upper sheath. Lemma glabrous, acutish or obtuse.

In dry sandy soil. Aug., Sept. Lake, Erie, Franklin.

2. Sporobolus vaginaeflorus Torr. Sheathed Rush-grass. An
annual grass with tufted, slender stems, ^-1^2 ft. high, and numerous
panicles partly included in the inflated sheath, the terminal one
exserted. Lemma acuminate, scabrous, shorter than the sharp-
pointed palet.

In dry soil. Aug., Sept. Auglaize, Madison, Warren, Vinton,
Athens.

3. Sporobolus neglectus Nash. Small Rush-grass. An annual
grass with erect slender often much branched, usually decumbent

300

OHIO BIOLOGICAL SURVEY

stems, 3^-1 ft. high, with inflated sheaths, and with the terminal
panicle usually more or less included in the upper sheath, the lateral
panicles enclosed in the sheaths. Lemma acute, glabrous, about
equalling the acute palet.

In dry and sandy soil. Aug., Sept. Cuyahoga, Wayne, Huron,
Auglaize.

4. Sporobolus cryptandrus (Torr.) Gr. Sand Dropseed. A per¬
ennial tufted grass with erect, simple stems, 1 3^-314 ft. high, or some¬
times branched at the base, and with an ample lead-colored, usually
open panicle included at the base in the upper sheath. Leaves long-
acuminate with a peculiar joint-like constriction about the middle
of the blade and a ring of long white hairs at its base. Lemma acute,
longer than the palet.

In sandy soil. Aug.-Oct. Lucas, Ottawa, Erie, Lorain.

5. Sporobolus heterolepis Gr. Northern Dropseed. A tufted
perennial grass with rather stout, wiry, erect stems, 1-3 ft. high, and
long exserted panicles with ascending branches. Lemma glabrous,
obtuse or subacute.

In dry soil. Aug., Sept. Franklin, Madison, Champaign.

36. Calamagrdstis Adans. Reed Bent-grass.

Tall often reed-like perennial grasses with flat leaf-blades, run¬
ning rhizomes, and panicles with many spikelets. Spikelets
1-flowered, rachilla prolonged behind the flower into a hairy bristle
or pedicle ; empty glumes subequal, keeled, membranous ; lemma
awned on the back, surrounded at the base with copious long hairs ;
palet shorter than the lemma, 2-nerved ; grain free, enclosed in the
flowering glumes.

1. Prolongation of the rachilla hairy its whole length; awn straight; panicle

open, its branches spreading or ascending^ usually loosely flowered.

C. canadensis.

1. Prolongation of the rachilla hairy only at the summit; panicle contracted,

its branches erect. C. cinnoides.

1. Calamagrostis canadensis (Mx.) Beauv. Bluejoint Reed
Bent-grass. A large grass with clustered, simple or somewhat
branched, erect, hollow stems, 2-5 ft. high, and a loose, usually
purplish panicle, the slender fascicled branches erect or ascending.
Spikelets with copious hairs on the callus, about as long as the
flowering glumes, and surrounding them ; lemma thin, erose-truncate,
bearing a delicate awn on the back.

An important and valuable meadow grass, good for forage and
hay. In swamps and wet soils. July-Sept. Northern Ohio, as far
south as Stark, Franklin, and Auglaize Counties.

GRASSES OF OHIO

301

2. Calamagrostis cinnoides (Muhl.) Scrib. Nuttall’s Reed
Bent-grass. A glaucous grass with stout, erect, simple stems, 3-5
ft. high, and contracted panicles with erect branches. Lemma acum¬
inate, awned above the middle ; callus-hairs about one-half the length
of the flowering glumes ; prolongation of the rachilla bearing a tuft
of terminal hairs.

In moist soil. July, Aug. No specimens.

37. Agrostis L. Bent-grass.

Annuals or perennials with flat or bristle-like leaves, membran¬
ous ligules, and open or contracted panicles. Spikelets 1-flowered ;
empty glumes subequal, membranous, keeled, acute; lemma broad,
obtuse, awnless or dorsally awned ; palet hyaline, shorter than the
lemma, sometimes minute or wanting; grain free, loosely inclosed
in the lemma.

1. Lemma with a prominent dorsal awn just above the middle; palet minute
or none. canina.

1. Lemma awnless or rarely with a short awn. 2

2. Palet conspicuous, at least half as long as the lemma, 2-nerved; panicle

open. A. alba.

2. Palet inconspicuous, minute and nerveless or wanting. 3

3. Stems weak, decumbent or prostrate at the base ; leaves lax. A. schweinitzii.

3. Stems and leaves erect. 4

4. Branches of the panicle not elongated, usually branching at or below the

middle. A. perennans.

4. Branches of the diffuse panicle long and capillary, usually branching above
the middle. A. hy emails.

1. Agrostis alba L. Red-top (Bent-grass). A perennial grass
with erect or decumbent stems, ^^-2^/2 ft. high, with creeping rhi¬
zomes or stoloniferous, and with a contracted or open usually purplish
panicle, the branches verticillate and ascending. Lemma nearly equal¬
ing the empty glumes, 3-nerved, rarely short-awned.

An important and valuable grass, extensively cultivated for pas¬
ture and hay. In flelds and meadows. July-Sept. General. From
Europe.

2. Agrostis schweimtzii Trin. Thin Bent-grass. A perennial
grass with weak, slender, simple or sparingly branched stems, 1-2^2
ft. high, from a decumbent or prostrate base, and with an open panicle
with widely spreading branches. Lemma shorter than the acuminate
unequal empty glumes; palet small or wanting.

In damp shaded places. July-Sept. Rather general.

3. Agrostis hyemalis (Walt.) B. S. P. Rough Bent-grass. A
perennial grass with very slender simple erect stems, 1-2 ft. high,
and loose purplish panicles, the whorled capillary branches spikelet-

302

OHIO BIOLOGICAL SURVEY

bearing- at the ends. Lemma awnless or rarely short-awned on the
back, palet very small or obsolete.

In dry or moist sandy or stony soil. June-Aug. Rather gen¬
eral, but no specimens from the northwestern nor southeastern
counties.

38. Apera Adans. Windlestraw.

Annual grasses with flat narrow leaves and diffuse panicles.
Spikelets 1-flowered, small, the rachilla prolonged beyond the flower
into a bristle ; empty glumes subequal, thin, keeled, acute ; lemma
membranous bearing a slender awn just below the 2-toothed apex;
palet nearly as long as the lemma, 2-keeled, 2-toothed ; grain nar¬
row, free.

1. Apera spica-venti (L.) Beauv. Silky Windlestraw. A grass
with tufted slender erect simple stems, 1-2 ft. high, and an open
panicle with verticillate capillary ascending branches, spikelet-bear-
ing near the ends. Spikelets shining; lemma scabrous, bearing a
dorsal awn.

In waste places. June, July. From Europe. Lake County.

39. Cinna L. Wood Reed-grass.

Tall perennial grasses with flat leaf-blades, conspicuous hyaline
ligules, and many-flowered nodding panicles. Spikelets 1-flowered,
the rachilla articulated below the empty glumes and prolonged be¬
hind the palet into a minute bristle ; empty glumes narrow, keeled,
acute ; lemma 3-5-nerved, short-awned from between the 2 minute
teeth of the apex; palet 1- or 2-nerved, the nerves close together;
stamen 1 ; grain narrow, free, enclosed in the flowering glumes.

1. Cinna arundinacea L. Wood Reed-grass. A tall grass with
erect, simple, often solitary stems, 2-5 ft. high, rather broad leaf
blades and a contracted panicle with slender ascending branches.
Lemma scabrous, usually bearing a minute awn from the 2-toothed
apex.

In moist woods and shaded swamps. Aug., Sept. General.

40. Ammophila Host. Beach-grass.

Coarse perennial grasses with long, creeping rhizomes, flat leaf-
blades, which are deeply grooved above and convolute on drying,
and with dense spike-like panicles. Spikelets 1-flowered, the rachilla
prolonged beyond the flower and hairy; empty glumes firm, com-
pressed-keeled, acute, the outer 1-nerved, the inner 3-nerved ; lemma
5-nerved, with short hairs at the base, 2-toothed and mucronate be-

GRASSES OF OHIO

303

tween the teeth; palet rather firm, its two nerves close together;
grain free, loosely enclosed in the flowering glumes.

1. Ammophila arenaria ( L.) Link. Sand Beach-grass. A glab¬
rous grass with erect, rigid, stout stems, 2-4 ft. high, from long
horizontal branching rhizomes, and with a dense, spike-like, com¬
pressed panicle. Lemma scabrous, usually with a minute awn just
below the apex.

On sandy beaches of the sea coast and the Great Lakes. July,
Aug. Erie County.

41. Alopecurus L. Foxtail.

Branching annual or perennial grasses with flat leaves, erect or
decumbent stems, and soft dense spike-like panicles. Spikelets 1-
flowered, flattened, falling from the axis entire; empty glumes usually
somewhat united at the base, keeled ; lemma broad, obtuse, 5-nerved,
with a slender dorsal awn on the back ; palet usually none.

1. Lemma shorter than the obtuse, long-ciliate empty glumes ; empty glumes

less than yg in. long. A. geniculatus.

1. Lemma as long as the acute long-ciliate empty glumes; empty glumes Yg-yi

in. long. A. pratensis.

1. Alopecurus geniculatus L. Marsh Foxtail. A grass, usually

glabrous, with slender, decumbent stems, simple or sparingly
branched, ft. high, and a slender spike-like panicle. Lemma

obtuse, glabrous, with an awn inserted about one-fourth above its
base.

In wet soil. June-Aug. Lake, Ottawa, Crawford, Auglaize,
Madison, Franklin, Perry. Introduced.

2. Alopecurus pratensis L. Meadow Foxtail. A glabrous grass
with slender, simple, erect stems, 1-2^4 ft. high, from short, creeping
rhizomes, and with a spike-like panicle. Empty glumes united at
the base for about one-fourth of their length ; lemma obtuse, its awn
inserted about one-fourth above the base.

In meadows and pastures. May, June. No specimens. From
Europe.

42. Heleochloa Host. Cat-tail-grass.

Low tufted branching annuals or perennials with flat leaves and
numerous spike-like panicles partly included in the inflated sheaths.
Spikelets 1 -flowered, flattened; empty glumes subequal, membranous,
acute, ciliate keeled; lemma 1-nerved; palet shorter, 2-nerved ; grain
free, oblong, loosely enclosed in the flowering glumes.

1. Heleochola schoenoides (L.) Host. Cat-tail-grass. A low,
cespitose, glabrous annual, usually almost prostrate, with branched

304

OHIO BIOLOGICAL SURVEY

stems, y2-\y2 ft. long, and with dense, oblong-ovoid, spike-like
panicles.

In waste places. July, Aug. Greene County. From Europe.

43. Phleum L. Timothy.

Erect simple-stemmed perennials with flat leaves and dense,
spike-like panicles. Spikelets 1-flowered, flattened; empty glumes
membranous, compressed, keeled, abruptly awn-pointed ; lemma
broad, truncate, hyaline, 5-nerved; palet narrow, hyaline; grain
ovoid, free, enclosed in the flowering glumes.

1. Phleum pratense L. Timothy. A glabrous grass with sim¬
ple, hollow, erect stems, 1-5 ft. high, from a swollen base, and with
a long-cylindrical, compact, spike-like panicle. Empty glumes short-
awned, ciliate on the keel ; lemma 5-nerved, truncate, hyaline. A
very important grass extensively cultivated for hay.

In fields and meadows. Also called herd-grass. June-Aug. Gen¬
eral. From Europe.

44. Muhlenbergia Schreb. Muhlenbergia.

Mostly perennial grasses with flat or involute leaves and small
spikelets usually in contracted panicles, and usually with scaly rhi¬
zomes. Spikelets 1-flowered, usually with a short barbate callus
below the flower; empty glumes membranous or hyaline, acute,
sometimes awned ; lemma narrow, 3-nerved, awned or awnless ; palet
thin, 2-keeled ; grain narrow, free, tightly enclosed in the flower¬
ing glumes.

1. Lemma not awned, but the empty glumes may be short awned. 2

1. Lemma long-awned. awn mostly twice as long as the body of the lemma. 4

2. Empty glumes not awned, broadly ovate, acute, about half as long as the

lemma. M. sobolifera.

2, Empty glumes long acuminate or short awned, as long or longer than
the lemma. 3

3. Empty glumes about equal in length to the lemma, long pointed. M. mexicana.

3. Empty glumes much longer than the lemma, usually about twice as long,

short-awned ; panicle usually dense. M. racemosa.

4. Empty glumes at least G as long as the lemma. 5

4. Empty glumes not more than the length of the lemma, the outer one

often wanting. M. schreberi.

5. Empty glumes lanceolate, about equaling the lemma. M. umbrosa.

5. Empty glumes broadly ovate, Yi-Yz as long as the lemma. M. tenuiUora.

1. Muhlenbergia sobolifera (Muhl.) Trin. Rock Muhlenbergia.
A glabrous grass with erect or ascending, sparingly branched stems,
2-3 ft. high, with numerous conspicuous rhizomes covered with
short, appressed scales, and with very slender, usually loose-flowered
panicles. Lemma acute, scabrous, 3-nerved.

In rocky woods. Sept., Oct. Highland, Wayne.

GRASSES OF OHIO

305

2. Muhlenbergia mexicana (L.) Trin. Mexican Muhlenbergia.
A glabrous grass with erect or often prostrate, much-branched
stems, 2-4 ft. high, with numerous conspicuous rhizomes, covered
with short appressed scales, and with a long contracted panicle,
usually partly included within the upper sheath, its branches spike¬
like and erect or appressed. Lemma acuminate, scabrous, sometimes
awned.

In swamps and borders of fields. Aug., Sept. General.

3. Muhlenbergia racemosa (Mx.) B. S. P. Marsh Muhlen¬
bergia. A grass with erect, usually branched stems, 1-3 ft. high,
with numerous conspicuous rhizomes covered with short, appressed
scales, and with dense spike-like panicles, interrupted at the base,
the branches erect or appressed. Empty glumes aristate; lemma
acuminate, awnless, its midrib strongly scabrous.

In wet places. Aug.-Oct. Summit, Wayne, Huron, Wyandot,
Champaign, Licking.

4. Muhlenbergia umbrosa Scrib. Wood Muhlenbergia. A grass
with erect branched stems, 1-3 ft. high, with numerous conspicuous
rhizomes, covered with short appressed scales, and with somewhat
lax panicles, the branches erect or ascending. Lemma strongly scab¬
rous, attenuate into a slender awn.

In moist woods and along streams. Aug.-Oct. Cuyahoga,
Champaign.

5. Muhlenbergia tenuiflora (Willd.) B. S. P. Slender Muhlen¬
bergia. A grass with slender, simple or sparingly branched stems,
2-3 ft. high, with numerous conspicuous rhizomes covered with short,
appressed scales, and with a loosely flowered, slender panicle with
appressed branches. Lemma scabrous, tapering into a slender awn.

In rocky woods and ravines. Aug., Sept. Portage, Wayne,
Fairfield, Madison, Greene.

6. Muhlenbergia schreberi Gmel. Spreading Muhlenbergia. A
diffusely much-branched grass with erect or ascending stems, 1-3
ft. high, from a decumbent base, often rooting at the lower nodes,
and with slender, somewhat lax panicles, the erect branches rather
densely flowered. Empty glumes minute, the outer one nearly ob¬
solete; lemma strongly scabrous, tapering into a slender awn.

In dry woods, hillsides, and waste places. Aug., Sept. General.

45. Brachyelytrum Beauv. Brachyelytrum.

A tall perennial with flat leaves, simple stems from short knotty
rhizomes, and a narrow, few-flowered panicle. Spikelets 1-flowered,
narrow, the rachilla prolonged into a slender naked bristle ; empty
glumes minute, unequal ; lemma rigid, 5-nerved, ending in a long

306

OHIO BIOLOGICAL SURVEY

straight awn ; palet rigid, 2-nervecl ; grain oblong, free, enclosed
in the flowering glumes.

1. Brachyelytrum erectum (Schreb.) Beauv. Brachyelytrum.
A grass with erect, slender, simple stems, 1-3 ft. high, with sparingly
retrorse-hispid sheaths, and with slender, narrow panicles. Empty
glumes unequal, the outer often obsolete ; lemma 5-nerved with an
erect awn.

In moist, rocky woods. July, August. Cuyahoga, Portage,
Lorain, Wayne, Highland, Franklin, Madison, Hocking, Adams.

46. Milium L. Millet-grass.

Annual or perennial grasses with flat leaves and terminal open
panicles. Spikelets 1-flowered ; empty glumes equal; lemma shining,
indurated, the margins inrolled ; palet similar, scarcely shorter; grain
free, tightly enclosed in the rigid, shining flowering glumes.

1. Milium effusum L. Tall Millet-grass. A glabrous perennial
grass with erect, simple stems, 2-6 ft. high, and a lax panicle with
long, slender, somewhat flexuous branches, in remote pairs or
fascicles, spikelet-bearing from about the middle. Empty glumes
equal ; lemma smooth, white.

In woods. June, July. Lake, Cuyahoga, Lorain, Wayne, Stark.

47. Oryzopsis Mx. Mountain-rice.

Tufted perennial grasses with flat or convolute leaves and panicu¬
late few-flowered inflorescences. Spikelets, 1-flowered, broad; empty
glumes rather broad, about equal, obtuse or acuminate; lemma some¬
what indurated, convolute, bearing a terminal deciduous slender awn ;
palet rather large ; grain free, tightly enclosed in the convolute lemma.

1. Oryzopsis racemosa (Sm.) Ricker. Black-fruited Mountain-
rice. A glabrous grass with erect, tufted stems, 1^-3 ft. high, and a
small panicle with nearly simple usually ascending branches. Empty
glumes about equal, acute ; lemma dark colored, acute, sparingly
pubescent, with a long awn.

In rocky woods. July-Sept. Geauga, Summit, Erie, Greene.
Highland.

48. Stipa L . Porcupine-grass.

Generally tall, tufted perennial grasses with convolute leaf-blades
and loose terminal panicles. Spikelets 1-flowered, narrow; empty
glumes narrow, acute or bristle-tipped ; lemma convolute, rigid, with

GRASSES OF OHIO

307

a hairy callus at the base, terminating in a simple, strong persistent,
bent, twisted awn ; palet 2-nerved, small ; grain narrow, free, tightly
enclosed in the fruiting lemma.

1. Stipa spartea Trin. Porcupine-grass. A tall, slender grass
with erect, simple stems, 2-4 ft. high, with involute leaf blades and
loose, narrow panicles, finally long-exserted. Empty glumes ac¬
uminate into long, slender points, glabrous ; lemma brownish, ap-
pressed-pubescent below with a long, rigid, scabrous, twisted, usually
bent awn ; callus acuminate, very sharp-pointed, densely clothed with
silky, appressed hairs. The awns are highly hygrometric, and the
sharp-pointed fruit is injurious to animals, by penetrating the skin.

On plains, prairies, and sandy places. June, July. Erie County,
where it occurs on Cedar Point.

