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THE AMERICAN MIDLAND NATURALIST
Monograph No. 6
THE AMERICAN MIDLAND NATURALIST
Monograph Series
EDITORIAL STAFF
John D. Muelle Zoology
Editor, University of Notre Dame, Notre Dame, Ind.
Edward A. Chapin Entomology
U. S. National Museum, Washington, D. C.
Albert L. Delisle Plant Morphology
University of Notre Dame, Notre Dame, Ind.
Carroll Lane Fenton Inrertebrate Paleontology
404 Livingston Ave., New Brunswick, N. J.
John Hobart Hoskins Paleobotany
University of Cincinnati, Cincinnati, Ohio
George Neville Jones Plant Taxonomy
University of Illinois, Urbana, 111.
Remington Kellogg Mammalogy
U. S. National Museum, Washington, D. C.
Jean Myron Linsdale Ornithology
Hastings Reservation, Monterey, Calif.
George Willard Martin Mycology
State University of Iowa, Iowa City, Iowa
Hugh M. Raup Plant Ecology
Harvard Forest, Harvard University, Petersham, Mass.
Karl Patterson Schmidt Ichthyology and Herpetology
Chicago Natural History Museum, Chicago, 111.
Harley Jones Van Cleave Invertebrate Zoology
University of Illinois, Urbana, 111.
0
■ ^l\
THE AMERICAN MIDLAND NATURALIST X)0/
Monograph No. 6
Edited by John D. Mizelle
Published by the University of Notre Dame
Notre Dame, Indiana
A Manual of the Mosses of Western Pennsylvania
and Adjacent Regions
SECOND EDITION
By O. E. JENNINGS
Director Emeritus, Carnegie Museum
Formerly Head, Department of Biology, University of Pittsburgh and
Curator of Botany, Carnegie Museum; Editor, The Bryologist 1913-1938
J
University of Notre Dame Press
Notre Dame, Indiana
September, 1951
Copyright, 1951
by
The American Midland Naturalist
University of Notre Dame
Notre Dame, Indiana
Preface to the First Edition*
The aim in the preparation of this Manual has been to make it a practical
handbook applying particularly to the region of western Pennsylvania and
embodying all that is at present known regarding the occurrence and distribu-
tion of mosses within that area. As a matter of fact, the Manual will be found
to apply also to the adjacent regions of central Pennsylvania, extreme south-
western New York, eastern Ohio, and northern West Virginia.
When the present writer took charge of the botanical collections in the
Carnegie Museum in 1904 he found that the Herbarium, aside from certain
specimens collected by Mr. D. A. Burnett in McKean County, a iew years
previously, contained but little to represent the rich flora of messes and liver-
worts to be expec.ed in the western end of Pennsylvania. One of the aims of
the Herbarium of the Carnegie Museum has been to assemble a very complete
and comprehensive collection of all the plants to be found in the general region
in which Pittsbur[;h is situated, and, in the prosecution of this v/ork, the
writer has been enabled to visit all of the counties in the \7estern half of
Pennsylvania and also adjacent portions of Ohio and \7est Virginia. Certain
localities in this general region have been made the subject of detailed ecologic
and systematc study and collection — particularly the peninsula of Presque
Isle, near Erie, Pennsylvania; the extensive Pymatuning Swamp in Crawford
County, Pennsylvania; the mountainous region in the vicinity of Ohio Pyle,
Fayette County; and the larger portion of Allegheny County, especially in the
vicinity of Pittsburgh. From these and other localities visited, extensive col-
lections of r-.osses have been made and the amount and representative nature
of the herbarium material thus available for study have becom.e such that it
has been deemed advisable to prepare a treatise embodying the results of the
work accomplished, thus placing within the reach of other students of the
mosses v/ithin the region a convenient m.eans of identifying and checkinrr their
collections. It is hoped that with all its faults this Manual may be to some
extent the m.eans of stimulating bryological study in a region of v.'hcse mosses
there is yet much to be learned.
In the preparation of this Manual the author has taken as the taxonomic
standard the monumental work of Warnstorf, Ruhland, and Brotherus, brought
to completion in 1909, in Engler iC Prantl's Die Natiirlichen Pjlanzenfa-milien,
Teil I, Abteilung III. In the characterization of the various orders, families, and
genera, these authors have been followed closely, and, while there is much to
be said against their arrangement of families in certain cases, it is nevertheless
very probable that their work will remain for a long time the standard and
that, from the standpoint of convenience at least, a similar sequence of families
in this Manual is justified. In the determination of the various species the
* This wor!- in a more condensed form was submitted as a major thesis in candidacy
for the degree of Doctor of Philosophy in the University of Pittsburgh, June, 1911.
Jennings: Manual of Mosses
author has. naturally, had recourse to the various works of Sullivant, Les-
quereux & James, Barnes SC Heald, Grout, and others among the American
bryologistb, and, among the European bryologists, particularly Dixon and
Jameson. In the determination of the Sphagnums the works of Warnstorf,
Roth, and Braithwa te were found particularly useful, while in the treatment of
synonymy the main reliance has been placed en the Index Bryologicus of
E. G. Paris.
In nomenclature the rulings of the International Botanical Congress, held
in Brussels in 1910, have been followed, taking as the starting point the Species
Muscorum of Hedwig (1801) and the three subsequent "Supplements" by
Schwaegrichen, Hedwig's having been the first comprehensive work to deal
with the messes in a modern way. In the present Manual the principle of
priority has been followed without exception, dating from Hedv»'i[T, and a few
new combinations have been found necessary. Plant names which have been
adopted from pre-Hedwigian sources without important changes in nature or
in status are indicated by a double citation of authors, the pre-Hedwigian
author being cited first, followed by a comma, and then the name of Hedwig
or Schwaegrichen or of the post-Hedv.'igian author, as the case may be. In
case the name of the plant has been derived from pre-Hedwigian sources, but
has been used in a different rank or, in the case of species, has been trans-
ferred from one genus to another, the name of the pre-Hedwigian author has
been enclosed in brackets.
So far as it has been possible to do so, the descriptions of the various
species are based entirely on specimens collected in the region covered by the
Manual. Where specimens of species reported as occurring in the region or
thought likely to be eventually discovered in the region, have not been avail-
able for description, the description has been in part compiled and in part
drawn up from specimens from other regions. It has been the aim to repre-
sent by original drawings, completely and in considerable detail every species
of which specimens collected in the region of the Manual have been available.
In the list of specimens, which, in the Manual, follows the description of each
species, the particular specimien figured has been so indicated and the fact that
the specimen is in the Herbarium of the Carnegie Museum. All drawings are
the work of the author alone, and, with the exception of a few of the larger
habit sketches, all drawings have been first traced by means of the camera
lucida, thus insuring a reasonable degree of accuracy in the relative position,
shape, and size of the various structures figured. The drawings of most of
the dissections have been made from permanent glycerine-jelly mica-covered
slides which are to be found in the Herbarium in the proper pocket with
the specimen.
Special acknowledgment should be here made to Dr. W. J. Holland,
Director of the Carnegie Museum, without whose generous and kindly support
the collections could not have been made and properly studied nor the Manual
prepared. To Professor J. C. Fettermann, of the University of Pittsburgh, is
due many thanks for suggestions and criticism, and to Mrs. O. E. Jennings
Preface to the Second Edition
is due much credit for assistance in the collection of specimens, in the prepara-
tion of the manuscript, and in the arrangement of the figures on the plates. —
O. E. Jennings, Carnegie Museum, September, 1912.
Preface to the Second Edition
In the thirty-eight years which have elapsed since the publication of the
first edition of this Manual, the mosses of the region have been much more
extensively collected and twenty-two species have been found which were not
previously known to occur in western Pennsylvania. Various friends, includ-
ing a number of the author's former students, especially Mr. Sidney K. East-
wood, have by their field-work extended our knowledge of the moss-flora of
our region. Very special acknowledgment is expressed to Mr. Charles M.
Boardman for contributing the results of his extensive field-work adding much
to our knowledge of the distribution of various species, and for adding several
species to the known flora of our region, including five species of Sphagnum.
To him is due in large part the revision of the treatment of the Sphagnales.
In the preparation of this second edition of the Manual, particularly with
reference to nomenclature and general range of species, frequent reference has
been had to the second edition of Engler fii Prantl, Die Natiirlichen Pflan-
::enjamilien (1924-1925); Brotherus, Die Laubmoos Fennoskandias (1923);
various articles in The Bryologist; and particularly the Moss Flora of North
America North of Mexico, A. J. Grout, Vols. I-III, (1928-1940). For
Sphagnum, further reference has been had to Sherrin, W. R., Ari Illustrated
Handbook of the British Sphagna (1927) and to Andrews, A. L., Sphag-
naceae, in Vol. 15, North Ajnerican Flora.
During the years which have passed since the first edition of the Manual
was prepared, bryology has advanced to the point that many changes in nomen-
clature have become necessary in order to bring this second edition up to date.
As in the first edition, all drawings are original and from specimens collected
in western Pennsylvania. — O. E. Jennings, Carnegie Museum, June 1, 1950.
To my wife
for assistance in field and herbarium
and for her never-ending encouragement
this book is affectionately dedicated
CONTENTS
Introduction 1
Directions for collecting, preparing and preserving specimens of mosses .. 7
Abbreviation of signs used in the manual 10
Analytical key to the genera of mosses of western Pennsylvania 10
Systematic Treatment
Order I. Sphagnales - 20
Order II. Andreaeales - 42
Order III. Bryales - 44
Acrocarpi. Fc-.milies 1-23 44-172
Pleurocarpi. Families 24-34 172-307
Glossary of bryological terms used in the manual 308
Explanation of the method of lettering the figures ., 324
Plates I-LXXII 325-326
Index 313
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County Map of Western Pennsylvania and Adjacent Areas. Mountain
Uplands, mostly 2,000-3,000 ft. altitude, swing across the southeastern quarter, with the
Ridge and Valley area to the southeast. The High Plateaus, mostly 1,800-2,400 ft. alti-
tude, occupy the northeastern area; plateaus cut by narrow deep valleys. The Glaciated
area in the northwest is bordered southwards by much dissected hilly country. A narrow
belt of sandy Lake Plain borders Lake Erie.
A Manual of the Mosses of Western Pennsylvania
and Adjacent Regions
Introduction
In a work containing keys and descriptions, so arranged as to make easier
the identification of the mosses of any region, it is desirable that a brief sketch
of the general life history of the mosses be included. In such a sketch it is
not necessary to enter upon a discussion of the many details of minute struc-
ture and behavior which, although interesting and important in themselves
and also for the light thus thrown upon genetic relationships, are yet of but
little practical value in a systematic manual where an easy and quick deter-
mination of the identity of the plant is the primary aim.
Speaking broadly, the life history of m.ost of our mosses begins with a
minute single-celled spore, usually spherical in shape, which, under suitable
conditions, germinates and grows out as a slender thread or filament, which
upon further grovv'th may form a matted felt like layer, or may flatten out
into a more or less lobed body spoken of as a thallus, or may simply form a
solid cell mass, sometimes consisting of but a few cells. In either case the
structure resulting from the growth of the germinated spore is termed the
protonema. The protonema usually gives rise to buds, which in most mosses
grow to be the green leafy shoots which are ordinarily known as moss plants,
after which the protonema usually disappears. In a few of the mosses the
protonema persists indefinitely as a green felt-like layer on the soil or other
substratum. The stems of the green shoots resulting from the growth of
protonemal buds usually send out hair-like rhizoids which function as roots in
holding ihe plants in place and sometimes act as absorbing organs. The
leaver on these green shoots are sessile and with the exception of the midribs
(costae) are almost uniformly of but one cell in thickness.
This whole phase in the life-history of a moss, beginning with the spore
and. including the protonema and the leafy shoot, is spoken of as the gameto-
phyte or se.\ual generation. The gametophyte is a sexual plant in that it
bears, in definite clusters surrounded by modified leaves called per^chaetial
leaves, the reproductive male and female organs which give rise respectively
to the sperm and egg. These clusters of reproductive organs surrounded by
more or less modified perichstial leaves are known as pertchaetia. When the
sperms and eggs are borne either in the same perich^tium or in different
perichaitia on the same plant the plant is spoken of as monoicoiis, but when they
are produced upon different plants, dioicous. When only male organs are in the
cluster the surrounding modified leaves are known as peri gonial leaves.
The sperms are borne in a globose or more or less club-shaped sac, usually
mounted upon a stalk, and this sac is termed the antheridmm. When ripe the
2 American Midland Naturalist Monograph No. 6
antheridia absorb water and the thin wall, consisting of but a single layer of
sterile cells, is ruptured, thus liberating the mass of fertile cells, each of which
immediately develops into a sperm. Each sperm consists of a more or less
oval or club-shaped and curved body, always free-swimming by means of two
long slender cilia attached at one end of the body.
The egg is borne in a special organ termed the archegonium. The arche-
gonium is usually more or less stalked and is differentiated into a swollen
basal portion termed the venter, which contains the one fertile egg cell, and
the more slender tapering neck, terminating the archegonium above and con-
taining an axial row of sterile cells termed the canal cells, the basal one of
which rests directly upon the egg cell. When the archegonium becomes ripe
the canal cells break down into a slimy mass of protoplasm, some of which may
escape at the tip of the neck. Sperms are attracted in some m.anner by the
slimy protoplasm thus escaping if there is a sufficient film of moisture present
so that they may swim about in the perichaetium or on the surface of the
plant. Having reached the apex of the archegonium the sperms may enter the
canal left open by the disintegration of the canal cells and eventually one of
the sperms may reach the eggs and, uniting with it, brings about fertilization.
After fertilization the egg immediately begins development as the sporo-
phyte but remains enclosed in the venter of the archegonium, which to a con-
siderable extent expands with the development of the young sporophyte but is
finally ruptured and usually carried upward on the tip of the sporophyte,
where it is then known as the calyptra or hood. The ultimate end of the
sporophyte is the production of spores which arise entirely by division of cells
and are thus known as asexual cells. The sporophyte is usually almost devoid
of chlorophyll and it develops at its base an absorbing organ termed the foot
through which its food is obtained from the gametophyte. The sporophyte
usually develops more or less of a stalk which is termed the seta and which
bears at the apex a globose to more or less elongated capsule in which the
asexual spores form. The method of opening (dehiscence) of the capsule and
the structures often associated with the dispersal of the spores are varied and
are so characteristic for the various systematic groups and species that the cap-
sule becomes highly important for the correct systematic placing of the plants.
Of the mosses there are to be distinguished three well-marked orders
known as the Sphagnales, the Andreaeales, and the Bryales. The order Sphag-
nales comprises the one genus Sphagnum. These mosses are known as peat
mosses or bog mosses, their most characteristic habitat being bogs and the
margins of ponds and small lakes. The general color is grayish green, the
stems are usually erect in dense tufts or mats and bear at intervals fascicles of
short and slender branchlets. The capsules are usually more or less chestnut-
colored and globose, while the leaves possess a peculiar and characteristic
structure consisting of a meshwork of slender green cells enclosing inflated
hyaline cells whose walls are more or less porose.
The Andreaeales contain the one genus Andreaea, all being small tufted
mosses growing on siliceous rocks in mountainous regions. The capsule splits
Jennings: Manual of Mosses 3
open by four, lateral, vertical slits which, however, do not reach the apex.
The Bryales comprise by far the greatest number of the mosses. The
capsule in the Bryales varies from globose to ovate or pyriform or elongated
cyhndric. The cells which give rise to the spores are known collectively as
sporogenous tissue and this tissue occupies but a small portion of the volume
of the capsule, being arranged in the form of a hollow tube or cylinder vertically
placed and open at both ends. The sterile tissues occupying the hollow part
of this tube constitute the cohunella. The outer wall of the capsule usually
contains more or less green chlorophyll and the middle portion of this wall is
more or less loosely arranged and contains hollow spaces. The capsule is
covered by an epidermis, perforated by stomata in most mosses. The stomata
are usually most highly developed on the rounded or tapering base of the
capsule which is often more cr less distinct and is known as the collum or neck-
In the ripening of the capsule the sterile tissues of the wall and of the columella
largely disappear, leaving the capsule filled with a mass of spores. In some
species the thin wall of the capsule bursts irregularly, this type of dehiscence
being known as cleistocarpous. In other species the top of the capsule sep-
arates as a ltd or operculum. The separation of the lid is often facilitated by
the modification of a series of epidermal cells termed the annulus, which usu-
ally becomes highly hygroscopic and is often deciduous. The sterile tissues
immediately beneath the lid are usually more or less highly modified to form
a single or double series of pointed structures known collectively as the
peristome. The pointed structures constituting the outer series in the double
peristome or the single series in a simple peristome are known as teeth, while
the inner and more delicate series of the double peristome are known as
segments. Between the individual segments in many species of mosses are very
delicate hair-like structures known as ctlia. Sometimes the cilia are in groups
of two or more alternating with the segments. The peristome is usually very
hygroscopic, curling inward and closing the mouth of the capsule in damp air
and opening outward and allowing the free dispersal of the spores in dry air.
Species whose capsules stand vertically are not so likely to have well-developed
peristomes as are species whose capsules are inclined or vertical, this variation
corresponding to the need for the regulation of spore dispersal.
The Sphagnales are most abundant in the cooler parts of the North Tem-
perate Zone, often constituting there large tracts of vegetation. By their
aquatic or semi-aquatic manner of life and their apical method of growth,
dying away below as they grow upward, they tend to form great tufts or
mats, often completely filling depressions and bogs and by the accumulation
of the encircling mats around ponds and small lakes tending to fill them also.
The mats hold water like a sponge and, being somewhat antiseptic, the dead
portions below the mat do not decay but become converted into peat, which,
especially in certain parts of Europe, has served a very important purpose as
fuel. A considerable number of Sphagnum bogs and Sphagnum cranberry
glades of limited area occur in the mountains of central Pennsylvania. The
only other bogs of any considerable extent in our region are those m the
4 American Midland Naturalist Monograph No. 6
northwestern part of Pennsylvania, particularly in the Pymatuning Swamp,
between Linesville and Hartstown, in Crawford County. Here, in places, the
Sphagnum and Tamarack formerly reigned supreme for acres in extent and
the peat deposits are apparently quite deep.* As may be seen in the treat-
ment of Sphagnum in this Manual, the Pymatuning Swamp has yielded a
goodly share of the species reported for our region.
The Andreaeales are represented in our region at only one station, although
they occur in the mountains both to the south and to the northeast. It is not
unlikely that at other stations Andreaea will yet be found to occur in the
mountains of central and western Pennsylvania upon sandstone ridges.
The Bryales include by far the greater number of moss species found in
Pennsylvania. Western Pennsylvania as a whole offers quite a variety of
habitats and its moss flora is fairly large, although, considering the area cov-
ered, there is a noticeable lack of certain species more or less peculiar to high
elevations, to outcrops of limestone, and to low-lying marshes and river swamps.
The northwestern part of Pennsylvania, in a triangular area extending as
far south as Beaver County and as far east as Warren County, was worked
over by the ice in the Glacial Period and is still in a rather youthful stage of
erosion, with a number of small lakes and ponds and considerable areas of
poorly drained lands. Occasional Sphagnum bogs occur here as well as
swamps along the flood-plains of some of the streams. Such conditions offer
suitable habitats for a number of aquatic and swamp-inhabiting species of the
Bryales which are not to be found at all or are quite rare in the rest of western
Pennsylvania. Presque Isle, near Erie, is a sandspit of about six miles in
length and over a mile in width at its outer extremity- and, containing as it
does a variety of ponds, lagoons, woodland swamps, marshes, and dry woods,
it affords certain habitats which are not duplicated anywhere else in our region.
The remainder of the region covered by this Manual is thei rath:r charac-
teristic hilly country of the Allegheny Plateau, ranging in altitude from about
700 feet above the sea, along the flood-plain of the Ohio River, to about 3,200
feet above the sea in the mountains of the southeastern part of our region. In
the western, southwestern and northeastern parts of our region the general
topography is that of an elevated tableland in an active state of erosion, the
rocks being largely sandstones and shales, and mainly non-calcareous. There
are many steep valleys and precipitous rock exposures with a minimum of
swampy areas or ponds. The flood-plains which have developed alona the
Ohio River, the Monongahela River, the lower Allegheny River, and the larger
tributaries of these streams have been so largely disturbed by the activities
of man that they now offer but few opportunities for collection in what must
have once been habitats rich in Bryales.
As the smaller streams in western Pennsylvania are ascended, however,
* The largest and most interesting part of this swamp has since been cleared and
Hooded to form the Pymatuning Reservoir below Linesville and the Wildlife Rufuge above.
Jennings: Manual of Mosses 5
the valleys often rapidly narrow to a more or less steep rock-walled canon
where erosion is highly active. In the narrow valleys the forest covering has
not been very extensively disturbed by man and the damp, cool, shaded
habitat with varying substrata of decaying wood, rich loam, shaly soil, bare
rock, or living bark, conduces to a rich and varied flora of the Bryales. Above
this area of active erosion there will usually be found, in the headwaters of the
streams, a region which has remained largely unaltered from a former ad-
vanced stage of physiographic development and which is characterized by wide
valleys with gently sloping soil-covered sides rising to broadly rounded and
soil-covered hills. These rounded hills, whose height above the bottoms of
the adjacent rounded valleys is rarely more than 300 to 350 feet, are in many
places still covered with the native forest consisting mostly of the white oak,
but the moss flora of these forests is poor.
Good collecting ground for the Bryales is also to be found in the moun-
tams of the eastern and southeastern parts of the region covered by this
Manual, particularly in the steep and rocky gorges which have been cut
through the sandstone ridges by the larger streams. Perhaps the best collect-
ing ground for the Bryales in our whole region is to be found in the vicinity
of Ohio Pyle, in Fayette County, where the Youghiogheny River and its
larger tributaries have cut out wild and rocky gorges sometimes a thousand
feet or more in depth. Somewhat similar and perhaps but little inferior to
the Ohio Pyle region are localities along the gaps cut through the ridges by the
Conemaugh and Loyalhanna rivers and the eastward-flowing Juniata and the
West Branch of the Susquehanna River.
The northeastern part of our region consists of an elevated plateau in part
still swampy or boggy but in part considerably dissected. The flora is largely
of a northern type, much of it having been originally of the northern mixed
hardwood forest of hemlock, birch, beech, and maple, but with areas more or
less dominated by white pine. This plateau region is a rich moss collecting
ground both for the Bryales and the Sphagnums.
In the present edition much the same sequence of families has been ob-
served as in the first edition. In classifying mosses, both gametophyte and
sporophyte characters are of value, and it is practically impossible to arrange
them in a satisfactory sequence. An arrangement based on either gametophyte
or sporophyte characters will cut across the other, and current practice varies
considerably. The old practice of grouping the mosses into acrocarpous, with
the sporophyte borne terminally on either the stem or evident branches, or
pleurocarpous, with the sporophyte borne laterally, is not accurate, but it is
very convenient for purposes of identification, particularly since it is desir-
able to identify so many specimens in which the sporophytes are not in proper
condition to study. As a matter of convenience these two groups are recog-
nized in the present work.
The following twenty- six species have now been definitely added to the
known moss-flora of western Pennsylvania since 1912 and are also described
and illustrated in this second edition.
American Midland Naturalist Monograph No. 6
Andreaed Rot bit
Astomum Muhlenbergianum
Atrichum crispum
Brachythecium oxycladon
Bruchia Sullivantii
Discelium nudum
Drepanocladus aduncus
Entodon compiessus
Ephemerum serratum
Grimmia pilifera
Hookeria acutijolia
Mnium spinulosum
OrthotTtchum obtusifolium
Plagiothecium Roeseanum
Astomum Muhlenbergianum
Ptychomitrium incitrvum
Rhacomitrlum heterostichum
var. gracilescens
Raphidostegium marylandicum
Rhytidium rugosum
Sphagnum fuscum
Sphagnum Girgensohnii
Sphagnum papillosum
Sphagnum Wulfiamim
Splachnum ampullaceum
Tetraplodcn angustatus
Thuidium pygmaeum
For some of the mosses included in the first edition no satisfactory material
from western Pennsylvania was then available for illustration. Specimens of
twenty of these species have since been collected in our region and have now
been illustrated in the present edition, as follows:
Bryhnia noyae-angliae
Dicranella rufescens
Dicranum rugosum
Drepanocladus fiuitans
Encalypta streptocarpa
Entodon compressus
Fissidens hyalinus
Fontinalis gigantea
Hygroamblystegium orthocladon
Isopterygium deplanatum
Isopterygium elegans
Isopterygium geophilum
Mnium hornum
Octodiceras debile
Plagiothecium Roeseanum
Pleuridium subulatum
Sphagnum squarrosum
Stereodon pratensis
Tortula papillosa
Trematodon ambiguus
Including the above additions the present edition contains 18 additional
plates covering forty-six species and bringing the number of individual figures
to a total of three thousand five hundred and seventeen.
In addition to the 243 kinds of mosses described and illustrated by original
drawings by the author from collections made in western Pennsylvania there
are included more than 100 descriptions of sf)ecies reported for the region by
various collectors years ago or which are known to occur in territory adjacent
to western Pennsylvania. Some of the species reported for our region by T. P.
James, Thos. C. Porter, and others are p>erhaps not now to be found here, while
others known from adjoining regions will undoubtedly eventually be found
here, although, unlike some of these localities, we do not have extended areas
of limestone outcrops such as occur to the east and west, on which certain
calciphilous species commonly occur.
The following new combinations occur in the second edition.
Sphagnum palustre var. squarrosulum (Nees &: Homschuch )
Sphagnum palustre var. brachycladum (Schliephacke)
Sphagnum plumulosum i. viride (Wamstorf)
Atrichum undulatum var. allegheniense (Jennings)
Eurynchium pulchellum var. praecox (Hedwig)
Pohlia nutans var. triciliata
Jennings: Manual of Mosses 7
The rotal number of genera, species, and varieties recognized in this
Manual as having been collected or authoritatively reported in western
Pennsylvania is as follows:
Genera Sfjecies Varieties
Sphagnales 1 24 7
Andreaeales 1 2 —
Bryales Ill 260 26
Total 113 286 33
Directions for Collecting, Preparing, and Preserving
Specimens of Mosses
For the benefit of those who may not be familiar with the usual methods
of collection and preservation of bryological specimens the following notes
may be of use.
Specimens of Bryales should be collected in fruit (rip>e sporophytes) as
far as possible. Specimens of Sphagnum are desirable in fruit, but deter-
mination is best made in this group from the vegetative characters. Specimens
of mosses when collected should be placed at once in envelopes or other suit-
able paper pockets and the data of collection, especially habitat, should be
written upon the envelope. Many collectors prefer to number the envelope
and under the corresponding number make note of the data in a note-book.
Collections may be carried home in a basket or regular tin collecting case and,
if carefully placed in the envelopes in the first place, the specimens need not
be taken out of the envelopes but the envelopes should be placed between
blotting papers or newspapers and subjected to a slight weight and so placed
that they will soon dry. A few books or two or three bricks are usually suffi-
cient weight for drying a package of mosses. Too much weight should be
guarded against, as the habit of the plant, i. e., the position assumed by leaves,
branches, etc., is often a great help in determining the species, and, if too much
weight is used in drying, the specimens will be so flattened as to destroy
these characters.
When dry, the specimens may be placed in paper pockets made from a
rectangular piece of paper by folding up the lower part of the rectangle to
within about one inch of the upper edge and then folding down this inch flap
over the first flap. The two ends should now be folded backward for about
one inch each and the pocket is then complete and ready for the reception
of the moss. The regulation method in most larger herbaria is to glue this
pocket in the middle of the back, midway between the two folded ends, to a
so-called "herbarium sheet" which is uniformly of white stiff paper measuring
115/2 by 16^2 inches. For small private collections smaller sizes are sometimes
used. On the lower right-hand corner of this sheet is written the name of the
species, and the number of specimens which such a sheet will accommodate is,
of course, restricted only by the space occupied by the pockets. The label for
8 American Midland Naturalist Monograph No. 6
each specimen should be fastened to the narrow flap at the upper edge of the
pocket and should always contain the name of the species, the exact locality
and habitat of the specimen, the name of the collector, and the date of the
collection. If material has been collected in sets for distribution the label
should so state and a number should be assigned to the various species so that
duplicate specimens reaching different botanists may be definitely correlated.
It is often advisable to add to the label also the name of the botanist who
identified the specimen, especially if he be a sp>ecialist.
For any extended study of the mosses, either Sphagnales or Bryales, it is
practically necessary to have at hand besides a pocket lens of some sort, a dis-
secting lens and a comjwund microscope. A dissecting lens may be rigged up
by providing some sort of a frame for holding the ordinary pocket lens at the
right distance above the table. This can be done by some such simple con-
trivance as knitting needles and corks, in the absence of anything better. The
writer has found very satisfactory the ordinary dissecting stand, which may be
obtained from any dealer in scientific apparatus, the stand preferably fitted with
a rack-and-pinion adjustment for focusing the lens. The writer has used with
good results a doublet lens (three-quarter inch) magnifying about four di-
ameters and a one-fourth inch aplanat lens magnifying about seven diameters.
The compound microscope should be fitted with a one-inch and also preferably
a two-inch eye-piece and the customary two-thirds and one-sixth objectives. A
sub-stage condenser is a great convenience and should be provided with dia-
phragms both above and below.
In preparing a moss for microscopic study the writer proceeds as follows:
A portion of the specimen, usually consisting of a whole plant, is selected and
soaked in water until it is soft and relaxed. A thin square of mica an inch
or more in width is prepared and placed on an ordinary glass miscroscope
slide, and upon it is placed a drop of a ten per cent solution of glycerine in
water which is kept already prepared in a small bottle with a medicine dropper
fastened into the stopper. The glass slide with the mica square and solution
in position are placed on the stand of the dissecting microscope. With small
forceps and with the aid of needles mounted in wooden handles the moss is
now carefully dissected and the parts suitably disposed on the mica square
in the film of ten per cent glycerine. It is usually best to place on the mica
square some thin cross-sections of the stem of the moss, cut with a scalpel or
kriife or fine scissors, some stem-leaves, some branch-leaves, some perichajtial
leaves or, better, the whole perichaetium dissected apart but not widely scat-
tered, and then the capsule so dissected as to show a patch of the epidermis
from the base of the capsule, the annulus, the peristome, both outer and
inner if they are present, and the spores.
Another thin mica square is now selected a little smaller than the first one
used and upon it is placed a small chunk of glycerine-jelly, which is melted
by holding the square in the forceps over a suitable source of heat — the writer
holds the square over the electric bulb of his desk light. The glycerine when
melted is smeared over the surface of the mica, which is then inverted and
Jennings: Manual of Mosses 9
quickly but carefully placed on the square on which the dissections are dis-
posed. To prevent the dissected objects from changing their position too
much, and to obviate the inclusion of air-bubbles, it is best to lay the square
which is to serve as cover so that one edge only comes into contact with the
other square and then let the cover settle down gradually, thus driving the air
out in front of the gradually advancing line of contact of the mica and mount-
ing medium. The slide is now ready for study under the compound micro-
scope and after this it may be placed in the paper pocket along with the speci-
mens from one of which the dissections were made. In order to insure greater
permanency of the slide, as thus made, some workers advocate sealing the slide
by running a little ring of Canada balsam* around the edge of the smaller
mica square, thus keeping the air away from the glycerine jelly and preventing
any further drying out. The object of placing the dissections in the ten per
cent solution of glycerine is to gradually allow the dissections to accommodate
themselves to increasing density of solutions; if the dissections were transferred
immediately from pure water to the melted glycerine jelly there would in
most cases be much shrinkage and curling, thus spoiling the slide for purposes
of study. In a few cases even the transferance from water to ten per cent
solution and thence to the jelly is too great a change and in such cases it is
necessary to pass the dissections through a series of solutions of increasing
glycerine per cent, up to a strong solution, before using the glycerine-jelly.
Another way is to place the dissections in weak glycerine solution and keep
adding more solution as the water evaporates from the first, thus gradually
increasing the density.
Another method has been described by Steyaert, R. L. (Science 105: 47-48.
1947) . Chloraphenol, consisting of two parts chloral hydrate and one part
phenol crystals,** is heated till liquefied and then kept in dropping bottles. This
liquid mixes with either water or Canada balsam. Dried material may be
placed and dissected directly in a mixture of two drops of chloraphenol to one
of balsam dissolved in xylol. After this, place the cover slip on and gently
heat to evaporate the chloraphenol, in the meantime placing droplets of balsam
at the edge of the cover slip so that it may flow under and ensure a sufficient
amount of balsam in the mount. The mount may then be sealed (ringed)
with varnish.
Or, material may be dissected in water which may then be drawn out by a
blotter while chloraphenol is run under the slip from the opposite edge.
Slightly warming the slide helps to get rid of air bubbles. The chloraphenol
may then similarly be drawn out with a blotter while balsam is drawn under
from the other side of the slip. When thoroughly filled with balsam the amount
may be permanently ringed with varnish.
* Before applying such a sealing medium, the edges of the slips should be dry.
Other sealing media often used are gum dammar, shellac varnish, gold size, or m.^rine
glue.
** Care must be taken not to let the fjhenol come into contact with the skin. It will
(a use a very severe bum.
10 American Midland Naturalist Monograph No. 6
Abbreviations and Signs Used in the Manual
cm Centimeter, equals the one-hundredth part of a meter, or about two-fifths of an inch.
mm — Millimeter, equals one-tenth of a centimeter; about one-twenty-fifth of an inch.
C.M.B.— Charles M. Boardman.
D.A.B.— D. A. Burnett.
S.K.E. — Sidney K. Eastwood.
G.K.J. — Grace K. Jennings (Mrs. O. E. Jennings).
O.E.J. — O. E. Jennings.
J.A.S.— John A. Shafer.
- The hyphen used between figures or betA-een words denotes either an intermediate state
or a variation from one to the other extreme.
Analytical Key to the Genera of Mosses
OF Western Pennsylvania
Order I. Sphagnales
Whitish mosses with fasciculate branches, mostly bog plants; leaf-cells of two kinds,
— large hyaline ones separated by narrow chlorophyllose ones; ecostate; opjerculate but
with no peristome Sphagnum, p. 20
Order II. Andreaeales
Dark colored or blackish, very fragile, alpine or subalpine plants growing in
cushions on granitic, hard sandstone, or slaty rocks; either costate or ecostate; leaf-
cells small and quite opaque; capsule dehiscing by four longitudinal slits, the valves
remaining united at the apex until quite old and empty Andreaea, p. 43
Order III. Bryales
Leaves various but not sphagnoid, costate or ecostate; capsule dehiscing irregularly
or, more often, by a deciduous operculum, often furnished with a peristome, never
four-valved as in Andreaea, plants largely green* I
H aplohymenium irisle is not known to fruit in the United States.
I. Sfxjrophyte borne at the apex of the main stem, sometimes appearing lateral by
the growth of a branch A. (AcROCARPi) p. 44
I. Sporophyte borne at the ajDex of a usually short lateral branch, or appearing
axillary B. (Pleurocarpi) p. 172
I. Sporophytes on rather vigorous, erect, often dendroid secondary branches which
rise from a rhizome-like primary stem see Thamnium or Climacium
A. AcROCARPI
1. Capsule non-operculate (or lid not readily separating) 2
1. Capsule operculate 10
2. Green protonema persistent; plants fruiting in autumn, minute
Ephemerum, p. 1 13
2. Green protonema persistent; plants fruiting in spring Acaulon, p. 87
2. Green protonema not persistent, plants fruiting mainly in spring 3
* Hookeria acutifolia has leaves 4-5 mm long, drying very thin, soft, fluffy, and pale
yellow or whitish.
Jennings: Manual of Mosses 11
3. Spores few, about 16 to 20, smooth, about 0.2 mm in diameter; leaves linear-
lanceolate, with percurrent costa Archtdium, p. 44
3. Spores numerous, rarely exreedrng O.Ci mm in diameter 4
4. Leaf-margms plane or involute 5
4. Leaf-margins more or less revolute - 9
5. Capsules pyriform, with a distinct neck 6
5. Capsules globose to ovoid '
6. Green protoncma occasionally abundant; neck none; capsule acute
SpoiUdera, p. 49
6. Green protonema sparse; usually none; neck more or less well developed;
capsule more or less rostrate Bruchia, p. 46
7. Leaves crisped when dry, strongly papillose on both sides; operculum rudi-
mentary but persistent .Astomum, p. 78
7. Leaves not crisp>ed when dry, smooth 8
8. Calyptra cucullatc (mitrate in Pleuridium palustre); leaves linear-lanceolate
to lanceolate-subulate Pleundium, p. 48
8. Calyptra campanulate; leaves lanceolate-ovate to lanceolate-obovate, dentate
or serrate Physcomitrella, p. 116
9. Capsule apiculate; leaves papillose Phascum, p. 88
9. Capsule not at all or faintly apiculate; leaves smooth Acaulon, p. 87
10. Protonema long persisting; stem-leaves minute; plant looks like a grain of
wheat sitting in a tuft of bristles Diphyscium, p. 155
10. Protonema brownish, persistent; plant looks like a shriveled, brownish grain
of wheat sitting obliquely on a stout seta about 1 cm high
Buxbaumia, p. 156
10. Protonema persistent; plants practically stemless, leaves ecostate; calyptra
splitting down one side and usually remaining attached to the seta
Discelitim. p. 113
10. Protonema jsersistent; plants practically stemless (1-2 mm); costa strongly
lamellate in apical p)ortion of the leaf Pogonatum, p. 165
10. Protonema not persistent; calyptra not as in Discelium. If with hypophy-
sis see Splachnaceae, p. 110 11
11. Peristome teeth none, or if present articulate, at least at base 12
11. Peristome teeth not distinctly articulate 60
12. Peristome present, sometimes imperfect 13
12. Peristome none 52
13. Leaves distichous, dorsally winged and clasping at the base 127
13. Leaves not distichously' clasping and dorsally winged 14
14. Leaves consisting of costa only, outer cells large and empty, inner small
and chlorophyllose Leucobryum, p. 66
14. Leaves with a lamina consisting mainly of one layerf of more or less uni-
form cells 15
15. Peristome single, 16- or 32-toothed; teeth without a median longitudinal line
on the exterior face 16
15. Peristome double, with 16 outer teeth and an inner variously segmented or
almost lacking membrane: teeth mostly with a median longitudinal line on
the exterior face 37
16. Capsules more or less octagonal, the angles with differentiated cells, when
dry 8-s:riate and furrowed Rhabdoweisia, p. 57
16. Capsule not octagonal, or, if plicate, the cells uniform 17
17. Exterior face of teeth longitudinally striate. (See also Ditrichum) 18
17. Exterior face of teeth smooth or papillose, not longitudinally striate. (See
also Seligeria ) 24
18. Leaves with differentiated, often enlarged, hyaline or brownish alar cells .... 12
18. Leaves without differentiated alar cells 19
19. Leaf-cells strongly and coarsely papillose. (See also Oncophorus)
Oreowfiria, p. 46
12 American Midland Naturalist Monograph No. 6
19. Leaf-cells smooth 20
20. Peristome of 16 broad solid teeth, sometimes somewhat bifid at the apex,
smooth and strongly hygroscopic Seligeria, p. 53
20. Peristome of 16 narrow, prolonged, more or less two-parted, minutely
striate or papillose teeth 21
21. Capsule cemuus, with a long and inflated neck; teeth cleft or perforate
Trematodon, p. 47
21. Capsule erect or inclined; neck short or* none; teeth cleft to the middle into
two (or three) filiform divisions; leaves not crisped when dry
DicraneV.a, p. 54
21. Capsule erect to horizontal; neck none; teeth cleft about half way; leaves
crisped when dry Oncophorus, p. 58
22. Monoicous; leaves tufted; capsules strumose, costa with distinctly hetero-
geneous tissues Oncophorus, p. 58
11. Dioicous; leaves flexuose-spreading or secund; capsule more or less distinct-
ly strumose 23
23. Basal auricles of leaves large and inflated; seta cygneous; peristome teeth
divided % to base Dicranodontium, p. 64
23. Basal auricles of leaves small or none; seta not cygneous; peristome teeth
divided about half way Dicranum, p. 59
24. Peristome distinctly twisted; teeth 32 25
24. Peristome not distinctly twisted; teeth 16 but often deeply^ cleft 27
25. Leaves large, oblong to sub-spatulate; costa with two median guides
Tortula, p. 91
25. Leaves small and narrow, linear-lanceolate; costa with from 4-8 median
guides 26
26. Leaf-margins plane, not revolute; cells papillose Tortella, p. 83
26. Leaf-margins revolute, at least below; cells smooth or papillose
Barhula, p. 85
27. Dark green to brownish or blackish mosses on rocks or sometimes trees; jseri-
stome single or none 28
27. Green plants, not blackish nor very dark 30
28. Calyptra campanulate, plicate, covering most of urn
OrthotTtchum, p. 103
28. Calyptra covering about half of the capsule, split and plicate; sfxsres large
Ptychomitrium, p. 95
28. Calyptra large cucullate; spores very large; teech wide, truncate; annulus
none Drummondia, p. 103
28. Calyptra short, not plicate; annulus usually present; spores medium size to
small 29
29. Teeth sub-entire, cribrose or irregular cleft in upjser half Grimtma, p. 95
29. Teeth cleft to near the base into filiform segments Rhacomitrium, p. 98
30. Calyptra mitrate, not folded nor torn, completely covering the capsule;
capsule erect; leaves ligulate to spatulate Encalypta, p. 93
30 Calyptra cucullate 31
31. Teeth merging below into a more or less continuous basal wall or membrane .. 32
31. Teeth not merging below into a continuous membrane 35
32. Upper leaf-cells papillose; costa with one or two rows o^ larger cells ven-
tral to the 2-4 median guides Desmatodon, p. 90
32. Upper leaf-cells papillose; costa with 4-6 median guide cells and a well
develofjed stereid region on ventral side Didymodon, p. 84
32. Leaves mostly smooth; peristome-teeth mostly medium length to long 33
33. Capsule inclined, distinctly plicate when dry; leaf-cells roundish-quadrate above
Ceratodon, p. 52
33. Capsule erect, nearly or entirely smooth when dry; leaf-cells elongated (2:1 or
more) above 34
34. Leaves glaucous (Saelania)
Jennings: Manual of Mosses 13
34. Leaves glossy-green ; -- Ditrkhum, p. 49
35. Plants living on trees Drummondia, p. 103
35. Plants living on earth or rocks ?o
36. Leaf-margins entire, distinctly involute above Weisia, p. 79
36. Leaf-margins minutely crenuh.te, plane Trichostomum, p. 82
37. Liner peristome distinct (or adhering to inner surface of teeth), lacking a
basal membrane 38
37. Inner penstome furnished with a distinct, often carinate basal membrane 41
38. Calyptra cucullate Funaria, p. 118
38. Calyptra mitrate or campanulate 39
39. Calyptra smooth, not plicate, extending below the capsule Encalypta, p. 93
39. Calyptra only partially enclosing the capsule, plicate and usually pubescent
40
40. Leaves crisped (except U. americana) when dry, base oval, neck of cap-
sule with stomata superficial Ulota, p. 107
40. Leaves not with an oval base and not crisped when dry; neck of capsule
with mostly immersed stomata Orthotrichum, p. 103
41. Capsule ribbed or furrowed when dry 42
41. Capsule smooth, not ribbed when dry 45
42. Capsule oblong, furrowed when dry; inner peristome of 64 filiform cilia
in groups of 4's Ttmmia, p. 154
42. Capsule ovoid-cylindrical; inner and outer peristomes of equal length
Aulacomrtium, p. 144
42. Capsule cylindric; inner peristome half as long as filiform teeth
Encalypta, p. 93
42. Capsule ovoid to sub-globose; inner peristome shorter than the outer 43
43. Cilia well developed Philonotis, p. 150
43. Cilia usually rudimentary or absent (see also Philonotis) 44
44. Leaf-cells papillose or mamillate Bartramia, p. 148
44. Leaf-cells smooth Plagiopus, p. 148
45. Upf>er leaf surface papillose Timmia, p. 154
45. Upper leaf surface not papillose 46
46. Segments 2 to 3 times as long as the teeth; cilia rudimentary
Meesia, p. 146
46. Segments and teeth of about equal length; cilia well developed or rudi-
mentary 47
47. Cilia distinctly appendiculate. (See also Pohlia) 48
47. Cilia not much if any appendiculate 49
48. Large stoloniferous, MniumAiVe plants with clustered capsules; leaf-cells
less than 4:1 Rhodobryum, p. 134
48. Smaller, non-stoloniferous mosses with the capsules borne singly; leaf-cells
less thn 4:1 Bryum, p. 127
48. Leaves linear-lanceolate, costa wide; leaf-cells above linear-rhomboid
Leptobryum, p. 121
49. Leaf-cells narrow, linear-rhomboidal above; leaves linear-lanceolate; costa wide
and indistinct Leptobryum, p. 121
49. Leaf-cells more or less linear; costa strong and fairly wide (see also Encalypta)
Pohlta. p. 122
49. Leaf-cells rhomboid-hexagonal, never linear 50
50. Large plants with the upfser leaves often bordered, ovate; leaf-cells not
more than twice as long as broad, rounded to hexagonal Mmum, p. 136
50. Smaller plants with linear-lanceolate upper leaves; leaf-cells more than
twice as long as broad 51
51. Annulus none; leaf-cells more or less rhombic-hexagonal, never linear, except at
margins; leaves pale glaucous green; plants in wide, loose tufts; stems up to
5 cm high; sporjgonia single Mniobryum, p. 126
51. Sp>orogonia single; stems not rising from rhizone-like stolons; annulus mostly
14 American Midland Naturalist Monograph No. 6
present; leaf-cells rectangular below, rhomboid-hexagonal above
Bryum, p. 127
5 1 . Sporogonia often several in a cluster; stems erect from rhizome-like stolons;
annulus present; median leaf-cells rhomboid-hexagonal with pitted walls
Rhodobryum, p. 134
52. Rock- or crevice-inhabiting mosses 53
52. Earth-inhabiting plants 58
53. Leaves on the sterile stems in two rows, forming a continuous marginal wing
Schistostega, p. 120
53. Leaves on the sterile stem not thus arranged 54
54. Leaves distichous, closely imbricate, carinate-plicate; stems radiculose and
bulbiform at base Bryoxiphium, p. 68
54. Leaves not as above; stem not bulbiferous at base 55
55. Leaves ecostate; seta less than 1 mm long; operculum apiculate
Hedwigia, p. 171
55. Leaves costate; operculum rostrate 56
56. Capsule smooth, terminating the main axis; plants often on calcareous
rocks 57
56. Capsule ribbed, borne on a short lateral branch;' plants not often on cal-
careous rocks Amphidium, p. 101
57. Leaf-margins revolute on one or both sides; upper leaf-cells clear and pellucid;
columella remaining attached to the operculum and falling away with it
Hymenostylium, p. 81
57. Leaf-margins always plane: upper leaf-cells densely papillose and obscure;
columella remaining in the capsule after the ojjerculum falls away
Gymnostomum, p. 80
57. Leaves canaliculate-subulate from a concave, lanceolate base; basal leaf-cells
thin-walled rectangular, upper cells isodiametric incrassate Seligeria, p. 53
58. Calyptra cucullate; cells isodiametric above the middle of the leaf (see
also Astomum) Pottia, p. 89
58. Calypra mitrate; cells elongated above the middle of the leaf 59
59. Capsule immersed and sessile, splitting equatorially and without very specially
modified cells at the line of dehiscence Aphanorhegma, p. 115
59. Capsule exserted on a lon^ seta, or, if immersed, ojxrculate with 1-3 rows of
denser cells below the line of dehiscence Physcomitrium, p. 117
60. Capsule symmetric; peristome single 61
60. Capsule unsymmetric; peristome double, the inner in the formi of a short
conical tube 64
61. Peristome teeth 4 Tetraphis, p. 158
61. Peristome teeth 32 or 64 62
61. Peristome teeth 16 Trichostomum, p. 82
62. Capsule usually square or hexagonal; calyptra densely hairy; teeth gener-
ally 64 PolytTtchum, p. 166
62. Capsule cylindrical; teeth 32 63
63. Calyptra densely hairy; leaves not crisped when dry; protonema persistent
Pogonatum, p. 165
63. Calyptra cucullate, almost smooth; leaves crisped when dry Atrichum, p. 159
64. Leaves green, costate, conspicuous; capsule sessile and immersed
• Diphyscium, p. 155
64. Leaves almost microscopic, colorless; capsule exserted on a thick, red stalk
Buxbaumia, p. 156
B. Pleurocarpi
65. Leaves distichous, dorsally wmged and clasping at the base Fissidens, p. 69
65. Leaves plun-seriate, or, if distichous not clasping at the base 66
66. Cilia none; segments none, irregular, or rudimentary, or filiform and not
Jennings: Manual of Mosses 15
split - ■ , • ^^
66. Cilia (none) or often present; segments carinate and often split along the
median line! (sometimes adhering to the teeth) 76
66. Inner peristome consisting only of a low papillose basal membrane
Litidbergia, p. 202
67. Segments quite rudimentary, sometimes with a distinct carinate basal mem-
brane; leaves smooth to usually more or less papillose "8
67. Leaves smooth V W
68. Leaves deltoid to round-ovate, spinulose-dentate to fimbriate; paraphyllia
none; costa single and usually half the length of the l:af Thtltj, p. 195
68. Leaves oval-elliptic to obovate, entire or cristate-serrate; costa double or
short or almost lacking Pterygynandrum, p. 278
68. Stem-leaves ovate, sub-cordate, apex abruptly long-acuminate; costa sub-
percurrent; branch-leaves narrower Leskeella, p. 205
69. Inner penstome lattice-like; leaf-cells narrow and prosenchymatous 70
69. Inner peristome not lattice-like, sometimes none; segments when present free,
sometimes rudimentary - 71
70. Ecostate; capsule immersed; plants aquatic Fontinalis, p. 173
70. Costate; capsule more or less exserted; plants growing on the base of
bushes and shrjbs Dkhelyma, p. 178
71. Leaves complanate, more or less transversely undulate Neckera, p. 187
71. Leaves more or less spreading, not transversely undulate 72
72. Inner peristome none, or, if present, the linear segments without a basal
membrane; teeth flat, thin, distantly articulated, approximate in pairs;
costa simple, often delicate and short, or none 73
72. Inner peristome lacking or the peristome double; basal membrane low 74
73. Costa mostly delicate and short; teeth approximate in pairs, broad and obtuse;
leaves serrate to ciliate-dentate Fabronia, p. 279
73. Costa strong, ending above the middle of the leaf; peristome deeply inserted;
teeth broadly lanceolate; segments shorter than the teeth, linear, almost
smooth Anacamptodon, p. 280
74. Calyptra cucullate; capsule exserted; segments of inner peristome rudi-
mentary and narrow or none or adhering to inner surface of teeth 75
74. Calyptra conical-campanulate; capsule immersed; segents linear or very
narrow-carinate (Cryphaea)
75. Ecostate Leucodon, p. 183
75. Costate Leptodon, p. 185
76. Leaves mostly rough-papillose. {Bryhnia spp. and Isopterygium geophilum
are papillose dorsally by projecting angles of cell-walls) 77
76. Leaves smooth (see also Helodium) , or sometimes slightly papillose at the
cell angles 89
77. Capsule erect and symmetric 78
77. Capsule usually arcuate, unsymmetric 84
78. Leaves costate, margins papillose-denticulate; cells pluripapillate; operculum
conic-rostrate Rauia, p. 206
78. Leaves costate, margin i:sually entire (see also Myurella) 79
78. Leaves ecostate 83
79. Costa not more than half the length of the leaf Haplohymenium, p. 197
79. Costa strong in our species, nearly reaching the apex of the leaf 80
80. Primary stems stoioniform, secondary stems bearing the sporophytes; stem-
leaves minute; paraphyllia none Anomodon, p. 198
80. Primary stems bearing the sporophytes and not stoioniform; branch and
stem leaves not markedly dissimilar; paraphyllia often present 81
81. Paraphyllia none; leaf-cells compact, oval-oblong to round-hexagonal, nearly
smooth; cilia none; teeth not hygroscopic; dioicous Leskeella. p. 205
81. Paraphyllia usually present; leaf-cells hexagonal to parenchymatous, mostly
unipapillate: autoicous - 82
16 American Midland Naturalist Monograph No. 6
82. Teeth with well developed lamellae; segments narrow-lmear Leskea, p. 20^
82. Teeth with low lamellae; segments none Lindbergia, p. 202
83. Cilia two; plants glaucous-green with closely imbricated leaves and julaceous
branches Myurella, p. 196
83. Cilia none; plants light green with loosely appressed leaves and more or less
flattened branches Schwetschkeopsis, p. 281
84. Plants creeping, one- to three-pinnate 85
84. Plants erect to ascending, simply pinnate, in large tufts 88
85. Plants small, delicate, one- to two-pinnate 86
85. Plants larger, one- to three-pinnate; m large flat mats Thuidium, p. 209
86. Costa of stem-leaves one-fifth to one-sixth oi the width of the leaf base
Thuidium, p. 209
86. Costa of stem-leaves one-tenth to one-twelfth of the width of the leaf base
(see also Thuidium) 87
87. Stem and branch leaves dissimilar; leaf-cells each with several minute papillae
Rauia, p. 206
87. Stem and branch leaves similar; leaf-cells each with but one or rarely two
papillae Haplochdium, p. 207
88. Stem and branch leaves rather similar in size and shape
(Helodium, Ivs. costate, p. 214
{Hookeria, Ivs. ecostate, p. 215
88. Stem and branch leaves dissimilar Rauia, p. 206
89. Stems dendroid, upright from a creeping base; capsules clustered 90
89. Stems prostrate or ascending with the capsules borne singly 91
90. Cilia none; capsules erect and symmetric Climacium, p. 180
90. Cilia well developed, appendiculate; capsules inclined, unsymmetric
PoTOtrichum, p. 189
91. Cilia none; capsule symmetric and erect, or nearly so (see also 120a) 92
91. Cilia generally present and well developed; capsule unsymmetric, generally
more or less arcuate and recurved, sometimes almost erect 9^
92. Branches strongly complanate; leaves cultriform Homalia, p. 188
92. Branches not strongly complanate; leaves ovate to lanceolate 93
93. Segments adhering to the teeth; basal membrane none or obscure 94
93. Segments free from the teeth 95
94. Leaves with short single costa; seta rough Homalotheciella, p. 286
94. Costa short and double or none; seta smooth ...Pylaisia, p. 226
95. Basal membrane broad and distinct Pylaisia, p. 226
95. Basal membrane none or narrow 96
96. Stem oval in cross-section; teeth of peristome not hyaline-margined
_ Entodon, p. 191
96. Stem rounded; teeth of peristome hyaline-margined Platygyrium, p. 229
96. Small, slender; leaves lance-linear to lanceolate; teeth not margined; cilia
rudimentary or none Amblystegiella, p. 223
97. Stem mostly woody, often stoloniferous, irregularly divided, the leafy branches
often more or less regularly pinnate; leaves erect-spreading to squarrose,
rarely imbricated; cells narrowly prosenchymatous, rarely parenchymatous,
towards the base more lax and often punctate; costa various, but rarely
almost percurrent; alar cells mostly rounded or oval — 4- to 6-sided, usually
forming a well defined but not markedly inflated group; teeth and segments
same length; basal membrane wide; cilia usually well developed; operculum
rounded, conical-obtuse to short rostrate {^Hypnaceae} 99
97. Characters not combined as above 93
98. Slender plants with creeping stems; leaves often secund or somewhat com-
planate; costa none or double and short; cells narrow, prosenchymatous;
alar cells 3 to 8, large, inflated and jjellucid; capsule small, oval to
oblong, exothecial cell-walls collenchymatous; operculum mostly long and
slenderly rostrate -.
Jennings: Manual of Mosses 17
(See also Hygrohypnum) Sematophyllum, p. 283
98. Stems round, creeping, procumbent or more or less erect, often irregularly
stoloniferous; leaves ascending to appressed, often complanate, rarely
secund: stem- and branch-leaves dissimilar in the stoloniferous species
only, ovate to lanceolate, mostly slenderly ecuminate; costa not often
reaching the apex but usually reaching to the middle at least; cells pro-
senchymatcus, long rhomboidal to linear-vermicular; operculum conical,
blunt to long rostrate [Bracbytheciaceae] 120
98. Branches somewhat complanate; leaves 4-5 mm long, ovate, soft, thin, dry-
ing whitish; leaf-cells parenchymatous Hookeria acutijolia, p. 215
Hypnaceae (99-119)
99. Costa single, reaching mid-leaf; stem-leaves lance-ovate, rugose, falcate-secund;
robust; ends of stem and branches hooked (falcate-secund): leaf-cells dor-
sally papillose; lid shortly rostrate Rhytidium, p. 254
99. If not as above and with costa in our species singk; extending to the middle
of the leaf or beyond; operculum never rostrate [Amblystegteae] 102
99. Costa double and usually short, or none; operculum sometimes rostrate 100
100. Stem and branch leaves usually distinctly dissimilar; leaves symmetric,
inserted at right angles to the axis of the stem; branches sometimes
terete or julaceous [Hylocomieae] 114
100. Stem and branch leaves quite similar; if minute plantd with median leaf-
cells oblong-hexagonal to rhomboidal (about 2-8:1); leaves lanceolate
to lance-subulate, see Amblystegiella; leaves often inserted obliquely and
unsymmetrically 101
101. Robust, stiff; branching complanately pinnate and plumose; leaves falcate-
secund; costa double and short or none Pttlium, p. 259
101. Leaves either symmetnc and normally inserted or unsymmetric and obliquelv
inserted and mostly falcate-secund and smooth; operculum sometimes ros-
trate {^Stereodoriteae^ 1 18
101. Leaves obliquely inserted, mostly two-ranked, mostly unsymmetric but little or
not at all secund; branches mostly complanate; operculum conic to short
rostrate, rarely long-rostrate [Plagiothecicae'] 119
102. Leaves margined SciaTomwm. p. 234
102. Leaves not margined 103
103. Costa strong, ending almost in the apex or sometimes excurrent 104
103. Costa not extending to the apex of the leaf 109
104. Paraphyllia numerous and polymorphic 105
104. Paraphyllia none or very few 106
105. Leaves deeply longitudinally folded, falcate-secund CratoneitTon. p. 233
105. Leaves not deeply longitudinally folded Hygroamblyslegutm, p. 230
106. Leaf-cells linear-vermicular to the leaf-base, mostly with blunt ends, the
alar cells forming a small but well defined group of quadrate or rec-
tangular cells Hygrohypnum, p. 243
106. Leaf-cells hexagonal and 2 to 6 times as long as wide, or prolonged-
linear and becoming wider and shorter towards the leaf-base, alar cells
forming a larger group often reaching to the costa 107
106. Leaf-cells elongate-hexagonal to rhomboid, about 2-6:1; often a few alar
cells inflated, aquatic or sub-equatic Hygroamblystegium, p. 230
107. Alar cells parenchymatous; leaves falcate-secund to cirtinate
Drepanocladus, p. 235
107. Alar cells prosenchymatous '08
108. Leaf-cells prolonged; leaves more or less concave Calliergon, p. 240
108. Leaf-cells prosenchymatous-hexagonal, 2 to 6 times as long as wide
Hygroamblystegium, p. 230
109. Leaves cordate- to ovate-lanceolate, more or less acuminate; costa weak, ending
18 American Midland Naturalist Monograph No. 6
at or beyond the middle of the hii\ cells rarely linear, mostly quadrate and
parenchymatous, or hexagonal and short-prosenchymatous
Amblystegium, p. 218
109. Aquatic; leaves mostly short and broad, rojnJed and LLint, soft; median leaf-
cells linear-vermicular, the alar forming a distinct group, sub-rectangular or
somewhat inflated Hygrohypnum, p. 243
109. Not with the above combination of chnracters 110
110. Leaf-cells narrowly linear; leaves broadly ovate to cordate or lanceolate,
prolonged into awl-shaped reflexed squarrose tips Campylium, p. 247
110. Not with above combination of characters HI
111. Leaves oval- to oblong-lanceolate, more or less long acuminate; leaf-cells nar-
row, prosenchymatous; plants shining Homomallium, p. 225
111. Not with characters combined as above 112
112. Plants very slender, not shining; leaves usually spreading in all directions,
lanceolate to linear-lanceolate; cells rhomboidal to long hexagonal, 2 to
4, or rarely 6-8, times as long as broad Amblystegiella, p. 223
112. Leaf-cells prolonged-linear, mostly very narrow 113
113. Leaves erect-spreading or imbricated, oblong-ovate to nearly circular, obtuse or
apiculate, often very concave; costa short and double, or none
Calliergonella, p. 242
113. Leaves more or less falcate-secund to circinate, from a mostly narrowed and
somewhat decurrent base, becoming ovate- to triangular- or cordate-lanceo-
late, more or less slenderly acuminate; cost^ weak and reaching above the
middle, or rarely even excurrent Drepanocladus, p. 235
1 14. Paraphyllia numerous; leaves more or less concave, from abruptly to
gradually acuminate, mostly plicate Hylocomium, p. 255
1 14. Paraphyllia none or few 115
1 15. Stem-leaves more or less squarrose-spreading to falcate-secund, acuminate 117
1 15. Stem-leaves more or less crowded, imbricate but with more or less spreading
or secund tips 116
1 15. Stem-leaves elliptic, concave, apex obtuse; branches julaceous Hypnum, p. 258
1 16. Stem-leaves turgidly imbricate and secund, rugose, narrowly lanceolate-
acuminate from a broadly oblong base, glossy; apex distinctly serrate;
cilia two; annulus present Rhytidium, p. 254
1 16. Stem-leaves close or loosely imbricate, not secund, broadly ovate or
rounded with an obtuse apex, olive or grayish-green, ap>ex faintly crenu-
late; cilia three; annulus none Hypnum, p. 258
116. Leaves broad, obtuse, concave; alar cells inflated Calliergonella, p. 242
117. Alar cells not at all or but very little differentiated; plants distantly and
irregularly pinnate; leaves squarrose or spreading-secund
Rhytidiadelphus, p. 252
117. Alar cells distinctly differentiated; plants closely pinnate; leaves circinate-
secund Ctenidium, p. 25 1
118. Plants large, to 15 cm, closely and regularly pinnate leaves linear-acumini-
nate from a broadly ovate base, stem-leaves plicate, falcate-secund;
cilia 3 or 4 Ptilium, p. 259
118. Plants large to quite slender, simple or pinnate, mostly irregularly pin-
nate; leaves ovate- to cordate-lanceolate, shortly to slenderly acuminate,
generally circinate-secund in two series Stereodon, p. 260
1 18. Characters not combined as above 128
1 19. Leaf-cells very narrowly prosenchymatous, alar cells mostly not differentiated;
leaves oblong to linear, short (winted, or ovate- to linear-hnceolate, acute to
long-acuminate or piliferous hopterygium, p. 269
1 19. Leaf-cells not so narrow, alar cells mostly broader, hyaline and thin-walled;
leaves broadly lanceolate to oval, more or less long-acuminate, decurrent
Plagiothecium, p. 274
Jennings: Manual of Mosses 19
BraCHY THECIACEAE ( 1 20- 1 26 )
120. Capsule erect to horizontal, symmetric to unsymmetric; basal membrane
low !20a
120. Capsule generally inclined or horizontal, unsymmetric; basal membrane
high - 121
120a. Calyptra hairy; segments adhering to peristome-teeth
Homa'otheciella, p. 286
120a. Calyptra smooth; segments not adhering to peristome-teeth (^ee also
Brachythecium ) Chamberlainia, p. 287
121. Leaves with several deep plications spoon-shaped; branches i'.daceous
C amptolhecium , p. 287
121. Leaves plane or but shallowly plicare 122
122. Operculum conic, sometimes short-rostrate; alar cells difFerenciited, quad-
rate; capsules usually short, thick, dark chestnut color. (If leaves
dorsally papillose, see Bryhnia) Brachythecium, p. 289
122. Operculum long rostrate; alar cells few or not differentiated 123
123. Autoicous; branches and leaves complanate, leaves mostly only slightly con-
cave, never sulcate, ovate to ovate-linceolate, more or less acuminate;
costa rarely ending in a spine on the back of the leaf; cells narrow, smooth;
seta smooth Rhynchostegium, p. 306
123. Dioicous; seta generally papillose; costa sometimes ending in a spine on the
back of the leaf; leaves not complanate 124
124. Leaves very concave, not at all or but weakly plicate, ovate to oblong,
more or less abruptly acuminate or piliferous; costa not ending in a
spine; cells mrrow and smooth Cirriphylltim, p. 293
124. Not with the characters combined as above 125
125. Stem and branch-leaves often dissimilar, stem-leaves ovate- to triangular-
cordate or rounded triangular-oval, obtuse to somewhat acuminate; costa
often ending in a spine; leaf-cells very narrow and smooth 126
125. Stem-leaves more or less concave, irregularly plicate; cells elongated-rhomboid
to elongated-hexagonal; branch-leaves rough on the back by papillae or
tooth-like projecting cell-angles; seta very rough Bryhnia, p. 304
126. Leaves not or very little concave, never plicate; seta smooth or rough
Oxyrhynchium, p. 300
126. Leaves more or less concave, indistinctly plicate; seta smooth
Eurhynchium, p. 302
127. Mostly not aquatic, sometimes submerged but yet floating Fissidens, p. 69
127. Aquatic, filiform, and floating Octodiceras, p. 75
128. Leaves obtuse 129
128. Leaves acute to acuminate - 130
129. Aquatic, usually m streams; leaves spreading or somewhat secund, soft and
thin; branching irregular Hygrohypnum, p. 243
129. In bogs or swamps; glossy; regularly pinnate; leaves loosely appressed or
descending Calliergoriella, p. 242
130. Leaves squarrose to squarrose-recurved (see also Plagiothecium)
Campylium, p. 247
130. Leaves otherwise 13 1
131. Alar cells inflated; leaves deirurrent Plagiothecium striatellum, p. 275
131. Alar cells not inflated 132
132. Median leaf-cells linear-hexagonal, about 4-8:1, the apical shorter and
sub-rhomboidal, the alar quadrate and green-opaque, incrassate: leaves
about 0.6-1.0 mm long Homomallium, p. 225
132. Leaves decurrent; median leaf-cells linear, 5-15:1 Plagiothecium, p. 274
132. Leaves not decurrent; median leaf-cells linear, 20:1
Isopterygium ptilchellum, p. 272
20 American Midland Naturalist Monograph No. 6
Order I. Sphagnales — Peat Mosses
Characteristic peat mosses, in bogs, usually either in water or water-soaked,
monoicous or dioicous, deeply cespitose, the tufts constantly growing upwards
at the same time that the plants are dying from below and often thus giving
rise to deep beds of peat, the tufts light grayish-green or sometimes yellowish,
often more or less tinted with red above: stems without rhizoids, usually com-
posed of an outer cuticular sheath consisting of one to three or four layers of
large lax cells, an intermediate hollow cylinder composed of prosenchymatous
cells with usually thickened walls, and a central pith of lax parenchymatous
cells; branches symmetrically fascicled, usually partly divergently spreading
and partly slender and appressed-pendent; leaves ecostate, unistratose, com-
posed of large, hyaline, more or less elliptic cells with usually perforated and
spirally thickened (fibrillose) walls and separated by narrow chlorophyllose
cells which meet at their ends to form a continuous network throughout the
leaf; stem-leaves usually different in form from the branch-leaves, remote,
often lacking entirely the pores and spiral fibrils, while the branch-leaves are
usually porose, fibrillose, and more or less densely imbricated; seta none but
the capsule is borne upon an outgrowth from the gametophyte termed a pseu-
dopodium; antheridial flowers usually at the ap£x of specialized branches of
the capitulum, the antheridia being pedicillate, globose, and solitary at the
base of the bracts; the archegonial flowers gemmiform, axillary in one of the
upper fascicles, only one of the three or four archegonia developing, as a rule:
capsule globose, castaneous, with a convex operculum, without annulus or
peristome; calyptra irregularly lacerate; spores developed from the amphithe-
cium, the columella from the endothecium.
This order is a peculiar one comprising but one family {S phagnaceae)
which contains but the one genus {S phagnum) with about 340 known species.
The Sphagnums are cosmopolitan in suitable habitats but are most abundant
in the cooler temperate regions of Europe and North America, in both of
these countries often forming bogs of large areas. In North America there
are known about 40 species, at least 30 species or varieties in our range.
The following treatment follows closely that of C. Wamstorf in Die Na-
tiirlichen Pjlanzenjamilien 1(3): 248-262. 1900. Also frequently consulted
was Sherrin, W. R. An Illustrated Handbook of the British Sphagna. 1927
and Andrews, A. LeRoy. Sphagnales. North American Flora 15:1-31. 1913.
I. Sphagnum [Dillenius] Hedwig
(Revised with the assistance of Charles M. Boardman)
Analytical Key to the Species
A. Branches in tufts of 6-12 with 3-5 of them spreading; branch-leaves narrowly ovate-
lanceolate, narrowed to an involute-tubulose point 6. S. Wulfianum
A. Branches in tufts of 2-6, with 2 or sometimes 3 of them spreading B
B. Cuticular cells of stems and branches spirally fibrose; branch-leaves cucullate at
the apex, which is obtuse and entire, rarely acute C. (Cymbifolia)
Jennings: Manual of Mosses — I. Sphagnales 21
B. Cuticular cells of stem and branches not fibrose G
c. Chlorophyllose cells of branch-leaves in cross-section exposed eithett dorsally or ven-
trally, or both D
c. Chlorophyllose cells of' branch-leaves centrally placed and entirely enclosed by the
hyaline cells 5. S. magellamcum
D. Lateral walls of chlorophyllose cells of branch-leaves facing into the hyaline cells
either more or less papillose or with comb-fibrils --E
D. Lateral walls of the chlorophyllose cells smooth F
E. Chlorophyllose cells of branch-leaves in cross-section more or less equilaterally tri-
angular and exposed only on the ventral (inner) side of the leaf or, in the upper
part of the leaf, usually broadly trap>ezotdal and exposed on both sides, and,
mainly in the lower part of the leaf, the walls facing into the hyalinf cells with
comb-fibrils 1. S. tmbrtcatum
E. Chlorophyllose cells of branch-leaves in cross-section narrowly to broadly spindle-
shaped or barrel-shaped, usually with thickened outer walls narrowly free on both
surfaces or nearly enclosed, the walls facing into the hyaline cells mostly minutely
papillose 2. S. papillosum
F. Chlorophyllose cells of branch-lea\es in cross-section very broadly triangular or
triangularly trapezoidal, and v.ith the broader face ventral 3. S. affine
F. Chlorophyllose cells narrowly triangular or trapezoidal, seen in cross-section not
more than one-half as wide as long, usually less 4. S. palustre
G. Chlorophyllose cells of branch-leaves in cross-section elliptical or more or less barrel-
shaped or rectangular, but not triangular or trapezoidal H
G. Chlorophyllose cells in cross-section triangular or trapezoidal with one or both faces
free I
H. Hyaline cells of stem-leaves non-fibrillose; chlorophyllose cells of branch-leaves in
cross-section enclosed on both surfaces, with the lumen sub-central, elliptic;
branch-leaves squarrose in their upper half; cuticular cells of branches with an
apical pore 7. S. compactum
H. Hyaline cells of stem-leaves fibrillose; chlorophyllose cells of branch-leaves free
on both surfaces, their hyaline cells with numerous pores N. (Subsecunda)
I. Face of chlorophyllose cells of branch-leaves in cross-section dorsally free; hyaline
cells strongly convex ventrally the walls adjoining the chlorophyllose cells smooth
or papillose J
I. Face of chlorophyllose cells free on the ventral (inner) surface of the leaf, the hya-
line cells dorsally strongly convex, the inner walls adjoining the chlorophyllose
cells smooth S. (Acut, folia)
J. Hyaline cells of median dorsal surface of branch-leaves with about 5 very large
pores, but these smaller towards the leaf-apex; chlorophyllose cells with the
exterior walls strongly thickened K. (Squarrofa)
J. Hyaline cells with pores on median leaf-surface (dorsal) none or very few;
chlorophyllose cells with the walls not strongly thickened L (Cuspidata)
K. Stem-leaves with a rounded hut narrow apex; branch-leaves at least 2 mm long
8. S. squaTTOSum
K. Stem-leaves not narrowed but rounded at apex; branch-leaves less than 1.8 mm long
9. S. teres
L. Cortical cells of stem small and thick-walled not much different from the inner
cells M
L. Cortical cells of stem distinctly larger in 1-3 layers, thin-walled
1 \. S. cuspida:um
M. Branch-leaves 1-2 mm long, when dry undulate and with more or less recurved tips
and with the dorsal pores of the hyaline cells more or less restricted to the cell-
angles 10. S. recur yum
M. Branch-leaves when dry but weakly undulate, scarcely recurved, about 1 mm long,
the upper hyaline cells with pores both in the cell-angles and along the sides
ICb. S. recurv. var. tenue
N. Cuticular cells of stem 2-3-stratose O
22 American Midland Naturalist Monograph No. 6
N. Cuticular cells of stem one-stratose P
O. Stem-leaves small, not more than 1 mm long, fibrtUose only towards the apex;
branch-leaves secund 12. S. contortum
O. Stem-leaves large, 1.5-2 mm long, fibrillose to the base or nearly so; branch-leaves
not secund 13. S. platyphyllum
P. Stem-leaves with the hyaline border strongly widened below, fibrils none or only
in the upp)er cells 15. S. subsecundum
p. Stem-leaves with a uniformly wide border and with fibrils more numerous Q
Q. Stem-leaves strongly auriculate, large, 1.5-2 mm long, fibrillose in the upper two-
thirds at least, or clear to the base 14. S. auriculatum
Q'. Stem-leaves not or but slightly auriculate at the base, usually of medium size, fibril-
lose in about the upjaer two-thirds R
R. Stem-leaves about 1-1.5 mm long, with the hyaline cells septate, and m the upper
half fibrillose 16. S. inundatum
R. Stem-leaves 1.3-1.5 (-2) mm long, very little septate, fibrillose in the upper two-
thirds, or to the middle 17. S. pungens
S. Stem-leaves erose or lacerate-fimbriate at the broadly rounded ap>ex, non-fibrillose T
S. Stem-leaves not fimbriate, but truncate or toothed at the ap)ex, usually more or less
fibrillose, at least above U
T. Stem-leaves spatulate, fringed at the ap>ex and around the upper half; plants deli-
cate, with slender branches and never' red 18. S. fimbrijtum
T. Stem-leaves lingulate, fringed only at the very broad apex; branches thicker than
in 5. fimbriatum 19. S. Ghgensohnii
V. Stem-leaves lingulate V
U. Stem-leaves more or less equilaterally triangular or triangular-lingulate W
V. Pigment red, never brown (green in var. ) 20. S. Warnstorfii
V. Pigment brown 21. S. fuscum
w. Branch-leaves five-seriate, when dry not lustrous 22. S. qititiqucfarium
w. Branch-leaves not distinctly 5seriate, when dry more or less lustrous X
X. Stem-leaves usually non-fibrillose and non-porose; branch-leaves usually lustrous
when dry 23. S. plumulosum
X. Stem-leaves usually fibrillose and porose; branch-leaves usually not glossy when
dry 24. S. acutifolium
Section I. Inophloea
Cuticular cells of stems and branches reinforced with spiral fibers and
porose.
Subsection I. Cymbifolia
Large, with the branch-leaves boat-shaped, cucullate and scabrous at the
back of the apex. Stem-leaves not bordered.
1. Sphagnum imbricatum (Homschuch) Russow
(S. Austini Sullivant)
Plate I
This species occurs in bogs and wet moors in Europe and Asia and in
North America from Labrador to Alaska and south to Louisiana. In our
region it is represented by the following variety. The typical form, as com-
pared with the following variety, has usually more yellowish or brownish
denser tufts with the shorter comal branches more erect and the divergent
branches more densely-leaved and more ascending; while the hyaline cells of
the stem-leaves are sparsely comb-fibrillose on the inside face of the lateral
walls; otherwise the characters of variety and species are identical; in fact it
Jennings: Manual of Mosses — I. Sphagnales 23
may be best to follow Andrews and regard species number three (S. affine) as
a variety of S. imhricatum, the latter possessing comb-ftbrils, while S. imbri-
catiim var. ajfine entirely lacks them.
la. Sphagnum imbricatum var. sublaeve Wamstorf
(S. Austini var. glaucum f. squarrosulum Roell)
Rather densely cespitose, large, usually more or less glaucous-green, gray-
ish or yellowish below; stems rather stout, with us about 4-8 cm long, the
wood-cylinder greenish or yellowish and surrounded by a cuticular sheath of
uaually four layers of thin-walled, large, fibrillose, and porose cells, the inner-
most largest; stem-leaves about 1-1.8 mm long, widely and blundy lingulate,
somewhat concave, the upper half rounded and with an erose-fimbriate margin,
the base more or less auriculate; hyaline cells of stem-leaves mosdy non-fibril-
lose and non-porose. a few often septate, the upper median more or less rcunded-
hexagonal, the basal elongate, the insertion-cells small and brownish-incrassate;
branches usually four, two or three spreading, tumid, about 1.5 cm long, the
rather shortly tapering apex pendent, the comal branches short and more or
less erect-spreading, often obtuse, the pendent branches closely applied to the
stem, very slender; branch -leaves 2-3 mm long, broadly ovate, very concave, th?
margins involute, the apex abruptly and bluntly tapering, cucullate and more
or less widely squarrose-spreading; the hyaline cells of the branch-leaves broad,
fibrillose, ventrally with a few large round median pores, with small pores in
the angles, dorsally with large round or elliptic pores at the cell-angles, the
large pores usually equalling about one-third the width of the hyaline cell; the
basal hyaline cells of the branch leaves are distinctly comb-fibrillose on the
inner lateral side of the wall adjoining the chlorophyllose cells; in cross-section
the chlorophyllose cells are widely trapezoidal, the ventral wall widest and
almost or quite as wide as the lateral walls, the dorsal wall exposed between
the highly convex dorsal walls of the hyaline cells and usually one-third to one-
half the width of the lateral wall; the cuticular sheath of the branches con-
sisting of one layer of rectangular, fibrillose, porose cells: fruit not seen, but
spores of S. imbricatum are stated to be yellowish, smooth, and about .025
mm in diameter.
This variety is probably well distributed in regions where the typical form
occurs.
Crawford Co.: Pymatuning Swamp, near Linesville, May 12, 1908. O.E.J, (fig-
ured). Fayette Co.: Ohio Pyle and Wiggins. C.M.B. Mercer Co.: Near Houston
Junction. July 12, 1902. J.A.S. Somerset Co.: Clear Run. C.M.B. Warren Co.:
Columbus, P. 168. Sept. 1948. C.M.B.
2. Sphagnum papillosum Lindberg
{S. cymbifolium var. papillosum Schimper)
Plate II
Vigorous, coarse, tufted, yellowish to ochraceous: stems reddish-brown,
almost 1 mm in diameter, inflated cuticular cells porose towards apex, scarcely
if at all fibrose; divergent branches 1 or 2, pendent 1 or 2; divergent branches
24 American Midland Naturalist Monograph No. 6
1-1.5 cm long, from a turgid 3 mm-thiclc portion (wet) rather suddenly taper-
ing to a thin point; branch leaves 2-2.5 mm long, broadly ovoid, cucullate, at
the margin with a border of about 2 rows of liner cells, of which a few project
outwards as minute teeth, the cucullate apex minutely dorsally scabrous by
erosion of the cell walls; median hyaline cells of branch-leaves about 3-4:1,
strongly fibrillose, somewhat pcrose dorsally, ventrally more so with distinct
pores, the chlorophyllose cells ovoid to triangular-ovoid with the ventral end
flush with the hyaline cells but dorsally being much exceeded by the convex
hyaline cells.
On the sides of the chlorophyllose cell-walls facing the hyaline cells there
are minute papillae in many of the leaves, but apparently not all. These
papillae seem best developed in the basal half of the leaf.
Crawford Co.: Pymatuning Swamp, Linesville. O.E.J. May 26-27, 1934 (Det.,
C.M.B. ). McKean Co.: Cathrine Swamp, on plateau, elevation 2080 ft. C.M.B., Sept.
2, 1948 (figured). Somerset Co.: Glade Run bog, Negro Mt. Elev. 2470 ft. C.M.B.,
July 15, 1949.
3. Sphagnum affine Renauld and Cardot
{S. imbricatum var. affine Warnstorf)
Plate I
Densely cespitose, usually bluish or glaucous-green above and more or less
yellowish below: stems robust, sometimes as much as a decimeter in length,
usually much less, densely branched; cuticular sheath distinct, three-layered,
the inner layer with the largest cells, the outer cells usually densely spirallv
fibrillose and 2-6-pored; stem-leaves large, 1.6-2.2 mm long, about two-thirds
as wide, widely spatulate, the rounded upper half somewhat concave, erose-
fimbriate; hyaline cells of stem-leaves usually weakly fibrillose in their upper
half, porose dorsally, the hyaline cells below non-fibrillose, the pores large and
few; branches usually 4, one or two of these very slender, pendent and rather
closely appressed to the stem, the divergent ones horizontally spreading, with
drooping tips, rather swollen below, tapering towards the apex, the branches
often 2 cm long; branch-leaves 2-3 mm long, broadly ovate, bluntly and cucul-
lately short-pointed, the whole leaf very concave and with more or less in-
volute margins, the apex dorsally scabrous by the erosion of the outer cell-
walls; hyaline cells of the branch-leaves rather wide, spirally fibrillose on both
sides, with large well-defined pores of one-third to one-half the cell-width and
confined mainly to the cell-angles; chlorophyllose cells in cross-section widely
trapezoidal, the wider face being ventrally exposed and more than or at least
half of the width (dorsal- ventral) of the smooth lateral walls, the dorsal face
exposed and rather narrow*; the cuticular cells of the branches porose and
densely fibrillose: fruit not seen.
In bcygs and swampy borders of ponds and streams. Europe and in North
America from Canada to Florida. Common in the more northern counties in
* The chlorophyllose cells are more nearly equilaterally triangular th.m appears in our
drawing in Plate I.
Jennings: Manual of Mosses — I. Sphagnales 25
ouj region and in the mountains, but mostly referred in the past to S. cymbi-
jolium.
Common and now known from 14 counties in western Pennsylvania. Specimen figured:
Open bog. Bear Meadows, Center Co., Pa., Sept. 21. 1909. O.E.J.
3a. Sphagnum afftne i. sqtiarrosula Warnstorf, is merely a strong grow-
ing form with the upper half of the leaves of the branches when dry strongly
squarrose. All gradations between the typical form and the squarrose form
are to be found in our region. A packet of specimens quite characteristically
of the squarrose form is as follows:
Cambria Co.: On boggy plateau near St. Lawrence, July 24, 1098. O.E.J. (Mixed
with S. recuryum Schwaegrichen ) .
4. Sphagnum palustre Linnaeus
{S. cymbifolium Ehrhart; S. latijolium of 1st edition)
Plate I
Densely cespitose, usually robust, bluish or glaucous-green to yellowish:
stems rather stout, usually 8-10 cm long, sometimes 2-3 dm long, the cuticular
sheath composed of 3-4 layers of inflated cells, the innermost of which are the
largest, the outer layer being rectangular, fibrillose and porose, the wood-
cylinder being usually yellowish or brownish; stem-leaves large, about 2 mm
long and 1.25 mm broad, sometimes 3 mm long, spatulate-lingulate, the
broadly rounded apex somewhat erose-fimbriate, below narrowly hyaline-bor-
dered, the insertion composed of brownish and incrassate cells; lower hyaline
cells of stem-leaves sometimes septate, non-porose, non-fibrillose, those of the
upper one-half or two-thirds of the leaf fibrillose and porose as are the branch-
leaves also, towards the apex the hyaline cells much broader relatively, often as
broad as long; branches in different plants variable, 1-2.5 cm long, more or
less turgid below, acutely tapering at the apex, usually tv/o spreading with
drooping tips and two pendent and closely appressed to the stem, the comal
short, ascending, more or less blunt; the branches in cross-section showing a
layer of inflated cuticular cells which are rectangular, porose, and fibrillose;
branch-leaves usually about 2 mm long, sometimes 3 mm, widely ovate, very
concave, the margins involute, the apex abruptly and bluntly tapering, cucul-
late, at back somewhat scabrous with the erosion of the outer cell-walls, when
dry the leaves being more or less closely imbricate; hyaline cells of branch-
leaves broad, fibrillose, ventrally porose with large lateral pores mainly con-
fined to the cell-angles, the pores often equalling one-third the width of the
cell, dorsally the pores somewhat smaller and more elliptic and lateral, mostly
in the cell-angles; in cross-section the chlorophyllose cells are narrowly barrel-
shaped or somewhat trapezoidal, exposed on both faces, being ventrally nearly
flush with the ventral surface of the hyaline cells but the latter dorsally very
convex and projecting much beyond the chlorophyllose cells, the lateral walls
of the chlorophyllose cells smooth; perichstial leaves very large, broad'y oval,
cucullate, hyaline-bordered, rounded obtuse at apex: capsule at maturity con-
siderably exserted above the comal tuft; spores yellow, .028- 033 mm, mature
in mid-summer.
26 American Midland Naturalist Monograph No. 6
In bogs, margins of quiet rivers and lakes, wet places in woods, etc.; a cos-
mot>olitan, occurring in North America from Labrador and Alaska south to
British Columbia, California, Texas, and Florida.
In our region thus far found in Allegheny, Bedford, Butler, Crawford, Erie, Fayette,
Lawrence, McKean, Warren, Westmoreland, and Somerset counties.
4a. Sphagnum palustre var. squarrosulum (Nees and
Homschuch) New Combination
(5. cymbijolium var. squarrosulum Nees and Homschuch; S. latifolium var.
squarrosulum (Nees and Homschuch) Jennings, Manual, 1913)
As compared with the typical species this variety has usually a darker or
more bluish-green color; the leaves have a more abruptly narrowed apex, the
apical third of the leaf especially in the comal branches being rather abruptly
squarrulose.
Probably with a world-wide distribution with the typical form but in our
region more common and apparently more partial to less decidedly boggy
situations.
Now known from the following counties: Beaver, Blair, Centre, Crawiford, Fayette,
Indiana, Somerset and Westmoreland. Specimen figured: In crevices of rock-bed of river
near fails, Ohio Pyle, Fayette Couny, Sept. 1-3, 1906. O.E.J, and G.K.J.
4b. Sphagnum palustre var. brachycladum (Schliephacke)
New Combination
(5. cymbijolium var. virescens f. brachycladum Schliephacke;
S. latifolium var. brachycladum (Schliephacke) Jennings)
Bluish-green or glaucous, yellowish below; branches short and closely
placed along a short stem, giving the plant a congested appearance; leaves
rather loosely imbricated and at their tips slightly squarrulose.
Centre Co.: In bog at Scotia, in the "Barrens," Sept. 22, 1909. O.E.J. Somerset
Co.: Along rivulets in swamp on e. flank of Negro Mt., 3 miles from Salisbury. Paul R.
Stewart. July 2, 1944.
5. Sphagnum magellanicum Bridel
(5. medium Limpricht; S. cymbijolium var. compactum Russow)
Plate III
Deeply cespitose up to 8-10 cm, gray-green to bluish-green, rose-red to
purple-red above, brownish or somewhat bleached below; stems rather densely
branched, the cuticular sheath consisting of 3-5 layers, distinct, the outer cells
smallest, porose and weakly fibrillose, the wood-cylinder castaneous to rose-
red, thick-walled; stem-leaves about 1.5 mm. long, (1-2 mm), broadly lingulate-
spatulate, the upper margins and the broadly rounded apex fimbriate; the
upper hyaline cells of the stem-leaves usually fibrillose and dorsally porose;
branches usually short, two slender and appressed to the stem, pendent, and
two horizontally spreading or somewhat up-curved, thick-fusiform, the comal
Jennings: Manual of Mosses — I. Sphagnales 27
and upper more or less obtuse, the lower short-pointed; cuticular cells of
branches densely fibrillose, porose; branch-leaves usually densely but some-
times loosely imbricated, 1.5-2.0 mm long, broadly ovate, very concave, cucul-
late, the apex dorsally rough by erosion of the cell-walls, the margin consisting
of one or two very narrow cells which are often eroded away and the edge
left more or less dentate; hyaline cells of the branch-leaves rather densely
fibrillose, dorsally with a few rather large pores usually confined to the cell-
angles; chlorophyllose cells in cross-section small, elliptic, central, enclosed
deeply on both sides by the hyaline cells, the lateral walls smooth: capsule
considerably exserted; spores stated to be .024-. 028 mm, somewhat rust-
colored, finely punctulate.
In bogs, etc. Almost cosmopolitan; in North America occurring from
Newfoundland to Alaska south to British Columbia and Florida.
Butler Co.: On wet roadside at edge of woods, Semiconon Run, lYi mi. n. of
Conoqjenessing. Sidney K. Eastwood, March 24, 1935. Centre Co.: In a sinkhole
pond in the Barrens, near Scotia, July 17 and September 22, 1909. O.E.J, (figured).
Also from mountain bogs and glades in McKean, Clearfield, Clinton, Somerset, Fayette,
Forest, Westmoreland, and Warren counties.
Section II. Lithophloea
Cuticular cells of the stems and branches not reinforced with spiral fibers.
Branch-leaves more or less truncate and toothed at the apex.
Subsection I. Polyclada
Branches in fascicles of 7-13. Wood-strand of stem reddish brown.
Chlorophyllose cells of branch-leaves elliptical, central, the thick end walls
very narrowly exposed on both sides of the leaf.
6. Sphagnum Wulfianum Girgensohn
Plate II
Loosely tufted, ours rather slender, 7-12 cm high, the uppermost short
branchlets congested to form a dense head; stem reddish-brown, in cross-
section with an outer layer of rows of somewhat inflated cells, underneath
which is a dense reddish-brown layer of small thick-walled cells inside of
which the central part of the stem consists of larger thin-walled cells; stem-
leaves lingulate-triangular, deflexed, non-fibrillose, border with a margin of
several rows of much narrower cells, the apex rather widely erose-denticulate;
branchlets in clusters of usually 3-5 spreading and 3-7 pendent, about 1.2-2 cm
long, the spreading branchlets slender and gradually tapering, the pendent
branchlets linear; their cortical cells non-fibrillose but with an apical pore;
branch-leaves about 1 mm long, when dry recurved-spreading, more imbricated
when wet but still with somewhat spreading points; branch-leaves ovate-lanceo-
late, often somewhat incurved-nibulose at apex, bordered with two or three
rows of linear cells, the apex often erose-bidentate, the median hyaline cells
fibrillose, about 8-10:1 at base, shorter above, dorsally with a few ringed pores:
chlorophyllose cells broadly elliptic, thick-walled, the lumen about central, but
28 American Midland Naturalist Monograph No. 6
their thick walls very narrowly exposed on both surfaces, exceeded dorsally by
the convex outer walls of the hyaline cells.
This is a far northern species with a reported range south to Connecticut,
New York, Minnesota, and British Columbia, hence this collection is a further
extension of range southwards.
Columbus Bog, northern Warren County, Pa., Charles M. Boardman, Sept. 1, 1948
(figured).
This species is easily distinguished by the fascicles of 6-12 branchlets and
also by the dense capitulum of short apical branchlets.
Subsection II. Rigid a
With short, densely placed branches, and forming dense tufts. Cortical
cells of the branches each with a pore at the upper end. Chlorophyllose cells
of branch-leaves small, elliptic, completely enclosed but nearer the dorsal
(outer) surface of the leaf. Branch-leaves ovate, ending in an involute, mostly
widely spreading or ascending apex.
7. Sphagnum compactum DeCandol'le
(S. rigidum Schimper)
Plate III
Densely cespitose, gray-green or glaucous-green, brownish above, below
whitish or grayish-brown compactly and closely short-branched; stems stout,
low, in ours 4-8 cm high, with a cuticular sheath of usually 3 layers of cells,
the outermost cells largest, non-fibrillose, the wood-cylinder decidedly castan-
eous or sometimes yellowish; stem-leaves very small, 0.6-0.8 mm long, broadly
to equilaterally triangular-lingulate, the apex concave and broadly rounded or
truncate, erose-dentate, the margins rather widely hyaline-bordered; hyaline
cells of stem-leaves broadly rhomboidal, non-porose, non-fibrillose; branches
short, usually not over 1 cm long, 3 or 4 to a fascicle, horizontally spreading
or somewhat upcurved, the others slender and appressed-pendent; branch-leaves
when dry with the upper half of the leaf more or less squarrose-spreading,
large, 2-3 mm long, ovate, concave, the margins narrowly bordered, the uppsr
margins involute and often slightly erose-ciliate or erose-dentate the apvex
erose-dentate and cucullate; hyaline cells of branch-leaves rather broadly
rhomboidal, fibrillose, dorsally with several large, round pores irregularly
scattered and also in the cell-angles, the pores about two-fifths as wide as
the cell, sometimes a few oval and lateral, ventrally the pores sm.all, oval,
an^. located in the cell-angles; in cross-section the chlorophyllose cells are
elliptic, enclosed both dorsally and ventrally by the moderately convex hyaline
cells; cuticuar cells of the branches arge, short-rectangular, with one large
apical pore; fruit not seen.
Our plants seem to be the variety squarrosum Russow (Roth. Die Euro-
paeischen Torfmoose, p. 14. 1906) (5. rigtdum var. squarrosum Russow.
Braithwaite. The Sphagnaceae or Peat Mosses of Europe and North Amer-
ica, p. 58. 1880) . In bogs and wet woods, widely distributed in the Northern
Hemisphere, in North America occurring from the Arctic regions south to the
northern part of the United States.
Jennings: Manual of Mosses — I. Sphagnales 29
Centre Co.: At margin of pond under Pinus rigida. "Barrens," near Scotia. Sept.
22, 1909. (O.E.J, (figured). Erie Co.: In black ash swamp at head of Conneautte
Lake. Edinboro. O.E.J, and J. C. Fetterman. June 2?, 1919. Westmoreiand Co.:
Laurel Hill Mt.. 9 mi. s.e. of Rector. Hugh Mozingo. Oct. 7, 1945.
Subsection III. Squarrosa
Branch-leaves squarrose-spreading, their chlorophyl'ose cells in cross-section
trapezoidal to rectangular or barrel-shape, or triangular towards leaf-base,
dorsally more widely exposed, thick-walled, their hyaline cells with large pores.
8. Sphagnum squarrosum [Persoon] Schwaegrichen
Plate LXX
Loosely cespitose, bluish- to yellow-green: stems long, loosely branched,
with wood-cylinder hyaline to greenish or yellowish, cuticular sheath distinctly
2- (3) -layered; stem-leaves broadly oblong-lingulate, the ape.x broadly rounded
and erose-fimbriate, the leaves very narrowly bordered, slightly auriculate, non-
fibrillose, the hyaline cells above short and broad; branches 4 or 5, two or three
tumid, horizontal, the leaves on the lower two-thirds of the divergent branches
with squarrose tips; branch-leaves lanceolate to lance-ovate, very concave,
acuminate, with involute margins and usually slightly erose, marginal cells
narrower, but not usually forming a hyaline border; hyaline cells of branch-
leaves richly fibrillose, on both sides with numerous large round pores of about
one- fourth to one-third the width of the cell; in cross-section the chlorophyllose
cells free on both surfaces, narrowly rectangular to trapezoidal, when trape-
zoidal with the wider face dorsal, the faces thick-walled, the lumen more or
less elliptic, the hyaline cells strongly convex on both surfaces: spores yellowish
and finely roughened, about .022-. 025 mm in diameter.
In usually shaded locations in swamps, boggy springs, along woodland
streams, etc., in Europe, and, in North America, from the Arctic regions to
the northern part of the United States.
Now known from the following eight counties in the northwestern and mountainous
parts of our region: Cambria, Carion, Elk, Erie, Huntingdon, McKean, Mercer, and
Somerset. Fipured from a specimen collected in a wooded swamp m Cook Forest, by Adam
M. Barker. Sept. 15. 1935.
9. Sphagnum teres (Schimper) Aongstroem
(S. squarrosum var. teres Schimper; S. porosum Lindberg)
This species is represented in our region by a plant in varying degrees
p>erhaps best regarded as the following variety, which differs from the typical
form of the species mainly in having the divergent branches more or less
squarrose-leaved rather than distinctly terete.
9a. Sphagnum teres var. subteres Lindberg
(.$". teres var. subsquarrosum Warnstorf)
Plate III
Weakly and loosely but quite deeply cespitose, yellowish-green to dis-
tinctly yellowish: stems up to 15 or even 20 cm long, slender, the cuticular
30 American Midland Naturalist Monograph No. 6
sheath usually three-layered, the outer cells perhaps a little the largest, non-
fibrillose, usually not distinctly porose, the wood-cylinder strong, yellowish or
rarely castaneous; stem-leaves large, about 1.5 mm long, broadly triangular-
lingulate, the margin narrowly hyaline-bordered, the rounded to somewhat
truncate apex erose-dentate, the base often slightly auriculate; hyaline cells of
stem-leaves non-fibrillose, non-f)orose, in the lateral portions of the basal half
of the leaf often septate, the upper hyaline cells about as broad as long;
branchs 3 to 5 to a fascicle, usually two appressed- pendent and very slender, the
others widely divergent but somewhat recurved, rather slender, about 1-1.5 cm
long; branch-leaves when dry imbricate but with the apical half of some of
them squarrose, the leaves usually 1.5 mm long, ovate, concave, the narrowly
hyaline-bordered margin involute towards the apex; hyaline cells of branch-
leaves short, wide, both ventrally and dorsally fibrillose, and with a few large
round pores about half as wide as the cell and usually located in the cell-
angles; in cross-section the chlorophyllose cells in the apical third of the leaf
trapezoidal to barrel-shaped and exposed both dorsally and ventrally, wider on
the dorsal face, towards the base of the leaf sometimes triangular and exposed
only dorsally; cuticular cells of branches rectangular: spores not seen but said
to be brownish, papillose, and about .025 mm in diameter.
In bogs, wooded swamps, etc., in Europe and, in North America, in Can-
ada and the northern United States, probably distributed widely with the
type form.
Crawford Co.: In tamarack bog one and one-half miles s.e. of Linesville, June 7,
1904 (figured) and June 12, 1905. O.E. This station is now submerged. Erie Co.: In
swamp near south shore of Presque Isle, August 3, 1935. Nelle Ammons. Somerset
Co.: Millers Run. O.E.J.
Subsection IV. Cuspidata
Chlorophyllose cells of branch-leaves triangular to trapezoidal, exposed on
the dorsal (outer) surface, sometimes narrowly on the ventral surface.
10. Sphagnum recurvum Beauvois
(S. intermedium Hoffman; S. apiculatum Lindberg)
Plate III
Loosely but deeply tufted, pale green to greenish- or whitish-yellow: stem
light green, slender, long, in our region often up to 3 dm long, the cuticular
sheath rather indistinct and consisting of 3 or 4 layers of small or medium-
sized rather thick-walled cells; stem-leaves small, about 0.5-0 8 (-1.0) mm long,
equilaterally triangular to ovate-triangular, obtuse and slightly erose- denticu-
late; hyaline cells of stem-leaves rather small, mostly non-fibrillose and non-
porose, towards the base on each side of the leaf more or less septate and nar-
rowing to form a very wide border, which abruptly narrows above but reaches
almost to the apex; branches usually 4, two very slender and appressed-pendent,
two somewhat larger and irregularly spreading; cuticular cells of the branches
elongate-rectangular, perforate and somewhat recurved at the apex, like those
of the stem non-fibrillose; branch-leaves lance-ovate, imbricate, in our region
Jennings: Manual of Mosses — I. Sphagnales 31
ranging from 1-2 mm long, when dry, with undulate margins, flexuose and with
a recurved apex, when moist straight and erect-appressed, tapering to a rather
narrowly obtuse apex with two or three teeth, the margin involute above; hyaline
cells of branch-leaves fibrillose and porose, above the midde rather narrow,
ventrally usually with large pores in the cell-angles of about one-third the width
of the cell, dorsally with small end-pores or sometimes a very few rather dis-
tinctly ringed lateral ones; in cross-section the hyaline cells are ventrally quite
convex, the chlorophyllose cells triangular or rarely trapedoidal, usually ex-
posed only on the dorsal face; perichsetical leaves large, broadly oval, concave,
pointed: spores smoothish, yellow, about .025 mm in diameter.
A cosmopolitan species occurring in North America from Newfoundland
to Labrador and south to the Gulf States.
This is probably our commonest sphagnum, occurring not only around the borders
of bogs but out in seepage areas in open] hillside fields. It is now known from seventeen
counties in western Pennsylvania, most of the records being from the southwestern and
northwestern parts of our area. The following variety amblyp.hyllum seems to be the com-
moner form in the central uplands and mountains. Figured from specimens collected May
29-31, 1910, around Mud Lake, Hartstown, Crawford County, O.E.J. & G.K.J.
10a. Sphagnum recurvum var. amblyphyllum (Russow) Wamstorf
(S. amblyphyllum Russow)
TTie variety has the stem-leaves more spatulate-triangular, with a more
rounded and somewhat erose-denticulate apex; the cuticular sheath is less
plamly differentiated and the cells are more incrassate than in the typical form
of the species. All possible intergradations seem to be represented among the
specimens examined. It may usually be recognized when dry by the strongly
undulate branch-leaves, these usually still remaining undulate when mounted
wet for examination under the microscope.
Now known from eight counties from Centre west to Crawford and Allegheny coun-
ties through the middle of the area.
10b. Sphagnum iRFXURVUM var. tenue Klinngraeff
(S. angustifolium Jensen; S. recurvum var. parvifolium Warnstorf; S. amblyphyl-
lum var. parvifolium Warnstorf; S. parrifolium (Sendt. ) Warnstorf)
Plate VI
Softly and loosely cespitose, yellowish- to grayish-green, or brownish above:
stems slender, usually at least 10-12 cm high, the wood-cylinder yellowish and
without any distinctly differentiated cuticular sheath: stem-leaves small, usually
0.5-0.7 mm long, equilaterally triangular to somewhat triangular- lingulate, the
apex rounded or somewhat truncate, erose-dentate, the hyaline border narrow
above and very wide below; hyaline cells of stem-leaves non-fibrillose, non-
porose, a few septate towards the base on each side of the median region;
branches 3-5, two being slender and appressed-pendent, two or three short, 5-9
mm long, divergent, recurved at the tips; branch-leaves lanceolate, about 1 mm
long, concave, the uniformly narrowly hyaline-bordered margin involute
towards the narrowed, slightly truncate-erose apex, leaves when dry more or
32 American Midland Naturalist Monograph No. 6
less undulate, loosely imbricate, with widely spreading or recurved tips; hyaline
cells of branch-leaves narrow, fibrillose, ventrally with rounded medium-sized
pores in the cell-angles, dorsally with single smaller round pores in the cell-
angles or sometimes in rows laterally; in cross-section the chlorophyllose
cells triangular and only dorsally exposed, or more usually trapezoidal and
free on both faces, the dorsal face wider, the hyaline cells more convex ven-
trally: fruit not seen.
In bogs, swamps, etc., probably widely distributed. In North America
extending from the Arctic south to northern United States.
In o-.ir region known from one locality. Blair Co.: Springy mountain slop>e, Rhodo-
dendron Park, Llyodsville, October 18, 1901. J.A.S. (figured).
11. Sphagnum cuspidatum Ehrhart
{S. virginianum Warnstorf)
Plate IV
Usually pale and slender messes of wet bogs or often submerged in pools.
Stem with 2 or 3 layers of non-porose, non-fibrillose, large and thin-walled
cortical cells clearly distinct from the wood-cylinder. Stem-leaves isosceles-
triangular, about 1-1.4 mm long and about 0.6-1.1 wide, inrolled above, slightly
toothed at the narrow truncate tip; the border strong, considerably widened
below; the hyaline cells fibrillose and porose. Branches usually 4, two of them
drooping or slightly spreading, the other two spreading linear-lanceolate to
elongated-lanceolate, about 1.5-3 mm long, the tip toothed, often falcate-
secund, the leaves often undulate when dry; the border of 2-4 (3-8) rows of
linear cells. Hyaline cells ranging from the basal linear-rhomboidal cells
about 15 times as long as wide upwards to those only about 5 times as long
as wide, all fibrillose and somewhat porose. Chlorophyllose cells trapezoidal
to 4-sided, both surfaces free, but the dorsal (abaxial) surface wider.
In wet bogs and pools, often submerged, Eurasia and from Georgia to
Newfoundland.
Allegheny Co.: On springy hillside about one mile west of Thornburg, July 12,
1922. Mrs. Alice B. Lord (figured).
Subsection V. Subsecunda
A difficult group of diverse forms. Branch-leaves mostly more or less
s.^cund, the pores of their hyaline cells mostly small and numerous, their
chlorophyllose cells truncately elliptic or trapezoidal, mostly central and ex-
posed on both surfaces.
12. Sphagnum contortum Schultz*
(S. subsecundum var. contortum Huebener; S. laricinum Spruce)
Loosely cespitose, green to brownish or yellowish, sometimes more or less
* Andrews, A. L. (N. Am. Flora 15: 21-22. 1913) treats S. contortum, S. platy-
phyllum, S. auriculdtum, S. subsecundum, and S. inundatum as being a single polymor-
phic species, S. subsecundum.
Jennings: Manual of Mosses — I. Sphagnales 33
purplish above: stem about 6-12 cm high, the wood-cylinder reddish to brown-
ish, surrounded by a distinct two-layered sheath of inflated cells; stem leaves
small, about 1 mm long, broadly lingulate or triangular-lingulate, the hyaline
border much broader towards the br.se, the apex broadlv rounded and mere or
less concave, cucullate, and erose-fimbriate; hyaline cells of stem-leaves in upper
third fibrillose, short and broad, ventrally with a few cells in the angles, dor-
sally with more numerous small ringed pores along the sides of the cell, very
few of the hyaline cells septate, the lower ones long and narrow; fasciculate
branches 3-5 to a fascicle, usually two slender and closely appressed pendent,
two divergent and recurved; branch-leaves about 1.5-2 mm long, broadly ovate
to lanceolate, more or less sharply acuminate, the upper margin involute and
narrowly hyaline-bordered, leaves when dry more or less subsecund and sub-
lustrous; hyaline cells richly fibrillose, slender, ventrally almost poreless, dor-
sally with small ringed pores more or less completely arranged in bead-like
rows, the pores most numerous towards upper margins of leaf; in cross-section
the chlorophyllose cells narrowly barrel-shaped, with both faces free and their
walls there somewhat thickened; cuticular cells of branches apical ly porose:
sp>ores not seen but reported as .020-. 030 mm in diameter, yellowish-brown,
finely roughened.
In swampy meadows, along ditches, margins of bogs, etc., in Europe and,
in North America from Greenland to Mexico, and along the Pacific Coast
Not heretofore reported from our region but a specimen collected by J. A.
Shafer, October 20, 1901, at Ohio Pyle, Fayette County, with stem-leaves
about .7-.8 mm long, with the margin uniformly hyaline-bordered and the
hyaline cells fibrillose to below the middle of the leaf is now referred here;
also a specimen from Centre Co., Neil D. Richmond, June 14, 1950.
13. Sphagnum platyphyllum (Sullivant) Warnstorf
(S. auriculatum Aongstroem; S. isophyllum Russow)
Plate V
Loosely cespitose, brownish- to grayish-green: stems in our region up to
10 cm high, slender, rather weak and sparsely branched; stem in cross-section
showing a usually brownish wood-cylinder, with a distinct cuticular sheath of
rather small, thin-walled, and usually uni-porose cells; stem-leaves large, usually
1.3-2.0 mm long, oval to oblong from an auriculate base, very concave, the
apex blunt and a little toothed or erose, the margin narrowly and uniformly
bordered; hyaline cells of the stem-leaves in lower half to two-thirds of the leaf
non-fibrillose and non-porose but some of them septate, in the upper half or
one-third of the leaf the hyaline cells fibrillose and on both sides with lateral
rows of small pores; branches usually 3, sometimes 4, usually spreading with
recurved tips, one or two being pendent and very slender; branch-leaves broadly
ovate, very concave, usually 2-3 mm long, the apex toothed, the margin more
or less incurved and with a narrow and uniform border; in cross-section the
chlorophyllose cells barrel-shaped, free on both surfaces, the hyaline cells
about equally convex on both sides; hyaline cells fibrillose, with numerous
small lateral pores on both sides; \\hen dry the leaves towards the base of the
34 American Midland Naturalist Monograph No. 6
spreading branches more or less sub-secund; spores stated by Warnstorf to be
.023-.028 mm in diameter, yellowish and finely papillose.
In turfy swamps and bogs in Europe and North America, extending in
the latter country from Massachusetts to Louisiana, also to Ohio.
Butler Co.: Open swampy pasture, Crider's Corners, Apr. 26, 1908. O.E.J.
14. Sphagnum auriculatum Schimper
(S. Gravetti Russow, p.p. — Warnstorf; S. subsecundum var. intermedium Warnstorf)
Plate V
Densely cespitose, grayish or glaucous green, light yellow below; stems
rather short (In our specimens about 5 cm), densely branched; branches in
fascicles 3-5, two or three of these drooping from a horizontally spreading base,
terete and rather thick, up to 1 cm long, the other one or two slender, and
rather closely appressed to the stem; in cross-section the cortical cells distinct,
in one layer, the outer cells of- the central axis much thickened and small;
stem-leaves large, 1.5-2.0 mm long, about half as wide, concave, from the dis-
tinctly auriculate base oval- Ungulate, the rounded apex narrowly toothed, some-
what cucullate, the margin narrow and of equal width from base to apex; cells
of the stem-leaves fibrillose to the base or nearly so, only rarely septate, ven-
trally with rather large poorly defined pores in the cell-angles, rarely none,
dorsally with numerous distinct pores along each side of the cell, the pores
circular to elliptic and about one- fourth the width of the cell; retort cells of
the branches with a distinct neck and terminal pore; lower branch-leaves large,
about 2 mm long, widely ovate, about 1.5 mm wide, very concave, the margins
more or less involute, the apex somewhat spreading, narrowly toothed, the
upper leaves more closely imbricated and lanceolate; cells of branch-leaves
ventrally with rather few large indistinct angle-pores, dorsally with numerous
distinct pores in a row along each side, as in the stem-leaves, the hyaline cells
usually with 8-10 spiral fibrils, the border of 2-4 thick-walled, linear-prosenchy-
matous cells; chlorophyllose cells in cross-section barrel-shaped with both
ends exposed: fruit unknown
In wooded swamps and wet shaded places, Europe and North America.
Known from the following collections in our region. Fayette Co. : Near Seaton's
Lake. Hugh N. Mozingo. April 7, 1946; and in wet cavities ini rocks, Ohio Pyle near
Falls, O.E.J., June 14, 1908 (figured). Somerset Co.: Shafer Run, 2 mi. n. of Baker-
vilie, Hugh N. Mozingo, Sept. 14, 1946. Venango Co.: Near Lisbon, Mrs. E. J.
Mason. Oct., 1947. Westmoreland Co.: Springy hillside, Mellon estate, "Rachel-
wood," New Florence, Sept. 9-11, 1907, O.E.J.
15. Sphagnum subsecundum Nees
Plate IX
Moderately densely cespitose, green to yellowish or brownish: stems 5-20
cm long, with a dark or purplish-brown wood cylinder, with a cuticular sheath
of one layer of moderately inflated cells; stem-leaves small, about 0.6-0.8 mm
long, broadly short-lingulate, somewhat auriculate, the margin broadly hyaline-
bordered below, the border narrowing and becoming fimbriate towards the
broadly erose-fimbriate apex, the upper half of the stem-leaves often distinctly
JtNNiNGS: Manual of Mosses — I. Sphagnales 35
concave and more or less cucullate; hyaline cells of the stem-leaves broad above,
usually all non-fibrillose, rarely a few septate, sometimes porose; of the 3-5
fasciculate branches two or three are variously divergent, short, usually 6-8 mm
long, slender and sometimes flagelliform; branch-leaves small, 1-1.5 mm long,
very concave, broadly ovate to lanceolate, acuminate to a narrowly
truncate or 3-5 toothed apex the margins uniformly narrowly hyaline-
bordered, involute, when dry closely imbricate to more or less sub-secund;
hyaline cells of branch-leaves narrow, richly fibrillose, ventrally non-porose,
or with a few small non-ringed pores in the cell-angles, dorsally with numerous
small ringed pores along the sides of the cells; in cross-section the chlorophyl-
lose cells narrowly barrel-shaped, relatively rather large as compared with the
hyaline cells, free on both faces, the hyaline cells but slightly convex on either
side: spores not seen from our region, finely papillose, yellowish, and .025-.028
mm in diameter.
In wet meadows, swamps, ditches, bogs, etc., in Europe and in Asia and,
in North America, from Newfoundland to Alabama. In our region frequent;
approaching the variety brachycladum Warnstorf in having stem-leaves more
or less cucullate and the divergent branches often only about 5 mm long.
Erie Co.: In bog ac south end of Cranberry Pond, Presque Isle. May S-'), 1906.
O.E.J, (figured). Also known from Elk, Centre, Clarion, Clearfield, Mercer, Butler,
Westmoreland, Fayette, Somerset, and Bedford counties, but not known from the south-
wfcstern border counties.
16. Sphagnum inundatum Russow, Warnstorf
Densely and deeply cespitose, gray or yellowish-green: stems usually 15-30
cm long, more or less completely submerged; branches with moderately
densely imbricate leaves; stem-leaves usually somewhat fimbriate at the narrow
apex, little or not at all auriculate, fibrillose only above the middle; branch-
leaves dorsally richly porose in lateral bead-like rows, ventrally with only a
few pores located in the cell-angles. Other characters are as described for
the variety auriculatiim.
In wet meadows, wooded swamps, bogs, etc. In cooler Europe, Asia, and
North America. In our region, so far as now known, represented only by
the following variety.
16a. Sphagnum inundatum var. auriculatum (Warnstorf) Roth
{S. contortum var. laxutn Roell)
Plate IX
Only moderately cespitose, green: stems in our specimens only about 6-8 cm
high, only occasionally completely submerged; wood-cylinder greenish, sur-
rounded by a cuticular sheath of one layer (ossacionally unsymmetrically two)
of inflated more or less distinctly porose cells; stem-leaves 1.2-1.5 mm long,
about three-fifths as wide, distinctly auriculate, towards the apex somewhat
concave, the margins narrowly uniformly hyaline-bordered and toward the
ap>ex involute, the narrow apex somewhat dentate but not fimbriate; the
hyaline cells of stem-leave.« broad, towards the lateral portions of the base
becoming narrower, usually septate, fibrillose at least as far down as the
36 American Midland Naturalist Monograph No. 6
middle of the leaf, or farther, and usually also fibrillose at the base of the
leaf, above ventrally with rather small distinct pores in the cell-angles and
usually other less distinct lateral pores, above dorsally with small pores in
cell-angles and numerously along the sides of the cells; of the usually 5 fascicu-
late branches two are pendent and the others short, usually 6-9 mm long, vari-
ously widely divergent; branch-leaves when dry very lax and widely divergent,
1.5-2 mm long, ovate, very concave, with involute, narrowly and uniformly
hyaline-bordered margins, the apex narrow and dentate-truncate; hyaline cells
of branch-leaves rather long and slender, richly fibrillose, dorsally with laterally-
placed bead-like rows of small pores about one-fifth as wide as the cell, ven-
trally with small ringed pores in the cell-angles, occasionally also a few laterally
arranged indistinct pores; cuticular cells of branches large with a short neck
and terminal pore; in cross-section the chlorophyllose cells narrowly elliptic
with about equally free and thickened faces: for the type of the species the
spores are stated to be yellow ,and about .030-. 03 5 mm in diameter; of the
variety the spores have not been seen.
Centre Co.: Headwaters of Laurel Run, Tussey Mt., above Shingletown, July 15,
1909. O.E.J. Fayette Co.: In pools and wet crevices in rocky bed of river above falls,
Ohio Pyle, September 1-4, 1906. O.E.J, and G.K.J, (figured).
17. Sphagnum pungens Roth
(S. contortum var. gracile Roeil)
Plate IX
Rather loosely cespitose, bluish-green, when dry sub-lustrous above, yellow-
ish or brownish below: stems rather stout, often forking, in our specimens up
to 6 or 7 cm high; wood-cylinder greenish or pale, enclosed in a one-layered
cuticular sheath which in places is unsymmetrically often two-layered; stem-
leaves broadly lingulate, about 1-1.5 mm long, at base about three-fifths as
wide, somewhat auriculate, the uniformly narrowly hyaline-bordered margin
somewhat erose-fimbriate towards the broadly rounded erose dentate apex; the
hyaline cells of stem-leaves broad, rarely septate, distinctly fibrillose in upper
two-thirds of leaf, ventrally with a few indistinct pores in the angles and along
the sides of the cell, dorsally with numerous small pores arranged in lateral
bead-like rows; of the usually 4 fasciculate branches, two are slender and ap-
pressed-pendent while the other two are horizontally divergent and recurved,
about 1-1.5 cm long, the lower and median leaves of the divergent branches
more or less widely squarrose, the upper ones imbricate so that the branch
ends in a sharply acuminate point; branch-leaves broadly ovate to lanceolate,
large, 1.8-2.6 mm long, concave, the uniformly narrowly hyaline-bordered
margins involute towards the acuminate few-toothed apex; hyaline cells of
branch-leaves narrow, long, richly fibrillose, ventrally with a few indistinct •
pores in the cell-angles, dorsally with numerous small ringed pores about one-
fourth to one-fifth as wide as the cell and arranged in bead like rows along
the sides of the cell; in cross-section the chlorophyllose cells relatively large,
narrowly barrel-shaped, free on both faces, the hyaline eel's not being markedly
convex on either face; cuticular cells of branches long-rectangular with a short
neck and a large apical pore: spores not known from our region.
Jennings: Manual of Mosses — I. Sphagnales 37
More or less intermediate between S. inundatum (Russow) Wamstorf,
and. S. auriadatum Schimper, and of doubtful status. Heretofore reported,
so far as known to the present writer, only from Europe, where it occurs in
swampy meadows.
Centre Co.: Bog in sink-hole in pine-barrens near Srotia, July 17, 1909. O.E.J,
(figured).
Subsection VI. Acutifolia
Branch-leaves small to medium size, lanceolate to lance-ovate, acute to
narrowly acute and truncate apex, their chlorophyllose cells in cross-section
triangular or elliptic-trapezoidal, nearer to and more or less widely exposed
on the ventral (inner) side and sometimes narrowly exposed dorsally between
the bulging hyaline cells.
18. Sphagnum fimbriatum Wilson
Plate VI
Loosely cespitose, grayish-green to yellowish brown: stems rather slender,
usually 4-5 cm high, sometimes much longer, in cross-section showing a
cuticular sheath of 2-3 layers of cells, the cells of the outer layer largest and
porose; stem-leaves very widely obovate-spatulate, about 0.7-0.8 mm, the upper
half broadly rounded and erose-fimbriate; hyaline cells of stem-leaves non-
fibrose, non-porose, very wide above the middle of the leaf, towards the base
often one- to several-septate, the hyaline border towards the base widening to
about one-third the width of the leaf on each side; fasciculate branches 3 or 4,
usually two slender, arcuate, and decurved, and up to 2.5 crr« long, the other
one or two pendent, rather closely appressed to the stem, filiform; branch-
leaves closely imbricated, shortly ovate-lanceolate below to slenderly lanceolate
above, concave, the upper margin incurved, the apex narrowly truncate and
dentate; hyaline cells rather small with four to six fibrils, ventrally with a few
round pores which are often almost as wide as the cell, dorsally with more
numerous lateral pores above one-third as wide as the cell; in cross-section the
chlorophyllose cells trapezoidal, free on both surfaces, the inner surface widest,
the hyaline cells extending convexly considerably beyond them on the dorsal
face; cuticular cells of branches without distinct necks; perichaetial leaves large,
obtusely ovate: spores stated to be smooth, yellowish-brown, about .025-.030
mm in diameter.
Usually in low-lying bogs and marshes, or along the borders of streams,
Europe, Asia, South America, and, in North America. frr>m the Arctic regions
through Canada to the northern part of the United States.
Now known from the following counties- Butler, Clearfield, Crawford, Erie, Elk,
McKean, Warren, Westmoreland and Somerset. Figured from specimen collected in
Pymatuning Swamp, near Linesville, June 7, 1904. O.E.J. Also Forest and Mercer counties.
C.M.B.
19. Sphagnum Girgensohnii Russow
Plate VII
A tall, slender, loosely cespitose, grayish green or translucent green species
with stems up to 10-15 cm in height. Stem in cross-section showing a cuticu-
lar sheath of 2-3 layers of thin-walled, non-fibrillose porose, and a zone of
38 American Midland Naturalist Monograph No. 6
much smaller dense cells which in turn surround a central core of larger
thinner-walled cells. Stem-leaves about 1-1.2 mm long, short-lingulate, often
almost as wide as long, the blunt broad apex partially erose-fimbriate, the
sides bordered above by a narrow margin of thick-walled linear cells, the
border widening at the base into a triangular patch, each patch about one-
fifth the width of the base; the upper middle hyaline cells about 3 times as
long as wide, non-fibrillose. Branches usually 4; two of them slenderly
attenuate and accumbent to the stem, the other two spreading, about 1.5-2.5
cm long, gradually acuminate, mostly spreading-falcate. Branch-leaves imbri-
cate and in the thicker part of the branch lanceolate to lance-ovate, about 1-1.4
mm long with the apex more or less squarrose-spreading, often involute tubular
and sometimes erose at the tip. Branch-leaves narrowly bordered, the median
cells ring-fibrillose and with many lateral pores between the fibrils. Chloro-
phyllose cells triangular-trapezoidal in cross-section, with the broader face
nearly flush with the upper (adaxial) surface of the leaf, the lower face
nearly enclosed between the strongly convex hyaline cells.
Widely distributed in boreal regions extending south in North America to
New Jersey, West Virginia, Ohio, Minnesota, and Oregon. Reported by
Boehner as common in swamps and bogs in Cattaraugus County, southwestern
New York.
Known/ from Elk, Fayette, McKean, Mercer, Somerset, and Tioga counties, in moun-
tain and upland swamps. Figured from sptecimen collected by Edmund W. Arthur, in a
wooded swamp. Swamp Root, Mercer Co., Sept. 20, 1946.
20. Sphagnum Warnstorfii Russow
(S. actttifolium var. gracile Russow)
In swampy meadows, margins of bogs, etc., in Europe and, in North
America, from Greenland to Pennsylvania and westward to the Pacific States
and Alaska. The sp>ecies varies from bright green to yellowish or from red
to purplish. The green variety has been found in our region but once, its
characters being as follows, but it is doubtfully different enough to merit
varietal status.
Crawford Co.: Pymatuning Swamp, Hartstown. O.E.J. May 29-30, 1915 (Det.,
C.M.B.). Fayette Co.: Cranberry-sphagnum glade, 1 mi. w. of Markleyfburg P.O.
O.E.J, and C.M.B. Aug. 25, 1949.
20a. Sphagnum Warnstorfii var. virescens Russow
Plate VI
Rather densely cespitose, bright green above, bleached or yellowish below:
stems in our specimens from about 5-12 cm high, the wood-cylinder green to
red and surrounded by a cuticular sheath of three layers of inflated cells, the
middle cells usually being the largest; stem-leaves about 1 mm long, broadly
lingulate, not auriculate, rather abruptly rounded to a narrow erose-dentate
somewhat concave apex, the margin very broadly hyaline-bordered below but
abruptly narrowing above and continuing rather narrow to the apex; hyaline
cells in upper half of stem-leaf broad, many of them once (or twice) septate,
Jennings: Manual of Mosses — I. Sphagnales 39
in the lower half of leaf the hyahne cells broad only in a narrow median strip
finked on both sides by narrow elongate cells, usually all hyaline cells of stem-
leaf non-fibrillose and non-porose; fascicu'ate branches usually 4, two very slen-
der and closely appressed pendent, and tv/o horizontally divergent, rather slen-
der, somewhat recurved, about 1-1.5 cm long, the comal branches short, obtuse,
ascending to erect; branch-leaves rather indistinctly five-ranked, when dry with
more or less spreading tips, ovate-lanceolate, concave, 1.5-2.0 mm long, the
margins uniformly narrowly hyaline-bordered and involute to the quite nar-
rowly acuminate and truncate-erose apex; hyaline cells of branch-leaves richly
fibrillose, ventrally with one or two large round m.edian pores of one-half to
two-thirds the width of the cell, these pores usually more numerous towards
the margin of the leaf, dorsally with quite numerous, small, elliptic, ringed
pores in the angles and along the sides of the cell; in cross-section the chloro-
phyllise cells narrowly trapezoidal with the ventral face wider, both faces
usually free, sometimes enclosed dorsally, the hyaline cells being dorsally quite
convex; the cuticular sheath of branches with Ions rectanc'ular cells with indis-
tinct necks and apical pores: spores for the species stated to be dark yellow,
minutely roughened, and about .025-. 030 mm in diameter.
Mercer Co.: Near Houston Jet. J.A.S. July 12. 1902 (figured).
