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MOSSES OF THE GREAT LAKES FOREST

by

Howard Crum

University of Michigan

CONTRIBUTIONS FROM THE UNIVERSITY OF MICHIGAN HERBARIUM
Volume 10, pp. 1-404, 1004 figures

University Herbarium, University of Michigan
Ann Arbor, Michigan
1973

CONTRIBUTIONS FROM THE UNIVERSITY OF MICHIGAN HERBARIUM

Editorial Committee

Howard Crum
Rogers McVaugh
Robert L. Shaffer

Mosses of the Great Lakes Forest

Volume 10, pp. 1-404, 1004 figures in text
Price: $6.00

Library of Congress Number 73-620129

Volume 10 is complete in this issue. Early issues of the Contributions were pub-
lished as Numbers 1-8 from 1939 to 1942; volume 9 appeared in parts from 1966 to 1972.
The title page and cumulative index for Numbers 1-8 were issued in 1966. For informa-
tion, address the Director, Herbarium, University of Michigan, Ann Arbor, Michi-
gan 48104.

Volume 9, No. 8 was published 31 August 1972

TABLE OF CONTENTS

TMUtTOC WG COMA yee eee tee aye Pa Pes a a eee Sa aa tei eee Sie Seah 1
Key to the Genera of Michigan Mosses ...........2. 0000 eee enue 3
SVSUCMIALIC A CCOUI Gs tcc cares te tee ite a5. earn teen ent ok te ema eke | OR ca ee tea 17
SPEHAGNOPRS DAC Es Rts trac atetnt ts 220-8 <0. eine oe MON eat aeiee fear Sea teal eres teas! 17
ANDREAEORSIDA, 2.402) 8 6. he, sh ee ee Ron eg ia Oe seem te a 50

BR ORS TD Ae oie eg etat,. ie ect ear tender mtr Ae aaa a ton ane $1
PiSSiGentaCe ae acer eta Ske kee Ne a ee a eee het Se cea ty 51

Bry oxiphiaceaei tors te ee eae enna date yon ee te, ets mn es 3 62
Selipenlaceae: 2 tr. hone ete aks, agen Mee eee een eat Ugh 62

DIET CH ACC AC 03a teks earner SEs sc (Ste gic. We aes eat ae Re ane as Tae 65
Dicranaceae ssc oo ewe Sie GA ee a ee a 74
ESUCODIVACEAe Soe eee 2 eel ee a Me Ror ett cht 92

Encaly ptacede tearm rare ic- aCe wi ces meg ies epeiee eo tee Seer sectors 94

POLUIA CC AG sear ata ear hg. 502s dhs op ea ee ee eee nines gee eon acon 97

GTI IACEAG? iter eee a) OS, cl ete OPA enh oe eare gee ten te eih fo? coils) ate fs 119

Hed wista Cede mk cp 4x2, erie May etme ee nae ret Eas Ah yaa vine 129

Flitlatia Cede sesey: Se: entre. Sepioe yee ee OR ae oh, dre Nr, 2 mes ase co Cp cee Va, ieee 130

phe merace dete cc ee ere h2, ax eg ali Nee ee the, Goad hl Sfop se a et 4 139
Splachnaceae: (etc - aie seedy ee ns eee ete a Sp clea horas 140
Schistostevacede ee inut carne teri eter nee tier A yen mes hain: Eyre 144

BSLV. ACE AG oy ea ete eekly cues Bae Onn earner Mate en eet eh a eae ares ee ae 148

MM ntace ae aire rec, col ie a eh RD a coe aes WERE) oceans 165
STATIN ACE AE eee che vas re ete erage pe ree whee Ae as ets etc ein eek har et fakes ae ay 179

A TIACOMIMIACEAG san tcrc ete epee reo ele, Mere Maan! Pat een Re cae uae 2 181

MGeSIACE RCIA) cx eer teetes ai er ee Pen OUI errs ey fry eketaiorsiaoam Ge yay 185
Catoscopiaceac t: at. yen yn arta se SNe Ge eee ee rie et ies oe Baise GE 189
Bartranniace ees ris. lee Cee Ws kee rey cake ie ta Seam mee MOIiee Ba Geel yeaah seid 190
Orthotrichaceae ects. sete oer eee ee ere eg Be ihe, ee a 194

Cry placacede tse carter ret tai prt, ea ape iirrg Peecth onsale ex 208

MeN COG ONLACEAR stare at tae, cerca Varney es ns se ig cay iia eat 209
Bontinalaceae: vsecas othe tie ower meray a ait ae age a ae piste als 213
INECKEGTACC AG Sas sa poe ee it tine ne a veri en a Ser fo olata cote 218

Bab foniaceae a wane hi ea eae ere este Cae kviee Greene as 221

WeskeaCeae ist iets etait eas Os ah teat eM eras ter Sioa So, TE Reale 7, 224

PAMIDIY SEER 1A COAG eset os race tents ages Peet cs eee eg es eae e att lire Fete ae fe tae Sens 249

Brachy the Glace ae viveguc eae eco es ho ee oe err eas® Gee Ja 280
Entodontaceaet s522 2 cgi ee eee ee ee ee ond Gala at a, aed 304
Plagiotheclaceaems.<:c.n.a0-eane cr oor ot mee ects ea anea es 2 octaves: 306

Sematophy llaceae testi, es pee nee esis ee tas cites es aos Gs S 310

Fy DNA CE AC reas erica ae Nos geste eg eg Un, ei a os Uae 314
Climaciaceae nas, 622 alone seer ere em omeigt cam nies ee Sat ns Gara ta 335
Hylocomiaceacis 2 cy spies nahn hae etn ee signe ore eee as a age ge ae eee 339

BU x DaUinia Ce ae in ye ay. aso ee ec er ease er iis ates Aocoed Tes ec Se ts 345

Diy SClACE AC eo etic eee ks mee tree aa een eg oes ees al gees aah foro 348
Metraphidaceae:- > 2. fasta wane eee wegeee sRte ss Fens ioe Bea 350
Polytrichaceses «242.50 oss 5 eee ne ete petro th hee ee cena 351

A Dictionary of Terms Relating to Mosses ..... 2. - ee ee eee ee ee es 365
Bibliography, G00 5 tee G ee Ace tne ws Syed encore eee el cae ait 379
F160), nme ee Bee fi nN Sink rng te eon Ear. \'c, coh Ner SamenR ar mee en ae Ree oe 399

ns | ae a - ee a ey Ss
- _ => : : - a a : oo 7 7

INTRODUCTION

The mosses of the Douglas Lake region have been studied with care and
enthusiasm for half a century, and more. Getting to know them has given me many
happy summers. Because of my efforts and the efforts of others, the flora has been
very completely inventoried. It is not the richest flora in the country by any means,
but the mosses are a conspicuous and attractive feature of the vegetation and
surprisingly numerous and interesting—yet not so numerous as to plague the student
with unusual problems in variation and intergradation. It is to the student that this
book is directed. It is to my family—to Irene, Mary, and Roger—that it is dedicated.

The local area has had rough treatment from man and nature. Scraped clean by
glaciation and flooded by post-glacial lakes, the land became available to plants only
about 10,000 years ago, some of it about 2500 years ago. Habitat diversity is limited.
The land has only slight relief, and the soils are mostly calcareous sands or lowland
peat. Scarcely any rock is exposed in Emmet and Cheboygan Counties, but in nearby
Mackinac and Presque Isle Counties limestone outcrops in occasional dry pavements or
low cliffs. Glacial erratics present some variety in substrate, of slight importance.
Streams are few, owing to porous sands and limey bedrock. The winters are harsh and
the summers short, providing comfort only to the hardy or the indifferent.

The flora is representative of the entire Great Lakes Basin and St. Lawrence
River Valley. This is the area occupied by the Great Lakes Forest, which in aboriginal
time consisted of a mixed forest of beech and sugar maple (interspersed with yellow
birch and hemlock) on better upland soils, various pines (jack, red, and white) on
sandy uplands, and swamp forests (elm-ash-red maple; spruce-tamarack, and white
cedar-fir) in lowlands. This forest complex, lying in a tension zone between the
oak-hickory and northern coniferous forests, occupies the northern half of Michigan’s
Lower Peninsula and some of the Upper Peninsula. It is Michigan’s cool North Woods.
In the Douglas Lake region the lowlands have escaped disturbance until very recently.
Extensive road building has had dire effects on drainage and altered swamplands of all
kinds, and the Dutch Elm disease has drastically changed the character of hardwood
swamps within the past 30 years. The uplands have been subject to misuse for a
century or more, in the form of lumbering, burning, farming, road-building, and
cottage living, not to mention the recreational assaults of recent years. The uplands are
now, except for limited acreage in farmland, in a process of regeneration from a
pioneering aspen-red maple stage through a red oak-pine mixture to a closed forest
dominated by beech and sugar maple.

This is a land of lakes. The most beautiful habitats are the many peat bogs at the
edges of pot-hole !akes, eventually transforming them into black spruce muskeg, the
forest climax on acid peat. Our bogs, fed by ground water and therefore minero-
trophic, are more properly considered rich or poor fens, depending on the stage of
succession toward drier and more acid conditions. Succession is often cyclic rather
than directional, and so the story-book progression from open water to muskeg is,
more often than not, only a theoretical possibility. Sphagnum creates an acid habitat
and controls the vegetation of bogs at every stage of development.

The Thuja swamps provide the richest habitat for mosses. Developed as the
climax forest on the calcareous soils wetted by a changing water table, they are
successional to sedgey fens, in stream valleys, at edges of lakes fed by streams, and
along the margins of beach pools at the sandy shores of the Great Lakes. A few species
of Sphagnum occur in open fens, and some characterize the woodland climax, but they
scarcely dominate or control any stage of succession. The beach pools harbor at their

margins an assemblage of rare northern disjuncts—the same group of species deposited
as fossils in the Cheboygan County Bryophyte Bed 13,000 years ago during the final
withdrawal of Pleistocene ice.

This book was written especially for students at the University of Michigan
Biological Station. It includes all the mosses from the area of the Straits of Mackinac,
arbitrarily delimited as Cheboygan, Emmet, Mackinac, and Presque Isle Counties
because of easy access from the station. The book is also tailored for use in a broader
area, in much of eastern North America, in fact. The introductory key includes all
Michigan genera, and the text gives incidental means of identifying all of Michigan’s
mosses and most of those of the entire Great Lakes-St. Lawrence River area. It
provides a personalized guide to the literature on the biology of mosses, with
references to the kind of thing that gives meaning to a field-oriented course in
taxonomy. The glossary is a complete dictionary of terms used in the literature to
describe mosses included here because good students always find use for other manuals
and specialized works on taxonomy. References are given to illustrations in manuals
likely to be available to students. Most of the illustrations in this book are my own. Some
were done by Michelle Smith, who also made the cover design. A few of the drawings
were adapted from pencil sketches made by Marilou Florian.

I e gratitude to the many people, teachers and students, who have left
collections in the herbaria of the University of Michigan, the Biological Station, and
elsewhere. Bryology at the Biological Station dates back to 1918 when George E.
Nichols visited there in search of a good source of Sphagnum for use in making
wartime surgical pads. He offered a course in bryology from 1920 until 1938. (An
announcement in The Bryologist that the course would be offered for the second time
in 1921 tells us that “despite unfavorable first impressions, Dr. Nichols found the
region bryologically quite rich.”) William C. Steere took over from 1939 to 1942 and
(after taking part in a Cinchona mission in South America) again in 1945 and 1946.
Margaret Fulford offered the course from 1947 to 1949 and 1951 to 1953. Paul M.
Patterson taught in 1950 and A. J. Sharp from 1954 to 1957 and 1959 to 1964. |
taught the course in 1958 (when I began this book) and from 1965 through 1972.
Dale H. Vitt replaced me during part of 1971 when I became ill.

I cannot thank every person who has helped me to know the mosses of the Great
Lakes Forest. I am, above all, grateful to William C. Steere, who directed me in
graduate study at the University of Michigan. He showed me far horizons and gently
pushed me toward them. His friendly concern has continued through the years. Lewis
E. Anderson has made many direct and indirect contributions to this study which is, in
a sense, a by-product of a larger flora which we are writing together. He spent a
summer with me at the Biological Station; then, as always, his light-hearted approach
to study made work fun. Wolfgang Maass also spent a summer at the station and
taught me much about Sphagnum. Harold Robinson, Norton G. Miller, Jerry A. Snider,
and Dale H. Vitt have been good friends and good companions in the field. Margaret
Fulford was my first teacher in bryology and first showed me the moss flora of
northern Michigan. A. J. Sharp found many rarities in the area and has been a constant
source of help and encouragement over many years. I got good ideas on resources in
the literature from Harvey A. Miller.

This work was done at the Herbarium of the University of Michigan, at the
Biological Station, and also at the National Museum of Canada.

I am indebted to the Rackham School of Graduate Studies, University of
Michigan, for two faculty research grants (NSF Institutional Grants 96 and 149) and
also, less directly, to the National Science Foundation for Grant GB-6095, which
provided funds for workshops in bryology at the Biological Station.

Publication has been made possible by a grant from the Donald Richards
Bryological Publication Fund and also from the Horace H. Rackham Publication Fund.

KEY TO THE GENERA OF MICHIGAN MOSSES

1. Leaf cells arranged in a network of narrow, green cells enclosing large hyaline cells
agnum, p. 17
1. Leaf cells not arranged in a network
. Costa covered by green lamellae
3. Leaves rigid wet or dry, broad and sheathing at base; costa broad, eae most of
the leaf above the shoulders, covered by 16-60 Bee calyptra densely hair

4. Capsules angled, cubic or prismatic, inclined, smooth Po ip trichum, ae 355
4. Capsules i angled, cylindric, ee erect, Snes and mammillose-roughened o
inclined, curved-asymmetric, and smooth Pogon ae P. 361
3A yee sane shriveled or otherwise eee when dry, not broadly sheathing at base
costa nar : lamellae 2-6, rarely 9; calyptra na

cells, doubly serrate, not ending in an awn; peristome of 32 short teeth See y
their tips to a membranous expansion of the tip of the columella Atrichum, p. 352
5. Plants small, scattered; leaves not iaeeaaee bordered, or serrate, ending in an awn;
peristome none (or rudimentary), the tip of the columella not expanded as
membrane Fe ee p. 119
2. Costa not lamellos
6. Leaves several ie rs thick, consisting mainly of costa (which is 1/3-2/3 or more the
widt i ae oa ee base and fills most of the subula), in section showing both hyaline
and g
le Plants aire leaves consisting almost entirely of costa, in section showing several
layers of hyaline cells enclosing a + median row of smaller, green cells; alar cells not

differentiated , Leucobryum, p. 92
7. Plants greenish or brownish; leaves consisting of a broad costa occupying 1/2-2/3 the
leaf base and most of the subula, in section showing hyaline cells on either side of a
median row of green cells (in our species also with green and hyaline cells on the
lower surface arranged in alternating longitudinal rows giving t es a striate
appearance under a lens); alar cells differentiated eee: p. 89
6. Leaves unistratose (or rarely + thickened at the apex or along the bn sta

narrower (or lacking), not showing hyaline and green cells in section
8. Plants dark (brown, reddish, or blackish), brittle when dry, growing on granitic rock;
capsules sessile, elevated at maturity on an extension of the stem, dehiscing by 4
longitudinal slits . Andreaea, p. 50
8. Plants generally not particularly aes or brittle, on various substrates; capsules, if
xserted, not elevated on an extension of the stem, operculate or indehiscent (not
s

10. Leaves appearing split at the base, consisting of 2 vaginant laminae which clasp the
stem and the base of the leaf above, with wing-like extensions apically and dorsally
Fissidens, p. 51
10. Leaves not differentiated into vaginant, apical, and dorsal laminae
11. Plants dimorphous (fertile plants lee ee crowded at the apex, the sterile
plants with leaves in 2 rows and c by broad decurrencies); leaves not
sheathing or subulate; plants of dae aie (especially caves), with a luminous
Schistostega, p. 144
11. Plants not dimorphous; leaves sheathing at base and narrowed to a linear subula
or awn; plants not growing in dark places, the protonema not luminous
12. Leaves contracted to a rough, linear subula; capsules exserted; peristome prese
Disti ae. p. 67
12. oe ending in oth awn Cee aaa longer toward the stem tips);
apsules eee rare) eens peristo Bryoxiphium, p. 62
9, eee inserted in more than 2 rows Riattiiy nea strongly complanate and
thus appearing 2-ranked
13. Leaves strikingly squarrose-recurved, in 5 distinct rows; cells mammillose on both
surfaces Paludella, p. 186
13. eee erect to wide-spreading, if + squarrose-recurved, not distinctly 5-ranked;
cells variously smooth, bulging, or papillose

5)

14. Peristome of 4 teeth (nearly always evident; sterile plants can be recognized by
cups in Tetraphis or protonematal flaps which are particularly obvious in
Tetradonti
a

m
15. Plants rather small, caespitose, nearly always on rotten stumps; sterile plants
ommon ndi a gemma cup; costa ie ee upper Pores

iso-
Tetraphis, p. 350
minute, scattered or Er eenous, on soil in ‘dark recesses, not gemmiferous;

t eX

sta weak, ending belo r even lacking; cells elongate; capsules
oblong-ovoid; operculum obliquely apiculat Tetradontium, p. 351
14. Peristome of more tha teeth, or none
16. Leaves fev all, evanescent (rare ); plants consisting ue
disproportionatey large,

ly seen :
strongly he ovoid-ventricose capsu . sto
pusdourit, 'p. 345
16. i ieee. persistent; an i qi mainly of sporophy
17. Leaves ligulate, rounded at

tip, with bulging cells in more than | hyer
eae se scarious, bristle- ee ae eae inclined, asymmetri
ovoid-ventrico Desens: '. 348
17. Leaves a cases ae as above
18.Stems erect, or sparsely

forked, generally in tufts; archegonia and
sporop ies fi (though sometimes appearing lateral because of innovat-

ing branche
19. Capsules ine operculate (rupturing wregularly); plants minute, ephemera
20. Leaf margins involute stomum, p. 102
20. Leaf margins not involute

21. Capsules emergent to shortly exserted, shortly pyriform because of a sto
Panes p. 78
c t a neck
22. Plants bulbiform (sometimes 3-angled); leaves erect-appressed, very bro

concave or concave-keeled ne p. 119

22. Plants not bulbiform or 3-angled; leaves erect to spreading, narrower
23. Leaves setaceous Pleuridium, p. 66
23. Leaves narrow to broad but scarcely setaceous

24. Plants from an abundant and spicuous protonema; leaves linear to
lanceolate, acute; margins plane; cells smooth or coarsely unipa
at the ends ie, p. 139
a scant, inconspicuous protonema; leaves broadly
Spt pa i to oblong-obovate, ending in a yellowish, smooth
. cae Dia

zascum, p. 112

oo p. 133
36, Exothecial cells firmer, evenly thickened or thickened at the co
27. Capsules dehiscing at the middle; exothecial cells Sa peal thick-
the corners he norrhegma, p. 133
OAs aes ms above the middle; exothecial cells at
shat ie immersum, p. 133
25, Plants ae to large (rarely minute), not ephemeral; capsules mostly
exs

28. aa on dung (or other organic matter of animal origin); capsules erect
and sterile

symmetric, with apoph on broader than the urn and
considerably eae in size, shape,

2
29. Apophysis much lar than the urn, ee lobose and wri a or
expanded and umbrel-tke, Sa ee colored: a
in pairs, chambered (as seen in sec

a chin, - 141
ae tiie aed som se ine than the urn, aa

ark-red; peristome teeth at first fused in 4’s, later 2’s,
28. Plants not on dung; capsules erect and symmetric or inclin

Te traplodon, p. 142
ed and
pe ae the neck sometimes well developed but not broader than the

30. Leaf cells. conspicuously bulging or mammillose (at least on the upper

31. Leaf cells mammillose on both surfaces; peristome single, with forke
teeth vertically pitted-striolate below D

d
chodontium, p. 90

31. Leaf cells bulging on the upper surface only; peristome none or double
the teeth not forked or vertically pitted-striolate
32. Leaves broadly lanceolate from a broader base, acute; capsules inclined
to pendulous; peristome double, the endostome consis
long-filiform cilia; calyptra often adhering to the tip of the seta after
falling from the mature capsule; brood bodies none Timmia, p. 180
32. Leaves See oblong-obovate, obtuse and apiculate; capsules erect;
peristome none; calyptra not adhering to the tip of the seta; brood
bodies often ee on are filaments Hyophila, p. 108
30. Leaf cells smooth or papillos
33. Leaf cells papillose & et longitudinally verruculose- striolate)
d

symmetric; peristome teet ed below the mouth, undivided;
in our species) longitudinally striolate icranowelsia, p. 92
35. Perichaetial leaves subulate; capsules suberect to inclined, some

to distinctly asymmetric; peristome teeth inserted at the mouth,
forked; upper cells coarsely and often irregularly papillose at ba
Dic

ck
cranum, in part, p. 79
34. Alar cells not noticeably differentiated

36. nee cells longitudinally striolate (Amphidium mougeotii, not yet
Bee Michigan, has + elongate papillae, rather similar to cuticular

Plagiopus, p. 191
36. wee cells papil

los
37. Stems repeatedly forked (and sometimes pleas +
in habit); sporophy tes often produced at inter

when dry; cells are capsules narrowly
urceolate and deeply furrowed when dry Amphidium pauses p. 208
s 2

pleurocarpous

38. sane mot much contorted when ft = with 1, some
i capsules ovoid to hemispheric, smo
39. Perch eres BRN ay ee at ae ena Lees
; leaves ecostate, nearly always ending in a broad
awn or at least a short tip; upper ails se 1 papilose
Hedwigia, p. 129
39. Ee cases leaves not ciliate; aes Bene poate ee
o hyaline-tipped o ned, the aw
a per and perichatit Hee sometimes + papillos
cere at back abov mia, in ee p. 121
37. Stems simple or sparsely Hee usually not savin sporophy tes
at interv

hyaline

Scalar

va
40. Plants in small, cushion-like tufts on trees and rocks; setae often
ee ene apsules often Tamers ee teeth 8 or
none or consisting ae 8 or lOc
a Gee cucullate, smooth, aa ie long-exserted; leaves
long-apiculate; brood re common in leaf axils or on ee
godon, p. 206
41. Calyptra mitrate, + plicate, often hairy; capsules im ee fe)
shortly exserted (rarely rather long-exserted); leaves Le short-
apiculate; brood bodies none or on lea
42. Leaves not or slightly ait when dry; basal cells oblong to
linear, moderately gaa not particu
obliquely arranged, the basal eae cells not much differenti-
fee capsules vanes immersed; stomata superficial
merse O
42. ie often crisped ie dry; basal cells linear, very
walled, yellow, arranged in rows fanning out from the costa;
ae ee nal cells short ae pale; capsules exserted; stomata
Ulo ta, p. 203
40. nee eee in more extensive tufts, nearly always or
rock; setae generally elongate and capsules distinctly ee
i th, if present, 16; endostome none or variously

e€

43. Leaves narrowed from a clasping base to a squarrose,
linear subula

43. Leaves not as above

rough,
Trichodon, p. 74

44. Leaves broad in proportion to their length, oblong, elliptic, or
base)

thin-walled with thickened, red-
c

S
walls; calyptra shorter than the capsule, cucullate, falling before
the operculum, peristome usually present, single, consisting of

peristome teeth long-filiform and closely wound together
above a high basal tube

ortula, p. 116
46. Cells of leat base less conspicuously differentiated; cells at
é wis

walls, forming a +
evident border; peristome (if sent) shorter, oblique
loosely spiraled above a low basal | hb ne
44. Leaves

ma
considerably longer than broad, lanceolate,
lanceolate or subulate (broadest near the bas €)

Weissia, p. 100
47. Leaf margins not eee

ending in a stalk bearing brood bodies a
its tip; 2 eed flexuose hae cells isodiametric, centrally

Au lacomnium, in part, p. 181

not ending in a gemmiferous stalk; costa

straight; cells isodiametric and indistinctly papillose over the

i bas

papillos
48. Sale —

u
; leaves lanceolate, vo Raia at
base; cells translucent, oie at 1 or bot S

ego p. 193
50. Plants of me aaa (especially on soil of banks
crevices of ape aves linear-subulate from a broad,
as sheathing _ base; obscure,
ends (see also Plagiopus which |
ruculose- Hae cells
49. Leaf cells no
narr

ngle

nas
) Bartramia, p. 191
t papillose at the ends; capsules mostly erect

ower and otherwise not as above; peristome none or
51. iS cells with 1 low, broad, often indistinct papilla over
each lumen
52. Capsules suberect to inclined, asymmetric, furrowed, som
times strumose; peristome teeth deeply — into
ee iat! pitted-striate vn eee p. 91
$2. Cap erect, symmetric, smooth, not oe
ane teeth split nearly to the base into 32 terete,
papillose divisions

53. Peristome consisting of filiform divisions Bile)
together

und
, in mers p. 108
33. aes consisting of slender but a filiform,

divisions idymodon, p. 106
SI. Leaf “cells pluripapillose

4. Leaf m margins ae geo on | or both sides

5 eaf

margins revolute side below; capsules
Si ne operculum remaining attached to
columella for a time after peat peristome no

ostomum recurviostrum, p. 99

55. Leaf margins revolute es “beth sides; oe
systylious; peristome none or well develo

56. Plants very small, scattered, ephemeral, on soll
disturbed places; leaves ending in a red apiculus or

ee awn; peristome none; calyptra finely papillos
oughout ottia, in pat, P. 112
56. ee small to fairly robust, + tufted, not ephem
eaves vue or hyaline-apiculate: Rene aah
calyptra smooth
57. Plants He reddish below; leaves entire; basal cells
firm-walled; peristome consisting of 32 long-filiform
divisions wound together in a spiral Barbuila, in cae p. 108
57. Plants prickicd below; leaves irregularly de ntate
he apex; basal cells lax and aa ikea peristome
ene of 32 shorter, erect o obliqu eee
Bebo HVT p. 105
54. Leaf margins plane or erect
8. Cells of leaf base large, lax, and hyaline, noticeably
extending beyond the shoulders as a V-shaped border
Tortella, p. 102
58. Cells of leaf base not extending up the margins above
the shoulders (or only slightly so
59. Leaves abruptly and irregularly toothed at ee of
hyaline basal cells and green upper cells ucladium, p. 100
59. mente entire or slightly and remotely ee
abov

60. Plants of moderate size; leaves concave, + tapered,
acute and apiculate; margins erect, + notched and

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scarcely tapered (with + parallel sides), bluntly acute;
margins Phe ree basal cells firm; peristome none
or greatly re
61. Annulus Soin peristome reduced to mere frag-
ments ‘'yroweisia, p. 99
61. Annulus and peristome non
'ymnostomum aeruginosum, p. 98
33. Leaf cells smooth
62. Plants in dark, green, brown, or blackish, often hoary tufts on rock;
leaves generally awned; cells often with sinuose-thickened walls
peristome single, consisting of 8 teeth (or rarely none) Grimmia, ’. 121
62. Plants = blackish or hoary, on various substrates; leaves not awned;
cell walls not SE ee peristome lacking, single, or double,
if present, the tee
63. ee minute, on limestone; peristome usually present, single, with
teeth undivide Seligeria, p. 63
63. Plants small to robust, on various substrates; peristome various, if
single, the teeth + divided
64. Capsules cylindric, erect or + curved from a long, narrow, ere
neck 1-2 times as long as the urn Tr eee p. 78
64.Capsules not long-necked
65. Leaves proportionately long (more than 5:1)

67. Capsules erect and symmetric, pyriform, smooth; peristome
teeth undivided or split only at the tips, smooth; plants on wet

rocks at edge of brooks Blindia, p. 65
67. Capsules mostly inclined, cylindric, + asym ric, somewhat

furrowed; peristome teeth forked, pitted- ne plants of dry

or moist habitats (not on wet rocks Dicranum, p. 79

66. Alar cells not particularly differentiated
68. Leaves + narrowed cea a broad, erect or sheathing base;
capsules strumose at bas

69. Leaves contorted wie dry; autoicous plants of woodland
habitats Oncophorus, p. 78
69. Leaves not or slightly contorted when dry; ee plants of +
wet, disturbed soil Dicraneila, in part, p. 74

68. Leaves more gradually tapered, not Sr ares eee ing at
base; capsules ee: distinctly strumo

70. Leaf cells long-rhomboidal to linear (ca. 6-12:1); capsules
Sa pyriform or oblong-pyriform; peristome doubl
71. Lea linear-subulate; costa filling about % the base
rae of the subula
71. Leaves oblong-lanceolate; costa narrow Pohlia, in part, p. 151
70. Leaf cells a ca ia erect or inclined, not pyriform;
peristome single o

72. Capsules ene porrent black, smooth, shiny; peristome
none; plants in very compact tufts in wet, cae ous fen
habitats ee = 189

72. Capsules not as above; peristome single; errr in ae
dense tufts, in moist or rarely wet habitats

73. Leaves ligulate, tounded-obtuse, peristome teeth not for

ked;
habdoweisia, p. 90
linear to ovate-lanceolate,
cleft or perforate; plants on soil (or
a variety of habitats in disturbed places
74. Peristome teeth flat, split % way down, pane vertically
pitted-striolate anella, in part, p. 74
74. Peristome teeth irregularly perforate or eee cleft into
terete, sometimes filiform divisions
75, Leaves narrowly lanceolate to subulate; le neither
revolute throughout nor toothed near the
Ditrichum, p. 71
75. Leaves lanceolate or ovate-lanceolate; margins revolute

; dioicous; penny

eed to the base but thickened and joined by cros
nections at the nodes below Ce ean p. 69
65. Leaves mre broad (less than 5:
77. Upper cells linear or long- arora (ca. 5-12:1)
78.Sporophytes from a short basal oy overtopped by a one
ae nov ee Pensome delicate and single ee of e
an i c elic hora, ha 165
78. oe terminal; peristome double, ee reeiee plan
of various substrates, not associated with c opper
Pohlia, in part, p. 151
77. Upper cells shorter, foes to pee Nap
79. Leaves bordered b row or linear cell
80. Leaf cells Hromboidal
81. ae connected by creeping, underground ‘ es er leave
n dense terminal rosettes adorn, p. 164
81. Some not connected by underground pear ee
conspicuous rosettes (though sometimes larger and more
ard the stem tips) eae p. 156
i agonal, rarely somewhat elonga

closing the capsule; costa bearing a few aa) pee
ventral lamellae

um crisp, 'p. 355
82. se nodding, broadly oblong- wine iene e double,
16 teeth, not attached to a central membrane; costa not

lose
83. Leaves entire or see ae and singly or doubly serrate;
endostome no longer than the exostome, consisting of free,
keeled and perforate pena and cilia from a high basal
membrane Mnium, p. 166

83. Leaves entire; endostome longer than the eee
consisting of a dome supported on narrow, flat seg
Cinclidium, p. 178
79. Leaves not bordered by narrow or linear cells
84. Leaf cells isodiametric
85. Leaves deeply concave, broadly oblong-ovate, unbordered,
strongly serrate; capsules curved and seria eat strongly ribbed
aco heterostichum, p. 182
85. Leaves not clea concave, ay ‘ellip tic to obovate,
if strongly serrate, istinctly bordered; capsules erect to
a

86.Plants on copper-bearing rock; leaves oblong-spatulate,
rounded at the apex; marginal cells somewhat thicker-walled
than the inner, often forming a distinct border
Scopelophila, p. 119
86. Plants not as abov
87. Plants very small: capsules erect, minute, obovoid, becom-
ing wide-mouthed when dry and e empty; peristome none

—_
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(or very rudimentary) Pottia truncata, p. 1

87. Plants larger; capsules erect to nodding, cylindric to
oblong-cylindric; peristome well developed

ed, entire and revolute at the

threads spirally twisted together above a basal tube
Tortula mucronifolia, p. 116
88. Leaves gered acute, plane and remotely and irregularly
ae d at

e argins; costa ending below the apex;
ome acu not twisted Mnium stellare, p. 167
84. Leaf real somewhat elongate and rectangular to short-
rhom
89. Plants iieeond silvery-green; upper cells not particularly
Bae short-rhomboidal, lax, hyaline; lower cells subquadrate,

Bryum argenteum, p. 159
89. Plants not julaceous, green or yellow; upper cells large and
es ice variously oblong, oblong-hexagonal or short-

al,

90. ees erect or strongly inclined from the base, the neck
broad and much shorter than the urn; peristome none,
single, or double (with the endostome reduce

91. Capsules erect and symmetric; peristome none
Physcomitrium, p. 131
91. Capsules inclined to horizontal, asymmetric; peristome
Funaria, p. 134

90. Capsules curved and inclined from a long, tapered, erect
neck; peristome double (the endostome delicate and irregu-

sharp point; cells lax, thin-walled; perichaetial leaves not
differentiated Amblyodon, p. 186