49. Aristida L. Triple-awn-grass.

Tufted annuals or perennials with narrow leaves and usually
with loose, narrow panicles. Spikelets 1 -flowered; empty glumes
unequal, narrow, acute or acuminate ; lemma somewhat indurated,
convolute, terminating in a three-branched awn ; palet thin, 2-nerved ;
grain elongated, free, tightly enclosed in the lemma.

1. Awns separate to the base, not articulated to the lemma. 2

1. Awns united below into a long twisted neck and articulated to the lemma.

A. tuberculosa.

2. Lateral awns much shorter than the middle one. 3

2. Lateral awns not much shorter than the middle one. 4

3. Middle awn coiled at the base. A. dichotonm.

3. Middle awn not coiled at the base. A. gracilis.

4. Outer empty glume 5-7-nerved; annuals. A. oliganfha.

4. Outer empty glume 1-3 nerved; perennials. A. piirpurascens.

1. Aristida dichotoma Mx. Poverty-grass. An annual tufted
grass with wiry much-branched stems, J^-2 ft. high, loose, glabrous
sheaths, much shorter than the internodes, and narrow, simple, few-
flowered, spike-like panicles, often reduced to racemes, the lateral
ones often sessile and partly enclosed in the sheaths. Lemma with
a horizontal awn, coiled at the base, with the two lateral awns erect
and very short.

In dry sandy soil. Aug., Sept. Scioto, Vinton, Eairfield.

2. Aristida oligantha Mx. Eew-flowered Triple-awn-grass. An
annual, tufted grass with wiry, branched, erect, solid stems, loose
sheaths, and few-flowered, lax, spike-like racemes or panicles. Empty
glumes unequal, awned ; lemma scabrous above with three nearly
equal, divergent awns.

In dry sterile soil. Aug., Sept. Cuyahoga County.

308

OHIO BIOLOGICAL SURVEY

3. Aristida gracilis Ell. Slender Triple-awn-grass. A glabrous
annual grass with slender, sparingly branched stems, ^-2 ft. high,
in small tufts or solitary, with rather close sheaths, and with a
spike-like, slender panicle or raceme. Lemma usually mottled, with
a horizontal middle awn and two much shorter, erect, lateral awns.

In dry sandy soil. Aug., Sept. Hamilton, Clermont, Athens,
Erie, Cuyahoga.

4. - Aristida purpurascens Poir. Purplish Triple-awn-grass. A
glabrous perennial grass, growing in small tufts, with erect, simple
or sparingly branched stems, 1-23/2 ft. high, and with a strict or
sometimes nodding purplish panicle with appressed branches. Lemma
with a horizontal middle awn and two somewhat shorter erect or
divergent lateral awns.

In dry sandy or gravelly soil. Sept., Oct. Wood, Fulton.

GRASSES OF OHIO

309

Tribe, Phalarideae. Canary-grass Tribe.

Grasses with laterally compressed spikelets jointed above the empty
glumes, arranged in panicles or racemes; spikelets with one perfect
dower and 2 staminate or vestigial dowers below, the vestiges sometimes
reduced to mere scale-like or bristle-like lemmas or obsolete.

50. Savastana Schrank. Vanilla-grass.

Fragrant perennial grasses with flat leaves and terminal panicles.
Spikelets 3-flowered, the terminal flower bisporangiate, the others
staminate or vestigial ; empty glumes subequal, acute, glabrous, and
shining; lemmas of the staminate flowers boat-shaped, indurated,
and hairy, each enclosing a 2-nerved palet and 3 stamens ; lemma of
the perfect flower similar but enclosing a 1-nerved palet and 2
stamens ; grain free, enclosed in the flowering glumes.

1. Savastana odorata (L.) Scrib. Vanilla-grass. A glabrous
grass with simple erect stems, 1-2 ft. high, from a creeping rhizome,
and with a pyramidal, usually compact panicle with slender droop¬
ing branches. Spikelets brownish ; staminate lemmas hispid-ciliate on
the margins and below the apex on the keel, awnless ; fertile lemma
hairy at the apex.

This sweet-scented grass is often strewn before churches in
northern Europe, whence it is often called holy-grass. In moist
meadows. May-July. Trumbull, Madison, Pickaway.

51. Phalaris L. Canary-grass.

Annuals or perennials with flat leaves and dense spike-like or
capitate panicles. Spikelets 1-flowered, laterally flattened ; empty
glumes equal, boat-shaped, strongly compressed and usually wing-
keeled ; sterile lemmas minute; fertile lemma indurated and shining;
palet faintly 2-nerved ; grain oblong, free, enclosed in the flowering
glumes.

1. Empty glumes not winged on the back; panicle narrow, branched.

P. anindinacea.

1. Empty glumes broadly wing-keeled on the back ; panicle very dense, appear¬
ing like a short spike. P. canariensis.

1. Phalaris arundinacea L. Reed Canary-grass. A glabrous
perennial with simple erect stems, 2-5 ft. high, and a dense panicle
with erect or slightly spreading branches. Spikelets lanceolate ;
sterile lemmas reduced to minute hairy scales; fertile lemma char-

310

OHIO BIOLOGICAL SURVEY

taceous with long appressed silky hairs. The common ribbon-grass
of gardens is the variety picta L.

In moist or wet soil. June-Aug. Rather general; no specimens
from the northw^estern nor from the southeastern counties.

2. Phalaris canariensis L. Canary-grass. An annual grass with
erect simple or branched stems, 1-3 ft. high, and an ovoid or oblong
head-like panicle. Spikelets broadly obovate, imbricated; empty
glumes wdiite wdth green veins ; empty lemmas thin-membranous,
sparingly hairy; fertile lemma brown, acute, with appressed silky
hairs.

This species furnishes the canary seed of commerce. In waste
places and roadsides. July, Aug. Cuyahoga, Montgomery, Hamil¬
ton. From Europe.

52. Anthoxanthum L. Vernal-grass.

Fragrant annual or perennial grasses with flat leaves and nar¬
row spike-like panicles. Spikelets 1-flowered, narrow, somewhat com¬
pressed ; empty glumes very unequal, acute or short-awned ; sterile
lemmas 2-lobed, awmed on the back ; fertile lemma, truncate, awnless ;
palet faintly 1-nerved; stamens 2; grain free, enclosed in the flower-
ing glumes.

1. Empty glumes pubescent; panicle 1^2-3 in. long; perennial. A. odoratum.

1. Empty glumes glabrous; panicle in. long; annual. A. piielii.

1. Anthoxanthum odoratum L. Sw^eet Vernal-grass. A per¬
ennial grass wdth simple or branched erect stems, 1-2 ft. high, and
spike-like panicles, the short branches erect or ascending. Spikelets
brownish green, spreading at flowering-time ; empty glumes sparsely
pilose ; first sterile lemma short-awned below the apex the second
bearing a strong bent awn near its base ; fertile lemma awnless.

In fields, pastures and waste places. May-July. Ashtabula,
Cuyahoga, Summit, Mahoning, Wayne, Franklin. From Europe.

2. Anthoxanthum puelii Lecoq & Lamotte. Long-awmed Ver¬
nal-grass. An annual grass with slender stems often branching above
the base, and with a dense spike-like panicle. Spikelets whitish green ;
empty glumes glabrous ; first sterile lemma with an awn twdce as
long as its body, pubescent below the middle, the second sterile
lemma with an awn 2-4 times the length of its body, also pubescent
below; fertile lemma awnless.

In waste places. May-Aug. Hamilton County. A native of
Europe*

GRASSES OF OHIO

311

Subfamily, Panicatae.

Tribe, Paniceae. Panic-grass Tribe.

Grasses having the 2-howered spikelets rounded or dorsally corn-
pressed, jointed below the empty glumes, and arranged in racemes or
panicles; lower dower staminate, vestigial or entirely absent; empty
glumes membranous ; dozvering glumes of the perfect dozver usually
awnless, indurated, drmly enclosing the free grain.

53. Panicum L. Panic-grass.

Annual or perennial grasses of various habit with the spikelets
in panicles. Spikelets 2-flowered, with 1 perfect flower and with
a staminate or vestigial one below ; empty glumes very unequal, the
outer often minute ; sterile lemma membranous often enclosing a
hyaline palet ; fertile lemma and palet indurated in fruit, margins of
the lemma thick and inrolled ; grain free, enclosed in the hardened
flowering glumes.

1. Basal leaves like those of the stem. 2

1. Basal leaves unlike those of the stem. 10

2. Basal leaf-sheaths compressed, often keeled. 3

2. Basal leaf-sheaths round, little flattened, never keeled. 4

3. Fruit stipitate ; spikelets conspicuously secund. P. stipitatum.

3. Fruit not stipitate; spikelets not conspicuously secund. P. agrostoides.

4. Leaf-sheaths smooth. 5

4. Leaf-sheaths pubescent ; annuals. 6

5. Outer empty glume about ^ as long as the spikelet ; tall perennials with

rhizomes. P. virgatum.

5. Outer empty glume about ^ as long as the spikelet; annuals.

P. dichotoinitlornm.

6. Spikelets ovate, more than % in. long; spikelets close. P. miliaceum.

6. Spikelets lanceolate or elliptic, less than % in. long; spikelets distant. 7

7. Panicle narrow, the branches ascending; spikelets less than ^ in. long.

P. dexile.

7. Panicle spreading when mature ; spikelet tV in. long or less. 8

8. Panicle very large, usually the length of the plant. P, capillare.

8. Panicle of medium size, usually the length of the plant. 9

9. Stem delicate; leaf-blade less than Y in. wide. P. philodclphicmn.

9. Stem stout; leaf-blade usually Y-ji in. wide. P. gattingeri.

10. Leaf-blades way up the stem less than ^2 in. wide, attenuate to cordate
at the base. 11

10. Leaf-blades Y2 way up the stem ^ in. or more wide, usually cordate clasp¬
ing at the base. 30

11. Spikelets Y in. long or more. 12

11. Spikelets less than Y in. long, usually about tV in. 22

12. Leaves less than in. wide and about 18 times as long as wide.

P. depaiiperatuin.

12. Leaves more than Ya in. wide, and not elongated. 13

13. Upper surface of the leaves glabrous. 14

13. Upper surface of the leaves pubescent. P. leibergii.

312

OHIO BIOLOGICAL SURVEY

14.

14.

15.

15.

16.

16.

17.

17.

18.
18.

19.

19.

20.

20.

21.

21.

22.

22.

23.

23.

24.

24.

25.

25.

26.
26.
27.

27.

28.

28.

29.

29.

30.

30.

31.

31.

32.

32.

33.

33.

34.

34.

Outer empty glume as long as the spikelet, short acute. P. scribnerianum.
Outer empty glume 3^ as long as the spikelet, long acuminate.

„ . , . P. xanthophysum.

Stem simple or with basal branches only. 16
Stem at length fasciately branched. 22

Hairs on the leaf sheath almost pg in. long or longer, spreading, usually
dense. ^ p Unearifolinm.

.9. hairs on the leaf sheath, or with hairs less than tV in. long, sometimes
cihate on the margin. 17

Leaf blade in. or less wide, usually 18 or more times as long as wide;
spikelets usually glabrous. 18

Leaf-blade usually much more than in. wide, never more than 8 times
longer than wide; spikelets more or less pubescent. 20.

Leaves much elongated; often 6 to 10 in. long, attenuate at the base.

P. werneri.

Leaves not elongated ; generally less than 33^ in, long, not attenuate at the
base, spikelet less than Yg in. long. P. bicknellii.

Spikelets roundish, not over tV in. long. 20

Spikelets oblong-elliptic or elliptic, usually in. or more long. 21
Nodes pubescent with appressed hairs, base of the leaves ciliate, panicle nearly
as wide as long. P. sphaerocarpon.

Nodes glabrous, base of the leaves not ciliate ; panicle not more than Yi

wide as long.

Stems pubescent.

Stems glabrous.

Spikelets glabrous. 23
Spikelets pubescent. 24
Nodes densely bearded.

Nodes not bearded.

Ligule at the top of the leaf sheath minute or absent. 25
Ligule present. 26

Nodes of the main stem glabrous or with a few hairs.

P. polyanthes.
P. tsugetorum.
P. boreale.

P. microcarpon,
P. dichotomum.

P.

P. boreale.

P. ashei.
lindheimeri.

Nodes of the main stem pubescent.

Upper sheaths glabrous, ciliate on the margin.

All of the sheaths pubescent. 27

Upper surface of the leaves glabrous, except for a few long hairs near the
base. P. tsugetorum.

Upper surface of the leaves pilose. 28

Outer empty glume acute; spikelet tV in. long, pubescence on the sheath more
than tV in. long. P. villosissimum.

Outer empty glume blunt ; spikelet less than in. long ; pubescence on the
sheaths not so dense and less than iV in .long. 29

Upper surface of the leaf-blade long-pilose; plants yellowish-green.

P. huachucae

Upper surface of the leaf-blade long appressed pubescent.

Spikelet 34 in. or more long. 32
Spikelet less than 34 In. long. 31
Spikelet less than in. long.

Spikelet more than tV in. long.

Leaf sheaths mostly papillose-hispid ; nodes glabrous or short pubescent.

P. eland estinum.

Leaf-sheaths glabrous or soft-pubescent. 33

Nodes glabrous. P. latifolium.

At least the lower nodes pubescent or bearded. 34

Panicle narrow, its branches appressed, rarely a little spreading; upper nodes
at least not bearded. P. xanthophysum.

Panicle open, its branches spreading; the nodes appressed-pubescent. P. boscii.

P. implicatum.

P. polyanthes.
P. commutatiim.

GRASSES OF OHIO

313

1. Panicum agrostoides Sprang. Agrostis-like Panic-grass. An
erect, rather stout, glabrous perennial, 1-3^ ft. high, with loose
sheaths and a purplish, oblong-ovate panicle, 6-12 in. long, the
branches stiffly ascending. Spikelets crowded on sparsely hairy, short
pedicels; lemma oval.

In wet ground and along shores. July-Sept. Portage, Erie.

2. Panicum stipitatum Nash. Long Panic-grass. A branched,
erect perennial with compressed stems, 3-5 ft. high, and leaf-blades
1 ft. or more long, often purplish and scabrous. Inflorescence a pyra¬
midal, purplish panicle ; spikelets secund, acuminate, crowded ; lemma
narrowly elliptic and raised on a delicate stalk.

In moist soil. July-Sept. Northeastern Ohio to Lorain, Fair-
field and Columbiana Counties.

3. Panicum virgatum L. Tall Smooth Panic-grass. A tall
glabrous perennial grass, 3-7^ ft. high, with creeping rhizomes, and
loose panicles with ascending or spreading branches. Spikelets ovate,
acute to acuminate ; outer empty glume about one-half as long
as the spikelet ; first lemma usually subtending a palet and staminate
flower. Second lemma shining, inclosing a perfect flower. An im¬
portant grass.

It makes fair hay when cut young. In dry or moist soil, es¬
pecially in prairies. Aug., Sept. General.

4. Panicum dichotomiflorum Mx. Spreading Panic-grass. A
glabrous branching annual grass with compressed decumbent and
geniculate stems, 4-6 ft. long, and a diffuse pyramidal panicle with
the lower branches at length widely spreading. Spikelets crowded,
lanceolate, acute, glabrous, sometimes purplish ; outer empty glume
one-fourth as long as the spikelet; fertile lemma elliptic, shining.

A weed. Cattle and horses are very fond of it. In wet soil and
cultivated fields. July-Sept. General.

5. Panicum miliaceum L. Millet Panic-grass. An erect or de¬
cumbent annual grass with rather stout stems, 1-2 ft. high, and a
dense panicle drooping at maturity. Sheaths papillose-hirsute. Spike¬
lets ovoid, acuminate, turgid ; outer empty glume ^ as long as the
spikelet.

In waste places. A good fodder grass. Sometimes cultivated
as a cereal. July-Sept. Lawrence, Richland, Erie. Introduced from
Europe.

6. Panicum capillare L. Tumble Panic-grass. A stout spar¬
ingly branched, erect or decumbent annual, l-5j4 ft. high, with
papillose-hirsute sheaths and a very large diffuse panicle with capil¬
lary branches, at first included in the sheath but later becoming
exserted and finally breaking off at maturity as a tumble grass.
Spikelets acute; outer empty glume 14-^4 as long as the spikelet.

314

OHIO BIOLOGICAL SURVEY

A bad weed. In dry soil and cultivated fields. July-Sept. Gen¬
eral and abundant.

7. Panicum gattmgeri Nash. Gattinger’s Panic-grass. An an¬
nual grass with widely spreading or decumbent stems, sometimes
forming mats, branching at all the nodes, 1-2 ft. long, and with
numerous exserted oval panicles, smaller and less diffuse than in
P. capillare. Sheaths hirsute ; spikelets elliptic, acute, glabrous, very
turgid.

In moist open ground. Aug.-Oct. Rather general.

8. Panicum flexile (Gatt.) Scrib. Wiry Panic-grass. A slender
erect annual grass, ^-2 ft. high with a few erect branches from the
base and with a narrowly oblong panicle with ascending branches,
usually one-half the length of the entire plant. Spikelets acuminate,
solitary at the ends of the branchlets ; outer empty glume ^.s long
as tlie spikelet.

In moist or dry sandy soil. Aug.-Oct. Adams, Champaign,
Madison, Franklin, Erie, Cuyahoga.

9. Panicum philadelphicum Bernh. Philadelphia Panic-grass.
A slender, erect or ascending, freely branching annual with zigzag
stems, 1-2 ft. long, and a rather few-flowered panicle about ^3 the
length of the plant. Spikelets elliptic, acute, smooth, usually in 2’s
at the ends of the divergent branchlets ; outer empty glume about
F3 the length of the spikelet.

In dry woods and thickets. Aug., Sept. Trumbull, Ottawa.

10. Panicum depauperatum Muhl. Starved Panic-grass. An

erect or ascending perennial ft. high, with erect elongated

leaves, and a few-flowered, much exserted, terminal panicle, the
lower panicles on very short basal branches. Spikelets glabrous,
acute ; outer empty glume the length of the spikelet.

In dry soil. June-Sept. Cuyahoga County.

11. Panicum linearifolium Scrib. Linear— leaf Panic-grass. A
densely tufted perennial grass with slender erect stems, 5^-lp2 ft.
high, and a rather few-flowered, loose panicle. Spikelets obtuse or
acutish, pubescent with spreading hairs; outer empty glume yi-Yz
as long as the spikelet.

In woods and hillsides. May-July. Rather general.

12. Panicum werneri Scrib. Werner’s Panic-grass. A smooth,
light green, tufted, sparingly branched or simple perennial grass,
Yz-^Yi ft. high, with erect, linear, acuminate leaves and a loose open
panicle, which is finally long-exserted. Spikelets oval, somewhat
pubescent ; outer empty glume Y ^-S long as the spikelet.

In dry woods, fields, and the drier parts of swamps. June, July.
Lake, Cuyahoga, Franklin, Athens.

GRASSES OF OHIO

315

13. Panicum bicknellii Nash. BicknelEs Panic-grass. A slen¬
der, usually stifif, erect or decumbent perennial grass, 5^-1 3^4 ft. high,
with the lower sheaths pubescent and a few-flowered panicle with
stiff, slender branches. Spikelets 'Oval or ovate, pubescent with
ascending hairs; outer empty glume the length of the spikelet.