21. Sphagnum fuscum (Schimper) Klingr
{S. acutifolium var. fuscum Schimper)
Plate VIII
Slender plants in dense, soft, deep, brownish tufts. Stems erect, about
5-10 cm tall. Branches usually 3, cf which two diverge almost at right angles,
then droop; the other branch hanging down alongside the stem. The diverg-
ent branches are about 1-1.5 cm long tapering to a slender attenuate tip. Stem
in cross-section showing 3-4 layers of larger, rounded-rectangular, thin-walled
cells, underlain abruptly by a layer of thick-walled, very much smaller reddish
brown cells, these grading abruptly into the central core of somewhat larger
colorless cells. Branch-leaves in lower third of divergent branches 1-1.4 (1.5)
mm long, somewhat spreading, lance-ovate, with upper margins incurved up to
a narrowly truncate toothed apex. Leaves in the attenuate tip lance linear and
clasping at base. Lower central hyaline cells of the branch-leaves about .115
mm long, transversely fibrillose, and with a few large dorsal pores, these being
mostly in pairs at the com.misures. Chlorophyllose cells narrowly triangular,
exposed, on both surfaces, but more so on the ventral surface, on the dorsal
surface deeply set between the strong convex hyaline cells. Stem-leaves nar-
rowly oblong-lingulate, not much, if any, v/idened at has, about 1.1-1.6 mm
long, about 2 J/2 to 3 times as long as wide, rounded and with a narrowly
erose apex. The border wide at the base, narrowing rapidly towards the apex.
In our region it was collectpd in Waterman Swamp, Cattarasjus Co., soithwestcrn
N. Y. (Boehner. Science Studies, Bonaventure College, 9: 4. 1941) and at Mt. Jewett,
Potter Co., northern Pennsylvania. O.E.J. Sept. 12. 1922 (figured). Cravcford Co.:
Pymatuning Swamp, May 29-31, 1915. O.E.J. McKean Co.: Cathrine Swamp. C.M.B.
Pi. 82. Sept. 2, 1948.
40 American Midland Naturalist Monograph No. 6
Springy places and swamps, Eurasia and North America, south to northern
Pennsylvania, Michigan, Minnesota, Colorado, and Washington.
The brown color of this moss constitutes the most obvious distinction be-
tween it and the usually red S. Warnsiorfi.
22. Sphagnum quinquefarium (Lindberg) Wamstorf
{Sphagnum acutifolium var. quinquefarium Lindberg)
Plate VI
Pale green or yellowish-green, reported as more or less rose-tinted above,
but in our region not rose-tinted so far as yet known, deeply and densely
cespitose: stems up to 10 cm, often forking, densely fasciculately branched, in
cross-section showing a yellowish or pale "wood-cylinder, the cuticular sheath
composed of 3 or 4 layers of large cells; stem-leaves lingulate-triangular from
a wide slightly auriculate base, rather large, about 1.2-1.8 mm long by about
three-fifths as wide, rounded above to a narrowly erose-truncate apex, the
margins narrowly hyaline-bordered and somewhat involute towards the apex,
towards the base widely bordered; hyaline cells of stem-leaves in median
portion and towards the apex widely rhomboid, in the upper half of the leaf
septate, usually faintly fibrillose and occasionally porose, in the lateral basal
portion septate, rapidly becoming very narrow outwards and merging there
into the broad hyaline border; brrnches usually 4 or 5 in a fascicle, usually 2 or
3 widely divergent, the comal short, dense, and widely ascending to erect;
branch-leaves oval to ovate, about 1.5 mm long, concave, with involute nar-
rowly hyaline-bordered margins, above quickly narrowed to a rather broad
dentate-truncate apex; hyaline cells of branch-leaves large, fibrillose, below
ventrally with a few small rounded pores in the cell-angles, the median lateral
cells with a few small rounded pores in the cell-angles, the median lateral
cells with a few larger indistinct pores, dorsally above with characteristic more
or less elliptic pores of about one-third the width of the cell and situated in the
cell-angles or along the sides; in cross-section the chlorophyllose cells rather
broadly triangular, ventrally free but dorsally enclosed between the highly con-
vex hyaline cells; cuticular cells of branches large, inflated, with a distinct neck
and apical pore: spores stated to be smooth, yellowish and about .021 -.025 mm
in diameter.
In bogs, etc., in Europe and, in North America, from Newfoundland to
New England and south along the mountains to the Carolinas.
Rare in our region. Clinton Co.: Along Hyner's Run above Hyner, July 14, 1908.
O.E.J, (figured).
23. Sphagnum plumulosum Roell
{S. subnitens Russow and Wamstorf; S. acutifolium var. subnitens Dixon)
Plate V
Densely cespitose, pale to grass-green, usually reddish to violet above:
stem in typical specimens 10-15 cm high, but in our region usually about 6-8
cm high, the wood-cylinder green to red, the cuticular sheath distinct, 2-4-
layered, with the outer cells largest: stem-leaves large, 1-1.5 mm long, broadly
Jhnnings: Manual of Mosses — I. Sphagnai.es 4i
triangular lingulate, the apex erose-truncate and toothed, the hyaline border of
margin narrow above, very wide below; hyaline-ceils of stem-leaves broadly
rhomboidal towards the apex and in median basal portion of leaf, towards
lateral basal portions rapidly much narrower and septate, all non-fibrillose and
non-porose; branches 3-5 in a fascicle, usually two of these variously divergent,
rather slender, terete, about 1-1.5 cm long, the others very slender and ap-
pressed-pendent; branch-leaves ovate, concave, about 1.5 mm long, the nar-
rowly hyaline-bordered margin towards the apex involute, the blade towards
the apex gradually narrowed towards an erose-dentate point, the leaves when
dry imbricate with more or less of a metallic lustre, not distinctly 5 seriate;
hyaline cells of branch-leaves fibrillose, rather broad, ventrally with usually
two or three median, large, round, ringed pores about one-third to one-half as
wide as the cell, occasionally a few pores also in the cell-angles, dorsally with
about 6-10 elliptic pores about one-third as wide as the cell and situated along
the sides and angles of the cell; in cross-section the chlorophyllose cells small
and shortly sub-rectangular to triangularly trapezoidal, situated much nearer
the ventral leaf-surface with the wider ventral face free, the narrower dorsal
face free or enclosed between the dorsally highly convex hyaline cells; cuticular
cells of branches inflated, short, with a distinct neck and terminal pore: spores
stated to be yellow, papillose, about .025- 030 mm in diameter.
In bogs, swamps, etc., widely distributed in the cooler parts of the
Northern Hemisphere, in North America occurring from Greenland and
Labrador south to New Jersey and Pennsylvania, and in Cal-fornia and Brit-
ish Columbia. In our region apparently represented only by the following
variety, more properly a form, as follows:
23a. Sphagnum plumulosum f. viride (Warnstorf)
New Combination
(S. subnitens var. viride Warnstorf)
Th:s form differs from the typical species in that the tufts are low and
entirely green or often bleached out below.
In deep, shaded swamps and bogs within the range of the type.
Crawford Co.: Shaded boggy margin of Mud Lake, Hartstown, May 29-31, 1909.
O.E.J, and G.K.J, (figured).
24. Sphagnum acutifolium Ehrhardt
(5. capillifolium (Ehrh. ) Russ. Sl Wamst.; S. capillaceum (Weiss) Schrank)
The typical form of this species has green to pale or variously reddish to
puiplish tufts with often short stout stems and a hyaline to yellowish or red-
dish wood-cylinder; the other characters are as described below for the variety
viride, to which some of our collections belong, although color differences are
here perhaps of not much taxonomic value.
Cambria Co.: Cresson, James. (Porter's Catalogue). Erie Co.: Lo • i le. L.K.H.
1950. HiTNTiNGCON Co.: Warrior's Ridge, Porter. (Porter's Catalogue). Warren Co.:
Columbus bog. C.M.B. 1948. It has been found also in Allegheny, Butler, Clearfield,
Crawford, Fayette, Forest, McKean, Mifflin, Somerset, and Westmoreland counties.
42 American Midland Naturalist Monograph No. 6
24a. Sphagnum acutifolium var. viride Warnstorff
(S. capillifolium var. viride Jennings)
Plate V
Rather densely cespitose, low, yellowish above, greenish to yellowish-green
below, lacking the reddish tinges so often characteristic of the species: stems
slender, in our region usually 5-8 cm long, in cross-section showing a yellov/ish
wood-cylinder and a distinct cuticular sheath of 2-4 layers of large but non-
porose cells; stem-leaves oval-triangular to lingulate-triangular, 1-2 mm long,
always widest at the base, towards the apex abruptly narrowed to a truncate
apex with a few teeth, the upper margin usually somewhat involute, the margin
narrowly hyaline-bordered, the border sometimes wider at the base; hyaline
cells of stem-leaves largely once-septate, especially below the middle, those of
the upper half of the leaf usually more or less completely fibrillose and some-
times distinctly laterally porose; branches fairly numerous, usually in fascicles
of four, two spreading-recurved and two appressed-pendent and very slender;
the cuticular sheath of branches composed of cells with a distinct neck and
terminal pore; branch-leaves 1-2 mm long, ovate-lanceolate, when dry hardly
secund but with slightly spreading tips, concave, with involute margins above,
uniformly narrowly hyaline-bordered, the narrow apex somewhat erose-dentate;
hyaline cells of branch-leaves rather slender, abruptly fibrillose, with small
somewhat elliptic pores at the cell-angles, sometimes also lateral pores of a
similar character between the angle-pores on both sides of the leaf, while towards
the margin of the leaf the pores are often larger and more numerous; in cross-
section the chlorophyllose cells are more or less trapezoidal, unusually short,
free on both surfaces but the hyaline cells projecting far beyond them both
ventrally and dorsally, especially dorsally; perichaetial leaves said to be very
large and broadly ovate: spores yellow, smoothish.
In cool, boggy situations in Europe, Asia, South America, and in the
regions of the South Pacific. In North America extending from Greenland
and Alaska south to Virginia, Pennsylvania, and Wisconsin.
Butler Co.: In boggy place in upland pasture 1 mi. n. of Smith School, Parker
Twp. Adam M. Barker. July 5, 1935. Fayette Co.: In hollows along rocky river-bed
above the falls, Ohio Pyle, July 4, 1908. O.E.J, (figured)
Order II. Andreaeales
Small, monoicous (or dioicous), dark brown to almost black, when dry
very brittle, mostly cespitose on granite or slate rocks : stems slender, radiculose
below, dichotomous, with fascicled branchlets, no central strand; leaves small,
crowded, erect-spreading to often falcate-secund, uni-stratose to partly bi-
stratose, thickish, often more or less papillose, costate to ecostate, very opaque;
cells small, incrassate: seta none, but represented by a pseudopodium from the
gametophore; capsule oval, opening by 4 (-8) vertical slits, the valves remain-
ing united both above and below; spores and columella derived from the en-
dothecium; no air-cavity between the spore-sac and the capsule-wall; calyptra
torn at the base, delicate; spores large, about .034 mm in diameter, chlorophyl-
lose.
Jennings: Manual of Mosses — II. Andreaeales 43
This peculiar order is represented by but one family, the Andreaeaceae,
which consists of only one genus, Andreaea [Ehrhart] Hedwig. There are
about 125 species, alpine and sub-alpine and widely distributed; 19 species
occur in North America, only 3 of which, however, are to be expected in our
general range.
I. Andreaea [Ehrhart] Hedwig
A. Leaves ecostate 1. A. rupestris
A. Leaves costate B
B. Leaf elongate acuminate from an oval base; costa filling about the middle two-
thirds of the leaf-apex, not excurrent 2. A. Rothti
B. Costa practically filling the whole apex of the leaf and excurrent
{A. Rothti var. crassinervia (Bruch) Moenkemeyer)
1. Andreaea rupestris Hedwig
(A. petTOphila Ehrhart)
Densely cespitose, dark brown to blackish: stems slender, about 1.5-2.5 cm
high, usually branching, more or less erect; leaves when dry very brittle,
crowded, small, ovate to lance-ovate, imbricated, often falcate-secund from an
erect base, usually c^tuse, entire, margin incurved; no costa; (the leaves are
so dense that they usually require bleaching in a solution of caustic potash
before the leaf -cells can be made out under the microscope); basal leaf -cells
narrow-rectangular, very incrassate, sinuose, above becoming shorter, the median
and upper cells rounded and angular-oblong, longitudinally seriate, dorsally
strongly papillose: fruit similar to that of Sphagtnim in being enclosed in the
perichaetium until mature, when it is quickly exserted on an outgrowth from
the tip of the leafy shoot similar in appearance to a short seta and termed
the pseudopodium; calyptra very thin and irregularly torn at base; capsule oval,
opening usually by four perpendicular slits along the sides but remaining
united at apex and base; columella persistent; spores smoothish, mature in late
spring.
In mountainous or hilly regions on non-calcareous rocks. In North Amer-
ica from the Arctic regions south to northern United States and south to
Georgia, Colorado, and California. Occurs in northeastern Pennsylvania and
in northern West Virginia.
2. Andreaea Rothii Weber and Mohr
(Andreaea rupestris Roth)
Plate LX
Small, blackish; stems 1-2 cm. long, branching to form tufts; leaves spread-
ing to falcate-secund, up to 1-1.5 mm long, suddenly elongate acuminate from
an oblong or ovate base, entire to faintly crenulate; costa strong, yellowish,
more or less plainly percurrent; alar cells rounded-quadrate, the median basal
ones variously either rounded or linear-oblong with irregular lumen, the upper
rounded, small, all densely incrassate; perichaetial leaves sheathing, suddenly
contracted into a short acuminate or linear point; capsule oblong-ovate, dark,
about .75 mm long, exserted on a twisted seta, splitting into four valves which
44 American Midland Naturalist Monograph No. 6
in old specimens finally break apart at the apex and coil over each other when
dry; spores .030-.040 mm, roughish, ripening in summer.
On non-calcareous rocks. Greenland to Michigan and Georgia; and in
Northwestern United States.
Fayette Co.: On rocks at White Rock, on Chestnut Ridge above Fairchance. Aug.
9, 1931. John L. Sheldon (figured).
Order III. Bryales — True Mosses
This order comprises numerous mosses of various habit: the endothecium
gives rise to the sporogenous tissue, which surrounds an inner sterile tissue,
loose in Arch.dmm, but forming the columella in the rest of the Bryales; the
spore-sac is separated from the wall of the capsule by a more or less highly
developed air-cavity; there is no pseudopodium but there is a more or less
elongated true seta; the outer wall of the archegonium after some growth is
ruptured, thus forming a basal vaginule and an apical calyptra; capsule cleisto-
carpous or, more usually, with a definite operculum and then often with a
single or double peristome: the order is conveniently divided, according to the
position of the sporogonium upon the leafy shoot of the gametophyte, into the
acrocarpous mosses (sporogonium at the apex of the leafy shoot) and pleuro-
carpous mosses (sporogonium lateral upon the leafy shoot) .
ACROCARPI
The acrocarpous mosses comprise about thirty families of the Bryales
widely distributed and numerous in number of species. For the analytical key
to the acrocarpous mosses see the general key to the genera of mosses at the
beginning of the book, page 10.
Family 1. Archidiaceae
Autoicous, sometimes paroicous or synoicous, rarely dioicous: small terres-
trial plants, closely gregarious and forming broad patches; stems erect, with
central strand, below bearing rhizoids; leaves of the shoots and also the basal
leaves minute, spreading, distant, linear-lanceolate, acuminate, flat, entire, the
costa ending in the point; perichastial leaves much larger, imbricated, more or
less linear-acuminate from a lanceolate base; leaf-cells smooth, prosenchymatous
or sometimes sub-vermicular to parenchymatous: capsule sessile, spherical,
terminal, non-operculate; columella none; spores commonly 16-20, about .2 mm
in diameter.
One genus only, the characters being as given for the family, comprising
about 25 species, distributed widely in the temperate zones. Six species are
native in North America, but only one is likely to be collected in our region.
1. Archidium Bridel
1. Archidium ohioense Schimper
Occurs on the ground in meadows and fields throughout eastern United
States from Quebec and Minnesota to Florida and Louisiana. Not yet re-
Jennings: Manual of Mosses — 2. Dicranaceae 45
p)orted in our region, but to be expected, as it occurs in eastern Pennsylvania
and in Ohio, and its type locality is Harper's Ferry.
Family 2. — DiCRANACEAE
Autoicous or dioicous; large to minute, mostly cespitose: stem with a
central strand, often thickly covered with rhizoids, mostly densely leafy,
branched; leaves often falcate-secund, mostly acuminate to narrowly linear
from a broader base, usually more or less smooth and shining, usually costate;
costa sometimes dorsally serrate, heterogenous; leaf-cells sometimes mammil-
late, the basal ones enlarged and mostly transparent, alar cells often much
larger and either hyaline or brownish, the central leaf-cells short to rounded,
mostly smooth; perichstial leaves often sheathing: seta usually erect and long;
capsule mostly unsymmetric, usually cernuous, when dry often curved and
sulcate; annulus present or absent; peristome simply or rarely none; when
present the peristome teeth are 16 in number, approximate, united below into
a basal membrane, usually two parted to the middle, or beyond, into linear
or awl-like divisions, no longitudinal lines, but the teeth minutely striate or
papillose on the dorsal face, rarely smooth, inner face yellow with one or two
longitudinal lines and with more or less projecting trabecube, operculum more
or less long-rostrate; calyptra usually cucullate.
Key to the Genera
A. Capsule with a long, slender, usually curved neck; leaves suddenly lanceolate or
subulate from a broad, clasping base 2. Trematodon
A. Capsule neck, if any, much shorter than the urn B
B. Cells of costa in cross-section homogeneous; peristome, if present, of 16 flat,
smooth, usually entire teeth 6. Seligeria
B. Cells of costa in cross-section heterogeneous; peristome none, or various C
C. Alar cells not conspicuously enlarged or inflated D
C. Alar cells conspicuously enlarged or inflated j
D. Leaf-cells smooth or essentially so in our species E
D. Leaf-cells more or less distinctly mamillose or papillose Q
E. Leaves not crisped when dry F
e. Leaves crisped when dry P
F. Costa more than one-half width of leaf-base Brothera
F. Costa less than half as wide as leaf-base G
G. Cleistocarpous H
G. With peristome I
H. Capsule ovoid, immersed, short-apiculate 3. Pleundium
H. Capsule pyriform with a short neck 1. Bruchia
r. Peristome-teeth unequally subulately 2-3 cleft to the middle or somewhat below,
papillose above 7. Dicranella
I. Peristome-teeth cleft to the base or nearly so into two, linear, filiform papillose divi-
sions N
J. Costa narrow, less than one-third as wide as leaf-base K
J. Costa at least one-third as wide as leaf-base L
K. Capsule mostly not strumose; peristome at base not forming a deeply inserted tube
10. Dicratium
K. Capsule strumose; jseristome teeth united at base into a deeply inserted tube
9. Oncophorus
L. No stereid bands in costa; costa filling most of the leaf above the middle M
46 American Midland Naturalist Monograph No. 6
L. Stereid bands above and below the guide-cells; peristome teeth separate to below
the mouth of the capsule 11. Dicranodontium
M. Alar cells not definitely inflated Brolhera
M. Inflated alar cells reaching the costa 10. Dicranum longifolium
N. Basal leaf-cells short-rectangular; the capsules sulcate 5. Ceratodon
N. Basal leaf-cells somewhat elongated-rectangular O
O. Capsules, when dry, sulcate 4. Ditrichum pallidum
O. Capsues not sulcate when dry 4. Ditrichum pnsillum and D. lineare
P. Peristome teeth divided one-half way down or more 9. Oncophorus Wahlenbcrgii
P. Peristome teeth not divided 8. Rhabdoweisia
Q. Cleistocarpous t 1. Bruchia
Q. With peristome {Oreoweisia) *
I. Bruchia Schwaegrichen
Autoicous or paroicous; gregarious: green protonema persistent but sparse;
stem short with a central strand; leaves long-canaHculate-subuIate, from an
oval to lanceolate base, erect to secund; costa broad and flat, filling the subulate
acumen; laminal cells rectangular; seta short; capsule pyriform, more or less
cernuous, rostrate; operculum none; calyptra covering one-third or more of the
capsule, mitrate, unsymmetrically cleft.
A widely distributed genus of about 25 species, 14 of these being found in
North America, one already found and another probably occurring in our
region.
Key to the Species
A. Collum less than one-half the whole length of capsule; seta usually longer than
capsule 1. B. ftcxuosa
A. Collum about as long as rest of capsule; seta usually shorter than capsule
2. B. Sullivantii
1. Bruchia flexuosa (Swartz) Mueller
Gregarious, the green protenema persistent but not very conspicuous: stems
about 2-4 mm long, curved to erect; leaves remote, small, lance-subulate, erect-
spreading from a concave base, somewhat serrulate at the apex; leaf-cells
rectangular, narrower at the margin, alar not much different; antheridia in
axils of comal leaves or in separate buds; seta short, stout, usually shorter than
the erect, ovoid-pyriform, partially exserted, apiculate capsule; calyptra nar-
rowly conic, mitrate; spores .030-. 040 mm in diameter, decidedly papillose,
mature about June.
This and the following species perhaps differ too little to be regarded as
distinct. On Clay soil m fields from Minnesota to New England and south
to the Gulf States. Occurs in eastern Pennsylvania and in Ohio.
* Oreoweisia serrulata extends south as far as Tennessee and Kentuclcy, but is reported
thus far in Pennsylvania only from the eastern part of the State. Moist, shaded, cool
ledges.
Jennings: Manual of Mosses — 2. Dicranaceae 47
Bruchia Sullivantii Austin
Plate LX
Very clojc to B. flexuosa, from which it differs mainly in having shorter
stems; the neck about as long as the spore-sac. Leaves suddenly canaliculate-
subulate from an ovate concave base, smooth or nearly so, basal cells thin-
walled, elongate rectangular, often very irregular, with an indistinct margin of
linear somewhat incrassate cells; cells at shoulder much shorter, polygonal,
thicker walled; costa strong percurrent: seta curved, 1-1.5 mm long; capsule
oblong-pyriform 1-2 mm long, acuminate, the neck about as long as the
spore-sac; calyptra about one-half the length of capsule, smooth; spores spinu-
lose, mature in June.
On clay soil in fields from New England to Minnesota, south to the Gulf
States. Occurs in eastern Pennsylvania and in Ohio.
Butler Co.: One mile north of Moniteau, Cherry Twp., on soil m old cornfield. Sid-
ney K. Eastwood. June 6, 1935 (figured).
2. Trematodon Richard
Autoicous, rarely dioicous; low, singly disposed: stem with a large central
strand and loose ground tissue; leaves yellowish-green, abruptly to gradually
lance-subulate from a broad clasping base, more or less crisped when dry; costa
ending below the apex or percurrent; cells thin-walled, loosely elongate-hexag-
onal to rectangular or, above, rhombic-pentagonal or -he.xagonal: seta yellow,
erect, rarely tortuous to cygneous; capsule with a long tapering neck, moder-
ately arcuate, the urn smooth, annulus differentiated; peristome-teeth united
below into a low basal tube, undivided and cribrose or two-parted to the base
into filiform divisions, articulate and longitudinally striate, peristome rarely
lacking; operculum as long as the urn, obliquely rostrate; calyptra inflated,
cucullate, not ciliate.
A cosmopolitan genus of about 70 species, of which about 10 occur in
North America, 2 of these in our region.
Key to the Species
A. Collum as long as urn of capsule 1. T. ambiguus
A. Collum twice as long as urn 2. T. longicollis
Trematodon ambiguus (Hedwig) Hornschuch
Plate LXXII
Gregarious, erect, simple or sparingly branched; stems 6-10 mm tall,
densely brownish radiculose below; leaves 4-8 mm long, from an ovate or ob-
long, concave, sheathing base abruptly narrowed to an equally long or some-
what longer linear-subulate, channelled apex, minutely serrulate at the tip;
costa at base thin, about one-fifth or one-sixth the width of the leaf, from
there percurrent and constituting most of the linear-subulate upf>er part of the
leaf; cells at base of leaf thin-walled, oblong-rectangular, somewhat inflated,
about .010-.02C mm wide by 2-5 times as long, at the margin a few rows much
narrower, the cells at the shoulder where the sheathing base suddenly tapers
48 American Midland Naturalist Monograph No. 6
into the narrow acumination much smaller, short-rectangular to rounded,
.003-.010 mm in diameter, thick-walled: seta bright yellow, slender, 1-2.5 cm
long; capsule about 2 mm long, often slightly curved, narrowly ovoid cylindric,
abruptly narrowed below into a slender, curved hypophysis, about as long or
slightly longer than the urn and strumose at the base with a goiter-like swell-
ing; operculum with a slender beak about two-thirds as long as the urn;
annulus large, revoluble peristome dark red, borne on a basal membrane
which projects above the mouth of the capsule, the teeth 16, awl-like, more or
less irregularly perforate or cleft or entirely divided, striate longitudinally;
capsule with a reddish rim of a few rows of small, incrassate, rounded cells,
below which the cells become linear and incrassate; spores rough-warty, ,023-
.024 mm in diameter, mature in June.
On clayey soil in open ground, Virginia to Newfoundland and Alaska.
Cresson, Cambria Co., James and Porter: and Westmoreland County, on recently dis-
turbed clay soil around sawmill, altitude 1300 ft., two miles north of Darlington, Chestnut
Ridge, May 29, 1949, O.E.J, (figured).
2. Trematodon longicollis Richard
Cespitose, light green to brownish-green; stems erect, usually about 5 mm
high; leaves abruptly linear-subulate from a concave ovate base, the subulation
canaliculate, minutely serrulate at apex; costa scarcely reaching the apex; leaf-
cells as in T. ambiguus; pcrichastial leaves quite gradually long-acuminate: seta
similar to T. ambiguus; collum twice as long as the urn; urn more strict'y
oblong-cylindric; peristome- teeth 16, narrow-subulate, nodosely articulate,
usually perforate rather than cleft. Otherwise very similar to T. ambiguus.
In old fields, etc., on sandy or clayey soil, in Europe, Asia, and in North
America, from Massachusetts, Pennsylvania, and Ohio southwards to Florida
and Mexico. Not yet reported in our region, but to be expected.
3. Pleuridium Bridel
Autoiccus or paroicous, rarely synoicous: weak, green or yellowish-green,
cespitose or gregarious: stem with a central strand, radiculose at base, perennial
by means of fertile shoots below the apex and by means of sterile flagella;
leaves mostly terminal, erect-spreading, sometimes secund, linear-subulate from
a broader base, upwards weakly denticulate, sometimes thickly imbricated;
costa varying from weak and ending below the apex to very broad and filling
the whole acumen, often rough-serrate dorsally; seta mostly very short and
erect, rather curved; capsule mostly immersed and oval to ovate-globose, short
pointed, sometimes obliquely so, cleistocarpous, without a collum; calyptra
cucullate, cleft almost to the apex on one side, covering scarcely half the
capsule.
About 30 species widely distributed, mainly in temperate regions, on soil.
Six species occur in North America, at least one in our region. The following
key is adapted from Grout's Moss Flora.
Jennings: Manual of Mosses — 2 Dicranaceae 49
Key to the Species
A. Calyptra mitrate; capsule with stomata immersed in middle of wall
(Sporledera palustrti (Br. &. Sch.) Hampe =^ P. palustre (Br. & Sch.) Bryol. Eur.)
A. Capsule split almost to apex B
b. Narrow upper part of leaf shorter than the broadened basal part C
B. Narrow part of leaf much longer than broad basal part D
C. Leaves gradually narrowed from lance-ovate base to serrulate apex
(P. Ravenellii Austin)
C. Leaves abruptly mucronate and smooth above from broad serrulate base
(P. Sulltvantii Austin)
D. Basal cells of leaf about 2-5:1 \. P. subiilatum
D. Basal cells of leaf about 5-8:1 (P. acuminatum Lindberg)
1. Pleuridium subulatum (Hedwig) Lindberg
(P. allernifoHum [Dickson; Kaulfuss] Rabenhorst;
Phascum subulatum Schreber)
Plate LX
Densely gregarious to cespitose, yellowish-green: stems usually simple,
about 2-6 mm high; lower stem-leaves lance-subulate, short; comal and peri-
chstial leaves m.uch longer, more or less erect or ascending, from a small oval
base gradually subulate-setaceous, canaliculate, nearly entirely to minutely
denticulate; costa wide, not very well defined, prartically filling the apex; basal
leaf-cells rectangular to more or less oblong-hexagonal, the upper cells often
becoming linear and forming a more or less distina margin to the costa: seta
shore, erect, about as long as the capsule; capsule oval or roundish, about 1 mm
long, obtusely apiculate, more or less castaneous or yellowish when mature;
calyptra cucullate, reaching about halfway down the capsule, short rostrate,
split almost to apex; spores large, mature from April to June, minutely rough-
ened; antheridia naked in axils of the upper or median leaves.
On moist clayey or sandy soil in old fields, along banks of ditches, etc., in
Europe, Asia and in North America, from New England to Wisconsin and
south to Alabama.
Butler Co.: In sandy meadow on south slope of hill two miles southwest of Glade
Mills. April 20. 1913. O.E.J, and A. R. Hillard. Washington Co.: On stony ground,
near Washington, Linn and Simonton, May 2, 1892, and May 6, 1893. Westmore-
land Co.: In sandy-clay meadow on gently sloping hillside east of Blackburn, April 24,
1913. O.E.J., G.K.J.. and R. J. Sim (figured).
4. DiTRlCHUM [Timm] Hampe
Dioicous or autoicous; mostly low, cespitose, not radiculose, green to yellow-
green, more or less shining: stem with a central strand, densely foliate, simple
or little branched; leaves with a broad base, not sheathing, mostly long-canalicu-
late-subulate, imbricated to erect- spreading or secund, when dry mostly a little
curved or straight; costa broad and flat, percurrent or excurrent, usually filling
the upper part of the acumen; leaf-cells rectangular, more or less prolonged,
alar cells not inflated: seta elongate, erect; capsule erect or a little cernuous,
symmetric or unsymmetric, sometimes weakly arcuate, mostly ovate to oblong-
50 American Midland Naturalist Monograph No. 6
cylindric, sometimes sulcata; peristome with a basal membrane, the 16 teeth
mostly cleft to the base or nearly so into two linear-filiform portions, papillose,
rarely weakly twisted to the left; articulations not projecting dorsally, some-
imes coupled at the base of the teeth; annulus mostly serrate, revoluble; oper-
culum mostly obliquely conic; calyptra reaching to below the middle of the
capsule.
A cosmopolitan genus of about 50 species, mostly growing on soil, about
10 species in North America, 3 of these occurring in our region.
Key to the Species
A. Dioicous: leaf-margins more or less recurved; seta castaneous B
A. Autoicous: awn serrulate; seta bright yellow 3. D. pallidum
B. Capsule somewhat unsymmetric, subsulcate, somewhat cernuous 1. D. lineare
B. Capsule symmetric, smooth, erect 2. D. pusilluin
1. DiTRICHUM LINEARE (Sw.) Lindberg
(D. tortile var. vaginans Grout; Trichostomum vaginans Sullivant;
Leptotrichum vaginans Schimjjer; Ditrichum vaginans Hampe )
Plate IX
Densely cespitose, yellowish-green, lustrous: stems erect, ascending, about
5 mm. high, usually with erect terete sterile branches, sometimes up to 1.5 cm
high; leaves 1-1.5 mm long, close, erect-appressed when dry, not much spread-
ing when moist, from an ovate concave base narrowed to a linear deeply
canaliculate acumination, margins narrowly recurved, usually entire; costa
strong, percurrent or rarely excurrent, comprising from one-third to one-half
of the width of the acumination; upper leaf-cells rectangular, mostly about 2:1,
rather dense and incrassate, smooth, the basal larger, elongate-rectangular up
to 6-8:1, moderately thin- walled, sub-hyaline or yellowish; perichastial leaves
larger, convolutely sheathing, above narrowing abruptly into a linear-subulate,
canaliculate, entire acumination, the basal cells larger and laxer than in the
stem- leaves: seta erect, flexuous, lustrous, yellowish to brownish, sinistrorse,
about 1-2 cm long; capsule brownish, about 1.5 mm long, narrowly to oblong-
cylindric, rounded at base, little changed when old; lid about one-fourth the
length of the urn, conic-rostellate obliquely, castaneous; annulus 2-3-seriate,
wide; peristome-teeth linear subulate, imperfect, forked to the base or often
united above, or irregularly cleft, deep castaneous, articulate; exothecial cells
yellowish-incrassate, irregularly oblong to rectangular, the 4 or 5 uppermost
rows much smaller, rounded and obscure; calyptra cucullate, covering about
one-half of the capsule; spores yellowish, smooth, about .01 5-. 018 mm, ma-
turing in late fall or winter.
Usually on sandy soil in hilly or mountainous districts. In Europe, and
in North America, from Maine to Missouri and North Carolina.
Not common in our region. Allegheny Co.: Powers Run, September 14, 1905 (fig-
ured). O.E.J, and G.E.K.; Wildwood Road, March 29, 1908. O.E.J, and G.K.J.;
Thornhill, December 29, 1908. O.E.J. Fayette Co.: Fort Necessity. H. N. Mozingo.
April 1, 1945. McKean Co.: West Branch, September 6, 1896 D.A.B. Westmore-
land Co.: One mi. n. of Darlington. C.M.B. Oct. 7, 1944.
Jennings: Manual of Mosses — 2. Dicranaceae 51
2. DiTRlCHUM PUSILLUM (Hedwig) E. G. Britton
{Leptotrichum pufillitm Hampe; Ditnchum tortile Brockmueller )
Plate X
Cespitose, yellowish-green, rather dull: stems short, about 5-10 mm high,
erect or erect-ascending from a radiculose base, mostly simple, reddish; leave^
about 2.5-3.5 mm long, closely appressed-erect to somewhat spreading, usually
somewhat secund or twisted, gradually lance-subulate and canaliculate from ai:
ovate-lanceolate concave base, margins more or less narrowly revolute, apex
usually denticulate; costa strong, less distinct at base, in the upper portion
constituting about one-third to one-half of the leaf-width, percurrent to slightly
excurrent; basal cells rectangular to linear-rectangular, alar not different, rather
thin-walled and hyaline, smooth, median smaller, mostly about 2:1, rectangular
to quadrate, smooth, the apical sometim.es bi-stratose; pericha:tial leaves more
or less sheathing, otherwise similar to the stem-leaves: seta reddish-brown,
shining, somewhat sinistrorse, erect, about 1 cm. long; capsule oblong to
oblong-cylindric, reddish to pale brown, smooth, non-sulcate, not constricted
below the mouth, abruptly narrowed to the seta at base, the urn about 1 mm
long; annulus uni-seriate; peristome single, rather low, reddish, the 16 teeth
cleft into linear-subulate, distinctly trabeculate, somewhat spirally twisted
divisions, at base united into a very low membrane; operculum conic-rostellate,
usually more or less oblique; calyptra cucullate, pale; spores rather thin-walled,
smooth, yellowish-pellucid, about .015-. 018 mm, mature in late fall or in
winter: dioicous.
On clayey soil in fields, along roadsides, etc., in Europe, Asia, northern
Africa, and in the eastern half of North America from Labrador to the Gulf
States.
On clay soil, roadside banks, etc., in Fayette, Greene, Allegheny, and southern Butler
counties in the extreme southwestern part of the state. Specimen figured: Keown Station,
O.E.J. November 14, 1909.
3. DiTRiCHUM PALLIDUM [Schreber] Hampe
Plate X
Loosely cespitose, bright green; stems about 5 mm high, more or less erect,
or with a creeping base; leaves erect-spreading, sometimes somewhat secund,
from a lance-ovate base, prolonged linear-subulate, concave, channeled towards
the apex; costa strong, long-excurrent, denticulate tov/ards the apex; basal leaf-
cells laxly oblong-hexagonal, thin-walled, hyaline, up to about .015-. 017 mm,
the median cells gradually much smaller, rectangular, forming but a narrow
margin to the costa; seta erect, yellow, slender, dextrorse and fliexuous when
dry, about 1-2 cm long; capsule ovate-oblong, yellowish-red, ascending to
horizontal, somewhat unsymmetric, usually somewhat strumose at base, about
2 mm. long, when dry and empty sub-arcuate and irregularly sulcate; peristome
single, the 16 teeth bifid deeply, united at base into a very low basal membrane,
the prongs cylindric, nodose-articulate, finely papillose, reddish, about 0.5 mm
long; annulus compound, deciduous, bordered by two or three rows of small,
rounded, reddish-pellucid cells; spores globose, papillose, about .017 mm, red-
52 American Midland Naturalist Monograph No. 6
dish-pellucid, mature in early summer; operculum conic-obtuse, about 0 6 mm
long; calyptra smooth, cucullate, slenderly straight-rostrate, about 2.5 mm
long, the beak reaching about 1.5 mm beyond the tip of the operculum; anther-
idial clusters gemmiform in axils of the perichaetial leaves.
On bare soil, usually in woods. Europe, Asia, northern Africa, and, in
North America, from Nova Scotia to Ontario south to the Gulf of Mexico
and westward to Kansas.
Common in our region, and now known from the following counties: Allegheny, Arm-
strong, Beaver Bedford, Butler, Centre, Crawford, Fayette, Lawrence. McKean, Washing-
ton, and Westmoreland. Specimen figured: Blackburn, Westmoreland County, O.E.J.
June 13, 1908.
5. Ceratodon Bridel
Dioicous, rarely autoicous; cespitose, green to brown or reddish-brown,
somewhat radiculose; stem 3-5-angled, with a central strand, thickly foliate,
often bushy-branched; leaves erect-spreading, appressed and more or less
twisted when dry, ovate to lance-linear, neither sheathing nor subulate-pointed,
margin revolute; costa strong, percurrent or long-excurrent, with median guides;
leaf-cells thick-walled, short-rectangular below, the upper quadrate to rounded,
smooth; perichaetial leaves distinctly sheathing; seta long and erect; capsule in-
clined to horizontal, elliptic-ovate to oblong, purplish to reddish-brown, shining,
when dry sulcate; annulus spirally deciduous, 2-4-seriate; peristome- teeth 16,
cleft nearly to the base into filiform divisions, united at the base into a tube,
the teeth closely articulated below, less closely above, papillose; operculum
conic, much shorter than the capsule; calyptra cucullate.
A cosmopolitan genus consisting of 27 * species; 4 species in North Amer-
ica, only one occurring in our region.
1. Ceratodon purpureus (Hedwig) Bridel
{Mnium purpureiim Linnaeus; Dicranum purpurascens Hedwig;
Dicranum purpureum Hedwig)
Plate X
Densely and often rather deeply brownish- or reddish-cespitose, mostly
green above and dark brown below: stems mostly branched, erect, about 1 cm
high, dying away below; leaves lanceolate to linear-lanceolate, carinate, the
margins revolute to near apex; costa strong, percurrent, at base about one-
sixth to one-fourth the width of the leaf; seta about 1.5 cm long, erect, dark-
castaneous, lustrous, twisted when dry; capsule oblong-linear, at first erect, later
inclined to horizontal and more or less curved, irregularly sulcate, strumose,
about 2.5 mm long, dark red-brown, lustrous, annulus distinct, revoluble; peri-
stome-teeth dark red below, basally confluent, papillose, weakly trabeculate to
a little above the middle, bordered, hyaline above; operculum conic-elongate,
about one-fourth the length of the urn, often somewhat curved, usually darker
* Brotherus (Pflanzenfamilien, 1924, 2nd. edit, p. 163) thinks these all can be
reduced to two species.
Jennings: Manual of Mosses — 2. Dicranaceae 53
brown than the urn; calyptra cucullatc; exothecial cells rather incrassate, irregu-
larly elongate-hexagonal or rectangular-oblong, two or three rows at the rim
much smaller and darker; spores smooth, rather thin-walled, yellowish-pellucid,
mature in May or June. Quite variable.
Cosmopolitan on burnt-over ground, roadsides, vacant lots, roofs, bare
clay soil, etc.
Very common in cur region. Now known from Allegheny, Armstrong, Beaver, Bed-
ford, Butler. Cambria, Centre. Clearfield, Clinton. Crawford, Ene, Fayette, Liwrence,
McKean, Somerset, Washington, and Westmoreland counties. Specimen figured: Sand-
plain, Presque Isle, Ene Co., May 8-9, 1906. O.E.J.
6. Seligeria Bryologia Europaea
Autoicous: minute, gregarious, or cespitose, rupestral: stem simple or
branched at base, rarely with long sterile shoots; leaves in 3 to 5 series, the
lower minute and distant, the upper abruptly larger and canaliculate-subulate
from a concave lanceolate base; costa often stronger above the base; alar cells
not differentiated: seta mostly erect, but little longer than involucral leaves;
capsule globose-pyriform, smooth; collum distinct; annulus none; peristome
deeply inserted, or rarely none; when present, teeth broadly lanceolate, usually
entire, truncate or acute; operculum obliquely rostrate; calyptra cucullate.
A widely distributed genus of about 20 species; 8 occurring in North
America; 2 in our region.
Key to the Species
A. Seta arcuate when moist; leaves with a long, acute, subulate acumination
1. S. recurvata
A. Seta erect when moist; leaves with a rather short, linear, sub-obtuse subulation
2. S. calcarea
1. Seligeria recurvata (Hedwig) Bryologia Europsa
(S. setccea [Wulfen] Lindberg)
Densely gregarious, dark green, very small: stems short, about 1 mm or
less, simple or forking; leaves up to 1.5 mm long, erect-spreading, flexuous,
with a long, canaliculate, acute subulation from a lance-ovate base, the margins
entire; costa long-excurrent, forming much of the subulation; basal leaf-cells
thin-walled, pellucid, irregular or rectangular, above becoming quadrate and
incrassate; perichajtial tubulose-sheathing at base, towards apex lance-subulate;
seta long, yellowish, arcuate when moist, but when old and dry often erect,
more or less flexuous; capsule oval to subglobose, erect, thin-walled, short-
necked, red-mouthed, turbinate when empty; exothecial cells lax; no annulus;
operculum about as long as urn (each about 0.4 mm), straight, subulate-
rostrate; peristome-teeth 16, lanceolate to linear, obtuse to acute, sometimes
irregularly bifid at apex, free, smooth, orange-pellucid, reflexed when dry.
On rocks in shade. Europe, Asia and, in North America, in Ontario,
Pennsylvania, New York, New Jersey and in the Rocky Mountains. Occurs
on limestone rocks in eastern Pennsylvania and may occur in similar habitats
in central Pennsylvania.
54 A.MERICAN Midland Naturalist Monograph No. 6
2. Seligeria calcarea [Dickson] Bryologia Europaea
Densely gregarious, dull, dark green: stems short, less than 1 mm, simple;
leaves short, less than 1 mm, the lower lanceolate, the upper from an oblong
concave base abruptly narrowed to a shorter, linear, obtuse or sub-obtuse,
entire subulation; costa rather flat, indistinct below, above obscure and filling
the whole apex; basal leaf-cells shortly rectangular, pellucid, thin-walled, above
becoming irregularly quadrate to rounded or hexagonal, obscure, chlorophyllose,
incrassate: seta straight, erect; capsule erect, oval-pyriform, turbinate when dry
and empty, brownish; peristome-teeth 16, broadly lanceolate, rather densely
articulate, flat, entire, smooth, orange-pellucid, reflexed when dry; lid subulate-
rostrate but considerably shorter than the urn; spores mature in spring or
early summer.
On calcareous or chalky rocks. Europe and, in North America, tn On-
tario, Lake Winnipeg, New York, and Ohio. Rare.
In our region reported but once. Huntington Co.: Warrior's Ridge. Porter.
(Porter's Catalogue).
7. Dicranella Schimper
Dioicous, rarely pseudautoicous : mainly small, gregarious, or cespitose,
terrestrial: stem erect, thickly foliate, sparsely radiculose; leaves somewhat
lustrous, from a sheathing base abruptly subulate and squarrose-spreading, or
from a non-sheathing base gradually linear to subulate and stiffly erect to
falcate-secund, mostly with plane edges; costa strong; m.ostly percurrent, often
filling the acumen; leaf-cells elongate-rectangular to linear: seta erect; capsule
cernuous, unsymmetric, short, short-necked, often strumate, or erect and
symmetric; peristome-teeth usually present, mostly unequally subulately 2-3-
cleft, papillose above, at the extreme base united to form more or less of a
basal membrane, exteriorly finely vertically striate; operculum conic-rostrate or
obliquely long-rostrate, sometimes as long or even longer than the urn.
A. large and cosmopolitan genus of about 60 species; about 30 species in
North America; at least 4 species in our region.
Key to the Species
A. Costa wide and flat and not well-defined at base; peristome weakly papillose; annu-
lus often differentiated B
A. Costa narrower and fairly well defined at base; jieristome distinctly striate-papillose;
annulus not differentiated D
B. Seta red (D. aispa [Ehrh.} Schimper)
B. Seta yellowish C
D. Capsules erect and symmetric 1. D. Fitzgeraldl
C, Capsules more or less cernuous 2. D. heteromalla
D. Leaves squarrose, with wide clasping base (D. Schreberi [Swartz] Schimper)
D. Leaves not as above E
E. Leaves squarrose, with wide clasping base (D. Schreberi [Swartz] Schimper)
E. Leaves not as above B
E. Capsule usually nodding 4. D. varia
E. Capsule erect 3. D. rufescens
Jennings: Manual op Mosses — 2. Dicranaceae 55
1. DiCRANELLA FiTZGERALDi Renauld and Cardot
(D. heteTomalla var. orthocarpa (Hedwig) Pans)
Plate X
Rather densely cespitose, yellowish-green: stem 5-10 mm long, mostly
simple, erect, leaves crowded, erect-spreading, sometimes subsecund, about
3-3.5 mm. long, up to 0.5 mm broad at base, from the lance-ovate base narrow-
ing above into a long, canaliculate-subulate, denticulate apex; costa at base
rather indistinct, about one fourth to one-third the width of the leaf, strong
above and constituting most of the acumination; basal leaf-cells elongate-
rectangular or sub-rectangular, reaching 8x55 microns, hyaline, a few in the
extreme alar portion often quadrate, median cells quadrate: seta erect, yellow-
ish-red, becoming quite dark brownish-red when old, about 7-8 mm long,
when dry sinistrorse in the lower half and dextrorse above; capsule erect, sym-
metric, about .6-. 9 x .25-. 3 mm, oblong, not constricted below mouth, smooth
or nearly so even when dry and empty, when ripe brown; peristome single, the
teeth very slightly united below, cleft about half-way into linear-subulate
prongs, sometimes sub-cribrose along the divisural, articulate, longitudinally
striolate-granulose, towards apex hyaline; spores minutely papillose, about
.014-. 017 mm, mature in late fall or winter; operculum low-conic with an
oblique rostrum.
On soil, soil-covered rocks, etc., in the eastern and southeastern parts of
the United States.
Allegheny Co.: Schenley Park, Pittsburgh, August 16, 1905, McKees Rocks,
August 27, 1905, and Fern Hollow, Pittsburgh, March 8, 1908 (figured). O.E.J.; Wild-
wood Road, March 29, 1908. O.E.J, and G.K.J. Armstrong'^Co.: On clay bank,
Logansport. C.M.B. Feb. 22, 1943. Butler Co.: Frazier's Mill Jefferson Twp., on
ground. S. K. Eastwood. Fayette Co.: Ohio Pyle "Peninsula." C.M.B. Oct. 13, 1935.
Indiana Co.: Clay roadside ditch, near Crete. O.E.J. &C G.K.J. Nov. 2, 1941.
2. DiCRANELLA HETEROMALLA [Dillenius] Schimper
{Dicranum heteTomallum Hedvvig)
Plate XI
Cespitose, bright yellowish to dark green; stem erect or ascending, 0.5-3.0
cm tall: leaves numerous, lance-subulate, concave, 2-3 mm long, denticulate
towards the apex, usually also denticulate dorsally towards the apex; costa
strong, one-fifth to one-third the width of the leaf at base, percurrent, bordered
towards the apex by a narrow margin of lamina; leaf-cells parenchymatous, at
leaf -base 2-5 times as long as wide, rectangular, brownish, narrower towards
the margin, the upper cells shorter and often obliquely quadrilateral; seta
1.5-2.5 cm long, greenish-yellow, dextrorse; capsule smooth, about 1.5 mm long,
oblong, castaneous to dark brown, more or less erect, usually slightly curved,
when dry bent and curved in at the upper part just below the rim on one side
in a very characteristic manner, furrowed; operculum hemispheric, with a linear
obliquely inclined beak about 1 mm long; peristome-teeth red, bifid to below
the middle or about to the middle, sometimes trifid, with somewhat projecting
trabecula?, articulate, minutely papillose-striate, hyaline and papillose at apex;
exothecial cells incrassate, irregularly elongate-rectangular to oblong-hexagonal.
56 American Midland Naturalist Monograph No. 6
the end-walls thinner than the lateral walls, two to four rows of cells at the
rim much smaller and rounded; spores .010-.015 mm, yellowish-incrassate,
mature in autumn.
Common, especially in hilly or mountainous districts, on rocks, clay banks,
soil-covered logs, etc. Europe, Asia, and, in North America, from Canada
to the Gulf States.
A very common moss on soil in the southwestern part of the State from Erie to
eastern Somerset and Greene counties (13 counties ) but as yet known only from Cam-
eron and Centre counties on the elevated plateaus and mountains. Specimen figured:
"Rachelwood," slope of Laurel Ridge, southeast of New Florence. Sept. 8-11, 1907,
O.E.J.
3. DiCRANELLA RUFESCENS [Dickson] Schimper
Plates XI and LX
Rather loosely cespitose, reddish- to yellowish-green: stems erect, in our
region generally very short, reddish, about 3 mm or less high, mostly simple;
leaves few, linear-lanceolate, sometimes reaching 1.5 mm long, gradually nar-
rowed, minutely denticulate towards apex, plane, erect-spreading or sometimes
sub-secund; costa narrow, about one-seventh to one fifth the width of the leaf-
base, percurrent; basal leaf-cells large, hyaline, smooth, thin-walled, quadrate-
rectangular to linear-rectangular, reaching 8-10 times as long as wide, median
cells shorter and smaller but similar, the percurrent costa margined by cells
similar to the median: seta erect, red, about 3-5 mm long, dextrorse when dry;
capsule globose-ovoid, erect, red, symmetrical, smooth to slightly wrinkled
when dry, the urn wide-mouthed and more or less turbinate; operculum
obliquely conic-rostrate, nearly as long as urn (0.5 mm) ; calyptra cucullate,
smooth, yellowish-red, narrowly conic, about 0.8 mm long; peristome pellucid
red-chestnut in color; teeth bifid about half-way, articulate, irregularly and
finely longitudinally striate-papillose; spores globose, smooth, orange-pellucid,
about .012-. 01 5 mm, mature in late summer or fall.
Usually on damp, bare soil, in Europe, Asia, and, in North America, from
Alaska to Nova Scotia and southwards to Virginia.
Rather common in our region. Now known from the following, counties: Allegheny,
Beaver, Butler, Cambria (Porter), Elk (Porter), Fayette, Huntingdon (Porter), Indiana,
McKean. Specimen figured: Clay soil on upper slope 'of Laurel Mt., above New Flor-
ence, Sept. 8-11, 1907. O.E.J. ; and for the perigonial shoots, clay bank of ditch, Elder's
Ridge, O.E.J. & G.K.J. Nov. 2, 1941.
4. DiCRANELLA VARIA [Hedwig] Schimper
Plate XI
Densely gregarious to cespitose, bright to yellowish-green; stems short,
usually about 5-7 mm high, ascending to erect, branching at base; leaves up to
2.5 mm long, linear-lanceolate, gradually narrowed to a long-linear acumina-
tion, spreading to recurved, not very secund, when dry somewhat flexuous,
margin narrowly revolute, entire, excepting sometimes at the very apex some-
what denticulate; costa wide and not well-delined, percurrent and comprising
a large portion of the acumen; basal leaf-cells rather thin-walled, rectangular or
with oblique end-walls, 2-6:1, gradually becoming smaller and narrov.'er above,
Jennings: Manual of Mosses — 2. Dicranaceae 57
the upper being about 2-4:1 and somewhat incrassate, all smooth and more or
less yellowish-pellucid: seta yeIlowish-brov»n to castaneous, ascending to erect,
about 5-8 mm long, sinistrorse; capsule ovate to oblong, more or less cemuous,
reddish to pale castaneous, curved, smooth, together with lid about 1-1.25 mm
long; lid about as long as urn, rostellate; peristome-teeth large, lance-subulate,
cleft to middle, strongly articulate, finely striate-papillose, rich castaneous
below, sub-hyaline above; spores yellowish, rather thick-walled, minutely rough-
ened, .020-. 024 mm, mature in late autumn or in winter.
On bare clay soil in fields, on ledges, etc. Widely distributed in the
Northern Hemisphere; in North America from Nova Scotia to Alaska and
south to Georgia and the subtropics.
Allegheny Co.: In niches on clifF, Powers Run, November 30, 1909 (figured).
O.E.J. Butler Co.: On moist shale in railroad cut, Wahlville. Sept. 22, 1935. Sidney
K. Eastwood. McKean Co.: Bennett Brook, October 23, 1897. D.A.B. Washington
Co.: On shale npar creek, west of Tayiortown. Nov. 5, 1892. A. Linn and J. S.
Simonton.
8. Rhabdoweisia Bryologia Europaea
Autoicous: low, densely cespitose: stem without central saand, in cioss-
section obtusely pentagonal, densely foliate, radiculose, branched; leaves v/hen
dry crisped, decurrent, linear to linear lanceolate, acute, plane-m.argined; costa
strong, disappearing below the apex; upper leaf-cells chlorophyllcse, quadrate
to rounded, smooth; basal cells rectangular and hyaline: S2ta straw-yellow,
erect; capsule erect, minute, symmetric, ovate to oblong, obtusely octagonal
with darker stride, 8-costate v/hen dry; annuli:s none; peristome rarely absent,
inserted on the rim; teeth arisina from lov/, broad, more or less united bases,
abruptly filiform or subulate, reddish-ye'Jow, trabeculr; prominent ventrally but
articulations scarcely projecting dorsally, surface of teeth non-papillose but
often obliquely minutely striate; operculum long-subu!ate, obliquely rostrate,
as long or longer than the urn; calyptra cucuUate, rostrate, reaching to the
middle of the capsule.
About 8 species of the Northern Hemisphere, inhabiting crevices of
silicious rocks; 5 species in North America; 2 species in cur region.
Key to the Species
A. Leaves entire or nearly so L R. der.tkulata
A. Leaves rather coarsely denticulate towards the apex 2. R. denticulata var. americana
1. Rhabdoweisia denticulata (Bridel) Bryologia Europcca
{Weisia fugax Am. Auth.)
Plate XI
Densely cespitose, usually dark green: stems short, in ours about 5 mm
high, radiculose at base; leaves lance-linear, recurved-spreading, numerous,
about 2-2.5 mm. long, usually somev/hat concave, acute to shortly acuminate,
margins plane, somewhat denticulate towards apex, leaves crisped when dry;
costa strong but not quite reaching apex; upper leaf-cells rounded-hevagonal,
about .010-. 014 mm, incrassate, chlorophyllose, papillose, rather yellow, ar-
58 American Midland Naturalist Monograph No. 6
ranged in rows, in the upper part of the leaf about 6 or 8 rows on either side
of the costa, the apical cells larger and more hyaline, the basal cells pellucid
and elongate-rectangular, about 2-8:1: seta erect, 2-3 mm high, yellowish;
capsule erect, symmetric, oval; the urn wide-mouthed, about 0.5-0.7 mm high,
brownish, when dry and empty 16-striate; operculum about as long as urn,
obliquely rostrate from a broad base; peristome-teeth rather abruptly subulate
from, a broad base, small, not very persistent, articulate, papillose; spores about
.018.020 mm, minutely roughened, yellowish-pellucid, maturing in mid-
summer; calyptra cucullate, covering about two-thirds of the urn; exothecal cells
yellowish, incrassate, irregularly oblong to rectangular, the upper two to four
rows much smaller and rounded.
In crevices in various kinds of rocks, in moist, shady cliffs, mountains
from Canada to North Carolina and Missouri.
Fayette Co.: On sandstone rocks near Bear Run falls, Oct. 30, 1935. Adam M.
Barker. Lawrence Co.: Slippery Rock Creek. Wm. James. (Porter's Catalogue). Mc-
Kean Co.: Sandstone rocks between Hawkins and Rutherford hollows, March 12, 1894;
and Toad Hollow, July 19, 1896, and August 1, 1897. D.A.B.
This moss occurs on the sandstone cliffs of Chestnut Ridge six miles south of the
Pennsylvania-West Virgmia state line. July 4, 1909. O.E.J. &i G.K.J, (figured).
2. Rhabdoweisia denticulata var. Americana Culman
Closely similar to the species but with the leaves narrower and narrowly
acute, entire or only slightly denticulate.
We have seen no sp>ecimens of this variety from our region although Grout
reports it as occurring with the species and refers most of the collections to it.
9. Oncophorus Bridel
Autoicous: rather large, cespitose in broad, soft, bright green or yellowish-
f^reen tufts, usually radiculose below: stems thickly foliate; leaves when dry
crisped, when moist ascending to squarrose, from a sheathing base more or
less abruptly long-acummate or subulate, concave, carinate; costa strong, per-
current or excurrent; cells in the sheathing base of the leaf long-rectangular,
translucent to hyaline, the alar sometimes somewhat differentiated, the laminal
cells small, mostly rounded-quadrate, at the margin bi-stratose; perichsetial
leaves sheathing to above the middle, abruptly subulate: seta long, erect; cap-
sule unsymmetric, strumose with a short collum, when empty more or less
weakly sulcate; annulus indistinct; peristome-teeth 16, deeply inserted, approxi-
mate, united below into a tube which is adherent to the wall of the capsule,
the teeth 2- (3) -divided to the middle, outwardly minutely papillose in longi-
tudinal lines, the inner surface with 1 (or 2) delicate longitudinal lines and
strongly projecting transverse plates; operculum at least half as long as the
capsule, obliquely rostrate; calyptra cucullate.
A genus of 5 species widely distributed on damp gravelly soil, on moist
non-calcareous rocks, or on decaying logs. Only one known species in our
range.
Jennings: Manual of Mosses — 2. Dicranaceae 59
1. Oncophorus Wahlenbergii Bridel
Plate XII
Densely cespitose, light or yellowish-green above, darker below: stem
ascending or erect, forking, up to 3 cm high, sparsely radiculose below; leaves
numerous, dense, much crisped when dry, abruptly flexuous-spreading when
moist, from a concave, widely obovate base abruptly contracting into a long,
carinate, linear-subulate, flexuous, rather acute portion which is low-serrate at
the apex both marginally and dorsally; costa strong, ending in the apex; leaf-
cells at base mostly pellucid and obliquely elongate-rectangular, about 3-10:1,
above at the shoulder and along the subulation quickly becoming much smaller,
incrassate, about .005-. 007 mm in diameter, smooth, sometimes faintly rounded
papillose: seta single, erect, flexuous, yellowish to brownish, when dry strongly
dextrorse, 1-1.5 cm long; capsule about 1.2 mm long, arcuate-cernuous, oblong-
cylindric, gibbous, distinctly sharply strumose, when old irregularly wrinkled;
peristome-teeth united at base into a rather deeply inserted tube, the teeth divid-
ed to the middle, lance-linear, castaneous-pellucid, very faintly dorsally articulate
below, strongly ventrally trabeculate in a double series separated by a more or less
zig-zag divisural line, at the base smooth, towards the middle minutely vertically
striate-papillose, at the apex sub-hyaline; annulus narrow with crenulate margin;
operculum obliquely rostrate; exothecial cells irregular, rather lax, with medium
walls, not much different towards the mouth; spores papillose, castaneous-
pellucid, about .028-. 030 mm, mature in spring.
On rocks, soil, or mostly on old logs, in cool and moist situations, usually
in the mountains in non-calcareous districts. Europe, Asia, and, in North
America, from Greenland to Alaska and south to Pennsylvania and Ohio;
and Wyoming.
Rare in our region. McKean Co.: Broadbow, D.A.B. (figured.)
10. DiCRANUM Hedwig
Autoicous or dioicous; mostly large and thickly tufted, often cushion-like:
stems mostly erect; leaves mainly falcate-secund, more or less subulate-acu-
minate from a concave, lanceolate base, and usually canaliculate to tubulose;
costa largely excurrent; alar leaf-cells mostly brownish and differentiated; inner
perichastial leaves elongate, involute-sheathing, the acumen often short or lack-
ing: seta erect, mostly twisted, sometimes 2 to 5 together in a pericha?tium;
capsule various from cylindric and erect to cernuous and arcuate or even rarely
strumose; operculum long-rostrate and by a differentiated annulus always
with a notched edge; peristome not inserted below the edge of the capsule;
teeth mostly 2-3-parted to the middle, vertically striate below, ventrally trabe-
culate; calyptra not ciliate at base.
A cosmopolitan genus as here treated of about 150 species, mostly on non-
calcareous sub-strata, in the tropics confined to the mountains and rather rare
in. the Southern Hemipshere. In North America about 40 species are known
and at least 6 species occur in our region.
60 American Midland Naturalist Monograph No. 6
Key to the Species
A. Capsule cernuous, unsymmetric B
A. Capsule erect, symmetric G
B. Leaf-cells (xsrose c
B. Leaf-cells very slightly or not at all porose (D. condensatum Hedw. )
C. Leaves transversely undulate; costa not reaching, or vanishing in apex D
C. Leaves not transversely undulate; costa percurrent to excurrent F
D. LJpper leaf-cells elongated 1. D. rugosum
D. LJpper leaf-cells iso-diametric E
E. Capsule solitary; costa and lamina dorsally smooth (D. Bergeri Bland.)
E. Capsules clustered; costa and lamina dorsally rough (D. Drummondii C. M.)
F. Capsules clustered; guides of costa in two rows; leaves up to 15 mm long
(D. majus Smith.)
F. Capsules solitary; guides of costa in one row; leaves up to 8 or 9 mm long
2. D. scoparium*
c. Costa v/ith median guides H
G. Costa without median guides, 2-4 stratose 6. D. longifolium
H. Entire lamina uni-stratose; costa percurrent I
H. LJpper lamina more or less bi-stratose; costa excurrent J
I. Upper leaf-cells short rectangular and mamillate dorsally 3. D. montanum
I. Uppwr leaf-cells less regular, not mamillate 4. D. flagellarc
J. Costa and margin entire, apex usually broken off 5a. D. fulvum var. viride
J. Costa and margin serrulate 5. D. fulvum
1. DiCRANUM RUGOSUM (Hoffm.) Bridel
(D. polysetum Swartz, Schwaegrichen; D. undulatum Ehrhart)
Plate LXI
Tall, up to 20 cm. or more, loosely cespitose: stems erect or decumbent,
densely radiculose below; leaves undulate, lustrous yellowish-green; 6-9 mm
long, lanceolate, the upper half spinosely serrate, the lower half with recurved
margin; costa strong, rather narrow, vanishing in the apex, with two serrate
dorsal lamellae above; alar cells distinct, brownish, not reaching costa, median
and upper leaf-cells elongate-elliptic to linear- fusiform, incrassate and porose:
seta long, reddish, usually 2-5 in a cluster; capsule arcuate-cernuous, rather
small, when dry and empty striate and brown; spores mature in late summer
or early fall.
On moist soil and on humus-covered rocks in m.oist and shady woods,
usually in hilly or mountainous regions. Europe, Asia, and, in North America,
in the northern United States and in Canada.
Apparently rare in our region. HuNTiNGTON Co.: Porter. (Porter's Catalogue).
Somerset Co.: 12 mi. s.w. of Somerset. C. M. Hepner, Dec. 1, 1933 (figured). Wash-
ington Co.: In sandy soils in woods, near Washington. A. Linn and J. S. Simonton.
* Dicranum Bonjeani DeNot., is regarded by Grout as a subsp)ecies of D. scoparium,
and separated as follows:
Leaf-apex slender, strongly serrate, strongly falcate-secund (except some varieties),
leaves not undulate 2. D. scoparium
Leaf-apex broad, often obtuse; leaves laxly spreading, rarely secund (except var.
alatum) , sometimes slightly undulate 2a. D. Bonjeani
Jennings: Manual of Mosses — 2. Dicranaceae 61
2. DicraNUM scoparium [Linnaeus] Hedwig
{Bryum scoparium Linnaeus)
Plate XII
Large, rather loosely tufted, glossy, yellowish-green, often brownish below:
stems growing upwards and dying away below, often 7 or 8 cm long, densely
felted-radiculose; leaves falcate-secund, often more or less tufted at the upper
end of the innovations, about 8-12 mm long, linear-subulate, not undulate,
concave, serrate towards apex, little changed when dry; costa strong, flat, one-
fourth to one-third the width of the leaf at base, above bearing four serrate
dorsal lamellae; leaf-cells at base enlarged, quadrate to rectangular, rather
thin-walled, orange-colored, the median elongate rectangular to somewhat linear,
incrassate, porose, the apical irregularly oblong, not porose: seta about 3 cm
long, erect-sinuose, yellowish to chestnut-brown, lighter below, usually sinis-
trorse, sometimes dextrose above; capsule 3.5-4 mm long, about 0.8 mm thick,
chestnut-brown, cylindric, arcuate, when dry furrowed and slightly constricted
below the mouth, tapering below into a short neck, exannulate; operculum
low-conic, subulate-rostrate the beak about 2.5 mm long; calyptra about 6-7
mm long, cucullate, conic-rostrate, peristome single; teeth pellucid, reddish-
brown, papillose above, below strongly articulate and vertically striate, divided
about one-half into 2 or 3 lance-subulate prongs, sometimes more or less crib-
rose; spores globose, slightly roughened, about .020-. 024 mm, mature in late
summer or early fall.
On soil, logs, rocks, etc., in woods. Europe, Asia, and, in North America,
throughout the cooler and temperate regions.
Quite common in our region. Known from Allegheny, Armstrong. Beaver, Bedford,
Butler, Cambria, Cameron, Centre, Clinton Erie, Fayette, Forest, Greene. Huntingdon,
McKean, Somerset, Venango, Warren, Washington, and Westmoreland counties. Speci-
men figured: Ohio Pyle, Fayette Co., Sept. 1-3, 1907. O.E.J. 6C G.K.J.
2a. Dicranum Bonjeani DeNot.
A specimen collected by C. M. Boardman on a talus slide one-half mile
north of Sulphur Springs, Bedford Co., Pa., approaches D. scoparium very
closely.
3. Dicranum montanum Hedwig
Plate XII
Densely cespitose, bluish to light yellowish-green, lustrous: stems erect,
short, up to 1 cm in our region, sparsely branching; leaves much crisped when
dry, in the same cushion some of the plants with equally-spreading leaves,
others with all secund leaves, from a wider base gradually narrowly linear-
lanceolate, up to 5 mm long, concave below and canaliculate above to near the
apex, on margin and back of costa serrulate above; costa rather strong, percur-
rent or almost excurrent, forming about one-fourth to one-fifth of the width
of the leaf at base; median leaf-cells shortly rectangular-quadrate to laterally
oblong, incrassate, yellov/ish, the upper somewhat smaller and rounded-quad-
rate, more or less distinctly papillose, the basal rectangular, thinner-walled, up
to 6:1, the alar not much larger but quadrate-inflated, all the basal cells often
62 American Midland Naturalist Monograph No. 6
more or less castaneous in color; perichaetial leaves similar to stem leaves; seta
single, erect, yellowish to brownish, about 1.5 cm high; capsule oblong-cyiindric,
slightly curved, yellowish to finally brownish, plicate when dry and empty, the
urn about 2.5 mm long; the lid conic, more or less obliquely rostrate, about
1.5 mm long, castaneous; annulus narrow; peristome-teeth cleft to below the
middle or nearly to the base into linear-subulate, deeply castaneous, articulate,
faintly trabeculate, striate-papillose divisions; exothecial cells yellowish-incras-
sate, irregularly oblong to rectangular, the upper 3-6 rows much smaller, more
deeply colored and incrassate, rounded-quadrate or hexagonal; spores smooth-
ish, yellowish, about .022-. 025 mm, not very thick-walled, maturing in early fall.
Mostly on rotten wood and on roots and trunks of trees, or on rocks.
Europe, Asia, and, in North America, from Newfoundland to Tennessee and
westward to the Rocky Mountains.
Now known from Bedford, Butler, Elk, Erie, Huntingdon, Indiana, Fayette, and Som-
erset counties. Figured from specimen from Ohio Pyle, on rotterx log, O.E.J, and G.K.J.
4. Dicranum flagellare Hedwig
Plate XII
Rather densely cespitose, bright green above, brownish below, tufts about
1 cm high : stem radiculose, often with flagellae in the axils of the upper leaves,
erect; leaves crisped and sub-secund when dry, falcate-secund when moist, from
an oblong base narrowed gradually into a subulate acumen, strongly involute to
near the apex, apex serrate; costa strong, about one-fourth to one-third the
width of the leaf-base, percurrent, serrate dorsally at the apex; alar leaf-cells
large, distinct, inflated-quadrate, rather thin-walled, colored, reaching nearly to
the costa, the leaf-cells above loosely elongate-rectangular, farther above be-
coming shorter, above the middle rounded-quadrate, incrassate; perichaetial
leaves shorter, abruptly subulate-acuminate from a sheathing base: seta erect,
sinistrorse when dry; reddish to yellowish-brown, about 2 cm. long; capsule
erect, cylindric, symmetric, reddish-brown, about 2.5 mm long, when dry striate
and often slightly curved; lid obliquely long-rostrate, lustrous, brown; peristome-
teeth trabeculate, articulate, confluent at base, cleft to two-thirds to three-
fourths, the lower two- thirds reddish and more or less vertically striate-papillose,
hyaline above; annulus delicate; exothecial cells elongate, strongly laterally
incrassate with thinner end-walls, several series at the rim much smaller and
rounded-quadrate; calyptra reaching to the middle of the capsule, fugacious;
spores globose, slightly roughened, yellow-incrassate, .018-.022 mm in diameter,
mature in summer.
On decayed logs and stumps and on bases of trees in moist woods. In
Europe, Asia, and, in North America, from Nova Scotia to British Columbia
and south to the Carolinas and Mexico.
■Rather common in our region. Known from Beaver, Bedford, Blair, Butler, Cambria,
Cumeron, Erie, Fayette, Forest. Huntingdon (Porter), McKean, Somerset, Washington,
and Westmoreland counties. Specimen figured: N^ellon Estate (Rachelwood) near New
Florence. Sept. 8-10, 1907. O.E.J.
Jennings: Manual of Mosses— 2. Dicranaceae 6^
5. Dicranum fulvum Hooker
(D. interriiptum Bryologia Europaea)
Plate XIII
Deeply bui rather loosely cespitose, fulvous to brownish-green: stem
ascending to erect, sparsely branching, radiculose at base; leaves numerous,
secund, somewhat crisped when dry, about 5-6 mm long, gradually narrowed,
from a concave lanceolate base to a linear-acuminate more or less concave to
canaliculate apex, the upper margin serrulate; costa strong, about one-third of
leat-width at base, usually somewhat excurrent, dorsally serrulate above, in
the long acumination occupying most of the leaf; median and upper leaf-cells
quadrate to shortly rectangular, strongly yellowish-incrassate, the lower rectan-
gular, not porose at base, becoming in the alar portion enlarged, inflated,
rectangular to quadrate, thin-walled, brownish, this alar area reaching usually
to the costa; perichaetial leaves linear-subulate from a broadly sheathing base:
seta single, rather stout, erect, flexuous, yellowish to dark with age, about 1-1.5
cm_ long, capsule erect, symmetric to slightly curved, the urn about 4 mm long,
cylindric, castaneous, sulcate when dry and empty; lid stoutly and more or less
obliquely rostrate and about 1.5 mm long; annulus rather narrow; exothecial
cells yellowish-incrassate, quadrate to rectangular or cblong-hexagonal, several
rows below the mouth much smaller and rounded-quadrate-hexagonal; spores
large, .024-. 030 mm, smoothish, rather thin-walled, mature in autumn.
Generally on non-calcareous rocks in moist woods among the hills or moun-
tains, often on the face of sandstone cliffs; Europe and North America from
Nova Scotia and North Carolina west to the Mississippi River.
Rather common in our region. Now known from Butler, Fayette, Greene, McKean,
Warren, and Westmoreland counties. Figured from specimen collected from rock in
woods, Ohio Pyle. Sept. 1-3, 1906. O.E.J. & G.K.J.
5a. Dicranum fulvum var. viride (Sull. 8c Lesq. Grout
(D. viride Lindberg)
Plate XIII
Densely cespitose, yellowish-green to dark green or sometimes almost
blackish: stems ascending, up to 2 cm high, simple or sparsely branching,
radiculose below; leaves 3-4 mm long, spreading or recurved, when dry crisped,
the apices usually found broken off, the leaves close, gradually linear-acuminate
from a lanceolate base, concave below, the acumination often concave or canalicu-
late, the margin entire or slightly denticulate at apex; costa strong, percurrent
or excurrent, at base comprising from one-fourth to one-third the width of the
leaf; median and upper leaf-cells more or less regularly quadrate, small, in-
crassate, towards base slightly larger, but not much longer than broad, non-
porose, suddenly becoming enlarged, thin-walled, brownish, and rectangular up
to 3:1, the alar inflated and sometimes extending to the costa: capsule oblong,
erect or slightly curved. Not seen in fruit in our region.
On decayed logs and on bases of trees in woods, rarely on rocks, in Europe,
Asia, and, in North America, from Newfoundland to the Rocky Mountains,
south to Pennsylvania and Ohio.
64 American Midland Naturalist Monograph No. 6
Known from the following counties: Allegheny, Butler, Cambria (Porter), Fayette,
McKean, Washington, and Westmoreland. Specimen figured: Base of Tilia amerkana.
Brush Creek Swamp, Crider's Corner's, Allegheny-Butler county line, April 26, 1908.
O.E.J.
6. DiCRANUM LONGIFOLIUM [Ehrhart] Hedwig
{Paraleucobryum longifolium (Hedwig) Loeske)
Plate XIII
Densely cespitose, pale green, glossy: stems more or less deeply castaneous,
ascending, geniculate at intervals, at least 3-5 cm long, sparingly brownish-
tomentose below; leaves lustrous, pale green, yellowish-green and hardly altered
when dry, falcate-secund, about 5-8 mm long, linear-subulate, from a short
lanceolate base about one fourth the length of the leaf, at the base reddish or
brownish, non-decurrent; costa wide, comprising about one-third the width of
the leaf-base, somewhat narrowed at insertion, the upper three-fourths of the
leaf consisting entirely of the linear-subulate, canaliculate, more or less spinose-
denticulate, excurrent costa; alar leaf-cells lax, rather thin-walled and hyaline,
sometimes brownish, rounded and extending to the costa, the laminal cells
immediately above with medium walls, obliquely oblong-angular, narrower
towards the margin and farther above becoming smaller and rhomboid-quad-
rate along the margin to elongate-rectangular near the costa: capsule cylindric,
erect, nearly straight, smooth, produced but rarely.
On tree-trunks and on non-calcareous rocks in hilly or mountainous re-
gions, in Europe, Asia, and, in North America, from Greenland and British
Columbia south to Colorado and North Carolina.
Rare in our region. McKean Co.: Bradford, 1896. Sterile. D.A.B. (figured).
Washington Co.: On root of beech tree near Washington. Dec, 1891. Linn Ac
Simonton.
11. DiCRANODONTlUM Bryologia Europaea
Dioicous: tall mosses mostly in dense tufts, the stems and often the basal
portion of the costa on the under side felted-radiculose: leaves weakly or not
at all auriculate, from the lanceolate base long-subulate, canaliculate-tubulose,
the acumen often plainly toothed on the margin and dorsal surface of the
costa by reason of the mammillate cells; costa broad and flat, long, excurrent,
and almost filling the acumen; alar cells reaching the costa, inflated, hyaline,
sometimes reddish, delicate, areolation above the alar cells widened towards
the costa and rectangular to long-hexagonal, at the margin usually united into
a more or less broad border; perichaetial leaves sheathing, abruptly long-subu-
late: seta arcuate, finally erect-flexuous; capsule symmetric, oblong-cylindric,
smooth; annulus not diff^erentiated; peristome inserted below the edge of the
capsule-mouth; teeth separate, two-parted deeply, or to the base, the divisions
filiform-subulate, below vertically and above obliquely striate-papillose; calyp>-
tra cucullate.
A cosmopolitan genus of 21 species; 5 species in North America; 3 species
occurring in our region.
Jennings: Manual of Mosses — 2. Dicranaceae 65
Key to the Species
A. Leaves smoothish to serrulate not over half-way down B.
A. Leaves serrulate to well below the middle (D. asperulum)
B. Peristome-teeth cleft to the base; leaves with somewhat widened aurirles
\. D. denudatum
B. Peristome-teeth not cleft to base; leaves not auriculate C
C. Leaves easily caducous; seta 1.5-2 cm; urn L5-2 mm long 2. D. virginkum
C. Leaves rather persistent; seta 5-8 mm; urn 1 mm long 3. D. Miltspaughu
\. Dicranodontium denudatum (Bridel) E. G. Britton
(Didymodon longirostris Starke)
Plate XIII
Densely and. softly cespitose, lustrous, pale green, when dried as in her-
barium-specimens often a lustrous yellowish-brown: stems erect or ascending,
up to 3 or 4 cm high, forking frequently, flexuous, radiculose below; leaves
rather numerous, often quickly deciduous, from a more or less sheathing oblong
concave base with more or less widened auricles gradually narrowed to a long,
flexuous-spreading or falcate-secund, linear-subulate or setaceous, tubulose
point, the margin entire to faintly denticulate towards the apex; costa strong,
one-fifth to one-third the width of the leaf at base, excurrent in the rough
subulation, in cross-section showing a median row of large hyaline cells bor-
dered on either side by minute incrassate cells; alar leaf-cells large, inflated,
hyaline to brownish, rectangular, above becoming incrassate and narrower, in
the oblong base the upper marginal cells elongate-linear and more or less
prosenchymatous, the median and upper rounded-quadrate, varying to short-
rectangular or oblong: seta cygneous, dextrorse; capsule oblongcylindric, small;
peristome-teeth cleft to the base or nearly so into two filiform divisions, in-
serted below the mouth of the urn, reddish; lid as long as the urn, subulate-
rostrate, i>traight; spores mature in late fall or in winter: dioicous.
On sandstone rocks, walls, turfy places, etc., usually in hilly or mountain-
ous regions. Europe, Asia, and, in North America, from New Brunswick and
Alaska south to Ohio and Pennsylvania.
Rare and usually sterile in our region, often on the vertical face of heavy conglomerate
boulders. Fayette Co.: Mouth of Cucumber Run, Ohio Pyle. C.M.B. June 22 and
Oct. 20. 1940. Forest Co.: Vertical face of s.s. block. Blue Jay Creek, 1 mi. n. of
Frost. C.M.B. May 28, 1946. McKean Co.: Rutherford Rocks, July 7. 1894. Haw-
kins, October 18, 1895, Langmade Rocks, April 16, 1896, all in the vicinity of Bradford.
D.A.B. (figured). Somerset Co.: Face of dry rock, Beck Spring, Laurel* Mt. C.M.B.
July 26, 1947.
2. Dicranodontium virginicum E. G. Britton
Lustrous, bright green: stems ascending to erect, below red-tomentose;
leaves erect-spreading to secund, variously straight to curled or twisted, often
5 mm. long, narrowly concave-subulate from a short, thick, non-auriculate
base, often caducous, the caducous leaves usually with smooth points, the per-
sistent ones with serrulate points; alar cells more or less hyaline, the median
and upper rectangular to quadrate, incrassate; seta appearing lateral by growth
of innovations, flexuous, up to 2 cm long, lustrous, yellow, arcuate to erect;
66 American Midland Naturalist Monograph No. 6
capsule cylindric, 1.5 to 2 mm long; peristome-teeth deep red, not deeply in-
serted, split about to the middle, papillose-striate at base, sub-hyaline above;
no annulus; lid subulate-rostrate, shorter than the urn, straight or curved;
calyptra cucullate, rostrate, covering only the upper third of urn; spores small,
mature in summer: dioicous, antheridia terminal.
At the southern border of our cegion, on sandstone boulder along wooded path, Tibbs
Run, Monongalia County, West Virginia. C. F. Millspaugh. Both Williams and Grout
regard this as the same as D. denudatum.
3. Dicranodontium Millspaughii E. G. Britton
Silky, cespitose, yellowish-green; stems rufous-tomentose at base, up to 3
cm long; leaves erect-spreading to secund, up to 5 mm long, from a broad,
concave, non-auriculate base narrowly tubulose-subulate; costa strong, excurrent
into a linear tip, dentate marginally and dorsally; alar leaf-cells large, hyaline,
mainly quadrate to shortly rectangular, extending to the costa, above quickly
smaller, incrassate, tending to fusiform-prosenchymatous towards the margin,
shorter to quadrate in the upper part of the lamina: seta cygneous, erect when
old, 5-8 mm long, stout; capsule pyriform-cylindric, smooth, the urn about 1
mm, long; peristome-teeth deeply inserted, red, confluent at base, split to the
middle or perforate to the base, papillose-striolate below, paler above; no
annulus but the rim of the urn dark colored; lid about as long as the urn
(1 mm), straight, subulate-rostrate; spores maturing in summer; dioicous.
At the southern border of our region on sandstone rock in deep woods along Tibbs
Run, Monongalia County, West Virginia. C. F. Millspaugh. Both Williams and Grout
regard this the same as D. denudatum.
Family 3. Leucobryaceae
Dioicous, rarely autoicous; densely cespitose and mere or less spongy like
Sphagnum, whitish to glaucous-green: stem without central strand, scarcely
radiculose; leaves pluriseriate, close, quite uniform in size; costa very broad,
constituting most of the leaf, sometimes narrow with a stereid-bundle, com-
posed of two kinds of cells, the outer large and parenchymatous with per-
forated inner walls, the inner smaller and chlorophyllose, the lamina hyaline,
usually very narrow and mainly basal: seta single, erect; capsule erect and
symmetric or inclined, unsymmetric and strumose; annulus none; peristome
usually inserted below the edge of the urn, the teeth m.ostly 16, sometimes only
8, lanceolate, articulate, entire or cleft to the middle; operculum conic, ros-
trate; calyptra cucullate or sometimes mitrate.
With the exception of the genus Leucobryiim the species of this family are
mostly tropical or sub-tropical in their distribution and occur mainly on
trees. In our region there occurs only the following genus:
1. Leucobryum Hampe
Dioicous: thickly to loosely cespitose; whitish or glaucous green, mostly
lustrous: leaves erect, when dry appressed and brittle, sometimes spiral, or
falcate, or squarrose-spreading, from an ovate base lanceolate- to subulate-
acuminate, canaliculate or sometimes almost tubulose above; costa flat, the
Jennings: Manual of Mosses — 3. Leucobryaceae 67
large parenchymatous outer cells 2-6-layered; lamina mostly narrow, often
vanishing below the apex, without a border; perichaetial leaves half-sheathing
and long-acuminate: seta terminal, or lateral by the growth of innovations,
long; capsule more or less arcuate, unsymmetric, often strumose, with 8 rib-
like projecting ridges; peristom.e on the edge of the urn, the teeth united at
base into a tube, cleft to the middle into two lance-subulate prongs, thickly
trabeculate, vertically striate and papillose; operculum subulate from a conical
base; calyptra inflated, cucullate, covering the urn.
About 100 species, mostly in the tropics, on trees, rocks, or on shaded
earth; 8 in North America; 2 species in our range.
Key to the Species
A. Leucocyst on the median line in 3 to 4 layers; leaves 3 to 9 mm long: capsules arcu-
ate, strumose 1. L. glaucum
A. Leucocysts on the median line m 2 layers in 4 to 14 series; leaves 1 to 4 mm long:
capsule almost erect, not strumose 2. L. albidiim
1. Leucobryum glaucum [Linnaeus] W. P. Schimper
{Dicranum glaucum Hedwig)
Pincushion Moss — White Moss
Plate XIV
In dense, rounded, spongy, whitish or glaucous tufts, often 6 or 7 cm deep,
only the upper 5 mm or thereabouts alive, the dead inner portion grayish-
brown and peaty: leaves crowded, in our region about 3-6 mm long, more or
less tubular som.ewhat more than halfway dov/n, acute, entire, ovate-lanceolate,
narrowed at base, erect-appressed, consisting almost wholly of the broad, thick
costa, the lamina extending about half-way up the leaf as a narrow margin of
2-5 rows of hyaline, thin-walled, long-rectangular to linear cells: seta about 10
mm, long, sinistrorse, castaneous, erect; capsule 1.5 to 2 mm long, castaneous,
when dry arcuate, oblong-cylindric, distinctly strumose, furrowed; lid long-
rostrate, nearly as long as the urn; calyptra longer than the capsule; peristome
slightly inserted, deep reddish-brown, dicranoid; spores rather thin-walled,
slightly roughened, .015-020 mm in diameter, slightly roughened, mature in
autumn. Capsules are produced infrequently.
Almost cosmopolitan on soil or on rocks in woods. In North America
it occurs from Newfoundland to Florida and westward to the Mississippi
River. Common in our region, especially preferring the somewhat acid soil of
exposed white oak wooded upper slopes, often thus associated with Kalmia
and some of the wild huckleberries.
Now known from 19 counties in western Pennsylvania and probably occurs in all.
Specimen figured; Barrens, Scotia, Centre County. Sept. 23, 1909. O.E.J.
2. Leucobryum albidum [Bridel] Lindberg
(/,. minus Hampe; Dicranum albidum Bride!)
Much smaller than L. glaucum: tufts very dense, about 1-3 cm deep; leaves
acute, narrower, shorter (about 1-4 mm long), closely imbricated and but little
68 American Midland Naturalist Monograph No. 6
spreading at the tip, the tubulose apex usually shorter than the broad base:
capsule almost symmetric, little or not at all inclined, slightly or not at all
strumose.
On stumps, logs, or on the ground, Europe and in the eastern part of
the United States.
Rare in our region. Huntingdon Co.: Porter. (Porter's Catalogue). Westmore-
land Co.: A sterile specimen! from near Bear Cave, Chestnut Ridge, Hillside. Septem-
ber 17, 1909. O.E.J, and G.K.J.
Family 4. Fissidentaceae
Autoicous or dioicous: minute to large, gregarious to cespitose, mostly
green: stem oval, mostly with central strand, basally radiculose, or with reddish
rhizoids from the leaf-axils; leaves distichous, mostly vertically placed, so that
they stand edgewise to the stem with a clasping sheath at the base, or extending
well up the leaf, and a dorsal lamina which is often somewhat decurrent, the
apical lamina being lacking in the perichaetial and lowest stem leaves and
little developed in Bryoxiphium; costa usually present; leaf-cells small, uniform,
rounded hexagonal, chlorophyllose: seta erect or cygneous, usually elongated;
capsule erect and symmetric, or cernuous and unsymmetric or curved, smooth,
collum present; annulus present or none; peristome present, except in Bryoxi-
phium, usually inserted, simple, red; teeth articulate, united at base, cleft to
the middle or below into two or three filiform divisions, trabeculate with two
serie:. of projecting transverse plates, yellowish; spores mostly small; operculum
more or less rostrate; calyptra small, narrowly conical, entire or cleft on one
side, rarely several times cleft, mostly smooth.
A family of over 700 species, largely tropical, with widely varied habitats,
represented in our range by three genera.
Key to the Genera
A. Dorsal lamina very narrow: peristome none: stem radiculose-bulbiform at base
1. Bryoxiphium
A. Dorsal lamina usually broad: peristome present: stem not radiculose-bulbiform at
base B
B. Mostly not aquatic; sometimes submerged, but floating 2. Fissidens
B. Aquatic, filiform, floating mosses 3. Octodiceras
1. Bryoxiphium Mitten
(Eustichia Bridel)
Slender, dioicous, more or less densely silky-cespitose, bright green or yel-
lowish: stem stiff, oval in cross-section, with central strand, radiculose at the
extreme base, upwardly flattened, with distichous, closely imbricated leaves,
simple or irregularly branched; leaves from a linear-lanceolate base, either
linear, with a small acumen, or rounded and abruptly more or less long-
subulate, denticulate above; costa percurrent, with a very narrow dorsal wing
which does not extend to the base of the leaf; basal leaf-cells hyaline, rectangu-
lar, upper cells chlorophyllose, triangular to irregularly trapezoidal, smooth,
towards the margin linear and forming a distinct border; perichaetium terminal,
with two concave, ovate, prolonged-acuminate, serrulate leaves with a complete
Jennings: Manual of Mosses — 4. Fissidentaceae 69
dorsal wing: seta shorter than the perichaetial leaves, flexuous or cygneous;
capsule spherical, oval or obovate, smooth; no peristome or annulus; spores
.015-.02C mm; operculum abruptly and irregularly rostrate; calyptra smooth,
covering, about one-third of the urn; antheridial plants similar in appearance
to the archegonial.
Three species; one in Mexico, one in Asia, and one in Europe, Korea, and
the United States, rare.
1. Bryoxiphium norvegicum [Bridel] Mitten
(Eustichia norvegica Mueller)
Plants 1-2.5 cm long, somewhat flexuous, flat, lustrous, yellow, fastened to
vertical sandstone cliffs by a rac'iculose bulbiform base; stems mostly simple;
leaves short-acuminate and as described for the genus; costa vanishmg at or
near the apex: seta rather thick, about 2 mm long; capsule obovate, pale yellow,
mouth reddish, peristome none; operculum reddish at base, attached to colum-
ella and long-persistent; calyptra cucullate, large, tipped with a slender beak.
On shaded vertical exposures of sandstone in Greenland, and in a few
widely separated localities from Minnesota to western Pennsylvania and
Tennessee.
Lawrence Co.: "Slippery Rock Creek, Lesquereux." (Porter's Catalogue). The
writer has not been able to find this species along Slippery Rock Creek, where Lesquereux
found it.
2. FissiDENS Hedwig
Autoicous or dioicous: stem short to long, erect to procumbent, more or
less branched or simple; leaves prominently winged, linear-obovate to lanceolate-
obovate; costa usually present; cells rounded-hexagonal, sometimes loosely
rhomboidal, rarely prosenchymatous, smooth or papillose: seta erect or ascend-
ing, lon-^ to short, mostly terminal, sometimes lateral; capsule mostly exserted,
erect or inclined, symmetric or unsymmetric; peristome mostly inserted below
the mouth of the urn, teeth cleft, exteriorly articulate, often striate-papillose;
spores m.ostly small; operculum conic to rostrate; calyptra entire to once or
rarely several times cleft, mostly smooth.
A. widely distributed renus of about 700 species, mainly tropical, on soil,
rocks, trees, humus, or in water. In our region at least 8 species.
Key to the Species
A.Costa none: minute plants 2-4 mm high L F. hyalinus
A. Costa well developed B
B. Leaves bordered, at least on the vaginant lamina of the perichaetial leaves, by a
band of linear cells C
B. Leaves not bordered, or at least the border not composed of linear cells G
C.Costa percurrent, confluent with border at apex and forming a mucro: capsule erect
2. F. hryoides
c. Costa not usually percurrent, or only so in upper leaves; border not usually reaching
apex: capsule curved or erect D
D. Leaves non-bordered, entire, obtuse 3. F. obtusifolius
D. Leaves bordered, at least on sheath of the perichaetial leaves, acute or apiculate E
E. Leaves bordered to near the apex F
E. Leaves usually bordered only on the sheath of the perichaetial leaves 5. F. exiguus
70 American Midland Naturalist Monograph No. 6
F Leaves broadly oblong-lanceolate: capsules usually more or less curved: plants
usually more than 2 mm 2. F. bryoides var. incurvus
F. Leaves narrowly oblong-lanceolate: capsules usually erect: plants often less than 2
mm high 4. F. minutulus
G. Leaves without ar marginal band of several rows of somewhat paler cells, the outer
row sometimes paler ^ I
G. Leaves with a marginal band of several rows of paler incrassate cells H
H. Upper median leaf-cells rather obscure, about .006-.010 x .006-.014 mm; border
distinct 6. F. cristatus
H. Upper median leaf-cells distinct, about .012-.016 x .015-.025 mm; border not
very distinct 7. F. adiantoides
I. Costa excurrent into the apiculus 8. F. taxifolius
I. Costa not quite reaching apex J
J. Leaves often apiculate: seta terminal: leaf-cells .010-.016 x .014-. 020 mm
9. F. osmundioides
J. Leaves more or less rounded at apex: seta lateral in basal half of stem: leaf-cells
about .007-.011 mm 10. F. subbasilaris
1. FiSSlDENS HYALINUS Hooker and Wilson
Plate LXI
Gregarious, pale green, minute, 2-4 mm high: stem usually simple, erect;
leaves in 3-5 pairs, soft, the upper much larger, lance-oblong, acute, non-costate,
margined by a single row of narrow elongate cells, the sheath hardly reaching
the middle of the leaf, margin entire; cells large, about .030-.045 x .060-. ICO
mm, thin-walled, irregularly elongate-hexagonal, hyaline: seta terminal, 1-2 mm
long, erect smooth; capsule oblong, erect, thin-walled; teeth closely articulate,
red, cleft to the middle; operculum rostrate; calyptra cylindric-conic and cov-
ering the rostrum only of the operculum; spores .014-.020 mm.
The original station of this rare moss was "moist, rocky ledges, Bank Lick,
on Cassidy's farm, near Cincinnati, Ohio," where it was first collected by T. G.
Lea, in 1839. This station has since been lost, but the moss has been found
elsewhere in Ohio: on ground in deep ravines near Plainesville, — H. C.
Beardslee, and later in Pennsylvania, as follows:
Washington Co.: On clay banks with Fissidens taxifolius on banks in ravines near
Washington, September and October, 1892, 1894, and 1898. Linn and Simonton. Bank
in shaded ravine near Monongahela Lock No. 4, Linn & Simonton. Sept. 6, 1895
(figured).
2. Fissidens bryoides [Linnaeus] Hedwig
(Hypnum bryoides Linnaeus)
Plate XIV
In loose tufts or densely gregarious, rather dark green: stems ascending or
erect, 5-15 mm high; leaves numerous, ascending, or the apical erect, oblong-
lingulate, usually abruptly and somewhat obliquely acuminate, the sheath
reaching about half way to the apex, the dorsal lamina gradually becoming
very narrow at base, the border strong and reaching the apex, where it becomes
confluent with the costa, margin entire or sometimes faintly denticulate at
apex; costa strong; leaf-cells rounded-hexagonal, somewhat incrassate, some-
what smaller at the apex of the sheath, becoming rectangular at the base, the
border consisting of two or three rows of linear-prosenchymatous incrassate
Jennings: Manual of Mosses — 4. Fissidentaceae 71
cells, the border of the vaginant lamina edged be!ow with a row of rectangular
cells about two to tnree times as long as wide: seta erect, flexuous, about 4-9
mm long, yellowish to reddish, smooth, slender, terminal: capsule typically
erect and symmetric, sometimes inclined, usually reddish-yellow, smooth,
oblong-oval, about 7-8 mm long; peristome-teeth red, the upper two-thirds split
into two awl-like prongs with spiral thickenings, pellucid, papillose, the teeth
inserted below the mouth; spores smooth, small, about .010-.012 mm in
diameter; operculum conic-rostrate. Mature in late fall or winter. Antheridial
flowers gemmiform, axillary.
Widely distributed in temperate regions on shaded soil, in our rejion
especially in and about greenhouses. Our specimens show considerable varia-
tion in the arrangement of the leaf-cells, either in rows or not so, and in the
capsule, the latter varying from erect and symmetric to arcuate. The spores in
our specimens are much smaller than is indicated in some descriptions. Most
of the specimens from our region have more or less unsymmetric or arcuate
capsules and belong to the following variety.
Allegheny Co.: In flower-f>ots, Phipps Conservatory, Schenley Park, Pittsburgh,
March 20, 1910. O.E.J, (figured). Elk Co.: James. (Porter's Catalogue).
2a. Fissidens bryoides var. incurvus (Weber & Mohr) Hiibener
(F. incurvus Schwaegrichen )
Typically this moss is about 2-6 mm high, with rather broadly oblong-
lanceolate leaves, which are obtuse-apiculate and narrowly bordered up to near
the apex: seta reddish, long, flexuous; capsule oval-cylindric, curved and
usually more or less inclined or cemuous; antheridial buds basal.
On rocks, or more rarely clay, usually in shaded brooks and ravines,
America from Greenland to Vancouver Island to Texas. Europe, Asia,
Africa, New Zealand.
Fayette Co.: On muddy rock in bed of mountain rivulet, Ohio Pyle, June 14, 1908.
O.E.J, and G.K.J. McKean Co.: Hunt's Run, April 28, 1893. D.A.B.
3. Fissidens obtusifolius Wilson
Plate XIV
Small, densely gregarious, sometimes forming cushions, usually growing
at right angles to the substratum, pale green: stems comparatively stout, in our
specimens about 3-6 mm long, mostly simple; leaves of fertile plants about 4-8
pairs, of sterile shoots about 6-12 pairs, distichous, vertical, in fertile shoots
closely placed, the lower small, obovate to oblong, the upper much larger,
oblong, ascending to erect, obtuse, the clasping portion extending above the
middle, non-margined except for a few elongate cells at the end of the sheath-
ing portion, entire, the apical leaves reaching to 1.5 mm long by 0.3 mm v/ide;
cells rounded to quadrate-hexagonal above, a few at the margin of the base
rectangular (up to 4:1), at the apex of the sheath a few marginal cells elongate
to linear, all incrassate; costa strong, disappearing shortly below the apex, the
dorsal lamina becoming narrow or disappearing at the base: seta comparatively
stout, erect, or upcurving, in ours about 1.5-2.0 mm long, brownish, smooth;
72 American Midland Naturalist Monograph No. 6
the capsule erect, oblong-oval to oblong- obovate, somewhat narrowed below
the mouth, smooth, brownish; operculum hemispheric-apiculate to very shortly
rostrate; peristome yellowish-pellucid, trabeculate, the teeth lanceolate, acumi-
nate; capsule walls with cells incrassate, quadrate to hexagonal; spores smooth,
.018-. 023 mm. Mature in autumn.
On wet rocks from New England to Minnesota, Colorado, Texas, and
Alabama.
In West Virginia on walls of Lock No. 9, Monongahela River, a short dis-
tance south of the West Virginia-Pennsylvania State Line, July 3, 1909. O.E.J.
Rare in our region. Beaver Co.: Gorge of Little Beaver Creek, on sides of large
sandstone rocks in dashing current and often inundated, Smith's Ferry, October 1, 1910.
O.E.J. (On the Ohio-Pennsylvania State Line.)
4. FissiDENS MiNUTULUS Sullivant
(f. incurvus var. minutuliis (Sullivant) Austin)
Plate XIV
Plants minute, 0.8-3.0 mm high, gregarious, green, erect: stem simple,
reddish; leaves 3-7 pairs, hardly imbricate, the uppermost much larger and
incurved-erect and up to 2.5 mm long, narrowly oblong-lanceolate, more or
less acute, the border narrow, ceasing below apex, widest at upper part of
sheath, margin entire or somewhat undulate, the sheath about one-half the
length of the leaf, the inferior lamina narrowing at base but hardly decurrent;
costa strong, ending usually a little below apex; leaf-cells incrassate, more or
less rounded to hexagonal, rather irregular, the basal becoming rectangular, the
border consisting of 1-3 rows of elongate-linear or ascending prosenchymatous
cells: seta reddish, smooth, erect, subflexuous, about 3-6 mm long; capsule
usually erect, subflexuous, about 3-6 mm long; capsule usually erect, symmetric,
0.7-0.9 mm long, yellowish to dark chestnut color, oval-oblong, tapering
abruptly at base; peristome rich red-chestnut, the teeth deeply forked into two
av.'I-like prongs with prominent spiral thickenings, teeth slightly inserted; spores
round to oblong, pellucid, pale yellow-red, smoothish, .014-. 017 mm in diam-
eter; operculum conic-rostrate. Mature in early autumn.
On damp stones and rocks in shady woods or in stream beds, Europe, and
from eastern Canada south to the Gulf States.
Now known from the following counties: Allegheny. Armstrong, Butler, Lawrence,
McKean, Warren, Washington, and Westmoreland. Figured from specimens from Mellon
Estate (Rachelwood), New Florence, Westmoreland Co., O.E.J. Sept. 8, 1907.
5. FiSSlDENS EXIGUUS Sullivant
Plate XV
Plants very small, gregarious, light green: stems, in our specimens, 1-2.5
mm, high, erect, or ascending; leaves usually 3-5 pairs, the lower minute, the
upper reaching 1.5 mm long, ascending to erect, oblong-lanceolate, acute, only
the sheath margined, entire, the dorsal lamina narrowing to none at the base,
the sheath about one-half the length of the leaf; costa stout, vanishing a little
below apex; cells in apical lamina quadrate to hexagonal, at base of leaf be-
Jennings: Manual of Mosses — 4. Fissidentaceae 73
coming rectangular, the sheath being bordered, especially in its upper part, by
a border one to four cells wide, of elongate and more or less prosenchymatous
cells, all cells moderately incrassate or more so in border of sheath: capsule on
an erect, somewhat flexucus, stout, reddish pedicel 2-5 mm long; capsule
oblong-oval, narrowed to pedicel at base, somewhat constricted below mouth,
smooth, reddish-yellow, about 0.5-0.7 mm long; operculum conic-rosrrate,
about two-thirds the length of the capsule; calyptra narrow, dimidiate; teeth
red, split to the middle into two awl-like prongs which have spiral thickenings,
closely infolded in wet specimens, inserted a little below edge of mouth;
spores sm.ooth, about .020 mm in diameter. Mature in September.
On stones and rocks in stream beds, especially in ravines. Southern Can-
ada, United States east of the Rockies, south to North Carolina, England.
Allegheny Co.: On sandstone rocks, ravine of Powers Run, November 30. 1909.
O.E.J. Fayette Co.: Cheat Haven, September 3-6. 1910. O.E.J, and G.K.J, (figured).
6. FiSSIDENS CRISTATUS Wilson
(f. decipiens De Notaris)
Plate XV
Usually tufted, branching from the base, green to dark green; stem erect,
1-2 cm high; leaves numerous, ascending, imbricate, the upper reaching 2.5 mm
long, oblong-lingulate, acute, crenulate below, irregularly serrate above, inferior
lamina narrowed and somewhat decurrent at base, sheath e.xtending half-way
to apex or a little above; costa strong, ending just below or in the apex, some-
times a little excurrent; leaf-cells irregularly angular to rounded-he.xagonal,
about .003-. 012 mm. in diameter, some of these ne.xt to the costa larger, the
marginal 3 or 4 rows paler and forming a rather obscure belt around the leaf,
all cells incrassate: seta ascending, usually about 1 cm high, smooth, light chest-
nut color, arising from the lower half of the stem; capsule oblong, smooth,
about 2 mm long, tapering to the seta, ascending to nearly erect, chestnut-
brown, constricted below the mouth at least when old; peristome bright red-
chestnut, the teeth split at one-third above the base into two very slender,
trabeculate, somewhat spirally papillose prongs; operculum conic, rostrate;
spores about .020 mm, in diameter, smooth, pale yellowish, globose. Mature
in winter or early spring.
On moist soil and stones or occasionally at base of trees. Nova Scotia to
the Gulf States and the Rocky Mountains, Europe, Asia.
Known from the following counties: Allegheny, Armstrong, Cameron, Crawford, But-
ler, Elk, Fayette, La.vrence, McKean, Somerset, Warren, Washington, and Westmoreland.
Specimen figured: On bark at base of black ash, Linesville, Crawford Co., June 11-12,
1907. O.E.J.
7. FiSSIDENS ADiANTOiDES [Linnaeus] Hedwig
This species differs chiefly from F. cristatiis in that the cells are larger, .012-
.016 X .015-. 025 mm, distinct; border none or rather indistinct: seta usually
longer than in F. cristatiis, about 1-2.5 cm long. The plants are often much
larger, 2-15 cm high, and are monoicous instead of dioicous, as in F. cristatus.
The two species apparently intergrade.
74 American Midland Naturalist Monograph No. 6
This spcies is reported as common in the eastern United States but, with
the following exceptions, all the specimens we have seen from our region
labeled as F. adiantoides we have referred to F. cristatus.
Washington Co.: Shaded ravine, McCracken Sta., Sept. 24, 1892; near Washing-
ton, Dec. 10, 1892; and ravine near Claysville, Dec. 17, 1892, all A. Linn and J. S.
Simonton.
8. FissiDENS taxifolius [Linnaeus] Hedwig
, (Hypnum taxifolium Linnaeus)
Plate XV
Plants gregarious, light green, branching at base, usually 5-10 mm high,
erect to ascending: stem rather stout and rigid; leaves close, imbricate, oblong-
ovate, apiculate, uniformly crenulate, non-bordered, ascending, the middle
leaves usually longest and up to 2 mm. long, the inferior lamina ending
abruptly ?.t the base, sheath extending to the middle or beyond; costa strong
and excurrent in the apiculus; leaf-cells rounded-hexagonal, about .010 mm in
diameter, incrassate, one or two rows next the costa larger, the marginal row
usually a little paler, the costa at the apex widening and consisting of elongate
parenchymatous cells: seta about 8-14 mm long, flexuous-ascending, smooth,
yellowish-castaneous, arising near the base of the plant; capsule varying from
sub-pendulous to erect, oblong, slightly inflated on the back, smooth, about 1.5
mm long, tapering abruptly to the seta, castaneous to dark brown; {peristome
bright red-chestnut, the teeth inserted a little below the mouth of the capsule,
forked to below the middle, the prongs very slender, trabeculate, somewhat
spirally papillose; spores smooth, about .016-. 017 mm in diameter, pale yel-
lowLsh-pellucid; operculum conic, obliquely rostrate to about half the length
of the capsule. Mature in late fall or winter.
On damp clayey soil, Canada and eastern United States, south to the
Gulf. Europe, Asia, Africa.
Known from Allegheny, Butler, Fayette, McKean, Lawrence, Tioga, and Washingon
counties. Specimen figured: Bennett, McKean Co., Oct., 26, 1898. D.A.B.
9. FiSSIDENS OSMUNDIOIDES [Swartz] Hedwig
(Dicranum osmund'wides Swartz)
Densely tufted, 1-5 (TO) cm high, dark green, tomentose below with
brown rhizoids: stems simple or sometimes branched basally, erect; leaves
numerous, close but hardly imbricated, the apical ones the largest, oblong-
lanceolate, serrulate towards the apex, non-bordered, usually rounded and
apiculate at apex, the sheath reaching from one-half to two-thirds the I'af-
length, inferior lamina often ceasing abruptly at base and not decurrent; costa
ending just below the apex; leaf-cells oval- or rounded-hexagonal, large, about
.010-.018 x .012-.025 mm, incrassate, a single row at margin often paler, pel-
lucid, and a little smaller: seta terminal, yellowish to chestnut-red, about 5-10
mm long; capsule narrow-oblong, sub-erect, to inclined, thick-walled, chestnut-
brown or darker; operculum conic with a needle-like usually straight bea'c nea ly
as long as the urn; calyptra cucullate or several-lobed at base; spores smooth
about .018-.025 mm. Mature in midsuinmer.
Jennings: Manual of Mosses — 4. Fissidentaceae 75
In swampy woods and along streams, quite widely distributed in the
cooler portions of the Northern Hemisphere, reaching the northern United
States and south to Missouri, Ohio, North Carolina, Georgia, and Tennessee.
It occurs in eastern Pennsylvania and in Ohio but has not yet been found in
western Pennsylvania.
10. FiSSIDENS SUBBASILARIS Hedwig
Plate XVI
Cespitose in wide mats, 5-10 mm high, erect or ascending, green, brownish
tomentulose at base: stems branching at base; leaves usually in 10-18 pairs,
crisped when dry, widely spreading to ascending, close, imbricate, those in
middle of stem often largest, the largest reaching about 1.5 mm, the sheath
reaching about three-fifths the length of the leaf, leaf oblong, rather obtuse,
but apiculate with a pointed cell, non-bordered, minutely crenulate below,
irregularly serrate above, the inferior lamina ceasing abruptly at the base; leaf-
cells incrassate, and rather obscure, small, about .007-. 01 2 mm, roundish-
hexagonal; the costa ending considerably below the apex: seta smooth, arising
from basal part of stem, ascending, usually about 3-5 mm long and reaching
about to the top of stem, light chestnut-color; capsule cylindric-oval, about 1.5
mm long, smooth, chestnut-color to dark brown, tapering at base, erect or very
nearly so; calyptra narrowly cucullate; operculum conic, obliquely rostrate to
about one-half the length of capsule; peristome rich chestnut-color, strongly
trabeculate, not papillose, the teeth slightly inserted, bifid to about the middle
into two slender prongs; spores smooth, pale yellowish pellucid, round, about
.016-. 018 mm in diameter. Mature in late autumn.
On earth and on rocks and bases of trees, Ontario and southwards
through our Eastern States to the Gulf.
Now known from the following counties: Allegheny. Butler, Fayette, Indiana, Mc-
Kean, Somerset, Tioga, Washington, and Westmoreland. Specimen figured: On base of
white oak, Douthett, southern Butler Co., O.E.J. Dec. 27, 1908.
3. OcTODiCERAS Bridel
{Conomitrium Montagne)
Plants slender, fasciculately branching, floating, filiform: leaves remote,
lance-linear, short-auriculate : flowers monoicous, the male axillary, the female
on elongated branchlets; seta short; capsule thin-walled, erect, very small,
without stomata; calyptra minute, conic, undivided, covering only the rostrum
of the operculum; opserculum conic-rostrate; peristome-teeth variously laciniate
or entire and evidently degenerate; annulus none; spores about .018-.025 mm.
This genus includes about 20 species of aquatic mosses more or less resem-
bling Fontinalis in general appearance, widely distributed over the earth, — 2
species occurring in eastern United States, one already found and another
likely to be found in our region.
Key to Species
A. Large much-branched plants, up to 15 cm. long: seta shorter than the capsule
1. O. debile
76 American Midland Naturalist Monograph No. 6
A. Small little-branched plants, up to 4 cm long: seta longer than the capsule
2. O. Hallianum
1. OcTODICERAS DEBILE (Schwaegrichen) Jennings
(Octodiceraf julianus Bridal; Cor.omitrium Julicinum Montagne;
Fontinalis Juliana Savi)
Plate LXI
Plants up to 15 cm long, flaccid, floating, blackish-green below, much
branched: leaves up to 5 or 6 mm long, distant, spreading, numerous, linear-
lanceolate, entire, non-bordered, vaginant lamina one-fourth to one-third the
length of the leaf, inferior lamina not quite reaching base, costa ending slightly
below the apex; leaf-cells irregular hexagonal or more quadrate below, about
.015-.020x .020-. 030 mm, somewhat incrassate, within the border at the lower
outer corner of the vaginant lamina several rows of elongated linear-oblong
cells: seta shorter than capsule, pale, fragile at base, elliptic, erect, scarcely
raised above perichaetical bracts, symmetric; operculum conic-rostrate and
about as long as urn; peristome-teeth short, imperfect, 16, yellowish-pellucid,
irregularly cleft and perforate in upper part; calyptra conic, dark, erose at
base; spores about .020-. 022 mm, mature in summer.
Almost cosmopolitan, but rather local, on stones and on wood in creeks
and springy swamps.
This species has not been collected in western Pennsylvania except as follows: BuTLER
Co.: Walley Mill region^ Parker Twp., on submerged rocks in brook. Sidney K. East-
wood, July 14, 1935 (figured). Crawford Co.: On clay bank along Shenango River,
west of Lineville. W. R. Van Dersal, Oct. 15, 1933. Huntingdon Co.: Porter.
(Porter's Catalogue.)
2. OcTODiCERAS Hallianum (SuIIivant and Lesquereux)
Jaeger and Sauerbeck
{Conomitrium Hallianum SuIIivant and Lesquereux; Fissidens Hallianus Mitten)
Plants smaller, up to 3-4 cm long, laxly tufted, dirty-green: stems sparsely
fasciculate-branching at base; leaves remote, narrowly linear-lanceolate, usually
in 5-10 pairs, entire, the sheath not reaching over one-fourth or one-third the
length of the upper pair of leaves, inferior lamina narrowing and reaching
almost to the base; cells irregularly hexagonal, tending to quadrate below,
about .015-. 022 mm: seta longer than capsule, pale; capsule pale, elliptic-oblong;
peristome-teeth undivided, reddish, subulate-lanceolate, articulate, inserted
below the mouth of urn, papillose; operculum acutely conic-rostrate and about
as long as urn; calyptra cucullate, covering the entire operculum; spores smooth,
about .018-.024 mm.
On wood and stones, in streams, swamps, etc., New Jersey, New York,
Illinois, and Idaho. Not reported for our immediate region, but perhaps
overlooked on account of small size.
Family 5. Pottiaceae
Autoicous or dioicous, rarely par-, syn-, or polyoicous: mostly small or
medium,-sized, more or less densely cespitose, rarely gregarious: stems mostly
Jennings: Manual of Mosses — 5. Pottiaceae 11
with central strand, radiculose below, thickly foliate, simple or more or less
branched; leaves pluri-seriate, rarely 3-seriate, lanceolate to broadly ovate or
obovate; costa heterogeneous, mostly percurrent, or excurrent, sometimes with
longitudinal lamellae or with green branched filaments on the ventral surface
above the middle; leaf-cells parenchymatous, the basal rectangular to elongate,
mostly pellucid, or hyaline, upper cells always chlorophyllose, on both sides
mostly warty papillose, loose, sometimes towards the apex 4-6-angled, or small
and rounded-quadrate; seta more or less elongate, mostly straight, rarely
almost lacking; capsule erect, symmetric, rarely slightly inclined, straight to
slightly arcuate, mostly oblong to cylindric, rarely oval to spherical; collum
short, rarely none; peristome various to none, mostly inserted on the mouth of
the urn, usually without projecting trabeculae; teeth 16, straight or spirally
twisted, often united at base into a tube, entire or 2-3-cleft into filiform-
subulate divisions, papillose; operculum mostly conic, rostrate; calyptra mostly
cucullate, smooth, rarely papillose or minutely bristly or short-hirsute.
A very large family, mainly confined to the temperate zones, occurring
almost entirely on soil or on rocks. The systematic relationships and the
scope of the family are variously treated by different bryologists who have
taken different characters as the basis for the various classifications. This
family was called Tortulaceae in the first edition, but for purposes of uniformity
it is here called Pottiaceae, following Brotherus (Pflanzenfamilien, 2nd edit.,
1924, and Grout, Moss Flora, 1938).
Key to Genera
A. Minute, bud-like plants with broadly ovate, concave or carinate leaves; cleistorarpous
9. Acaulon
A. Not as above B
B. Leaves mostly narrow, often linear-lanceolate, never broadest above the middle
except some occasionally Imgulate in Gymnostomum calcareum; costa with sev-
eral guides, no accompanying cells, but 2 stereid bands, rarely longe.xcurrent C
B. Leaves mostly broad, ovate-oblong ct spatulare or lingulate; costa with 2 median
guides, with accompanying cells, and 1 stereid band, mostly more or less lono-
excurrent M
C. Plants minute; areolation dense, strongly papillose: capsules cleistocarpous
\. Astomum
C. With deciduous operculum D
D. Peristome none E
D. Peristome present, rudimentary or well-deveIop>ed F
E. Operculum deciduous with the columella detached 4. Hymenostylium
E. Columella remaining in the urn after the falling away of the operculum
3. Gymnostomum
F. The exterior surface of the teeth more strongly developed and with projecting
plates; leaves crispate when dry; seta long 2. Weisia
F. Both surfaces of the teeth equally well-developed and no projecting plates G
G. Perichaetial leaves long-convolute-sheathing 8. Barbida
G. Perichaetial leaves not or but little convolute-sheathing H
H. Leaves more or less lingulate, margins plane; cells smooth 7. Didymodon
H. Leaves more or less lanceolate j
J. Leaf-margins plane or involute; cells papillose K
I. Leaf-margins more or less revolute; leaf-cells nearly smooth or papillo.e L
K. Divisions of p>eristome erect or slightly dextrorsely twisted 5. Trkhostomum
73 American Midland Naturalist Monograph No. 6
K. Divisions of peristome distinctly sinistrorsely twisted ...6. Torlella
L. Peristome-teeth 16, more or less 2-cleft or perforate, erect or', dextrorsely ascending
7. Didynwdon
L. Peristome-teeth 32, filiform, strongly twisted sinistrorsely 8. Barbula
M. Cleistocarpous; capsule spherical to oval, apiculate; leaves ovate to broadly lan-
ceolate 10. Phascum
M. Operculate N
N. Peristome-teeth 16 or none O
N. Peristome-teeth 32, filiform, sinistrorsely twisted, with a high basal membrane;
leaves oblanceolate to obovate or spatulate 13. Tortula
O. Peristome-teeth none, or rudimentary from a low basal membrane 11. Pottia
O. Peristome-teeth small, separate to the base, more or less divided into two or three
slender prongs 12. Desmatadon
I. AsTOMUM Hampe
Autoicous, rarely polyoicous: small, gregarious to cespitose, dull green:
stem with a few-celled central strand, radiculose, thickly foliate; upper leaves
tufted, when dry mostly crisped, keeled, from a broad base lanceolate to sub-
ulate-lanceolate, margin plane to involute, entire; casta strong, percurrent or
excurrent; leaf-cells in upper part of leaf small, rounded-quadrate, papillose
both sides, the lower cells elongate-quadrangular, thin-walled and hyaline:
capsule mostly immersed, almost spherical to oblong-elliptic, m.ostly with a
small, elongate-conic op)erculum, which, however, is rarely deciduous; calyptra
cucullate, rarely mitrate, smooth.
A widely distributed genus of 21 terrestrial species; 3 species occurring in
North America; 2 (3) species in our region.
1. AsTOMUM Muhlenbergianum (Swartz) Grout
(Astomum Sullivantii Schimper; A. crispum Am. Auth.)
Plate LXII
Densely cespitose, pale to dark green: stem about 5 mm high, usually
branched above, erect; leaves numerous, close, when dry crispate, the stem-
leaves small, lance-linear, the comal and perichaetial much larger, up to 4 mm
long, elongate-linear from a narrowly oblong, concave, whitish base, usually
narrowly involute or canaliculate above, the apex abruptly acute; costa strong,
acutely and shortly excurrent-mucronate, sometimes upturned so as to make
the leaf somewhat cucullate; basal leaf-cells laxly and irregularly lon^-'ectangu-
lar, hyaline, upper leaf-cells sub-quadrate, densely chlorophyllose, papillose:
seta erect, shorter than the capsule; capsule immersed, ovoid, small, brownish,
about 1 mm long; lid finally distinct but not separating from the urn of its
own accord, minute, obliquely conic-apiculate to short rostellate; exothecial
cells laxly hexagonal to oblong-hexagonal, one to three rows of cells being
somewhat smaller at the junction of the lid; calyptra cucullate; spores papil-
lose, .021 -.027 mm, mature in spring: autoicous; the immature capsule fully
grown by late November.
In old sandy or clayey fields and along roadsides, principally in non-
calcareous districts, temperate Europe, Japan, Algeria, and, in North America,
from Saskatchewan to Massachusetts and southward to the Gulf States.
Jennings: Manual of Mosses — 5. Pottiaceae 79
Beaver Co.: James. (Porter's Catalogue). Indiana Co.: James. (Porter's Cata-
logue). About 3 miles northeast of Saltsburg along abandoned road above Black Leg
Creek, Nov. 23, 1941. O.E.J, and G.K.J, (figured). Washington Co.: Snake Woods,
near Washington. Mar. 10, 1894. Linn and Simonton.
2. AsTOMUM NITIDULUM Bryologia Europaea
{Systegium nitidulum (Jaeger)
The moss reported in Porter's Catalogue as collected by James in Indiana
County, and listed as Astomum nitidulum Bry. Eur., is doubtful. It is sug-
gested that A. nitidulum is a hybrid of Astorrmm Muhlenbergianum and
Weisa viridula, or that it is A. phascoides (Hook.) Grout from Ohio west-
ward. This latter species has stouter leaves than A. Muhlenbergianum with
longer seta and a rostrate lid.
Indiana Co.: Derry. James. (Porter's Catalogue).
2. Weisia Hedwig
Autoicous, rarely paroicous, polyoicous, or dioicous: low, cespitose, freely
branching: upper leaves much larger, relatively to the lower, erect-spreading,
crisped when dry, carinate, elongate-lanceolate; costa strong, cuspidate-excurrent;
basal leaf-cells rectangular, hyaline, the upper small, rounded, low-papillose on
both surfaces: seta erect or sometimes curved, mostly longer than the peri-
chaetial leaves; capsule erect and symmetric or a little inclined and swollen
dorsally, round-ovoid to cylindrical, narrow-mouthed, finally usually some-
what plicate, the urn at the rim being several cells thick and the insertion of
the peristome thus considerably removed from the exterior border of the rim;
peristome-teeth short, often rudimentary, undivided, papillose, the exterior
layer more strongly developed and with projecting bars; lid obliquely long-
rostrate; calyptra cucullate.
A widely distributed genus of 27 terrestrial species; 7 species occurring in
North America; only one in our range.
1. Weisia viridula [Linnaeus] Hedwig
Plate XVI
Densely cespitose, yellowish-green: stem erect, often branching, up to 5
mm. tall; leaves erect-spreading, the upper much larger and up to 3 mm long
and 0.5 mm wide, lance-linear, tapering to an acute or acuminate apex, the
margin strongly involute, entire, leaves crispate when dry; costa strong, about
.030-. 040 mm wide at base, excurrent into a shoit and more or less hyaline
point; upper leaf-cells roundish-hexagonal, strongly papillose, obscure, the basal
more or less elongate-rectangular and hyaline: seta slender, up to 1 cm long,
lustrous, yellowish, faintly sinistrorse; capsule erect, ovoid, symmetric, slightly
narrowed at mouth, reddish-brown, about 0.9 mm long, somewhat plicate when
dry and empty; exothecial cells rather thin-walled, irregularly oblong to hexag-
onal or rounded, those at the mouth in 3-5 rows, much smaller, quadrate and
darker in color; peristome-teeth 16, more or less rudimentary, short, irregularly
linear, divided, or truncate, papillose; lid conic, obliquely long-rostrate, alto-
gether nearly as long as the urn; calyptra cucullate, covering about t\'.o-thirds
80 American Midland Naturalist Monograph No. 6
of the capsule; spores orange-pellucid, papillose, about .016-.019 mm in
diameter, mature in spring.
Almost cosmopolitan on bare earth in fields, excavations, along road-
sides, etc.
Allegheny Co.: On shale cliffs, Little Sewickley Creek, 2 mi. n. of Edgeworth,
C.M.B. Bedford Co.: On shale cliff, Gravel Pit Sta. C.M.B. June 15, 1941. Fayette
Co.: Ohio Pyle, on clay bank, September 1-3, 1906. O.E.J, and G.K.J, (figured.)
McKean Co.: Bradford, Nov. 21, 1896, Divide, Bolivar and Bennett, Dec. 15, 1896,
and Quintuple, Mar. 20, 1898. D.A.B. Washington Co.: Linn and Simonton. (Por-
ter's Catalogue). Westmoreland Co.: Clay bank, Darlington, C.M.B. May 19, 1945.
3. Gymnostomum Hedwig
Dioicous: densely cespitose, rusty in color below: stem thickly foliate,
sparsely radiculose, in cross-section circular, the central strand few-celled, the
branching dichotomous; leaves erect-spreading, more or less carinate, not
crisped but when dry, more or less appressed and curved; elongate-lanceolate
or subulate to sub-lingulate, margin plane; costa strong, vanishing below the
apex; upper leaf- cells rounded-quadrate, small, thickly papillose on both sides,
as is also the costa, lower cells rectangular, the walls yellow to hyaline; peri-
chaetial leaves somewhat sheathing at the base; seta long, erect; capsule erect,
symmetric, oval or oblong, when ripe smooth and shining, the wall of the
capsule not distinctly thickened at the mouth; peristome none; operculum
conic, long rostrate, easily deciduous; calyptra narrowly cucullate, covering
about half of the urn.
A widely distributed genus of II species, mainly occurring on calcareous
rocks; 3 species occurring in North America; two in our region.
Key to the Species
a. Stems usually less than 2 cm; at least some of the leaves obtuse, narrowly lingulate
to ligulate 1. G. calcareum
A. Stems from 1 to 10 cm; leaves oblong to linear-lanceolate, more or less acute
2. G. aeruginosum
1. Gymnostomum calcareum Nees and Hornschuch
(G. viridulum Bryol. Eur.; G. tenue of L. and J. Manual)
Plate XVI
Densely cespitose, yellowish-green: stems erect, branched, up to 10 mm
high; leaves about I mm long, spreading, somewhat recurved, elongate-oblong-
lanceolate, somewhat concave, usually larger and tufted at the apex of the
stem, obtuse, plane-margined; upper leaf-cells densely papillose, small, incras-
sate, obscure, the interior basal cells hyaline, rectangular or up to 2-3:1; costa
strong, ending below the apex: seta erect; capsule oblong, often somewhat con-
stricted below the mouth when dry and empty, erect, symmetric, tapering
below; lid conic, obliquely rostrate, the beak one-half to two-thirds as long as
the urn; calyptra cucullate; annulus none; peristome none; exothecial cells
rectangular to quadrate, at the mouth becoming smaller, darker and quadrate
in 3 to 5 rows; spores smooth, .008-.0I1 mm in diameter, mature in summer.