92. Plants coarse, more rigid; leaves erect or wide-spreading in
ows, not shrunken when dry, oblong-lanceolate to
ligulate, * rotadadobtee or acute; cells firm; perichaetial
ee elon ngate Meesia, p. 186
18.Stems creeping or ascending, freely branched, usually = interwoven mats;
archegonia and spor are lateral (or at the ends of branches)
93. Leaf cells with nodulose-wavy walls; peristome teeth ais nearly ie the base
into 2-3 filiform divisions Rhacomitrium, p. 126
93. Leaf cells not nodulose; peristome teeth (if present) split to i base into
ns

apillos
95. Leaf cells aeTioe at back because of projecting upper angles
96.Stem and branch leaves obviously differentiated in size and shape

97. Plants 1-pinnate; paraphyllia very few and polymorphous; upper leaf
cells hexagonal to short-rhombic Heterocladium, p. 230
97. Plants 2-3-pinnate in flat, layered fronds; paraphyllia felting stems and
branches, filiform-branched; upper cells linear-rhomboidal
Hylocomium splendens, p. 342
96. Stem and branch leaves not conspicuously differentiated
8. Branches smoothly julaceous wet or dry; leaves crowded, tightly
appressed, rounded-ovate, rounded and sometimes apiculate at the apex
Myurella bg a 229
98. Branches not particularly julaceous; leaves not notably crowd
appressed, + acute
99. Costa single, extending beyond the leaf middle
100. Leaves secund, 3-5 mm. long, both rugose and plicate Rhytidium, p. 345
100. Leaves not or ae secund, up to 1.2mm. long, not rugose,
sometimes + biplica
101. Plants on soil; eee (especially on stems) pein seattle
with sharply serrate, plane margins; costa not promin t back
ie not particularly opaque, oblong to inearhomboia pata-
Ilia non ryhnia, p. 292
101. Plants on rock; leaves decurrent, with entire or + serrulate, wee
revolute m ; costa prominent; cells opaque, odiametric to
eas paraphyllia present, few to moderately numero
Pecan ted bh p. 226
99. Costa double and usually very short, or none
102. Plants robust, coarse, erect; leaves wide-spreading, oe
sah extending 2 or more the leaf length; upper cells ir
inose at back Rhytidiadelphus a p. 340
102. ae smaller, creeping; leaves erect or secund, oblong-elliptic

103. Plants of medium size, pinnately branched in a triangular ies
leaves falcate-secund, abruptly acuminate from -

decurrent base tenidom, ». 332
103. Plants small and slender, sparsely ae a ae lea
oblong-elliptic, broadly acute, not decurrent Pte fer aes St p. 229

95. Leaf cells with 1 or more papillae over each ee
104. Costa very pe and double or none
105. Leaves ct-spreading, moderately concave, ovate-lanceolate,
a se pets all around; papillae small and indistinct
chwetschkeopsis, p. 223
10S. Leaves rounded or Pane deeply concave, abruptly apiculate
o short-acuminate; papillae dis Myurella abinee p. 229
104. Costa single, ending at or above de middle
106. Branches julaceous; leaves rounded, deeply concave; cells with a single,
illa

large, simple and curved or intricately branched pa helia, p. 236

106. Branches often terete but not julaceous: leaves ovate lanceolate,
moderately concave; cells with 1-several, not intricately branched
apillae

narrowed to a short, yellowish capone cells rather coarsel
ipapi reels p. 238
108. Plants in dense mats on rock or bases of trees; leaves not squar
when moist, gradually narrowed to a hyaline easel cells oT
papillose Anomodon ee p. 233
107. Leaves not ending in a hairpoint
109. Plants irregularly branched; Si a none or few and lanceolate
110. Leaf cells cere anak

111. Leaves usually broken off near ce middle Haplohymenium, p. 236
111. Leaves not br pices Anomodon, p. 232
110. Leaf nee unipapillose (the papillae often broad, blunt, and

indis
112, ae none or few; leaves broadly lanceolate or ovate,
obtuse or bluntly and often obliquely pointed; margins plane;

1]

cells isodiametric, rather eee papillose; capsules erect or +
curved, cylindric; alee on Les skea, p. 225

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upper ends (sometimes indistinctly papillose, or smooth); capsules
inclin a and asymmetric, oblong-cylindric, + tapered at base;
lan Pseudoleskea, P. 226
109. Plants ime branched, usually evenly so and frondose
paraphyllia numerous, in most cases felting stems and branches,
i h

Se;

113. Paraphyllia filiform-branched, matted on stems, less abundant on
branches, also attached to leaf bases; plants loosely 1-pinnate (some-
times unequally so); cells somewhat elongate; papillae single,
central or near the upper en Helodium, p. 247
113. Paraphyllia shorter, abundant but not matted or noticeably at-
ee to leaf bases; plants 1-3-pinnate, usually evenly so, some-
s frondose; papillae single and + central or
ee Plants evenly 1-3-pinnate, often frondose; apical cells of branch
leaves truncate and pluripapillose; cells uni- or pluripapillose
Thuidium, p. 240
114. tea loosely subpinnate, not frondose; apical cells of es ch
leaves not truncate, ending in a single papilla; cells unipapillo
Hap p. 239
94. Leaf cells smooth Sa mk Myurella julacea)
115. Paraphyllia abun
116. Plants dendroid, een by a creeping, underground “rhizome”
Climacium, p. 336
116. Plants not dendroid or rhizomatous
117. Plants of calcareous seeps; leaves somewhat to strongly falcate- eee
costa single, strong, ending well above the midleaf or even excu

paraphyllia very few to numerous, unbranched C euron, . 250
117. Plants of moist or wet woodlands, not in seepage; costa single

ouble, more slender ay ending far below the ‘bat apex; paraphyllia

abundant, filiform-branched ‘ylocomium, in part, p. 342

115. Paraphyllia few o ne
118. Plants submerged at flood time, otherwise stranded; leaves long, narrow
setaceous, strongly falcate-secund, keeled, 3-ranked Dichelyma, '». 216
118. Plants of varying habitats (some submerged or emergent, but not
normally stranded, except in some species of Fontinalis); leaves neither
keeled nor 3-ranked (or if so, not falcate-secun
119. Costa typically single (though + variable in Forsstroemia, Homali
Pseudoleskeella, and Hygrohypnum), extending to the leaf middle ar

eyon
120. Plants normally submerged
121.' Leaves + falcate- aenaTt stems and branches hooked at the t
DreerrecnaLG p. 263
121. Leaves not falcata: secund; stem and branch tips not hooked
122, Leaves wide-spreading and loosely complanate
Leptodictyum riparium, p. 256
122, Leaves erect or ST ee

123. Leaves lanceolate, acuminate; upper cells ae or
an owee ae, Heer: (ca. 2-6:1); costa sto sub
percurrent to excurrent Hygro Seana ae p. 258

123. ae eroatee broadly pointed or rounded he x; cells

at pe
thin- or firm-walled, long-rhomboidal; costa relatively strong but

ent
124. Leaves rounded-ovate, acute or obtuse, serrulate all around;
apical cells short and rhombic, noticeably shorter than the
upper median; alar cells scarcely differentiated
urhynchium he chaaete p. 298
124. Leaves cordate-ovate, broadly rounded at ee apex, entire
cells not differentiated; alar cells large, lax, hyalin
een in part, p. 271

120. Plants in dry to wet habitats, not normally submerged
125. Plants on tree trunks; stems long, creeping; branches crowded,
earing sporophytes at their tips; leaf cells see eine nh oe
green throughout; peristome single, of 16 very short, truncate teeth
Drummondia, p. 207
125. Plants on various substrates; stems variously creeping to -
ascending, with branches horiz 1 to ascending; sporophytes on
lateral buds, not at tips of - Sates branches; upper cells shortly
rhomboidal to linear; peristom

126. Plants of seepy, calcareous habitats fasually emergent from shallow
e

osta various; upper cells
sean ~~ cells scarcely differentiated to inflated in well-
arked groups
127. Branches stoutly and softly julaceous; leaves broad ae ve ed
, abruptly contracted to a sle

rip ne p. 294

e€
128. Leaves long- -lanceolate, slenderly acuminate, falcate-secund

stata P. 263
a ea ovate or lanceolate, not or rarely subsecund (o
ong-lanceolate and falcate secund, tomentose, with ee
: eaves
129. Upper cells short (up to 5:1)
130. Leaves serrulate above the middle; rou,

ee Slate p. 284
130. — entire or slightly denticulate ae below o the

131. Bees wide-spreading from the insertion
Leptodictyum, in part, p. 256
131. Leaves erect or appres

sed
132. ae pert bearing dense clusters of bud-like brood
branc eaf
133. ois ee filling the oo acumen and usually ex-
current; peristome teeth Leskeella, p. 226
133. Costa variable, usually ae Ae ending at or eee the
midleaf, sometimes longer, sometimes nearly lac
eristome teeth cross-striolate Ps Sie ae 'p. oad
7 Plants without brood branches
ce ge variable in length, sometimes double or nearly
ing
135. pee growing on trunks of trees, erect and loosely
subpinnate from a wiry, creeping primary stem; peri-
Sedo large, sheathing; capsules immersed to shortl

For sstroemia, p. 208
135. Plants on rock, irregularly branched; perichaetia not

Pse oe p. 227
134. Costa single, not particularly variabl
oe in knotholes and other protected crevices

capsules erect and symmetric, great

con-
ae below the mouth when dry; peristome te

eth
recurved when dry nacamptodon, p. 222

136. Plants not on _ trees; capsules horizontally inclined,
curved and asymmetric, less i

coarse, sometimes robust, growing on wet r
beside brooks

eae p. 258

137. Costa slender, ee at or below the leaf middle or
umen; plants small and slender,
swampy habitats Am Heer p. 261
129. Upper cells longer
138. Leaves conspicuously complanate (and appearing to be 2-
ranked), eerie elliptic, rounded and often apiculate at the
a tric omalia, p. 220
138. Leaves not or loosely complanate (not appearing 2-ranked),
lanceolate to cordate-ovate, rounded at the apex to acute or

cuminate
139. Leaves loosely and often indistinctly complanate, acute or
acuminate
140. Plants _ swampy habitats, particularly in places subject os
flooding; leaves entire, not twisted at the tips; costa no
ee as a tip at back Lep es p. 256
140. Plants of various woodland habitats; leaves serrulate to
serrate, often twisted at the tips; costa usually projecting as
a tooth at back
141. Leaves oblong-lanceolate; alar cells somewhat pan
entiated in small groups; seta oy opercu ae nic,
achy a orn p. 290
141. ae oblong-ovate; alar cells aneih tfereniated seta
mooth; operculum obliquely long-rostra
Rhyn eae p. 299
139. noe not at all complanate, rounded to acuminate a

142. aA cells considerably differentiated, larger and laxer or
conspicuously hyaline and inflated
143. Leaves elliptic to suborbicular, broadly rounded (and
sometimes apiculate) at the apex Calliergon, in ae p. 271
143. Leave ae o lanceolate, acuminate to a narrow

r blunt t
144. Ppt Pea branched and often + dendroid in dense,
tufted mats; stem leaves broadly decurrent with alar cells
relatively large; leaves sharply pointed
Brachythecium rivulare, p. 291
144, are phe branched, in loose masses; leaves with +
ted alar cells, sometimes auriculate but not ae
Hause: as ae leaves acuminate to a narrow, round-
ee a 267, 273
142. Alar cells not much differentiated
145. Leaves rounded at the apex
146. Costa relatively strong and single or double on oe same
lants; plants growing on wet rocks in or beside streams
Hygrohypnum, in ee p. 270
146. Costa generally single; plants of wet, sedgey, calcareous
ens Calliergon trifarium, p. 273
145. Leaves acute or acumin
147. Leaves broadly See to orbicular, concave and
usually cucullate at the apex; costa single or double,
often on the same plants, parte ee ly well developed
rohypnum, in spa p. 270
147. ee peel ai less Br ciatetlyy concave, not cucul-
late
148. ee sides saree to + squarrose-recurved o
d Campylium, in oe pacol
148. Leaves erect or erect-spreadin
149. Sone cee paaae in dense tufts, brown-tomentose
ith us wide-spreading branches below, un-
eee eee leaves conspicuously eg e
sedan p. 281
149, ears creeping or ascending, not i sean ely
and irregularly branched; leaves smooth or eee

150. Plants very slender, in wet habitats; leaves not
plicate, abruptly toothed near the base; ee bodies
often borne at back of costa near the leaf t
Rhynchosteiel, ag 300
150. Plants more robust, small to large, in wet or
dry habitats; leaves mostly + plicate, variously yee
or serrulate but not abruptly toothed at base; brood
bodies none
151. wae acute or obtuse; apical cells elie short-
than the median; operculum long- gee
E urhynohiun, P. 296
151. Leaves aoeaere ge cells similar to the media
operculum merely apiculate Br 7 ene 'p. 283
119. Costa short and ios or none
152. Plants often very long, trailing from a point of attachment, sub-
merged in flowing water (or rarely stranded except at flood time)
Fontinalis, p. 213
152, Plants not submerged (or otherwise as above)
153. Leaf cells less than 6:1
154. Leaf cells squarrose-recurved from a suberect, + clasping
Cam

mpylium ie p. 251
154. Leaves not squ

155. Leaves ee ‘cells thick-walled; capsules immersed; perichaetial
leaves long-sheathing; plants coarse, relatively robust Leuwcodon, p. 211

155. Leaves not plicate; cells thin-walled : peri
aan leaves not particularly elongate or sheathing; plants small

ute
156. Plants minute; leaves coarsely dentate; cells rhombic; exothecia
cells with wavy walls; tele single Fabronia, — 223
156. Plants minute or small; leaves entire or nearly so; cells rhombic
or shortly eee Sec cells straight-walled;
]

157. Leaves only 0.15-0.5 mm. long, + flat; alar cells siete
oe ciaale or subquadrate in small and inconspicuo
oups Paatne: Dol?
157. Leaves 0.5-0.8 m long, concave; alar cells subquadrate in
ngular nn Seer 1/3 up the margins Homomallium, p. 318
153. Leaf ee long
158. Plants ‘tyes ely producing dense, axillary clusters of bud-like
brood branches (especially noticeable at branch tips)
Platygyrium, p. 315
158. Plants without brood aoe
159. opi distinctly complan
160. Leaves conspicuously ae: plants on trunks of trees tetas
on roc
160. Leaves not undulate; plants on various substrates but ae
trunks of trees
161. Leaves very shiny, eas falcate and serrate at the tips;
alar — s inflated in a group of 34 across the a and 34
ar baie = 313
161. Las + shiny but not ota neither falcate nor serrate
the tips; alar cells not inflate
162. Leaves decurrent
162. Leaves not decurrent
163. Leaves broadly oblong-ovate, erect-spreading, entire;
cells loosely quadrate; capsules erect and symmetric;
peristome teeth smooth, papillose, or variously striolate;
dostome with a very low basal membrane, ies segments,
dn on, in part, ’D. 304
ate, wide-spreading, mostly + serrulate; alar
ells sca rc ay: differe ntiated; capsules mostly alin ed and
c; peristome perfect and “hypnaceous”
164. ee rigidly a ee and pectinate in ap-
earance; cortical cells large, thin-walled; leaves entire; alar

Ae pal

Plagiothecium, in part, p. 307

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cells scarcely differentiated; capsules + erect and symmetr
Isopte pasion, p. 3
164. Leaves not or + complanate, not rigid or pectinate; cortical
cells not much differentiated; leaves mostly + eae oe
eae een + differentiated; capsules inclined and asym-

tro
Ww

165. ee serrulate all around; apical cells grade than the
median yllum, p. 32
165. Leaves entire or serrulate only near the ape eee cells
not differentiated sopterygium, p. 322
159. Leaves not at all or only loosely and indistinctly complanate
166. Alar cells inflated
167. ents broadly rounded and usually apiculate at the apex; al
lis in large and conspicuous auricles allierg Se p. 277
167. Leave acute or acuminate; alar cells in + conspicuous decur-

w

168. Hence terete-foliate or loosely and irregularly complanate-
foliate (sometimes on the same plants); margins + revolute,
entire or sometimes serrulate at the extreme tip; alar cells
somewhat inflated in narrow decurrencies

lagiothecium cavifolium, p. 307

168. Branches not terete; leaves wide-spreading Or squarrose (and
occasionally somewhat comp aes margins plane, serrulate

in the upper % or more; alar cells lax and inflated in

conspicuous hyaline to red-yellow fone

Herzogiella striatella, p. 332
ed

166. Alar cells not inflated, though sometimes + enlarged or other-

wise differentiated
169. Leaves wide-spreading to squarrose, not at all complanate or
secund
170. Plants of open fens; leaves eee Laie when dry;
alar cells enlarged pylium stellatum, p. 2
170. Plants of se woodlands or open ie ie but not fens;
mae smooth or plicate, not striolate; alar cells not or
what differentiated tidiadelphus, in part, p. 340
169. eee not wide-spreading to squarrose, sometimes + com-
planate or secund
171. Leaves falcate-secund
172, Cortical cells of stems and branches large and hyaline
ae off with leaves on dissection); alar cells large, lax,

Nn
B=

173. Plants on wet rocks in or near streams; leaves not com-
but a or eee oe
th a % or mo
the leaf length, double or forked, ee sin oe
Hyg Pri eee p. 271
173. Plants oe various substrates in tee or swampy habitats
bu mally on rock in stream beds; leaves iver
Be pene acuminate to an acute point, the
acumen not twisted; costa short and double
num, in part, p. 324
172. Cortical cells not large and pees alar cells various
174. Plants robust and tumid or stoutly julaceous, floa or
mergent from the shallow ee of calcareous a eae
aes and + undulate when d
Scorpidium scorpioides, p. 278
174. ae on moderate size, more slenderly julaceous, i
t floating or emergent, not growing in fen habitats
175. ee are branched, on wet rocks in or near
brooks; leaves broad, ‘cucullate at te broad apex; costa
variable and single or double, often well developed
Hygro fet eT in part, p. 270
175. oe usually pinnately branched, in mois dland
bitats; leaves lanceolate or ovate and teas: not
setae at the apex; costa short and double or none

2

176. ety very regularly pinnate and frondose; ae strong-

plicate Ptilium, p. 334
176. oe irregularly to regularly pinnate; leaves sa rare ely
+ plicat Hypnum, p. 324
171. Leaves not falcate-secund (though sometimes + homomallous,
especially when
177. Plants stoutly julaceous, emergent from shallow water of
calcareous fens; leaves rounded and +
cucullate at the apex
177. Plants more slenderly julaceous, if at all, not emerge
not found in fens; leaves narrowly or broadly pointed,
sometimes + cucullate
178. a is abruptly inflated and usually yellow in a
cuo w of 34 cells at the insertion (rarely 2 rows);
ace cells with thickened corner aS ies p. 313
178. Alar and exothecial cells not as abov
179, Plants on wet rocks in or near streams; leaves broadly
ointed to rounded at the apex, concave and often
cucullate; costa single or double and ie in length but
usually well developed and '%4-% the leaf len:

con-

perohypnum, in part, Ba 270
179. Plants of drier habitats; leaves Pars to acuminate, scarce

y ae tinctly concave; costae very short and ao or

lac
180. Plants on tree trunks; leaves homomallous when dry;
ranches short, equal, hea er and conspicuously curved
in the same direc tion when dry; alar small and
eee p. 316
e trunks; _leaves not homomallous

the tip s serrulate in the upper
1/2- 2/3: alar cells scarcely ‘lifferentiated

Herzogiella turfacea, p. 332

181. ence broader, not at all to diene complanate
ins entire or somewhat serrulate near the

ae cells differentiated
182. Plants + erect, subp

innate; stems orange; branc
julaceous, not at all flattened; alar cells shortly ve
with thick, orange walls Pleu ne ‘aN 301
182. sae oe irregularly branched; branches + tere
re us and slightly flattened: alar s sub-
ee erect and symmetric, cylindric; peristome
double but not hypnaceous, the teeth not cross-
striolate); branches julaceous, blunt; leaves + asym-
metric, elliptic, abruptly acute to apiculate
ntodon, in part, P. 304
183. eee suberect, + curved and asymmetric, oblong
ric; peristome hypnaceous, the teet OSs-
ht Sue oe oming broken); ie terete,
eaves symmetric, lanceolate,

radually
ae ee a ee p. 310

SYSTEMATIC ACCOUNT
Class SPHAGNOPSIDA
A small class consisting of a single genus.
Sphagnum L.

Robust mosses developing from a thallose protonema, growing in wet habitats
(usually in peat bogs), often forming large, deep hummocks because of continued
apical growth; rhizoids lacking after the initial growth stages. Stems erect, sparsely
forked, consisting of a central, thin-walled parenchyma enveloped by a pigmented
wood cylinder of thick-walled prosenchyma which is usually surrounded by | or more
layers of hyaline, thin-walled, parenchymatous cortical cells. Branches usually in
fascicles spirally disposed around the stem, crowded at the stem tips in a head-like tuft
(the capitulum). Branch leaves spirally arranged, composed of linear, green cells in a
network enclosing large, empty, rhomboidal, hyaline cells nearly always with large,
rounded or elliptic pores (often bordered by a fibril ring) on both surfaces and nearly
always reinforced within by annular or spiral fibrils. Stem leaves less crowded than
branch leaves and usually clearly differentiated in size and shape, with a lesser
development of pores and fibrils, often showing extensive resorption of the walls of
the hyaline cells. Dioicous or monoicous; antheridia and archegonia always on separate
branches; paraphyses none; antheridial branches single or in fascicles of 2-3, scarcely
differentiated except that the leaves associated with antheridia may be highly colored;
antheridia globose or oval, long-stalked, each borne singly at the side of a leaf;
archegonial branches single or rarely paired, bearing 1-5 archegonia at the apex;
perichaetial leaves much larger and otherwise conspicuously differentiated, enclosing
the sporophyte until maturity. Sporophytes consisting of capsule and foot, elevated
beyond the perichaetial leaves at maturity by the elongation of the tip of the fruiting
branch as a “‘pseudopodium” (derived from the receptacle); capsules globose (but
urceolate to oblong-cylindric when dry and empty), dark-brown or black, operculate,
without annulus or peristome, usually with numerous pseudostomata in the lower half
or nearly throughout; endothecium giving rise only to the columella which is
overarched by the spore cavity. Spores tetrahedral, relatively large (18-42 u), explo-
sively discharged from the capsule. Calyptra a delicate, hyaline membrane investing’ the
capsule until maturity and finally irregularly ruptured. The name comes from a Greek
designation for some plant now indeterminable. Sphagnum is commonly known as
PEAT MOSS, less often as BOG MOSS.

The genus has a reputation for being very difficult and has been shunned even by
professional bryologists seemingly in agreement with W. J. Hooker, who said (in
Smith’s English Flora in 1844), “Linnaeus was perhaps correct in supposing that there
was but one species of Sphagnum.... It is very certain that the limits of the species, if
such they may be called, cannot be defined.” Our species are not that difficult, by any
means. If one recognizes the major groups, as outlined below, each species can be
recognized in turn by a few key characters, both macroscopic and microscopic. With
some practice, most of our species can be recognized even in the field. The structure
and taxonomy of the genus is admirably explained by Blomquist (Jour. Elisha Mitchell
Sci. Soc. 54: 1-21. 1938).

T enus may be divided into two easily recognized subgenera: The
INOPHLOEA include those species with robust, turgid branches and broad, concave,

17

18

obtuse, and cucullate branch leaves which are roughened at back near the apex because
of extensive “resorption” of the walls of hyaline cells and bordered by a row of cells
which appear to be linear because of resorption along their outer margins (looking like
a furrow in cross-section); the cortical cells of stems and branches are reinforced on
the inside by spiral fibrils. (Our species are S. magellanicum, S. centrale, S. papillosum,
and S. palustre.) The LITOPHLOEA are a larger and more diverse group. They have
more slender branches with narrower, mostly pointed leaves which are not cucullate
and not roughened at the back of the apex; the border of the branch leaves consists of
linear cells in two or more rows (or exceptionally of a single row of cells resorbed at
their margins); the cortical cells of stems and branches are not fibrillose. This is an
artificial grouping because of the intermediate nature of the S. compactum group,
which has branch leaves with broad, though truncate tips and a border formed by
resorption, although the cortical cells are not fibrillose. For this reason it is useful to
divide our species into smaller, more homogeneous sections: PALUSTRIA or, more
correctly, SPHAGNUM (S. magellanicum, S. centrale, S. papillosum, and S. palustre).
RIGIDA (S. compactum). POLYCLADA (S. wulfianum). SQUARROSA (S. squarrosum
and S. teres). CUSPIDATA (S. riparium, S. recurvum, sensu lato, S. pulchrum, S.
cuspidatum, S. majus, and S. annulatum var. porosum). SUBSECUNDA (S. sub-
secundum, sensu lato). ACUTIFOLIA (S. fimbriatum, S. girgensohnii, S. russowii, S.
warnstorfil, S. fuscum, §. capillaceum, and S. subnitens).—Modifications of this
subdivision appear, for example, in Andrews (Bryol. 14: 72-75. 1911; North American
Flora 15(1): 1-31. 1913; Ann. Bryol. 9: 3-12. 1937) and Isoviita (Ann. Bot. Fenn. 3:
199-264. 1966). Because of nomenclatural problems I have avoided assigning names of
ranks for the subdivisions inserted as conveniences in the text.

The importance of the genus in bog formation is well known, but little use has
been made of the distinctive spores of many species in palynological studies. Reference
may be made to papers on the morphology of spores by Terasmae (Bryol. 58:
306-311. 1955) and Tallis (Trans. Brit. Bryol. Soc. 4: 209-213. 1962). The latter
includes a key which is also used in Savicz-L. & Smirnova’s Handbook of Sphagnaceae
of the USSR, pp. 105-109 (in Russian).

Krutsch (Atlas der mittel- und jungtertiaren dispersen Sporen ... 1963) was able
to distinguish fossil spores of Sphagnum and microspores of Selaginella and also to
assign the Sphagnum spores to taxonomic sections.

Phagnum spores are discharged by an air gun mechanism resulting from an
internal pressure of 4-6 atmospheres—a pressure similar to that carried in the huge tires
of heavy trucks! It is small wonder that the spores are emitted in audible puffs, like
the snapping of a wheat straw, and hurled a distance of about three centimeters (and
no doubt more). Spores discharged by such an explosive means are more likely to be
stopped in an ecologically suitable place close at hand rather than face the uncer-
tainties of long distance dispersal. Another advantage may be a greater likelihood that
both sexes will be established close at hand.

Sphagnum is exceedingly important in nature. It controls and impedes drainage
of the vast North. Its effect on permafrost and maneuverability in the arctic is
immense. It controls some kinds of hydrarch successions, not only by its remarkable
ability to soak up water but also by creating an acid habitat for itself and other plants
active in the development of bogs. This is done by an exchange of hydrogen ions for
other cations (especially Catt and Mgtt, less readily Kt and Nat) so that the species
can flourish even in proximity to highly basic waters. The first invaders are usually
sedges, but the acidity which gives direction to bog succession is provided by
Sphagnum, with different species in control at each stage of succession toward drier
and more acid conditions. Peat mosses are of basic importance in the mountains in
controlling erosion of slopes and flooding of valleys. They are important in the seeding
of trees and other plants and in the developmental succession of forests.

19

Carex and other bog plants are usually more important than Sphagnum in
providing the bulk of peat cut for fuel. (Irish immigrants to North America were said
to bring with them some of the old sod of home. The Emerald Isle, of course, provides
just the right ‘“‘Atlantic” type of climate for the abundant growth of Sphagnum and
the formation of peat. The fragrance of peat fires is said to be one of Ireland’s
attractive features.) Sphagnum steeped in tar was used in the Scottish Highlands for
caulking boats, and in northern Europe Sphagnum moss was stuffed between timbers
of houses to deaden sound. Sphagnum also has importance in potting, mulching, and
packing. During the First World War dried Sphagnum was used extensively for surgical
dressings, thus freeing cotton for the manufacture of gun powder; see Hotson (Science,

S. 48: 203-208. 1918; Puget Sound Biol. Sta. Bull. 2: 211-247. 1919; Bryol. 24:
74-78, 89-96. 1921), Nichols (Bryol. 21: 53-56, 81-83. 1918 and Publ. Smithsonian
Inst. 2558: 221-234. 1920), and Porter (Intern. Jour. Surgery 30: 129-135. 1917).
Other uses, direct or indirect, have included bedding, food and litter for livestock,
medicinal baths, and the production of gunpowder, fireworks, paint, paper, insulation,
fabrics, charcoal, nitrates, ammonium compounds, sugar, alcohol, and dyes; see Davis
(U.S. Dept. of Interior Bureau of Mines Bull. 16: 1-214. 1911), Thieret (Econ. Bot.
10: 75-91. 1956), and Welch (Proc. Indiana Acad. Sci. 58: 3146. 1948)

It is said that Eskimos use Sphagnum in lieu of socks and diapers. Stark, in A
Popular History of British Mosses, quaintly records information on uses of Sphagnum
in Lapland (taken from Linnaeus’ Flora of Lapland): “The Lapland matrons are well
acquainted with this moss. They dry and lay it in their children’s cradles, to supply the
place of bed, bolster, and every covering; and being changed night and morning, it
keeps the infant remarkably clean, dry, and warm. It is sufficiently soft of itself,
but the tender mother, not satisfied with this, frequently covers the moss with the
downy hairs of the reindeer, and by that means makes a most delicate nest for the
new-born babe

Even recently the Chippewa Indians of Lac Vieux Desert in Michigan’s Upper
Peninsula put dry moss or pulverized rotten wood between the thighs of babies to
keep them clean.

Good accounts of bog ecology are presented by Transeau (Bot. Gaz. 40
(July-Dec.): 351-375, 418-448. 1905; 41 (Jan.-June): 17-42. 1906), Rigg (Bot. Rev. 6:
666-693. 1940; 17: 109-131. 1951), Gates (Ecol. Monogr. 12: 213-254. 1942), Potzger
(Bogs of the Quetico-Superior Country... 1950), Dansereau & Segadas-Vianna
(Canad. Jour Bot. 30: 490-520. 1952), Heinselman (Ecol. Monogr. 33: 327-374. 1953),
Newbould (New Biol. 26: 88-105. 1958), Ketchledge (N.Y. State Conserv. June-July
1964, 4 pp.). Other papers of worth include several listed in the bibliography, by
Conway, Lange, Osvald, Rose, and Sjors.

Useful keys have been published by Duncan, Fearnsides, Gams, Nyholm, and
Proctor. Andrews’ monograph in the North American Flora (1913) is the best
reference for North America as a whole and useful for the entire Northern Hemisphere.

1. Branches stout and tumid; branch leaves very broad, broadly pointed or porcane pe
cucullate-concave, not bordered by linear cells but with a marginal resorption furr
2. Cortical cells of stems and Reapers without spiral fibrils; leaves broadly eared at the
apex, concave but not cucullat 5.S. compactum
2. Cortical cells fibrillose; leaves eee and cucullate at the apex
3 cells of branch leaves triangular in section, with the base exposed on the inner
surface 4. S. palustre
3. Green cells of branch leaves oval to elliptic in section, equally eee or included
4.Green cells truncately elliptic or trapezoidal, exposed on both surfaces or more
broadly on the inner surface; walls of hyaline cells usually pense on the inside
where adjacent to green cells 3. S. papillosum
4. Green cells central; walls of hyaline cells smooth

5. Plants generally reddish; green ee ied elliptic, central and entirely included;
hyaline cells not or very slightly c

. S. magellanicum
5. Plants whitish-green; green cells aaa long-elliptic or lenticular, central but with
thickened end wa which reach both surfaces; hyaline cells slightly convex,
especially on the outer surfac 2. S. centrale
1, Branches more slender; branch leaves narrower, acuminate, concave or involute but n
cucullate at the apex, bordered by linear cells, without a resorption fu
6. Stems hard and brittle; branches crowded,
rounded . 8. wulfianum
6.Stems more flexible; branches not particularly crowded, in fascicles of 6 or less:
nde

in fascicles of 6-12; capitulum large an

n of perforations, the apex of the leaf not or only
c

ells of branch leaves typically very finely
lose on inner walls adjacent to green cells
8. Plants ee stem cortex in 2-3 layers; branch leaves ovate-hastate, squarrose from
broad, base . S. squarrosum
8. Plants ae stem cortex in 3-4 layers; branch leaves ovate-lanceolate, aybricate or
preading, especially when dr

e
d the leaf apex nearly
the apex nearly entire or rent dow
apex conspicuously lacerate-fringed); apical bud mostly small,
prominent; hyaline cells of branch leaves not papillose

Ty 8. S. teres
eaves es ica resorbed (usually not much resorbed on the outer

but some large and

Stem leaves with a single, deep, conspicuous rent or perforated area at the middle

the apex .S. riparium
9. Stem leaves entire or lacerate but not deeply rent or perforate at the middle of tl

apex

10. Stem leaves entire or nearly so at the apex; green cells of branch leaves exposed only
or more broa

dly on the outer surface or, if + equally exposed, the pigment brown
11. Green cells of branch leaves triangular in .