On dry, wooded hills. July, Aug. Gallia County.

14. Panicum sphaerocarpon Ell. Round-fruited Panic-grass. A
dull green, usually erect, simple or somewhat branched perennial,
3^-2 ft. high, with somewhat pubescent nodes and an ovoid, long-
exserted, loosely flowered panicle with viscid spots on the axis and
ascending branches. Spikelets nearly spherical, obtuse, usually pur¬
ple ; outer empty glume broadly ovate, very short.

In dry or sandy soil. July-Sept. Cuyahoga, Summit, Trumbull,
Hocking, Scioto.

15. Panicum polyanthes Schultes. Many-flowered Panic-grass.
A light green perennial grass with erect, simple stems, 1-3 ft. high,
with glabrous nodes and very long sheaths with finely ciliate margins,
and with a densely flowered, elliptic panicle with slender ascending
branches. Spikelets obovoid or nearly spherical ; outer empty glume
minute.

In damp ground and woods. July-Sept. Fairfield, Hocking,
Jackson.

16. Panicum dichbtomum L. Forked Panic-grass. A glabrous,
often purplish, perennial grass with erect, dichotomously branched
stems, j4-2 ft. high, from short, knotted rhizomes, with light green
to purplish leaves, and with a much exserted terminal panicle with
lax spreading branches and few spikelets. Spikelets usually glabrous,
ellipsoid ; outer empty glume minute.

In woodlands and thickets. May-Aug. Rather general ; no
specimens from the northwestern counties.

17. Panicum microcarpon Muhl. Small-fruited Panic-grass. A
perennial grass, at first erect but later with densely branched, spread¬
ing or prostrate stems, 2-3 ft. long, strongly barbed on the nodes,
and with a long-exserted, rigid primary panicle and small few-
flowered secondary panicles. Spikelets purplish, glabrous ; outer
empty glume Ys as long as the spikelet.

In moist soil and swampy places. June-Aug. Cuyahoga, Lorain,
Erie, Fairfield, Hocking, Jackson, Adams.

18. Panicum boreMe Nash. Northern Panic-grass. An erect
or decumbent, somewhat branched perennial, 1-2 ft. high, with a
narrow, loosely flowered panicle, the slender branches ascending or
spreading. Spikelets ellipsoid, pubescent ; outer empty glume minute.

In moist, open ground or woods. June, July. Fulton County.

316

OHIO BIOLOGICAL SURVEY

19. Panicum lindheimeri Nash. Lindheimer’s Panic-grass. An
erect or spreading dichotomously branched perennial grass l-ly^ ft.
high, with swollen nodes and a loosely flowered papicle with ascend¬
ing or spreading branches, the primary one long-exserted. Spikelets
pubescent, obovoid ; outer empty glume minute.

In dry sandy woods and open ground. June-Aug. Ashtabula,
Hocking.

20. Panicum huachucae Ashe. Hairy Panic-grass. A profusely
branched erect or somewhat decumbent, hairy perennial grass, ^-1^
ft. high, with a rather densely flowered, ovoid panicle with ascending
or spreading branches. Spikelets pubescent ; outer empty glume
minute.

In dry soil and on prairies. June-Sept. General.

21. Panicum villosissimum Nash. Villous Panic-grass. A vil¬
lous, olive-green perennial grass with tufted, at length branched,
erect or ascending stems, 1-2 ft. high, and a loosely flowered, long-
exserted primary panicle. Spikelets obovate to elliptic, densely
pubescent with short spreading hairs ; outer empty glume a little
less than ^ as long as the spikelet.

In dry sandy soil and open woods. June-Aug. Cuyahoga, Erie,
Licking.

22. Panicum implicatum Scrib. Slender-stemmed Panic-grass.
A slender, tufted, more or less pubescent, at length much branched
perennial, j^-2 ft. high, with densely papillose-hirsute sheaths and
an open, ovate panicle with widely spreading flexuous branches.
Spikelets obovoid, obtuse, papillose-pilose ; outer empty glume almost

' p2 as long as the spikelet.

In wet soil. July-Sept. Gallia County.

23. Panicum tsugetorum Nash. Hemlock Panic-grass. A bluish-

green or sometimes purplish perennial with slender, tufted, branched,
decumbent or prostrate stems, ft. long, and a rather loosely

flowered panicle with ascending or spreading branches. Spikelets
(broadly ovate, pubescent ; outer empty glume ys as long as the
spikelet.

In sandy woods. June-Sept. Defiance, Summit.

24. Panicum leibergii (Vasey) Scrib. Leiberg’s Panic-grass.
A pubescent perennial grass, 1-234 ft. high, with slender, erect, and
at length branched stems, with papillose-hispid sheaths and an ob¬
long panicle with erect or ascending branches. Spikelets oval, papil¬
lose-hirsute, outer empty glume the length of the spikelet.

On prairies and dry soil. June, July. No specimens.

25. Panicum scribnerianum Nash. Scribner’s Panic-grass. An
erect perennial with simple or dichotomously branched stems, p2-2

GRASSES OF OHIO

317

ft. high, and a small, ovoid, short-exserted panicle. Spikelets obtuse,
very turgid, slightly pubescent; outer empty glume minute.

In dry or moist sandy soil or on dry prairies. June-Aug. Cuya¬
hoga, Erie, Wood, Franklin.

26. Panicum xanthophysum Gr. Slender Panic-grass. A
tufted, yellowish green perennial grass with simple, erect or ascend¬
ing stems, 1-2 ft. high, and a few-flowered, very narrow, finally long-
exserted panicle with erect branches. Spikelets broadly obovate,
very turgid, usually pubescent ; outer empty glume nearly as long
as the spikelet.

In dry soil. June-Aug. Lake County.

27. Panicum ashei Pear. Ashe's Panic-grass. An erect, stiff,

sparingly branched, purplish ‘ perennial grass, ft. high, with

a small panicle, its branches ascending. Spikelets elliptic, obtuse,
purplish ; outer empty glume minute.

In dry woods. May-Aug. Cuyahoga, Lake, Trumbull, Fairfield.

28. Panicum commutatum Schultes. Variable Panic-grass. An
erect, rather slender, finally dichotomously branched perennial grass,
1-2J4 ft. high, with firm leaf-blades, cordate clasping at the base,
and a lax panicle with spreading branches. Spikelets elliptic, obtuse ;
outer empty glume minute.

In dry woods and thickets. June-Aug. Lawrence, Gallia, Fair-
field, Wayne.

29. Panicum latifolium L. Broad-leaf Panic-grass. An erect,
nearly smooth perennial grass with simple or branched stems, 1-3
ft. high, with broad leaf-blades cordate clasping at the base, and a
long-exserted panicle with ascending few-flowered branches. Spike¬
lets ovate-elliptic, obtuse ; outer empty glume about Yz the length
of the spikelet.

In rocky woods and on sand dunes. July, Aug. General.

30. Panicum bdscii Poir. Bose’s Panic-grass. A glabrous or
minutely pubescent perennial with erect, simple or somewhat
dichotomously branched stems, l-2j^ ft. high, with broad leaves and
a slightly exserted panicle with few-flowered spreading or ascend¬
ing branches. Spikelets ovate ; outer empty glume nearly ^4 as long
as the spikelet.

In woods. June-Aug. Warren, Adams, Jackson, Belmont.

The variety P. boscii molle (Vas.) Hitch. & Ch. is downy-
pubescent thruout and usually not so tall. Reported from Hamil¬
ton, Lawrence, and Cuyahoga Counties.

31. Panicum clandestinum L. Hispid Panic-grass. An erect
or ascending, rather stout perennial with at first simple but later
much branched stems, 1J4-4 ft. high, with broad cordate clasping,
glabrous leaf-blades and rather long-exserted primary panicles with

318

OHIO BIOLOGICAL SURVEY

ascending branches, the secondary panicles usually included in the
sheaths. Spikelets elliptic, pubescent; outer empty glume ^ as
long as the spikelet.

In moist ground and thickets. June, July. General.

54. Leptoloma Chase. Witch-grass.

Tufted perennial grasses with flat leaves and very diffuse ter¬
minal panicles which break away at maturity and become tumble-
grasses. Spikelets with 1 perfect flower, solitary on long capillary
pedicels ; outer empty glume very minute or wanting, inner empty
glume 3-nerved, nearly as long as the 5-7-nerved sterile lemma ;
fertile lemma indurated, papillose, with hyaline margin not inrolled ;
lemma of similar texture ; grain free, enclosed in the flowering
glumes.

1. Leptoloma cognatum (Schultes) Chase. Fall Witch-grass.
A perennial slender tufted grass with erect or decumbent stems, 1-2
ft. high, usually much branched at the base, and with a loose panicle
bearded in the axils, the lower branches at first erect and included
in the upper sheath but finally exserted and widely spreading.
Spikelets lanceolate, acuminate, on capillary pedicels ; lemma lanceo¬
late.

In dry soil. July-Sept. From the West. Lake County.

»

55. Syntherisma Walt. Crab-grass.

Annual, mostly weedy grasses with branched stems, thin flat
leaves, and an inflorescence of secund, spike-like racemes which are
digitate or approximate at the ends of the branches. Spikelets with
1 perfect flower and a vestige of a second one, lanceolate-elliptic,
sessile or short pediceled ; empty glumes 1-3 nerved, the outer some¬
times obsolete ; sterile lemma 5-nerved ; fertile lemma chartaceous,
its margins hyaline and not inrolled; palet of similar texture; grain
free, enclosed in the flowering glumes.

1. Rachis of the raceme branches with the angles wingless ; outer empty glume
wanting or minute ; raceme branches 2-5 ; stems erect. S. filiforme.

1. Rachis of the raceme branches with the lateral angles broadly winged; stems
spreading. 2

2. Pedicels terete, glabrous or nearly so ; outer empty glume wanting or minute ;

raceme branches 2-6. Y. ischaeninm.

2. Pedicels sharply 3-angled, the angles hispidulous; outer empty glume present,
small, raceme branches 3-12. S. sanguinale

1 . Syntherisma filiforme (L.) Nash. Slender Crab-grass. A
slender usually tufted grass with erect stems, 1-4 ft. high, and a
panicle of 1-5 spike-like racemes. Spikelets elliptic, pubescent; outer

GRASSES OF OHIO

319

empty glume usually obsolete, the inner one and the sterile lemma
villous between the nerves with gland-tipped hairs ; fertile lemma
dark-brown, acute.

In dry sandy soil. July-Sept. No specimens.

2. Syntherisma ischaemum (Schreb.) Nash. Small Crab-grass.
A glabrous grass with spreading or decumbent much-branched
stems, 5^-2 ft. long, and a panicle of 2-6 aggregated spike-like racemes.
Inner empty glume and sterile lemma densely short-villous between
the nerves ; fertile lemma dark-brown.

In cultivated soil and waste places. July-Sept. Lorain, Wayne,
Auglaize, Fairfield. From Europe.

3. Syntherisma sanguinale (L.) Dulac. Large Crab-grass. A
grass with branched spreading stems, 1-5 ft. long, usually decumbent
at the base and often rooting at the lower nodes. Panicles of 3-12
narrowly linear, spike-like racemes, digitate or approximate in
whorls ; spikelets elliptic-lanceolate, acute ; outer empty glume usually
present, minute ; fertile lemma pale or grayish.

In cultivated and waste ground. A very troublesome weed.
Makes good hay and pasturage. July-Sept. General. Naturalized
from Europe.

56. Echinochloa Beauv. Barnyard-grass.

Usually coarse annual grasses with long broad leaf-blades, com¬
pressed sheaths, and terminal panicles of stout one-sided racemose
branches. Spikelets with 1 perfect terminal flower and a staminate
or vestigial one below ; empty glumes unequal, spiny hispid on the
nerves, mucronate ; sterile lemma similar, usually awned from the
apex, its palet hyaline ; fertile lemma and palet chartaceous, acu¬
minate ; margin of the lemma inrolled ; grain free, enclosed in the
flowering glumes.

1. Leaf sheaths glabrous ; second and third glumes more or less awned.

E. crus-galli.

1. Leaf sheaths, at least the lower ones densely papillose-hirsute; second and

third glumes long-awned. E. walteri.

1. Echinochloa crus-gMli (L.) Beauv. Common Barnyard-grass.
A rather succulent grass branching from the base, with ascending
or erect stems, 2-7^3 ft. long, and a dense panicle of numerous erect
or spreading, or sometimes reflexed branches. Spikelets ovate, green
or purple, densely crowded in rows on one side of the rachis ; empty
lemma more or less awned ; fertile lemma ovate, abruptly pointed.

A common weed. Sometimes cultivated for hay. A variety is
sometimes advertised as “Billion Dollar Grass.” In cultivated and
especially in manured soil and waste places. Aug.-Oct. General
and abundant. Naturalized from Europe.

320

OHIO BIOLOGICAL SURVEY

2. Echinochloa waited (Pursh.) Nash. Marsh Cockspur-grass.
A grass with robust more or less succulent stems, 3-6 ft. long, and
a long more or less drooping panicle with compact ascending or
spreading branches. Spikelets long-awned, ovate-lanceolate, on one
side of the rachis, brownish-purple; inner empty glume and empty
lemma awned ; fertile lemma, ovate-lanceolate, acuminate.

In marshes and ditches. Aug.-Oct. Erie, Lorain, Shelby, Au¬
glaize, Licking.

57. Paspalum L. Paspalum.

Perennial grasses of various habit with the spikelets in spike¬
like, one-sided racemes, digitate or racemose at the tip of the main
stem or branches. Spikelets with 1 perfect flower and a vestige,
plano-convex, nearly sessile, placed with the back of the fertile
lemma toward the rachis; empty glumes very unequal or the outer
entirely obsolete; lemma and palet indurated, margins of the lemma
inrolled ; grain free, enclosed in the flowering glumes.

1. Rachis of the racemes very broad, membranous with inrolled margins, about
twice the width of the spikelets ; rachis not extending beyond the spikelets.

P. dissectum.

1. Rachis narrower than the spikelets not inrolled on the margins. 2

2. Stems simple, axillary peduncles none; racemes stout; sheaths glabrous or

nearly so. P. laeve.

2. Stems branched, the upper sheaths containing from 1-3 branches ; racemes
slender. 3

3. Spikelets somewhat pubescent, in. long. P. setaceum.

3. Spikelets glabrous, nearly % in. long. P. muhlenbergii.

1. Paspalum setaceum Mx. Slender Paspalum. A slender grass
with erect or ascending stems, ^-2^4 ft. high, and with an inflor¬
escence of 1-2 spike-like racemes. Spikelets ovate, somewhat pube¬
scent, about 1/16 in. long; one of the empty glumes 3-nerved, the
other 2-nerved.

In dry fields. Aug.-Oct. Cuyahoga County.

2. Paspalum muhlenbergii Nash. Muhlenberg’s Paspalum. A
tufted branching pubescent perennial grass with spreading or re¬
clining stems, 1-2^ ft. long, and an inflorescence of 1-2 spike-like
racemes. Spikelets glabrous, oval or broadly obovate, in pairs ;
lemma with inrolled margins ; empty glumes usually 3-nerved.

In fields and sandy soil. Aug.-Oct. Cuyahoga, Erie, Warren,
Hamilton, Scioto, Guernsey.

58. Chaetdchloa Scrib. Foxtail-grass,

Annual or perennial grasses usually with erect stems and flat
leaf blades, the inflorescence in spike-like panicles containing nu¬
merous persistent awn-like bristles on the rachis below the articu-

GRASSES OF OHIO

321

lation of the spikelets. Spikelet 2-flowerecI, one perfect, the other
vestigial or rarely staminate; empty glumes very unequal; sterile
lemma often enclosing a hyaline palet; fertile lemma and palet char-
taceous or indurated in fruit ; grain free enclosed in the flowering
glumes.

1. Bristles of the inflorescence downwardly barbed ; comparatively inconspic¬
uous. . C. verticillata.

1. Bristles upwardly barbed. 2

2. Bristles at the base of the spikelets 5-16, involucrate, yellowish-brown ; in¬
florescence racemose ; inner empty glume shorter than the spikelet. C.glauca.

2. Bristles at the base of the spikelet 1-3, not involucrate; inflorescence pan-

iculate; inner empty glume as long as the spikelet. 3

3. Inflorescence 1-314 in. long^ J4 in. thick or less; bristles green. C.viridis.

3. Inflorescence 4-9 in. long, ^-2 in. thick; bristles usually purple. C. italica.

1. Chaetochloa verticillata (L.) Scrib. Verticillate Foxtail-
grass. A tufted annual grass with more or less branched, erect or
decumbent stems, 1-2 ft. high, and a dense spike-like green panicle
which is somewhat compound and interrupted at the base. Involucral
bristles of the inflorescence retrorsely barbed ; outer empty glume
1-nerved, less than one-half as long as the spikelet; fertile lemma
abruptly apiculate and obscurely transverse-rugose.

In waste places and yards. July-Sept. From Europe. Cuya¬
hoga, Wayne, Jefferson, Franklin, Ross, Montgomery, Warren, Ham¬
ilton. ,,,

2. Chaetochloa glauca (L.) Scrib. Yellow Foxtail-grass. An

annual grass with compressed, erect or ascending stems, branching
at the base, 1-4 ft. high, with glaucous leaves, and a yellowish, com¬
pact, spike-like panicle. Involucral bristles upwardly barbed, yellow¬
ish-brown ; spikelets oval, much shorter than the bristles ; fertile
lemma striate, undulate-rugose, very convex,

A weed. In cultivated fields and waste places. July-Sept. Gen¬
eral. From Europe.

3. Chaetochloa viridis (L.) Scrib. Green Eoxtail-grass. An

annual tufted grass with simple or branched, erect or ascending
stems, l-Sys ft. high, and a rather thick spike-like green panicle with
slender upwardly barbed greenish bristles. Spikelets elliptic, much
shorter than the bristles ; outer empty glume less than one-half as
long as the spikelet; fertile lemma only moderately convex, obtuse,
striate and faintly wrinkled.

Often a bad weed. In cultivated fields and waste places. July-
Sept. General. Naturalized from Europe.

4. Chaetochloa itMica (L.) Scrib. Italian Millet. An annual

grass with erect stems, 2-5 ft. high, and thick, compact, compound,

nodding panicles, interrupted at the base. Involucral bristles up¬
wardly barbed, usually purplish ; spikelets elliptic, shorter than the

322

OHIO BIOLOGICAL SURVEY

bristles; outer empty glume less than one-half as long as the spikelet ;*
fertile lemma only moderately convex, striate, finely and faintly trans¬
verse-rugose or pitted.

Several varieties are extensively cultivated for hay. If fed in
large quantities and too frequently, the hay is injurious to horses.
In cultivated fields and waste places. Also called German or Hun¬
garian millet. July-Sept. Rather general. Escaped from cultivation.

Pennisetum glaucum (L.) R. Br. Pearl-millet. A coarse annual grass,
with a solid stem, 3-8 ft. high, bearing cylindrical, spike-like panicles, 6-12 in.
long. Spikelets with 1 perfect flower and an additional vestigial lemma, sur¬
rounded at the base by a cluster of bristles. Empty glumes small and hyaline.
Cultivated for fodder and for the grain which is used as food. Other related
species are cultivated as ornamental grasses.