Jennings: Manual of Mosses — 5. Pottiaceae 81
Cosmopolitan on damp limestone rocks and boulders, but rare in our
region.
Bradford Co.: On moist shaly limestone cliff. June 25, 1935. S. K. Eastwood.
Fayette Co.: On wet cliff, one mile up Meadow Run, Ohio Pyle, C. M. Boardman,
lune 23, 1940. Lawrence Co.: Gorge near Rock Point, June 26, 1909 (figured).
Sterile. O.E.J.
2. Gymnostomum aeruginosum Smith
(G. TUpestre Schleich)
Forms dense cushions; stems slender, branched, 1 to 3 or 4 or occasionally
even to 10 cm long; leaves oblong- to linear-lanceolate, about 1-1.5 mm long:
capsule thin-walled, brownish; spores about .010-. 012 mm in diameter, mature
in summer.
On moist limestone rocks, from southern Canada through eastern United
States in Texas
Huntingdon Co.: Spruce Creek, T. C. Porter.
4. Hymenostylium Bridel
Dioicous: densely and deeply cespitose, green to rusty or yelIowish-['reen:
stem densely foliate, sparsely radiculose, without a central strand, triangular in
cross-section; leaves erect spreading, rarely squarrose-recurved, when dry in-
volute, sometimes somewhat twisted when dry, not crisped, more or less
carinate, elongate-lanceolate, acuminate; costa mostly ending below the apex;
laminal leaf-cells thick-walled, smooth or papillose: seta long, erect; capsule
erect, symmetric, obovate, firm, when empty smooth and pyriform; peristome
none; lid obliquely long-rostrate from a broad base, remaining attached to the
columella and deciduous thus attached; calyptra cucullate, covering about
half of the urn.
A widely distributed genus of about 25 species, occurring mainly on cal-
careous rocks; 8 species in North America; only one occurring in our region.
1. Hymenostylium recurvirostrum (Ehrhart, Hedwig) Lindberg
(^Gymnostomum curvirostrum Bridel; Weisia curvirostris Mueller;
Gymnostomum recurvirostrum Hedwig 1801)
Plate XVI
Closely cespitose, 2-4 cm high, bright green above, darker and more or less
ferruginous below: leaves little or not at all twisted when dry, erect to recurved-
spreading when moist, narrowly lanceolate acuminate, 1-1.5 mm long, apex
acute, base sub-clasping, margin entire but papillose, as are also the entire
upper and lower surfaces of the lamina and costa, one or both lower margins
recurved; costa strong, vanishing just below the apex, at base occupying about
one-eighth the entire width of the leaf; upper leaf-cells rounded to sub-quad-
rangular, the lower towards the costa becoming elongate-rectangular: seta 8-10
mm long, lustrous, castaneous; capsule about 1 mm long, rounded ovate, lus-
trous, castaneous, widest towards the mouth, when dry and empty decidedly
82 American Midland Naturalist Monograph No. 6
urceolate; peristome none; operculum v/ith a long and oblique rostrum at least
two-thirds the length of the urn, the operculum often remaining attached to
the columella for some time after the spores have been shed; spores yellowish,
moderately incrassate, smooth, about .014-017 mm in diameter, mature in
September or October.
Not uncom.mon on wet cliffs, principally calcareous, in Europe, Asia,
northern Africa, and, in North America, from Alaska to Labrador south to
California and the Carolinas.
Allegheny Co.: Guyasuta Hollow, Aspinwall, on wet cliff near waterfalls, October
12 and 25 (figured) 1908, and September 8, 1909. O.E.J. Butler Co.: On cliff, Win-
field Jet., Buffalo Creek. June 8, 1940. C.M.B. Lawrence Co.: On wet face of
exposure of the Homewood Sandstone, near Rock Point, October 15, 1910. O.E.J, and
G.K.J. Somerset Co.: Vicinity of Trent. Aug., 1932. Chas. M. Hepner.
5. Trichostomum Bruch
Dioicous, rarely autoicous: densely cespitose, medium size, green to yel-
lowish-green: stem with central strand, erect, radiculose, rarely felted, densely
leaved, mostly dichotomously branching; leaves spreading, mostly crisped when
dry, upper leaves much the larger, long and narrow, more or less concave to
canaliculate, margins mostly erect to involute, often undulate, mostly entire;
costa well-developed, sometimes ending below the apex or excurrent; upper
leaf-cells small, rounded, chlorophyllose, papillose on both faces, towards the
base elongated-rectangular, mostly hyaline: seta long, erect; capsule erect,
rarely inclined, mostly symmetric, oblong-cylindric to cylindric, short-necked,
rarely strumose; basal membrane of peristome low or none, the teeth 16, erect,
smooth or papillose, red or yellow, undivided or cleft into two filiform non-
articulated divisions which are sometimes approximate in pairs; spore small;
lid conic, rostrate, the exothecial cells of the base in vertical series or rarely
dextrorsely ascending; calyptra cucullate, smooth.
A genus of about 80 species, v/idely distributed on earth and rocks. Sev-
eral species in North America; only one in our region.
1. Trichostomum cylindricum (Bruch) C. Mueller
(Didymodon cylindricus Bryologia Europaea; T. tenuirostre Lindberg)
Plate XVI
Rather loosely and softly cespitose, yellowish, dark below: stems erect
branching, rather flexuous, reaching to 1.5-2 cm in height; leaves about 2-3 mm
long, narrowly linear-lanceolate, when dry crisped and contorted, when moist
spreading or flexuous, gradually acuminate or sometimes rather abruptly nar-
rowed on an acute apex, the margin papillose-sinuate, plane or involute; basal
leaf-cells elongate-rectangular or more or less angular-oblong, somewhat in-
flated, hyaline in a broad band that does not extend up the margin, above
rather abruptly becoming much smaller, incrassate, quadrate to rounded-
hexagonal, the median and upper rounded-quadrate to rounded-hexagonal or
transversely oblong, densely papillose, much incrassate; costa strong, usually
forming the apex of larger pellucid cells: seta single or sometimes in pairs,
Jennings: Manual of Mosses — 5. Pottiaceae 83
slender, erect, about 1.5 mm long, yellow; capsule linear-cylindric, brownish;
lid conic and obliquely rostrate; peristome-teeth short, untwisted, linear-subu-
late, fragile, usually more or less irregularly cleft or perforate; spores about .012
mm in diameter, mature in autumn: dioicous: fruit produced but rarely.
On wet non-calcareous stones in brooks or at the base of cliffs in hilly or
mountainous districts, in Europe, Asia, South America, and, in North Amer-
ica, from Greenland to Manitoba and southward in the mountains to North
Carolina.
Rare in our region. Huntingdon Co.: Alexandria. Porter. (Porter's Catalogue).
McKean Co.: Toad Hollow, Bradford, July 19, 1896. Sterile. D.A.B. (figured).
6. Tortella (C. Mueller) Limpricht
Dioicous; rarely autoicous: widely and deeply cespitose, the cushions often
yellowish-green outside, brov/nish inside: stem erect, mostly without a central
strand, felted-radiculose: leaves tufted at the apex of the stem, widely spreading
to recurved-squarrose from a whitish and shining base, cirrhate-crispate when
dry, elongate-lanceolate to subulate, margin undulate, entire, usually involute
above; costa strong, ending in the apex or excurrent; basal leaf-cells differen-
tiated, hyaline, elongate-rectangular, extending up the margins and forming a
Y-shaped area, smooth; upper cells green, small, rounded- quadrangular, thickly
papillose on both sides: seta red, long, erect; capsule erect to inclined, oblong
to cylindric; annulus rarely differentiated; peristome attached below the rim
of the urn, the basal membrane low, teeth 32, filiform, sinistrorsely wound,
papillose; spores small; lid small and elongate-conic; calyptra cucullate, smooth,
long-rostrate.
A cosmopolitan genus, the 37 species mainly occurring on soil or on rocks;
5 species in North America; 2 in our region.
Key to the Species*
A. Dioicous: leaves long-acuminate 1. T. tortuosa
A. Autoicous: leaves linear-lanceolate to oblong-lanceolate, costa shortly excurrent as an
abrupt mucro 2. T. humilis
1. Tortella tortuosa [Linnaeus] Limpricht
(Bar bill J tortuosa Weber and Mohr; Tortula tortuosa Ehrhari)
Densely cespitose in rounded tufts, yellowish or pale green above, brown-
ish below: stems stout, branching, up to 6 cm high, red-brown-radiculose;
leaves crowded, usually 4-6 mm long, lance-linear, tapering to a gradually
acuminate apex, flexuous-spreading, margin crenulate-papillose, more or less
undulate, plane at the apex; leaves when dry strongly spirally crispate-con-
torted; costa strong, pale, excurrent into the fine and sometimes denticulate
acumen; basal leaf-cells thin-walled, hyaline, extending obliquely up the mar-
"^ Tortella nitida (Lindb.) Brotherus, with leaves (when dry) curling in circles at the
tip and with the costa lustrous dorsally; and also T. fragilis (H. & W. ) Limpricht, with
leaves curled only slightly at the tip when dry and frequently broken off, may be expected
in the northern part of our range.
84 American Midland Naturalist Monograph No. 6
gin, above becoming abruptly smaller, chlorophyllose, rounded, incrassate,
papillose: seta 1-3 cm long, reddish below, paler above; capsule cylindric,
2.5-3.5 mm long, usually somewhat curved, almost erect; lid obliquely and
slenderly conic-rostrate, at least one-half as long as urn; peristome-teeth long
and from a low basal membrane, two or three times dextrorsely twisted; spores
mature in late spring or early summer.
On rocks, usually calcareous, in hilly or mountainous districts, Europe,
Asia, northern Africa, and, in North America, from Greenland to the Gulf
and from Idaho to Vancouver Island.
Apparently rare in our region. Cambria Co. : Cresson. James. (Porter's Catalogue).
2. ToRTELLA HUMILIS (Hedwig) Jennings
(Tortula caespitosa Hooker and Greville; Barbula caespitosa Schwaegrichen )
Plate XVII
Loosely cespitose, green to yellowish-green, about 5 mm high; leaves
crispate when dry, erect-spreading when moist, oblong-lanceolate and about 2
mm long below, the upper linear-lanceolate and up to 3.5 mm long, somewhat
concave, the margin plane or sometimes involute, the perichaetial leaves simi-
lar and sheathing; costa strong, excurrent-cuspidate; the lower one-fourth of
the leaf has a large V-shaped patch of hyaline rectangular cells reaching about
.018 X .085 mm, the median cells rounded-hexagonal, papillose, rather opaque,
much smaller, about .007-.008 mm in diameter, the upper similar: seta yel-
lowish-brown, 15-20 mm long, erect, dextrorse; capsule yellowish-brown; ovoid-
cylindric, about 2-2.5 mm long, 0.5 mm thick, erect, symmetric, sometimes
arcuate, tapering at the base; peristome single, of 32 filiform, papillose, articu-
late teeth about 0.6-0.8 mm long, two or three times dextrorse, arising from a
low membrane scarcely exserted above the mouth of the capsule; spores globose,
somewhat papillose, about .008-.011 mm, mature in early summer; operculum
narrowly conic-rostrate; calyptra smooth, cucullate, rostrate, covering about
one-half of the capsule.
Almost cosmopolitan in temperate or sub-tropical regions on earth and on
the roots of trees in the woods.
Allegheny Co.: Coraopolis, September 11, 1905, and near Carnot, October II,
1908. O.E.J.) (figured). Cambria Co.: James. (Porter's Catalogue). Fayette Co.:
Ohio Pyle, September 1-3, 1906. O.E.J, and G.K.J. Huntingdon Co.: On limestone
rocks, Pennsylvania Furnace, July 13, 1909. O.E.J. Lawrence Co.: On VanPort lime-
stone, Squaw Run, n. of EUwood City. C.M.B. Nov. 26, 1948. McKean Co.: Bolivar
Run, September 6, 1897. D.A.B. Somerset Co.: Vicinity of Trent. Aug., 1932. C.
M. Hepner. Washington Co.: At root of tree, Snake Woods near Washington. Nov.,
1891. Linn & Simonton. Westmoreland Co.: Chestnut Ridge, s.e. of Torrance.
C.M.B. June 13, 1943.
7. DiDYMODON Hedwig
Dioicous, rarely synoicous; paraphyses filiform: mostly slender plants, red
or brown, cespitose: stem with central strand, thickly foliate, radiculose, the
branches reaching to about the same height; leaves more or less keeled, erect-
spreading, mostly lanceolate from a broad base, the margin revolute; costa
Jennings: Manual of Mosses — 5. Pottiaceae 85
well developed, upwards cylindrical, rarely excurrent; leaf-cells small, rounded-
quadrangular, often smooth, sometimes the basal elongate and pellucid: seta
long, erect; capsule erect, oblong to cylindric, sometimes slightly arcuate,
short-necked, smooth; no annulus; peristome inserted on the edge of the urn,
papillose, the trabeculae projecting; teeth 16, plane, narrow, undivided, or
perforate, or cleft to the base into filiform parts approximate in pairs; spores
small; operculum conic-rostrate; calyptra smooth.
A widely distributed genus of 90 species, on soil or rock, mainly in tem-
perate regions; 17 species in North America; only one in our region.
1. DiDYMODON RECURViROSTRis [Hedwig] Jennings
(D. rubellus Bryologia Europaea; Barbula rubellct Mitten;
Weisict recurvirostra Hedwig)
Cespitose in large, soft patches, bright green above, rusty red below; stems
erect, branched, usually 2-5 cm high, radiculose below; leaves when dry flexu-
ous and somewhat curled, when moist somewhat recurved-spreading from the
appressed and whitish base, narrowly lance-linear, the comal longer, abruptly
acute, margin narrowly revolute to near apex, apex obscurely denticulate; costa
either ending in the apex or minutely apiculate-excurrent; basal leaf-cells
elongate, rectangular, pellucid, medium-walled, the median and upper much
smaller, papillose, rather obscure, quadrate; perichaetial bracts long-sheathing:
seta long, red, slender, sinistrorse; capsule erect, oblong-cylindric, becoming
reddish-brown, smooth; annulus fragile, revoluble; peristome-teeth 16, united
at base into a very low membrance, linear from a wider base, nodose-articulate,
reddish, minutely roughened, with the median line but rarely divided; lid
short, obliquely conic-rostrate; spores mature in summer or in early autumn:
paroicous or synoicous.
On wet, usually calcareous rocks, stones, walls, etc., widely distributed in
the Old World and, in North America, occurring from Greenland to Alaska
and south to the northern United States. Although not yet recorded from
our region this species is to be expected here.
8. Barbula Hedwig
Dioicous; paraphyses filiform: more or less slender and densely and deeply
cespitose, the tufts green to brownish: stems with central strand, thickly-
leaved, forked; leaves erect-spreading, rarely recurved-squarrose, keeled, oblong
to prolonged linear-lanceolate; with mostly revolute margins; costa strong,
ending in the point, or excurrent; leaf-cells very small, thickened and opaque,
papillose both sides; basal leaf-cells enlarged, quadrate to rectangular, colored:
seta long, erect; capsule erect, rarely a little inclined, oblong to cylindric,
straight or rarely a little arcuate; annulus distinct or none; peristome rarely
rudimentary, or none; the teeth united below into a rather low basal membrane
which is rather deeply inserted, the 32 teeth spirally one to several times dex-
trorsely wound, filiform; operculum conic-rostrate; calyptra cucullate, long-
rostrate, reaching to about the middle of the urn; spores small.
86 American Midland Naturalist Monograph No. 6
A genus of 300 species distributed over the whole earth, on soil and rocks,
many of these species are poorly defined and probably synonymous; about 20
species in North America; only two species definitely known from our region.
Key to the Species
A. Perichaetial leaves high-convolute-sheathing: seta yellow or later reddish
3. B. convoluta
A. Perichaetial leaves not as above: seta red or brown B
B. Stem-leaves obtuse, costa shortly mucronate-excurrent 2. B. unguiculata
B. Stem-leaves acute, costa not mucronate-excurrent C
C. Costa .070 mm wide at base and tapering gradually; leaves widely spreading or but
little recurved 1. B. acuminata
C. Costa .050 mm wide at base and of equal breadth to the middle; leaves strongly
recurved or squarrose (B. reflexa Bridel )
1. Barbula acuminata Hedwig
{B. fallax Hedwig)
Loosely and widely cespitose, brownish dull green: stems fastigiately
branched, slender, 1-5 cm high; leaves somewhat distant, recurved-spreading
or arcuate, appressed and slightly twisted when dry, lanceolate-acuminate from
the base, the base ovate, the leaves carinate and often faintly plicate below,
the margin revolute in the lower half at least, entire; costa strong, ending in
the apex; upper leaf-cells small, rounded to hexagonal, incrassate, strongly
papillose, gradually larger below, and at the lowest part of the base a few
elongate-rectangular and pellucid: seta reddish, capsule brownish, long-ovoid
to sub-cylindric, mostly symmetric and erect; lid long, often as long as the
urn, acutely rostrate-subulate; peristome-teeth reddish, long, filiform, dex-
trorsely much twisted, united at base into a low membrane; annulus none;
spores mature from late fall to spring: dioicous.
On moist earth, rocks, walls, etc., usually on calcareous substrata, in
Europe, Asia, northern Africa, and, in North America from the Arctic region
south to Virginia and Iowa. To be expected in northern Pennsylvania.
2. Barbula unguiculata [Hudson] Hedwig
Plate XVII
Densely cespitose, yellowish-green: stems erect, somewhat branching, usu-
ally about 1 cm high; leaves about 2 mm long, erect-spreading, somewhat
recurved, when dry spirally imbricate and twisted, oblong-lanceolate, sometimes
lingulate, obtuse, mucronate, entire, the margin recurved below, plane above;
costa strong, excurrent and thus forming the rounded mucro; upper leaf-cells
small, about .008-.010 mm, rounded-quadrate, incrassate, strongly papillose,
obscure, the basal elongate-rectangular, yellowish-pellucid to more or less hya-
line, the marginal not different; perichaetial leaves longer and more erect:
seta erect, castaneous, lustrous, about I cm high, when dry sinistrorse; capsule
oblong-cylindric, deep-castaneous, the urn about 1.8 mm long, erect, exannu-
late, rather smooth when dry and empty; lid about one-third as long as urn,
conic-rostrate, slightly curved or straight; the 16 peristome-teeth castaneous,
pellucid, papillose, cleft to the base into 32 filiform divisions, from a narrow
Jennings: Manual of Mosses — 5. Pottiaceae 87
membrane at the base twisted into about two turns dextrorsely; spores smooth-
ish, yellowish, about .009-.0I2 mm, mature from November to spring: dioicous.
A quite variable species occurring on moist earth, banks, stones, walls, etc.,
in Europe, Asia, northern Africa, and throughout southern Canada and
northeastern United States.
Allegheny Co.: Schenley Park, Pittsburgh, August, 1905 (figured); Fern Hollow,
Pittsburgh, January 21, 1906, and Powers Run, November 30, 1909. O.E.J. Butler
Co.: T. P. James. (Porter's Catalogue). Centre Co.: T. C. Porter. (Porter's Cata-
logue). Huntingdon Co.: T. C. Porter. (Porter's Catalogue). McKean Co.: West
Branch Swamp, Bradford, April 10, 1894. D.A.B. Washington Co.: Grove Sta..
near Washington. Nov. 5, 1892. Lmn & Simonton. Westmoreland Co.: T. P.
James. (Porter's Catalogue).
3. Barbula convoluta Hedwig
Plate XVII
Densely cespitose, yellowish-green: stems 1-3 cm high, usually about 1-1.5
cm, erect, branching; leaves about 1-1.5 mm long, when dry crisped, when
moist erect-spreading, often somewhat recurved, lance-oblong to lance-linear
or Ungulate, rounded to obtuse, sometimes sub-acute, concave, the margins
mostly plane or slightly recurved on one side at base, mostly minutely crenu-
late with bifid papillae; basal leaf-cells elongate-rectangular, pellucid to hyaline,
rather incrassate, smooth, median and upper leaf-cells elongate-rectangular,
pellucid to hyaline, rather incrassate, smooth, median and upper leaf-cells
small, sub-quadrate, densely papillose, strongly incrassate, often rather ob-
scure; costa strong, yellowish-pellucid, ending below apex or rarely shortly
apiculate-excurrent; perichaetial leaves high-convolute-sheathing, the inner ecos-
tate: seta erect, about 1.5 mm long, yellow, or reddish when old, sinistrorse
below, dextrorse above; capsule small, symmetric, erect, reddish-brown, narrow-
ly oblong, the urn about 1.5 mm long; lid conic-rostrate, oblique, about 1 mm
long, the cells spirally arranged; exothecial cells narrow, elongate-rectangular,
brownish or yellowish pellucid, two or three series at the rim much smaller,
sub-quadrate and darkly obscure; annulus distinct and narrow; peristome-teeth
consisting of 32 filiform articulate divisions several times dextrorsely twisted
from a low basal membrane; spores brownish-pellucid, medium-walled, smooth-
ish, about .016-. 018 mm, mature in spring: dioicous.
On soil, especially in calcareous districts, Europe, Asia, northern Africa,
and from southern Canada to Florida, Kansas, and California. In our region
occurring at Latshaw, southern New York, near the Pennsylvania State-line
(figured) and as follows, — not common:
Lawrence Co.: Enon Valley. T. P. James. (Porter's Catalogue). Lycoming Co.:
McMinn. (Porter's Catalogue ) .
9. Acaulon C. Mueller
(Sphaerangium Schimper)
Dioicous: minute, bud-like, brown, gregarious: stem very short and few-
leaved, without central strand, unbranched, green protonema persistent; leaves
erect-spreading, broadly ovate, keeled or concave, above with revolute and
88 American Midland Naturalist Monograph No. 6
sinuate-denticulate margin, or plane and entire; costa more or less excurrent-
cuspidate; upper cells short-rhombic, below rhomboidal, dorsally strong thick-
ened, smooth, rarely with a few high papillae, the lowest thin-walled, hyaline
and rectangular: seta very short; capsule sub-globose, cleistocarpous, immersed;
calyptra very small and delicate, conic-mitriform 3-5-cleft; spores small, brown,
subglobose, minutely granulose.
A widely distributed genus, on soil, mostly in the temperate zone. Fifteen
species in all, 3 in North America, 2 in our region.
Key to the Species
A. Uppermost and perichaetial leaves sharply carinate; costa strongly excurrent
1. A. triquetrum
A. Uppermost and jserichaetial leaves concave; costa rarely excurrent 2. rufescens
1. AcAULON TRIQUETRUM (Spruce) C. Mueller
{Sphaerangium triquetrum Schimper)
Plants minute, about 1 mm high, bulbiform, pale green or yellowish, dis-
tinctly triquetrous; protonema green, persistent; lower leaves small, rounded,
the middle leaves larger, broadly ovate, deeply concave, convolute, the upper
largest (perichaetial) broadly ovate, deeply carinate, triquetrous, all upper
leaves with reflexed margins, erose-denticulate above; costa strong, in upper
leaves excurrent in a recurved apiculus; basal leaf-cells elongate-hexagonal,
lax, thin-walled, hexagonal to ob long- hexagonal : seta about as long as capsule,
arcuate; capsule globose, smooth; calyptra minute, mitrate, cleft-lobed, cover-
ing only very apex of capsule; spores papillose, about .025-. 030 mm, mature
in early spring.
On clayey or sandy soil in fields or on banks, Europe, Algeria, and, in
North America, from v/estern Canada to New England south to the Caro-
linas; occurs in Ohio and in Pennsylvania but not yet reported from our range.
2. AcAULON RUFESCENS Jaeger
(Phascum rufescens Kindb. )
Plants minute, bulbiform, about 1 mm high, yellowish-green, not markedly
triquetrous: protonema persistent; lower leaves very small, ecostate, the upper
much larger, broadly ovate^ deeply concave, convolute, plane-margined, erose-
denticulate at apex; in larger leaves the costa thick, excurrent into a squarrose-
recurved apiculus; leaf -cells about as in A. triquetrum: seta flexuose or arcuate;
capsule globose, smooth; calyptra and spores similar to those of A. triquetrum.
On bare clayey or sandy soil in moist fields, eastern and central North
America.
Indi.ana Co.: Derry. James. (Porter's Catalogue, as Sphaerangium muticum [Schre-
ber] Schimper).
10. Phascum [Linnaeus] Hedwig
Autoicous or synoicous: very small, closely gregarious: stem short, without
central strand, erect, simple or bushy-branched; leaves mostly ovate-lanceolate
to elongate-lanceolate, mostly with entire and revolute margins, the upper
Jennings: Manual of Mosses — 5. Pottiaceae 89
mostly with a strong excurrent costa; upper leaf-cells quadrate to hexagonal,
warty-papillose on both sides, rarely smooth; basal leaf-cells rectangular and
hyaline: seta very short, sometimes curved; capsule immersed or slightly
emergent, sometimes two in a perichaetium, mostly globose and obtusely
apiculate, with no indication of an operculum; calyptra cucullate or rarely
mitrate, small, conic.
A widely distributed genus of 15 species, on soil; 5 occurring in North
America, one in our range.
1. Phascum cuspidaTum [Schreber] Hedwig
(P. acaulon Linnaeus)
Cespitose, deep green: stems short, 1-2 mm high, simple or forked; leaves
crowded, erect, the comal longest, oblong-lanceolate, acuminate, more or less
carinate, entire, revolute towards middle; costa excurrent; basal cells lax, hy-
aline, the upper rectangular to hexagonal, .015-.030 mm, finely dorsally papil-
lose: seta short, straight or curved; capsule globose, sometimes two or three
on the same plant, immersed or rarely emergent, obtusely apiculate, about 1
mm long, cleistocarpous; calyptra cucullate, conic, covering only the upper
portion of the capsule; spores large, .028-.035 mm, yellowish-pellucid, finely
roughened, mature in spring: autoicous or paroicous, antheridia clustered in
the axils of the upper leaves.
On soil in old fields, pastures, etc., usually preferring a sandy soil, in
Europe, Asia, Algeria, South America, and from Ontario to Virginia and
west to Arizona.
Rare in our region. Beaver Co.: T. P. James. (Porter's Catalogue).
1 1 . Pottia Ehrhart, Fuernrohr
Autoicous or paroicous, rarely synoicous or dioicous: small, gregarious to
cespitose, green to brownish or whitish: stem with central strand above, often
simple, radiculose at base, leafy; leaves tufted above, spreading to imbricate,
carinate to deeply concave, oblong to elongate-lanceolate, or spatulate, acu-
minate to piliferous, rarely obtuse, margin revolute or plane; costa without
lamellae, complete to excurrent, rarely incomplete; lower leaf-cells elongate,
pellucid, smooth, the upper rounded-quadrate or rounded-hexagonal, mostly
papillose on both sides: seta mostly long and straight; capsule exserted or
rarely immersed, erect, symmetric, short-necked; annulus none or deciduous
or remaining attached; peristome often none or rudimentary, when present of
16 perforate or upwards 2-3-cleft teeth upon a basal membrane, articulate;
operculum mainly obliquely rostrate, rarely conic-obtuse, sometimes not de-
ciduous; calyptra cucullate, papillose or smooth, usually falling away with the
operculum; spores large, variously papillose or pitted.
A genus of about 50 species widely distributed, on soil or soil-covered
rocks, mainly in the temperate zones; 12 species in North America, 1 species
in our region.
90 American Midland Naturalist Monograph No. 6
1. PoTTiA TRUNCATa [Hedwig] Fuernrohr
(P. truiicatiila Lindbsrg; GymnoUomum truncjtum Hedwig)
Plate XVII
Densely cespitose, dull green: stems simple or sparingly branched, erect,
about 2.5 mm high, radiculose at base; leaves numerous, the upper much larger
than the lower, obovate to oblong-spatulate, about 1.5-2.5 mm long, soft,
spreading, the margins plane, minutely crenulate with the projecting trans-
verse cell-walls, the leaves when dry become twisted, apex abruptly acute, costa
strong and excurrent into a short point; basal leaf-cells quadrate to rectangular,
large, lax, hyaline, more or less inflated, above becoming gradually smaller,
the median and upper medium- to thin-walled, smooth, hexagonal: seta erect,
about 3-4 mm high, mostly yellowish; capsule broadly oval or turbinate, erect,
symmetric, about 0.6-0.8 mm high, exannulate, more or less castaneous, when
dry and empty smooth and turbinate-hemispheric; lid broadly convex to flat-
tish with a beak about one-half as long as the urn; exothecial cells medium-
walled, castaneous, pellucid, irregularly quadrate to rectangular, the upper two
or three rows at the rim much smaller, rounded-quadrate, obscure; spores
orange-pellucid or brownish-pellucid, minutely punctulate, large, .026-030 mm,
mature from autumn to spring.
On moist soil in grasslands, along streams, etc., Europe, Asia, northern
Africa, and from Ontario to New England and Maryland, Pennsylvania,
and Michigan.
Rare in our region. McKean Co.: Corydon Street, Bradford. D.A.B. (figured).
12. Desmatodon Bridel
Autoicous: slender plants in mostly low, soft, green to yellow-green tufts,
dense to loose: stem mostly with central strand, thickly foliate, forking; leaves
when dry appressed and more or less plicate, when moist erect-spreading, cari-
nate to concave, obovate to ovate or lance-linear, mostly with recurved mar-
gins below, plane above, often serrate, sometimes margined; costa ending below
the apex or less often mucronately or aristately excurrent; leaf-cel's loose, thin-
walled, above rounded-quadrate or more or less hexagonal or rhomboidal,
mammillose to papillose, below rectangular and long-hexagonal, hyaline, smooth:
seta elongate, mostly straight; capsule erect, inclined, or even pendent, mostly
symmetric, ovate to cylindric; annulus persistent or falling away in pieces;
peristome inserted below the rim of urn, the basal membrane forming a tube
which is slightly exserted from the urn, thickly articulate, the 16 teeth rather
broad, divided to the base into two or three flat, filiform, papillose, divisions,
united here and there, usually twisted; lid stoutly and obliquely rostrate, with
the cells more or less spirally arranged; calyptra cucullate, smooth, long-
rostrate; spores large.
A. genus of 8 species, mainly on rich humus-soil in the mountains or on
mostly limestone rocks; one species in our re2ion.
Key to the Species
A. Costa of at least the upper leaves excurrent into a long smooth point
- (D. piimhobius)
Jennings: Manual of Mosses — 5. Pottiaceae 91
A. Leaves obtuse to short apiculate B
B. Leaves bordered with a band of lighter colored, elongate cells (D. Porferi)
B. Leaves not bordered L D. obtusijoLus
I. Desmatodon obtusifolius (Schwaegrichen) Jur.
{Desmatodon arenaceus Sullivant; D. ohioensis Schimper;
Didymodon areriaceus Kindberg)
Plate XVII
Gregarious to loosely cespitose, bright yellowish-green: stems short, in our
specimens about 3 mm. long, radiculose at base; leaves erect-spreading when
moist, crisped when dry, very small below but increasing to form a comal
tuft above, from ovate to lance-ovate, the comal 2-3 mm long, bluntly acute,
short-apiculate, the margin entire and more or less revolute; costa strong,
reaching the apex or extending into the apiculation; upper leaf-cells opaque,
incrassate, papillose, from rounded to hexagonal or quadrate, towards the base
of the leaf becoming elongate, thin-walled and hyaline: seta erect, 6-8 mm
high, sub-lustrous, sinistrorse, castaneous; capsule dark-castaneous, oblong,
the urn 2-4 mm long; peristome-teeth yellow, slender, divided almost to the
base into two slender, minutely-papillose prongs; annulus distinct, revoluble;
operculum bluntly and obliquely conic-hexagonal, immediately below the
annulus being smaller and incrassate; spores smoothish, yellowish, .01 2-. 01 5
mm, mature in spring.
On sandy soil, rocks, etc., mainly confined to the drainage-system of the
Ohio River.
Rather rare in our region. Crawford Co.: Linesville, May 12, 1908. O.E.J, (fig-
ured). Fayette Co.: Along river-bank at Ohio Pyle, September 1-3, 1907. O.E.J, and
G.K.J. McKean Co.: Near Bradford. December 15. 1894. D.A.B.
13. ToRTULA Hedwig
Autoicous or dioicous, rarely svnoicous or polyoicous: small to robust,
in green to brown tufts or cushions: stems mostly with a central strand, below
brownish- or red-radiculose, simple or branched; leaves mostly larger at the
ends of the shoots, often appearing rosette like, when dry not crispate but
somewhat twisted and contorted, when moist erect-spreading, mostly keeled,
cbovate or spatulate, rounded at the apex or rarely short-acuate, commonly
bordered, usually entire; costa strong, often cylindric, often mucronate-excur-
rent or, more commonly, excurrent into a hyaline hair-like awn; upper leaf-
cells rounded-hexagonal, loose, chlorophyllose, papillose, grading below into
the rectangular to elongate hyaline basal cells: seta long, erect; capsci'e erect,
cylindric, symmetric, short-necked, straight or sometimes slightly arcuate;
annulus present; peristome single, rarely none, basal membrane low to high,
teeth 32, filiform, equally spaced, mostly once to twice dextrorsely wound,
papillose and transversely striate, articulate; operculum conic, obliquely ros-
trate; calyptra cucullate, reaching to the middle of the urn; spores small.
A large genus of more than 200 species, widely distributed in the temper-
92 American Midland Naturalist Monograph No. 6
ate regions, mainly on calcareous rocks or soils; about 40 species occurring in
North America; only one species thus far reported in our region.
Key to the Species
A. Small; leaves when dry contorted and twisted; basal membrane low B
A. Medium to robust; basal membrane high and tessellated D
B. Teeth rather short, erect or slightly wound C
B. Teeth long, once to several times wound (T. muralis [Linnaeus] Hedwig)
C. Cells of leaf-margin not distinctly differentiated into a border
(T. plinthobia [Sull.] Broth.)
C. Cells distinctly differentiated at margin into a border
(T. Porteri [James and Aust.] Broth.)
D. On trees; leaves deeply concave, margins involute; costa spinulose-aristate
1. Tortula papulosa
D. On soil or stones; leaves not deeply concave; margin not revolute; costa smooth-
cuspidate (T. ruralis [Hedwig] Smith)
1. Tortula papillosa Wilson, mss.. Spring
(Barbula papillosa C. Mueller)
Plate LXII
Loosely cespitose, green, brownish in drying: stem short, up to 1 cm;
leaves erect-spreading, when dry appressed but scarcely twisted, broadly obo-
vate-spatulate, sometimes fiddle-shaped (panduriform), with margins involute,
the apex rounded to short-acute; costa thick and spongy, dorsally papillose,
above ventrally often bearing numerous shortly pedicellate multicellular
gemmae, excurrent-mucronate or cuspidate; basal leaf-cells rectangular, a few
hyaline, upper leaf-cells pellucid, incrassate, more or less collenchymatous,
large, ventrally smooth, dorsally simply papillose: capsule, known thus far
only from Australia and New Zealand, reddish-brown, short, with a short seta.
On tree-trunks (in America often on elms), rarely on rocks in open places.
South America, New Zealand, Australia, Europe, and, in North America,
ranging south in the Atlantic States to North Carolina and west to Michigan.
Rare and always sterile in our region. Blair Co.: Tyrone, T. P. James. (Porter's
Catalogue). Washington Co.: On bark of elm tree near Washington. Linn &: Simon-
ton. April 7, 1894 (figured) and March 16, 1894.
Family 6. Encalyptaceae
Autoicous, rarely dioicous: robust, usually densely cespitose, bright green,
the inside of the cushion rust-colored: stem 3-5-angled with little or no central
strand, erect, brown-radiculose, thickly-leaved, branched dichotomously; leaves
erect-spreading, when dry folded and twisted, more or less lingulate, acute to
obtuse, margins plane to undulate; costa highly developed, usually percurrent
to very shortly excurrent, prominent dorsally and dorsally papillose or toothed;
cells in upper two-thirds of leaf rather symmetrically hexagonal, chlorophyllose,
opaque, thickly papillose on both sides, in the lower third the cells much
larger, without chlorophyll, rectangular to rhombic, hyaline or slightly colored,
smooth, bordered by a few rows of narrow, elongate, and yellowish cells: seta
0.5-2.5 cm long, erect; capsule erect, symmetric, cylindric, smooth or plicate,
mostly with a short neck; annulus present; peristome varying from none to
Jenning: Manual of Mosses — 6. Encalyptaceae 93
well-developed, usually of 16 teeth; operculum from a conic base very long
and slenderly erect-rostrate; calyptra cylindric-campanulate ("extinguisher-
like"), long-rostrate, straight, completely enclosing the capsule, the border
fringed; spores large and papillose.
A world-wide family mostly on soil and rocks, occurring in the tropics,
however, only on the higher mountains. At least 30 species; 2 in our region.
The family embraces but one genus, with characters as given for the family.
1. Encalypta Schreber, Hedwig
Key to the Specihs
A. Monoicous: no gemmae; peristome single; capsule smooth; leaf-apex with distinct
point 1. E. ciliata
A. Dioicous: clusters of slender brown gemmae in axils of leaves: peristome double;
capsule spirally striate; leaf-apex obtuse to bluntly mucronate 2. E. streptocarpa
1. Encalypta ciliata Hedwig
(Leersia laciniata Hedwig; Leersia ciliata Hedwig)
Loosely cespitose, bright green: stems branched, 1-2.5 cm high, densely
radiculose below; leaves large, broadly obovate-oblong to Ungulate, rounded
at ape.x, apiculate, plane-margmed and narrowly recurved below, spreading
when moist, crisped and incurved when dry; costa yellov.'ish, ending just below
apex or percurrent; basal cells lax, hyaline, rectangular, walls red, the marginal
paler and narrower in several rows, the upper leaf -eel Is opaque, densely papil-
lose, hexagonal-quadrate, the walls pellucid, cells about .015 mm across: seta
long, erect, yellowish to reddish, dextrorse; capsule cylindric, reddish-brown,
smooth, constricted below mouth and smooth when dry, at base abruptly
tapering into the seta; peristome single, the teeth 16, lanceolate, reddish, in-
serted below the rim, irregularly divided in some specimens, papillose, articu-
late, strongly incurved when moist; annulus none; exotherial cells smaller in
several rows at the rim; lid erect, nearly as long as urn, narrow, slenderly ros-
trate-clavate; calyptra straw-colored, mitrate, cylindric, slenderly rostrate, ex-
tending below the base of capsule, the lower margin fringed with a row of
narrowly lanceolate teeth; spores roughened, mature in late summer or early
fall: autoicous.
In crevices or shaded places on rocks and walls, almost cosmopolitan in
mountainous or hilly regions; in North America from the Arctic region south
to the northern United States. Reported from the adjacent states of New
York and Ohio and to be expected from the northern part of our range.
2. Encalypta streptocarpa Hedwig
(E. contort a Lindberg)
Extinguisher Moss
Plate LXII
Densely cespitose, dull or yellowish-green; stems up to 3 to 6 or 7 cm
high, branched, densely radiculose at base; leaves rather crowded, spreading,
when dry twisted and crisped, large, 5-6 mm long, oblong-lingulate, sometimes
94 American Midland Naturalist Monograph No. 6
narrowed in the middle, more or less undulate, plane-margined, often incurved
and sub-cucullate at apex; costa strong,- reddish, ending just below apex, dor-
sally scabrous; basal leaf-cells hyaline, rectangular to elongate rectangular,
1-4:1, the marginal often forming a distinct border of a few rows of linear-
elongate cells, upper leaf-cells hexagonal-quadrate, about .015 mm, multi-
papillose, incrassate-pellucid, usually regularly seriate; perichaetial leaves
oblong, abruptly long lanceolate-acuminate: seta long, red; capsule long, cylin-
dric, dextrorsely orange-striate, furrowed when dry; peristome double, the teeth
filiform, papillose, articulate, red, the inner peristome of 16 or 32 filiform pale
segments half as long as the teeth and adherent to the latter by the broad
puncticulate basal membrane; lid narrow, long, rostrate, erect; calyptra very
long, cylindric, scabrous at the tip of the long and slender beak, extending
considerably below the capsule and laciniate at the border; spores mature in
late summer; dioicous.
On rocks, walls, earth, etc., usually in crevices on calcareous substrata,
mainly confined to rough country. Europe, Asia, and from Ontario and Vir-
ginia to British Columbia. Not yet found fruiting in North America.
Rare in our region. Cambria Co.: Cresson. T. P. James. (Porter's Catalogue).
NX'estmoreland Co.: In limestone rock on Chestnut Ridge, 2 miles east of Hillside,
Charles M. Boardman, May 30, 1936 (figured).
Family 7. Grimmiaceae
Autoicous or dioicous: cespitose, dark green to blackish: stem mostly
without central strand, radiculose only at the base, branches mostly of equal
height, leaves often hyaline-pointed, often piliferous, but rarely crispate, when
damp more or less spreading, rarely secund, mostly lanceolate, rarely and then
only upwards denticulate, towards the apex and at the margin two to several
cells thick, sometimes papillose; costa complete or nearly so; cells small, often
sinuate-walled; above mostly rounded-quadrate, towards the base inflated and
mostly pellucid, rectangular to elongate, rarely linear throughout the whole
leaf: seta rarely shorter than the capsule; capsule mostly symmetric, globose
to cylindric, mostly smooth, often immersed or emergent; annulus present, or
none at. all; teeth 16, mostly separate to the insertion, red to orange, papillose,
plane, undivided or cleft or cribrose, rarely divided to the base into filiform
parts, trabeculae mostly projecting only outwards; operculum mostly rostrate,
sometimes deciduous with the attached columella; calyptra mostly small,
mitrate or cucullate, glabrous, sometimes campanulate and plicate.
A large family, world-wide in distribution, but most abundant in sub-arctic
and temperate regions, mostly on stones or rocks, rarely on soil or trees. In
Grout's Moss Flora the Hedwigiaceae are included as a sub-family of the
Grimmiaceae.
Key to the Species
.■\. Costa with basal guides, or homogeneous; calyptra rarely campanulate; spores small
to medium-sized B
A. Costa with several median guides; spores small; calyptra campanulate; basal leaf-
1. Ptychomitrium
Jennings: Manual of Mosses — 7. Grimmiaceae 95
B. Teeth undivided, cribrose, cleft in upper half, or none; branches! as high as the
stem; basal leaf-cells mostly smooth-walled 2. Grimmia
B. Teeth divided almost to the base into two filiform divisions; branches irregular
short; basal leaf-cells with nodulose or sinuose walls 3. RhacomitTtum
1. Ptychomitrium Fuernrohr
Autoicous: cespitose in loose yellowish-green to brownish or blackish
cushions; stem with central strand, erect or ascending, radiculose at the base,
thickly-leaved; leaves long, narrow, the points not hyaline, crispate when dry,
spreading when moist; costa strong, percurrent or ending below the apex; cells
not with sinuose walls, smooth, upwards small and rounded-quadrate, below
linear to more or less loosely rectangular; perichaetial leaves not sheathing: seta
straight, more or less elongate, mostly two or more to a perichaetium; capsule
smooth, erect, symmetric, mostly oval to oblong-elliptic; annulus wide, decidu-
ous, rarely none; peristome inserted below the mouth; teeth 16, papillose,
usually deeply divided into two subulate prongs, trabeculae more or less dis-
tinct; spores small; operculum conic with a long, fine, straight beak; calyptra
campanulate, plicate and lobed, reaching about halfway down the capsule.
A widely distributed genus of 62 species, of which at least 9 occur in
North America and one in our region. Occurring on rocks and stones, —
rarely on trees.
1. Ptychomitrium incurvum (Muhlenberg) Sullivant
Plate LXIII
Densely cespitose, dark green to brownish: stems about 5 mm high, erect;
leaves erect-spreading when moist, sometimes incurved, twisted-crispate when
dry, the lower small, increasing in size upwards, linear-lanceolate, obtuse, thick,
opaque, the margin plane; costa broad, ending in apex; basal leaf-cells rectangu-
lar, pellucid, the upper m.uch smaller, rounded to quadrate, incrassate, dense;
seta about 2-3 mm high, erect; capsule erect, oval; peristome-teeth 16, long-
subulate, articulate, papillose; lid erect, conic-subulate, about as long as urn;
calyptra long-rostate, mitrate, plicate-lobed to base of beak, covering a little
more than half of the urn; spores mature in spring.
On more or less exposed calcareous rocks from Connecticut to Georgia
and Texas. Not uncomm.on in eastern Pennsylvania, northern Ohio, and
western New York.
Westmoreland Co.: On limestone rock, l'/2 miles east of Hillside. Chestnut Ridge,
C.M.B. May 30. 1936 (figured).
2. Grimmia Ehrhart, Hedwig
Autoicous or dioicous: forming cushions and mats, slender, often hoary
by reason of the hyaline leaf-apices: stem erect or ascending, mostly with a
central strand, radiculose mainly at the base, thickly-leaved; leaves imbricate
when dry, rarely crispate or spirally appressed, spreading to recurved-squarrose
when moist, lower often small and bract-like, the upper often suddenly larger,
often hyaline-piliferous, carinate, concave, sometimes canaliculate, mostly
96 American Midland Naturalist Monograph No. 6
lanceolate from an oblong or ovate base, acuminate, entire, margins plane or
revolute; costa complete or extending to the base of the piliferous apex; upper
cells small, rarely papillose, rounded-quadrate, often opaque, looser towards
the middle, the basal linear to rectangular, mostly smooth-walled, and some-
times forming a colored border; perichaetial leaves mostly larger, more or less
sheathing, areolation looser: seta sometimes shorter than the capsule, rarely
much longer than the perichaetial leaves, arcuate or straight, mostly yellow,
twisted when dry, capsule mostly symmetric, smooth to ribbed, globose to
cylindric; annulus persistent or curling off, sometimes none; peristome rarely
absent, when present inserted below the mouth; teeth reddish-brown, broad to
subulate, entire to cribrose, sometimes cleft to the middle, the trabeculae pro-
jecting outwards; operculum often rostrate, never longer than the urn; calyptra
lobed-mitrate to cucullate, long-rostrate, smooth; spores small, .010-. 012 mm
diameter.
A large genus of world-wide distribution, but mainly confined to the moun-
tains of the tropics, occurring on rocks and stones. About 230 species, of
which more than 90 occur in North America and at least 4 in our region.
Key to the Species
A. Seta not longer than the capsule; operculum mostly falling with the columella still
attached; hyaline leaf-points decidedly spinulose B
A. Seta longer than the capsule F
B. Apices of upper leaves with short hair-points C
B. Apices of upper leaves long-piliferous E
C. Lower and median leaf-cells with sinuose walls 4. G. laevigata
C. Lower leaf-cells not with conspicuously sinuose walls D
D. Slender small plants in dense cushions; central strand in stem; teeth decidedly
cribrose or cleft 2. G. confertj
D. More robust, loosely cespitose; with indistinct central strand; teeth slightly cribrose
or entire - 1. G. apocarpa
E. Leaf-cells without sinuose walls; capsule oblong; calyptra often cucullate,
( G. ambigua [Sullivant] Sullivant )
E. Leaf-cells with somewhat sinuose walls; capsule oblong-ovate 3. G. pilifera
F. Capsule distinctly ribbed; seta curved; leaf-margins of a single layer of cells
(G. Olneyi Sullivant)
F. Capsule smooth, seta straight; upper leaf-margins of more than one layer of cells ....G
G. Leaves lanceolate, tapering; basal leaf-cells thin-walled and elongate-rectangular,
about 1:4 to 1:8, alpine (G. obtusa Schwaegrichen; G. Doniana Smith)
G. Leaves ovate to oblong-lanceolate, the long apex rough, piliferous; basal leaf-cells
more or less quadrate 4. G. laevigata
1. Grimmia apocarpa [Linnaeus] Hedwig
Plate XVIII
Loosely cespitose, more or less erect, branching rather freely, about 2 cm
high, dull olive-green, drying stiff and ncn-crisped: leaves ovate-lanceolate,
erectinp-spreading, strongly costa te and usually more or less carinate, margin
narrowly recurved, apex narrowly obtuse, leaves about 2 mm long; perichaetial
leaves similar but somewhat longer and thinner; costa ending in or just below
the apex, terete dorsally; basal cells rectangular, about .008-010 x .015.030 mm,
upper basal cells quadrate, and in our specimens usually sinuose-walled, the
Jennings: Manual of Mosses — 7. Grimmiaceae 97
median and upper cells rounded and .005-. 009 mm in diameter, all cells in-
crassate and more or less opaque: seta erect, stout, about 0.5 mm long; capsule
immersed, oval-oblong, about 1 mm long, reddish-brown, rather thick-walled,
smooth; calyptra short, lobed; operculum low-conic, rostrate; peristome single,
teeth 16, lance-linear, trabeculate, somewhat cribrose, reddish-brown, faintly
papillose, when dry reflexed-revolute; spores reddish-brown, in our specimens
about .012-. 018 mm in diameter; columella falling away with the operculum
and remaining attached to it; spores mature in late spring.
On stones, hard earth, etc., with a wide distribution over the colder regions
of the earth. In America occurring from Alaska and Newfoundland to the
Northern States and south in the mountains to Georgia.
G. apocarpa var. graciln (Schleicher) Weber and Mohr occurs on rocks
from lower Canada to West Virginia and Tennessee. It grows in loose, slender
mats up to 10 cm long; leaves lanceolate, acuminate, 1.5-2 mm long.
Now known from the following counties: Butler. Erie, Fayette, Greene, McKean
(Porter), Washington (Porter), and Westmoreland. Specimen figured: Shaly bank of
stream. Shades Ravme, east of TrafFord, Westmoreland Co., March 25, 1910. O.E.J.
2. Grimmia conferta Funck
(G. apocarpa var. conferta (Funck) Sprengel)
Densely cespitose, in gray-green rounded cushions: stems slender; leaves
1 mm or less long, keeled, margins revolute, lance-ovate to oblong, acuminate,
opaque, apex hyaline, denticulate; costa strong, dorsally prominent, ending at
apex; basal leaf-cells rectangular to quadrate, the upper smaller and rounded,
all incrassate and dense: seta short; capsule immersed, ovate-globose, wide-
mouthed, hemispheric and somewhat wrinkled when dry; peristome-teeth light
reddish-brown to orange, fragile, markedly cribrose; annulus said to be none;
lid wide, low-convex, apiculate; spores mature in spring.
On rugged exposed rocks, Europe, Asia, and Africa, and, in North Amer-
ica, from Nova Scotia to British Columbia south to Idaho and North Carolina.
Washington Co.: Linn and Simonton. (Porter's Catalogue).
3. Grimmia pilifera Beauvois
{^Grimmia pennsylyanica Schwaegrichen)
Plate LXIII
Densely cespitose, dark green: stems 1 to 3 cm high, robust, rigid, branch-
ing; leaves close, narrowly ovate-lanceolate, acuminate, concave, the hyaline
point usually distinctly spinulose; margins somewhat recurved below and
thickened above; basal leaf-cells linear-rectangular 3-6:1, thin-walled, hyaline
to yellow-pellucid, shortly above base the cells incrassate-sinuous, short-r-rctan-
gular, the upper rounded-quadrate to hexagonal, small, piliferous: seta about
half as long as urn, capsule more or less completely immersed, oblong-ovate,
smooth, even when dry, lid conic-rostrate, about three-fifths as long as urn,
erect; annulus large; peristome-teeth large, broadly lanceolate, irregularly split
and cribrose to about the middle, castaneous pellucid; calyptra lobed, mitrate;
spores mature in the autumn but often not shed till spring: dioicous.
98 American Midland Naturalist Monograph No. 6
On moist rocks in woods, Japan and, in North America, from Nova
Scotia to Georgia and Minnesota, and in Mexico. Reported from Painesville,
Ohio (W. C. Werner) and in Pennsylvania.
Fayette Co.: Meadow Run, Ohio F*yle. June 23. 1940. C. M. Boardman (figured).
Washington Co.: On sandstone rock on high hill, near Grier's Station. A. Linn and
J. S. Simonton. Oct. 5, 1898.
4. Grimmia laevigata (Bridal) Bridel
(G. campestjis Burch.; G. leucophaea Greville)
Cespitose loosely in wide, dull gray-green tufts; hoary above: stems stout;
leaves close, larger towards top of stem, when dry imbricate-appressed, very
concave, oblong-oval to rather widely ovate, 1-1.5 mm long, plane-margined,
at the apex abruptly terminating in a hyaline, flanened, finely denticulated
hair; the smaller lower leaves acuminate but without the hair-point; costa nar-
row, ending in the apex; basal leaf -cells quadrate, except a few rectangular ones
near the costa, the upper smaller and rounded, all incrassate, non-sinuose, the
upper quite chlorophyllose: seta erect; capsule included, or emergent, 1-2 mm
long, elliptic, broadly oblong, brownish smooth when dry; annulus large; lid
conic-rostellate, short, peristome-teeth cleft to about the middle, cribrose below,
castaneous-pellucid; calyptra mitrate, lobed; spores mature in spring.
On rocks, mainly non-calcareous, often granite or sandstone, almost cos-
mopolitan. In North America from New York and Pennsylvania to Alabama
and northwestward to Oklahoma; also in the west. Rare in our region.
Blair Co.: Tyrone, T. P. James. (Poner's Catalogue).
3. Rhacomitrium Bridel
Dioicous: robust plants, loosely and widely cespitose, the mats green to
yellowish or blackish-green: stem without central strand, procumbent to erect,
radiculose at the base only, uniformly foliate, often with numerous short
branches giving the shoot a nodose appearance; leaves spreading to recurved-
spreading or sometimes secund, when dry appressed, from an ovate to oblong
base mostly lanceolate to lance-linear, more or less long-acuminate, often
piliferous, sometimes lingulate and obtuse, margins sometimes 2-layered and
sometimes recurved; costa mostly broad flat, and complete; cells nearly all with
sinuose or nodulose walls, often papillose, towards the base or sometimes all
over linear: seta long, straight, rarely curved, twisted; capsule erect, oblong to
cylindric, narrow-mouthed, smooth: annulus broad, curling off; teeth united
at the base but mostly cleft deeply into 2 (-3-4) filiform divisions, often very
long, trabeculate; spores small; operculum conic with a long subulate apex
from one-third to more than the length of the urn: calyptra mitrate, lobed,
not folded, subulate-rostrate, glabrous or rough.
A world-wide genus ol: about 90 species, mostly on siliceous rocks: about
10 in North America; probably 2 species in our region.
Key- to the Species
A. Upper leaf-cells quadrate, lower ones linear; shoots not appearing nodose by arrangj-
ment of short lateral branches B
Jennings: Manual of Mosses — 7. Grimmiaceae 99
A. Upper leaf-cells elongate; shoots appearing nodose with short lateral branchlets D
B. Leaves lingulate; leai-apex broad and rounded 1. R. aciculare
B. Leaves lanceolate-acuminate C
C. Leaf-apex hyaline-acuminate 2. R. microcarpum
C. Leaf-apex obtuse, not hyaline 3b. R. heterostichum var. gracilescens
D. Leaf-apex acute, not hyaline; seta 4-5 mm long (R fasciculart)
D. Upper leaves with hyaline points; seta 4-10 mm long
3a. R. heterostichum var. ramulosum
1. Rhacomitrium aciculare [Linnaeus] Bridel
(Dicranum aciculare Hedwig)
Plate XVIII
Cespitose in coarse tufts, dark dull green to blackish; stems long, up to 6
or 8 cm long, stout, branching by short innovations, radiculose below; leaves
erect-spreading, stiffly imbricate when dry, large, up to 1 mm wide by 2.5 mm
long, broadly ovate-oblong, usually somewhat plicate at the base, the margin
usually revolute, the ape.x broadly obtuse, denticulate to entire, non-hyaline,
the upper margin usually thickened; costa strong, ending below apex; leaf-cells
densely yellowish-pellucid, incrassate, sub-papillose, the upper sub-quadrate to
rounded-hexagonal and in two layers at the margin, the basal elongate-rectan-
gular to linear, markedly sinuose-incrassate, at the margin sub-quadrate, the
alar a little larger, quadrate: seta erect, straight, about 10-12 mm long; capsule
dark brown, erect, oblong-cylindric to elliptic, smooth, with narrow mouth;
peristome-teeth cleft to below the middle into 2 or 3 unequal divisions; lid
aciculate-rostrate or subulate, almost as long as urn; calyptra smooth, long-
rostrate, mitrate, lobed, covering only the top of capsule; annulus rather large,
revoluble; spores mature in spring; fruit rarely found.
On wet, shaded, non-calcareous rocks in hilly or mountainous country, in
Europe, Africa, and, in North America, from Alaska and Labrador south to
California and Alabama. Occurs in northern West Virginia and Pennsylvania.
Now known from Butler, Cambria (Porter), Clearfield, Fayette, Somerset, and West-
moreland counties. Specimen figured: Ohio Pyle, Fayette Co., in crevices of rock along
the river near the falls. Sept. 1-3, 1906. O.E.J. &C G.K.J.
2. Rhacomitrium microcarpum (Hedwig) Bridel, not Schrader
{Rhacomiirtum sudeticiim Bryologia Europaea; Trichostomum microcarpum
Hedwig; R. heterostichum var. sudeticnm ( Funck )
Loosely cespitose, dull or yellowish-green above: stem slender with ascend-
ing branches, 2-5 cm high; leaves divaricately spreading, erect when dry, nar-
rowly lanceolate, linear-acuminate, apex hyaline, denticulate, margin more or
less bistratose above, usually revolute below; costa strong, ending in apex;
basal leaf-cells linear, sinuose, incrassate, upper leaf-cells rounded-quadrate: seta
short, light-colored, often curved or flexuose; capsule relatively very small,
elliptic, pale brown, thin-walled; annulus large, revoluble; peristome-teeth
brownish, irregularly divided; lid conic-rostrate, shorter than the urn; spores
mature in spring.
On wet rocks or cliffs, principally granite or gneiss, Europe, Asia, and, in
100 American Midland Naturalist Monograph No. 6
North America, from Greenland to British Columbiaj^ south to Oregon and
northeastern United States. Perhaps to be expected in the eastern or north-
eastern part of our region.
3. Rhacomitrium heterostichum var. ramulosum (Lindberg) Jones
(R. ericoides (Schwaegrichen) Jennings; Trichostemum ericoides Schwaegrichen )
Cespitose in low, green to yellow-green tufts: stems slender, markedly
nodose with obtuse lateral innovations; leaves crowded, spreading to falcate-
secund, lanceolate from an ovate base, acuminate, the apex hyaline, flat, den-
ticulate, not very narrow, margin revolute, not thickened; costa strong, ending
in the apex; leaf-cells all linear, the upper 3-6:1, incrassate, sinuose, the mar-
ginal shorter and in the alar portion often a few larger, rectangular, pellucid,
and not sinuose-walled: seta yellowish, short; capsule small, elliptic-cylindric
to oblong, pale yellow-brown, thin-walled; annulus large, revoluble; peristome-
teeth short, divided almost to the base; calyptra somewhat papillose at the
apex; lid short-rostrate; spores mature in spring.
On exposed rocks and stones in hilly or mountainous regions, Europe, and,
in North America, from Greenland to British Columbia south to Oregon,
eastern Pennsylvania and Georgia. Perhaps will be found in the eastern part
of our region.
3b. Rhacomitrium heterostichum var. gracilescens Bry. eur.
Plate LII
When dry forming crisped, blackish-green, dense tufts about 1 cm deep,
the prostrate stems 1-3 cm long and sending up short, irregularly spaced
upcurvied branchlets which, when dry have closely appressed leaves and are
less than 1 mm thick; when wet the leaves spreading-ascending or somewhat
recurved- spreading; leaves about 2 mm long, concave, from a broader base
slenderly lanceolate to a narrow, obtuse, entire apex, the base sub-clasping, the
margins entire, thickened and imperfectly bi-stratose, narrowly revolute; costa
50-60 jW, thick, prominent dorsally, percurrent into the somewhat cucullate leaf-
apex, bi-stratose with usually 2-3 smaller interior cells; basal cells enlarged,
somewhat colored in a narrow basal band, with a few somewhat larger rounded
alar cells, above these the cells rapidly becoming strongly incrassate and sinu-
ose, the median lower about 3-3.5 by 8-10 /<,, these at the apex roughly rounded
and about 2.5/j, in diameter. Some of the upper cells appear faintly papillose.
The specimen here described and figured was collected by Charles M.
Boardman on rocks at Beck Spring, Laurel Ridge, Somerset Co., s.w. Pa.,
July 26, 1947, and seems best placed under the polymorphic R. heterostichum,
nearest to var. gracilescens, although were the cells papillose it might as well be
referred to R. protensum Braun. The variety gracilescens is a very rare alpine
moss found in the U. S. on Bald Mountain, near Camden, Maine.
Family 8. Orthotrichaceae
Dioicous or autoicous, rarely heteroicous or polyoicous: cespitose, light
green to yellowish- or blackish-green outside the tufts, inside brown to black:
Jennings: Manual of Mosses — 8. Orthotrichaceae 101
stem mostly with no central strand, erect to ascending, or creeping with erect
or ascending branches, radicuiose below, or along the creeping stem, with
reddish or brownish filam.ents; leaves spreading to squarrose, carinate, mostly
from a decurrent base more or less lanceolate, sometimes oblong-lingulate to
linear, mostly entire, usually papillose both sides; costa strong, sometimes
excurrent-aristate or piliferous; upper cells generally rich in chlorophyll, round-
quadrate to round-hexagonal, basal mostly elongate rectangular to linear, pellu-
cid; perichaetial usually more or less sheathing: seta erect; capsule erect, sym-
metric, collum distinct, oval to pyriform or cylindric, smooth or striate, deeply
plicate when dry and erripty; annulus persistent; peristome mostly double,
rarely none; teeth 16 united or approximate in 8 pairs, lanceolate, flat, whitish
to yellowish or reddish, reflexed when dry, exteriorly papillose or transversely,
obliquely, or longitudinally striate, rarely smooth, rather delicately trabeculate;
inner peristome of 8 or 16, filiform, or lanceolate; glabrous or papillose seg-
ments alternating with the teeth; spores small to very large; operculum long-
rostrate; calyptra cucullate and sometimes smooth to campanulate, often plicate
and hairy.
A rather large family of mostly tree-inhabiting species, mostly of tem-
perate regions.
Key to the Genera
A. Calyptra cucullate, not plicate; stems erect or creeping B
A. Calyptra mostly campanulate, plicate; stems mostly erect C
B. Stem erect: peristome none; leaves crispate when dry 1. Amphidium
B. Stem creeping with erect or ascending branches: peristome single 2. Drummondia
C. Leaves when dry crispate, at the base ovate and mostly with a hyaline border; cap-
sule exserted in our species 4. Ulota
C. Leaves not ovate at the base nor with hyaline border: capsule immersed or emergent
in our species 3. Orthotrichum
1. Amphidium (Nees) Schimper
{Amphoridium Schimper; ^ygodon Authors)
Autoicous or dioicous: tufts cushion-like, soft, often extensive, yellowish
olive-green to blackish, inside rusty brown: stem furcately branching, uniformly
foliate, radiculose to the apex with smooth filaments; leaves linear-lanceolate,
papillose both sides, when dry contorted or crispate; costa practically complete,
with median guides; leaf-cells thick-walled, angular to roundish, green, towards
the base generally elongate, rectangular, thin-walled to hyaline; perichaetial
leaves erect, longer, sheathing or half-sheathing: seta short, generally erect,
thickening above and grading into the long collum; capsule emergent to ex-
serted, mostly erect, pyriform, with 8 projecting, reddish-brown ribs, when
empty much widened at the mouth and urceolate; annulus none; peristome
none; operculum obliquely rostrate froin a low-conic base; calyptra cucullate,
glabrous, not plicate.
A world-wide genus of 12 species, on mainly non-calcareous rocks; 5 species
in North America; 2 in our range.
102 American Midland Naturalist Monograph No. 6
Key to t^e Species
A. Autoicous: leaf -margins plane: seta 1.5 mm long or less; beak of operculum shorter
than radius of capsule 1. A. lapponicum
A. Dioicous: leaf-margins recurved below: seta about 3 mmi long; beak of operculum
longer than radius of capsule 2. A. Mougeotii
1. Amphidium lapponicum (Hedwig) Schimper
(Anictangium lapponicum Hedwig; Z.ygodon lapponicus Bryologia Europaea)
Densely cespitose, dark olive-green above, blackish below: stems dichoto-
mously branching, usually 1-3 cm high; leaves flexuous-spreading, lance-oblong
to lance-linear, 1.5-2.0 mm long, margins plane, acute, crisped when dry; costa
strong, ending below apex; basal cells large, thin-walled, pellucid to hyaline,
the upper small, quadrate-hexagonal, rather obscure, incrassate, papillose; peri-
chaetial leaves sheathing: seta very short, usually not over 1.5 mm; capsule
oval with distinct collum about as long as sporangium, partially immersed,
brownish, when dry constricted below mouth, urceolate, with 8 reddish striae,
the mouth forming a deep red, thickened rim; no peristome; lid red, lustrous,
small, low-conic, obliquely rostellate, the length of the beak not more than
one-half the diameter of the capsule; calyptra cucullate, small, reaching about
half-way down the urn, brownish; spores mature in early summer: autoicous,
the antheridial flowers axillary along the stem.
In crevices of shaded rocks, rarely in calcareous habitats; Europe, Asia,
and, in North America, from Greenland to British Columbia south to northern
United States and to California.
Rare in our region. Huntingdon Co.: Porter. (Porter's Catalogue).
2. Amphidium Mougeotii (Bryologia Europaea) Schimper
{Z.ygodon Mougeotii Bryologia Europaea; Anoectangiitm Mougeotii Lindberg)
Densely cespitose, in large tufts, yellowish-green above, rusty to blackish
below: stems usually 2-6 cm high, dichotomously branching; leaves spreading
or erect-spreading, crisped when dry, elongate lance-linear, acuminate, carinate,
margin narrowly revolute below, slightly irregular above; costa strong, vanish-
ing at apex; basal leaf-cells narrowly rectangular, rather thick-walled, above
shorter, sub-quadrate to rounded, incrassate, hardly papillose, pellucid; peri-
chaetial leaves sheathing only the base of seta: seta about 2-3 mm long; cap-
sule shortly exserted, narrowly oval with a distinct neck, when dry urceolate,
only slightly contracted belov/ mouth, 8-striate: peristome none; lid low,
obliquely rostrate, the length of the beak at least one-half the radius of the
capsule: calyptra cucullate; spores mature in summer or early autumn: dio-
icous: fruit rare.
On damp, shaded, usually non-calcareous rocks, in mountains or hilly re-
gions; in Europe, Asia, and in North America, from Newfoundland to Alaska
south to Alabama and Oregon. To be looked for in the eastern part of
our range.
Jennings: Manual of Mosses — 8. Orthotrichaceae 103
2. Drummondia Hooker
Autoicous or dioicous: slender, in low, dense, green, scarcely shining, often
extensive mats: stem long, creeping, brown-radiculose, thickly covered with
short, erect, simple or furcate branches; leaves when dry stiffly appressed, when
moist erect-spreading to spreading, ovate-lanceolate to linear-oblong, acute or
obtuse, entire; costa strong, almost percurrent; cells uniformly rounded, smooth,
chlorophyllose: seta erect, long; capsule erect, symmetric, oval, smooth, when
dry shriveled; annulus none; peristome simple, inserted below urn-mouth; teeth
16, very short, truncate, entire, smooth, densely trabeculate; spores very large
(.08-. 10 mm); round or oval, several-celled, green, smoothish; operculum
obliquely rostrate; calyptra cucullate, large.
A small genus of 7 species, on trees, rarely on rocks; mostly Asiatic, one
in our region.
1. Drummondia prorepens [Bridel] Jennings
(Anodontiitm prorepens Bridel; Gymnoslomurn proreper^s Hedwig; Hypnum
clavellatum DiUenius; Orthotrtchuin clavellatum Hooker)
Plate XVIII
Stems creeping, radiculose on the under side, with numerous short, erect
branches, 6-10 mm, forming dark green or blackish tufts: leaves erect to
spreading, oblong to ovate-lanceolate, 1 to 1.5 mm long, obtuse to acute, con-
cave, carinate, firm; costa strong, almost percurrent; cells small, thick-walled,
rounded; the alar often quadrate-inflated and hyaline in the stem-leaves, the
whole lower fourth of the perichaetial leaves elongate-rectangular and hyaline:
seta erect, about 2 5 mm. long, sinistrcrse; capsule ovate-globose, smooth, about
1 mm high; operculum low-conic, obliquely long- rostrate; peristome of 16 very
short, wide, truncate, smooth, trabeculate teeth, often more or less confluent;
annulus none; calyptra, at first conic, large, cucullate; spores minutely rough-
ened, chlorophyllose, about .080-095 mm, moderately incrassate, mature in
summer.
On tree-trunks in woods, Japan, and in North America from New England
to Alabama, Missouri, and Ontario.
Erie Co.: Presque Isle. May 8-9, 1906. O.E.J, (figured). McKean Co.: Quin-
tuple, Bradford, November 10, 1893. D.A.B. Washington Co.: Linn and Simonton.
(Porter's Catalogue).
3. Orthotrichum Hedwig
Autoicous, rarely dioicous: cespitcse in cushions sometimes on rocks,
mostly on trees: stems erect and ascending, radiculose at the base, thickly
leaved, branched; leaves when dry never crispate but straight and appressed,
ovate-or linear-lanceolate, mostly acute, margins usually revolute; costa quite
strong, mostly not quite percurrent; basal leaf-cells rectangular to elongate,
pellucid to hyaline, the marginal often shorter and green: seta generally shorter
than the scarcely or non-sheathing perichaetial leaves; capsule oval to cylindric,
usually with 8 or 16 colored striae, when dry usually 8 (-16) -furrowed; an-
nulus persistent; peristome mostly double, sometimes single, rarely none,
usually with 16 broadly lanceolate teeth in pairs, and 8 to 16 filiform seg-
104 American Midland Naturalist Monograph No. 6
ments; operculum conic to convex, rostrate; calyptra campanulate, plicate,
covering most of the urn, glabrous, hirsute or papillose.
A cosmopolitan genus of about 200 species, on trees or rocks, rare, however,
in the Tropics; about 60 species occur in North America; at least 5 in our
region.
Key to the Species
A. Peristome simple, teeth 16, erect or erect-spreading when dry B
A. Peristome double, teeth more or less reflexed when dry D
B. Capsule half-emergent, 16-striate (O. cupulatum [Hoffmann] Schwaegrichen)
B. Capsule immersed to half-emergent, 8-striate C
C. Capsule ovate-cylindric, half-emergent when leaves are dry 1. O. strangulatum
c. Capsule much shorter, ovate-globose, practically immersed in the dry leaves
2. O. Lescurii
D. Capsule smooth when dry, immersed E
D. Capsule plicate when dry F
E. Teeth 16, in pairs; segments 8, short, filiform ( O. pusillum Mitten)
E. Teeth 16, not in pairs; segments 16, comparatively broad
(O. elegans Hooker & Greville)
F. Leaves more or less obtuse at the extreme apex G
F. Leaves usually distinctly acute at the extreme apex !
G. Capsule strongly contracted below thg mouth when dry and very decidedly plicate
with reddish-brown folds 3. O. stellatum
G. Not very strongly contracted nor very decidedly plicate H
H. Stomata immersed; leaf-margins revolute 4. O. ohioense
H. Stomata not immersed; leaf-margins erect; leaves short and broad, oblong-ovate to
lingulate, obtuse, usually with brood-bodies 6. O. obtusijolium
I. Capsule but little contracted below mouth when dry, ribs orange, segments 8; leaves
oblong-lanceolate 5. O. pumilum
I. Capsule strongly contracted under the mouth when dry J
J. Capsule with very prominent reddish-brown ribs when dry, half-emergent
3. O. stellatum
J. Capsule with less prominent light colored ribs, usually immersed (or emergent)
(O. sard dum Sullivant)
L Orthotrichum strangulatum Schwaegrichen
(O. Porteri Austin; O. cupulatum var. Porteri Venturi)
Plate XVIII
Densely cespitose, about 1 cm high: stems densely leaved, branched; upper
leaves lanceolate, about 3 mm long, the lower ovate, shorter, acute, margins
entire or papillose, more or less revolute, lamina often somewhat bi-stratose
at margins and apex; costa strong, almost percurrent; basal leaf-cells quadrate
at margin to rectanoular (2:1) towards costa, smooth, hyaline, the median
rounded-hexagonal, dense, papillose, becoming towards apex densely incrassate-
rounded, arranged in quite regular rows: seta short, about 0.5 mm, capsule
about 1.5 mm long, oblong-cylindric, tapering rather gradually to the seta,
when dry often only partly immersed, when wet always immersed, dark reddish-
brown, deeply 8-costate and 8-furrowed, the costae with about 3 rows of rectan-
gular cells with thicker longitudinal walls, stomata few, immersed, calyptra
mitrate, quite densely erect-hirsute; operculum low with a rounded apiculation;
peristome single, teeth paired, papillose, erect-spreading when dry, irregularly
Jennings: Manual of Mosses — 8. Orthotrichaceae lO'?
triangular-lanceolate, about 8-articulate, the divisural distinct and the teeth
often split; the three or four upper rows of cells of the capsule densely incras-
sate, brown, pellucid like the costal and laterally oblong; spores globose, papil-
lose, .01 7-. 020 mm, mature in summer.
On rocks, mostly limestone, from Alabama to Missouri, Minnesota, and
New England.
Rare in our region. Cambria Co.: Cresson. James. (Porter's Catalogue). Centre
Co.: On limestone 2 mi. w. of Scotia, Sept. 22, 1909. O.E.J, (figured). Westmore-
land Co.: On rocks. Chestnut Ridge above Hillside. Sept. 23, 1910. O.E.J. &C G.K.J.
2. Orthotrichum Lescurii Austin
(O. cupulatum var. minus Sullivant)
Plate XIX
Rather densely cespitose, about 5 mm high: stems thickly-foliate, branched;
leaves lanceolate or som.e of the lower ovate, the upper about 3 mm long, the
lower shorter, acute, the margins entire, revolute, strongly costate nearly to
the apex; basal leaf-cells quadrate to e'ongate-rectangular, smooth, hyaline,
above becoming rounded-hexagonal, sub-opaque, densely papillose, smaller and
quite regularly he.xagonal at the apex; seta very short, 0 5 mm, about one-half
enclosed in the involucre; calyptra narrowly campanulate, plicate, hirsute with
erect hairs; lid mamillate, rounded but flattened; capsule oblong-cylindric and
rather suddenly tapering to the seta, about 1.3 mm high and 0.5 mm in thick-
ness, when moist globose-oblong and 8-striate, when dry deeply 8 furrowed and
sometimes contracted below the mouth; capsule always about the sam.e length
as the upper leaves, or sometimes slightly exserted when dry; peristome single,
the teeth 8, short, equidistant but leaning towards each other in pairs, triangu-
lar-lanceolate, papillose, articulate, the divisural usually complete and often
split, teeth when dry erect or incurved; spores mature in spring, .014-. 017 mm.
On rocks, usually granite or trap; from New England to Ontario south
to Missouri and Pennsylvania, and in the Rocky Mountains to British
Columbia.
Rare in our region. Westmoreland Co.: On sandstone rocks at mouth of Bear's
Cave, on slope of Chestnut Ridge above Hillside, September 16, 1910. O.E.J, and
G.K.J, (figured).
3. Orthotrichum stellatum Bridel
(O. Braunn Bryologia Europaea; O. strangulatum Sullivant)
Plate XIX
Sparsely cespitose to scattered, less than 5 mm high, dark green: stems
sometimes creeping at base, erect-spreading, simple or branched; leaves spread-
ing when moist, the upper somewhat clasping, when dry appressed, not crisped,
concave, ovate to lance-ovate, the margins more or less revolute, apex sub-acute
to obtuse, sometimes erose-denticulate and sometimes hyaline; costa strong,
sub-percurrent basal leaf -cells hyaline, smooth, at margin quadrate, about .016
mm in diameter, toward the costa rectangular and reaching about .090 x .016
mm, median cells papillose, opaque, rounded, about .020 mm in diameter, the
apical smaller, rounded and less papillose; capsule oblong-oval, about 1.2 mm
106 American Midland Najuralist Monograph No. 6
long tapering abruptly into a seta about two-thirds as long, 8-costate, when
dry much constricted below the mouth and very deeply 8-plicate, somewhat
sinistrorse, reddish-brown; peristome-teeth lighter in color, granulose, with
distinct divisurals, when dry closely reflexed, when moist erect, segments linear,
shorter than teeth; calyptra narrowly conic-mitrate, non-hirsute and plicate;
spores globose, somewhat papillose, incrassate, about .017 mm in diameter.
On bark of living trees (often on apple and butternut trees. — Strout);
Europe, Asia, northern Africa, and from Nova Scotia to Georgia and Iowa.
Scarce in our region. Allegheny Co.: On base of Quercus imbricaria in mixed pine
and oak woods at Dutil Church, near Douthett, December 29, 1908. O.E.J, (figured).
McKean Co.: Bradford. D. A. Burnett. (Porter's Catalogue).
4. Orthotrichum ohioense Sullivant
(O. canadense Sullivant, not Bryologia Europaea)
Plate XIX
Densely cespitose, yellowish green above, dark or brownish below: stems
freely branching, about 6-10 mm high; leaves lanceolate from an oblong base,
about 1.5-3 mm long, spreading to ascending, bluntly acute to rounded-obtuse,
papillose with entire and revolute margins; costa strong, ending at a little
below the apex; median leaf-cells quadrate to rectangular, moderately incras-
sate, towards the margins and upwards becoming smaller, more incrassate,
quadrate, sub-papillose, the upper small, rounded, densely papillose, incrassate:
seta shorter than the urn; capsule more or less completely immersed, ovate-
oblong when moist to somewhat narrower and pyriform-campanulate when dry,
symmetric, when dry 8-striate, pale yellow, tapering at base, slightly con-
stricted below the mouth; exothecial cells at mouth in one to three rov/s, small,
quadrate, below abruptly rounded and strongly incrassate, on the main body
of the urn rectangular and much smaller; stomata immersed, the outer peri-
stome of 8 double teeth, yellowish-pellucid, densely puncticulate, triangular-
lanceolate, 5-7-articulate; segments of inner peristome of 8 short, linear-subu-
late segments of two rows of cells; calyptra conic-campanulate, yellowish,
plicate, densely erect-hairy; operculum low-convex, apiculate rostrate; spores
yellowish-brown, pellucid, densely papillose, .01 8-. 020 mm, mature in spring. —
about April: autoicous, antheridial clusters axillary.
On bark of trees. New Brunswick to Ontario and south to Georgia.
Probably rather common in our region. Ashtabula, Ohio, and as follows: Erie Co.:
On bark of Populus deltoides, Presque Isle, September 20-22, 1906. O.E.J. McKean
Co.: Langmade, Bradford, May 8, 1898. D.A.B. (figured). Washington Co.: On
bark of fallen willow. Hackney Station, A. Linn and J. S. Simonton, April 27, 1894.
4a. Orthotrichum ohioense var. citrinum (Austin)
Lesquereux and James
(O. citrinum Austin)
Leaves dark green, narrowly lanceolate; capsule thin, yellow.
On bark of trees and with about the same range as the species.
Occurs in our region at Painesville, Ohio, and as follows: Westmoreland Co.: T.
P. James. (Porter's Catalogue).
Jennings: Manual of Mosses — 8. Orthotrichaceae 107
5. Orthotrichum pumilum Dickson
Densely cespitose, dark green, tufts less than 1 cm high; stems only a few
mm high; leaves erect-spreading, lance-oblong, when dry imbricated and
straight, obtuse to short-acuminate, margin recurved; upper leaf-cells rounded-
hexagonal, rather thin-walled for the genus, relatively rather large, minutely
papillose, the basal cells larger, rectangular and smooth; costa strong, ending a
little below the apex: seta very short, capsule immersed, small, when moist
oval-oblong with a distinct neck, when dry narrower, constricted below the
mouth, 8-plicate, yellowish to orange; stomata immersed; exothecial cells quad-
rate to rectangular-hexagonal, becoming at the mouth much smaller, rounded,
and rather obscure; peristome-teeth 8, bigeminate, yellowish-pellucid, triangu-
lar-lanceolate, papillose, when dry reflexed, the segments alrriost as long, lance-
subulate, 8 in number; calyptra smooth, short, inflated, lustrous, with few or
no hairs, light yellow; spores .012-. 015 mm, mature in spring: autoicous.
On trunks of trees; Europe, Asia, Algeria, and, in North America, from
southeastern Canada to Tennessee and westward to Idaho and Utah.
Rare in our region. Washington Co.: Linn and Simonton. (Porter's Catalogue).
6. Orthotrichum obtusifolium (Schrader) Bridel
Plate LXIII
Small yellowish green tufts up to about 8-10 mm high; leaves appressed
when dry, spreading when moist, about 1.5-2.5 mm long, oblong-ovate to
lingulate, entire, decurrent, trough-shaped with plane margins, obtuse to
broadly acute; costa strong but ending below apex; thick-walled brood-bodies
abundant on apical upper surface of some leaves; upper leaf-cells thick-walled,
rounded, basal marginal short-rectangular, basal median somewhat larger and
elongate-rectangular, middle and upper cells with low blunt papillae.
On deciduous tree bark, from southern Canada and New England to
Maryland and from Alaska to our Southwest.
Washington Co.: "On fallen ash tree," near Washington, Linn and Simonton, Feb.
10 and Mar. 16, 1894 (figured).
4. Ulota Mohr
Autoicous, rarely dioicous: mostly forming cushions on living trees: stems
often creeping with erect or ascending branches, radiculose; leaves mostly
(except U. americana) crisped or contorted when dry, mostly spreading to
squarrose, from a broadly concave base lance-linear, carinate, with margins
mostly revolute below; costa percurrent or nearly so; basal cells narrowly linear,
yellowish but with a margin of one to several rows of hyaline, thin-walled,
rectangular to quadrate cells: capsule erect, exserted, with long, tapering neck,
symmetric, 8-plicate when dry, stomata superficial; annulus persisting; peri-
stome mostly double, segments usually 8, rarely 16 or none; lid convex or
conic, rostrate; calyptra mitrate, with 10-16 obtuse folds, incised-lobed at base,
hirsute with shining golden-yellow hairs or rarely almost glabrous.
A world-wide genus of about 50 species, most numerous in America; in
North America about 15 species; in our region three species.
108 American Midland Naturalist Monograph No. 6
Key to the Species
A. Rupwstral; leaves not crispate when dry 1. (J. americana
A. Arboreal: leaves more or less crispate when dry B
B. Capsule smooth, slightly plicate below the distinctly narrowed mouth and at the
neck 2. U. Ludwigii
B. Capsule wide-mouthed, distinctly plicate c
C. Capsule constricted below the mouth, gradually narrowed at base to the long neck:
teeth confluent 3. U. crispa
C. Capsule not distinctly constricted below the mouth, abruptly narrowed to the shorter
neck; teeth separated at apex 3a. U. crispa var minus
I. Ulota AMERICANA [Beauvois] Limpricht, not Mitten
(U. Hutchinsiae Hammar)
Plate XIX
Rather loosely cespitose, blackish with greenish tips, about 1 cm high, or
less: stems creeping, sparingly branched with erect branches, when dry the
leaves appressed and straight; leaves often with hair-like paraphyllia at base,
lance-ovate or linear-lanceolate from an ovate base, carinate, concave at least
below, margins usually revolute in lower half, costa and base of lamina pellucid-
castaneous, apex sub-acute; costa strong, sometimes percurrent; apical and
median leaf-cells incrassate, papillose, rounded-quadrate, rather opaque, the
basal marginal rounded-quadrate to rounded-rectangular, hyaline, towards the
costa becoming linear, more or less vermicular and occasionally anastomosing,
much incrassate and markedly pellucid-castaneous : seta about 2 mm long,
smooth, with a distinct smooth volva at base; capsule yellowish, cylindric-
oblong, basally tapering, when dry 8-costate and with more or less distinct
intermediate costae at mouth, the neck and seta decidedly dextrorse; stomata
immersed; peristome-teeth 16, more or less paired, articulate, granular, when
dry strongly reflexed, segments 8, about one-half as high, bi-seriate below;
calyptra yellowish, mitrate, plicate, incised-Iobate at base, densely clothed with
erect to spreading slender hairs; lid conic-rostrate; spores globose, papillose,
incrassate, about .016-.018 mm, mature in spring.
On non-calcareous rocks, mainly in hilly or mountainous regions; Europe,
Asia, and from New Brunswick to Georgia and west to the Rocky Mountains.
Common in our region. Cambria Co.: Cresson. T. P. James. (Porter's Catalogue).
Centre Co.: Dry rocks at top of Bald Eagle Mt., near Mattemville, July 14, 1909.
O.E.J. Fayette Co.: On rocks in bed of Youghiogheny River at Ohio Pyle, Septem-
ber 1-3, 1906. O.E.J, and G.K.J. Huntingdon Co.: T. C. Porter. (Porter Cata-
logue). McKean Co.: On rocks, Rutherford, May 13, 1898. D.A.B. Somerset
Co.: On dry, crumbling shale, Ursina, May 12, 1905. O.E.J, (figured).
2. Ulota Ludwigii (Bridel) Bridel
{Weissia coarctata Lindberg)
Plate XIX
Loosely cespitose, yellowish green: stems more or less creeping with erect
shoots often 1 cm high, usually shorter, somewhat branched below; leaves
erect-spreading but slightly twisted when dry, lance-ovate to lanceolate, concave
at base, often carinate-concave in upper third, acuminate above but the extreme
Jennings: Manual of Mosses — 8. Orthotrichaceae 109
apex rather obtuse, the margin entire and often recurved; costa strong, reddish,
sub percurrent; basal leaf-cells at margin quadrate, hyaline, towards the costa
rectangular to linear-vermicular, reddish-pellucid, the median cells rounded-
quadrate, incrassate, slightly papillose, the apical cells similar; capsule pyriform,
tap>ering into a slender dextrorse seta, seta and capsule together about 3.5 mm
long, capsule strongly costate but with a very small mouth and, even when
dry, smooth and plicate only immediately below the mouth, pale yellowish-
brown, stomata superficial at the base of the urn; calyptra narrowly conic-
mitrate, hairy; lid rosteliate; peristome single, or rarely with rudimentary seg-
ments, teeth somewhat paired but split apart above, when dry erect, narrowly
triangular, granulose, distinctly articulate, with a distinct divisural; spores
papillose, globose, about .020-. 022 mm in diameter, mature in summer.
On tree-trunks in woods, usually in mountainous or hilly country; Europe,
and in North America from the Gulf of St. Lawrence to Ontario and south
to North Carolina.
Rather uncommon in our region. Centre Co.: Bear Meadows. T. C. Porter. (Por-
ter's Catalogue). McKean Co.: Rutherford, March 6, 1893. D.A.B. (figured). Som-
erset Co.: Bark of Butternut tree, Fall Run, near Barronville, John F. Lewis, May 17,
1930. Washington Co.: Linn and Simonton. (Porter's Catalogue).
3. Ulota crisp a [Linnaeus] Bridel
{Ulota ulophylla Brotherus; Orthotrichum crispum Hedwig)
Plate XX
Densely cespitose, yellowish-green above, darker below, the tufts about 8
mm high: stems sparingly branched, growing perpendicular to the bark on
which it is found, sometimes decumbent at base; leaves straight and erect-
spreading when moist, when dry much crisped, narrowly lance-ovate to sharply
acute to acuminate at the apex, concave and more or less carinate, often
marginally revolute; marginal basal leaf-cells hyaline, the inner basal pellucid,
linear, often somewhat vermicular, the median cells incrassate, rounded, bluntly
papillose, the apical cells smaller and less papillose; costa strong, sub-percurrent;
seta and capsule together about 4 mm long, capsule ovate-globose when wet,
about 1 mm long, tapering rather gradually into the neck and seta, when dry
somewhat constricted below the m.outh, with the neck and seta dextrorse, the
costa brownish-pellucid; annulus brown, pellucid, of about 3 series of small,
close-set, rounded, cells; teeth triangular-lanceolate, united into 8 pairs, when
dry reflexed, each pair confluent and cribrorse at apex, often split along the
divisural below; segments 8, consisting of two rows of cells nearly up to the
apex, a little shorter than the teeth; spores globose, about .023-.026 mm,
mature in summer.
On trees in woods; Europe, Asia, Tasmania, Alaska, and from Newfound-
land to Georgia and west to Minnesota.
Fairly common in our region, especially on black oak trunks. Now known from Alle-
gheny, Butler, Cameron, Centre, Crawford. Fayette, McKean, Potter, Washington, and
^X'estmoreland counties. Specimen figured: on black oak truunk. Bald Eagle Mt., near
Matternville, Center Co., Sept. 22, 1909. O.E.J.
110 American Midland Naturalist Monograph No. 6
3a. Ulota crispa var. minus (Schwaegrichen) Jennings
{U. crispula Bruch)
With shorter stems and leaves than in the species and a capsule which
rather abruptly narrows into a long neck, and with a sub-globose to oval urn,
which, when dry and empty, is more or less open-mouthed and turbinate,
with little or no constriction below the mouth.
This variety is reported with a general range similar to that of the species
but we have as yet seen no typical specimens of it from our region. Grout
(Moss Flora) does not regard minus as sufficiently distinct from crispa.
Porter's Catalogue lists it trom several counties in Eastern Pennsylvania and
from McKean County, D. A. Burnett; but a specimen in the Herbarium of the
Carnegie Museum collected by Burnett, at Langmade, May 29, 1898, McKean
County, is evidently purely U. ulophylla.
Family 9. Splachnaceae
Autoicous or dioicous, rarely pseudautoicous : annual or perennial cespitose
bog or alpine mosses, usually living on decaying animal or vegetable matter,
the tufts green to yellow-green, inside more or less red-radiculose, sometimes
blackish: stem delicate with a large central strand; leaves mostly distant, flaccid,
more or less broad; costa mostly not quite percurrent, usually with two basal
guides; leaf-cells loose, parenchymatous, 4-6-sided, elongate towards the base,
sparingly chlorophyllose, often inflated at the margin of the leaf: seta erect,
sometimes very long; capsule erect, symmetric, with a long collum or with a
large colored hypophysis: usually brownish or red when mature, operculate,
and with a peristome of 32 divisions joined together to form 16 teeth (except
Tayloria splachnoides) and often grouped in twos or fours, and usually con-
sisting of three layers. There is a pre-peristome present in a few species. The
peristome teeth more or less hygroscopic, vertically striate, trabeculate, punc-
tate, mostly golden-brown. Annulus usually none; spore-sack surrounded by
a cavity; columella strong; spores small to large; operculum convex to umbonate
or long-conic, rarely none; calyptra small, either cucullate and united into a
tube below or conic and almost entire to lobed.
There are 5 genera and more than 60 species of these peculiar plants,
widely distributed in mostly cold northern and alpine regions.
Key to the Genera
Hypophysis much wider than the capsule 1. Splachnum
Hypophysis not much if any wider than the capsule 2. Tetraplodon
1. Splachnum Linnaeus
A genus of 7 or 8 species of mostly cold northern bogs and on decaying
cattle dung. Only the following species known from our region.
1. Splachnum ampullaceum Linnaeus
Plate LII
Loosely caespitose, up to about 1 cm high, in low tufts matted together
below by reddish-brown filaments; monoecious, the perigonial branches arising
Jennings: Manual of Mosses- -9. Splanchnaceae 111
from within the matted material. Leaves ot the fertile stem few, about 1 mm
long, tufted, thin, obovate to broadly lanceolate, abruptly acuminate, costate
into the apex, usually with a few sharp teeth towards the apex; leaf-cells large,
lax, irregularly rhomboid-hexagonal but longer towards the margin and base
and smaller above; perigonial branches up to about 1 cm high much exceeding
the perichaetial, sparsely leafy with lanceolate slenderly pointed, above sharply
serrate blades; antheridial flowers capitulate to discoid, surrounded by squar-
rose very slenderly acuminate squarrose perigonial leaves up to 3 mm long: seta
erect, somewhat twisted when dry, reddish; hypophysis much wrinkled when
dry, when wet about 3 mm long by 2 mm thick, pyriform, more slenderly
tapering below, when ripe red-purple above, somewhat lilac below, composed
of loose tissue and with stomata surrounded by a ring of stellately radiating
cells; capsule oval-oblong, about 1 mm high; operculum convex, obtusely mam-
iliate; peristome yellowish, consisting of 32 divisions united to form 16 three-
layered teeth forming 8 triangular groups, very hygroscopic, inflexed when wet
and abruptly strongly reflexed when dry, the areolation of the middle layer of
cells strongly predominating when viewed under the microscope and being
quite irregular in pattern: pre-peristome consisting of 16 very thin, oblong-
lingulate teeth about one-fifth the height of the main peristome; annulus none;
areolation of the capsule walls of rounded, small, strongly incrassate cells;
columella much exserted from dry capsule dilated at the apex; spores smooth,
6-7 /x in diameter.
Mostly on cow dung. New Jersey, Pennsylvania, and Ohio to Wisconsin,
and north to Newfoundland, Quebec, and Ontario.
These specimens are smaller in practically all parts than stated in most
descriptions.
Elk Co.: Midmont Swamp. C.M.B. July 1, 1948. Elev. 1940 ft. Somerset Co.:
Open sphagnum bog. near Mt. Davis. Elev. 2400 ft. C.M.B. & O.E.J. July 15. 1949
(figured).
2. TeTRAPLODON Bruch & Schimper
A genus of at least 9 species, mostly on decaying animal excreta of cold
northern and alpine regions and resembling Splschnum excepting for the nar-
rower leaves, hypophysis little if any Vv'ider than the capsule, and teeth first in
fours then in pairs, and consisting of two layers of cells.
1. TeTRAPLODON angustatus (Hedwig) Bryologia Europaea
{Splachnum angustatum [Linnaeus f.} Hedwig)
Plate LXXII
In dense tufts; stems erect, about 8-12 mm tall, pale translucent-yellowish,
with dense reddish tomentum below; leaves few, rather distant except for
about four to .seven in a comal tuft, these being about 3-5 mm long by 0.5-0.8
mm wide, narrowly lanre-oblong, tapering into a very long slender acumination,
usually with a few, scattered, .sharply spreading, one-celled teeth; costa loosely
cellular, at base about one-foiarth to one-third the width of the leaf, tapering
gradually into the long, somewhat reddish, flexuous, excurrent point; leaf-cells
112 American Midland Naturalist Monograph No. 6
lax, thin willed, ratlier irregular rectangular, two to six times as long as wide,
narrower along the margin : seta pale yellowish, somewhat translucent, about 4-8
mm long in our specimens and exceeding the leaves, abruptly tapering above
into a pale yellowish hypophysis with a dark brownish center; hypophysis when
moist about 0.7-0.9 mm long, smooth, pyriform-oval, when dry shrunken and
deeply wrinkled and corrugated: capsule when moist about as long as the
hypophysis, narrowly cylindric-oval, about twice as high as thick, when dry
much constricted in the middle, reddish purple towards the rim; peristome
teeth 16 (32 divisions), adhering in 8 groups which, when mature and dry are
abruptly reflexed and occa-^ionally split apart into 16 teeth, each of the eight
groups triangular, blunt, about twice as high as wide at the base; operculum
low-hemisperic, distinctly bluntly apiculate; calyptra narrowly conical; spores
smooth, thin-walled, globose to ellipsoid, about .007-. 009 mm in diameter.
Usually occurring on animal droppings and heretofore reported as ranging
from the Catskills to Minnesota and the Rockies, north to Newfoundland,
Athabasca, and British Columbia.
Somerset Co.: Open sphagnum bog, near Mt. Davis, at an altitude of Z'lOO ft.,
O. E. Jennings, Aug. 21, 1936; and July 15, 1949, Charles M. Boardman (figured). In
one case the plants were intermingled in the same tuft with Splachnum ampulldceum.
This station marks an extension of more than 250 miles south of the pre-
viously known range but *"he plants were here in company with other northern
bog plants such as Menyanthes which is here also at its southernmost frontier.
The leaves are somewhat less serrulate than is generally described for T. angus-
tatus, the spores slightly smaller, and the fallen capsules which were shaken
out of the tufts scarcely appeared to be capable of entirely covering the
capsule. Otherwise the specimens seem typical.
Family 10. DiSCELIACEAE
Dioicous; gregarious, annual, with persistent protonema: stem very short
with gemmiform foliation; inner leaves largest, ovate to lance-oblong, obtuse
to acute, with plane margins, entire or irregularly crenulate at apex, faintly
costate towards apex; cells irregularly loosely rhomboid-hexagonal, thin-walled,
somewhat pellucid, elongated below, smooth, very sparsely chlorophyllose: seta
elongate, 2-3 cm, stiff, slender, pellucid, red or castaneous, sinistrorsely twisted;
capsule minute, cernuous or horizontal, globose-ovate, smooth, with a very
short collum; annulus of one (or two) series of cells, falling away in pieces;
peristome inserted below the mouth, simple; teeth lanceolate, acute, red, the
lower half usually perforate or split, the exterior usually vertically striate but
with no median line, not papillose, the interior with papillae and projecting
trabeculae: spores medium size; operculum reddish or orange, large, convex,
umbonate; calyptra split down on one side and usually remaining attached to
the seta by the constricted base.
A peculiar family consisting of but one genus with only the following
species. Occurring on bare soil in northern Europe, Asia, and, in North
America, in Illinois, Ohio, eastern Pennsylvania, and New Jersey.
Jennings: Manual op Mosses — 11. Ephemeraceae 113
1. DiSCELIUM Bridel
1. DiSCELIUM NUDUM (Dickson) Bridel
{Discelium mcarnatum (Schwaegrichen) Jennings; D. nudum Bridel;
Weisia incarnata Schwaegrichen )
Plate LXIII
Characters as for the f.Tnily.
The Ohio station for this species is near Ashtabula, only a few miles from
the northwestern corner of Pennsylvania.
Allegheny Co.: Four miles east of Monongahela City, C. M. Hepner, April 8 and
April 16 (figured), 1934; roadside ditch 1 mile north of Indianola, C. M. Boardman,
Nov. 24, 1935, and May 10, 1936.
Family 11. Ephemeraceae
Autoieous or dioicous, rarely polyoicous or synoicous; minute, about 1-2
mm high, singly disposed or gregarious, mostly stemless; protonema persistent;
leaves minute, rosette-like, spreading or erect-spreading; leaf-cells mostly lax,
more or less elongate below, rectangular to rhomboid-hexagonal, above shorter,
usually smooth; costa none to excurrent, usually present: seta none or short;
capsule immersed, sub-globose, c'.eistocarpous (or with dehiscence line in Natio-
mitrium); operculum sometimes differentiated but rarely deciduous of itself;
spores mostly large and papillore; calyptra mostly small, delicate, mitrate-
campanulate.
Minute plants growing on soil.
1. Ephemerum Hampe*
Dioicous, rarely polyoicous: minute plants with abundant and persistent
green protonema: upper leaves elongate-lanceolate to linear; costa none or vari-
ously developed; leaf-cells lax, mostly thin-walled, rhomboidal: seta rudi-
mentary or none; capsule mostly globose and apiculate, cleistocarpous, walls
(exothecium) of two layers of cells with stomata; spores large, up to .08 mm
in diameter, warty; calyptra campanulate, delicate, torn at the base or some-
times only on one side.
A cosmopolitan genus of about 30 species; most abundant in North Amer-
ica, perhaps several of these occurring in our region, but on account of their
minute size not yet collected.
Key to the Species
A. Costa none B
A. Costa more or less complete, or vanishing towards the base C
B. Leaves lanceolate, erect-patent, spores .060-. 080 mm
1. E. seTTatum [Schreber] Hampe
* Nanomttnum Lindberg (Micromitrium Austin) differs from Ephemerum m having
a rudimentary but not deciduous operculum and the capsule without stomata and with a
wall but one cell thick; leaves ecostate and calyptra minute.
Leaves entire N. synoicum (James) Lindberg
Leaves serrulate N. Austinii (Sullivant) Lindberg
114 American Midland Naturalist Monograph No. 6
D. Leaves linear-lanceolate, often secund; spores smaller
(E. serratum var. angustatum Bryologia Europaea)
C. Costa percurrent or vanishing near the apex D
C. Costa excurrent and quite strong G
D. Upper leaves broadly lanceolate to elongate-lanceolate E
D. Upper leaves linear to linear-lanceolate, long-acuminate F
E. Capsule short, obtuse; costa effaced at base, upwards towards the afsex continuous
2. E. cohaerans
E. Capsule acutely beaked; costa loosely areolate, scarcely distinct except towards the
short, entire, pointed apex (£. pallidum Schimper)
F. Calyptra smooth G
F. Calyptra papillose (£. crassinervium var. papillosum (Austin) R. &C C.)
G. Leaves linear-lanceolate; costa excurrent, weak at base, serrate at ajjex
3. E. crassinervium
G. Leaves linear; costa usually strong and wide at base, decidedly excurrent and strongly
spinulose (£. spinulosum Schimper)
1. Ephemerum serratum (Hedwig) Hampe
(Phascum serratum Schreber)
Plate LXIV
Protonema abundant, branched, persistent, alga-Iike, growing on wet soil.
Plants minute, under the hand-lens looking like little buds. Leaves up to 1 or
1.5 mm long, ecostate, ovate to ovate-lanceolate, sharply serrate down to the
middle or beyond; leat-cells irregularly rhomboid-hexagonal to rectangular and
up to 6:1. Antheridia usually 3 or 4, narrowly obovoid, about half the length
of the longest leaves. (Capsule not seen but described as "shining globular
to kidney-shaped, warty, maturing winter to spring." — Grout) .
On open, wet soil. Rare. New York to Ohio and North Carolina;
California.
Erie Co.: Around margins of Old Fog Horn Pond, Presque Isle. July 27, 1933.
Nelle Ammons (figured).
2. Ephemerum cohaerans (Hedwig) Hampe
{Phascum cohaerans Hedwig)
Plants minute, 1-1.5 mm high, densely gregarious or somewhat cespitose:
protonema persistent, green or yellowish with age; leaves lance-ovate to lance-
oblong, erect-spreading, acute, serrate above; costa thin, stronger above, ending
in the apex or just below; leaf-cells lax, oblong-hexagonal, rather thin-walled:
capsule sub-sessile, castaneous, sub-globose, obtusely apiculate, bearing stomata
all over; calyptra more or less lobed or torn at base; spores large, .060-.080
mm, coarsely tuberculate, mature in late autumn: dioicous.
On moist sandy or clayey soil, Europe and, in eastern North America,
south to Louisiana. Not yet collected in our region but occurring in eastern
Pennsylvania and in Ohio.
3. Ephemeru.m crassinervium (Schwaegrichen) C. Mueller, not Hampe
{Phascum crassinervium Schwaegrichen)
Plants minute, not over 1 mm high, gregarious: green protonema persistent;
leaves erect-spreading, flexuous, linear-lanceolate, slenderly long-acuminate,
Jennings: Manual of Mosses — 12. Funariaceae 115
rather coarsely serrate above, marginally plane; costa flat, faint below, stronger
above, excurrent, denticulate dorsally above; leaf-cells more or less rectangular
to oblong-hexagonal, thin-walled: capsule with a very short seta, immersed,
globose, apiculate; the capsule about half-covered by the cleft-lobate, mitrate-
conic calyptra; spores large, papillose, mature in late fall to early spring.
On moist earth, often in swamps, eastern North America from Connecti-
cut to Florida and Illinois It occurs in central Ohio and eastern Pennsyl-
vania and is, probably, the plant collected by James in Indiana County as
reported in Porter's Catalogue, but specimen not seen.
Family 12. FuNARIACEAE
Autoicous or paroicous, rarely dioicous or synoicous: annual or rarely
biennial, low, mostly light green, gregarious or loosely cespitose: stem mostly
with a central strand, radiculose only at the base; leaves soft, wide, the upper
larger and forming a rosette, concave, margin plane to involute, entire or den-
ticulate upwards, sometim.es bordered; costa delicate, rarely excurrent, with
two large basal guides, rarely lacking; leaf-cells large, parenchymatous, thin-
walled, never papillose, but slightly chlorophyllose, oblong-rectangular below,
rhombic-hexagonal above: seta mostly erect and red, twisted; capsule either
erect, symmetric and globose to pyriform, or cemuous and arcuate-pyriform;
collum mostly distinct; annulus rarely present; peristome inserted back of the
periphery to the distance of the thickness of several cells, simple or double,
rudimentary or none; teeth, if present, 16, obliquely dextrorse, strongly trabecu-
late; segments, if present, 16, opposite the teeth, with no basal membrane;
columella mostly thick; spores mostly medium-sized; operculum mostly weakly
convex, sometimes umbonate or none; calyptra various, often inflated, usually
rostrate and cucullate.
Key to the Genera
A. Capsule immersed - B
A. Capsule exserted C
B. Capsule more or less regularly dehiscing at about the equator 1. Aphanorhegma
B. Capsule with distinct operculum, smaller than urn 2. Physcom.tTium
C. Capsules unsymmetric, peristomate, usually with a double peristome; seta much
longer than stem 3. Funaria
1. Aphanorhegma Sullivant
Paroicous, rarely synoicous: low, gregarious to almost cespitose. pale gr:en;
stem radiculose at base, loosely foliate below, densely foliate above; leaves
spreading or the upper almost erect, obovate to oblong or spatulate-lanceolate,
acute, serrate in the upper half; costa ending below the apex; leaf-cells lax, the
basal rectangular, the upper oblong-hexagonal, the marginal forming a narrow
uniseriate border: sera rudimentary; capsule spherical, without a collum, laxly
areolate; annulus none; peristome none; spores large, densely spinulose; oper-
culum half-spherical, of same size as urn, obtusely apiculate; calyptra ccnic-
mitrate, lobed, glabrous.
116 American Midland Naturalist Monograph No. 6
A genus of tliree species, on damp soil. One in Cuba and the following
two in temperate North America:
Key to the Species
A. With strongly collenchymatous exothecial cells; capsule rather regularly dehiscing
1. A. serratum
A. Exothecial cells not collenchymatous, thin-walled; capsule not regularly dehiscing
2. A. patens
1. Aphanorhegma serratum (Hooker, f. and Wilson) Sullivant
Plate XX
Gregarious, light green: stems erect, simple or forking, 1-5 mm high, radicu-
lose at base; leaver* small and lance-oblong below, rapidly becoming larger up
to 3-5 mm long, narrowly lance-obovate above, the lower widely spreading and
flexuous, the inner erect-spreading, thin, slightly serrate above the middle, apex
acute to acuminate: costa medium, ending in or just below the apex; the
median basal cells thin-walled and more or less inflated, rectangular, the mar-
ginal narrower, a few quadrate at the base, becoming linear-rectangular above
the base, in the upper part of the leaf their tips extending as low serrations,
the median rhomboid to short rectangular with walls medium, the apical
longer and narrower: seta very short and stout; capsule brown when ripe, glo-
bose to depressed-globose, about 0.75 mm in diameter, smooth to apically
papillose, splitting in the middle along a line of one or two rows of small
more or less orange-pellucid cells, the upper half of the capsub (operculum)
apiculate-rostrate; exothecial cells of capsule quadrate, conspicuously collen-
chymatous; calyptra hyaline, conic-mitriform, 4-6-lobed, covering the upper
half of the operculum; spores globose, about .030 mm in diameter, orange-
pellucid or even darker, mature in autumn.
On damp clayey soil in the northern and middle United States, in our
region usually along streams where submerged during periods of high water.
Now known from Allegheny, Butler, Clarion, Fayette, Greene, Potter, Washingon,
and Westmoreland counties. Specimen figured: Cheat Haven, Fayette Co., Sept. 6, 1910.
O.E.J, and G.K.J.
2. Aphanorhegma patens (Hedwig) Lindberg
(Physcomitrella patens [Hedw.] Bryologia Europaea; Phascum patens Hedwig)
Gregarious, pale green: stem distinct but very short, about 2 mm; leaves
lance-ovate to oblong or oval, usually shortly and bluntly acuminate, the
upper often obovate-acuminate and larger, forming a rosette, all serrate above;
costa narrow, ending below the apex: leaf-cells lax, widely rectangular to hex-
agonal: seta short, capsule globose, thin-walled, usually splitting equatorially,
brownish, immersed to slightly emergent, obtusely apiculate; spores papillose,
.025-. 030 mm, mature in autumn: paroicous; antheridia sessile in upper leaf-
axils.
On wet clayey or sandy soil in fields, along sides of pools, river banks, etc.
Europe, Asia, and, in North America, from Quebec to the northern part of
eastern United States. Not uncommon in Ohio and also reported from Lan-
caster County, Pennsylvania. Not yet reported from our region.
Jennings: Manual of Mosses — 12. Funariaceae 117
2. Physcomitrium (Bridel) Fuernrohr
Autoicous: mostly minute, densely gregarious to cespitose, green, mud-
inhabiting mosses: stem erect, simple, radiculose below, loosely foliate; leaves
flaccid, mostly appressed when dry, spreading when moist, concave, obovate to
oblanceolate or spatulate, mostly not margined, more or less serrate, obtuse
to acuminate; costa mostly strong, incomplete to excurrent; areolations lax:
seta mostly long; capsule erect, symmetric, globose to short-pyriform, with lax
areolation; collum short and thick; annulus small-celled and persistent or large-
celled and disappearmg in pieces; gymnostomous; spores large, papillose; oper-
culum broad, conic-onvex, umbonate or apiculate; calyptra long and erect-
rostrate, mitrate, lobed to the base of the beak, covering one-half or less of the
capsule.
A cosmopolitan genus of about 75 species; about 20 species in North
America, at least 2 sF>ecies in our range.
Key to the Species
A. Seta very short; capsule immersed, wide-mouthed; calyptra small .1. P. immersum
A. Seta longer; capsule exserted, not wide-mouthed; calyptra larger 2. P. tuTbtnatum
1. Physcomitrium immersum Sullivant
(Gymnostomiim immersum Sullivant)
Plate XX
Plants small, erect, gregarious, light green, simple or branching, 3-8 mm
high; leaves 1.5-3.5 mm long, obovate to oblanceolate, serrate above the mid-
dle, spreading to ascending; costa strong, ending a little below the apex;
leaf-cells parenchymatous, rather large and thin-walled, the basal rectangular,
about 2-5:1, the upper irregularly oblong, the marginal narrower and in the
alar region a few much shorter: capsule immersed, globose to pyriform-globose,
0.6-0.9 mm in diameter, apiculate-rostrate, about the upper two-fifths consti-
tuting the operculum, yellow-brown when ripe; seta considerably shorter than
capsule and stout; exothecial cells irregularly quadrate to hexagonal, somewhat
mcrassate, the annulus consisting of one to three rows of much smaller, orange-
pellucid, to brown-pellucid cells, the cells of the wall usually laterally elon-
gated for one or two rows above and below the annulus; calyptra mitrate, the
basal margin 4-5-lobed, covering about one-half of the operculum; spores d nse-
ly papillose, orange to brownish-pellucid, globose, in our specimens about .035
mm in diameter, mature in autumn.
Usually on clayey or sandy flood-plains where submerged in time of
freshets. Quebec to Colorado, Missouri, r.nd the Carolinas, but not commonly
collected, probably on account of its small size and special habitat.
Allegheny Co.: Bare clay bank of creek, Darlington Hollow, Aspinwall. Oct. 25,
1908. O.E.J. ; Mud cracks m dry pond basin, Glenshaw. Oct. 30, 1932. J. L. Cart-
ledge. Beaver Co.: Clay bank of Little Beaver Creek, New Galilee, Sept. 10, 1906.
O.E.J.; Bank of Ohio River, Smith's Ferry, Oct. 1, 1910. O.E.J, (figured). Butler
Co.: On ground in cornfield, Millinger School, Dec. 2, 1934. Sidney K. Eastwood.
McKean Co.: East Branch, Bradford. June 15. 1895. D.A.B.
118 American Midland Naturalist Monograph No. 6
2. Physcomitrium turbinatum (Michaux) Bridel
{Phasciim strangulatum Kindberg; P. Hookeri Macoun)
Turk's-Cap Moss; Urn Moss
Plate XX
Gregarious, often densely so, light green; stems 3-5 mm high, erect, usua'Iy
simple; leaves 3-5 mm long, lance-oblong to obovate-lanceolate, slightly ser-
rulate above the middle, flat and spreading when moist, somewhat crisped and
incurved when dry; capsule erect, 1.5-2 mm high, globose to pyriiorm, whan
dry becoming turbinate and constricted below the mouth and at the base, final-
ly becoming brown and urn-shaped; exothecial cells slightly incrassatc, rhom-
boid to hexagonal, the mouth bordered by about 9-12 rows of laterally
somewhat elongated cells and a narrow fringe of orange-pellucid and much
smaller cells in 1-3 rows; operculum convex, bluntly mamillate to sub-rostrate;
calyptra somewhat oblique, rostrate, unequally split at base into 3-5 lobes;
spores decidedly papillose, orange-pellucid, in our specimens measuring about
.016-.040 mm, mature in May and June, occasionally later: autoicous.
Common on bare earth in fields, along roadsides, etc., from Quebec to
Florida and v/est to the Rocky Mountains, and also in California.
Known from Allegheny, Armstrong, Beaver, Butler, Crawford, Fayette, Lawrence,
McKean, Somerset, Venango, Washington, and Westmoreland counties. Specimen fig-
ured: Douthett, Allegheny County, June 5, 1909. O.E.J.
3. Fun ARIA Schreber, Hedwig
Autoicous: the antheridial inflorescences discoid, terminal, the archegoniil
on innovations: gregarious to cespitose: stem usually simple, radiculose at
base; lower leaves distant, small, the upper becoming much larger, those at the
apex more or less upright and tufted or gemmiform, entire or serrate, more
or less acute; costa incomplete to excurrent; areolation lax, elongate-rectangular
to rhombic, at the margin sometimes longer and narrower, forming a border:
seta elongated, erect or cygneous at fruiting time, later erect and twisted;
capsule with a thick colliim or elongate-pyriform, symmetric to oblique^ arcu-
ate, with a narrow mouth, smooth to plicate, erect to cernuous; annulus large-
celled, revolute or none; peristome deeply inserted, double in our species; teeth
lance-subulate, reddish to brownish-red, obliquely ascnding to the right; seg-
ments as long or shorter, yellow, papillose, opposite the teeth; spores medium;
operculum flat or convex; calyptra long-persistent, inflated-cucullate, long-
rostrate, smooth, entire.
A large cosmopolitan genus of about 200 species (including Entosthodjn) ,
on soil; about 25 species in North America, 3 in our range.
Key to the Species
A. Body of capsule neither striate nor plicate; no annulus 1. F. americanj
A. Capsule striate and more or less plicate; annulus curlmg off B
B. Leaves long acuminate; costa very often excurrent; segments less than Yl length of
teeth 2. F. flavicans
B. Leaves short acuminate; costa mostly percurrent; segments more than Yi ^^^ length
of teeth 3. F. hygrometrica
Jennings: Manual of Mosses — 12. Funariaceae 119
1. FuNARiA AMERICANA Lindberg
(F. Muhlenbergii Hedwig, — mainly plate, not description, — Lindberg)
Small, gregarious to loosely cespitose: stems very short; leaves erect-
spreading, ovate-oblong, long-acuminate, somewhat concave, plane-margined,
entire; costa strong, excurrent into a hair-point, leaf-cells lax, moderately thin-
walled, the basal quadrate-hexagonal to rectangular, the upper elongate rectan-
gular: seta slender, rather short, up to 1.5 cm long, when dry dextrorse below,
sinistrorse above; capsule erect, sub-cemuous, pyriform-oblong, the mouth
tilted to one side, when dry the long tapering neck rugulose, the urn smoothish
and constricted below mouth; no annulus; peristome-teeth lance-linear, dex-
trorsely tilted, castaneous-pellucid, papillose, articulate, with divisural, strongly
trabeculate; segments about as long and opposite teeth, pale pellucid and papil-
lose; lid conic, obtuse, calyptra inflated, long-rostrate, cucullate; spores
papillose, mature in May: autoicous.
On bare ground, or among grass, eastern Pennsylvania to Ohio and Mirme-
sota, south to Georgia, and m California, but not often collected, — pverhaps to
he expected in our region.
2. FuNARiA FLAVICANS Richardson, Michaux
Loosely cespitose: stems erect, smaller than F. hygrometrica; lower leaves
small, the upper leaves larger and tufted, oblong-spatulate to obovate. concave,
plane-margined, entire, at apex long-acuminate; costa percurrent or excurrent;
leaf-cells large, lax; seca long, erect, capsule oval-pyriform to globose-pyriform,
more or less horizontal or downward curved, dark reddish when mature, with
mouth less oblique and smaller than in F. hygrometrica, gradually attenuate
below into the seta, not much furrowed when old; lid low-convex, not apicu-
late; spores about .025 mm in diameter, mature in May or June: autoicous.
On bare moist earth, usuallv clay, in eastern United States from Connecti-
cut and New York south and west.
Lawrence Co.: T. P. James. (Porter's Catalogue).
3. Fun ARIA hygrometrica [L. — Sibthorp] He:'.wig
The Cord Moss
Plate XXI
Loosely cespitose, rathsr light green: stems about 3-10 mm high, erect,
radiculose at base, simple or basally divided; leaves erect to appressed, con-
cave, forming a bulbiform tuft, oblong-ovate, acute or shortly acuminate,
entire or slightly crenate, larger leaves 2-4 mm long by three-fifths as wide,
strongly costate to the apex or percurrently costate; cells rectangular to hex-
agonal, narrower towards margin, above more or less quadrate-hexagonal, the
lower more or less inflated, above becoming more or less incrassate: seta about
2-5 cm high, erect, sinistrorse, flexuous, lustrous, chestnut-brown, paler above;
capsule unsymm.etric, arched and turgid on upper side, 2-3 m.m long, strongly
incurved at mouth, deeply sulcate when dry, pvriform, yellowish to brown
120 American Midland Naturalist Monograph No. 6
when old, usually more or less horizontal but the upper part of seta often vari-
ously bent and curved and strongly hygroscopic; mouth about 0.6-0.8 mm
wide, annulus revoluble, deep castaneous; operculum low-convex; peristome-
teeth castaneous-pellucid, papillose, strongly trabeculate, spirally twisted,
united at apex; segments about three-fourths as long, papillose; spores smooth,
round, about .014-.017 mm; mature in May or June; calyptra cucullate, long-
rostrate, early deciduous: autoicous.
Widely distributed over the earth; throughout North America. Common
in our region on earth, burnt-over spots, etc. (Quite variable in size and leaf-
characters but we have not been able to recognize any of the several described
varieties in our region.)
Known from the following counties: Allegheny, Beaver, Butler, Cambria, Crawford,
Erie, Fayette, Greene, Huntingdon, Lawrence, McKean, Mercer, Somerset, Union, Venan-
go, Washington, and Westmoreland. It probably occurs in every county. Specimen fig-
ured: Ligonier to Donegal, Westmoreland Co., June 23, 1904. O.E.J.
Family 13. Schistostegaceae
This family consists of only the genus Schistostega whose characters are
those of the family.
1. Schistostega Mohr
Dioicous: inflorescences gemmiform, terminal, paraphyses none: minute
and slender mosses in holes in earth, in caves, etc.: annual, gregariousi on an
abundant persistent protonema, which is more or less luminous by reflected
light; sterile stems from the middle upwards with distichous, basally confluent
leaves; fertile stems with a terminal 5-seriate tuft of leaves; leaves entire, uni-
stratose, ecostate; cells prosenchymatous, lax-rhombic, sparsely chlorophyllose :
seta thin, erect, almost hyaline; capsule minute, 0.5 mm long, erect, symmetric,
globose, without stomata, annulus, or peristome; operculum small, convex and
with a red border; calyptra very small and fugacious, mitrate, covering only
the operculum, smooth, and naked; propagation often by brood-bodies on the
protonema.
One species only, in crevices and caves in non-calcareous districts, in
Europe, and, in North America, from Ontario, New England, New York
and Ohio to British Columbia.
1. Schistostega pennata [Hedwig] Hooker and Taylor
{Gymnostomitm pennatum Hedwig, Schistostega osmundacea Mohr)
The Luminous Moss
With characters as given for the genus. Not vet known in our region.
Family 14. Bryaceae
Dioicous, autoicous, paroicous, or synoicous, sometimes heteroicous; an-
theridial inflorescences with paraphyses; cespitose, persistent, mostly on soil
or rocks, sometimes on trees or rotting wood; stem usually rounded-pentagonal.
Jennings: Manual of Mosses — 14. Bryaceae 121
with central strand, radiculose at leas: at base; leaves in several series, below
mostly small and remote, above larger and often tufted, often bordered: costa
mostly with 2-5 median guides, often excurrent; cells never papillose, upp:r
prosenchymatous, mostly rhomboidal or rhombic-hexagonal, rarely linear or
vermicular, basal rectangular to quadrate: seta elongate, erect, smooth, more
or less curved; capsule cernuous to pendulous, sometimes erect, mostly sym-
m.etric, rarely arcuate, neither striate nor plicate, ovate or pyriform, rarely
almost globose; collum evident, usually wrinkling when dry; annulus usually
present, large-celled, spirally deciduous; peristome rarely none, or simple,
mostly double, the 16 teeth often bordered, hygroscopic, papillose on the
exterior, C!>pecially towards the apex, divisural line evident, trabeculae promi-
nent; segments alternating with teeth, delicate, yellowish or hyaline, often
with cilia, often united below into a basal membrane; spores small to medium;
operculum conic to convex, umbonate to apiculate or rarely short-rostrate;
calyptra cucullate, small, fugacious.
A large and cosmopolitan familv of about 15 genera and possibly 1,000
species.
Key to the Genera
A. Leaf-cells narrow, upwards narrowly rhombic to linear B
A. Leaf-cells lax, upwards rhombic to hexagonal, never linear C
B. Leaves long-subulate; cilia prommently appendiculate L Leptobryum
B. Leaves linear-lanceolate; cilia non-appendiculate, often rudimentary or none
2. Pohlia
C. Annulus mostly none; leaves decurrent, non-bordered; costa ending below apex
3. Mmobryum
C. Annulus usually present; leaves often bordered; costa usually percurrent or slightly
excurrent D
D. Sporogonia single; stem without rhizome-like stolons 4. Bryum
D. Sporogonia often several together; stems erect from rhizome-like stolons
5. Rhodobryum
1. Leptobryum (Schimper) Wilson
Synoicous or dioicous; paraohyses of the antheridial inflorescence with an
acuminate end-cell; no paraphyses in the archegonial inflorescence: weak, cespi-
tose in low, soft, lax, yellowish-green tufts; stem erect, thin, brown-radiculose
at base; lower leaves remote, small, lanceolate, uppermost leaves much larger,
tufted, erect to spreading, elongate-subulate from a lanceolate base, canalicu-
late and often distinctly toothed towards the apex; costa broad, flat, incom-
plete or percurrently filling the apex; cells very narrow and long, in the
subulation linear, the basal rectangular-elongate: seta short to long, very
thin, tortuous, twisted when dry; capsule cernuous to almost pendent, with a
thin, long, somewhat arcuate, pyriform collum, lustrous, narrow-mouthed;
annulus narrow, deciduous; peristome-teeth pale yellow, the upper part subu-
late and bordered; segments about as long, fenestrate, the lower third forming
a basal membrane, the cilia mostly long-appendiculate; spores of medium size;
operculum small, convex, and mostly umbonate.
A genus of four species; one in Tasmania, two in South America, and th?
following, almost a cosmopolitan:
122 American Midland Naturalist Monograph No. 6
1. Leptobryum pyriforme [Linnaeus] Schimper
{Webera piriformis Hedwig)
Plate XXI
Densely cespitose in light yellowish-green, soft, lustrous tufts: stems .5-1.5
cm high, slender, erect, reddish, brown-radiculose at base; leaves mostly erect-
spreading, flexuous, the upper forming a comal tuft, linear-setaceous, up to
4-5 mm, long, the basal portion lanceolate, the upper portion flexuous, with
plane margin, denticulate above; costa strong but rather wide and indistinct,
occupying most of the upper portion of the leaf and somewhat excurrent;
leaf-cells narrow and linear-prosenchymatous, or below elongate and paren-
chymatous, at base rectangular and larger, all thin-walled; perichaetial bracts
linear from a wider base: seta slender, flexuous, orange to brown, about 1-1.5
cm long; capsule inclined to pendulous, pyriform with a long narrow nrck,
altogether about 2.5 mm long, the neck much wrinkled when old, and at least
as long as the globose-oval part of the capsule, which is a lustrous orange- to
dark chestnut-brown, the mouth rather wide; annulus wide; peristome-teeth
yellowish, linear-lanceolate, the upper third suddenly narrower and sub-hyaline
and papillose, trabeculate, lamellae and divisural evident; segments about as
long, carinately split and sometimes gaping; cilia 3, strongly appcndicuate,
about as long as segments, basal membrane one-third to almost one-half the
height of the teeth; operculum convex-apiculate : spores smoothish, about .012-
.015 mm: usually synoicous: mature in June or July.
On moist shaded soil, old walls, shaded cliffs and rocks n'-ar trickling
water, etc. Cosmopolitan.
Now known from Allegheny, Butler. Erie, MrKean, Tioga, Washington, and West-
moreland counties. Specimen figured: On stone wall, Perrysville Ave., North Side, Pitts-
burgh. May 26, 1909.