12. Branch leaves broadly ovate, abru narrowed to a broad, channeled, cuspidate
apex; green cells broadly triangular in eae with the apex of the triangle

aes)
, gradually narrowed to a slightly channeled o
nearly a e€ apex; a en cells isosceles-triangular, with the apex reaching i inner
surface

early $ 0. S. reeurvum

ll. Gn cells ac elliptic to trapezoidal in sectio

13. Green ae of branch leaves ae-aietts elliptic ‘ trapezoidal in section with
exposure equal or somewhat broader on the outer surface; hyaline cells of este

leaves on ae outer surface with aie pores arranged like beads along the co

mi

. 5. an
13. nage cells of branch leaves triangular to trapezoidal, with broader exposure on
outer ee hyaline cells of branch leaves with much fewer pores, not
anged like

re Hyaline cells of each leaves on the outer surface with few, small, ee pore
i corners S. ae
14. eas cells of branch leaves on the aie surface with rather numerous pores

in rows along the middle of the cell

15. Hyaline cells of branch leaves on the outer surface with numerous, large pores
eae in a single +

+ median row and also smaller pores in the corners, on the
ner surface with pores few or Jackin 3. S. majus
15 ane cells of branch leaves with numerous small to medium-sized pores on

the outer surface and few to several, minute ringed pores on the inner surfac

annulatum var. pou
10. Stem leaves entire or lacerate at the ie green cells of branch ia exposed only
or more the inner bee ce or, if + equally exposed, the pigment red*
t to ae pani cortical cells of stems with a
single pore at the upper end
Ste

m leaves aR eee fimbriate - the sides and across the apex; ag

of aa leaves divided 1 or more tim a doa

17. Stem leaves + fringed across the apex or only at the middle of the a hyalin
cells of stem leaves not (or only ae divided

*See also S. quinquefarium, p. 364.

18. Stems not at all red below the tips; stem leaves fringed across a broad, truncate
pex, with mid-basal cells greatly resorbed in a triangular, sieve-like area
17. S. girgensohnii
18. Stems often reddish (at least near the tips); stem leaves + erose at the middle of
a rounded apex, the mid-basal cells not conspicuously ea etais 18. S. russowii
16. Stem leaves not Pea or lacerate; cortical cells of stems without
19. Hyaline cells of branch ne with very small, round, strongly Se pores

on
the outer surface in the apical part 19. S. warnstorfii

19. Hyaline cells with larger, elliptic pores on the outer surface

20. Plants brown; stem leaves broadly pointed, nearly flat 20. S. fuscum

20. Plants green or (more commonly) red or red-tinged; stem leaves involute-pointed

21. Hyaline cells of branch leaves not conspicuously convex on the outer surface
(bulging % or less their diameters); stem leaves with hyaline cells + sama
not divided or 1-divided (and rarely subdivided); branch leaves a lustrou

when dry aS. capilaceu

21. Hyaline cells of branch leaves bulging on the outer surface about one

diameters; stem leaves with hyaline cells not fibrillose, mostly once to severa

times divided; branch leaves with a metallic luster when dry 22. S. subnitens

Cortical cells of stems and branches eee: those of the br sce porose but not

retort- shaped; leaves lacking a border of narrow ce ith hee dae furr em and branch
leaves similar, cucullate, roughened near the apex at back ee cause of extensive ene of hyaline
cells; perichaetial leaves with a lower median zone of unifor narrow cells surrounded at the

sides and apex by a network of fibrillose and porose hyaline cells in a network of green cells.

1. Sphagnum magellanicum Brid.—Robust, usually reddish or purplish plants
(pale-green in the shade), hummock-forming. Wood cylinder red; cortical cells of stems
in 3-4 layers, fibrillose, with 1 pore at the upper end of nearly every outer cell. Stem
leaves large, lingulate, with hyaline cells not divided, not fibrillose or sometimes only
near the leaf apex, mostly resorbed on the outer surface. Branches in fascicles of 4-5
(2 spreading), with cortical cells in 1 layer, fibrillose, often with 1 large pore. Branch
leaves broadly ovate, concave-cucullate, denticulate, with a resorption furrow; hyaline
cells plane or nearly so on both surfaces, on the inner surface with a few corner pores,
on the outer surface with numerous, large, elliptic pores in the corners and along the
commissures; in section, green cells small, shortly elliptic, central, entirely included.
Dioicous. Spores 22-31 wu, minutely or obscurely papillose.—n = ca. 17-19, 19, 38+3.

Breen, Mosses of Florida, Pl. 125 (fig. a Darlington, Mosses of Michigan, fig. 3. Jennings,
Mosses of Miao Rees (ed. 1) Pl. 2 (as S. medium), (ed. 2) Pl. 3. Welch, Mosses of
Indiana, fig. 10.—Fig. 1-8.

Typically in open parts of pene relatively oligotrophic. Greenland; Labrador to Alaska, south
to the Gulf of Mexico and California; Guatemala and Costa ica; i Jamaica and Santo Domingo;
South America to Tierra del Fuego; "Falkland Islands; Europe and A

CHEBOYGAN CO.—Hermits Bog, Iron Bridge (Carp Creek), Gleasons Bog, Bryants Bog,
Gates Bog, Nichols Bog, Livingston Bog, Mud Lake, Hebron Swamp, Little Lake 16, Weber Lake.
i ar

Bay. PRESQUE ISLE CO.—Hammond Bay, Loon Lake, Kelsey ae Clinton ae

The only one of our INOPHLOEA developing a red coloration, this common
species is a significant hummock-former in open bogs. It is well characterized by the
structure of its branch leaves as revealed in cross-section: the hyaline cells are nearly
plane on both surfaces, and the green cells are small, elliptic, central, and completely
included. Sphagnum centrale is a whitish-green, shade species with somewhat bulging
hyaline cells and apparently central green cells which are, however, longer and
lens-shaped and narrowly exposed on both surfaces because of thickened end walls.

Sphagnum magellanicum. 1. Stem cortex. 2. Branch cortex. 3. Stem leaf. 4. Branch leaf.
5. Back of branch leaf tip showing resorption. 6. Cross-section of branch leaf. 7. Cells of upper median
portion of branch leaf, outer surface. 8. Cells at margin of branch leaf near its apex, outer surface.
Sphagnum centrale. 9. Cross-section of branch leaf.
phagnum papillosum. 10. Cells of branch leaf, inner surface. 11. Cells of branch leaf, outer
surface. 12. Cross-section of branch leaf.
Dhagnum palustre. 13. Cross-section of branch leaf.

2. Sphagnum centrale C. Jens.—Robust, usually whitish-green or yellowish to
pale-brownish plants, never red. Wood cylinder green or brownish; cortical cells of
stems in 3-4 layers, fibrillose, the nile cells with 1-3 (sometimes 4 or more) pores.
Stem leaves large, oblong- obovate, concave; hyaline cells undivided, not fibrillose.
Branches in fascicles of 4-5 (2 spreading), with cortical cells in 1 layer, fibrillose,
usually with a single pore. Branch leaves broadly ovate, concave-cucullate, denticulate,
with a resorption furrow; hyaline cells slightly convex on the inner surface, somewhat
more so on the outer, on the inner surface with 2-6 round to elliptic, ringed pores at
the ends and along the commissures, on the outer surface similar, with 2-3 often
grouped at contiguous corners; in section, green cells slender and lenticular to elliptic,

23

usually nearly reaching both surfaces or the end walls thickened and reaching both
surfaces. a coeace dioicous.] Spores about 26-30 y, finely roughened or nearly
smooth.—n = 38+?

Darlington, Mosses of Michigan, fig. 4.—Fig. 9.

In shady places, especially in bog forests but also found on open mats among scattered
spruce and tamarack trees. Circumpolar; widespread in northern North America, south to New

York and Indiana

CHEBOYGAN CO.—Reeses Bog, Hermits Bog, Gorge of Carp Creek, Iron Bridge (Carp
Creek), Livingston Bog, Gates Bog, Ingleside, Mud Lake, Little Lake a Grass Lake, Weber Lake,
und. EMMET CO.

Pinegrove State Forest Campgro —Pleasa ntview Swamp, Stutsmanville, Galloway
og, between Levering and ie Lake, Ca Conway Bog, Bear ek Township, Wycamp
Lake, Big Stone Bay NA oe ee Bay, Hog Island Point, Epoufette, Hiawatha Sports
Club, Peggley Lake, Rexton, McN a Pond, Little Dollar Lake. Fee oe ISLE CO.—Hammond

y Lak
Bay, Evergreen Beach, te Isle aes Kelsey Lake, Clinton L

A whitish-green species of shaded habitats much resembling S. magellanicum in
typical form because the green cells of the branch leaves appear to be central and
included, but actually they are elongate and have thickened end walls which provide a
narrow exposure on both surfaces. Sphagnum centrale is not an entirely satisfactory
species. The green cells may be narrowly triangular-trapezoidal and cause confusion
with a form of S. papillosum lacking papillae (and this in turn approaches S. palustre
in cross-section). The hyaline cells of the stem leaves of S. papillosum are subdivided
(at least in the lower portions of leaves), and those of the branch leaves have numerous
fibrils and few pores; in S. centrale, the hyaline cells of the stem leaves are undivided,
and those of branch leaves have few fibrils but numerous pores.

Several local collections of S. centrale and S. cuspidatum have noticeably small
capsules with small, spherical, reticulate or papillose spores (about 11-13 yw). The spores
actually belong to a fungus, Tilletia sphagni Naw. (which is a conidial stage of the
discomycete Helotium schimperi Naw.). Similar spores in several members of the
ACUTIFOLIA were erroneously considered by Warnstorf (Hedwigia 25: 89-92. 1886)
to be microspores resulting from a subdivision of spore mother cells forming 16 rather
than the normal four spores. (I have also seen Tilletia spores in capsules of S.
perichaetiale Hampe from Virginia.

3. Sphagnum papillosum Lindb.—Plants slender to robust, usually yellowish or
brownish, hummock-forming. Wood cylinder brown or blackish; cortical cells of stems
in 3-4 layers, thin-walled, weakly fibrillose, the outer cells rarely with more than 1-2
pores. Stem leaves lingulate, short or long; hyaline cells frequently divided, not or
somewhat fibrillose (if not, the outer walls largely resorbed). Branches in fascicles of
4-5 (2 spreading), with cortical cells in 1 layer, fibrillose, often with a single large pore
at the upper ends. Branch leaves imbricate or slightly spreading, broadly ovate,
cucullate-concave, denticulate at the margins, with a resorption furrow; hyaline cells
somewhat convex on the outer surface, finely papillose on the inside where adjacent to
green cells (or the papillae often reduced or lacking), on the inner surface with many
small, ringed pores in the angles and often in apical and lateral regions of the leaf with
1-5 large, round pores along the cell middle, on the outer surface with 4-10 rounded or
elliptic pores in the corners and along the commissures; in section, green cells
truncately elliptic to trapezoidal or nearly triangular, + equally exposed or exposed
more broadly on the inner surface (the sides + curved). Dioicous. Spores 31-36 y,
papillose.—n = 38+4

Darlington, Mosses of Michigan, fig. 5. Jennings, Mosses of Western Pennsylvania, (ed. 2) PI.
2.—Fig. 10-1

In wet bog situations, but typically in the open, especially at or near the margins of bog
lakes; RS A or relatively eutrophic. Greenland; oi ador to North Carolina and the Great
Lakes; Alaska and British Columbia; Brazil; Europe and A

CHEBOYGAN CO.-—Reeses Bog, Iron Bridge ats Creek), Gates Bog, Nichols Bog, Mud
Lake, Topinabee, Little Lake 16, Weber Lake. EMMET CO.—Gladys Bog, Galloway Bog, Conway
Bog, Stutsmanville. MACKINAC CO.—Rexton tae, Little Dollar Lake, McNamara Pond,
Cranberry Lake. PRESQUE ISLE Co.—Evergreen Beach.

Plants typically growing in wetter, exposed, and mineral rich parts of bogs and
recognizable in the field, with reasonable certainty, because of a brownish tinge.
Microscopically, the species offers problems. The green cells of the branch leaves, as
viewed in section, are more or less trapezoidal to nearly triangular, and they are
usually finely and densely papillose on inner walls where they are adjacent to hyaline
cells. Often, however, in the so-called var. Jaeve Warnst., the papillae are difficult to
demonstrate or even lacking, and confusion with S. palustre and sometimes S. centrale
is common. Cardot (Bull. Soc. Roy. Bot. Belg. 25(1): 19-136. 1886) commented that
S. papillosum without papillae is no more than S. cymbifolium (= S. palustre). Meylan
(Rev. Bryol. et Lichén. 7: 184-187. 1935) came to the conclusion that S. papillosum,
though generally recognizable by its brown color, is really only a variety of S. palustre,
because of a similarity in leaf sections and variability in papillosity

However, the hyaline cells of the stem leaves, frequently divided in §. papillo-
sum, are usually quite undivided in S. palustre, S. icontrale: and S. magellanicum. S.
centrale is usually not too difficult to recognize because the green cells of the branch
leaves have thickened end walls exposed on both surfaces. The green cells of S.
palustre are really triangular and not triangular-trapezoidal. The differences are tedious
but cause little trouble locally, as S. palustre is very rare!

An important hummock former in Europe and some other parts of the United
States is a similar species offering similar problems in identification, S. imbricatum
Hornsch. ex Russ. (It has green cells which are equilateral-triangular in section; the
“fringe fibrils” on the inner walls of hyaline cells adjacent to the green cells may be
entirely lacking, poorly developed, or very obvious.) The marvelous scanning electron
photomicrographs published by Mozingo ef al. (Bryol. 72(4): 484488. 1969) are
illustrative of all the INOPHLOEA. The details of the hyaline cells and pores of branch
leaves are beautifully shown.

4. Sphagnum palustre L.—Plants low to robust, green or uncommonly brownish-
tinged. Wood cylinder brown; cortical cells of stems in 3 layers, thin-walled, fibrillose,
the outer cells with 1-4 (and often more) irregularly rounded pores. Stem leaves large,
long-lingulate to subspatulate; hyaline cells not divided, often somewhat fibrillose
above, resorbed on the outer surface. Branches in fascicles of 4-5 (2 spreading), with
cortical cells in 1 layer, fibrillose, frequently with 1 large pore. Branch leaves imbricate
to squarrose, cucullate-concave, broadly ovate, denticulate, with a resorption furrow;
hyaline cells somewhat convex on the outer surface, on the inner surface with elliptic
to round pores in the corners and a few large, round pores at the middle, more
numerous (2-8) near the leaf margins, on the outer surface with large, round pores at
the ends and numerous (3-10), smaller, elliptic, ringed ones along the commissures, in
section, green cells isosceles-triangular and nearer the inner surface). Dioicous. Spores
about 24-29 u, finely roughened or nearly smooth.—n = 38+?, 38+4.

25

Breen, Mosses of Florida, Pl. 126. Darlington, Mosses of Michigan, fig. 6. Jennings, Mosses of
Western Pennsylvania, Se 1) Pl. 1 (as S. latifolium), (ed. 2) Pl. 1 (as S. ‘palustre). Welch, Mosses of
Indiana, fig. 11.—Fig. 13

shade in bog forests or sometimes in semi-open parts of bog mats, tolerating a wide range
of fe conditions. ee lear through the Great Lakes and south to the Gulf of Mexico;
Mexico; Europe and Asi

CHEBOYGAN CO.-—Vincent Lake, Weber Lake.

Andrews considered S. palustre the most common of the INOPHLOEA in the
Northern Hemisphere and said that anyone collecting Sphagna in North America would
want directions for avoiding it! However, his concept (Bryol. 25: 7-8. 1912) included
the typical form with green cells of branch leaves unmistakably triangular in section
and other forms with more or less triangular-trapezoidal cells. S. palustre is actually
uncommon in Michigan and very rare locally. The form with trapezoidal or somewhat
triangular green cells is common, however, and is considered here as a non-papillose
form of S. papillosum.

II. Cortical cells of stems and branches without fibrils, some or all of those of branches

perichaetial leaves consisting of uniformly narrow cells in the border and usually also in the apex or
nearly throughout.

ortical cells of branches all + uniform and somewhat retort-shaped; branch leaves with a
resorption peas lacking a border of narrow cells.

5. Sphagnum compactum DC.—Plants usually short and compact, dirty-white
(especially when dry), sometimes pale-green or brownish-tinged. Wood cylinder brown,
cortical cells of stem in 2-3 layers, thin-walled, without fibrils or pores. Stem leaves
minute (less than 1mm. long), concave, bluntly triangular-lingulate, sometimes +
lacerate at apex; hyaline cells not divided, without fibrils. Branches short, crowded, in
fascicles of 4-5 (2 spreading or ascending), with cortical cells in 1 layer, without fibrils,
the apex of each cell projecting as a slight neck ending in a pore. Branch leaves
imbricate to squarrose, large, involute-concave, broadly ovate-hastate, ending in a
broad, truncate, toothed apex which is concave but not cucullate, slightly denticulate
at the margin, with a resorption furrow; hyaline cells somewhat convex on the inner
surface, not or slightly so on the outer surface, on the inner surface with ringed pores
in the corners (usually 3 per cell and occurring as 3’s at adjoining comers), on the
outer surface with 4-8 large, round pores mostly enclosed within a subdivision limited
by a fibril thickening and also with numerous similar subdivisions lacking a perforation,
thus forming pseudopores; in section, green cells elliptic, central or nearer the outer
surface, included. Monoicous. Spores 25-33 4, very finely roughened or nearly
smooth.—n = 19+2, 19+2-4.

eee Mosses of Michigan, fig. 7. Jennings, Mosses of Western ou ae (ed. 1) Pl. 2
& (ed. Oe Pl. 3 (as var. squarrosum). Welch, Mosses of Indiana, fig. 13.—Fig. 14-2

In moist to relatively dry habitats, especially at the outer margins of open Chamaedaphne
bogs and sometimes on banks of ditches in boggy areas; oligotrophic. Greenland; Labrador through
the Great Lakes and au to the Gulf of Mexico;.Alaska and British Columbia; Azores and
Madeira; Europe and Asi

CHEBOYGAN CO.-—Near Douglas Lake, Gates Bog. EMMET CO.-—French Farm Lake,
between Levering and Carp Lake.

| \ ' ‘ap , ) ’ \ / f\ 7 {
_ he \ Oy Ce | rt |
| | i | \
15 | |
\ |
a at —
‘ a Q
14 ve 9
| Sh |
\ ae OK
\ \__ / wi y =
\ We / p——
| \ 16 / ; ‘ \ f : e
- ay vA \ fs / 4 ( \ }
\ : i / C7
\ i i O)
\ 10S aa 21
| WA \ Mo .
| Jt y, “Av 7 Sw,
—~ { 20
AA i \
i 7 ( : Ay \ |
| heh \ Ae try

mM TY oe 24 i a

Sphagnum compactum. 14. Stem cortex. 15. Branch cortex. 16. Stem leaf. 17. Branch leaf.
18. Cells at margin of branch leaf apex showing resorption. 19. coals of branch leaf. 20.
Cells of branch leaf, inner surface. 21. Cells of bran Bs leaf, outer é

Sphagnum wulfianum. 22. Stem cortex. 23. Branch re in aie ce and in profile (right).
24. Stem leaves. 25. Branch leaves. 26. Cross-section of branch leaf. 27. Cells of branch leaf, inner
surface. 28. Cells of branch leaf outer surface

Although the leaves are margined by a resorption furrow, as in the INOPHLOEA,
they are more or less truncate at the apex and smooth at the back, and the cortical
cells of stems and branches are not fibrillose. The compact, pin-cushion manner of
growth, relatively large and fleshy-looking leaves, as well as a washed-out whitish color
are suggestive of the genus Leucobryum. The species is further distinguished by branch
leaves with nearly central green cells and hyaline cells with large pores arranged in 3’s °
at adjacent corners on the inner surface. Plants growing in the shade have a looser
habit and spreading leaf tips.

Ze

Cortical cells of branches usually of two types, the retort cells only in axils of leaves (1-2
per ae axil) or apparently lacking; branch leaves usually bordered by narrow cells, only rarely with
a resorption furrow.

. Stem leaves much shorter than branch leaves. Branches 6-12 in a fascicle, crowded in
large capitulum; green cells of branch leaves truncately elliptic in section, + equally exposed on
both surfaces

6. Sphagnum wulfianum Girg.—Plants loosely tufted, rather robust, dark-green
tinged with brown. Stems dark, stiff and brittle. Terminal bud not conspicuous. Wood
cylinder red-brown; cortical cells of stem in 24 layers, rather small, with moderately
thick walls, without fibrils or pores. Stem leaves very small, oblong-triangular, slightly
concave, involute to an abruptly pointed apex, narrowly bordered; hyaline cells
without fibrils, often divided. Branches crowded in fascicles of 6-12 (3 or more
spreading), with cortical cells in 1 layer, without fibrils; retort cells with a short neck.
Branch leaves with points spreading when dry, small, narrowly ovate-lanceolate,
involute-tubulose to a sharp point toothed across the apex, entire, bordered by 2-3
rows of narrow cells; hyaline cells very slightly convex, more so on the outer surface,
very finely papillose on the inside where overlying green cells, on the inner surface
with few (rarely 3-4), inconspicuous pores in the ends and corners, on the outer
surface with small, ringed pores along the commissures (4-8 per cell in the upper half
of the leaf, generally fewer below); in section, green cells truncately elliptic, equally
exposed on both surfaces or slightly more broadly on the outer surface (the lumen
lenticular, central). Dioicous or sometimes monoicous. Spores 18-24 y, finely granu-
lose-roughened.—n = 19+

Darlington, Mosses of Michigan, fig. 8. paren Mosses of Western Pennsylvania, (ed. 2) Pl.
2. Sullivant, ene Muscorum Supplement, Pl. 9.—Fig. 22-28.

In bog forests, usually on low ridges resulting from logs in an advanced state of decay, rarely
open bogs. Gr eenland: Eee om Canada to Connecticut and the Great Lakes; Alberta and
Britich Columbia; Europe ‘and A

CHEBOYGAN CO.—Reeses Bog, Hermits Bog, Gleasons Bog, Gates Bog, Livingston Bog.
EMMET CO.—Antique Bog, Stutsmanville, Wycamp Lake, Conway Bog. MACKINAC CO.-—Prentiss
Bay, Epoufette, Rexton Lake, Hog Island Forest Campground, Hendricks, McNamara Pond.
PRESQUE ISLE CO.—Hammond Bay, Clinton Lake

The plants are loosely tufted and appear top heavy because of an exceptionally
large, rounded capitulum resembling a head of clover (resulting from an unusually high
number of branches in each fascicle as well as a crowding of fascicles). Further
distinctive traits are furnished by dark, woody, brittle stems and tips of branch leaves
nicely curved outward.

b. Terminal bud large. Stem leaves with a narrow border only toward the base, with
extensive resorption especially on the dorsal surface resulting in a perforated or somewhat lacerate

broadly on the outer surface, the hyaline cells usually finely papillose on the inner walls adjacent
to the hyaline cells.

7. Sphagnum squarrosum Crome—Plants robust, bright- or pale-green to yellow-
ish, ee a prominent terminal bud. Wood cylinder green to red-brown; cortical cells of
stem 2-3 layers, thin-walled, without fibrils or pores. Stem leaves large, long,
ior qin nistes slightly concave, indistinctly bordered, weakly lacerate at the apex;

28

hyaline cells mostly undivided and without fibrils, with membrane pleats, the walls
almost completely resorbed on the outer surface. Branches in fascicles of 5 (2
spreading), with cortical cells in 1-2 layers, without fibrils, the retort cells with an
inconspicuous neck. Branch leaves wide-spreading or squarrose from an erect base, large
(2-2.8 mm. long), ovate-hastate, abruptly narrowed to an involute, acute, toothed apex,
bordered by linear cells, entire; hyaline cells convex on both surfaces, more so on the
inner, often papillose on the inside where overlying green cells, on the inner surface

with 4-10 round pores at the ends and near the commissures, on the outer surface with
1-3 per cell (except in the median basal region where they are large and up to 10 or
more); in section, green cells triangular with the base exposed on the outer surface or
trapezoidal and narrowly exposed on the inner surface as well. Monoicous. Spores
17-27 wu, nearly smooth or very finely granulose.—n = 19, 19+2, 19+2-4, 19+4, 38+2-4.

gton, Mosses of Michigan, fig. 9. Grout, Mosses with Hand-lens & Microscope, fig. 10a.
Jennings, ees of Western Pennsylvania, (ed. 2) Pl. 70.—Fig. 29-34.

In depressions in bog forests; eutrophic. Greenland; Labrador to Saskatchewan on Montana,
south to New Jersey, Ohio, and Colorado; Alaska to California; Arizona; Europe and A

CHEBOYGAN CO.—Reeses Bog, Gorge of Carp Creek, Iron Bridge aa ee aaa as
Lake, Bryants Bog, Nichols Bog, Livingston Bog, Little Lake a Grass Lake. EMMET C na
Beach, Wycamp Lake, between Levering and Carp Lake, Larks Lake, Cecil a Big ze nn
MACKINAC CO.—Mackinac Island, Epoufette, Hog Is a Forest oo Hendricks,
McNamara Pond, Hessel. PRESQUE ISLE CO.—Hammond Bay, Evergreen Beac

A woodland species recognizable at a glance because of its large size, bright-green
color, and branch leaves abruptly narrowed and widely spreading. Like S. feres, to
which it is closely related, it has a prominent terminal bud.

8. Sphagnum teres (Schimp.) Angstr. ex Hartm.—Plants rather slender, mostly
yellowish- or brownish-green, with a prominent terminal bud. Wood cylinder brownish
or red-brown; cortical cells of stem in 3-4 layers, thin-walled, without fibrils or pores
(but on staining a large rounded or elliptic pore-like thin spot can be demonstrated in
each cell, or sometimes | at either end). Stem leaves large, + elliptic, flat, narrowly and
indistinctly bordered even at base, + perforate or weakly fringed at the apex because

leaf), without fibrils, the walls almost entirely resorbed on the outer surface, with

membrane pleats. Branches in fascicles of 5 (2-3 spreading), with cortical cells in 1

layer, the retort cells with an inconspicuous neck. Branch leaves imbricate or

sometimes + spreading or squarrose at the tips, small (1-2 mm. long), ovate-lanceolate,

strongly involute toward the toothed apex, bordered by linear cells, entire; hyaline
the i

trapezoidal, exposed exclusively or more broadly on the outer surface. Dioicous.
Spores 20-26 y, finely granulose.—n = 19+2, 19+4.

Darlington, Mosses of Michigan, fig. 10. Jennings, Mosses of Western ae Loe 1) Pl.
2 & (ed. 2) Pl. 3 (as var. subteres). Sullivant, cones Muscorum Supplement, Pl. 4.—Fig. 35-42.

n wetter parts of floating mats, often in wet sedge mats near open water and also in alder
thickets nearby; distinctly eutrophic. Greenland; Pye to Alaska and south to Pennsylvania, the
Great Lakes, Colorado, and California; Europe and A

BOYGAN CO.-—Reeses Bog, Nichols Bog, Mud Lake, Little Lake 16, Weber Lake.
ENMET co. —Carp Lake. MACKINAC CO.-Peggley Lake, Rexton, cial Hog Island Forest
Campground, Hiawatha Sports Club. PRESQUE ISLE CO. _Hammond B

(/ aw
! | UA JP WAN Np
L ZN A\ Sf NAS aa ‘i
AAR \) Wf NA \ \
ia
f NN fe - BM >
A Dt. CK
41 7 WRA
A a

Sphagnum squarrosum. 29. Stem cortex. 30. Stem leaf. 31. Branch leaves. 32. Cross-section
of branch oe 33. Cells of branch leaf, inner surface. 34. Cells of branch leaf, outer surface.

Sphagnum teres. 35. Stem cortex. 36. Stem leaves. 37. Cells at apex of stem leaf. 38. Branch
cortex. 39. Branch lea . Cross- ee of branch leaf. 41. Cells of branch leaf, inner surface.
42. Cells of branch leaf, outer surfac

Sphagnum teres is recognized in the field more by arenes than by special
characteristics. The plants are slender and yellowish and grow in very wet, mineral-rich
habitats; they have a large pela bud, relatively stiff, Meade! branches with leaves
appressed when moist but more or less spreading to squarrose when dry, and
red-brown stems. When peri in the shade, they tend to have more spreading leaf
tips, and occasional robust forms may be difficult to distinguish from S. squarrosum
(at least as dried specimens). Ordinarily the two species are not difficult to distinguish,
however; S. squarrosum is stouter and bright-green, with ovate-hastate rather than
ovate-lanceolate branch leaves which have smaller and more numerous pores. (A further

30

point of contrast is that S. squarrosum fruits commonly and abundantly.) Both species
are most clearly differentiated from others of the LITOPHLOEA by their large,
long-lingulate stem leaves with short, broad, and efibrillose hyaline cells extensively
resorbed (but especially so on the outer surface) resulting in perforation and slight
laceration of the apex. In keying, S. teres can be confused with S. girgensohnii because
of a degree of fringing at the apex of stem leaves and because of membrane thinnings
in the stem cortex resembling pores on staining. The membrane thinnings, however, are
eroded and torn areas not bounded by a distinct circular wall as in S. girgensohnii.
Sphagnum girgensohnii can also be distinguished from S. teres by elliptic pores on the
outer surface of branch leaves, strongly bordered stem leaves with most hyaline cells
divided and erosion in a conspicuous triangular area in the basal median portion, and
(usually) isodiametric cells of the stem cortex. Broader exposure of green cells on the
outer surface and the fine papillae on the inner walls of hyaline cells further aid in
distinguishing S. teres.

c. Stem leaves usually bordered along the sides by several rows of elongate cells, with
resorption on the inner surface, or less often on both surfaces and then lacerate-fringed; perichaetial
leaves usually showing resorption on the inner surface; branch leaves often flat and + wavy at the
margins when dry; green cells of branch leaves usually with exclusive or greater exposure on the
outer surface.

9. Sphagnum riparium Angstr.—Plants robust, dark green, hard and shiny in
appearance when moist. Terminal bud very large; young pendent branches in pairs
between rays of the capitulum. Wood-cylinder green; cortical cells of stems scarcely
differentiated, long, narrow, thick-walled, without pores or fibrils. Stem leaves rather
large, broadly oblong-triangular to elliptic, deeply rent or conspicuously perforate at
the middle, bordered by 4-6 rows of narrow cells; hyaline cells often divided once or
twice, especially toward the leaf apex, without fibrils, the membrane on the inner
surface almost entirely resorbed. Branches in fascicles of 4 or 5 (2 spreading), their
cortical cells in 1 layer, nonfibrillose, of 2 kinds, the retort cells with an inconspicuous
neck. Branch leaves when dry slightly undulate with tips reflexed, ovate-lanceolate,
involute to a narrow, toothed apex consisting entirely of green cells, bordered by 2-5
rows of linear cells, without a resorption furrow; hyaline cells somewhat convex on the
outer surface, nearly plane on the inner, and on the inner surface with rounded pores
mostly of nearly cell width at the ends and elsewhere, on the outer with small pores at
ends and corners (3-5 per cell in the upper portion of the leaf); in section, green cells
trapezoidal with broader exposure on the outer surface, the lumen triangular. Dioicous.
Spores about 20-30 uw, minutely roughened.—n = 19+4.

Fig. 43-50.
re, in wet, relatively acid habitats, especially at margins of ponds and in sedge mats near

a
open water; mesotrophic or perhaps oligotrophic. Circumpolar; Greenland; Labrador to Alaska,
south to New Hampshire, New York, Michigan, and Washington.

EMMET CO.-—Stutsmanville Bog.