59. Cenchrus L. Sandbur-grass.

Annual or perennial grasses with flat leaves and spikate inflor¬
escence, the small clusters of spikelets covered by a prickly, bur-like
involucre which is deciduous with them at maturity. Prickles ret-
rosely barbed. Involucres with 2 or more spikelets. Spikelets 2-
flowered usually with one perfect flower and a second staminate or
vestigial flower; empty glumes shorter than the lemmas, unequal;
lemma and palet of the perfect flower chartaceous, the lemma not
inrolled at the margins ; grain free, inclosed in the glumes and the
prickly involucre.

Cenchrus tribuloides L. Sandbur-grass. An annual ascending
or trailing grass with flattened, much-branched solid stems, ^-3 ft.
long, and a stout spike-like raceme of prickly burs, often partly en¬
closed in the upper leaf sheath. Prickly involucres pubescent, the
prickles retrorsely barbed ; spikelets enclosed in the involucre ; fertile
lemma not inrolled at the margin ; somewhat indurated.

A disagreeable, troublesome weed. In sandy soil and cultivated
fields. July-Sept. Lucas, Wood, Ottawa, Erie, Lorain, Cuyahoga,
Eranklin, Highland, Gallia.

GRASSES OF OHIO

323

Tribe, Oryzeae. Rice Tribe.

Grasses ivith monosporangiatc or bisporangiatc, more or less laterally
compressed spikelets in loose panieles; raehilla artieulated helozv the
empty glumes zvhich are often minute or obsolete; stamens often 6;
hilum linear.

60. HomalocenchrUs Mieg. Cut-grass.

Perennial marsh grasses with flat narrow rough and cutting leaf-
blades and with paniculate inflorescence. Spikelets apparently 1-
flowered, strongly flattened laterally, those of the open part of the
panicle usually sterile, those inclosed in the sheath cleistogamous
and fruitful ; empty glumes none ; flowering glumes boat-shaped,
somewhat indurated, awnless; stamens 1-6; grain ovoid, free.

1. Branches of the rather simple panicle stiffly spreading; spikelets in. long

or less. H. virginicus.

1. Branches of the diffuse panicle generally lax; spikelets Y'Y i^i- lo^g-

H. oryzoides.

1. HomalocenchrUs virginicus (Willd.) Britt. Virginia Cut-
grass. A grass with branched, weak, ascending stems, 1-3 ft. long,
with clustered scaly rhizomes, and with simple panicles finally long-
exserted, the slender branches stiffly spreading and usually naked
below the middle. Lateral panicles usually included ; spikelets ob¬
long, appressed ; lemma hispid on the keel ; stamens 2.

In swamps and wet woods. Aug., Sept. Rather general.

2. HomalocenchrUs oryzoides (L.) Poll. Rice Cut-grass. A
grass with rather stout, branching stems, 1-4 ft. long, ascending from
a decumbent base with slender creeping rhizomes, with very rough
leaves and a diffusely branched, lax panicle with ascending or
spreading branches. Lateral panicles usually included. Spikelets
elliptic ; lemma hispid, strongly bristly ciliate on the keel.

In swamps and along ditches and streams. Aug., Sept. Rather
general.

61. Zizania L. Wild-rice.

A tall monecious hydrophytic annual grass with long flat leaves
and large terminal panicle, the upper part consisting of carpellate
spikelets, the lower of staminate, with some perfect spikelets be¬
tween the two regions. Spikelets on club-shaped pedicles, readily
deciduous, apparently 1-flowered, probably consisting of 2 coalescent
flowers ; empty glumes membranous, very minute or nearly obsolete.
Carpellate spikelet with a long-awned flowering glume and 6 vestigial

324

OHIO BIOLOGICAL SURVEY

stamens; staminate spikelet with 6 stamens and a vestigial gynecium,
its glumes short-awned or awn-pointed ; spikelets between the stam¬
inate and carpellate parts of the inflorescence more or less perfect,
with 6 stamens ; grain cylindrical, closely enveloped in the flower¬
ing glumes.

1. Zizania aquatica L. Wild Rice. A large annual grass with
an erect stem, 3-10 ft. high, broad flat leaves, and a large loose
panicle, the upper part carpellate with erect branches, the lower
staminate with ascending or spreading branches. Staminate spikelets
with acute awn-pointed glumes ; carpellate spikelets linear, the lem¬
ma long-awned ; staminate spikelet with 6 stamens and a vestigial
ovulary ; carpellate spikelet with 6 vestigial stamens.

The grain was used by the Indians for food. In swamps, often in
rather deep water. July, Aug. Erie, Licking, Perry, Hocking.

GRASSES OF OHIO

325

Tribe, Andropogoneae. Beard-grass Tribe.

Grasses usually having the spikelets in pairs on the branches of the
inflorescence, the one sessile and perfect, the other pediceled and perfect,
staniinate, or vestigial; often arranged in rows; empty glumes more
or less indurated, the dowering glumes hyaline; lemma of the perfect
flower usually awned; spikelets articulate belozv the empty glumes,
dor sally compress or round.

62. Holcus L. Sorghum.

Annual or perennial grasses with solid stems, long broad flat
leaves and large terminal panicles. Spikelets 1-flowered, in pairs at
the nodes or in 3’s at the ends, the one sessile and bisporangiate
the other pedicellate and staminate or entirely vestigial; empty
glumes of the sessile spikelet indurated and usually shining, ob¬
scurely nerved; flowering glumes hyaline, lemma awned. palet small
or sometimes absent ; grain free, more or less enclosed in the flower¬
ing glumes.

1. Panicle lax the branches with spikelets nearly to the base; stem usually not

over ii^- thick; perennial. H. halapense.

1. Panicle large and compact or with very long branches naked below; stem

usually y2-iy2 in. in diameter; annual. H. sorghum.

1. Holcus halapensis L. Johnson-grass. A coarse perennial
grass with solid, simple or sometimes branched stems, 3-5 ft. high,
and an open panicle, usually with whorled, spreading branches, naked
toward the base. Sessile spikelet, ovate-lanceolate, usually purplish,
pubescent with appressed hairs ; awn of the fertile lemma more or
less bent ; pedicellate spikelets with membranous glumes, sometimes
enclosing a staminate flower but usually vestigial.

Cultivated as a forage grass. Escaped. In cultivated fields and
waste places. A troublesome weed in the South. July-Sept. Native
of Europe. Cuyahoga, Erie, Eranklin, Madison.

2. Holcus sorghum L. Common Sorghum. A coarse annual
grass with thick solid stems, 5-12 ft. high, usually with rather dense,
much-branched panicles, but in some varieties with the naked part
of the branches greatly elongated. Sessile spikelets ovoid or ellip¬
tical, with a large usually exposed grain ; pediceled spikelets vestigial.

Cultivated in many forms for fodder, grain, syrup, and broom-
corn under various names as sorghum, durra. Kafir-corn, broom-corn,
feterita, milo-maize, etc. July-Sept. Volunteer in Adams County.

Siidan-grass is an annual slender-stemmed grass, 4-10 ft. high, recently
introduced as a forage crop. It is without rhizomes and is apparently a variety
of the common sorghum.

326

OHIO BIOLOGICAL SURVEY

63. Sorghastrum Nash. Indian-grass.

Tall stout perennial grasses with hollow stems, long narrow flat
leaves, and terminal open panicles. Spikelets in pairs, the sessile
spikelet with 1 perfect flower, the pedicellate spikelet vestigial; empty
glumes of the sessile spikelet indurated and shining, flowering glumes
hyaline, the lemma usually awned ; palet sometimes wanting; grain
free.

1. Sorghastrum nutans (L.) Nash. Indian-grass. A large grass
with simple, erect hollow stems, 3-8 ft. high, and a narrowly oblong
panicle with slender, erect or spreading branches. Spikelets lanceo¬
late, yellowish or reddish brown and shining, clothed especially
toward the base with fawn-colored hairs, at length drooping; lemma
with a twisted awn.

A valuable pasture and hay grass. In dry soil especially prairies.
Aug., Sept. Ashtabula, Cuyahoga, Erie, Wyandot, Auglaize, Frank¬
lin, Madison, Adams.

Erianthus ravennae (L.) Beauv. Phime-grass. A robust perennial grass,
with solid stems. 5-10 ft. high, with a plume-like, silky panicle, 1-2 ft. long.
Spikelets with 1 perfect flower and a sterile lemma, in pairs, one sessile, the other
pedicellate, with a ring of long hairs at the base. Cultivated for ornament.

64. Miscanthus Anderss. Plume-grass.

Tall erect perennials with flat leaves and ample terminal, com¬
monly hairy panicles. Spikelets in pairs unequally pedicellate, with
1 perfect flower ; empty glumes membranous, blunt at the apex ;
sterile lemma thinner ; fertile lemma thinly hyaline, 2-toothed at the
apex and awned ; palet thin, hyaline ; grain free.

1. Miscanthus sinensis Anderss. Chinese Plume-grass. A
large perennial grass with stems 3-9 ft. high, with long, slender
leaves and a panicle of numerous erect or ascending branches.
Spikelets yellowish brown, surrounded at the base with long, white
or purplish hairs ; lemma hyaline with a spirally twisted awn aris¬
ing from between the two teeth of the apex.

The variety with banded leaves is commonly cultivated for
ornament in gardens and parks and occasionally escapes into waste
places. Aug., Sept. An escape in Lake County.

65. Andropogon L. Beard-grass, Bluestem.

Tall usually tufted perennials with narrow leaves and terminal
and axillary racemes consisting of spike-like branches often more
or less digitately arranged and with long silky hairs on the rachis
and pedicels. Spikelets in pairs at each node of the jointed rachis
one sessile and bisporangiate, the other pedicellate and either stam-

GRASSES OF OHIO

327

inate or vestigial ; empty glumes of the sessile spikelet subequal,
indurated, the outer one dorsally flattened with a strong nerve near
each margin, the inner one keeled above; first lemma hyaline; fertile
lemma membranous or hyaline, awned, its palet hyaline, sometimes
obsolete; grain free.

1. Inflorescence of 2 or more racemose branches ; joints of the rachis not
clavate. 2

1. Raceme simple; joints of the rachis clavate, with a cup-shaped depression at

the top. A. sc Oparins.

2. Raceme branches 2-6 on a long peduncle ; rachis with short inconspicuous

hairs ; tall coarse grasses. A. fnrcatus.

2. Raceme branches 2 or 3, subtended by a foliaceous sheath ; rachis covered
with long white hairs. A. virginicus.

1. Andropogon furcatus Muhl. Big Bluestem. A large grass
with solid, robust stems simple at the base but branching from the
upper nodes, S-Sj/s ft. high, with a sweetish sap and with a more or
less digitate panicle of 2-6 spike-like racemes, usually purplish.
Rachis joints and pedicels hairy on the sides and at the summit;
pedicellate spikelet staminate; sessile spikelet perfect; fertile lemma
with a delicate awn.

A characteristic and important prairie grass valuable for hay
and pasture. In dry or moist soil. Aug., Sept. Rather general.

2. Andropogon virgmicus L. Virginia Beard-grass. A grass
with rather slender stems sparingly branched above, 1^-4 ft. high,
and with a loose and elongated inflorescence of 2-4 spike-like racemes
protruding from the sides of the sheaths. Hairs of the inflorescence
long and silky ; sessile spikelet perfect, its lemma with a nearly
straight awn ; pedicellate spikelet vestigial.

In dry or moist open fields and hillsides. Aug., Sept. Gallia,
Jackson, Meigs, Athens, Vinton, Hocking, Fairfield, Belmont.

3. Andropogon scoparius Mx. Little Bluestem. (Schizachyrium
scoparium (Mx.) Nash.) A tufted grass with rather slender and
stiff, simple or branched stems, 1-4^2 ft. high, and with loose racemes
on long-exserted, slender peduncles. Joints of the rachis and pedicels
ciliate with spreading hairs; sessile spikelet perfect, the fertile lemma
with a bent and twisted awn ; pedicellate spikelet reduced to a single
awn-pointed glume.

An important hay and forage grass in the West. On dry prairies
and in sandy fields. Aug.-Oct. Rather general.

328

OHIO BIOLOGICAL SURVEY

Tribe, Maydeae. Corn Tribe.

Monecioiis grasses having the staniinate and carpellate spikelets in
different inflorescences or in different parts of the same inflorescence,
often with highly specialized structures; spikelets articulated below the
empty glumes, round or dorsally compressed, in pairs, normally 2-
flowered, some of the flowers usually more or less vestigial or obsolete
and the staminate and carpellate flozvers usually showing vestiges of
the opposite organs.

66. Tripsacum L. Gama-grass.

Tall stout monecious perennials with solid stems, thick creeping rhizomes,
broad flat leaves, and a panicle of spikate or racemose branches, the joints of
the fruiting rachis separating at the nodes, each joint containing a grain and the
remains of the spikelet, the upper part of the inflorescence staminate, lower part
carpellate. Staminate spikelets in pairs, the flowers containing vestigial ovularies ;
carpellate spikelet solitary imbedded in the rachis joint, with one carpellate flower
which contains vestigial stamens and a vestigial flower with only a vestigial
ovulary; outer empty glume of the carpellate flower large and cartilaginous, the
inner coriaceous ; flowering glumes hyaline; empty glumes of the staminate
spikelets subcoriaceous, the flowering glumes membranous or hyaline ; grain
free enclosed in the pocket of the rachis joint and the glumes.

Tripsacum dactyloides L. Gama-grass. A monecious, perennial grass
with thick rhizomes and solid stems, 1-7 ft. high. Inflorescence consisting of a
number of spike-like racemes with paired staminate spikelets above and single
carpellate spikelets below. Spikelets 2-flowered, the carpellate spikelet, with one
carpellate and one vestigial flower, contained in the deeply channeled joints of
the rachis which separate spontaneously at maturity, by cleavage planes.

Euchlaena mexicana Schrad. Teosinte. A monecious annual grass, 8-15
ft. high, of much value for forage in the south. The carpellate inflorescence
is surrounded by a peculiar husk, and the grains are enclosed in the flattened
rachis joints much as in Tripsacum.

67. Coix L. Job’s-tears.

Coarse much-branched annuals with large broad leaves and a
monecious inflorescence, the staminate spikelets covered by a modi¬
fied leaf-sheath which becomes bony and bead-like in fruit, the stam¬
inate inflorescence projecting thru this sheath. Staminate spikelets
2-flowered, the flowers with vestigial ovularies; carpellate spikelets
2-flowered, the outer flower entirely vestigial with a lemma, the inner
containing a normal gynecium and 3 vestigial stamens ; two or more
vestigial structures usually present in the sheath besides the carpel¬
late spikelet ; empty glumes thick-membranous ; flowering glumes
hyaline ; grain free, enclosed in the glumes and the bony sheath.

1. Coix lacryma-jobi L. Job’s-tears. A much-branched annual
grass with rather thick stems, 2-4 ft. high, and a pecular inflores-

GRASSES OF OHIO

329

cence, the carpellate spikelet covered by a hardening bead-like sheath,
bluish-gray in color and shining when ripe, the staminate inflores¬
cence small and projecting on a slender pedicel from the bead-like
sheath.

Grown for the ornamental fruits which are used as beads and
for rosaries. In waste places around gardens. July-Sept. Persistent
in Franklin County.

68. Zea L. Indian-corn.

Tall coarse annual grasses with solid stems, large broad flat
leaves, prominent prop roots, and a monecious inflorescence, the
staminate spikelets in an ample panicle at the summit of the stem,
and the carpellate spikelets on an ear covered with husks at the
side of the main stem. Staminate and carpellate, spikelets sometimes
in the same panicle. Staminate spikelets 2-flowered, in pairs, one
sessile and the other pediceled ; carpellate spikelets always in double
rows on the ear, usually 2-flowered, the outer flower vestigial ; style
of the gynecium a very long ‘‘silk” ; empty glumes thick-membranous
or coriaceous ; flowering glumes hyaline ; grain large, free.

1. Zea mays L. Indian-corn. A tall, coarse, annual grass
with broad leaves and a thick, solid, erect stem, 3-15 ft. high, de¬
veloping prominent prop-roots at the basal joints, an ample staminate
panicle at the top, and one or more peculiar, compact, carpellate in¬
florescences at the side, called ears, covered with peculiar leaf-
sheaths or husks ; grain large, exposed, with a very long style or silk.

Extensively cultivated in many varieties for grain, green corn,
and fodder. Many products are made from the grain. The stigmas
of the silk are used in medicine. In fields and waste places. June-
Sept. Also called maize. Spontaneous in Brown, Adams, Scioto,
Fayette, Monroe, Erie, and Hancock Counties.

THE LISRARY OF THE

NOV 22 1938

university of ilunois

INDEX

Agropyron .

Agrostideae .

Agrostis .

Apolecuriis .

Ammophila .

Andropogoneae .

Andropogon .

Anthroxanthum

Apera .

Aristida .

Arrhenatherum ..

Aspris .

Atheropogon .

Avena .

Aveneae .

Barley . ......

Barnyard-grass ...

Beach-grass .

Beard-grass .

Beckmannia .

Bent-grass .

Bermuda-grass ...

Blue-grass . .

Blue-stem .

Bottle-brush-grass

Bouteloua .

Brachyelytrum ...

Brome-grass .

Bromus .

Broom-corn .

Calamagrostis .

Canary-grass . .

Capriola . .

Cat-tail-grass .

Cenchrus . .

Chaetochloa . .

Cheat .

Chess .

Chlorideae .

Chloris .

Cinna .

Cockspur-grass ....

Page

. 291

. 299

. 301

. 303

. 302

. 325

. 326

. 310

. 302

. 307

. 287

. 289

. 297

. 288

. 287

,291, 294

. 319

. 302

325, 326

. 296

299, 301

. 297

. 278

. 327

. 294

. 298

. 305

. 272

. 272

. 325

. 300

. 309

. 297

. 303

. 322

. 320

. 273

. 272

. 296

. 296

. 302

. 320

Page

Coix . 328

Corn . 328

Cortaderia . 286

Couch-grass . 291

Crab-grass . 318

Cut-grass . 323

Cynosurus . 286

Dactylis . 281

Danthonia . 287

Darnel . 291

Deschampsia . 289

Dogtail-grass . 286

Dropseed . 299

Durra . 325

Eaton-grass . 283

Echinochloa . 319

Eleusine . 297

Elymus . 293

Eragrostis . 281

Erinanthus . 326

Euchlaena . 328

Ealse-oats . 288

Eescue-grass . 272, 275

Eestuceae . . . 272

Eestuca . 275

Eeterita . 325

Eoxtail . 303

Foxtail-grass . 320

Gama-grass . 328

Grama-grass . 297

Graminaceae . 257

Hair-grass . 289

Heleochloa . 303

Holcus . 325

Homalocenchrus . 323

Hordeae . 291

Hordeum . 294

Hystrix . 294

Indian-grass . 326

Indian-corn . 329

Johnson-grass . 325

Job’s-tears . 328

Kafir-corn . 325

INDE X- - - Continued

Koeler-grass .

Koeleria .

Korycarpus .

Leptoloma .

Lolium .

Love-grass .

Maize .

Maydeae . .

Manna-grass ..
Meadow-grass

Melica .

Melic-grass .

Mesquite-grass

Milium .