2. Pohlia Hedwig
(Webera Hedwig)
Mostly paroicous or dioicous: paraphyses mostly present and filiform:
robust to weak, gregarious, or cespitose: stem mostly red; leaves more or less
tufted on the fertile shoots, linear-lanceolate to lanceolate, non-bordered,
towards apex more or less toothed; costa mostly incomplete; cells narrowly
rhomboid-hexagonal to linear, the basal slightly more lax: seta long, slcnd'r,
tortuous and twisted, at apex hooked or curved; capsule cemuous or pendu-
lous, rarely erect, with short collum, obovate to oblanccolate or long-clavate;
nnnulus mostly biseriate; peristome inserted near the mouth; teeth yellowish,
papillose, with border narrow or none; segments mostly about as long, rarely
rudimentary, often with a low basal membrane, often narrow, usually split bu:
not fenestrate, cilia non-apf>endiculate, often rudimentary or lackina; spores
mostly small; operculum convex-conic, umbonate or apiculate.
A world-wide genus of about 120 species, inhibiting soil, rocks, and decay-
ing wood. About 40 species in North America; at least 5 species in our
range.
Jennings: Manual of Mosses — 14. Bryaceae 123
Key to the Species*
A. Upper leaf-cells ver>i narrow: inner peristome with a low basal membrane, usually
complete narrow segments, and cilia various, often rudimentary or none B
A. Leaf-cells linear to moderately narrow: basal membrane comprising one-third to one-
half the height of the inner peristome; segments split, cilia well developed C
K. Paroicous; capsule long and slender with a long slender collum 1. P. elongata
B. Polyoicous; capsule oblong to pyriform with a rather short collum 2. P. cruact
C. Paroicous; not bearing gemmae: costa hardly reaching the serrate apex D
C. Dioicous; often bearing gemmae: costa incomplete or percurrent E
D. Cilia two, not sub-appendiculate. articulate 3. P. nutans
D. Cilia three, distmctly sub-appendiculate 3a. P. nutans var. triciHata
E. Costa not reaching apex except irj uppermost leaves; capsule small and very short;
stem not reddish 4. P. pulchella
E.Costa percurrent: capsule larger and elongate-pyriform; stem reddish below
5. P. annotina
1. POHLIA ELONGATA Hedwig
( Webera elongMa Schwaegrichen )
Gregarious to respitose, bright pale green: stems erect, up to 2 cm high,
branching towards base; leaves crowded and larger in the comal tufts, lanceo-
late, erect-spreading, thin, the margin recurved below, the apex gradually
narrowed, serrate; costa brownish, vanishing below or at the apex; leaf-cells
narrow, linear-rhomboidal and vermicular above, hexagonal-rcctangulsr below:
seta long, slender, 2-4.5 cm high; capsule sub-erect to horizontal, narrowly
elliptic, pale, 2-5.5 mm long, the neck slender and longer than the rest of th;
capsule, when dry and empty the capsule constricted below the mouth; oper-
culum conic-acuminate or acutely rostellate; outer peristome yellowish, the
inner with a basal membrane about one-third as high as the teeth; cilia two.
more or less weli-devcloped but always short ?.nd never appendiculate; paroi-
cous; antheridia in pairs in axils of upper leaves: mature in August.
On earth and among rocks, in crevices, etc., where moist, usually in the
mountains. Europe, northern Africa, Asia, North America from G;eenland
to the northern United States and southwards, in the mountains, west to
British Columbia. Rare in our range.
We have seen no specimens from Pennsylvania, but it is reported as follows: Mc-
Kean Co.: Bradford. (Porter's Catalogue).
2. PoHLlA CRUDA [Linnaeus] Lindberg
{Webera cruda Bruch)
Robust, up to 6 or 7 cm high, glaucous green and shining above, brownish
below: st.:ms red, simple, cespitosc; the leaves below ovate, becoming linear-
lanceolate in the comal tuft, erect-spreading, serrate towards che moderately
acute apex, margin plane, rather rigid; costa reddish at base, not re?_ching apex;
leaf-cells linear-prosenchymatous above, larger and rectangular at base, where
* Pohlia carnea (L.) Lindberg has now been found in New York, Ohio, Illinois, and
Iowa. It has widely hexagonal-rhomboid, thin-walled cells up to .018-. 025 x .100 mm,
narrowed towards edge; no annulus; capsule dark red, very short (1.5 mm), oval; seta
thickened at top. The leaves are small and narrowly lanceolate.
124 American Midland Naturalist Monograph No. 6
often reddish: seta long; capsule oblong, with inconspicuous neck, often unsym-
metric. mostly horizontal, castaneous or red-brown, when dry and empty
ventricose at base and constricted at the mouth; operculum conic-apiculate;
peristome yellowish, basal m.embrane not more than one-third as high as teeth,
rilia two or three, well-developed; usually autoicous, sometimes synoicous or
dioicous: spcres mature in summer.
On shaded earth, clefts in rocks, etc., usually in mountainous regions.
Cosmopolitan but local in its distribution. In North America it etxends from
North Carolina and Tennessee to the far North. In our region rare, being
unknown from western Pennsylvania, but reported from the eastern part of
Pennsylvania and from Painesville, northeastern Ohio.
3. POHLIA NUTANS [Schreber] Lindberg
{Webera nutans Hedwig)
Plate XXI
More or less densely cespitose, usually dark green: stems about 1-2 cm
high, branching by lateral innovations, or from the, base, erect, matted with a
castaneous tomentum below, reddish; leaves ovate below to much longer and
linear-lanceolate in the comal tuft, hardly decurrent, the comal long-acumi.iate,
the margin often somewhat recurved below, denticulate towards apex, leaves
somewhat shrunken, twisted and lustrous when dry; costa strong, reddish,
ending in or a little below apex; leaf -cells long-rhomboid and more or less
pointed and prosenchymatoiis above, rectangular below, slightly narrower
towards the margin: seta slender, flexuous, usually 2-3 cm long, lustrous,
castaneous below, often yellowish above; capsule horizontal to sub-pendulous,
oblong to obovate, usually about 3-4 mm in length, with a distinct but short
neck, often gibbous when dry and then contracted below the wide mouth,
yellowish to brown in age; operculum convex-mammillate; peristome-teeth orange-
yellow below, paler and papillose in the rather abruptly narrowed upper half,
strongly trabeculate, lamellate, divisural zigzag and distinct; segments about
as long, carinately split and gaping below but remaining unsplit at apex; cilia
nearly as long, two in number, filiform, articulate, basal membrane half as
high as teeth; annulus wide, revoluble; spores smoothish, yellowish-pellucid,
about .014-.016 mm, mature in early summ.er: autoicous; antheridia in axils
of upper leaves.
On various habitats in moist pbces or swampy fields and woods. Common
and nearly cosmopolitan.
This species is now known from the following counties: Allegheny, Armstrong, Beaver,
Cambria, Centre, Crawford, Erie, Fayette, Indiana, Lawrence, Mercer, Montour, McKean,
Somerset, Washington, and Westmoreland counties. Specimen figured: Presque Isle, Erie
County, May 8-9, 1906. O.E.J.
3a. Pohlia nutans var. triciliata New Combination
{W. nutans var. triciliata Jennings)
Plate XXII
Plants laxly to densely cespitose, shining, dark green to yellowish: stem
simple or sparsely branched, erect, castaneous, at the base reddish-radiculose,
Jennings: Manual of Mosses — 14. Bryaceae 125
about 6-15 mm high; lower leaves short, about 0.5-1.0 mm long, costate
almost to the apex, ovate, above the leaves becoming relatively longer and
ovate-lanceolate, denticulate towards the apex; upper leaves clustered, erect-
spreading, 2.5-4.0 mm long, lanceolate, sub-decurrent, denticulate at apex,
acuminate, non-margined, strongly percurrently to excurrently costate; peri-
chaetial leaves elongate-lanceolate to linear, long-acuminate, denticulate at apex,
excurrently costate; cells of the lower and median leaves incrassate, above the
middle oblong-hexagonal to rhomboidal, about .010-.015 x .035-. 065 mm
elongate-rectangular at base where about .008-. 020 x .0^0-. 100 mm; cells of
the coma! and perichaetial leaves incrassate, elongate to linear-prosenchyma-
tous, about .008-. Oil x .040-. 065 mm, towards the margin gradually narrower
and there reaching .003 -.006 x .080-. 100 mm, at the base elongate-rectangular:
inflorescence paroicous, terminal; antheridia in the axils of the comal leaves:
pedicel solitary, slender, lustrous, castaneous, erect, flexuous, about 4-6 cm
high; capsule horizontal to sub-pendulous, smooth, castaneous to yellowish-
brown, ovate-oblong, often very slightly curved, 2.5-3.5 mm long, when dry
and empty contracted under the mouth, the basal third narrowed into a collum;
exothecial cells incrassate, yellowish-pellucid, irregularly sub-quadrate to oblong-
hexagonal or elongate-rectangular, about .025-. 035 x .035-. 505 mm, in three to
five rows under the mouth abruptly smaller, sub-quadrate, somewhat opaque,
and about .006-. 010 x .012-. 018 mm; annulus broad, revoluble; operculum
rather wide, conic-mamillate; teeth of peristome linear-lanceolate, yellowish,
articulate, strongly trabeculate, narrowly margined above, sub-hyaline and
papillose at apex; segments of inner peristome nearly as long as teeth, hyaline,
granular, carinately split and gaping, cilia three, as long as segments, filiform,
strongly articulate and often sub-appendiculate, hyaline, granular; basal mem-
brane reaching to middle of teeth; spores minutely roughened, yellowish-
pellucid, .012-.015 mm in diameter, mature in Tune.
On earth with more or less humus. Thus far known only as follows:
Butler Co.: On earth under pines on rocky hillside. West Winfield, May 26, 1906.
O.E.J. Cranx'FORd Co.: On hummocks of earrh with Polytrichum, near Hartstown, July
26, 1908. O.E.J. Type Specimen (figured).
4. PoHLlA pulchella (Hedwig) Lindberg
{Bryum LescHrianum Sullivant; Webera pulchella Schimper)
Plate XXII
Gregarious to loosely cespitose, pale green: stems not red, ascending, usu-
ally simple, usually 1-1.5 cm long; leaves small and remote below, gradually
increasing in size and number above, the upper lanceolate, the comal linear-
lanceolate, up to 2.5 mm long, long-acuminate at the serrulate apex, the mar-
gins more or less recurved, the base non-decurrent; costa strong, reddish,
ending below apex; leaf-cells elongaterhomboid-hexagonal, prosenchymatous,
rather thick-walled, and basal often reddish and tending to rectangular, the
marginal slightly narrower: seta erect, 1-1.5 cm long, yel!ov/ish-brown, lus-
trous, slender flexuous; capsule horizontal to abruptly pendent, short, 1.5-2 cm
long, yellowish-brown, the short tapering neck darker brown, capsule pyriform
in general shape, when dry and empty widely flaring at the mouth; operculum
126 American Midland Naturalist Monograph No. 6
conic-apiculate to mammillate; annulus revoluble; peristome rather short, teeth
Hnear-lanceolate, yellowish-pellucid, abruptly narrowed above the middle to a
sub-hyaline papillose apex, divisural and lamellae present, trabeculae strong,
often a few connected by oblique or vertical bars; segments of inner peristome
a little shorter than teeth, carinately split and gaping, cilia usually two, some-
times one, articulate, shorter than segments; basal membrane one-third the
height of teeth; spores minutely roughened, about .015-. 018 mm, mature in
May: dioicous.
On wet clay or sandy soil, Eastern Canada to District of Columbia, Penn-
sylvania and Michigan.
Probably not rare in our region. Allegheny Co.: Power's Run, May 7, 1905.
O.E.J. McKean Co.: Quintuple, May 7, 1896. D.A.B. Westmoreland Co.: On
damp clay with Pogonotum, slope of Chestnut Ridge, Hillside, May 22, 1909. O.E.J,
(figured).
5. PoHLiA annotina (Hedwig) Loeske*
{Webera annotina Bruch)
Loosely cespitose, light green: stems short, 1-2 cm, branching with slender
stiff innovations from the base; leaves below small, lanceolate, non-decurrent,
the upper longer, narrow-lanceolate, acuminate, margins somewhat recurved,
serrulate at apex; costa nearly or quite percurrent, often reddish at base; leaf-
cells rather thick-walled, narrowly rhomboid, small: seta red, flexuous: capsule
small, about 2 mm long, castaneous, the neck about as long as the rest of
capsule, tapering, the whole capsule oval-pyriform, inclined to horizontal;
annulus broad, revoluble; operculum conic-apiculate; mouth wide; peristome-
tceth yellowish, segments widely carinately gaping, cilia in pairs, articulate;
exothecial cells more or less collenchymatous: the sterile stems bearing in the
axils of most of the leaves greenish, sub-sessile, clustered, ovate to ovoid
gemmae with short non-twisted points: dioicous.
Moist, sandy soil, especially among rocks in mountains. Europe, Algeria,
Asia, and, in North America, from Greenland to British Columbia and south
to New England, Pennsylvania, and Kansas.
Rare in our region. Beaver Co.: Lesquereux. (Porter's Catalogue).
3. Mniobryum (Schimper, ex parte) Limpricht
Dioicous, rarely polyoicous: weak to robust, loosely cespitose in brownish
to whitish-green tufts, or gregarious: stems erect, red, radiculose at base; leaves
erect to erect-spreading, the upper lancolate to lance-linear, the apex acute and
distantly serrulate; costa mostly incomplete; cells lax and thin-walled; seta
elongate, when dry sinistrorse, more or less hooked or curved at the top;
capsule more or less pendent, usually short-pyriform, wide-mouthed, almost
turbinate, exothecial cells mostly hexagonal and often broader than high;
annulus none in our species; peristomes equal in length; teeth lanceolate, finely
* Grout, in the Moss Flora of North America pointf. o it thnt Pohlia proligera Lind-
berg is more northern. The moss of our region formerly referred to that :pecies is P
annotina.
Jennings: Manual of Mosses — 14. Bryaceae 127
papillose, not prominently bordered; inner peristome yellowish, the basal mem-
brane constituting one-half or more of its height; segments split, cilia 2-3,
well developed, weakly articulate; spores medium size; operculum medium
size, quite convex, often apiculate.
A genus of about 20 species, distributed over the whole earth, five of these
being in North America, one in our range.
1. Mniobryum Wahlenbergii [Weber and Mohr] Jennings
(M. albicans Limpricht; Webera albicans Schimpcr; Hypniim Wahlenbergii
Weber and Mohr; Pohlia Wahlenbergii Andrews)
Plate XXII
Cespitoss in soft, large, glaucous or whitish-green tufts: stems usually 2-6
cm long, more or less chestnut-red, especially in the older portion, slender,
flexuous. branched and matted tosether with a brownish tomentum at base;
leaves remote below, in the upper portion rather remote, about 2.5 mm long,
when dry somewhat shrunken but hardly twisted, spreading, widely ovate-
lanceolate, at the base narrowed and somewhat decurrent, the margin plane
and serrulate towards the obtusely acute apex; costa strong, reddish, ending a
little below apex; leaf-cells rhombic-hexagonal, pellucid, about .01 5-. 025 mm
wide, slightly narrower towards margin, tending to become inflated and rectan-
gular at base but hardly distinctly so, the lowermost often reddish: seta erect-
flexuous, 2-4 cm high, slender, yellowish to reddish-brown, abruptly hooked at
the summit; capsule pendent, shortly wide-pyriform, about 2.5 mm long, red-
dish-brown when ripe, the neck short and wide, when dry and empty the cap-
sule wide-mouthed; annulus none; peristome-teeth brownish-yellow, pellucid,
strongly trabeculate, the trabeculae often with oblique connections, the lamellae
and divisural indistinct, teeth lance-linear, papillose and sub-hyaline at apex;
segments equal in length to teeth, narrow, carinately split, the cilia 2-4, some-
times more or less connected at apex, nearly as long as segments, papillose,
not appendiculate; basal membrane nearly reaching middle of teeth; spores
smoothish, rather thin-walled, about .01 8-. 024 mm; operculum convex-apicu-
late; exothecial cells irregularly quadrate-hexagonal, yellowish-pellucid, rather
thin-walled, 2-3 rows at mouth much smaller and darker: dioicous; antheridial
flower terminal, discoid, the perigonial bracts wide-spreading; mature in our
region in May.
Almost a cosmopolitan in ditches, springs, or on wet clay banks, etc.
Rarely fruiting but rather common sterile.
Known from Allegheny, Beaver, Butler, Erie, McKean, Washington, and Westmore-
land counties. Specimen figured: Springy places. Quintuple, McKean Co., May 17, 1895.
D.A.B.
4. Bryum [Dillenius] Schimper
Mostly synoicous: paraphyses present, filiform: perennial, small, robust,
rarely gregarious, usually more or less densely cespitose: stem upright to as-
cending, often red, branching below the inflorescence, radiculose; lower leaves
remote, upper leaves tufted, mostly erect-spreading, concave, oval or ovate to
lanceolate, or elliptic to spatulate, mostly acute, often narrowed and decurrent
128 American Midland Naturalist Monograph No. 6
at base, mostly bordered, entire or toothed towards the apex; costa mostly
strong, often excurrent, projecting dorsally, provided with median guides;
leaf-cells mostly rhombic- to rhomboid-hexagonal, the basal parenchymatous,
quadrate to elongate-rectangular; perichaetial leaves narrower and smaller
inside: seta long, reddish to brown, hooked or arcuate at apex, capsule cernu-
ous to pendent, rarely horizontal, the collum distinct, pyriform to cylindric,
rarely ovoid to globose, symmetric to slightly curved, the curve sometimes
being entirely in the collum, phaneropore, annulus present, large-celled, plur-
iseriate, revoluble; the two peristomes of nearly equal length, teeth confluent
at their insertion, lanceolate to linear-subulate, often abruptly narrowed above
the middle, yellowish to orange, often hyaline at apex and sometimes with a
hyaline border, dorsally minutely papillose, trabeculae sometimes united by
cross-partitions; segments mostly free, basal membrane usually high, outwardly
carinate, segments narrowly linear to lanceolate-subulate, split along the keel
and more or less fenestrate or gaping, rarely entire; cilia filiform, rarely short
or lacking, often appendiculate; spores .010-. 050 mm; operculum conic to con-
vex-umbonate or rarely quite apiculate.
A large and difficult genus of about 800 species, of wide distribution; about
175 species in North America, of which there are 9 or 10 species in our
general range.
Key to the Species
A. Leaves usually distinctly bordertd B
A. Leaves not distinctly bordered, at least not above I
B. Costa vanishing below the apex H
B. Costa percurrent to excurrent C
C. Leaves short- to long-decurrent; costa short-excurrent; synoicous or dioicous
8. B. bimum (and 7 B. pseudotriquetrum)
c. Leaves not or but slightly decurrent D
D. Costa fsercurrent to short-excurrent K
D. Costa long-excurrent E
E. Peristome-teeth with the "trabeculae connected by various cross partitions
L B. angustirete
E. Peristome-teeth not so connected f
F. Dioicous G
F. Synoicous 4. B. cusp-datum
F. Mostly autoicous; antheridia at ap)ex of lateral innovations 5. B. pallescens
G. Leaves ovate-lanceolate, with rhomboidal cells 6. B. caespiticium
G. Leaves rounded to widely ovate; cells short-hexagonal 11. B. cap Hare
H. Leaves blunt at apex 3. B. tortifolium
H. Leaves short-acuminate, cuspidate 11. B. capillare
1. Costa long-excurrent 4. B. cuspidatum
I. Costa vanishing below middle of leaf; foliage usually white or silvery
10. B. argenteum
K. Cilia none, or short and coherent 2. B. uliginosum
K. Cilia well developed, appendiculate 9. B. bicolor
1. Bryum angustirete Kindberg
(B. pendulum Schimper; B. cernuum Bryologia Europaea)
Plate XXII
Densely cespitose, usually darkish-green: stems in our region short, about
6-9 mm, erect, sparsely branched, matted below with a castaneous tomentum,
Jennings: Manual of Mosses — 14. Bryaceae 129
stem reddish; leaves rather lax, somewhat tufted, close, erect-spreading, ovate-
lanceolate, long-cuspidateacuminate, more or less recurved on the borders,
reddish at base, usually obscurely denticulate at apex, when any rigid, shrunken
and somewhat twisted; costa stout, reddish, long-excurrent; leaf cells with thick-
ened corners rather small, rhomboidal to somewhat elongate above, at base thin-
walled, rather inflated, rectangular, at margin linear-prosenchymatous in 2-4
rows, forming a weak border: seta usually 3 cm long, slender, flexuous, lus-
trous, castaneous; capsule pendulous, elongate oval-pyriform, usually 4-5 mm
long, tapering below into a neck at least 1.5 mm long, brownish, hardly con-
tracted below the mouth except when dried prematurely; annulus 2-3 seriate,
revoluble; operculum rather small, conic-apiculate; peristome-teeth linear-
triangular, yellowish-pellucid below, sub-hyaline and papillose above, strongly
trabeculate and with prominent oblique or vertical connections between the
plates, the lamellae and divisural indistinct, the inner peristome more or less
closely adherent to the teeth, the segments narrow, the cilia 2-3 and rudi-
mentary, the basal membrane about two-fifths the height of the teeth; spores
large, .024. 030 mm, yellowish-pellucid, irregularly rounded-quadrate to hex-
agonal, the upper four or five rows much smaller, rounded to transversely
elongate, reddish-pellucid: synoicous: mature in June.
On earth, rocks, walls and decaying logs. Temperate regions and moun-
tains of Europe, Algeria, Asia, and North America from Greenland to Alaska
and south to the northern United States, as far as District of Columbia,
Pennsylvania, Ohio, etc.
Rare in our region. Allegheny Co.: Sloping shaly hillside. Fern Hollo.v, Pitts-
burgh, June 8. 1909. G.K.J, (figured).
2. Bryum uliginosum (Bridel) Bryologia Europaea
(DidymoJon cernuum Swartz; Bryum cernuiim Lindberg)
Loosely tufted, green to somewhat brownish: stems branched, more or less
erect, slender; leaves up to 4 or 5 mm long, narrowly lance-ovate, acuminate,
erect-spreading when moist, closely compacted and wavy when dry, only very
slightly decurrent; costa percurrent to short-excurrent; cells rhomboidal-hexag-
onal, thin-walled, with a distinct margin of thickened cells: autoicous, an-
theridial flowers gemmiform, on short branches, inconspicuous: seta slender,
erect, up to 5 cm long; capsule erect to sub-pendulous, more or less arcuate,
oval-pyriform, neck often longer than rest of capsule, slenderly tapering, the
mouth small and oblique; operculum small, low convex-conical, peristome
teeth pale yellow, broad below, slenderly acuminate, papillose, trabeculae dis-
tinct; basal membrane high, segments pale yellow, papillose, gaping; cilia none
or rudimentary; spores castaneous, papillose, about .025 mm in diameter,
mature in summer.
Boggy or sandy places in Eurasia; in North America from northern Canada
south to New York, Ohio, Illinois, and the Southwest. Not yet reported from
our region.
3. Bryum tortifolium Funck
(Bryum obtusifolium Lindberg; B. cyclophyllum Bruch and Schimper)
In low soft tufts, rarely over 2 cm high; leaves distant, scarcely narrowly
130 American Midland Naturalist Monograph No. 6
decurrent, crisped and curled when dry, entire, obovate to sub-orbicular, with
a narrow base; costa slender and banishing below the apex; cells wide, thin-
walled, towards edge forming an indistinct margin: dioiccus: seta short; cap-
sule short-pyriform with wide mouth. A very distinct species on account of
the leaf characters.
Wet places, Eurasia; and from Arctic America south to Pennsylvania,
New Jersey, and northwest. Not yet reported from our region.
4. Bryum cuspidatum Schimper
(B. affine Lindberg)*
Plate XXIII
Rather densely cespitose, becoming dark green: stem short, in ours usually
1-2 cm, occasionally longer, with slender innovations, somewhat matted with
a brownish tomentum, dark brown; leaves rather numerous, somewhat clasping
and short-decurrent, the margins revolute to near the apex where the leaves
are slenderly acuminate and more or less serrulate, the leaves ranging from
oblong-lanceolate below to elongate ovate-lanceolate above and on the branches;
when dry the leaves are moderately shrunken and twisted; costa strong, reddish,
long-excurrent; leaf-cells rhomboid-hexagonal above, to thin-walled, reddish,
and more or less inflated-rectangular at the base, the marginal in two to five
rows of linear-prosenchymatous more or less yellowish-pellucid cells forming
a strongly marked border: seta slender, flexuous, lustrous-castaneous, about
2-4.5 cm in height; capsule about 3 mm long, elongate oblong-pyriform, with
a tapering neck a little shorter than the rest of the capsule, yellowish-brown,
finely deep brown, when dry and empty constricted below the deeper-colored
mouth, more or less pendulous; operculum wide, convex, mamillate; annulus
wide, revoluble; peristome-teeth linear-triangular, yellowish-pellucid below, sub-
hyaline and papillose ^bove, strongly trabeculate, lamellate, the zig-zag divis-
ural usually distinct: basal membrane of inner peristome about half the length
of the teeth, the segments carinately split, hyaline and papillose, a little shorter
than the teeth, the three filiform, appendiculate cilia somewhat shorter than the
segments; spores yellowish-pellucid, minutely roughened, usually about .010-
.014 mm; exothecial cells irregularly quadrate to rectangular-hexagonal, incras-
sate, three or four rows below the mouth being much smaller, rounded-quad-
rate and reddish-pellucid; synoicous: spores mature in summer.
Usually on or between wet rocks. Widely distributed in Europe, Asia, and
northern North America.
Known from the following counties: Allegheny, Beaver, Cambria, Erie, Fayette, Law-
rence, McKean, Somerset, and Westmoreland. Specimen figured: Presque Isle, Erie Co.,
May 8-9, 1906. O.E.J, (figured).
5. Bryum pallescens [Schleicher] Schwaegrichen
Plate XXIII
Sub-cespitose, yellowish-green: stems short, 4-9 mm, sparsely branching,
reddish, somewhat reddish-tomentose below, erect; leaves small and remote
* Also B. intermedium of the first edition of this Manual, B. intermedium being an
Jennings: Manual of Mosses — 14. Bryaceae 131
blow, tufted above, ovate below to ovate- or linear-lanceolate above, non-de-
current to sub-decurrent, erect-spreading, when dry more or less shrunken,
twisted, and appressed, reddish at base, margins recurved, apex obscurely den-
ticulate, cuspidate-acuminate; costa strong, reddish, long-excurrent; leaf-cells
rhomboid to elongate, thin-wal!ed and rectangular at base, the border wide
and formed of several rows of linear-prosenchymatous incrassite yellowish
cells: seta erect, slender, flexuous, lustrous-castaneous, in our specimens about
2 cm long: capsule oblong-pyriform, about two fifths neck, horizontal to sub-
pendulous, contracted below the mouth when dry, brown; annulus wide, revol-
uble; lid conic-apiculate; teeth of the peristome yellowish-pellucid, towards
apex sub-hyaline and papillose, linear-triangular, strongly trabeculate, lamellae
and divisural rather indistinct; segments of inner peristome slightly shorter,
carinately split, cilia 3, stronqly appendiculate, slightly shorter than segments,
basal membrane one-half height of teeth; spores yellowish-pellucid, smoothish
or minutely roughened, about .014-016 mm; exothecial cells incrassate, rectangu-
larly quadrate or hexagonal, the upper three or four rows much smaller and
rounded to transversely elongate and darkly reddish-pellucid: gonioautoicous,
— antheridia in apex of the lateral innovations: mature in May or June.
In crevices of walls and sandstone rocks. Europe, Asia, northern Africa,
North America from Greenland to British Columbia and south to the northern
United States, Ohio and New York.
Rare in our region. Allegheny Co.: On debris in rock-crevices along sandstone clifF
facing the Allegheny River at Power's Run, April 28, 1907. O.E.J, (figured).
6. Bryum caespiticium [Linnaeus] Hedwig
Plate XXIV
Densely cespitose, yellowish-green: stems erect, rarely more than 1 cm high
with us, branching by lateral innovations, brownish-tomentose below, reddish
above; leaves remote and small below, the upper densely tufted, ovate to
lanceolate or narrower within, concave, narrowly acuminate, borders recurved,
the apex slightly denticulate or sometimes entire, the base often reddish, the
comal reaching 3-3.5 x 1 mm, when dry but little shrunken or twisted; costa
strong, reddish, long-excurrent; leaf-cells narrow-rhomboid, becoming larger
and rectangular at base, the marginal in one to three rows of linear-prosen-
chymatous incrassate cells but not forming a very pronounced border; seta
erect, slender, flexuous, lustrous-castaneous, about 2-4 cm long, rarely more;
capsule oblong-pyriform, 3-3.5 mm long, yellowish-brown to darker with age,
the neck comprising almost one-half the capsule, horizontal to pendulous,
sometimes unsymmetrically up-curved, constricted below the mouth when dry
and empty: the mouth darker colored; peristome-teeth yellowish-pellucid below,
paler and minutely papillose above, linear-lanceolate; segments of inner peri-
stome almost as long, somewhat yellowish, carinately split and gaping, cilia as
long as segments or almost so, strongly appendiculate, basal membrane about
two- fifths the height of teeth; spores about .012-017 mm, smoothish, yellowish-
pellucid; exothecial cells incrassate, rectangular below to irregularly quadrate-
hexagonal above, the upper three or four rows much smaller and rounded-
quadrate to laterally elongate and colored; operculum usually orange-brownish,
132 American Midland Naturalist Monograph No. 6
mamillate to conic-apiculate : dioicous: mature in May to June or July.
A cosmopolitan common on earth in pastures, etc., also on dry banks,
stones, walls, etc.
Known from the following counties: Allegheny, Butler, Centre, Clinton, Erie, Hun-
tingdon, McKean, Mercer, Washington, and Westmoreland. Specimen figured: Chestnut
Ridge above Hillside, Westmoreland Co., May 23, 1909. O.E.J.
7. Bryum pseudotriquetrum [Hedwig, p. p.] Schwaegrichen*
(B. ventncosum Dickson)
This species is practically similar to Bryum bimum in everything except
that it is dioicous. According to Dixon and Jameson's Handbook the paler,
more lax-leaved, and more flaccid plants usually belong to B. bimum while
the more rigid and compact specimens are B. pseudotriquetrum, — but this is
not always the case.
This species has much the same habitat and the same range as does B.
bimum, but in our region seems to be rare. In Porter's Catalogue it is reported
from Cresson, Cambria County, by James, and in the Carnegie Museum are
two specimens from two localities in McKean County which were distributed
as this species, but which prove to be synoicous and typically B. bimum.
8. Bryum bimum [Schreber] Bridel
{Mnium bimum Bridel)
Plate XXIII
Rather loosely but deeply cespitose and matted together with a chestnut-
colored tomentum: stems usually 3-6 cm high, rather sparsely branching; leaves
more or less long decurrent, 2-3 mm long, elliptic to oblong-lanceolate, acu-
minate, the margins revolute almost to apex, entire or serrulate above; costa
reddish, strong, percurrent to excurrent; leaf-cells rhomboid-hexagonal, the
basal reddish and inflated-rectangular, the marginal in three or four rows
linear-prosenchymatous and more or less yellowish-pellucid, forming a distinct
border; leaves when dry more or less shrunken, twisted, and appressed: seta
2.5-5.5 cm long, slender, flexuous, lustrous, castaneous; capsule 3-5 mm long,
pendulous, brown, subcylindric, tapering to a neck which is but slightly shorter
than the rest of capsule, slightly constricted below the mouth when dry and
empty, sometimes unsymmetrically up-curved; operculum broad, convex-mam-
illate; annulus large, revoluble; mouth deep chestnut, pellucid; peristome teeth
linear-triangular, yellowish-pellucid below, sub-hyaline and papillose above,
strongly trabeculate, lamellate, divisural zig-zag; basal membrane of inner
peristome half the height of teeth, the segments a little shorter, hyaline, carin-
ately split, cilia 3, strongly appendiculate; spores yellowish, .014-.016 mm,
minutely "punctulate" or granular: synoicous: mature in July.
On wet soil, rocks, or decaying wood, in swamps or other wet places.
Cosmopolitan; in our region more common in the mountains and in the swampy
glaciated regions towards the northwestern corner of Pennsylvania.
* Andrews in Grout (Moss Flora), regards B. bimum and B. pseudotriquetrum as
belonging to one variable species. According to the International Rules B. pseudotrique-
tium has precedence, as to name.
Jennings: Manual of Mosses — 14. Bryaceae 133
Known from the following counties: Allegheny, Butler, Cambria, Centre, Crawford,
Erie, Fayette, Indiana (Porter), McKean, and Washington. Specimens figured: Flinton,
July 23, 1908, and St. Lawrence, Cambria Co., July 24, 1908.
9. Bryum bicolor Dickson
(Bryum atTopuipureum Authors, not Weber and Mohr)
Gregarious, low, dark green; stems erect, reddish; leaves compact, im-
bricate when dry, the upper larger, concave, ovate-acuminate, entire, non-
decurrent; costa usually excurrent in the comal leaves; cells rhomboidal, rather
thick-walled, not forminp a distinct border; dioicous; seta short, about 1 cm,
dark purplish red; capsule pendulous, small, 2 3 mm long, shortly oblong to
obovate, passing quickly into the seta, when ripe dark red; operculum wide, low
conic, apiculate, shining, red; peristome teeth castaneous, strong, rather abrupt-
ly long acuminate, papillose; segments papillose pale yellow, with high basal
membrane; cilia appendiculate; spores described as small, about .010 mm in
diameter, ripening in late spring or early summer.
Said, to occur on moist clay or sandy ground. Northern part of the Old
World and in North America from southern Canada to the Gulf States and
California. Not yet reported from our region.
10. Bryum argenteum [Linnaeus] Hedwig
Plate XXIV
More or less densely cespitose, more or less whitish and silvery green:
stems short, rndiculose, with numerous lateral innovations; leaves closely im-
bricated, deeply concave and so numerous that the branches are terete and
julaceous, leaves small, about 1 mm long, widely ovate or obovate, slightly or
no'; at all decurrent, margins plane, entire, acute to long-acuminate, when dry
silvery shininp and hardly altered in shape; costa thin, wide, disappearing in
upper third of leaf; leaf-cells rhomboid-hexagonal above, below rectangular, all
somewhat pellucid and incrassate, the lower half of the leaf more or less chloro-
phyllose, the upper half colorless: seta s'ender, lustrous, usually chestnut-
colored below, pale above, often dark when old, flexuous, 1-1.5 cm long; cap-
sule about 1.5-2 mm long, oblong, the neck short and hardly tapering, by a
quick turn at the apex of the seta pendent and often touching the seta at its
wider part, somewhat constricted below the rr.outh when dry and empty, dark
brown when old; annulus wide, revoluble; peristome-teeth linear lanceolate,
yellowish-pellucid, hyaline at ap>ex, trabeculate, lamellate, divisural zigzag; seg-
ments nearly as long, carinately split and gaping, faintly yellowish-pellucid,
cilia as long as segments, three in number, appendiculate, basal membrane half
ar high as teeth; exothecial cells quadrate to hexagonal, densely incrassate and
orange-pellucid, the upper eight to ten rows smaller, less densely incrassate,
rounded-quadrate to laterally elongate; operculum convex, apiculate, orange;
spores .010-.014 mm, smoothish, yellowish-pellucid: dioicous: mature mostly
from October to November.
Cosmopolitan, comm.cn on dry earth, crevices of brick or stone pavements
and walls, soil-covered rocks, etc
134 American Midland Naturalist Monograph No. 6
Known from the following counties: Allegheny, Beaver, Crawford, Erie, Fayette, Hun-
tingdon, Lycoming, McKean, Somerset, Washington, and Westmoreland. Sp>ecimen fig-
ured: Mellon's summer home. Laurel Hill Mts., New Florence, Westmoreland Co.,
September 8-11, 1907. O.E.J.
H. Bryum capillare [Linnaeus] Hedwig
(Mnium capillare Linnaeus)
Plate XXIV
Rather densely cespitose, soft, light green: stems low, in our specimen
about 5 mm high, reddish below, radiculose at base, erect, rather stout, some-
times branching at base: leaves rather dense, spreading, not forming a very dis-
tinct comal tuft, soft, widely obovate-spatulate or rounded with a narrowly
oblong base, the apex abruptly acuminate, margins plane; costa rather wide,
reddish at base, in the lower leaves and younger plants ending below the apex
but in upper leaves of older plants excurrent-acuminate to piliferous; leaf-cells
rhomboid-hexagonal, thin-walled, the marginal in one to several rows elongated
and narrow, forming a rather indistinct border, the upper marginal projecting to
form low denticulations, the basal parenchymatous, rectangular, those near
the costa in the middle of the leaf more or less inflated: seta rather long, up to
3 cm; capsule rather large (5 mm) with a distinct neck comprising about one-
third the length of the capsule, which is subcylindric, usually symmetric, hori-
zontal to sub-pendulous, reddish to chestnut-color; operculum conic-apiculate,
reddish-orange; peristome large, reddish; typically dioicous: mature in July
or August.
On leaf-mould and loamy soil in woods, often on bases of trees and on
ledges, — almost cosmopolitan.
McKean Co.: Quintuple, July 15, 1896. D.A.B. (figured). Washington Co.:
Linn and Simonton. (Porter's Catalogue).
5. Rhodobryum (Schimper) Hampe
Dioicous or rarely polyoicous: very robust plants of Mnium-like aspect,
gregarious to loosely cespitose: stem ascending from subterranean rhizome-like
stolons; lower leaves remote, mostly scale-like and imbricated, comal leaves
large, spatulate, bordered, sharply doubly serrate above, forming terminal
rosettes; costa broad, narrowing above and disappearing just below apex in
most species; leaf-cells rhombic- to elongate-hexagonal, at the base elongnte-
rectangular; perichaetial leaves smaller, lanceolate, long-acuminate: seta single
or in twos or threes, elongate, brownish, more or less hooked at apex; capsule
horizontal to pendent, oblong-cylindric, slightly arcuate, collum short; annulus
broad and revoluble or narrow and deciduous in pieces; peristome- teeth con-
fluent at their insertion, lanceolate to linear-subulate, yellowish- to reddish-
brown, hyaline above, somewhat bordered, and finely papillose; segments free,
yellowish, broadly lance-subulate, fenestrate to gaping along the keel; basal
membrane high and carinate outwards; cilia filiform, long-appendiculate; spores
.014-.024 mm; operculum convex-apiculate.
A widely distributed genus of about 5C species; 7 species occur in North
America, one being in our range.
Jennings: Manual of Mossns— 15. Mniaceae 135
1. Rhodobryum roseum (B. &: S.) Limpricht
(Bryum onturiense Kindberg)*
Plate XXIV
Gregarious to loosely cespitose, deep green: stems erect from long creeping
rhizome-like stolons, 2-5 cm high, stout, with minute appressed bract-like
leaves up to the summit, where the leaves suddenly enlarge to form a con-
spicuous rosette about 1 cm across; comal leaves numerous, obovate-spatulate
from a narrow base, the apex suddenly narrowed and acuminate and more or
less twisted, the margin revolute for about three-fourths the length of the leaf
and in the upper part prominently sharply spinulose dentate; costa strong,
ending below the apex but mostly plainly excurrent; leaf-cells rather large,
elongate-hexagonal, the walls medium, towards the base larger, thinner-walled,
more or less hyaline, rectangular: setae 1-3 to a perichaetium, erect, lustrous,
castaneous, 2-4 cm long: capsule pale brownish, oblong-cylindrical, about 4-5
mm long, incurved, somewhat constricted below the mouth when empty, at
the base having a narrow incurved collum about one-third the length of the
rest of the capsule, the capsule horizontal to sub-pendulous; peristome-teeth
large, linear-lanceolate, narrowly bordered, yellowish, hyaline and papillose
above, strongly trabeculate, lam.ellate with distinct divisural; segments about
four-fifths as long, carinately split and gaping; cilia 3, about as long as seg-
ments, strongly appendiculate; basal membiane about two-fifths the height of
teeth; spores yellowish, minutely roughened, about .020 mm; operculum convex-
apiculate; exothecial cells incrassate, rectangular to irregularly rounded, towards
the mouth in several rows very much smaller, very strongly incrassate and
darker; dioicous: mature in September and October.
On rotten logs and rich humus in woods, sometimes on stones. South-
eastern Canada and northeastern United States. Rarely found in fruit.
This interesting moss is now known from 12 counties in western Pennsylvania and
probably will be found in all. Specimen figured: Moon Twp., Allegheny Co., 1889.
J..A.S.
Family 15. Mniaceae
Synoicous or dioicous, rarely autoicous; male flowers disk-like with club-
shaped paraphyses: female flowers bud-like with filiform paraphyses: mostly
robust, cespitose: stem with a central strand, radiculose below, mostly erect,
frequently stoloniferous; comal leaves large and mostly spreading in a terminal
rosette, lower and stoloniferous leaves smaller and somewhat dissimilar: costa
strong, broad at base, tapering upwards and ending below or in the apez, rarely
toothed dorsally; cells parenchym.atous, mostly hexagonal or rounded, smooth,
uniform in size or gradually smaller in size, non-margined, costate: seta long,
stiff, smooth, mostly shortly hocked above; capsule mostly cernuous or pendent,
rarely erect, symmetric, oblong-ovoid to cylindric, rarely globose, sometimes
arcuate, collum short; annulus mostly biseriate and revoluble; peristome double
* The American Rhodobryum ontanense (Kind.) Paris is supposed to differ from
R. roseum in that the latter has the costa ending below the apex of the leaf. Andrews, in
Grout's Moss Flora, is probably correct in not regarding our American plants as specifi-
cally different from the European R. roseum.
136 American Midland Naturalist Monograph No. 6
and mostly complete as in Bryum; spores mostly large; operculum convex to
obliquely rostrate; calyptra cucullate, narrow, mostly fugacious, smooth.
Distributed over the whole earth, most abundant in damp woods and
swamps, on earth, bark of trees, or rocks, in the temperate zones. Five genera,
of which but one occurs in our region.
1. Mnium Linnaeus, Hedwig
Synoicous or dioicous, rarely autoicous: mostly robust, cespitose in bright
green to dark green or later brownish tufts; stem erect, often stoloniferous,
often bearing creeping flagelliform branches; leaves bract-like and remote below,
increasing upwards to the terminal rosette, broadly ovate, obovate, or oblong,
to spatulate from a narrow decurrent base, when dry contorted to crispate, when
wet erect-spreading to recurved, mostly with a border of 1-3 layers of elongate
prosenchymatous colored cells, each layer of the border usually sharply serrate;
costa stout; cells rounded to hexagonal, often collenchymatous and punctate,
uniform or smaller towards the margin: seta single or clustered, long; capsule
cernuous to pendent, rarelv erect, mostly oblong-ovoid, rarely arcuate; exothe-
cial cells rounded, annulus revoluble; teeth 16, strong, separate at base, green-
ish-yellow to reddish-brown, more or less papillose, the zigzag divisural line
distinct, the dorsal plates low, the trabeculae numerous, often united by spor-
adic cross-walls; inner peristome mostly yellowish-red, the basal membrane
half-v/ay to the apex and sometimes perforate; segments usually as long as the
teeth, lanceolate, mostly abruptly subulate, usually fenestrate and finally gap-
ing; cilia usually in 3's com.plete, mostly articulate; operculum convex to conic
and rostrate; calyptra narrowly cucullate.
About 80 species, cosmopolitan, on various sub-strata, usually in moist or
shaded situations; about 30 species occurring in North America, about 11
species in our range.
Key to the Species
A. Leaves not cfistinctly bordered B
A. Leaves distinctly bordered C
B. Margin with a single series of low irregular teeth in the upper half; cells incras-
sate 10. M. stellare
B. Margin not distinctly toothed; cells thin-walled 12. M. cinclidioides
C. Leaves with entire or almost entire margin D
C. Leaves with serrate margin G
D. Border indistinct and of one series of cells only 12. M. cinclidioides
D. Border of 1-4 series of cells in 1-4 layers E
E. Lid acutely rostrate; leaves obovate F
E. Lid conic-apiculate; oblong to oval or sub-orbicular 9b. M. affine var. rugicum
F. On stones; leaves usually minutely apiculate and percurrently costate
1 1. M. punctatnm
F. In swamps; leaves not usually apiculate and costa not usually reaching apex;
often very large 11. M. piinct. var. elatum
G. Leaves serrate with a single row of teeth H
G. Leaves serrate with a double row of teeth K
H. Leaves serrate only in upper two-thirds O
H. Leaves serrate to the base or very nearly so I
I. Teeth slender and usually of 2-4 cells 9a. M. affine var. ciliare
I. Teeth usually of but one cell and not so slender J
Jennings: Manual of Mosses — 15. Mniaceae 137
J. Leaf broadly obovate, ajjex bluntly rounded, abruptly apiculate or cuspidate; teeth
blunt 6. M. Tostratum
J. Leaf apex more or less acute and cuspidate; teeth sharp 8. M. medium
K. Leaves lanceolate to elliptic-ovate; costa incomplete, dorsally toothed 1. M. hornum
K. Leaves oblong-ovate or wider than lanceolate; costa usually complete in upper leaves
at least L
L.Costa toothed dorsally M
L.Costa not toothed dorsally N
M. Cells not collenchymatous, about .014-. 020 mm 2. M. orthorrhynchum
M. Cells follenchyamtous, about .020-. 030 mm in diameter M. lycopodioides*
N. Cells rounded and strongly collenchymatous 3. M. serratum
N. Cells angled and not collenchymatous 4. M. spinulosum
O. Cells thin-walled, hexagonal, not collenchymatous 5. M. Drummondii
O. Cells somewhat incrassate, round-hexagonal, collenchymatous 7. M. ciispidatum
1. Mnium hornum Linnaeus, Hedwig
(Astrophyllum hornum Lmdberg)
Plate LXV
A robust species in dense tufts, up to 6 or 8 mm high with, erect un-
branched stems and terminal rosettes of leaves which reach a length of 3-5 mm
but are narrowly elliptic-lanceolate to elliptic-ovate, acute, sharply apiculate,
all leaves with a reddish, thickened border, sharply doubly spinosely serrate in
the upper half; the costa ending below the apex and spinose dorsally above;
leaf-cells angular, not very regularly seriate, rather small: seta solitary, 2-3 cm
long; capsule subpendulous, finally horizontal, ovate- elliptic and tapering to a
distinct neck, when old pale yellowish with a red mouth; operculum conic-
apiculate: dicicous, the antheridial flowers being disc-like. The leaves are
proportionally narrower than the other species of the genus and the calyptra
often remains for a tim.e clasping the seta just below the capsule, mature in
late spring or summer.
On shaded soil and rocks and banks of streams: Europe, Algeria, Japan,
North America from Newfoundland to Georgia and west to Ohio and Ten-
nessee. (Lesquereux and James in their manual say: "More generally on
quartz or schistose rocks.") but in our region usually among rocks on sand
where often subject to flooding.
Fairly common in the mountains of Pennsylvania » especially in the Ohio Pyle region
of Fayette County, on sandy soil along the river. Now known from Bedford, Cambria
(Porter), Clarion, Clearfield, Elk, Fayette, Forest, Somerset, Venango, Warren, and
Westmoreland counties. Specimen figured: One mile up Meadow Run, Ohio Pyle. June
23, 1940. Chas. M. Boardman.
2. Mnium orthorrhynchum Bridel
(Astrophyllum orthorrhynchum Lindberg)
Quite similar to M. serratum but the leaf-cells only about .01 5-. 020 mm;
densely tufted; leaves close, oblong-ovate, doubly spinose-serrate from below
the middle- costa usually ending in the apiculation, toothed dorsally above; leaf-
* Mnium lycopodiaides (Hooker) Schwaegrichen, as reported from Blair and Elk
counties in our region (Porter's Catalogue), has the stem winged with decurrent leaf-
bases; strong reddish, dorsally toothed costa; cells hexagonal and de?.nitely collenchyma-
tous. Eurasia, and rather rare in the northernmost United States.
138 American Midland Naturalist Monograph No. 6
cells angular, hexagonal to quadrate, non-collenchymatous : seta solitary, red;
capsule elliptic-oblong, abruptly contracted into the neck, straight and more or
less horizontal, brownish; operculum shortly rostrate: dioicous, antheridial
flower discoid. Mature in late summer.
On moist rocks, usually calcareous, along cool shaded ravines and streams:
Europe, Asia, North America from Greenland to British Columbia and south
through the northern part of the United States to North Carolina.
One report in our region. Blair Co.: Porter. (Porter's Catalogue).
3. Mnium serratum Schrader, Schwaegrichen
(M. marginatum Beauvois; Astrophyllum marginatum Lindberg)
Plate XXV
Loosely cespitose in soft tufts, rather dark green: stems and lower leaves
often deep reddish tinged, stems slender, rather short, usually 1.5-3 cm in our
spcimens, simple or branched below with erect branches; leaves rather remote,
strongly decurrent, the lower ovate-lanceolate, the upper oblong spatulate-
lanceolate, all acuminate, the strong red border sharply doubly serrate, the
leaves when dry more or less twisted but hardly crispate; costa in upper leaves
confluent with the border in the apiculus but in the middle and lower leaves
and often even the upper leaves of sterile shoots the costa ends below the apex,
not spinose; leaf-cells from .020-.035 mm in diameter, irregularly rounded,
somewhat incrassate, strongly collenchymatous, the basal elongate: seta mostly
single; capsule horizontal, yellowish to brown, oval-oblong, tapering at neck;
peristome yellow or sometimes brown, inserted, the teeth lance-linear, pellucid
yellowish-brown, papillose above, strongly trabeculate, divisural faint; segments
a little shorter than teeth, papillose above, slender, cilia 3 (2), the basal mem-
brane reaching somewhat above the middle; spores rounded, about .025-.030
mm; operculum stoutly short- rostrate; synoicous; mature in spring.
Usually near streams on shaded banks or rocks, or in crevices of rocks
where moist, in Europe, northern Asia, and, in North America, from Anticosti
to Alaska and south to Tennessee, Missouri, and the Southwest.
Now known from the following counties: Allegheny, Armstrong, Butler, Elk, Fayette,
McKean, Somerset, Washington, and Westmoreland. Specimen figured: Hawkins and
Quintuple, McKean Co. D.A.B. Aug. 2, 1895.
4. Mnium spinulosum Bryologia Europaea
Plate LXV
Tufted, erect, 1-1.5 cm tall, drying a bright emerald green; stems reddish
and radiculose below; similar in many respects to Mnium serratum,, the lower
leaves small and scale-like, the middle and upper abruptly larger, elliptic to
obovate or spatulate at the apex of the stem, decurrent, acute, sharply doubly
serrate on the thickened reddish border in the upper two-thirds, not crisped
when dry; costa strong, sometimes reddish, percurrent, not dorsally toothed,
often ending below the apex in the lower and middle leaves; leaf-cells about
.020-. 030 mm, angled hexagonal, or below rectangular, incrassate, non-collen-
chymatous: synoicous: sporophytes single (or clustered), seta erect, usually
reddish yellow, 1.5 to 2 cm high, abruptly hooked at the top and abruptly
Jennings: Manual of Mosses — 15. Mniaceae 139
enlarging into the short neck of the cylindric-eHiptic capsule; operculum with
an upturned slender beak about as long as half the diameter of the capsule;
capsule light yellow, 2-3 mm long with red rim at mouth; peristome teeth
reddish, lanceolate, minutely papillose, strongly and closely trabeculate below,
inner peristome orange; basal membrane nearly one-half height of teeth; seg-
ments broad reaching to about three-quarters height of teeth, irregularly and
shortly carinate; cilia 3 (2) almost equalling teeth, nodose; spores variable,
rather thick-walled, smoothish, elliptic to spherical, .015-.024 mm, ripening in
late spring or early summer.
On the ground in evergreen woods, usually in mountainous or hilly regions.
Europe and northern North America, from Nova Scotia to Alaska and south
to the northernmost United States. It is reported from eastern Pennsylvania
and Maryland, and Ohio, but only once from our region.
Clearfield Co.: One mile north of Clearfield, Lawrence Twp., on humus soil in
hemlock woods. Sidney K. Eastwood, July 4, 1936 (figured). Capsules deoperculate but
still full of spores.
5. Mnium Drummondii Bruch and Schimper
Bright green, loosely tufted, about 2 cm high, brownish radiculose below;
sterile stems more or less stoloniform; leaves slightly crisped when dry, decur-
rent, broadly obovate-spatulate, acute to short-acuminate, bordered by 1-4 rows
of cells, serrate only above, the teeth single and sharp; costa percurrent; leaf-
cells thin-walled, non-coUenchymatous, hexagonal, about .035-. 040 mm; seta
slender, reddish; capsules 1-3 (or 4) from flower, pendulous, oblong, yellow-
ish; operculum convex-apiculate; outer peristome yellow, papillose; segments
about as long; cilia 2 or 3, somewhat appendiculate; spores roughened, yellow-
ish, ripening in late spring or early summer.
Shaded moist rocks or soil; northern Eurasia, and from southern Canada
south to Maryland and Pennsylvania. Not yet reported from our part of the
State.
6. Mnium rostratum Schrader, Schwaegrichen
{Astrophyllum rostratum Lindberg )
Large, loosely cespitose, stoloniferous: stems erect, short, the sterile shoots
creeping or arched; leaves broadly oblong or obovate, rounded at both ends,
tapering but little at base, at the apex very broadly rounded or almost truncate,
short apiculate, the border strong, brownish, serrate in at least the upper half
with a single row of short obtuse or almost obsolete teeth; the comal leaves
large, up to 5 mm long, those of the sterile shoots complanate-two-ranked;
costa excurrent in the short apiculus; leaf-cells incrassate, collenchymatous,
about .025-.030 mm, rounded-hexagonal, not radiating in rows from the costa
as in ajfirte var. rugJcum, which in the sterile condition it closely resembles:
capsules usually 1-3, clustered, sub-pendulous to horizontal, yellowish, oper-
culum long-rostrate; peristome-teeth papillose, yellowish, the inner peristome
orange; synoicous: mature in spring to early summer.
On wet rocks and. earth in woods: almost cosmopolitan in the temperate
zones, in North America from central and southern Canada south to Virginia,
Pennsylvania, Ohio. Montana and Oregon, but apparently rather rare.
140 American Midland Naturalist Monograph No. 6
All Pennsylvania specimens in the Carnegie Museum Herbarium which
were labeled M. rostratum are non-collenchymatous and the leaf-cells are defi-
nitely arranged in series radiating from the costa.
The specie'^ is reported from our region as follows: Cambria Co.: James. (Porter's
Catalogue). Lycoming Co.: McMinn. (Porter's Catalogue).
7. Mnium cuspidatum Linnaeus, Hedwig
(Mnium sylvaticum Lindberg)
Plate XXV
Loosely cespitose in large light to dark patches: stems branching with
sterile shoots prostrate or sub-erect, in our specimens usually about L5-3 cm
high, reddish, radiculose below; leaves decurrent, oblong-oval, acute, the upper
tending to obovate, those on the branches more rounded or oval, all shortly
cuspidate and serrate in the upper half or two-thirds with a single row of short
one-celled teeth, occasionally some teeth two-celled, the border of 3-5 rows of
incrassate, linear, yellowish-pellucid cells; costa confluent with the border in
the apiculate apex or ending a little below the apex; leaf-cells about .020-.025
mm, incrassate, collenchymatous, hexagonal to somewhat rounded, the basal
tending to rectangular: seta solitary, pale yellowish or brownish, erect; capsule
pale yellowish or brownish, sub-pendulous, oblong-oval, rather abruptly nar-
rowing to the seta, the base and mouth brown; operculum conic-obtuse; teeth
yellow, lance-linear, papillose above, divisural indistinct; inner peristome a little
shorter, the basal membrane extending to the middle or a little above, the
basal part of the segments more or less irregularly fenestrate with rounded
holes, the upper part of the segments finally gaping or breaking apart; cilia
three, Unear, somewhat appendiculate, the inner peristome brownish-pellucid,
the tips of the segments and the cilia being paler and papillose; spores rounded,
faintly papillose, yellowish, about .025 mm in diameter: synoicous, mature
in May.
In moist woods on earth, stones, rotten logs, etc. Common and widely
distributed over the temperate parts of Europe, Asia, and of North America.
Common and now known from seventeen counties in western Pennsylvania and prob-
ably occurs in all. Specimens figured: Ohio Pyle, Fayette Co., May 30, 1908. O.E.J.
8. Mnium medium Bryologia Europaea
(Astrophyllum medium Lindberg)
Plate XXV
Widely and rather loosely cespitose, large, light to dark green: stems
erect, up to 5 cm in our specimens, branching at the base, densely covered with
a brovm felted tomentum, sterile shoots long and prostrate or ascending; leaves
distant, shriveled when dry, ovate to oblong, somewhat narrowed and slightly
decurrent at base, rather obtuse at apex, cuspidate, narrowly margined all
around, sharply serrate from near the base with mainly one-celled teeth, the
comal leaves rosulate, and up to 5 x 15 mm; costa reddish, strong, percurrent
cuspidate; leaf-cells large, rounded above to elliptic-hexagonal towards base,
the margin consisting of about two rows of linear, much incrassate, more or
less colored cells, the laminal cells all incrassate and collenchymatous and in-
Jennings: Manual of Mosses — 15. Mniaceae 141
creasing in size towards the costa: synoicous: capsules clustered, occasionally
single, on erect stout setae, pendent, oblong; operculum convex, rostrate-apicu-
late: mature in May.
Mostly on wet rocks and shaded damp earth and logs; cooler Europe and
Asia, and, in North America, from Greenland to Alaska and south to New
Jersey, Pennsylvania, Minnesota, Idaho to California.
Armstrong Co.: Ravine, west bank of Allegheny River, 1 mi. north of Foxburg.
June 10, 1934. Chas. M. Boardman. Beaver Co.: South branch, Brady Run, 3 mi.
west of Fallston. Jan. 28, 1934. Charles M. Boardman. Butler Co.: On wet rocks,
Sawmill Run, Butler Twp. Nov. 4, 1934. Sidney K. Eastwood. McKean Co.: On
leaf-mold, etc., at headwaters of Marilla Brook in wet, springy places, September 24.
18Q4 (figured), West Branch Swamp, May 26. 1895, Bradford, November 2, 1898. All
D.A.B.
9. Mnium affine Blandow, Schwaegrichen
(Astrophyllum cuspidatum Lindberg)
As Grout points out in his "Mosses v/ith Hand-lens and Microscope,"
the true Mnium ajfine Blandow is rare in eastern United States. Andrews,
in Grout's Moss Flora, regards M. affine as so variable as to make it impos-
sible to distinguish the various published varieties. It has the capsules usually
clustered, 2-4 together, and the teeth of the leaves shorter than in the variety
ciliare. Its general range is Europe, Asia, and North America south to New
Jersey, West Virginia, and Washington.
Now known from Erie. Forest, and Washington counties. Specimens figured: In ravine,
Snake Woods near Washington, May 6, 18934 and Nov. 11, 1893. A. Linn and J. S.
Simonton.
9a. Mnium affine var. ciliare (Greville) C. Mueller
{Astrophyllum ciliare Lindberg; Bryum cilure Greville)
Plate XXVI
Moderately large, loosely cespitose, pale to dark green with age: stems
erect, usually about 3 cm high, reddish-brown, rather stout, radiculose below,
with long, slender sterile shoots which are prostrate or arched; stem-leaves
ovate, varying to oblong-elliptic or at the apex rosulate and obovate to narrow
spatulate, somewhat acute, apiculate, up to 6-10 mm long, decurrent, margined,
serrate down to the narrowed base v/ith long slender teeth of 2-4 cells; costa
excurrent-apiculate, strong; leaf-cells large, .020-040 (-.070) mm in diameter,
angled, somewhat incrassate, hexagonal to irregularly somewhat elongate rectan-
gular, especially towards the base, hardly collenchymatous, marginal cells
prosenchymatous-linear and cartilaginous pellucid, often yellowish to reddish:
seta single, erect flexuous, strong, reddish, about 2.5 cm long; capsule pendent,
elliptic-oblong, about 4 mm Icna, narrov/ed to a short drrker c'^rer' neck,
yellowish-brov/n; lid conic-apiculate; peristome teeth pale pellucid, strongly
trabeculate, the divisural rather faint, finely papillose above; inner peristome
brownish pellucid, the basal membrane reaching about half-way, non- fenestrate,
the segments and usually three cilia finely papillose above and often exceeding
the teeth; spores round, yellowish-pellucid, finely papillose, about .028-.030 mm;
dioicous; nntheridial flower terminal-discoid; mature in May.
142 American Midland Naturalist Monograph No. 6
On rocks and soil in swamps and. moist woods, Asia, Europe, and in
North America through southern Canada south to Georgia, Louisiana, Mis-
souri, Montana, and California.
Known from the following counties: Butler. Cameron, Clarion, Crawford, Erie, Fay-
ette, McKean, Somerset, Venango, Warren, Washington, and Westmoreland. Specimen
figured: Ohio Pyle, Fayette Co., September 1-3. 1906. O.E.J, and G.K.J.
9b. Mnium affine var. rugicum (Laurer) Bryologia Europaea
{Astrophyllum rugicum Lindberg)
Plate XXVI
Darker green than true affine, almost blackish: stems short, usually simple;
leaves oblong to broadly oval or sub-orbicular, the apex blunt and rounded
with an apiculation or almost entire, the margin little or not at all serrate:
capsule much as in affme var. ciliare but usually smaller. The leaves often
very closely resemble those of M. rostra turn but Grout says the leaf-cells have
thinner walls in rugicum and also radiate in more or less definite series from
the costa, while in ro stratum the thick- walled cells are irregularly arranged, or
at least not in radiating series.
In cool, shaded ravines and swamps; Europe, and, in North America, from
Greenland to Alaska and locally south to Louisiana and Colorado.
In our region not known to range more than fifty miles east of the western State line.
Allegheny Co.: Power's Run, April 18, 1906, and June 17, 1909, O.E.J.; Wildwood
Hollow, March 29, 1908, and Coraopolis, September 14, 1905. O.E.J, and G.E.K. All
sterile. Beaver Co.s Beaver Falls, May 11, 1907. O.E.J. Sterile. Butler Co.: On
wet log, 5 mi. north of Zelienople, March, 1927. L. K. Henry. Erie Co.: Damp woods
near Erie. Agnes E. Hartman. July 30, 1927. Fayette Co.: Cheat Haven, September
3-6, 1910. O.E.J, and G.K.J, (figured). Ohio Pyle, September 1-3, 1907. O.E.J, and
G.K.J. (Both sterile). Washington Co.: In ravine, Snake Woods near Washington,
June 8, 1895, and under waterfalls below Taylorstown, Nov. 16, 1895. A. Linn and
J. S. Simonton.
10. Mnium stellare [Reichenbach] Hedwig
Plate XXVI
Densely cespitose, soft, deep or bluish-green: stems erect, usually 1-3 cm
high, branching at base; leaves gradually larger above, elliptic-oblong, to sub-
orbicular below, slightly decurrent, rounded and acute at apex to obtuse-apicu-
late, non-bordered, the upper part of the leaf obtusely irregularly short serrate;
costa thin, ending considerably below the ap>ex, smooth on back; leaf-cells in-
crassate, angular, irregular to hexagonal or subquadrate, fairly uniform in size,
about .020. 030 mm; seta solitary; capsule horizontal to inclined, oblong; lid
conic-convex; peristome yellowish; dioicous; antheridial flower discoid: mature
in late spring or early summer.
At the base of trees or on rocks in swampy woods or on humus, in tem-
perate Europe, Asia, and North America, through lower Canada and north-
eastern United States. This species rarely fruits and all specimens from
Pennsylvania thus far have been sterile.
Allegheny Co.: Under side of rocks in crevices, Fern Hollow, Pittsburgh, March
9, 1908. O.E.J. McKean Co.: Rutherford Run, March 12, 1894, and Qumtuple, Sep-
Jennings: Manual of Mosses — 15. Mniaceae 143
tember 9, 1894, and November 13. 1896. D.A.B. Washington Co.: North Branch
of Maple Creek, above Charleroi, April 24, 1908. O.E.J, (figured).
Now known also from Erie and Westmoreland counties.
11. Mnium PUNCTATUM [Linnaeus] Hedwig
Plate XXVII
Rather large., dark green, erect, loosely tufted, 1-3 inches high: stems rigid,
dark, densely tomentose nearly to the apex; dioicous; leaves remote, forming
at the apex a rosette and largest there, spreading, the lower smaller, rounded-
ovate, the terminal about 4-5 x 6-9 mm, broadly obovate, all narrowed to a few
cells at the base, entire, apex usually apiculate, often somewhat emarginate-
apiculate, bordered by a cartilaginous-thickened purplish-brown-pellucid rim of
about 2-5 rows of elongate incrassate cells; costa strong, usually terminating or
percurrent in the apiculus. or sometimes ceasing just below the apex; median
cells rounded- to elongate-hexagonal, about .030-. 040 x .0;0-.085 mm, incras-
sate, often in obliquely ascending series from costa to border, the basal rather
larger, rectangular, slightly inflated, the apical smaller, irregularly angular:
seta 2-3 cm long, erect, f^exuous, purplish-brown, rather lustrous; capsule sub-
pendulous, oval-oblong, yellowish to brown when old, finally when dry some-
what sulcate; operculum conic, acutely rostrate; peristome-teeth yellowish-brown,
pellucid, papillose, trabeculate; segments nearly as high, yellow-pellucid, finely
papillose, the basal membrane reaching to one-third the height, cilia usually
three, slightly shorter than segments; spores smoothish, round, about .030-040
mm; fruiting in spring rather early (April), and sometimes with two or three
capsules to a plant; usuallv dioicous.
On soil and rocks in damp woods, ravines, swamps, etc. Rather common.
Europe, .A.sia, and North America down to middle United States.
Known from the following counties: Allegheny, Armstrong. Butler. Cameron, Clear-
field, Clinton, Crawford, Elk (Porter), Fayette, Forest, Huntingdon (Porter), Lawrence,
MfKean, Mercer, Somerset, Tioga, Venango, Warren, Washington, and Westmoreland,
and probably occurs in all. Specimen figured: Shades, above Blackburn, Westmoreland
Co.. March 25, 1910. O.E.J.
Ua. Mnium punctatum var. elatum Schimper
Plate XXVII
This variety differs typically from true punctatutn in that it grows in muddy
shaded places and swamps, is much larger, — in our specimens reaching a height
of 7 or 8 cm and with leaves up to 10 or 11 mm long, the leaves are rounded
and usually non-apiculate at apex, the border consisting of usually but one
layer of cells, and the costa ending below the apex.
In swamps and muddy shaded places: Europe, Asia, and, in North
America, from the Arctic regions south to Virginia, Michigan, and Idaho. In
Pennsylvania most of the specimens of punctatum show some of the characters
of the variety but we have found no specimen which clearly possesses all the
characters attributed to the variety. The following specimens more or less
closely approach the variety:
Known from the following counties: Allegheny. Clarion. Crawford, Fayette, Lawrence.
144 American Midland Naturalist Monograph No. 6
McKean, Mercer, and Westmoreland. Specimen figured: Houston Junction, Mercer Co.,
July 12, 1902. J.A.S.
12. Mnium cinclidioides (Blytt) Huebener
Plate XXVII
Large, loosely cespitose, bright green, becoming dark when old: stems rigid,
under exceptional conditions reaching 15 or 20 cm or more, our specimens
sterile and about 4-8 cm high, stems dark brownish; leaves remote, thin, large,
the lower ones ovate to oblong and not at all decurrent, the upper rosulate
from a narrow base, widely oblong-lingulate or obovate, rounded and obtuse
v/ith a minute apiculus, more or less undulate, up to 7 or 8 mm long and 4
mm wide in our specimens, margin non-bordered, entire with the exception of
occasionally projecting marginal cells; costa ending considerably below the
apex; leaf-cells rhomboid-hexagonal, arranged in series radiating from the costa,
the marginal gradually becoming linear and parallel to the margin, all rather
thin-walled, chlorophyllose, the larj^est up to about .030 x .100 or .110 mm:
seta long, rather slender; capsule abruptly pendent, shortly oval; lid conic-
apiculate; peristome brownish: dioicous: mature in late spring or early summer.
In bogs, pools, and swamps in the cooler parts of Europe, Asia, and North
America down in glaciated regions to New Jersey and Pennsylvania and south-
ward along the upland plateaus and mountains; generally sterile.
Butler Co.: Wet bank of brook, north of Dougherty's Mills. July 28, 1935. Sid-
ney K. Eastwood. Crawford Co.: In Pymatuning Swamp, Linesville, June 12, 1905.
O.E.J, (figured). Sterile. Elk Co.: Wet wood in swamp. Sept. 2, 1935. Sidney K.
Eastwood. McKean Co.: Sphagnum Swamp, West Branch, July 5, 1896, and July 22,
1894. D.A.B. Sterile. Somerset Co.: Cranberry Glade Run, Laurel Hill mt. Swampy
woods. Elev. 2300 i ft. C.M.B. June 28, 1942. Vena::co Co.: Tarl;iln Run. near Van.
John Wurdack. Aug. 29, 1936.
Family 16. AuLACOMrnACEAE
Dioicous, rarely autoicous: robust to slender, more or less high-cespitose :
stem mostly with a central strand, with one to three innovations below the
apex, also with slender sterile shoots from the older portions; leaves 8-seriate,
gradually larger above, carinate or concave, ovate or oblong to lanceolate or
lance linear, acute to obtuse, non-bcrdered, mostly toothed above; costa mostly
incomplete, with median guides; areolation small, rounded, incrassate, mostly
papillose: sporogonia solitary; seta usually long, erect; capsule cernuous, rarely
erect, oblong to cylindric, with a short coUum, more or less 8-striate, plicate
when dry; annulus present; exothecial ce'ls elongate to rectangular, the longi-
tudinal walls thickened; phanerophore, stomata in the collum only; peristomes
free and essentially as in Byrum; spores .008-. 014 mm; operculum conic to ros-
trate; calyptra narrowly cucullate, Icng-rostrate, split on one side, fugacious.
Inhabiting the colder and temperate parts of the world, in moist habitats
on soil, rocks, trees, etc. The genus Leptotheca with species in the south tem-
perate zone and the following:
1. Aulacomnium Schwaegrichen
With characters as for the family, the stem sometimes bearing flagelliform
pseudopodia, which are leafless or nearly so and bear a cluster of gemmae at
Jennings: Manual of Mosses — 16. Adi.acomniaceae 145
the tips; leaves crowded, decurrent, erect-ascending, the margins more or less
revolute; costa ending below apex; cells each with a central papilla; capsule
somewhat arcuate; annulus 2-4-seriate, revoluble; teeth lance-linear and sub-
ulate-acuminate, yellow to rusty, the divisural zigzag, finely papillose, with
numerous articulations; inner peristome delicate, hyaline; basal membrane one-
half height of teeth, segments lance-subulate, gaping; cilia well developed,
delicate, mostly only weakly articulate.
Nine species widely distributed; five in North America, two in our range.
Key to the Species
A. Autoicous; leaves strongly serrate from the middle upwards 1. A. heterostichum
A. Dioicous; leaves merely serrulate near the apex 2. A. palustrc
1. Aulacomnium heterostichum (Hedwig) Bryologia Europaea
(^AnhenopteTum heterostichum Hedwig)
Plate XXVIII
Rather loosely cespitose, pale to yellowish-green: stems branching by ter-
minal, annual innovations, the annual growth in our specimens being usually
about 8-10 mm, stems brown-radiculose below; leaves usually about 8-10 mm,
stems brown-radiculose below; leaves obovate below to oblong or elongate-
ovate above, often somewhat unsymmetrically inclined, the leaf plane above,
coarsely repand denticulate in the upper half, mostly apiculate; costa strong,
yellowish-brown, ending just below apex; leaf-cells incrassate, median and
apical rounded-quadrate, about .008-.015 mm in diameter, basal similar, quad-
rate to rectangular and 3:1: seta about 6-15 mm long, erect, flexuous, reddish-
brown, smooth, little or not at all twisted; capsule about 2.5 mm long, oblong-
cylindric, arcuate, inclined, reddish-brown, striate, when dry 8-plicate, tapering
below into a short collum: annulate; doubly peristomate; teeth inserted on the
capsule-rim, lanceolate, about 25-30 articulate, distinct to the base, yellowish-
pellucid, rather indistinctly finely horizontally striate-papillate below, seg-
ments of same length or a little shorter, hyaline, more or less completely
carinate-cleft in median portion, united in the lower third with the cilia into a
basal membrane: cilia 3 (2), somewhat shorter, somewhat articulate; spores
pellucid-yellowish, not distinctly papillose, about .010-.014 mm; mature in
May to June; operculum convex, obtusely short-rostrate; calyptra long-rostrate,
cucullate.
On shaded, moist, earthy banks, bases of trees, etc., Japan and in North
America from Ontario to Minnesota, and Texas to Florida.
Known from the following counties: Beaver, Bedford, Butler. Cameron, Clarion, Clear-
field, Erie, Fayette, Green, McKean, Somerset, and Washington. Specimen figured: Ohio
Pyle, Fayette Co., J.A.S. June 15, 1902.
2. Aulacomnium palustre (Weber and Mohr) Schwaegrichen
(Hypnum palustre Weber and Mohr)
Plate XXVIII
Robust, densely cespitose mosses of bogs and moist places on soil or rotten
wood; tufts often 2-3 inches deep, light yellowish-green above, below darker
146 American Midland Naturalist Monograph No. 6
and stem covered with a reddish-brown tomentum; leaves oblong to lanceolate,
about 4 mm. long, minutely denticulate towards the apex, carinate, rather cris-
pate when dry; costa strong, ending just below apex; upper cells small, round
incrassate, unipapillate, basal cells elongate-rectangular or hexagonal, thin-
walled: seta erect, tortuous, in ours about 3 cm long, upper part dextrorse,
lower part sinistrorse; capsule sub-cylindrical, 4-5 mm long, when dry strongly
sulcate, arcuate, constricted below mouth; annulus high, colored at the base;
teeth lance-Iinear, subulate-acuminate, yellowish, trabeculae sometimes united
by oblique walls, divisural zigzags; segments delicate, slightly shorter, hyaline,
cilia about 3, equally long, weakly articulate; spores small, smooth, .008-. 012
mm; mature in early summer; operculum long-conic, straight or sometimes
somewhat recurved.
Cosmopolitan. In swampy woods and bogs. In North America from the
Arctic regions south over most of the United States.
Rather common in the northern part of our region. Known from the folio ving coun-
ties: Allegheny, Armstrong, Beaver, Butler, Cameron, Centre, Clarion, Clearfield, Clinton,
Crawford, Erie, Fayette, Greene, Lawrence, McKean, Mercer, Somerset, Washington, and
Westmoreland. Specimen figured: Presque Isle, Erie Co., May 8-9, 1906. O.E.J.
2a. AuLACOMNlUM PALUSTRE var. IMBRICATUM Bryologia Europaea
Leaves more ovate than lanceolate; imbricated when dry, not crispate or
contorted.
From lower Canada down into eastern United States.
Family 17. MeesiacEAE
Synoicous, autoicous, dioicous, or polycoicous: robust to slender, cespitose:
stem with a central strand, elongate, leaves 3-8 seriate, moderately close, mostly
from an erect base erect-spreading to recurved-squarrose, lance-ovate to lance-
acuminate, non-bordered, sometimes toothed at the apex; costa strong, without
guides, mostly incomplete; cells mostly parenchymatous and smooth, upper
firm-walled, rectangular to rounded 4-6-sided, the basal often thin-walled,
elongate-rectangular and hyaline: seta mostly long and slender, erect, tortuous;
capsule erect, from a long collum elongate arcuate-pyriform, the mouth small
and oblique, never constricted below the mouth; annulus small-celled, 1-2-seri-
ate, loosening itself here and there, rarely persisting; teeth mostly much shorter
than the segments, truncate, more or less completely confluent, with straight
divisural and thin rectangular dorsal plates, the inner layer with low lamellae;
inner peristome with a carinate basal membrane united to the teeth; segments
narrowly linear, alternating with the teeth, cilia rudimentary or none; spores
.032-. 056 mm, mostly finely granulate; operculum small, conic, obtuse; calyptra
small, cucullate, smooth, fugacious.
A small family (3 genera) of mostly bog mosses of the cooler parts of
the northern hemisphere. One genus represented in our range.
1. Meesia Hedwig
Characters mainly as for the family; the tufts green to yellowish-green,
inside brown to blackish: leaves more or less decurrent, acute or obtuse, mostly
Jennings: Manual of Mosses — 18. Bartramiaceae 147
entire; upper cells mostly rectangular, thick-walled, the lower elongate and
hyaline: seta usually very long, inner peristome with a very low basal mem-
brane; segments 2 to 4 times the length of the teeth, often more or less united
at the tip; cilia short or rudimentary, often being represented by a chain-like
series of thickenings on the persisting wall of the inner peristome.
Ten species in bogs and wet places; four species in North America; two
species may be looked for in bogs and swamps in the northern part of our
region.
Key to the Species
A. Leaves three-ranked, serrate; dioicous 1. M. triquetra
A. Leaves 5-8-ranked, entire; synoicous (M. longiseta Hedwig)*
1. Meesia triquetra [Linnaeus] Aongstroem
(M. tristicha (Funck) Bryologia Europaea)
Loosely cespitose, dark green: stems elongate, radiculose below, sparingly
branching; leaves three-ranked, distant, lanceolate to ovate-lanceolate, acute,
strongly iquarrose from a concave strongly decurrent half-clasping base, the
spreading portion carinate, the margins sharply serrate; costa strong, ending in
the apex or just below; upper leaf-cells rectangular to hexagonal, incrassate,
the lower hyaline, elongate-rectangular; perichaetial leaves larger, about six in
number: seta long; capsule pyriform, curved from a long erect collum, when
dry and empty more or less wrinkled and twisted; peristome-teeth 16, short,
unequal, bifid; segments alternate, 16, about three times as long as teeth, united
below into a low basal membrane, yellowish, linear, irregularly articulate and
appendiculate; exothecial cells at mouth very small and in several rows, darker;
lid convex-conic; spores large, ripening in summer.
In bogs and swampy woods, Europe, Asia, and, in North America, from
New Jersey, eastern Pennsylvania, Ohio and Lake Superior, north and west
to Arctic America and the Pacific Ocean.
Family 18. Bartramiaceae
Dioicous or synoicous, rarely paroicous or autoicous: slender to very
robust, cespitose: stems with central strand, erect, dichotomous or more often
with whorled "sub-floral" innovations, leaves 5-8-seriate, little or not at all
decurrent, lance-ovate to lance-subulate, non-bordered, serrate marginally above
and often also on the back of the costa; costa mostly strong, with median
guides, ending below or in the apex or excurrent in a serrate arista; cells paren-
chymatous, round-quadrate to elongate-rectangular, rarely linear, mostly thick-
walled, mostly mamillate on both sides; basal cells either not wider, or lax,
wider, and hyaline, mostly smooth, alar cells rarely differentiated: seta usually
long and straight, little or not at all twisted when dry; capsule erect to cernu-
ous, rarely pendent, more or less globose, darkly striate, collum rarely distinct,
mouth oblique or rarely symmetrical, exothecial cells rectangular to hexagonal,
several series at the mouth laterally elongate; annulus none or very incomplete;
peristome mostly double or sometimes single or rudimentary, or lacking alto-
* In swamps and sphagnum bogs as far south as New York and Ohio.
148 American Midland Naturalist Monograph No. 6
gether; always inserted back from the exothecium by the width of several cells,
peristome-teeth dagger-shape, golden brown to reddish-yellow, mostly non-
bordered, inner peristome mostly shorter, carinate, the basal membrane one-
fourth to one-half the height of the inner peristome; segments at first carinately
gaping, then divergently parted, cilia 1-3, rarely well-developed, sometimes
none, non-articulate; spore-sac very small; lid small, short-conic, rarely ros-
trate; calyptra small, cucullate, smooth, fugacious; spores large, round to oval
or reniform, papillose.
A large family of nine genera; three genera in our region.
Key to the Genera
A. Cilia usually well-developed: stem with a whorl of sub-floral shoots 3. Philonotis
A. Cilia poorly developed or none )i
B. Leaf-cells mamillate or papillose; leaves 5-seriate or pluriseriate 2. Bartramtj
L. Leaf-cells smooth; stem triangular and le.wes 3-seriate I. Plagiopus
1. Plagiopus Bridel
Synoicous: quite slender, laxly to densely cespitose, dull green, becoming
brownish: stem triangular, erect or ascending, the outer layer of cells lax,
hyaline, inflated, the central strand poorly defined, branching above the base,
the shoots of about equal height; leaves spreadina to recurved, somewhat,
twisted but not crispate when dry, from a non-sheathing base narrowly lance-
olate, acuminate, sharply carinate above, the margin usually revolute, doubly
serrate above; costa strong, percurrent, or shortly excurrent, dorsally projecting
and simply serrate upwards; leaf-cells incrassate, smooth, the upper minute,
shortly rectangular and quadrate, basally more elongate and pellucid, the alar
slightly more lax and quadrate: seta 1-L5 cm long, erect, castaneous, not twisted
when dry; capsule erect, somewhat inclined when dry, globose, slightly arcuate,
brown, finely striate, when dry somev/hat shortened at the base and mouth,
slightly curved and stronf^ly plicate; peristome double, the teeth smooth, nar-
rowly dagger-like, reddish-brown in the upper half, with inter-lamellar thicken-
ings, the inner peristome shorter and pale yellow, cilia none; lid small, short-
conic; spores mostly uniform, warty.
A genus of three species: one in New Zealand, one in Java, and the
following:
L Plagiopus Oederi [Gunnerus] Limpricht
(Biyum Oederi Gunnerus; Bartramia Oederi Schwaegrichen;
Bartramia grandiftora Schwaegrichen)
With characters essentially as given above for the genus. The spores
mature in spring.
On moist soil and rocks in shady woods, mainly in non-calcareous and hilly
or mountainous districts; Europe, Asia, and, in North America, from Canada
to eastern Pennsylvania, Illinois, and west to the Rocky Mountains. It may
eventually be found in our region.
2. Bartramia Hedwig
Synoicous, paroicous, autoicous, or dioicous: slender to robust, laxly to
densely cespitose, the tufts often blue-green above, brownish-yellow inside:
Jennings: Manual of Mosses — 18. Bartramiaceae 149
stem with central strand usually distinct, erect, monopodial or dichotomous,
branches not whorled; leaves mostly 8-striate, from a mostly half-sheathing
base gradually or abruptly subulate-linear, serrate upwards and often on the
back of the costa; lamina upwards, sometimes only at the margin, two-layered;
costa strong, projecting dorsally, incomplete to excurrent; leaf-cells small, in-
crassate, rectangular, mamillate on both sides, the basal elongate rectangular
to linear, smooth, pellucid to hyah'ne: seta mostly 1-2 cm long, rarely very
short, mostly straight; capsule mostly inclined, somewhat arcuate, with mouth
oblique, globose, no collum, when dry mostly sulcate, more or less shrunken
in the middle and flattened on the ends; peristome double or single, rarely
none, teeth not united at the apex, neither with inter-!amel!ar thickenings; cilia
mostly none; lid small, inflated to short-conic.
A cosmopolitan genus of more than 100 species, on earth or rocks in dry
or moderately moist habitats; at least 15 species in North America; two
species in our range.
Key to the Species
A. Leaf-base neither sheathing nor conspicuously scarious, margin revolute
1. B. pomiformis
A. Leaf-base scarious and sheathing, margin plane 2. B. ithypbylla
1. Bartramia pomiformis Linnaeus, p.p., Hedwig
Apple Moss
Plate XXVIII
Rather densely cespitose, soft, yellowish-green: stems about 1.5-3 cm long,
erect, densely reddish-brown-felted below; leaves about 4-6 mm long, the lance-
subulate part spreading rather abruptly from a more or less erect and concave
but scarcely sheathing lance-ovate base, more or less crisped when dry. the
margin revolute in the basal half at least, serrate above, the costa rather nar-
row and distinct, e.xcurrent in a spinulose-serrate subulation; basal leaf-cells
smooth, hyaline, often reddish-brown and pellucid at insertion, elongate-rectan-
gular, the marginal shorter in a few rows, median cells rounded-quadrate,
incrassate, papillose: seta about 5-10 mm long, erect or curved-ascending,
smooth, reddish-brown; capsule globose, about 1.5 mm in diameter, striate,
unsymmetric, reddish-brown when ripe, globose to oblong or narrowly oblong,
when dry deeply sulcate, cernuous, occasionally strumose, often somewhat
arcuate; peristome double, teeth reddish-brown, narrowly triangular-lanceolate,
faintly papillose, prominently articulate, sub-trabeculate, divisural faint, zigzag;
segments two-thirds as long as teeth, carinately split, the cilia two or three and
rudimentary, or none; lid convex, bluntly umbonate; calyptra narrowly cucul-
late, about 2 mm, long; spores reddish-brown, pellucid, coarsely papillose,
mature in May or June.
Cosmopolitan on rocks or swampy soil in moist and shady woods; in
North America from the Arctic regions to Florida and Colorado.
This is an easily recognized moss. The leaves look like tufts of green
wool and are surmounted by slender stemmed fruits like miniature apples.
Common in our region — known from 15 counties in western Pennsylvania and prob-
ably occurs in all. Speimen figured: On clay roadside ban'<, Hartstown. Crawford Co.,
May 29-31, 1909. O.E.J. & G.K.J.
150 American Midland Naturalist Monograph No. 6
la. Bartramia pomiformis var. crispa (Swartz) Bryologia Europaea
This variety is taller and looser than the species: leaves longer, more dis-
tant, when dry more crispate; the innovations are long, often longer than
the seta.
In moister or more shaded situations but with much the same general
distribution as the species.
McKean Co.: D.A.B. (Porter's Catalogue).
2. Bartramia ithyphylla [Haller] Hedwig
Densely cespitose, silky, glaucous-green or yellowish: leaves closely, rigidly
divergent from, a white, scarious, erect-appressed glossy base, when dry quite
straight and more or less erect, the spreading lamina linear-subulate, abruptly
contracted from the obovate base, margin plane, sharply denticulate above;
costa strong but not very distinct above, excurrent into the denticulate
subulation; basal leaf-cells linear, 4-10:1, hyaline, the median and upper
papillose, obscure, about 3-6:1: seta long; capsule similar to that of B.
pomiformis, globose-oblong, when dry curved and deeply furrowed; peristome-
teeth reddish-brown, apically bifid or irregularly perforate; segments yellowish,
cleft, much shorter than the teeth: synoicous: spores large, mature in summer.
On moist earth or in moist fissures of rocks, mainly in alpine regions, in
Europe, Asia, and in Arctic and temperate North America.
Rare in our region. Fayette Co.: Layion's, Rev. S. W. Knipe. (Porter's Cata-
logue), and Knight. (Lesquereux and James).
3. Philonotis Bridel
Dioiccus, rarely autoicous: very slender to robust, cespitose, bright green
to yellowish-green or bluish-green: stem with a distinct central strand, erect,
more or less elongate, usually with whorled sub-floral shoots; leaves erect-
spreading to secund, uniform, or dimorphic, lance-ovate, mostly acute, dentate
or serrate, mostly with lamina one-layered; costa percurrent to excurrent, rarely
incomplete, cells of the apex elongate to shortly rectangular, sometimes rhom-
boidal, rarely parenchymatous and 5-6-sided, mostly ventrally or on both sides
mamillate, rarely so only dorsally, or rarely smooth, basal cells more lax:
sporogonia solitary, seta erect, long; capsule inclined to horizontal, globose,
unsymmetric, with mostly short coUum, striate, when dry sulcate and mostly
constricted in the middle, rarely drying erect and smooth; peristome mostly
double, the inner one rarely lacking; teeth 16, generally with interlamellar
thickenings; basal membrane high; cilia distinct, except in P. Muhlenbergii;
lid mostly low-convex to short conic.
A. large and cosmopolitan genus of about 175 species, on earth and rock
in swamps and springy places and on dripping ledges; probably about 30
species in North America; five species in our general region.
Key to the Species
A. Perigonial bracts mostly obtuse, widely spreading from an erect base; median leaf-
cells abojt .006-.010 mm wide; leaves dimorphic: cilia well-developed ....4. P. fontana
A. Perigonial bracts acute ot acuminate B
B. Perigonia gemmiform, often apparently lateral; autoicous; costa percurrent to
Jennings: Manual of Mosses — 18. Bartramiaceae 151
rather long-excurrent 1. P. lorigiseta
B. Perigonia terminal and discoid or rosette-like C
C. Leaves neither plicate nor with either or both margins revolute D
C. Leaves plicate and more or less revolute 5. P. calcarea var. occidentalu
D. Costa weak but more or less spinulose-excurrent; leaf-cells linear; cilia well devel-
oped 2. P. marchica
D. Costa only percurrent to shortly excurrent; leaf-cells linear-oblong to oblong; cilir,
short and rudimentary ?. P. Muhlenbergit
1. Philonotis LONGISETA (Richard) E. G. Britton
{BdTtramia longisetj Richard; Bartramij radicalis Beauvois)
Stems rather slender, yellowish to green, up to 2 cm high; leaves ascending
to spreading, lanceolate to somewhat triangular-lanceolate, about 1-1.5 mm
long, margins bluntly serrate and revolute; costa weak but percurrent to dis-
tinctly e.\'current; leaf-cells papillose, rectangular to linear above, larger below;
autoicous, perigonia often lateral, gemmiform, perigonial bracts lanceolate from
an ovate to oblong clasping base; perichaetial bracts slenderly lanceolate acu-
minate from an ovate or oblong base, with costa excurrent; seta up to about 2
cm long, castaneous; capsule tilted or slightly drooping, about 1.5-2 mm long,
mouth somewhat oblique; lid low-conic; teeth reddish, lance-acuminate, papil-
lose, strongly trabeculate; segments shorter, the cilia usually adherent; spores
about .025 mm, papillose, and said to be oblong or reniform.
From Pennsylvania and Ohio south to the Gulf States. Was collected
by Wolle at the Nockamixon Cliffs, Bucks County, and in Lehigh County,
but not yet reported from our region.
2. Philonotis marchica (Willdenow) Bridel
{Leskea marchica Willdenow)
Tufted, bright green, the older parts tom.entose, up to 7 or 8 cm high;
leaves ascending to patent, lance-ovate to triangular lanceolate, serrate, some-
times revolute; costa slender but percurrent to spinulose-excurrent; cells oblong-
linear above, wider below, mostly with sharp papillae at the upper end: dio-
icous; perigonia large, terminal, their bracts lance-acuminate and squarrose
from a broad sheathing base, serrate; perichaetial bracts lance-ovate, tapering
from an oblong base; seta up to 3 or 4 mm long; capsule tending to subglobose,
tilted to drooping, with small oblique mouth; lid rounded-convex; teeth reddish,
strongly trabeculate, papillose; segments somewhat shorter, yellowish, papillose
in lines; cilia various, rarely well developed and separate above; spores oblong-
reniform, rough, reddish.
Europe; in America from eastern Canada to Minnesota, Georgia, and
Texas.
Collected along Slippery Rock Creek by Lesquereux. (Porter's Catalogue).
3. Philonotis Muhlenbergii (Schwaegrichcn) Bridel
{P. marchica Sullivant)
Plate XXVIII
Rather densely cespitose, light yellowish to bluish-green: branches reddish,
whorled from below the archegonal clusters, erect to ascending, reddish-tomen-
152 American Midland Naturalist Monograph No. 6
tose below about 1-3 cm high, slender; leaves of fertile stems 1-2 mm long,
rather distant, lance-ovate, acute, ascending to appressed, when dry somewhat
crispate, carinate, with revolute margins but not plicate, more or less spreading,
serrulate in apical half, scarcely decurrent; costa strong, brownish, percurrent
to somewhat excurrent; leaf cells mostly parenchymatous, rectangular to hexag-
onal to oblong above, incrassate, median cells strongly papillose on upper end,
about .003-. 006 X .015-030 mm, elongate-rectangular, apical cells narrower and
tendinjT to vermicular-hexagonal, basal cells looser, more or less rectangular,
up to .012 X .040-. 060 mm, smooth; inner perichaetial leaves ovate-triangular at
base with the costa excurrent into a subulate apex, the margin entire, the cells
rather lax; perigonial leaves erect-spreading, long-acuminate, serrate; seta about
2.5-3 cm long, erect, smooth, shining, reddish-brov.n, when dry flexuous; cap-
sule globose to ovoid-globose, faintly striate, about 2-2.5 mm long, brownish,
when dry sulcate and variously wrinkled, arcuate, cemuous, the neck sunken in,
about 4-6 rows of cells at the mou'.h of the capsule laterally elongate; peristome
double, the teeth 16, narrowly triangular-lanceolate, prominently articulate,
pellucid, oranoe to reddish-brown, divisural zigzag, distinct below; segments
narrow, about four-fifths as high as the teeth, mostly split apart; cilia three,
very short and rudimentary, the basal membrane comprising more than half
the height of the inner peristome, the segments and the upper part of the
membrane orange-pellucid, papillose-striate; spores globose, papillose, pellucid,
orange to reddish-brown, .01 8-. 025 mm, mature in June.
On dripping rocks along streams, wet places, etc., from Virginia to Maine.
Uncommon in our region. Allegheny Co.: In crevices of rocky bed of stream, ravine
of Power's Run, May 14, 1908. O.E.J, (figured). Lawrence Co.: Wet rocks in deep
lavine near Rock Point. June 26, 1909. O.E.J.
4. Philonotis fontana [Linnaeus] Bridel
(Mnium fontanum Linnaeus; Bartramia fontana Swartz)
Plate XXIX
Cespitose, yellowish-green, sometimes quite glaucous, loose above but inter-
woven belov/ with a reddish-brown felt-like tomentum: stems erect, reddish,
slender, usually 2-6 cm high, densely fulvous-radiculose below, the innovations
usually whorled and giving the plants the appearance of being pleurocarpous;
leaves about 1.5-2 mm long, lance-ovate, acuminate, appressed when dry,
usually quite plicate on each side of the costa near the base, serrate above,
usually more or less revolute towards the base; costa strong, often percurrent
or even excurrent, usually reddish; basal cells elongate-rectangular to elongate-
hexagonal, loose, pale pellucid, about .008-. 012 (-.015) mm wide, the end-
walls often papillose, the cells in the acumen linear-vermicular, incrassate and
more or less papillose at both ends; perigonial leaves spreading, broadly tri-
angular-ovate, the inner often obtuse and rounded at the apex, the costa not
reaching the apex: seta dark red, 2-4.5 cm long; capsule ovate-globose, large,
brownish, thick-walled, striate, oblong, when dry and empty arcuate and ir-
regularly ribbed; operculum conic-convex, acute; peristome-teeth reddish-brown,
p>ellucid, lanceolate; peristome-segments nearly as long as teeth, narrow, carinate-
!y gaping, cilia three (two) about as long as segments; spores very slightly
Jennings: Manual of Mosses — 18. Bartramiaceae 153
papillose, incrassate, yellowish-brown, about .01 9-. 025 mm, usually mature
in June.
Water-loving mosses usually avoiding calcareous habitat*,, on dripping
rocks or in swamps and wet places. Cosmopolitan and occurring in North
America throughout, from Canada to Florida, more abundant in the cooler
and more mountainous regions.
Common but only occasionally fruitmg in our region. Known from Allegheny, Arm-
strong, Centre, Clinton, Fayette, Huntingdon, Indiana, Lawrence, Lycoming, McKean,
Washington, and Westmoreland counties. Thus far this moss has not been found in
western Pennsylvania north of the terminal glacial moraine. Specimen figured: Flood-
plam of Brush Creek, Douthett. Allegheny Co., April 26, 1908. O.E.J, and G.K.J.
4a. Philonotis fontana var. falcata Bridel
Leaves falcate-secund; branches hooked towards the apex.
Centre Co.: Matternville Gap, Bald Eagle Mt., July 15, 1909. O.E.J.
5. Philonotis calcarea (Bryologia Europaea) Schimper,
f. occidentalis Flowers
Plate XXIX
Densely and softly cespitose, bright green, more or less glaucous above,
brownish below: stems long, up to 10-12 cm, slender, erect in the dense tufts,
red-brown and densely felted-tomentose below; branches in whorls; leaves
dimorphic, stem-leaves ovate-lanceolate, acuminate, about 2-3 mm long, deeply
concave, plicate, towards the apex sharply serrulate, towards the base the basal
papillae of the cells forming rounded projections, especially on the revolute
margins, the margins revolute narrowly towards the clasping and sub-decurrent
base, the leaves erect-spreading to secund when moist, shrunken and sub-cris-
pate when dry; branch-leaves when moist usually more or less falcate-secund,
ovate-lanceolate and narrowly acuminate, about 1-1.5 mm long, by 0.5 mm
wide, when dry somewhat shrunken and twisted; costa in both forms of leaves
strong, excurrent, dorsally papillose, basal leaf-cells rather thin-walled, rectan-
gular, up to .060. 080 x .01 5-. 018 mm, pale, pellucid, towards the margins and
upwards lecominp shorter, more incrassate, papillose at the ends, the median
and. upper leaf-cells becoming quadrate to 2-4 times as wide as long, strongly
papillose at their upper ends, incrassate, pellucid: capsule not seen but said to
be large and similar to that of P. fontana ■' perigonial leaves oblong-lanceolate:
spores mature in summer, but the capsules rather rarely produced. In vegeta-
tive characters this species is difficult to differentiate from forms of P. fontana.
Flowers, in Grout (Moss Flora), doubts the occurrence of typical P. calcarea
in North America and establishes forma occidentalis to cover our American
specimens which seem more or less intermediate between calcarea and fontana.
Our specimens seem to belong to the form.
Uncommon in our region. Clinton Co.: In roadside ditch, north of Renovo, July
15, 1908. O.E.J, (figured). Huntingdon Co.: Warrior's Ridge, T. C. Porter. (Por-
ter's Catalogue).
154 American Midland Naturalist Monograph No. 6
F.-i-.i'.'y P. Timmtaceae
Dioicous or autoicous; robust, in more or less high, lax, dull-gieen to yel-
lowish-green tufts, brownish inside, with a brown tomentum below: stem erect
or procumbent, with central strand, densely-leaved, simple or dichotomous;
leaves 8-seriatc, of uniform length, from a half-sheathing, non decurrent base
spreading to recurved, elongate lance-linear, carinate; lamina unistratose, chan-
neled to concave, non-bordered, serrate; costa strong, percurrent, often dorsally
toothed above, with several median guides; leaf-cells green, small, rounded to
4-6-sided, ventrally mamilkte; cells of the sheathing part without chlorophyll,
sometimes dorsally papillose, elongate-rectangular to linear, narrow towards
the margin: sporogonia solitary; seta long, erect; capsule cernuous to almost
pendent, from a short collum oblong-oval, brown, not or but slightly striate,
when dry ribbed; annulus revoluble; peristome inserted back from the cdr;e of
the mouth, always double, the inner as long as the outer, when dry the parts
sharply bulged outwards in the middle; teeth 16, confluent at the base, broadly
lance-linear, rarely split, plane, below yellowish and transversely striate-punc-
tate, above whitish and vertically papillose-striate, divisural zigzag, dorsal
plates low. sometimes cut by cross-walls; inner peristome free, yellow, basal
membrane high, carinate, transversely striate, dividing into 64 filiform, papil-
lose cilia, united apically into groups of fours, generally appendiculate on the
inner side: spores .012-. 023 mm, yellow, almost smooth; operculum hemi-
spheric, often apiculate; calyptra cucullate, long and narrow, often remaining
on the seta.
One genus with characters as for the family; 8 species; 4 in North Amer-
ica, one in our range
1. TiMMiA Hedwig
1. TiMMiA cucuLLATA Richard*
(T. megapoiitana American authors, in part; T. megapolkana
var. cucullata (Richard) Sayre
Plate XXIX
Loosely cespitose, bright green above, brownish below: stems erect, spar-
ingly branched, radiculose below; leaves lanceolate to lance linear, spreading
from a concave appressed and more or less sheathing base, acute to subacute,
the margins serrate almost to the sheathing base, the spreading portion of the
leaf about as wide as the sheath, concave, smooth on back or more or less
involute; costa rather narrow, strong, ending in the apex; basal leaf-cells elon-
gate-rectangular, rather thin-walled, hyaline, hardly inflated, in upper part of
sheathing base becoming shorter to quadrate, and incrassate, the outer walls
bulging so as to appear slightly papillose, about .010 mm in diameter: seta
* Timmia megapoiitana Hedwig ranges south to No v Jersey, New York, and Mis-
souri, and might occur in our region. Its lea\es are narrowed gradually to the apex
instead of suddenly as in T. cucullata; the mouth of the capsule when dry not wider than
rest of the capsule in megapoiitana, but flaring in cucullata. — See Sayre in Groit's Moss
Flora.
Jennings: Manual of Mosses— 20. Buxbaumiaceae 155
about 2 cm long, erect; capsule inclined to cernuous, oblong, when dry and
empty unsymmetric, strongly curved, and somewhat wrinkled and tapering
gradually from the wide mouth to the seta; lid rounded and apiculate; annulus
revoluble, pluriseriate; peristome double, the teeth 16, lance-linear, yellowish-
pellucid, trabeculate on inner side, articulate and with a divisural on outer
surface, inner peristome with high basal membrane and 64 filiform cilia united
into groups of four each, opposite to and about as long as teeth; calyptra
cucullate; spores smoothish, mature usually in May.
In shade on moist banks, or bases of trees, mostly in calcareous districts;
Europe, and, in North America, from Newfoundland to Virginia, Kentucky,
and Missouri.
McKean Co.: Riverside swamp, ten .miles north of Bradford, on base of old elm,
August 19, 1896. D.A.B. Sterile (figured). Washington Co.: Bank of Buffalo
Creek, Taylorstown, May 18, 1895. Linn &C Simonton.
Family 20. Buxbaumiaceae
Autoicour or dioicous: perennial, low, gregarious to laxly cespitose, dark
green, finally brownish: protonema more or less persistent; stem without cen-
tral strand, mostly very short, erect, thickly foliate to almost leafless: seta
ranging from almost none up to 5-20 cm long, erect; capsule proportionally
large, sub-erect to inclined, often finally more or less horizontal, dorsiventrally
unsymmetric, flattened above, ovate to oblong or ovate-conic, narrowed to a
very small mouth; annulate; peristome double or, apparently, single, the inner
consisting of a membraneous plaited cone with an apical opening, the teeth
originating from one to four concentric rows of cells, faintly barred; operculum
conic, glabrous, smooth; spores very small.
A very small and rather primitive family of mosses, growing on earth or
decayed wood. Theriota, in Korea, and the following two genera:
Key to the Genera
A. Seta almost none; capsule immersed in the fringing bristles of the penchaetial leaves
I. Diphyscium
A. Seta 5-20 cm long, thick, red or reddish-brown; leaves none at the time of ripening
2. Buxbaumia
1. Diphyscium [Ehrhart] Mohr
{Webera Ehrhart, not Hedwig)
Dioicous: perennial, mostly low and densely gregarious; protonema long-
persisting; stem without central strand, short, erect, radiculose, thickly-leaved,
simple, rarely longer and branched; leaves twisted or crispate when dry,
spreading when moist, the lower lingulate or elongate-spatulate, obtuse or
acuminate, entire; costa strong, without guides in distal part, ending below
apex; lamina 2- (3-) stratose; leaf-cells on both sides mamillate to smooth,
rounded to 4-6 sided, incrassate, often widened transversely, in the basal por-
tion uni-stratose, pellucid to hyaline, lax, elongate 4-6 sided with the transverse
walls incrassate, smooth; perichaetial leaves much larger, erect, whitish, mem-
branaceous, e'ongate, lanceolate to linear, the apex usually fringed and the
156 American Midland Naturalist Monograph No. 6
costa long-aristate-excurrent: seta very short, without central strand; capsule
immersed, obliquely ovate-conic, gibbous, without collum, mouth very small;
annulus present; outer peristome rudimentary or none, when present consisting
of 16 triangular teeth; inner peristome pale, membranaceous, 16-plaited, papil-
lose, short-conic; operculum small, acute-conic, falling away attached to the
upper part of the fleshy columella; calyptra very small, conic, glabrous.
A genus of about 15 species, only one in North America.
1. DiPHYSCiUM FOLIOSUM (Hedwig) Mohr
{Buxbaumia foliosa Hedwig; Webera sessilis Lindberg)
Plate XXIX
Small, widely cespitose, very short-stemmed (1-2 mm), the general af>-
pearance being that of a grain of wheat sitting in a tuft of bristles: stem-leaves
minute, the largest being about 3 mm long, linear-oblong, obtuse, somewhat
concave, crisped when dry; costa broad, nearly reaching apex; perichaetial leaves
4-7 mm long, lance-linear, the costa one-third the width of the leaf at base,
brownish-yellow, excurrent as a spinulose arista which often reaches one-half
the whole length of the leaf, the apex of the lamina acute and entire or lacerate-
ciliate; cells at base of the stem-leaves and of nearly the whole lamina of the
perichaetial leaves hyaline, thin-walled, irregularly elongate-rectangular to
hexagonal, the costa in the lamina of the perichaetial leaves being bordered on
either side by several rows of smaller, chlorophyllose, quadrate, and somewhat
opaque cells, the lamina of the stem-leaves also being opaque with very small
round-hexagonal cells: capsule 4-6 mm high, ventricose, ovate-conic, yellowish-
green; operculum acute-conic; calyptra small but covering the operculum;
annulus and outer peristome more or less rudimentary; inner peristome conic,
whitish, membranaceous, papillose, 16-carinate; spores moderately thick- walled,
.007-.010, papillose, mature from mid-summer to early fall.
Widely distributed in the Northern Hemisphere. In North America it
occurs from Alabama to Nova Scotia and Ontario, on moist shaded banks and
clayey hillsides, and, in our region, especially along rather bare banks along
paths in black oak — scarlet oak woods in the mountains. Usually associated
with white flecks of Cladonia lichen thalli.
Butler Co.: Roadside bank, McKelvy School, Brady Twp., April 26, 1936. Sidney
K. Eastwood. Cameron Co.: Canoe Run, Lumber Twp., Sept. 1, 1935. A. M. Barker.
Centre Co.: Tussey Mt., Shingletown, July 15, 1909. O.E.J. Fayette Co.: Wooded
roadside bank, Sugar Loaf Mt., Sept. 1-3, 1906 (figured), and near Lover's Leap, Ohio
Pyle, Sept. 4, 1906. O.E.J. & G.K.J. Somerset Co.: Moist bank, Laurel Ridge, 2 mi.
s. of Miller School, Oct. 6, 1935. C.M.B. Westmoreland Co.: Roadside bank,
"Rachelwood," New Florence, Sept. 9, 1907. O.E.J. Alt. 2,100 ft.
2. Buxbaumia Haller, Hedwig
Dioicous; antheridial plants microscopic on the green protonema; arche-
gonial plants with a short stem, a few small leaves, and one or two archegonia
but no paraphyses: isolated or gregarious, annual: stem barely 1 mm high,
simple, with hyaline rhizoids; leaves ovate to lance-ovate, the basal portion
green and its cells growing out into long brown filaments during the develop-
Jennings: Manual of Mosses — 21. Tetraphidaceae 157
ment of rhe sporogonia, finally covering the stem and vaginule with a thick
tomentum; leaf-cells lax, long-hexagonal; perichaetial and upper leaves soon
disappearing: seta 5-20 mm long, thick, red-brown, warty, the central strand
being surrounded by an air-space; capsule obliquely ascending, the upper sur-
face flattened, the urn brownish and with a short erect collum and narrowed
above to a very small mouth; a low pseud-annulus present consisting of a
number of layers of cells; outer peristome of one to several rows of short
and irregular teeth; inner peristome a pale, plaited, membranous truncated cone,
as in Diphyscium, 32-carinate; spore-sac small, surrounded by a large air-
space; spores small; operculum small, conic-obtuse, erect, falling tardily with
the upper part of the columella attached; calyptra small, covering only the
operculum, glabrous, fugacious.
A rather widely distributed genus of 5 species, 3 or which occur in North
America, one in our region.
1. Buxbaumia aphylla Linnaeus, Hedwig
"Bug-on-a-Stick Moss"
Plate XXX
Plants minute on a thick, brownish, felted protonema and after the devel-
opment of the sporophyte usually completely obscured by a dense growth of
protonemal filaments: seta erect, stout, rough, about 1 cm high, castaneous;
capsule when ripe, lustrous, castaneous, ovate-acuminate in outline, flattened
obliquely in the upper two-thirds, with angular edges, smoothish, about 5-7
mm long, resembles a shriveled grain of wheat; operculum about 1.5 mm high,
oblong-conic, disproportionately small; calyptra falling early, conic, covering
only about one-half of operculum; peristome consisting of an outer series of
papillose short, slender teeth, and a longer, papillose, conical, plaited cone;
spores smooth, spherical, about .007 mm in diameter, mature in late fall and
early spring.
On clayey banks in woods or, particularly, along neglected roadsides, often
with thalli of Cladonia. Rarely en decayed wood. Europe, Asia, and, in
North America, from Canada to Virginia and Washington State. Found
commonly in Butler and adjacent counties, but to find it requires a proper
recognition of its habitat and a trained eye. It is usually on clay roadside
banks mixed with white flecks of some Cladonia lichen.
Collected by Sidney K. Eastwood from more than thirty localities in Butler County
(Eastwood, S. K. Bryologist 39: 127-129. 1936). Now known from the following coun-
ties: Allegheny, Beaver, Butler, Cimeron (Porter's Catalogue), Clarion, Clearfield, Fay-
ette, Huntingdon (Porter's Catalogue), Indiana, Lawrence, Lycoming, and Westm.orelind.
Specimen figured: On clay bank with decayed wood, 3 mi. n. of Wurtemberg. G.K.J.
Oct. 16, 1910.
Family 21. Tetraphidaceae
( Georgijceae )
Autoicous: slender to very small, dull, gregarious to cespitose, bright to
brownish-green; stem erect, leaves 3-5 seriate, unistratose, costa obscure to
well-developed and ending just below the apex; cells parenchymatous, thick-
158 American Midland Naturalist Monograph No. 6
walled, smooth; perichaetial leaves longer: seta long, erect, straight or genicu-
late in the middle; sinistrorse below, dextrorse above; capsule erect, symmetric,
smooth, oval or cylindric; annulus none; peristome inserted below the m^outh,
consisting of the entire mass of tissue enclosed within the epidermal operculum,
this splitting by two planes vertically into four solid, three-angled, elongate-
pyramidal teeth; columella reaching only to the level of the mouth; spores
.008-.015 mm, smooth; operculum conic, unistratose, cleft on one side; calyptra
conic, glabrous, plicate, the margin lobed.
A very small family of 5 species, occurring on rocks, rotten wood, soil, etc.,
in Europe, Asia, and North America. Two genera; T etradontium and the
following:
1. Tetraphis Hedwig
{Georgia Ehrhart)
Slender plants, more or less densely cespitose in wide soft tufts, bright
green to brownish, radiculose-tomentose below: stems to 3 cm long, with
central strand, three-angled, branched, with distant, scale-like, ecostate leaves
below; upper stem-leaves abruptly larger, approximate, ovate-lanceolate, acute,
margins plane and entire, with costa ending below, or in the apex, 4-5-stratose
at base, cells incrassate, uniform; leaf-cells incrassate, round-hexagonal, wider
transversely, elongate in the leaf-apex, rectangular at the leaf-base: seta 1-1.5
cm long, often two together; capsule erect, symmetric, greenish, when empty
brownish and weakly dextrorse, without stomata; calyptra enclosing the upper
one-third of the urn, its apex carinate-toothed; gemmae lenticular, borne in a
cup formed of four or five broadly cordate bracts at the apex of the more
slender and flexuous gemmiferous stems.
Four species, all occurring in North America, only the following one in
our range:
1. Tetraphis pellucida [Linnaeus] Hedwig
(^Georgia pellucida Rabenhorst)
Plate XXX
Loosely cespitose in wide yellowish-green tufts: stems erect, about 1 cm.
high, densely felted-radiculose at the base, reddish blow; basal-leaves minute,
upper leaves larger, tufted, ovate-lanceolate, margin entire; certain stems
bearing at the apex gemmae-cups about 1 mm in diameter, the surrounding
leaves being broadly obovate to reniform, truncate or apiculate at the apex;
perichaetial leaves linear-lanceolate, up to 4.5 mm long; costa of stem leaves
wide, ending below apex, in perichaetial leaves often percurrent; aerolation
dense, rounded, the cells of the perichaetial laves irregularly elongate at base;
the cups enclosing small, many-celled, lenticular gemmae: seta yellowish to
reddish, erect, dextrorse above, about 1-15 cm long; capsule cylindric-lanceolate,
erect to ascending, reddish, about 2-2.5 mm long; annulus none; peristome
consisting of 4 linear-triangular thick teeth, reddish to brownish, comprising
about one-fifth of the length of the capsule; operculum lustrous, conic, acute;
calyptra whitish and lacerate below, plicate, enclosing the whole capsule, at
apex solid, acute, rough; spores about .010 mm, thin-walled, slightly papillose,
mature in summer or early fall; capsules persistent.
Jennings: Manual of Mosses — 22. Polytrichaceae 159
Widely distributed, on peaty soil, decayed logs, etc., Europe, Asia, and,
in North America, in Canada and northern United States.
Common in our region and very often found on the cut end of a partially decayed
stump or log. Now known from 16 counties in western Pennsylvania and probably occurs
m all. Specimen figured: "Rachelwood," New Florence, Westmoreland Co., Sept. 8-11,
1907. O.E.J.
Family 22. Polytrichaceae
Dioicous, rarely paroicous or synoicous; antheridial flower terminal, large,
discoid, generally bearing a shoot in its middle; archegonial flowers terminal,
bud-like: perennial, mostly very large, mostly cespitose, with a long horizontal,
subterranean, triangular, blackish, branched, radiculose rhizome: stem erect
with lower leaves none or remote, leaves weakly costate, three-seriate, without
lamellae, red to hyal.'ne, small and scale-like; upper part of stem five-or-more-
angled, with specialized central strand; stem structure complex; upper leaves
larger, the sheathing base usually yellowish to hyaline, lamina more or less
spreading or recurved, when dry mostly erect, sometimes convolute to crispate,
mostly lanceolate to lance-subulate, sometimes Ungulate, mostly sharply
toothed, mostly plane with erect edges, uni-stratose or with two-stratose zone
next the ccsta, rarely two-stratose to the margin, with narrow, vertical, green,
longitudinal, m.ostly uni stratose lamellae on the ventral surface of the costa
and of the bi-stratose lamina; costa narrow to wide, incomplete to aristate-
excurrent, dcrsally often toothed and rarely lamellate, complex in structure;
leaf-cells parenchymatous, small, the basal rectangular to linear and narrower
towards the margin: seta elongate, mostly solitary, often flattened and weakly
sinistrorse; capsule first erect, later inclined to horizontal or pendent, cylindric
to prismatically 4-6-sided or cubic, collum various; annulus none or uniseriate
with three or four transitional bordering cells; peristome rarely lacking, mostly
simple v/ith 32 to 64, rarely 16, lingulate, short, unbarred teeth, triangular in
cross-section, rising from a basal membrane, the teeth hyaline, often with a
colored median line, incurved to meet the disk-like apex (epiphragm) of the
columella; spores mostly small, .00S-.012 or .014-.021 mm, mostly smooth;
operculum, apiculate to rostrate from a conic to convex base; calyptra cucullate,
rarely glabrous, mostly spinulose to long, villous and felted.
A cosmopolitan family, mostly on siliceous or other non-calcareous soils: in
colder reoioiis often forming laroe masses ur sods.
N3
Key to the Genera
A. Capsules cylindric - B
A. Capsules four-angled or six-angled 3. Polytrichum
B. Leaves not crisped when dry; calyptra hairy; protonema persistent 2. Pogonatum
B. Leaves crisped when dry; calyptra not hairy 1. Atnchum
1. Atrichum Beauvois
{Catharinaea Ehrhart )
Stems of moderate height, in loose tufts or gregarious, dark green to bronze
when old. central strand generally well developed; rhizome creeping, branched,
hearing loosely- to thickly-leaved erect shoots, densely radiculose at the base;
160 American Midland Naturalist Monograph No. 6
leaves Ungulate to ovate-oblong, not sheathing nor narrowed above the base
but slightly embracing the stem, margined, serrate; crisped when dry, costa
narrow, on the upper side with 1-12 narrow lamellae, ending below or in the
apex, towards the apex often spinose-serrate; cells chlorophyllose, the upper
rounded-hexagonal, smooth, the basal mostly rectangular: seta long, erect;
capsule smooth, cylindric to oval, rarely obovate, often more or less curved;
operculum long-rostrate; peristome of 32 teeth with pale borders and a median
orange to reddish-brown line, the basal membrane narrow and reddish-brown
or orange; calyptra smooth except at the apex, where it is sometimes spinulose:
mostly dioicous.
A cosmopolitan genus comprising about 40 species, growing on earth,
mostly in the temperate zones; about 20 species have been reported for
North America; four species in our region.
Key to the Species
A. Upper leaf-cells .015 to .025 mm, in diameter; costa and lamellae not exceeding one-
fifth of the width of the median part of leaf B
A. Upper leaf-cells .007 to .015 mm in diameter; costa and lamellae e.xtending over
one-fifth to two-thirds of the width of median part of leaf C
B. Lamellae 3-6 in number and 3-6 cells in height, covering about 1/8 to 1/11 of the
median lef-width 2. A. undulatum
B. Lamellae 5-7 in number and 4-7 cells in height, covering about 1/7 to 1/3 of the
median leaf-width 2a and 2b. A. undulatum var. minus
B. Lamellae 0-4 in number, 1-3 cells high; leaves crispate when dry 1. A. crispum
B. Lamellae 4-6 in number, 6-13 cells in height - {A. undulatum var. Sdwyni)
C. Lamellae 4-7, 6-9 cells high, covering 1/3 to 1/4 of median leaf-width
3. A. angustaturn
C. Lamellae 6-8, 8-14 cells high, covering about 1/5 to 1/3 of median leaf-width;
upper cells papillose 4. A. papillosum
C. Lamellae 7-12, 8-14 cells high, covering about 1/2 to 2/3 of median leaf-width
5. A. angustaturn var. plurilamellatum
1. Atrichum crispum (James) Sullivant
(Catharinaea critpa James)
Plate LXIV
Gregarious, yellowish-green, becoming brownish, especially below; branches
simple, erect from a branched underground stem; leaves crisped when dry,
when wet ascending, lance oblong to oblanceolate, or obovate, up to 8 mm
long by 1.2 wide, flat, or but faintly undulate, acute, marginally spinulose-
serrate in upper half, bordered from near base to tip with colored, incrassate,
narrowly linear cells, 2 or 3 cells wide and usually 2 cells thick; costa strong,
percurrent (or excurrent), about 1/10 the width of the leaf, with rarely one
or two teeth dorsally near the apex, frequently with from one to three lamella
on upper side of costa in upper part of leaf, the lamellae smooth and usually
2-3 cells high; leaf-cells smooth, the basal thin-walled and semi-inflated,
rounded-oblong to rectangular, 1-4:1, the marginal much narrower, above base
the cells become incrassate, rounded to quadrate-hexagonal. Not seen in
Jennings: Manual of Mosses — 22. Polytrichaceae 161
fruit in our region, but has seta 15-20 mm tall; capsule terete, more or less
erect, with urn 2-3.5 mm long, about 3-4:1, lid with beak half as long as urn.
Stream banks or other moist situations; N.E. to Ontario and Tennessee;
also in the Northwest.
Butler Co.: On ground in wet ditch, Semiconon Run, lYi mi. north of Conoquenes-
sing. Sidney K. Eastwood. Sept. 22, 1935. Fayette Co.: Partly buried in sand around
banks of river, around "Peninsula," Ohio Pyle. Charles M. Boardman. Four collections.
Oct. 12, 1935, figured). Somerset Co.: Beck Spring. Mill Creek, Laurel Hill Mt.
C.M.B. July 26, 1947. Westmoreland Co.: On wet clay in roadside ditch, Laiael
Hill Mt., 9 mi. s.e. of Rector. H. N. Mozingo. Oct. 7, 1945.
2. Atrichum undulatum [Linnaeus] Beauvois
{Bryiim unditlatiim L. )
Plate XXX
Loosely cespitose, dull, dark green: stems erect, ranging from 1.5-6 cm
long, usually about 3-4 cm, mostly simple, more or less gray-radiculose below,
arising from a rhizone-like base; lower leaves minute, increasing in size up-
wards, the upper leaves lanceolate-lingulate, much crisped when dry, trans-
versely undulate when moist, sub-acute to obtuse, about 6 8 mm long, 1 mm
wide, serrulate to the middle or slightly below, the uppermost teeth double,
strong, being inserted in a border of 1-3 rows of brownish, pellucid to hyaline,
incrassate, narrow cells; the crests of the undulations on the back of the leaf
also often spinose in upper part of the leaf; leaf-cells elongate-rectangular at
base, reaching about .017 x .033 mm, becoming quadrate towards leaf-middle,
towards apex hexagonal and somewhat longer transversely and about .01 7-. 024
mm; costa strong, ending just below apex, sharply dorsally toothed, ventrally
with 3-6 longitudinal lamellae which each consist of 3-6 rows of cells similar
to those of the leaf-blade, the costa and its lamellae covering rarely m,ore than
one-fourth of the total leaf-width (in our region sometimes even narrower;
"1 11-1, 7 of median width" — Frye: seta erect, flexuose, somewhat sinistrorse,
smooth, lustrous castaneous, 2-5 cm long; capsule lustrous, becoming dull with
age, castaneous, cylindrical, arcuate to almost straight, inclined, smooth, about
4-5 x 1-1.3 mm; peristome single, the 32 teeth linear-lanceolate, obtuse, about
C.3 mm high, orange-pellucid along the median line, united in the lower third
into a reddish-orange basal membrane, the teeth covered (especially along the
margins) with a hyaline, densely but minutely papillose layer which, during
the winter, becomes deciduous, thus leaving the teeth perfectly smooth; spores
smooth, orange, spherical, about .016-.019 mm in diameter; mature in late
fall, operculum conic, curved linear- rostrate, about 2.5-3 mm long; calyptra
pale, roughened towards apex, covering about one-half to one-third of urn.
Widely distributed throughout the North Temperate Zone on earth, par-
ticularly partly shaded clay banks; in North America extending as far :outh
as South Carolina and California.
Common in our region — known from Allegheny, Beaver, Bedford, Butler, Clinton,
Crawford. Elk, Erie, Fayette, McKean, Somerset, Washington, and Westmoreland coun-
ties. Specimen figured: Hyner Creek, above Hyner. Clinton Co., July 15. 1908. O.E.J.
162 American Midland Naturalist Monograph No. 6
2a. Atrichum undulatum var. minus (Lamarck and DeCandolIe)
Weber and Mohr
Plate XXXI
Differs from the species in having the stem, leaves and sub-erect capsule
shorter; lamella covering 1/7-1/5 of median part of leaf.
Erie Co.: Presque Isle, May 8-9, 1906. O.E.J, (figured). Fayette Co.: Ohio
Pyle, May 30-3!, 1908. O.E.J.
2b. Atrichum undulatum var. allegheniense (Jennings)
New Combination*
Plate XXX
Similar to the species in general habit and appearance but usually some-
what smaller and more slender: the lamellae 5-7 in number, usually 6, ranging
from. 4-7, usually 6, cells in height, the costa and lamellae together occupying
from one-sixth to one-third of the width of the upper part of the leaf; the
upper leaf -cells about .016-.018 mm in diameter.
This variety is much more abundant in the Pittsburgh district than is the
species, evidently preferring habitats with shales and sandstones such as those
of the Carboniferous.
Now known from the following counties, thus far apparently avoiding the High
Plateau: Allegheny, Armstrong, Beaver, Butler, Crawford, Erie, Fayette, Lycoming,
Somerset, Washington, and Westmoreland. Type specimen: Power's Run. across from
Verona, Allegheny County. O.E.J. April 18, 1906 (figured).
3. Atrichum angustatum (Bridel) Bryologia Europaea
{Polytrichum angustatum Schwaegrichen; Catharinaea angustata Bride! )
Plate XXXI
Loosely cespitose: stems erect, about 1-3 cm high, or more; lower leaves
minute, the size of leaves increasing upwards, dull green, much crisped when
dry, lance-linear, doubly serrate from about the middle upwards along the
margin and dorsally towards the apex; costa with about 5-8 ventral lamellae
above, the costa and lamellae together occupying about one-fourth to one-half
of the width of the leaf; lamellae 6-8 cells high, cells equal in size; basal leaf-
cells elongate- rectangular, rather incrassate, about .010-. 025 mm in diam-
eter, above becoming quadrate, towards the apex slightly smaller more
or less hexagonal with the longer diameter transverse, about .010-. 015
mm: seta erect, about 1.5-3.5 cm long, smooth, lustrous, castaneous, somewhat
sinistrorse; capsule lustrous, castaneous, linear-cylindric, usually slightly curved,
abou: 5-8 x 1 mm; peristome-teeth obtuse, about 2.5 mm long; calyptra cucul-
late, slenderly rostrate, about 4-7 mm long; operculum hemispheric, slenderly
rostrate, about 2.5 mm long and more or less abruptly divaricately bent; spores
about .012-.018 mm, pale, orange-pellucid, smooth to minutely roughened,
somewhat incrassate, mature in late fall and winter.