Large, dark-green, hard and shiny when moist, S. riparium is much stouter than
S. recurvum and otherwise distinguished by a large terminal bud and turgid branches
with pointed tips and broad, imbricate leaves narrowed to a small, recurved tip and not
undulate. The leaf tip consists of uniformly narrow, green cells rather than the usual
network of green and hyaline cells. The most striking feature of the species is the large
stem leaves which are deeply and conspicuously perforate or split at the middle of the
apex. The pairing of young pendent branches, observable between the rays of the
capitulum, is a good field character for S. riparium, S. recurvum, and S. pulchrum.

ie AAS
YN Ye Af iah\
| Wy \ 44 \
| \ : | if We \\
Veal | / /
| |

55 rpgeee 5S. mo 56 ee. lea a ;

Sphagnum riparium. 43. Stem cortex. 44. Stem leaves. 45. Cells at apex of stem leaf. 46.
Branch cortex. 47. Branch leaves. 48. Cross-section of branch leaf. 49. Cells of branch leaf, inner
ace

ace,

agnum recurvum. 51. Stem cortex. 52. Stem leaves. 53. Branch cortex. 54. Branch leaves.
55. Cross- jane of branch leaf. 56. Cells of branch leaf, inner surface. 57. Cells of branch leaf,
outer surfac

10. Sphagnum recurvum P.-Beauv., sensu Jato—Plants usually rather robust,
bright-green or (more commonly) tinged ‘with yellow or brown. Stems often reddish
toward the tip; wood cylinder yellow-green; cortical cells of stem not or only slightly
differentiated, without fibrils or pores. Stem leaves very small, triangular to
oblong-triangular, the border poorly developed or occupying nearly the width of the
base; hyaline cells not divided, without fibrils or pores (or rarely with traces of fibrils).
Branches in fascicles of 5 (2 spreading) with cortical cells in 1 layer, the retort cells
with an inconspicuous neck. Branch leaves undulate and spreading at the tips when

a2

dry, lanceolate, somewhat channeled toward the toothed apex, bordered by linear cells,
entire; hyaline cells slightly convex on the inner surface, and on the inner surface with
4-7 large, round pores in the angles and near the commissures (sometimes visible only
on strong staining), on the outer surface with end pores throughout and in the apical
portion with 2-6 small, ringed pores also in the angles and near the commissures; in
section, green cells isosceles-triangular with the base exposed on the outer surface of
the leaf, its apex reaching the inner surface or nearly so. Dioicous. Spores 24-27 y,
finely-roughened to nearly smooth.—n = 19+?, 19+2, 19+4.

en, Mosses_ of Florida, Pl. 128 (fig.8-14). Darlington, Mosses of share fig. 12A
(as var. ape Jennings, Mosses of Western Pennsylvania, (ed. 1), Pl. 2, Pl. 3 (as S. See Het
(ed. 2) Pl. 3, Pl. 6 (as var. ns Welch, Mosses of Indiana, fig. 14.—Fig. 51-

Very common, usually in mesotrophic habitats, especially in the hollows of open
aes mats or Vaccinium heaths or in the transition between bog and bog forest.
Greenland, Newfoundland, Alaska, south to California, Colorado, Nebraska, Missouri, and the Gulf of
Mexico; Panama; South America; Europe and Asia

CHEBOYGAN CO.-—Reeses Bog, Iron Bridge (Carp Creek), Gleasons Bog, Bryants Bog,
Nichols Bog, Maple River, Livingston Bog, Gates Bog, Vincent Lake, Mud Lake, Grass Bay, Little
ke 16, Pine Grove State Forest Campground, Weber Lake. EMMET CO.—Near Levering, between

Hendricks, Little Dollar Lake, one Pond, Peggley Lake, Hiawatha Sport ae “PRESOUE
ISLE CO.—Loon Lake, Hammond B

Locally very common and easy to recognize (in a broad sense) in a dry condition
because of soft, nearly flat, broadly pointed leaves recurved at the tips and undulate at
the margins. The leaves are concave and spirally ranked when moist. The young
pendent branches ee in pairs between the rays of the capitulum are useful in
distinguishing S. recurvum and its relatives (S. pulchrum and S. riparium) from
members of the ACUTIFOLIA. Other useful characters include the poorly
differentiated cortex, relatively small stem leaves, and the brown-tinged antheridial
“catkins

The name recurvum may be in question. The Europeans have tended to shun it,
because it is American and, from their point of view, geographically unlikely. Shunning
the oldest name without explaining its relationship to the complex is scarcely a
satisfactory solution. I prefer to follow Andrews, knowing that his judgment was
invariably good (although his concepts were over-inclusive).

Our Michigan material falls into three categories, as follows:

Var. recurvum [S. apiculatum H. Lindb., S. fallax (Klinggr.) Klinggr., S.
mucronatum (Russ.) Zick.]—Plants of medium size, usually yellowish or brownish.
Stems not red-tinged. Stem leaves small (ca. 0.8-1 mm. long), about as broad as long,
deltoid, rather sharply pointed, broadly bordered at base. Branch leaves not distinctly
in rows, the hyaline cells on the outer surface with rounded pores at the upper ends
(and sometimes with smaller pores in the corners) and on the inner surface with
rounded end and corner pores (often visible only on strong staining).

Var. tenue Klinggr. [S. angustifolium (Russ.) C. Jens., S. parvifolium (Warnst.)
Warnst.] —Plants relatively slender, usually brownish or yellowish Stems sometimes
red-tinged. Stem leaves ca. | mm. long (as broad as long), + deltoid, broadly pointed or
bluntly obtuse, concave at the tip, not strongly bordered at base. Branch leaves on the
outer surface with few end and corner pores, on the inner surface with numerous end
and corner pores.—n = 19+

Andrews’ rie znd of the var. fenue as slender plants with small capitula
and smaller, not or scarcely undulate branch leaves does not work out convincingly.

n

33

Var. amblyphyllum (Russ.) Warnst.—Plants relatively stout, pale green or yellow.
Stems without red tinges. Stem leaves oblong- or ovate-deltoid, usually somewhat
longer than broad (ca. 1-1.2 X 1 mm.), somewhat to considerably eroded because of
extensive resorption at the broad, rounded, flat apex; border not sharply
differentiated at base. Branch leaves 5-ranked when moist, with margins reflexed when
dry.

This variety can be confused with S. pulchrum because of its relatively large size
with branch leaves (especially in the capitulum) somewhat spirally ranked when moist
and leaf margins more or less reflexed.

The scarcity of pores in the leaves of S. recurvum and related species makes
them less absorbent, and hence during the First World War they were little used in the
production of surgical dressings. Some of the INOPHLOEA, especially S. papillosum,
were prized because of numerous large pores in the branch leaves and also in the
greatly differentiated cortex. Nichols (Bryol. 21: 53-56. 1918) reported that good
Sphagnum pads could absorb 16-22 times their weight as compared with an absorption
of 5-6 times by cotton pads.

The use of Sphagnum in surgical dressings has an incidental advantage because of
a built-in antibiotic property. Sphagnum is by nature somewhat antiseptic. This
property, plus acidity and anaerobic conditions, make possible the accumulation of
peat. Mosses in general have few parasites, either plant or animal, and for that reason
require no special protection in the herbarium. McCleary and Walkington (Rev. Bryol.
et Lichén. 34: 309-314. 1966) tested extracts from 50 species in 33 genera of mosses
and found that 18 species show strong antibiotic properties. The antibiotically active
mosses were found in the genera Sphagnum, Dicranum, Mnium, Atrichum, and
Polytrichum.

Magdefrau (Zeitsch. Bot. 29: 337-375. 1935) established the fact that in mosses
both external conduction by capillarity and internal conduction through the central
strand are important. He found that at 90% relative humidity internal conduction
alone suffices to maintain turgor, but at lower humidities both types of conduction are
necessary. He also (Ann. Bryol. 10: 146. 1938) compared the effectiveness of internal
and external conduction in various mosses (in grams/0.2 grams dry weight in 24 hours
at 70% relative humidity): .

Total conduction Internal conduction
Sphagnum recurvum 6.54 0.07
Mnium undulatum 2.22 1.13
Thamnium alopecurum 0.019 0.007
Drepanocladus vernicosus 22.73 0.79
Rhytidiadelphus triquetrus 1.67 0.11
Polytrichum commune 3.32 2,24

Magdefrau also showed an unmistakable relationship of growth form to conduc-
tion. Very likely the denser tufts of Drepanocladus vernicosus account for greater
conduction. In Sphagnum only 1% of the conduction was internal, compared with 50%
in Mnium undulatum and 66% in Polytrichum commune. A comparison of S. recurvum
with some of the INOPHLOEA would be particularly informative.

The report of S. lindbergii Schimp. in Darlington’s Mosses of Michigan (without
collection data) must be rejected. S. lindbergii is a northern relative of S. recurvum
with stem leaves lacerate across a broad apex.

11. Sphagnum pulchrum (Lindb. ex Braithw.) Warnst.—Plants rather robust,
bright-green or brownish, shiny when dry. Apical bud small; young pendent branches
in pairs between the rays of the capitulum. Wood cylinder yellow-brown; cortical cells

34

somewhat differentiated (but with thick walls) in 1-2 layers, without fibrils or pores.
Stem leaves small (about | mm. long), triangular, concave, involute-pointed, the border
occupying nearly the width of the base; hyaline cells not divided, rarely fibrillose.
Branches in fascicles of 4 or 5 (2 spreading); cortical cells in 1 layer, the retort cells
with conspicuous necks. Branch leaves somewhat reflexed, undulate-reflexed at the
margins and somewhat lacquered in appearance when dry, 5-ranked when moist (and
spirally arranged wet or dry), broadly ovate, abruptly narrowed to a rather long, broad,
channeled, toothed, cuspidate point, bordered by linear cells, entire; hyaline cells
somewhat convex on the inner surface, and on the inner surface with 3-6 rounded
pores at the corners, on the outer surface with 2-5 small, ringed pores at ends and
corners and along the commissures; in section, green cells broadly triangular, with the
base exposed on the outer surface and the apex usually reaching only % or less the
distance to the inner surface. Dioicous. Spores 25-28 y, slightly roughened [not seen. ]

Darlington, Mosses of Michigan, fig. 13.—Fig. 58-63.

Rare; in open, wet portions of bogs, at the margins of bog lakes but not normally

submerged; ‘mesotrophic. Europe; Newfoundland to northern Ontario, south to Wisconsin, northern

Michigan, and New Jersey (an Atlantic species in coastal areas of northwestern Europe, ‘also more
common along the Atlantic coast of North America).

MACKINAC CO.—Rexton Lake, Little Lake.

Not always easy to distinguish from S. recurvum var. amblyphyllum but usually
recognized as follows: generally robust and often brownish, with cortical cells small
and thick-walled but somewhat more differentiated. Branch leaves often in five spiral
ranks, broad, appearing abruptly apiculate because of involute apical margins, the
margins otherwise reflexed and somewhat undulate. The green cells of the branch
leaves are equilateral-triangular and do not reach the upper surface of the leaves.

12. Sphagnum cuspidatum Ehrh. ex Hoffm.—Plants weak and flexuose (especially
when wet), slender to large, often very long and sometimes plumose, whitish-green to
yellowish (sometimes mottled with brown because of colored antheridial branches).
Apical bud small. Wood cylinder yellow-green; cortical cells of stem clearly
differentiated but rather small, in 2-3 layers, without fibrils or pores. Stem leaves
small, triangular-ovate, bluntly pointed, strongly concave, slightly toothed at the apex,
weakly bordered; hyaline cells not divided, with or without fibrils in the apical
portion. Branches generally in fascicles of 4 (2 spreading), with cortical cells in 1 layer,
the retort cells with conspicuous necks. Branch leaves + undulate and falcate when dry,
narrow and long-lanceolate (about 4-5 : 1), especially long and cohering at branch tips,

omewhat involute, bordered by linear cells, and entire below the toothed apex;
aie cells slightly convex on the inner surface, and on the inner surface with few,
small, and often inconspicuous pores in the ends or corners, those on the outer surface
few, + ringed, somewhat larger; in section, green cells trapezoidal with broader
exposure on the outer surface. Dioicous. Spores 33-42 u, + granulose.—n = 1942,
19+3-4

reen, Mosses of Florida, Pl. 129 (fig. 1-6). Darlington, Mosses of Michigan, fig. 14.
Jennings, Mosses of Western Pennsylvania, (ed. 2) Pl. 4.—Fig. 64-70.

In limp, loose masses meee 7 Perse of water (at least during part of the season), in
relatively acid situations, at the edge of bog lakes or in drainage channels in older parts of the mat,
also in depressions in boggy heaths. gana to Florida and the Great Lakes; Europe and
Asia.

zie
ey
2y

LD.

patesas

dT
S

L

r
\
Pi
a ee a
a

Litt

Hebe L—~/ 67

hagnum pulchrum. 58. Stem cortex. 59

=
68
A Ono Eee

Hf 60 m

13
|

o { \
cat fA
| cs ee 8

| . \ |

@
ey
a Sy a
a
Col ose

Gx

EC [
:g 5
oid
oy

a

Fa

Stem leaves. 60. Branch leaves. 61. Cross-section

Sp ;
of branch leaf. 62. Cells of branch leaf, inner surface. ee Cells of branch leaf, outer surface.

num cuspidatum, 64. Stem cortex

eaf. 66. Branch cortex. 67. Branch leaf.

Spha . 65. Ste
68. Cross-section of branch leaf. 69. Cells of Beach ae inner surface. 70. Cells of branch leaf,

outer surface

CHEBOYGAN CO.-—Iron Bridge (Carp Creek), Bryants Bog, Gates Bog, Nichols Bog, Mud

Lake, Topinabee, Weber Lake. EMMET CO.
Stutsmanville, Big

—Maple River,
Stone Bay. MACKINAC CO.—Rexton Lake, Little Dollar Lake, Hendricks,
Cranberry Lake. PRESQUE ISLE CO.—Loon Lake.

between Levering and Carp Lake,

Plants of particularly wet habitats (usually submerged and appearing weak and
limp). As compared with S. majus, the branch leaves are relatively long and narrow
(usually about four times as long as broad) and stick together like the point of an
artist’s brush when drawn from the water. The pores on the outer surface of the

branch leaves are few or no
subme

none; those on the inner surface are more numerous (but in
rged forms sometimes entirely lacking). S. cuspidatum commonly grows barely

submerged in shallow pools and drainage channels of open bogs. As such puddles dry
out, it assumes a form that is often confused with S. recurvum. Typically, the branch

36

leaves are longer, often nearly linear, frequently without undulations, and their cells
are narrow with pores only at the ends. In questionable cases, a number of characters
can be used: In S. cuspidatum, the pendent branches are poorly differentiated and do
not conceal the stems; the young pendent branches are not paired; the stem leaves are
somewhat fibrillose above; the stem cortex consists of about two layers of fairly
good-sized, rather thin-walled cells; and the green cells of the branch leaves are
trapezoidal in section. In S. recurvum, the pendent branches (paired in the capitulum)
are long and appressed to the stem; the stem leaves are not fibrillose above; the stem
cortex is less differentiated; and the green cells of the branch leaves are triangular in
section. The long-tapered branches of $. cuspidatum are also useful in identification.

13. Sphagnum majus (Russ.) C. Jens.—Plants medium-sized to large, light-brown
or brownish-green (and somewhat mottled because of darker shades within the
capitulum when moist), usually shiny when dry; terminal bud scarcely discernible to
moderately enlarged. Wood cylinder yellow-green; cortical cells of stem poorly
differentiated in 1 layer of rather small cells without fibrils or pores. Stem leaves
rather small, oblong-triangular, + concave and rounded at the apex, the border very
broad and increasing in width toward the base; hyaline cells not divided, with
rudimentary fibrils in the apical portion or none. Branches in fascicles of 4-5 (2
spreading), with cortical cells in 1 layer, the retort cells with inconspicuous necks.
Branch leaves not or sometimes slightly undulate, usually subsecund, long,
ovate-lanceolate (broader than in S$. cuspidatum), somewhat involute-pointed, bordered
by linear cells and entire; hyaline cells on the inner surface somewhat convex and with
pores lacking (or occasionally with a few, larger, unringed pores restricted to corners in
the apical portion uf the leaf), on the outer surface with numerous, large pores often
in a single + median row (% or more the width of the cell) and also smaller pores in
the lateral corners (the median pores sometimes few or lacking); in section, green cells
mostly trapezoidal with broader exposure on the outer surface. Dioicous. Spores
24-38 uw, somewhat roughened to nearly smooth.—n = 38.

Darlington, Mosses of Michigan, ae 15 (as S. dusenii). Sullivant, Icones Muscorum
Supplement, PL 2 (as S. cuspidatum var.).—Fig. 71-75.

Submerged or emergent, in relatively eutrophic habitats, in open sedgey bogs and commonly
in the moat surrounding bog 1 Labrador to Alaska and British Columbia, south to Virginia and
ae Great Lakes; Europe and Asi

CHEBOYGAN CO.-—Iron Bridge (Carp Creek), pees Bog, Gleasons Bog, Bryants Bog,
Vincent Lake, Gates Bog, Nichols Bog, Mud Lake, Topinabee, Little Lake 16. EMMET CO.—
Stutsmanville, Cecil Bay, Big Stone Bay, Carp Lake, between Levering and Carp Lake. MACKINAC
CO.—Little Dollar Lake.

Sphagnum majus is best distinguished from S. cuspidatum by the rather large and
numerous pores on the outer surface of the branch leaves (which may, unfortunately,
be reduced or lacking in submerged forms). The macroscopic differences are sometimes
helpful: S. majus is stronger, stouter, essentially erect, and nearly always brownish;
when moist, it seems brownish-green mottled because of darker shading within the
capitulum. The branches of the capitulum are less pointed and the branch leaves
broader and not undulate. S. annulatum var. porosum has a large terminal bud and
hyaline cells of branch leaves with pores on both surfaces (those on the inner surface
few and exceedingly small).

14. Sphagnum annulatum Warnst. var. porosum (Schlieph. & Warnst. ex Warnst.)
Maass & Isov. ex Maass—Plants medium-sized to large, green or brown, + glossy when

\ > 8 |
| Wes
\ d iG
\ ae:
ALS
JG
nf ¢
LON
a a aa \
{ |
te
78

Sphagnum majus.
leaf, inner surface. 75. Cells of branch leaf, outer surface.
S

phagnum annulatum var. porosum. 76. Cells of branch leaf, inner surface. 77. Cells of
branch leaf, outer surface.

phagnum subsecundu
Cross-section of branch leaf. 81. Cells of branch leaf, outer surface.

71. Stem cortex. 72. Stem leaves. 73. Branch leaves. 74. Cells of branch

m var. subsecundum. 78. Cross-section of stem. 79. Branch leaves. 80.

dry, with the terminal bud large (especially prominent and noticeable when moist).
la 1-

hyaline cells not divided, without pores, fibrillose nearly throughout the leaf. Branch
leaves + spreading at the tips and undulate at the margins when dry, often somewhat
secund, about 2.3-3 mm. long, ovate, acuminate; hyaline cells with numerous rather
small to medium-sized, round pores in a median row or sometimes in 2 irregular rows
on the outer surface (occasionally also with a few scattered, small, ringed pores) and

few to several minute, ringed pores on some but not all cells on the inner surface

(easily overlooked); in section, green cells tending to be triangular, exposed on the
outer surface.

aass, we es he Pl. 69 (fig. 2), Pl. 72 (fig. 7-10), Pl. 73 (fig. 16, 18, 19), Pl. 76 (fig.
40-41). sere 76-7

Very rare. In very wet, open sites, and submerged or emergent, at the very margins of bog
lakes or in sh drainage channels in open bogs in moderately minerotrophic habitats.
Circumpolar; in North America, known from Alaska, ROR Ec District of the Canadian North-
west, British Cena Alberta, Saskatchewan, Manitoba, Ontario, Wyoming, and Michigan.

CHEBOYGAN CO.-—Livingston Bog, Nichols Bog, Mud Lake.

Distinguished from S. majus by a large terminal bud and smaller pores on the
outer surface of branch leaves, as well as few, scattered, minute, ringed pores on the
inner surface (easily overlooked except on heavy staining). Rarely the pores may be
medium-sized and unringed but in that case too numerous for S. majus. Also in
contrast to S. majus, S. annulatum var. porosum may be found prostrate, owing to
weaker stems.

d. Hyaline cells of branch leaves with numerous, small, elliptic pores arranged in a bead-like
ieee aon the commissures; green cells exposed equally on both surfaces or more broadly on
the

15. Sphagnum subsecundum Nees ex Sturm, sensu lato—Plants small to medi-
um-sized, green or tinged with orange. Wood cylinder brown; cortical cells of stem in 1
layer, thin-walled, without fibrils or pores. Stem leaves small (0.1-6 mm. long), +
triangular, blunt and cucullate at the apex, the border narrow, slightly wider below;
hyaline cells rarely divided, with fibrils none or only at extreme apex, with pores none
or rarely few to fairly numerous on the outer surface near the apex, abundant on the
inner surface. Branches in fascicles of 2-6 (2-3 spreading), the spreading ones curved,
with cortical cells in 1 layer, without fibrils, the retort cells with inconspicuous necks.
Branch leaves + curved-secund, not undulate, small (1.1-1.4 mm. long), normally ovate
(but varying from lanceolate to suborbicular), generally involute to the toothed apex,
the border of linear cells, entire; hyaline cells only slightly convex (somewhat
more so on the inner surface), on the inner surface with few or no pores on the
outer surface numerous, in rows like a string of beads along the commissures, ellip-
tic, strongly ringed (sometimes pseudopores); in section, green cells truncately
elliptic with equal exposure to trapezoidal with broader exposure on the outer surface
(and sometimes nearly included on the inner surface). Dioicous. Spores 33-36 yu, finely
granulose.—n = 1942, cf

Breen, ae of Florida, Pl. 130 (figs. oe Darlington, Mosses of Michigan, fig. 16.
Jennings, Mosses of Western Pennsylvania, (ed. 1) P ots (as S. platyphyllum, S. auriculatum, S.
susecundu, S inundatum var. auriculatum, and 2 wens), (ed. 2) Pl. 5 (as S. platvphyllum

oe oo Pl. 9, 10 (as S. inundatum var. ata Welch, Mosses of Indiana, fig.
16.- Fig. 78-

In minerotrophic habitats, in burned-over, sedgey bogs, wet sedge meadows, and floating
mats, often at the edge of water or in the moat surrounding bogs, commonly in very wet or even
submerged sites. Greenland; Labrador to Saskatchewan and Montana, south to the Gulf of Mexico;
Washington to California; Costa Rica; Europe and Asia

The outstanding character of this species complex is the bead-like arrangement of
pores along the margins of the hyaline cells of the branch leaves (and sometimes also
the stem leaves). A number of segregates can be recognized locally:

Var. subsecundum.—Plants usually yellowish to orange-brown, shiny when dry,
with a scarcely discernible terminal bud and branches of the capitulum irregularly
curved because of curved-secund leaves; cortical cells of stem in 1 layer; stem leaves
about 1mm. long and shorter than branch leaves, oblong-triangular, blunt and
cucullate, with fibrils and pores only near the apex.—Livingston Bog, Gleasons Bog,
Nichols Bog, Gates Bog, Vincent Lake, Topinabee, Little Lake 16 and Weber Lake in
Cheboygan County; Galloway Lake, Carp Lake and between Levering and Carp Lake,
and Wilderness State Park in Emmet County; Peggley Lake, Little Dollar Lake, Rexton
Lake, Hiawatha Sports Club, and Cranberry Lake in Mackinac County.

39

Var. contortum (Schultz) Htib.—Terminal bud small; branches of capitulum
curved; cortical cells of stems in 2-3 layers; stem leaves oblong-triangular, 1-1.6 mm.
long and shorter than branch leaves, with few pores on the outer surface of hyaline
cells, numerous on the inner; branch leaves 1.3-2 mm. long, not particularly crowded,
rather indistinctly curved-secund when dry.—Bois Blanc Island, Peggley Lake, and
Ryerse Lake in Mackinac County and Dow Lake in Emmet County.

Var. platyphyllum (Lindb. ex Braithw.) Card.—Plants large and coarse (especially
when wet), with limp stems, a large terminal bud, and 1-3 fat sausage-like branches at
each fascicle (or nearly simple); capitulum scarcely differentiated and not star-shaped;
pendent branches poorly differentiated and not concealing the stem; cortical cells of
stem 2-4-layered; stem leaves oblong, 1.5-3 mm. long and similar to branch leaves in
shape, size, and structure, or larger, with hyaline cells fibrillose and porose throughout,
with many pores on the outer surface and rather few pores on the inner; branch leaves
about 2-2.5 mm. long, loosely erect and dull on drying.—Weber Lake, Topinabee, and
Nichols Bog in Cheboygan County and Dow Lake in Emmet County.

Var. rufescens (Nees, Hornsch. & Sturm) Htib. (S. auriculatum Schimp.)—Plants
sometimes rather robust, with a small terminal bud and pendent branches obscuring
the stems; cortical cells of stem 1-layered; stem leaves oblong, 1.2-2.0 mm. long, more
than 4 as long as the branch leaves, concave, with hyaline cells fibrillose in the upper
Y% or more (often nearly throughout), with pores many on the outer surface, rather
numerous on the inner.—_n = 38+4.—Found in nearby Chippewa County and at Little
Lake 16 and Livingston Bog, Cheboygan County.

Var. inundatum (Russ.) C. Jens.—Cortical cells of stem in 1 layer (or irregularly
2-layered); stem leaves oblong-triangular, 0.8-1.4 mm. long, only about % as long as the
branch leaves, with hyaline cells fibrillose in the upper 4%, with pores none or few on
the outer surface, many on the inner.—n = 38+2+, ca. 42.-Found nearby in Chippewa
and Antrim Counties.

Andrews’ inclusive view of the species was perhaps overdone, as is the recent
tendency to recognize a number of technical species in spite of intergradation. In view
of several studies showing that the variations in the SUBSECUNDA are particularly
influenced by fluctuations in habitat, such as seasonal flooding, I prefer to segregate
varieties rather than species. Significant ecological variations have been related to
various taxonomic units by Aberg (Ark. Bot. 29A(1): 1-78. 1937), Jelenc (Rev. Bryol.
et Lichén. 37: 17-32. 1970), and Magdefrau & Winkler (Zeitschr. Wiss. Pfl.-physiol. 54:
174-182. 1966). Rahman (Jour. Bryol. 7: 169-179. 1973) used methods of numerical
taxonomy in defining variants in Great Britain. Jensen (Bot. Tidsskr. 13: 199-210.
1883: translated by Gravet, Rev. Bryol. 14: 33-42. 1887) analyzed the kinds of
variations which recur in various unrelated species of Sphagnum and attributed such
analogous forms to environmental factors, such as moisture, less commonly light, and
occasionally temperature:

1. Fo. immersae. Submerged plants characterized by long stems, poorly marked capitula,
widely spaced branch tufts with branches + uniform, and not ey the stem, long

eee remote from the s

2s mpactae et ae sree of cold or dry habitats, growing in low, compact tufts
with ice erect ee and imbricated leaves
3. Fo. squarrosulae de forms with robust stems and bright-green leaves with squarrose-

reading tips; a ae branches become somewhat elongated and occur at intervals
well below the stem ti

40

4. Fo. homophyliae. Plants of dry (or sometimes aquatic) laa in which the stem leaves
develop a form and structure similar to that of the branch leave

5. Fo. laxae. Plants with widely spaced leaves

6. Fo. tenellae. Plants of slender

7

. Po. falcatae. Plants with curved branches (especially at the stem tips) and correspondingly
curved-secund leaves.

All these expressions can be demonstrated especially well in the Sphagnum
subsecundum complex. Isophylly (see 4, above) is common here and, for example, in
the complex centered around S. capillaceum.

Plants often red; stem leaves with resorption mostly on the upper surface; perichaetial
leaves San without resorption on oa surface; green cells of branch leaves with exclusive or
eater exposure on the inner surface

16. Sphagnum fimbriatum Wils. ex J. D. Hook. & Wils.—Plants slender and rather
flexuose, pale, green or yellowish, with a very large, conic terminal bud. Wood cylinder
green; cortical cells of stem in 1-3 layers, thin-walled, without fibrils, the outer cells
with a single large pore in the upper end. Stem leaves broadly spatulate, lacerate-
fringed at the broad apex and along the sides because of extensive resorption, with a
triangular area of resorption at the middle of the base; hyaline cells divided | or more
times, without fibrils, resorbed almost entirely on both surfaces. Branches usually in
fascicles of 5 (2-3 spreading), with cortical cells in 1 layer, without fibrils, the retort
cells with inconspicuous necks. Branch leaves imbricate, small, narrowly ovate,
involute-concave, bordered by linear cells, entire except at the toothed apex; hyaline
cells convex, especially on the outer surface, on the inner surface with 4-6 large, round
pores nearly as wide as the cell, on the outer surface with more numerous, smaller,
elliptic pores near the commissures, strongly ringed in the upper half of the leaf; in
section, green cells trapezoidal with slightly broader exposure on the inner surface.
Monoicous. Spores 20-26 yu, nearly smooth to finely roughened.—n = 19+2, 19+4,
19+6-7, 38+4.

Darlington, Mosses of Michigan, fig. 17. Jennings, Mosses of Western Pennsylvania, (ed. 1) PL.
3, (ed. 2) Pl. 6. Welch, Mosses of Indiana, Fig. 17.—Fig. 82

In ela tively rel places in the open (in Chamaedaphne bogs) or in the brushy transition
betw bog and forest, in low hummocks supported by the bases of Chamaedaphne bushes;
ae a ese Greentand: Labrador to Alaska, south to Maryland, the Great Lakes,
Missouri, and California; South America; Europe and Asia.

CHEBOYGAN CO.—Reeses Bog, Mud Lake, Grass Bay, Little me 16. EMMET CO.—
Between Levering and Carp Lake, Larks Lake. MACKINAC CO.—Rexton Lak

The plants, slender and bright-green when moist, are grayish when dry, and the
large terminal bud is grayish-cobwebby because of extensive fringing of the leaves
enveloping it. The branches of the capitulum are slender and the spreading branches
below the capitulum long and stringy. As in S. girgensohnii and S. russowii, the
cortical cells of the stem are porose. The stem leaves are conspicuously fringed-lacerate
at both sides and apex and therefore bordered only at the basal corners. The plants
commonly fruit. (In arctic tundra, plants may be smaller, more compact, and tinged
with brown.)

7. Sphagnum girgensohnii Russ.—Plants usually fairly robust, green or some-
times slightly yellowish or brownish (not at all reddish). Stems woody but flexuose;
terminal bud moderately large. Wood cylinder green to brown, not at all reddish;

*See also S. quinquefarium, p. 364.

ae Ns
| / J y j \\
ra / W

I( = vi
| \ ;
\ 3 / \ \
: <
r 4 8 2
{\ A 85
PX /\
vA ‘ ‘ fp \
ey aes
| \/ \
\
> : Ad Pry,
& ie] f
\_£ Se 4 \
\ \
86 \ \
I \
89 :
Ven we
\ \ | ]
Pui:
ane \ F

Sphagnum billed atum. 82. Stem cortex. 83. Stem leaves. 84. Cells at apex of stem leaf. 85.
Branch leaves. 86. Cross-section of branch leaf. 87. Cells of branch leaf, inner surface. 88. Cells of
branch leaf, outer su me ce.

phagnum girgensohnii. 89. Stem cortex. 90. Stem leaves. 91. Branch leaves. 92. Cross-
section of eae leaf. 93. Cells of branch leaf, inner surface. 94. Cells of branch leaf, aes
surf;

cortical cells in 2-4 layers, isodiametric, thin-walled, without fibrils, the outer cells with
a single, large, round pore. Stem leaves large, broadly lingulate, slightly concave,
shortly lacerate at the broadly truncate apex, with a strong border broadened toward
the base; mid-basal cells greatly resorbed in a triangular, sieve-like area; hyaline cells
not divided (or a few near the upper margins often divided), without fibrils (except in
hemi-isophyllous forms), greatly resorbed on both surfaces. Branches in fascicles of 4-5
(2 spreading), with cortical cells in 1 layer, without fibrils, the retort cells with
conspicuous necks. Branch leaves imbricate or rarely + squarrose, mostly small,
narrowly ovate, involute, bordered by linear cells, entire except at the apex; hyaline
cells not or slightly convex on the inner surface, decidedly so on the outer, on the
inner surface with 3-5 large, rounded pores, on the outer surface with 4-12 elliptic
pores along the commissures; in section the green cells trapezoidal to triangular with
broader exposure on the inner surface. Dioicous (and rarely also monoicous). Spores
18-24 uw, very finely papillose.—n = 19+2, 19+4

42

Darlington, Mosses of Michigan, fig. 18. Jennings, Mosses of Western oo ee 2) PL.
7, Welch, Mosses of Indiana, fig. 18. Sullivant, Icones Muscorum Supplement, Pl. 5.—Fig. 89-94.

shade in bog forests or Thuja swamps; eutrophic to mesotrophic. Greenland; Labrador
eaten the Great Lakes = south to North Carolina and Tennessee; Alaska to Oregon, Idaho and
Montana; Europe and Asi

CHEBOYGAN CO.—Hermits Bog, Reeses Bog, Gorge of Carp Creek, Iron Bridge (on Carp
Creek), Bryants Bog, Livingston Bog, Mud Lake, Gates Bog, Grass Bay, Grass Lake. EMMET
CO.—Conway Bog, Stutsmanville, Wycamp Lake, Wilderness State Park. MACKINAC CO,—Hiawatha
Sports Club, Hog Island Forest Campground, Prentiss Bay, Epoufette, Rexton, Garnet, Little Dollar
Lake, Hendricks, McNamara Pond. PRESQUE ISLE CO.—Hammond Bay, Evergreen Beach.

The capitulum is conspicuously star-shaped because the spreading branches are
usually long and drooping in five rows. The stem leaves are moderately fringed across
the broad apex but entire and bordered along the sides. A triangular sieve-like area of
large, open, resorbed cells at the middle of the base of stem leaves is a striking feature
of the species and its relative, S. fimbriatum.

Cryptothallus mirabilis Malmb., a saprophytic liverwort of northern and western
Europe, should be sought in our area. It grows in woodlands in moist, acid litter,
covered by bryophytes, of which species of Sphagnum are most prominent. In Norway
it is most often associated with Sphagnum girgensohnii, whereas in Great Britain it
favors S. fimbriatum, S. palustre, and S. recurvum. It has also been found in the
absence of Sphagnum, under other bryophytes such as Pellia epiphylla, for example. It
has an endotrophic mycorrhizal relationship similar to that known for Riccardia
pinguis, which it greatly resembles except in its yellow-white color. The mycorrhizal
relationship is apparently shared by Betula (see Williams, Trans. Brit. Bryol. Soc. 5
357-366. 1950, and Dickson & Holman, Blyttia 26: 8-10. 1968).