Millet .

Millet-grass ... .

Milo-maize .

Miscanthiis .

Mountain-rice .
Muhlenbergia ..
Nothoholcus ...

Oats .

Oat-grass .

Orchard-grass

Oryzeae .

Oryzopsis .

Pampas-grass .
Panic-grass .....

Paniceae .

Panicum . .

Panictilaria . .

Paspalum .

Pearl-millet .

Pennisetum .

Phalarideae .

Phalaris .

Phleum .

Plume-grass .

Poa .

Porcupine-grass
Poverty-grass ...

Phragmites .

Purple-top .

Ray-grass .

Red-top .

Page

. 284

. 284

. 284

. 318

. 291

. 281

. 328

. 328

. 276

. 278

. 275

. 275

. 298

. 306

. 321

. 306

. 325

. 326

. 306

. 304

. 290

287, 288

. 287

. . 281

. 323

. 306

. 286

. 311

. 311

. 311

. 276

. 320

. 322

. 322

. 309

. 309

. 304

. 326

. 278

. 306

. 307

.........286

. 285

. 291

. 301

Page

Reed Bent-grass . 300

Reed-grass . 286

Rice . 323

Rush-grass . 299

Rye . 292

Rye-grass . 291

Sandbur-grass . 322

Sand-grass . 285

Savastana . 309

Schizachyrium . 327

Secale . 292

Slough-grass . 296

Sorghastrum . 326

Sorghum . 325

Spartina . 296

Spear-grass . 278

Sphenopholis . 283

Sporobolus . 299

Stink-grass . 282

Stipa . 306

Sudan-grass . 325

Syntherisma . 318

Teosinte . 328

Timothy . 304

Tridens . 285

Triplasis . 285

Triple-awn-grass . 307

Tripsacum . 328

Trisetum . 288

Triticum . 292

Vanilla-grass . 309

Velvet-grass . 290

Vernal-grass . 310

Wheat . 292

Wheat-grass . 291

Wild-oat-grass . 287

Wild-rice . 323

Wild-rye . 293

Windlestraw . 302

Witch-grass . 318

Wood Reed-grass . 302

Yard-grass . 297

Zea . 329

Zizania . 323

Volume II, No. 6

Bulletin No. 1 0

OHIO BIOLOGICAL SURVEY

THE ASCOMYCETES OF OHIO IV

THE LECIDEACEAE

By BRUCE FINK

(wiveasmr of umis

1923

THE ASCOMYCETES OF OHIO V

THE PELTIGERACEAE

By LEAFY J. CORRINGTON

Published by

THE OHIO STATE UNIVERSITY
Columbus,

1921

1 O B S

THE ASCOMYCETES OF OHIO IV*

The Lecideaceae.

Bruce Fink.

GENERAL CONSIDERATIONS

It was stated in the second paper of this series that the disposition of
the Lecideaceae in an early paper of the series would show what slight
changes are needed in treating lichens as we treat other ascomycetes. It
is hoped that this paper has accomplished this in phraseology intelligible
to those acquainted with the present-day language of systematic mycology.

The Lecideaceae form a well-defined family of lichens, the affinities of
which seem plainly marked. In apothecial structure, and so far as known,
in structure of the sexual reproductive areas, the family seems to be
closely related to the mainly non-lichen Patellariaceae and to such lichens
as the Gyalectaceae, the Lecanactidaceae, the Collemaceae, the Baeomycetaceae,
and the Cladoniaceae.

Following the commonly-accepted theory that the lichens have been
evolved from non-algicolous fungi, the origin of the Lecideaceae and
related lichens from Patellaria-\\ke ancestors is a reasonable supposition,
though the relative rank of the various related families named in the last
paragraph is not easy to decide. Within the Lecideaceae, the line of evolu¬
tion seems to have been in the direction of a well-developed exciple and
from simpler to more complex spores. With the advance in these two
directions has gone a slightly increased development of the tballus.

In structure, the thallus is crustose, and the thalli vary from inconspic¬
uous, evanescent conditions to those which are conspicuous and sometimes
even subsquamulous. Rarely the thallus extends upward as a veil which
surrounds the apothecia laterally and suggests how the thalloid exciple of
higher families probably arose. As usual in crustose forms, the thalli are
composed of hyphae which are densely disposed toward the upper, exposed
surface and more loosely disposed toward the lower surface (Fig. 2).

The apothecial evolution passes Trom forms with weak, light-colored
exciples and soft texture (Fig. 10) to those with strong, dark exciples,
which are firm in texture (Fig. 11). The superficial apothecial characters
are so much alike in many of the species that one cannot always feel
certain even of the genus of unfamiliar forms until he has studied them
microscopically.

•Contributions from the Botanical Laboratory of Miami University. — XVIII

334

335

5 ”7 o.sr

0 H

Wb. 10,

THE ASCOMYCETES OF OHIO IV

The paraphyses are commonly distinct in young apothecia, but in
mature apothecia they are usually more or less gelatinized and coherent.
In some of the species, they become so gelatinized that they form a
homogeneous mass about the asci, in which the individual paraphyses are
no longer discernible. When distinct, the paraphyses are sometimes
branched, most commonly toward their apices (Fig. 1 and 12).

There is great diversity with respect to spore development, the whole
range of spore structure, from minute, simple, hyaline spores to those
which are large, brown, and muriform being found within the family
(Figs. 3, 4, 5, 6, 7, 8, 9, and 13). This condition makes it appear quite
possible that the family may be polygenetic.

The genus, Biatorella, contains non-lichen forms and is probably as a
whole more closely related to the Patellariaceae than to the Lecidiaceae.
However, our two species, both of which are lichens, are herein admitted
to the latter family. Through one or more species with larger spores
than are usually found in this genus, Biatorella approaches Lecidea.
Starting with Lecidea, we have a natural series in spore development with
intermediate conditions difficult to place The series runs thus: Lecidea
with simple hyaline spores (Fig. 3); Biatorina with two-celled, hyaline
spores (Fig. 4); Bilimbia with several-celled, hyaline spores, not much
narrowed (Fig. 5); ^xvA Bacidia with several-celled, hyaline, acicular spores
(Fig. 6). Buellia and Rhizocarpon are aberrant genera, brown-spored, and
closely related among themselves (Figs. 8, 9, and 13). Through Buellia,
the two genera are related to Rinodina of the Physciaceae. The two aber¬
rant genera are like other members of the Lecideaceae with respect to thallus
development and general apothecial characters, the aberrancy being with
respect to the spores, on which account the two genera are placed in an¬
other family, the Buelliaceae, by some workers, perhaps with sufficient reason .

The algal host is Pleurococcus-\\]^e (Fig. 2, c) in nearly all species of
the Lecideaceae; but the host cells are so hypertrophied and distorted that
their generic rank is often difficult to ascertain, except by cultivation out¬
side of the lichen thallus. The algal-host cells are few in number in some
of the species and are sometimes absent during a portion of the life history
of the lichen. The host is usually found throughout the superficial portions
of the thallus, except near the upper surface, from which portion the algae
are usually absent, except in a dead or dying condition, difficult to detect.

The writer has collected the Lecideaceae, with other fungi, in Butler
County for fifteen years, and has worked for the Ohio Biological Survey in
Preble, Warren, Highland, Fairfield, Adams, Hocking, and Lake counties.
Besides these collections made by the writer, a few specimens were examined
from Champaign, Hamilton, Wayne, Morgan, Madison, Muskingum, Frank¬
lin, Vinton, and Summit counties. Of the 37 species treated in this paper,
24 had not been reported from Ohio previously.

336

OHIO BIOLOGICAL SURVEY

Systematic Account.

LECIDEACEAE

Thallus crustose, without plectenchymatous cortex (Fig. 2, a), vary¬
ing from granulose and often evanescent to conspicuous, areolate, or even
subsquamulose conditions, attached to the substratum by hyphal rhizoids
(Fig. 2, d), and in a few instances extending up as a veil and surrounding
the apothecia laterally, the hyphae densely interwoven toward the upper
surface, but more loosely disposed below (Fig. 2, a and b); apothecia usu¬
ally minute or small, commonly rounded, the exciple weak and obscure
(Fig. 10, d), or more strongly developed when conspicuous and much darker
in color (Fig. 11, b); hypothecium varying from hyaline to dark brown
(Fig. 10, band Fig. 11, c); hymenium almost always lighter and commonly
hyaline (Figs. 10 and 11, a); paraphyses usually simple, but branched forms
to be found frequently (Figs. 1 and 12), pale throughout or darkened
toward the sometimes enlarged apex, commonly more or less coherent and
indistinct at maturity; spores simple and hyaline to muriform and brown
(Figs. 2, 3, 4, 5, 6, 7, 8, 9, and 13).

KEY TO THE GENERA

Spores minute, numerous in each ascus . . . . Biatorella, p.

Spores larger, usually 8 in each ascus,

Spores hyaline.

Spores one-celled (simple) . Lecidea, p.

Spores more than one-celled (compound).

Spores 2-celled . Biatorina, p.

Spores 4- to several-celled.

Spores ellipsoid, fusiform, or dactyloid . Bilimbia, p.

Spores acicular . Bacidia, p.

Spores brown, or becoming brown.

Spores 2-celled . Buellia, p.

Spores 4-celled and becoming muriform . Rhizocarpon, p.

Biatorella De Not. Giorn. Bot. Ital. 21. 192. 1846.

Thallus granulose to verrucose and subareolate, sometimes inconspicu¬
ous and evanescent; apothecia minute to middle-sized, adnate or more or
less immersed, exciple usually prominent and persistent, but sometimes
becoming covered, disk flat to convex; hypothecium and hymenium pale
to brown; spores simple, hyaline, minute, numerous in each ascus.

KEY TO THE SPECIES OF BIATORELLA

The whole apothecium dark colored . 1. B. simplex

The disk of the apothecium white-pruinose . . . 2. B. pruinosa

1 Biatorella simplex (Dav.) Br. & Rostr. Bot. Tidssk. 3: 241 1869.
Lichen simplex Dav. Trans. Linn. Soc. Lond. 2: 283. pi. 28. f. 2. 1794.
Thallus thin and smooth or thicker and roughened, sometimes subar¬
eolate, ash-white to green-gray and darkening, rarely disappearing; apothecia
minute to middle-sized, 0.2 to 0.8 mm. in diameter, adnate, scattered or

THE ASCOMYCETES OF OHIO IV

337

crowded, rounded or variously irregular, black but usually dark red when
damp, flat or slightly convex, the thin exciple raised and persistent;
hypothecium light or darker brown; hymenium pale or tinged brown;
paraphyses semi-distinct to coherent-indistinct; asci cylindrico-clavate;
spores oblong-ellipsoid, 3 to 5 mic. long and 1 to 1.5 mic. wide.

Collected in Butler, Adams, Montgomery, Hocking, and Ross counties.
On limestone. Not previously reported from Ohio, but probably frequent
where there is limestone, though inconspicuous and easily overlooked.

2. Biatorella pruinosa ( J. E. Smith) Mudd Man. Brit. Lich. 191. 1861.

Lichen pruinosus J. E. Smith in Sowerby, Eng. Bot. 32: pi. 2244 1811.

Thallus light colored, usually thin and smooth, rarely disappearing;
apothecia minute to middle-sized, 0. 2 to 1 mm. in diameter, adnate
scattered or crowded, flat or slightly convex, the disk pruinose, and the
exciple persistent; hypothecium lighter or darker brown; hymenium usually
pale; paraphyses coherent and becoming indistinct; asci cylindrico-clavate;
spores oblong-ellipsoid, 3 to 5 mic. long and 1 to 1.5 mic. wide.

Collected in Butler and Adams counties. On limestone. Not pre¬
viously reported from Ohio, but often occurring with the last in limestone
areas.

Lecidea Ach. Meth. Lich. XXX. 32. pi. 2. f. 1, 2. 1803.

Thallus smooth, roughened, or verrucose, in some species chinky to
areolate, or even subsquamulose, rarely rudimentary and evanescent;
apothecia minute to middle-sized, usually adnate, but rarely sessile or
immersed, with' pale to black, and flat to strongly convex disk; exciple and
hypothecium from pale to dark brown in section; hymenium lighter,
most commonly pale; spores simple, hyaline, 8 in each ascus.

KEY TO THE SPECIES OF LECIDEA
Exciple soft, usually light colored.

Apothecia usually surrounded by a thalloid veil . coarciata

Apothecia not surrounded by a thalloid veil.

Exciple becoming covered.

Hypothecium pale or pale yellow.

Apothecia always minute.

Spores 5 to 7 mic. long . 2. L. intropallida

Spores 7 to 15 mic. long . 3. L. varians

Apothecia reaching middle size . 4. L. rupestris

Hypothecium light-brown to dark brown.

Thallus gray -green or lighter . 5. L. viridescens

Thallus darker from the first or becoming so.

Thallus minute and evanescent . 6. L. humicola

Thallus well developed and persistent.

Thallus of raised granules . 7. L. uliginosa

Thallus of flat granules . . 8. L. salvicola

Exciple persistent . 9. L. flexuosa

Exciple horny, dark colored.

Disk usually convex, commonly on wood . 10. L. enteroleuca

Disk flat or less commonly convex, on rocks.

Disk usually white-to rusty-green-pruinose . 11. L. albocaerulescens

Disk black, scarcely pruinose. . 12. L. platycarpa

338

OHIO BIOLOGICAL SURVEY

1. Lecidea coarctata (J. E. Smith) Nyl. Act. Soc. Linn. Bord. 21: 358. 1856.

Lichen coarctatus J. E. Smith in Sowerby, Eng. Bot. 8: pi. 534. 1789.

Thallus of minute, scattered or clustered, rounded, angular, or min¬
utely and irregulary crenate, green-gray, pale brown, or more commonly
ash-white granules, sometimes passing into a subcontinuous, chinky or are-
olate crust; apothecia minute to small, 0.2 to 0.4 mm. in diameter, adnate,
from flesh-colored to black, commonly concave or flat, sometimes difform,
frequently surrounded laterally by a thalloid veil; hypothecium and
hymenium pale to pale brown; paraphyses distinct; asci clavate or cylindri-
co-clavate; spores ovoid to ellipsoid, 13 to 23 mic. long and 7 to 10 mic.
wide.

Collected in Lake, Ross, Hocking, and Preble counties. Also exam¬
ined from Lawrence County. On rocks and old bricks. Not previously
reported from Ohio. Widely distributed in the State, but rare, except in
Lake County, where this fungus was unusually common.

2. Lecidea intropallida sp. nov.

Thallus a continuous, smooth or slightly roughened, ash-gray and dark¬
ening crust; apothecia minute, 0.15 to 0.25 mm. in diameter, adnate or
partly immersed, flesh-colored to yellow-brown, flat to slightly convex, the
concolorous and inconspicuous exciple soon covered; hypothecium and
hymenium pale; paraphyses sometimes distinct, but more commonly coher¬
ent-indistinct; asci clavate; spores simple, hyaline, ellipsoid, 5 to 7 mic. long
and 2.5 to 3 mic. wide.

Collected near Painesville in Lake County. On pebbles in a moist wood.
The type specimen is deposited in the writer’s herbarium, and a cotype
may be seen in the State Herbarium.

3. Lecidea varians Ach. Syn. Meth. Lich. 38. 1914.

Thallus of very minute, raised or flattened, green-gray to yellow-green
granules, these forming a thin but continuous, smooth or granulate-
rugose, often chinky crust, usually bordered and often decussated by black
lines; apothecia minute, 0.12. to 0.25 mm. in diameter, often clustered
or even conglomerate, adnate, from pale yellow to brown and Anally black,
flat with a thin exciple to convex with covered exciple; hypothecium pale
to pale yellow; hymenium pale below, but often yellow or blue-violet above;
paraphyses usually coherent, distinct or indistinct; asci clavate; spores
oviod-ellipsoid, 7 to 15 mic. long and 5 to 7. 5 mic. wide.

Collected in Adams County. On maple bark. Also reported from
Franklin County. The plant is so minute and inconspicuous as to be
very difficult to detect and is probably distributed widely in the State.

4. Lecidea rupestris (Scop.) Ach. Meth. Lich. 70. 1803. (See Fig, 10),

Lichen rupestris Scop. FI. Cam. ed. 2. 2: 363, 364. 1772.

Thallus a continuous, moderately thick, smooth or more or less
roughened, often chinky to subareolate, ash-gray, yellow-green, or darken-

THE ASCOMYCETES OF OHIO IV

339

ing crust; apothecia small to large, 0.4 to 1.3 mm. in diameter, at first im¬
mersed but becoming adnate, yellow to yellow or red-brown, flat to strongly
convex and the exciple covered; hypothecium pale or pale yellow; hyme-
nium pale; paraphyses coherent, semi-distinct to indistinct; asci clavate;
spores ellipsoid, 10 to 15 mic. long and 5 to 7 mic. wide.

Collected in Adams Country. On calcareous rocks. Not previously
reported from North America.

5. Lecidea viridescens (Schrad.) Ach. Meth. Lich. 62. 1903.

Lichen viridescens Schrad. Spic. Fi. Germ. 88. 1794.

Thallus of very minute, smooth or delequescent and powdery, ash-
grey to grey-green granules, spread over the substratum as a thin or rarely
thicker crust; apothecia minute to small, 0.2 to 0.5 mm. in diameter,
adnate, frequently clustered or even conglomerate, becoming black, from
flat with the thin livid or darker exciple visible to convex with the exciple
covered; hypothecium pale or darker brown; hymenium pale to pale
brown; paraphyses coherent, semi-distinct to indistinct; asci clavate; spores
oblong-ellipsoid, 9 to 12 mic. long and 4 to 5.5 mic. wide.

Collected on Little Mountain in Lake County, and in Hocking County.
On logs in woods. Not previously reported from Ohio, and probably rare
in the State.

6. Lecidea humicola (Ach.) comb. nov.

Lecidea uliginosa humicola Ach. Meth. Lich. 43. 1903.

Thallus of very minute inconspicuous and evanescent, brown-black
granules; apothecia minute, 0.2 to 0.4 mm. in diameter, adnate, dark brown
to black, scattered or clustered, plain with a thin concolorous exciple vis¬
ible, to convex with the exciple finally covered; hypothecium dark brown;
hym.enium pale brown; asci clavate; paraphyses coherent-indistinct; spores
oblong-ellipsoid, 9 to 15 mic. long and 5 to 7 mic. wide.

Collected in Hocking County. On soil in a moist wood. Not pre¬
viously reported from North America.

7. Lecidea uliginosa (Schrad.) Ach. Meth. Lich. 43. 1803.

Lichen uliginosus Schrad. Spic. FI. Germ. 88. 1794.

Thallus of scattered, clustered, or even heaped, irregular and minute,
green-olive to rust-brown, or even brown-black, somewhat raised and rarely
coralloid granules, these forming a scattered or continuous crust; apothecia
minute to small, 0.2 to 0.4 mm. in diameter, closely adnate or more or less
immersed, often clustered, brown to black-brown, flat with the thin lighter-
colored or black exciple visible, or becoming strongly convex, with the
exciple finally covered; hypothecium light or darker brown; hymenium
tinged yellow or brown; paraphyses closely coherent, but usually remaining
distinct; asci long-clavate; spores oblong-ellipsoid, 8 to 14 mic. long and 4
to 7 mic. wide.