■'' Frye, in Grout's Moss Flora, regards this as synonymous with the preceding A.
undulatum var. minus.
Jennings: Manual of Mosses— 22. Polytrichaceae 163
Occurring on wooded shaly or clayey banks; Europe, Asia, and, in North
America, from Newfoundland and Ontario to the Gulf States. In our region
not very common, seemingly preferring steep slopes of ravines, and there often
under hemlocks. Quite variable and often approaching closely the two species
next following.
Now known from the following counties: Allegheny, Armstrong, Beaver, Butler, Craw-
ford, Ene, Greene. Lawrence, McKean, Washington, and Westmoreland. Specimen fig-
ured: Hartstown, Crawford County, May 29-31, 1909. O.E.J.
4. Atrichum papillosum (Jennings) Frye*
(Catharinaea papillosa Jennmgs)
Plate XXXI
Plants loosely cespitose, dark green, dioicous: stems simple or at the base
sparsely branched, towards the base radiculose, erect, about 2 cm high, chest-
nut-brown to purple; lower leaves dark green, short, oblong, above gradually
longer, erect-spreading, the upper leaves tufted, erect-spreading to erect, oblong-
linear, 3-5 mm. long, 0.7-1.0 mm wide, margined, above chlorophyll-bearing,
obtuse to somewhat acute, towards the apex dorsally serrate-spinulose, in the
margin above the middle more or less doubly serrate-spinulose, when moistened
slightly undulate, when dry crisped and circinate, lamellate ventrally along
the percurrent costa; lamellae 4-8, mostly 6-14 cells high, smooth or minutely
and sparsely papillose above the middle, extending over 1/5 to 1/3 the median
width of the leaf; leaf-cells on both sides usually slightly minutely papillose,
the lower cells hyaline, hexagonal-rectangular, about 10-15 by .030-. 045 mm,
the median more or less quadrate, the upper cells quadrate-orbicular, chloro-
phyll-bearing, about .008-.015 mm, the lower border cells linear, incrassate, in
two to three series, more or less two-layered, towards the apex gradually becom-
ing rectangular, in the teeth triangular and sometimes .125-. 140 mm long;
perichaetial leaves similar to the stem-leaves; the perigonial leaves of the
antheridial plants from an ovate-orbicular concave base abruptly linear-acumin-
ate, about 2 mm wide and 3 mm. long, towards the apex canaliculte, lam.ellate
marginally and dorsally serrate-spinulose and usually sparsely papillose; lamel-
lae usually papillose, 4-6, only 3-7 cells high, disappearing quickly below the
base of the acumen: flowers dioicous or rarely arising from the center of the
masculine flower of the preceding year: seta solitary, erect, flexuous, slightly
sinistrorse, about 2 cm long, smooth, sub-lustrous, chestnut-brown; capsule
linear-cylindric, 3.5-6 mm long, 0.5-0.7 mm in diameter, erect-arcuate, tapering
abruptly at the base, smooth, chestnut-brown; cells of the capsule rectangular,
their lateral walls much incrassate, in a series of 5 or 6 cells under the mouth
smaller, quadrate, dark-incrassate; peristome teeth 32, linear-oblong, about 0.3
mm. high, in the median line reddish-orange, towards the sides hyaline, in the
margins a little dark and densely although minutely papillose, forming a basal
membrane in the lower third; spores smooth, incrassate, orange-pellucid, glo-
bose, about .COS-. 01 1 mm; calyptra about 5 mm long, narrowly cucullate,
* Grout (Moss Flora 1: 251. 1939) regards this as synonymous with A. Macmillani
(Holz) Frye, of eastern United States, common in the southeast.
164 American Midland Naturalist Monograph No. 6
much shorter than the capsule, towards the apex spinulose-hairy; operculum
hemispheric-conic, shining, dark chestnut-brown, terminating in an oblique
linear-subulate rostrum 1.8 mm long. Known from the following ocalities:
In western Pennsylvania this moss is thus far known only from the extreme south-
western counties, as follows: Allegheny Co.: Along a shaly roadside bank at the west
end of Fern Hollow Bridge, Pittsburgh. O. E. Jennings, March 8, 1903. Type (figured).
Two pockets of specimens deposited in the Pennsylvania Herbarium of the Carnegie Mu-
seum). Same locality March 26, 1910. O.E.J.; Stoops Ferry, October 7, 1905. O.E.J,
and G.E.K.; Library P.O., Apnl 29, 1906. O.E.J. Beaver Co.: Beaver Falls, May 11,
1907. O.E.J. Bedford Co.: Wills Mt., near Hvndman, October 9, 1904. O.E.J. BuT-
inp Co.: Valencia, September 27, 1905. O.E.J. Fayette Co.: Ohio Pyle, May 30-31,
1908. O.E.J. Greene Co.: 2 mi. e. of Waynesburg. C.M.B. Apr. 23, 1938. Law-
pi-, nce Co.: Ravine below Graceland Cemetery, New Castle. 1906. Susan Gageby.
Washington Co.: Hanlin, May 23, 1908, and Charleroi, June 24, 1908. O.E.J.
Westmoreland Co.: "Shades," east of Blackburn, March 25, 1910. O.E.J, and G.K.J.
5. Atrichum angustatum var. PLURILAMELLATUM (Jennings) Frye
{Catharinaea plurilamellata Jennings)
Plate XXXI
Loosely cespitose, dark green, dioicous: stems simple or sparsely branched
below, slightly radiculose towards the base, about 1.5 cm tall, erect, flexuous,
reddish-brown; lower leaves dark green to purplish, short, becoming abruptly
longer above; upper leaves clustered, erect-spreading to erect, linear-lanceolate
4-7 mm long, .9-1.3 mm broad, margined, above with chlorophyll, obtuse to
sub-acute, toothed on the back towards apex, more or less doubly serrate in the
apical third, not very strongly undulate when damp, crisped and circinate
when dry; lamellae 7-12 in number, 8-14 (usually about 11) cells high, smooth
or sometimes sparsely minutely papillose, usually occupying from 1/2 to 2/3
of the width of the median part of the leaf; lower cells rectangular, about
.010-.016 X .025-.035 mm, the upper rounded-quadrate, about 007-.013 mm,
the lower marginal cells linear, incrassate, 2- to 3 -seriate and more or less
bi-stratose, towards the apex becoming rectangular, with triangular teeth about
.025-. 040 mm long; perichaetial leaves similar: seta solitary, erect, flexuous,
somewhat sinistrorse, about 1.5-2 cm long, smooth, shining, reddish-brown;
capsule oblong-cylindric, 4-5 mm long, 0.6-0.9 mm in diameter, erect, arcuate,
abruptly tapering at the base, reddish-brown, smooth, shining when fresh;
peristome teeth 32, linear-oblong, about .25 mm high, reddish-orange along
median portion, hyaline towards margins, when young with densely minutely
papillose margins, the lower one-fourth portion united into a basal membrane;
spores smooth, incrassate, orange-pellucid, globose, about .008- .011 mm; calyp-
tra about 5-6 mm long, narrowly cucullate, extending to about the middle of
capsule, spinulose-hairy on apex; operculum hemispheric-conic, shining reddish-
brown, terminating in a linear-subulate oblique beak about 2 mm long.
Known only as follows: Allegheny Co.: Powers Run, September 21, 1905, May
30-31, 1908. O.E.J. Fayette Co.: On shaded woodland bank, Ohio Pyle. O.E.J, and
G.E.K., September 10, 1905 (figured). Type. (Deposited in Pennsylvania Herbarium of
the Carnegie Museum). Westmoreland Co.: Miss K. R. Holmes, 1902.
Jennings: Manual of Mosses— 22. Polytrichaceae 165
2. PoGONATUM Beauvois
Dioicous: gregarious to weakly cespitose: fertile stems arising from a
creeping underground stem or from a radiculose protonema, erect, stiff, short
and simple or some longer and branched; leaves gradually longer upwards,
erect-spreading to recurved, more or less clasping at base, stiffened by mostly
numerous lamellae, especially towards the apex, the margins more or less dis-
tinctly spinulose, leaf-blade smooth dorsally, usually for the most part bi-
stratose; the lower part of costa narrow and plane, dorsally towards the apex
toothed; cells of the leaf-blade small, incrassate, in the unistratose border
mostly quadrate or transversely elongate, basal cells elongate to linear, yellow-
ish to hyaline, thinner walled: seta solitary, sometimes more, castaneous; cap-
sule erect, straight or curved, cylindrical, without stomata; peristome-teeth 32,
pale to yellowish-brown with a darker axis; operculum convex and more or less
long-rostrate; calyptra mitrate, densely hirsute, more or less shaggy.
A large genus of about 160 species, growing on earth, widely distributed;
about 45 species reported for North America; only one species yet found in
our region.
Key to the Species
A. Leaf-margins entire; lamellae 25-35 (P. brachyphyllum [Richard] Schwaegrichen) *
A. Leaf-margins more or less serrate; lamellae 10-15 pensilvanicum
1. PoGONATUM PENSILVANICUM (Hedwig) Paris
(Polytrichum pensilvanicum Hedwig; Pogonatum breyicaule
Beauvois; P. tenue E. G. Britton)
Plate XXXII
Plants scattered on a green felt-like persistent protonema which covers the
moist, bare clay: stems very short, usually about 1-2 mm long; leaves lance-
olate-subulate, the lower shorter and more ovate, narrowing abruptly to an
acuminate apex, margins serrulate in the upper half; upper leaves lanceolate
with a long-acuminate, serrulate apex, appressed or somewhat spreading; lamel-
lae 10 to 15, 5 or 6 cells high, terminal cell orbicular to ovoid in cross-section,
smooth: seta slender, smooth, yellowish to reddish; calyptra light yellow, very
hairy and more or less shaggy, completely covering the capsule; capsule erect,
symmetric, long cylindric, minutely papillose, slightly or not at all constricted
below the lid, yellowish to reddish, about 4 mm x 0.8 mm; lid obtuse to trun-
cate, abruptly tipped with a beak about 0.4 mm long; spores maturing in our
region about November.
A common moss on bare clay banks, especially if somewhat moist and
shaded, from Nova Scotia to Missouri and south to Alabama.
Usually found on clay roadside banks where the soil is covered with the
dense felt of protonema above which are borne the sporophytes with their
light gray shaggy calyptra.
Now known from 14 counties in western Pennsylvania and probably occurs in all.
Specimen; figured: Darlington Hollow, Allegheny Co., Oct. 12, 1908. O.E.J. &.' G.K.J.
* On clay banks, from the Gulf States extending north to New Jersey and Penn-
sylvania.
166 American Midland Naturalist Monograph No. 6
3. PoLYTRiCHUM Dillenius, Hedwig
Hair-Cap Moss
Dioicous; antheridial flowers cup-shaped, sprouting from the middle:
robust, stiff, in green to bluish-green, mostly high tufts; stem with a complete
central strand, rising from a subterranean rhizome, often whitish tomentose,
mostly simple; leaves dense, erect spreading to recurved, drying stiffly erect,
from a sheathing scarious base elongate-lanceolate to linear-subulate, non-bor-
dered, with margin plane to involute, sharply serrate, the sheathing base uni-
stratose and hyaline, at least at the angles; lamina bi-stratose except at the
margin; costa narrow and flat below, above stronger, dorsally toothed towards
apex, mostly excurrent as a colored, toothed awn; lamellae erect, high, numer-
ous, covering the costa and the bi-stratose lamina ventrally; cells in the sheath-
ing base elongate-rectangular to linear, narrower marginward; laminal cells
small, incrassate, quadrate-hexgonal : sporogonia solitary; seta long, stiff yellow-
ish-red to purplish, often drying flat and sinistrorse; capsule first erect, finally
inclined or horizontal, mostly prismatic, 4-6-angled, oblong to cubic, collum
hemispheric or disk-like, with stomata; spore-sac free; teeth 64, with a colored
axis, basal membrane colored; operculum large, conic to convex, rostrate;
calyptra cucullate and with the long, shaggy hairs completely enclosing the
capsule.
About 100 species, ma-'nly on soil in the cooler parts of the globe; about
20 species in North America; at least 5 species in our range.
Key to the Species
A. Exothecial cells of the capsule not pitted B
B. Exothecial cells of the capsule pitted C
B. Marginal cells of the lamellae like the others in size and cell-walls 1. P. gracile
B. Marginal cells of the lamellae thicker walled and in cross-section obversely pyri-
form 2. P. ohioense
C. Leaf-margin entire, broad and infle.xed; marginal cells of lamellae not emarginate in
cross section D
c. Leaf-margin sharply serrulate, not inflexed; marginal cells of lamellae emarginate in
cross-section 6. P. commune
D. Excurrent costa long, hyaline; plants low (about L5-2 cm tall), simple
- 3. P. piliferum
D. Excurrent costa red, short; plants larger E
E. Stems not or but slightly tomentose; capsules oblong-tetragonal 4. P. juniperinum
E. Stems whitish-tomentose below; capsules more or less cubical 5. P. alpcstre
1. POLYTRICHUM GRACILE Dickscn
Plate LXXII
Tufted, dark green, erect; stems 3-10 cm tall, matted together below with
brownish tomentum; leaves erect-spreading when moist, somewhat curled and
twisted when dry, the upper ones longer, 7-10 mm long, the lower one- fourth
to one-third portion of the leaf oblong, about 1.5 mm wide, abruptly tapering
upwards into a long, very slenderly acuminate, serrulate-spinose tip, the basal
sheathing portion of the leaf with thin rectangular cells .035-.070 mm long,
one-half to one-third as wide, with narrower cells in a few marginal rows, the
Jennings: Manual of Mosses— 22. Polytrichaceae 167
cells in the lamina at the middle of the leaf in 4-9 rows, thick-walled, rounded-
quadrate, .009-,014 mm in diameter, the lamina above the sheathing base in-
creasingly serrulate with one-celled somewhat spreading teeth; costa at base
about one-fourth the width of the leaf, narrowing slenderly above into a sharply
serrulate, red, awl-like point about .007-. 010 mm long; lamellae numerous,
4-6 cells high, the cells equal in size, smooth, rounded, about .014-. 015 mm in
diameter, the terminal one slightly obtusely pointed: seta erect, slender, 4-6 cm
long, pale yellowish-red; capsule more or less horizontal, ovoid-cylindric, 4-5
mm long and half as thick, rather irregularly obtusely 4-6-angled, considerably
shrunken and wrinkled when dry, the apophysis about half as wide as the
capsule and only moderately developed; calyptra reddish-brown, about covering
the capsule; operculum low-hemispheric, long-rostrate; peristome teeth 64, pale
yellowish-red, with a median darker line and several indistinct, longitudinally
parallel atriae: capsule wall of thick, irregularly rectangular or rounded-hexag-
onal cells .035-.040 mm in diam.eter; spores oreenish-yellow, thin-walled, faintly
warty, .018-.020 mm in diameter, mature in early summer.
Widely distributed in bogs, heaths, and boggy woods, northern Eurasia
and North America; New Zealand.
Our only collection made by Charles M. Boardman, July 15, 1949, in Mt. Davis bog,
Somerset Co., Pa., altitude 2400 feet (figured).
2. PoYYTRlCHUM OHIOENSE Renaud and Cardot
Ohio Hair-Cap Moss
Plate XXXII
Erect, loosely cespitose, about 2.5-7 cm high (gametophyte), olive-green:
stems wiry, slightly or not at all tomentose below, chestnut-brown; leaves
widely spreading when moist, erect-appressed when dry, or with the tips flexu-
ous-spreading, lower leaves small and linear, the upper about 8-12 mm long, the
linear-lanceolate limb spreading from an oblong sheathing base about 2-3 mm
long, the limb serrate and ending in a spinulose, stiff, pellucid acumen, very
strongly costate, the costa with about 40-50 lamellae; lamellae 4-6 cells high,
the terminal cell wider but not longer and not bi-cuspidate or retuse; cells in
alar portion of sheathing base of leaf elongate-rectangular, about .010-.015 x
.100-. 130 mm, prosenchymatous or parenchymatous, in middle of sheathing
base narrower and proportionally longer, somewhat incrassate, in limb rounded,
incrassate and pellucid-opaque; perichaetial bracts similar but with a longer
more hyaline sheathing base and a narrower hmb: seta 4-8 cm long, wiry,
fle.xuous, lustrous, chestnut-brown shading to golden above, erect; capsule erect
soon becoming more or less horizontal, acutely 4- (5) angled, narrowed
towards the base, about 2-3 by 4-6 mm, yellowish-brown, hypophysis small but
distinct; peristome-teeth pale yellow with a darker median portion, about
0.2-02.5 mm long; spores round, smooth, .015-. 017 mm, mature in midsummer;
cells of exothecium about .007-.010 by .010. 015 mm, quadrate to hexagonal,
incrassate, non-p>orose; operculum with a rostrum about equal in length to the
diameter of the capsule; calyptra yellowish, exceeding the capsule.
168 American Midland Naturalist Monograph No. 6
On earth in moist woods, Alaska to Labrador, south to Missouri and
Alabama, also in northern Europe.
Common in old fields, open woods, etc., esfsecially on peaty soil and clay hummocks
in the northern and upland counties, where it forms dark patches. Now known from 26
counties in western Pennsylvania and probably occurs in all. Specimen figured: Ohio
Pyle, Fayette Co., Sept. 1-3, 1907. O.E.J, and G.K.J.
3. PoLYTRlCHUM PILIFERUM Schreber, Hedwig
Plates XXXII, LXIV
Rather loosely cespitose, light green, rather glaucous: stems simple, erect,
1-4 cm high, purplish-brown, radiculose slightly at the base, leafy only in the
upper 1 cm or thereabouts; leaves when moist ascending, when dry imbricate-
appressed, base hyaline, rounded-oblong, about 1.5 mm long, the limb narrowly
lanceolate, about 3 mm long with wide margins inflexed and in the upper part
meeting or overlapping, the apex abruptly terminating in a hyaline, linear,
dentate arista about 1 mm long; costa wide, with about 25-35 lamellae ven-
trally, dorsally more or less papillose or dentate; lamellae usually of 4-7 cells,
the terminal one slightly wider and apically abruptly elongate; leaf-cells in
alar region of sheathing base quadrate to rectangular or hexagonal, hyaline,
slightly incrassate, in middle of sheathing part larger, rectangular, about .015-
.018 X .030-.040 mm, somewhat brownish-pellucid, incrassate, at base of limb
abruptly passing into rather opaque or brown-pellucid, much incrassate,
rounded cells, about .010-. 015 mm in diameter, in reflexed margin of limb
larger and irregularly obliquely rhombic: seta about 2-3.5 cm long, erect, flexu-
ous, lustrous, light chestnut-brown to paler above; capsule small, about 2-2.5
mm long, tetragonal -oblong to almost cubic, sharply angled, erect to pendulous,
usually horizontal in age; operculum shortly rostrate; calyptra covering whole
capsule; cells of exothecium hexagonal with a large oblong pore one-half the
diameter of the cell; peristome-teeth rather hyaline, about 0.2 mm high; spores
round, smooth, about .010-.012 mm, mature in mid-summer.
In dry. sandy soil, heaths, etc., in cooler regions over almost the whole
earth. In North America ranging from the Arctic regions south to California.
Erie Co.: On sand-plain, Presque Isle, Sept. 20-22, 1906. O.E.J. Fayette Co.:
Upper Cucumber Run, Ohio Pyle, June 23, 1935. C.M.B. (Plate LXIV) McKean
Co.: Bradford, Dec. 23, 1896. D.A.B. (Plate XXXII) Washington Co.: Near
Washington. Linn & Simonton. (Porter's Catalogue).
4. POLYTRICHUM JUNIPERINUM Willdenow, Hedwipr
Juniper Hair-Cap Moss
Plate XXXIII
Rather loosely cespitose, erect, light green and somewhat glaucous: stems
slightly tomentose at base, in our specimens about 5-9 cm high, brown; leaves
rather crowded, when moist spreading, when dry erect-appressed, or in the older
stems somewhat spreading, the base oblong, sheathing, the limb lance-linear,
5-6 mm long, the margin entire or crenulate and inflexed, the costa strong and
excurrent into a reddish dentate arista; cells at base of sheathing portion of
Jennings: Manual of Mosses — 22. Polytrichaceae 169
leaf lincar-rectanguIar, mainly parenchymatous, moderately incrassate, in ;niddle
portion of sheath relatively wider, about .010 x .040-. 090 mm, both prosen-
chymatous, in inflexed margin of limb obliquely quadrate-rectangular-elongate,
decidedly incrassate; lamellae about 30-40, usually 6-7 cells high, the terminal
cell somewhat broader and with an abruptly narrowed shortly prolonged apex;
perichaetial leaves more hyaline with a longer sheathing base and a considerably
longer slightly dentate arista: seta erect, flexuous, lustrous, about 4-6 cm high,
somewhat sinistrorse; capsule tetragonal-oblong, about 3-5 x 2-2.5 mm, sharply
angled, reddish to dark chestnut-brown when old, apophysis short but rather
distinct, capsule p>endulous to horizontal; spores round, smoothish, about .008-
.011 mm, mature in midsummer; cells of exothecium elongate-hexagonal to
quadrate-hexagonal, the perforation linear-oblong, about half as long as cell.
Heathlands, dry pastures, thin woods, etc., in hilly or mountainous regions
the world over. It occurs from Arctic America south to North Carolina.
Now known from eighteen counties in western Pennsylvania. Specimen figured: On
ground in dry pasture, near Kittanning, Armstrong Co., Sept. 24, 1904. O.E.J.
5. PoLYTRlCHUM ALPESTRE Hoppe, Schwaegtichen*
(Polytrichum strictum Banks, Menzies)
Plate XXXIII
Large mosses forming dense tufts up to 20 cm deep, tufts deeply matted
with a dirty-white tomentum: stems branching, rather slender, often matted
tomentose to within 2-3 cm of the apex; leaves more or less rigidly appressed-
imbricate when dry, when moist with the lance-linear limb spreading and 4-5
mm long, from an oblong sheathing base about 1.5 mm long, margin of limb
entire, inflexed, the apex rather abruptly narrowed into a linear, reddish-
pellucid, slightly serrate acumen, leaves dorsally serrulate nearly to the sheath-
ing base, the costa bearing ventrally about 25-35 lamellae; lamellae 5-8 cells
high, the terminal cell broader and abruptly narrowing to an obtuse short
acumen, as seen in cross-section; areolation of sheathing base almost hyaline,
not so markedly incrassate, the median basal rectangular, 4-8 times as long as
broad, towards the margin and upwards in the sheathing portion narrower,
longer, either prosenchymatous or parenchymatous, cells of inflexed margin of
limb obliquely quadrate or rectangular to linear-oblong in the border, incras-
sate; perigonial leaves of male plant obovate-orbicular, the costa broad, lamel-
late on the uppver half, percurrent in a broadly acute acumen: seta erect, slen-
der, wiry, sinistrorsly flexuous, 4-8 cm long, lustrous, rich chestnut-brown
below and lighter above; capsule more or less cubic, 2-3 mm long, acutely
angled, papillose, yellowish to chestnut-brown, apophysis distinct; cells of
exothecium more or less hexagonal, the central pore round to oblong, often
one-third the diameter of the whole cell; peristome about 0.2 mm high, teeth
rather slender; calyptra yellowish-brown, covering the capsule; operculum flat-
tened, the rostrum about 0.5 mm long. Evidently mature in midsummer.
* Frye, T. C, m Grout's Moss Flora, reduces this to a variety of P. jumperirium, i.e.
P. juniperinum var. alpestre Bryologia Europaea.
170 American Midland Naturalist Monograph No. 6
From the Arctic regions of North America south to North CaroHna;
southern South America; northern Europe and Asia. In bogs or boggy woods.
Rar^ in our region. Crawford Co.: In small Cassandra bog near Linesville, May
28, 1908. O.E.J, (figured). Washington Co.: Hanlin, in tuft of Leucobryum. May
21, 1908. O.E.J.
6. PoLYTRlCHUM COMMUNE Linnaeus, Hedwig
Plate XXXIII
Large, erect, 10-20 cm high, loosely cespitose in large masses, rather dark
olive green: stems simple, flexuous, woody, slightly or not tomentose at base,
chestnut-brown, rather densely foliate above; lower leaves small, linear, becom-
ing gradually larger above up to about 15 mm long, the limb linear-lanceolate
from an oblong sheathing base, when moist spreading or recurved, when dry
appressed-erect, serrate to the sheathing base, the apex linear, serrate, pellucid;
areolation at base of sheath parenchymatous, rectangular, above becoming
linear-prosenchymatous, abruptly grading at base of limb into rounded incras-
sate cells about .0I0-.015 mm in diameter, towards apex becoming elliptic
with the longest diameter transverse, all pellucid to more or less opaque;
lamellae 40-60, 4-9 cells high, the terminal cell broader and retuse to bi-cus-
pidate at apex, the lamellae extending almost to the base of limb; perichaetial
bracts up to 20 mm long, with a longer sheathing portion and few or no
lamellae; antheridial flowers conspicuous, cup-shaped, the broadly obovate
shortly acuminate bracts which form the cup being about 4 mm long, the costa
broad and weak below but stronger and bearing numerous lamellae in the
upper half; the successive annual growths of the male plant taking place from
the center of the antheridial flower of the preceding season: seta wiry, flexuous,
6-10 cm long, lustrous, chestnut to light golden-brown; capsule erect, but later
inclined, and, when old and empty, cemuous, light to deep chestnut-brown,
more or less cubial or shortly rectangular, apophysis discoid, distinct; capsule-
urn about 3-5 mm long; operculum low-conic, the beak about 1 mm long,
straight or curved; peristome-teeth about .25 mm high, more or less reddish-
pellucid; exothecial cells hexagonal, the outer face convex and with a rounded
to elliptic pore; spores round, smooth, about .008-. 010 mm, mature in mid
summer — calyptra covering the whole capsule, rather lustrous, yellowish-brown.
Cosmopolitan; in North America almost throughout, in marshy places,
pastures, woods, etc.
Now (1942) known from every county in western Pennsylvania except Blair, Clarion,
Forest, Fulton, Himtingdon, Indiana, and Venango, but probably occurs in all. Sp>eci-
men figured: Near Hartstown, Crawford Co., in wet meadow, June 28, 1908. O.E.J.
This is the form which some authorities recognize as var. uliginosttm Huebener, as
follows:
6a. PoLYTRiCHUM COMMUNE var. ULIGINOSUM Huebener
Plate XXXIII
In this variety the stems are less strong and rigid than in the species, and
the leaves in the dry specimens are wide-spreading to recurved. It is rather
rare in the eastern part of the United States. Perhaps not sufficiently distinct.
Jennings: Manual of Mosses — 23. Hedwigiaceae 171
Cambria Co.: Cresson, May 18, 1904. O.E.J. ; upland plateau near St. Lawrence,
July 24, 1908. O.E.J. Crawford Co.: Near Linesville, May 12, 1908; and Harts-
town, June 28, 1908. O.E.J, (figured). (Near Linesville this variety forms quite exten-
sive heaths in the low-lying peaty pastures around the Pymatuning Swamp, especially on
the low mounds of peaty soil formed by the uprooting and subsequent decay of trees).
6b. POLYTRICHUM COMMUNE var. PERIGONIALE (Michaux)
Bryologia Europaea
Lamellae 6-9 (instead of 4 9 cells high), inner perichaetial bracts much
longer than foliage leaves; operculum with straight beak. Arctic America
south to North Carolina. Not yet known from our region.
Family 23. Hedwigiaceae
Autoicous; paraphyses long, yellow, filiform: more or less robust, stiff,
cespitose; stem without central strand, irregularly to almost pinnately branched,
rarely with long, pendent, 2-3-pinnate branches, densely-leaved, radiculose
below, sometimes stoloniferous; leaves about 8-seriate, spreading, drying im-
bricate, broad, thin, ecostate, concave, sometimes plicate, papillose; lamina one-
layered, golden-brown at base, cells incrassate, punctate, non-margined, with
several rows of small quadrate cells in the alar portion, or margined with the
alar portion concave, sharply differentiated by large, colored, 4-6-sided cells;
leaves on stolons recurved-squarrose, from a wider base suddenly long pilifer-
ous-acuminate; perichaetial leaves erect, longer than the stem-leaves, with
ciliate margins at apex: seta various; vaginula ciliate; capsule short, erect,
shortly and thickly collumate; annulus none; spores large; operculum low,
convex to rostrate; calyptra minute and mitrate to large and cucullate.
A small but widely distributed family of six genera, only one genus in
our region.
1. Hedwigia Ehrhart, Hedwig
Autoicous: laxly cespitose, glaucous-green: rarely stoloniferous, erect to
ascending, irregularly branched; leaves concave, ovate, tipped with a hyaline,
serrate to ciliate acumination, margins revolute, entire, non-bordered; leaf-cells
two- to several-papillose, papillae on both sides, the upper cells oblong, the
lower elongate, the median basal yellow, linear, becoming quadrate and brown-
ish towards the angles; perichaetial leaves larger, the upper margins furnished
with long, sinuose, articulate, sometimes toothed cilia: seta about 5-8 mm long,
yellow, thicker upwards; capsule immersed, obovate to globose, smooth, pale
brown, the mouth red and wide; spores .028-.032 mm, yellow with vermiform
lines; operculum plano-convex, red, sometimes unbonate; calyptra minute,
conic-mitrate, fugaceous, covering only the apex of the operculum.
A cosmopolitan genus of one very variable species, occurring on non-
calcareus rocks.
1. Hedwigia ciliata [Ehrhart] Bryologia Europaea
(H. albicans Lindberg; Fontinalis albicans Weber; Anictangium ctliatum Hedwig)
Plate XXXIV
In patches of varying size up to quite large, blackish or brownish below,
172 American Midland Naturalist Monograph No. 6
glaucous-green above, more or less hoary, especially in late summer or in
autumn, owing to the colorless tips of the leaves: stems from 2 or 3 up to 8 or
10 cm long, rather slender, irregularly forking and branching, the branches
usually rather short; leaves more or less secund on the procumbent stems,
when dry imbricated but with recurved apex, when moist spreading, concave,
ovate, 1.5-3 mm long, the apex sub-obtuse to long-acuminate, papillose-denticu-
late to spinulosely denticulate, more or less hyaline; costa none; the median
basal leaf-cells yellowish pellucid, not papillose, narrowly linear, incrassate,
porose, towards the margin and in upper part of leaf the cells sub-quadrate or
rectangular, with more or less sinuose walls, the cells in the angles often brown-
ish and larger, the median and upper cells prominently papillose, longitudinally
seriate, varying from quadrate to rounded or hexagonal; perichaetial leaves
prominently ciliate towards the apex, not plicate: seta practically none: capsule
sub-sessile, immersed, globose-oblong, about 0.6-0.9 mm in diameter, wide-
mouthed ?nd truncate when dry and empty, red-rimmed, the urn castaneous;
lid convex, sometimes mamillate, about three-fourths as wide as the median
diameter of the urn; calyptra small, sub-cucullate and fugacious; annulus none
but one or two rows of exothecial cells at the rim of the urn smaller, laterally
elongate, and castaneous-pellucid; peristome none; spores mature in spring,
minute, shallowly pitted, pale, thin-walled, about .025-.028 mm: autoicous.
On dry rocks, boulders, stone-walls, etc., in non-calcareous habtiats; almost
cosmopolitan; in North America occurring from the Arctic regions to Mexico.
Common in our region. Now known from Allegheny, Beaver, Bedford, Fayette, For-
est, McKean, Somerset, Washington, and Westmoreland counties. Specimen figured: Ohio
Pyle, Fayette Co., May 30-31, and July 4, 1908. O.E.J. Not yet known from the glaci-
ated region of western Pennsylvania.
la. Hedwigia ciliaTa f. LEUCOPHAEA (Bryologia Europaea) Jones
(H. albicans var. leucophaea Limpricht)
Very hoary; more robust than the species: leaves more falcate, wider, the
hyaline base of the piliferous acumination occupying about the whole upper
third of the leaf.
With the type and in the same general habitat.
Huntingdon Co.: Stone Creek, T. C. Porter. (Porter's Catalogue). Westmore-
land Co.: T. P. James. (Porter's Catalogue).
PLEUROCARPI
In the following families the flowers, as a general rule are borne in the
axils of leaves along the side of the stem. The antheridial flowers are enclosed
in an involucre of modified leaves, the perigonial bracts; the archegonial flow-
ers have a similar involucre of perichaetial bracts; and the sporophyte thus is
borne laterally on the stem. Most of the pleurocarpous mosses have a creep-
ing habit.
Family 24. Fontinalaceae
Dioicous or autoicous: filiform paraphyses few: slender to robust, aquatic,
floating, blackish-green or reddish-brown: stem without central strand, 3-5-
Jenn.ngs: Manual of Mosses — 24. Fontinalaceae 173
angled, or round, much branched but bare below, fastened by a cushion of
rhizoids at the base; leaves 3- and 5-seriate, ovate-acute to lance-subulate,
carinate to concave or plane, mostly decurrent, rarely winged, entire or dentate
at apex; lamina uni-stratose above, bi- to tri-stratose below, with single costa
or none; median leaf-cells mostly elongate prosenchymatous, smooth, the basal
orange, laxer, rarely loosely rhombic hexagonal: seta rudimentary or normal:
capsule erect, non-collumate, without annulus, without stomata; peristome
none, single, or double, teeth when present 16, hygroscopic, as long as or
shorter than the segments; mostly linear, orange- to brown-pellucid, non-bor-
dered, mostly papillose, ventrally with projecting transverse trabeculae; inner
peristome without basal membrane, segments filiform, 16, usually more or less
united into a carinate, trellis-like cone, rarely free and appendiculate; lid short-
conic to rostrate; calyptra small and conic or cucullate and reaching to below
the capsule.
A family of six genera, confined almost exclusively to the temperate and
colder parts of the Northern Hemisphere; two of the genera in our range.
Key to the Genera
A. Leaves ecostate; calyptra short 1. Fontinalit
A. Leaves costate; calyptra enclosing the whole capsule 2. Dichelyma
1. FoNTiNALis Linnaeus, Hedwig
Water Mosses
Dioicous: floral branches apparently axillary, very leafy; antheridial clus-
ters short and obtusely gemmiform; archegonial branches elongate and acute:
slender to very robust: stems sharply 3-angled to round, much branched; leaves
of one form, 3-seriate, otherwise as for the family, ecostate; perichaetial leaves
larger, almost enclosing capsule, broadly obovate, obtuse, lacerate when old:
vaginule and seta rudimentary; capsule oval to ovate, mostly delicate; peristome
double, inner and outer of same length, teeth 16, linear-lanceolate, orange to
brownish, plane, papillose, mostly apically united in pairs, the divisural zigzag,
articulations prominent, the trabeculae projecting both ventrally and laterally;
segments 16, filiform, united by lateral processes into a plaited cone, rarely free
and appendiculate; spores irregular in size, mostly green, almost smooth; lid
conic, calyptra reaching but little below the operculum, the base lacerate
when old.
A genus of about 60 species; about 30 reported for North America; at
least five occurring in our region.
Most of the specimens of this genus in our collections have been verified
by Winona H. Welch.
Key ro the Species
A. Older stem-leaves carinate or keeled-conduplicate 1. F. antipyretica v.ir. gigantea
A. Older stem-leaves not as above B
B. Leaves essentially one form, not dimorphic - D
B. Leaves dimorphic C
C. Vernal leaves lance-ovate to siiborbicular, summer leaves narrowly lanceolate; apical
cells reminding one of Sphagnum 2. F. biformis
C Stem-leaves lance-ovate to lanceolate, branch leaves narrowly lanceolate, both acumi-
174 American Midland Naturalist Monograph No. 6
nate 3. F. Sultivanti
D. Stem-leaves rather firm, more or less distinctly concave E
D. Stem-leaves plane or nearly so, mostly distinctly flaccid F
E. Leaves narrowly to oblong lanceolate, long-acuminate 4. F. dalecarlica
E. Leaves ovate-lanceolate, apex usually denticulate, broadly obtuse or apiculate
5. F. novae-angliae
F. Alar cells enlarged, forming distinct auricles, and leaves clasping the stem
10. F. Lescursi
F. Leaves not clasping the stem G
G. Majority of leaves 4-8 mm long by 0.5-1.5 mm wide, very long and narrowly acu-
minate; auricles distinct 7. F. flaccida
G. Majority of leaves 2.4-5.5 mm long _ H
H. Apex of leaf usually acuminate, long and narrow 6. F. hypnoides
H. Apex of leaf usually shorter and broad i
1. Leaf-auricles usually small but distinct 9. F. Duriaei
I. Leaf-auricles none or quite indistinct 8. F. nitida
1. Fontinalis antipyretica Linnaeus, Hedwig
Leaves lance-ovate to lanceoval, 5-8 x 2-4 mm; median leaf-cells about
6-15:1; perichaetial leaves obtuse; peristome teeth papillose. — Apparently most-
ly represented in eastern United States by var. gigantea.
1. Fontinalis antipyretica var. gigantea Sullivant
[Fontinalis gigantea Sullivant)
Plate LXIV
Floating, long, dark, brownish-green or golden green: stems denuded below,
slender, up to sometimes 6 or 8 dm. long, irregularly divided; the branches
turgidly three-cornered and sometimes 2 or 3 dm long; leaves deeply concave,
carinate, up to 6-8 mm long, 3-6 mm wide, broadly ovate or lance-ovate to
almost orbicular, entire, mostly broadly obtuse at the tip; median leaf-cells
about 7-11:1, linear-rhomboid and more or less vermicular, the apical and
basal shorter and wider, the alar sub-rectangular and somewhat inflated, form-
ing auricles which are convex upwards (adaxially) ; perichaetial leaves closely
imbricated, the upper sheathing, truncate, rounded, entire, or lacerate at the
apex: capsule rather small, sub-sessile, usually only the rostrate calyptra and
the conic lid emergent from the sheathing perichaetial leaves when mature,
about 2:1, m.ore or less turgid-oblong; lid reddish; peristome usually a bright
coral color, smoothish, the inner peristome united at the apex and sometimes
well down towards the middle into a lattice-work, the bars incomplete below;
spores mature in summer.
In cool streams and in ponds, on stones or on wood; Europe, and from
Canada through the northern United States to Alabama. Scarce and confined
to the mountains in our region.
Blair Co.: T. P. James. (Porter's Catalogue). Cambria Co.: T. P. James. (Por-
ter's Catalogue). Cameron Co.: Submerged rocks in brook. Tannery School, Lumber
Twp., Sept. 1, 1935. S. K. Eastwood. Lycoming Co.: In spring. Bald Eagle Mt.,
south of Williamsport, Feb. 15, 1949. Harry Roslund. McKean Co.: D.A.B. (Por-
ter's Catalogue). Somerset Co.: Glade Run Swamp, eastern border of county. June 28,
1942. O.E.J. Westmoreland Co.: Big Springs, west of Bakersville, Oct. 31, 1933.
C. M. Hepner (figured).
Jenn.ngs: Manual of Mosses — 24. Fontinalaceae 175
2. FONTINALIS BIFORMIS SuIIivant
Yellowish green to dirty green: stems long, much-branching; leaves of two
kinds; the vernal large, soft, lance-ovate, concave, blunt to acute, when fresh
and moist quite prominently three-ranked, and rather widely spreading, the
apical cells somewhat like Sphagnum cells; the summer leaves much smaller,
narrower, convolute and tubulose above, rigid, covering the younger branches;
median leaf -cells of the vernal leaves linear, the apical broadly rhomboidal and
sphagniform, the angular quadrate-oblong, much larger, forming small de-
current auricles; costa none; archegonial clusters rare, situated towards the base
of the stems; antheridial clusters usually 2 to 4 together and long-stipitate:
capsule oblong-oval, enfolded by the suborbicular inner perichaetial leaves;
lid conic, rostrate; peristom.e-teeth lance-linear, about 20-articulate, cilia tessel-
late and united at the apex, papillose.
In woodland rivulets and streams, Ohio and Indiana.
Rare in our region. Portage County, Ohio, and McKean Co.: (Porter's Catalogue).
3. FoNTlNALlS SULLIVANTI Lindberg
(F. Lescurii var. gracilescens SuIIivant)
Quite similar to F. Lescurti but smaller and more slender: very slender,
regularly pinnate with remote and attenuate branches; leaves distant, the stem-
leaves lanceolate, soft, narrowly long-acuminate, somewhat concave, acute to
somewhat obtuse, entire or sub-denticulate, yellowish, about 5 mm long, the
branch-leaves about half as long, more rigid, more concave, acuminate; peri-
chaetial leaves rather short as compared with F. Lescurii, not undulate at apex;
median leaf-cells linear-flexuous, the apical shorter and broader, the basal
shorter and broader, the alar much larger, inflated-oblong forming auricles:
capsules sessile, cylindric; lid conic, long-acuminate; peristome teeth papillose,
the inner peristome as in F. Lescurii; spores minutely roughened.
In stagnant water, swamps, etc., on bushes, tree roots, etc., eastern United
States. Not reported from our region.
4. FoNTlNALlS DALECARLICA Bryologia Europaea
Plate XXXIV
Stem.<: slender, much-branched, naked below, 1-3 (4) dm long, attenuate,
dark-castaneous, sub-lustrous; leaves somewhat close, erect-spreading to some-
what imbricate, more or less glossy, lance-oblong to narrowly lanceolate, acu-
minate, 2-3 mm long, often slightly toothed at apex, margins usually somewhat
involute, concave, sometimes very slightly auricled at base; leaf-cells prosen-
chymatous, rather incrassate, linear-oblong, about 10-18:1, the marginal slightly
narrower, the alar rectangular to irregularly quadrate-hexagonal, considerably
larger, usually slightly colored; p>erichaetial leaves apiculate, the apex finally
lacerate: capsule immersed, about 2 mm long; peristome orange to brownish,
the teeth slender, granulose, with about 14-22 lamellae, the inner peristome
with an imperfect lattice; spores muriculate, about .025-032 mm, mature in
summer.
176 American Midland Naturalist Monograph No. 6
In rapidly flowing streams, occurring from Greenland to Georgia, Tennes-
see, and Wisconsin; also in Europe.
Cameron Co.: On rocks in bottom of creek 3 mi. w. of Truman. June 22, 1935,
and Tannery School, Sept. 1, 1935. Sidney K. Eastwood. Centre Co.: In swiftly run-
ning mountain-stream about three miles south of Boalsburg, Sept. 22, 1909. Sterile.
O.E.J, (figured); Bear Meadows. T. C. Porter. (Porter's Catalogue); Stream, Tussey
Mt., near Shingletown, O.E.J. July 15, 1909. Fayette Co.: Sheepskin Run, one-half
mi. e. of Ohio Pyle. C.M.B. Nov^ 6, 1943. Somerset Co.: Clear Run, Shafer Run,
and Blue Hole Creek, all Laurel Ridge, various dates. C.M.B. Venango Co.: On sub-
merged rock in creek near St. George. Aug. 4, 1935. Sidney K. Eastwood. Westmore-
land Co.: On rocks in swift water. Rock Run, Laurel Mts., Aug. 31, 1925. O.E.J, and
G.K.J.; Fish Hatchery trough. Rector. W. R. Witz, April 4, 1937; Rock Run, Forbes
Forest. C.M.B. Sept. 16, 1934.
5. Fontinalis novae-angliae SuIIivant
Plate XXXIV
Rather bright green, fairly firm: stems usually 3-4 dm long, wiry, purplish-
black, slender, rather freely branching, naked below but quite densely foliate
towards the apex; leaves rather close, somewhat appressed, thin, entire, some-
times faintly serrulate at apex, the main branches with leaves about 2-2.5 mm,
broad when moist, the branches more or less linear-attenuate; leaves about
3-4.5 mm long, broadly ovate-lanceolate, concave, the margins somewhat
revolute, the apex cucullate, rounded-obtuse; median leaf-cells linear-vermicular
to somewhat narrowly linear-oblong, prosenchymatous, incrassate, about 6-15:1,
the alar cells forming a quite distinct group, quadrate to oblong, moderately
enlarged, somewhat incrassate and colored; capsule sub-cylindric to oblong-
oval, near base of stem, closely invested by the ovate-sub-orbicular perichaetial
leaves, which are lacerate when old; capsules rare; pristome-teeth colored, linear-
lanceolate, 18-20 articulate, slightly papillose; cilia tessellate and united at
apex only, minutely papillose; spores smooth.
In brooks and swift-running streams from Newfoundland to Ontario and
North Carolina.
Butler Co.: On rocks in swift stream, 5 mi. north of Zelienople. Oct. 3, 1925 and
4 mi. n.e. of Harmony, May 3, 1930. L. K. Henry; Walley Mill, Parker Twp. July 14,
1935. Sidney K. Eastwood. Clearfield Co.: In rapid stream north of Mahaffey, May
30, 1937. Sidney K. Eastwood. Fay'ette Co.: Cucumber Run, Ohio Pyle. July 18,
1895. C.M.B. Forest Co.: Buzzard Swamp, June 30, 1948. C.M.B. Huntingdon
Co.: Spruce Creek, T. C. Porter. (Porter's Catalogue). Venango Co.: Submerged
rocks in stream near St. George. Aug. 4, 1935. Sidney K. Eastwood. Westmoreland
Co.: Creek below Hillside Station, September 17, 1909. O.E.J, and G.K.J, (figured).
6. Fontinalis hypnoides Hartman*
Rather delicate, soft, slender, pinnately divided plants; leaves distant, flac-
cid, narrowly lanceolate to lance-ovate, gradually acuminate, 3-6 mm long,
usually entire; median leaf cells linear attenuate to linear-rhomboidal, the alar
more or less enlarged but indistinct, not forming distinct auricles; capsule about
* The( descriptions of F. hypnoides, nitida, and jiaccida have been largely compiled
from Welch's treatment in Grout s Moss Flora.
Jenn.ngs: Manual of Mosses — 24. Fontinalaceae 177
half emersed, oval; peristome teeth linear, papillose; trellis of inner peristome
perfect with appendiculate cross-pieces; spores more or less muricate, about
.012-.015 mm in diameter.
Various kinds of water habitats, United States and Canada. Not yet
reported from our region.
7. FONTINALIS FLACCIDA Renauld and Cardot*
Soft, delicate, yellowish green, slender, pinnately divided plants up to 35
or 40 cm long; leaves distant (1.5 mm apart), flaccid, concave at base, less so
above, narrowly lanceolate to lanceolate, 4-8 mm long, very slenderly acuminate,
the apex itself often obtuse to truncate, often denticulate; alar cells much en-
larged, hyaline to yellowish-brown, forming quite evident auricles: capsule
immersed, subcylindric, 2 mm long; teeth of peristome linear, slightly papil-
lose; trellis of inner peristome imperfect, muricate; spores smoothish.
Reported variously in stagnant or running water in various parts of eastern
United States. Not yet reported from our region.
8. FONTINALIS NITIDA Lindberg*
Soft, delicate, slender, pinnately divided plants up to 20-40 cm long; leaves
about 1 mm apart, flaccid, usually flat, lance-oblong to lance-ovate, gradually
acuminate but usually with a short and broad acumen; alar cells enlarged and
fairly distinct and often forming auricles: the oval capsule usually immersed;
peristome teeth linear, papillose; trellis complete and perfect, muricate, with
cross-bars appendiculate; spores finely muricate.
Asia; and in fresh (non-stagnant) waters, United States and Canada.
Not yet reported in our region.
9. FoNTlNALls DURIAEI Schimper
Plants soft and weak, green to olive- or yellow-green; stems slender, 10-30
cm long, pinnately but bushily branched, often denuded below; leaves about
0.5-1.5 mm apart, spreading to erect-spreading excepting at ends of branches
where imbricate, plane to sub-concave, lance-oblong to broadly lance-ovate,
rather abruptly and broadly acuminate, 3-5 (-7) mm long by 1-2.5 (-4) mm
wide; apex acute to sub-obtuse, denticulate to entire; median leaf-cells linear-
attenuate to linear-rhombic, .0085-. 017 mm wide; 6-18:1 alar cells more or less
enlarged, hyaline to yellowish-brown, rectangular to quadrate or sub-hexagonal,
auricles none or indistinct: upper perichaetial leaves oval-suborbicular, broadly
rounded at apex to apiculate, lacerate when old: capsules immersed, oval to
short-oblong, about 2 mm long with conical op>erculum; peristome brownish
orange, the teeth linear, papillose, 0.75-0.9 mm long, frequently united in pairs
at apex, with 15-35 lamellae; trellis perfect, muricate, with transverse strands
complete; spores minutely muricate or smooth, .01 5-. 020 mm in diameter,
mature in summer.
* The descriptions of F. hypnoides, nitida, and flaccida have been largely compiled
from Welch's treatment in Grouts Moss Flora.
178 American Midland Naturalist Monograph No. 6
In streams and falls, occasionally in swamps. Europe, Africa, Canada
and the United States.
Known in our region only from a creek west of Linesville, Crawford County, Pa., Aug.
4, 1909. O.E.J. (Det. by Winona H. Welch).
10. Fontinalis Lescurii Sullivant
Loose, soft, green to glossy golden-green: stems long, reaching sometimes
3 or 4 dm naked and blackish below, dividing and branching irregularly except
sometimes at the apex, where the branches may be arranged pinnately; leaves
erect-spreading, soft, obscurely three-ranked, concave, clasping at the base,
lance-ovate to lance-oblong and rather acuminate to a somewhat obtuse apex,
slightly denticulate at the apex, usually about 4-6 mm long; median leaf-cells
about 12-15:1, elongate-linear, flexuous, the apical and basal shorter and
broader, the alar enlarged oblong, inflated, forming quite distinct auricles;
perichaetia numerous towards the base of the stems, perichaetial leaves sheath-
ing, the inner rounded-obtuse, broadly oval, reaching nearly to the apex of
the mature capsule: capsule short, sub-cylindric, enclosed by the closely fold-
ing perichaetial leaves until almost mature, about 2.5:1; lid long-conic; peri-
stome-teeth red-orange, papillose, about 20-25-articulate, the inner peristome
more or less compeltely united into a lattice-work at the apex but free and
merely appendiculate below; spores mature in summer.
On stones in streams from Nova Scotia to Georgia and west to Tennessee
and Oklahoma.
Rare in our region. Huntingdon Co.: T. C. Porter. (Porter's Catalogue). -Mc-
Kean Co.: Bradford. D.A.B. (Porter's Catalogue).
2. DiCHELYMA Myrin
Dioicous; antheridial shoots small, gemmiform; archegonial shoots long;
slender to robust, shining, green to golden-brown, blackish below; branching
various, the branches recurved at the apex; leaves 3-seriate, falcate-secund to
circinate, lance-subulate from a slightly decurrent base, carinate-plicate, weakly
serrate; costa complete to long-excurrent; median leaf-cells linear, narrow, the
alar not wider; inner perichaetial leaves long, tubular, sinistrorsely wound
around the seta: seta long; capsule ovate, soft, brownish; peristome-teeth 16,
lance-linear, obtuse, papillose, spreading either when damp or when dry, often
more or less cleft or divided along the median line, trabeculae low and dis-
tant; inner peristome longer and sometimes falling away with the operculum,
segments filiform, more or less united; lid about as long as urn, conic, mostly
oblique and curved; calyptra enclosing the whole capsule, split along one side,
sinistrorse; spores small and uniform in size.
A rather widely distributed genus of 5 or 6 species; 4 species occurring in
North America; 2 species in our region.
Key to the Species
A. Leaves subulate; costa long-excurrent 1. D. capillaceum
A. Leaves acute; costa complete or almost so, subpercurrent to slightly excurrent
2. D. pallescens
Jenn.ngs: Manual of Mosses — 24. Fontinalaceae 179
1. Dichelyma capillaceum [Dillenius] Bryologia Europaea
(D. pallescens Sullivant and Lesquereux; Fontinalis capillacea Hedwig)
Yellowish above, brownish to blackish below; stems slender, often 10-15
cm long, with a few distichous, divaricate, or one-sided branches; leaves erect-
spreading, secund to falcate-secund, long-linear from a lance-oval base, about
5-7 mm long, serrulate towards the apex; costa long-excurrent; perichaetial
leaves linear, thin, ecostate, pale and twisted and reaching above capsule; leaf-
cells narrow, linear-rhomboid: seta short, slender; capsule small, pale yellowish,
thin-walled, ovate, the urn truncate and about 1.5-2:1, finally laterally emerging
from the perichaetium; lid high-conic; peristome double, the teeth shorter than
the inner peristome, narrowly linear, densely papillose, segments longer than
teeth, constricted at the articulations, pale yellow, papillose, forming a con-
nected lattice-work only above; spores mature in late summer.
On sticks and the bases of bushes in and around the edges of slow streams,
ponds, and swamps; Europe and from New Brunswick and Ontario southward
to North Carolina and Tennessee. Yot yet recorded in our region.
2. Dichelyma pallescens Bryologia Europaea
(Fontinalis capillacea Hooker)
Plate XXXV
Slender, light yellowish-green, sometimes glossy: stems usually about 5-10
cm long, the branching sub-distichous; leaves secund, more or less falcate, the
ends of the branches and stems appearing hooked, leaves oblong-lanceolate,
about 3-5 mm long, gradually long-acuminate, complicate-carinate, nearly
entire; or denticulate above, plane-margined, acute to obtuse; costa percurrent
or nearly so; median leaf-cells rhomboid-linear, prosenchymatous, about 8-15:1,
rather incrassate, the basal colored and somewhat shorter, a few alar indistinctly
wider and oblong, incrassate, the apical shorter; perichaetial leaves about as
long or usually longer than the seta and capsule together: seta about 4 mm
long, slender, enclosed in the perichaetium; capsule small, thin, ovate, yellow-
ish, about 1 mm long, truncate by the falling away of the lid, emerging later-
ally from the perichaetum; lid high-conic; peristome-teeth linear, rather rudi-
mentary, pale, castaneous-pellucid, with distinct divisural and lamellae, and
about 10-12 castaneous-pellucid, low ventral trabeculae; segments filiform,
longer than teeth, united only at the summit or entirely free, sometimes re-
maining on the ripe capsule only as short, filiform, cilia-like structures between
the teeth; exothecial cells rounded, castaneous-pellucid, incrassate-collenchymat-
ous, the upper laterally oblong and smaller; spores mature in summer, cas-
taneous-pellucid, incrassate, minutely papillose, varying from about .016-
.025 mm.
More or less inundated on sticks and the bases of bushes along creeks and
around ponds; New Brunswick to Minnesota and Pennsylvania.
Not yet found in our region, excepting along the northern border. McKean Co.:
Bradford. D.A.B. (Porter's Catalogue); Riverside, New York, a few miles north of
Bradford. D.A.B. October 18, 1897 (figured).
180 American Midland Naturalist Monograph No. 6
Family 25. Climaceae
Dioicous; flowers on secondary stems and at base of branches; gregarious,
large and stately, growing in swamps: stems rhizome-Iilce, subterranean, radicu-
lose, with smooth, branched, reddish-brown rhizoids, secondary shoots 3- to
several-angled, erect, with more or less tree-like branching, with central strand;
branches leafy, cylindric, simple, pinnate or bi-pinnate; paraphyllia numerous;
leaves dimorphous, the rhizome and lower part of stem having scale-like and
appressed colored leaves, the upper stem and branches having green leaves;
leaves plicate; costa simple, homogeneous, ending below the apex, at the base
widened by two or three layers of laminal cells; leaf-cells smooth, upwardly
narrow-rhombic, downwards linear, the basal orange, the alar hyaline, lax, thin-
walled and forming a distinct group; perichaetial leaves numerous, long and
slender; sporogonia often aggregated: seta long, erect, stiff, sinistrorse; capsule
erect and symmetric in Cltmacium; exannulate; peristome double with the parts
of equal length; teeth confluent at base, reddish-brown, articulate, papillose,
ours not transversely striate, the lamellae numerous; inner peristome yellow,
papillose, with more or less of a basal membrane, the segments carinate, more
or less gaping along the keel, cilia none; spores medium size; operculum ros-
trate from a convex base; calyptra cucullate.
Two genera: Pleuroz'opsis, with one species, in our regions bordering the
North Pacific, and the following:
1. Cltmacium Weber and Mohr
Mostly as characterized in the description of the family: branches simple,
or sometimes almost pinnate, unequal, attenuate; branch-leaves lance-ligulate
from a decurrent, auricled base, bluntly to sharply acute, sharply serrate above;
inner perichaetial leaves abruptly acuminate, entire, short-costate; costa of the
leaves strong, ending below the apex, dorsally toothed above: seta 15-45 mm
long, stiff, castaneous; capsule erect, symmetric, almost cylindric, castaneous;
teeth lance-linear, acuminate, with a dark red border, with low papillose dorsal,
plates, and with closely placed trabeculae; inner peristome orange, vertically
striate-papillose, segments linear, carinately gaping, finally divided; spores .015-
.020 mm, rusty, warty; calyptra long, narrow, enclosing whole capsule, cleft
on one side to apex, sometimes twisted.
A widely distributed genus of about 5 species: 3 occurring in North
America and extending into our region.
Key to the Species
'i. Plants of dendroid habit B
A. Plants not distinctly dendroid, median leaf-cells about 2-3:1 3. C Kindbergii
B. Auricles not prominent; median leaf-cells 8-10:1 1. C. dendroides
B. Auricles broad, median leaf-cells about 5-7:1 2. C. americamim
1. Climacium dendroides [Linnaeus] Weber and Mohr
(Hypnum dendroides Linnaeus; Leskea dendroides Hedwig)
Dendroidal, robust, bright or yellowish-green; the primary stems under-
ground, creeping, divided; the secondary stems rising to a height of sometimes
Jennings: Manual of Mosses — 25. Climaceae 181
10 cm, leafless below, bearing at the summit numerous erect-spreading, flexuous,
usually straight branches; leaves large; stem-leaves broadly amplexicaul, with a
more or less rounded and cucullate or apiculate apex; branch-leaves imbricated,
giving to the branches a thick and turgid appearance, about 2 x 0.7 mm, lance-
oblong to lingulate-oblong, denticulate at base but quite sharply serrate above,
plicate, somewhat cordate but little auriculate at the base; costa nearly reaching
apex; median leaf-cells about 6-10:1, linear-rhomboidal to linear-hexagonal,
shorter and wider towards the apex and towards the base, the alar somewhat
lax, wider, hyaline, few, forming small auricles; perichaetial leaves entire, non-
plicate, the inner sheathing: seta deep red, about 2.5-3 cm long; capsule erect,
castaneous, oblong-cylindric, about 4 mm long, about 3-4:1; lid often remaining
attached to the columella, straight, acutely rostrate; calyptra reaching to below
the capsule; peristome large, the teeth forming a cone when moist but usually
curved in between the segments when dry; spores mature in fall, green in color.
On wet ground in marshes, at borders of streams, margins of swamps and
lakes, etc.; Europe, Asia, and from Arctic America south to New Jersey and
the Northwest. It may eventually be found to occur in the northern part
of our region.
2. Climacium americanum Bridel
American Tree Moss
Plate XXXV
Loosely cespitose, robust, yellowish-green, lustrous: primary stems creeping
stolon-like in the leaf-mould, throwing up at intervals dendroidal secondary
stems to a height of 5-8 cm; secondary stems with large, scattering to closely
imbricate, ovate, scale-like leaves below, above bearing a closely tufted group
of branches; branches ascending, 1.5-2.5 cm long, terete, acute to obtuse;
branch-leaves 1.7-2.2 mm long, about two-fifths as wide, broadly lanceolate,
sharply serrate in upper half, often denticulate below, acute, broadly auriculate,
erect-spreading, when dry imbricate, strong bi-sulcate; costa strong, ending just
below apex; median leaf-cells oblong-hexagonal with more or less rounded or
truncate ends, 5-7:1, rather incrassate, those of the auricles quadrate along the
margin to diamond-shaped towards the interior, the apical and the upper
marginal larger and rhombic-oblong: seta erect, stout, dextrorse above, sinis-
trorse below, castaneous, about 1-1.6 cm long; capsule castaneous, narrowly
cylindric, about 5-6 mm long, 5-6:1, erect to slightly curved, slightly con-
tracted below the mouth when dry, nearly smooth; annulus none; operculum
conic-rostrate, about 1 mm long; peristome-teeth shallowly inserted, orange-
castaneous, non-striate but decidedly papillose, strongly and rather densely
trabeculate, the lamellae and divisural not very distinct, the teeth slender and
often perforate below; segments usually longer than teeth, linear, arising from
a very narrow and often somewhat perforate basal membrane which is usually
inserted entirely below the rim of the urn, the segments yellowish, granular-
papillose, perforate-cleft in a ladder-like manner along the median line; cilia
none, or sometimes represented by mere stubs rising from the basal membrane;
exothecial cells heavily incrassate, castaneous-pellucid, oblong, the upper
rounded-quadrate, those at the rim smaller and transversely oblong, darkly
182 American Midland Naturalist Monograph No. 6
incrassate; spores yellowish, minutely roughened, about .016-.018 mm, the
walls moderately incrassate.
In damp, shady woods on rotten logs, stumps, wet soil, rocks, etc., often
in swamps. From New Brunswick to the Carolinas and Alabama and west to
the Rocky Mountain region.
Not uncommon in our region but rather rarely found in fruit. Known from Allegheny,
Aimstrong, Bedford, Butler, Cameron, Clarion, Crawford, Elk, Erie, Fayette, Forest,
McKean (Porter), Somerset, Tioga, Venango, Warren, Washington, and Westmoreland
counties. Specimen figured: Moon Township, Allegheny Co., 1889. J.A.S.
3. Climacium Kindbergii (Renauld and Cardot) Grout
(C. americanum var. Kindbergii Renauld and Cardot)
Plate XXXV
Dark yellowish-green to almost black, usually rather densely cespitose:
secondary stems about 3-6 cm tall, sometimes indistinctly dendroidal, stout,
castaneous, bearing along the stem rather scattered widely ovate leaves about
3-4 mm long, leaves not sulcate, plane-margined, sub-clasping at base, acute
and almost entire at apex, strongly costate into the apex; basal cells in a wide
area, rather thin-walled, large, rectangular to rhombic-oblong, often somewhat
brownish, above quickly passing into linear prosenchymatous cells about 10-
15:1, the apical cells shorter and rhombic oblong; branches densely tufted,
ascending to widely spreading, 1.5-2.5 cm long; branch-leaves about 1.5-2.5 mm
long, broadly lance-ovate, somewhat clasping by the auriculate base with
rounded auricles, strongly sulcate, apex obtuse to acute, margin plane, serrate
in upper half, strongly costate almost to the apex; median leaf-cells of the
branch-leaves oblong-hexagonal, 2-3:1, somewhat incrassate, the basal short
rhombic to quadrate-rectangular in the auricles, the median basal longer with
rounded ends, incrassate and more or less castaneous-pellucid : sporogonium not
seen from our region but described as having the seta more flexuous and con-
siderably longer than in C. americanum; capsule 4-6 mm in length; peristome-
teeth perforate.
In swamps and pools in woods from New England to Indiana and the
Gulf States. According to Grout, less common inland.
Bedford Co.: Raystown Branch, lYi mi. s. of Schellburg. C.M.B. July 19, 1941.
Butler Co.: Outlet of West Liberty Bog. C.M.B. Nov. 26, 1948. Crawford Co.:
Twin Lakes, Pymatuning Swamp. C.M.B. July 5, 1947. Erie Co.: Among shrubs in
swamp, Presque Isle. Nelle Ammons. Aug. 3, 1935. Fayette Co.: Along margin of
densely shaded mountain stream. Meadow Run Valley, four miles south of Ohio Pyle,
September 1-3, 1906. O.E.J, and G.K.J, (figured); Near Falls, Ohio Pyle, Oct. 12,
1935. C. M. Boardman. Mercer Co.: One mile north of North Liberty, on tree root
in wet woods. May 30, 1935. C.M.B.; In swamp, North Liberty, April 14, 1935. Sid-
ney K. Eastwood.
Family 26. Leucodontaceae
Dioicous, rarely autoicous: antheridial shoots gemmiform, axillary; arche-
gonial clusters terminal on short perichaetial branches; both kinds on secondary
Jennings: Manual of Mosses — 26. Leucodontaceae 183
shoots: paraphyses few, filiform: plants more or less stiff and robust, laxly
cespitose, mostly shining; stem cylindric, central axis rudimentary or none;
main stem creeping, branched, radiculose with brownish radicles; secondary
stems numerous, erect or ascending, rarely pendent, thickly-leaved, simple or
branched; leaves pluri-seriate, decurrent, often plicate, ovate to lanceolate,
abruptly to slenderly acute, non-bordered, one layered; costa double or simple
or none; leaf-cells incrassate, mostly smooth, rhombic above, below elongate
along the middle of the leaf, towards the margin rounded-quadrate in many
series; capsule erect, symmetric, oval or ovate to oblong-cylindric; annulus pres-
ent; peristome double, teeth lanceolate to lance-subulate, densely articulate,
non-bordered, mostly papillose, mostly without projecting lamellae, rarely with
cross-striae on basal plates; basal membrane of inner peristome low, segments
rudimentary and narrow or none, as long or shorter than teeth, cilia none; lid
conic, obliquely rostrate; calyptra cucullate; spores medium to large.
On rocks and trees, mainly confined to temperate regions; 7 genera; only 2
genera in our region.
Key to the Genera
A.Costa double or none 1. Leucodon
A. Costa single 2. Leptodon
1. Leucodon Schwaegrichen
Dioicous: blackish to yellowish or brownish-green, dull or lustrous: pri-
mary stems very long and branched; secondary stems usually simple, equally
high, sometimes more or less pinnate, thickly leafy; leaves drying appressed,
straight or secund, mostly pluri-plicate, when moist spreading, ovate-acuminate
to short-acute, entire or apically serrate, ecostate; median leaf-cells smooth, ob-
long-rhombic, the basal reddish-yellow; inner perichaetial leaves high-sheathing,
long-acuminate: seta mostly more or less elongate, reddish; capsule mostly
exserted, oblong to oval (or globose), reddish brown to blackish, with a small
mouth and short collum, stomata none; annulus present; peristome double with
the inner peristome rudimentary or apparently lacking; teeth whitish to yel-
lowish, mostly gaping in the middle or divaricately cleft; lid conic, constricted
at the base, sometimes obliquely rostrate; calyptra smooth, cucullate, envelop-
ing the capsule and upper end of seta; spores .025-. 03 5 mm, yellowish-green,
finely warty.
A widely distributed genus of about 30 species, occurring on trees and
rocks; 8 species reported for North America; 3 species, probably, in our region.
Key to the Species
A. Leaves plicate; secondary stems well developed: seta about 2-3.5 mm long, with cap-
sule emergent but shorter than the perichaetial leaves 1. L. brachypus
A. Secondary stems less developed: capsule long-exserted B
B. Leaves ovate-elliptic, rather abruptly and shortly acuminate, scarcely plicate
2. L. julaceui
B. Leaves lance-ovate, long and slenderly acuminate, much plicate 3. L. scitiroidcs
184 American Midland Naturalist Monograph No. 6
1. Leucodon brachypus Bridel
Plate XXXV
Moderately robust, brownish to light green, loosely tutted: stems usually
at least 5-6 cm long, with rather numerous secondary simple or branched divi-
sions; leaves about 2 mm long, ovate, bluntly acute to short-acuminate, ob-
scurely more or less secund, usually plicate with two folds, entire to serrulate
above; costa none; median leaf-cells linear-fusiform and castaneous pellucid at
base, the interior median rhombic, about 5-8.1, grading to oval at the apex,
the marginal basal rounded-quadrate to transversely oblong, all cells incras-
cate; penchaetial leaves loosely appressed-sheathing, non-plicate, the inner
surpassing the capsule; seta about 2-4 mm long, wrapped in the perichaetial
leaves; capsule oval-oblong, about 1.2-2 mm long, about 2:1 castaneous, small-
mouthed, dark-rimmed; lid conic, obliquely short-rostrate; peristome-teeth
rather broad, irregular, pale to whitish, papillose, often bifid at apex, the inner
peristome very thin, narrow, and without segments or cilia; spores mature in
winter or late fall, pale, rather thin-walled, granular.
On trees and rocks in hilly or mountainous regions; from Nova Scotia to
Kansas and south to the Gulf States.
Cambria Co.: Cresson. T. C. Porter and T. P. James. (Porter's Catalogue). Craw-
ford Co.: On bark at base of Fraxinus nigra, near Linesville, June 11-12, 1907. O.E.J.
McKean Co.: Quintuple, November 11, 1893, (approaching L. sciuroides in arumina-
tion of leaf-apex) and Langmade, near Bradford, August 11, 1895. D.A.B. (figured).
Washington Co.: Linn and Simonton. (Porter's Catalogue).
2. Leucodon julaceus [Linnaeus] Sullivant
(Hypnum julaceum Linnaeus; Pterigynandrum julaceum Hedwig)
Plate XXXVI
Resembling the preceding in habit but with shorter secondary stems and
distinctly terete branches, which are julaceous when dry: leaves crowded, closely
appressed-imbricate when dry, scarcely secund, ovate-elliptic, abruptly short-
acuminate, entire or slightly serrulate at apex, the margins often recurved,
blade concave, scarcely plicate, the base rounded and sub-clasping, mammillose
at back of apex; leaf-cells mainly as described for the genus, but the upper
much shorter and broader than in the other species, in the median upper third
rhombic-oblong, incrassate^ about 2-3:1, seriate; the marginal rounded-hexag-
onal but towards the base usually densely transversely oblong-hexagonal, the
basal median linear-vermicular and much incrassate, those above becoming
shorter; costa none; perichaetial leaves linear-oblong, filiform-acuminate, reach-
ing well up to the capsule: seta slender, partly exserted; capsule turgid-oval,
castaneous, about 0.5-0.7 x 1 mm; annulus none; lid obliquely short-rostrate,
about half as long as the urn; peristome closely similar to that of L. brachypus,
the teeth apically bifid; spores mature in fall.
In woods on tree-trunks, often mixed with other mosses, from New Eng-
land to Minnesota and south to Florida and Texas.
As yet known only from the southern third of western Pennsylvania. Ai LEGHENY
Co.: On base of white oak tree. Library, Apnl 29, 1909. O.E.J, (figured). Bedford
Jennings: Manual of Mosses — 26. Leucodontaceae 185
Co.: Roystown Branch, lYi mi. s. of Schellsburg. C.M.B. July 19, 1941. Cambria
Co.: T. P. James. (Porter's Catalogue). Indiana Co.: T. P. James. (Porter's Cata-
logue). Washington Co.: On bark of log, May 8, 1891, and on rotten wood, Nov.
5, 1892. Near Washington. Linn & Simonton.
3. Leucodon SCIUROIDES [Linnaeus] Schwaegrichen
(Hypmim sciuroides Linnaeus; Fisstdens schnoides Hedwig)
Rigidly cespitose, brownish to olive-green: secondary stems terete and jula-
ceous, more or less curved-ascending at the ends, usually 3 or 4 cm long; leaves
densely crowded, slightly secund, closely imbricate when dry, more or less
open-spreading when moist, lance-ovate, long and slenderly acuminate, entire,
usually about 5-plicate, somewhat decurrent; costa none; leaf-cells about as for
L. brachypus; perichaetial leaves pale, non-plicate: seta about 7 or 8 mm long,
rather stout; capsule oblong-elliptic, brown, exserted; lid conic, same color as
urn; peristome-teeth slender, pale to whitish, remotely articulate, entire or split
towards the base; annulus simple, falling away in fragments; calyptra yellowish-
brown apical ly, reaching to the base of capsule; spores mature in spring but
capsules very rarely found.
On trunks of trees, or very rarely on rocks, in woods; Europe, and from
lower eastern Canada through the northeastern United States to Pennsylvania
and Iowa. Not yet found in our region.
2. Leptodon Mohr
(Forsstroemia Lindberg)
Autoicous, rarely dioicous; quite robust to slender, green to brownish-
green, mostly dull: leaves drying imbricate and non-plicate or indistinctly pli-
cate, when moist erect-spreading, ovate to oblong, short acute, also ovate and
acuminate, margin more or less revolute, entire or ape.x serrate; costa rather
narrow, ending about the middle; apical and median cells elliptic or oval, the
angular rounded quadrate to transversely oblong; inner perichaetial leaves
sheathing, long and narrowly pointed, costate or ecostate: seta short, 2-5 mm,
straight, red to yellowish; capsule mostly exserted, ovate to oval, pale or red-
dish-brown; annulus narrow or none; peristome-teeth lance-linear, mostly yel-
lowish, pellucid, densely articulate, finely papillose above, sometimes broken
through on the divisural; inner peristome none or very rudimentary; spores
.020-. 035 mm, yellowish-green, finely papillose; lid conic, narrowly acuminate
to shortly rostrate; calyptra cucullate with erect hairs, rarely smooth.
A widely distributed genus of about 20 species, mostly arboreal in habitat;
4 species in North America; 1 species in our region.
1. Leptodon trichomitrion (Hedwig) Mohr
(Pterigynandnim trichomitrium Hedwig; Forsstroemia
trichomitrid (Hedwig) Lindberg)
Plate XXXVI
Broadly cespitose, rather rigid, yellowish-green; primary stems creeping,
filiform, the secondary stems numerous and abundantly branched; leaves close,
loosely erect-spreading, lance-ovate, shortly acuminate to acute, entire, when
186 American Midland Naturalist Monograph No. 6
dry somewhat plicate, about 1.5-2 mm long, the extreme apex rather blunt, the
base concave, the margins reflexed; perichaetial leaves loose in texture, the inner
sheathing, reaching to the base of the capsule or a little higher: seta short,
slightly longer than the capsule; capsule ovate-cylindric, thin-walled, rather
gradually narrowed below, about 3-4:1, about 1.5 mm long; exothecial cells
rather incrassate, irregularly polygonal to rectangular-oblong, several rows at
the narrowed mouth smaller, rounded-quadrate and dark-castaneous; peristome-
teeth whitish, lance-linear, rather remotely articulate, sometimes perforate along
the divisural, the inner peristome entire to more or less torn, adhering to the
ventral surface of the teeth; lid short-rostrate; spores mature in winter, orange-
incrassate, almost smooth, about .023-. 025 mm.
In woods on trees, rarely on rocks; Asia, and from New England to On-
tario and the Gulf States.
Common in eastern Pennsylvania but rare in our region. Beaver Co.: About 8 feet
up on elm trunk, along Raccoon Creek, one mile south of Traverse Creek. April 1, 1934.
Chas. M. Boardman. McKean Co.: Near Latshaw, N. Y. north of Bradford, August
25, 1895. D.A.B. (figured).
Family 27. Neckeraceae
Dioicous, rarely autoicous or synoicous; sexual clusters only on secondary
shoots and their branches, with filiform, often yellowish paraphyses: slender
to robust, mostly stiff, laxly cespitose: stem somewhat dorsiventrally flattened,
with oi without a rudimentary central strand: primary stem more or less creep-
ing, mostly filiform, mostly sparsely fasciculately radiculose; secondary stems
more or less elongate and ascending or much elongated and pendent, mostly
distantly or symmetrically pinnate, thickly-leaved, julaceous or flattened; leaves
nearly always pluri-seriate, uni-stratose, of various forms; costa mostly delicate,
homogenous, simple or double or none; median cells mostly prosenchymatous,
the apical sometimes parenchymatous, the basal often colored, the alar some-
times differentiated: capsule mostly erect and symmetric, peristome mostly
double, teeth yellowish to brownish, lance-linear, dorsally sometimes abnor-
mally thickened, ventrally trabeculate; the inner peristome with mostly low
carinate basal membrane, rarely rudimentary or none, segments linear to fili-
form, often fenestrate, rarely cleft the whole length, cilia mostly none; lid
conic, erectly to obliquely rostrate; calyptra mitrate to cucullate, mostly hirsute;
spores of varying size.
A large family, occurring mainly on trees in warmer regions, often forming
a conspicuous part of the vegetation; about 20 genera, of which but three
occur in our region.
Key to the Genera
A. Secondary stems flattened, ascending or pendent; leaves mostly conspicuously unsym-
metric B
A. Secondary stems mostly erect and branched in a tree-lil;e manner; leaves not com-
planately disposed 3. Porotrichum
B. Exannulate; basal membrane of inner peristome low, cilia none, segments narrowly
linear 1. Neckera
B. Annulus 2-seriate; basal membrane conspicuous, cilia rudimentary and soon disap-
pearing or well-developed, segments about ss broad as teeth 2. Homalia
Jennings: Manual of Mosses — 27. Neckeraceae 187
1. Neckera Hedwig
Autoicous or dioicous, rarely synoicous: mostly more or less robust, ces-
pitose, green to yellowish or brownish, somewhat lustrous: primary stems often
stoloniferous, paraphyllia mostly none; leaves on the filiform shoots small,
ecostate, sym.metric, concave; normal leaves either 1 -seriate, the dorsal and
ventral alternately turned to the side, the lateral spreading, or 4-seriate, the
dorsal and ventral series lacking, leaves often transversely undulate, unsym-
metric, rugose, more or less spatulose from a broader and shortly decurrent
base, acute to obtuse or truncate; costa various, upper leaf-cells rounded to
rhombic, the lower linear, the alar differentiated, small and quadrate; peri-
chaetial leaves high-sheathing, narrow, long-acuminate: capsule oval or elliptic,
immersed to exserted; annulus none; peristome double, inserted far back;
peristome-teeth lance-linear, often basally striate, low-trabeculate, sometimes
split along the divisural; basal segments 16, the basal membrane mostly very
low; cilia none; calyptra mostly cucullate and with erect hairs; spores medium,
mostly brownish, papillose.
A widely distributed genus of about 130 species; about 20 species reported
for North America; two species definitely known for our region.
Key to the Species
A. Leaves rounded and usually apiculate at the apex ....B
A. Leaves strongly undulate, acute to acun-.inatc; perichaeti'jm longer than seta and cap-
sule I. N. pennata
B. Leaves undulate L (N. pennata var. oligocarpa)
B. Leaves not undulate C
C. Plants rather robust; leaves complanate; seta 7-10 mm long, exceeding the peri-
chaetium 2. N. complanata
C. Plants very slender; leaves not complanate 3. N: gracilis
1. Neckera pennata [Linnaeus] Hedwig
(Fontinalis pennata Linnaeus)
Large with primary stems creeping, often stoloniferous, the secondary stems
6 to 8 or 10 cm long, erect, pinnate or nearly simple; leaves lance-ovate, acute
to acuminate, more or less undulate above, the margins entire or slightly den-
ticulate; costa short and faint, more or less bi-striate and wrinkled; median
leaf-cells linear at base, towards the apex the upper marginal and apical
broadly rhomboid; inner perichaetial leaves entire, half-sheathing, elongate-
lanceolate, reaching somewhat beyond the capsule; seta very short; capsule
immersed, yellowish, oblong-oval, brown when old, about 2.5:1; lid acute-conic
or acuminate; calyptra very small and covering only the operculum; peristome
double, teeth irregularly divided, subulate-linear from a lance-linear base, some-
times apically coherent, the segments rudimentary and very short; spores in
summer.
On trees or on moist rocks in cool, moist woods, usually on the trunks of
deciduous trees; widely distributed in temperate regions, in North America
extending from lower Canada south to North Carolina. Probably rather
common in the eastern part of our region.
Cambria Co.: Crefson. T. C. Porter. (Porter's Catalogue). McKean Co.: D. A.
Burnett. (Porter's Catalogue) .
188 American Midland Naturalist Monograph No. 6
2. Neckera complanata [Linnaeus] Huebener
(Hypnum complanatum Linnaeus; Homalia complanata DeNotaris)
Yellowish to pale green, in rather large and dense tufts, soft: stems long,
often reaching 8 or 10 cm, branchlets pinnately arranged; complanate, some-
times more or less flagelliform; leaves oblong-lingulate, compressed, complanate,
usually rounded at the apex and short-apiculate, sometimes acute or acuminate,
those at the tips of the branches often more or less deflected and falcate, the
margin usually inflexed at base on one side, serrulate at apex; costa double,
very short and faint, or none; median leaf-cells linear-vermicular, the apical
shorter and wider, rhomboidal, the angular quadrate-oval and yellowish-pel-
lucid; perichaetia borne along the sides of the stem, the leaves long-sheathing:
seta yellow, about 1 cm long; capsule oval to elliptic-oblong, pale, orange-
yellow or castaneous, about 2:1, small-mouthed; lid subulate-rostrate, usually
oblique; calyptra cucullate, reaching to about the middle of the urn, one-half
length of urn; peristome-teeth long, pale, narrow, the segments about half as
long, filiform from an enlarged base; spores mature in spring but capsules
rarely produced.
On bark of trees, rarely on rocks; Europe, Asia, northern Africa, and from
Labrador to Tennessee.
Rare in our region. Reported from "Allegheny Mountains in Pennsylvania" in
Lesquereax and James' Manual.
3. Neckera gracilis (James) Kindberg
{Homalia gracilis James)
Minute, slender, irregularly branched plants with stems and branches
mostly flagelliform at the ends, complanately foliate, and together with the
leaves not much more than 1 mm wide; leaves oblong-lingulate, about 0.1 mm
long, not undulate, rounded-obtuse or apiculate, somewhat serrulate at the
apex; costa very short, often double; upper leaf-cells rhomboidal, about 8-12 jx
wide and but little longer, the basal longer in the median and smaller in the
marginal portion.
"On locks, usually in elevated regions, New Jersey, New York and
Vermont. "^ — Grout.
2. Homalia (Bridel) Bryologia Europaea
Dioicous or autoicous: slender to robust, in wide, more or less lustrous,
dark colored, matted tufts: primary stems with stolons; secondary stems mostly
irregularly dichotomous, non-flagellate; leaves 4-seriate, complanately spread-
ing, not transversely undulate, unsymmetric, spatulate to Ungulate from a
slightly decurrent base, rarely rounded, obtuse, non-bordered, with apex entire
or serrulate; costa simple, incomplete or none; upper leaf-cells rounded to
hexagonal, lower elongate, at least the median so, rarely all linear; inner
perichaetial leaves, short-sheathing, lanceolate, acute; seta long, mostly smooth;
capsule mostly erect to cernuous, oblong from a narrowed base, when old
sometimes arcuate, red-brown, rarely almost pendent and short-oval; annulus
Jennings: Manual of Mosses — 27. Neckeraceae 189
2-senate; peristome double, inserted at the mouth; teeth linear-subulate from
a broader base, yellow to brownish, apically hyaline, mostly transversely striate
and with well-developed lamellae; inner peristome yellow, papillose, marked
with fine transverse lines on outer surface of basal plates; basal membrane
high, carinate, segments longer and almost as broad as the teeth, broken
through in places along the keel, cilia mostly rudimentary and fugaceous,
sometimes well developed and appendiculate: lid conic, obliquely rostrate;
calyptra cucullate, mostly glabrous; spores small, brownish.
About 20 species on trees, rocks, and stones, mostly in temperate regions;
4 species in North America; one species in our region.
1. HoMALiA Jamesii Schimper
In straggling tufts, shining yellow-green, repeatedly distichous, stolonifer-
ous: stems slender, interruptedly foliate by the numerous innovations; the
branches strongly complanate-foliate; leaves cultriform, sub-falcate, oblong,
obtusely apiculate, minutely serrulate above the middle, striolate lengthwise
when dry; costa faint, slender, reaching half-way or more; lower median leaf-
cells linear-fusiform, the apical and marginal about 1.5-1:1, about as broad as
long, rhomboidal: seta about 1.5 cm long, slender; capsule erect to cernuous,
oblong-cylindric, about 2.5:1, symmetric, when dry scarcely constricted below
the mouth; peristome double, teeth long, yellowish, confluent at base; inner
peristome about as long as the teeth, the segments narrow, sub-linear, more or
less carinately perforate, cilia rudimentary and solitary or none; annulus pres-
ent; spores mature in fall but capsules rarely found.
On rocks and in crevices, in mountainous or hilly districts; from New-
foundland and Nova Scotia to Pennsylvania, also in Washington State. Pos-
sibly will be found to occur in the eastern part of our region.
3. PoROTRiCHUM Bridel
(Thamnntm Bryologia Europaea )
Dioicous or, rarely, autoicous; mostly robust to very robust, with a long,
creeping primary stem: the primary stem has scale-'ike leaves, and is more
or less densely brown-radiculose; the secondary stem erect to ascending, without
branches below, stoloniferous, somewhat dendroid in habit; branches spreading,
flattened, obtuse; leaves erect-spreading to spreading, plane to concave, smooth
to plicate, not rugose, non-decurrent, unsymmetric, mostly oblong to ovate or
ovate-lingulate, the apex obtuse to acute, serrate; costa strong, mostly incom-
plete; median leaf-cells parenchymatous, the basal sometimes linear; inner
perichaetial leaves lanceolate-acuminate and spreading from a half-sheathing
base: seta various, mostly 10-15 mm long, in certain species not more than 4
mm, in others up to 4 cm. long, red, smooth; capsule inclined to horizontal,
arcuate, gibbous, rarely erect, symmetric, and oval; annulus revolute; peristome-
teeth lanceolate to linear, subulate-acuminate, yellowish, bordered, with a
zig-zag divisural; inner peristome pale yellow, basal membrane prominent,
segments broad, carinately split and gaping; cilia often appendiculate; spores
small; lid conic, rostrate; calyptra cucullate, glabrous.
190 American Midland Naturalist Monograph No. 6
About 30 species in temperate and warm regions; 7 species reported for
North America; one species in our region.
1. PoROTRiciiUM allegheniense (Mueller) Grout
(Hypnum allegheniense C. Mueller; Thamnium allegheniense Bryologia Europaea )
Plate XXXVI
Large, dendroidal in habit, bright to pale green, usually rising to a height
of 4-7 cm; leaves of the branches and branchlets up to 3 or 3.5 mm long,
rather lustrous and sub-plicate when dry, erect-spreading, oblong-elliptic, short-
pointed, concave, the base somewhat narrowed but scarcely concave, the apex
broadly acute, the plane margin strongly serrate above; costa strong, extending
to near the apex; leaf-cells incrassate, the median shortly rounded- or rhom-
boid-hexagonal, about 2:1, the basal becoming elongate-oblong, varying to
elongate-rectangular, the lower marginal and angular, scarcely wider but sub-
rectangular to quadrate; perichaetial leaves erect, sheathing, narrowly acumin-
ate: seta lustrous, of a rich castaneous color, usually about 1 cm long, smooth,
arcuate; rapsule oblong-cvlindric, castaneous and rarely somewhat wrinkled
when dry, about 2-2.5:1, about 2 to 2.5 mm long, nearly symmetric but by
the curving of the pedicel inclined or horizontal, sometimes curved; lid conic,
long- and stout-rostrate, the whole lid being about one-half to one-third as
long as the urn; peristome normally hypnoid, large; teeth lance-subulate, dis-
tinctly but finely cross-striate in at least the lower half, hyaline and papillose
above, castaneous-pellucid below, the dorsal lamellae and the divisural dis-
tinct, the trabeculae well developed; segments papillose, pale yellowish, about
as long as teeth, cleft carinately between the articulations; basal membrane
one-third as high as teeth; cilia 2-3, sub-appendiculate, almost as long as seg-
ments; annulus narrow, revoluble, simple; spores mature in late fall or early
winter, smooth, castaneous-pel'ucid, medium-walled (about .009 mm Grout) .
On dripping rocks and ledges along streams in the hills or mountains from
Nova Scotia to Minnesota and south to the Gulf States.
Cambria Co.: Cresson. T. P. James. (Porter's Catalogue). Huntingdon Co.r
T. C. Porter. (Porter's Catalogue). McKean Co.: On stones in or at rhe edge of
streams, Hedge-hog Hollow, March 18, 1894, Bennett Brook, April 9, 1893 (figured),
and Limestone Creek, N. Y., all near Bradford.
Family 28. Entodontaceae
Autoicous or diocious: slender to quite robust, mostly stiff, laxly cespitose,
mostly lustrous; central strand none or but few-celled; stem thickly-foliate,
julaceous or complanate; leaves pluri-seriate, uni-stratose, often unsymmetric;
costa delicate, homogeneous, never complete, or double and very short, or
none; leaf-cells mostly prosenchymatous, the alar differentiated, being quadrate
or transversely widened: capsule exserted, mostly erect and symmetric, never
plicate; peristome mostly double, the inner rarely lacking; teeth yellow to
castaneous, with divisural, trabeculate, mostly papillose; segments narrow or
lance-subulate, often split carinately, the basal membrane low, carinate, the
Jennings: Manual of Mosses — 28. Entodontaceae 191
cilia rudimentary or none; spores mostly small; lid conic, short- to long-ros-
trate; calvptra cucullate, glabrous.
Mostly in warmer and temperate regions, on trees, sometimes on rocks or
on soil: 14 genera, one genus definitely known from our region.
Key to the Genera
A. Leaf-cells smooth 1 . Entodon
A. Leaf-cells more or less strongly papillose (Pterigynandrum)
1. Entodon C. Mueller
Autoicous, rarely dioicous: green to golden-brown: stem prostrate to
ascending, complanate-leaved, rarely julaceous, thickly pinnately branched,
mostly short, simple, ascending or spreading; stem-leaves compressed, slightly
decurrent, concave, the dorsal and ventral imbricate, the lateral spreading,
oval, from an ovate base obtuse or apiculate or rarely slenderly acuminate,
entire or apically serrate; costa double and very short, or none; median leaf-
cells narrowly linear, smooth, and basal lax and incrassate, the alar laxly
quadrate and sometimes slightly inflated, forming a distinct hyaline group:
seta mostly 1-3 cm long, red or yellow, twisted when dry; capsule erect,
straight or weakly curved; collum short; annulate or exannulate; teeth inserted
below the mouth, lance-linear, acuminate, thin, plane, mostly non-margined,
orange to castaneous, articulate, mostly low-trabeculate; inner peristome with-
out prominent basal membrane, segments linear, carinate, yellow, as long as or
shorter than the teeth, cilia none; spores .012-. 020 mm.
Nearly 150 species, on trees and on calcareous rocks, in temperate and
warmer regions; about 33 species occurring in North America; 3 species in
our region.
Key to the Species
A. Leaves narrowly gradually accminite; segments adhering to teeth (£. brevisetus)
A. Leaves acute, abruptly acuminate-apiculate, or almost obtute; segments free B
B. Teeth with noi less than twenty articulations; leaves acute to almost obtuse; but
not apiculate. complanate L E. compressui
h. Teeth with less than twenty articulations C
C. Leaves acuminat?-apiculate: teeth 15-20 articulate: capsule less than 4.5:1; leaves
complanate 2. E. cladorhizam
C. Leaves abruptly apiculate: teeth less than 10-articulate; capsule about 5:1: leaves
not complanate 3. E. seductriy
1. Entodon compressus C. Mueller
(Cylindrotheciiim compressum Bryologia Europaea)
Plate LXXI
Widely and flatly cespitose, glossy yellow-green, with compressed stems
and branches: considerably more slender than E. cladorh'.zans, but quite simi-
lar in general appearance: leaves about 1.0-1.1 x 0.4-0.5 mm, quite concave,
broadly oblong-ovate, obtuse to broadly acute, entire at apex; median leaf-
cells linear, the alar quadrate and numerous reaching almost across the base
cf the leaf, a few on the corner often somewhat inflated; costa none or very
rudimentary: seta erect, 1-1.4 cm. long; capsule erect, ovoid to elliptic, narrow-
192 American Midland Naturalist Monograph No. 6
mouthed, about 2.5 x 0.6 mm; lid rather long and with a slender, curved
rostrum; annulus large, rather persistent; peristome-teeth long, lance-linear,
closely articulate; the segments somewhat shorter, linear-subulate, free from
the teeth, these latter densely papillose; spores mature in fall or early winter.
In shaded situations on soil and decaying logs and on bases and roots of
trees in moist situations, often near water-courses: northern Eurasia and from
New Jersey to Missouri and northwards to Rhode Island and Nebraska.
Rare in our region. Beaver Co.: On base of elm tree along Raccoon Creek one mile
south of Traverse Creek. April 1, 1934. C.M.B. (figured). Washington Co.: On
root in damp spot. Taylorstown. Linn and Simonton. Apr. 29, 1899.
2. Entodon cladorhizans Schimper
{Cylindrothecium cladorhizans Schimper; Neckera cladorhizans Hedwig)
Plate XXXVI
Cespitose in wide tufts, brightly lustrous, yellowish-green: stems com-
pressed, somewhat pinnately branched, rather acuminate and sometimes up-
curved at the apex; branches complanate and spreading widely from the stem,
more or less acuminate to attenuate at the apex, where sometimes rooting;
leaves loosely imbricate, very concave, non-plicate, narrowed at the apex,
margin plane or narrowly revolute below, apex sub-acute, faintly serrulate,
usually turned slightly backwards; leaves ovate to oblong, about 1-2 mm long
by one-half as wide; costa double, short and indistinct, or none; median leaf-
cells long-linear, prosenchymatous, smooth, with firm and hyaline walls, the
alar hyaline to somewhat reddish, incrassate, quadrate-rectangular in a tri-
angular patch of 6-8 rows in depth, bordered by a few intermediate, sub-quad-
rate to sub-vermicular cells, the apical cells shorter and rhombic: seta erect,
smooth, sinistrorse, rich castaneous in color, lustrous, about 8-12 mm long;
capsule about 4 6:1, oblong-cylindric, tapering abruptly to the seta, smooth,
not sulcate when dry, castaneous, narrowed somewhat at the mouth, 2.5-3.5
mm long; annulus early deciduous, large, pluri-seriate with incrassate quadrate
cells; exothecial cells yellowish with thin walls, rectangular to oblong, towards
the rim suddenly much smaller and incrassate, more or less laterally oblong
under the annulus; operculum conic-rostrate, about 0.4-0.6 mm, long, often
apiculate; peristome double, deeply inserted, teeth light-castaneous, about 16-
20-articulate, below lightly papillose-striate in variously divergent or radiating
lines, not finely transversely striate as in most hypnaceous peristomes, some-
times perforate, (lacunose) above; segments distinct, linear, very narrow,
carinate, hyaline, very slightly granulose-roughened, entire, nearly as long
as the teeth, arising from a very narrow basal membrane; cilia none; spores
papillose, incrassate, castaneous, about .016-020 mm, mature in late autumn
or early winter.
On leaf -mould, rotting logs, bases of trees, etc.; Europe, and, in America,
from New Bruswick to Ontario and south to the Gulf States.
Common in our region. Known from Allegheny, Armstrong, Beaver, Bedford, Butler,
Crawford, Fayette, Indiana, Lawrence, McKean, Somerset, Washington, and Westmore-
land counties, nearly all in the southwestern corner of the State. Specimen figured: Wild-
wood Hollow, Allegheny Co., March 29, 1909. O.E.J. SC G.K.J.
Jennings: Manual of Mosses— 29. Leskeaceae 193
3. Entodon seductrix (Hedwig) C. Mueller
(Neckera seJuclrtx Hedwig; Cylindrothecium seductrix SuUivant )
Plate XXXVII
Robust, widely cespitose in glossy yellowish-green mats: branches sub-
pinnately arranged, slender, julaceous, up to 2 or 2.5 cm long; leaves about
0.8-1.4 mm long, broadly oblong-elliptic to ovate, imbricate, deeply concave,
short-apiculate with the apiculation often reflexed, margin plane, entire, or
sometimes slightly serrulate, at base often slightly reflexed, base of leaf slightly
narrowed; costa short and double; median leaf-cells linear-to oblong-prosen-
chymatous, alar cells quadrate, slightly incrassate, forming a distinct group
sometimes extending along the m.argin for one-fourth the length of the leaf;
perichaetial leaves with a slender acumen, narrower and reaching a length of
about 3 mm: seta glossy, red-castaneous, erect, sinistrorse, about 1.5 cm long;
capsule 2-3 mm long, castaneous, about 5-6:1, cylindric, erect, symmetric or
slightly curved; exothecial cells yellowish with medium walls, rectangular to
irregularly oblong, towards the rim smaller, quadrate to laterally oblong in-
crassate, and formmg a rather indefinite annulus of 2 or 3 series; peristome-
teeth few articulate above, deeply inserted, lance-linear, rather short, bordered,
not transversely striolate but irregularly papillose, dorsal lamellae and divisural
strongly marked: segments nearly as long as teeth, narrowly linear-carinate,
free from teeth, arising from a very narrow basal membrane, smooth, cilia
none; operculum conic-rostrate, usually somewhat oblique, about 0.5-0.8 mm
long; calyptra small, enclosing only about half of the capsule; spores yellow-
ish-incrassate, about .014-018 mm in diameter, minutely roughened, mature
in late summer. Variable.
On rotten logs, earth, rocks, roots of trees, etc.; from New England to
Minnesota and south to the Gulf States.
Common in our region. Known from Allegheny. Armstrong. Beaver, Butler, Craw-
ford, Erie, Fayette, Greene, Huntingdon, Indiana (Porter), La;\Tence, McKean, Wash-
ington, and Westmoreland counties. Specimen/ figured: Charleroi. Washington Co., Oct.
13, 1906. O.E.J. & G.K.J.
3a. Entodon seductrix var. minor (Austin) Grout
Differs from the type in size, being only about one-half to two-thirds as
large, usually darker in color: capsule about 3-4:1, about 2 mm long; spores
usually about .010-. 015 mm in diameter.
Allegheny Co.: Bark of decaying log, mixed oak and pine woods, Dutil Church,
Douthett, December 29, 1908. O.E.J.
Family 29. Leskeaceae
Autoicous or dioicous: slender to robust, mostly stiff, cespitose, bright or
dark green, when old brownish, dull or rarely sub-lustrous: stem mostly with-
out central strand, the primary stems mostly creeping, simple, pinnate, or vari-
ously branched, often stoloniform with distant and minute leaves; secondary
stems mostly erect simple, pinnate, or variously branched, both main and
194 American Midland Naturalist Monograph No. 6
secondary stems stoloniferous; paraphyllia mostly present; leaves rarely uni-
form, usually differentiated into basal and foliate leaves, the latter again into
stem-leaves and branch-leaves; basal leaves distant, small, delicate, pale,
smooth, ecostate; foliate leaves pluriseriate, dense, spreading, rarely secund,
drying appressed to imbricate, symmetric, apex sometimes one-sided, mostly
acuminate, mostly concave, often with two short folds at base, unistratose,
mostly papillose; costa mostly simple and strong, rarely short, double, delicate,
or forked; cells richly chlorophyllose, mostly parenchymatous, small, often
oblong to linear in the middle of the base, or up to the middle of the leaf;
branch-leaves usually shorter and narrower than the stem-leaves; perichaetial
leaves delicate, hyaline, much elongate, ecostate or weakly costate: seta
plicate; annulus usually present; peristome double, the teeth mostly basally
straight and long; capsule erect and symmetric to cernuous and arcuate, non-
confluent, prominently articulate and trabeculate or dorsally uniformly papil-
lose, with weak ventral plates, whitish to red or brownish, often quite red at
the insertion; inner peristome carinate, with basal membrane, segments, and,
rarely, with cilia; peristome often degenerate in the species with erect cap-
sules; lid conic or convex-conic and rostrate; calyptra cucullate; spores mostly
small.
As here treated this is a large family of about 30 genera sometimes, as by
Brotherus, separated into three families: Leskeaceae, Theliaceae, and
Thuidiaceae.
Mostly in temperate and tropic regions, mainly on trees and rocks.
Eleven genera occur in our general region.
Key to the Genera
A. Archegonial clusters borne on the branches: primary stems stoloniform with minute
leaves; costa simple; capsule erect, symmetric; segments filiform or rudimentary;
cilia usually none B
A. Archegonial flowers on the stem; stem not stoloniform C
B. Very slender; costa not reaching above the middle of the leaf: peristome-segments
none 3. Haplohymenium
B. More or less robust; costa ending in or just below apex; peristome-segments fili-
form 4. Anomodon
C. Costa short, simple, forked, double, or none, never reaching much above middle of
leaf D
C. Costs simple (except Pseiido-Leskeella) , elongate, ending a little below the af>ex, or
excurrent E
D. Stem creeping, densely simply pinnate, costa short, simple or forked; teeth non-
bordered, non-trabeculate 1. Thelia
D. Stem ascending to erect, irregularly bushy-branched; costa indistinct or none; teeth
bordered, trabeculate 2. Myurella
E. Leaves of stem and branches alike; stem creeping with ascending or erect, short,
blunt branches F
E. Stem and branch-leaves unlike; stem 1-3 -pinnate, often quite fern-like in general
form H
F. Teeth without distinct lamellae; segments filiform 7. Leskeella
I . Teeth distinctly lamellate; segments narrowly linear or none G
G. Teeth with well-developed lamellae; segments narrowly linear 6. Leskea
G. Teeth with distinct but low lamellae; segments none 5. Ltndbergia
H. Cilia 3, smooth; cells of stem-leaves elongate-hexagonal to almost linear; stem and
branch-leaves similar; paraphyllia felted along stem and leaf-base ....11. Helodium
Jennings: Manual of Mosses — 29. Leskeaceae 195
H. Cilia 2-4, nodose to app)endiculate: cells of stem-leaves rounded-angular to long-
hexagonal; I
1. Operculum merely sharply acute: costa of stem-leaves p)ercurrent to excurrent; leaf-
margins indistinctly serrate above; apical cell of branch-leaf with a single terminal
papilla 9. Haplocladium
I. Operculum distinctly rostrate: costa incomplete to excurrent; stem- and branch-
leaves dissimilar; paraphyllia not so felted nor attached to leaf-base J
J. Margin of stem-leaves entire, base not decurrent; cells uniform, rounded-angular,
the median with 2-6 papillae on each side 8. Rauia
J. Margin of stem-leaves entire or toothed above, the base somewhat decurrent; cells
mostly uniform, rounded- to oval- or oblong-hexagonal, the median ranging
from dorsally unipapillose to both sides pluri-papillose 10. Thiiidtum
1. Thelia Sullivant
Dioicous: more or less slender, densely cespitose, yellowish to blue-green,
dull; stem elongate, creeping, more or less brown-radiculose, densely-leaved,
thickly pinnately branched; branches short, julaceous, obtuse, erect to ascend-
ing; paraphyllia various; leaves densely imbricate, either dry or moist, more
or less decurrent, spoon-like, broadly ovate, abruptly subulate-acuminate, the
margins plane, mostly ciliate-serrate to laciniate; costa short, simple (or
forked); cells rhombic, each dorsally with a high and one- to several-pointed
papilla; median basal leaf-cells elongate, the alar in several series almost
quadrate; inner perichaetial leaves larger, delicate, erect, oblong and subulate-
acuminate, with long marginal cilia, costa ending in mid-leaf, areolation
elongate, the upper cells uni-papillose above: seta 5-15 mm long, thin, drying
twisted, smooth and red; capsule symmetric, erect, oblong to cylindric, golden-
brown; annulus none; peristome-teeth basally confluent, narrowly linear-lance-
olate, pale, non-bordered, finely papillose, distantly articulate, non-trabeculate;
inner peristome pale, papillose; basal membrane low, segments very short or
rudimentary, cilia none; lid conic, short-rostrate; calyptra cucullate, smooth;
spores small.
A North American genus of but four species; three species in our region,
more common southwards.
Key to the Species
A. Papillae on dorsal surface of leaf long, curved, with one fxjint 1. T. hirtella
A. Papillae on dorsal leaf-surface lower, each with two or more points B
B. Papillae usually two-pointed; leaves ciliate; mostly on trees 2. T. asprelLi
B. Papillae usually 3 or 4 pointed; leaves non-ciliate; mostly on rocks or sand
3. T. LescuTti
1. Thelia hirtella (Hedwig) Sullivant
(Pterigynandrum hirtellum Hedwig; Hypnum hirlellum C. Mueller)
Plate XXXVIII
Light green to glaucous-green, small, forming thin and loosely adheren'"
mats: primary stems creeping, felted with a reddish-brown tomentum, pinnate
with numerous short, crowded, julaceous secondary stems and branches; leaves
sub-orbicular, deeply concave, abruptly and narrowly acuminate, decurrent at
base, dorsally papillose, the margins plane, spinulose-dentate above, at least
in the upper half, fimbriate-ciliate below with usually upturned cilia; costa
196 American Midland Naturalist Monograph No. 6
slender, reaching about to the leaf-middle; median leaf-cells pellucid, rhom-
boid-elliptic, with long, slender, simple dorsal papillae; apical leaf -cells linear,
the basal larger and looser, the alar quadrate to rectangular, almost smooth,
rather incrassate; perichaetial leaves numerous, the inner lance-oblong, nar-
rowly acuminate, ciliate-fimbriate in the upper part: seta about 1 cm long;
capsule narrowly oblong-cylindric, about 2.5 x 0.5 mm, erect, symmetric, thin-
walled; peristome-teeth linear, distinctly lamellate, the inner basal membrane
truncate and about one-third as high as the teeth, peristome whitish; spores
pale yellow, mature in fall, thin-walled, about .012-.015 mm, smooth.
On trunks and roots of trees and on stumps, in woods; from New England
and Ontario to Kansas and the Gulf States.
Not often collected in our region. Erie Co.: In oak woods, Presque Isle. May 8-9,
1906. O.E.J. McKean Co.: On trees, near the ground. Gates Hollow, Bradford, April
18, 1897. D.A.B. (figured). Washington Co.: On wood, near Washington, 1892. A.
Linn & J. S. Simonton. Westmoreland Co.: T. P. James. (Porter's Catalogue).
2. Thelia asprella Sullivant
(Lesked asprella Bryologia Europaea
Plate XXXVIII
In most respects quite similar to T. hirtella, but more glaucous-green:
densely interwoven into mats up to 1.5 mm thick; leaves bordered nearly all
around by somewhat longer cilia, and the papillae on the dorsal surface of the
leaf more or less branched or stellate: peristome-teeth longer with nodose
articulations; spores mature in early fall.
In the same habitat as the last species and often mixed with it; ranging
from New England to Ontario and Minnesota and south to the Gulf States.
Known from Bedford, Butler, Cameron, Centre, Clarion, Clearfield, Erie, Fayette,
McKean (Porter), Somerset, Washington, and Westmoreland counties. Specimen figured:
In oak woods, Presque Isle. Erie Co., May 8-9, 1906. O.E.J.
3. Thelia Lescurii Sullivant
Closely resembling T. asprella but with the stouter stems fasciculately
branched, whitish or light glaucous-green in color: leaves deltoid-ovate with a
shorter acumen than in T. asprella, not so distinctly ciliate-fimbriate; the pap-
illae usually stellately 3- or 4-lobed; the capsule relatively more slender and
longer and on a longer seta; the teeth shorter and only sub-nodosely articu-
late, the irmer membrane longer and with short segments; spores mature in fall.
On flat rocks, ledges, or on dry, sandy soil; from New England to Mis
souri and the Gulf States. In eastern Pennsylvania and may occur in our
region.
2. Myurella Bryologia Europaea
Dioicous: slender, forming cushions or loose tufts, soft (stiff when dry),
light to bluish-green, dull to sub-lustrous: in thick tufts the stems are upright, in
loose tufts ascending, irregularly bushy-branched with small-leaved stolons,
basally bushy-radiculose; branches obtuse, sometimes apically flagelliform;
paraphyllia none; leaves 5-seriate, more or less appressed-imbricate, round-
JtNNiNGS: Manual of Mosses — 29. Leskeaceae 197
ovate, obtuse to abruptly apiculate to acuminate, spoon-shaped, marginally
plane to involute, serrate to dentate; costa mostly very short and delicate,
simple or forked; sometimes costa none; median leaf-cells small, incrassate,
elliptic, some rhomboid, at base short-rectangular to quadrate, smooth or
papillose by the thickening of the cell-angles, rarely dorsally mamillate; inner
perichaetial leaves red-brown, elongate-lanceolate, long-acuminate, plane-mar-
gined, serrate, ecostate, with linear cells: seta 10-20 mm long, thin, drying
twisted, red, smooth; capsule erect, somewhat inclined when empty, symmetric,
oblong-oval, short-necked, yellow-brown, finally constricted below the mouth;
annulus present; peristome-teeth basally confluent, lance subulate, yellow or
pale, bordered by the broader dorsal layer, lamellae numerous; inner peristome
finely papillose, hyaline to pale yellow, basal membrane yellow, carinate, seg-
ments lance-subulate, same length as teeth, cilia mostly shorter, filiform; lid
brightly colored, conic, acute to obtuse; calyptra fugaceous, small; spores small.
A genus of six species occurring in Europe, Asia, and in North America;
one species m our region.
1. Myurella Careyana Sullivant
(M. gracilis (Weinmann) Lindberg)
Pale glaucous-green, loosely cespitose, interwoven with long radicles below:
stems slender, creeping to ascending, stoloniferous, fasciculately branching; the
branches julaceous; leaves loosely imbricate, open-erect, wide-ovate, narrowly
long-acuminate, spinulose-dentate all around, very shortly costate or ecostate;
leaf-cells large, pellucid, elliptic-rhomboid, dorsally with large papillae as in
Theha asprella: perichaetial leaves sheathing, lanceolate, filiform-acuminate,
dentate: capsules sub-erect, small, inflated, oval-oblong to obovate-oblong; seta
up to 1 cm long; peristome normally hypnoid, with articulate, yellowish, trans-
versely-striate teeth, entire segments and cilia two, somewhat shorter than
the teeth.
Mainly in crevices and hollows in moist, shaded limestone rocks in hilly
or mountainous regions; Europe, Asia, and from Nova Scotia to northwestern
Canada, south to North Carolina and Tennessee.
Rare in our region. Huntingdon Co.: Alexandria. T. C. Porter. (Porter's Cata-
logue) .
3. Haplohymenium Dozy and Molkenboer
Dioicous: slender, stiff, forming mats, dull, dark green to yellowish- or
brownish green : stems filiform, creeping, widely radiculose, here and there in
fascicles, more or less pinnately branched, branches spreading, short, obtuse;
paraphyllia none; lower leaves smaller, somewhat secund, abruptly lance-subu-
late and recurved-circinate from a broadly ovate base; costa very short or none;
upper leaves spreading to squarrose-spreading, imbricate when dry, from a
concave ovate base more or less abruptly lingulate, obtuse to short-acute, non-
plicate, margin plane and entire; costa delicate and reach ng to mid-leaf, or
stronger but not reaching apex; median leaf-cells turgid, thin-walled, rounded-
hexagonal, with mostly several papillae over the lumen, the marginal smaller,
transversely broader, in many rows towards the basal margin transversely
198 American Midland Naturalist Monograph No. 6
rectangular or hexagonal, only in middle of base oblong and pellucid: seta ?--4
mm, thin, drying twisted, reddish or yellowish, smooth; capsule erect, oval,
smooth, brownish, broadly annulate; peristome-teeth basally confluent, lance-
linear, yellowish, distantly articulate, split apart above, the ventral layer
broader, hyaline, non-trabeculate, but with papillae-like irregular processes;
inner peristome smooth, the basal membrane very low, with no segments nor
cilia; lid conic, obliquely short-rostrate; calyptra inflated-cucullate, furnished
with a few long, erect hairs; spores .020-.025 mm, papillose.
About 20 species, mostly living on tree-trunks, rarely on rocks; one species
occurring in North America and reaching our region.
1. Haplohymenium triste (Cesati) Kindberg*
{Leskea tristis Cesati; Anomodon tristis Suilivant)
Plate XXXVIII
Small, very slender, dull dirty-green, loosely, thinly, and intricately ces-
pitose: stems prostrate, sometimes pendent, branching with irregularly or pin-
nately arranged branches; branchlets erect or curved-ascending; leaves about
0.5-0.8 mm long, appressed when dry, more or less squarrose-spreading when
moist, mainly lanceolate from an ovate-base, concave, sub-clasping, crenulate
on the plane margins by the large and protuberant cells, apically acute to
short-apiculate or obtuse, the apex of the leaf very often broken oflF in the
dried specimens; costa slender, ending in the middle of the leaf; median leaf-
cells oblong-rectangular, about .011-.014 mm in diameter, thin-walled, pellucid,
the upper more or less rounded-hexagonal, the lower marginal transversely
oblong-hexagonal, the lower median often radiating from the basal part of
the costa in a characteristic manner: capsule unknown: leaf -cells turgid and
bearing several large papillae on each surface.
On bases of trees and on steep, sunny rocks; Europe, Asia, and, in the
eastern United States. In the Lesquereux and James Manual the habitat is
stated as particularly on the Hornbeam.
Rare in our region. Clearfield Co.: T. P. James. (Porter's Catalogue). McKean
Co.: Gates Hollow, Bradford, July 8, 1895. D.A.B. (figured).
4. Anomodon Hooker and Taylor
Dioicous: more or less robust, stiff, loosely cespitose, bright to blue-green,
dull, later yellowish to brownish, the mats mostly ochraceous inside: stem far-
creeping, stoloniform, small-leaved, radiculose, bearing ascending to erect,
often basal ly-stoloniferous secondary stems; all leafy shoots having rather uni-
form leaves, the branches sometimes flagelliform; foriage-leaves 5-seriate,
dense, rarely secund, when dry mostly imbricated, little different when moist,
Ungulate from a broadly ovate or oblong and little or not at all decurrent
base, or the upper part lanceolate to subulate, margins plane and entire; costa
strong, smooth, mostly ending below the apex; median leaf-cells rounded-
hexagonal, on both sides densely papillose with one- and two-pointed papillae,
rarely unipapillose over the lum.en, only the median basal elongate, smooth,
* Perhaps better included in the genus Anomodon.
Jennings: Manual of Mosses — 29. Leskeaceae 199
rarely rhombic; inner perichaetial leaves elongate, sheathing, above similar to
the foliage-leaves, or pale spreading, lance-subulate, with elongate cells: seta
more or less elongate, drying twisted, smooth, straight; capsule erect, sym-
metric, oblong-cylindric, rarely curved, not narrowed below the mouth; peri-
stome-teeth lance-linear, either pale, papillose, distantly articulate, non-trabe-
culate, or yellowish, striate and weakly trabeculate; inner peristome finely
papillose, with basal membrane low, carinate segments filiform, entire, often
attached apically to the columella, cilia rudimentary or, mostly, none; lid
conic, obtuse, acute, or rostrate; calyptra cucullate, smooth; spores small.
About 20 species confined to the northern Hemisphere; 10 reported for
North America; 5 species in our region.
Key to the Species '
A. Slender and flagelliform branches present: annulus rudimentary or none, teeth striate
4. A. atteniiatus
A. Slender and flagelliform branches none: annulus present or absent .. R
B. Teeth striate: leaves with a hyaime, piliferous-subulate acumination .-..5. A. rostratus
B. Teeth papillose, non-striate: leaf-acumination not piliferous-acuminate C
C. Leaves with rounded, fimbriate-papillose basal auricles, apex apiculate 1. A. Rtigelii
C. Leaves not auricled D
D. Leaves not secund, the upper half of leaf oblong-lingulate: peristome segments
short or none 2. A. minor
D. Leaves more or less secund, lance-lingulate: peristome segments at least '/2 the
length of teeth 3. A. riticulosus
I. Anomodon Rugelii (C. Mueller) Keissler
(A. apiculatus Bryologia Europaea; Leskea apxulata W. P. Schimper;
Hypnum Rugelii C. Mueller)
Plate XXXVIII
Cespitose in tangled mats, glaucous-green, reddish or brownish when old:
stems creeping, divided, the secondary stems and branches straight or ascend-
ing; leaves 1.5-1.8 mm long, more or less obscurely two-ranked, abruptly
oblong-lingulate from an ovate or oblong-ovate and broader base, the base
non-decurrent but with very large and broadly rounded fimbriate papillose
auricles, the apex often apiculate, the upper margin broadly incurved, the
leaves when dry crispate; costa pellucid, ending considerably below the apex;
leaf-cells opaque, chlorophyllose, minute, rounded, papillose on both faces, the
median basal elongate, smooth, the alar somewhat larger, rounded-quadrate;
inner perichaetial leaves long-sheathing: seta erect, about 5-7 mm long, dex-
trorse above, sinistrorse below; capsule erect or somewhat inclined, symmetric,
ovate-cylindric, about 2-2.5 x 0.5 mm thick-walled, castaneous, longitudinally
many-plicate when dry; annulus none; lid conic-acuminate, small; peristome
double, the teeth lance-linear, nodose-articulate, faintly papillose, the dorsal
lamellae and divisural usually very faint or invisible, the segments rudi-
mentary, or very short, from a low basal membrane; spores mature in autumn,
medium-walled, brownish, papillose, about .009-. 012 mm.
On shaded rocks and bases of trees or on decayed logs, in woods, mainly
in mountainous districts; Europe, Asia, and from New England to Ontario
and Georgia. Rare in our region.
200 American Midland Naturalist Monograph No. 6
McKean Co.: Bennett Brook, Bradford, November 7, 1897, and Limestone Creek,
Bradford, October to December, 1896. D.A.B. (figured). The latter mixed with Grout's
No. 134, North American Musci Pleurocarpi.
2. Anomodon minor (Hedwig) Lindberg
(Neckera viticulosa var. minor Hedwig; A. obtusifolius Bryologia Europaea)
Plate XXXVIII
Loosely widely cespitose, glaucous-green, brownish when old; primary
stems creeping, flagellate, robust, with numerous, more or less erect, secondary
stems and branches usually up to height of 2-4 cm; leaves somewhat com-
planate, broadly Ungulate-obtuse from a broadly ovate base, thick, opaquely
chlorophyllose, entire, very densely papillose on both sides; costa pellucid,
rather strong, vanishing below apex; leaf-cells minute, about .009-. 012 mm,
rounded-hexagonal, the median basal elongate and non-papillose, the alar
scarcely different from the upper; perichaetial leaves sheathing: seta erect,
about 1 cm high, sinistrorse; capsule erect, castaneous, symmetric, oblong-
cylindric, about 3:1, the mouth small; lid conic-acuminate, about two-fifths
as long as urn; annulus present, large; peristome-teeth narrowly lance-linear,
hyaline, faintly papillose, about 8-10-nodose-articulate, the divisural and dorsal
lamellae x'ery faint or not visible; segments very short or rudimentary, or none,
from a very low basal membrane; exothecial cells rather thin-walled, irregularly
quadrate to oblong-rectangular; spores maturing in late fall or in winter,
medium to thin-walled, brownish, papillose, .009-.012 mm in diameter.
On rocks and trees, in woods, usually at the base of trees; Asia and from
New Brunswick to Dakota and south to the Gulf States.
Rather common in our region. Allegheny Co.: Near Montrose, September 21,
1905. O.E.J. Beaver Co.: On log along Raccoon Creek, 3 mi. s. of Traverse Cr.,
April 1, 1894. C.M.B. Butler Co.: On base of oak tree 1 mi. s. of Evans City. Jan.
13, 1935. Sidney K. Eastwood. Cambria Co.: T. P. James. (Porter's Catalogue).
Clearfield Co.: Phillipsburg. T. P. James. (Porter's Catalogue). Fayette Co.:
Laurel Run, 2 mi. e. of Wymps Gap, on tree. C.M.B. Aug. 24, 1940. McKean Co.:
Lewiston Creek, November 21, 1897. D.A.B. (figured). Washington Co.- Linn and
Simonton No. 43. Bark of tree near Washington, March 24, 1893.
3. Anomodon viticulosus [Linnaeus] Hooker and Taylor
Large, widely tufted, dark green above, yellowish within: stems creeping,
long, sending up secondary stems and branches, the secondary stems 4-10 cm
long, sometimes becoming more or less geniculate by repeated innovations:
leaves more or less crisped when dry, sub-falcate, secund, lance-lingulate from
an ovate base, frequently serrulate at the apex, apex bluntly acute; costa
strong, pellucid, ending a little below the apex; median, upper, and lower
marginal leaf-cells opaque, minute, rounded-quadrate, the median basal some-
what elongated, cells minutely papillose; perichaetial leaves long, linear-acu-
minate from an ovate base: seta twisted when dry, erect; capsule oblong-
cylindric, symmetric or slightly curved, about 3:1; lid narrowly conic; peris-
tome-teeth lance-linear, more or less irregular, yellowish, the inner peristome
consisting of a very low basal membrane and irregular segments up to 1/2
Jennings: Manual of Mosses — 29. Leskeaceae 201
or 2/3 the length of the teeth; annulus double; spores mature in winter or
early spring.
Mostly on shaded rocks, sometimes on trees; Europe, Algeria, Asia, and
in lower Canada and eastern United States south to Virginia. Occurs in
Pennsylvania at least as far west as Franklin County, and is to be looked
for in our region, particularly on calcareous habitats.
4. Anomodon attenuatus [Schreber] Huebener
(Leskea attenuata Hedwig; . HypnMw attenuatum Schreber)
Plate XXXVIII
Slender, loosely and widely tufted, with the secondary stems fasciculately
branched and with numerous slender flagelliform. branches; leaves spreading
to secund, concave, usually more or less distinctly homomallous when dry,
about 0.8-1.2 mm long, broadly lanceolate from an ovate base which is plainly
narrowed to the insertion, the insertion somewhat excavate and decurrent, the
apex acute and minutely apiculate and often with a very few teeth near the
apiculation, the leaf-margins plane below, and usually minutely crenulate by
reason of the projecting papillae; costa strong, ending a little below the apex;
areolation densely papillose on both sides, irregularly hexagonal to quadrate,
opaque, rather thin-walled, a few of the median basal elongate-rectangular to
oblong, pellucid; perichaetial leaves lance-acuminate from an ovate base: seta
about 2 cm long, twisted; capsule long, cylindric, straight or slightly curved,
lustrous; castaneous lid long-rostrate; teeth of peristome narrowly lanceolate;
segments filiform, irregular fragi'e, nearly as long as the teeth; annulus narrow;
spores macure in fall.
On bases of trees, stumps, and rocks, in woods; Europe, Asia, and from
Newfoundland to British Columbia and south to Florida and Cuba.
Common in our region but usually sterile. Known from 16 counties and probably
occurs in all. Specimen figured: Wildwood Hollow. Nov. 19, 1908. O.E.J. &: G.K.J.
5. Anomodon rostratus (Hedwig) Schimper
{Leskea rostrata Hedwig)
Plate XXXVIII
Densely cespitose, tufts bright green above, yellowish inside: primary
stems slender, creeping, fasciculately branched with short, slender julaceous
secondary stems and branches; leaves densely-imbricate, ovate and concave at
base, narrowly lanceolate above with a long and hyaline piliferous acumina-
tion, more or less indistinctly two-ranked, the margin crenulate-papillose, often
recurved towards the middle; leaf-cells minute, chlorophyllose, opaque,
rounded-quadrate to oblong-hexagonal, pluri-papillose on both faces, the me-
dian marginal rounded-quadrate, about .008-.010 mm, the median interior
about as wide but more oblong, about 2:1, the median basal longer, hyaline
and non-papillose or but slightly so, the apical long and linear, smooth; costa
strong and ending a little below the apex; perichaetial leaves long, pale, ecos-
tate, the inner with a filiform and often reflexed point about as long as the
202 American Midland Naturalist Monograph No. 6
main portion of the leaf: seta short, about 7-10 mm long, erect, sinistrorse,
richly castaneous; capsule about 2 mm long, oval-oblong, about 2.5:1, erect,
symmetric castaneous; lid conic, obliquely rostrate, about one-half to three-
fifths as long as the urn; teeth small, lance-linear, the divisural and dorsal
lamellae indistinct, the teeth with about 15 to 18 nodose articulations, pale,
papillose; segments about as long as the teeth, linear, rising from a basal mem-
brane about one-third as high as the teeth, the cilia solitary and rudimentary
or none; exothecial cells medium-walled, oblong-rectangular to oblong-he.xag-
onal, becoming quadrate above, about two rows at the rim much smaller and
heavily castaneus-incrassate; spores mature in fall, thin-walled, nearly smooth,
slightly brownish, about .010 mm in diameter.
In moist places, on rocks or more usually on the bases of trees; Europe,
Asia, and from Canada to the Gulf States and to the West.
Very common in our region, especially on the bases of white oak trees. Known from
collections from the following counties: Allegheny, Beaver, Bedford, Butler, Campron,
Centre, Clarion, Crawford, Erie, Fayette, Greene, Huntingdon, McKean, Somerset, Wash-
ington, and Westmoreland. Figured from specimens collected four miles up the valley of
Meadow Run, Ohio Pyle, Fayette Co., May 30-31, 1908, and Sept. 1-3, 1906. O.E.J.
5. Lindbergia Kindberg
(Fabroleskea Grout)
Autoicous: rather softly and loosely cespitose, bright to brownish-green,
dull: stem elongate, creeping, radiculose, densely-leaved, branched with
elongate mostly irregularly pinnate branches; branchlets short or unequal in
length, obtuse; dry leaves imbricate, when moist spreading to almost squarrose-
spreading, somewhat concave, more or less decurrent, ovate to lance-ovate,
abruptly subulate-acuminate, non-plicate, margins entire or rarely indistinctly
apically serrulate; costa strong, incomplete; median leaf-cells lax, round-oval
or rhombic hexagonal, smooth, or unipapillose, the marginal smaller and quad-
rate or transversely broader, the basal marginal in many rows quadrate to trans-
versely broader; inner perichaetial larger, thin, erect, from a sheathing base
subulate-acuminate, entire or serrulate; costa shorter: seta 5-10 mm long,
straight, thin, red, smooth; capsule erect, symmetric, oval-oblong, rarely slightly
curved, brown, small-mouthed and short-necked; annulus present or none;
peristome deeply inserted, teeth lanceolate, obtuse, basally confluent, pale to
yellow, non-striate, more or less papillose, divisural zigzag, low-trabeculate;
inner peristome papillose with a very low basal membrane, no segments, no
cilia; lid conic-obtuse; calyptra cucullate; spores .025-. 030 mm.