18. Sphagnum russowii Warnst.—Plants slender to moderately robust, green or
mottled-red. Terminal bud small to moderate in size. Wood cylinder green; cortical
cells of stem in 2-4 layers, thin-walled, without fibrils, the outer cells mostly with a
single, rather large, rounded pore near the upper end. Stem leaves rather large,
oblong-lingulate (usually noticeably enlarged toward the apex), nearly flat or some-
times slightly concave above, slightly (and often inconspicuously) erose at the middle
of a broad, + rounded apex, the border often much broadened near the base; hyaline
cells mostly undivided (but some are divided in each leaf), often with traces of fibrils,
nearly all resorbed on the inner surface, mostly with membrane pleats (only rarely
with a few pores and considerable development of fibrils). Branches in fascicles of 4-5
(2 spreading), with cortical cells in a single layer, without fibrils, the retort cells with
conspicuous necks. Branch leaves imbricate, small, ovate-lanceolate, involute, bordered
by linear cells and entire except at the apex; hyaline cells very slightly convex on the
inner surface, more so on the outer, on the inner surface with 3-6 large pores nearly as
wide as the cells, on the outer surface with 4-8 elliptic pores along the commissures; in
section the green cells triangular with the base exposed on the inner surface, the apex
reaching the inner surface. Dioicous or monoicous. Spores 20-26 uw, finely granulose.—n
= 19+5-11, 38+4, 38+5-11, 38+8.

Darlington, Mosses of Michigan, fig. 19 (as S. robustum).—Fig. 95-101.

common in hummocks in open bogs and in the marginal forests; oligotrophic.
Greenland; Labrador to the Great Lakes and New York; North Carolina; western United States;
Europe and Asia.

CHEBOYGAN CO.~—Reeses Bog, Gorge of Carp Creek, Iron Bridge (Carp Creek), Gates Bog,
Mud Lake, Livingston Bog, Little Lake 16, Grass Lake. EMMET CO.—Antique Bog, Stutsmanville,

43

Carp Lake, between Levering and Carp Lake. MACKINAC CO.—Bois Blanc Island, Epoufette,
Rexton Lake, Little Dollar Lake, Scotty Bay, Prentiss Bay, McNamara Pond, Hendricks, Hog Island
Forest Campground, Hiawatha Sports Club. PRESQUE ISLE CO.—Evergreen Beach, Hammond Bay.

Sphagnum russowii is only medium-sized. The name robustum often applied to
the species is inappropriate and, fortunately, nomenclaturally unacceptable. In contrast
to S. fimbriatum and S. girgensohnii, which also have porose cells of the stem cortex,
S. russowii has only an inconspicuous terminal bud. The plants are often red-mottled;
even in green shade forms, red tinges can be found in streaks on the stem near the
capitulum or in leaves protecting the antheridia on male branches. The young spreading
branches of the capitulum are relatively coarse and + clavate. The broad, lingulate stem
leaves are slightly fringed at the middle of the apex but by no means approach the
laceration of S. girgensohnii. Confusion is much more likely with S. capillaceum var.
tenellum, which, of course, lacks pores in the stem cortex. It should be noted that the
pores are occasionally hard to demonstrate. In such cases it is helpful to remember
that the stem leaves of S. russowii have only few hyaline cells divided, whereas most of
them are divided in S. capillaceum var. tenellum.

19. Sphagnum warnstorfii Russ.—Plants slender and delicate, green or red. Wood
cylinder usually reddish (sometimes brown); cortical cells of stem in 2-4 layers,
thin-walled, without fibrils, the outer cells without pores. Stem leaves of medium size,
oblong-lingulate, slightly concave, the border strong, somewhat to strongly broadened
near the base; hyaline cells mostly 1-, rarely 2-divided without fibrils (or rarely with a
few vestigial fibrils), mostly resorbed on the inner surface, with membrane pleats.
Branches in fascicles of 3-5 (2 spreading), with cortical cells in 1 layer, without fibrils,
the retort cells with conspicuous necks. Branch leaves usually in 5 spiral ranks, slightly
recurved and often + secund at the tips when dry, small, lanceolate, involute, bordered
by linear cells; hyaline cells slightly convex on the inner surface, much more so on the
outer, on the inner surface with 2-6 pores at the ends and sometimes also at the side
corners, on the outer surface in the upper % with 2-8 very small, strongly ringed,
rounded or + elliptic pores mostly near the commissures; in section, the green cells
triangular or trapezoidal with broader exposure on the inner surface. Dioicous. Spores
17-20 y, finely papillose.—n = 19+2, 19+2-4, 19+4.

Breen, Mosses of Florida, Pl. 132 (fig. 1-7). Aria se ae of Michigan, fig.
Jennings, Mosses of Western Pennsylvania, (ed. 1) Pl. 3 & (ed. . 6 (as S. warnstorfii var.
virescens).—Fig. 102-107

calciphile, comm in open, rich fens and in bog forests (such as Thuja swamps).
eRe Labrador to ees south to the Great Lakes and Colorado; Florida; Europe and Asia.

CHEBOYGAN CO.-—Reeses Bog, Hermits Bog, Iron Bridge (Carp Creek), Livingston Bog,
Blanchard Lake, Mud Lake, Indian River, Hebron Swamp, Pine Grove State Forest Campground,
Little Lake 16, Grass Lake. EMMET CO.—Galloway Bog, Pleasantview Swamp, Wycamp Lake, Cecil
Bay, en Creek Gin ser: Conway Bog, Ramona Beach. MACKINAC CO.—Mud Lake, McNamara

ond, watha Sports Club. PRESQUE ISLE CO.—Hammond Bay, Evergreen Beach,
ee Isle ees Clinton Lake.

The plants are generally red or even purplish. Even in green shade forms, the red
branch axis can be seen showing through the leaves. The branch leaves, arranged in five
distinct rows, curved outward when dry. The exceedingly small, rounded, and strongly
ringed pores are discernible only as pepper-dots under the low power of a microscope.

44

52 errs OK : ) S

a aoe ee E Jer Sen :

eS re | = pa 7 =< 3
SOR eto ®

ODT Tes SHE |
L < LDS

45

20. Sphagnum fuscum (Schimp.) Klinggr.—Plants usually slender and delicate, in
very compact hummocks, usually brown (but occasionally, in shade, merely tinged
with brown or completely green). Wood cylinder brown; cortical cells of stem
thin-walled, without fibrils, the outer cells without pores. Stem leaves medium-sized,
lingulate, slightly concave, with a strong border, strongly broadened near the base;
hyaline cells mostly 1-, sometimes 2-divided, without fibrils, nearly resorbed on the
inner surface. Branches slender and thread-like, in fascicles of 3-5 (2 normally
spreading), with cortical cells in 1 layer, without fibrils, the retort cells with
conspicuous necks. Branch leaves imbricate or with somewhat spreading tips, small,
lanceolate, involute, bordered by linear cells and entire except at the toothed apex;
hyaline cells not or slightly convex on the inner surface, strongly so on the outer, on
the inner surface with pores at the ends and sometimes also in the side corners (24
per cell), on the outer surface with 3-8 small, elliptic pores near the commissures in the
apical part of the leaf, increasing in size toward the base; in section the green cells
triangular or rarely trapezoidal with broader exposure on the inner surface. Dioicous.
Spores 20-24 uw, finely papillose or nearly smooth.—n = 19+2, 19+3-4, 19+4.

Darlington, Mosses of Michigan, fig. 20. Jennings, Mosses of Western Pennsylvania, (ed. 2) Pl.
8.—Fig. 108-115.

Capping hummocks in older, drier, more acid parts of open bogs; distinctly oligotrophic.
Greenland; es to Alaska, south to Virginia; the Great Lakes area, Colorado, and Washington;
Europe and A

CHEBOYGAN CO.—Reeses Bog, Iron Bridge Sa ie Gleasons Bog, Livingston Bog,
Nichols Bog, Mud Lake, Little Lake 16, Weber Grass Lake, Pine Grove State Forest:
Campground. EMMET CO.—West of Pellston, eee nee Bog, Conway Bog, Bear
Creek Township, Cecil Bay, Wilderness State Park. MACKINAC CO. hen tiss Bay, Hessel, Mud
Lake, Little Dollar Lake, Peggley Lake, Hendricks, Hog Island Forest Campground, Rexton Lake.
PRESQUE ISLE CO.-Hammond Bay, Evergreen Beach, Loon Lake, Kelsey Pond, Clinton Lake.

Sphagnum fuscum is recognized by its brown color and compact growth at the
top of hummocks. Unusually dark red-brown expressions of S. capillaceum and its var.
tenellum may confuse the unwary; but even in occasional green shade forms, the stems
and wood cylinders of S. fuscum are definitely brown, not red. The species is much
like S. capillaceum var. tenellum in the strongly bordered, lingulate, nearly flat stem
leaves, with divided, efibrillose hyaline cells. Its chief character is admittedly the brown
pigment, and yet the compact hummocks and thread-like branches interwoven inside
the hummocks have a characteristic “look”? which defies description. A distinctive
habitat dignifies its claim to specific rank.

Another brownish species, S. subfulvum Sjors, has been found in nearby Luce
and Chippewa Counties. As compared with S. fuscum, the tufts are looser and, when
dry, shiny. The branch leaves are larger and broader, and the stem leaves more
obviously pointed. The border of the stem leaves is only moderately expanded at base,
and the hyaline cells are often twice divided. Another means of distinguishing S.
subfulvum is provided by much larger pores on the outer surface of branch leaves.

Sphagnum russowii. 95. Stem cortex. 96. Stem leaves. 97. Upper cells of stem leaves. 98.
Branch leaves. 99. Cr aaa of branch leaf. 100. Cells of branch leaf, inner surface. 101. Cells
of branch leaf, outer surfac

m trae 102. Stem leaves. 103. Upper cells of stem leaves. 104. Branch leaves.
105. Cross-section of branch leaf. 106. Cells of branch leaf, inner surface. 107. Cells of branch leaf,
outer surface.

Sphagnum fuscum. 108. Stem cortex. 109. Stem leaves. 110. Upper cells of stem leaves. 111.
Branch cortex. 112. Branch leaves. 113. Cross-section of branch leaf. 114. Cells of branch leaf,
inner surface. 115. Cells of branch leaf, outer surface

46

21. Sphagnum capillaceum (Weiss) Schrank (S. nemoreum Scop.)—Plants general-
ly short and compact, green or more often red or red-tinged, not shiny; capitula +
rounded. Wood cylinder reddish; cortical cells of stems in 2-4 layers, thin-walled,
without fibrils or pores. Stem leaves oblong-obovate, somewhat pointed and concave at
the apex, the border usually poorly differentiated at base; hyaline cells + fibrillose at
least in the apical part of the leaf, with many cells 1-, occasionally 2-divided, rarely
with some development of membrane pleats, usually mostly resorbed on the inner
surface, often with few to numerous large, rounded membrane gaps on the inner
surface. Branches in fascicles of 3-5 (2 normally spreading), with cortical cells in 1
layer, without fibrils, the retort cells with conspicuous necks. Branch leaves imbricate
or slightly spreading, small, lanceolate, involute, with a border of linear cells, entire
except at the apex; hyaline cells slightly convex on the inner surface, strongly so on
the outer, on the inner surface with pores at the ends in the apical part of the leaf, in
the lower side regions with other pores nearly as wide as the cells, in all about 2-5, on
the outer surface with 5-12 relatively large, elliptic pores along the commissures; in
section, green cells triangular to trapezoidal with broader exposure on the inner
surface. Usually monoicous. Spores 20-29 yu, very finely papillose-roughened.—n =
19+2, 19+4, 19+4-9.

at

Darlington, ae of Michigan, fig. 22A. Jennings, Mosses of Western Pennsylvania, (ed. 1)
Pl 4 & (ed. 2) Pl. 5 (as S. capillifolium var. viride). Welch, Mosses of Indiana, fig. 19.—Fig.
6-122.

ery common, in open or shady, relatively acid situations. Greenland; Labrador to Alaska
and south to fag th Carolina, the Great Lakes, Arkansas, Colorado, and Washington; Mexico.
Europe and A

CHEBOYGAN CO.—Reeses Bog, Hermits ae Douglas Lake, Hogback Bog, Bryants Bog,
Gleasons Bog, Gates Bog, Mud Lake, Little Lak , Grass Lake, Weber Lake, Pine Grove State
Forest Campground. EMMET CO.-—Gladys Bop, ae Creek Township, Stutsmanville, between
Levering and Carp Lake, Wilderness State Park. MACKINAC CO.—Rexton Lake, Cranberry Lake,
Hog Island Forest Campground, Hendricks, McNamara Pond, Cut River, Epoufette, Bois Blanc
Island, Mackinac Island, Prentiss Bay: PRESQUE ISLE CO.- Aanmond Bay, Clinton Lake.

The acute, more or less involute-pointed stem leaves with hyaline cells less
commonly divided and somewhat fibrillose aid in distinguishing this from the var.
aetias Stem leaves even more concave and pointed and with abundant pores and
fibri weak border indicate the var. fenerum. Actually var. tenellum, var.
aale and var. fenerum represent degrees of an isophyllous tendency and are not
as distinct as one might wish.

Var. tenellum (Schimp.) Andr. (S. rubellum Wils.)—Slender, often red plants with
+ flattened capitula. Stem leaves oblong or oblong-lingulate (with parallel sides),
rounded at the apex, nearly flat, with the border abruptly and conspicuously
broadened at base, the hyaline cells mostly 1-2-divided, not or sometimes slightly
fibrillose. Branch leaves often curved and subsecund. Generally dioicous.—n = 19+2,
19+4

Darlington, Mosses of Michigan, fig. 22B.—Fig. 123-124.

pen, distinctly acid situations, at the tops of hummocks, more characteristic of raised
bogs ra eee fens. Ranging with the species but relatively uncommon in our essentially
minerotrophic peatlands.

CHEBOYGAN CO.—Reeses Bog, Iron nae eae Creek), Bryants Bog, Livingston Bog,
Mud Lake, Gates Bog, Little Lake 16, Grass Lake. EMMET CO.—Stutsmanville, Conway Bog, Big
Stone Bay, French Farm Lake. MACKINAC CO.—Rexton Lake, Hendricks, Ozark, ae
PRESQUE ISLE CO.—Hammond Bay, Evergreen Beach.

47

)]

i)

at

x

Sphagnum capillaceum. 116. Stem cortex. 117. Stem leaves. 118. Upper cells of stem leaf.
119. Branch leaves. 120. Cross-section of branch leaf. 121. Cells of branch leaf, inner surface. 122.
Cells of branch leaf, outer surface. Var. tenellum. 123. Stem leaf. 124. Upper cells of stem leaf.
Var. tenerum. 125. Stem leaves. 126. Upper cell of stem leaf, outer surface.

The capitula are flattened and not so nicely rounded as in var. capillaceum and
var. tenerum. The flat, lingulate stem leaves rounded at the apex and usually lacking
fibrils in the hyaline cells provide the most tangible distinctions from the species
proper. The leaves are often somewhat secund when dry. The hyaline cells of the stem
leaves are more commonly divided than in S. russowit.

Osvald (Acta Phytogeog. Suecica 13: 43. 1940) said that the differences between
S. capillaceum and the var. tenellum seem less distinct in America than in Europe.
Andrews once told me the same thing.

Var. tenerum (Sull. & Lesq. ex Sull.) Crum (S. tenerum Sull. & Lesq. ex
Sull.)—Rather slender, greenish to pinkish plants with small, rounded capitula. Stem
leaves relatively large, oblong-ovate, somewhat narrowed and concave at the tip, with
the border scarcely to somewhat broadened at base and hyaline cells commonly
1-(rarely 2-)divided, fibrillose nearly throughout the leaf, with numerous large, rounded
pores on the inner surface (3-5 per cell) and more numerous elliptic pores along the
commissures on the outer surface. Branch leaves imbricate; hyaline cells bulging up to
Y% their diameters on the outer surface.—n = 19+

ay Mosses of Florida, Pl. 133 (fig. 1-7), as S. tenerum.—Fig. 125-126.—The illustrations

Dar n’s Mosses of Michigan (fig. 24a) are not very convincing, and the collections
fa pee to in the text are misidentified.

48

and more open situations, such as blueberry or tamarack-
Ce aa a . ng re ‘Eastern cay tend Bee Newfoundland and Labrador to Florida and
the Gulf States, less commonly inland, in Que w York, Ontario, oe North Carolina,

Tennessee, Missouri, and, Perce in ie yarn and California; Denmar

CHEBOYGAN CO.-—Livingston Bog, Mud Lake, Gates Bog, Pine Grove State Forest
Campground. EMMET CO.—Gladys Hee ee Bog, Stutsmanville. MACKINAC CO.-—Little
Dollar Lake, Ryerse Lake, Hessel, Prentiss

—

he plants, variable in overall Leah most often resemble var. capillaceum
in miniature, because of dainty, rounded capitula. The var. tenerum differs from the
var. capillaceum in the more isophyllous nature of the stem leaves which approach the
branch leaves in shape (being somewhat pointed and concave at the tip and not ver
strongly bordered at the base) and, more so, in structure (having hyaline cells fibrillose
nearly throughout the leaf and porose on both surfaces). The stem leaves of var.
capillaceum also show a degree of isophylly, but the fibrils are not so well or so
extensively developed, and the pores, if present at all, are large, rounded, and restricted
to the inner surface.

In extreme form, the var. temerum can scarcely be confused with the var.
capillaceum; even in less convincing forms, it is a somewhat better segregate than the
var. tenellum. The habitat preferences need investigation. (The kinds of variations
expressed by this complex are discussed under S. subsecundum, above).

22. Sphagnum subnitens Russ. & Warnst. ex Warnst.—Rather robust plants in
loose, soft mats, yellowish-brown with pale, pinkish, bluish, or purple-brown tinges,
somewhat lustrous when dry. Wood cylinder red-brown; cortical cells of stem in 3-4
layers, large and thin-walled, without fibrils or pores. Stem leaves large, oblong or
oblong-triangular, broadly acute and abruptly involute-cuspidate, the border not much
broadened at the base; hyaline cells without fibrils (or with occasional traces here and
there), mostly 1(2)-divided, often with membrane pleats, greatly resorbed on the inner
surface. Branches in fascicles of 3-5 (2 spreading), their cortical cells in 1 layer,
non-fibrillose, the retort cells with + inconspicuous necks. Branch leaves lanceolate,
involute to a narrow, toothed apex which is + spreading when dry, bordered by 2-3
rows of linear cells, without a resorption furrow; hyaline cells strongly convex on the
outer surface (bulging about 2/3 their diameter), only slightly so on the inner, on the
inner surface with pores none or very few, on the outer surface with 2-5 large, elliptic
pores at ends and corners; in section, green cells triangular or more often trapezoidal,
with broader exposure on the inner surface, the lumen triangular. Monoicous. Spores
about 22-28 yu, finely papillose.—_n = 19+2, 19+4, 19+5-8

pane

Darlington, Mosses of Michigan, fig. 23 (as S. aan Jennings, Mosses of Western
aa (ed. 1) Pl. 4, (ed. 2) PL. 5 (as S. plumulosum).—Fig. 127-133.

I g mats and also in bog forests, especially in areas subject to flooding; meso-eutrophic.
Circumpolar; Azores; en British Columbia to California; Labrador to New Jersey and
Michigan.

MACKINAC CO.—Epoufette; Peggley Lake.

Best recognized by its soft sheen and reddish to bluish tinges. The narrow,
spreading leaf tips aid in distinguishing this from S. subfulvum Sjors (fig. 134-139),
which has been collected in Luce and Chippewa Counties. S. suwbfulvum is pale-brown
(never red). The scleroderm is dark-brown. The stem leaves are 0.8-1.5 mm. long,
oblong-lingulate, and rather broadly pointed (not particularly pinched or concave-
cuspidate at the tip); and the border is moderately broadened at base; the branch leaves

/ 130

1g is 133

fs ap
Re f_] 07
a Ape
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is
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a a
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136 Vi

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666: ‘

Sphagnum subnitens. 127. Stem cortex. 128. Stem leaves. 129. Upper cells of stem leaf. 130.
Branch leaves. 131. Cross-section of branch leaf. 132. Cells of branch leaf, inner surface. 133. Cells
of branch leaf, outer surface.

Sphagnum subfulvum. 134. Stem leaves. 135. Upper cells of stem leaves. 136. Branch
leaves. 137. Cross-section of branch leaf. 138. Cells of branch leaf, inner surface. 139. Cells of

branch leaf, outer surface.

are appressed, ovate-lanceolate, and shortly and broadly pointed. S. subnitens is

variegated with yellow-brown, pinkish, or violet. Its scleroderm is red-brown. The stem

leaves are 1.3-1.7 mm. long, oblong-triangular, broadly acute and abruptly concave-

cuspidate, and the border is only slightly broadened at base. The branch leaves are
¥

lanceolate, with long, narrow tips which are somewhat spreading when dry.
A certain glossiness is found in all Sphagna with hyaline cells convex on the

outer, visible surface of branch leaves. The metallic luster of S. subnitens and S.

subfulvum is seen only in the dried condition.

Class ANDREAEOPSIDA

Protonema normally thallose. Gametophytes small, dark-brown, reddish, or
blackish, rigid and brittle, usually growing on rocks. Stems without a central strand, its
cells uniformly stereid, containing large oil globules. Leaves various in shape, mostly
concave and oblong-ovate or subpanduriform from an erect base; costa single or
lacking; cells incrassate and porose, containing oil globules, often papillose. Perichaetial
leaves often large and convolute-sheathing. Setae lacking, the sporophyte consisting of
foot and capsule, elevated at maturity by a stalk-like pseudopodium (derived from the
archegonial stalk); capsule without a peristome, dehiscing by means of 4 (or sometimes
4-8, rarely 10) longitudinal valves; spore sac surrounding the columella, not separated
from the capsule wall by air spaces; both sporogenous tissue and columella derived
from the endothecium.

A small class of 1 family and 2 genera, neither represented in the local flora, but
Andreaea rupestris Hedw. (with ecostate, fiddle-shaped leaves—fig. 140-144) and A.
rothii Web. & Mohr (with costate, narrow, tapered leaves—fig. 145-146) occur on acid
rocks in the Upper Peninsula—The genus Andreaea was named for J. G. R. Andreae,
an apothecary of Hannover and a friend of Ehrhart’s.

(Qw™«ua
\y we /}
WE < yp if
in|
f~ x \ / | |
fj \ | |
| | | |
i | i Pa A
\ |] \] | ain
Li vi .
Wey | Vf Ho |
\ \ 64 | \u
| Su Vy ey
| 7 / 145
/\ . 142
/ \ fI
/
\ / | \ | | \ Q
\ y | \ | | | x
VfL IY q
\ \ g Ae
SOROaAyOrar |

Andreaea rupestris. 140. Habit. 141. Dry capsule after dehiscence. 142. Leaves. 143. Upper
cells of leaf in surface view. 144. Upper cells of leaf in profile.
Andreaea rothii, 145. Leaves. 146. Cells at leaf shoulders.

5]

Spores of Andreaea commonly begin germination by undergoing their first
division before being shed from the capsule. Such precocious germination (similar to
that of many leafy liverworts) may have survival value for mosses of rocky exposures
where conditions for ecesis are hazardous.

Andreaea rothii has a chromosome number of n = 10, 11; A. rupestris, n = 10.

Class BRYOPSIDA

Protonema filamentous. Gametophytes consisting of erect to prostrate leafy
stems attached to the substrate by non-living, hair-like rhizoids with + oblique
cross-walls. Branches not grouped at nodes in fascicles. Leaves inserted in 2 or, more
often, many ranks, often toothed at the margins but not (or very rarely) lobed, stem
and branch leaves often + dimorphous, rarely dorsal or ventral leaves differentiated
from lateral leaves, often costate (with 1 or frequently 2 nerves of varying length).
Male and female sex organs borne in bud-like inflorescences on the same or separate
plants (sometimes mixed together in the same inflorescence); archegonia narrowly
flask-shaped; antheridia vesiculose and cucumber-shaped; paraphyses associated with
antheridia, often also with archegonia; perichaetial and perigonial leaves often greatly
differentiated. Sporophytes consisting of foot, seta, and capsule, the seta sometimes
very short and virtually lacking but usually clearly elongate at maturity, not delicate or
short-lived. The capsule dehiscing by means of a lid-like operculum (or rarely
indehiscent), the mouth generally fringed by a peristome of 1 or 2 series of teeth in
multiples of 4 (usually 16); stomata generally present at or near the base of the
capsule; spore sac surrounding a central columella and not overarching it (the columella
in Archidium lacking); both spore sac and columella derived from the endothecium.
Developing sporophyte protected by a calyptra formed by the upper part of the
archegonium which continues growth after fertilization and is eventually torn from the
vaginula and carried upward as a cap over part or all of the capsule as the seta
elongates.

FISSIDENTACEAE

Plants minute to fairly large, gregarious or tufted, mostly erect, simple or
sparsely branched, radiculose only at base. Leaves narrow, lingulate to lanceolate,
distichous and complanate, usually crowded or overlapping, seemingly split to the costa
on the adaxial side into 2 vaginant laminae which clasp the stem and often the base of
the leaf above; costa single and generally well developed (rarely lacking), ending
somewhat below the apex to shortly excurrent; cells mostly small, usually unistratose,
hexagonal or + rounded-hexagonal, rarely shortly rhomboidal, smooth to mammillose
or papillose, sometimes variously differentiated at the margins. Setae terminal or lateral,
elongate, erect or flexuose, often geniculate at base; capsules erect to inclined,
symmetric or somewhat curved, often constricted below the mouth, smooth;
operculum generally rostrate; peristome teeth 16, red or red-brown, inserted at or near
the mouth, forked in the upper 1/3-2/3 (very rarely entire or with rudimentary forks).
Spores spherical. Calyptra nearly always cucullate.—A family of about 700 species in
one genus (Fissidens) and a few species in three segregate genera.

Fissidens Hedw.

With the characters of the family._The name means split tooth, referring to the
peristome teeth which are generally forked. The equitant leaves (like a rider on

nw

horseback, divided below into vaginant laminae sheathing the stem and usually the
next leaf above and thus resembling the leaf of an /ris) provide the best means of
recognition. By the structure of the costa and other evidence, Salmon (Ann. of Bot.
13: 103-130. 1899) demonstrated that the vaginant laminae constitute the true leaf,
and the apical and dorsal laminae can therefore be regarded as outgrowths from it.
More recently, however, Robinson (Rev. Bryol. et Lichén. 37: 941-947. 1972)
considered the vaginant laminae the true leaf and the apical lamina a part of the true
leaf re-oriented vertically rather than an outgrowth from it. The spirally ridged tips of
the peristome teeth of most species are also a unique feature of the family.

1. Plants aquatic, coarse, ae robust; leaf cells in 2-3 | . F. grandifrons
1. Plants often in moist or e

en wet places but not aba: ie small to medium- sized;
cells unistratose (or ie a es in small, scattered patche
)

er of linear cells (the Barer sometimes very short and

ees leaves broadly acute and often apiculate;
8.F.

3. Border limited to the vaginant laminae of uppe

bryoides
rounded a

r and perichaetial leaves; leaves ge ly
t the apex; costa ending 2-6 cells ee the apex 7. F. obtusifolius
2. Leaves not fee by linear cells
4. Leaves coarsely and irregularly serrate toward ek apex, usually bordered by 3-5 rows of
pale cells similar in shape to the inner green c
5. Leaf cells 6-9, rarely 12 u, rounded, bulging, + aioe here and there, dark-green and
obscure; pale margin usually very distin F. cristatus

c :
5. Leaf cells 9-20 u, hexagonal, flat or somewhat convex, unistratose, not eg p
margin slightly to moderately differentiated

eee
4. Leaves subentire to crenate or evenly crenate-serrulate, not or only indistinctly paler at
the margins

6. Setae terminal; costa ending 4-11 cells below the apex 4. F. osmundoides
6. Setae lateral; costa ending in or near the apiculus

+ brown, ending in a stout, cuspidate apiculus; cells of
5.

F. prs
7. Plants dioicous; costa pellucid, ending at or near a minute, pellucid apiculus; cells o
vaginant lamina smooth or inconspicuously 2-3-papillose - bushi
|. Fissidens grandifrons Brid.—Plants aquatic, robust, rigid, dark- or blackish-
_ green, often lime-encrusted, freely and irregularly branched, 1.5-10 cm. high. Leaves
3-4.5 mm. long, crowded and overlapping, rigidly erect-spreading, scarcely altered on
drying, oblong-linear or oblong-lanceolate, bluntly acute or gradually tapered to a
narrow, rounded tip, crenulate all around, unbordered; dorsal lamina scarcely tapered,
usually not reaching the stem; costa ending near the apex, covered by short cells; cells
opaque, pluristratose (2-3-stratose near the costa above, thinner toward the margins)
irregularly rounded-hexagonal, + incrassate, bulging. Inflorescences and sporophytes not
seen.—n = 16.
Conard, How to Know the Mosses, (ed. 2) fig. 35a-b. Grout, Mosses with Hand-lens and
Wisin, fig. 28. Fig. 147-150.

Attached to limestone rocks or pebbles and submerged in running water or springs.
Circumpolar; widespread in North America, south to Guatemala.

HEBOYGAN CO.—Mill Creek, Sturgeon River, Pigeon River. EMMET CO.—Ramona Beach
MACKINAC CO.—Mackinac Island.

Distinctive in habitat, aspect, and structure. The leaves are composed of several
layers of cells making them opaque and rigid. The plants rarely fruit but apparently

reproduce by means of axillary buds which become radiculose and are readily detached
in flowing water (see Hill, Bryol. 5: 56-58. 1902).

Fissidens fontanus (B.-Pyl.) Steud. (fig. 151-153) has long, narrow, flaccid leaves
of one layer of cells. According to Mrs. Britton (Bryol 5: 83. 1902), the fruits fall off

Ke

ig

MLGEE Ss
SK

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SS

ESE
h lowe <
(Le

ss<S=

SS

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EE

we

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saa

HEE EE
ee

—afe
SSS
WS
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MOE
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SSN =S\,

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UN/148

I A Jud ti yn |
(SAILS WAAL A pk
AW H

I]

Leaf. 149. Cells at leaf apex. 150. Cross sections of

Fissidens grandifrons. 147. Habit. 148.
Fissidens fontanus. 151, Habit. 152. Leaf. 153. Cells at tips of two leaves (at different

leaf.

magnifications).

just before maturity and float on the surface, many of them quite green, with the
calyptra still attached. She illustrated such a calyptra from which a new plant had
sprouted. /. fontanus occurs in its usual habitat in southern Michigan, submerged in
brooks or in places periodically inundated, such as hardwood swamps, and has been
reported from deep waters of Lake Michigan and Lake Erie (at 18 to 34 feet) and

from bottom samples of undetermined depth in Lake Superior and Lake Ontario
t. 5: 62-63. 1966). I have, subsequent to that report, seen a

(Hiltunen, Mich.

specimen taken at 35 feet in Lake Superior.

illiam Ellison recently found a few mosses on the bottoms of lakes in the
Douglas Lake region, none at great depths: Calliergon giganteum at 1m., Scorpidium

54

scorpioides at 1-2 m., and Drepanocladus aduncus at 3 m. in Arnott Lake, Emmet Co.;
Drepanocladus exannulatus at 1m. in a ditch near Sherett Lake, Emmet Co.;
Drepanocladus aduncus up to 2m. in Lancaster Lake, Cheboygan Co.; and Fissidens
grandifrons at 6-l0m. in Lake Huron. He found no bryophytes in Douglas, Burt,
Mullett, Munro, and Vincent Lakes, all in Cheboygan Co.

Lake Tahoe, on the border between California and Nevada, is one of the deepest
lakes in the world, with a maximum depth of 1645 ft. and an average depth of 1027
ft. It is also remarkable for extreme light penetration and low primary productivity. It
is, in fact, ultra-oligotrophic. Yet Frantz & Cordone (Ecol. 48: 709-714. 1967) found
algae, mosses, and liverworts as low as 500 ft. Their ee records are given here,
with depths in feet:

MUSCI

Fissidens adiantoides, 244 A, luridum, 290
F. grandifrons, 244, 329, 400 Leptodictyum riparium, 100408
Brachythecium sp., 342 L. riparium fo, fluitans, 251
Eurhynchium s 329, 392, 400 Porothamnium bigelovii, 329, 392
Hygrohypnum sp., 400 Fontinalis nitida, 400
H. molle, 225, 283, 285, 299

HEPATICAE
Blepharostoma arachnoideum, 329, 362 Chiloscyphus fragilis, 199, 342, 362, 400

Thamnobryum alopecurum was found on a bit of limestone at a depth of 200 ft.
in Lake Geneva (Schnetzler, Bot. Centralbl. 23: 330-331. 1885). In Crater Lake,
Oregon, Hasler (Jour. Wildl. Manag. 2: 94-103. 1938) recorded Fontinalis and
Drepanocladus at “astonishing depths of 394 feet.” Depth records from other lakes are
less notable. For example, Williams (Bryol. 33: 32. 1930) and Juday (Ecol. 15: 325.
1934) reported three bryophytes—Drepanocladus fluitans var. submersus, Fontinalis
antipyretica, and Chiloscyphus rivularis, at depths of 44-66 feet in Wisconsin lakes.