340 OHIO BIOLOGICAL SURVEY

Collected in Preble, Butler, Warren, Adams, Fairfield, and Lake coun¬
ties. On dead wood. Widely distributed in Ohio.

8. Lecidea sylvicola Koerb. Syst. Lich. 254. 1855.

Thallus of minute, irregular, somewhat flattened or more rarely hemis¬
pherical, green-gray, olive-brown, or darker granules, these forming a thin,
continuous, or rarely scattered, subleprose, verrucose, or even subareolate,
wide-spread crust; apothecia minute to small, 0.2 to 0.5 mm. in diameter,
adnate or rarely more or less immersed, dark brown to black, flat to con¬
vex, the black exciple soon becoming covered; hypothecium brown to
black-brown; hymenium pale or tinged brown; paraphyses coherent, semi-
distinct to indistinct; asci clavate; spores ellipsoid, 5 to 9 mic. long and 2.5
to 4 mic. wide.

Collected in Lake, Ross, Preble, Hocking, and Butler counties. On
various rocks. Not previously reported from Ohio, and apparently new to
America under this name. Widely distributed in Ohio.

For possible relationship to Lecidea myriocarpoides Nyl. See “The
Lichens of Minnesota” (Cont. Nat. Herb. 14: 74. 1910).

9. Lecidea flexuosa (Fr.) Nyl. Act. Soc. Linn. Bord. 21. 356. 1856.

Biatora flexuosa Fr. Vet. Akad. Handl. 1822: 267. 1822.

Thallus of small or minute, flattened or rugose, scattered or clustered,
ash-grey to green-gray granules, these bursting into sorediate heaps, or
forming a moderately thick, areolate crust; apothecia minute to small, 0.2
to 0.4 mm. in diameter, adnate, black, and flat, the thin, livid or darker,
persistent exciple becoming flexuous; hypothecium pale or darker brown;
hymenium tinged brown; paraphyses coherent, semi-distinct to indistinct;
asci cylindrico-clavate; spores oblong-ellipsoid, 5 to 10 mic. long and 3 to
5 mic. wide.

Collected in Preble, Adams, Ross, and Butler counties. On dead wood.
Not previously reported from Ohio, and rare, though probably distributed
widely in the State.

The spores are slightly below normal size in our specimens.

10. Lecidea enteroleuca Ach. Lich. Univ. 177. 1810.

Thallus thin or becoming moderately thick, smooth or more often
granulate, chinky or areolate, the granules or verrucae rarely becoming
heaped in the thicker forms, ash-to green-gray, occurring in rounded areas,
or irregularly and often widely spread over the substratum; apothecia
minute to middle-sized, 0.35 to 1.2 mm. in diameter, adnate, black, flat to
more commonly convex, the frequently flexuous exciple often becoming
covered; hypothecium pale to dark brown; hymenium pale below, but
usually more or less colored above; paraphyses distinct, but often more or
less coherent; asci clavate; spores ovoid-ellipsoid, 8 to 17 mic. long and
5 to 9 mic. wide (Fig. 3).

THE ASCOMYCETES OF OHIO IV

341

Collected in Lake, Adams, and Hocking counties. On bark and rocks.
Not previously reported from Ohio. Rare, but doubtless distributed widely
in the State.

11. Lecidea albocaerulescens (Wulf.) Schaer. Lich. Helv. Spic. 3: 142.
1828.

Lichen albocaerulescens^ vM.m Jacq. Coll. Bot. 2: 184. pi. 5. f. 1. 1788.

Thallus smooth or somewhat rough, more or less chinky or becoming
obscurely small-areolate, ash- to green-gray, or becoming olivaceous, spread¬
ing over the substratum as a continuous, moderately thick crust; apothecia
small to large, 0.5 to 1.5 mm. in diameter, adnate or more or less immers¬
ed, usually flat, almost always white or rusty-green pruinose, the black
exciple rarely becoming covered; hypothecium brown to black-brown;
hymenium commonly pale; paraphyses distinct, but usually coherent; asci
clavate to inflated-clavate; spores ovoid-ellipsoid, 15 to 24 mic. long and 7
to 10 mic. wide.

Collected in Preble, Hocking, and Lake counties. Also examined from
Lawrence County. On rocks other than calcareous. Not previously
reported from Ohio. Rare, but apparently distributed widely in the State.

12. Lecidea platycarpa Ach. Lich. Univ. 173. pi. 2. f. 5. 1810.

Thallus a thin, obscurely or more or less plainly roughened, usually

chinky to subareolate, ash- to green-gray, continuous or more or less scat¬
tered, sometimes disappearing crust; apothecia small to middle-sized or
even larger, 0.4 to 1.5 mm. in diameter, commonly scattered, brown-black
to black, rarely and obscurely white-pruinose, adnate to sessile, rounded to
flexuous, flat or finally convex, the raised exciple sometimes becoming
covered; hypothecium dark brown; hymenium pale below and colored above;
paraphyses distinct or coherent-semidistinct; asci clavate; spores ovoid-to
oblong-ellipsoid, 14 to 20 mic. long and 6 to 10 mic. wide.

Collected in Ross and Hocking Counties. On rocks. Not previously
reported from Ohio.

Biatorina Mass. Ric. Lich. 134. f. 262-271. 1852.

Thallus commonly granulose, and often passing into verrucose and
chinky conditions, but scarcely ever areolate, sometimes scant and evane¬
scent; apothecia usually minute or small, and commonly adnate, exciple
weak and often becoming covered; hypothecium and hymenium passing
from pale through shades of brown, the former becoming darker than the
latter, this rarely tinged blue or violet above; spores hyaline, 2-celled.

KEY TO THE SPECIES OF BIATORINA

Growing on another lichen . . 1. B. heerii

Growing on wood or on rocks.

On old wood . 2. B. prasina

On rocks.

Exciple strong and seldom becoming covered . 4. B. chalybeia

Exciple weak and usually becoming covered . 3. B. lentibularis

2 O B S

342

OHIO BIOLOGICAL SURVEY

1. Biatorina heerii (Hepp) Fink Cont. Nat. Herb. 14: 83. 1910.

Biatoro heerii Hepp, Spore Flecht. Eur. pi. 16. f. 135. 1853.

Thallus of very minute, rounded and frequently heaped granules, some¬
times visible under a hand lens, but often seen only in sections of the sub¬
stratum, rarely disappearing; apothecia minute, 0.1 to 0.3 mm. in diameter,
adnate to sessile, flesh-colored and blackening, flat to slightly convex, the
concolorous or darker exciple commonly persistent; hypothecium and
hymenium pale to light brown; paraphyses distinct to coherent-indistinct;
asci clavate; spores ellipsoid, 7 to 12 mic. long and 3 to 3.5 mic. wide.

Collected in Butler County. On the thallus of Peltigera canina. Not
previously reported from Ohio. So minute as to be difficult to detect.
Consequently nothing further is known of its distribution in the State.

2. Biatorina prasina (Fr.) Fink Cont. Nat. Herb. 14: 84. 1910.

Micarea prasina Fr. Syst. Orb. Veg. 257. 1825.

Thallus of minute, closely clustered or even heaped granules, these
forming a wide-spread, frequently subleprose, green-gray to dark-olive
crust; apothecia minute to small, 0.2 to 0.5 mm. in diameter, adnate, com¬
monly carneous or darkening, more or less convex and usually becoming
convex with the exciple Anally covered; hypothecium pale or pale brown;
hymenium pale below and commonly darker above; paraphyses coherent,
semi-distinct to indistinct; asci clavate; spores oblong-ellipsoid, 8 to 12 mic.
long and 3.5 to 5 mic. wide.

Collected on Little Mountain in Lake County. On a rotten log. Not
previously reported from Ohio, and evidently rare in the State.

Simple spores were seen in the specimens collected, but they were
supposed to be immature.

3. Biatorina lentibularis (Ach.) Koerb. Syst. Lich. 191. 1855.

Lecidea lentibularis Ach. Syn. Meth. Lich. 28. 1814.

Thallus a thin, smooth or subtartareous, rarely rimose-areolate, ash-
white to brown-gray, wide-spread and continuous or Anally disappearing
crust; apothecia minute to small, 0.2 to 0.5 mm. in diameter, adnate, black,
from flat becoming convex and often irregular, the inconspicuous exciple
then becoming covered; hypothecium pale to darker brown; hymenium
pale or tinged brown; paraphyses distinct to coherent-indistinct; asci clavate;
spores oblong-ellipsoid, 8 to 11 mic. long and 2.7 to 4 mic. wide.

A single collection was made in Highland County. On exposed cal¬
careous rocks. Not previously reported from Ohio, and doubtless rare in
the State.

An occasional 4-celled spore was seen, a transitional character pre¬
viously noted by Th. M. Fries. The plant is closely related to the next
below, from which it may not be distinct.

THE ASCOMYCETES OF OHIO IV

343

4. Biatorina chalybeia (Borr.) Mudd, Man. Brit. Lich. 180. 1861.

Lecidea chalybeia Borr, in Sowerby, Eng. Bot. Suppl. 1: pi. 2687. f. 2. 1831.

Thallus a thin, smooth or roughened, ash-gray and darkening crust,
forming a continuous layer, becoming inconspicuous and rarely disappear¬
ing; apothecia minute to small, 0.3 to 0.5 mm. in diameter, adnate to
sessile, concave to slightly convex, black, the exciple concolorous, promi¬
nent, and rarely becoming covered; hypothecium dark brown; hymenium
pale below and* pale brown above; paraphyses wide and strong, distinct to
coherent-indistinct; asci clavate; spores oblong-ellipsoid, 8 to 12 mic. long
and 3.5 to 4.75 mic. wide (Fig. 4).

Collected in Butler County. On calcareous rocks. Not previously
reported from Ohio, and probably rare in the State.

The spores are somewhat larger than in European specimens.

Bilimbia De Not. Giorn. Bot. Ital. 21: 190. 1846.

Thallus usually composed of minute granules, these often run together
to form a leprose or verrucose and rarely areolate or even subsquaniulose
crust, rarely disappearing; apothecia minute or small, usually adnate, with
a weak and often covered exciple; hypothecium pale to dark brown;
hymenium pale or tinged brown; spores hyaline, usually fusiform or dacty-
loid, varying from 4- to 9-celled.

KEY TO THE SPECIES OF BILIMBIA

On rocks.

Apothecia and spores smaller . . . 5. B. microcarpa

Apothecia and spores larger . 6. B. trachona

On other substrata.

On mosses . . 2. B. hypnophila

On wood or bark.

Spores becoming more than 4-celled . . 3. B, naegelii

Spores not more than 4-celled.

Apothecia flesh-colored to dark brown . . 1. B. sphaeroides

Apothecia black . . . 4. B. melaena

1. Bilimbia sphaeroides (Dicks.) Koerb. Syst. Lich. 213. 1855.

Lichen sphaeroides Dicks. PI. Crypt. Brit. 1: 9. pi. 2. f. 3. 1785.

Thallus of minute, gray-green, often heaped granules, these forming
a continuous, thin or thicker crust; apothecia minute to small, 0.2 to 0.4
mm. in diameter, adnate, flesh-colored to red-brown, flat to convex and
subglobose, the inconspicuous, concolorous exciple soon covered; hypo¬
thecium and hymenium pale; paraphyses usually coherent-indistinct; asci
clavate; spores fusiform-ellipsoid, 4-celled, 12 to 20 mic. long and 4 to 6
mic. wide.

Collected on Little Mountain, in Lake County. On a rotten log in a
wood. Rare in Ohio, and its distribution unknown.

The plant is typical internally, but is young with small, flat or slightly
convex, light-colored apothecia.

344

OHIO BIOLOGICAL SURVEY

2. Bilimbia hypnophila (Ach.) Th. Fr. Nov. Act. Reg. Soc. Sci. Ups.
III. 3; 283. 1861.

Lecidea hypnophila Ach. Lich. Univ. 199. 1810.

Thallus of minute, usually crowded, sometimes confluent granules,
these forming an ash- or green-gray, thin, leprose or subgranulose, some¬
times scattered and disappearing crust; apothecia minute to middle-sized,
0.2 to 0.75 mm. in diameter, light brown to black, adnate to sessile, scat¬
tered or clustered, becoming strongly convex and the exciple becoming
covered; hypothecium pale or darker brown; hymenium pale, or tinged
brown below and more plainly brown above; paraphyses coherent, semi-
distinct to indistinct; asci clavate or long-clavate; spores ellipsoid to fusi¬
form, 4- to 8-celled, 16 to 35 mic. long and 4 to 8 mic. wide.

Collected in Preble, Hocking, and Adams counties. Over mosses on
rocks or bases of trees; or rarely on rocks, soil, bark, or wood. Not pre¬
viously reported from Ohio, and not a common fungus in the State.

3. Bilimbia naegelii (Hepp) Zwackh. Flora. 45: 505. 1862.

Biatora naegelii Hepp, Spor. Flecht. Fur. pi. 4. f. 1. 19. 1853.

Thallus of usually flattened granules, these commonly running together

to form a moderately thin, more or less roughened, often chinky, ash- or
green-gray, or darkening, limited or rarely wide-spread crust; apothecia
minute to middle-sized, 0.2 to 0.9 mm. in diameter, adnate or rarely sessile,
flesh-colored to dark brown, scattered or clustered, flat with the thin exciple
visible to strongly convex with the exciple covered; hypothecium pale or
tinged brown; hymenium pale throughout or tinged brown above; para¬
physes coherent, semi-distinct to indistinct; asci clavate; spores fusiform-
ellipsoid, 4- to 8-celled, 18 to 25 mic. long and 3 to 4 mic. wide.

Collected in Highland County. On bark. Not previously reported
from Ohio, and doubtless rare in the State.

The usual width given for the spores is 4 to 6 mic., and our plant is
placed here provisionally.

4. Bilimbia melaena (Nyl.) Th. Fr. Lich. Scand. 383-385. 1871.

Lecidea melaena Nyl. Bot. Not. 1853: 182. 1853.

Thallus of minute, olive-green to black-brown granules, these forming
a thin, granulose or scurfy, sometimes disappearing crust; apothecia minute
to small, 0.25 to 0.55 mm. in diameter, black-brown to black, sessile, occur¬
ring singly or in clusters, strongly convex to subglobose, the exciple soon
covered; hypothecium pale brown to red-brown; hymenium pale or tinged
brown; paraphyses coherent, semi-distinct' to indistinct; asci clavate to in-
flated-clavate; spores oblong-ellipsoid or dactyloid, 2- to 4-celled, 12 to 22
mic. long and 4 to 6 mic. wide.

Collected in Lake County. On an old log in a wood. Not previously
reported in Ohio, and rare in the State.

THE ASCOMYCETES OF OHIO IV

345

Nylander called the apothecium pale within, but forms with red-brown
hypothecia are admitted by later writers.

5. Bilimbia microcarpa Th. Fr. Bot. Not. 1863: 8. 1863.

Bilimbia obscurata microcarpa Th. Fr. Nov. Act. Soc. Sci. Ups. HI. 3:
183. 1861.

Thallus of minute ash-gray or green-gray granules, these rarely form¬
ing a thin or moderately thick, subcontinuous, verrucose crust, but more
often scattered or disappearing entirely; apothecia minute to small, 0.25
to 0.7 mm. in diameter, scattered or conglomerate, dirty brown to black,
soon becoming convex and subglobose, with the pale exciple then covered;
hypothecium pale to pale red-brown; hymenium pale; asci clavate to inflat-
ed-clavate; paraphyses coherent-indistinct; spores fusiform, 4-celled, 16 to
25 mic. long and 4 to 6 mic. wide.

Collected in Hocking County. On shaded sandstone. Not previously
reported from North America.

6. Bilimbia trachona (Ach.) Oliver Lich. France 38,39. 1903.

Verrucaria trachona Ach. Meth. Lich. Suppl. 16. 1803.

Thallus thin and granular, passing into smooth or leprose conditions,
thence to thickened and subareolate states, ash-colored to dark brown-
green, usually continuous over considerable areas; apothecia minute to
middle-sized, 0.4 to 0.1 mm. in diameter, from brown-black with lighter
exciple to wholly black, adnate or somewhat immersed, flat or finally convex
with the exciple at length covered; hypothecium pale brown to black-brown;
hymenium pale or rarely pale brown; paraphyses distinct to coherent semi-
distinct; asci clavate; spores fusiform-dactyloid, 4-celled, 12 to 20 mic. long
and 2.5 to 4.5 mic. wide (Fig. 5).

Collected in several localities in Preble, Highland, and Adams counties.
On rocks, usually limestone. Also reported from Cuyahoga and Ottawa
counties. Not common, but doubtless distributed widely in the State.

Bacidia De Not. Giorn. Bot. Ital. 2: 189. 1846.

Thallus granulose, passing into chinky, verrucose, subareolate and
subsquamulose conditions, seldom or never disappearing; apothecia minute
to large, adnate or rarely immersed more or less, exciple usually weak and
becoming covered; hypothecium commonly some shade of yellow or brown;
hymenium pale to light brown; spores hyaline, acicular, varying from 4- to
16-celled, often curved or variously twisted, usually 8 in each ascus.

KEY TO THE SPECIES OF BACIDIA

On rocks.

Spores hamate or spirally twisted . 7. B. umbrina

Spores straight or only slighly curved.

Thallus ash- or green-gray . 5. B. inundata

Thallus olive or darker . . . 1. B. egenuloidea

346

OHIO BIOLOGICAL SURVEY

On bark.

Spores less than 40 mic. in length . . 6. B. incompta

Spores 40 to 70 mic. long.

Apotheci a flesh-yellow to red-brown . 2, B. rubella

Apothecia brown to black.

Apothecia usually brown with a striate, usually

pruinose margin . . 3. B. fuscorubella

Apothecia usually black or dark brown, without striate

and pruinose margin . . 4. B. schweinitzii

1. Bacidia egenuloidea sp. nov.

Thallus of minute, crowded granules, forming a rather thick, conspicu¬
ous, rugose and obscurely chinky, dirt-olive and darkening, wide-spread
crust; apothecia minute to small, 0.25 to 0.4 mm. in diameter, yellow-brown
and darkening, adnate-sessile, flat with an elevated, darker exciple; hypo-
thecium and hymenium pale or tinged brown; paraphyses coherent, semi-
distinct; asci clavate; spores hyaline obscurely several-celled, variously
curved, 25 to 40 mic. long and 0.75 to 1.25 mic. wide.

Collected in Preble County. On granite in a damp fleld near West
Alexandria. The type specimen is deposited in the writer’s herbarium, and
a cotype may be found in the State Herbarium.

2. Bacidia rubella (Hoffm.) Mass. Ric. Lich. 118. f. 231. 1852

Verrucaria rubella Hoffm. Deutsch. FI. 2: 174. 1795.

Thallus of minute, scattered or crowed granules, these frequently
becoming compacted into a subleprose or more or less verrucose or chinky,
ash- to green-gray, moderately thick or thinner, continuous or sometimes
scattered and disappearing crust (Fig. 2); apothecia small to large, 0.5 to
1.35 mm. in diameter, sessile to adnate, flesh-yellow to red-brown, flat
with a rather thick and lighter-colored exciple, or becoming convex with
the exciple Anally covered; hypothecium pale yellow to brown; hymenium
pale yellow; paraphyses coherent, semi-distinct to indistinct; asci long
clavate; spores about 8- to 16-celled, 45 to 65 mic. long and 3 to 4 mic.
wide.