As here limited the genus consists of 13 species, occurring on tree- trunks
in the Northern Hemisphere; three species in North America; one species
in our region.
1. Lindbergia Austini (Sullivant) Brotherus
(Fdhroleskea AuUini Grout; Leskea Austini Sullivant; Lindbergia
brachyptera var. Austini (Sullivant) Grout)
Medium size, intricately matted; stems irregularly divided, the branches
usually quite unequal; leaves spreading to more or less squarrose when moist-
Jennings: Manual of Mosses — 29. Leskeaceae 203
ened, imbricate when dry, ovate, long and slenderly acuminate, strongly pap-
illose, entire; costa ending above the middle; leaf-cells unipapillate, elliptic-
rhomboid above, the basal marginal thick, rounded-quadrate; perichaetial
leaves longer, lance-acuminate: seta short, erect; capsule erect, oval-cylindric
with a small mouth; teeth broadly lanceolate, deeply inserted, opaque, papil-
lose on both surfaces, the inner peristome consisting merely of a low basal
membrane scarcely exceeding the rim of the urn; annulus none; lid short-
conic; spores mature in summer.
On tree-trunks and on rocks or stone-walls; southern Canada to South
Carolina and Arizona. In Porter's Catalogue the habitat is given as juniperiis
virginianiis. Not yet collected in our region.
6. Leskea Hedwig
Autoicous: rather slender, usually weak, loosely cespitose, dull, dark to
sooty-green: stems creeping, sparsely radiculose, rather thickly-leaved, more or
less pinnately branched, with short, erect or ascending branches; leaves when
dry imbricate, when moist erect-spreading to spreading, sometimes sub-secund,
from a somewhat decurrent, cordate-ovate base narrowed to an acute or obtuse
apex, sometimes apiculate, shortly two-plicate, revolute on one or both lower
margins, rarely indistinctly serrate at apex; costa strong, incomplete; median
leaf-cells either thin-walled, rounded-hexagonal, one- to several-papillose, at the
base almost quadrate, in the middle rhomboidal, or more or less thickened,
with oval to oblong acumen; branch-leaves smaller; inner perichaetial leaves
pale, sheathing, abruptly to slenderly acuminate, entire or serrulate, at the
apex, delicately and incompletely costate: seta long, thin, red, smooth; capsule
erect, oblong-cylindric, sometimes slightly curved and weakly inclined, yellow-
ish, finally light brown and plicate; annulus revoluble; teeth drying strongly
incurved, linear, acuminate, entirely separate, non-bordered, pale yellow, at the
base transversely dorsally striate, thickly trabeculate, papillose above; inner
peristome papillose, basal membrane low, segments linear, carinate, as long or
shorter than the teeth, cilia rudimentary or none; lid acute-conic; calyptra
cucullate, glabrous; spores small.
A widely distributed genus of about 15 species; 5 species reported for
North America; 2 (or 3) in our region.
Key ro the Species
A. Branch-leaves more or less secund, lanceolate; capsule sometimes slightly curved,
usually straight 1. L. polycarpa*
A. Branch-leaves not secund, ovate, acute to obtuse; cupsule erect, straight B
B. Branch-leaves two-plicate, symmetric, the margin often revolute 2. L. gracilescens
B. Branch-leaves non-plicate, often unsymmetric, plane-margined 3. L obscura
* Leskea arenicola Best has pinnately branched stems with distinct, small, central
strand; stem-leaves lance-ovate to ovate and secund; branch-leaves smaller, but scarcely
secund; costa ending in the rather blunt acumination; capsule curved; segments as long as
teeth; basal membrane low: lid long-conical.
Bases of trees, rotten wood, etc. New England to Virginia and west to the Mississippi.
Differ from polycarpa in more curved capsules and long conic opercukim.
204 American Midland Naturalist Monograph No. 6
1. Leskea polycarpa Ehrhart, Hedwig
(L. polycarpa Ehrhart; Hypnum medium Dickson)
Slender, thinly tufted: stems prostrate, pinnately to bi-pinnately branched,
2-4 cm long, with short curved or erect branches, intricately matted into close
but thin patches; central strand distinct; leaves erect-spreading to secund,
loosely appressed-imbricate when dry, about 0.4x1.0 mm, lanceolate and
gradually acute from a slightly decurrent, sub-cordate, more or less ovate base,
entire, usually two-plicate, acute to acuminate; costa ending a little below the
apex; median leaf-cells thin-walled, pellucid, more or less dorsally papillose,
hexagonal, about .007-.008 mm wide, with one or two papillae on each sur-
face; branch-leaves smaller and more obtusely pointed: seta about 1 cm. long,
reddish; capsule cylindric, narrow, basally tapering, straight or almost so,
reddish-brown, constricted below the mouth when dry; lid short-conic, acute;
peristome-teeth long, whitish, narrowly linear, connivent when dry; segments
about as long, from a low basal membrane narrowly linear, scarcely carinately
split: cilia rudimentary or none; spores mature in early summer.
On roots, bases of trees, stones, or decaying wood in wet situations; Asia,
and from Newfoundland to British Columbia and southward. Not yet found
in our region.
2. Leskea gracilescens Hedwig
(L. obscura Lesquereux and James, p. p.; Hypnum gracilescens Beauvois)
Intricately cespitose in thin mats: stems prostrate, pinnately branched with
numerous simple, erect, somewhat julaceous branchlets; central strand indis-
tinct or none; paraphyllia usually few, lanceolate; stem-leaves erect-spreading
when moist, appressed-imbricate when dry, about 0.4-0.5 x 0.7-0.9 mm, ovate,
acute or somewhat acuminate, entire, margins more or less revolute, some-
times more quickly tapering to a blunt point, somewhat bi-plicate; costa sub-
percurrent; branch-leaves hardly different but scarcely plicate; median leaf-
cells usually uni-papillate on dorsal surface, smooth on ventral, quadrate-
hexagonal, about .008-.010 mm, the apical more rounded, the basal somewhat
quadrate; capsule erect, basally tapering, oblong-cylindric; peristome-teeth
whitish, lance-linear, lamellate, about 0.4 mm long; the linear segments shorter,
carinate, sometimes more or less rudimentary; basal membrane I/4 as high as
teeth; cilia none; lid conic, acute to obtuse; spores mature in summer.
On the base of trees, roots, and on rotten logs, etc.; from eastern lower
Canada to the Gulf States and westward to the Rocky Mountains. Common-
est in northeastern United States. Reported from Middle Wheeling Creek,
W. Va., A. J. Sharp and Miss Gail Holliday, June 11, 1935; and:
Washington Co.: Linn and Simonton. (Porter's Catalogue).
3. Leskea obscura Hedwig
{L. nervosa Sullivant; L. microcarpa Schimp)er)
Plates XXXIX, LXVII
Small, loosely and intricately cespitose, dark green: stems prostrate, rather
irregularly divided, sparingly branched, the branches short, plump, erect; cen-
Jennings: Manual of Mosses — 29. Leskeaceae 205
tral strand small but distinct; leaves incurved-appressed when dry, spreading
when moist, about 0.8-1.2 mm long, from an ovate base narrowed above to a
rather blunt apex, concave, margins plane or incurved, entire or serrulate;
costa ending a little below the apex; median leaf-cells quadrate-hexagonal,
about .C08-.010 mm wide, with several small papillae on the lower surface,
on the upper surface less papillose or almost smooth, apical and basal cells
somewhat wider and shorter, the alar oblong-quadrate, longer towards the
costa; branch-leaves similar; perichaetial leaves long-sheathing, rather laxly-
celled, costate; seta about 1.5-2 cm long; capsule erect, straight, short-cylindric,
sometimes slightly curved, more or less wrinkled and contracted below the
mouth when dry; lid conic, rather obtuse; peristome-teeth yellowish, papillose;
segments linear, slender, partly carinately cleft between the articulations,
shorter than the teeth, arising from a basal membrane about one-fifth the
length of the teeth; spores mature in early summer; annulus 2-seriate.
On stones, roots of trees, logs, etc., often where sometimes overflowed;
Japan, and from New Brunswick to Ontario and southwards through the
eastern and central part of the United States. Probably fairly common in
our region, but sometimes difficult to distinguish from L. polycarpa or L.
gracilescens.
Allegheny Co.: On bark of white oak at three feet from ground. Fern Hollow,
Pittsburgh, March 8, 1908, and at base of trees in swampv woods near Douthett, Decem-
ber 29, 1908. O.E.J, (figured Plate XXXIX). Beaver Co.: On decaying bark, 3 mi.
east of Legionville, along Sewicklev Creek. May 5, 1918. O.E.J. Blair Co.: Tyrone.
T. P. James. (Porter's Catalogue). Cranxford Co.: On bark, Ulmtif americana. Con-
neaut Outlet. C.M.B. Sept. 23, 1949. McKean Co.: Tuna Creek. Bradford. Dec. 21,
1895. D.A.B. Tioga Co.: On large Elm tree, Goodrich Settlement, Tioga. June 24,
1935. Sidney K. Eastwood. Washington Co.: On tree, subject to inundation, near
Washington. May 28, 1892; near Arden, July 20, 1891; and at Hackberry Sta., June
16, 1894. All by Linn &: Simonton. Westmoreland Co.: Chestnut Ridge, s.e. of
Torrance. C.M.B. June 13, 1943.
7. Leskeella (Limpricht) Loeske
Dioicous: slender, in flattened, wide-spreading mats, dark green to brown-
ish, dull; stem widely creeping, fasciculately yellowish-red-radiculose, densely-
leaved, numerously-branched, with erect and short branches; leaves drying
imbricate, when moist trect-spreading to secund, more or less abruptly long-
acuminate from a decurrent, doubly-plicate, cordate base, margins narrowly
revolute below but plane in the acumen, entire; costa strong, yellow-brown,
ending in the acumen; cells rounded-hexagonal, in leaf-middle oval and oblong,
in middle of base rectangular, the alar quadrate; branch-leaves smaller with
plane margins and delicate and shorter costa; perichaetial leaves pale, from the
erect and half-sheathing base abruptly long-acuminate, delicately costate to
the acumen: seta elongate, stif?, dark chestnut-color, smooth; capsule erect,
symmetric, cylindric or oblong, rarely weakly curved, finally rust-colored to
brown; annulus rather persistent, deciduous in sections; peristome-teeth erect
when dry, confluent at base, subulate, bordered, yellowish, cross- and obliquely-
striate, smooth or papillose above, not distinctly trabeculate; inner peristome
finely papillose, basal membrane moderately prominent, segments irregular, in
206 American Midland Naturalist Monograph No. 6
nodose projections, filiform, etc., sometimes carinate, cilia mostly none; lid
convex, obliquely thick-rostrate; calyptra glabrous, cucullate and reaching to
base of capsule: spores small.
A small genus of 4 species; only the following in North America:
1. Leskeella nervosa [Bridel] Loeske
{Leskea nervosa Myrin; Lescuraea rigidula Kindberg;
Hypnum nervosum C. Mueller)
Plate XXXIX
Slender, in thin and appressed tufts, dark green to brownish, the older
parts almost black: stems with central strand distinct, creeping, up to 5 or 6
cm long, pinnately divided and again branched into numerous, crowded, short
and erect or longer and creeping branches, often with numerous gemmiform
branches towards the apex; stem-leaves close, broadly ovate, sub-cordate, open-
spreading when moist, imbricate when dry, slightly decurrent, about 0.4-0.5 x
1-1.2 mm, suddenly long-acuminate, the acumen recurved, the margins plane,
sub-sinuate, the blade concave and deeply plicate; costa almost percurrent,
slender; branch-leaves considerably narrower, lanceolate, more rigidly erect-
spreading, smaller, up to about 0.6-0.7 mm long; leaf-cells oblong to oval-
hexagonal, ranging from 1 to 3:1, the alar quadrate to transversely oval-hex-
agonal in about 4-6 rows extending well up the margins and becoming rounded;
cells smooth to lightly papillose, incrassate; inner perichaetial leaves long-
sheathing, long-acuminate: seta 10-12 mm long; capsule erect, sub-cylindric,
symmetric, small, castaneous; lid narrowly conic to short-rostrate; peristome
short, the teeth whitish, lance-linear; segments shorter, irregular, subulate,
basal membrane ^ the length of teeth; cilia none; annulus narrow; spores
mature in summer, rough.
Mostly on bases of trees, especially maples, in our region: Europe, and
in the northeastern United States south to Pennsylvania.
Not common in our region. Crawford Co.: Bark of Black Ash. Conneaut Outlet.
C.M.B. Sept. 23, 1949. Fayette Co.: Cheat Haven, September 3-6, 1910. O.E.J, and
G.K.J. McKean Co.: Bennett Brook, Bradford, July 12, 1896 (figured), and Bolivar
Run, 1896. D.A.B.
8. Rauia Austin
Autoicous: quite slender, dull, bluish-green to brownish-green: stems
thickly-leaved, simple or divided, more or less regularly pinnately branched;
branchlets short, ascending, julaceous, obtuse; leaves dimorphic, drying im-
bricate, when moist erect-spreading; stem-leaves plicate, triangular to cordate-
ovate, tapering to a lanceolate or lance-subulate acumination, the margins en-
tire; costa strong, incomplete; median leaf-cells rounded-angular, with low and
usually numerous papillae on both sides; branch-leaves lance-ovate, short-
pointed, the costa dorsally somewhat rough; inner perichaetial leaves appressed,
pale, lance-oblong, slenderly acuminate, entire, incompletely costate, and with
elongate, smooth cells: seta slender, 10-15 mm long, reddish, smooth; capsule
from nearly upright to horizontal, oblong-cylindric, mostly weakly curved,
light brown, drying more or less constricted below the mouth; annulus revol-
Jennings: Manual of Mosses — 29. Leskeaceae 207
uble; peristome-teeth lance-subulate, bordered, transversely striate, numerously
trabeculate; inner peristome yellow, finely papillose, basal membrane promi-
nent, carinate, the segments about as long as the teeth, carinately split, the
cilia 2 or 3, more or less complete, nodose; lid conic, short-rostrate; calyptra
cucullate; spores .009-. Oil mm.
A genus of about 14 species, widely distributed in both hemispheres; 4
species in North America; 1 species in our region.
1. Rauia SCITa (Beauvois) Austin
(Thuidium scitum Austin; Hypnum scttum Beauvois)
Plate XXXIX
In appressed, green or yellowish-brown mats, medium-sized: stem prostrate,
castaneous, 2-3-times divided; densely pinnately branched; branchlets short
and slender, usually about 2-3 mm long, smooth, paraphylla present; stem-
leaves broadly cordate-deltoid, acuminate, about 0.5-0.6 mm long, margins
plane, finely papillose-denticulate; branch-leaves ovate-cordate, smaller, shorter
acuminate, both kinds of leaves concave, erect-spreading; costa pellucid, broad,
about three-fifths to four-fifths as long as the leaf; median leaf-cells rounded-
hexagonal, minute, .006 mm diameter, with 2-5 small bead-like papillae on
each surface, incrassate, rather obscure, the basal median oblong, paraphyllia
numerous, linear to ovate, more or less branched, occurring on both stem and
branches; irmer perichaetial leaves elongate-lanceolate, filiform-acuminate,
somewhat longitudinally plicate: capsule sub-erect, about 1.3-1.5 mm long,
rather thin-walled, when old and empty more or less wrinkled, urn cylindric,
straight or slightly curved, more or less twisted when old, the seta about 1.5
cm long; lid conic-rostrate, curved upward; annulus large; exothecial cells
rather thin-walled, mainly quadrate to rectangular; peristome-teeth lance-subu-
late, shallowly inserted, castaneous and transversely striolate below, hyaline
and papi'lose above; lamellate and trabeculate; segments nearly as long as
teeth, carinately partially split, the basal membrane about two-fifths as high,
the cilia usually three, nodose; spores medium-walled, castaneous-pellucid,
papillose, about .010-.013 mm, mature in fall and winter; autoicious.
On the bases of trees fnd on stones in woods; from Ontario to Missouri,
eastward to the Atlantic Ocean and southward to North Carolina.
Rare in our region. Allegheny Co.: Moon Twp., April, 1902. J. A. Shafer. Mc-
Kean Co.: On base of trees, Rutherford, August 4, 1897, Bradford, October, 1897;
Gates Hollow, July 28, 1895, and Limestone Creek, Bradford, October to December,
1896. D.A.B. (figured). The last named specimen issued with Grout's No. 134, in part,
North American Musci Pleuiocarpi.
9. Haplocladium (C. Mueller) C. Mueller
Autoicous: slender, forming mats, yellowish-green to brownish-yellow,
dull: stems creeping, elongate, with brownish rhizoids, variously pinnate with
branches mostly ascending, julaceous, short, obtuse and simple, or somewhat
longer, acute and pinnate with scattering short branchlets; leaves more or less
uniform, drying appressed, sometimes weakly secund, when moist erect-
208 American Midland Naturalist Monograph No. 6
spreading; stem-leaves more or less doubly plicate, from a more or less broadly
ovate base, lanceolate to lance-subulate, the margin revolute at base, the upper
margin indistinctly serrulate to entire; costa strong, sometimes percurrent,
sometimes excurrent, mostly smooth; cells more or less pellucid, oval to oblong-
hexagonal, with one papilla over the lumen, the alar quadrate; branch-leaves
narrower at the base, shorter- pointed, plane- margined, more or less distinctly
serrate, with a single terminal papilla on the apical cell; costa shorter, cells
mostly opaque; inner perichaetial leaves erect, pale, plicate, from a lanceolate
or linear base long-acuminate, incompletely costate: seta 1.5-2.5 cm long,
red-castaneous, smooth; capsule inclined, oblong-cylindric, drying more or less
horizontal and arcuate, v hen old and empty contracted below the mouth;
annulus present; peristome-teeth lance-subulate, yellow, bordered, transversely-
striate, dorsally lamellate; inner peristome yellowish, the basal membrane wide
and carinate, the segments lance-subulate, of same length as teeth, carinate,
entire or narrowly split, cilia complete, 2 or 3, slender, nodose or appendicu-
late; lid convex-conic; acute; calyptra cucullate; spores .008-.012 mm.
A genus of nearly 50 species, mostly occurring in eastern Asia and in
South America; only the following in our region.
Key to the Species
A. Stem-leaves rounded-ovate, short-acuminate; margins erose-serrate 1. H. virgin'tanum
A. Stem-leaves ovate, more or less long-acuminate; entire to crenulate serrate
2. H. m'crophyllum
1. Haplocladium virginianum (Bridel) Brotherus
(Thuidium virginianum Lindberg; T. gracile var. lancastriense Cardot; Hypnum
gracile var. lancastriense Sullivant and Lesquereux)
Plate XXXIX
Small to medium-sized, appressed-cespitose, dark to dirty green: stems
diffusely divided, the branches short and erect or ascending pinnately; leaves
of the stems rounded-ovate, concave, narrowed to the base but scarcely decur-
rent, about 0.6-0.8 mm long, abruptly acuminate, costate into the acumen,
serrulate above, erose-dentate below, appressed when dry, loose when moist;
costa ending in the apex; median leaf-cells quadrate-hexagonal, uni-papillate,
rather incrassate, the lower marginal more or less transversely oblong-quadrate
or hexagonal; branch-leaves about 0.4-0.6 mm long, broadly and shortly acu-
minate with a serrulate margin above; perichaetial leaves long, pale, up to 2.5
mm long: seta slender, about 2-2.5 cm long, rather richly castaneous, dextrorse;
capsule oblong-cylindric, castaneous, curved, more or less horizontal, often
when old more or less pendent by the curving of the upper part of the seta,
annulate, the urn about 2.5:1, about 2 mm long, constricted below the mouth
when dry and empty; lid obtusely short-beaked, about one-third as long as the
urn; peristome normally hypnoid, yellowish, the 16 teeth lance-linear, dorsally
cross-striate, with zigzag divisural line, distinct dorsal lamellae, and about
35-40 closely placed trabeculae; segments about as long as teeth, carinate;
cilia (1-) 2-3, nodose-articulate, the basal membrane about two- fifths the
height of the peristome; exothecial cells incrassate, quadrate to oblong-hexag-
Jennings: Manual of Mosses — 29. Leskeaceae 209
onal, about three series in the rim much smaller and rounded; spores incras-
sate, yellowish, faintly papillose, about .011-. 014 mm, mature in spring.
On the ground or on roots of trees in rather open woods; from New Eng-
land to Minnesota and Mexico, also in Europe.
Rather common in our region. Now known from Beaver, Blair, Cambria (Porter),
Centre, Fayette. Huntingdon, and Westmoreland counties, all in the southern half of our
region. Specimen figured: Warrior's Ridge above Huntingdon, July 20 1908. O.E.J.
2. Haplocl.^dium microphvllum (Swartz; Hedwig) Brotherus
(Hypnum gracile Bruch and Schimper; Thuidium microphyllum Best )
Plate XXXIX
Medium size, pale green to yellowish, appressed-matted : stems diffusely
divided, densely pinnately branched; stem-leaves broadly ovate to lance-ovate,
up to 1.2 mm long, concave, long and narrowly acuminate, the margins entire
or sinuately denticulate; ccsta almost percurrent; leaf-cells quadrate-hexagonal,
somewhat incrassate, the apical and a very few of the basal elongate, all
strongly uni-papillate; branch-leaves narrower and smaller, up to 1 mm long;
paraphyllia numerous and branched on the stem but simple and few or none
on the smaller branches; inner perichaetial leaves high-sheathing, long-acu-
minate, costate, up to 2.5 mm long: seta up to 2 or 2.5 cm long, castaneous,
twisted, somewhat arcuate above; capsule turgid-oblong, about 2-2.5:1, about
2 mm long, dorsally somewhat turgid, when mature somewhat inclined or
pendent by the curving of the upper part of the seta; lid short-conic, mamil-
late; peristome normally hypnoid, the teeth yellowish, densely trabeculate, dor-
sally with rather indistinct lamellae and divisural, finely cross-striate, narrowly
hyaline-margined; segments carinate, about as long as teeth; cilia 2 to 3, a
little shorter than the teeth, strongly nodose to shortly appendiculate, the basal
membrane about one-third as high as teeth; annulus simple; spores mature in
summer, somewhat incrassate, pale brownish-pellucid, very slightly roughened.
On earth, rotten wood, etc., often on bases of trees; Europe, Asia, and
from southern Canada to the West Indies.
Rather rare in our region. Bedford Co.: Raystown Branch, Juniata River, l^/i mi. s.
of Schellsburg. July 19, 1941. C.M.B. Butler Co.: On roadside bank. Ribold.
C.M.B. June 2, 1945. Erie Co.: On dead log, Presque Isle. Nelle Ammons. July 13,
1935. McKean Co.: On hillside at mouth of Langmade Hollow, Bradford, November,
1895 (figured) and Limestone Creek, N. Y., near the Pennsylvania State line north of
Bradford, October 16, 1896. D.A.B. Washington Co.: On rotten log. Snake Woods,
near Washington. Aug. 13, 1892. Linn H. Simonton.
10. Thuidium Bryologia Europaea
Fern Mosses
Autoicous or dioicous. slender to robust, mostly stiff, dull, forming
greenish to yellowish or brownish mats or cushions: stem with a few-celled
central strand, spreading to ascending or rarely erect, radiculose here and there
in fascicles, not much divided, once to thrice pinnately branched, flattened
like the frond of a fern; leaves dimorphic, when dry incurved or appressed,
210 American Midland Naturalist Monograph No. 6
when moist erect-spreading or open-erect but never secund; stem-leaves plicate,
from a narrowed and decurrent base, mostly with revolute margin, entire or
apically toothed; costa strong, mostly incomplete, rarely excurrent, sometimes
dorsally rough, leaf-cells rather uniform, rounded to oval- or oblong-hexagonal,
both sides numerously papillose or uni-papillose dorsally or on both sides;
branch-leaves of first order often similar to stem-leaves, those of the second or
third order smaller, mostly lance-ovate, with the costa weaker and shorter;
inner perichaetial leaves pale, appressed, mostly lanceolate and plicate, often
with a prolonged and filiform apex, sometimes with ciliate margins, costa in-
complete, cells elongate, smooth: seta elongate, castaneous or red, smooth or
rough; capsule inclined to horizontal, oval-oblong to cylindric, more or less
arcuate, brown to yellow, drying constricted below the mouth; annulus some-
times persistent; peristome-teeth basally confluent, lance-subulate, bordered,
cross-striate, numerously trabeculate; inner peristome yellow to orange, smooth
or finely papillose, with prominent carinate basal membrane, segments as long
as teeth, carinately split at least in part; cilia 2-4, nodose to appendiculate, or
sometimes rudimentary, or none; lid convex-conic, obliquely rostrate; calyptra
cucullate, mostly smooth; spores .007-.010 mm or .012-. 016 mm.
A widely distributed genus of about 160 species, on tree-trunks, rocks, or
earth; about 25 species reported for North America; at least 4, probably more,
in our region.
Key to the Species
A. Delicate, small, not over 5 cm; 1-2 pinnate B
A. Larger, up to 10 cm; 1-3 pinnate C
B. Branchlets papillate; leaf-cells about .006 mm: seta 1-1.5 cm long ....2. T. pygmaeum
B. Branchlets smooth; cells about .009 mm: seta 2-3 cm long 1. T. minutulum
C. Stems simply pinnate; plants ascending in tufts 3. T. abietinum
C. Stems 2-3 pinnate, forming flat mats d
D. Leaf-margin revolute; costa not filling entire apex of leaf E
D. Leaf-margin plane; costa of stem-leaves filling the entire ap>ex; perichaetial leaves
not ciliate 4. T. recognitum
E. Stem-leaves lance-acuminate, from a triangular-ovate base; perichaetial leaves ciliate
7. T. delicatulum
E. Stem-leaves long-lance subulate; perichaetial leaves not ciliate
6. T. Philiberti
E. Stem-leaves shortly and broadly acuminate; perichaetial leaves not ciliate 5. T.Alleni
1. Thuidium minutulum [Hedwig] Bryologia Europaea
(Hypnum minutulum Hedwig)
Plate XXXIX
Small, slender, simply pinnate; stems irregularly divided, not over 3 or 4
cm long, both stems and branches smooth, bearing rather few linear-oblong
simple paraphyllia only about 2 to 5 cells high, notched at apex; stem-leaves
distant, deltoid, acuminate or apiculate, somewhat revolute on the borders,
rather opaque, about 0.6 0.8 mm long; costa strong, ending near the apex;
median leaf-cells irregularly polygonal to quadrate-hexagonal, the marginal
somewhat larger and sometimes transversely elongate, all leaf-cells incrassate,
pluri-papillose, the apical cell with 2 to 5 marginal papillae; branch-leaves
Jennings: Manual of Mosses — 29. Leskeaceae 211
ovate-acuminate, about 0.2-0.3 mm long, concave and with a shorter costa;
perichaetial leaves erect, slenderly lance-acuminate, the acumen more or less
reflexed: seta about 2-2.5 cm long, slender, dark yellow or brown and sinis-
trorse when old; capsule yellowish, oval-oblong, cernuous to horizontal, the
urn about 2 mm long, when dry somewhat constricted below the rim; lid
obliquely subulate-rostrate and about 1 mm long; peristome castaneous, the
teeth slender, densely trabeculate, the dorsal lamellae cross-striate and pro-
jecting to form a narrow margin, the divisural distinct; segments as long as
the teeth, split carinately, arising from a basal membrane about one-third as
high, cilia usually 2, articulate, nearly as long; spores about .010-.012 mm, pale
brown, medium- walled, slightly roughened, mature in fall.
On rotten logs and stumps and at the base of trees in woods. Europe. In
North America from Florida and Mexico north to New Brunswick and
Minnesota.
In our region known from Allegheny, Blair, Fayette, Huntingdon, McKean and Wash-
ington counties. Specimen figured: On stump. Meadow Run Valley, 4 mi. s. of Ohio
Pyle. Sept. 1-3, 1906. O.E.J. & G.K.J.
2. Thuidium pygmaeum Bryologia Europaea
Plate LXVI
Minute, rather harsh, in dull, olive or yellowish green, thin, interwoven
mats; bi-pinnate, 1-2 cm long; branches capillary; with insignificant simple 2-5-
celled paraphyllia; stem-leaves triangular-ovate but acuminately narrowed, the
lower margins often more or less reflexed, median cells pluripapillate, incras-
sate, hexagonal-quadrate, about .006 mm in diameter; primary branch-'eaves
long, ovate with obtuse or sub-acute apex; seta about 1.5 cm long; capsule in-
clined to horizontal, unsymmetrical, wide-mouthed; annulus conspicuous; lid
conic, obliquely rostrate; peristome complete; spores ripening in fall.
Mostly on stones (or soil) in calcareous districts. So stiff and intricately
interlaced that it is difficult to separate out individual plants. First described
from specimens collected at Columbus, Ohio. Range now extended to Penn-
sylvania, New Jersey, and Canada.
Washington Co.: Linn and Simonton. On stones, in a shaded ravine, McCracken
Station, vicinity of Washington. Sept. 24, 1892 (figured).
3. Thuidium abietinum [Linnaeus] Bryologia Europaea
(Hypnitm abietintim Bridel )
Large, yellowish-green above, brownish below, in dense tufts or mats; stem
5-12 cm long, regularly pinnate, stiffly ascending-erect, the branches 2-10 mm
long, unequal, attenuate, slender, terete when dry; stem-leaves crowded, ovate,
acuminate, 0.85 mm wide by 1.3 mm long, deeply bi-plicate with margins pap-
illose-serrulate; costa strong ending well above the middle; median cells oval-
rhombic, uni-papillate, about .010 mm in diameter; branch-leaves smaller, with
one or two papillae on the apical cell; inner perichaetial leaves plicate, with
toothed but not ciliate margin; seta 1.5-3 cm long; capsule cylindric, curved,
212 American Midland Naturalist Monograph No. 6
suberect, tapering at base; annulus distinct; operculum long-conic, acute;
spores in spring.
Mostly in dry places, on earth, stones, or rocks, especially in calcareous
habitats. Arctic America south to Virginia and Colorado. Collected by Por-
ter on a cliff near Easton, Pa.
4. Thuidium recognitum [Hedwig] Lindberg
{Hypnum recognitum Hedwig; T. delkatidum Bryologia Europaea)
Plate XL
Yellowish-green, not as bright-colored as some of the other Thuidiums,
quite similar to the preceding but mostly bipinnate: the primary branches are
nearly equal in length, thus making the general outline of the frond more
linear-oblong; stem-leaves usually about 0.6 to 0.8 mm long, broadly triangu-
lar, auriculate-cordate, abruptly acuminate, recurved-spreading when moist,
the apex acute and often very slender, the serrulate margins usually plane, the
leaves sulcate when dry; costa sub-percurrent, somewhat broadened at apex;
leaf-cells incrassate, and each with a long, slender, upcurved dorsal papilla, the
median rhombic-oblong to rounded-quadrate, the apical somewhat narrower;
branch-leaves with apical cells with 2-4 papillae; perichaetial leaves up to 4.5
mm long, slenderly long-acuminate, non-ciliate: seta about 2-2.5 cm long,
slender, castaneous, lustrous, dextrorse above; capsule oblong-cylindric, arcuate,
more or less inclined, the urn castaneous, 2.5-3 mm long, when dry quite
sharply bent and constricted at the mouth; peristome as in T. dehcatulum;
annulus large, deciduous, pluri-seriate; lid short- rostrate; spores with yellowish,
medium-thick, granular walls, about .012-.014 mm, mature in midsummer.
On the ground on rocks or on rotten wood in moist, shaded woods;
Europe, Asia, northern Africa, and from northern Canada southeastward
to Florida.
Allegheny Co.: On clay bank under hemlocks, Wildwood Road Hollow, March
29, 1908. O.E.J, and G.K.J.; Guyasuta Hollow, October 25, 1908. O.E.J. Armstrong
Co.: Kittanning, September 24, 1904. O.E.J, (figured). Clinton Co.: Between Reno-
vo and Haneyville, July 15, 1908. O.E.J. Erie Co.: Moist woods. Presque Isle. Nelle
Ammons. July 2, 1935. McKean Co.: Langmade Hollow, May 3, 1896, Toad Hollow,
June 17, 1896, and Bolivar Run, July 17, 1897, all near Bradford. D.A.B. Washing-
ton Co.: On base of beech tree. Claysville. Linn &C Simonton. Dec. 17, 1892.
5. Thuidium Alleni Austin
Forming rather loose and thick, wide mats; stems more or less ascending,
somewhat irregularly bipinnate; paraphyllia numerous; stem-leaves ovate,
broadly acuminate, with apex acute to obtuse, sometimes more slenderly taper-
ing; margins of leaves crenulate-papillose; costate to well above the middle;
on each surface; paraphyllia few or none on branches; branch-leaves acuminate
median leaf -cells incrassate, rather oblong, 1-2:1, with one or more low papillae
from a concave, broadly ovate base, acute to obtuse; perichaetial leaves not
ciliate. According to Grout {Moss Flora) abundant in Florida. Reported
from other states as far north as Pennsylvania and Connecticut. "Mature
sporophytes unknown" ^ — Grout.
Jennings: Manual of Mosses — 29. Leskeaceae 213
6. Thuidium Philiberti Limpricht
(Thiddium intermedium Philibert, not Mitten)
Grout (Moss Flora) regards this as "A poorly delimited subspecies of T.
delicatulnni." Stems to 12 cm or more, bi- to tri-pinnate; branches up to 1
cm; stem leaves cordate-lanceolate, narrowed into a filiform acumination of a
single row of 3-8 linear cells, about 1.5-1.8 mm by 0.6 mm; costa strong ending
at or in the long acumination; median cells incrassate, roundish-oval, towards
th basal margin much more rectangular; branch-leaves ovate, acuminate, about
0.6 mm long by 0.4 mm wide, margins plane, apical cell with 2 or 3 points;
inner perichaetial leaves with the slender acumination about three times the
length of the body of the leaf mostly without cilia; seta 1.5-2.5 cm long;
capsule inclined, curved, cylindric, about 3 mm by 0.8 mm, castaneous; lid
cbliquely-rostrate; spores ripe in fall and winter.
In wet swampy situations on ground or bases of trees. New Jersey, Penn-
sylvania, and in the West. Not yet known from our region.
7. Thuidium delicatulum [Linnaeus] Mitten
Common Fern Moss
Plate XL
The "Common Fern Moss." — Bright green above, darker below, large,
forming rather large and intricately woven mats, when dry rather stiff and
harsh; stems elongate, procumbent or arched and alternately rooting, often
reaching a length of 10 or 12 cm; the branching is twice or thrice pinnate,
very regular and fern-like; stem-leaves triangular-ovate, somewhat cordate,
gradually acuminate, about 1 m.m long, erect-spreading, appressed when dry.
somewhat serrate and marginally m.ore or less recurved; leaf-cells uni-papillate
on both rides, rather incrassate, the median quadrate-oblong to rhombic-oval
or rounded-quadrate, about .007 to .008 mm across; costa strong and ending
in the acumen; branch-leaves much smaller, broadly ovate, acuminate, the
apical cells with 2 (to 4) papillae, usually appearing bifid; inner perichaetial
bracts with a very few, filiform, articulate cilia on the margins: seta about 2
to 3 cm long, rather stiff, richly castaneous, somewhat dextrorse; capsule large,
the urn about 3.5 to 4 mm long, strongly inclined to horizontal, arcuate, nar-
rowly oblong, yellowish to castaneous; lid slenderly conic-rostrate, about 1.5
mm long; annulus narrow, usually 2-seriate; peristome large, reddish, the teeth
strongly and numerously trabeculate, often split at the hyaline and papillose
apex, dorsally cross-striate below, the divisural and lamellae distinct; segments
about as long as teeth, carinately split, papillose above; the 2 or 3 slender
nodose cilia rising from a basal membrane about one-third as high as the
peristome; spores rather thin-walled, pale-castaneous, very slightly roughened,
about .014-. 017 mm, mature in winter.
On the ground, stones rotten wood, stumps, etc.; in moist woods; Europe,
Asia, and from Labrador to the Rocky Mountains and south to the West
Indies and South America. Very common in the woods of our region.
Known from 18 coiintieri in western Pennsylvania, and probably occurs in all. Speci-
men figured: Power's Run, Allegheny Co., Sept. 14, 1905. O.E.J, and G.K.J.
214 American Midland Naturalist Monograph No. 6
11. Helodium (Sullivant) Warnstorf
(Elodtium (Sullivant) Warnstorf)
Autoicous or rarely dioicous: more or less robust, in deep, soft, slightly
lustrous, green to yellowish-green or sometimes brownish tufts: stems elliptic
in cross-section, without central strand, mostly simple, thickly-foliate, some-
what distichously pinnate; branchlets filiform; paraphyses small, branched,
numerous; leaves all similar, when dry appressed, when moist erect-spreading,
or erect, concave, with a dorsally projecting main plication; stem-leaves from a
narrowed base suddenly lance-ovate, acuminate, the margin almost wholly
revolute, mostly entire, sometimes apically serrate; costa incomplete, mostly
weak; cells pellucid, elongate-hexagonal to almost linear, smooth or unipapil-
late over the lumen or in the cell-angle, the basal cells laxly rectangular;
branch-leaves smaller; perichaetial leaves erect, pale, plicate, delicate, narrowly
acuminate, incompletely costate: seta 2-5 cm long, smooth; capsule inclined
to horizontal, oblong-cylindric, brown, more or less arcuate; annulus revoluble;
peristome-teeth broadly lance-subulate, yellow, basally confluent, transversely
striate, apically almost smooth, hyaline-bordered, high-trabeculate, the plates
numerous, and often forked or with transverse walls; inner peristome yellowish,
almost smooth, with high basal membrane, carinate; segments as long as teeth,
lance-subulate, entire or very narrowly carinately split, cilia 3, complete, deli-
cate, smooth; lid convex-conic, acute; calyptra cucullate, glabrous; spores about
.010-.016 (-.024) mm.
A genus of five species, at least one of these in our region.*
1. Helodium paludosum (Sullivant) Austin
{Hypnum paludosum Sullivant; Thuidium paludosum Jaeger and Sauerbeck)
Plate XL
Yellowish-green, irregularly pinnate: primary stems creeping, branchlets
distichous, unequal: stem leaves somewhat rigid, about 1-1.5 mm long, erect-
spreading to somewhat appressed, lance-oblong, acuminate, somewhat cordate
at base, concave below, reflexed on the borders, smooth on both faces, at the
base bearing 1 to 3 paraphyllose branched filaments, the base decurrent, plicate-
striate; costa sub-percurrent; median leaf-cells shortly linear-oblong to linear-
rhomboid, usually smooth, sometimes dorsally lightly papillose at the distal
end; stems and branches vvith numerous filamentous and branched paraphyllia;
branch-leaves narrower, smaller, usually 0.6-0.8 mm long; inner perichaetial
leaves oblong, gradually slenderly acuminate, up to 3 mm long, longitudinally
plicate: -eta about 1.5-3 cm long, slender, red-castaneous, dextrorse above;
capsule oblong-r\'Iindric, curved, strongly inclined to almost horizontal, about
3:1, the urn about 3-3.5 mm long; lid conic, apiculate; peristome normally
* Helodium Blandowii (Weber & Mobr) Warnstorf, is larger than paludosum,
having stems up to 10 cm longi and very regularly pinnate; the stem-leaves quickly nar-
row to a somewhat clasping base bearing paraphyllose appendages; leaf-cells distinctly
papillose dorsally at the distal end. — On wet marshy ground, Eurasia, and from New
Jersey, Ohio, and Colorado northwards.
Jennings: Manual of Mosses — 30. Hookeriaceae 215
hypnoid, large, the teeth tather broadly lance-acuminate, densely trabeculate,
the lower trabeculae often forked and thus united by obliquely transverse bars,
the dorsal lamellae numerous and below densely cross-striate, yellowish; seg-
ments as long as teeth, carinate but rarely split, the basal membrane about
one-third as high as teeth, the cilia 3, nodose or appendiculate above, nearly
as long as segments; spores mature in winter, about .018-. 022 mm, medium-
walled, yellowish, granular; annulus large.
In wet, grassy fields, swamps, and bogs; Asia, and from New England to
Ontario and south to Illinois and probably rather common in the northern
part of our region.
Allegheny Co.: Swampy ground near Douthett, about on boundary line of Butler
and Allegheny counties, April 26, 1908. O.E.J. Bedford Co.: In swamp along Rays-
town Branch, Juniata River, south of Schellsburg. July 19, 1941. C.M.B. Butler Co.:
Svv?mpy ground near Crider's Corner's April 26, 1908. O.E.J. Crawford Co.: Pyma-
tuning Swamp, near Linesville, June 12, 1905. O.E.J, (figured). Erie Co.: On decay-
ing log in swamp, Presque Isle. Nelle Ammons. Aug. 3, 1935.
la. Helodium paludosum var. HELODioiDES (Renauld and Cardot) Best
{Thuidtum elodioides Renauld and Cardot)
Leaves smaller with margins dentate-serrate, the cells more or less strongly
and often sub-centrally papillose; darker green; cells shorter, elliptic or oval.
In swampy meadows, swamps, bogs, etc.; from New York to Ohio and
Indiana. Apparently rare m our region.
Family 30. HoOKERIACEAE
Soft, often complanately foliate; leaf-cells smooth, thin-walled, mostly
parenchymatous, alar not differentiated.
A large family of about 700 species mostly tropical and on trees or decay-
ing wood. Only Hookeria in our region, with one species.
1. Hookeria acutifolia Hooker
(Pterygophyllum lucens SuUivant (1864), not Bridel (1819);
Pterygophyllum acuminatum Paris)
Plate LXVII
In large, soft, light green mats with somewhat complanate foliate branches
5-10 mm wide, in ours about 2-3 cm long. When dry, the leaves are thin,
soft, translucent, lustrous, and yellowish. Leaves ovate to oblong-ovate, acute,
4-5 mm long, entire, slightly decurrent, the lateral somewhat unsymmetric, the
apex often slightly eroded, sometimes producing protonemal filaments; leaf-
cells somewhat irregularly oblong-hexagonal, somewhat shorter at basal angles
and at apex and slightly narrower along the margin, the median about 3-5:1.
Not found fruiting in our region.
On earth and rocks in cool wet situations. Connecticut to Ohio and the
southern Appalachians.
Fayette Co.: Ohio Pyle on rock below Cucumber Falls, wet with spray. O.E.J. &C
G.K.J, and Dr. and Mrs. W. H. Emig, May 12, 1917 (figured); on dripping shaded
216 American Midland Naturalist Monograph No. 6
rocks, Meadow Run ravine, Sullivant Moss Society Foray. John Churchill and C. M.
Boardman, June 23, 1940.
Family 31. Hypnaceae
Autoicous or dioicous, rarely pseudautoicous or polyoicous: antheridial
clusters gemmiform, small, archegonial clusters on short mostly rooting peri-
chaetial branches: slender to robust, variously cespitose, rarely floating, dull to
lustrous: stem without central strand, mostly woody, often stoloniferous, mostly
irregularly pinnate, but the branches often regularly pinnate; leaves pluriseriate,
unistratose, erect-spreading to squarrose, rarely densely imbricate, often secund
or circinate, of various forms, sometimes unsymmetric; costa homogeneous,
mostly thin and rather short, simple, double, forked, or none, rarely strong
and complete to excurrent; leaf cells mostly narrowly prosenchymatous, rarely
parenchymatous, at the base looser, the alar mostly differentiated into a dis-
tinct group, rounded to oval or 4-6 sided, small to inflated, mostly hyaline:
seta elongated, mostly smooth; capsule mostly inclined to horizontal, mostly
arcuate, rarely pendent, or erect, mostly smooth; collum scant; peristome
double, both parts of same length, teeth lance- subulate, mostly strongly hygro-
scopic, mostly confluent at base, rarely separate, yellow, red-brown to purple,
mostly transversely striate with divisural zigzag, with trabeculae numerous and
well-developed; basal membrane of inner peristome wide, segments keeled,
mostly lance-subulate, cilii mostly complete, filiform, nodose to articulate,
rarely rudimentary or none; lid usually conic-convex, in our species obtuse to
acute or only very shortly rostrate; spores small.
As here treated, a large and cosmopolitan family, distributed on all kinds
of substrata. The limitations of both the family and the genera are treated
variously by the authorities.
Key to the Genera
A. Costa in our species single, usually extending to leaf-middle or beyond; lid never
strongly rostrate; plants not complanately foliate; capsules mostly curved-cylindric,
or subcylindric d. (Amblystegiae)
A. Costa short, double or single or none, rarely single in Homomallium and Hygro-
hypnum; lid sometimes rostrate B
B. Stem-leaves and branch-leaves usually distinctly dissimilar, at least as to size, sym-
metric and normally inserted O
B. Stem- and branch-leaves more or less closely similar, although often secund or
falcate, or inserted obliquely and unsymmetricallv C
C. Leaves either symmetric and normally inserted or unsymmetric and obliquely in-
serted; lid sometimes rostrate T
C. Leaves more or less obliquely inserted and apparently two-ranked, mostly unsym-
metric; branches mostly complanate; lid conic to short-rostrate, rarely long-
rostrate u
D. Leaves bordered 7. Sciaromium
D. Leaves non-bordered E
E. Leaves mostly large, broad, and obtuse or sometimes apiculate 9. Calliergon
E. Leaves not as above F
F. Costa strong, sub-percurrent, or sometimes excurrent C
F. Costa not reaching leaf-apex K
G Paraphyllia often present, polymorphic; leaves non-plicate, mostly erect-spreading;
more or less aquatic; basal leaf-cells mostly rectangular, mcrassate, pellucid
Jennings: Manual of Mosses — 31. Hypnaceae 217
6. Hygroamblystegium
G. Not as above; paraphyllia none or scarce H
H. Leaf-cells linear-vermicular to the leaf-base, mostly with blunt ends, alar cells
forming a small, distmct, fairly well-defined group of quadrate or rectangular
cells 1 1. Hygrohypnum
H. Leaf-cells hexagonal and 2-6 times as long as wide or prolonged-linear and be-
coming wider and shorter basally, alar cells, forming a group which is large
and often extends to the costa -.I
I. Ends of shoots usually circinate. Leaves falcate to circinate; alar cells parenchyma-
tous; mostly enlarged and mflated 8. Drepanocladus
I. Alar cells more or less prosenchymatous or rounded J
J. Median leaf-cells prolonged-linear; leaves concave, large, ovate to oblong or cir-
cular 9. Calliergori
J. Median leaf-cells prosenchymatous-hexagonal, 2-6 times as long as wide; aquatic;
leaves mostly erect-spreading 6. Hygroamblystegium
K. Leaves cordate to ovate-lanceolate, acuminate; costa weak; reaching the middle of
leaf or beyond; leaf-cells rarely linear, mostly parenchymatous and 4-sided or
prosenchymatous and 6-sided; small plants; capsules relatively large
1. Amblyslegium
K. Characters not combined as above L
L. Leaf-cells narrowly hnear; leaves broadly ovate or cordate, with reflexed-squar-
rose and subulate-acuminate tips 12. Campylium
L. Leaf-cells and leaves not as above M
M. Small plants; leaves ovate to oblong-lanceolate, long-acuminate, 0.7-1.0 mm long;
cells narrowly prosenchymatous; plants shining 3. Homomjllium
M. Not as above N
N. Slender, dull; leaves spreading, lanceolate to lance-linear; median leaf-cells rhom-
boidal to oblong-hexagonal, 2-4 or rarely 4-8 times as long as broad; oper-
culum not rostrate 2. Amblystegiella
N. Median leaf-cells prolonged-linear, mostly very narrow O
o. Leaves erect-spreading to imbricated, oblong-ovate to rounded, obtuse or apiculate,
often deeply concave and cucullate; costa short and double or none; no para-
phyllia 10. Calliergonella
O. Leaves more or less falcate-secund to circinate, from a mostly narrowed and some-
what decurrent base becoming ovate- to triangular- or cordate-lanceolate, more or
less slenderly acuminate, costa weak, reaching somewhat above the leaf-middle or
even in some cases excurrent; no paraphyllia 8. Drepanocladus
O. Not as above P
P. Paraphyllia numerous; leaves more or less erect, spreading to squarrose-recurved,
from abruptly to shortly acuminate, mostly plicate, concave 16. Hylocomium
P. Characters not combined as above Qi
Q. Leaves falcate-secund or circinate R
Q. Stem-leaves not secund, imbricate, broadly ovate or rounded and with an obtuse
apex; apex finely crenulate; annulus none; cilia three 17. Hypnutn
R. Very robust; stem-leaves turgidly imbricate but strongly rugose and falcate-
secund, lustrous, narrowly lance-acuminate from a broadly oblong base
15. Rhytidium
R. Not as above S
S. Plants distantly and irregularly pinnate; leaves squarrose or spreading and secund;
alar cells little or not at all differentiated 14. Rhytidiadelphus
S. Closely pinnate; leaves circinate secund; alar cells quadrate to rectangular and nu-
merous but not inflated 13. Ctenidium
T. Aquatic or subaquatic on stones and rocks in running water; leaves mostly oblong-
ovate, flaccid, obtuse, rounded, often somewhat secund Il< Hygrohypnum
T. Not as above V
u. Leaf-cells very narrowly prosenchymatous, alar cells mostly not differentiated;
218 American Midland Naturalist Monograph No. 6
leaves oblong to linear, short-pointed, ovate to linear-lanceolate, acute to long-
acuminate or piliferous, not decurrent 20. Isopterygium
U. Leaf-cells often wider, alar cells broader proportionally, hyaline and thin-walled;
leaves broadly lanceolate to oval, more or less long-acuminate, distinctly decur-
rent 21. Plagiothecium
V. Terrestrial or sub-aquatic; leaves mostly squarrose or recurved-squarrose, ovate to
lanceolate, acuminate, decurrent; capsule curved 12. Campylium
V. Not as above w
w. Capsule erect and symmetric or nearly so X
w. Capsule more or less cernuous or horizontal, and unsymmetric bb
X. Leaves closely imbricate when dry, concave 22. Pterygynandrum
X. Leaves erect-spreading or but slightly secund Y
X. Leaves falcate-secund or circinate aa
Y. Alar cells quadrate not inflated Z
Y. Some alar cells inflated and enlarged 19. Stereodon
2. Median leaf-cells about 2-4:1; leaves lanceolate to lance-subulate ....2. Amblystegiclla
Z. Median leaf-cells about 6-9:1, leaves ovate to lance-oblong 5. Platygyrium
AA. Segments attached to a wide basal membrane; cilia rudimentary or none; branches
short, erect or ascending 4. Pylaisia
AA. Not as above 19. Stereodon
BB. Stem-leaves decurrent, plicate, cordate-auriculate at base and abruptly long and
slender acuminate; quadrate alar cells numerous, none inflated 13. Ctenid'tum
BB. Not as above cc
CC. Robust to slender, simple or pinnate, mostly irregularly pinnate; leaves ovate- to
cordate-lanceolate, shortly to slenderly acuminate, generally circinate secund in 2
series; capsule erect to symmetric to cernuous and unsymmetric 19. Stereodon
CC. Not as above DD
DD. Slender; lustrous; leaves ovate to lance-oblong, 0.7-1.0 mm long
3. Homomallium
DD. Robust plants EE
EE. Branching plumosely complanate-pinnate; median leaf-cells about 10-15:1 ....18. Ptilium
EE. Branching irregularly- to bi- or tri-pinnate; stiff; median leaf-cells not more than
10:1 16. Hylocomium
1. Amblystegium Bryologia Europaea*
Autoicous: usually more or less slender, in thin and spreading mats:
stem creeping to ascending or even erect, irregularly to pinnately branched, the
branches mostly more or less erect; stem-leaves similar to branch-leaves, erect-
spreading to squarrose, mostly shortly decurrent, cordate- to ovate-lanceolate,
long-acuminate, rather concave, non-plicate, entire to serrate; costa thin, simple,
reaching to the middle of the leaf or beyond, rarely complete; cells paren-
chymatous; and rectangular to elongate-prosenchymatous and hexagonal, rare-
ly linear, smooth, the alar quadrate to rectangular; the inner perichaetial
leaves erect, broadly lanceolate, mostly costate: seta long, thin, reddish to cas-
taneous, flattened when dry; capsule, from an erect collum, curved to oblong
or cylindric, smooth when dry, constricted below the expanded mouth, annu-
late; peristome-teeth basally confluent, yellow to orange, lance-subulate, bor-
dered, dorsally cross-striate, above pale and papillose, densely trabeculate
* Species with thinner-walled and longer leaf-cells; with costa single and well devel-
oped; and leaves spreading to erect-spreading are treated under the genus Leptodictyiim in
Grout's Moss Flora.
Jennings: Manual of Mosses — 31. Hypnaceae 219
below; inner peristome yellowish, basal membrane high; segments carinate,
entire, or slightly gaping along the keel; cilia complete, nodose, rarely appen-
diculate; lid conic, obtuse to acute; spores small.
A genus of about 50 species occurring mainly in temperate regions, on
various sub-strata; about 20 species in North America; at least five in our
range.
Key to the Species
A. Leaves erect-spreading (widely spreading in A. Juratzkinum); median leaf-cells
about 2-6:! B
A. Leaves mostly loosely and widely to squarrosely spreading; median leaf-cells mostly
4-8:1, rarely 10-15:1 F
B. Cells in middle of leaf about 2-4:1 C
B. Cells in middle of leaf about 4-6(-8):l B
c. Very slender; costa thin, ending near the middle of the leaf I. A. serpens
C. Less slender; costa stronger, almost reaching ap>ex D
D. Stem-leaves ovate-acuminate, acute, slenderly acuminate; entire or nearly so G
D. Stem-leaves ovate-cordate, abruptly narrowed to a rather blunt acumination
3. A. OTthocladon
E. Costa reaching to about three-fourths the length of the leaf 4. A. ]uratzkanum
E.Costa reaching about to the middle of the leaf Campyliutn mdicale, p. 2°' I
F. Slender: median leaf-cells prosenchymatous, hexagonal to hnear, 4-8(-10):l
6. A. Kochii
F. Rather robust: median leaf-cells elongate-prosenchymatous to linear, 5-10(-15):l;
leaves very slenderly acuminate 7. A. nparium
G.Costa reaching apex or almost so; median leaf-cells averaging about 3:1
2. A. varium
G. Costa a little shorter; median leaf-cells averaging about 4:1 5. A. trichopodium
Amblystegium serpens [Linnaeus] Bryologia Europaea
(Hypnum serpens Linnaeus)
Plate XLI
Dull, more or less yellowish-green, very small and slender, forming thin,
soft, densely interwoven mats: stems prostrate, radiculose, irregularly branch-
ing, the branches ascending or spreading or erect; leaves rather crowded, when
moist variously spreading, when dry more or less appressed and imbricate;
stem-leaves lance-ovate to ovate-acuminate, usually long-acuminate, the largest
about 0.8-1.0x0.4-0.5 mm, often much smaller, narrowed and decurrent at
base, denticulate or entire, somewhat concave, the margins plane; costa usually
reaching about to the middle of the leaf or above, often quite faint and indis-
tinct; branch-leaves similar but smaller and narrower, usually more lanceolate;
median leaf-cells oblong- to rhomboid-hexagonal, about 2-4:1, the basal
broader and more rectangular, the alar quadrate to transversely elongate but
not forming a well-defined group, some of the apical considerably longer; peri-
chaetial leaves lanceolate, thin, plicate, up to 1.5 mm long: seta rather slender,
1-3 cm long, reddish, dextrorse; capsule cylindric, the urn below 1.5 mm long,
strongly curved, cemuous, constricted below the mouth when dry; lid convex-
conic, rather obtusely apiculate; peristome rather large for the capsule, typical-
ly hypnaceous, teeth pale castaneous, strongly trabeculate, below dorsally
cross-striolate, the dorsal lamellae projecting to form a more or less crenate
220 American Midland Naturalist Monograph No. 6
hyaline margin; segments about as long as teeth, carinately split, rising from
a basal membrane about two-fifths as high, the cilia 1 (sometimes 2 or 3), as
long as segments, slender, nodose to appendiculate; annulus 2-3-seriate; npores
papillose, when mature brownish or yellowish, medium-walled, about .014-. 018
mm, mature in spring: autoicous.
On bases and roots of trees, decaying logs, soil, rocks, etc., in moist woods;
cosmopolitan; in North America occurring from the Arctic regions to the
Gulf of Mexico.
Fairly common in our region. Now known from Allegheny, Beaver, Cambria, Craw-
ford, Erie, Fayette, McKean, and Washington counties. Specimen figured: Meadow Run
Valley, four miles south of Ohio Pyle, Fayette Co., Sept. 1-3, 1906. O.E.J. &: G.K.J.
2. Amblystegium varium (Hedwig) Lindberg
{Leskea varia Hedwig; Stereodon varius Mitten; Hypniim debile Bridel)
Plate XLI
More or less loosely cespitose, green to light-green above, darker below,
the stems and branches similar but larger than A. serpens; leaves rather close
together, erect- to widely-spreading, the stem-leaves ovate-acuminate, the
largest about 1-1.5 mm x 0.5-0.7 mm, usually long-acuminate, somewhat con-
cave, the margins entire or very slightly denticulate, plane, the base very
slightly decurrent; branch-leaves similar but smaller and more lance-ovate,
usually about 0.6-0.8 x 0.3-0.4 mm; costa strong, more or less colored, usually
yellowish or brownish, reaching usually into the acumen; medium leaf-cells
rhomboid-hexagonal, usually about 2-4:1, somewhat incrassate, rather regularly
arranged, the basal larger and more incrassate, sometimes yellowish, short-
rectangular, the basal marginal distinctly quadrate; inner perichaetial leaves
slenderly lance-triangular, about 1.6 mm long; seta reddish, slender, dextrorse,
varying from 1-2 cm in length; capsule reddish-yellow, about 4-6:1, cylindric,
arcuate, the urn about 1.3 mm long, rather smooth, even when dry and empty;
annulus 2-3-seriate; peristome typically hypnaceous, similar to that of A. ser-
pens, the teeth basally confluent, dorsally cross-striolate below, hyaline-papil-
lose above, strongly and closely trabeculate; the segments about as long, slightly
carinately cleft, the basal membrane about two-fifths as high, the cilia 1 or 2,
nodose to shortly appendiculate; lid conic-acute; spores about .012-.018 mm,
slightly papillose, medium-walled, mature in late spring: autoicoiis.
On bases of trees, soil, rocks, rotting wood, etc., in moist woods; Europe,
and, in North America, from Canada to the Gulf of Mexico. Very common
in our region. Quite variable.
Now known from Allegheny, Beaver, Butler, Crawford, Erie, Fayette, Lawrence, Mc-
Kean, Mercer, Somerset, Washington, and Westmoreland counties. Specimen figured:
Slope of Chestnut Ridge above Hillside, Westmoreland Co., September 16-17, 1909.
O.E.J, and G.K.J.
2a. Amblystegium varium var. ovatum Grout
Slender, more or less julaceous; stem-leaves smaller, 1.2-1.5 mm long,
concave, round-ovate, abruptly shortly acuminate; leaf-cells 2:1, the quadrate
Jennings: Manual of Mosses— 31. Hypnaceae 221
alar cells numerous and extending 1/2 the way up the margin of the body of
the leaf. — St. Louis, Mo., and Bushkill, eastern Penna.
3. Amblystegium orthocladon (Beauvois) Jaeger
{Hypnum orthocladon Beauvois; A. varium var. orthocladon Husnot)
Plates XLI, LXVI
In deference to Dr. Grout's extensive work on this group of mosses I am
following him in placing this species under the genus Hygroamblystegiuni,
which see, page 233.
4. Amblystegium juratzkanum Schimper
Plate XLI
Light yellowish-green, small: stems prostrate, rooting, slender, the branches
irregularly disposed, often ascending to erect, and rising to a height of 1-1.5
cm; leaves when dry usually squarrose-spreading and shriveled, ovate-lanceo-
late, gradually acuminate, about 1 to 1.4 mm long by 0.5 mm wide but quite
variable, almost entire to minutely denticulate, plane, the base narrowed, decu.--
rent and slightly concave; costa yellowish, fairly strong, reaching to i.omswhit
above the middle; median leaf-cells prosenchymatous, linear-hexagonal, about
4-8:1, moderately incrassate, hyaline, the apical similar, the basal tending lo
sub-quadrate or shortly rectangular, the alar forming a rather distinct ^roup,
sub-pellucid, 2-3 times as wide as the median cells, decidedly incrassate, and
always as long or longer than wide; perichaetial leaves 1.5-2 mm long, acumi-
nate, thin, plicate: seta castaneous, smooth, about 2-2.5 cm high, when dry
flattened, flexuous, dextrorse; capsule unsymmetric, cernuous, decidedly arcu-
ate, often describing a half-circle, about 1.5-2.0 mm long, smooth, reddish
when dry and empty much contracted below the wide mouth; peristome lypi-
cally hypnoid; teeth reddish, pellucid, strongly articulate and trabeculate, con-
fluent slightly at base, hyaline-margined, divisural zigzag, dorsal cross-striae
evident; segment as long as the teeth, sub-entire, reddish-yellow, carinate, not
at all or but slightly split, cilia 1-3, of equal length, or some shorter, nodose,
united a little below the middle with the segments to form the basal mem-
brane; spores rather clear, minutely papillose, medium to rather thin-walled,
mature in spring, .010-. 012 mm in diameter.
On moist soil and stones; Europe, Asia, and, in North America, from
Canada to the Gulf of Mexico. Rather common in our region. This species
is intermediate between A. serpens and A. Kochii, but from the former differs
in the more squarrose-spreading leaves, longer alar cells, and stronger costa,
while from the latter it differs mainly in smaller size and longer-pointed leaves.
Allegheny Co.: Douthett, June 5, 1909, Fern Hollow, Pittsburgh, April 25, 1909,
Power's Run, May 7, 1905, Nine-Mile Run, May 17, 1907. O.E.J. ; Moon Township,
May 18, 1902, and Laschell Hollow, June 15, 1902. J.A.S. Crawford Co.: Linesville,
in Pymituning Swamp, June 11-12, 1907, and May 12, 1908. O.E.J, (figured). Erie
Co.: Presque Isle, June 9-11, 1905. O.E.J. Fayette Co.: Ohio Pyle, September 1-3,
1906. O.E.J.
222 American Midland Naturalist Monograph No. 6
5. Amblystegium trichopodium C. Hartman
{Amblyslegium riparium var. trichopodium Bruch and Schimper;
Leptodictyum trichopodium Warnstorf)
Loosely matted; stems 2-4 cm long; branches often ascending; stem-leaves
loosely to widely spreading, ovate to lance-ovate, abruptly long-acuminate,
about 1.5 mm long, plane, sometimes sub-serrulate; costa strong almost reach-
ing apex; median leaf -cells elongated hexagonal to rhomboidal, about 4:1,
rectangular to oblong at base; branch leaves smaller and more lanceolate: seta
about 3 cm long; capsule ovoid to somewhat cylindric, cemuous and curved;
peristome complete; spores ripe in spring.
Frequent across northern North America, but not yet reported for our
region.
6. Amblystegium Kochii Bryologia Europaea
{Leptodictyum trichopodium var. Kochii (Bruch and Schimper) Brotherus)
Plate XLII
Stem prostrate with short erect or ascending branches, the branches not
usually reaching more than 5 or 6 mm long, the general color of the loose
mats being pale green to deep green: stem- and branch-leaves very similar,
spreading rather widely or almost squarrose, erect-spreading when dry, cordate-
ovate, narrowed but scarcely decurrent at base, the apex long and slenderly
acuminate, the leaves sometimes narrower and more lanceolate but always
long-acuminate, usually 1-1.5 mm long, entire to faintly serrulate, plane-
margined; costa medium strong, yellowish, usually about three-fourths as long
as the leaf; median leaf-cells more or less chlorophyllose, thick-walled, rhom-
boid-hexagonal, the ends blunt or parenchymatous, about 4-6:1 or longer, the
basal wider, the alar rectangular to rounded-quadrate, quite densely incras-
sate, hyaline or colored, but scarcely forming distinct alar patches; perichaetiai
leaves up to 2 mm long, lance-linear, long-acuminate: seta about 1.5-3 cm
long, castaneous, flexuous, dextrorse; capsule hypnoid, similar to that of A.
serpens, the urn oblong-cylindric, inclined to cernuous, arcuate, about 2-2.5
mm long, contracted below the mouth when dry; peristome-teeth brownish or
yellowish, hyaline and papillose above, cross-striolate below, hyaline-margined,
strongly and closely trabeculate, the dorsal lamellae and divisural plain; seg-
ments about as long as the teeth, slightly carinate, split, the basal membrane
about two- fifths as high; cilia usually 3, pale, papillose, some of them as lona
as the teeth, nodose; annulus rather large, two-seriate; upper exothecial cells
small, rounded-hexagonal or quadrate, below becoming elongate-hexagonal or
oblong-rectangular; spores in late spring or early summer, somewhat incrassate,
castaneous, minutely roughened, about .015-.018 mm.
On moist earth in swampy or marshy places; Europe, Asia, and probably
throughout temperate North America. Fairly common in our region but in
its smaller sizes difficult to satisfactorily distinguish from A. ]iiratzk<inum.
Grout (Moss Flora) treats this as a variety of A. trichopodium.
Now known from Allegheny, Centre, Crawford, Fayette, Lawrence, McKean, Wash-
ington, and Westmoreland counties. Sfsecimen figured: On root of black walnut, tree,
Hanlin. May 21, 1908. O.E.J.
Jennings: Manual of Mosses — 31. Hypnaceae 223
7. Amblystegium riparium [Linnaeus] Bryologia Europaea
(Hypnum riparium Linnaeus; H. laxi folium Bridel; Stereodon
riparium Mitten; Leptodictyum riparium Warnstorf)
Plate XLII
Loosely cespitose, yellowish-green, the flat tufts soft: stems creeping, sub-
pinnate, the branches usually 2 or 3 cm long, spreading to horizontal, the
stems sometimes floating and reaching a length of 8 or 10 cm; stem-leaves
2-4 mm long, rather widely spreading or almost squarrose both wet and dry,
often somewhat complanate, at tips of branches more or less secund, widely
lance-ovate to oblong-lanceolate, gradually tapering to a fine, flat, non-chan-
neled acumination, shortly decurrent, rounded at base, non-auriculate, some-
what excavate; branch-leaves similar but smaller, all leaves entire and plane-
margined; costa fairly strong, reaching from one-half to three-fourths the
length of the leaf; median leaf-cells linear-rhomboid, prosenchymatous, usual-
ly 8-12(-15):l, thin-walled, chlorophyllose, towards the base lax and sub-
rectangular, at the angles often somewhat larger, rectangular, and sub-inflated,
but not forming very distinct nor hyaline patches: seta usually 1-2 cm long;
capsule rather turgid, oblong-cyilndnc, arcuate, inclined; peristome hypnoiJ
but relatively rather large; teeth dark orange, cilia 2 or 3, appendiculate, about
as long as the entire or slightly parted segments, the basal membrane :-each-
ing to about two-fifths as high as the peristome; annulus 2-3-seriate; exothecial
cells very much smaller at rim, below becoming irregular to rectangular,
medium-walled; spores minutely roughened, .Oil-. 014 mm, mature in spring:
autoicous.
In swamps, springs, brooks, etc., on bases of trees, roots, stones, etc.,
sometimes floating; almost cosmopolitan; in North America ranging from the
Arctic regions to Louisiana and Cuba.
Probably common in our region, in suitable habitats. Now known from Allegheny,
Beaver, Butler, Centre, Crawford, Erie, Indiana, and Lawrence counties. Figured from
specimen collected at Beaver Falls, Beaver Co., O.E.J. , May 14, 1907.
The specimen from Centre Co., in swampy spot in gap of Bald Eagle Mt.,
near Matternville, O.E.J., Sept. 20, 1909, has slenderly acuminate leaves
approaching longijolium (Schultz) Bryologia Europaea.
7a. Amblystegium riparium var. flaccidum (Lesquereux and
James) Renauld and Cardot
Plate XLII
Smaller ?.nd of a much more slender habit; leaves more distant and tend-
ing to narrowly lanceolate and slenderly acuminate.
McKean Co.: East Branch swamp, near Bradford, June 15, 1895. D.A.B. (figured).
2. Amblystegiella Loeske
Autoicous or dioicous: very slender, stems filiform, mostly creeping, irregu-
larly branched; leaves rather laxly disposed, erect-spreading or rarely weakly
secund, lanceolate to lance-subulate from a sometimes somewhat decurrent
224 American Midland Naturalist Monograph No. 6
base, slightly concave, non-plicate, margin plane and entire; costa none or
very short and weak; median leaf-cells rhomboid-hexagonal or oblong-hexag-
onal, 2-4 (-8) :1, the basal rather lax, parenchymatous, the alar quadrate; inner
perichaetial leaves erect, basally sheathing, lanceolate to lance-oblong, long-
acuminate, ecostate, or with the costa ending in or above mid-leaf: seta 5-12
mm long, drying flattened, yellowish-red to castaneous; capsule mostly erect
and symmetric, rarely secund and cemuous, obovate to oblong-cylindric, when
dry and empty constricted below the wide mouth, smooth; annulus present;
peristome-teeth narrowly lance-ovate, basally confluent, yellowish, bordered,
dorsally cross-striate, above pale and papillose, densely trabeculate below; inner
peristome pale or yellow, basal membrane high, segments entire or but slightly
split, cilia rarely 1-3 and complete, mostly solitary and rudimentary or none,
non-appendiculate; lid high-convex, obtuse to acute; spores small.
A genus of 9 species, confined to the Northern Hemisphere, occurring
on trees and rocks; 5 species in North America; two sp>ecies occurring in our
range.
Key to the Species
A. Leaves not narrowed to insertion; inner perichaetial leaves irregularly toothed
-... 1. A. mtnutissima
A. Leaves narrowed to the insertion B
B. On tree bases; leaves lanceolate to linear-lanceolate; inner perichaetial leaves entire
3. A. subtilis
B. On rocks; leaves ovate to lance-ovate; inner perichaetial leaves denticulate above
2. A. confervoides
L Amblystegiella MiNUTissiMA (Sullivant and Lesquereux)
Nichols
(Hypnum minutissimum Sullivant and Lesquereux;
Amblysteginm minutissimum Jaeger)
Minute, pale green: stems prostrate, short, up to about 1 cm long, with
radicles in fascicles, the branches occurring sub pinnately and spreading to
erect; leaves loose, narrowly triangular-lanceolate, broadest and not narrowed
at base, 0.3-0.4 mm long, more or less serrulate, ecostate or very faintlv marked
with striae; leaf -cells large, oblong, about 4-8:1, the marginal alar cells about
2:1; capsule minute, about 0.5 mm long, ovoid, symmetric or slightly curved,
constricted below the mouth and turbinate when dry and empty, thin-walled,
yellowish; seta slender, 4 or 5 mm long; teeth yellowish, hyaline-bordered;
cilia as long as segments and 1 or 2 in number; annulus 2-seriate, persistent;
lid conic, apiculate-rostrate, about one-half to nearly as long as urn; spores
ripe in summer.
On rocks and stones in shaded ravines, said to prefer limestone, from New
Jersey and Pennsylvania westward to Illinois, Ontario, the Rocky Mountains
and British Columbia. Rare in our region.
Rare in our region. Huntingdon Co.: Alexandria. T. C. Porter. (Porter's Cata-
logue ) .
Jennings: Manual of Mosses — 31. Hypnaceae 225
2. Amblystegiella confervoides (Bridel) Loeske
{Hypnum confervoides Bridel; Hypnum conferva Schwaegrichen;
A conferva (Schwaegrichen) Jennings)
Dark green, minute; stems irregularly branching, about 0.5-1.0 cm long;
leaves very small, about 0.1-0.3 mm. long, rather distant, more or less appressed
both wet and dry, entire or almost so, ovate, acuminate, ecostate; leaf-ccHs
irregularly quadrate-rhomboid to oblong-hexagonal, ranging from 1-3:1, some
of them wider transversely, the apical shorter than the median, usually not
more than twice as long as wide, the alar numerous and quadrate to trans-
versely elongate, extending up to the edge of the leaf: capsule cernuous,
reddish-brown, more or less curved, oblong, minute; peristome perfect with
double cilia or sometimes 3; spores mature in summer to autumn: autoicous.
Mainly on shaded ledges of limestone; Europe, Asia, and, in North Ameri-
ca from New Brunswick to southeastern Pennsylvania and westward to the
Rocky Mountains.
Rare in our region. Huntingdon Co.: On limestone rocks, one mile south of
Pennsylvania Furnace, July 13, 1909. O.E.J.
3. Amblystegiella subtilis (Hedwig) Loeske
(Leskea subtilis Hedwig; Amblystegium subtile Bryologia Europaea)
Small, slender mosses in dark green, thin mats; stems about 2-3 cm long,
with numerous but irregular branches; leaves lanceolate to lance-linear, nar-
rowed at the base, and with long and slender acumination, usually not much
over 0.5 mm long, non-decurrent, entire; costa faint or none; median cells
oblong-he.xagonal, 2-3:1, the alar quadrate and often wider than long: inner
perichaetial leaves entire; seta about 1 mm long; capsule slightly longer, oblong-
cylindric, usually erect; annulate; lid convex to conic; cilia rudimentary or none;
spores ripe in late summer or early fall.
Bases of hardwood trees in moist, cool woods. Southeastern Canada to
Minnesota and south to Illinois, Ohio, Pennsylvania, and New Jersey. Not
yet reported from our region.
3. Homo M allium (Schimper) Loeske
Autoicous: slender, rarely somewhat robust, light to brownish or yellowish-
green, more or less shining; stems creeping, divided and irregularly pinnately
branched, with the branches short, erect, and more or less curved; leaves erect-
spreading or secund above, the lower mostly straight, the upper often curved,
concave, non-plicate, lance-ovate to ovate, the base narrowed and but little
decurrent, apex elongate-subulate, the margins plane, entire or serrate at apex;
costa none or short, thin, and double, or longer and sometimes forked; leaf-
cells narrowly prosenchymatous, smooth or with projecting ends, towards the
base shorter and a little wider, the alar numerous, small, quadrate, green,
passing rapidly into the narrower cells above; inner perichaetial leaves almost
sheathing, abruptly acuminate: seta 1-2 cm long, thin, compressed, reddish;
capsule inclined to horizontal, oblong, when dry and empty strongly curved
and narrowly constricted below the mouth; annulus revoluble; peristome-teeth
226 American Midland Naturalist Monograph No. 6
yellow, basally confluent, dorsally cross-striate, bordered, pale and papillose
above, trabeculae numerous and closed below, above strongly projecting; inner
peristome yellowish, papillose, and with a high basal membrane, segments
keeled, split, cilia 2-3, papillose, nodose; spores small; lid shortly and acutely
rostrate.
A genus of about 10 species, occurring on rocks and tree-trunks; 2 species
in North America; 1 species in our region.
1. HoMOMALLiUM ADNATUM (Hedwig) Brotherus
{Hypnum adnatum Hedwig; Amblystegiella adnata Nichols;
Stereodon adnatum Mitten)
Plate XLIII
Widely cespitose in thin, closely adherent mats, pale green, or yellowish-
green, darker below: stems irregularly branching, creeping, the branches close,
short; leaves close, erect-spreading, lanceolate to ovate or oblong, shortly and
widely acuminate to slenderly acuminate, entire or nearly so, concave, ecostate
or slightly bistriate at base, the margins often more or less recurved below, the
leaves 0.6-1.0 mm long; median leaf -cells somewhat pellucid, sub-rhomboidal,
prosenchymatous, about 4-8:1, the apical often shorter, the alar numerous,
smaller, more incrassate and opaque, quadrate and extending along the margin
to one-fourth or one-third the length of the leaf; outer perichaetial broadly
ovate, narrowly gradually acuminate, spreading, the inner oblong, erect, more
abruptly acuminate, dentate, and costate nearly to the middle: seta erect, 1.5-
2 cm long, dextrorse; capsule arcuate, oblong, narrowed to a distinct neck,
cernuous, reddish or yellowish, when dry constricted below the mouth but not
wrinkled; lid paler, acutely conic; annulus present; exothecial cells rounded-
hexagonal near the rim, rectangular below; p>eristome perfect, the teeth promi-
nently and numerously trabeculate, hyaline and papillose apically, hyaline-
margined and dorsally cross-striolate below, the segments entire and very
slightly split, about as long as the teeth, the cilia 1-2, about as long, hyaline
and slightly papillose, the basal membrane about two-fifths as high; spores
rather incrassate, pale-castaneous, papillose, .009-.012 mm, mature in summer.
On rocks and on bases of trees in woods; Asia and from lower Canada to
West Virginia and Texas.
Fairly common in our region. Now known from Allegheny, Butler, Fayette, Hun-
tingdon, McKean, Washington, and Westmoreland counties. Specimen figured: Lime-
stone rocks, Pennsylvania Furnace, Huntingdon Co., July 13, 1909. O.E.J.
4. Pylaisia Bryologia Europaea
( Pylaisiella Kindberg )
Autoicous; slender to rather robust, lustrous, in flat, thin tufts: stem
creeping, long, unsymmetrically pinnate; branches short, ascending to erect,
often curved, in cross section appearing appressed: leaves homogeneous, more
or less imbricate, when moist erect-spreading, often secund, somewhat decur-
rent, concave, non-plicate, ovate to lance-oval, more or less long-acuminate,
mostly plane and entire; costa double, very short or none; leaf-cells linear-
Jennings: Manual of Mosses — 31. Hypnaceae 227
rhombic, smooth, alar numerous, quadrate: seta 1-2 cm long, castaneous, dry-
ing twisted, smooth; capsule erect, symmetric, rarely somewhat curved, oval
to oblong-cyHndric, collum short; annulus small-celled or none; peristome
deeply inserted; teeth lance-subulate, at the ap>ex often irregular and some-
times remaining in the lid or attached to the segments, yellowish, hyaline-
bordered, striate, densely articulate and trabeculate; basal membrane low, seg-
ments narrowly lance-subulate, as long as the teeth or shorter, sometimes two-
cleft, the divisions remaining attached to the teeth; cilia mostly rudimentary;
spores small to large; lid conic to rostrate.
About 15 species, mainly on trees, in temperate regions; about 7 or 8
species in North America, probably four species in our region.
Key to the Species
A. Segments completely adherent to the teeth _ 1. P. intricata
A. Segments free, at least in the upper third B
B. Annulus 2-3-seriate, large-celled; spores about .017-. 024 mm 2. P. Selwynit
B. Annulus uni-seriate;; spores .010-. 016 mm _-C
C. Operculum rostrate; cilia none; spores .009-. 012 mm 3. subdenticulatu
c. Operculum merely conic; cilia single, short or rudimentary; spores .012-. 016 mm
4. P. polyantha
1. Pylaisia intricata (Hedv/ig) Renauld and Cardot
(P. velutina Bryologia Europaea; Pylainella velutina Kindberg,
Pterygynjndrum intricatum Hedwig)
Plate XXXVII
Similar in appearance to P. Selwynii with which it is often confused and
with which it grows, light-green, glossy, in closely entangled mats: branches
ascending or erect, when dry usually more or less hooked at the end; leaves
lanceolate, long-acuminate, usually falcate-secund, about 0.8-1.2 mm long,
about 0.2-0.3 mm wide; leaf-cells similar to those of P. Selwynii but with a
smaller group of incrassate, quadrate, obscure alar cells; median leaf-cells about
6-10:1, sub-vermicular, about .004-. 005 mm wide; costa none: seta straight,
smooth, about 4-5 mm long; capsule ovoid-cylindric, about 2 mm long, erect,
symmetric, castaneous; lid long-conic, about 0.5 mm long; peristome-teeth
closely trabeculate, dorsally distinctly lamellate and with divisural, finely cross-
striate; segments very delicate, split and adherent to the teeth throughout their
whole length, basal membrane indistinct or none; spores densely incrassate,
castaneous-pellucid, finely papillose, in our specimens about .018-. 030 mm in
diameter, mature in fall.
On bases of trees or on stumps or logs, usually in mountainous or hilly
regions; Newfoundland to Ontario and Minnesota, south to North Carolina.
Rare in our region. Butler Co.; On log m woods, Millingar School, Oakland Twp.,
Dec. 2, 1934. Sidney K. Eastwood. McKean, Co.: Bennett Brook, October 23, 1897,
and Limestone Creek, near Bradford, October to December, 1896 (figured). The latter
mixed with Grout's No. 134. North American Musci Pleurocarpi. Washington Co.:
On bark of fallen tree, near Washington, Aug. 4, 1892. A. Lmn and J. S. Simonton.
228 American Midland Naturalist Monograph No. 6
2. Pylaisia Selwynii Kindberg
(P. intncata Bryologia Europaea; Pylaisiella intricata Grout;
Pylaisia Schimperi Cardot)
Plate XXXVII
In thin, densely interwoven mats, dark-green, glossy; rather closely and
regularly pinnate: branches more or less ascending to erect, usually about
3-4 mm long, when dry decidedly curved or hooked at the end; leaves close,
imbricate at the base, prominently falcate-secund, especially so when dry,
lance-ovate, about 0.7-1.0 mm long by about one-third as wide, rather long-
acuminate, sub-serrate to entire, rounded at the base, concave, the margin
plane and non-bordered; costa short and double, or none; median leaf -cells
about 6-10:1, usually .003-.004 mm wide, linear-prosenchymatous, the apical
i;horter and wider, the alar distinct, numerous, quadrate to transversely rectan-
gular, yellowish-incrassate, forming a triangular group extending up along
the leaf-margin to one-third the length of the leaf; perichaetial leaves similar
but longer, up to 2.5 mm, and more slenderly acuminate: seta about 1.5 cm
long, lustrous, red-castaneous, dextrorse above; capsule ovoid-oblong castane-
ous, about 2 mm long, about 2.5:1, erect, symmetric, small-mouthed; peri-
stome-teeth narrowly triangular lanceolate, closely trabeculate, the dorsal
lamellae narrow, numerous, finely cross-striate, pale yellow, bordered up to
two-thirds or three-fourths by the linear, adherent, hyaline and somewhat papil-
lose segments, which are usually united at the tip but widely split below; cilia
none; basal membrane very narrow or none; annulus 2-3-seriate, narrow; cxo-
thecial cells yellowish, somewhat incrassate, irregularly round-hexagonal to
oblong-rhomboidal, below the annulus several series being much smaller nnd
transversely rhomboid-oblong; lid about 0.5 mm high, conic-obtuse, often
somewhat oblique; spores densely chlorophyllose, densely incrassate, castaneous-
pellucid, papillose, about .018-. 025 mm, mature in September or October.
On stones and bark of trees; Siberia, and from Canada to Florida.
Common in our region. Known from Allegheny, Butler, Clearfield, Elk, Erie, Fayette,
Indiana, Lawrence, McKean, Washmgton, and Westmoreland counties. Specimen figured:
Ohio Pyle, Fayette Co., Sept. 1-3, 1906. O.E.J, .ind G.K.J.
3. Pylaisia subdenticula Bryologia Europaea
(Pylaisia denticulata SuUivant)
Intricately cespitose, glossy, yellow-green; stems creeping with erect or
ascending branches about 5-6 mm long; branch-leaves subfalcate, secund, erect-
spreading to imbricate when dry, lance-ovate, entire below, sub-denticulate
above, acuminate, concave, ecostate or faintly costate at base; leaf-cells Hnear-
rhomboidal, 6-8:1, quadrate alar cells numerous, incrassate, extending up the
margin: seta short, erect; capsule oblong, about 2.5-3.5 mm long, about 3-4:1,
erect; lid shortlv rostrate; annulus narrow; peristome-teeth lance-linear, seg-
ments free, basal membrane distinct, cilia none; spores about .008-. 012 mm in
diameter, mature in autumn.
On bases of trees and on rocks, in woods, ranging irregularly from New
Jennings: Manual of Mosses — 31. Hypnaceae 229
York and New Jersey to Illinois and Minnesota, south to North Carohna
and New Mexico. Net yet found in our region.
4. Pylaisia POLYANTHA [Schreber] Bryologia Europaea
(Leskea polyantha Hedwig)
Intricately matted, yellowish-green: stems prostrate, rooting on bark, not
stoloniform, up to 6 or 8 cm long, with numerous erect or ascending, curved
branches about 0.5-1 cm lon^r; branch-leaves erect and secund or poi itin'^ up-
wards, when dry loosely imbricate, small, ovate-lanceolate, rapidly narrowed
into a tapering acumination of about same length as the body of the leaf,
entire, slightly concave, non-plicate, plane-margined, ecostate or with a very
short and faint double or single nerve; median leaf-cells thin-walled, about
6-10:1, the alar few, quadrate, pellucid, rather broad and distmct; stem-leaves
somewhat broader and more abruptly acuminate: seta about 1.5 cm high; cap-
sule oblong-cylindric, about 3.5-4:1, about 2.5 mm long; lid conic, acute, short;
annulus single, narrow; peristome-teeth lance-linear, closely articulate, some-
what granular above, segments about as long as teeth, lance-linear, granulose,
somewhat split when old; cilia single, usually rudimentary; spores mature in
fall or winter, about .01 2-. 01 6 mm.
On tree trunks and in hedges, etc.; Europe, Asia, and in lower Canada and
the northern United States.
Apparently rare in our region. McKean Co.: Bradford. D. A. Burnett, (Porter's
Catalogue ) .
5. PlaTYGYRIUM Bryologia Europaea
Dioicous, medium size, flatly cespitose, green to golden or brownish-green,
lustrous: stem elongate, creeping, ventrally densely radiculose, thickly-leaved
and unsymmetrically pinnate; branches julaceous, mostly short, simple; leaves
imbricate when dry, moist spreading, decurrent, non-plicate, ovate to oblong-
lanceolate, sharply acute, smooth, margins revolute; costa short and double or
none; apical cells rhomboid, linear below, alar quite large, numerous, quadrate;
seta 8-15 mm, sometimes 20 mm, smooth, castaneous; capsule erect, .-.ymmetric
or slightly arcuate, narrowly oblong to almost cylindric; annulus broad, pluri-
seriate, revoluble entire or sometimes in pieces; peristome inserted on the
mouth, double; teeth lance-linear, yellow, broadly bordered, non-striate, trabec-
ulae thickened; basal membrane not prominent, segments narrowly linear, cari-
nately cleft, cilia none; spores .012-018 mm, lid conic, shortly and obliquelv
rostrate. A widely distributed genus of about 4 species; one species in North
America.
1. PlaTYGYRIUM repens [Bridel] Bryologia Europaea
{Pterogonium repens Schwaegrichen; Anomodon repens Fuernrohr;
Cylindrothecium repens DeNotaris; Entodon repens Grout )
Plate XXXVII
Densely but thinly matted, bright to dark green, pinnately branching:
leaves ecostate, crowded, erect-spreading when moist, imbricate when dry, con-
230 American Midland Naturalist Monograph No. 6
cave, subscarious, lustrous, ovate to long-lanceolate, about 0.7-0.9 mm long,
acuminate, the margin entire and recurved below; leaf-cells all medium-walled,
at apex rhomboidal, the median linear-rhomboidal prosenchymatous, about
6-9:1, the alar distinct, quadrate and relatively large, extending up the mar-
gin; inner perichaetial leaves about twice as long as the branch-leaves, ecostate,
more acuminate: seta erect, 10-15 mm long, smooth, lustrous, dark-castaneous,
sinistrorse; urn of capsule about! 1:0-2.5 mm long, erect, symmetric, oblong-
cylindric, castaneous, not narrowed below the mouth when dry; op>erculum
about two-fifths the length of the urn, slenderly and obliquely but bluntly
rostrate; annulus persistent, large, 2-3-seriate, and appearing like modified
upper exothecial cells; peristome -teeth rather deeply inserted, linear-lanceolate,
light yellowish-brown, strongly about 15-18 trabeculate, widely hyaline-bor-
dered, papillose below in irregular and often radiating lines, but not cross-
striate below as in most hypnaceous peristomes, lamellae and divisural line
rather indistinct; segments about two-thirds as long as teeth, linear, narrow,
arising from a very low basal membrane, more or less carinately cleft; cilia
none; exothecial cells quadrate to irregular or oblong-hexagonal, yellowish;
spores about .014-. 018 mm, yellowish, minutely roughened, medium-walled,
mature in autumn; gemmae often abundant in the axils of the upper leaves.
On bark at base of trees, on decaying logs, stumps, and in woods; widely
distributed in the Northern Hemisphere; in North America from New Bruns-
wick to the Pacific and south to the Gulf of Mexico.
Very common in our region at lower altitudes, rarely found in the mountains or
plateau uplands. Known from the following counties: Allegheny (more than 50 collec-
tions), Armstrong, Beaver, Butler, Crawford, Erie, Fayette, Greene, Somerset, Washing-
ton, and Westmoreland. Sf>ecimens figured: On rotten log, oak woods, Keown, Alle-
gheny Co., Nov. 14, 1909. O.E.J.
6. Hygroamblystegium Loeske
Autoicous or dioicous: aquatic or sub-aquatic; slender to quite robust,
mostly stiffly cespitose, dark-green to blackish-green, dull: stem more or less
elongate, mostly floating, rarely more or less erect, mostly rather regularly
pinnate, with forward-directed, rarely erect, mostly simple branches; leaves
close, spreading to secund, concave, non-plicate, not at all or but slightly
decurrent, rarely long-decurrent, mostly ovate to oblong-lanceolate, long-acu-
minate, margins plane, entire or remotely indistinctly denticulate; costa strong,
short or percurrent, sometimes thickly excurrent; cells green, prosenchymatous,
hexagonal, 2-4(-6) :1, alar cells more or less plainly differentiated; perichaetial
leaves elongate-lanceolate, costa complete or sub-percurrent: seta elongate,
castaneous; capsule inclined to horizontal, early symmetric or somewhat dor-
sally gibbous, oblong-cylindric, later more or less arcuate, when dry and empty
constricted below the mouth; peristome-teeth dark-yellow to orange, more or
less basally confluent, lance-subulate, broadly bordered, dorsally cross-striate,
apically pale and papillose, the margin step-like, the trabeculae strongly pro-
jecting; inner peristome yellow, finely papillose, with high basal membrane,
segments mostly carinately split, cilia complete, nodose to short-appendiculate;
lid high-convex and apiculate or acute; spores small.
Jennings: Manual of Mosses — 31. Hypnaceae 231
A genus of about 10 species, in very damp or frequently submerged places
or in water, mostly in temperate or cooler regions; 5 species occur in North
America; at least 5 species in our region.
Key to the Species
A. Leaves non-decurrent, entire or indistinctly and remotely serrate B
A. Leaves mostly decurrent, mostly with small but distinct teeth C
B. Stem-leaves lance-oblong to oblong-ovate with rather obtuse point 1. H. fluviatile
B. Stem-leaves triangular lanceolate to triangular ovate, sub-obtuse to acuminate D
C. Costa sub-percurrent to percurrent; leaves with decurrent auricles of inflated cells
4. H. filicimim
c:. Costa excurrent; leaves non-decurrent, non-auriculate, not basally excavate
3. H. Jioterophilum
D. Stem-leaves more or less acuminate 2. H. irriguum {^ tenax)
D. Srem-leaves sub-obtuse to acute 3. H. orthocladon
1. Hygroamblystegium fluviatile [Swartz] Loeske
(Amblystegium fluviatile Bryologia Europaea; Hypnum fluviatile Swartz)
Plate XLIII
Robust, aquatic, floating in flat and elongated tufts, soft, olive- to dark-
green, devoid of leaves below: stems with few branchlets, long, the branchlets
more or less parallel and scarcely pinnate; leaves oblong-lanceolate to oblong-
ovate, not markedly narrowed below, rather remote, erect-spreading, especially
when dry, non-decurrent, gradually tapering to a short, blunt point, entire or
very faintly serrulate, very concave, the margins more or less recurved at base;
costa thick and strong, yellowish, ending in the apex; median leaf-cells loose,
he.xagonal-rhomboid, about 3-6:1, the basal cells rectangular, pellucid, some-
times somewhat opaque, strongly incrassate, not forming auricles, sometimes
quite orange; perichaetial leaves erect, strongly costate: seta about 1.5 cm long,
castaneous, dextrorse; capsules about 2.5 mm long, oblong-cylindric, sub-erect,
sub-arcuate, rather thick-walled, yellowish-brown, when dry and empty strongly
arcuate and constricted below the mouth; below the 2-3-seriate annulus ihe
exothecial cells small and rounded-quadrate; peristome slightly irserted, teeth
strongly confluent at base, dorsally cross-striolate, brownish below, apically
hyaline and papillose; segments about as long as teeth, carinately spht, the
three nodose cilia about as long, the basal membrane about two-fifths to one-
half as high as teeth; spores medium-walled, minutely papillose, brownish,
about .016-.019 mm, mature in early summer.
On earth and on rocks and stones in running water, usually m non-cal-
careous districts; Europe, and, in North America, from Newfoundland to
New Jersey and westward to the Mississippi. Most of our specimens seem to
belong to forma brevijoliuvi Boulay, with concave, oblong-ovate leaves, sub-
obtuse at apex, and with colored opaque basal cells.
Now known from Allegheny, Beaver, Butler, Cameron, Fayette, McKean, and West-
moreland counties, all in the non-glaciated area. Specimen figured: Shades Ravine, Black-
burn, Allegheny Co.. June H, 1908. O.E.J.
232 American Midland Naturalist Monograph No. 6
2. Hygroamblystegium irriguum (Wilson) Loeske
(Hypnum irrimium Wilson; Amblystegium irriguum Bryologia Europaea; Hypnum
tenax Hedwig — fid. Cardot; Hygroamblystegium tenax Jennings)
Plate XLIII
Dark green, aquatic, cespitose: stems rigid, irregularly pinnate, long,
denuded at the base, usually with a few paraphyllia at the nodes; stem-leaves
ovate to widely triangular lanceolate to triangular — about 1-1.5 mm long, gradu-
ally acuminate, acute or sub-acute, narrowed at the base, sub-decurrent, spread-
ing and sub-secund, or on the longer branches erect-spreading, entire to
sub-serrulate, plane-margined; branch-leaves narrower and tending to lance-
ovate; costa thick and wide, yellowish-brown, narrowing and becoming indis-
tinct in the acumen but often reaching the apex; median-leaf-cells hexagonal-
rhomboid, about 3-6:1, incrassate, often sub-opaque, smaller in the apex, at the
base one or two rows usually somewhat enlarged, rectangular, incrassato, often
colored, a few rows above these shorter, quadrate, but no distinct auricles
being formed: seta about 1.5-2.5 cm long, smooth, castaneous, dextrorse; cap-
sule oblong-cylindric, the urn 2-2.5 mm long, 5ub-cernuous and sub-arcuate
before ripening to strongly arcuate when dry, smooth, constricted bslow the
mouth, bro'ivnish; annulus 3-seriate; lid convex-conic, apiculate; peristome-teeth
basally confluent, orange-pellucid and dorsally cross-striolate below, bordered,
strongly tra'reculate; the segments slightly shorter than the teeth, carinately
split but scarcely gaping, yellowish-hyaline, the 3 cilia nodose, hyaline-papillose,
about as lone, as the segments, the basal membrane about two-fifths as high;
exotheciai cells small and rounded, hexagonal to transversely rounded at rim
but soon becoming rather elongate oblong-hexagonal or rectangular below;
spores mature in late spring or early summer, brownish, medium-walled, papil-
lose. .016-.019 znm.
Or\ stones and earth in very wet situations or in water, usually in non-
calcareous districts; Europe, Asia, northern Africa, and, in North America,
from Ont.irio to California, south to Georgia and Arkansas.
Now known from Allegheny, Bedford, Butler, Cambria (Porter), Crawford, Erie,
Fayette, Huntingdon (Porter), Washington, and Westmoreland counties, and apparently
avoiding the upland plateau region. Specimen figured: Hillside, Westmoreland Co., May
23, 1908. O.E.J.
2a. Hygroamblystegium irriguum var. spinifolium (Schimper) Grout
(H. falldx var. spinifolium Warnstorf; Amblystegium irriguum var. spinifolium
Schimper; A. fallax var. spinifolium Limpricht)
This variety differs from the species in being more robust, with longer
stems, longer and narrower leaves, the leaves reaching nearly 2 mm in length
and with a strongly excurrent and stout costa: Grout states the upper leaf-cells
to be about 6-8:1, and the basal cells more lax.
Usually in and around calcareous springs and probably distributed mainly
as is the .species. Ohio, Pennsylvania, New Jersey, and Ontario.
Butler Co.: On wet cliffs, Winfield Junction. C.M.B. June 8, 1940. Crawford
Co.: Pymatuning Sv-famp, Linesville, May 12, 1908. O.E.J.
Jennings: Manual of Mosses — 31. Hypnaceae 233
3. Hygroamblystegium orthocladon (P.B.) Grout
(Hypnum orthocladon P. Beauvois; Amblystegium orthocladon (P.B.) Jaeger)
Plates XLI, LXVI
This plant is regarded by Grout after extensive study as belonging to
Hygroamblystegium, intermediate between H. irrtgutim and the wider and
more obtuse-leaved variety brevifolium of H. fluviatde. In deference to Dr.
Grout's extensive studies in this group I am here placing it under Hygro-
amblystegium.
Rather dark green, sometimes olive-green, rather stiff when dry, commonly
tufted; stems irregularly branched, the branches of plants in the denser tufts
often erect and sometimes 2 cm tall, but usually less than 1.5 cm long; leaves
up to 1 mm long, broadly cordate-ovate, usually rapidly and uniformly nar-
rowed to an acute or sub-obtuse apex, rounded to a narrower base, slightly
concave, slightly decurrent, widely spreading both wet and dry, the margins
plane and entire; branch-leaves smaller, narrower, and more acute; costa strong,
wide at base, yellowish, usually extending up into the apex; leaf-cells sub-
incrassate, the median oblong-rhomboidal to elongate-hexagonal, Vv'ith rounded
ends, .010-. 020 mm long, about 2-4 (-6) :1, the apical similar, th? mec'ian basal
oblong-rectangular, the cells of the angles somewhat wider, varying to short-
rectangular or quaorate, incrassate, often op::que or colored: seta ur.uaHy about
1-1.5 cm long but occasionally reaching 3-4 cm; castaneoas; capsule ca3taneous,
oblona-c/lindric, arcuate, cernuous, constricted below the rim -when diy, nar-
rowed at base to a distinct neck, the urn about 1.6-1.9 (to 3) mm long; peri-
stome typically hypnoid; basal membrane high, cilia about equalling seginents;
operculum short-conic; spores mature in spring, somewhat incrassate, smoothish,
about .010-018 mm.
On stones, rotten wood, bases of trees, etc., along or in brook; or in moist
situations in woods; Europe, and probably widely ranging through more ele-
vated parts of eastern United States. Grout {Moss Flora') treats it as a sub-
species of Hygroamblystegium irriguum. The specimen figured from \Vash-
ington County is unusually large.
Now known from the following countias: Allegheny, Butler, Crawford, Favette,
Greene, McKean, Warren, Washington, and Westmoreland. Specimens figured: Cheat
Haven, Fayette Co., Sept. 26, 1910. O.E.J, dc G.K.J. (Plate XLI).; On stonos in Calcar
Spring, Snake woods, near Washington, Washington Co., Dec. 12, 1897, A. Linn and
J. S. Simonton (Plate LXVI).
4. Hygroamblystegium filicinu.m [Linnaeus'] Loeske*
(Amblystegium filiciniirn DeNotaris; Stereodon filiciniif Mitten; Hypnum
compressum Bridel, Cratoiieuron flic'inum Roth)
Variable, forming loose to dense tufts, rather rigid, bright or golden yel-
low: stems usually densely brownish tomcntose, especially en the prostrate or
procumbent forms, rather regularly pinnately branched, with usually numer-
ous oval to lanceolate, laciniate oaraphyllia; brarxhes slend:r, short, stiff, non-
tadiculose, with few or no paraphvllia, usually hooked at the ape."-:; stem-leaves
* Grout {Moss Flora) treats this as a species of Critoncuron.
234 American Midland Naturalist Monograph No. 6
cordate-triangular, finely and gradually acuminate, varying from erect-spread-
ing to sub-secund; branch-leaves rather narrower, more usually strongly falcate-
secund; all leaves rigid, altered but little in drying, not plicate, markedly decur-
rent, the base cordate and narrowed, the margin plane or recurved at the base
and closely and finely serrulate from base to apex; costa strong, usually ending
in the apex; median leaf-cells elliptic-hexagonal to elongate rectangular, mostly
about 3-6:1, usually obtuse at the ends, the alar abruptly inflated, hyaline or
colored, forming well-defined auricles of sub-rectangular cells, these cells reach-
ing to the base of the costa or nearly so; perichaetial leaves erect, strongly
costate but scarcely plicate, denticulate: seta long, reddish, flexuous, up to 3-5
cm long, flattened and twisted; capsule reddish, sub-cylindric, rather turgid,
arcuate, when dry and empty constricted below the mouth and more or less
sulcate; lid conic, acute, or apiculate; peristome hypnoid, the segments more or
less cleft cn.inatcly, cilia 2 or 3, nearly as long as the segments and teeth;
annulus simple, narrow; spores mature in spring.
On earth, stones, etc., in or near springs, streams, or swamps, principally in
calcareous districts; Europe, Asia, northern Africa, and, in North America,
from the Arctic regions south to the southern part of the United States.
Rare in our region. Huntingdon Co.: Spruce Creek. T. C. Porter. (Porter's Cat-
alogue).
5. Hygroamblystegium noterophilum (Sullivant) Warnstorf
(Hypntim noterophilum Sullivant; Amblystegium noterophilum Holzinger;
Hypnum irriguum spinifoliiim Lesquereux and James)
Larger than fluviatile, dull dark green in older parts, rather stiff, irregularly
branching; stem with strong central strand and thick-walled cortical cells; leaves
entire, lance-oblong when aquatic, when emersed ovate and up to 2.5 mm long;
basal and costal part of leaf bi-stratose; capsule highly stomatiferous; spores
mature in early summer.
In springs and running water in calcareous regions. Ontario and northern
United States, the type locality in Franklin County, Pennsylvania.
7. SCIAROMIUM Mitten
Mostly dioicous: more or less robust, stiff, cespitose, dull, dark green
to blackish: stem long, floating, sparsely radiculose, with irregularly and
sometimes rather fasciculately arranged branches mostly directed forwards
and mostly long and simple; leaves close, spreading to secund, concave-
carinate, non-plicate, not at all or but slightly decurrent, ovate to lance-
oblong, sub-acute to acuminate, plane-margined, mostly entire, broadly and
thickly bordered; costa strong, ending apically in the border or excurrent;
median leaf-cells chlorophyllose, strongly incrassate, rather opaque, prosen-
chymatous-hexagonal, 2-4 (-6) :1, the basal cells more lax, the alar somewhat
differentiated, the marginal slender, strongly incrassate, hyaline, in several
layers; costa ending in the border at the apex: seta 1-3 cm long, castaneous
below, more yellowish above; capsue inclined, unsymmetric, oblong, when dry
somewhat constricted below the mouth, annulate; peristome-teeth yellow, bor-
Jennings: Manual of Mosses — 31. Hypnaceae 23'>
dered, cross-striate, apically pale and papillose, the margin step-like, trabeculae
numerous; inner peristome yellowish, with high basal membrane, segments
keeled, narrowly carinately split, cilia 1-3, shorter than the segments and
nodose; lid high-convex, apiculate; spores small.
A genus mainly confined to South America and embracing about 20
species; only 1 species occurs in North America and this occurs rarely in our
region.
1. SciAROMiUM Lescurii (Sullivant) Brotherus
(Hypnum Lescurii Sullivant; Amblysiegium Lescuni Jaeger)
Plate XLIII
Loosely cespitose, dull, dark green to blackish-green: stems closely and un-
equally branched, the branches as described for th: genus, but often with
short branchlets, 1-1.5 cm long, more or less erect, and pinnately disposed;
leaves of the stem thick, rather opaque, erect-spreading, entire below to sub-
serrulate all around, broadly ovate-cordate to oblong-ovate, 1-1.3 mm long,
abruptly short-acuminate, the branch-leaves similar but more lance-ovate;
median leaf-cells prosenchymatous, hexagonal to oblong, about 1-3:1, nor
much differentiated except for the yellowish or cactancous border which is
composed of 4 or 5 rows of linear, prosenchymatous, fle.xuous, highly incras-
sate cells, the border cells in the alar region becoming short and rectangular or
obliquely quadrilateral; costa very strong, castaneous or yellowish, merging
at apex into the border: seta 1-3 cm long, reddish; capsule reddish, short-
necked, the urn about 2.5 mm long, oblong, cernuous, somewhat arcuate; the
teeth confluent at base, hyaine-papillose above, yellowish below, dorsally lam-
ellate and cross-striate, numerously trabeculate, hyaline-margined; segments
yellowish, carinately split and about as long as the teeth, the basal membrane
about two-fifths as high; cilia 3 (or 4), pale, papillose, nearly as long as seg-
ments; annulus compound; spores mature in late spring or early summer,
castaneous, medium-walled, smoothish, about .012-.015 mm; lid conic-apiculate.
On stones and rocks in streams, usually in mountainous or hilly regions;
occurring from New England to Ontario, Alabama p.nd Georgia.
Rare in our region. Fayette Co.: Ohio Pyle, May 30-31. 1908. 6.E.J. (figured).
8. Drepanocladus (C Mueller) Roth
Dioicous, rarely autoicous: mostly robust, often densely cespitose, green to
yellowish or brownish, lustrous: stem procumbent to erect, often floating, vari-
ously pinnate, the ends of the shoots usually circinate: leaves usually more or
less circinate-secund, rarely erect to squarrose, more or less concave, from a
mostly narrowed and decurrent base ovate- to triangular- or cordate-lanceolate,
acute to prolonged acuminate, entire or serrulate; costa mostly simple and
thin, ending usually above the middle of the leaf, sometimes strong and p>er-
current or even excurrent; leaf-cells mostly long-linear, smooth, in the more or
less excavate angles parenchymatous, thin-walled, hyaline and inflated or thick-
walled and colored, usually forming a well-defined group sometimes reaching
to the costa; inner p)erichaetial erect, mostly plicate, elongate-subulate: seta
236 American Midland Naturalist Monograph No. 6
long to very long; capsule inclined to horizontal, cylindric, arcuate, when dry
constricted below the mouth, smooth, annulate; lid convex, apiculate.
A difficult genus of over 40 species of water-mosses, quite largely swamp-
mosses, — often formmg quite large masses of vegetation, — almost exclusively
confined to temperate and cold regions; about 22 species occur in North Amer-
ica, perhaps the following 8 to be included in our list. Species not well-
defined and extremely variable. See Grout's Moss Flora for descriptions of
many forms and varieties.
Key to the Species
A. Stem without central strand or inflated cuticular cells; leaves without inflated alar
cells 2. D. vernicosus
A. Stems with central strand B
B. Stem in cross-section displaying cortical layer of enlarged, thin-walled, hyaline
cells E
B. Stem without enlarged cuticular cells C
C. Leaves usually entire ;— F
C. Leaves serrulate, ends of stems and branches hooked D
D. Costa usually less than three-fourths length of leaf; alar group of cells not reach-
ing over to the costa 7. D. ftuitans
D. Costa extending well up to the apex of leaf; alar group of cells large, excavate,
and extending over to the costa 8. D. exannulatus
E. I eaves strongly plicate, 3-5 mm long I. D. uncinatus
E. Leaves not plicate, about 2 mm long 3. D. intermedius
F. Ends of stems and branches more or less strongly hooked (See also forms of
D. Kneiffti) G
F. Ends of stems and branches not or but moderately hooked (certain forms strongly
hooked); leaves broad-lanceolate to ovate-oblong 5. D. Kneiffti
G. Stem-leaves falcate-secund; enlarged, inflated, thin-walled, auricular cells usually
reaching nearly to the costa; costa usually about to middle of leaf 4. D. adur.cus
G. Enlarged, medium thick-walled, auricular cells not reaching costa; costa usually
extending into the acumen 6. D. Sendlneri
1. Drepanocladus uncinatus [Hedwig] Warnstorf
{Hypnum uncinatum Hedwig; Amblystegium aduncum Lindberg)
Rather slender and loosely interlaced, pale green or golden green: stems
distantly and irregularly pinnately branched, 2-10 cm long, in cross-section
showing a layer of large hyaline cortical cells and a central strand; leaves 3-5
mm long, rather crowded, regularly falcate to sub-circinate, little altered when
dry, spirally fllcxuous at the points in the younger and softer branches, narrov/-
ly elongate-lanceolate, strongly plicate both wet and dry, gradually very long
and slenderly acuminate, usually denticulate above, texture very thin; costa
narrow, about 0.30-0.35 mm at base, extending well into the acumen; leaf-cells
very long, linear- flexuous, thin-walled, pointed, uniform to the base and apex,
the alar forming a rather small and indistinct group of slightly enlarged and
slightly mflated cells, and extending decurrently below and marginally a short
distance above; perichaetial leaves erect, straight, long, plicate, sheathing:
seta variable, but usually 2-3 cm high; capsule cylindric, arcuate, orange-red,
darker when old, when dry and empty somewhat constricted below the mouth,
Jennings: Manual of Mosses — 31. Hypnaceae 237
smooth; annulus broad, 3-seriate; lid high-convex, conic-acuminate; peristome
hypnoid, teeth orange-ycIIow, paler above, segments somewhat carinately split,
cilia 2, slender and about as long as segments; spores mature in late spring or
early summer.
On earth, decaying wood, stones, etc., bordering streams or in wet situa-
tions in the shade, mainly in hilly or mountainous regions almost the world
over; in North America from Arctic regions south to the Gulf States. Ex-
tremely variable with many named varieties.
Rare in our region. Cambria Co.: T. P. James. (Porter's Catalogue). Hunting-
don Co.: T. C. Porter. (Porter's Catalogue).
2. Drepanocladus vernicosus (Lindberg) Warnstorf
{Hypnum vernicosum Lindberg)
Deeply tufted, yellowish-green, darker or brownish below; stems slender,
more or less regularly pinnate, hooked at the ends; central strand none but
outer cuticular cells small and somewhat incrassate; stem-leaves plicate, neither
auricled nor decurrent lanceolate to ovate-lanceolate, strongly circinate or fal-
cate-secund, gradually long acuminate and entire; costa usually reaching beyond
the middle of leaf; median leaf-cells linear-flexuous, about 10-12:1; about 2
or 3 rows of the basal cells incrassate, reddish or brownish; alar not differen-
tiated: inner perichaetial leaves erect, acuminate, costa long: seta 4-5 cm long,
reddish; capsule castaneous, cylindric-ovoid, cemuous, curved, narrowed under
mouth when dry; lid conic-apiculate; annulus 3-seriate; spores in May or June.
Rarely fruiting.
In usually non-calcareous regions in wet places and swamps. Eurasia, and
from Canada south to New Jersey, Pennsylvania, and Ohio.
3. Drepanocladus intermedius (Lindberg) Warnstorf
{Hypnum intermedium Lindberg)
Yellowish-green to brownish tufts; stems slender, showing central strand
and inflated hyaline cuticular cells, ascending to erect, 8-15 cm long, irregular-
ly pinnate; leaves non-plicate, non-decurrent, from an ovate to oblong base
long-acuminate, about 2 mm long, entire; costa thin, disappearing above the
middle; leaf cells very narrow, 2-4 in the angles thin-walled and slightly
enlarged, forming a faint group; inner perichaetial leaves erect, long and slen-
derly acuminate, delicately costate; seta 4-5 cm long, reddish; spores ripe in
May or June.
In bogs, marshy places, or on wet rocks, often forming masses of \'egeta-
tion. Possibly to be expected in the northern part of our region.
4. Drepanocladus aduncus (Hedwig) Warnstorf
(Hypnum aduncum Hedwig. not Linnaeus)
Plate LXVII
An extremely variable moss of which many varieties and forms have been
named. Yellowish- green mats (or tufts), brownish below; stems 2-3 cm long
238 American Midland Naturalist Monograph No. 6
or in forms up to 30 or 40 cm, bushy branched with erect branches or floating;
stem-leaves triangular ovate to lance-ovate and acuminate or lanceolate and
very slenderly acuminate when in water, often secund or falcate at ends of
branches, non-plicate, entire, the large inflated, thin-walled cells at the exca-
vated auricles decurrent, usually hyaline and reaching well towards the costa;
costa disappearing at or above the middle; m.edian; median leaf-cells linear-
flexuous (or wider in some varieties) ; branch-leaves smaller and narrower, and
often more falcate or even falcate-secund; seta about 2-4 cm long, rather slen-
der; capsule about 2-2.5 mm long, curved, cernuous; spores in late spring or
early summer; lid conic-apiculate; annulate. Not often found in fruit.
Eurasia, northern Africa, New Zealand, and widely distributed in colder
and temperate parts of North America.
Crawford Co.: Pymatuning Swamp, Linesville. O.E.J. Aug. 19, 1904 (figured).
Erie Co.: Numerous collections on Presque Isle by Nelle Ammons. Summer of 1935.
Lawrence Co.: In moist field with Gentiana crinita, Frew Mill Road, e. of New Castle.
C.M.B. Sept. 18, 1948. Warren Co.: Iron Spring bog, Columbus. C.M.B. Sept. 1,
1948. Westmoreland Co.: On rock in stream, Tannery Hollow, Chestnut Ridge.
C.M.B. Apr. 30, 1944. Elev. 1700 ft.
5. Drepanocladus Kneiffii (Schimper) Warnstorf
{Hypnum aduncum var. Kneiffii Schimper; Amblyslegium Kneiffii
Bryologia Europaea)
Stems slender, long, flexuous, prostrate or ascending, more or less pinnate-
ly branched, the cross-section showing a central strand, but not a distinct corti-
cal layer of enlarged hyaline cells; leaves distant, broadly lanceolate to ovate-
oblong, costate to the middle at least, usually not secund nor falcate except
sometimes at the end of the branches, the acumen flat and entire, the lower
leaves usually shorter and wider; basal leaf-ceils much as in D. aduncus, the
alar large, inflated, and extending about half-way to the costa. Closely related
on the whole to D. aduncus, and by some bryologists regarded as merely a
variety of that species.
Along streams and ditches, about as widely distributed as is D. aduncus.
Butler Co.: In swamp among grasses, 1 m. west of West Liberty, June 28, 1941.
Charles M. Boardman. Erie Co.: Cranberry Pond, Presque Isle, Erie, July 30, 1935,
Nelle Ammons; on base of Alnus, May 8-9, 1906. O.E.J.
6. Drepanocladus Sendtneri (Schimper) Warnstorf
{Hypnum Sendtneri Schimper)
Rather strong plants in yellowish tufts, brownish below; stems 10-15 cm
long, erect, sparsely irregularly pinnately divided; leaves close, strongly falcate
or circinate, hooked at the ends of stems and branches, lanceolate to lance-
ovate, about 10-11 mm long, long-acuminate, entire, non-plicate, short-decur-
rent; costa strong and wide, extending into the leaf-tip; median leaf-cells
narrowly linear to somewhat oblong, about 6-10:1, basal larger, incrassate, the
alar forming a well-defined group of colored, incrassate, excavate, inflated
cells; dioicous; seta 3-4 cm, reddish, slender; capsule resembling that of
Jennings: Manual of Mosses — 31. Hypnaceae 239
aduncus. Grout suggests that Sendtneri is a calcicolus form of adiincus, which
it parallels in many of its forms.
In wet Hmy bogs, swamps, etc. Eurasia, and, in North America extending
from Canada south to North CaroHna, and in the west to California.
7. Drepanocladus fluitans [Linnaeus] Wamstorf
{Hypnum ftuitans Linnaeus; Amblystegium fiuilans DeNotaris)
Plate LXIX
Loosely and softly cespitose, yellowish to dark brown, irregularly to regu-
larly pinnately branched: leaves more or less secund or falcate, narrowly lan-
ceolate to oblong-lanceolate, tapering gradually into a very slender flexuose
acumination, the branch-leaves somewhat narrower than the stem-leaves but
quite similar, all denticulate, excavate at the base, sometimes reaching a length
of 4 mm, decurrent; costa not markedly wide, reaching into the ap>ex or at least
nearly so; leaf-cells about 20-30:1, long, reaching to .100 mm or more,
pointed, narrow, somewhat incrassate, the alar, hyalme or colored and often
forming more or less distinct auricles reaching sometimes to the base of the
costa, somewhat inflated: seta red, long, up to 5 or 6 cm or sometimes much
longer, flexuous, strongly dextrorse; capsule more or less inclined, curved,
rather thin-walled, with a distinct collum, about 3-4:1; lid high-convex, bluntly
apiculate; peristome-teeth rather short, segments rarely carinately split, cilia
usually 1 or 2, usually considerably shorter than the segments: annulus none;
spores mature in early summer. Very variable and split up into many forms
and varieties by various authors.
In ditches, swamps, bogs, stagnant pools, among willows, cat-tails, etc.,
often immersed or floating, almost cosmopolitan in temperate and cold regions;
in North America, throughout Canada and the northern United States.
Centre Co.: Bear Meadows. T. C. Porter. (Porter's Catalogue). Erie Co.: En-
tirely submerged in lily pond. Presque Isle. Nelle Ammons. July 13, 1935. McKean
Cc: In stagnant water. West Branch Swamp, Bradford. Among willows. D.A.B. May
26, 1895 (figured) and June 9, 1895. Somerset Co.: Pleasant Union, Sept. 7, 1942;
and Mt. Davis Fire Tower, eiev. 3210 ft., Sept. 21, 1947. C.M.B.
8. Drepanocladus exannulatus (Guembel) Wamstorf
{Hypnum exannulatum Bryologia Europaea; Amblystegium
exannulatus DeNotaris )
Plate XLIV
Typically more rigid, compact, and more completely pinnate than D.
fluitans. the leaves more falcate, usually serrulate, frequently striate, especially
when dry: the costa reaching well towards the apex and rather stronger than
in D. fluitans, biconvex; the alar cells hyaline and much enlarged, forming
an excavate and well defined excavate auricle extending across to the costa.
In our region the specimens show the following characteristics: yellowish-
brown, floating, the stems up to 8 or 10 cm long, the tips of stems and
branches hooked; leaves rather remote, reaching 4 mm long, irregularly and
widely spreading, not definitely circinate or secund, except at the tips of stems
240 American Midland Naturalist Monograph No. 6
and branches, slenderly acuminate into a sub-channeled acumen, entire, the
base rounded to somewhat excavate and decurrent auricles, so that the inser-
tion is more or less of a semi-circle; median leaf-cells linear, rather incrassate>
about 10-15:1, reaching 0.3 mm or even longer, towards the base rapidly be-
coming shorter and quickly passing into large, hyaline, oblong, much-inflated
cells, thus forming a distinct patch reaching to the costa and, below, passing
abruptly into the narrowly linear epidermal cells of the stem; in cross-section
the stem may be seen to have the 3 or 4 outer layers small and very thick-
walled.
In bogs and wet places, usually in cool or alpine regions; northern and
temperate Europe and Asia and, in North America, from Greenland to Alaska
south to the northern United States.
Crawford Co.: In pools, Pymatuning Swamp, Linesville, August 19, 1904. Sterile.
O.E.J, (figured). Elk Co.: Midmont Swamp. Elev. 1940. C.M.B. July 1, 1948. Erie
Co.: Among cattails in swamp. Presque Isle, Nelle Ammons. July 22, 1933. McKp.an
Co.: Cathrine Swamp. C.M.B. Sept. 2. 1948.
9. Calliergon (SuIIivant) Kindberg
Mostly dioicous: more or less robust, stiffly and loosely cespitose, gi-eenish
to brownish or yellowish, rather lustrous; stem long, in v/ater and in deep
swamps not bearing rhizoids but assuming a more or less erect habit, in dry
places procumbent and bearing rhizoids, irregularly to regularly pinnatelv
branched; stem-leaves large, erect-spreading to imbricate, concave, rarely some-
what plicate, ovate to oblong or almost circular, the apex broadly rounded to
cucullate, the margin plane and entire or rarely so.mewhat revolute below;
costa mostly strong and almost complete, sometimes indistinctly forked at the
end; leaf-cells elongate, linear-hexagonal, shorter below, the alar forming! a
distinct group of large, quadrate, rectangular, and polygonal cells, at first thin
and hyaline but later colored and incrassate, the alar portion of the leaf exca-
vate; branch-leaves smaller, narrower, the apex often canaliculate; the inner
perichaetial leaves erect, more or less long-acuminate, mostly non-plicate, with
a simple costa: seta mostly very long, drying flat, red to castaneous; capsule
inclined to horizontal, thickly oblong to oblong-cylindric, more or less dorsally
gibbous, drying arcuate, smooth; annulus none to broad; peristome normally
hypnoid; lid convex, acute to obtuse-conic.
A genus of about 15 species of aquatic, largely swamp- inhabiting; mosses,
confined to temperate and cold regions: 8 species occurring in North .America;
1 species within our range and others to be expected.
Key to the Species
A. Costa weak; alar cells incrassate 4. C. trifarium
A.Costa strong; alar cells thin-walled and inflated B
B. Costa extending to the middle or a little above 3. C. stramineum
B. Costa sub-percurrent C
C. Plants slender, simple or but sparingly branched; alar cells gradually enlarged and
long-decurrent 1. C. cordifolium
c. Robust and profusely branched; alar cells abruptly enlarged and inflated
2. C. giganteum
1
Jennings; Manual of Mosses — 31. Hypnaceae 241
1. Calliergon cordifolium [Hedwig] Kindberg
(Hypnum cordifolium Hedwig; Amblystegium cordifolium DeNotaris)
Plate XLIV
Slender, tall, loosely and softly cespitose, green: stems brownish. 10-
0 cm in length; when growing in swamps, more or less erect; when in
dryer situations, more procumbent, and furnished with rhizoids; sparsely
branched, the branches more or less pinnately branched or simple, cuspidate
at the tips; leaves distant, erect-spreading, to spreading, thin, shrinking
when dry, large, 2-5 mm long, concave, cordate- to oblong-ovate, entire,
the apex rounded and sometimes cucuUate, the base decurrent: costa
slender, reaching nearly to the apex; median leaf-cells large, about
the apical and upper marginal short and wide, the cells towards th: base
gardually becoming large, wide and more or less hyaline-inflated, rounded-
hexagonal to rectangular, forming a wide but not distinctly bounded group or
band reaching clear across the base of the leaf and quite strongly decurrent;
perichaetial leaves erect, sheathing, from an ovate base long-acuminate, up to
2.5-3 mm long: seta erect, flexuous, usually 4.5-8 cm long, castaneous, when
dry flattened and dextrorse; capsule oblong-cylindric, about 3 mm long, rather
turgid-arcuate, inclined to horizontal, castaneous, slightly constricted below
the mouth when dry, exannulate; peristome-teeth pale yellow, rather thin,
rather long, hayline-margined, strongly trabeculate, the dorsal lamellae hyaline
snd papillose above, the basal portion rather irregularly striate, the teeth
confluent at base; the segment entire or but slightly carinately split, about as
long as the teeth; cilia 2 or 3, slender, nodose, about as long as the segments;
the basal membrane about one-half as high as the teeth; exothecial cells incras-
sate, rounded-quadrate to rounded-hexsgonal; lid conic, acute to apiculate;
spores mature in late spring or early summer, about .012-.015 mm, yellowish,
smooth, rather thin-walled.
In swamps, margins of pools, marshy places, etc.; Europe, Asia, New Zea-
land, and in North America from the Arctic region south to New Jersey,
Pennsylvania, and Ohio.
Fairly common in our region and now known from the following counties: Allegheny,
Butler. Cambria, Crawford. Erie. Fayette, McKean, Somerset, Warren, and Westmore-
land.. Specimen figured: Pymatuning Swamp, near Linesville, Crawford Co., May 18,
1905. O.E.J.
2. Calliergon giganteum (Schimper) Kindberg
(Hypnum giganteum Schimper)
Usually robust mosses in wet places or even in deep water, up to 10-30 cm
long; rather regularly densely pinnate with irregular branches which are often
sharp pointed; stem-leaves up to 4 x 2 mm, plicate and lustrous when dry,
decurrent, cucullate at apex, entire; costa wide, vanishing in the apex; median
cells linear-fle.xuose, only about .007 mm wide, apical cells wider, alar forming
wide abruptly inflated auricles; branch-leaves narrower; seta 5-6 cm long, red;
exannulate; ripe in May or June.
In cold swamps, Eurasia, and from Canada south to eastern Pennsylvania
and New Jersey.
242 American Midland Naturalist Monograph No. 6
3. Calliergon stramineum (Dickson; Bridel) Kindberg
(Hypnum stramineum Dickson)
Very slender yellowish-green mosses of cold bogs, erect or ascending in soft
tufts; stems up to 20 cm long; stem-leaves oblong-lingulate, concave, cucullate
at apex, entire, up to 2 x 1 mm; alar cells abruptly enlarged forming decurrent
auricles, apical tending to rounded-quadrate; costate to about % length of leaf.
Northern Eurasia and from Arctic America south to New Jersey and eastern
Pennsylvania (Pocono Mts.). Not reported from our region.