Persson (Bot. Notis. 1942: 308-324. 1942) reported 21 bryophytes from the
bottoms of several lakes in northern Sweden. In one lake as many as 15 species were
found in luxuriant masses and especially around beds of water lilies (both Nymphaea
and Nuphar). An association with water lilies suggests to me a preference for
calcareous and eutrophic conditions, as does Persson’s belief that bryophytes are
favored in lakes richly sedimented owing to fluvial or glacial influences. His records,
with depths in meters, follow:

MUSCI
Fissidens adiantoides, 2-5.5 Calliergon megalophyllum, 0.5-5
Bryum pseudotriquetrum, 1-6 C. trifarium, 1-
Cinclidium Wik um, 3-5 Calliergonella cuspidata, 2-5
Meesia triquetra, 1-3 Scorpidium scorpioides, 0.5-5.5
Fontinalis Lime 1-5.5 Drepanocladus aduncus, 2.5-5
F. hypnoides, 0.5-6 D. exannulatus, 0).5-5
Eurhynchium riparioides, 1-5 .5 D. sendtneri, 0.5-5.5
Campylium helodes, 3-5 D. trichophyllus, 0.5-3
C. polygamum, 0.5-4
HEPATICAE
Riccardia sinuata, 34 C. polyanthus, 34

Chiloseyphus ee 2-5

A specimen of Scorpidium scorpioides (in herb. MICH) from the Tuktoyatuk
Peninsula in Canada’s Northwest Territories, was collected by J. H. Dickson, submerged
at 50 ft. in a Carex cordorrhiza mire

It is curious that many of the bryophytes found on lake bottoms are not
normally aquatic or amphibious.

Be)

2. Fissidens adiantoides Hedw.—Plants about 10-35 (rarely 90) mm. high. Leaves
1.5-4.5 mm. long, broadly oblong, acute or occasionally obtuse, + wavy, crenulate
below, unevenly serrate at apex, usually + bordered by 4-5 rows of paler cells; dorsal
lamina + rounded and undulate at base; costa pellucid, ending 2-3 cells below the apex,
often somewhat sinuose above; cells 10-16 4, hexagonal, often rounded at the corners,
somewhat convex or nearly plane, usually not particularly obscure, unistratose.
Dioicous. Setae lateral, 7-15 (rarely 23) mm. long, straight or flexuose, reddish;
capsules 1-1.5(2.5)mm. long, suberect to inclined, often curved, often constricted
below the mouth when dry; annulus of 1-2 rows of small, persistent cells; operculum
0.7-1 mm. long, long-rostrate, sometimes oblique; peristome teeth finely papillose in
vertical rows below, + papillose at the tips. Spores 16-26 y, green, very finely papillose.
Calyptra cucullate._n = 12, 19, 20, 21, 24

Breen, Mosses of Florida, Pl. 1 (fig. 7).—Fig. 154-155.

On r humus, rocks, logs, stumps, or bark of exposed roots or tree bases, in woods.
Europe; Greenland Alaska to Oregon and Wyoming and Labrador to Minnesota and south to the
Gulf of Mex

CHEBOYGAN CO.—Douglas Lake, Reeses Bog, Carp Creek, Gorge of Carp Creek, Iron
Bridge, eee Run, near Levering, Mill Creek, Duncan Bay, Grass Bay, Stewart Creek (Black
Lake). MET CO. Amott Lake, Maple River, is Pane Swamp, Cecil Bay, Big Stone Bay,
caus aes Bear Creek Township, Wycamp Creek. MACKINAC CO.—Bois Blanc Island,
Hendricks. PRESQUE ISLE CO.—Mast Point, Hammond Bay, Evergreen Beach, Grand Lake,
Ocqueoc Falls, Clinton Lake.

The cells of the leaf margins are only moderately differentiated from the inner
cells which are similarly plane or nearly so. In F. cristatus the inner cells are smaller,
bulging on both surfaces, and bistratose here and there, resulting in an opaque tissue
contrasting with the border of flat, translucent cells. Identification is easy in most
instances, although there is a degree of intergradation with F. cristatus, especially in
cell size. But the resemblance to F. cristatus should not be over-emphasized. Anderson
and Bryan (Rev. Bryol. et Lichénol. 25: 254-267. 1956) produced cytological evidence
of a rather remote relationship (in spite of the fact that the chromosome numbers, in
some populations, suggest a polyploid series).

3. Fissidens cristatus Wils. ex Mitt.—Plants 10-25 mm. high. Leaves 1.5-3.5 mm.
long, broadly oblong, acute to obtuse, sometimes broadly apiculate, evenly crenulate
below, unevenly serrate at apex, clearly bordered by 3-5 rows of pale cells, often
somewhat wavy; dorsal lamina abruptly rounded and undulate at base; costa pellucid,
ending 2-6 cells below the apex, often + sinuose above; cells small, 6-9 or sometimes
12 uw, obscure, rounded, bulging, irregularly bistratose. Dioicous. Setae lateral, 5-10 mm.
long, reddish; capsules 1.2-1.9 mm. long, suberect to inclined; annulus of 1-2 rows of
small, persistent cells; operculum 1-1.5mm. long, erect or oblique, long-rostrate;
peristome teeth finely papillose in vertical lines below, papillose above (the tips with
prominent projections at the articulations). Spores 14-20 y, green, finely papillose.
Calyptra cucullate.—n = 12, 12+1, 13+2, 16.

Breen, Mosses of Florida, Pl. 1 (figs. 5-6). Conard, How to Know the Mosses, (ed. 2) fig.
38a-d. Darlington, Mosses of Michigan, fig. 27. Grout, Moss Flora of North America 1: Pl. 6 left.
Jennings, Mosses of Western Pennsylvania, (ed. 1) Pl. 11, (ed. 2) Pl. 15. Welch, Mosses of Indiana,
fig. 43.—Fig. 156-158.

On soil or humus, bark of exposed roots or bases of tree, rotten wood, and rocks, in woods.
Europe, Asia Minor, and eastern Asia; Philippine Islands; eastern North America from Nova Scotia
to Minnesota and south to the Gulf of Mexico; Arizona; Aleutian Islands.

56

CHEBOYGAN CO.—Gorge of Carp Creek, Reeses Bog, Wolfs Bog, Mud Lake. EMMET
CO.—Arnott Lake, Galloway Bog, Pellston, Maple River, Pleasantview Swamp. MACKINAC
CO.—Mackinac Island, Gros Cap, Ozark, Caffey Corner. PRESQUE ISLE Co.—Rainy River Falls,
Clinton Lake.

The specific epithet refers to the differentiated leaf margin resembling a crest
(such as a rooster’s comb) made conspicuous by irregular serrations and pale cells
contrasting with the small, dark, bulging cells within.

. Fissidens osmundoides Hedw.—Plants 5-12 (rarely 40)mm. high. Leaves
1-2. i mm. long, oblong, broadly acute to obtuse or rounded, sometimes + apiculate,

apex, + sinuose above; cells 10-14(18) u, irregularly hexagonal, + thin-walled, not or
slightly bulging. Dioicous. Setae terminal, 3-7 mm. long, straight or flexuose, red;
capsules 1-1.5 mm. long, suberect or inclined; annulus of 1 row of small, persistent
cells; operculum 1.1 mm. long, erect, long-rostrate; peristome teeth finely papillose in
vertical rows below, with straight, vertical ridges at the tips. Spores 13-18 yu, green,
smooth. Calyptrae mitrate-rostrate.-n = 11, 12, 16.

Grout, Mosses with Hand-lens and Microscope, fig. 25. Welch, Mosses of Indiana, fig.
40.—Fig. 159.

, humus, rocks, or logs, in woods. ore Greenland to Alaska, south to Colorado in
the a ae ] Georgia and Arkansas in the Ea

CHEBOYGAN CO.—Reeses Bog, Iron Bridge, Mud Lake, Duncan Bay, Grass Bay. EMMET
CO.—Levering, Arnott Lake, Pleasantview Swamp, Cecil Bay, Big Stone Creek, Conway Bog.
MACKINAC CO.—Bois Blanc Island, St. Martins Point, McKays Creek, Peggley Lake, McNamara
Pond. PRESQUE ISLE CO.—Hammond Bay, Evergreen Beach, Clinton Lake.

The evenly crenulate leaf margins as well as the costa ending well below the leaf
apex adequately characterize this species. (The leaf margins are often rather pale but
by no means as well differentiated as in F. cristatus or F’. adiantoides.) The leaves of
many species of Fissidens give a neatly pinnate effect suggesting a fern frond and
accounting for such names as osmundoides, adiantoides, asplenioides, and poly-
podioides.

5. Fissidens taxifolius Hedw.—Plants small, 4-8 mm. high. Leaves 1.5-2.2 mm.
long, oblong, obtuse or broadly acute, abruptly cuspidate, + wavy, evenly crenate-
serrulate, rarely faintly pale-margined; dorsal lamina rounded and + undulate at base;
costa pellucid, disappearing in the cuspidate tip; cells 7-9(11.5) u, rounded-hexagonal,
thin-walled, bulging; cells of vaginant laminae bluntly and inconspicuously unipapillose.
Autoicous; perigonium on a short, basal branch. Setae lateral, 8-11 mm. long, reddish,
straight or flexuose; capsules 1.2-1.5 mm. long, inclined, slightly curved, constricted
below the mouth; annulus of 1-2 rows of small, persistent cells; operculum 1 mm. long,
obliquely long-rostrate; peristome teeth finely papillose (sometimes vertically papillose-
striolate below, papillose at the tips). Spores 13-19 uy, green, very finely papillose.
Calyptra cucullate.—_n = 9, 10, 12, 12+1, 24.

Breen, Mosses of Florida, Pl. 8 (figs. 6-7). as How to Know the Mosses, (ed. 2) fig.
ae peek Bais with Hand-lens and Microscope, fig. 26, on Flora of North America, 1: Pl.
8 upper left. Sara Mosses of Western Pennsylvania, re 1) Pl. 11, (ed. 2) Pl. 15. Welch, Mosses

of Indiana, oo 38 160-161.

ff

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long, with a dark, often brownish costa

growing on soil and rock. Europe, eastern Asia, and the Caribbean; British
disappearing in a stout, cuspidate apiculus, and the cells of the vaginant laminae are

Columbia and Washington and widespread in eastern North America from Nova Scotia to South

Dakota, south to Florida and Texas.
—Cecil Bay.

Mi

idens adiantoides, 154. Leaf. 155. Cells at leaf tip.
issidens cristatus. 156. Habit. 157. Leaf. 158. Cells at leaf tip.

issidens subbasilaris. 164. Leaves. 165. Cells at leaf tip.

SS

F

F
The leaves are about 1.5-2.2 mm.

A. calciphile
EMMET CO.

58

inconspicuously unipapillose. These features contrast with those of F. bushii, in which
the leaves are only about 1-1.5 mm. long, with a pellucid costa ending short of the
small apiculus but similar to it in color and thus appearing to be excurrent, and the
cells of the vaginant laminae are smooth or rarely and indistinctly 2-3-papillose.

6. Fissidens bushii (Card. & Thér.) Card. & Thér.—Small plants 4-7 mm. high.
Leaves 1-1.5 mm. long, oblong, broadly acute to rounded-obtuse, abruptly narrowed to
a small, pellucid apiculus, crenate-serrulate all around, serrulate at the tip, often faintly
pale-margined; dorsal lamina abruptly narrowed or rounded at the base; costa pellucid,
ending near the apiculus but similar to it in color and thus appearing excurrent; cells
7-9 wu, green and somewhat obscure, irregularly bistratose, rounded-hexagonal, thin- or
firm-walled, distinctly bulging, those of the vaginant laminae not or rarely very
indistinctly roughened by 2-3 low, blunt papillae. Dioicous; both male and female buds
basal. Setae 6-10mm. long, red-yellow, erect or flexuous; capsules 1 mm. long,
suberect to inclined, + curved, somewhat constricted below the mouth when dry;
annulus of 1-2 rows of small,persistent cells; operculum 1 mm. long, obliquely rostrate;
peristome teeth finely papillose in vertical rows below, indistinctly spiral-thickened
above. Spores 14-18 y, finely papillose. Calyptrae cucullate.

Conard, How to Know the Mosses, (ed. 2), fig. 39c. Grout, Moss Flora of North America,
vol. 1, Pl. 20. —Fig. 162-163.

On soil and occasionally on rocks, in open woods or in exposed, disturbed places; a
ci eas Widespread in eastern North America from Maine and Ontario south to Florida and
Texas

CHEBOYGAN CO.-—E. of Mullet Lake,

The costa ends somewhat short of the small apiculus but appears to be joined to
it because of a similar whitish-pellucid color. In Fissidens taxifolius, by contrast, a
dark-green or brownish costa nearly fills the stout, cuspidate apiculus and appears to
be excurrent.

Though not likely to be confused with this, or any of the local species, Fissidens
subbasilaris Hedw. (fig. 164-165), found only once in southern Michigan but widely
distributed elsewhere in eastern North America, deserves mention. The plants are small

d dark-green, and the upper portion of the costa is covered by short, green cells and
therefore appears dark and dull, even as viewed with a lens.

7. Fissidens obtusifolius Wils.—Small plants. Leaves 1.5-3.5 mm. long, oblong-
lingulate, broadly rounded at the apex, subentire, the perichaetial leaves bordered by
linear cells, sometimes very shortly and indistinctly but sometimes nearly the length of
the vaginant laminae; dorsal lamina broad, tapered or ending rather abruptly near the
leaf base; costa ending well below the apex; cells smooth, very irregularly hexagonal,
thin-walled to + incrassate. Dioicous. Setae terminal, 3mm. long, rather stout, +
flexuose; capsules 0.5-0.7 mm. long, erect; annulus of 1-2 rows of small, persistent
cells; operculum conic, 3 mm. long; peristome teeth finely papillose below, spirally
aici above. Spores 16-18 wu, greenish, very finely papillose.—n = 6

en, Mosses of Florida, Pl. 5 (fig. cin Conard, How to Know the Mosses, (ed. 2) fig.
36a-b, cats Mosses with Hand- lens and Microscope, fig. 24, Moss Flora of North America 1: Pl.
8, left middle. Jennings, Mosses of Western a (ed. 1) Pl. 10, (ed 2) Pl. 14. Welch,
Mosses of Indiana, fig. 35.—Fig. 166-167

On wet limestone (and perhaps on other kinds of oo as well) in or along streams. Arizona,
Michigan, Minnesota, and Indiana. Endemic to North Am

MACKINAC CO.—Bois Blanc Island.

Fissidens obtusifolius. 166. Habit. 167. Upper cells of leaf.
Fissidens exilis, 168. Habit. 169. Cells at leaf tip. 170. Cells at base of vaginant lamina.

The small, very flat, shiny plants with rounded leaf tips can be taken for some
kind of leafy liverwort. The species has not yet been found in our area, although it
occurs along the Tahquamenon River in Luce County. Material previously reported
from Mackinac Island as the var. apiculatus Grout is acutely F. exiguus (a form of F.
bryoides with leaves bordered only toward the base). The var. apiculatus differs from
the typical variety in having leaves with a longer costa, broadly acute and sometimes
more or less apiculate apex, and smaller, more irregular cells. It has been found in
southern Michigan.

Fissidens obtusifolius is often associated with Didymodon rigidulus and Hyophila
involuta. The latter (see fig. 300-304) has not been found in our area although it may
be expected in our few wet limestone habitats..

Fissidens exilis Hedw. (fig. 168-170), an extremely rare species, has been found
on moist soil near Grand Ledge in Eaton County. It is recognized by its small size and
bluntly serrate vaginant laminae which have a suggestion of an intramarginal border
formed of elongate inner cells. I have seen other collections from northern and central
Europe and American localities in Maryland, New York, Ohio, and Pennsylvania, where
it grows as scattered plants from an abundant protonema on wet, bare clay in
disturbed places, such as roadbanks.

8. Fissidens bryoides Hedw.—Small plants 2-7 mm. high. Leaves 1-2 mm. long,
oblong to oblong-lingulate, obtuse to gradually or rather abruptly acute, sometimes
cuspidate; margins typically bordered by hyaline, linear cells extending to the apex or

60

nearly so but occasionally limited to the vaginant laminae or extending varying
distances upward; dorsal lamina tapered, often ending above the leaf insertion; costa
ending slightly below the apex, percurrent, or frequently extending into the apiculus
sometimes appearing confluent with the border; cells (6)9-13(16) u, hexagonal,
thin-walled, smooth and + pellucid or somewhat bulging and rather obscure,

same collections, dioicous, synoicous, or autoicous (with od buds basal or
axillary). Setae terminal, 3-7 mm. long, straight or + flexuose, often abruptly bent at
the base, yellow or yellow-brown; capsules erect or + curved and inclined, constricted
below the mouth when dry; annulus of 1-2 rows of small, persistent cells; operculum
conic or conic-rostrate; peristome teeth bifid, minutely papillose below, spirally ridged
above. Spores 11-21 wu, green, very finely papillose. Calyptra cucullate.—n = 5 (also as
F. minutulus and F. viridulus), 10 (also as F. minutulus), 11 (as F. exiguus and F.
minutulus), 12.

een, Mosses of Florida, Pl. 3 (as F. bryoides, FP. viridulus & var. texanus, F. pusillus, & F.

bryoides & F. minutulus), 15 (as F. exiguus). Welch, Mosses of indiana, fig. 32 (as F. minutulus),
33, 34 (as F. exiguus).—Fig. 171-177.

n moist soil and rock, ee in or along brooks, rarely on exposed roots. Circumpolar, very
widely distributed in North Amer

Typical F. bryoides does not occur in our area. In the local flora are three recognizable
forms which could be referred to F. viridulus (Sw.) F. minutulus Sull., and F. ext ull.
CHEBOYGAN CO.—Mill Creek (“F. exiguus’’). EMME T CO, —Brutus (“F. viridulus”), Galloway
Bog, Big Stone Creek (“F. viridulus”) MACKINAC CO.—Mackinac Island, Bois Blanc Island,
McKays Creek, Gros Cap, Caffey Corner (“‘F. viridulus’’; some material from Mackinac Island is “F.
minutulus,”” verging on as viridulus’’).

I take an extremely conservative view of F. bryoides. It is possible to
demonstrate, sometimes in the same collections, apparently dioicous plants, as well as
autoicous and synoicous ones. At its best development, F. bryoides has small, rather
convex, and relatively obscure leaf cells, whereas F. viridulus usually has larger, flatter,
more pellucid cells, but there is no constancy in cell characteristics in any of the forms
included here. The relative length of the leaf border and development of the apiculus
are equally worthless because of gradation from F. bryoides (with a strong border often
confluent with a strong apiculus—fig. 171-172), to F. viridulus (with the border ending a
few cells below the apex—fig. 173), to F. minutulus (with the border slightly shorter and
with an acute or sometimes bluntly apiculate apex—fig. 174), and through F. pusillus (fig.
175) to F. exiguus (with a short border ending near the apex of the vaginant lamina
but frequently extending well above it in some leaves—fig. 176-177). Apparently
differences do exist, in habitat and times of fruiting, but the variants cannot be
distinguished by purely taxonomic means and should be investigated ecologically,
cytologically, and otherwise for biological rather than morphological differences.

Smith, studying a part of the problem, in Great Britain (Trans. Brit. Bryol. Soc.
6(1): 56-68. 1970), considered F. minutulus a synonym of F. viridulus. I certainly
agree, although the problem is geographically and taxonomically greater than that. His
paper serves as a model of techniques useful in analyzing species of variable and
overlapping characteristics.

According to W. J. Hooker (in Smith’s English Flora, 1844), “The moss which
engaged Mungo Park’s attention so much in Africa as to revive his drooping spirits

aa ———

ee a SS
fr is GERBER
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pment of the leaf border in

Fissidens bryoides, sensu lato, showing variation in the develo

175. Cells at leaf apex (“F. pusillu
of vaginant lamina (“F. exiguus’’).

when sinking under fatigue is this species,

as I have ascertained by means of original

specimens given to me by his brother-in-law, Mr. Dickson.”

oO i
5 ON
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fot 8)
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Mungo Park was an African explorer who was ca

escape after four months and made his wa
after 19 months of hardship. Sick, clothed i

n rags, and destitute, he faced a journey of

pass, and, in fact, all but shirt, trousers

, in his Travels in the Interior of Africa

2

, ‘I considered my fate as certain

hat. He wrote

62

and that I had no alternative but to lie down and perish... . At this moment, painful
as my reflections were, the extraordinary beauty of a small moss in fructification
irresistibly caught my eye. I mention this to shew from what trifling circumstances the
mind will sometimes derive consolation; for though the whole plant was not larger
than the top of one of my fingers, I could not contemplate the delicate conformation
of its roots, leaves, and capsula without admiration. Can that Being (thought 1), who
planted, watered, and brought to perfection in this obscure part of the world a thing
which appears of so small importance, look with unconcern upon the situation and
sufferings of creatures formed after his own image? Surely not! Reflections like these
would not allow me despair. I started up, and disregarding both hunger and fatigue,
travelled forwards, assured that relief was at hand.”
Stark, in his Popular History of British Mosses, moralized:

Sad, faint, and weary, on the sand
Our traveller sat him down; his hand
Cover’d his burning head.

Above, beneath, behind, around,

No resting for the eye he found;

All nature see’d as dead

One tiny tuft of Moss alone

Mantling with freshest green a stone
Fix’d his delighted gaze;

Through bursting tears of joy he smiled,
And while he raised the tendril wild,
His lips o’erflow’d with praise.

O! shall not He who keeps thee green,
Here in the waste, unknown, unseen,
Thy fellow-exile save?

He who commands the dew to feed
Thy gentle flower, can surely lead

Me from a scorching grave.

When Park returned to Africa in 1805, his luck ran out. He perished in rapids on
the Niger during an attack by natives.

Somewhat like the Fissidentaceae because of crowded and distichous leaves, are
the Bryoxiphiaceae, consisting of a single rare species, Bryoxiphium norvegicum (Brid.)
Mitt., the SWORD MOSS (fig. 178-180). It has not been found in Michigan although it
occurs in many scattered localities in the eastern United States. Because it is most
often found on ancient sandstone outcrops—in Hocking County, Ohio, and at Turkey
Run in Indiana, for example—Steere (Ecol. 18: 346-358. 1937) suggested that it may
be a preglacial and interglacial relic in scattered areas that escaped glaciation. He thus
related its distribution to that of Cordilleran disjuncts far eastward in the Keweenaw
Peninsula of Michigan and the Shickshock Mountains of the Gaspé, which Fernald
publicized as possible nunataks. (The idea has failed to gain acceptance, although the
extensive literature on the nunatak hypothesis is still interesting and worthwhile.)

SELIGERIACEAE

Plants minute and gregarious or larger and tufted, simple or forked, not
tomentose. Leaves + subulate from a narrow base; costa flat below, terete above and
often filling most of the subula, subpercurrent to excurrent; cells smooth, small and
generally short above, oblong-rhomboidal below, usually not differentiated at the basal
angles. Setae usually elongate, erect or somewhat curved; capsules mostly exserted,

63

erect and symmetric, cylindric or shortly pyriform, sometimes wide-mouthed when dry
and empty; operculum rostrate; peristome teeth usually present, 16, lanceolate, entire
or sometimes slightly perforate or split at the tips. Calyptrae mostly cucullate.

Seligeria BSG

Very small or minute, gregarious plants, erect, simple or forked. Leaves small
below, longer and more crowded above, lanceolate to linear, usually long-subulate from
a lanceolate base; costa subpercurrent to excurrent, often filling most of the subula;
cells smooth, short above, rhomboidal to rectangular or linear below, not differentiated
at the basal angles. Autoicous. Setae + elongate, straight or + curved, sometimes
cygneous, yellow or yellow-brown; capsules exserted, erect, subcylindric, subglobose, or
shortly obovoid, usually with a short neck, smooth; annulus none; operculum obliquely
rostrate; peristome teeth usually present, generally inserted below the mouth, acute or
truncate to emarginate, smooth. Spores spherical, brownish. Calyptrae cucullate,
smooth, naked, brown.—Named for Ignaz Seliger, a Silesian pastor, this genus is
remarkable for the exceedingly small size of both gametophytes and sporophytes.

Peristome lacking; setae erect or + flexuose; capsules wide-mouthed when dry and empty
1. S. donniana

Peristome present; setae usually + curved or sinuose; capsules subcylindric when dry and
empty 2. S. campylopoda

|. Seligeria donniana (Sm.) C. M.—Plants up to 1.5 mm. long. Leaves erect, not
much altered on drying, 0.2-1 mm. long (including the perichaetial leaves), linear from
a narrowly lanceolate base, acute, irregular at the margins; costa filling most of the
fleshy subula; cells green and subquadrate to short-rectangular above, pale and
oblong-linear below. Perichaetial leaves 0.7-1 mm. long, ovate-lanceolate, serrulate at
the shoulders. Setae 1.5-3mm. long, yellow, erect or + flexuose; capsules
0.35-0.55 mm. long, shortly obovoid, hemispheric or wide-mouthed when dry and
empty; operculum 0.15-0.35 mm. long; peristome none. Spores 8-11 u, smooth.—n =
12+1

Grout, Moss Flora of North America, 1: Pl. 35 upper left.—Fig. 181-182.

On calcareous rock, in moist, shaded places, generally on cliffs. Europe and northern Asia;
Alberta, Michigan, and Ontario.

MACKINAC CO.—Mackinac Island.

In spite of its extremely small size, this rare species is easily recognized by its
wide-mouthed capsules lacking a peristome.

2. Seligeria campylopoda Kindb. ex Macoun & Kindb.—Plants 1-2 mm. high.
Leaves erect or erect-spreading and + curved when dry, spreading when moist,
0.7-1mm. long (including perichaetial leaves), linear or linear-lanceolate (not
setaceous), gradually and broadly acuminate, blunt or sometimes acute; margins entire
or + irregular above, often + recurved below; costa not filling the subula, percurrent or
ending well below the apex; cells small and incrassate, rounded-quadrate or shortly
oblong above, usually short below, rectangular or oblong-lienar near the insertion.
Perichaetial leaves ovate-lanceolate, gradually or rather abruptly acuminate. Setae
yellow, 1.5-3mm. long, usually + curved (sometimes arcuate or somewhat sinuose),
sometimes nearly erect; urn 0.45-0.6 mm. long, subcylindric, brownish (yellow when

Seligeria
Seliger

moist); peristome inserted below the mouth, lanceolate, acute or truncate, brownish or
red-brown, 130-170 u high. Spores 9-10 4, smooth or nearly so.

Conard, How to Know the Mosses, (ed. 2) fig. 107a-c. Grout, Moss Flora of North America,
1: Pl. 35A.—Fig. 183-184.

On limestone, particularly on cliffs but also frequently on pebbles, in dry or moist, shaded
places. Northern Europe; British Columbia, Washington, Alberta, and Montana and in the East from
New Brunswick through the Great Lakes, south to Iowa and Pennsylvania.

CHEBOYGAN CO.—Grapevine Point, Colonial Point, Mill Creek. EMMET CO.—Near Mack-
inaw City, Big Stone Bay. MACKINAC CO.—Bois Blanc Island, Mackinac Island, Gros Cap.

Only a bryologist is likely to see the tiny Seligerias, and yet they can be easily
found by the informed collector in moist, calcareous habitats, such as shaded cliffs,
especially in the Great Lakes-St. Lawrence area and the northern Rocky Mountains. S.

65

compylopoda is rather common in our area. A huge glacial erratic boulder along the
road at Big Stone Bay used to present a vulgar display of thousands of plants of S.
campylopoda, but a sign attracting tourists to a rock millions of years old soon
brought doom to the colony!

Seligeria calcarea (Hedw.) BSG (fig. 185-186) has been collected in nearby
Alpena County. It differs from S$. campylopoda most notably in having capsules widest

at the mouth (especially when dry), setae straight or nearly so when moist, and leaf
cells firm- but not thick-walled.

Elsewhere, in the Upper Peninsula, are S. recurvata (Hedw.) BSG and S. pusilla
(Hedw.) BSG.—S. recurvata (fig. 187-188) has setae curved to cygneous when moist
and capsules subcylindric and not widest at the mouth even when dry; it differs from
S. campylopoda in having setaceous, acute leaves with the costa essentially filling the
subula and excurrent. S. pusilla (fig. 189-190) has setae erect or nearly so when moist
and capsules widest at the mouth, especially when dry and empty; it resembles S.
calcarea but has leaves gradually narrowed to a setaceous, acute subula which is not or
only slightly fleshy at the tip.

Another member of the family found in wet crevices of siliceous rocks in the
Upper Peninsula is Blindia acuta (Hedw.) BSG. The plants are larger (1-7 cm. high) and

dioicous, with swollen and conspicuous male buds, and the leaves have colored and
inflated alar cells.

DITRICHACEAE

Plants minute or rather small, gregarious or tufted. Stems erect, simple or
forked. Leaves mostly lanceolate, acuminate or subulate, rarely sheathing at base,
rarely distichous; costa single, subpercurrent to ee -excurrent; cells smooth or rarely
roughened because of thickened cross-walls, firm-walled or incrassate, subquadrate or
short-rectangular to linear, not differentiated at the basal angles or forming a marginal
border. Setae usually elongate; capsules sometimes immersed to emergent and sub
globose but usually long-exserted and + cylindric, erect to strongly inclined, often +
curved or asymmetric; operculum and annulus usually differentiated; peristome teeth
present (except in cleistocarpous genera), 16, usually variously split or perforate nearly

to the base (or rarely irregularly subdivided into 8 groups). Spores spherical. Calyptrae
cucullate (or rarely mitrate).

1, Plants minute (rarely as high as 7 mm.), ephemeral; capsules immersed; operculum an
peristome none 1. Pleuridium

1. Plants larger (generally more than 1 cm. high), persistent; capsules exserted; Pires ee
and peristome differentiated

2. Leaves distichous, sheathing at base and abruptly narrowed to a long, slender, rough
subu 2. Distichium

2. Leaves in many rows, not notably sheathing at base or abruptly narrowed to the acumen
smooth

3. Leaves subtubulose above; margins erect; cells rounded or oblong-elliptic above 5. Ditrichum

below; upper cells subquadrate [see also species

4. Plants bluish-green or glaucous, autoicous; perichaetial leaves scarcely differentiated;
capsules erect and symmetric, brownish, irregularly wrinkled-striate when dry;
peristome teeth bifid nearly to the base, the forks not thickened or connected at fee

S 4, Saelania
4.Plants not bluish or glaucous, dioicous; perichaetial leaves broadly pointed and
strongly sheathing; capsules strongly inclined, asymmetric, dark, purplish-red, sulcate;
peristome teeth bifid nearly to the base, thickened and generally connect ted at the

nodes below 3. Ceratodon

1. Pleuridium Brid.

Very small, gregarious or loosely tufted ephemerals. Stems erect, simple or
forked. Leaves usually erect-spreading or subsecund (or occasionally imbricate below a
comal tuft), scarcely altered on drying, gradually or sometimes abruptly elongate
toward the stem tip, mostly narrow and subulate, sometimes abruptly subulate from a
broad base, the lower leaves sometimes differentiated and ovate-lanceolate; costa
generally filling most of the awn and usually excurrent, at least in upper and
perichaetial leaves; cells smooth, short to linear above, oblong-rhomboidal and rec-
tangular below, not differentiated at the basal angles. Setae very short, erect; capsules
immersed, subglobose or broadly ellipsoidal to ovoid, bluntly mucronate to apiculate
or sometimes rostrate; operculum and peristome not differentiated. Spores spherical,
brownish, papillose. pe ie cucullate or rarely mitrate, smooth.—The name means at
the side because the short but curved seta of some species makes the sporophyte
appear lateral in position.

Pleuridium subulatum (Hedw.) Rabh.—Plants in loose, yellow-green or yellow-
brown tufts 3-7 mm. high. Leaves 1.5-4mm. long, loosely erect or erect-spreading,
rather gradually acuminate to subulate from an oblong or ovate base, channeled or
subtubulose above; margins erect or incurved above, entire or + sinuose-serrulate at the
shoulders (or sometimes irregularly dentate at shoulders of upper and perichaetial
leaves); costa filling most of the subula and percurrent or excurrent; cells firm-walled
and linear or linear-flexuose above, oblong-rhomboidal or linear near the shoulders,
rather laxly short- to long-rectangular at base. Autoicous and paroicous, sometimes on
the same stems, the antheridia in small axillary buds, sometimes weakly 1-2-bracteate
or even naked in axils of upper leaves. Setae 0.2-1 mm. long; capsules 0.8-1.1 xX

brown, often shiny, stomatose at extreme base. Spores 24-41 y, ae or
orange-brown as seen in mass, finely papillose. Calyptrae cucullate.—n = 13, 2

en, Mosses of Florida, Pl. 10, figs. 4-9. Conard, How to Know the Mosses, (ed. 2)
es 102d (as P. acuminatum). Darlington, Mosses of Michigan, fig. 28. Grout, Mosses ee
Hand-lens and Microscope, ei 31 (as P. subulatum, above, and P. alternifolium, bela w), Moss
Flora of North America 1: 13C, figs. 1-2, figs. 34 (as P. acuminatum). Jennings, Mosses of
Western Pennsylvania, (ed. 2) a 60. Welch, Mosses of Indiana, fig. 47.—Fig. 191-194.