Collected in Butler, Highland, Adams, and Preble counties. Also
examined from Franklin County. On bark. Widely distributed in Ohio,
but not common.

3. Bacidia fuscorubella (Hoffm.) Arn. Flora 54: 55. 1871.

Verrucaria fuscorubella Hoffm. Deutsch. FI. 2: 175. 1795.

Thallus of minute, crowded or scattered granules, these forming a
usually conspicuous and often rugose and chinky, green-gray or darker,
frequently wide-spread, rarely disappearing crust; apothecia small to large,
0.6 to 1.5 mm. in diameter, pale to darker brown and Anally black, adnate
or sessile, flat with an elevated, and sometimes transversely striate, and
usually pruinose exciple, less frequently becoming convex with the exciple
rarely becoming covered; hypothecium yellow to yellow-brown; hymenium

THE ASCOMYCETES OF OHIO IV

347

pale yellow; paraphyses coherent, semi-distinct to indistinct; asci long-
clavate; spores about 7- to 14-celled, 40 to 70 mic. long and 3 to 5 mic. wide.

Collected in Butler and Adams counties. Also reported from Cham¬
paign and Hamilton counties. On bark. This fungus appears to be rare
in Ohio.

In one specimen, some of the disks are partly or wholly pruinose, but
the plant seemed nearer to this than to Bacidia sujfusa (Fr.) Fink.

4. Bacidia schweinitzii (Tuck.) Fink Cont. Nat. Herb. 14: 89. 1910.

Biatora schweinitzii Tuck, in Dari. FI. Cestr. ed. 3. 447. 1853.

Thallus thin and inconspicuous, or becoming thick and more prom¬
inent, composed of rounded and often crowded or even heaped granules,
these frequently compacted into a continuous or scattered, verrucose and
often chinky, green-gray to olivaceous crust; apothecia small to large, 0.6
to 1.75 mm. in diameter, dark brown to black, adnate or sessile, flat or
slightly convex, the concolorous or lighter exciple frequently becoming
flexuous; hypothecium pale yellow to dark brown; hymenium pale yellow;
paraphyses coherent, distinct to semi-distinct; asci long-clavate; spores
about 7- to 15-celled, 40 to 70 mic. long and 2.5 to 3.5 mic. wide.

Collected in Fairfleld, Hocking, and Adams counties. On bark. Evi¬
dently a rare fungus in Ohio.

5. Bacidia inundata (Fr.) Koerb. Syst. Lich. 187. 1855.

Biatora inundata Fr. Vet. Akad. Handl. 1822: 270. 1822.

Thallus of minute granules, these usually compacted into a thin or rarely
thicker, granulate, chinky, or subareolate, ash- or green-gray or darkening,
commonly wide-spread, continuous or scattered crust; apothecia minute to
middle-sized, 0.2 to 0.75 mm. in diameter, pale brown to Anally black, adnate
or rarely more or less immersed, usually flat and bordered by the commonly
lighter colored exciple, rarely becoming convex, the exciple then Anally
covered; hypothecium pale to brown; hymenium pale to pale brown; para¬
physes coherent, semi-distinct to indistinct; asci clavate to long-clavate,
spores 4- to 8-celled, 15 to 40 mic. long and 1.5 to 2.6 mic. wide.

Collected in Butler, Preble, Highland, Adams, Warren, and Lake coun¬
ties. On various rocks in shaded or open moist places, and also about the
moist shaded bases of rocks in dry fields. Also reported from Cuyahoga
county and doubtless common in all parts of the State.

6. Bacidia incompta (Borr.) Anzi, Cat. Lich. Sondr. 70. 1860.

Lecidea incompta Borr. in Sowerby, Engl. Bot. Suppl. 2: pi. 2699. 1834.

Thallus of very minute granules, these forming a continuous or more

or less broken, wide-spread, sometimes thick and rugose or rarely even
subareolate, or again thin, smooth, more or less mealy, light or darker
green-gray, rarely disappearing crust; apothecia minute to middle-sized,
0.3 to 0.75 mm. in diameter, dark brown to black, adnate to sub-sessile.

348

OHIO BIOLOGICAL SURVEY

flat or becoming convex, with a thin and frequently flexuous exciple; hypo-
thecium pale brown to brown; hymenium pale below and pale brown above;
paraphyses coherent, semi-distinct to indistinct; asci long-clavate; spores
4- to 12-celled, 18 to 35 mic. long and 1.5 to 3 mic. wide.

Collected in Adams County. On bark. Not previously reported from
Ohio, and doubtless rare in the State.

7. Bacidia umbrina (Ach.) Br. & Rostr. Bot. Tidssk. 3: 235. 1869.

Lecidea umbrina Ach. Lich. Univ. 183. 1810.

Thallus a rather thick and continuous, or rarely thinner and scattered,
subleprose, chinky, rugose-granulate or subareolate, green-gray to dark
olive-brown, sometimes largely disappearing crust; apothecia minute to
small, 0.25 to 0.6 mm. in diameter, light brown to black, adnate to some¬
what immersed, at first flat with a commonly paler exciple, becoming con¬
vex with the exciple sometimes covered; hypothecium pale or darker brown;
hymenium pale throughout, or tinged brown above; paraphyses coherent,
semi-distinct to indistinct; asci long-clavate, or inflated-clavate; spores
hamate, or more or less spirally twisted, about 4- to 8-celled, 18 to 30 mic.
long and 2 to 3 mic. wide (Fig. 7).

Collected in Preble, Lake, Hocking, and Adams counties. Also ex¬
amined from Wayne County. On various rocks. Not previously reported
from Ohio, but evidently distributed widely in the State.

Buellia De Not. Giorn. Bot. Ital. 21; 195. 1846.

Thallus granulose, verrucose, or areolate, rather better developed than
those of the preceding genera as shown in the more frequent verrucose and
areolate conditions; apothecia minute to large, sessile to immersed, the
disk and the exciple usually black; hypothecium usually brown; hymenium
pale to light brown; paraphyses usually distinct; spores brown, 2-celled.

KEY TO THE SPECIES OF BUELLIA

On rocks . 3. B. turgescenioides

On wood, or on bark.

On dead wood . . 1. B. myriocarpa

On bark . . 2. B. parasema

1. Buellia myriocarpa (Lam. & DC.) Mudd. Man. Brit. Lich. 217. 1861.

Patellaria myriocarpa Lam. & DC. FI. ed. 3. 2: 346. 1805.

Thallus a thin and scurfy, smooth or chinky, or thicker and roughened-
verrucose, ash- to green-gray, or darkening crust, irregularly spread over
small areas, and rarely disappearing; apothecia minute to small, 0.2 to 0.6
mm. in diameter, often numerous, black, adnate, flat and bordered by an
exciple, or becoming convex with the exciple sometimes covered; hypothe¬
cium dark brown; hymenium pale, or pale below and pale brown above;
paraphyses distinct, but sometimes loosely coherent; asci clavate; spores
oblong-ellipsoid, 7 to 16 mic. long and 4 to 7.5 mic. wide.

• THE ASCOMYCETES OF OHIO IV

349

Collected in Butler and Lake counties. On dead Vv^ood, especially posts
and boards. Also reported from Cuyahoga County. An inconspicuous
fungus, doubtless distributed widely in the State.

2. Buellia parasema (Ach.) Koerb. Syst. Lich. 228. 1855.

Lichen parasemus Ach. Lich. Suec. 64. 1798.

Thallus usually continuous and smooth, but sometimes becoming
thicker and roughened, granulate, chinky, or finally areolate, ash- to green-
gray, and darkening, or even yellow-green, usually bordered wholly or in
part by a black margin; apothecia small to large, 0.4 to 1.3 mm. in diameter,
black, adnate to sessile, or rarely more or less immersed, flat with a prom¬
inent, concolorous, sometimes flexuous exciple, or sometimes becoming
convex, with the exciple often covered (Fig. 11); hypothecium dark brown;
hymenium pale below and pale brown above; paraphyses distinct (Fig. 12),
but sometimes loosely coherent; asci clavate (Fig. 13), or rarely inflated
clavate; spores oblong to ellipsoid, 10 to 18 mic. long and 5 to 9 mic. wide,
rarely 3-celled (Fig. 13).

Collected in Fairfield, Lake, Adams, Highland, Hocking, and Butler
counties. Also examined from Morgan, Madison, and Muskingum coun¬
ties. On bark. Generally distributed in Ohio.

3. Buellia turgescentoides sp. nov.

Thallus a thick, continuous or scattered, flat or verrucose, areolate or
subareolate, dull olive-brown, and darkening crust, covering small areas
or spreading widely over the substratum, the marginal areoles sometimes
lobulate; apothecia minute to small, 0.2 to 0.5 mm. in diameter, im.mersed
to adnate, scattered or clustered, black, flat with the thin concolorous
exciple visible, or convex with the exciple covered; hypothecium pale or
darker brown; hymenium pale; paraphyses stout, distinct, but often loosely
coherent; asci clavate or inflated-clavate; spores brown, 2-celled, oblong to
oblong-ellipsoid, 8 to 13 mic. long, and 4 to 6 mic. wide, 8 in each ascus.

Collected in Lake County. On exposed igneous rocks. The type
specimen is deposited in the writer’s herbarium, and a cotype may be found
in the State Herbarium.

This species is a coarser plant than Buellia turgescens (Nyl.) Tuck.,
with much stronger, darker thallus and apothecia on the whole larger.

Rhizocarpon Ram. in Lam. & DC. FI. Fr. ed. 3. 2: 365. 1805.

Thallus usually verrucose, areolate or subareolate, tending toward
squamulose conditions, better developed than in other members of the
family, scarcely ever showing granulate conditions, and never disappearing
entirely; apothecia also larger than in the other genera, adnate to immersed,
usually black, but rarely white-pruinose; hypothecium usually dark brown;
hymenium pale to light brown; spores 4-celled to muriform, and pale to
brown, various conditions of septation and coloration sometimes appearing
in the same hymenium.

350

OHIO BIOLOGICAL SURVEY

KEY TO THE SPECIES OF RHIZOCARPON

On bark . 2. R. alboatrum

On rocks.

Spores smaller and 4-celled . 1. R. vernicomoideum

Spores larger and becoming muriform . 3. R. petraeum

1. Rhizocarpon vernicomoideum sp. nov.

Thallus of minute, rounded, scattered or sometimes clustered, straw-
colored granules, covering small areas, and usually resting on and limited
wholly or in part by a black hypothallus; apothecia minute to small, 0.2 to 0.6
mm. in diameter, black, semi-immersed to adnate, at first flat with a thin
somewhat raised exciple, becoming convex with the exciple finally covered;
hypothecium brown; hymenium pale or tinged brown below and light
brown above; paraphyses coherent, distinct or semi-distinct; asci clavate;
spores brown, 4-celled, becoming slightly constricted at the septa, 15 to 18
mic. long and 5 to 7 mic. wide, 8 in each ascus.

Collected at Cantwell Cave in Hocking County. On shaded sand¬
stone, intermingled with an ash-gray, crustose thallus, which appeared
like a sterile Pertusaria. The type specimen is deposited in the writer’s
herbarium, and a cotype may be seen in the State Herbarium.

The plant resembles Buellia vernicoma Tuck.

2. Rhizocarpon alboatrum (Hoffm,) Th. Fr. Nov. Act. Reg. Soc. Sci.
Ups. HI. 3: 337. 1861.

Lichen alboater Hoffm. Lich. Icon. 30. 1784.

Thallus ash-gray varying toward white, commonly spread widely over
the substratum as a continuous or rarely scattered or disappearing, smooth,
chinky, verrucose-areolate, or sometimes mealy crust: apothecia small to
middle-sized, 0.35. to 1 mm. in diameter, adnate or immersed, dull black
and often more or less white-pruinose, flat with the black exciple visible,
or convex when the exciple often becomes covered; hypothecium brown
to black-brown; hymenium pale or tinged brown; paraphyses distinct, but
sometimes coherent; asci clavate; spores oblong-ellipsoid, brown, 4-celled
to muriform, 12 to 22 mic. long and 4 to 9 mic. wide (Fig.8j, 8 in each
ascus.

Collected in Butler, Preble, Ross, and Highland counties. On bark,
especially elm bark. Also reported from Ottawa County. Rare but doubt¬
less distributed widely in the State.

3. Rhizocarpon petraeum (Wulf. ) Koerb. Syst. Lich. 260. 1855,

Lichen petraeus Wulf. in Jacq. Coll. Bot. 3: 4. pi. 6. f. 2a. 1789.

Thallus an ash or green-gray crust, or varying toward brown or brown-
black, smooth to more commonly roughened, chinky to areolate, continuous
or scattered, of moderate thickness, often widely and irregularly disposed
on the substratum; apothecia small to large, 0.5 to 1.3 mm. in diameter,
immersed to adnate, black-brown to black, flat with the concolorous

THE ASCOMYCETES OF OHIO IV

351

exdple visible, or becoming somewhat convex, with the exciple often
covered; hypothecium dark brown; hymenium pale, or tinged brown, espec¬
ially above; paraphyses coherent, semi-distinct; asci clavate or inflated-cla-
vate; spores oblong-ellipsoid, 4 celled to muriform, hyaline to finally brown,
15 to 40 mic. long and 7 to 18 mic. wide. 8 in each ascus (Fig. 9).

Collected in Lake, Hocking, and Ross counties. Also examined from
Summit, Vinton, and Ashtabula counties. On rocks. Rare but widely
distributed in the State

352

OHIO BIOLOGICAL SURVEY

EXPLANATION OF PLATE XIII

Fig. 1. Five paraphyses of Rhizocarpon alboairum to illustrate types of simple and
branched forms found in the same hymenium. X 450.

Fig. 2. A section of the thallus of Bac/W/fl rubella and two cells of the woody sub¬
stratum: a, the upper densely interwoven portion of the thallus; b, part of
the less densely interwoven portion below; c, the algal-host cells; d, one of
the cells of the woody substratum and three hypal rhizoids within it. X 450.

Fig. 3. Spores of Lecidea enteroleuca to illustrate the simple, hyaline type. X 760.

Fig. 4. Spores of Biatorina chalybeia to illustrate the 2-celled, hyaline type. X 760.

Fig. 5. Spores of Bilimbia trachona to illustrate the several-celled, hyaline, fusiform
or dactyloid type. X 760.

Fig. 6. Spores of Bacidia fuscorubella to illustrate the several-celled, hyaline, acicular
type. X 760.

Fig. 7. Spores of Bacidia umbrina to illustrate the several-celled, hyaline, hamate or
spirally twisted type. X 760.

Fig. 8. Spores of Rhizocarpon alboatrum to illustrate the several-celled to many-
celled and muriform, hyaline to brown type. X 760.

Fig. 9. Spores of Rhizocarpon petraeum of the same type as those shown in the last
figure, but larger, and usually composed of more cells. X 760.

Fig. 10. A vertical section through an apothecium of Lecidea rupestris: a, the hyme¬
nium, composed of asci and paraphyses; b, the hypothecium; c, the mycel¬
ium, the cells of the algal host, and particles of the limestone on which the
plant was growing; d, the weak, light-colored, covered exciple. X 79.

Fig. 11. A vertical section through an apothecium of Buellia parasema, the thallus
below, and a portion of the woody substratum: a, the hymenium, composed
of asci and paraphyses; b, the strongly developed, dark colored exciple; c,
the dark colored hypothecium; d, the thallus, composed of interwoven
hyphae, and enclosing cells of the algal host, a portion of the woody
substratum. X 79.

Fig. 12. One branched and one unbranched paraphysis of Buellia parasema. X 450.

Fig. 13. An ascus of Buella parasema, containing 8 spores. X 450.

The figures were outlined with camera lucida and drawn on the table, close to the base of the

microscope, 100 mm. below the stage. They were reduced one-half in making the plate. Figures 2,

10, and 11 are partly diagrammatical.

OHIO BIOLOGICAL SURVEY— VOL. 11.

PLATE XIII.

353

354

OHIO BIOLOGICAL SURVEY

THE ASCOMYCETES OF OHIO V*

The Peliigeraceae.

Leafy J. Corrington.

Two genera, Peltigera and Nephroma, constitute the Peliigeraceae as
represented in the flora of Ohio. The thallus is plainly foliose with the
margins of the lobes usually ascending and is gray-green to brown in color.
The lower surface is often conspicuously veined. There are two pronounced
distinctions between the two genera. Peltigera has a well-developed
cortex on the upper side of the thallus only (Fig. 1), while in Nephroma
there is a well-developed cortex on both upper and lower sides (Fig. 2).

The position of the apothecia constitutes another distinction. In both
genera the apothecia are marginal or submarginal on the lobes, which
are usually narrow and somewhat extended; but in Peltigera they are immer¬
sed in the upper surface, while in Nephroma they are imbedded in the lower
surface.

Peltigera furnishes seven species for Ohio, while only one species of
Nephroma has thus far been found in the State.

The algal hosts are usually Daciylococcus or Polycoccus, and both
hosts are sometimes found in the sam.e thallus. The chains of cells are
usually badly broken up, and the nature of the algal host is, therefore,
difficult to distinguish. Other algae doubtless sometimes occur in the
thalli of Peltigerae.

Nephroma with cortex on both sides, is to be regarded higher than
Peltigera, which has the cortex on the upper side only. The family is
most closely related to the Stictaceae, from which family it is kept distinct
on account of the absence of cyphellae and the difference in disposition of
the apothecia.

The collecting on which this paper is based was partly by Bruce Fink
in connection with general collecting of fungi in Butler County and in
collecting in Adams, Warren, Fairfleld, Preble, Ross, Highland, and Lake
counties for the Ohio Biological Survey. However, a considerable amount
of material found by other collectors and previously reported from Ohio
was examined. Hence, the collecting for the Ohio Biological Survey added
little to knowledge of the Pcltigeraceae of Ohio, except in way of addition to
distribution in the State.

Systematic Account
PELTIGERACEAE

Thallus foliose, with plectenchymatous cortex above (Fig.5), or both
above and below (Fig. 2), with medulla of loosely interwoven hyphae,
trichomatic hyphae, usually present, attached to the substratum by com-

*Contributions from the Botanical Laboratory of Miami University— XIX

THE ASCOMYCETES OF OHIO V

355

pound rhizoids; apothecia of considerable size, commonly on extended
lobes, usually imbedded in the tissues on the upper side, or more rarely
on the lower side; exciple inconspicuous; hypothecium usually light or
darker brown; hymenium usually pale below and brown or tinged brown
above; paraphyses simple or branched, distinct, seldom gelatinized or
coherent; spores hyaline or brown, 4- to several-called, elongated.

KEY TO THE GENERA

1. Cortex developed on the upper side of thallus only,

spores hyaline . Peltigera.

2. Cortex developed on both upper and lower sides of

thallus, spores brown . Nephroma.

Peltigera Willd. FI. Berol. Prodr. 347. 1787.