4. Calliergon trifarium (Weber and Mohr) Kindberg
(Hypnum trifarium Weber and Mohr)
When dry, in rather stiff and lustrous tufts, yellowish-green above to brown
below; stems and branches julaceous both wet and dry; stem-leaves appressed,
concave, broadly ovate to almost orbicular, obtuse, entire, scarcely decurrent;
costa thin, vanishing at about the middle or little above; median leaf-cells
linear-flexuous, shorter but scarcely wider above, the basal and alar thick-
walled, shorter and broader, but not much different. Cold bogs, especially if
calcareous, Eurasia and from Arctic America, south to Ohio and Connecticut.
Not yet found in our region.
10. Calliergonella Loeske
(Acrocladium Mitten, in part)
Autoicous or dioicous: robust, rather stiffly but loosely cespitose, lustrous,
green to yellowish or brownish; stems long, densely foliate, the apex of the
shoots rigid and acuminate by reason of the convolute apical leaves, the stems
erect, not bearing rhizoids, or procumbent, here and there with fascicles of
rhizoids, irregularly branched; stem-leaves appressed, smooth, drying somewhat
imbricate, when damp erect-spreading, concave, often cucullate. from a narrow
and sub-decurrent base broadly oblong-ovate, obtuse, rarely apiculate, entire,
the margin apically more or less involute; branch-leaves smaller and propor-
tionally narrower than stem-leaves; costa short and double or none; leaf-cells
narrowly verm.icular, smooth, wider and porose towards the base, in the exca-
vate alar portions lax,, oval-4-6-sided, hyaline, thin-v/illed, forming a d'stinct
auricular group; inner perichactial leaves erect, entire: seta 3-7 cm high,
twisted, reddish; capsule horizontal from an erect collum, oblong to cylindric,
drying arcuate and dorsally gibbous, smooth or plicate, little narrowed below
the mouth; peristome normally hypnoid with appendiculate cilia; lid convex-
conic.
As here recognized the genus consists of one species.
1. Calliergonella cuspidata [Linnaeus] Loeske
(Hypnum cuspidatum Linnaeus; H. flexile Bridel; Calliergon cuspidatum
Kindberg; Acrocladium cuspidatum Lindberg)
Plate XLIV
Tall and moderately robust with characters mainly as outlined for the
Jennings: Manual of Mosses — 31. Hypnaceae 243
genus: leaves usually bright, glossy, yellowLsh-green, or almost pure green;
stem-leaves broadly elliptic-oblong, up to 2.5 mm long, concave-cucullate,
entire, the apex often apiculate, ecostate or the costa short .ind double, leaves
crowded, usually more or less erect-spreading when moist, towards the tips of
the stems and branches imbricate-convolute so as to make th? tips cuspidate;
branch-leaves smaller and relatively narrower; median leaf-cells linear-vennicu-
lar, about 10-15:1, the alar suddenly inflated, thin-walled, hexagonal, hyaline
or colored, forming a very distinct group, the apical rather abruptly shorter,
rounded, and incrassate: seta 4-6 cm long; capsule reddish-brown; peristome-
teeth orange, hyaline-bordered, the margins step-like above; cilia 3, appendicu-
late, slightly shorter than the narrowly cleft segments; spores mature in
summer, the large capsules being but rarely produced; annulus 3-seriate.
In marshy places, swamps, and bogs; Europe, Asia, northern Africa, the
Argentine, and, in North America, through Canada and the northern part of
the United States south to New Jersey, Pennsylvania, and Iowa. Rather
uncommon in our region.
Erie Co.: Under Cephalanthus thicket, border of Cranberry Pond, Presque Isle.
C.M.B. Sept. 3, 1934. McKean Co.: East Branch, Tema Swamp, north of Bradford,
January 18, 1895. D.A.B. Snyder Co.: In bog between Shamokin Dam and Richfield,
July 17, 1908. O.E.J, (figured).
11. Hygrohypnum Lindberg
Autoicous or dioicous: slender to robust, in flattish or cushion-like tufts,
lustrous, green to yellowish-green or golden-green: stem long, procumbent, with
few or no rhizoids, remotely and irregularly branched; branches ascending;
leaves spreading to secund or imbricate, concave, smooth to weakly plicate,
more or less decurrent, lance-ovatc, and acuminate or mostly broadly oval and
obtuse to roimded, sometimes almost orbicular, margins plane, entire or ser-
rate; costa mostly unequally forked, short, weak, rarely simple and long; leaf-
cells to the base uniformly narrowly linear-vermicular, mostly with obtuse ends,
smooth, the apical often shorter and rhombic, the basal yellow to orange, the
alar portions little or not excavate but with wider, quadrate to rectangular,
hyaline to colored cells forming a small but often well-defined auricular group;
inner perichaetial leaves erect, elongate, plicate, costa simple or forked, short:
seta long, reddish, drying flattened and twisted; capsule inclined to horizontal,
mostly oval to oblong, dorsally gibbous, drying arcuate and mostly constricted
below the mouth, annulate; peristome normally hypnoid; lid convex-conic.
A genus of about 25 species in wet or moist places in cool regions; in
North America about 12 species have been reported; in our region at least 5
species, probably others to be expected.
Key to the Species
A. Epidermal stem-cells hyalme and enlarged; leaves usually distinctly falcate-secund
3. H. ochraceum
A. Epidermal stem-cells not as above B
B. Leaves widely spreading, broadly ovate to almost orbicular, and harsh when dry
5. H. dilatjtum
244 American Midland Naturalist Monograph No. 6
E. Leaves otherwise C
C. Costa single and reaching midleaf or beyond D
C. Costa none or short and double E
D. Leaves less than 1 mm long 4. H. Closteri
D. Leaves more than 1 mm long 1. H. luridum
E. Leaves entire, concave, often sub-tubulose above H. luridum
E. Leaves denticulate or serrulate towards apex F
F. Leaves widely spreading; costa single, reaching to midleaf or above
4a. H. Closteri f. serrulatum
F. Costa none, or short and double or forked G
G. Terminal leaves secund; stems and branches rather julaceous; alar cells suddenly in-
flated forming distinct auricles 2. H. eugyrium
G. Stems and branches attenuate and stoloniferous at ends; alar cells only somewhat
enlarged, rectangular to quadrate; leaves sub-clasping 6. H. novae-caesareae
\. Hygrohypnum luridum [Hedwig] Jennings
(Hypnum palustre Hudson; Amblystegtum palustre Lindberg; Hypnum
luridum Hedwig; Calliergon palustre Kindberg)
Yellowish-green, or dark -green, irregularly cespitose in low patches: stems
denuded below, long, divided irregularly, the branches erect to ascending, often
more or less hooked at the tip; leaves close, either imbricated or more or less
falcate-secund, always concave, the margins strongly incurved towards the
summit, narrowly oval- to broadly ovate-oblong, entire, about 1-L5 mm long,
the apex variable, either obtuse or acute or rounded and apiculate; costa usually
single or forked and reaching about half way up the leaf, but variable; leaf-
cells rather lax, about 5-10:1, usually linear-rhomboid, rather opaque, some-
what shorter towards the apex and towards the base, the alar few, quadrate,
sub-opaque, somewhat inflated, forming small, ill-defined auricles which are
somewhat decurrent: seta about 1-2 cm long; capsule oblong or oval-oblong,
orange-brown, arcuate, rather short and thick, more or less horizontal, dark
when dry, exannulate: lid orange-yellow, conic, obtuse to apiculate; peristome
normally hypnoid, teeth yellowish, segments scarcely carinately cleft, a little
longer than the 2 or 3 cilia; spores mature in summer.
On wet, cold rocks, where often overflowed, especially in calcareous dis-
tricts: Europe, Asia, and the northern United States and Canada, south to
New Jersey and Pennsylvania.
Rare in our region. Huntingdon Co.: T. C. Porter. (Porter's Catalogue). Mc-
Kean Co.: D.A.B. (Porter's Catalogue)
2. Hygrohypnum eugyrium (Brvologia Europaea) Loeske
{Hypnum eugyrium Bryologia Europaea; Amblystegium eugyrium Lindberg;
Calliergon eugyrium Kindberg)
Widely cespitose in low, dense, usually sand-filled tufts, lustrous, green to
reddish to brownish: stems prostrate, often leafless below; branches numerous,
erect or procumbent, usually from 0.5-1.0 cm long; leaves wide -spreading when
moist, distinctly falcate-secund towards ends of branches, when dry imbricate-
erect and concave, thus giving the branches a turgid appearance, oval-oblong
or lance-oblong, narrowed to the base, slightly denticulate towards the shortly
Jennings: Manual of Mosses — 31. Hypnaceae 245
acuminate acute apex, the margins incurved towards the apex; costa short, in-
distinct and double, or none; median leaf-cells linear, somewhat incrassate,
often somewhat obtuse at ends, about 8-10:1, shorter at the apex, the alar
abruptly much enlarged and inflated, the marginal thin-walled, the inner ones
incrassate, hyaline to yellowish-brown, forming well-defined and somewhat in-
flated auricles; perichactial leaves whitish, the outer with flexuous spreading
tips, the inner erect, long-acuminate, often crose-denticulate at the apex, pli-
cate: capsule short, oval to oblong, cemuous, turgid, yellowish-brown; peri-
stom.e-teeth yellowish, slender, strongly trabeculate; segments carinately cleft
and about equalled in length by the 2 or 3 granulose and nodose cilia; lid
conic-convex; annulus usually 3-seriate; spores mature in spring.
On rocks in streams or along the banks where kept wet, in hilly or moun-
tainous and usually non-calcareous regions; Europe, and from Newfoundland
to Alaska and south to Georgia and Colorado.
Some specimens from McKean County shew intergradations with the fol-
lowing variety.
Fayette Co.: On rock in stream-bed, Cucumber Run, above Falls, Ohiopyle. June
24, 1934; and on rock in stream, Sheepskm Run, Ohiopyle, Nov. 6, 1943. C.M.B.
Westmoreland Co.: On rock in stream one mi. above Darlington, May 19, 1945.
C.M.B.
2a. Hygrohypnum eugyrium var. Mackayi (Schimper) Brothcrus
(Hypnum eugyrium var. Mackayi Schimper; Hygrohypnum Mackayi Loeske;
Hypnum Mackayi Breidler)
Plate XLV
Leaves about 1-1.5x0.6-0.7 mm, broadly oblong, distinctly serrulate at
apex, sub-clasping and auriculate at base, less strongly falcate than in the
species; perichaetial leaves hyaline, plicate, the inner reaching 3 mm in length:
seta about 2 cm long, castaneous, smooth, somewhat flexuous, dextrorse above;
capsule with urn 2-2.5 mm long; exothccial cells rounded-hexagonal, somewhat
incrassate-collenchymatous, rather uniformly seriate; peristome-teeth about as
long as the slender carinate segments, the basal membrane about two-fifths as
high; spores minutely papillose, rather thin-walled, faintly yellowish, about
.024-. 027 mm, mature in late spring or early summer
When sterile it is very difficult to distinguish this moss from Sematophyl-
liivi marylandicum. In the Sematophyllum the walls of the outer alar cells
?rc not much thinner than are the walls of the inner alars. Sematophyllum also
sometimes has a faint double costa.
On stones in streams in hilly or mountainous regions and with about the
same general distribution as the sptcies.
Rare in our region. McKean Co.: On stones in brook at head of Bennett Brook,
Bradford, August 26, 1894, November 2, 1896, and July, 1897 (figured), the latter
issued as Grout's North American Musci Pleurocarpi. No. 129. Also Limestone Creek,
Bradford, July 7, 1895. All D.A.B. Also Tionesta Tract, Wetmore Twp., C.M.B.
Sept. 23, 1939.
246 American Midland Naturalist Monograph No. 6
3. Hygrohypnum oCHR^CEUM (Turner) Loeske
(Hypnum ochrdceum Turner; Amblystegium ochraceum Lindberg;
Limnobium ochraceum Bryologia Europaea)
Plate XLV
Yellowish or rusty green, softly cespitose in wide tufts: stems up to 8 or 9
cm long, ascending or horizontally floating, sparsely and irregularly pinnately
branched, without rhizoids, the stems and branches somewhat hooked at the
apex, the cortical cells of the stem very large and thin-walled; leaves falcate-
secund, concave, plicate, widely lance-oblong to ovate-oblong, somewhat
rounded at the base, the margins plane, entire excepting for slight serration at
the rather widely sub-obtuse apex; costa single or double, often reaching half
the length of the leaf; median leaf-cells linear-vermicular, about 8-14:1, fair-
ly thick-walled, usually rounded at the ends, the apical oval-rhomboid and
much shorter, the basal l.-'.rger and towards the angles of the leaf forming dis-
tinct decurrent auricles of abruptly enlarged, hyaline, inflated, rectangular cells:
perichaetial leaves ecostate, lance-acuminate: seta slender, flexuous, erect; cap-
sules sub-erect to cernuous from a short erect coUum, oblong, arcuate; lid
convex, mamillate; peristome hypnoid, the teeth yellowish, broadly margined,
rather distantly trabeculate, equalled in length by the carinately split segments,
the cilia shorter, unequal, nodose, two or three in number; annulus large, usual-
ly 3-seriate; spores mature in spring or early summer.
On rocks in streams or on dripping ledges, in the mountains of northern
and temperate Europe and Asia, and, in North America, from the Arctic
regions south to the latitude of New Jersey and West Virginia.
Although rare in this district, so far as now known, this species may eventually be
found to be not uncommon in cool, rocky streams in the more mountainous parts of our
region. Fayette Co.: On rock in streim, Bl'e Hole Creek. C.M.B. lulv 5, 1948.
Westmoreland Co.: In mountain rivulet, Mellon's estate, Laurel Hill Mt., New Flor-
ence, September 8-10, 1907. O.E.J. Sterile (figured).
4. Hygrohypnum Closteri (Austin) Grout
(Hypnum Closteri Austin)
Slender, in loose tufts, green to yellowish; stems creeping, irregularly
branched; leaves rather widely spaced, spreading, not or rarely secund, leaves
ovate or oblong-ovate, rarely reaching 1 mm long, flat apex obtuse, margin
entire, only very slightly decurrent; costa normally single, reaching to mid-
leaf or beyond; median leaf-cells linear, somewhat opaque, about 3-5:1, apical
shorter; basal shorter and wider, alar but little enlarged: seta 6-8 mm long,
castaneous; capsule, brown, ovoid, cernuous, arcuate, much constricted below
mouth when dry and old; annulate; lid low-conic, apiculate; spores ripe in
spring.
On stones in cool streams. Vermont to eastern Pennsylvania and Virginia,
but not yet found in our region.
4a. Hygrohypnum Closteri f. serru latum Grout
Lseaf-cells more hyaline; leaves often serrulate above.
McKean Co.: Bolivar Run, Burnett No. 3208 (Type). September II, 1898.
Jennings: Manual of Mosses— 31. Hypnacfae 247
5. Hygrohypnum dilatatum (Wilson) Loeske
{Hypnum dilatatum Schimper)
Yellowish-green above, blackish below, the tufts stiff and harsh when dry;
stems ascending towards tip and with erect, short, blunt branches; leaves widely
spreading, somewhat secund, from a narrower, somewhat decurrent base broad-
ly oval-elliotic to almost orbicular, obtuse to apiculate, up to 2 mm long,
entire or tamtly denticulate at apex; costa usually faint, forked, rarely reaching
mid-leaf, occasionally single; leaf-cells incrassate 10-15:1, towards base 20:1,
at apex 2-4:1, the alar cells form a weak group of oval-angular to rectangular,
yellowish, thick-walled: seta 1-2 cm; capsule oblong from a narrow neck, 2 mm
long, castaneous, constricted below the mouth when dry and empty; annulus
2-seriate; lid orange, high convex, with red apiculus; spores ripe in summer.
On non-calcareous stones in swiftly flowing mountain brooks. Eurasia,
and in North America from Canada south to West Virginia and the South-
west. To be exoected in our mountains also.
6. Hygrohypnum novae-caesareae (Austin) Grout
{Hypnum micaris Wilson, not Swartz; Rhytichostet^ium novae-caesareae Austin;
Raphidostegium novae-caesareae Renauld and Cardot)
Plate LIII
Small, yellowish-green, glossy, forming wide, thin mats: stems prostrate,
very slender, sparsely branching, often flagelliform, the branches short, simple
or sparsely branched, sub-erect; leaves spreading or the upper sometimes sub-
secund, sub-orbicular, apiculate to shortly acuminate, 0.6-0.8 mm long, serru-
late above, concave, the margins somewhat reflexed below; costa double and
very faint; median leaf-cells linear, flexuous, about 6-10:1, the apical rhom-
boid-oblong, rather incrassate, much smaller than the median, the basal a little
shorter and wider than the median, the alar region with about 6 to 10 larger,
quadrate to rectangular, rather incrassate cells and with the outermost one to
three cells much larger and more or less inflated: the capsules of this species
have thus far been found but once, — on damp rocks along Stony Creek, Car-
bon County, Pennsylvania, by Francis WoIIe: capsules small with a shortly
rostrate lid, the exothecial cells non-col lenchym.atous: dioicous.
On damp rocks in cool and moist mountain ravines, Eurasia, and from
Vermont to western Pennsylvania and Georgia, in the mountains.
Rare in our region. Favette Co.: Beck Spring, Laurel Ridge, July 26, 1947, and
clinging to rock. Blue Hole Creek, Laurel Ridge. July ■>. 1948. C.M.B. McKean Co.:
Bennet Brook, July 10, 1898. (figured).
12. Campylium (Sullivant) Bryhn
Mostly dioicous: slender, rarely robust, mostly stiffly cespitose, green to
yellowish or brownish, drying more or less lustrous; stems creeping to ascend-
ing or erect, bushy to variously pinnate; leaves from a shortly decurrent base
broadly ovate or cordate, gradually or abruptly narrowed into a long, slender,
canaliculate acumination which is mostly strongly squarrose-reflexed, margin
248 American Midland Naturalist Monograph No. 6
plane, mostly entire; costa various, mostly short; cells narrowly rectangular-
oblong to linear-prosenchymatou5, smooth; alar cells forming a distinct group,
yellowish, incrassate to inflated, small, quadrate: seta long, drying twisted,
reddish to yellowish-red; capsule inclined to horizontal, sub-cylindric, arcuate,
annulate; peristome normally hypnoid; lid convex, acute to conic-obtuse; spores
small.
About 30 species in both dry and wet habitats, mainly confined to the tem-
perate regions; about 20 species reported for North America; 4 species now
known in our region.
Key to the Species
A. Costa simple, thin, ending in about the middle of the leaf, or somewhat above the
middle C
A. Costa none or very short B
B. Stem slender, creeping; leaves finely serrulate all around; alar cells small, quadrate
1. C. hispidulum
B. Stem usually erect or ascending; leaves entire; alar cells dilated, sub-rectangular
4. C. stellatum
C. Leaves strongly squarrose; alar cells scarcely enlarged 2. C. chrysophyllum
C. Leaves spreading-erect; alar cells enlarged D
D. Stem-leaves very slenderly acuminate from a broadly lanceolate blade 3. ..C folyganum
D. Stem-leaves abruptly acuminate from a cordate-ovate blade 5. C. radicalc
1. Campylium hispidulum (Bridel) Mitten
(Hypnum hispidulum Bridel; Chrysohypnum hispidulum Roth; Stereodon
hispidulus Mitten; Amblystegium hispidulum Kindberg)
Plate XLV
Slender, interlaced in bright green tufts more or less yellowish below;
stems creeping radiculose, abundantly but irregularly branching, the branchlets
slender and erect or ascending; leaves widely spreading to distinctly squarrose,
about 0.5-0.8 m.m long, triangular-cordate, abruptly acuminate, the slender
acumen about one-third as long as the main body of the leaf, the leaf slightly
concave, decurrent, sub-serrulate all around, excavate at the base; costa double
and very short, or none; median leaf-cells about 3-6:1, with .005-. 006 mm
wide, prosenchymatous, elongate-oblong with blunt ends, the alar numerous,
sub-rectangular to quadrate, granulose, up to twice as wide as the median cells:
seta pale castaneous to yellow, about 1.5-2 cm long, slender dextrorse; capsule
small, oblong, more or less incurved, yellowish-brown, wide-mouthed, the urn
about 1.2-1.4 mm long, when dry furrowed and narrowed below th? mouth;
annulus uni-seriate; lid convex-conic with an upturned apiculation; peristome
normally hypnoid, the segments slightly cleft and almost equalled in length
by the nodose to sub-appendiculate cilia; spores mature in summer, yellowish,
medium-walled, minutely papillose, about .011-. 014 mm.
On the bases and roots of trees, on decaying wood, on humus, etc., always
near the ground in moist shaded places; in Europe, Asia, and, in North Amer-
ica from Canada to the southern part of the United States.
Rather common in our region. Now known from .Allegheny, Cambria (Porter), Cen-
tre, Erie, Indiana (Porter), Fayette. McKean, Washington, and Westmoreland counties.
Specimen figured: Edge of pond at Scotia, Centre Co., Sept. 20, 1909. O.E.J.
Jennings: Manual of Mosses — 31. Hypnaceae 249
2. Campylium chrysophyllum (Bridel) Bryhn
{Hypnnm chryfophyllurn Bridel; Chrysohypnum chrysophyllum
Loeske; Amblystegtum chrysophyllum De Notaris)
Plate XLV
Ce^pitose in low, lax, or dense, bright golden-green tufts or mats; stems
slender, rather long, prostrate, more or less regularly pinnate, the branchlets
erect or spreading; leaves close, small, 1-1.5x0.4-0.8 mm, squarrose-spreading
from the sub-clasping base, sometimes secund, stem-leaves ovate-cordate to tri-
angular-cordate, decurrent, narrowed abruptly to a long somewhat channeled
acumination, entire or very slightly denticulate at base; branch-leaves similar
but smaller and narrower; costa single, reaching about to the middle or higher;
median leaf-cells about 5-10:1, about .005-. 010 mm wide, rather incrassate,
the alar forming a group of small, incrassate, sub-opaque, sub-quadrate cells:
seta castaneous, about 2-2.5 cm long, slender, fle.xuous; capsule oblong-cylindric,
inclined to horizontal, arcuate, castaneous to orange; lid conic-apiculate; annu-
lus large, compound; peristome normally hypnoid, the teeth yellowish, hyaline-
margined, strongly trabeculate, dorsally lamellate, cross-striolate below, hyaline
and papillose above; the segments not usually carinately split, the cilia stout,
nodose, 2 or 3, and about as long as segments, basal membrane one-half as
high as sigments; spores in early summer, light brown, smooth, .010-. 012 mm:
dioicous.
On earth, stones, roots of trees, etc., in moist places; Europe, Asia, and, in
North America, from Canada to the southern and southwestern United States.
Common in our region excepting on the High Plateau.
Now known from Allegheny, Beaver, Butler, Crawford, Erie, Fayette, Indiana, Law-
rence, McKean, Washington, and Westmoreland counties. Specimen figured: P/matun-
mg Swamp, Linesville, May 12, 1908. O.E.J.
3. Campylium polygamum (Schimper) Bryhn
{Hypnum polygamum Wilson; Chrysohypnum polygamum Loeske)
Plate XLVI
Moderately robust, yellowish-green to golden, low, cespitose: stems erect
to ascending, about 3-6 cm long, divided and with rather numerous, irregularly
pinnate, rather crowded, and erect or ascending branchlets; stem-leaves lance-
ovate, 2-2.5 mm long, moderately close, erect-spreading both wet and dry,
with an ovate or oblong base narrowed above into a long, gradually tapering,
channeled acumination, entire, the base rounded and clasping, somewhat decur-
rent; branch-leaves elongate-lanceolate, with the sides tapering in a straight line
from the rounded-ovate base, the leaves averaging about 3 mm long; median
leaf-cells narrowly linear, about 8-12:1, in the older leaves somewhat incras-
sate, towards the base often porose, the alar sub-rectangular, somewhat enlarged,
distinct, forming often orange-pellucid auricles; costa not very strong but dis-
tinct and usually reaching somewhat above the middle of the leaf: seta slender,
flexuous, about 3-4 cm long; capsule oblong-cylindric, curved; lid conic-apicu-
250 American Midland Naturalist Monograph No. 6
late; peristome normally hypnoid, cilia well developed; annulus present; spores
mature in summer.
In moist places in meadows and swamps and said to prefer sandy soils;
Europe, Asia, and from Arctic North America south to New Jersey, Virginia,
and Pennsylvania.
Rare in our region. Allegheny Co.: Schenley Park, Pittsburgh, August 26 1906.
O.E.J. (?). Crawford Co.: Near Linesville in the Pytnatuning Swamp, May 10-11,
1906. O.E.J, (figured). Erie Co.: On soil among cattails in swamp, Presque Isle,
Nelle Ammons. July 18, 1933. McKean Co.: Wildcat Hollow, Marvin Creek. On
twig in stream. C.M.B. Sept. 1, 1935.
4. Campylium stellatum [Schreber] Lange and C. Jensen
{Hypnum stellatum Schreber; Chrysohypnum stellatum Loeske;
Amblystegium stellatum Lindberg)
Plate XLVI
Robust, densely tufted, soft, lustrous, bright to golden green: stems stout,
usually ascending, up to 8 or 10 cm long, irregularly divided, the branchlets
sub-pinnate and more or less crowded and erect; leaves close, from 1-3 mm
long, from a widely squarrose to an erect-spreading and more or less cordate
base narrowed, often rather abruptly, to a gradually long-acuminate, acute,
ascending or usually squarrosely spreading acumen, the base entire or some-
times slightly denticulate, slightly excavate and with rounded and subdecur-
rent auricles, the upper part of the leaf more or less channeled; costa none or
very short, either single, forked, or double, but usually appearing only as yel-
lowish or brownish striae; median leaf -cells narrowly linear, about 8-15:1, in
the older leaves rather incrassate and blunt at the ends, the basal often porose,
the alar somewhat enlarged, sub-rectangular, incrassate, sometimes rather
opaque usually more or less orange-pellucid, forming distinct, often somewhat
decurrent auricles: seta rather long, castaneous, up to 4.5 cm long, stout, dex-
trorse above, flexuous; capsule oblong-cylindric, about 3-4.5:1, arcuate, the urn
2-2.5 mm long, inclined to horizontal, sulcate and constricted below the mouth
when dry and empty, brownish; lid highly convex-acuminate; annulus 2-3-
seriate; peristome normally hypnoid, large, the teeth trabeculate, confluent at
base, the lower part orange-colored and dorsally cross-striolate, the lamellae
and divisural distinct, the upper part paler and papillose; segments somewhat
shorter and slightly carinately split; cilia 2 (or 3), nodose, about as long as
the segments; spores mature in spring (?) or summer, minutely papillose,
rather thin-walled, pale yellowish, .011-.014 mm.
On wet banks and tufts in cool swamps and bogs. Europe, Asia, and
from Arctic America southwards to Pennsylvania and Ohio; and the West.
BuTi.ER Co.: West Liberty Bog. O.E.J. June 28, 1941. Erie Co.: Presque Isle.
O.E.J. May 8-9, 1906. Lawrence Co.: Frew Mill Road, m patch of Gentiana crinita,
Sept. 18, 1948; and also on wall of old Van Port limestone quarry Yl f"'- ^- o^ New
Castle, Nov. 26, 1948. C.M.B. The Presque Isle specimen was figured.
Jennings: Manual of Mosses — 31. Hypnaceae 251
5. Campylium radicale (Beauvois) Grout
(Hypnum radicale Beauvois; H. bergenense Austin;
Amblystegium radicale Mitten)
Plate XLII
Loose, slender, pale-green, little branched, the branches often erect and up
to 2 cm or more long; branch-leaves distant, more or less widely spreading,
lanceolate- to ovate-cordate, up to 1.5x0.7 mm, concave at base, entire or
almost so, dccurrent, abruptly slenderly acuminate and somewhat channelled;
m.edian leaf -cells about 4-8:1, sometimes longer, medium-walled, the alar some-
times more abruptly enlarged and hyaline; costa well developed, orange, up to
two-thirds or three fourths as long as the leaf; perichaetial leaves slenderly
acuminate and up to 2.5 mm long, plicate, erect: seta up to 3 cm long, cas-
taneous, strong, dextrorse; capsule yellowish, the urn about 2.7 mm long,
arcuate, oblong-cylindric, contracted below the mouth when dry; peristome-
teeth strong, castaneous, strongly trabeculate, hyaline-margined, the dorsal
lamellae cross-striolate below, papillose and hyaline above; segments entire or
nearly so, about as long as teeth, the basal membrane about two-fifths as high
as teeth; cilia 2 or 3, usually one, at least, as long as the segments, nodose,
hyaline, minutely papillose, medium-walled, about .016-.019 mm, mature in
spring.
On rotten logs, roots of trees, wet soil, etc., in wet, sh.qded places: Europe,
Asia, and apparently well distributed throughout temperate North America.
The species occurs in eastern Pennsylvania and has been found twice in our region.
Butler Co.: West Liberty Bog. C.M.B. Aug. 29, 1931. McKean Co.: In springy
places near Bradford, May 17, 1895. D.A.B. (figured).
13. Ctenidium (Schimper) Mitten
Usually dioicous, mostly slender, soft, cespitose, green to yellowish or
golden-brown, lustrous: stem long, here and there with clusters of rhizoids,
more or less regularly pinnate, branches short and mostly horizontally spread-
ing; leaves falcate to circinate-secund, decurrent, more or less plicate, mostly
abruptly lance-subulate from a broadly cordate base, serrate (C. procerrimum
is entire) ; costa double and very short or none; cells narrowly linear, the upper
angle usually ending in a forward-projecting papilla, the alar cells distinct,
quadrate and rectangular, the angles sometimes weakly e.xcavate; branch-leaves
smaller: seta 1-2.5 cm long, red, smooth or nearly so; capsule inclined to
nearly horizontal, thickly oval, dorsally gibbous, not constricted below the
mouth; annulus broad, revoluble; peristome normally hypnoid; lid long-conic,
acute or obtuse; calyptra mostly more or less hairy.
A genus of 30 species occurring mainly on trees and rocks in temperate
and warm regions; 4 species in North America, the following species in our
range.
1. Ctenidium molluscum [Hedwig] Mitten
{Hypnum molluscum Hedwig; H. compressum Roth)
Plate XL VI
Very densely cespitose, soft, lustrous, golden-green, rather robust: stems
252 American Midland Naturalist Monograph No. 6
reaching 10 cm in length, prostrate or ascending, closely regularly pinnate,
plumose; leaves densely imbricated, falcate-secund to circinate, stem-leaves 1.8-
2.5 mm long, when dry usually plicate, and, especially towards the points,
more or less undulate and crisped, from a cordate-acuminate, plane-margined,
auriculate base rather abruptly and slenderly long-acuminate, plane-margined,
strongly serrate, especially at the base, somewhat decurrent; costa very short
and double or none; median leaf -cells about 8-15:1, the corners somewhat pro-
jecting dorsally, gradually towards the angles becoming irregularly quadrate-
hexagonal, shorter and wider, pellucid, forming poorly defined auricles of
about the same color as the rest of the leaf; branch-leaves considerably smaller
and narrower, not cordate-auriculate; perichaetial leaves slenderly lance-acumi-
nate; paraphyllia ovate, mostly at the base of the branches: seta brownish,
flexuous, slender, about 1.0-2.5 cm long, castaneous; capsule-urn about 2.5 mm
long, oblong to oval, slightly curved to almost straight, from the curved apex
of the seta mostly horizontal, not constricted below the mouth when dry; lid
conic-acuminate; annulus broad; peristome normally hypnoid, teeth yellowish,
segments carinately cleft, about as long as teeth, the cilia 2 or 3, stout, about
as long as the segments, the basal membrane about one-half the height of the
peristome; calyptra somewhat hairy when young; spores mature in summer,
smooth, yellowish-incrassate, about .01 5-. 018 mm.
On moist, shaded earth and rocks, or bases of trees, in woods, more partic-
ularly in hilly or mountainous districts; Europe, Asia, northern Africa, and
from Newfoundland to the Rocky Mountains and south to Georgia and
Oklahoma.
Armstrong Co.: Crooked Creek, one mi. s.w. of Tunnelton. C.M.B. Aug. 18,
1935. Beaver Co.: On ground. Raccoon Creek, 2 mi. w. of Little Traverse Creek.
C.M.B. Sept. 15, 1935. Blair Co.: A. P. Garber. (Porter's Catalogue). Butler Co.:
On wet roadside bank, Semiconon Run, 2]/2 mi. w. of Conoquenessing. Sidney K. East-
wood. March 24, 1935. Cambria Co.: Cresson. T. C. Porter. (Porter's Catalogue).
Elk Co.: McMinn. (Porter's Catalogue). McKean Co.: On rich, shaded bank of
stream, Langmade, April 3, 1897, and April 25, 1897 (figured), and on rocks bordering
rivulets, head of Gates' Hollow, Bradford, October 27, 1895. D.A.B. Washington
Co.: Linn and Simonton. (Porter's Catalogue).
14. Rhytidiadelphus (Lindberg) Warnstorf
Dioicous: more or less robust, stiff, loosely cespitose forming loose, wide,
stiff, green to yellowish or grayish, and rather lustrous mats; stem angled, long,
without rhizoids, simple to regularly or irregularly pinnate; branches partly
short and obtuse, partly long and acuminate, and often curved above; upper
half of the leaf spreading-squarrose to reflexed-squarrose, sometimes circinate-
pecund, mostly plicate, scarcely decurrent, from an ovate or cordate base more
or less long-acuminate, plane-margined, rather sharply serrate; costa reaching
above mid-leaf, or short, double, or sometimes none; cells narrowly linear,
smooth, or the upper angle projecting dorsally as a tooth, the basal wider,
shorter, more or less incrassate and porose, colored, the alar mostly not differ-
entiated; seta 2-4 cm long, castaneous; capsule horizontal to pendent, from a
very short neck thickly oval, dorsally gibbous, when dry and empty plicate,
Jennings: Manual of Mosses — 31. Hypnaceae 253
but not constricted below the mouth, annulate; peristome normally hypnoid;
hd convex, conic-acute.
A genus of 6 species of forest and meadow in the temperate and cold
regions of the Northern Hemisphere; 4 species in North America; 2 species
in our region.
Key to the Species
A. Cells smooth both sides; stem-leaves not plicate, squarrose-recurved, long and slender-
ly acuminate 1. R. squarrosus
A. Cells dorsally spinose; stem-leaves strongly plicate, spreading, short-acuminate
2. R. triquetnis
1. Rhytidiadelphus squarrosus [Linnaeus, Hedwig] Wamstorf
(Hypnum squarrosum Linnaeus; Hylocomium squctrrosum Bryologia Europaea)
Widely and softly cespicose, bright green, lustrous: stems robust, but
slender, up to 10 or even 15 cm long, procumbent or more or less ascending
to erect at the ends, the branchlets rather distant, flexuous, unequal, attenu-
ated and more or less sub-flagelliform; stem-leaves crowded, about 3 mm long,
abruptly squarrose from a cordate-ovate more or less erect-sheathing base, not
secund, imbricated, the squarrose portion long and gradually tapering and
channeled, denticulate above, the apical leaves somewhat stellately spreading,
branch-leaves smaller but otherwise very similar to stem-leaves; costa short,
double, faint; median leaf-cells smooth dorsally, about 8-10:1, narrowly-linear,
the alar gradually rectangular-hexagonal, larger, short, opaque to pellucid,
numerous, but not forming abruptly differentiated auricles; perichaetial leaves
squarrose, the inner linear-acuminate and apically serrate: seta usually 3-4 cm
long, flexuous; capsule castaneous, short, ovoid, dorsally gibbous, inclined to
horizontal, or even pendent by the cur\'ing of the upper part of the seta; lid
convex-conic, rather acute; annulus 2-seriate; peristome normally hypnoid, seg-
ments carinately split between the articulations, cilia 3; spores mature in winter
or early spring.
On soil, rocks, or logs, m cool places in moist or wet meadows and borders
cf woods in grassy places; Azores, Europe, Asia, and, in North America,
from the Arctic regions to the northern United States as far south as Penn-
sylvania and Tennessee.
Rare in our region. Cambria Co.: Lesquereux, at Cresson. (Porter's Flora). Somhr-
SEtCo.: Beck Spring, Laurel Ridge. C.M.B. July 26, 1947 (figured).
2. Rhytidiadelpus triquetrus [Linnaeus, Hedwig] Wamstorf
(Hypnum triquetrum Linnaeus; Hylocomium tr'iquetrum Bryologia Europaea)
Plate XLVII
Very robust, stiff, elastic, bright to yellowish-green, bushy-cespito:-e: stems
long, up to 15 or 18 cm, branching unequally and irregularly, sometimes
more or less pinnately, reddish, woody, ascending or sometimes erect; stem-
leaves large, 4-5 mm long, stiff, scarious, divaricately or horizontally spreading
both wet and dry, widely cordate- to deltoid-triangular, widely rounded-
254 American Midland Naturalist Monograph No. 6
auriculate at base, the insertion narrow and decurrent; leaves plicate, denticu-
late, papillose dorsally, gradually tapering above to a sub-acute apex; branch-
leaves narrower and smaller towards the ends of the attenuate branches; costa
forked, or of two parallel divisions reaching about three-fourths the length of
the leaf; perichaetial leaves non-costate, the acuminations squarrose; median
leaf-cells linear, about 8-10:1, at the angles oblong-hexagonal, pellucid, not
usually forming distinct auricles, the upper cells dorsally forming spinulose
papillae: seta 1.5-3.5 cm long, rather rigid, lustrous, castaneous; capsule turgid-
oblong, large, castaneous, about 3 mm long, dorsally gibbous, inclined or more
nearly horizontal by a curve in the upper part of the seta, when dry more or
less plicate and constricted below the mouth; the exothecial cells rounded-
hexagonal, rather small, incrassate, castaneous; lid conic, acute; annulus 2-3-
seriate; peristome normally hypnoid, the teeth orange-yellow, strongly trabecu-
late, dorsally lamellate, the lamellae papillose but non-striate, projecting to
form a distinct border; the segments carinately split, the cilia 2 (or 3) and
about as long as the segments, stout, the basal membrane reaching about one-
half the height of the peristome; spores medium-walled, smooth, yellowish,
.018-. 025 mm, ripening in winter or early spring.
On shaded banks, humus, or rotten logs in cool woods with a moderate
amount of moisture, or in swamps; Europe, Asia, northern Africa, and, in
North America from the Arctic regions south to the northern United States,
and along the mountains in North Carolina, in ravines and cool, rocky woods.
Cambria Co.: T. C. Porter. (Porter's Catalogue). Elk Co.: McMinn. (Porter's
Catalogue). McKean Co.: On decaying leaves under hemlocks, Marilla Brook, Brad-
ford, June 5, 1895 (figured), and September 29, 1894; Bennett Brook, July 15, 1893.
D.A.B. Washington Co.: Linn and Simonton.
15. Rhytidium (Sullivant) Kindberg
Dioicous: very robust, in wide, loose, yellowish or brownish-yellow tufts
or mats: when dry stiff and lustrous; stems long, tumid, with hooked tips,
prostrate to ascending or erect, with few or no rhizoids, simple to regularly
pinnate, or bipinnate, rarely bushy; branches 2-seriate, short and thick, or
longer, acuminate and downwardly arcuate; leaves crowded, imbricate, falcate-
secund, concave, plicate, rugose, scarcely decurrent, lance-ovate to oblong-
ovate, narrowed into a long, canaliculate, lance-subulate, sharply-toothed
point, the margin more or less revolute; costa simple, thin, reaching; to mid-
leaf; median leaf-cells narrowly vermicular, with dorsally (sometimes a few
ventrally, also) projecting and forward-pointing teeth-like papillae at the upper
end of the cell, towards the costa at base the cells more lax, rectangular, porose,
incrassate, the alar region not excavate, the alar cells forming a distinct longi-
tudinal band of small, quadrate and polygonal, yellowish, incrassate, numerous
cells; inner perichaetial leaves elongate-lanceolate, slenderly acuminate, plicate,
serrate, ecostate: seta 2-5 cm long, castaneous, when dry twisted; capsule in-
clined to horizontal, elliptic to sub-cylindric, dorsally gibbous, when dry arcu-
ate and constricted below the mouth, brownish; annulus 3-seriate, remaining
attached to the operculum; lid convex-conic, shortly and obliquely rostrate;
Jennings: Manual of Mosses — 31 Hypnaceae 253
peristome normally hypnoid, teeth msty-yellow, segments broadly split, cilia 2,
as long as the segments; spores in summer but capsules very rarely produced.
One species, as follows, on exposed sunny rocks and ledges, and in dry,
grassy places; Europe, Asia, and from Arctic America south through Canada
to North Carolina. Usually in hilly or mountainous regions on calcareous
substrata. Rare in our region.
1. Rhytidium rugosum [Ehrhart, Hedwig] Kindberg
{Hylocomium rugosum DeNotans; Hypnum rugosum Ehrhart)
Plate LXVI
Stems reaching 8 or 10 cm or more, the branches tumid and sometimes 4-6
mm in diameter; the leaves 3 mm long or more, sometimes costate above the
middle, margins narrowly reflexed.
Beaver Co.: On ground in Pinus virginiana forest, along Service Creek, one mile
west of Raccoon Creek, C. M. Boardman, Feb. 10, 1935 (figured). Huntingdon Co.:
T. C. Porter. (Porter's Catalogue).
16. Hylocomium Bryologia Europaea
Dioicous, more or less robust, stiffly and laxly cespitose in green or yellow-
ish and more or less lustrous tufts: stem mostly very long and procumbent
or ascending, more or less arcuate, once to three times pinnate; paraphyllia
numerous, much-branched; leaves more or less spreading, concave, mostly pli-
cate, oblong to cordate, long-acuminate, plane-margined, serrate; costa thin,
double, sometimes reaching mid-leaf; cells linear, mostly smooth, basally shorter
and laxer, colored, incrassate, porose, alar not differentiated; inner perichaetial
leaves with reflexed-squarrose acuminations: seta more or less elongate, red;
capsule inclined to horizontal, thickly ovate or oblong-oval, somewhat dorsally
gibbous, with neck short and narrowed into the seta, drying mostly smooth
and scarcely constricted below the mouth, annulate; peristome normally hyp-
noid; lid convex with a conic-acute point or shortly and obliquely rostrate.
A small, variously delimited genus mainly inhabiting forests in temperate
and cold regions; 3 species in our region.
Key to the Species
A. Leaves at base semi-amplexicaul, with very large and rounded auricles; stems erect
or arched 3. H. brevirostre
A. Leaves with broad insertion but not with rounded auricles B
B. Stem closely 2-3 pinnate; leaves obscurely bi-costate, rarely reaching midleaf
1. H. splendens
E. Stem irregularly or distantly 1-2 pinnate; costa double and reaching to mid-leaf
or more 2. H. umbratum
1. Hylocomium splendens (Hedwig) Bryologia Europaea
{Hypnum splendens Hedwig; Hylocomium proliferum Lindberg )
Plate XLVII
Widely cespitose in loose mats, lustrous, yellowish to brownish or olive-
green: stems long, trailing, red, with green, branched paraphyllia, stems some-
256 American Midland Naturalist Monograph No. 6
times up to 15 or 20 cm long, divided, the fern-like shoot of each year ascend-
ing from the side of the upper third of the preceding year's shoot, the divisions
usually complanately and loosely bi- to tri-pinnate; stem-leaves crowded, erect-
spreading to loosely imbricate, broadly ovate to ovate-oblong, 2-3 mm long,
the insertion wide, the upper portion of the leaf abruptly acuminate into a
slender, transversely undulate and flexuous point, or sometimes shorter and
obtuse, the leaves basally plicate, sub-decurrent, somewhat concave, recurved
at margin below, denticulate and dorsally spinulosely papillose above; branch-
leaves usually acute, smaller and non-plicate, concave, elliptic-oblong; costa
double and reaching to one-fourth or one-third the length of the leaf, but
faint; median leaf -cells linear-flexuous, about 8-10:1, the lower more or less
porose, the basal orange-pellucid, incrassate and larger, but not forming dis-
tinct auricular groups; perichaetial bracts long, the inner erect, narrowly acumi-
nate and sheathing: seta about 1.5-2.0 cm long, red, usually stiff, curved,
when dry wrinkled and sometimes sinistrorse; capsule oblong-ovate, orange-
brown, somewhat turgid, usually horizontally inclined, the urn about 2-3 mm
long; peristome hypnoid, the teeth basally confluent, trabeculate, lamellate,
dorsally striolate below, papillose above, brownish; segments about as long,
widely carinately gaping, yellowish, the three slender, nodose cilia about as long,
the basal membrane about two-fifths as high as the teeth; lid rostrate; exothecial
cells brownish, rather thin-walled, rectangular to hexagonal, several rows at the
rim much smaller; spores smooth, medium-walled, .010-.014 mm, mature in
spring.
On stones and logs m cool, rich, moist mountain woods and ravines;
Europe, Asia, northern Africa, and, in North America from the Arctic regions
south to North Carolina.
Not common in our region. Bedford Co.: On big blocks of "gannister" on the open
rock talus slope in gap west of White Sulphur Springs. O.E.J. June 15, 1941. Blair
Co.: T. C. Porter. (Porter's Catalogue). Clarion Co.: Thorns Run, Cook Forest. S.
K. Eastwood. Sept. 15, 1935. Elk Co.: McMinn. (Porter's Catalogue). Jefferson
Co.: Kate Stoy. McKean Co.: On logs and on ground over leaves, Rutherford Run,
April 25, 1893, West Branch Swamp, on logs, October 15, 1893, and on rich, shaded
banks over leaves, Manila Brook, June 30, 1895 (figured), all Bradford. D.A.B. Tioga
Co.: On wet cliff near Leetonia. S. K. Eastwood. Sept. 6, 1936. Washington Co.:
Linn and Simonton. (Porter's Catalogue).
2. Hylocomium umbratum [Ehrhart] Bryologia Europaea
{Hypnum umbratuin Ehrhart)
Plate XLVIII
Slender, not so large and not complanately branched as in H. splendcns,
more erect and forming loose, green tufts often 12 or 15 cm high, sometimes
yellowish, somewhat lustrous: stems rigid, pinnately or bi-pinnately, bushily
branched, the branchlets unequal, often drooping, sometimes distinctly flagel-
liform, the stems reddish, bearing numerous conspicuous and branched para-
phyllia; stem-leaves quite broadly triangular-ovate, rather distant, rather
spreading, about 2 mm long, acute to long-acuminate, decurrent, strongly pli-
Jennings: Manual of Mosses — 31. Hypnaceae 257
rate, undulately strongly dentate all around, the teeth sometimes recurved, no
papillae on back of leaf; branch-leaves more ovate and smaller; costa double
and strong, reaching to about mid-leaf; median leaf-cells about 8-10:1, linear,
not forming distinct auricles, the extreme basal castaneous-incrassate, rounded;
perichaetial leaves broad, apically spreading: seta slender, 3-4 cm long, flexu-
cus; capsule short, about 2:1, turgid-ovate, more or less horizontally inclined,
somewhat plicate and constricted below the mouth when dry and empty; peri-
stome normally hypnoid, segments carinately split, the cilia usually 2, about
as long as segments; annulus none; lid conic, shortly apiculate; spores mature
in winter or early spring.
Over rocks, logs, and woods-humus, in cool mountain woods and ravines;
Europe, Asia, and, in North America, from Newfoundland to Alaska south
to Ohio, and southwards in the mountains to North Carolina.
Rare in our region. McKean Co.: In deep, densely shaded ravines, altitude 1700
feet along Marilla Brook one-half mile above Bradford, April 21, 1879. D.A.B. (fig-
ured ) .
3. Hylocomium brevirostre [Ehrhart] Bryologia Europaea
{Hypnum brevirostre Ehrhart; H. interruptum Bridel )
Plate XLVIII
Robust, rigid, forming large, swollen tufts of a dark but glossy yellow-green:
stems much-divided, up to 12-15 cm long, erect to arched-procumbent, irregu-
larly pinnately branched, the branches unequal, attenuate, not complanately
arranged, but bushy, paraphyllia rather small, branched, stems reddish-brown;
stem-leaves somewhat crowded, spreading to squarrose, about 2-4 mm long,
cordate-ovate to triangular-ovate, abruptly narrowed to a rather long channeled
acumen, the base notably with large rounded, sub-clasping, and somewhat
decurrent auricles, the margins finely and regularly denticulate, the leaves when
dry much plicate: branch-leaves smaller, narrower, more ovate, less squarrose;
perichaetial leaves sheathing at base, subulate-acuminate, squarrose, apical'y
serrate; costa of stem- and branch-leaves double and reaching to about one-
third the length of the leaf; median leaf-cells linear, about 5-8:1, rather incras-
sate, basal cells shorter and wider, colored, with pitted walls, the alar not
differentiated: seta flexuous, about 2 cm long, dextrorse and arcuate above,
castaneous; capsule turgidly ovoid-oblong, horizontally inclined, the urn about
2 mm long, castaneous, arcuate and constricted below the mouth when dry;
lid conic-acuminate, about 1 mm long; annulus usually 2-seriate, rather wide;
peristome hypnoid, teeth orange-yellow, dorsally lamellate, cross-striolate
below, papillose above, moderately trabeculate, confluent and inserted at base;
segments slender, about as long at teeth, yellowish, carinately gaping, finely
spinose-papillose above, the basal membrane about two-fifths as high; cilia
usually short, nodose-appendiculate; exothecial cells laterally strongly cas-
taneous-incrassate, rounded-hexagonal to rounded-rectangular; spores about
.021 -.024 mm, oblong to round, castaneous, moderately incrassate, somewhat
papillose, mature in winter or early spring.
In cool, deep, shaded ravines and in swamps on soil, rocks, or at the bases
258 American Midland Naturalist Monograph No. 6
of trees, usually confined to mountainous regions; Europe, Asia, northern
Africa, and, in North America, from Nova Scotia to Ontario and south in the
mountains to Georgia; Missouri.
It is probable that deforestaion and lumbering activities have largely so
modified the environment that this species is becoming rare. Most of the col-
lections recorded were made prior to 1900.
Rather common in our region. Known from Beaver (Porter), Blair (Porter), Cam-
bria (Porter), Clinton, Elk (Porter), Fayette, McKean, Somerset, Warren, and Wash-
ington (Porter) counties. Sfjecimen figured: Densely shaded rocks, Marilla Brook,
Bradford, McKean Co., April 25, 1895. D.A.B. (This was distributed in Grout's
N. Am. Musci Pleurocarpi, No. 44).
17. Hypnum Linnaeus, Hedwig
Dioicous: robust, stiff, deeply and loosely cespitose, dark to pale green or
almost straw-colored, more or less lustrous: from a decumbent base ascending
to erect, with straight pointed ends and rather regularly pinnate; branches
mostly spreading and 2-seriate, usually slenderly attenuate, sometimes thick,
julaceous, and obtuse; no paraphyllia; leaves crowded, imbricately appressed,
spoon-shaped, more or less distinctly plicate, scarcely decurrent, broadly ovate
to ovate-oblong, apex blunt, the margin often narrowly revolute below and
broadly involute upwards, at the very apex only weakly crenulate or serrulate;
costa indistinct, or very thin, short and double; median leaf-cells narrowly
prosenchymatous, smooth, the basal shorter, laxer, porose, incrassate, yellowish
to orange-red, the alar abruptly enlarged, quadrate to shortly rectangular, or
several-angled, incrassate, colored, the alae more or less excavate; perichaetial
leaves sheathing, lance-oblong, rather abruptly acuminate, indistinctly costate:
seta 2-4 cm long, sinistrorse, tortuous, yellowish-red to red; capsule cernuous,
2-2.5 mm long, usually horizontal, symmetric, or dorsally somewhat gibbous,
drying arcuate, slightly constricted below the mouth, brownish, smooth; annu-
liis none; lid high-convex, acute or conic-obtuse.
The genus is variously delimited by different authors; as here restricted it
contains only the following species.
1. Hypnum Schreberi Willdenow, Schwaegrichen
{H. parietinum Linnaeus; H. muticum Swartz; Stereodon Schreberi
Mitten; Hylocomium parietinum Lindberg)
Plate XL VIII
Usually bright yellowish-green: stems up to 12 or 15 cm long, bright red;
stem-leaves 1.5-2.5 mm long; median leaf-cells about 10-15:1, the apical
shorter: cp.psules produced rather infrequently; exothecial cells transversely
oblong-hexagonal, laterally strongly castaneous-incrassate; peristome-teeth slen-
der, strongly trabeculate, dorsally lamellate, faintly transversely papillose-
striolate, margined, yellowish, confluent below; segments broad, nearly as long
as the teeth, widely carinately gaping, yellowish and papillose; cilia sub-
appendiculate, about as lona as the segments, usually single; the basal mem-
Jennings: Manual of Mosses— 31. Hypnaceae 259
brane about two-fifths as high as teeth; spores about .014-. 018 mm, smooth,
moderately incrassate, cast?.neous, ripe in autumn.
Grout, in his Moss Flora of North America, places this species, together
with Callkrgon cuspidatum, in Loeske's genus Calliergonella.
Mainly over humus, etc., in moist, shaded woods, but occurring in moist
pastures, dry open woods, and bogs as well; Europe, Asia, and from Arctic
America to the northern United States; as far south as Virginia in the
mountains.
Not uncommon in the elevated plateaus and m the mountains of western Pennsyl-
vania. Now known from Bedford, Somerset, Fayette, Huntingdon, Cameron, Elk, Forest,
McKean, Warren, Washington, and Westmoreland counties. Specimen figured: On de-
caying log, West Branch Swamp, Bradford, McKean Co., April 5, 1895. D.A.B.
18. Ptilium (Sullivant) DeNotaris
Dioicous: robust, stiff, la.xly cespitose, plume-like, yellowish-green, in shade
bright green, lustrous; stem 5-20 cm long, ascending to erect, simple or 2-3-
divided, regularly and densely complanately pinnate with derse complanate
branches; branches horizontally spreading, circinate at the apex, of nearly equal
length below, rapidly becoming shorter at the apex; leaves crowded, circinate
to almost coiled, deeply pluri-plicate, long-lance-subulate from a broadly ovate
and scarcely decurrent base, plane-margined, finely serrulate above the middle;
costa none, or double and short; median leaf-cells very narrowly linear, ver-
micular-prosenchymatous, smooth, the basal shorter, wider often and porose, a
few alar indistinctly differentiated, quadrate to shortly rectangular: seta 3-5
cm long, tortuous, red, drying dextrorse above; capsule cernuous to horizontal,
arcuate, about 2 mm long, castaneous, cylindric, smooth; annulus narrow,
2-seriate; lid dome-like, shortly apiculate.
The genus contains only 1 species, as follows:
1. Ptilium crista-castrensis [Linnaeus] DeNotaris
{Hypnum crista-castrensis Linnaeus; Stereodon crista-castrensis Mitten)
Plume Moss
Plate XLIX
An easily recognized, rigid, robust, plume-like, bright yellowish-green
species: stem-leaves about 2-3 mm long; median leaf-cells about 10-20:1,
branch-leaves not usually reaching 2 mm in length; e.xothecial cells strongly
castaneous-incrassate, small and rounded in several series at the rim, below
becoming oblong-rectangular; peristome-teeth castaneous, large, strongly trabec-
ulate, lamellate, crosswise faintly and finely dorsally papillose-striolate, con-
fluent below; segments as long as teeth, yellowish, papillose; the basal
membrane about one-half as high; cilia 2-4, slender, hyaline, about as long as
the segments, nodose-appendiculate; spores smoothish, castaneous, medium-
walled, about .010-.014 mm, usually mature in early autumn.
On woods-humus, rotten logs, and m.oist earth, in cool woods, usually in
mountainous regions; Europe, Asia, and from Arctic America south to t-h<»
260 American Midland Naturalist Monograph No. 6
northern United States and southwards in the mountains to North CaroHna.
Not uncommon in the more mountainous portions of our region. Now known from
Bedford, Somerset, Fayette, Washington, Blair, Cambria, Clinton, Elk, Clarion, McKcan,
and Warren counties. Specimen figured: Allegheny Mts., August 17, 1875. B. H. Pat-
terson.
19. Stereodon Bridel, Mitten
Mainly dioicous: robust to quite slender, green to yellowish-green or
golden brown, lustrous: stems elongate, decumbent or ascending, rarely erect,
mostly non-stoloniferous, simple or divided, irregularly or rarely regularly pin-
nate, the shoots mostly with hooked or circinate ends; leaves entire or serrulate,
often apparently 2-seriate, in most species falcate-secund, non-decurrent or but
slightly so, rather concave, ovate- to cordate-lanceolate, acuminate to more or
less subulate-acuminate; costa short and double or none; leaf-cells narrowly
prosenchymatous, smooth en both sides (except S. molluscum) , the basal
mostly incrassate and porose, parenchymatous in the mostly somewhat excavate
angles; inner perichaetial leaves plicate, lance-subulate: seta long, drying
twisted; capsule cemuous to horizontal, rarely erect, oblong to cylindric, arcu-
ate or rarely straight, mostly smooth and annulate; lid convex-conic, umbonate
to apiculate, or sometimes short-rostrate.
A large genus mainly confined to the temperate regions. Many authors
now include most of the species under Hypnum (See Brotherus and Grout).
This genus makes up a large part of the mossy mantle over old logs, roots,
bases of trees, roots, and earth in our woods, particularly in the southwestern
counties.
Key to the Species
A. Leaves neither distinctly falcate nor secund B
A. Leaves more or less distinctly falcate or secund C
B. Leaves entire 9. S. H aidant anus
B. Leaves distinctly serrulate, at least towards the apex S. nemorosus*
C. Leaves usually distinctly entire D
C. Leaves serrate to serrulate at least towards the apex F
D. Usually distinctly and regularly pinnate 7. S. Patientiae
D. Not distinctly regularly pinnate E
E. Alar cells gradually enlarged, not sharply differentiated; p>erichaetial leaves plicate;
capsule, when dry, curved, not plicate 8. S. pratensis
E. Quadrate alar cells numerous, inflated ones few; jserichaetial leaves not plicate; cap-
sules unsymmetric 5. S. cupressiformis
F. Alar cells strongly inflated M (see also l)
F. Alar cells not much inflated, or, if so, very few in number; or none G
G. Not regularly pinnate; leaves usually entire
5. S. ciipressiformis (See also S. pratensis)
G. Rather regularly pinnate; leaves serrate or serrulate, at least above H
H. Capsule symmetric or nearly so; a few alar cells somewhat inflated I
H. Capsule more or less curved or arcuate J
"■•" S. nemorosus (Koch) Lindberg is a southern Appalachian species extending north
to Virginia. It might occur in the mountains of Fayette or Somerset counties. Once
reported in eastern Pennsylvania.
Jennings: Manual of Mosses — 31. Hypnaceae 261
I. Capsule 3-4 mm long with apiculate or subrostellate lid
4. S. tmponens (Sc2 also S. cupressiformis)
I. Capsule 2 mm long; lid Yz as long as um 12. S. tenuirostris
}. Quadrate alar cells very numerous; 6-10 along the margin K
J. Quadrate alar cells few or none L
K. Leaves little or not serrulate; inflated alar cells few; dioicous 5. S. cupressiformis
K. Leaves distmctly serrate; no inflated alar cells; monoicous S
L. Monoicous; capsules often not plicate when dry and empty; a few subquadrate
alar cells and a few inflated angle cells 3. S. fertilis
L. Dioicous; capsules strongly plicate when dry and empty; alar cells not quadrate
but broader, incrassate, often colored 6. S. curvijolius
M. More or less regularly and evenly pinnate Q
M. Not regularly pinnate N
N. Operculum not rostrate O
N. Operculum distinctly rostrate ..-- P
O. Leaves entire or serrulate at apex; decurrcnt cells inflated; leaf-apex gradually loni-
acuminate 6. S. curyif alius
O. Leaves entire; a few alar cells abruptly mflated and hyaline; leaf-apex broadlv acu-
minate to acute or blunt 7. S. Patientiae
P. Lid about '/2 as long as the urn 10. S. recurvans
P. Lid about as long as the urn H. S. delicatulus
Q. Slender and sparsely branched; lid about j/2 as long as the urn; cilia rudimentary
or none 12. S. tenuirostris
Q. More robust and abundantly branching; lid relatively short; cilia well developed R
R. Leaves entire, broadly acuminate to acute or blunt; a number of alar cells abrJDt-
ly inflated and hyaline 7. S. Patientiae
R. Leaves entire or serrulate near apex; only a few enlarged or inflated alar cells L
S. Capsule more or less arcuate; quadrate alar cells very numerous and extending up
along the margin 10 or more cells 1. S. reptilis
S. Capsule erect and symmetric or nearly so; quadrate alar cells few; in the mountains
at higher altitudes 2. S. palles:cns
1. Stereodon reptilis (Richard) Mitten
{Hypniim reptilis Richard)
Plate XLIX
Smp.ll, dark or yellowish-green, lustrous, widely and loosely cespitose:
.^rems more or less regularly pinnate, closely interwoven; stem-leaves crowded,
lance-acuminate from an ovatc-oblong base, about 1 x 0.4-0.5 mm, falcate-
secund, slightly decurrent, serrate above, serrulate to entire below, the margins
usually revolute; median leaf-cells linear-rhomboidal to linear-flexuous, about
8-12:1, shorti^r and more or less colored towards the base, the alar cells numer-
ous, quadrate, much incrassate, sub-opaque, none inflated; branch-leaves similar
but proportionally much narrower; paraphylla small and few; costa double,
short, yellowish, or none; inner perichaetial leaves long-acuminate; usually
faintly bi-costate, apically serrate, strongly plicate: seta castaneous, lustrous,
about 1.5 cm long, when dry dextrorse above; capsules about 2.5 mm long,
cylindric, yellowish, mostly abruptly arcuate jujt below the mouth so that the
lid often points at right angles to the direction assumed by the base of the
capsule, when dry the um more or less wrinkled and narrowed below the
mouth; lid yellow, rather large, high-convex, narrowly obliquely rostrate; peri-
stome hypnoid, the teeth subulate-acuminate, orange-yellow and dorsally cross-
262 American Midland Naturalist Monograph No. 6
striolate at base, hyaline and papillose above; segments about as long as teeth,
carinately cleft between the articulations; cilia usually 2, articulate, slightly
shorter than the segmeints; annulus large, compound; spores rather strongly
incrassate, yellowish-brown, papillose, about .014-.017 mm, mature in mid-
summer; usually fruits abundantly.
On bases of trees, roots, logs, etc., in dense woods, especially in spruce
woods and mainly confined to hilly or mountainous regions; Europe, Asia,
and from Canada south in the mountains to North Carolina and in the West
to Arizona.
Erie Co.: Eight collections on Presque Isle. Nelle Ammons, various dates summers
of 1933 and 1935. McKean Co.: Several collections of D. A. Burnet, in various locali-
ties of northern McKean Co., 1895-1897. Somerset Co.: On tree root, hemlock woods,
near Buckstown, Oct. 7, 1934, and on logs and tree base, Shafer Run and Clear Run,
Sept. 14, 1946, and Oct. 6, 1935. C.M.B.; on base of dead tree, near Bakersville. C.
M. Hepner. Oct. 21, 1933. Warren Co.: Bark of sugar maple, near Big Bend.
C.M.B. Sept. 2, 1935. Washington Co.: On rotten wood, Snake Woods. Linn and
Simonton. Nov. 1892.
2. Stereodon pallescens (Hedwig) Lindberg
{Leskea pallescens Hedwig; Rhynchostegium Jamesii Sullivant; Hypnum
pTOtuberans Brotherus; Hypnum pallescens Bryologia Europaea)
Slender, in thin silky mats, more or less pinnate; stem-leaves weakly
falcate, loosely arranged, from an ovate somewhat decurrent base narrowly and
long acuminate, about 1 mm long, plane, weakly apically serrulate; costa short
and double or none; a few of the alar cells quadrate and at least the marginal
hyaline: seta 6-10 mm long, reddish; capsules about 1.5-2x0.5 mm, castane-
ous, nearly symmetric, not shrinking below the mouth when dry and empty;
lid reddish yellow, slightly rostrate; spores ripe in mid- to late summer.
Mostly on trunks and roots of coniferous trees, or on rotten wood; Europe,
and south in the mountains from Canada to Tennessee. Not yet found in
western Pennsylvania, but to be expected.
3. Stereodon fertilis (Sendtner) Lindberg
(^Hypnum fertile Sendtner)
Plate XLIX
Yellowish-green, lustrous, usually darker below, densely interwoven: stems
prostrate or ascending, from 3-10 cm long, scarcely branching, castaneous,
densely and rather regularly, complanately and somewhat plumosely pinnate
with short branchlets; stem-leaves 1.5-2.0 mm long, concave, falcate-secund,
scarcely complanate, from an oblong-ovate base slenderly acuminate, the base
not decurrent, sub-auriculate, somewhat excavate, the margin plane, entire
below, serrulate ibove; costa usually bi-furcate or double, rarely none; branch-
leaves similar to stem-leaves but smaller and narrower, strongly falcate to
circinate-secund; inner perichaetial leaves lance-acuminate, plicate, faintly bi-
costate, serrulate above; median leaf-cells linear-vermicular, about 1:15-20,
rather incrassate, sub-acute, median basal cells strongly incrassate, castaneous-
Jennings: Manual of Mosses — 31. Hypnaceae 263
pellucid, porose, the alar portion with several oblong inflated hyaline cells,
above which is a patch of about 9-15 smaller, incrassate, quadrate to oblong-
hexagonal cells often more or less castaneous-pellucid: seta 1.5-2.0 cm long,
de.xtrorse, castaneous, lustrous; capsule about 1.8 mm long, oblong, inclined
to horizontal, arcuate, bright castaneous with a darker and lustrous rim, when
dry narrowed below the mouth and sub-costate or smooth; lid short, conic-
acute; peristome-teeth lance-subulate, brownish-pellucid, more or less narrowly
hyaline-marained, dorsally cross-striolate below, the divisural line and the
lamellae fairly distinct, above hyaline and papillose, the trabeculae numerous
and strong; segments pale, about as long as the teeth, carinately split between
the articulations, the basal membrane about two-fifths as high, the cilia usually
2, hyaline, nodose, somewhat shorter than the segments; exothecial cells rather
thin-walled, quadrate-hexagonal to oblong-hexagonal, at the mouth smaller
and deeply castaneous; spores medium-walled, somewhat brownish, smooth or
very nearly so, about .015-.018 mm, m.-^.ture in summer.
On decaying logs in moist and cool places, usually in hilly or mountainous
regions; Europe, Asia, .='.nd, in North America, from New Brunswick to
British Columbia and southwards to the northern United States.
Armstrong Co.: West bank Allegheny River, Foxburg. June 10, 1934. C.M.B.
Fayette Co.: Laurel Run, 2 mi. e. of Wymp's Gap, elev. 2100 ft. Aug. 24, 1940.
C.M.B. McKean Co.: Leer's Run on decaying log, August 5, 1895 (figured). Lang-
made Hollow, August 11, 1895. Gates' Hollow, October 27, 1895, and Bennett Brook,
August 8, 1897. D.A.B. Somerset Co.: Cranberry Glade Run, Laurel Hill. On log
in swampy woods. June 22, 1947. C.M.B.
4. Stereodon imponens (Hedwig) Lindberg
(Hypnum imponens Hedwig; H. cupressiforme Hooker)
Log Moss
Plate XLIX
Robust in flat, thin, widely interwoven tufts of a yellow-green color: stems
rigid, reddish-brown, with numerous paraphyllia, closely, rather regularly and
more or less complanately pinnate, prostrate or sub-erect, sometimes reaching
more than 10 cm in length; stem-leaves usually somewhat complanate-secund.
from a triangular-oblong base gradually long-acuminate, the base not e.xcavate,
scarcely decurrent, the acumen strongly falcate-secund, the whole leaf about
2 x 0.5-0.7 mm, serrulate above, the margin often narrowly recurved below;
branch-leaves narrower, otherwise similar to the stem-leaves, about as long;
costa short and double or none; median leaf-cells linear- vermicular, about 10-
15:1, the basal broader and more or less orange-pellucid, the alar cells sub-
quadrate, a few somewhat inflated at the extreme angles, forming a small but
quite distinct auricular patch of an orange-brown color, all cells rather incras-
sate; perichaetial leaves plicate, ecostate, spinose-serrulate above: seta about
3 cm long, castaneous, sinistrorse when dry; capsule cylindric, nearly erect,
slightly curved, about 3-4 mm long, castaneous, about 4-6:1; lid convex at
base with an oblique long-acuminate point; peristome normally hvpnoid, the
teeth strongly trabeculate, the trabeculae often dividing, the lamellae and
264 American Midland Naturalist Monograph No. 6
divisural distinct, cross-striolate below; the segments about as long, slightly
carinately split, the basal membrane reaching about two-fifths as high, the cilia
articulate and usually single; annulus compound, adherent; exothecial cells
yellowish-pellucid, laterally quite incrassate, oblong-quadrate to long-rectangu-
lar; spores yellowish, medium-walled, mmutely roughened, about .013-.015
mm, mature in late autumn or early winter.
This is commonly sold in the Pittsburgh markets at Christmas time in
sheets and called "Log Moss."
On earth, stones, roots, logs, etc., with us mainly on humus or rotten
wood, in moist woods; Europe, Asia, and from Canada southwards to Cali-
fornia and Georgia.
Very common in our region. Now known in 15 counties in western Pennsylvania and
probably occurring in all. It is particularly abundant in the southwestern counties grow-
ing in sheets over decaying logs. Specimen figured: Wildwood Hollow, Allegheny Co.,
Nov. 19. 1908. O.E.J.
5. Stereodon cupressiformis [Lmnaeus] Lindberg
{Hypnum cupressiforme Linnaeus; H. compressum Schuitz)
Widely cespitose in flat, soft tufts, usually yellowish or brownish-green:
stems up to 10 cm long, greenish, procumbent, irregularly pinnate, the branches
spreading or ascending, usually curved; leaves closely imbricate, concave at the
base, not decurrent, falcate-secund, lustrous, oblong- to ovate-lanceolate, nar-
rowed rather abruptly to a long acumination, plane-margined, typically entire,
sometimes denticulate towards the apex; costa none or very short and double;
median leaf-cells about 10:1, linear-vermicular, rather obtuse, the angular
sub-quadrate, numerous, rather opaque, a few at the extreme angles larger,
scarcely inflated, somewhat orange-pellucid, or hyaline, rather incrassate, the
auricles not well-defined; perichaetial leaves denticulate, not plicate: seta red,
about 3-4 cm long; capsule sub-erect, curved, sub-cylindric or oblong, castane-
ous, somewhat constricted below the mouth when drv; lid convex at base, with
an acuminate or sub-rostrate apex; peristome normally hypnoid; cilia usually
one or two; spores mature in late autumn or early winter.
On soil, rocks, roots, and bases of trees, in moist woods or ravines; practi-
cally cosmopolitan, — in North America occurring from the Arctic regions to
the Gulf States. Rare in our region. Quite variable. There are a number of
named varieties.
Cambria Co.: T. P. James. (Porter's Catalogue). McKean Co.: Three pockets
so-labeled in the Carnegie Muscim Herbarium are apparently typical S. fertilis and the
Porter Catalogue record is probably founded upon some of the same Burnett collections.
5a. Stereodon cupressiformis var. filiformis (Bridel) Jennings
(Hypnum cupressiforme var. filiforme Bridel)
Plate L
More slender, distantly pinnate, the branches long, very slender and almost
filiform; leaves very small, falcate-secund, more or less regularlv and neatly
imbricated in two rows.
Habitat and range as for the species.
Jennings: Manual of Mosses — 31. Hypnaceae 26'>
Rare in our region. McKean Co.: Rutherford Rocks, on moist and densely shaded
fragments of sandstone at base of cliff, May 5, 1895, and Hawkin's Hollow, August 2,
1895. D.A.B. (figured).
6. Stereodon curvifolius (Hedwig) Mitten
{Hypnum curvifolium Hedwig)
Plate L
Robust, lustrous, yellowish-green in large flat mats: stems prostrate, rather
regularly pinnately branched, the branchlets short and unequal, the whole
plant complanately secund; leaves crowded, imbricate in two rows, falcate-
serund, thus giving to the plants a plaited appearance from the dorsal view-
point; stem-leaves about 0.7-0.8x1.4-1.8 mm, oblong-ovate to elongate and
triangular-ovate, long-acuminate, plane-margined, crenulate-serrulate about to
the middle, and at the angles, concave, at the base abruptly narrowed and
cordate or sub-cordate, somewhat decurrent; costa none or double and faint;
branch-leaves similar but proportionally shorter and narrower, about 0.4-0. 5-x-
1-1.5 mm; median leaf-cells about .005-.007 x .035-.050 mm, linear-vermicular,
basal median cells incrassate, porose, more or less vermicular to linear-oblong,
a few of the alar cells sub-quadrate, yellowish or brownish and incrassate,
about .020-.025 mm in diameter, the decurrent cells enlarged, thin-walled, and
hyaline; perichaetial leaves erect, whitish, numerous, the inner sheathing, pli-
cate, reaching 4-5 mm long: seta about 2.5 cm long, dextrorse above, sinis-
trorse below, castaneous, sub-lustrous; capsule about 2.5 mm long, pale cas-
taneous, constricted below the mouth when dry and empty and then also
strongly plicate, the urn oblong, arcuate, cemuous; lid conic, apiculate; peri-
stome normally hypnoid, teeth yellov/ish pellucid, slender, strongly trabecu-
late, the lamellae and divisural distinct, the apical portion of the teeth hyaline
and papillose, the lower dorsal surface cross-striolate; segments about as long
as the teeth, slender, slightly carinately cleft, pale yellowish-pellucid, cilia two
or three, about as long as segments, articulate, hyaline, papillose; annulus 3-
seriate, revoluble; spores yellowish, medium-walled, granulose, about .019-.023
mm, mature in early spring.
On soil, rocks, or more usually on decaying logs in moist, cool woods;
Asia, and from Arctic America south to Georgia, Missouri, and Colorado.
Common in our region. The type was from Lancaster, Pennsylvania.
Now known from the following counties: Allegheny, Armstrong, Beaver, Butler, Cam-
bria (Porter), Crawford, Erie. Fayette, McKean. Mercer, Somerset, Washington, and
Westmoreland. Specimen figured: Pymatuning Swamp near Linesville, Crawford Co.,
May 12, 1908.
7. Stereodon Patientiae Lindberg
{Hypnum arcuatum Lindberg; H. LinJbergii Mitt.; H. Patientiae
Lindberg; Stereodon arciiatus Lindberg)
Plate L
Robust, widely cespitose in yellowish-green mats, usually more or less
regularlv pinnate, lustrous at least on the younger parts: stems prostrate,
usually sparsely branched; stem-leaves about 1.6-2.2 mm long, complanately
falcate-secund, lustrous, ovate-oblong, decurrent, the margins plane and entire
266 American Midland Naturalist Monograph No. 6
or sub-denticulate at the apex, the leaves rather shortly and broadly acuminate,
the tip flat and widely acute to somewhat obtuse, the decurrent auricles are
made up of large, oblong, inflated, thin-walled, and hyaline cells bordered
above by about two series of smaller, quadrate, usually brownish-pellucid,
thicker-walled cells, these latter grading quickly into linear-vermicular median
cells about 12-20:1, the apical cells oblong-rectangular or obliquely more or
less rhomboidal; costa very short, and double or none; branch-leaves similar;
perichaetial leaves sheathing, the inner lanceolate to lance-linear, up to 6 or 7
mm long, plicate, entire, acuminate; seta about 2.5-3 cm long, dextrorse, lus-
trous, castaneous; capsule about 2-2.5 mm long, about 4-5:1, oblong-cylindric,
erect at base but arcuate so that the lid usually points more or less horizon-
tally, when dry plicate but scarcely narrowed below the mouth; annulus large,
revoluble; lid conic, apiculate, scarcely longer than wide; peristome hypnoid.
the teeth lance-linear, dorsally cross-striolate, yellowish-pellucid below, hyaline
and papillose above, the divisural and dorsal lamellae prominent, the trabeculae
.<:trong and numerous; segments rising from a basal membrane about two-fifths
the height of the teeth, the segments about as long as teeth, narrow, somewhat
carinately split; cilia 1-3, shorter, nodose, hyaline-papillose, often joined to-
gether above; spores smoothish, yellowish, moderately incrassate, about .014-
.018 mm, mature in late spring or early summer.
On the ground in woods and wet, grassy places in swamps, around springs,
etc.; Europe, Asia, and from Arctic America to the northern United States
and south, in the East, to Florida.
Now known from 13 counties in western Pennsylvania and probably occurs in all.
Specimen figured: Kittanning, Armstrong Co., May 28, 1907. O.E.J.
8. Stereodon pratensis (Koch) Wamstorf
(Hypnum pratense Koch; Isopterygium pretense Lindberg)
Plate LXX
Softly and flatly cespitose, bright green, complanately flattened: stems
prostrate to sub-erect, non-radiculose, irregularly sub-pinnate, branchlets rather
sparse; leaves sub-secund on the larger branches and on the stems, plane to
somewhat concave, entire; costa double and very faint and short or none;
median leaf-cells narrowly rhomboid-vermicular, the alar rather gradually
enlarged, fewer, less enlarged and less difl^erentiated than in S. Patientiae; peri-
chaetial leaves plicate, the inner long-lanceolate and shortly acuminate: pedicel
long, twisted in two directions; capsule non-plicate, oblong to turgid-ovate,
cemuous, arcuate when dry; lid convex-conic; annulus 3seriate; peristome
normally hypnoid, the cilia 3, about as long as the segments; spores mature in
spring. The capsules are rarely produced.
Specimens sometimes show an abruptly enlarged group of alar cells typical
of S. Patientiae in the larger stem-leaves, while less vigorous leaves are of the
S. pratensis type, as in the Linn & Simonton specimen, see figure, Plate LXX.
In open swamps and marshy meadows, Eurasia, and from Pennsylvania
and New Jersey to Arctic America. Varieties of it range farther south.
Jennings: Manual of Mosses — 31. Hypnaceae 267
Cambria Co.: Wiltmore. T. P. James. (Porter's Catalogue). McKean Co.: Near
Bradford. D.A.B. WASHINGTON Co.: McCracken Sta., W. & W. Ry., Oct. 21, 1893.
Linn & Simonton (figured).
9. Stereodon Haldanianus (Greville) Lindberg
(Hypnitm Haldianum Greville; Heterophyllon Haldani
Kindberg; Hypnum pulchrum Hooker)
Plate L
Widely and loosely cespitose, dark to brownish-green: stems long, creeping,
irregularly pinnate, the branchlets unequal and disposed much as in some of
the Brachytheciae; leaves loosely and more or less evenly imbricate to loosely
spreading; stem-leaves usually decurrent, about 0.7-1.5 mm long, oblong-ovate
to somewhat lanceolate, rapidly narrowed to a short and acute apex, entire,
plane-margined, concave; branch-leaves lance-ovate to lanceolate, about 0.5-1.5
mm long, short-acuminate, otherwise similar to the stem-leaves; median leaf-
cells linear-flexuose, about 12-20:1, prosenchymatous, the alar inflated, rather
incrassate, large, forming a quite distinct auricle, bordered above by a few
considerably smaller and chlorophyllose cells; costa rudimentary or none, or
double and short; perichaetial leaves spreading, abruptly filiform-acuminate,
the inner non-plicate; paraphyllia large and numerous: seta about 2 cm long,
lustrous, castaneous, when dry sinistrorse; capsule long-cylindric, dull-castane-
ous, curved, sub-erect to more or less inclined, about 4-6:1, urn about 2.5-3
mm long, the lid conic and obliquely short-rostrate; peristome-teeth confluent
at base, transversely striolate and yellowish below, strongly trabeculate, the
divisural and the dorsal lamellae usually faint; segments slightly carinately
cleft, below more or lei>s faintly transversely striolate-papillose, above papillose,
about as long as the teeth; cilia usually single and shorter, sometimes two and
rudimentary, or sometim.es none, articulate; spores granulose, yellowish-brown,
somewhat incrassate, about .015-.016 mm, mature in late fall or winter.
On earth, humus, rocks, rotten logs, etc., in woods; Europe, Asia, and, in
North America, from Nova Scotia to Montana and southwards to the Gulf
States.
Common in our region. Known from numerous collections from the following coun-
ties: Allegheny, Armstrong, Beaver, Butler, Cambria (Porter), Crawford. Elk (Porter).
Erie, Fayette, McKean, and Somerset. Specimen figured: Wildwood Road Hollow, Alle-
gheny Co., Nov. 19, 1908. O.E.J, and G.K.J.
10. Stereodon recurvans [Richard] Brotherus
(Hypnum recurvans Schwaegrichen: Sematophyllum recurvans E. G. Britton;
Leskea recurvans Richard; Rhapidostegium recurvans Bryologia
Europaea; Brolherella recurvans Fleischer)
Plate LI
Very glossy, widely cespitose in flat tufts, usually yellowish-green: stems
prostrate, reddish, irregularly pinnate; leaves about 1.2-1.5 mm long, strongly
complanately falcate-secund, close, imbricate at base, soft, thin, more or less
268 American Midland Naturalist Monograph No. 6
concave, lance-ovate, slenderly long-acuminate, non-decurrent, more or less
sharply serrate at the apex, the margin often narrowly recurved below; costa
obsolete, or very short and double; perichaetial leaves gradually long-acumi-
nate, serrate at apex; median leaf-cells linear-flexuose, the basal yellowish or
brownish, shorter, wider, the alar consisting of a group of 4 to 8 hyaline or
colored, much inflated and enlarged cells forming a group bordered above by
a few sub-quadrate and smaller cells: seta about 1.5-2 cm long, lustrous, cas-
taneous, somewhat sinistrorse; capsule oblong-oval, slightly curved, obliquely
inclined to almost horizontal, the urn about 3-4:1, about 1.5-2 mm long, light
castaneous, the tapering base darker, the urn when old strongly arcuate; annu-
lus present; lid conic and together with the slender beak about one-half the
length of the urn; peristome hypnoid, the teeth large, strongly trabeculate, the
divisural faint, the thin dorsal lamellae transversely papillose-striolate; segments
about as long as teeth but usually not splitting, the basal membrane about two-
fifths as long, the cilia usually one, sometimes two, slightly appendiculate,
somewhat shorter than the segments; spores .016-.018 mm, medium-walled,
granulose, brownish, mature in late fall, the capsules often remaining in good
condition until early spring: dioicous.
On soil, humus, bases of trees, logs, etc., in moist woods, mainly in moun-
tainous or hilly regions; from Newfoundland to Manitoba and south in the
mountains to Georgia.
Common in our region. Known from eleven counties in western Pennsylvania and
probably occurs in all, although most abundantly collected m the southwestern counties.
Specimen figured: As to old capsules and peristome, Ohio Pyle, Fayette Co., May 30-31,
1908. O.E.J. As to other figures, Ohio Pyle, Sept. 1-3, 1906. O.E.J. & G.K.J.
11. Stereodon delicatulus (James) Brotherus
(Hypnum laxepatulum Lesquereux and James; Rhynchostegium delicatulum
James; Sematophyllum delicatulum E. G. Britten; Rhapidostegium
deliculatum Paris; Brotherella delicatula Fleischer)
Small, depressed cespitose, dark green, scarcely lustrous: stems slender,
prostrate, subpinnately branching; leaves rather open, mostly falcate-secund,
two- ranked, sharply serrulate towards the apex; costa none or very short and
double; leaves concave, ovate, narrowed into a long acumination; a few of the
extreme alar cells much enlarged and inflated as in S. recurvans, colored or
hyaline, bordered by a few sub-quadrate and smaller, the median linear-flexu-
cus, prosenchymatous; perichaetial leaves non-plicate, sharply serrate above:
seta shorter than in S. recurvans; capsule about 1-1.5 mm long, ovoid-oblong,
almost erect and only slightly curved, about 2-3:1, urn about equalled in length
by the slenderly long-rostrate lid; peristome hypnoid, segments entire, cilia
usually one or two; spores mature in late fall.
On rotten wood, or soil, or at the base of trees, mainly in the mountains
from New England to Alabama.
Rare in our region. McKean Co.: Bradford. D. A. Burnett. (Porter's Catalogue).
Jennings: Manual of Mosses — 31. Hypnaceae 269
12. Stereodon tenuirostris (Bruch and Schimper) Brotherus
(Sematophyllttm temthostTe E. G. Britton; Hypnum cylindrocarpum
C. Mueller; Rhaphidostegmm cylmdricarpum Jaeger)
Plate LI
Flatly and broadly cespitose in thin intricate mats, slender: stems prostrate,
reddish, or green, pinnately branched, branches few, slender; stem-leaves sub-
lustrous, about 1.5 mm long, falacte-secund but not complanate, narrowly
lance-oblong, non-decurrent, acuminate, concave, apically serrate, marginally
somewhat refle.xed to the base of the acumen; median leaf-celLs linear-prosen-
chymatous, the apical usually a little larger, the alar few in number, inflated,
sub-quadrate, bordered above by a few small quadrate, sub-opaque, often
transversely elongated cells; branch-leaves similar to the stem-leaves, some-
times a little larger; costa very short and double or none; perichaetial leaves
erect, the inner plicate and gradually narrowed to a very slender serrate point,
with a very short and double costa or none: seta about 5-7 mm long, sinis-
trorsc above, lustrous, castaneous; capsule cylindric to lance-oblong, the urn
about 1-1.5 mm long, erect to somewhat inclined, symmetric; annulus no.ne;
e.xothecial cells somewhat collenchymatous, brownish, oblong-rectangular, the
upper 3 or 4 rows rounded-quadrate; peristome-teeth yellowish, lance-subulate,
finely cross-striolate, strongly trabeculate, the dorsal lamellae projecting to
form a rather conspicuous hyaline border; segments about thre;-fourths as long,
slender, carinately split between the articulations, the basal membrane about
one- third as high as the teeth, cilia none or very rudimentary; lid conic and
with a slender rostrum about one-half as long as the urn; spores in late fall
to early spring, about .014-. 018 mm, smoothish, brownish, rather thinly in-
crassate.
On rotten logs and bases of trees and on rocks in dark woods, in the moun-
tains from North Carolina and Georgia north to New York and Indiana.
Allegheny Co.: Wildwood Road Hollow, side of ravine under dense shade of
hemlocks, November 19, 1908. O.E.J. Cameron Co.: On wet rocks 3 mi. w. of Tru-
man. June 22, 1935. Sidney K. Eastwood. Fayette Co.: On rock in shaded woods in
valley of Meadow Run, four imles south of Ohio Pyle, September 1-3, 1906. O.E.J, and
G.K.J. Westmoreland Co.: "Shades," near Blackburn, March 25, 1910. O.E.J, and
G.K.J, (figured).
20. IsoPTERYGiUM Mitten
Autoicous or dioicous: mostly slender to verv slender, cespitose. soft,
mostly bright or yellowish-green and glossy: stem creeping to ascending, up-
right only in the thick mats, mostly irregularly branched; leaves uniform, most-
ly complanately and obliquely inserted, smooth, usually more or less two-
seriate, from a narrow and little or not at all decurrent base oval to oblong
and short-pointed or else ovate to lance-oblong and acute to piliferous, margins
plane and entire to serrate; costa double, very short, or none; cells prosen-
chymatous, smooth or papillose in the upper angle, the basal shorter, the alar
not usually differentiated: seta long, sm.ooth, mostly drying twisted; capsule
sub-erect to cernuous or horizontal, with a collum, oval to oblong or cylindric,
270 American Midland Naturalist Monograph No. 6
almost symmetric or weakly gibbous, when dry only rarely arcuate and nar-
rowed below the mouth, mostly smooth; annulu5 present or none; peristome-
teeth basally confluent, subulate, yellowish, mostly hyaline-bordered, with
divisural zigzag, cross-striate, apically hvaline and papillose, lamellae numer-
ous; cilia 1-2, nodose, rarely 3 and appcndiculate; lid conic-convex, sometimes
tostrate.
A genus of world-wide distribution and containing about 170 species, most-
ly occurring on decaying wood; about 25 species reported for North America;
about 7 species in our region.
Key to the Species*
A. Leaves entire or only obscurely serrulate at apex B
A. Leaves usually distinctly serrate or serrulate at least in upper half E
B. Leaves perfectly entire C
B. Leaves obscurely serrulate at apex D
C. Cells about .005-.008 x .075-.160 mm, about two rows at the base shorter and sub-
oval; cortical stem-cells not enlarged and hyaline 5. /. pulchellum
C. Cells about .003-.005 x .080-.100 mm, hardly different at base; cortical cells of stem
enlarged and hyaline 1. /. Muellerianum
D. Alar cells few, quadrate, forming a small group 6. /. micans
D. Alar cells very mdistinctly sub-rectangular, not forming a well defined group
2. /. elegans
D. See also /. pulchellum var. nitidulum
E. Leaves shortly bi-costate; annulus large and compound; leaf-cells papillose by pro-
truding cell angles 4. /. geophilum
E. Leaves ecostate or obsoletely costate F
F. Leaves serrulate to the base or nearly so G
F. Leaves not serrulate below the middle /. micans
G. Operculum conic; leaf-cells not papillose dorsally towards leaf-apex; leaves compla-
nately spreading 7. /. turfaceum
G. Operculum short-rostrate; leaves closely complanately overlapping 3. /. deplanatum
1. Isopterygium Muellerianum (Schimper) Lindberg
(Plagiothecium Muellerianum Schimper)
Yellowish-green, laxly cespitose; the branches long, flattened, straggling
out into flagella or stolons or forming flattened strands, very slender; the
stems and branches thick, often more than .150 mm in diameter, with very
large and thin-walled outer cells which are 3 or 4 times as wide as the lower
cells of the leaf and usually about .015-.025 mm wide; leaves rigid, not much
different when dry, the p)oints directed forwards and upwards so that the
dorsal surface of the branch is concave, lance-ovate, long-apiculate, concave,
non-decurrent, plane-margined, entire; costa double, very faint and short;
median cells linear, narrow, up to .100 mm long, about 20-25:1, the alar and
basal scarcely different; perichaetial leaves ovate-oblong, acuminate, entire: seta
rather short, cistaneous; capsule smooth, small, long-necked, obovoid, erect to
inclined, when dry pale brown, wide-mouthed and campanulate; lid conical,
''' Also /. subfalcatum Austin, with sub-cultirform leaves turned backwords, non-decur-
rent, apically serrate. In cracks and crevices of rocks, Pennsylvania to New England.
See Sullivant's Icones Muscorum. Supplement, plate 67.
Jennings: Manual of Mosses — 31. Hypnaceae 271
rostellatc; peristome-teeth with rather distant articulations, ciha short and
unequal; annulus 1-seriate; spores mature in autumn; dioicous.
On moist earth and rocks in cool ravines, etc., mostly in hilly or moun-
tainous regions; Europe, Asia, and from northeastern Canada to Ohio and
Minnesota, and southward in the mountains to North Carolina. Not yet
reported from^ our region but to be expected, — especially in the more moun-
tainous portions.
2. IsoPTERYGiUM ELEGANS [Hooker] Lindberg
(Hypnum elegan<: Hooker; Isothecium elegans Bridel;
Plagiothecium elegans Suliivant)
Plate LXVIII
Small, densely cespitose, thin, pale shining green: stems branching irregu-
larly, the branches slender, partly procumbent, partly ascending, numerous,
usually pointing one way; usually there are also axillary, gemmiferous branch-
lets; leaves complanate, sub-distichous, the points usually pointing downwards,
lustrous, little changed when dry, about 1-1.5 mm long, lance-oblong to ovate-
oblong, rather gradually narrowed from about the middle, then abruptly nar-
rowed to a fine, short acumen, the base rounded, non-decurrent, plane-mar-
gined, entire except at the acumen where a little denticulate; costa double and
short, faint or sometimes reaching one-third the length of the leaf; median
leaf -cells narrowly linear, about 20-30:1, about .004-. 007 mm, pointed, hardly
differentiated at the angles; perichaetial leaves lance-acuminate: capsule turgid-
ovoid, slightly inflated at the curved neck, nearly symmetric but horizontal or
sub-pendent by a curve in the upper part of the seta, when dry and empty
somewhat wide-mouthed, turbinate, costate; peristome hypnoid, yellow, teeth
broadly lanceolate, blunt, segments entire, cilia 3, rather slender, as long as
the segments; annulus simple; lid conic, obtusely pointed; spores mature in
spring.
On rocks or earth usually in moist, cool crevices of ledges, etc., in hilly
or mountainous regions in woods: Europe, Asia, and from Arctic America to
northern United States and south in the mountains to North Carolina and
Tennessee.
Armstrong Co.: At small waterfall, west bank Allegheny River. 2 mi. n. of Parkers
Landing. C.M.B. June 10, 1934. McKean Co.: D. A. Burnett. (Porter's Catalogue).
Westmoreland Co.: On rock clifF, Meadow Run at Jacobs Creek. C.M.B. Aug. 18,
1945 (figured).
3. IsoPTERYGlUM DEPLANATUM (Sullivant) Mitten
(Hyprium deplanatum Sullivant; Rhynchostegium deplanatum Schimper);
Plagiothecium deplanatum Grout)
Plate LXIX
Golden-green, lustrous, small, flattened: stems prostrate, irregularly pin-
nately branching; leaves 2-ranked, imbricate, giving stems and branches a
plaited appearance, thin, concave, ovate-lanceolate, gradually long-acuminate,
s-errulate all around, more sharply so above, plane-margined; median leaf-cells
272 American Midland Naturalist Monograph No. 6
linear, flexuous, prosenchymatous, the basal somewhat larger but very similar,
a small group of short-rectangular or quadrate cells at the angles; costa none
or but very faint: capsule oval-oblong, cernuous, arcuate, plicate when dry
and then constricted below the mouth; peristome hypnoid, segments narrow,
cilia 2 or 3, about as long as the segments, unequal; annulus none; spores in
autumn, but capsules rarely produced.
Over earth, stones, and rotten wood and humus, in woods; from Nova
Scotia to Manitoba and southward to Missouri and Tennessee.
Rare in our region. McKean Co.: D. A. Burnett. Porter's Catalogue). Washing-
ton Co.: On rock, near Washington. December 5, 1891 (figured). A. Linn and J. S.
Simonton.
4. IsoPTERYGiUM GEOPHILUM (Austin) Jaeger
(Rhynchostegium geophilum Austin; Plagiothecium geophilum Grout;
Hypnum depressum Sullivant and Lesquereux)
Plate LXVIII
Dark green, very glossy, thinly, softly, and loosely matted, leafy branches
about 2.5-3 mm wide; stems prostrate, irregularly divided, more or less com-
pressed; leaves long, somewhat concave, distichous, rather distant, widely
spreading, ovate to oblong-lanceolate, gradually and symmetrically narrowed
to a somewhat blunt af>ex, serrulate above, rounded at the base, non-decurrent;
costa short, double, rather distinct or none; median leaf-cells linear, prosen-
chymatous, flexuous, about 8-12:1, a few alar sub-rectangular, thick-walled,
only a little enlarged and not forming a distinct auricle, upper cells shorter,
papillose dorsally by projecting cell-wall angles; many branches of our speci-
mens (July 26, 1947) were tipped with capitate clusters of linear, often forked
gemmae: capsule small, ovate, gibbous, thin-walled, unsymmetric, inclined;
urn about 1 mm long; peristome normally hypnoid, teeth yellowish, segments
narrow, linear, the cilia 2 or 3, some as long as segments; annulus large, 2-
seriate; lid conic, obliquely long-rostrate; spores mature in spring or summer.
On moist earth or stones, usually near water in lowlands or in shady
ravines; occurring from New York to Wisconsin and south to Maryland and
Georgia.
Cambria Co.: Cresson. T. C. Porter. (Porter's Catalogue). Somerset Co.: Becic
Spring, Laurel Ridge. C.M.B. July 26, 1947 (figured).
5. IsoPTERYGiUM PULCHELLUM (Dickson) Jaeger
(Plagiothecium pulchellum Bryologia Europaea; P. pseudo-latebricola
Kindberg; Leskea pulchella Hedwig)
Slender, in prostrate and straggling tufts, bright glossy metallic green:
stems creeping, usually not much more than 1 cm long, the branches numer-
ous and slender, erect or curved-ascending; leaves subdistichous, about 1 mm
long, more or less falcate at tips of stems and branches, very glossy, hardly
altered when dry, entire, plane-margined, narrowly lance-ovate, from near the
base rounded but not decurrent nor excavate; costa usually none; median leaf-
cells narrowly linear, about 12 25:1, .0O5-.(X)8 mm wide, pointed, the basal
Jennings: Manual of Mosses — 31. Hypnaceae 273
suboval and wider and shorter but not differentiated otherwise at the angles:
monoicous; seta reddish, slender, about 1-1.5 cm long; capsule small, rather
variable, ranging from oblong and tapering below into the neck to short and
ovoid, and from erect and symmetric to curved and more or less horizontal,
when dry usually wide-mouthed and constricted below the rim, ranging from
greenish-brown when young to castaneous when old; lid conic, apiculate; peri-
stome-teeth densely barred, cilia 2, a little shorter than the segments; annulus
2-seriate; spores mature in early summer.
On rocks and roots of trees and on rotten wood, in moist woods; Europe,
Asia, and from Arctic America to the northern United States as far south as
Pennsylvania.
IsOPTERYGlUM PULCHELLUM var. NfTlDULUM (Wahlenberg) Brotherus
(Hypnum nitidulum Wahlenberg)
Larger plants in thinner mats, with often stoloniferous stems and more
complanate branches; leaves with very slender acuminations, often serrulate
above; capsule almost horizontal; annulus 1 -seriate often falling still attached
to the operculum. Range about the same as the species.
Elk Co.: Benezette. McMinn. (Porter's Catalogue).
6. IsOPTERYGlUM MICANS (Swartz) E. G. Britton
(Hypnum albulum C. Mueller; H. micans Swartz; Sematophyllum micans
Braithwaite; Plagiothecium micans Paris)
Small, thinly matted, loose, glossy, whitish-green to yellowish-green: stems
prostrate, rooting, irregularly branching; leaves loose, erect-spreading to secund
and pointing downwards, very small, about 0.8-1.2 mm long, ovate-lanceolate,
gradually long-acuminate, serrulate above, thin; costa usually none; perichaetial
leaves abniptly acuminate, the inner apically serrate; median leaf-cells linear,
prosenchymatous, flexuous, about 15:1, at the base a row considerably
enlarged and broad, at the angles a few sub-quadrate: seta long and slender;
capsule very small, ovate-oblong, light castaneous, constricted below the mouth
when dry and empty, slightly incurved; peristom.e-segments not split, about as
long as teeth, the cilia 1 or 2, short, nodose; annulus none; lid conic, apiculate
to short-rostrate; spores mature in mid-winter.
On earth and rotten wood in moist woods; mainly along the eastern United
States from New York southwards.
Rare in our region. Centre Co.: Bear Meadows. T. C. Porter. (Porter's Cata-
logue ) .
7. IsOPTERYGlUM TURFACEUM (Lindberg) Lindberg
{Hypnum turfaceum Lindberg; Stereodon turfaceuf Mitten;
Plagiothecium turfaceum Lindberg)
Plate LI
Small, light green to yellowish-green, loosely matted: stems prostrate, more
or less pinnately branching with short branches, rooting at the perichaetia and
274 American Midland Naturalist Monograph No. 6
at the main forks and usually quite difficult to separate from the substratum
without breaking in pieces; leaves about 1.5 mm long, complanately arranged,
lance-ovate, long-acuminate from an ovate-oblong base, sharply serrate above
the middle, margins plane, serrulate, or entire towards the base; costa none
or very short and faint; oerichaetial leaves ovate, basally concave, abruptly
short-pointed, dentate at apex; median leaf-cells fusiform to broadly linear,
prosenchymatous, about 8-12:1, the basal a little shorter and wider, the alar
either not differentiated or a few sub-quadrate to rectangular and incrassate:
seta slender, about 1.5 cm long, castaneous, somewhat twisted; capsule oblong,
about 2-3.1, about 2 mm long, slightly curved and somewhat inclined when
young, when old and empty arcuate, horizontal, plicate, castaneous, and con-
stricted below the mouth; annulus large, double; lid conic; exothecial cells
small and rounded in three or four series at the rim, gradually becoming
oblong-rectangular or irregular-oblong below, the upper more or less distinctly
rastaneous-collenchymatous; peristome small, the teeth lance-subulate, papil-
lose above, dorsally transversely striolate below, strongly trabeculate and
lamellate, slightly confluent at base; segm.ents nearly as long, narrow, not split,
papillose, yellowish, basal membrane about two-fifths as high; cilia strong,
nodose, often about as long as the segments, 1 or 2 in number; spores more
or less greenish-yellow, about .008-.011 mm, papillose, rather thin-walled,
mature in summer.
On rich woods-humus in moist woods or in peat bogs; Europe, and from
Canada to Georgia and Texas.
Cambria Co.: Ebensburg. T. P. James. (Porter's Catalogue). Crawford Co.:
In swamp near Hartstown, May 29-31, 1909. O.E.J, and G.K.J, (figured). McKean
Co.: East. Branch Swamp, Bradford, July 1, 1896. D.A.B.
21. Plagiothecium Bryologia Europaea*
Autoicous or dioicous, rarely polyoicous: usually more or less robust, most-
ly softly cespitose, bright to yellowish or whitish-green, lustrous: stems creep-
ing to ascending, or in thick cushions erect, with ascending and small-leaved
stolons, mostly irregularly branched; branches often elongate-flagelliform;
paraphyllia none; leaves uniform, obliquely inserted, non-plicate, some species
complanate-distichous, concave from a narrow and more or less decurrent base,
broadly lanceolate to ovate, acuminate, mostly plane-margined and entire,
sometimes serrate; costa short, mostly double, sometimes none; median leaf-
cells chlorophyllose, elongate-rhom.boid to linear, thin-walled, the basal shorter
and wider, the alar lax and hyaline: seta long, reddish, drying twisted; capsule
erect to cernuous, with collum, oblong to cylindric, symmetric to weakly dor-
sally gibbous, drying wrinkled or smooth and often arcuate; annulus mostly
revoluble; peristome-teeth yellowish, confluent basally, lance-subulate, mostly
* Plagiothecium and the next following genus (Isopterygium) are probably best treated
as one genus. Grout (Moss Flora, 1932) unites them under Plagiothecium. Brotherus
(Die Naturlichen Pflanzenfamilien. 2nd edit. 1925) recognizes Plagiothecium and Isopter-
ygium as here treated except that he places deplanatum and geophilum in a third genus
Taxiphyllum.
Jennings: Manual of Mosses — 31. Hypnaceae TT^
hyaline-bordered, the divisural zigzag, the teeth dorsally cross-striate, numer-
ously lamellate; lid convex-conic, acute to rarely rostrate.
A genus of about 70 species, mostly growing on rocks and stones, rare in
the tropics; about 17 species in North America; at least 4 species in our region.
Key to the Species
A. Leaves spreading more or less uniformly in all directions B
K. Leaves distinctly complanately disposed C
B. Leaves erect-spreading; alar cells not inflated 2. P. Roeseanum
B. Leaves squarrose, more or less serrulate; alar cells inflated L P. striatellum
C. Costa forked, often reaching to the middle of the leaf; capsule usually striate when
dry 3. P. sytvaticum
<:. Costa thin, short and double, or none; capsule usually smooth when dry; lid conic
to short-rostrate 4. P. denticulatum
1. Pl.agiothecium striatellum (Bride!) Lindberg
(Hypnum MuehlenbeckH Schimper; Plagiothecium Muehlenbeckit
Bryologia Europaea; Leskea striatella Bridel)
Plate LI
Slender, dense, dark green, lustrous: stem prostrate, branches crowded,
erect or ascending, straight or slightly curved; leaves about 1-1.3 mm long,
crowded, sub-complanate, the branch-leaves squarrose-spreading, ovate-lanceo-
late or triangular-lanceolate with a long slender and flexuous acumen, plane-
margined, serrulate above at least, the base strongly decurrent: costa double
and faint; median leaf-cells linear-fusiform, flexuous, rather short, about 6-
10:1, the basal somewhat larger, the alar abruptly very much enlarged, inflated,
hyaline to colored, and forming the much decurrent and plainly distinct
auricles; inner perichaetial leaves half-sheathing, the apex filiform- flexuous and
usually recurved: capsules about 2 mm long, slightly curved, oblong-cylindric
with a tapering neck, distinctly striate when dry; lid conic, rather obtuse;
annulus large, compound; exothecial cells minute and rounded in three to five
series at the rim, below rapidly becoming irregularly oblong and more or less
incrassate; teeth short, yellowish, papillose above, dorsally cross-striolate below,
lamellate, strongly trabeculate, slightly confluent at base; segments about as
long, slender, pale, granulose, only slightly carinate cleft; basal membrane only
about one-fourth to one-third as high as the teeth; the cilia 1 to 3, a little
shorter than the segments; spores mature in late spring or early summer,
yellowish, papillose, rather incrassate, .007-.010 mm.
On earth, rocks, and rotten logs, in woods, usually in non-calcareous habi-
tats; Europe, and from Arctic America south to North Carolina.
Butler Co.: One mi. e. of Mercer Road Sta. C.M.B. Oct. 4. 194L Cambria
Co.: Ebensburg. T. P. James. (Porter's Catalogue). Favette Co.: Cucumber Falls,
Ohiopyle, June 22, 1940, and Laurel Run, Chestnut Ridge, Dunbar Twp., June 16,
1940. C.M.B. Lawrence Co.: Slippery Rock Creek, 1906. Miss Susan Gageby. Mc-
Kean Co.: On sandstone rocks, Rutherford Rocks, altitude 2000 feet, July 7, 1894,
Divide between Hawkins and Rutherford Hollows, April 25, 1893, and March 12, 1894,
and Langmade. May 9, 1896 (figured). All near Bradford. D.A.B.
276 American Midland Naturalist Monograph No. 6
2. Plagiothecium Roeseanum (Hampe, mss.) Bryologia
{Hypnum Sullivantiae Schimper; H. Roeseanum Hampe)
Plate LXXI
Compactly cespitose, pale green to yellowish-green, somewhat shining; stems
more or less erect, sparsely branched, radiculose at base; leaves about 2 mm
long, somewhat crowded, sub-imbricate, ovate-oblong, abruptly and very shortly
acuminate, rarely slightly serrulate towards the apex, thin, concave, the leaves
hardly complanate, but the branches appearing julaceous; costa various, often
bifid, rather strong and long; median leaf-cells narrowly linear, about 10-15:1,
towards the base gradually becoming shorter and broader; perichaetial leaves
erect, the inner oblong, narrowly acuminate: capsule cylindric-oblong, erect to
sub-inclined, smooth and constricted at the neck when dry; lid conic, obliquely
i-hort-rostrate; annulus large, simple; peristome hypnoid, the two cilia strong
and about as long as the entire segments; spxjres mature in summer.
This species apparently intergrades with P. denticulatum. The stems are
quite brittle and it is difficult to separate the leaves from them without break-
ing them.
On stones and earth in moist or swampy woods; Eurasia, and from Nova
Scotia to Alaska and south to Georgia.
Allegheny Co.: Powers Run, O.E.J. &C G.K.J. May 25, 1904 (named by F. E.
Wynn. 1944). Fayette Co.: On face of large rock, Laurel Run, Chestnut Ridge,
Dunbar Twp. C.M.B. June 16, 1940 (figured).
3. Plagiothecium sylvaticum [Hudson, Bridel] Bryologia Europaea
{Hypnum silvaticum Bridel; H. denticulatum C. Mueller)
PlatP LI
Both this and P. Roeseanum are perhaps but varieties of P. denticulatum,
but until better known should probably be kept apart as separate species.
Tufts large, dull or but slightly glossy, deep olive-green to yellowish-green;
stems prostrate, stoloniferous; leaves rather soft, large, concave about 2-3 mm
long, not very regularly complanate, widely sperading, shrinking and somewhat
twisted when dry, broadly ovate-lanceolate, narrowed considerably towards the
decurrent base, tapering abruptly to the acute, entire or obsoletely denticulate
apex, plane-margined: costa rather faint, double, often reaching one-third the
length of the leaf: median leaf-cells about 8-10:1, about .016 mm wide, large,
hexagonal-rhomboid, the alar cells numerous and quadrate-oblong, sub-inflated,
hyaline and decurrent; perichaetial bearing rhizoids at base, about 3 mm high,
the leaves sheathing with a flexuous acumen, non-costate; seta castaneous, slen-
der, about 2-4 cm long, dextrorse above when dry; capsule yellowish, about 2
mm long, long-cylindric from a tapering neck, inclined, arcuate, smooth, but
when dry and empty somewhat striate; lid conic, acuminate to sub-rostrate,
about one-half as long as the urn; peristome-teeth bright orange at base, pale
above, lance-subulate, confluent at base, closely trabeculate, the dorsal lamellae
finely cross-striolate; segments slender, about as long as the teeth, narrowly
Jennings: Manual of Mosses — 31. Hypnaceae 277
carinately gaping, the basal membrane about one-third as high, the cilia very
slender, about as long as the segments, nodose, usually 3 in number; exothecial
cells moderately incrassate, small and quadrate at rim, larger and oblong-
rectangular, to rounded-hexagonal below; annulus large, revoluble, 2-seriate;
spores pale yellowish, smooth, rather thin-walled, .006-. 010 mm, mature in
mid-summer.
On humus, rocks, rotten logs, etc., in woods; Europe, Asia, northern
Africa, and from southern Canada to Alabama and from Alaska to Oregon.
Known from Allegheny, Armstrong, Butler, Crawford, Erie, McKean, Washington,
and Westmoreland counties. With the exception of the McKean County station these
records are all from the western part of our region and seem to indicate that this species
is at least not common in the? mountains and plateau counties. Specimen figured: Wild-
wood Hollow, Allegheny County. June 11. 1908.
4. Plagiothecium denticulatum [Linnaeus] Bryologia Europaea
(Hypnum denticulatum Linnaeus)
Plate LIII
Variable, in flattened tufts of a pale and lustrous green, moderately robust,
the more or less ascending and elongate branches complanate; leaves rather
close, complanate and sub-distichous, rather spreading, when dry little changed,
glossy, 2-3 mm long, sub-concave, oval to lance-oblong, shortly and sometimes
almost apiculately acute, usually slightly denticulate at the apex, the lower
margins entire and often narrowly recurved, the base narrowed to a rather wide
and strongly decurrent insertion; costa thin, variable, usually short and double,
sometimes forked and reaching almost to the middle, sometimes none; median
leaf-cells rhomboid-hexagonal, rather large, about 10-15:1, about .010-015 mm
wide, thin-walled, chlorophyllose, gradually becoming laxer, pellucid, and more
or less rectangular at base, the alar more hyaline, sub-rectangular, sub-inflated,
and still somewhat larger but not forming clearly differentiated auricles, strong-
ly decurrent; the apical leaf-cells much smaller, incrassate, rhomboid: seta
about 2.5-4 cm long, flexuous, dextrorse above when dry, slender; capsule-um
about 2.5 mm long, sub-erect to horizontal, cylindric and with a distinct neck,
arcuate to nearly symmetric, when dry and empty sometimes striate; lid conic,
obtusely acuminate to long-acuminate, about one-third as long as the urn;
annulus usually 2-seriate, large, revoluble; exothecial cells rounded-quadrate,
small at rim, gradually larger and more oblong below, all incrassate; peristome-
teeth lance-.subulate, hyaline and papillose above, yellowish and dorsally cross-
striolate below, strongly lamellate and trabeculate, confluent at base; segments
slender, as long as the teeth, basal membrane one-third as high, cilia 2 or 3,
usually 3, slender, nodose; spore3 smooth, medium-walled, yellowish, .008-011
mm, mature in summer.
On stones, humus, and rotten wood, in moist forests; cosmopolitan in tem-
perate and cooler regions; in North America from the Arctic regions to the
northern United States and southwards in the mountains to Georgia.
278 American Midland Naturalist Monograph No. 6
This is a common and quite variable spwcies now known from numerous localities in
13 counties and probably occurring in all. Specimen figured: Falls Creek, Jefferson Co.,
July 18, 1904. O.E.J.
Other mosses closely related to P. denticulatum are P. Ruthei Limpricht,
with asymmetric leaves each clasping the base of the leaf above something like
Fissidens, and occurring in swamps from New England to Georgia and Mary-
land, and Minnesota. Also P. laetum Bryologia Europaea, with slenderly
acuminate leaves, scarcely decurrent, and with capsules almost erect and sym-
metric; no cilia. This species occurs from eastern Pennsylvania to Canada in
rock crevices in mountain areas.
22. Pterygynandrum Hedwig
Dioicous; slender to quite robust, variously cespitose, green to yellowish-
green, dull or lustrous, primary stem stolon-like, irregularly radiculose; second-
ary stems secund, filiform-julaceous, the base stolon-like, radiculose, often
flagelliform, densely-leaved, ascending, forked, bushy or pinnately branched;
leaves imbricate, sometimes secund, somewhat decurrent, non-plicate, deeply
concave, short-acuminate to somewhat obtuse, margins narrowly revolute to the
middle or above, entire or serrulate upwards; costa usually very short and thin,
forked or double, rarely single and reaching to the middle of the leaf; cells
narrow linear-vermicular to rhombic-hexagonal, strongly and sharply dorsally
papillose, the median about 3-5:1, basal cells wider and longer, alar quadrate
in several series; inner perichaetial leaves thin, broadly lanceolate, sheathing,
acute, the margins entire and plane: seta 8-15 mm long, red or yellow-red,
drying twisted; capsule erect, mostly symmetric, cylindric, yellow to brown;
annulus 2-seriate, narrow; peristome inserted near the mouth; teeth lanceolate,
confluent at the base, yellowish, below transversely and obliquely striate, above
smooth, distantly articulate, non-trabeculate; inner {peristome hyaline, smooth,
with quite low basal m.embrane, the segments very narrow, short, or sometimes
almost as long as the teeth; cilia none; spores .010-.018 mm; lid conic, shortly
and mostly bluntly rostrate.
Three species; only the following in North America:
1. Pterygynandrum filiforme [Timm] Hedwig
(Leskea cylindrica Bridel)
On base-s of trees and on rocks, in woods, widely distributed in the North-
ern Hemisphere, — in North America, extending from Greenland to British
Columbia and southwards to the northern United States. Occurs in the
Pocono region of eastern Pennsylvania and, possibly, will be found in the
northern or northeastern part of our region.
Family 32. Fabroniaceae
Autoicous or dioicous: slender to very slender, weak, cespitose, mostly
bright or light green, mostly lustrous: stem without central strand, weak,
creeping, thin, with red, fasciculate radicles; the secondary stems densely-
leaved, simple or branched, erect; leaves 5-8-seriate, drying appressed, spread-
Jennings: Manual of Mosses— 32. Fabroniaceae 279
ing when moist, rarely secund, more or less concave, unistratcse, non-decurrent,
ovate to lanceolate, non-bordered, non-plicate; costa simple, delicate and short,
rarely ecostate; median leaf-cells mostly prosenchymatous, smooth, mostly
thin-walled, towards the basal angles quadrate to rectangular: capsule exserted,
erect, symmetric, oval to sub-cylindric, drying often longitudinally wrinkled
and constricted below the mouth, the collum short and thick; peristome deeply
inserted, single or double; teeth plane, distantly articulate, non-lamellate, in
our genera non-bordered, teeth rarely none; inner peristome none or consisting
generally of subulate segments; lid broad, mostly conic and rostrate; calyptra
cucullate, naked, smooth, small, fugaceous; spores small.
Mostly occurring on tree-trunks in warm regions; only 3 genera withm our
range.
Key to the Genera
A. Costa none or very rudimentary; leaves serrulate the whole length ....3. Schwetschkeopsis
A. Costa plainly evident B
B. Inner peristome none, teeth short, broad, and blunt; leaves denticulate to ciliate-
dentate 1- Fabroniu
B. Peristome double; teeth broadly lanceolate; leaves essentially entire
2. Anacjmptodon
1. Fabronia Raddi
Autoicous, rarely dioicous: stem creeping, partly stoloniform, rarely erect,
irregularly branched; branches often partly stoloniform and partly leafy; leafy
branches thickly julaceous, the leaves often drying imbricate, sometimes secund,
ovate to ovate-lanceolate, mostly subulate-acuminate or piliferous, entire to
serrate or even ciliate-Iaciniate; costa mostly delicate and short, sometimes
indistinct; median leaf-cells elongate-rhomboid to elongate-hexagonal, the alar
quadrate in several series, sometimes not differentiated; inner perichaetial leaves
sheathing, subulate-acuminate, ecostate: seta mostly 1-7 mm long, thin, pale
yellow, smooth, drying twisted; capsule erect, symmetric, ovate to pyriform,
with a short neck, drying plicate, the collum shrinking and the capsule becom-
ing cup-shaped to hemispheric, light brown, wide-mouthed; annulus none;
peristome simple, rarely none, teeth very hygroscopic, at first united in pairs,
later separating, broad, obtuse, often cleft or perforate divisurally, brown, non-
bordered, longitudinally striate-papillosc, non-trabeculate; lid conic-convex to
low convex, mostly short-rostrate.
A genus of about 100 species, widely distributed in warm regions, mostly
arboreal in habitat, rarely on rocks; 13 species reported for North America;
two species in eastern Pennsylvania and perhaps reaching our region.
Key to the Species
A. Leaves obscurely serrate 1. R. Ravenelii
A. Leaves ciliate-dentate 2. F. ciliarif
1. Fabronia R.^venelii Sullivant
( F . cjToltniana Sullivant )
Very small, delicate, loosely cespitose, bright green: stems creeping with
280 American Midland Naturalist Monograph No. 6
more or less erect branches; leaves loose, elongate-lanceolate, subulate-acumi-
nate, concave, costate to the middle, entire or but obscurely serrate; median
leaf-cells linear-fusiform, the basal and alar quadrate; inner perichaetial leaves
erostate, oblong, short-acuminate: capsule more or less pyriform; the teeth of
the peristome brown, 16, approximate in pairs, orange-pellucid, acuminate-
deltoid; lid conic, obtuse; spores about .017 mm in diameter.
On bark of trees, rotten logs, etc., extending from the South up into south-
eastern Pennsylvania.
2. Fabronia ciliaris (Bridel) Bridel
(F. octoblepharis Schwaegr.; F. pusilla Schwaegr.;
Pterogonium octoblepharis Schleicher)
Small, delicate, thinly cespitose: stems creeping with erect branches; leaves
lance-ovate, filiform-acuminate, thin, green, spreading, sometimes 2-ranked,
coarsely and irregularly lacerate-dentate on the border the acuminate tip con-
sisting of a single elongated cell, costate to considerably above the middle,
non-plicate, plane-margined, closely imbricate when dry; median leaf-cells thin-
walled, linear-rhombic to hexagonal, about 3-5:1, the basal clear across the
lower one- fourth or one-fifth of the leaf quadrate or sub-quadrate: seta rather
long; capsule oval, neck rather distinct, the urn erect, symmetric, more or less
contracted below the mouth when dry and empty; peristome single, with the
teeth united in pairs, dark brown, recurved when dry, when old more or less
bifid; spores mature in spring, about .015 mm in diameter.
On rocks and trees from New Jersey throughout the Central States to
Minnesota and southwards. Occurs in southeastern Pennsylvania and may
reach the southern part of our region.
2. Anacamptodon Bridel
Autoicous: mostly densely cespitose, dark green, when old brownish to
yellowish, lustrous: stem long-creeping, densely radiculcse; the branches short,
densely-leaved, erect to ascending; leaves spreading, often secund, ovate to oval,
long-acuminate, entire; costa strong, ending above the middle of the leaf; leaf-
cells rich in chlorophyll, elongated rhombic-hexagonal, the basal rectangular;
inner perichaetial leaves elongate, not sheathing, generally acuminate, thinly
costate; seta 5-8 mm long, quite thick, straight, smooth, red to dark castaneous,
drying twisted; capsule erect, symmetric, oval, short and thick-necked, drying
strongly constricted below the mouth, smooth; annulus broad, delicate but
persistent; peristome double, deeply inserted, the teeth strongly hygroscopic,
apically united in pairs, broadly lanceolate, pale brown, divisural line almost
straight; teeth articulate below, densely finely papillose: basal membrane of
inner peristome none, the segments filiform, somewhat shorter than the teeth,
non-carinate, brown, almost smooth; lid conic-convex, straight or obliquely
rostrate; spores about .008. 010 mm.
A genus of 7 species of which 5 are confined to Asia. The following
occurs in Europe and eastern North Ameria:
Jennings: Manual of Mosses — 32. Fabroniaceae 281
1. Anacamptodon splachnoides [Froelich] B.idel
{Campylodontium hypnoides Schwaegrichen; Neckera splachnoides Schwaegrichen)
Knot-Hole Moss
Plate XXXVII
Dark-green or bluish-green, small, delicate, thinly tufted: stems creeping,
with erect branches; leaves lance-ovate, up to 1.5 mm long, acuminate, entire,
plano-concave, chlorophyllose, soft, closely imbricated when dry, non-decurrent,
non-plicate; costa rather slender, reaching to above the middle of the leaf;
median leaf-cells rhombic-hexagonal, about 3-5:1, with a few quadrate and sub-
inflated cells at the base; perichaetial leaves few: seta about 6-8 mm long, sinis-
trorse when dry; capsule erect and symmetric, about 2:1, oval-oblong, thick-
necked, constricted below the mouth when dry; peristome double, teeth approx-
imately in pairs and reflexed when dry; 16 in number, lanceolate, pale, articu-
late, the divisural zigzag; segments filiform, about one-half to two-thirds as
long as teeth; no basal membrane; annulus none; exothecial cells castaneous-
incrassate, rectangular or irregularly oblong, above smaller and quadrate, those
at the rim minute and rounded; lid short-rostrate from a conic base, more or
less oblique, one-half to two-thirds as long as the urn; calyptra whitish, cover-
ing only the upper part of the urn; spores about .010 mm, minutely papillose,
yellowish-green, medium-walled, mature in June.
In moist cavities in decaying wood, knot-holes in trees, in forks of tree-
trunks, etc.; Europe, Asia, and from New England to Illinois and south.
Collected but seldom in our region, and then only in small quantities. Fayette Co.:
In knot-hole 20 feer up in a beech tree, Tates Hollow. Dec. 1, 1933. John Lewis. In-
diana Co.: T. P. James. (Porter's Catalogue). McKean Co.: Bradford, different
dates, in cavities in decaying wood. D.A.B. (figured). Issued as No. 148. Grout's
North American Musci Pleurocarpi. Somerset Co.: Near Trent. Aug. 19, 1933.
Chas M. Hepner. Washington- Co.: In knot of a beech log, near Washington, Sept.
15, 1891. Linn &C Simonton.
3. Schwetschkeopsis Brotherus
Dioicous: slender, soft, forming tufts, green to yellowish: stem long, creep-
ing, radiculose, mostly densely and symmetrically pinnately branched; branches
somewhat julaceous to complanately-leaved, obtuse, short to long, ascending,
simple or branched; branch-leaves when dry imbricate, when moist erect-spread-
in, non-decurrent, concave, plicate, lance-ovate, acuminate to subulate-pointed,
serrulate, plane-margined; costa none; leaf-ceils oblong-oval to oblong-linear,
dorsally papillose above, alar quadrate and numerous, chlorophyllose, occupy-
ing most of leaf-base: seta up to 7 mm long, slender, tortuous, yellowish-red,
smooth, when dry twisted; capsule oblong-ovoid, mostly erect and symmetric,
shortly collumate; exannulate, peristome double, teeth lanceolate, yellow, with
zigzag divisural, densely transversely striate, closely tra'^eculate; in:'er peri-
stome hyaline, basal membrane one-third as high as teeth, smooth, segments
about as long as teeth, broad, split along keel, finely papillose, cilia rudimen-
tary or none; spores about .015 mm; lid obliquely rostrate.
282 American Midland Naturalist Monograph No. 6
Three species, on trees: one in Japan and Korea, one in Nepal, and the
following:
1. SCHWETSCHKEOPSIS DENTICULATA (Sullivant) Brotherus
(Leskea denticulate Sullivant)
Plate XXXVI
Light green, soft, silky: stems usually 2-3 cm long, sometimes more, irregu-
larly branched, paraphyllia none; stem-leaves erect-spreading, close, concave,
ovate, somewhat decurrent, abruptly and narrowly acuminate, 0.4-0.9 mm long,
0.3-0.4 mm wide, plane-margined, sometimes slightly striate, marginally
undulate-denticulate; ccostate; apical leaf-cells dorsally uni-papillate, the
median oblong-oval to elongate-rhomboidal, sometimes vermicular, about 4-8:1,
about .005-.008 mm wide, the marginal uni-seriate and curvi-linear, the alar
forming a large group of quadrate incrassate cells; branch-leaves smaller and
less abrupdy acuminate, with more oblong and shorter cells: seta yellowish-
red, slender, tortuous, erect; capsule erect or nearly so, oblong, about 2-3 : 1 ;
operculum conic-rostrate, about two-thirds as long as the urn; peristome about
the same width as the teeth; no cilia; no annulus; fruit rarely found.
Mostly on bases of trees, rarely on rocks, occurring in Asia and from Con-
necticut to the Mississippi River and south to the Gulf.
Apparently rare in our region. Butler Co.: On base of tree, 2 mi. s.e. of Browns-
dale, Aug. 18, 1935. Sidney K. Eastwood. McKean Co.: Lewis's Run, Bradford, No-
vember 24, 1895, and Limestone Creek, Bradford, December, 1896. D.A.B. (figured).