A 8 sae igre (rarely found in summer), on soil in bare patches in second-year stubble
fields ee la oadsides, pastures. Europe, North Africa, Asia, and Madeira; Nova Scotia to
eo . ee . "Florida and Texas; also in Arizona.

EMMET CO.—Brutus.

Because of variable sexual conditions, the paroicous P. acuminatum Lindb. can be
included in P. subulatum. The single collection known from the Straits region is autoicous
and, therefore, clearly P. suwbulatum by any definition.)

Mrs. Britton (The Observer, vol. 5-7. 1894-96) referred to the pygmy mosses
(Pleuridium, Phascum, Bruchia, Ephemerum, and Nanomitrium) as brownies (an
American equivalent of elves and pyxies, immortalized by Palmer Cox). Our “brown-
ies’ are ephemerals of weedy habitats. They are able to complete their short life cycles
in the early spring and late fall when they escape competition from larger weeds
dominating the habitats at other times. They are commonly found in the second year
of old-field succession (the first year after cultivation) and not in subsequent years.

A natural hybrid between Pleuridium subulatum and Ditrichum pallidum (Hedw.)
Hampe has been described (from Maryland) as Pleuriditrichum marylandicum, by
Andrews and Hermann (Bryol. 62: 119-122. 1959)

67

Pleuridium palustre (Bruch & Schimp.) BSG, widely distributed in the East but
not recorded from the Great Lakes area, may be autoicous or paroicous. The calyptra
is mitrate, and the capsules are strongly apiculate or even shortly rostrate, with
stomata just below the middle

2. Distichium BSG

Slender but often elongate plants in dense, green, tomentose tufts. Leaves
distichous, with an erect, oblong, sheathing base abruptly narrowed to a rough,
spreading, linear subula; costa very rough, excurrent, filling most of the subula; lower
cells linear or oblong-linear, smooth and pellucid; upper cells small, subquadrate or
short-rectangular, green and obscure, roughened because of thickened cross-walls.
Autoicous; perigonia axillary; perichaetial leaves not notably differentiated. Setae
single, straight, elongate; capsules cylindric or ovoid-cylindric, erect and symmetric or
strongly inclined and + asymmetric, finely wrinkled when dry; annulus large, deciduous
in fragments or + persistent; operculum low-conic; peristome inserted somewhat below
the mouth, dark- or red-brown, consisting of 16 equidistant, lanceolate teeth variously
split or perforate, or of 8 groups of about 4 irregular, filiform divisions. Spores
spherical. Calyptra cucullate, smooth, entire—-The name refers to the arrangement of
leaves in two rows.

ie cylindric, erect and symmetric; spores 16-25 yu 1. D. capillaceum
Capsules ovoid-cylindric, horizontal and asymmetric; spores 32-48 yw 2. D. inelinatum

1. Distichium capillaceum (Hedw.) BSG—Plants 1-6 cm. high. Leaves (1.5)2.5-
4.5mm. long. Setae 8-18 mm. long, red or reddish-brown or sometimes yellowish;
capsules 1-2 mm. long, cylindric, erect, brown; peristome teeth 16, equidistant,
lanceolate, split or perforate nearly to the base into 2 + equal divisions, finely
papillose-roughened or nearly smooth except for faint, oblique striolations. Spores
16-25 yw, densely and finely papillose-roughened.—n = 14,

Conard, How to Know the Mosses, (ed. 2) fig. 32a-c. Darlington, Mosses of Michigan. fig. 31.
Grout, ee with Hand-lens and pees Re. fig. 34 (as Swartzia montana).—Fig. 195-197

Typical of damp, shaded, calcareous rocks and cliffs, but also found on soil or humus, and
sometimes on bark at base Ne trees or shrubs ripe swamps. In North America, from Ala ska to
Greenland, south to Colorado and Arizona in te West and New aes to Iowa in the East; Mexico
to Patagonia; New Zealand, ete and New Coe Europe and Asi

CHEBOYGAN CO.-Iron Bridge. EMMET CO.-—Cecil Bay, Big Stone Creek. MACKINAC
CO.—Mackinac Island, Ozark. PRESQUE ISLE CO.—Evergreen Beach

Very pretty, slender mosses in dense, soft, often silky tufts, easily known by
distichous leaves with shiny, white, clasping bases contrasting with dull, rough, subulate
points.

2. Distichium inclinatum (Hedw.) BSG—Plants 1.5-3 cm. high. Leaves 2-3 mm.
long. Setae (7)12-20 mm. long; capsules 1-1.5 mm. long, horizontal, ovoid-cylindric,
brown; peristome teeth 16, equidistant, lanceolate, divided by perforations into 2-3
narrow divisions, weakly oblique- -striolate. Spores 32-48 uw, densely and finely papillose-
roughened.—n = 13, 14

Grout, Moss Flora of North America 1: Pl. 18C.—Fig. 198.

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LIL / 203 aye

Pleuridium subulatum. 191. Habit. 192. Capsule with calyptra. 193. Leaf, 194. Cells at leaf
shoulder.

Distichium capillaceum, 195, Habit. 196. Leaves. 197. Cells above leaf shoulder.

Distichium inclinatum. 198. Capsule.

Ceratodon purpureus. 199. Habit. 200. Leaves. 201. Cells at margin of upper portion of leaf.
202. Young and old capsules.

Saelania glaucescens. 203. Leaves. 204. Cells at margin of upper part of leaf. 205. Dry
capsule. 206. Portion of peristome.

69

cally uncommon. On damp rocks and cliffs, also on soil or humus of sheltered banks;
cai a calciphile. Europe and Hane Alaska to Greenland, south to Montana and the Great
Lakes and eastward to Newfoundlan

CHEBOYGAN CO.-—Gorge of Carp Creek. EMMET CO.—Big Stone Creek

The shorter, horizontal, asymmetric capsules and much larger spores distinguish
this from D. capillaceum. (The cylindric and symmetric capsules of D. capillaceum are
occasionally somewhat inclined, causing possible confusion between the two species.)

3. Ceratodon Brid.

Rather small plants in dense, dull tufts, often forked. Leaves contorted when
dry, lanceolate, gradually acuminate, usually acute, sometimes + awned, keeled;
margins usually strongly reflexed to revolute, notched or irregularly serrulate near the
tip; costa subpercurrent to excurrent, prominent at back; cells subquadrate, thick-
walled, and smooth above, short-rectangular below. Dioicous; perichaetial leaves
differentiated, sheathing. Setae erect, elongate; capsules exserted, suberect and nearly
symmetric to strongly inclined and asymmetric, + strumose at base, + sulcate when
dry; annulus large and revoluble; operculum conic; peristome teeth 16, lanceolate, split
nearly to the base into 2 terete divisions which are thickened and connected at the
joints toward the base, bordered, papillose. Spores spherical. Calyptra cucullate,
smooth, entire.-The name refers to the forked peristome teeth which resemble horns
(of a goat, for example).

Ceratodon purpureus (Hedw.) Brid.—Plants 0.5-2.5 cm. high, dirty-green, yellow-
brown, or reddish, often in dense tufts. Leaves 1.8-2 mm. long, acute or sometimes +
awned; margins strongly reflexed or revolute nearly to the apex, irregularly notched or
serrulate near the tip; costa percurrent to excurrent; upper cells 7-9 uw wide. Perigonia
swollen, conspicuous; perichaetial leaves elongate, erect and sheathing, the inner leaves
obtuse or shortly and broadly pointed. Setae 8-30 mm. long, dark, purplish-red (or
rarely + yellow or orange); capsules 1.25-2 mm. long, ovoid-cylindric, inclined to
horizontal, asymmetric, strumose at base, dark-red, deeply furrowed when dry;
operculum about 0.5 mm. long. Spores 11-15 , yellow or greenish-yellow, smooth or
nearly so.—n = 13.

Conard, How to Know the ie (ed. 2) fig. 127. Darlington, Mosses of Michigan, fig. 32.
Grout, Mosses with Hand-lens and Microscope, figs. 36-37. Jennings, Mosses of Western Pennsyl-
vania, (ed. 1) Pl. 6 (as C. purpurascens), (ed. 2) Pl. 10. Welch, Mosses of Indiana, fig. 49.—Fig.

99-202.

A weed, exceedingly common on sterile soil and sometimes rock or old wood, usually
dry, open, were places, often on old roofs. Siar ae though largely replaced in RS
latitudes by closely related species, varieties, or form

CHEBOYGAN CO.—Douglas Lake, Reeses Bog, Hermits Bog, Gorge of Carp Creek, lron
Bridge (Carp Creek), Vincent Lake, Smiths Bog, Bryants Bog, Colonial Point, Ellis Township,
Inverness Township, Duncan Bay, Little Lake 16, Weber Lake. EMMET CO.—Pellston, Brutus,
Alanson, Maple River, Arnott Lake, Pleasantview Swamp, Pellston ion Stutsmanville, Carp Lake,
Wycamp Lake, Cecil Bay. MACKINAC CO.-Bois Blanc Island, Mackin c Island, St. Martins Bay,
St. Martins Point, McKays Creek, Gros Cap, Epoufette, Ozark, Pr se Bay. PRESQUE ISLE

O.—Evergreen Beach, Hammond Bay, Huron Beach, Mast Point, Presque Isle lighthouse, Ocqueoc
Falls, Rainy River Falls.

Surely no other moss causes the trouble in identification that Ceratodon
purpureus does. Karl Miiller (von Halle) said it was created to weary the bryologist! Its

70
sterile forms are protean and occasionally fool the most seasoned bryologist, because
iv

Ditrichaceae. To the practiced eye, however, the leaves present a microscopic appear-
ance which is nearly unmistakable, because the margin is strongly reflexed or revolute
nearly to the apex where it becomes irregularly notched or serrulate and also because
of the characteristic squarish, firm, smooth upper cells. From descriptions Bryoery thro-
phyllum recurvirostrum might seem similar, but its leaves have papillose upper cells and
large, lax, thin-walled basal cells.

The peristome is particularly beautiful. Its morphological development is
described by Evans and Hooker (Bull. Torrey Bot. Club 40: 97-109. 1913). The
annulus, of the revoluble type, is good for demonstration partly because the species is
always available (being common, ubiquitous, and, in fact, a rank weed even in the
ae of cities) and also fruits freely.

e many weeds, thriving in areas disturbed or even polluted by man, Ceratodon
is a rains as evidenced also by its characteristic occurrence on sea cliffs on which
birds roost or nest. It is also an early invader after fire. Its success as a pioneer on bare
soils, in — areas, for example, is partly owing to underground gemmae produced
on rhiz

eae burning, nitrogen, potassium, and iron are soon leached away, and more
acid-loving species (such as Polytrichum) will replace the original invaders, such as
Ceratodon purpureus, Funaria hygrometrica, and Marchantia polymorpha. According to
Uggla (Acta Phytogeogr. Suec. 41: 1-116. 1958), Ceratodon purpureus occurs in great
abundance in burned-over areas in northern Scandinavia, where it reaches a maximum
after some decades and then declines rapidly. (In a 1933 burn, it was bees common in
1945 and 1947 but rare in 1957.)—For further information on bur e Funaria
hygrometrica, Buxbaumia aphylla, Pleurozium schreberi and pene ig juniperinum,

elsewhere in this book.

Gates (Bryol. 49: 66-71. 1946; Papers Mich. Acad. Sci., Arts & Lett. 37: 31-45.
1951) and Duncan (Mich. Bot. 12: 167-177. 1973) followed 40 years of succession on
dry sandy soil once occupied by a laboratory building at Douglas Lake, Michigan. The
area was first available for revegetation in 1930. Ceratodon purpureus was first noted
in 1934. By 1940 it occupied 46% of the plot, and 94.5% by 1950. Polytrichum
juniperinum appeared in 1942; by 1971 it and P. piliferum had almost entirely
replaced Ceratodon and were accompanied by nine species of Cladonia. Leach (Jour.
Ecol. 19: 98-102. 1931) also found species of Polytrichum following Ceratodon on
British sands. According to Watson (Structure and Life History of Bryophytes), burned
soils in Great Britain may be pioneered by Funaria and later succeeded by Ceratodon,
Polytricha, and lichens. (Funaria, like Marchantia, favors wet soils. Ceratodon grows
equally well on wet or dry, sterile or fertile soils.

Unlike Ceratodon, most mosses are restricted in distribution, both ecologically
and geographically, even though their spores may be exceedingly numerous and easily
wafted about by winds. Ingold (Trans. Proc. Bot. Soc. Edinb. 38: 76-88. 1959)
estimated the number of spores produced by four capsules of Eurhynchium confertum
as 280,000, 480,000, 500,000, and 700,000. Kreulen (Jour. Bryol. 7(1): 61-74. 1972)
made the following computations (of spores per capsule):

Archidium a 1-20 Grimmia pulvinata 200,000
Atrichum undulatum 450,000 Hypnodendron auricomum 1,500,000
Brachythecium ane 400,000 Hypopterygium cf. tenellum 240,000
Buxbaumia aphylla 5,500,000 Leucobryum glaucum 80,000

‘eratodon purpureus 120,000 Mnium hornum 120,000
Dawsonia lativaginata 80,000,000 Ochrobryum kunzeanum 22,500
Dicranella heteromalla 80,000 Orthodontium lineare 70,000
Eurhynchium confertum 210,000 Orthotrichum cupulatum 470,000

Funaria hygrometrica 350,000 Phascum cuspidatum 5,500

Ta

Physcomitrium pyriforme 9,000 Rhacopilum spectabile 600,000
Pohlia nutans 150,000 Schistostega pennata 36,000
Polytrichum formosum 2,200,000 Tortula muralis 1,000,000
P. piliferum 1,400,000 (and, from capsules of different size

Pottia truncata 7,500 and habitat, 500,000-700,000)

It can be seen, by comparing Ceratodon with Buxbaumia, for example, that the
number of spores is not necessarily correlated with abundance of individuals or their
overall distributions. The factors controlling distributions of spore-bearing plants,
including dispersal mechanisms, are discussed by Crum (Jour. Hattori Bot. Lab. 35:
269-298. 1972).

4. Saelania Lindb.

Rather small plants, bluish-green and glaucous because of a very fine, whitish,
mealy, and filamentous covering on the leaves. Leaves somewhat contorted when dry,
erect or erect-spreading when moist, narrowly lanceolate, gradually acuminate, keeled;
margins irregularly (and often doubly) serrate in the upper %4, often on both surfaces,
usually + reflexed at least below the middle; costa percurrent or shortly excurrent,
prominent and + toothed at back; upper cells subquadrate or short-rectangular,

times elsewhere). Autoicous; perigonia terminal on well-developed branches; perichae-
tial leaves not much differentiated. Setae erect, elongate; capsules cylindric, erect and
symmetric, irregularly wrinkled-striate when dry; annulus of 2-3 rows of large, pale
cells, persistent; operculum rostrate; peristome teeth 16, inserted near the mouth, split
nearly to the base into 2 terete forks, densely papillose. Spores spherical. Calyptra
cucullate, smooth, entire.—Named for a Finnish botanist, Anders Thiodolf Saelan.

Saelania glaucescens (Hedw.) Bom. & Broth.—Rather small plants (0.4-1.5 cm.
high). Leaves 1-2.5 mm. long, + reflexed in the lower half or more; costa percurrent or
shortly excurrent; upper cells 10-11 u wide, subquadrate or short-rectangular, bistratose
in spots (with occasional cells projecting as teeth). Setae 4-13 mm. long; capsules 1.5
to 2.5mm. long, yellow or yellow-brown; operculum rostrate, , about 0.8-1 mm. long;
peristome teeth red-brown. Spores 16-20 y, finely papillose .—

Darlington, Mosses of Michigan, fig. 33. Grout, Moss Flora of North America 1: Pl. 18F.—
Fig. 203-206.

soil (often calcareous), typically on steep banks, particularly in the protection of
erence turf, frequent at roadsides, also on rocks or soil in rock crevices. Europe, Asia, New
Zealand; Greenland to Alaska and south to Arizona, Colorado, Iowa, Michigan, and New

CHEBOYGAN CO.-—Lake bluff near Bryants Bog and on bank of road on Grapevine Point
and in “Manville” on the campus of the Biological Station (all at Douglas Lake).

The remarkable bluish cast of the plants is unmistakable. The leaf structure is
similar to that of Ceratodon, although one would scarcely confuse the two genera. The
plants are pretty when growing, but dried specimens are likely to seem infected with a
mold which is actually a cobwebby covering normal to the leaves and accounting for
their waxy bloom.

5. Ditrichum Hampe

Rather small, loosely tufted plants, simple or forked. Leaves lanceolate or
ovate-lanceolate, mostly gradually acuminate or subulate, often curved and sometimes

72

secund; costa subpercurrent to excurrent, the lamina narrow above and sometimes
absent above the shoulders; upper cells subquadrate to linear, smooth (or very rarely +
papillose); lower cells usually oblong to linear. Perichaetial leaves not much differ-
entiated but sometimes enlarged and clasping at base. Setae elongate, erect or flexuose;
capsules cylindric or ellipsoidal, suberect or inclined, often curved and asymmetric and
frequently + striate or furrowed when dry and empty; annulus large, compound,
sa gues or + persistent; operculum conic or conic-rostrate; peristome inserted near

outh, the teeth 16, usually split to the low basal membrane into 2 filiform,
a ee papillose divisions, suberect, or somewhat oblique or weakly spiraled.
Spores spherical. Calyptrae cucullate, smooth, entire.-The name refers to the
peristome teeth, each usually consisting of two hair-like divisions.

Ditrichum flexicaule (Schwaegr.) Hampe—Plants in yellow-brown or dark-green,
dull or sometimes + shiny tufts 1.5-6 or more cm. high, usually + tomentose below,
often repeatedly forked, occasionally (in reduced forms) bearing stiffly erect branches
with ovate-lanceolate, erect leaves. Leaves 2-7 mm. long, erect-spreading and flexuose
(and sometimes moderately twisted at the apex when dry, sometimes contorted or
circinate), lanceolate, gradually long-acuminate, subtubulose above, usually irregularly
serrate at and occasionally below the tip, smooth at back; costa subpercurrent to
rather long-excurrent; cells smooth and incrassate throughout, rounded or oblong-
elliptic and dark-green above; basal cells variable, subquadrate to shortly rhomboidal in
reduced forms with short leaves, oblong to linear in larger plants, a few marginal cells
at the shoulders nearly always obliquely short-rhomboidal and somewhat paler forming
a suggestion of a border. Dioicous. Setae 8-25 mm. long, dark-red, somewhat flexuous;
capsules 1.5-2 mm. long, cylindric, erect and symmetric or nearly so, only slightly
wrinkled or furrowed when dry and empty, chestnut-brown; operculum 0.7-0.9 mm.
long, high-conic or conic-rostrate, blunt; peristome teeth bifid nearly to the basal
membrane, densely papillose or spiculose, light-brown at base, whitish above. Spores
9-11 uw, smooth or nearly so, yellowish in mass.

ngton, Mosses of Michigan, fig. 35. Grout, Moss Flora of North America 1: Pl. 29A,
30A nee = 207- 209.

n indicator of basic conditions extremely common in the North and in the Rock
Mountains, typically on cliffs but also common on soil; locally on calcareous sand along the shores
of Lake Michigan and Lake Huron. Circumpolar; Guatemala and Colombia; New Zealand; across
North America south to Vermont, Michigan, lowa, and Colorado.

CHEBOYGAN CO.—Duncan Bay. EMMET CO.—Cecil Bay. MACKINAC CO.—Mackinac Island,
Bush Bay. PRESQUE ISLE CO.—Mast Point, Hammond Bay, Evergreen Beach, Hoeft State Park.

This species of extraordinary variability can be recognized by small, dark upper
leaf cells with thick walls and rounded ends and by a subtle border of short, oblique
cells at the leaf shoulders. Small plants having short leaves with the costa only slightly
excurrent have been called the var. densum (BSG) Braithw.; this form frequently
produces stout, erect, somewhat reduced branches (somewhat like those of Dicranum
flagellare). Robust plants of moist habitats (especially on cliffs) with long, flexuose, +
twisted leaves with long-excurrent costae have been given a variety of names, including
var. Sterile De Not. and, most appropriately, D. giganteum Williams. These two
extremes, completely intergrading, are directly related to conditions of moisture and
exposure. All our material can be assigned to D. flexicaule, in the typical sense, or to
the var. densum.

Ditrichum lineare (Sw.) Lindb. (found in Chippewa County on roadside sand at
Trout Lake and on Sugar Island) is smaller and bears straight innovations with crowded

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Ditrichum flexicaule. 207. Habit. 208. Leaf. 209. Cells at margin of shoulders of two leaves.

Ditrichum pusillum. 210. Leaves. 211. Cells of leaf tip. 212. Cells at margin of upper median
portion of leaf. 213. Capsule. 214. Portion of peristome.

Ditrichum pallidum. 215. Leaves. 216. Capsule (moist).

and imbricate, blunt, cucullate leaves; the leaf cells are linear and often bistratose in
spots along the margins. It is probably an acidophile. Also found on Sugar Island (and
elsewhere in the state) is D. pusillum (Hedw.) Hampe, which has chronically suffered
from confusion with D. lineare. In D. lineare, the plants normally bear straight, terete
innovations with short, blunt, cucullate, appressed leaves. The leaf margins are entire or
nearly so, plane (or rarely somewhat recurved in places), and not thickened. The leaf
cells are linear. The peristome teeth are rather finely papillose and irregularly split by
perforations into two coherent divisions. In D. pusillum (fig. 210-214), the plants are
sometimes forked but do not produce especially differentiated innovations. The leaf
margins are irregularly thickened and serrulate and also recurved. The leaf cells are
subquadrate to short-rectangular. The peristome teeth, split nearly to the base into two
usually free divisions, are obliquely striolate and often rather lightly papillose as well.
Both species are found as pioneers on roadbanks of sandy or silty soil. Chromosome
numbers reported for D. lineare are n = 14, for D. pusillum, n = 13, 13+1, 14.

Ditrichum pallidum (Hedw.) Hampe (fig. 215-216), common in southern Mich-
igan and southward in oak-hickory woodlands, is best recognized by its long, slender,
yellow setae (n = 13). Ditrichum rhynchostegium Kindb. has an orange-yellow seta. It
differs also in having the costa not quite filling the leaf subula and therefore narrowly
margined by short cells, and its spores are smaller (only 11-13 #) and nearly smooth.

Trichodon cylindricus (Hedw.) Schimp. is a rare western disjunct known from
Mt. Bohemia in Keweenaw County. Like a Ditrichum in most respects, the plants are
small and slender and have leaves abruptly narrowed from a sheathing base to a
wide-spreading, linear, roughened subula largely filled by the costa.

DICRANACEAE

Plants small to large, usually in dense, often radiculose tufts. Stems erect, simple
or forked. Leaves in many rows, often falcate-secund, mostly narrowly long-lanceolate
and often subulate; costa single and usually strong, ending near the apex to excurrent:
upper cells variously shaped, smooth or rarely mammillose or papillose; lower cells
elongate, often clearly differentiated at the basal angles. Setae mostly elongate;
capsules erect or inclined, sometimes asymmetric; operculum nearly always differ-
entiated, typically + obliquely rostrate from a conic base; peristome teeth usually
present and well developed, mostly lanceolate and usually deeply split into 2(3)
divisions, mostly vertically pitted-striolate below. Calyptra cucullate.

1. Alar cells clearly differentiated
2, Leaves consisting almost entirely of costa, in cross-section showing a median layer of
green cells enclosed dorsally and ventrally by hyaline cells (and sometimes with green
cells mingled with the dorsal hyaline cells); stereids lacking 5. Paraleucobryum
osta narrower, consisting of median guide cells and 2 stereid bands 4. Dicranum
1. Alar cells scarcely differentiated
3. Neck of capsules very long and narrow (at least as long as the urn); peristome teeth
vertically barred below 2. Trematodon
3. Neck not conspicuously differentiated; peristome teeth pitted-striolate below
4. Leaves abruptly narrowed from a strongly sheathing base to a wide-spreading subula
strongly crisped and curled when dry; autoicous 3. Oncophorus
4. Leaves gradually or abruptly narrowed, mostly not sheathing at base, not much
contorted when dry; dioicous

’

1. Dicranella

1. Dicranella Schimp.

Small plants in loose, green, yellowish, or brownish tufts. Stems sparsely
radiculose below, becoming forked by innovation. Leaves usually small below, larger
and more crowded above, erect-flexuose to secund, narrowly lanceolate or subulate,
gradually tapered from a narrow, nonsheathing base or + abruptly narrowed and +
squarrose from a broad, obovate, sheathing base; costa subpercurrent to excurrent: cells
smooth, subquadrate to linear in the upper median region, larger and oblong toward
the insertion, not differentiated at the basal angles. Dioicous; perichaetial leaves usually
similar to the upper stem leaves, sometimes more sheathing at base. Setae single,
elongate, erect or flexuose; urn of capsule erect or inclined, symmetric or asymmetric,
sometimes strumose, smooth or furrowed, subglobose to oblong-cylindric; annulus
lacking or present, often compound; operculum long-rostrate, inclined or curved;
peristome teeth 16, sometimes short and irregular but usually well developed and
divided about 4 way down into 2 papillose forks, nearly always vertically pitted-
striolate below. Spores sphericalThe name meaning little Dicranum derives from
some resemblance to that genus in the appearance of the leafy plants, especially in
species with secund leaves.

1. Setae yellow; capsules suberect, furrowed when dry and empty and contracted below the
strongly oblique mouth . D. heteromalla

1. Setae reddish; capsules curved and inclined, smooth (or somewhat plicate), not notably
oblique at the mouth or contracted below it

ies

2. Leaves erect-spreading or secund, gradually narrowed from a lanceolate base to a
linear-lanceolate point, bistratose and irregularly recurved at the margins 3. D. varia

2. Leaves es -spreading or squarrose from an obovate, sheathing base ae gradually

arrowed from the shoulders to a lanceolate point; margins plane and unistrat
3. Leaves ae margins entire except at the extreme apex; median cells 7-9 wide

D. schreberiana

3. Leaves usually blunt; margins bluntly and irregularly serrulate to serrate; media an cells
-15 uw wide la. D. schreberiana var. robusta

1. Dicranella schreberiana (Hedw.) Schimp.—Plants small, yellowish, in cushions
5-20 mm. high. Leaves up to 2.5mm. long, with a flexuose-spreading to squarrose

plane and entire or sometimes + toothed at the extreme tip; costa percurrent,
becoming slightly excurrent in upper and perichaetial leaves, sometimes + toothed at
back near the tips, about 60 uw wide at base; cells distinct, thin-walled, rectangular or
rather irregular, the upper median 7-9 uw wide, 2-3:1. Seta red, 7-12(25) mm. long;
capsule curved and nodding, asymmetric, usually somewhat strumose, irregularly ribbed
or occasionally smooth, 0.8-1 mm. long; annulus none; operculum long-rostrate, curved,
equal to the urn; peristome teeth 400-455 wu high, forked less than ’-way from the tip.
Spores smooth, greenish-yellow, 14-18 w.—n = 14, 15.

Conard, How to Know the Mosses, (ed. 2) fig. 132 (as D. schreberi). Grout, Moss Flora of
North America, 1: Pl. 37 middle and lower left (D. schreberi & D. grevilleana).

On soil (often clay) on banks of roads, ditches, and streams. Circumpolar; Alaska to
New Pere south to New Jersey and the Great Lakes region

MET CO.-—Cecil Bay. MACKINAC CO.—Mackinac Island. PRESQUE ISLE CO.—Mast
Point, cee Beach.

At least some of the local material could be referred to D. ueeny (Brid.)
Schimp. because of clearly striate capsules, but the variation in this and other
characteristics are such that I cannot recognize D. grevilleana as distinct from D.
schreberiana.

la. Var. robusta Schimp. ex Braithw.—Rather robust, yellow, tufted plants
0.5-5.5, rarely 8 cm. high. Leaves 1-3 mm. long, not crowded, laxly squarrose from an
obovate, sheathing base, gradually narrowed from the shoulders and tapered to a blunt
or acute apex; margins plane, bluntly and irregularly serrulate to serrate, costa 25-50 yu
wide near the base, percurrent, nearly always + toothed at back near the tip; upper
median cells very distinct, large and rectangular, 11-15 wide, about 3:1. Seta red or
reddish-yellow, 9-11 mm. long; urn about 1 mm. long, not plicate or strumose, rather
broad in proportion to length; peristome teeth 360 uw high. Spores 13-16 4, smooth or
nearly so.

On wet soil in roadside ditches and in creek beds. Europe; British Columbia and Alberta; »
Maine to one south to Michigan and New York.

CHEBOYGAN CO.-—Douglas Lake, Gorge of Carp Creek, Point Nipigon. EMMET CO.—Maple
River at Brutus Road. MACKINAC CO.-—Bois Blanc Island. PRESQUE ISLE CO.—Hammond Bay.

Smooth capsules and leaves with gradually tapered bases, blunt points, strongly
serrate margins, and shorter, broader cells characterize the variety.

Dicranella heteromalla (Hedw.) Schimp.—Small plants in yellow to dark-green,
rather ae tufts 1 (rarely up to 4) cm. high. Leaves erect to falcate-secund, 2-3 mm

76

long, gradually narrowed from a lanceolate base to a subula largely filled by the costa;
margins erect, serrulate in the upper half of the limb; costa excurrent, about 100 p
wide near the insertion, about 1/3 the width of the leaf base; cells in 1-3 rows above
the middle of the leaf, rather short-rectangular, 4-7 X 11-14. Setae 5-15 mm. long,
yellowish, capsules 1-1.5 mm. long, inclined to horizontal or more often suberect,
cylindric, often curved, asymmetric, when dry usually furrowed and contracted below
the oblique mouth; annulus consisting of 1 row of small, persistent cells; operculum
Imm. long, long-rostrate; peristome 400-450 yu, divided about % way down. Spores
14-18 w, very minutely roughened.—n = 11, 12+1, 13, 13+2-3, 14.

See ard, How to Know the Mosses, (ed. 2) fig. 131. Darlington, Mosses of sien a fig. 38.
Gro Mosses with Hand-lens and Microscope, fig. & Plate 11. Jennings, Mosses of Western
Pennsvania, (ed. 1) Pl. 7, (ed. 2) Pl. 11. Welch, Mosses of Indiana, fig. 53, 54 (as var. eee
Fig. 217-218

soil in disturbed places, especially on roadbanks and along trails in hardwood forests,
sometimes on low mounds caused by windthrows. Circumpolar; on South America; Alaska to
California: Newfoundland to ares south to the Gulf of Mexic

CHEBOYGAN CO.-—Grapevine Point, Colonial ee Vincent Lake, Lancaster Lake, Munro
Lake, Mud Lake, Mullett Lake, Little Lake 16. EMMET CO.—Pellston Hills, Cross Village.
MACKINAC CO.—Cut River, Ozark, Hendricks. SRE SOU ISLE CO.—Lamprey peel
Station, Posen.

The setaceous leaves and the shape of the empty capsules, with somewhat
oblique furrows, tapering neck, and very oblique mouth (appearing chucked under the
chin), are distinctive.

Dicranella cerviculata (Hedw.) Schimp., known from nearby Alpena, Luce, and
Alger Counties, is characterized as follows: Small plants with leaves erect- spreading and
+ flexuose or secund, narrowly lance-subulate, acute or + blunt; margins plane, entire
or slightly serrulate at the tips; costa about 4 the leaf base; upper cells long-rectangular
(S-10:1). Setae yellowish; capsules nodding, asymmetric, strumose, becoming furrowed
when dry and empty; annulus of 1 row of cells

Dicranella subulata (Hedw.) Schimp., which has been found in the Upper
Peninsula, has stem leaves lance-subulate; costa long-excurrent; cells long-rectangular
(6-10:1); perichaetial leaves abruptly subulate from a sheathing base and flexuose-
spreading to squarrose; seta reddish; capsules inclined, asymmetric but not strumose,
furrowed; annulus compound.

3. Dicranella varia (Hedw.) Schimp.—Small plants in dull, dirty- to light-green or
yellowish tufts 6-15 mm. high. Leaves erect-spreading or + falcate-secund, about 1 mm
long below, becoming as much as 2mm. long above, gradually narrowed from a
lanceolate base to a linear-lanceolate, + keeled limb, blunt or acute at the narrow,
irregularly denticulate tip; margins bistratose and irregularly recurved above the basal
portion of the leaf, entire except at the extreme tip; costa percurrent, about 45-60 yu
wide near the insertion, 1/5-1/4 the width of the leaf base; upper median cells
oblong-linear, 5-10 X 40-50 u, mostly about 6-9:1. Setae 5-8(16) mm. long, reddish;
urn of capsule 0.7-1 mm. long, asymmetric, short-ovoid, variable in the same collec-
tions, sometimes erect but usually inclined to horizontal and curved, smooth, not
strumose, contracted below the mouth; annulus none; operculum 0.5- 0.7 mm. long,
stoutly rostrate; peristome teeth 360 A10 u long, divided about 1/3 down. Spores
14-20 p, indistinctly roughened.—n = 14,

Dicranella heteromalla, 217. Habit. 218. Leaf.
Dicranella varia, 219. Leaf. 220. Capsule. 221. Peristome tooth (taken from an unpublished
drawing Ethel Hutchinson).
matodon ambiguus. sts Habit, with dry capsule (detached, upper right).
seen flexuosa, 223. Habit
Oncophorus wahlenbergii. 224, Habit. 225. Leaf.