Thallus foliose, usually adnate toward the center, with the lobes more
or less ascending at the margins, green-gray varying toward brown, the
upper surface sometimes bare, or again clothed with trichomatic hyphae,
giving it a downy appearance, or bearing cephalodia or isidioid branchlets,
the lower surface usually conspicuously veined, with tufted -rhizoids
descending from the veins, color of these light or dark; cross section
showing two distinct layers, the upper plectenchymatous cortex composed
of 2 to 4 layers of meshes, and the medulla, composed of densely interwoven
and irregularly disposed hyphae; lower cortex lacking, but the hyphae of
the lower portion in some instances more or less horizontally arranged and
produced into hyphal rhizoids, thus serving for support and protection
much like a true plectenchymatous cortex; apothecia usually orbicular,
frequently revolute, imbedded in the upper surface of the lobes; exciple
plectenchymatous (Fig. 4); hypothecium of interwoven hyphae, usually
tinged brown; hymenium commonly pale below and brown above; par¬
aphyses usually simple, but some branched ones present in all of the
species, hyaline in the main, but usually enlarged and tinged brown at the
apex; asci usually cylindrico-clavate; spores hyaline, fusiform to acicular,
sometimes curved, 4 to 8-celled, 8 arranged parallel in the asci.

The algal host cells lie in the medulla, just below the upper cortex.

2. P.

KEY TO THE SPECIES OF PELTIGERA

Upper surface of the thallus bearing cephalodia
Upper surface of the thallus devoid of cephalodia.

Thallus bearing trichomatic hyphae above.

Upper surface bearing isidioid branchlets or lobules
Upper surface devoid of isidioid branchlets or lobules.

Orbicular sorediate areas on the upper surface of the thallus 3
Soredia lacking on the upper surface.

Lower surface of the thallus of light color
Lower surface partly or wholly dark
Thallus devoid of trichomatic hyphae.

Apothecia orbicular and revolute, spores 4-to 8-celled
Apothecia usually transversely oblong, spores 4-celled

1. P. aphthosa

5.

6. P.

7. P.

praetextata

P. sored iata

4. P. canina
P. rufescens

polydactyla

horizontalis

356

OHIO BIOLOGICAL SURVEY

1. Peltigera aphthosa (L.) Willd. FI. BeroL Prodr. 347. 1787.

Lichen aphthosus L. Sp. PI. 1148. 1753.

Thallus closely attached to the substratum at the center, the lobes
ascending, 6 to 8 cm. in diameter, smooth and devoid of trichomatic hyphae
above, cephalodia more or less rounded and irregularly scattered over the
surface (Fig. 3), the lobes broad and rounded with crenate margins,
brown above, the lower surface having numerous veins, these forming a
dark brown nap at the center, the veins distinct and light toward the
margin, dark rhizoids extending from the veins; medulla composed of thick-
walled, densely interwoven hyphae, irregularly disposed; apothecia on
extended lobules, orbicular and frequently revolute, the disk red-brown, 2
to 5 mm. in diameter, the margin entire or crenulate; hypothecium pale
brown; hymenium hyaline to pale brown above; asci cylindrico-clavate;
spores acicular, straight, 4-to 6-celled, 47 to 66 mic. long and 4 to 5 mic.
wide (Fig. 8, d).

Examined from Clark County. Also reported from Champaign
County. On earth and often on humus-covered rocks. Rare in Ohio.

2. Peltigera praetextata (Sommerf.) Fink. Proc. Ind. Acad. Sci. 1918:
267. 1918.

Peltigera ulorrhiza praetextata Sommerf. Lapp. Suppl. 123. 1826.

Thallus adnate toward the center, more or less ascending toward the
margins, 7 to 15 cm. in diameter, the upper surface having isidioid branch-
lets or lobules scattered more or less thickly, the lobes broad, wavy,
crenate, with frequently isidioid, lobulate margins, trichomatic hyphae
often present, usually green-gray toward the center, becoming brown toward
the margin, the lower surface light with numerous dark veins and bearing
rhizoids of the same color, the veins and rhizoids becoming light colored
toward the margin; medulla of densely interwoven and irregularly dis¬
posed hyphae; apothecia on narrow, somewhat extended lobes, the disk
brown to brown-black, revolute, 2 to 5 mm. in diameter, hypothecium
light to darker brown; hymenium pale below and brown above; asci cylin¬
drico-clavate; spores sub-fusiform to acicular, usually straight but some¬
times slightly curved, 4-to 6-celled, 42 to 56 mic. long and 3 to 5 mic. wide.

Examined from Eranklin, Adams, Butler, Marion, Jefferson, and Preble
counties. On soil, old logs, and moss in woods. Not previously reported
from Ohio, but included under Peltigera canina and Peltigera rufescens.
Evidently widely distributed and frequent in the State, but seldom fruited.

3. Peltigera sorediata (Schaer. ) Eink comb. nov.

Peltigera canina spuria sorediata Schaer. Enum. Lich. Eur. 21. 1850.

Thallus small, composed of scattered lobes, these 1 to 4.5 cm. in length,
adnate with slightly ascending rounded, and crenate margins, the upper
surface usually deep gray at the center, becoming lighter toward the
margin, thickly covered with trichomatic hyphae, orbicular sorediate areas

THE ASCOMYCETES OF OHIO V

357

scattered over the upper surface, the lower surface ash-white to cream-
colored, with a network of veins of the same color, with similarly colored
rhizoids extending downward; medulla of small, densely interwoven and
irregularly extending hyphae; apothecia somewhat digitately clustered on
the narrow lobes, small, 1. 3 to 3 mm. in diameter, orbicular, flat or semi¬
revolute, dark brown; hypothecium light brown; hymenium hyaline below
and brown above; asci long-clavate; spores acicular, 6-to 8-celled, 53 to 66
mic. long and 3 to 3.5 mic. wide (Fig. 8, b).

Examined from Butler and Lake counties. On damp earth and
mossy rocks. Not previously reported from Ohio, and probably not
widely distributed in the State. Surely rare.

4. Peltigera canina (L.) Hoffm. Deutsch. FI. 2: 108. 1795.

Lichen caninm L. Sp. PI. 1149. 1753.

Thallus closely adnate toward center, the lobes more or less ascend¬
ing, 6 to 15 cm. in diameter, the upper surface for the most part thickly
covered with trichomatic hyphae, generally giving it a downy appearance
under the lens, the lobes numerous and usually broad and rounded, with
entire or crenate and much crisped margins, usually green-gray but some¬
times becoming brown, below almost white, netted with light brown or
gray veins, these bearing rhizoids of the same color; medulla (Fig. 5) of
densely interwoven and irregularly disposed hyphae; apothecia on narrow,
extended lobes, often erect, orbicular, usually revolute, 2 to 7 mm. in
diameter, the disk dark brown; hypothecium (Fig. 7) pale brown; hy¬
menium (Fig. 6) pale below and brown above; asci long-clavate; spores
acicular, straight or sometimes curved, 4- to 8-celled, 30 to 65 mic. long
and 3 to 5 mic. wide (Fig. 8, d).

Examined from Butler, Franklin, Ashtabula, Green, Seneca, Summit,
Lorain, Preble, Brown, and Adams counties. On soil or mosses in woods.
Generally distributed and frequent in Ohio.

5. Peltigera rufescens (Neck.) Hoffm. Deutsch. FI. 2: 107. 1795.

Lichen rufescens Neck. Meth. Muse. 79. 1771.

Thallus closely adnate at the center with ascending lobes, 5 to 15 cm.
in diameter, the upper surface smooth and devoid of trichomatic hyphae
for the most part, but the margins sometimes sparingly covered with them,
green-gray to brown, the lobes crowded, rather small with crenate, much
crisped, elevated margins, the lower surface usually becoming dark brown
except at the margins, and thickly reticulated with brown veins, from
these numerous rhizoids of similar color extending; medulla of densely
interwoven variously disposed hyphae; apothecia numerous on narrow,
extending lobes, the disk brown to black-brown, revolute, 4 to 7 mm. in
diameter; hypothecium pale brown; hymenium hyaline to pale brown below
and dark brown above; asci long-clavate; spores acicular, straight or curved,
4- to 8-celled, 45 to 68 mic. long and 3.5 to 5 mic. wide.

358

OHIO BIOLOGICAL SURVEY

Examined from Butler, Preble, Clark, Adams, and Summit counties.
Also reported from Champaign County. On earth and mosses, commonly
about trees. Widely distributed in Ohio, but infrequent.

6. Peltigera polydactyla ( Neck. ) Hoffm. Deutsch. FI. 2: 106. 1795.

Lichen polydactylon Neck. Muse. 85. 1771.

Thallus adnate at the center with ascendant margins of the lobes, 6 to
11 cm. in diameter, the upper surface smooth and shining, devoid of trichc-
matic hyphae, the lobes broad with crisped, crenate margins, except those
bearing the apothecia, these much narrower and more elongated and usually
digitately clustered, brown in color for the most part, the lower surface
showing through a reticulation of dark veins as small light-colored spots,
numerous dark rhizoids extending downward from the veins; medulla of
densely interwoven and irregularly disposed hyphae; apothecia orbicular,
and usually revolute, the disk dark brown, 3 to 10 mm. in diameter;
hypothecium pale brown; hymenium pale below and dark brown above;
asci clavate to cylindrico-clavate; spores acicular, straight or slightly
curved, 4- to 6-celled, 42 to 70 mic. long and 3 to 4 mic. wide.

Examined from Clark, Fairfield, and Morgan counties. On earth.
Rare in Ohio.

7. Peltigera horizontalis (L.) Hoffm. Deutsch. El. 2: 107. 1795.

Lichen horizontalis L. Mant. PI. 2: 132. 1771.

Thallus mostly adnate, the margins scarcely ascending, 6 to 20 cm.
in diameter, the upper surface smooth and shining, devoid of trichomatic
hyphae, green-gray to brown, the lobes broad and rounded with entire or
crenate margins, the lower surface covered with numerous veins, these
giving a dark coloration toward the center and becoming light colored
toward the margins, numerous dark rhizoids extending down from the
veins; medulla of thick-walled, densely and irregularly disposed hyphae;
apothecia on somewhat narrowed lobes, transversely oblong or infrequently
orbicular, the disk red-brown, concave, 2 to 4 mm. in diameter; hypothe¬
cium light brown; hymenium pale below and dark brown toward the upper
surface; asci cylindrico-clavate; spores fusiform to long-ellipsoid, straight to
curved, 4-celled, 26 to 40 mic. long and 5 to 6.5 mic. wide. (Fig. 8 a).

Examined from Lake and Fairfield counties. On earth and mossy
rocks. Rare in Ohio.

Nephroma Ach. Lich. Univ. 101. 521. pi. 11. f. 1. 1810.

Thallus foliose, but smaller and thinner than that of Peltigera, and
devoid of trichomatic hyphae, more or less closely attached to the sub¬
stratum by rhizoids; cortex well developed on both upper and lower sides;
medulla well developed (Fig. 2); apothecia confined to the lower side of the
thallus, marginal on narrow, slightly elongated lobes, thalloid margin per¬
sistent and crenate; hypothecium usually some shade of brown; hymenium

THE ASCOMYCETES OF OHIO V

359

usually pale below and brown above; paraphyses simple or branched; spores
brown, 4-celled, 8 in each ascus.

The algal-host cells occur as in Peltigera

1. Nephroma Helvetica Ach. Lich. Univ. 523. 1810.

Thallus adnate, rather closely attached to the substratum by numer¬
ous short, hyaline, thick-walled rhizoids, irregular or sometimes orbicular
in form, 6 to 10 cm. in diameter, green-gray to brown above, smooth or
bearing tooth-like branchlets, narrowly and laciniately lobed, the margins
of the lobes serrate or crenate, slightly ascending, beneath finally tomentose,
and brown or black-brown; plectenchymatous cortices well developed above
and below; medulla of narrow, thin-walled, densely, variously disposed hy-
phae; apothecia numerous, the disk red-brown to almost black, 1.3 to 3 mm.
in diameter; hypothecium of interwoven hyphae, pale brown; hymenium
pale brown below and darker above; asci clavate; paraphyses simple or
branched, slightly swollen and brown at the apex; spores brown, ellipsoid,
4-celled, 15 to 21 mic. long and 5.5 to 8 mic. wide.

Examined from Butler and Champaign Counties. On trunks and
mossy rocks. Rare and usually sterile in Ohio.

360

OHIO BIOLOGICAL SURVEY

Fig. 1

Fig. 2
Fig. 3

Fig. 4

EXPLANATION OF PLATE XIV

A section through the thallus of Peltigera canina, showing the cortex above
and the medulla below, the medullary hyphae of the lower portion running
in a somewhat horizontal direction. The algal cells shaded. X 380.

A section through the thallus of Nephroma helvetica, showing the cortices, upper
and lower, and the mycelial medulla within. The algal cell shaded. X 760

A section through a cephalodium of Peltigera aphthosa; a, the surrounding
cortex; b, the internal hyphae and the cells of the algal host; c, the support¬
ing hyphae from the thallus below. Partly diagramatic. X 48.

A small portion of a section through the exciple of Peltigera canina, showing
the plectenchymatous structure. X 380.

OHIO BIOLOGICAL SURVEY— VOL. IL

PLATE XIV.

361

362

OHIO BIOLOGICAL SURVEY

EXPLANATION OF PLATE XV

Fig. 5. A section through the cortex of Peltigera canina, showing its relation to the
medullary hyphae. X 760.

Fig. 6. A portion of a section of the hymenium of Peltigera canina, showing two asci
containing spores, two asci with protoplasmic contents, and five paraphyses.
X 760.

Fig. 7. A portion of a section through an apothecium of Peltigera canina, showing
part of the hymenium of interwoven hyphae below and the bases of three
paraphyses above. X 760.

Fig. 8. Types of spores found in the Peltigeraceae: a, 4-celled spores of Peltigera
horizontalis; b, 6- to 8-celled spores of Peltigera sorediata; c, 4-celled spores
of Peltigera aphthoisa; d, 4- to 8-celled spores of Peltigera canina. X 380.

The drawings were made with camera lucida and were reduced one-half in making the plates.

OHIO BIOLOGICAL SURVEY— VOL. IL

PLATE XV.

363

INDEX

Bacidia . 345

Bacidia egenuloidea . 346

Bacidia fuscorubella . 346

Bacidia incompta . 347

Bacidia inundata . 347

Bacidia rubella . 346

Bacidia schweinitzii . 347

Bacidia umbrina . 348

Biatorella . 336

Biatorella pruinosa . 337

Biatorella simplex . . 336

Biatorina . 341

Biatorina chalybeia . 341

Biatorina heerii . 343

Biatorina lentibularis . 342

Biatorina prasina . 342

Bilimbia . 343

Bilimbia hypnophila . 344

Bilimbia melaena . 344

Bilimbia microcarpa . 345

Bilimbia naegelii . 344

Bilimbia sphaeroides . 343

Bilimbia trachona . 345

Buellia . .. 348

Buellia myriocarpa . 348

Buellia parasema . 349

Buellia turgescentoides . 349

Lecideaceae . 336

Lecidea . . . . 337

Lecidea albocaerulescens . 341

Lecidea coarctata . 338

Lecidea enteroleuca . 340

Lecidea flexuosa . 340

Lecidea humicola . 339

Lecidea intropallida . 338

Lecidea platycarpa . 341

Lecidea rupestris . 338

Lecidea sylvicola . . 340

Lecidea uliginosa . 339

Lecidea varians 338

Lecidea viridescens . 339

Nephroma . 358

Nephroma helvetica . 359

Peltigeraceae . 354

Peltigera aphthosa . 356

Peltigera canina . 357

Peltigera horizontalis . 358

Peltigera polydactyla . . . 358

Peltigera praetextata . 356

Peltigera rufescens . 357

Peltigera sorediata . 356

Rhizocarpon . 349

Rhizocarpon alboatrum . 350

Rhizocarpon petraeum . 350

Rhizocarpon vernicomoideum . 350

364

L. I (31? Al? I /i N
UN I V. OF ILLINOIS
UI?I3 ANA ill N

The Ohio State University Bulletin

Volume XXV May 28, 1921 Number 26

OHIO BIOLOGICAL SURVEY

BULLETIN W OF Si.!:;"’;

VOLUME II, No. 6 ‘

JUL 3 ^ 1923

: y,

THE ASCOMYCETES OF OHIO IV

BRUCE FINK

THE ASCOMYCETES OF OHIO V

LEAFY J. CORRINGTON

MAY. 1921

PUBLISHED BY THE UNIVERSITY AT COLUMBUS. OHIO

Entered as second-class matter November 1 7, 1905, at the Postoffice, at
Columbus, Ohio, under Act of Congress, July 16, 1 894.

OHIO BIOLOGICAL SURVEY

Herbert Osborn, Director

OHIO STATE UNIVERSITY IN CO-OPERATION WITH
OTHER OHIO ■ COLLEGES AND UNIVERSITIES

Administrative

Board consisting of Representatives from co-

operating institutions.

E. L. Fullmer,

Bald win- Wallace University,

Berea, 0.

A. B. Plowman,

University of Akron,

Akron, 0.

L. B. V/alton,

Kenyon College,

Gambler, 0.

Bruce Fink,

Miami University,

Oxford, 0.

.F. 0. Grover,

Oberlin College,

Oberlin, 0.

E. L. Rice,

Ohio Wesleyan University,

Delaware, 0.

F. H. Herrick,

Western Reserve University,

Cleveland, 0.

C. G. Shatzer,

\

Wittenberg College,

Springfield, 0.

ANNOUNCEMENT

The Bulletins of the Ohio Biological Survey will be issued as work
on any special subject is completed, and will form volumes of about
500 pages each.

They will be sent to co-operating institutions and individuals,
libraries and colleges in Ohio and to such surveys, societies and other
organizations as may offer suitable exchange material.

Additional copies of each Bulletin and of completed volumes
will be sold at such price as may cover the cost of publication.

Special rates on quantities to schools for classes.

Subscription for entire volumes . $2.00

Price of this number . 50

All orders should be accompanied by remittance made payable to
Ohio Biological Survey and sent to the Director.

Correspondence concerning the Survey, applications for exchanges
and purchase of copies of Bulletins should be addressed to the
Director — Professor Herbert Osborn, Columbus, Ohio.

Bulletins Ohio Biological Survey

I. Outline of Biological Survey Plan

Syrphidae of Ohio by C. L. Metcalf . $ .50

II. Catalog of Ohio Vascular Plants

by John H. Schaffner . 50

III. Botanical Survey of the Sugar Grove Region

by R. F. Griggs . . . 50

IV. The Euglenoidina of Ohio

by L. B. Walton . 50

V. The Ascomycetes of Ohio. — I

by Bruce Fink

The Ascomycetes of Ohio. — II

by Bruce Fink and C. Audrey Richards . 50

VI. Qualities and Uses of the Woods of Ohio

by Wm. R. Lazenby . 50

VII. The Physiographic Ecology of the Cincinnati Region

by E. Lucy Braun . 50

VIII. The Tingitoidea of Ohio

by Herbert Osborn and Carl J. Drake . 50

IX. The Grasses of Ohio "

by John H. Schaffner . 50

X. The Ascomycetes of Ohio. — IV and V

by Bruce Fink and Leafy J. Corrington . 50

0

OF iu.iaoiuc.T;a

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