The latter issued as Grout's No. 134, North American Musci Pleurocarpi.
Family 33. Sematophyllaceae
Autocoius or dioicous; antheridial clusters gemmiform, small; archegonial
clusters on very short, usually rooting, perichaetial branches: slender to robust,
cespitose, green to yellowish or brownish, often lustrous: stem without central
strand, creeping to ascending, mostly irregularly branched, sometimes more or
less regularly pinnate; paraphyllia none; leaves pluriseriate, mostly uniform
and symmetric, of various forms; costa double, very short or none; cells mostly
prosenchymatous, smooth or papillose, in the leaf-angles one row being oblong,
inflated, thin-walled: capsule exserted, mostly cernuous to pendent, mostly oval
to oblong, usually unsymmetric, collum weak; exothecial cells collenchymatous;
annulus none; peristome-teeth as long as the segments, the latter rarely lacking,
the teeth mostly entirely separate, mainly dorsally striate, lamellae mostly well-
developed, inner peristome free, basal membrane high, segments mostly carinate
and lance-subulate, rarely filiform, cilia usually present; spores mostly small;
!id from a convex-conic base slenderly rostrate; calyptra mostly cucullate and
glabrous.
A rather large family almost exclusively of tropic and sub-tropic distribu-
tion and mostly living on trees; in our region there occurs but one genus, as
follows:
Jennings: Manual of Mosses— 33. Sematophyllaceae 283
I. Sematophyllum Mitten
(Rhaphidostegium (Bryologia Europaea) DeNor.)
Usually autoicous: mostly slender, mostly densely and widely cespitose,
dark to pale green or yellowish to brownish: stem creeping, more or less elon-
gate, regularly pinnately branched or irregular, with branches horizontally
spreading to erect, rather julaceous; leaves uniform, non-plicate, concave, entire,
ovate to oblong or oblong-elliptic, obtusely to piliferous-acuminate; usually
ecostate, rarely obsoletely bi-costate; cells narrowly prosenchymatous, the apical
sometimes rhombic, the basal golden-yellow, narrowly rectangular, incrassate
and porose, the alar oblong, inflated, hyaline to yellowish or red-brown and
forming a small, non-excavate group bounded above by small quadrate cells:
seta long, mostly smooth; capsule sub-erect or horizontally inclined, oval to
oblong, smooth; peristome hypnoid, teeth lance-subulate, with divisural zigzag,
hyaline-bordered, prominently lamellate, especially so in the upper third; peri-
stome-segments yellowish, carinate, with a high basal membrane, mostly split,
cilia 1 or 2, nodose, or sometimes rudimentary; spores small, lid slenderly
subulate-rostrate; calyptra glabrous.
A genus of about 100 species of temperate and warmer regions, occurring
mainly on trees and rocks; 3 species in our region.
Key to the Species
A. Capsules erect and symmetric or nearly so: branches curved at ends and with apical
leaves second 3. S. adnatum
A. Capsules more or less inclined or horizontal; leaves erect-spreading in all directions,
or somewhat secund B
B. Leaves usually more than 1.5 mm long 2. S. marylandicum
B. Leaves usually less than L5 mm long 1. S. carolinianttm
1. Sematophyllum carolinianum (Mueller) E. G. Britton
(Hypnum carolinianum C. Mueller; H. demissum var. carolinianum
Lesquereux and James; Rhaphidostegium carolinianum Jaeger)
Plate LIII
Rather dirk green, drooping-cespitose, lustrous: stems irregularly branch-
ing, often buried in the sand and then more or less leafless and with erect to
ascending simple branchlets about 1-1.5 cm long; leaves imbricate, more or less
secund or complanate above, non-plicate, concave, lance-ovate or lance-oblong,
more or less sub-serrulate at apex, shortly acute, the margin often rather broad-
ly reflexed; costa none, or faintly indicated by striae; median leaf-cells linear-
flexuous, small, incrassate, about 8-10:1, towards the base shorter and broader,
the alar abruptly much enlarged and inflated to form a group of 2-8 pellucid
and hyaline or colored cells; perichaetial leaves rather closely imbricate, lance-
oblong, acuminate: seta erect, sinistrorse below, castaneous, about 1 cm long;
capsule curved and inclined, constricted below the mouth when dry and empty,
the urn about 1.2-1.5 mm long, oblong-pyriform, yellowish; exothecial cells
rounded-hexagonal, collenchymatous; peristome orange-yellow, the teeth with
distinct divisural and lamellae, dorsally cross-striolate, hyaline-margined, strong-
284 American Midland Naturalist Monograph No. 6
ly trabeculate; segments about as long, slender, rarely split, the cilia 1 (or 2),
about one-half to two-thirds as long as segments, the basal membrane about
two-fifths the height of the peristome; lid comparatively large, the beak oblique,
subulate, and about two-thirds to three-fourths as long as the urn; spores
smooth, yellowish-incrassate, usually chlorophyllose, about .014-.018 mm,
mature in summer or early autumn.
On wet non-calcareous rocks, mainly in ravines in hilly or mountainous
districts; Asia, and from Newfoundland southwards to Georgia.
Probably not common in our region. Allegheny Co.: Haysville Hollow, September
20, 1908. O.E.J.; on damp rocks under hemlocks, Wildwood Road, November 19, 1908.
O.E.J, and G.K.J. Fayette Co.: On damp rocks in deep hollows and ravines, Ohio
Pyle, September 1-3, 1906, and September 1-3, 1907. O.E.J, and G.K.J, (figured); also
May 30-31, June 13, and July 4, 1908. O.E.J.; Ohio Pyle, June 15, 1902. J.A.S.
Somerset Co.: Near Trent. C. M. Hepner, July, 1932; and Clear Run, Laurel Ridge,
C.M.B. Oct. 6, 1935. On wet rock in stream. Westmoreland Co.: Four collections
in Laurel Hill and Chestnut Ridge Mts. C.M.B. 1934-45.
2. Sematophyllum marylandicum (Mueller) E. G. Britton
{Rhaphidostegium marylandicum Jaeg. & Sauerb.; Hypnum marylandicum C. Mueller)
Plate LII
Dark green loosely interwoven, matted mosses with often subsecund at
stem- and branch-tips, oblong-ovate, shortly acuminate to acute, 1.5-2 mm
long, concave, entire, margin not reflexed; median leaf-cells linear, about 8-
12:1, shorter, incrassate, and pellucid at base, alar cells inflated in a small
group, apical cells about 2-4:1, rhomboid-oblong; spores ripe in spring.
On wet stones and ledges in cool mountainous habitats from New Eng-
land to Georgia.
This species is very difficult to distinguish from Hygrohypmim eugyrium
var. Macktyt, but the Hygrohypnum has usually a faint double costa, some-
times lacking, and the walls of the outer alar cells are much thinner than are
the walls of the inner cells.
McKean Co.: Cathrine Swamp. C.M.B. Sept. 2, 1948 (figured). Somerset
Co.: Bluehole Creek, Laurel Ridge. C.M.B. On rock in stream. July 5, 1948; and H.
N. Mozingo, wet rock in stream, near Mt. Davis, Aug. 26, 1945.
3. Sematophyllum adnatum (Richard, Michaux) Britton
(Lesked adnata Richard; Rh. microcarpum Jaeger; Leskea microcarpj
Bridei; Rhaphidostegium adnatum Bryologia Ejropaea)
Small, in tangled, thin, green to golden-green mars: stems prostrate, with
short and incurved branches; leaves rather closely imbricate when dry, sub-
homomallous, the upper usually distinctly secund, narrowly oblong-lanceolate,
the apex rather shortly acuminate, subserrulate to entire, margins quite broad-
ly reflexed; costa double but very short and faint; median leaf-cells linear-fusi-
form, flexuous, about 8-12:1, shorter and wider at the base, tov/ards the angles
a border of sub-rectangular and scarcely inflated cells and at the extreme angle
a few distinctly inflated alar cells: seta short and smooth, abou.t 5-8 mm long;
ca lyptra more or less persistent, cuculiate, reaching to a little below the mouth
Jennings: Manual of Mosses — 34. Brachytheciaceae 285
of the urn; capsule castaneous, about 1-1.3 mm long, oblong to oblong-cylin-
dric, about 2-2.5:1, erect and symmetric or nearly so, thin-walled but with
collenchymatous exothecial cells, slightly constricted below the reddish rim
when dry and empty; annul us none; lid obliquely subulate-rostrate from a coni-
cal base, about as long as the urn; oeristome-teeth with an unusually distinct
and heavy divisural, cilia single and usually about half as long as the usually
entire segments, basal membrane reaching to about one-third the height of the
inner peristome; spores mature in late summer to fall.
On base of usually living trees; in moist woods from southern New Eng-
land to Ohio and southwards to the Gulf States. Not yet found in our
region.
Family 34. Brachytheciaceae
Autiocous or dioicous; paraphyses filiform; antheridial cluster^ gemmiform;
archegonial clusters on very short, rooting branches: slender to robust: stem
with central strand, creeping to ascending, or rarely erect, often interruptedly
sloloniferous, fasciculately radiculose, mostly irregularly pinnate; branches
mostly acute, often flagelliform and rooting at the ends; leaves unistratose,
pluriseriate, erect-spreading or appressed, rarely homomallous, dimorphic in
the stoloniferous species; cordate-oblong to lance-ovate or lanceolate, acumi-
nate or rarely obtuse; costa mostly incomplete; median leaf-cells prosenchyma-
tous, elongate-rhomboid to linear-vermicular, smooth or rarely papillose
towards the upper end of the cell, the basal cells lax and often porose, the
alar usually differentiated, being quadrate, green or hyaline, sometimes in-
flated: seta elongate, often rough; capsule cernuous to horizontal, mostly
short, ovoid or oblong and dorsally gibbous, when dry and empty more or less
arcuate, rarely erect and symmetric, oval to oblong-cylindric, never pendent,
smooth; collum faint; exothecial cells collenchymatous; peristome hypnoid,
imp>erfecr in some species with capsules erect; teeth lance-subulate, mostly
strongly hygroscopic, basally confluent, yellow or orange to red-brown, with a
zigzag divisural, dorsally cross-striate, lamellae numerous and well-developed;
inner peristome mostly free, with a high basal membrane, carinate segments
which are lance-subulate, cilia mostly complete, rarely none or rudimentary;
lid conic, obtuse to acute, often long-rostrate; calyptra cucullate, early decidu-
ous, mostly glabrous.
A large and cosmopolitan family on various substrata, containing about 20
genera with more than 500 species.
Key to the Genera
A. Capsule erect to suberect, and somewhat unsymmetric to symmetric; basal membrane
mostly low H
A. Capsule cernuus to horizontal, unsymmetric; basal membrane mostly high R
B. Leaves mostly with several deep plications 2. Camptothecium
B. Leaves not deeply plicate c
C. Lid conic, sometimes acute; alar cells differentiated 4. BTachythecium
C. Lid long-rostrate, alar cells few or none D
D. Cells narrow, dorsally smooth E
D. Cells oblong-rhomboid to rhomboid-hexagonal, those of the branch-leaves more or
286 American Midland Naturalist Monograph No. 6
less dorsally rough 8. Bryhnia
E. Leaves complanate (Also see Oxyrrhynchium) 9. Rhynchostegium
E. Leaves imbricated or spreading F
F. Leaves deeply concave, spoon-like abruptly piliferous-acuminatc 5. Cirriphyllum
F. Leaves plane or somewhat concave, acute or gradually acuminate, not piliferous G
G. Leaves not much concave, non-plicate 6. Oxyrhynchium
G. Leaves mostly distinctly concave and plicate 7. Eurhynchiutn
H. Calyptra hairy; segments adhering to the peristome-teeth 1. Homalotheciella
H. Calyptra smooth; segments not adhering to the peristome-teeth. (See also some
species of Brachythecium) 3. Chamberlainia
1. Homalotheciella (Cardot) Brotherus
Autoicous: slender, soft, laxly cespitose, green, lustrous: stem creeping,
elongate, beset thickly with obtuse, short, ascending to erect, densely-leaved
branches which are often arcuate when dry; paraphyllia none; when dry the
leaves imbricate, when moist erect-spreading, non-decurrent, non-plicate, con-
cave, oval to oblong, the apex acuminate to lancc-subulate, upper half of leaf
serrulate to entire; coGta simple, sometim.es reaching to mid-leaf; median leaf-
cells oblong-elliptic, thin, smooth, the alar green, numerous, quadrate; inner
perichaetial leaves abruptly serrate-subulate from a sheathing base; seta about
7 mm long, castaneous, rough; capsule erect to almost horizontal, more or less
unsymmetric, oblong, drying somewhat constricted below the mouth and often
sub-arcuate; annulus present; inner peristome much shorter than the outer,
teeth basally confluent, lance-linear, dorsally cross-striate, apically papillose,
lamellae laterally projecting; inner peristome somewhat united with the outer,
yellow, smooth, with low basal membrane, segments short, narrow, adherent
to the teeth; cilia none; lid long-rostrate; calyptra slightly hairy at base.
A small genus of three (4) North American species, one species occurring
in our range.
1. Homalotheciella subcapillata (Hedwig) Cardot
(Pterigynandrum suhcapillatum Hedwig; Homalothecium subcapillatum
SuUivant; Platygyrium brachycladon Kindberg)
Forming light green, thin, glossy mats: stems prostrate, irregularly branch-
ing; leaves loosely imbricate when dry, elliptic-oblong, abruptly long-acumin-
nate, more or less serrate above, about 0.9-L2 mm long, concave, non-plicate,
not papillose; costa usually reaching about to the middle of the leaf; median
leaf-cells, about 8-10:1, fusiform-elliotic, towards the apex somewhat shorter,
the alar quadrate, numerous and forming a group which extends upwards
along the margin to often one-third the length of the leaf; inner perichaetial
leaves sheathing, long-acuminate: seta rough, about 6-9 mm long, slender;
capsule about 2-3.5:1, sub-erect, slightly incurved, dorsally somewhat gibbous,
slightly constricted below the mouth when dry; operculum conic or convex;
annulus 2-seriate; peristome-teth confluent at base, dark red, with a broad
pellucid central stripe marked by a delicate medial line, the segments adhering
to and lining the teeth inside, forming a hyaline border; spores mature in
autumn, about .025 mm.
Jennings: Manual of Mosses — 34. Brachytheciaceae 287
On bark of tree^ and on fallen trunks in woods; in the eastern United
States from New England to North Carolina.
Rare in our region. Elk Co.: McMinn. (Porter's Catalogue).
2. Camptothecium Bryologia Europaea
Dioicous and pseudautoicous: slender to robust, widely ccspitcse, mostly
yellowish-green, drying st'ift, mostly lustrous: stem elongate, procumbent to
ascending to erect, thickly-leaved, sometimes stoloniferous, more or less regu-
larly pinnate; leaves erect-spreading, sometimes weakly secund, non-decurrent,
slightly concave, strongly plicate, lance-oval, subulate-acuminate, serrulate all
around; costa simple, ending near or in the apex; median leaf-cells prosenchym.-
atous, 10-20:1, vermicular, thin, smooth, or with weakly projecting upper
angles, the basal lax, yellow, porose, the alar numerous, quadrate: perichaetium
not rooting, inner perichaetial leaves much elongate and abruptly subulate:
seta moderately long, castaneous, mostly rough, drying twisted; capsule cernu-
ous to horizontal, dorsally gibbous, oblong to oblong-cylindric, more or less
curved; annulus present; peristome-teeth basally confluent, linear-subulate, bor-
dered, dorsally cross-striate, thickly lamellate; inner peristome of same length,
free, the segments broad and carinately split, cilia strong and nodose; lid conic-
acute to thickly short-rostrate; calyptra glabrous.
A genus of about 15 species, confined mainly to temperate regions on soil,
bark of trees in woods, or in swamps; a number of species occur in the West,
but in our region only the following:
1. Camptothecium nitens [Schreber] Schimper
(Hypnum nitens Schreber)
A Striking species by reason of its bright yellow or golden color, silky
lustre, strongly plicate leaver, and stems densely covered by a felt of reddish
radicles: the stems often reach a length of 10 cm, strong; the elongate-lanceo-
late leaves entire, stronglv plicate, marginally revolute, gradually and evenly
narrowed to the slender apex, reaching usually over 3 mm long; median leaf-
cells linear, the basal shorter with very thick and porose walls, the alar broader
and short-rectangular to sub-quadrate but rather few in number and not form-
ing a very distinct auricle: seta smooth; capsule cylindric, arcuate, contracted
below the mouth when dry and empty; peristome well developed; cilia long;
spores mature in spring.
In wet meadows, bogs, and swamps; Europe, Asia, and from Arctic Amer-
ica to northern United States. Occurs in eastern Pennsylvania but not yet
reported in our region.
3. Chamberl.ainia Grout
Cespitose, mostly glossy green, variously branching; .stem-leaves ovate;
branch-leaves erect-spreading, imbricate when dry, lanceolate to ovate-lanceolate,
concave, more or less serrulate, costate to above middle; median leaf-cells
288 American Midland Naturalist Monograph No. 6
linear to elongate rhomboidal, alar quadrate, often numerous: seta 1-2 cm
high, smooth; capsule erect, symmetric, cylindric; operculum conic, apiculate
to short rostrate; annulus none; peristome cilia none or single and rudimen-
tary; spores ripe in autumn.
This genus has been segregated from. Bracbythecium by Dr. Grout and
named in honor of E. B. Chamberlain (1878-1925), former secretary-treasurer
of the SuUivnnt Moss Society. Three species; two in our region.
1. Chamberlainia cyrtophylla (Kindberg) Grout
{Brachythecium cyrtophyllum Kindberg)
Plate LIII'
Cespitose, lustrous, dark green: stem irregularly branching to sub-pinnate,
creeping, up to 4 to 6 cm long; stem-leaves broadly ovate, up to 1 mm long;
branch-leaves similar but narrower and smaller, lance-ovate to lanceolate, acute
to short-acuminate, 0.6-0.8 x 0.3 mm, rather close, loosely appressed when
dry, serrulate at least in the upper half, marginally reflexed at base, not plicate,
not decurrent, when moist more or less spreading; costa stout, reaching about
two-thirds the length of the leaf; median leaf-cells rhomboid-fusiform, about
4-8:1, the alar sub-quadrate, numerous, sub-inflated, somewhat chlorophyllose;
perichaetial leaves ecostate, half-sheathing: seta about 2-2.5 cm long, dextrorse
above, erect, flexuous; capsule erect, cylindric, sometimes slightly curved, from
1.5 to 3 mm long, castaneous, smooth; annulus none; peristome-teeth slender,
pale castaneous, confluent at base, hyaline and papillose above, the dorsal
lamellae closely cross-striolate below, the trabeculae close and strong; segm.ents
nrarly as Icn*] a3 the teeth, slender, pale yellowish, more or less carinately
split, the cilia rudimentary or none; basal membrane about one-fourth the
height of the teeth; lid high-conic, usually acutely apiculate; spores papillose,
brownish, medium- to thick-walled, .012-.016 mm, mature in autumn. Very
closely related to the following species, which it apparently replaces to the west
and northwest of our region.
On roots and bases of trees and on old logs, in woods from our region
northwestward to Minnesota and Ontario, south to North Carolina.
Uncommon in our region. Allegheny Co.: Fern Hollow, Pittsburgh, on old logs
in ravine, January 21, 1906 (figured), and March 8, 1908. O.E.J. McKean Co.:
Bradford. D.A.B. (Porter's Catalogue).
2. Chamberlainia acuminata (Hedwig) Grout
(Leskea acuminata Hedwig; Hypnum acuminatum Beauvois;
Brachythecium acuminatum Kindberg)
Plate LIV
Widely and somewhat densely cespitose, dark to yellowish-green, glossy:
stems slender, prostrate, up to 5 to 8 cm long, bearing rhizoids, at least near
the perichaetia, rather distantly and unequally branched, the branches two-
ranked, plumose to sub-julaceous, acute, not usually more than 1 cm long:
stem-leaves close, erect-spreading, lance-ovate to ovate, about 1-1.5 mm long,
Jennings: Manual of Mosses — 34. Brachythfciaceae 289
acuminate, concave, with the borders reflexed below, the upper half serrulate,
the leaf non-plicate or but slightly plicate; narrowed and somewhat decurrent
at the base; costa usually reaching beyond the middle of the leaf; branch-leaves
similar to the stem-leaves but relatively narrower and smaller; median leaf-cells
linear- flexuose, about 51-0:1, medium-walled, prosenchymatous with rounded
ends, apical cells a little shorter, the basal sub-quadrate or sub-rectangular, the
alar numerous and sub-quadrste to quadrate, rather thin-walled and sub-
inflated: seta erect, castaneous, flexuous, about 1-1.5 cm long; capsule castane-
ous, erect, the urn 1.5-3 mm long, symmetric or sometimes slightly curved,
cylindric, about 3.5-4.5:1, tapering at base; lid high-conic, acute to apiculate;
exothecial cells densely yellowish-incrassate, small, rounded but varying to
quite irregular in size and shape but with rounded comers; peristome-teeth
narrow, castaneous, margined, numerously trabeculate, hyaline and papillose
above, dorsally cross-striate below, the lamellae distinct, teeth confluent at
base; segments about as long as the teeth, narrow, carinately split, cilia rudi-
mentary or none, the basal membrane only about one-fourth as high as the
teeth; annulus none; spores castaneous, papillose, medium-walled, about .014-
.018 mm, mature in l.ate fall or in winter.
On earth, woods-humus, roots and bases of trees, stones, and very often
en rotten logs, forming wide mats, in woods from the southeastern part of
Canada to the Gulf States and Minnesota.
Rather common m our region. Allegheny Co.: Thirteen pockets determined from
various localities, mainly on old logs in ravines. O.E.J, and G.K.J. ; Fern Hollow, Janu-
ary 21, 1906. O.E.J, (figured). Beaver Co.: Eight feet up the trunk of elm tree. Rac-
coon Creek, 1 mi. s. of Traverse Creek. C.M.B. April 1, 1934. Clearfield Co.:
Phillipsburg. T. P. James. (Porter's Catalogue). McKean Co.: Gate's Hollow, Brad-
ford, April 29, 1898. D.A.B. Issued as Grout's No. 116, North American Musci
Pleurocarpi. Washington Co.: On decayed wood, near Washington. Linn 8C Simon-
ton, No. 24. Oct., 1891. Westmoreland Co.: Near Apollo, 1902. Miss K. R.
Holmes; Greensburg, T. P. James. (Porter's Catalogue).
4. Brachythecium Bryologia Europaea
Autoicous or dioicous: slender to robust, mostly widely and flatly cespitose,
green or yellowish to whitish, sometimes lustrous: stems creeping or procum-
bent, sometimes more or less erect, thickly-leaved, irregularly divided, inter-
ruptedly pinnate, stolon-like at the apex; stem and branch-leaves unlike, stem-
leaves erect-spreading to spreading, more or less concave, mostly plicate,
narrowly lanceolate from a narrowed, ovate or triangular-cordate and decur-
rent base, acuminate, marginally plane, serrate all around or only towards the
apex, rarely entire; costa simple, usually long but rarely complete; median
leaf-cells narrow to moderately wide, elongate-rhomboid to linear, smooth, the
basal more lax, and shorter, the alar quadrate to rectangular or oblong-hexag-
onal, forming a rather indefinitely bounded group; branch-leaves mostly
shorter, narrower, with a somewhat weaker costa; inner perichaetial leaves slen-
derly and finely acuminate: seta more or less long, smooth to rough; capsule
cernuous to horizontal, rarely erect, mostly short-ovoid and dorsally gibbous,
rarely oblong-cylindric, slightly arcuate when dry and empty; usually annu-
290 American Midland Naturalist Monograph No. 6
Iste; peristomc-teeth strong, basally confluent, dorsally cross-striate, apically
papillate, thickly lamellate; inner fjeristome about the same length, yellow to
orange, free, with wide basal membrane, the segments broadly lanceolate, long-
acuminate, carinately split and often gaping, cilia complete, nodose to appen-
diculate, rarely rudimentary or lacking; lid conic-convex, obtuse to acute;
calyptra glabrous.
A genus of about 225 species, occurring on various substrata, mostly con-
fined to temperate regions; at least 12 species in our region.
Key to the Species
A. Seta smooth; annulus often present; cilia well-developed B
A. Seta rough at least in part E
B. Capsules sub-erect, narrowly cylindric-oblong; usually more than 3 : 1
1. B. oxycladon
B. Capsules cernuous, usually less than 3:1 C
C. Leaves narrowed gradually from base to acuminate ap>ex, non-plicate 5. B. dcutum
C. Stem-leaves ovate-lanceolate, more or less plicate D
D. Stem-leaves broad, about 1 mm at base, not falcate-secund 3. B. salebromm
D. Stem-leaves narrow, about 0.5-0.6 mm at base, very gradually narrowed to the
apex 4. B. flexicaule
E. Seta rough only above F
E. Seta rough throughout H
F.Costa percurrent or very nearly so 11. B. populeum
F. Costa ending about in middle of the leaf G
G. Stem-leaves plicate: cilia non-appendiculate 2. B. campestre
G. Stem-leaves non-plicate: cilia appendiculate 12. B. flagellare
H. Cilia appendiculate I
H. Cilia non-appendiculate J
r. Costa percurrent or sub-percurrent 8. B. reflexum
I. Costa distinctly incomplete 9. B. Starkei
J. Stem-leaves very short-acuminate, alar cells abruptly inflated 7. B. rivulare
J. Leaves gradually acuminate; alar cells not very abruptly enlarged and inflated K
K. Slender; leaves lanceolate, often secund 10. B. velutinum
K. Robust; leaves ovate to lance-ovate, not secund 6. B. rutabulum
\. Brachythecium oxycladon [Bridel] Jaeger and Sauerb
(B. laetum Bryologia Europaea; Hypnum oxycladon Bridel)
Plate LXIX
Cespitose, bright or yellowish-green: stems prostrate, branching unequally
and irregularly, the branchlets attenuate at the apex and erect; leaves close,
loosely imbricate, ovate in the stem-leaves (2-2.5 mm) and more lance-ovate
in the branch-leaves (1.5-2 mm) rather abruptly acuminate, concave, plicate,
finely serrulate all around; costa rather narrow, extending about to mid-leaf
or somewhat farther; median leaf -cells long, narrow, about 8-10:1, flexuous,
the basal more or less quadrate, the alar numerous, small, rather incrassate,
the alar portion strongly decurrent: seta about 2.5 cm long, flexuous, flattened
and dextrorse when dry; capsule sub-erect, about 4:1, 3-4 mm long, oblong-
cylindric, when dry somewhat arcuate and often inclined; lid conic-acuminate;
Jennings: Manual of Mosses — 34. Brachytheciaceae 291
annulus none; peristome parts of about equal length, hynoid, the ciha some-
what appendiculate, usually 2 in number; spores mature in fall.
On earth, rocks, roots and bases of trees, in woods, but not so frequently
occurring on rotten logs as do some of the other species. From eastern Can-
ada to Tennessee and westward.
Erie Co.: Entrance to Lily Pond, among trees. Presque Isle, July 13, 1925 and Sand
Bank Trail, July 25, 1925, Nelle Amnions, Washington Co,: On decayed wood near
Washington. Oct. 3, and 21 (figured), 1891. A Linn and J. S. Simonton.
2. Brachythecium campestre (Bruch) Bryologia Europaea
{Hypniirn campestre Bruch)
Plate LIV
Very closely resembling B. salebrosum, but differing in having the seta
smooth at base and slightly rough above; the capsule longer and the leaves
longer-acuminate. Otherwise the characters are as given for B. salebrosum.
On moist earth, often in grassy places, rocks, or on rotten logs, usually
preferring a non-calcareous habitat. Spores mature in autumn or early winter.
Europe, Asia, northern Africa, and, in North America from Canada to the
northern United States and south to Pennsylvania and New Jersey.
Infrequent in our region. Allegheny Co.: Darlington Hollow, Sharpsburg, Octo-
ber 25, 1908, and Power's Run, on shaded rock. November 30, 1909 (figured). O.E.J.
Fayette Co.: Ohio Pyle, along Meadow Run Valley, four miles south of village, Sep-
tember 1-3, 1906. O.E.J, and G.K.J. McKean Co.: Bennett Brook, August 26, 1894,
and Quintuple, September 9, 1896. D.A.B. Both near Bradford.
3. Brachythecium salebrosum [Hoffmann] Bryologia Europaea
{Hypnum salebrosum Hoffmann)
Plate LIV
Widely cespitose in glossy, dark yellow-green mats: stems usually 5 or 6
cm or more long, creeping and irregularly branching; stem-leaves lance-ovate,
about 1.5-2.5x0.6-1.1 mm, in our region apparently somewhat smaller than
most descriptions call for; branch-leaves similar, lanceolate, about 1.8-2.2x0.5-
0.6 mm, abruptly slenderly acuminate, serrate above, entire or sub-serrulate
below, concave, the lower margins narrowly reflexed, the narrow insertion
decurrent, both kinds of leaves plicate and erect-spreading; costa thin, usually
reaching to the middle or a little above; median leaf -cells linear-fusiform, flexu-
cus, about 8-12:1, the basal shorter and broader, usually two or three rows of
lax, rather large, oblong or sub-quadrate cells across the whole base of the leaf,
the alar more numerous, lax, sub-quadrate, rather thin-walled, the alae quite
strongly decurrent; perichaetial leaves filiform-acuminate, ecostste or nearly so:
seta smooth, castaneous, about 2-2.5 cm long, flexuous, flattened and twisted
when dry; capsule oblong-ovoid, dorsallv turgid, inclined to horizontal, usually
arcuate, about 2-3:1, castaneous, the urn about 2-2.5 mm long; the lid conic-
acuminate, about 1 mm long; annulus narrow; e.^othecial cells rounded-quad-
rate at the rim, larger and irregularly oblong or elliptic below, all strongly
292 American Midland Naturalist Monograph No. 6
yellowish-incrassate; peristome-teeth slender, confluent at base, closely trabecu-
latc and lamellate, dorsally cross-striolate and brownish below, hyaline and
papillose above, rather prominently margined; segments about as long as the
teeth, finely papillose, carinately split and usually gaping; cilia a little shorter,
hyaline, nodose, 1 to 3 in number; basal membrane about one-third as high
as the teeth; spores mature in late fall or winter, about .015-.020 mm, the walls
medium-incrassate, brownish, and somewhat papillose: autoicous.
On earth, stones, roots and bases of trees, rotten wood, etc., in moist, shady
woods; said to be especially common in pine or hemlock woods; Europe, Asia,
northern Africa, and from eastern Canada southward to North Carolina and
Missouri.
Common in our region. Now known from the following counties: Allegheny, Bedford,
Butler, Crawford, Clinton, Elk (Porter), Erie, Greene, Indiana, Lawrence, McKean, Som-
erset, Washington, and Westmoreland. Specimen figured: Douthett, Allegheny Co.,
April 26, 1908. O.E.J.
4. Brachythecium flexicaule Renauld and Cardot
Plate LV
Widely cespitose, yellowish-green: stems usually at least 3-6 cm long,
creeping, irregularly pinnate; leaves plicate, erect-spreading, the stem-leaves
lanceolate, about 1.8-2.5x0.6-0.9 mm; branch-leaves narrower, up to 2.4 x 0.5-
0.7 mm, gradually slenderly acuminate from a deeply concave, somewhat
decurrent plicate base with often narrowly reflexed basal margins, the margins
serrulate above; costa extending to above the middle of the leaf; median leaf-
cells linear-fusiform, prosenchymatous, flexuous, about 8-15:1, rather incras-
sate, the apical shorter, the basal rather abruptly shorter and wider with two
to four rows of large oblong to rounded-quadrate cells across the whole median
base, the alar cells sub-quadrate, rather incrassate, numerous, the wings decur-
rent; perichaetial leaves up to 3 mm long with slender flexuous acuminations,
partly sheathing, erostate or nearly so: seta smooth, castaneous, usually sinis-
trorse, 1.5-2.5 cm long; capsule about 3-4:1, oblong-cylindric, inclined to
nearly horizontal, dorsally gibbous, sub-arcuate, pale-castaneous, slightly nar-
rowed below the rim when dry, the urn from 2-4 mm long; lid conic-acumi-
nate, about 1-3 mm long; exothecial cells small and rounded at the rim, below
larger and oblong to linear-oblong, all sharply yellowish-incrassate; annulus
indistinct; peristome-teeth confluent at base, castaneous and dorsally cross-
striolate below, closely trabeculate and lamellate, margined, hyaline towards
apex; segments very slender, about as long as teeth, carinately cleft and gaping
in median portion, yellowish, papillose, basal membrane one-fourth to one-
third as high, the cilia somewhat shorter than the segments, filiform, nodose,
hyaline-papillose; spores rather incrassate, smoothish, brown-walled, .013-. 016
mm. According to Grout this is probably B. salebrosum variety densum Bry-
ologia Europaea. In most characters it is quite similar to typical salebrosum
but differs in having narrow leaves with evenly narrowed and very slender
acuminations.
Jennings: Manual oi Mosses — 34. Brachythfciaceae 293
Ranging from Newfoundland, New England and the Adirondacks to New
Jersey and Pennsylvania, and occurring also in British Columbia.
Rare in our region. Allegheny Co.: Guyasuta Hollow on clay and stones, Octo-
ber 12, 1908. O.E.J. Erie Co.: On log in hemlock woods. Presque Isle. Nelle Am-
mons. McKean Co.: Bennett Brook, May 3, 1893. D.A.B. (figured).
5. Brachythecium ACUTUM (Mitten) Sullivant
(Hypnum acntum Mitten )
Loosely ccspitose, bright glossy green: stems long, fle.xuous, creeping, basal-
ly radiculose, sparsely branched; branchlets short, sometimes reflexed; leaves
loose, open-spreading, more imbricate when dry, lanceolate to lance-ovate, ncn-
striate, slightly decurrent, plane-margined, scarcely concave, obscurely serru-
late or almost entire, short auriculate at base, the margins tapering gradually
and almost in a straight line from base to apex; median leaf-cells linear-vermic-
ular, about 10:1, the basal lax, the alar sub-quadrate, small, numerous and
extending down to form a rather strong decurrent portion; costa reaching to
somewhat above the middle; stem-leaves wider, triangular-ovate, reaching 2.5 x
1 mm, long and slenderly acuminate: seta smooth, about 1.5-2.5 cm long,
flexuous; capsule ovoid-oblong, dorsally turgid, inclined to horizontal, usually
slightly arcuate, about 2-3:1; annulus narrow; peristome hypnoid, the cilia 2
or 3, strongly nodose to sub-appendiculate; lid conic-acuminate; spores mature
in late fall or winter.
In moist woods on rotten logs and earth; Canada and the northern United
States, south to New Jersey, Pennsylvania, and Ohio.
Rare in our region. McKean Co.: D.A.B. (Porter's Catalogue).
6. Brachythecium rutabulum [Linnaeus] Bryologia Europaea
{Hypnum rittabulitm Linnaeus )
Plate LV
Widely and loosely cespitose. yellowish-green, glossy: stems prostrate,
creeping, often stolonifeious at the end, the branchlets more or less erect and
attenuate; stem-leaves large, cordate-ovate to more or less deltoid, or narrower
and lance-ovate, the wider ones abruptlv and rather shortly acuminate, the
narrower ones slenderly acuminate, the leaves varying in size up to 2.5 x 0.7-
1.5 mm, decurrent; the branch-leaves ovate to lance-ovate, about 1.7-2.0x0.6-
1.0 mm., concave, decurrent, the margin slightly serrulate all around, when
dry more or less reflexed at base and the leaves then somewhat plicate; costa
thin, reaching to the middle or beyond; median leaf-cells acutely elongate-
rhomboid or linear-rhomboid, usually about 10-20:1, the apical som.ewhat
shorter, the basal shorter and wider, incrassate especiallv in the stem-leaves,
the alar similar, except that a few are more enlarged, inflated, and oblong-
quadrate, but scarcely forming distinct auricles; perichaetial leaves up to 2.5
mm long, slenderly acuminate: seta 2-3 cm long, rough throughout, drving
flattened and twisted, castaneous, sinistrorse except sometimes at the very apex;
capsule about 2-3 x 1 mm, oval-oblong to sub-cylindric, unsymmetric, inclined
294 American Midland Naturalist Monograph No. 6
or more usually nearly horizontal, dorsally gibbous, arcuate, dark-castaneous;
lid conic to conic-acuminate; annulus broad, 2-3-seriate; peristome-teeth slender,
castaneous below, the apex hyaline and papillose, basally confluent, the lamel-
lae and trabeculae closely placed, teeth dorsally cross-striolate, margined;
segments slender, about as long as the teeth, yellowish, carinately split; basal
membrane about one-half as high as the segments, some of the cilia usually as
long as segments, hyaline, nodose, usually 2 or 3; spores usually minutely
roughened, somewhat incrassate, brownish, about .01 6-. 020 mm, maturing in
early winter.
In wet places on earth, stones, rotten wood, bases of trees, etc., in shady
woods and thickets; Europe, Asia, northern Africa, and, in North America,
from Canada to New Jersey and Pennsylvania.
Rather common in our region. Collected in 1 1 counties widely distributed in western
Pennsylvania: Allegheny, Armstrong, Butler, Erie, Fayette, Greene, Lawrence, McKean,
Warren, Washington, and Westmoreland. Specimen figured: Wildwood Road Hollow,
Allegheny Co., O.E.J. &C G.K.J. Nov. 19, 1908.
7. Brachythecium rivulare (Bruch) Bryologia Europaea
{Hypnum rivulare Bruch)
Plate LVI
Robust, cespitose in wide and thick mats, pale golden green, shining, rigid:
stems hard and woody, prostrate, filiform, leafless when old; branches irregu-
lar on the ascending or sub-erect and somewhat dendroid secondary stems
which usually reach a height of 3 or 4 cm; stem-leaves broadly ovate, rather
regularly imbricate wh'^n dry, erect -spreading or more open when moist, rather
distant, abruptly short-acuminate or acute, concave, decurrent, plicate, denticu-
late, reaching about 1.8-2.5x1.0-1.4 mm; branch-leaves similar to the stem-
leaves but usually wider, ovate to lance-ovate, decurrent, about 1.5x0.7 mm,
quite concave, dentate above, the margins plane or reflexed below, often some-
what plicate; median leaf-cells linear, about 10-15:1, prosenchymatous with
rounded ends, rather incrassate, the apical shorter, the basal abruptly laxer,
shorter, wider, the median basal usually with incrassate and porose walls, the
alar abruptly differentiated, more or less enlarged, inflated, hyaline to orange-
pellucid, forming distinct and widely decurrent auricles; costa often forking,
reaching to the middle or above; seta 1.5-2.5 cm long, strongly papillose
throughout, castaneous; capsule castaneous, turgid- to oblong-ovate, about 2-
3x1 mm, more or less arcuate, inclined to more or less horizontal; lid conic-
acuminate; annulus 2-seriate; exothccial cells at rim small and rounded, below
larger and rounded-oblong; peristome-teeth castaneous below, apically hyaline
and papillose, basally confluent, strongly trabeculate, distinctly margined by
the projecting edges of the cross-striolate dorsal lamellae; segments nearly as
lone, carinately split and gaping, yellowish, the basal membrane about one-half
as high, cilia 2 or 3, nodose, slender, about as long as the segments; spores
smoothish, the walls somewhat incrassate and greenish-brown, about 0.16-0.20
mm, maturing in fall.
Jennings: Manual of Mosses — 34. Brachytheciaceae 295
On wet rocks in or at the margin of streams, swamps, or in wet pLices in
ravines, usually where often submerged; Europe, Asia, and from Canada to
Missouri and Virginia. Rather rare in our region. Quire variable with a
number of named varieties.
Now known from the following counties: Allegheny, Beaver, Butler, Cambria (Por-
ter), Crawford, Erie. Fayette. McKean, Somerset, Warren, Washington, and Westmore-
land. Specimen figured: Pymatuning Swamp, Lmesville, Crawford Co., May 10-11, 1906.
O.E.J.
8. Br.^chythecium reflexum [Starke] Bryologia Europaea
{Hypnum reflexum Starke; H. subtenue ]ames\ Thuidium laxifolium Macoun )
Very slender, dark green, densely intertwining to form low, flat patches:
the branches short, delicate, often curved, more or less pinnatelv arranged;
stem-leaves cotdate-triangular, quickly narrowed to a fine, long, often twisted
acumen, strongly decurrent, minutely serrulate all around; branch-leaves nar-
rower, lanceolate, strongly decurrent, serrulate, smooth to faintly plicate,
margins plane to very narrowly recurved, when dry spreading or imbricate and
rendering the branches rather julaceous; costa strong, reaching to apex or even
mto the acumen; leaf-cells short and broad, about 3-8:1, rhomboid-fusiform,
sub-obtuse, rather incrassate, towards the basal angles becoming gradually
shorter and broader, the alar large, pellucid, rounded-quadrate to rounded-
rectangular, numerous and extending up the sides of the leaf but not forming
very clearly distinct auricles: seta slender, about 1-1.5 cm long, rough; capsule
small, about 2 mm long, about 2:1, ovate-globose, curved, dorsaliy turgid,
abruptly horizontal, almost black when old; lid convex-conic, apiculate; annu-
lus narrow, 2-seriate; cilia slender and appendiculate; spores mature in winter:
autoicous.
On rocks and tree-trunks in mountainous or hilly regions; Europe, Asia,
and from Garrett Countv, Maryland (J. Donnell Smith), and westward.
Rare in our region. McKean Co.: Bradford. D. A. Burnett. (Porter's Catalogue).
9. Brachythecium Starkei [Bridel] Bryologia Europaea
(Hypnum Starkei Bridel)
Plate LVI
Dark green, widely and thinly cespitose, the plants usually quite distinctly
complanate: stems slender, creeping, radiculose, pinnate v/ith short, curved-
sscending, rather distant, slender branches; branch-leaves loose, divergently
spreading, often somewhat secund, those from the middle of the branches
broadly ovate to broadly triangular-cordate, abruptly and usually rather shortly
slender-acuminate, apically twisted, broadly decurrent, marginally serrate
above, denticulate below; costa variable but usually about three-fourths as long
as the leaf; median leaf -cells about 8-15:1, fusiform-hexagonal to fusiform-
rhomboid, sometimes shorter, somewhat incrassate; the basal in one or two
rows more or less rectangular-oblong, the alar rather numerous, sub-rectangular,
with thick and often brownish or yellowish walls, forming quite disitnct auri-
296 American Midland Naturalist Monograph No. 6
cles; stem-leaves usually smaller than branch-leaves, broadly ovate and broadly
lon^-acuminate : seta papillose, about 2 cm long, flexuous, slender, castaneous,
rough; capsule small, turgid-oval, often blackish when ripe, the urn about 2.5-
3x1 mm, dorsally gibbous, abruptly more or less horizontal, sub-globose when
empty; annulus large; exothecial cells rounded-quadrate and small at the rim,
oblong-rectangular and a little larger below, all strongly castaneous or yellow-
ish and incrassate; peristome-teeth castaneous below, set far back from the
edge of rim, margined, rather widely confluent at base, lamellate, cross-striolate
dorsally below, hyaline and papillose at apex, strongly trabeculate; segments
slender, nearly as long as teeth, carinately split and often widely gaping in the
middle, yellowish; basal membrane about two-fifths as high as the teeth, the
cilia 2 or 3, strongly appendiculate, hyaline granular, a little shorter than the
segments; spores about .012-. 01 5 mm, greenish-yellow or brownish, slightly
roughened, medium.-walled, mature in winter.
On moist, rotten wood, stumps, bases of trees, earth, in moist woods in
hilly or mountainous regions; Europe, and from Canada to northern United
States as far south as New Jersey and Pennsylvania.
Rather rare in our region. Elk Co.: Benezette. McMinn. (Porter's Catalogue).
Lawrence Co.: Kennedy's Mills. Kellar Shelar. Nov. 20, 1931. McKean Co.: On
shaded banks along Marilla Brook, Bradford, April 25, 1897. D.A.B. (figured).
Washington Co.: On stone near creek, Snowden Sta. Dec. 3, 1892. Linn & Simon-
ton, No. 96.
10. Brachythecium veluTINUm [Linnaeus] Bryologia Europaea
(Hypnum velutinum Linnaeus; H. declivum Mitten)
Plate LVI
Slender and usually in low, soft, silky mats, bright or yellowish-green,
prostrate: stems radiculose; branches numerous, short, in our specimens the
branches usually less than 5 mm long, crowded, irregular or curved, more or
less sub-pinnate; branch-leaves loosely erect-spreading to falcate-secund at tips
of branches, more widely spreading when drv, lanceolate to lance-ovate, in
ours mainly 1-1.5 mm long, tapering to a long acumination, serrate, apically
often twisted, shortly decurrent, faintly plicate, glossy when dry, marginally
plane; costa slender, reaching somewhat beyond the middle; median leaf-cells
narrow-linear, rather obtuse, about 8-15:1, the apical similar but a little shorter,
the basal shorter, the alar few, rather opaque, incrassate, sub-quadrate; the
stem-leaves similar but usually not so large as some of the branch-leaves; p>eri-
chaetial leaves erect, slenderly acuminate, up to 1.8 mm long; seta about 1.5
cm long, very rough, castaneous, often flattened and twisted when dry; cap-
sule about 2-2.5 mm long, 2-3:1, turgid-oblong, dorsally gibbous to sub-
arcuate, castaneous, inclined to horizontally spreading; exothecial cells small
and rounded-quadrate at rim, oblong-rectangular below, all densely incrassate;
peristome-teeth slender, castaneous and confluent at base, apicallv hyaline and
papillose, dorsally cross-striolate, closely trabeculate and lamellate; segments
as long as the teeth, slender, carinately split between the nodes, yellowish, the
Jennings: Manual of Mosses--34. Brachytheciaceae 297
basal membrane one-third to two-ftfths as high; ciha 2 or 3, nodose, hyaHne,
somewhat shorter than the segments; hd conic-acuminate, about 0.5-0.8 mm
long; annulus large; spores mature in winter, faintly roughened, medium-
walled, brownish, .013-.016 mm in diameter.
On earth, rocks, bases of trees, rotting wood, etc., in rather dry woods,
often on knolls; Europe, Asia, and from Canada south to New Jersey and
Pennsylvania and also in the Pacific States.
Thus far reported but once in our region. McKean Co.: Langmade, near Bradford,
April 25, 1898. D.A.B. (figured).
11. Brachythecium populeum (Hedwig) Bryologia Europaea
(Hypnum populeum Hedwig)
Slender, densely cespitose in small yellowish-green tufts, lustrous: stems
procumbent, branched with numerous, more or less pinnately-arranged, erect
or curved-ascending branches; leaves of stem and branches similar e.xcept that
the branch-leaves are narrower and lanceolate; stem-leaves rather closely imbri-
cated, erect to erect-spreading when dry, ovate-lanceolate, serrate to nearly
entire, slenderly and gradually acuminate, non-striate, shortly decurrent; costa
strong and reaching the apex; median le=>-f-cells about 5-8:1, sometimes rela-
tively longer, the basal more or less rectangular, the alar numerous, often
yellowish but rather opaque; seta rough except towards the base, where nearly
smooth, dark brown; annulus persistent, simple, narrow; capsule about 2:1,
cernuous, turgid-ovate to ovoid, mostly dorsally gibbous, glossy, constricted at
the mouth when dry; lid shcrt-acuminate; peristome norm.al, cilia short, usually
1 or 2 and unequal, appendiculate; spores mature in winter; autoicous.
On roots of trees, stones, sometimes on bases of trees, in shadv woods, said
to be somewhat partial to pine woods; Europe, northern Africa, and from
Nova Scotia to North Carolina and in British Columbia.
Rare in our region. McKean Co.: "B. populeum rufescens." Bradford. D.A.B.
(Porter's Catalogue).
12. Brachythecium flagellare (Hedwig) Jennings
{Hypnum flagellare Hedwig; Hypnum plumosum Swartz;
B. plumosum (Sw. ) Bryologia Europaea)
Plate LVII
Robust in loose, wide, green mats, brownish below: stems prostrate, up to
5 or 6 or more cm long, with rather densely pinnate branches; the branches
j-tout, ascending to erect, somewhat tumid with the closely imbricate, concave
leaves; leaves crowded, erect-spreading when moist, imbricated when dry, often
quite strongly falcate-secund, the branch-leaves lanceolate to broadly lance-
ovate, abruptly slenderly acuminate, about 1.3-2.0x0.4-0.9 mm, decurrent,
serrate above to nearly entire, the base very concave som.ewhat excavate at the
298 American Midland Naturalist Monograph No. 6
alae, narrowed, sometimes striate when dry, margin plane or slightly recurved
at the base; costa reaching to the middle or a little farther; median leaf-cells
narrow to linear, about 8-15:1, the apical shorter, the basal shorter, the median
basal enlarged, rounded to oblong ,incrassate, sometimes porose, the alar some-
what smaller, oblong to sub-quadrate, incrassate and somewhat opaque; stem-
leaves similar, rather scattered, usually smaller and narrower, about 1.5x0.6-
0.7 mm, narrowly triangular-ovate; perichaetial bracts more or less erect, partly
sheathing: seta papillose in the upper half, brown to blackish, stout, 1.5-2.0
cm long, sinistrorse below, sometimes dextrorse above; capsule about 1.5-2.5 x 1
mm, turgidly oval-oblong, blackish when old, dcrsally gibbous, horizontal to
sub-erect, somewhat unsymmetric; lid conic-acute about 0.6-0.8 mm long; annu-
lus simple, persistent; peristome-teeth castaneous, confluent at base, strongly
irabeculate and lamellate, prominently margined by the projecting lamellae,
dorsally cross-striolate below, hyaline and papillose at apex; segments narrow,
carinately split but usually not widely gaping, yellowish, nearly as long as
teeth, the basal membrane about one-third as high; cilia 2, nodose, hyaline,
appendiculate below, about as long as the segments; spores smooth, medium-
walled, brownish, about .013-.017 mm, mature in autumn.
In streams, or on moist rocks, in non-calcareous habitats; Europe. Asia,
Hawaiian Islands, and from Newfoundland to British Columbia and south in
the mountains to Alabama; Florida. Very common in our region.
Now known from the following counties: Allegheny, Armstrong, Beaver. Bedford,
Butler, Centre, Erie, Fayette, Greene, McKean, Washington, and Westmoreland. Speci-
men figured: Wildwood Hallow, Allegheny Co., Nov. 19, 1908. O.E.J, and G.K.J.
12a. Brachythecium flagellare var. homomallum
(Bryologia Europaea) Jennings
(B. plumosum var. homomallum Bryologia Europaea)
Plate LVII
This variety differs from the type of the species in having the leaves dis-
tinctly falcate-secund and branches curved at tip. It is said to be generally
smaller with narrower leaves and with the capsule small and ovate. In the
same pockets with typical B. flagellare can often be found specim.ens with
characters approaching more or less closely the variety. The following pocket
of specimens perhaps typical of the variety:
McKean Co.: Gate's Hollow, Bradford, April 18, 1897. D A.B. (figured).
5. Cirriphyllum Grout
Dioicous: robust, widely cespitose, whitish to yellowish-green, rarely darker,
mostly lustrous: stem creeping to ascending, often stolon-like, pinnately to
fasciculately branched, often with flagellae; branches ascending to erect, more
or less densely-leaved and julaceous; leaves uniform, often spreading, often
drying imbricate, very concave, somewhat weakly plicate, ovate to oblong from
a somewhat narrowed and decurrent base, more or less abruptly lanceolate to
JtNNiNGS: Manual of Mosses — 34. Brachytheciaceae 299
pilifcrous at the apex, plane-margined, serrate to entire; costa simple, ending
at or above the middle of the leaf, never ending in a dorsal spine; median
leaf-cells narrowly prosenchymatous, smooth, the basal, shorter, thickened, and
porose, the alar more or less numerous, short-rectangular to quadrate, mostly
green; inner perichaetial leaves from a sheathing base abruptly long and finely
acuminate: seta elongate, mostly rough; capsule cemuous to horizontal, oval
to oblong-oval, more or less dorsally gibbous, rarely erect and sub-cylindric;
annulus present; peristome as in Brachythecium; lid usually more or less long-
rostrate from a conic base.
A small genus of about 15 species, mostly in temperate regions on rocks
and earth; 4 or more species in North America; 2 species in our region.
Key to the Species
A. Stems without stolons, almost regularly pmnate; the acumination about one-half as
long as the body of the leaf: seta rough 1. C. ptliftrum
A. Stem with stolons, irregularly branched; the leaf-acumination short: seta smooth
2. C. Boscii.
1. CiRRlPHYLLUM PILIFERUM [Schreber] Grout
(Hypnum piliferum Schreber; Eurynchium piliferum Bryologia Europaea )
Robust, in loose straggling patches, glossy yellow-green: stems elongate,
up to 10 or 15 cm long, prostrate, creeping, radiculose, more or less pinnate;
the ends of the stems and branches of a paler shining green; leaves concave,
widely oblong-ovate, spoon-shaped, abruptly hair-pointed from the rounded
apex, the piliferous acumination often reaching one-half the length of the main
portion of the leaf, towards the apex of the stems and branches the leaves m.ore
closely imbricate and forming cuspidate terete points, but with the piliferous
leaf-tips flexuous-spreading, leaf-margin usually denticulate, plane or inflexed;
when dry the leaves striate; median leaf-cells about 10-15:1, the basal more
lax, shorter and wider, the angular forming a well-defined patch, large, oval-
rectangular; the branch-leaves somewhat smaller, narrower and more gradually
pointed; costa broad at base, reaching to about three-fourths the length of the
leaf: seta about 2.5 cm long, rough; capsule ovoid-oblong to turgid, somewhat
arcuate, when dry and empty strongly arcuate and constricted below the mouth,
about 2 mm long; lid conic with a subulate beak about as long as urn, 2 mm;
peristome large, teeth long, the segments about as long, the cilia nodose to
sub-appendiculate, 2 or 3, about as long as the segments; spores mature in fall
but capsules rarely found.
In wet woods and swampy meadows, on the ground or on the bases of
trees; Europe, and from New Brunswick to New Jersey, Pennsylvania, and
Ohio; Washington.
Rare in our region. Elk Co.: Benezette. McMinn. (Porters Catalogue). McKean
Co.: D. A. Burnett. (Porter's Catalogue).
300 American Midland Naturalist Monograph No. 6
2. Cirriphyllum Bosch (Schwaegrichen) Grout
{Hypnum Boscii Schwaegrichen; Eurynchium Boscii Jaeger).
Spoon-Leaved Moss
Plate LVII
Loosely cespitose in large, golden-green mats, the older portions blackish,
robust: stems up to 8-10 cm long, prostrate, somewhat pinnately branching,
the branches mostly simple, erect, turgid-terete; leaves closely to loosely imbri-
cate, large, about L5-2.5 mm long, spoon-shap>ed, abruptly acuminate, the
acumination filiform and twisted, the leaves oblong-ovate, scarious, shining;
costa double and short, or simple and reaching to the leaf-middle or beyond;
meadian leaf-cells narrowly linear-rhomboid, the marginal shorter and mainly
rhomboid, the basal short, wide, yellowish-brown, pellucid, irregularly oblong
to rectangular, larger but shorter, the alar incrassate, quadrate, forming an
indistinct group, the apical shorter and wider than the median, the median
about 6-10:1; perichaetial leaves narrowly long-acuminate, the inner erect:
seta smooth; capsule oblong, about 2.5-3:1, the urn about 2 mm. long, inclined,
sub-arcuate; lid sharply obliquely rostrate, about 1 mm long; annulus 2-seriate;
peristome normally hypnoid with somewhat split segments and cilia 3, about
as long as segments, nodose; spores mature in fall, about .016 mm.
On earth or rocks in moist woods, often at the edges of the woods, or even
in the fields; from New England to Florida and westward to Missouri.
Probably fairly common in our region. Cambria Co.: (Porter's Catalogue), Hun-
tingdon Co.: Pennsylvania Furnace, July 13, 1909. Washington Co.: Linn and
Simonton. (Porter's Catalogue). Westmoreland Co.: Hillside, May 22, 1909. O.E.J.
(figured ) .
6. Oxyrhynchium (Bryologia Europaea) Wamstorf
Mostly dioicous: slender to robust, laxly to densely cespitose, dark to yel-
lowish-green, drying soft or stiff, dull to lustrous: stem creeping or ascending,
often stolon-like, often bearing rhizoids, irregularly pinnate to fasciculately
branched; branches mostly complanately-leaved, stem-leaves and branch-leaves
sometimes different, sometimes similar except in size, non-plicate, little or not
concave; stem-leaves erect-spreading to squarrose, from a somewhat narrowed
and sometimes decurrent base ovate to triangular oval, with short and broad or
somewhat longer apex, plane-margined, somewhat serrate; costa simple, ending
at or above the leaf-m.iddle, often ending in a dorsal spine; median leaf-cells
narrowly prosenchymatous, smooth, the basal shorter, mostly incrassate and
porose, the alar differentiated: seta elongated, mostly red, quite thick, mostly
rough; capsule cemuous to horizontal, sometimes sub-erect, thickly oval to
oblong-ovate, dorsally gibbous; annulus present; peristome as in Brachythe-
cium; lid long and obliquely subulate-rostrate; calyptra glabrous.
A genus of about 20 sp)ecies, on damp and shaded rocks, stones, or some-
times in water, mostly in temperate regions; 2 species in our region.
Key to the Species
A. Aquatic: alar leaf-cells forming a slightly differentiated group: seta smooth
1. O. riparioides
A. Terrestrial : alar-cells not differentiated: seta roughly papillose 2. O. hians
Jennings: Manual of Mosses — 34. Brachytheciaceae 301
1. OxYRHYNCHiUM RiPARioiDES [Hedwig] Jennings
(Hypnum ntfciforme Necker; Emyncbium rusciforme Milde; Hypiium
riparioides Hedwig; Rhynchostegium rusciforme Bryologia Europaea )
Plate LVIII
Robust, in large tufts, dark to blackish below: stems prostrate, woody, and
usually denuded below; branches suberect, or ascending, usually more or less
rigid and harsh, especially when dry; leaves ovate, loosely ascending or erect-
spreading, scarcely decurrent, about 2-2.5x1.5 mm, obtuse to acute, plane-
margined, denticulate nearly to the base; costa thick below, reaching to one-
half or two-thirds the length of the leaf, or occasionally even sub-percurrent,
often ending in a dorsal spine; median leaf-cells incrassate, linear-fusiform,
about 10-12:1, the apical and basal shorter and broader, but no alar group
differentiated, the median and upper slightly dorsally spinose- seta smooth,
about 1.5 cm long, castaneous, slightly twisted when dry; capsule castaneous,
ovoid-oblong, somewhat constricted below the mouth when dry, about 2-3:1,
dorsally turgid but scarcely curved, inclined or nearly horizontal, the urn about
1.5-2 mm long; lid obliquely slenderly rostrate from a conic base, about two-
thirds as long as the urn; annulus revoluble, usually 2-seriate; exothecial cells
vellowish-incrassate, at the rim small and rounded-quadrate, below rather large
and irregularly oblong-rectangular; peristome-teeth slender, apically hyaline-
papillose, strongly trabeculate, dorsally plainly lamellate and finely cross-
Jtriolate, margined, confluent at base; segments about as long, usually carinate-
ly widely gaping but remaining unsplit at apex, the basal membrane about one-
half as high; cilia 2-3, subulate, nodose to sub-appendiculate, somewhat shorter
than the segments; spores weakly papillose, medium-walled, yellowish, about
.010-. 01 3 mm, mature in early fall.
On wet or submerged rocks in streams and rivulets; Europe, Asia, northern
Africa, and from Newfoundland to Ontario and southwards in the mountains
to Georgia.
Common in our region. Now known from the following counties: Armstron'^, Bed-
ford, Butler, Cambria (Porter), Cameron, Centre, Fayette, Lawrence, McKean, Somer-
set, Warren, Washington, and Westmoreland. Specimen figured: Rachelwood, Mellon's
Estate near New Florence, Laurel Hill Mts., Westmoreland Co., Sept. 8-11, 1907.
O.E.J.
2. OXYRHYNCHIUM HIANS (Hedwig) Jennings
(Hypnum hians Hedwig; Eurynchium htans Jaeger and Sauerbeck; Hypnum
praelongum C. Mueller; Pterygynandrum apiculatum Bridel )
Plate LVIII
Rather slender, depressed, cespitose, somewhat shining: stems creeping,
rather sparsely branched, slender, usually not over 3 or 4 cm long, the branches
short and more or less distichouslv arranged; leaves of the stem and longer
branches rather distant, on some of the short branches sometimes more or less
imbricated-julaceous, the stem-leaves about 1-1.6 mm long by three-fourths as
wide, ovate, the apex abruptly acute to shortly acuminate, the base clasping
but not decurrent, margins sharply serrulate nearly to the base; branch-leaves
302 American Midland Naturalist Monograph No. 6
closely similar; costa distinct, reaching to one-half to four-fifths the length of
the leaf, dorsally ending in a spine; median cells about 5-8:1, prosenchymatous,
medium-walled, the apical rhomboid, shorter, about 2-4:1, the basal shorter
and incrassate, the alar forming an indistinct group of thick-walled quadrate
to rectangular cells; perichaetial leaves up to 2 mm long, ovate-oblong, sheath-
ing, acuminate, serrate above: seta dark-castaneous, stout, strongly papillose,
1-1.5 cm long; capsule inclined to horizontal, arcuately oblong-cylindric, nar-
rowed below the rim but slightly when dry, the urn about 2 mm long by 1
mm thick, castaneous; operculum conic and slenderly rostrate, yellowish, about
1 mm long; exothecial cells yellowish-incrassate, at the rim rounded-quadrate
in about two series, below larger oblong-rectangular; annulus narrow, 2-
seriate; peristome-teeth castaneous, slender, hyaline-papillose at apex, strongly
trabeculate, narrowly margined, the dorsal lamellae often in three series
towards the base, striolate in various directions; segments about as long as
teeth, slender, narrowly carinately gaping between nodes, the basal membrane
about two-fifths as high as teeth, the cilia usually t\vo, slender, nodose to short-
ly appendiculatc, nearly as long as segments; spores slightlv papillose, yellow-
ish, medium-walled, about .011-.015 mm in diameter, mature in late fall or
early winter.
On the ground in moist, shady places in woods, etc., in Europe, Asia, and,
in North America from Nova Scotia south to the Gulf east of the Mississippi.
Apparently not common in our region. Allegheny Co.: Clay stream-bank, South
Park. H. N. Mozingo. March 11, 1945. Erie Co.: In woods near Lagoon Boathouse,
Presque Isle, Nelle Ammons. July 22, 1933. Fayette Co.: Ohio Pyle, September 1-3,
1906. O.E.J, and G.K.J. McKean Co.: On shaded banks of rivoilet, Bennett Brook,
April 9, 1893, Marilla Brook, September 29, 1894 (figured), and on ground over leaf-
mold, April 19, 1897. All near Bradford. D.A.B. Washington Co.: Shady bank,
Oak Grove Station, Nov. 5, 1892; on the ground. Oak Grove Station, Dec. 5, 1891;
and Snake Woods, Nov. 19, 1892,^ in the vicinity of Washington; and on stone, Clays-
ville, Dec. 17, 1892 all A. Linn and J. S. Simonton.
7. EuRHYNCHlUM Bryologia Europaea
Dioicous and pseudoautoicous : slender to robust, laxly or densely cespitose,
green to yellowish, drying stiff and more or less lustrous: stem creeping to
ascending, often more or less stolon-like, here and there fasciculate, often bear-
ing flagellae, pinnate to fasciculate or even dendroid; branches more or less
densely-leaved; leaves often dimorphic, mostly plicate; stem-leaves spreading
to squarrose, more or less concave, ovate-cordate to triangular-cordate from a
narrowed and more or less decurrent base, margins plane, serrate, the apex
short and broad to long and narrow; costa simple, more or less elongate, often
ending as a dorsal spine; median leaf-cells smooth, prosenchymatous, narrow,
at base shorter and usually incrassate and p)orose, the alar differentiated; inner
perichaetial leaves with squarrose-reflexed, subulate tips: seta mostly smooth;
capsule cernuous, sometimes horizontal, ovate to sub-cylindric, more or less
dorsally gibbous; peristome as in Brachythecitim; lid long and finely rostrate;
calyptra glabrous.
A genus of about 14 species, on rocks, earth, or bark, almost entirely in
Jennings: Manual of Mosses— 34. Brachytheciaceae 303
temperate regions; about 6 species in North America; probably only one species
in our region.
1. EuRHYNCHiUM PULCHELLUM (Hedwig) Jennings
(Hyptiiim pulcheUum Hedwig; H. strigosum Hoffmann; Eurhynchitim
strigosum Bryologia Europaea)
Specimens collected by Nelle Ammons on Presque Isle may belong to this
species, but our region is represented mostly by the following varieties of this
species.
la. EuRHYNCHiUM PULCHELLUM var. ROBUSTLJM (Roell) Jennings
(E. slTtgosum var. rohtisliim Roell; Hyprium strigosum Drummond )
Common Beaked Moss
Plate LVIII
Loosely and widely matted or densely tufted, bright and shini;-ig green:
stems stoloniferous, creeping, with distant leaves; secondary stems prostrate to
erect, often curved, rather robust; leaves on the middle of the branches erect-
spreading, lance-ovate, scarcely decurrent, reaching about 1-1.2 ,x 0.4-0.5 mm,
acute to widely obtuse, plane-margined, sharply serrate above, concave, some-
what plicate, costate to about two-thirds, the costa usually ending in a dorsal
spine; median leaf -cells about 8-10:1, linear to linear-rhomboid, the apical
becoming rhomboid-oblong and about 2-3:1, the basal somewhat shorter than
the median, the alar few, rectangular to quadrate or oval; stem-leaves decurrent,
rather long-acuminate from an ovate to triangular-ovate base, somewhat larger
than the branch-leaves, reaching about 1.2-1.5 mm long, serrate nearly to the
base, costate to about two-thirds; paraphyllia small, rounded-ovate; leaves on
the stolons ecostate, triangular-ovate, small, acuminate: seta castaneous,
smooth, about 1-1.5 cm long, drying dextrorse above; capsule yellowish-brown,
cblong-ovate, about 2-3:1, more or less dorsally turgid or sub-arcuate, drying
slightly constricted below the mouth, inclined or almost horizontal, the urn
about 2 mm long; annulus 2-3-seriate; lid convex, slenderly rostrate, about 1.5
mm long; exothecial cells rounded-quadrate at rim, oblong-hexagonal to rec-
tangular below, incrassate; peristome-teeth hyaline and papillose at apex, below
dorsally cross-striolate, margined, plainly lamellate, strongly trabeculate, con-
fluent at base; segments narrow, nearly as long as the teeth, carinately split
between the nodes, yellowish, the basal membrane about two-fifths as high:
cilia 3, slender, hyaline, noc'ose, usually one or two of them nearly as long
as the segments; spores yellowish, incrassate, papillose, about .012-.014 mm,
mature in autumn.
The species occurs on gravelly or sandy soil, rocks, roots of trees, etc , in
open woods in Europe, Asia, northern Africa, and from Arctic America to
northern United States. The variety rohuslum occurs from eastern Canada
south to Louisiana.
Elk Co.: McMinn. (Porter's Catalogue). McKean Co.: Si.x pockets of specimens
304 American Midland Naturalist Monograph No. 6
collected on the ground, or on rocks in woods, near Bradford, May 13, 1893, to Septem-
ber 29, 1896 (figured). D.A.B.
lb. Eurhynchium pulchellum var. praecox (Hedwig) New Combination
(£. stngosum var. praecox Husnot; Hypnum praecox Hedwig)
Loose yellow-green mats; creeping and often stoloniferous; branches erect,
julaceous about 4-5 mm long; branch-leaves erect-ascending when moist, im-
bricate dry, decurrent ovate-cordate, acute to bluntly obtuse, serrulate all
around, more or less plicate; costa to % length of leaf, ending in dorsal spine;
median leaf-cells 6-8:1, apical short and wide, basal and alar numerous and
quadrate: seta about 1 cm long; capsule ovoid, somewhat curved, horizontal,
about 2:1; lid long-rostrate, 2/3 length of urn; cilia 1-3, nodose; spores .010-
.012 mm, ripe in autumn.
Moist shady soil or rocks. New York, New Jersey, and eastern Pennsyl-
vania.
Bedford Co.: Sulphur Springs. C.M.B. Sept. 29, 1940.
8. Bryhnia Kaurin
Dioicous: more or less slender, weak, widely and laxly cespitose, more or
less dark green, when old yellowish or brownish, rather dull: stem elongate,
procumbent, rhizoids fascicled, branching interruptedly pinnate, some of the
shoots in the middle of the tufts often erect and tree-like but later procumbent
and giving rise to new shoots; branches usually spreading to recurved, thm,
acute, mostly laxly-leaved; paraphyllia none; stem-leaves loosely imbricate,
more or less concave, irregularly plicate, triangular-cordate to lance-ovate from
a widely decurrent and non-auriculate base; shortly or more slenderly pointed,
plane-margined, finely serrate all around; costa simple, ending in or over the
leaf-middle, smooth; median leaf-cells incrassate, green, oblong-rhomboid to
oblong-hexagonal, the basal lax, a few alar rectangular; branch-leaves mostly
dorsally rough by projecting cell-angles, sharply serrate all around; costa often
ending dorsally in a spine; inner perichaetial leaves oblong, abruptly narrowed
to a reflexed-squarrose, long, serrate acumination: seta 8-15 mm, dark red,
very rough; capsule cernuous to horizontal, dorsally gibbous, oval to oblong-
cylindric; annulus present; peristomes of equal length, the teeth basally con-
fluent, dorsally cross-striate, normally lamellate, apically papillose; inner peri-
stome yellow, finely papillose, basal membrane high, segments lanceolate, long-
subulate, split and finally gaping along the keel, cilia well-developed; lid more
or less plainly and thicky sub-rostrate from a conic base; calyptra glabrous.
A small genus of 10 species, occurring on various substrata, confined to the
Northern Hemisphere; 3 species in North America; 2 species in our region.
Key to the Species
A. Branch-leaves acute to short-pointed, the apex mostly twisted 1. B. novae-angliai
A. Branch-leaves acuminate, the apex not twisted 2. B. graminicolor
Jennings: Manual of Mosses — 34. Brachytheciaceae 305
1. Bryhnia NOVAE-ANGLIAE (Sullivant and Lesquereux) Grout
(Hypnum novae-angliae Sullivant and Lesquereux; Brachythtcium
novae-angliae Jaeger and Sauerbeck)
Plate LXIX
Widely and loosely matted, bright green outside, dirty green inside, rather
rigid: stems prostrate, irregularly sub-pinnately branched, sometimes more or
less dendroidal in appearance; branches often indistinctly julaceous; branch-
leaves rather loosely imbricate when dry, erect-spreading when moist, broadly
ovate, acute to shortly acuminate, concave, decurrent, serrulate, up to 1-1.2 x
0.5-0.6 mm, dorsally papillose by reason of the projecting cell-angles, the leaf-
apex often twisted about half-around; median leaf-cells about 5-6:1, oblong-
hexagonal, somewhat shorter and broader below and at the basal angles; peri-
chaetial leaves ovate, abruptly long-acuminate, faintly costate; costa of branch-
and stem-leaves reaching to the middle or slightly beyond; stem-leaves similar
to the branch-leaves but more broadly triangular-ovate and with more distinct-
ly quadrate alar cells, occasionally some inflated, and somewhat excavate: seta
."-hort, very rough, dark castaneous; capsule dark-castaneous, blackish when old,
about 4-5:1, reaching about 3.5-4.5 mm in length, oblong, erect, slightly
curved; lid conic-acuminate or sub-rostellate; peristome normal; annulus
double, large; spores mature in winter.
On the ground and on stones in swamps and wet, shady places; Europe,
Asia, and from eastern Canada to Maryland and Pennsylvania.
Allegheny Co.: Swampy spot. Deer Creek at Middle Road. C.M.B. Oct. 29,
1949. Butler Co.: Swampy soil, Criders Corner. O.E.J. Dec. 29, 1908 (figured):
West Liberty Bog outlet. C.M.B. Nov. 26. 1948. Fayette Co.: Hollow below Cu-
cumber Falls, Ohiopyle. C.M.B. June 22, 1940. Somerset Co.: Wet rock by spring,
1 mi. s. of Bakersville. Aug. 10, 1945; and Beck Spring, Laurel Hill. C.M.B. Aug.
23, 1946; Cranberry Glade Run. C.M.B. June 28, 1942. Warren Co.: South side
Allegheny River, 2 mi. s. of Big Bend. C.M.B. Sept. 2, 1935. Westmoreland Co.:
On rock, in Meadow Run. Jacobs Creek. C.M.B. Aug. 18. 1945; and Chestnut Ridge s.
of Torrance. C.M.B. June 13, 1943.
2. Bryhnia graminicolor [Bridel] Grout
{Hypnum graminicolor Bridel; H. Sullivanlii Spruce;
Eurhynchium graminicolor Paris)
Plate LIX
Small, much more slender than the preceding species, densely to loosely
cespitose, pale green, yellowish below: stem.s slender, red, usually not over 1-2
cm long, rather irregularly branched with erect branches; branch-leaves reaching
about 0.8 x 0.8-0.3 mm, narrowly lance-ovate, long-acuminate, concave, serru-
late to the base, marginally refiexed below, the base decurrent, the back strong-
ly papillose by reason of the projecting cell-angles, the costa reaching to above
the middle; stem-leaves larger, up to 0.8-1.0x0.4-0.5 mm, with a somewhat
more slender acumen; median leaf-cells linear-flexuous, incrassate, varying from
■^-8:1, obtuse, the alar sub-quadrate, thin-walled, pellucid; perichaetial leaves
cblong, basally sheathing, filiform-acuminate, very faintlv costate: seta about
306 American Midland Naturalist Monograph No. 6 .
1 cm long, rough throughout; capsule oval to oblong or turgid-ovate, dorsally
somewhat gibbous about 2-3:1, inclined, about 2 mm long; annulus simple,
persistent; lid conic to short- rostrate; peristome normally hypnoid, segments
shorter than the teeth, carinately split, the cilia 2, somewhat shorter; rather
uncommon, capsules rarely produced.
In moist woods and shady places on rocks or earth; from New Brunswick
to Minnesota and south to Georgia.
Rare in our region. Huntingdon Co.: Alexandria. T. C. Porter (Porter's Cata-
logue). McKean Co.: Bolivar and Bennett divide on shaded dripping rocks, April 21,
1895 (figured), and on perpendicular faces of rocks, Lewis Run, April 25, 1895. D.A.B.
Warren Co.: South Fork of Ten-Mile Creek, 2 miles southwest of Jefferson. March 15,
1942. C.M.B.
9. Rhynchostegium Bryologia Europaea
Autoicous: more or less robust to quite slender, mostly soft, cespitose, pale
green to dark green, rarely yellowish to golden-brown, more or less lustrous:
stem creeping, bearing rhizoids, sometimes stolon-like, irregularly to pinnately
branched; branches more or less thickly-leaved, often complanate; leaves spread-
ing, rarely imbricate, shortly or non-decurrent, mostly a little concave, non-
plicate, ovate to lance-ovate from a narrowed base, with a short or long point,
mostly serrulate, the margin basally reflexed; costa simple or rarely forked,
ending in about the middle of the leaf; median leaf-cells mostly narrowly
prosenchymatous, smooth, the basal shorter and wider, the alar not differen-
tiated, sometimes short-rectangular or quadrate; inner perichaetial leaves sheath-
ing, abruptly subulate and reflexed from the middle: seta more or less elon-
gate, smooth; capsule cemuous to horizontal, oval and weakly gibbous dorsally
to oblong or oblong-cylindric and almost symmetric, often constricted below
the mouth when dry and empty: annulus present; peristome as in Brachythe-
cium; lid long-rostrate from a convex-conic base; calyptra glabrous.
About 130 species, occurring on earth and stones, mostly in the temperate
and sub-tropic regions; about 10 species in North America; probably only the
following in our region:
1. Rhynchostegium serrulatum (Hedwig) Jaeger
(Hypnum serrulatum Hedwig; Eurynchium serrulatum Lindberg)
Plate LIX
Loosely matted, bright yellowish-green, when dry sub-lustrous: stems creep-
ing, sub-pinnately branched v/ith long and more or less 2-ranked branches;
branch-leaves complanate, 1.5-2 mm long, thin, concave, ovate-lanceolate, acu-
minate, serrulate from usually below the middle, thin-costate to the middle or
beyond, the apex often twisted, the margin plane and not bordered; perichaetial
leaves similar but more oblong; stem-leaves similar but relatively wider and
more cordate and with more distinct alar cells; median leaf-cells linear, prosen-
chymatous, about 8-10:1, at base somewhat broader and shorter, the alar not
differentiated: seta about 2.5 cm long, smooth, castaneous, sinistrorse when
dry; capsule light yellow to dark castaneous, oblong, cernuous, incurved, when
Jennings: Manual of Mosses — 34. Brachytheciaceae 307
dry contracted below the mouth; lid conic, slenderly rostrate, the beak long
and recurved; peristome-segments nearly as long as teeth, cilia usually 3, about
as long as segments, nodose to weakly appendiculate; basal membrane reaching
almost to middle of inner peristome; teeth narrowly lanceolate, yellowish-
brown, with distinct divisural, moderately trabeculate; annulus large; exothecial
cells rectangular to hexagonal, yellow-incrassate, or brownish; spores yellowish-
incrassate, finely papillose, about .009-.012 mm in diameter, mature in Sep-
tember and October.
In shaded woods on leaf -humus, old logs, etc., from New England to
the Gulf States and west to Kansas. It often hangs down in wide, thin mats
from overhanging ledges.
This is one o^ the most commonly collected mosses of our area, now represented in
the herbarium by specimens from more than si.xty localities from 16 counties. It probably
occurs in all.
308 American Midland Naturalist Monograph No. 6
Glossary of Bryological Terms Used in the Manual
Acaulescent , stemless.
Acrocarpous, with the fruit terminal on the stem or branch.
Acumen, a slenderly tapering apex, — acumination.
Acuminate, narrowly and slenderly tapering at the apex.
Acute, rather abruptly sharply pointed.
Alar, applied to the cells at the basal angles of the leaf.
Angular, applied to the alar group of cells.
Annulus, the ring of specialized cells often occurring between the rim of the capsule and
the op>erculum.
Antheridium, the male reproductive organ. See Introduction.
Apiculate, ending in a sharp and short point or apiculus.
Apophysis, the hypophysis or swelling of the seta just below the capsule.
Appendiculate, with reference to the cilia, with short transverse bars.
Arcbegonium, the more or less flask-shap>ed female organ.
Arcuate, bent like a bow.
Areolation, the cellular mesh or network of the leaf.
Aristale, awn-like or bristle-like.
Articulate, jointed, or with cross-bars.
Attenuate, long drawn out.
Auriculate, furnished with more or less ear-like lobes at the basal angles, applied to the
leaf.
Autoicous, having the archegonia and antheridia in separate clusters on the same plant.
Axillary, situated in the axil or upper angle of the insertion of a leaf.
Beak, the prolonged narrow apex of the operculum.
Bicostate, having a double costa or midrib.
Bifid, two-cleft.
Bifurcate, forked.
Bi-stratose, with two layers of cells.
Bi-striate, with two parallel lines or striae.
Calyptra, the thin and usually more or less membranous hood or cap on top of the
capsule.
Campanulate, bell-shaped.
Canaliculate, channeled.
Cancellate, (teeth) lattice-like.
Capsule, the spore-case or so-called "fruit" of a moss.
Carinate, keeled.
Caulescent, furnished with a stem.
Castaneous, chestnut-brown in color.
Central Strand, a central bundle of narrow and elongated cells found in some moss-stems.
Cernuous, somewhat drooping, nodding.
Cespitose, forming mats or tufts.
Cblorophyllose, containing chlorophyll or the green coloring matter of leaves.
Cilia, fine hair-like processes, usually applied to the hair-like structures often occurring
between the peristome-segments.
Ciranate, coiled inward from the apex.
Cirrate, curling up in drying.
Clavate, club-shaped.
Cleistocarpous, applied to a capsule which bursts open irregularly.
Collum, the more or less tafjering neck or base of the capsule.
Columella, the central axis of the capsule around which the spores are produced.
Comose, tufted at the apex, in a coma.
Complanate, flattened.
Jennings: Manual of Mosses 309
Confluent, merging together.
Constricted, contracted somewhere below the top or apex.
Cordate, heart-shaped.
Cortex, the outer baric or specialized layer.
Cortical, referring to the cortex.
Costa, the midrib or mid-vein of the leaf.
Crenate, with rounded teeth.
Cribrose, perforated more or less sieve-like.
Crispate, variously curled and bent.
Cuctillate, hood-like.
Cultrijorm, curved like a short, wide scimitar.
Cuneate, wedge-shaped.
Cuspidate, tipped with a sharp and rigid point.
Cuticular, belonging to the outermost skin.
Cygneous, abruptly down-curved like a swan's neck.
Cymbiform, the whole leaf more or less boat-shaped.
Decumbent, reclining but with the apex ascending.
Decurrent (leaves) with the borders extending down the stem below the insertion.
Dehiscent, splitting open.
Dendroid, tree-like in form.
Dentate, toothed with outwardly directed teeth.
Denticulate, minutely toothed.
Deoperculate, (capsule) with the lid fallen off.
Dextrorse, twisted to the right as the threads of the ordinary screw or bolt, used in the
opposite sense by some authors.
Dimidiate, split on one side.
Dimorphous, with two forms.
Diotcous, with the antheridia and archegonia on separate plants.
Discoid, disk-shaped as in some male inflorescences.
Distichous, in two opposite rows, two-ranked.
Divaricate, widely diverging or spreading.
Divisural (Line), the median line running up and down the teeth of the peristome and
often zigzag.
Ducts, applied to the narrow chlorophyllose cells in the leaves of the Sphagnums.
Ecostate, without a costa.
Emarginate, apically notched.
Emergent, applied to capsules rising slightly above the perichaetial leaves.
Exannulate. with no annulus.
Erose, irregularly notched.
Excavate, applied to leaf-insertions hollowed out in a more or less definite curve.
Excurrent, with the costa extending beyond the apex of the leaf.
Exothecial, the outer layer of cells of the capsule-wall.
Exserted, projecting beyond, as a capsule rising beyond the perichaetial leaves.
Falcate, scythe-shaped, flat, gradually tapering and curved.
Falcate-secund, falcate and turned to one side of the stem.
Fasciculate, m close and usually short clusters; usually applied to short, unequal, lateral,
bunched branches.
Fastigiate, with branches erect, near together, and more or less equal in height.
Fenestrate, furnished with openings.
Fibrtllose, applied to hyaline cells of Sphagnum in which the walls are lined with fine
fibrils or filaments.
Filiform, thread-like.
Fimbriate, fringed.
Flagelltjorm, lash-like or whip-like.
310 American Midland Naturalist Monograph No. 6
Flexuose, wavy or bending alternately back and forward.
Frondose, bearing fronds or frond-like.
Fugacious, falling away very early.
Fusiform, spindle-shaped.
Gametophyte, the sexual stage in the life-history of the moss and resulting from the germi-
nation of a spore. Usually begins with a filamentous protonema which eventually
gives rise to leafy stems, which finally bear the sexual organs (archegonia and
antheridia) and, upon the fertilization of the archegonium, there is produced the
other alternating phase, the sporophyte.
Gemmae, small more or less bud-like bodies capable of reproducing the plant.
Gemmiparous, producing gemmae.
Geniculate, bent like a knee.
Gibhous, swollen on one side.
Glabrous, with a smooth surface.
Glaucous, covered or whitened with a bloom.
Granulose, finely roughened as with grains of sand.
Gregarious, growing near together or in groups but not forming tufts or mats.
Gymnostomous, with the mouth of the capsule devoid of peristome.
Hamate, hooked.
Heteroicous, with two or more forms of inflorescence in the same cluster.
Hispid, beset with stiff hairs.
Hispidulous, minutely hispid.
Homomallous, (leaves) bent or curved to one side, all in the same direction.
Hyaline, transparent and colorless like water.
Hygroscopic, altering form or position with changes in moisture.
Hypophysis, an enlarged of the seta immediately below the capsule.
Imbricated, overlapped like the shingles on a roof.
Immersed, (capsule) concealed within the leaves of the perichaetium.
Incrassate, thickened, or thick-walled (cells).
Indehiscent, not splitting open.
Inflorescence, the clusters of reproductive organs, usually with enclosing bracts.
Innovation, a young offshoot from the stem.
Insertion, the point of attachment of the leaf to the stem or branch.
Involucre, a whorl of leaves or bracts around the flower.
Jutaceous, worm-like or catkin-like.
Laciniate, deeply slashed or cut into narrow lobes.
Lamellae, thin plates, particularly the flat plates on the dorsal surface of many peristome-
teeth; also on ventral surface of many leaves.
Lamina, the leaf-blade.
Lanceolate, lance-shaped.
Lid, the covering of the mouth of the capsule, the operculum.
Ligulate, strap>-shaped.
Linear, long and narrow with parallel sides.
Lingulate, tongue-shaped.
Lumen, the cavity of a cell.
Mamillate, tipped with a nipple-shaped projection.
Margin, (of a leaf) a bordenng band of peculiar shape or color.
Mitriform, mitre-shaped, or like a peaked cap, symmetric.
Monoicous, with the antheridia and archegonia on the same plant.
Mucronate, with the costa percurrent as a short small abrupt tip, tipped with a mucro.
Muricate, with the surface roughened with short, hard pwints.
Muticous, not pointed.
Jennings: Manual of Mosses 311
Neck, the collum.
Nodose, (cilia) with knots or swollen articulations.
Ob-, a prefix often used to convey the sense of inversion.
Obconic, inversely conic.
Obcordate, inversely cordate.
Obovdte, inversely ovate, narrowed towards the base.
Obsolete, scarcely apparent.
Operculum, the lid covering the mouth of the capsule.
Ovate, more or less egg-shaped, with the broader end downward.
Ovoid, more usually applied to a solid with an egg-like outline.
Panduriform, fiddle-shap>ed.
Papillae, minute nipple-shaped protuberances.
Papillose or Papillate, covered with papillae.
Paraphyllia, minute thin leaves or branched organs scattered imong the leaves.
Paraphyses, jointed and hvaline hair-like structures growing among reproductive organs.
Parenchymatous, compnased of broad cells joined end-to-end with square ends, not dove-
tailed.
Paroicous, having the antheridia and archegonia in the same cluster but not mixed, the
antheridia being in the axils of the perichaetial leaves below the archegonia.
Patent, spreading.
Pectinate, branched or divided like a comb.
Pedicel, the seta or stalk of the capsule.
Pedicellate, furnished with a pedicel.
Pellucid, translucent but scarcely hyaline.
Pendulous, drooping rather more than when cernuous, hanging down.
Percurrent, (costa) running through the whole length of the leaf.
Penchaetium, the involucre of bracts around the female flower and thus also around the
base of the seta or sessile capsule.
Perigonium, the whorl of bracts around the male or antheridial flower.
Peristome, the fringe of teeth, etc., at the mouth of the capsule.
Persistent, not easily nor early deciduous.
Pinnate, with the branches more or less equidistant and arranged on both sides of the
stem like a feather.
Piliferous, bearing a hair-like prolongation.
Plane, flat.
Pletirocarpous, with the flowi-rs more or less axillary and the fruit laterally borne.
Plicate, folded longitudinally.
Plumose, plume-like.
Pluriseriate, arranged in several or many series, as of leaves on the stem.
Polygamous, with the antheridia and archegonia variously disposed on the same plant.
Porose, pierced with small holes or pores.
Procumbent , trailing along on the ground.
Proliferous, bearing abnormal shoots, often from the flower cluster.
Prosenchymatous, composed of narrow cells whose ends dove-tail past each other, as op-
posed to the square-ended parenchymatous cells,
Protonema, the green filamentous phase of the gametophyte which is derived directly
from the germination of the spore, and sometimes persisting.
Pseudopodium, in Sphagnum the false seta bearing the capsule; in Aulacomnuim, etc., a
leafless seta-like branch bearing gemmae.
Punctate, marked with dots.
Pyriform, pear-shapied.
Quadrate, square.
Radicles, rootlets or rhizotds growing out from the base of the Stem.
312 American Midland Naturalist Monograph No. 6
Radiculose, covered with radicles.
Ramose, branching.
Ramulose, bearing smaller branchlets.
Repand, undulatelv or wavy-margined.
Reticulate, in the form of a net-work.
Retort Cells, cuticular cells of Sphagnum having an outward-curved apex.
Retuse, with the obtuse apex slightly indented.
Revolute, rolled backward from the matgin.
Revoluble, curling off, as does the annulus of many mosses.
Rhomboid, diamond-shaped.
Rostellate, short-beaked.
Rostrate, with a more or less long beak.
Rugose, wrinkled.
Rupestral, inhabiting rocks.
Scabrous, rough.
Scarious, thin, dry, membranous, but not green.
Secund, turned to one side.
Segments, the main divisions of the inner peristome.
Serrate, with forward-projecting teeth.
Serrulate, minutely serrate.
Sessile, not stalked.
Seta, the stalk or pedicel bearing the capsule,
Setaceous, bristle-like.
Sheathing, applied to pierichaetial leaves which wrap around the seta or ordinary leaves
wrapping around the stem.
Sinistrorse, twisted to the left, as is the case with the threads of the rather-rare "left-
handed" screw or bolt. By some authors used in the opposite sense.
Sinuose, wavy.
Spatulate, spatula-like, bluntly and narrowly obovate and quite attenuate downwards.
Spinulose, furnished with small spines.
Sporangium, usually synonymous with capsule.
Sporophyte, the spore-bearing generation of the moss arising from the fertilization of the
archegonium and known also as the sporogonium, — usually consisting of foot, seta,
and capsule.
Squarrose, spreading abruptly and widely.
Squarrulose, a lesser degree of squarrose.
Stegocarpous, with the capsule operculate.
Stipitate, mounted on a short stalk.
Stoloniferous, bearing slender, creeping and usually minutely-leaved secondary stems or
branches.
Slomata, breathing pores, or openings, in the epidermis.
Stomatose, bearing stomata.
Striate, marked with fine longitudinal lines or ridges.
Striolate, being very finely striate.
Strumose, furnished with a struma or unsymmetrical swelling at the base of the capsule,
goitre-like.
Sub-, as a prefix commonly used to denote the idea of somewhat or slightly.
Subulate, awl-like.
Sulcate, longitudinally grooved.
Synoicous, with the antheridia and archegonia mixed together in the same flower.
Terete, cylindrical or tapering.
Terrestrial, growing on earth.
Tessellate, checkered.
Tomentose, covered with soft matted hairs or tomentum.
Trabeculae, the more or less projecting plates or the inner side of the peristome-teeth.
Jennings: Manual of Mosses
313
Trabeculate, furnished with trabeculae.
Truncate, with the apical portion more or less squarely cut off.
Ttibtilose, tube-like.
Tumid, swollen, turgid.
Turgid, more or less rigidly swollen as from internal pressure, tumid.
U mbonate , with a slight projection in the center like the boss of a shield.
Uncinate, hook-shaped.
Unilateral, one-sided.
Unistratose, (cells) m one layer.
Urceolate, urn-like, contracted at or below the mouth.
Utricles, applied to the large hyalme cells of the leaves of Sphagnum.
V aginate, surrounded by a sheath.
Vaginule, a small sheath, the modified remams of the lower part of the archegonium sur-
rounding the base of the seta.
Ventral, the surface of the leaf facing the stem, as ordinarily situated.
Ventricose, bulgmg on one side.
Vermicular, worm-shapied.
Verrucose, minutely warty.
Verticil, a whorl.
Verticillate, whorled.
Vesiculose, more or less bladdery, like inflated air-spaces, vesicular.
Villous, covered with long, soft hairs.
Index of Plant Names
Acaulon 87
rufescens 88
triquetrum 88
Acrocarpi 44
Acrocladium 242
cuspidatum 242
Amphidium 101
lapponicum 102
Mcnjgeotii 102
Amphoridium _ 101
Amblystegiella 225
adnata 226
conferva 225
confervoides 225
minutissima 224
subtilis 225
Amblystegium 2 18
aduncum 236
chrysophyllum 249
cordifolium 241
eugyrium 244
exannulatus 239
fallax 232
fiiicinum 233
fluitans 239
fluviatile 231
hispidulum 248
var. spinifolium 232
Juratzkanum 221 [41]
Kneiffti 238
Kochii 222 [42]
Lescurii 235
minutissimum 224
noterophilum 234
ochraceum 246
orthocladon 221, 233, [41, 66]
palustre 244
radicale - 25 1
riparium 223 [42]
var. flaccidum 223 [42]
var. trichopodtum 222
serpens 219 [41]
stelLitum 250
subtile 225
trichopodium 222
varium 220 [41]
var. orthocladon 221
var. ovatum 220
American Tree Moss 181
Anacamptodon 280
splachnoides 281 [37]
Andreafacfae 43
* Numbers in brackets refer to plates. Synonyms are italicized.
314
American Midland Naturalist Monograph No. 6
Andreaea 43
petrophila 43
Rothii 43 [60]
var. crassinervia 43
rupestris 43
Anictangh'.m ciliatum 171
lapponicum 10'
Anodontium prorepens 103
Anoectangium Mougeottii 102
Anomodon 193
apiculatum 199
attenuatus 201 [38]
minor 200 [38]
obtusifolius 200
rostratus 201 [38]
repens 229
Rugelii 199 [38]
tristis 198
viticulosus 200
Aphanorhegma 1 1 5
patens 1 16
serratum 116 [20]
Archipiaceae 44
Archidium 44
ohioense 44
ArrherwpteTum heterostichum 143
Astomum 78
crispum 78
Mulilenbergianum 78 [62]
nitidulum 79
Stillivantii 78
AstTOphyllum ciliare 141
cuspidatum 141
hornum 137
marginatum 138
medium 140
orthorhynchum 137
TOftratum 139
Tugicum 142
Atrichum 159
angustatum ..162 [31]
var. plurilamellatum 164 [31]
crispum 160 [64]
Macmillani 163
papillosum 163 [31]
undulatum 161 [30]
var. allegheniense 162 [30]
var. minus 162 [31]
var. Selwyni 160
AULACOMNIACEAE 144
Aulacomnium 144
heterostichum 145 [28]
palustre 145 [28]
var. imbricatum 146
Barbula 85
acuminata 86
caespitosa 84
convoluta 87 [17]
fallax 86
papulosa 92
rubella 85
tortuosa 83
unguiculata 86 [17]
Bar TRAMIACEAE 147
Bartramia 148
fontana 1 52
grandi flora 148
ithyphylla 150
longisetd 151
Oederi 148
pomiformis 149 [28]
var. crispa 150
radicalis 151
Beaked Moss, Common 303
Brachytheciaceae 285
Brachythecium 289
acuminatum 288
acutum 293
campestre 291 [54]
cyrtophyllum 288
flagellare 297 [57]
var. homomallum 298 [57]
flexicaule 292 [55]
laetum 290
novae-angltae 305
oxycladon 290 [69]
plumosum 297
var. homomallum 298
populeum 297
reflexum 295
rivulare 294 [56]
rutabulum 293 [55]
salebrosum 291 [54]
Starkei 295 [56]
velutinum 296 [56]
Brotherella delicatula 268
recurvans 267
Bruchia 46
flexuosa 46
Sullivantii 47 [60]
Bryaceah 120
Bryhnia 304
graminicolor 305 [59]
novae-angliae 305 [69]
Bryoxiphium 68
norvegicum 69
Bryum 127
affine 130
angustirete 128 [22]
atropurpureum 133
argenteum 133 [24]
bicolor 133
bimum 132 [23]
caespiticium 131 [24]
Jennings: Manual of Mosses
315
capillare 134 [24]
cermium 128, 129
ctliare 141
cuspidatutn 130 [23]
cyclophyllum 129
intermedium 130
LescuTianum 125
obtttsifolium 129
Oederi 148
onlariense 135
pallescens 130 [23]
pendulum 128
pseudotriquetrum 132
pulchellum (Pohlia pulchella) 125
scoparium 61
tortifolium 129
uliginosum 129
undulatum 161
ventricosum 132
Bug-on-a-Stick Moss 157
BUABAUMIACEAE 155
Buxbaumia 156
aphylla 157 [30]
foliosa
.156
Calliergon 240
cordifoliLim 241 [44]
cuspid at um 242
eugyrium 244
giganteum 241
palustre 244
strammeum 242
trifarium 242
Calliergonella 242
cuspidata 242 [44]
Campylium 247
chrysophyllum 249 [45]
hispidulum 248 [45]
polygamum 249 [46]
radicale 251 [42]
stellatum 250 [46]
Campylodontium hypnoides 281
Camptothecium 28^
nitens 287
Catharinaea 159
angustata 162
crispa 160
papulosa - 163
plurilamellata 1 64
undulata 16!
var. allegheniensis 162
var. minor 162
Ceratodon 52
purpureas 52 [10]
Chamberlainia 287
acuminata 288 [54]
cyrtophylla 288 [53]
C.hiysohypnum chrysoph .Hum 249
hisptdulum 248
polygamum 249
stelldtum 250
Cirriphyllum 298
Boscii 300 [57]
piliferum 299
Climaceae 180
Climacium 180
americanum 181 [35]
var. Kindbergii 182
dendroides 180
Kmdbergii 182 [35]
Conomitnum 75
Hallianum 76
julianum 76
Cord Moss 119
Cratoneuron filicinum 233
Ctenidium 251
mollusrum 251 [46]
Cyltndrothectum cladorhizans 192
compressus 191
repens 229
seductrix 193
Desmatodon 90
arenaceus 91
obtusifolius 91 [17]
ohioensis 91
plinthobius 93
Porteri 91
Dichelyma 178
capillaceum 179
pallescens 179
pallescens 179 [35]
DiCRANACEAE 45
Dicranella 54
crispa 54
Fitzgeraldi 55 [10]
heteromalla 55 [11]
var. oTthocarpa 55
rufescens 56 [11, 60]
Schreberi 5 t
varia 56 [11]
Dicranodontium 64
asperulum 65
denudatum 65 [13]
Millspaughii 65
virginicum 65
Dicranum 55
aciculare 99
albidum 6"
Bergeri 60
Bonjeanii 61
condensatum 6)
Drummondii 60
flagellare 62 [12]
fulvum 63 [13]
316
American Midland Naturalist Monograph No. 6
var. viride 63 [13]
glaucum 67
heteromaltum 55
interruptum 63
longifolium 64 [13]
majus 6U
Millspaughii 66
montanum -.61 [12]
osmundioides 74
polyselum 60
purpuritscenf 52
purpureum 52
rugosum 60 [61]
scoparium 61 [12]
undulatum 60
ytride 63 [13]
Didymodon 84
arenaceus 91
cernuum 129
cyUndricus 82
recurvirostris 85
longiroslris 65
rubellus 85
Diphyscium 155
foliosum 156 [29]
DiSCELIACEAE 112
Discelium 1 13
incaTtiatum 1 13
nudum 113 [63]
Ditrichum 49
lineare 50 [9]
pallidum 51 [10]
pus.llum 51 [10]
tortile 51
var. vaginans 50
vaginans 50
Drepanocladus 235
adimcus 237 [67]
var. Kneiffii 238
exannulatus 239 [44]
fluitans 239 [69]
intermedius _ 217
Kneiffii 238
Sendtneri 238
uncinatus 236
vernicosus 237
Drummondia 103
prorejiens 103 [18]
Elodium paliidosum = Helodium 214 [40]
var. elodioides 215
Encalyptaceae 92
Encalypta ciliata 93
contortd 93
streptocarpa 93 [62]
Entodontaceae 190
■ Entodon 191
brevisetus 191
cladorhizans 192 [36]
compressus 191 [71]
re pens 229
seductrix 193 [37]
var. minor 193
Ephemeraceae 113
Ephemerum 113
cohaerans 114
crassinervium 114
var. papillosum 114
pallidum 114
papillosum 114
serratum 114 [64]
var. angustatum 114
spinulosum 1 14
Eurhynchium 302
Boscii 300
gTaminicoloT 305
hians 301
piliftmm 299
pulchellum 303
var. praecox 304
var. robustum 303 [58]
rusciforme 301
serriilatum 306
strigosum 303
var. robustum 303
var. praecox 304
Eustichia norregica 69
Extinguisher Moss 93
Fabroleskea Austini 202
Fabroniaceae 278
Fabronia 279
caroitniana 279
ciliaris 280
octoblepharis 280
pusitla 280
Ravcnclii 279
Fern Moss, Common 213
Fern Mosses 209
Fissidentaceae 68
Fissidens 69
adiantoides 73
bryoides 70 [14]
var. incurvus 71
cristatus 73 [15]
decipiens 73
exiguus 72 [15]
Hallianus 76
hyalinus 70 [61]
incurvus 71
var. minutulus 71
minutulus 72 [14]
obtusifolius 71 [14]
osmundioides 74
sciuroides 185
Jennings: Manual of Mosses
317
subbasilaris 75 [16]
taxifolius 74 [15]
FONTINALACEAE 172
Fontinalis 173
albicans 171
antipyretica var. gigantea 174 [64]
biformis 175
capillacea 179
dalecarlica 175 [34]
Duriaei 17/
flaccida 177
gigantea 174 [64]
hypnoides 176
Juliana 76
Lescuni 178
var. gracilescens 175
nitida 177
novae-angliae 176 [34]
pennata 187
Sullivantii 175
Forsstroemia trichomitria 185
FUNARIACEAE 115
Funana 118
americana 1 19
flavicans 1 19
hygrometrica 119 [21]
Muhlenbergii 1 19
Georgiaceae 157
Georgia pellucida 158 [30]
Glyphomitnum ( Ptychomitrium ) 95
Grimmiaceae 94
Grimmia 95
ambigua 96
apocarpa 96 [18]
var. conferta 97
campestris 98
conferta 97
Doniana 96
laevigata 93
leucophaea 98
obtusa 96
Olneyi 96
pennsylvanicd 97
pilifera 97 [63]
Gymnostomum 80
aeruginosum 81
calcareum SO [16]
curviTOstrum 81
immersum 1 17
pennatum 120
prorepens 103
TecuTvirostTum 81
rupestre 81
tenue 80
truncatum 90
vtridulum 80
Hair-Cap Moss 166
Haplocladium 207
microphylium 209 [39]
virginianum 208 [39]
Hapiohymenium _ 197
triste 198 [38]
Hedwigiaceae 171
Hedwigia 171
albicans var. leucophaea 172
ciliata 171 [34]
f. leucophaea 172
Helodium 2 14
paludosum 214 [40]
var. helodioides 215
Heterophyllum Haldani 267
Homalia 188
complanata 188
gracilis 188
Jamesii 189
Homalotheciella subcapillata 286
Homomallium 225
adnatum 226 [43]
Hookeriaceae 215
Hookeria acutifolia 215 [67]
Hygroamblystegium 230
jallax var. spintfolium 232
filicinum 233
fluviatile 231 [43]
irriguum 232 [43]
var. spinifolium 232
noterophilum 234
orthocladon 233 [41, 66]
tenax 232
Hygrohypnum 243
Closteri 246
f. serrulatum 246
dilatatum 247
eugyrium 244
var. Mackayi 245 [45]
luridum 244
Mackayi 245
novaecaesareae 247 [53]
ochraceum 246 [45]
Hylocomium 255
brevirostre 257 [48}
parietinum 258
proliferum 255
rugosum 255
splendens 255 [47]
squarrosum 253
triqiietrum 253
umbratum 256 [48]
Hymenostylium recurvirostrum 81 [16]
Hypnaceae 216
Hypnum 258
abietinum 2 1 1
acuminatum 288
acutum 293
318
American Midland Naturalist Monograph No. 6
adnatum 226
aduncum 237
var. Kneiffii 238
albulum 273
allegheniense 190
aTcuatum 265
attenuatum 201
bergenense 25 1
BoscK. 300
brevirostre 257
campestre 291
carolinianum 283
chrysophyllum 249
clavellatum 103
Closteri 246
complanatum 188
compressum 25 1
conferra 225
conferyoides 225
cordifolium 241
compressum 233
crista-castrensis 259
cupressiforme 263, 264
cuTvifolium 265
cuspidatum 242
cylindrocarpum 269
debtle 220
decUvum 296
delicatulum 212
dendroides 180
demissum var. carolinianum 283
denticulatum 276, 277
deplanatum 27 1
depressum 272
dilatatum 247
elegans 27 1
eugyrium 244
var. Mackayi 245
exannulatum 239
fertile 262
ftagellare 297
flextle 242
fluttans 239
fluviatile 23 1
giganteum 241
gracile 209
gracilescens 204
graminicolor 305
Haldanianum 267
hians 301
hirtellum 195
hisprdulum 248
imponens 263
intermedium 237
interruptum 257
irriguum 232
var. spinifolium 234
julaceum 184
laxepatulum 268
laxifolium 223
Lescurii 235
Lindbergii 265
luridum 244
Mackayi 245
rnarylandicum 284
medium 204
micans 247, 273
minutissimum 224
minutulum 210
molluscum 25 1
Muehlenbeckii 275
muticum 258
nervosum 206
nitens 287
nitidulum 273
noterophilum 234
novae-angliae 305
ochraceum 246
orthocladon 233
oxycladon 290
pallescens 262
paltidosum 214
palustre 244
parietinum 258
Patientiae 265
piliferum 299
plumosum 297
populeum 297
polygamum 249
praecox 304
praelongum 301
pratense 266
protuberans 262
pulchellum 303
pulchrum 267
radicale 25 1
recognitum 212
recurvans 267
reflexum 295
reptilis 261
riparioides 301
riparium 223
rivulare 294
Roeseanum 276
Rugelii 199
riparium 223
rusci forme 301
rutabulum 293
salebrosum 291
Schreberi 258 [48]
scitum 207
sciuroides 185
Sendtneri 238
serpens 22 1
serrulatum 306
silvaticum 276
Jennings: Manual of Mosses
319
splendens 255
squarrosum 253
Starkei 295
stelbtiim 250
stTdmineum 242
strigosum 303
subtenue 295
Stillivantiae 276
taxijolium 74
tenax 232
tTtfarium 242
tTiqitetTum 253
turfaceiim 273
iimhTdtum 256
loiciriatiim 236
velutinum 296
vernicosum 237
Wahlenbergii 127
Isopterygium 269
deplanatum 271 [69]
elegans 271 [68]
geophilum 272 [68]
micans 273
Muellerianum 270
pretense 266
pulchellum 272
turfaceum 273 [51]
Isotheciiim elegans 271
Juniper Hair-Cap Moss 163
Knothole Moss 281
maTchica _ 15 1
micTOcarpd 284
nervosa 204
nervosa 206
obscura 204 [39, 67]
pallescens 262
poly ant ha 229
polycarpa 204
pulchella 272
recurvans 267
rostrata 201
striatella 275
subttlis 225
tristis 198
yaria 220
Leskeella 205
nervosa 205 [39]
Leucobryaceae 66
Leucobryum 66
albidum 67
glaucum 67 [14]
minus 67
Leucodon 183
brachypus 184 [35]
julaceus 184 [36]
sciuroides 185
Leucodontaceae 182
Limnobtum ochraceum 246
Lindbergia 202
Austini 202
brachypterd var. Austini 202
Log Moss 263
Luminous Moss 120
Leersia cilidtd 93
Idciniatd 93
Leprobryum 121
pyriforme 122 [21]
Leptodictyum 218
riparium 223
trichopodium 222
var. Kochii 22?
Leptodon trichomitrion 185 [36]
Leptotrichum pusillum 51
Vdgindns 50
Lescurede rigiduld 206
Leskeaceae 193
Leskea 203
dcumiridtd 288
ddndtd 284
apiculata 199
arenicola 203
alteniidta 201
Austini 202
cylindrtca 278
dendroides 180
denticulata 282
gracilescens 204
Meesiaceae 146
Meesia 146
longiseta 147
triquetra 147
tnstichd 147
Micromitrittm 1 13
Mniaceae 135
Mniobryum 126
dlbicdns - 1 27
Wahlenbergii 127 [22]
Mnium 136
affine ." 141
var. ciliare 141 [26]
var. rugicum 142 [26]
bimum 132
Cdpillare 134
cinclidioides 144 [27]
cuspidatum 140 [25]
Drummondii 139
fontanum 152
homum 137 [65]
lycopodioides 137
marginatum 138
medium 140 [25]
320
American Midland Naturalist Monograph No. 6
orthorhychum 137
purpureum 52
punctatum 143 [27]
var. datum 143 [27]
rostratum 139
serratum 138 [25]
spinulosum 138 [65]
stellare 142 [26]
Myoireila 196
Careyana 197
gracilis 197
Nanomitrium 1 13
Neckera 187
complanata 188
gracilis 188
Octodiceras 75
debile 76 [61]
Hallianum 76
Julianum 76
Oncophorus 58
Wahlenbergii 59 [12]
Oreoweisia 46
serrulata 46
Orthotrichum 103
Braunii 105
canadense 106
citrinum 106
clavellatum 103
CTtspum 109
cupulatum 104
var. Porteri 104
var. minus 105
elegans 104
Lescuni 105 [19]
ohioense 106 [19]
var. citrinum 105
obtusifolium 107 [63]
Porteri 104
pumilum 107
pusillum 104
sordidum 104
stellatum 105 [19]
strangulatum 104 [18]
Oxyrhynchium 300
hians 301 [58]
riparioides 301 [58]
Paraleucobryum longifolium 64
Peat Mosses 20
Phascum 83
acaulon 89
cohaerans 1 14
crassinervium 1 14
cuspidatum 89
Hookeri 1 18
patens 1 16
Tufescens 88
serratum 1 14
strangulatum 1 18
subulatum 49
Philonotis 150
calcarea f. occidentalis 153 [29]
fontana 152 [29]
var. falcata 153 [29]
longiseta 151
marchica 15 )
Muhlenbergii 151 [28]
Physcomitrella patens 116
Physcomitrium 1 17
immersum 117 [20]
turbinatum 118 [20]
Pincushion Moss 67
Plagiopus Oederi 148
Plagiothecium 274
denticulatum 277 [53]
deplanatum 271
elegans 27 1
geophilum 272
laetum 278
micans 273
Muehlenbeckii 275
Muellerianum 270
pseudo-latebricola 272
pulchellum 272
Roeseanum 276 [71]
Ruthei 273
striatellum 275 [51]
sylvaticum 276 [51]
turfaceum 273
Plates, Explanation of 313
Platygyrium 229
repens 229 [37]
Pleuridium 4)
acuminatum 49
alternifolium 49
palustre 49
Ravenelii 49
subulatum 49 [63]
Sullivantii 49
Pleurocarpi 172
Plume Moss 259
Pogonatum 16)
brachyphyllum 165
brevicaule 165
pennsylvanicum 165 [32]
temie 165
Pohlia 122
annotina 126
camea 123
cruda 123
elongata 123
nutans 124 [21]
var. triciliata - 124 [22]
Jennings: Manual of Mosses
321
proligera 126
pulchella 125 [22]
W ahlenhergii 127
POLVTRICHACEAE 159
Polytrichum 166
alpestre 169 [33]
angustatum 162
commune 170 [33]
var. perigoniale 171
var. uliginosum 170 [33]
gracile 166 [ll]
juniperinum 168 [33]
var. alpestre _ 169
ohioense 167 [32]
pensilranicum 165
piliferum 16S [32, 64]
strict um 169
Porotrichum 189
allegheniense 190 [36]
POTTIACEAE 76
Pottia 89
truncata 90 [17]
truncatula 90
Pterogonium repens 229
octoblepharis 280
PterygophyUum acuminatum 215
lucens 2 15
Pterygynandrum 273
apiculatum 301
filiforme 278
hirtellum 195
intricatum 227
julaceum 184
subcapitellatum 286
tnchomitrium 185
Ptilium crista-castrensis 259 [49]
Ptychomitrium mcurvum 95 [63]
Pylaisia 226
denticulata 228
intricata 228
intricata 227 [37]
polyantha 229
Schimperi 228
Selwynii 228 [37]
subdenticiilata 223
velutina 227
Pylaisiella intricata 228
velutina 227
Rauia 206
scita 207 [39]
Rhabdoweisia denticulata 57 [11]
var. americana 58
Rhacomitrium 98
aciculare 99 [18]
ericoides 100
fasciculate 99
heterostichum var. sudeticum 99
var. ramulosum 100
var. gracilescens 100 [52]
microcarpum 99
sudeticum 99
Rhaphidostegium 283
adnatum 284
carolinianum 283
cylindricarpum 269
delicatulum 268
marylandicum 284
microcarpum 284
novae-caesareae 247
recur vans 267
Rhodobryum 134
ontariense 135
roseum 135 [24]
Rhynchostegium 306
delicatulum 268
deplanatum 271
geophilum 272
Jamesii 262
novae-caesareae 247
rusciforme 301
serrulatum 306 [59]
Rhytidiadelphus 252
squarrosus 253
triquetrus 253 [47]
Rhytidium 254
rugosum 255 [66]
SCHISTOSTEGACEAE 120
Schistostega 120
osmundacea 120
pennata 120
Schwetschkeopsis 281
denticulata 282 [36]
Sciaromium 234
Lescuni 235 [43]
Seligena 53
calcarea 54
recurvata 53
setacea 53
Sematophvllaceae 283
Sematophylliim 283
adnatum 284
carolinianum 283 [53]
delicatulum 268
marylandicum 284 [52]
micans 273
recurvans 267
tenuirostre 269
Sphaerangium 87
muticum 88
triquetrum 88
Sphagnaceae 20
Sphagnales 20
Sphagnum 20
acutifolium 41
322
American Midland Naturalist Monograph No. 6
var. fuscum 39
var. gracile 38
var. quinquefarium 40
var. subnitens 40
van viride 42 [5]
affine 24 [1]
amblyphyllum var. parvifolium 31
angustifolium 31
apiculatum 30
atinculatum 33
auriculatum 34 [5]
Austini 22
var. glaucum 23
capillaceum 41
var. viride 42
compactum 28
var. squarrosum 28 [3}
contortum 32
var. gracile 36
var. laxum 35
cuspidatum 32 [4]
cymbifolium 25
var. compactum 26
var. papillosum 23
var. squarrosulum 26
var. virescens i. brachycladon 26
fimbriatum 37 [6}
fuscum 39 [8]
Girgensohnii 37 [7]
Gravetii 34
imbricatum 22 [1]
var. affine 24
var. sublaeve 23
intermedium 30
inundatum 35
var. auriculatum 35 [9}
isophyllum 33
laricinum 32
latifolium 25
var. brachycladum 26
var. squarrosulum 26
magellanicum 26 [3]
medium 26
palustre 25 [1]
var. brachycladum 26
var. squarrosulum 26
papillosum 23 [2]
parvifolium 3 1
platyphyllum 33 [5]
plumulosum 40 [5]
f. viride 41
pungens 36 [9]
quinquefarium 40 [6]
recurvum 30 [3}
var. amblyphyllum 31
var. parvifolium 3 1
var. tenue 31 [6]
rigidum 28
squarrosum 29 [70]
var. teres 29
subnitens 40
var. viride 41
subsecundum 34 [9]
var. contortum 32
var. intermedium 34
teres 29
var. subteres 29 [3]
virginianum 32
Warnstorfii 38
var. virescens 38 [6]
Wulfianum 27 [2]
Splachnaceae 110
Splachnum 110
ampullaceum 110 [52]
angustatum 1 1 1
Spoon-Leaved Moss 300
Sporledera palustris 49
Stereodon 260
adnatum 226
arcuatus 265
crista-castrensis 259
cupressiformis 264
var filiformis 264 [50]
curvifolius 265 [50]
delicatulus 268
fertilis 262 [49]
filicinus 233
Haldanianus 267 [50]
hispidulus 248
imponens 263 [49]
nemorosus 260
pallescens 262
Patientiae 265 [50]
pratensis 266 [70]
recurvans 267 [51]
reptilis 261 [49]
riparium 223
Schreberi 258
tenuirostris 269 [51]
turfaceus 273
varius 220
Systegium crispum 78
nit-dulum 79
Sullivant.i 78
Tetraphidaceae 157
Tetraphis 158
pellucida 158 [30]
Tetraplodon 1 1 1
angustatus Ill [72]
Thamnium 189
allegheniense 190
Thelia 195
asprella 196 [38]
hirtella 195 [38]
Lescurii 196
Jennings: Manual of Mosses
323
Thuidium 209
abietinum 2 1 1
Alleni 212
delicatidum 212
delicatulum 213 [40]
elodioides 2 1 5
gTJctle var. lancastriense 208
intermedium 213
Idxi folium 295
micTophyllum 209
miniitulum 210 [39]
paludosum 2 14
Philiberti 2 1 3
pygmaeum 211 [66]
recognitum 212 [40]
scitum 207
virginiiinum 208
Timmia 154
cucullata 154 [29]
megapolitana 154
var. cucullata 154
Tortella 83
fragilis _ 83
humilis 84 [17]
nitida 83
tortuosa 83
Tortula 91
caespitosa 84
muralis 92
papillosa 92 [62]
plinthobia 92
Porteri _ 92
ruralis 92
tortuosa 83
Tree Moss, American 181
Trematodon 47
ambiguus 47 [72]
longicollis 43
Trichostomum 82
cylindricum 82 [16]
ericoides 100
micTOcarpum 99
tentiirostre 82
yaginans 50
True Mosses 44
Uloca 107
americana 108 [19]
cnspa 109 [20]
var. minus 110
crispula 1 10
Hutchinsiae 108
Ludwigii 108 [19]
ulophylla 109
Water Mosses 173
Webera 155
Webera^ Pohlia 122
albicans 127
annotina 126
carnea 123
cruda 123
elongata 123
Lescuriana 125
nutans 124
var. triciliata 124
proligera 126
pulchella 125
pyriformis 1 22
sessiiis 156
Weisia 79
coarctata 108
crispa (see Astomum ) 78
currirostris 81
fugax 57
incarnata ....1 13
recuTvirostra 85
vindula 79 [16]
White Moss 67
Zygodon 101
lapponicus 102
Mougeotii 102
Plates
Explanation of the Method of Lettering the Figures
To facilitate reference and comparison of the various plates a series of
letters and figures has been used which applies uniformly to all plates.
a — Apex
an — Anniilus
B — Branch
b — Base
be — Basal median
c — Central or Median
ci — Cilia
CT — Cross-Section
d — Dorsal
g — Gemmae
Im — Lower Margin
m — Margin
o — Lid or Operculum
P — Plant, habit sketch of, or
of a plant
of
a portion
sp — Spores
S — Stem
s — Peristome-segments
t — Teeth
upm — Upper Margin
-.■ — Ventral
1 — Stem-Leaf
2 — Branch-leaf
3 — Perichaetial Leaf
A — Paraphyllia
5 — Seta
6 — Capsule
7 — Calyptra
8 — Peristome
The above letters and figures are used in various combinations. \a, for
instance, indicates the apex of a stem-leaf; \bc, the median basal portion of a
stem-leaf; lad, dorsal view of branch-leaf; 3upm, upper margin of perichaetial
leaf, etc.
324
Plate I
325
Plate II
Sphagnum papillosum
326
Plate III
Sphagnum recurvum g^ iSphagnum teres v/i». subteres
327
Plate IV
Sphagnum cuspioatum
328
Plate V
Sphagnum platyphyllum
Sphagnum auriculatum
329
Plate VI
S. RECURVUM VAR. TENUE
Sphagnum fimbriatum
mm
Sphagnum warnstorfii vap.virescens Sphagnum guinquefarium (M
330
Plate VII
Sphagnum Girgensohnii
331
Plate VIII
Sphagnum fuscum
3?2
Plate IX
Sphagnum pungens
333
Plate X
Ceratodon pupureus
DiCRANELLA FITZGERALDI (Sj^
334
Plate XI
DiCRANELLA VARIA
Rhabdoweisia DENTICULATA
335
Plate XII
DiCRANUM MONTANUM
336
Plate XIII
DiCRANUM LONGIFOLIUM
DlCRANODONTIUM DENUDATUM '"ff
337
Plate XIV
338
Plate XV
FiSSlDENS TAXIFOLIUS
FiSSIDENS CRISTATUS
339
Plate XVI
Gymnostomum calcareum
Trichostomum cylindricum
340
Plate XVII
Barbula convoluta
POTTIA TRUNCATA DESMATODON OBTUSIFOLIUS
341
Place XVIII
Ik ^'m
*
OOrP
IScPn
ONgaa
rBssd
Grimmia apocarpa
Rhacomitrium aciculare
Mm
Qrummondia prorepens
:.'tH
Orthotrichum strangulatum "^
/4.J.
342
Plate XIX
P Orthotrichum ohioense
1<: * ■'i'
Ulota ludwigii
343
Plate XX
i*;o It x'i
Ulota crispa
^aX*
icAZJ-
Aphanorhegma serratum
Physcomitrium immersum
Physcomitrium turbinatum
344
Plate XXI
M5
Plate XXII
MnIOBRYUM WAHLCNeCRGII
la
lh^76- Bryum angustirete
346
Plate XXIII
mm
Bryum cuspidatum Bryum pallescens
347
Plate XXIV
mm /##
a-
Bryum c/espiticium I 'f*^ Bryum argenteum ^
Bryum capillare
348
Plate XXV
H9
Plate XXVI
Mnium AFFINE var. rugicum
350
Plate XXVII
MnIUM CINCLIDIOIDES
351
Plate XXVIII
Bartramia pomiformis
Philonotis muhlenbergii
352
Plate XXIX
'■■MrK.
m
r
TiMMIA CUCULLATA
353
Plate XXX
lb '^"0
Atrichum undulatum
Atrichum undulatum VAR. allegheniense
354
Plate XXXI
Atrichum undulatum var. m;nug
Atrichum angustatum
88888 ,
Atrichum papillosum
Atrichum angustatum var. plurilamellatum
355
Plate XXXII
POLYTBICHUM PILIFERUM
POLYTRICHUM OHIOENSE CfU--
356
Plate XXXIII
357
Plate XXXI\'
FONTINALIS NOVAC-ANGLIAE
338
Plate XXXV
W^'i-IM^:^
Climacium kindbergii
359
Plate XXXVI
POROTRICHUM ALLEGHENI ENSE
Entodon CUADORHIZANS
360
Plate XXXVII
Pylaisia intricata
Pylaisia Selwynii Anacamptodon splachnoides
361
Plate XXXVIII
■^^'^tvv i: \' ■ - -■ i-.i'*-' I 'f^/ /(' ./-
Anomodon rugeli
Anomodon minor
2fc. -J
Anomodon rostratus
362
Plate XXXIX
Haplocladium
virginianum
Haplocladium
microphyllum
Thuidium
minutulum
?63
Plate XL
Helodium paludosum
364
Plate XLl
vcjvv 0'" I'l^fia'' ''^-^ H YGROAMBLYSTEGI''
ySp'' 4/ ;®v ORTHOCUADON
^ ■ ' / :.,^^:
Amblystegium serpens ~ ''' " ■""'
0..0 I ll
1 l\ <^^".■^^^■^k'^
.Jjj^
jj Amblystegium juratzkanum
2ft . '0
Amblystegium varium
i__iJ
365
Plate XLII
Amblystegium riparium
Amblystegium riparium var flaccidum
366
Plate XLIII
367
Plate XLIV
Calliergoneula cuspidata
368
Plate XLV
Campylium hispidulum
CAmpylium chrysophyllum
369
Plate XLVI
Campylium polygamum
\
Campylium stellatum
A
i,\'
Ctenioium molluscum
370
Plate XLVII
Hylocomium splendens
£
371
Plate XLVIII
372
Plate XLIX
Stereodon fertilis
Stereodon imponens
373
Plate L
Stereodon Patientiae
Stcreodon haldanian
us f
374
Plate LI
375
Plate LII
Sematophyllum marylandicum
■75
Rhacomitrium
heterostichum
var. gracilescens
376
Plate LIII
377
Plate LIV
378
Plate LV
Brachythecium flexicaule
Brachythecium rutabulum
379
Plate LVI
Brachythccium rivulare
Brachythecium velutinum
380
Plate LVII
CiRRIPHYLLUM BOSCH
381
Plate LVIII
EURHYNCHIUM PU LCHELLUM yx/r R08USTUM
382
Plate LIX
mmm
i Mil lU I' '•7/'"7 ,/u 7 'r
Rhynchostegium scrrulatum
383
Plate LX
PUEURIDIUM SUBULATUM
OlCRANELLA RUFESCENS ^
384
Plate LXI
s7S
DiCRANUM RUGOSUM
margin
OCTODICERAS DEBILE «'
385
Plate LXII
a/26 Jlnr, ^15
TORTULA PAPILLOSA
Encalypta STREPTOCARPA^
386
Plate LXIII
387
Plate LXIV
6 upm
POLYTRICHUM PILIFERUM
tern aSS
FONTINAUIS ANTIPYRETICA VAR. GIGANTEA
388
Plate LXV
389
Plate LXVI
390
Plate LXVII
Plate LXVIII
ISOPTERYGIUM GEOPHILUM
y.325
392
Plate LXIX
Brachythecium OXYCLADON
DRCPANOCLADUS FLUITANS
393
Place LXX
Stereodon pratensi
394
Plate LXXI
395
Plate LXXII
■ 230 \\^^mm^
Trematodon ambiguus I
Tetraplodon angustatus
396