Grout, Mosses with Hand-lens and Microscope, Pl. 13. Jennings, Mosses of Western
Pennsylvania (ed. 1) Pl. 7, (ed. 2) Pl. 11. Welch, Mosses of Indiana, Pe 52.—Fig. 219-221.

On disturbed, calcareous soil (especially clay), on moist banks of ditches and along roads, in
open situations. Circumpolar; Alaska to Nova Scotia, south to California and the Gulf of Mexico.
Cuba, Jamaica, Haiti; Mexico and Guatemala.

CHEBOYGAN CO.-—Colonial Point, Point Nipigon, Mill Creek, Cheboygan, Duncan Bay.
MACKINAC CO.—Mackinac Island, Nunns Creek. PRESQUE ISLE CO.—Mast Point, Posen.

The short, oval, and asymmetric capsules with disproportionately large teeth
provide a means of quick identification. Microscopically this species is recognized by
the narrowly recurved margins on at least some leaves of each plant, usually more
obvious along the shoulders. The median cells are long and narrow, 6-9:1, and the
upper margins bistratose.

2. Trematodon Mx.

Plants small, gregarious to loosely tufted, light-green or yellowish. Stems erect,
usually simple. Leaves + contorted when dry, oblong-ovate and clasping at base,
gradually to abruptly narrowed to a lanceolate or subulate point; costa subpercurrent
to excurrent; cells smooth, small and obscure, subquadrate or short-rectangular, longer
at base, not differentiated at the basal angles. Setae elongate, erect, yellow; capsules
erect Or somewhat curved, cylindric, the slender, conspicuous neck as long as the urn
or longer, often strumose at base; annulus simple or compound; operculum slenderly
and obliquely long-rostrate; peristome lacking or single, with 16 undivided, perforate,
or forked teeth, vertically barred on the outer surface. Calyptra smooth, moderately
inflated.—The name refers to the perforate peristome teeth. The outstanding feature of
the genus is the long neck of the capsule and the odd sculpturing of the peristome.

Trematodon ambiguus (Hedw.) Hornsch.—Stems 4-7 mm. high. Leaves 2.5-S mm.
long, oblong and clasping at base, very abruptly contracted to a long awn, erect and
entire at the margins, blunt at the tip; costa filling the awn; cells of the leaf base lax
and yellowish, narrowly long-rectangular to rhomboidal, shorter and broader at the
shoulders. Autoicous. Setae slender, bright-yellow (5)11-20(27) mm. long; capsules
light-brown, the urn 1-2.5 mm. long, the neck usually about as long as or slightly
longer than the urn, strumose at base; operculum 1-1.5 mm. long; annulus compound
and revoluble; peristome teeth 270-310 w high, perforate or forked nearly to the low,
well-developed basal membrane, with vertical bars below and fine papillae at the pale
tips. Spores 23-30 yp, subreniform, papillose.

ard, How to Know the Mosses, (ed. 2) fig. 125e. Darlington, Mosses of Michigan, ae oe
Grout, ee with Hand-lens and Microscope, fig. 38, Moss Flora of North America 1: PI.
Jennings, Mosses of Western Pennsylvania, (ed. 2) Pl. 72.—Fig. 222.

n moist soil along roads and on banks of ditches or streams in more or less open, disturbed
situations. Europe; eastern Asia; British Columbia; Newfoundland to Michigan and New York.

CHEBOYGAN CO.—Colonial Point. MACKINAC CO.—McNamara Pond.

This species has been called the LONG-NECKED MOSS in reference to its most
striking feature. The bright yellow setae are particularly noticeable in the field.

Trematodon longicollis Mx., a more easterly species found once in southern
Ontario, is distinguished by a much longer neck of the capsule and gradually tapered
leaves with the costa not filling the subula and margins more or less recurved and
bluntly serrulate above.

The genus Bruchia might be expected in ephemeral situations such as old fields
during the winter and spring. B. flexuosa (Sw. ex Schwaegr.) C. M. (Fig. 223) has been
found in southern Michigan. It is a pygmy moss with erect capsules emergent or barely
exserted and shortly pyriform in shape because of the broad, tapered neck which is
about as long as the rest of the capsule. (The shape is suggestive of an observation
balloon.) The capsules are indehiscent, lacking an operculum and, of course, a
peristome.

3. Oncophorus (Brid.) Brid.

Plants in dense, dull, green to yellow-brown tufts, somewhat radiculose below.
Leaves usually strongly crisped when dry, lanceolate and gradually acuminate with an

th)

erect base or linear-subulate from an obovate, clasping base; margins usually + toothed;
costa narrow, subpercurrent to slightly excurrent, with 2 stereid bands; upper cells
small, subquadrate or short-rectangular, smooth, generally bistratose at least at the
margins; lower cells rectangular to linear, firm- to thick-walled, the alar cells not or
very poorly differentiated. Autoicous; perigonia axillary; perichaetial leaves not much
differentiated. Setae single, elongate, erect; urn of capsule cylindric, curved and

m
urn; peristome teeth 16, inserted at the mouth, red, lanceolate, split about 4% way
down, vertically pitted- -striolate nearly to the tips. Spores spherical. Calyptra smooth,
entire at base.—The name refers to a goiter-like swelling at the base of the capsule.

Oncophorus wahlenbergii Brid.—Plants in light or yellowish-green tufts. Leaves
(3)5-6 mm. long, strongly crisped and curled when dry, obovate or oblong and clasping
at base, not decurrent, abruptly narrowed to a long-linear (rarely broader), flexuose,
subtubulose subula; margins erect (or rarely slightly recurved), + serrulate near the tip
or as much as 4 way down, usually bistratose above (sometimes indistinctly so); costa
percurrent or somewhat excurrent, often somewhat roughened at back above; upper
cells quadrate to short-rectangular, sometimes quite thick-walled; cells of the sheathing
base pale, rectangular to linear with firm to incrassate walls; alar cells not noticeably
differentiated. Seta 8-15(21)mm. long, yellowish (sometimes becoming reddish);
capsule 1-2 mm. long, strongly inclined to horizontal or sometimes cernuous, strumose
at base, moderately furrowed when old or occasionally nearly smooth; operculum
usually about 1 mm. long; peristome conspicuously red. Spores 14-30 yu, green, smooth
or nearly so.—n = 12+2, 14.

Conard, How to Know the Mosses, (ed. 2) fig. 115. Darlington, Mosses of Michigan, fig. 42.
Grout, Mosses with Hand-lens and Microscope, fig. 35: Jennings, Mosses of Western Pennsylvania,
(ed. 1) Pl. 8, (ed. 2) Pl. 12—Fig. 224-225

Typically on moss-covered, rotten logs in moist forests. Circumpolar; Greenland to Alaska,
south to British Columbia, Colorado, Michigan, and New York.

CHEBOYGAN CO.-—Reeses Bog, Hermits Bog, Gorge of Carp Creek, Iron Bridge (Carp
Creek), Livingston Bog, Duncan Bay, 3 miles north of Cheboygan, Grass Bay. EMMET CO.—Arnott
Lake, Maple River, Pleasantview Swamp, Carp Lake, Cecil Bay, Conway Bog. MACKINAC
Co. Bois Blanc Island, Mackinac Island, St. Martins Point, Epoufette, Ozark, McNamara Pond.
PRESQUE ISLE CO.—Hammond Bay, Evergreen Beach, Clinton Lake.

A beautiful moss growing in small, green tufts. The leaves are strongly crisped
and contorted when dry, and the pale capsules, with a curious goiter at base, offer a
striking contrast to the Fuchsia-red peristome. The species is often confused with O.
virens (Hedw.) Brid., which occurs in Michigan but not in our area: Oncophorus
wahlenbergii has leaves distinctly clasping at the oblong or obovate base and abruptly
narrowed to a subtubulose acumen; the leaf margins are plane (or rarely indistinctly
revolute at the middle). O. virens has leaves erect but not clasping at base,
ovate-lanceolate, and gradually narrowed to the acumen which is keeled (and, when
dry, folded); the leaf margins are distinctly revolute near the middle

4. Dicranum Hedw.

Plants small to large, in dense, + tomentose tufts. Stems simple or forked. Leaves
generally lanceolate (rarely ovate), gradually acuminate, keeled or subtubulose, usually
secund, sometimes crisped when dry, not bordered (but occasionally bistratose at
margins); costa well developed, narrow, ending near the apex to shortly excurrent, with

80

2 stereid bands, often ridged and serrate at back; cells smooth or papillose, elongate

incrassate, and porose throughout or short and unpitted above and usually elongate
and + porose below; alar cells well differentiated, usually yellow-brown near the
margins but hyaline within. Dioicous; male plants similar to the female plants or
dwarfed and growing on the tomentum or leaves of female plants, minute and bud-like
or sometimes somewhat elongate with as many as 3 perigonial buds; perichaetial leaves
convolute-sheathing, abruptly subulate. Setae elongate, slender, erect; capsules long-
exserted, ieee = foc lacing nearly straight and suberect to curved and inclined
or horizontal, + furrowed when dry and empty, not or slightly strumose; operculum
long-rostrate; eee teeth 16, inserted at or near the mouth, lanceolate, split 1/2 or
2/3 down into 2 or rarely 3 rather unequal divisions, red-brown or brown, vertically
pitted-striolate below, papillose at the pale tips. Spores SS finely and indis-
tinctly papillose. Calyptra smooth, entire at base.—The refers to the forked

of pe eth. The ae pitted- pen peceee teeth and
differentiated alar cells are other generic features. Secund or contorted leaves are
characteristic of most of the species. One of the “FORK MOSSES’’—perhaps Dicranum
elongatum Schleich.—is reported in Franklin’s lh to the Polar Sea (Appendix,
1824) as growing in dense cushions called “WOMEN’S HEADS” by Arctic- American
natives because “when you kick them they never get out of the way.” (None of the
Dicrana are so densely tufted or so troublesome.)

1. Upper leaf cells elongate and porose; annulus n

2. Leaves wide-spreading, strongly undulate; Sent broadly recurved on 1 or ra sides;
costa with 2 serrate ridges at back; setae clustered D. polysetum
2. Leaves erect and typically falcate-secund, not or very slightly undulate; margins n
recurved; costa with 4 serrate ridges at back; setae single 5. D. scoparium
1. Upper cells short, not or indistinctly porose; annulus ae
3. Leaves fragile, bistratose above 1. D. viride
3. Leaves not fragile, unistratose (or occasionally + bistratose at margins)

4. Leaves concave or subtubulose above

5. Plants not bearing brood branches; leaves cabo) ovate at base, gradually narrowed to
a broad acumen; capsules inclined and asymm . D. spurium
é ches in axils ‘of upper leaves; leaves slender and

: branches stout, rigid and terete, with small, ovate, appressed leaves; ae cells
oo

D. flagellare
6. pie a weak and slender, with linear, spreading, greatly contorted lea es (the
ee sually consisting almost entirely of such branches); leaf cells + eee es at
2. D. montanum
4. oie + keeled above

7. Leaves not undulate; In
patches); male plants large D. fuscescens

7. Leaves usually + undulate above; margins unistratose; male plants dwarfed and resting
on the leaves or tomentum of female la

8. Leaves not much contorted when ce aes undulate above, broadly acute or
obtuse; cost aa: itl below the pea

i n ntorted when ce ae or only slightly undulate, acuminate

costa percurrent to dies excurrent

9. Leaves strongly contorted when dry; leaf cells in conspicuous longitudinal rows
separated by well-marked pellucid lines; setae mostly clustere 6. D. dr

eaves curved or somewhat crisped when dry; leaf cells less acai in ro
" setae single

margins bistratose above (sometimes indistinctly or eo i

ees
: “Bulcoron

|. Dicranum viride (Sull. & Lesq.) Lindb.—Rather small plants in dull, dark-green
to yellowish-brown, + tomentose tufts 1.5-3.5 cm. or less high. Leaves erect-spreading,
flexuose, occasionally + falcate-secund,
acuminate, endin

glassy-looking, channeled above, entire or only occasionally very slightly serrulate near
the apex; costa up to 4% the leaf base, 90-150 u wide near the base, long-excurrent

8]

completely me the deciduous leaf tip; upper cells mostly subquadrate, 8-10 u wide,
bistratose and obscure, smooth; lower cells short-rectangular, thick-walled, + cole
alar cells extending to the costa, brownish at the margins. Male and female plants
similar. Seta 10-17 mm. long; urn 2.5-3mm. long, erect and symmetric, cylindric
smooth or obscurely furrowed when dry and empty; annulus of mostly 2 irregular
rows of larger cells, persistent; operculum 1.25-2 mm. long. Spores 16-21 u

onard, How to Know the Mosses, (ed. 2) fig. 58d (as D. fulvum var. viride). Grout, Mosses
with Tandon and Microscope, fig. 48. Jennings, Mosses of Western Pennsylvania, (ed. 1 PLY
(ed. 2) Pl. 13 (as D. fulvum var. aN Welch, Mosses of Indiana, fig. 58.—Fig. 226-227

the bark of trees, saat near the base. Circumpolar; Nova Scotia to Minnesota, south
to Iowa, eee and North Caro

CHEBOYGAN CO.—Reeses Bog, Hermits Bog, Gorge of Carp Creek, Iron Bridge, Sedge
Point, Livingston Bog, Colonial Point, Mud Lake, Black Lake. EMMET CO.—Arnott Lake, Galloway
ae Pleasantview Swamp, Pellston Hills, Cross Village, Middle Village, Five Mile Creek, Conway

a Beach. MACKINAC CO.-—Bois Blanc Island, Mackinac Island, Epoufette, Ozark,
eee PRESQUE ISLE CO.—Evergreen Beach, Rainy River Falls

Dicranum fulvum Hook. has been needlessly confused with this species. It is
larger, coarser, brownish-green to yellow-brown, and grows on granitic or siliceous
rocks (and therefore not to be expected in our area). D. viride is green, grows on bark,
and has fragile leaf tips which are smooth and entire or nearly so, longer basal cells,
and a narrower costa.

Dicranum fragilifolium Lindb., a rare species found on Isle Royale and elsewhere
in the Lake Superior region, has fragile leaves which are pale green to yellow, shin
and scarcely contorted when dry, with longer, oblong-linear basal cells. The male plants
are dwarfed; and the capsules are curved and inclined.

2. Dicranum montanum Hedw.—Plants very small to moderate in size, in dense,
brown, yellow-brown or more often dark-green, dull, tomentose tufts 0.5-3 cm. high,
often bearing small, strongly crisped leaves on short, weak branches near the apex but
more frequently consisting almost entirely of such branches. Leaves not secund,
strongly crisped when dry, 2-3.5 mm. long (or in reduced forms, only 0.4-0.5 mm),
narrowly lanceolate and gradually acuminate, acute, + rough at back above, concave
but usually not particularly subtubulose above, serrulate about % down; costa
percurrent or shortly excurrent; upper cells 8-11 4, subquadrate, obscure, + mammil-
lose at back; lower cells oblong and incrassate; alar cells brown to hyaline, rather
poorly differentiated. Male and female plants similar. Seta 6-14mm. long; urn
1.5-1.75 mm. long, erect but usually slightly curved, slightly furrowed when dry;
annulus deciduous in fragments, consisting of 1-2 irregular rows of large cells;
operculum 1.2-1.5 mm. long. Spores 16-21 uw.—n = 12, 14

, Moss Flora of North America, 1: 44E. Je ae Mosses of Western Pennsylvania,
(ed. 1) Pl. - (ed. 2) Pl. 12. Welch, Mosses of es fig. 55.— 228.

common on rotten logs, stumps, and bark of trees, particularly on their bases or
exposed roots. Circumpolar; British Columbia to Newfoundland, south to Arkansas and South
Carolina; Arizona.

CHEBOYGAN CO.—Reeses eee cae Bog, Iron Bridge, Maple River, Colonial Point,
Vincent Lake, Livingston Bog, Mud Grass Bay. EMMET CO.—Arnott Lake, Maple River
Township, Pellston Hills, See is tee Village, Middle Village, Five Mile Creek, Ramona
Beach, Cecil Bay, Big Stone Creek. MACKINAC CO.-—Bois Blanc Island, Mackinac Island, St.
Martins Point, McKays Creek, Cut River, Epoufette, Ozark, Hendricks. PRESQUE ISLE CO.—Ever-
green Beach, Hammond Bay, Mast Point, Ocqueoc River, Rainy River Falls, Clinton Lake.

82

Dicranum montanum occurs as a scurfy, dull, dark-green growth on bark at the
base of trees and sometimes on rotten wood. It is related to D. flagellare, but it is
much smaller and instead of producing stout, axillary branchlets with short, appressed
leaves, it bears weak branchlets with strongly crisped and spreading, roughened leaves
differing from those of ordinary stems mainly in size. Generally whole colonies of this
moss appear to consist of such leafy branchlets or of plants reduced and resembling
hem)

3. Dicranum flagellare Hedw.—Plants in compact, dull, green or brownish tufts
(0.5)1-3(5.5) cm. high, tomentose below, nearly always bearing in the axils of upper
leaves clusters of stout, stiff, terete branchlets with minute, appressed leaves. Leaves
2-3 mm. long, falcate-secund or equally spreading and moderately crisped when dry,
lance-acuminate or sometimes ovate-lanceolate (rarely ovate), acute to obtuse (or rarely
broad and rounded at the tips), strongly subtubulose above, serrulate at margins and +
rough at back above; costa percurrent or slightly excurrent; upper cells mostly
subquadrate, 8-11 4 wide, smooth, rather thick-walled; lower cells rectangular and
elongate; alar cells well differentiated, usually brownish, scarcely extending to the
costa. Seta 10-20 mm. long; urn 2-3 mm. long, erect, straight or nearly so, irregularly
furrowed when dry; annulus of 2 irregular rows of cells, + deciduous; operculum
1.2-2 mm. long. Spores 14-16(21) w.—n = 12, 23.

Con se How to Know the Mosses, (ed. 2) fig. 119. Grout, Mosses with Hand-lens and
Microscope, fig. 46c, c’ d, Moss Flora of North America 1: Pl. 44D. Jen ae Mosses of Western
eee tod. iy, Pl. 8, (ed. 2) Pl. 12. Welch, Mosses of Indiana, fig. 56.—Fig. 229.

common in the Straits region on rotten logs and stumps, rarely on fies soil.
Circumpolar; British Columbia to Newfoundland, south to Louisiana and Florida; Mexic

CHEBOYGAN CO.—Reeses Bog, Hermits Bog, Gorge of Carp Creek, Iron Bridge, Gleasons
Bog, Rs Lake, Wolffs Bog, Livingston Bog, Mud Lake, Grass Bay, rk Lake 16, Weber Lake.
EMMET CO.—Arnott Lake, Galloway Bog, Pleasantview Sw ons Wycamp Lake, Five Mile Creek.
Tia CO.—Bois Blanc Island, McKays Creek, Cut Riv ene e, Ozark, Little Dollar
Lake. PRESQUE ISLE CO.—Lamprey Experiment Station, ees Beach, Ocqueoc River, Kelsey
Pond, Loon Lake.

Easily recognized by subtubulose leaves and axillary branchlets with short,
appressed leaves. The branchlets, too stout and stiff to be considered flagellate, as the
name suggests, are usually produced in great numbers. When too few to be seen easily,
they can be detected in the field by gently rubbing them off in the palm of one’s
hand. Similar branchlets are produced in other genera of Dicranaceae, in species of
Holomitrium and Campylopus, for example.)

4. Dicranum polysetum Sw.—Plants large, in light-green to golden, usually glossy,
densely tomentose tufts 2-15 cm. high. Leaves spreading and flexuose, rarely secund,
not much altered when dry, 8-10 mm. long, lanceolate, gradually long-acuminate,
acute, keeled and + folded when dry, usually undulate, sharply toothed on the margins
and at back of the costa about % down, with margins + broadly recurved below the
middle on 1 or both sides; costa very narrow, vanishing below the apex, with 2 serrate
ridges at back (the teeth on each ridge in 1 or rarely 2 rows); cells elongate and porose
throughout, smooth; alar cells clearly differentiated, golden-brown. Male plants
dwarfed. Setae usually clustered, 1-5(7) per perichaetium, 20-37 mm. long; um
2.5-4 mm. long, strongly curved and inclined to horizontal, asymmetric, furrowed when
dry and empty, not strumose; annulus none; operculum 2.5-4mm. long. Spores
16-27 u.—n = 10+1, 11, 12, 12+1, 13, 13+1, 14.

mpl | } } | | /232

Ir LP }]
ats I

|
|
i ||

| Oke

a

SERS
SHR,
1Yy
f aN

( &

: Se)

at FERS K

Dicranum viride. 226. Leaves. 227. Cells of upper part of leaf.

Dicranum montanum ae Been of sterile and fruiting plants.

Dicranum flagellare. 229,

Dicranum polysetum. 230. ane 231. Leaf. 232. Upper cells of leaf.

Dicranum scoparium, 233. Habit. 234. Leaves. 235. Cells of upper portion of leaf. 236.
Peristome tooth.

onard, How to Know oe bee (ed. 2) fig. 116e-f (as D. rugosum). Grout, Mosses eh
i a ns and Microscope, fig. a’ (as D. undulatum), Moss Flora of North America 1: Pl. 4
(as D. undulatum). Jennings, tee a Western Pennsylvania, (ed. 2) Pl. 61 (as D. rugosum). oe
230-232.

Very common on humus, sometimes on soil (often on thin soil or humus over rocks), in
more or le haded, dry and moist woods. Circumpolar; Yukon to Nova Scotia, south to
Washington, Missouri, ene and Nort Carolina.

84

CHEBOYGAN CO.—Reeses Bog, Hermits Bog, Carp Creek, head of Gorge of Carp Creek,
Iron Bridge, Burt Lake, Pine Point, Bryants Bog, Livingston Bog, Mud Lake, Grass Bay, Little Lake
16, Weber Lake. EMMET CO.—Arnott Lake, Pleasantview Swamp, Stutsmanville, Galloway Bog,
Cecil Late Big Stone Bay, Waugoshance Point, French Farm Lake, Conway Bog. MACKINAC
CO.-—Bois Blanc Island, Mackinac Island, St. Martins Point, McKays Creek, Prentiss Bay, Hog Island
a nee Ozark, Hendricks, Pepeley Lake. PRESQUE ISLE CO.—Presque Isle lighthouse,
Huron Beach, Mast Point, Lamprey Experiment Station, Evergreen Beach, Clinton Lake.

Our largest and showiest Dicranum. The leaves glisten because of undulations
which reflect light like a rippled mirror. The clustered sporophytes and wide-spreading
leaves with long, porose cells are among many well-marked differences between this
and D. undulatum, a bog species with undulate leaves called D. bergeri Bland. in most
books. It is unfortunate that, for nomenclatural reasons, the name D. undulatum goes
with a less common, less undulate species, and both D. undulatum and D. rugosum, so
suitably applied to this species in the past, must give place to D. polysetum.

Mrs. Britton (The Observer 6: 18. 1895) commented that the Dicrana always
seem to be fruited and abundantly so. She related this phenomenon (true of some but
not all our members of the genus) to the fact that the plants grow in dense tufts that
present an appearance of crowded abundance and also that the cycle from fertilization
to spore dispersal takes 17 to 22 months, with the result that overlapping stages of
development add to an illusion of abundance. Such an overlap is exemplified by many
mosses, of course, because of a predominance of species of perennial growth.

It goes without saying that mosses that are monoicous—with both sexes on the
same plants—have the best chance of accomplishing fertilization and forming sporo-
phytes. All the Dicrana are dioicous, but those which fruit most commonly may be
thought of as functionally monoicous, as the male plants are minute buds resting on
the tomentum or leaves of the much larger female plants. Such a dwarf-male,
quasi-autoicous condition facilitates fertilization and provides the genetic advantages of
outbreeding, lacking in a truly autoicous situation. Dwarf male plants are easily
demonstrated in Dicranum polysetum, D. scoparium, D. drummondii, D. sabuletorum,
and D. undulatum. These are the species in our flora that fruit in abundance

An unpublished dissertation by Loveland on “Sexual Dimorphism in the Moss
Genus Dicranum Hedw.” presents the view that dwarf males in Dicranum result from a
chemical influence of female plants on spores, rather than a chromosomal or genetic
factor. Wallace (Abstr. XI International Botanical Congress, p. 232. 1969, and
“Developmental Morphology and Sexual Dimorphism..., 1970) found that in
Camptothecium megaptilum the majority of male plants are reduced and epiphytic on
older leaves of female plants: Only one male plant in 200 was large and independent in
occurrence, but spores germinated in culture all developed into large plants, and dwarf
males removed from the leaves of female plants became transformed into large plants.
She concluded that there is a hormonal control of sexuality but also thought that a
slight differentiation of spores into size classes suggests incipient heterospory. Other
discussions of sexual dimorphism (and the differentiation of sex chromosomes and
anisospores sometimes associated with such dimorphism) include Allen (Proc. Amer.
Phil. Soc. 58: 289-316. 1919), Dening (Flora 30: 57-86. 1935-36), Ernst-
Schwarzenbach (Verh. Schweiz. Naturf. Ges. Solothurn 1936: 310-311. 1936 and Ann.
Bryol. 11: 46-55. 1938), Fleischer (Ber. Deutsch. Bot. Ges. 38: 84-92. 1920), Miyoshi
(Hikobia 3: 294. 1963), Khanna (The Nucleus 14: 14-23. 1971), Ramsay (Bryol. 69:
293-311. 1966), Vitt (Mich. Bot. 7: 195-202. 1968), and Woesler (Planta 24: 1-13.
1935). Epiphytic dwarf male plants have been found in a Japanese species of Mniaceae,
Mnium minutulum Besch. (Koponen, Hikobia 6: 47. 1971).—See also discussions of
sexuality in Burhynchium pulchellum and Pleurozium schreberi, elsewhere in this book.

Reviews of the genetics of bryophytes were presented by Allen (Bot. Rev. 1:
269-291. 1935, and 11: 260-287.

85

Maturation cycles in mosses present a fertile field for investigation. Published
reports indicate that mosses of temperate regions require about six to 20 months to
complete the maturation of sporophytes (most species falling in the range of seven to
15 months). Papers on phenology and especially maturation cycles include Arnell
(Bryol. 8: 41-44. 1905), Barkman (Buxbaumia 15: 21-30. 1961), Benson-Evans &
Brough (Trans. Cardiff Nat. Soc. 92: 4-23. 1966), Briggs (New Phytol. 64: 366-386.
1965), Clarke & Greene (Trans. Brit. Bryol. Soc. 6(1): 114-128. 1970 and 6(2):
278-295. 1971), Forman (Bryol. 68: 289-300. 1965), Greene (Trans. Brit. Bryol. Soc.
3: 736-745. 1960), Grimme (Hedwigia 42: 1-75. 1903), Longton (Bull. Brit. Antarct.
Surv. 10: 85-95. 1966, 27: 51-96. 1972), Longton & Greene (Phil. Trans. Roy. Soc.
London, B, 252: 295-322. 1967 and Ann. Bot. 33: 83-105. 1969), Towle (Bryol. 8:
44-45. 1905; 9: 54-56. 1906; 11: 53-54. 1908), Towle & Gilbert (Bryol. 7: 35-36.
1904), and van der Wijk (Buxbaumia 14: 25-39. 1960). Newton (Jour. Linn. Soc.
London, Bot. 65: 189-209. 1972) studied the effect of photoperiod on some aspects
of the life cycle.

5. Dicranum scoparium Hedw.—Plants usually coarse and fairly robust, in glossy,
yellowish or green or sometimes dull, dirty-green or brownish tufts (1)2-8(12) cm.
high, densely tomentose. Leaves exceedingly variable, typically falcate-secund, but
often erect-spreading or sometimes appressed, not much altered on drying, sometimes
slightly undulate, 3.5-8(12) mm. long, narrowly or broadly lanceolate, gradually
acuminate, narrowly to broadly acute, + keeled above, rarely subtubulose, typically
strongly serrate in the upper 1/3; costa 75-120 u wide near the base, ending near the
apex to slightly excurrent, typically strongly (2)4-ridged at back and serrate above;
cells smooth, elongate and porose throughout (sometimes relatively broad and ob-
scurely porose), often linear and more elongate; alar cells well differentiated, yellow-
brown, not extending quite to the costa. Male plants usually dwarfed but rarely large
and similar to the female plants. Setae single (rarely paired), 18-35 mm. long; urn
2-3.5 mm. long, + curved, suberect to horizontal, nearly smooth or somewhat furrowed
when old; annulus none; operculum 2.5-3 mm. long, often longer than the urn. Spores
14-24 w.—n = 10, 1041, 11, 12, 12+1, 13, 14.

Dale ae to Know the Mosses, (ed. 2) fig. 117a-d. aoe pr with Hand-lens and
ieee 42a, a’, 43, 44b, Moss Flora of North America 1: 46C. Jennings, Mosses of
Western eee es (ed. 1) Pl. 8, (ed. 2) pl. 12. Welch, Mosses of eee fig. 60.—Fig. 233-236.

Pike ommon throughout the Straits region on various substrata, but typically on soil in
dry t ane oist forests. Circumpolar; Australia and New Zealand; Greenland to Alaska and
south me California, Colorado, Louisiana, and Florida.

CHEBOYGAN CO.—Douglas Lake, Reeses Bog, Hermits Bog, Gorge of Carp Creek, a
Bridge, Colonial Point, Grass Bay, Weber Lake. EMMET CO.—Arnott Lake, Pleasantview Swam
Carp Lake, Wilderness State Park, Cecil Bay. MACKINAC CO.-—Bois Blanc Island, Mackinac Island,
St. Martins Point, McKays Creek, Prentiss Bay, Cut River, Epoufette, Ozark, Little Dollar Lake.
PRESQUE ISLE CO. —Presque Isle lighthouse, Mast Point, Clinton Lake.

Often called the BROOM MOSS, owing to secund leaves which give the tufts a
swept appearance. The elongate, porose leaf cells and costa with four serrate ridges at
back (or rarely and atypically only two) are characteristic features of the species. In D.
polysetum, the costa has only two ridges at back of the costa.

Although the name is occasionally useful in an ecological context, Dicranum
bonjeanii De Not. ex Lisa, in North America at least, is no more than a pigeon-hole for
convergent forms of D. scoparium and D. polysetum with rather broadly pointed,
erect, shiny leaves. Plants which could be referred there have been found on sand
among jackpines near the shore of Lake Huron at Evergreen Beach (Presque Isle Co.).

86

Large forms of D. scoparium are often confused with D. majus Sm., a species of
more oceanic climes, the Canadian Maritime Provinces, for example. It has setae in
clusters, leaves up to 12 mm. long, and upper leaf cells in bistratose streaks above,
forming indistinct, intramarginal borders of inconspicuous ridges toward the apex. It
does not have the well-developed serrate ridges at the back of the costa so
characteristic of D. scoparium.

6. Dicranum drummondii C. M.—Rather robust plants in dull, green or yellow-
brown tufts 4-8 cm. high, densely rusty-tomentose below. Leaves flexuose-spreading
and sometimes + secund, strongly contorted when dry, 5-9 mm. long, lanceolate,
gradually long-acuminate, acute, strongly keeled and + undulate above, sharply serrate
in the upper 1/2 or 2/3, plane-margined; costa 115-150 u wide just above the alar
region, subpercurrent to very slightly excurrent, serrate and densely papillose at back
above, distinctly ridged at back, usually laterally spurred; upper cells 7-10(14) u,
subquadrate or short-rectangular, small and dark, only moderately incrassate, in +
regular longitudinal rows, usually + papillose at back, some cells usually projecting as
scattered teeth at the undulations; lower cells elongate, slightly porose; alar cells
yellow-brown, not extending to the costa. Male plants dwarfed. Setae usually clustered,
1-3 per perichaetium, 20-27 mm. long; capsules 2-2.5 mm. long, horizontal, strongly
curved and asymmetric, furrowed, not strumose; annulus of 2 rows of cells, fragment-
ing; operculum 2.5-3 mm. long. Spores 14-23 yp

Conard, How to Know the Mosses, (ed. 2) fig. 60a-c. Grout, Mosses with Hand-lens and
Microscope, fig. 46b, b’, Moss Flora of North America 1: Pl. 48A (fig. 1), 50C.—Fig. 237-238

humus, rarely on rotten wood, in dry coniferous forests, such as jackpine barrens, rarely
in aspens. Endemic to North America; Alberta and Montana to Newfoundland, south to Wisconsin,
Ohio, and North Carolina.

CHEBOYGAN CO.—Reeses Bog, head of Gorge of Carp Creek, Iron Bridge, Bryants Bog.
EMMET CO.—Stutsmanville, Cecil Bay, Big Stone Bay, Conway Bog. PRESQUE ISLE CO.—Ever-
green Beach.

Unmistakable in typical form, this species may sometimes be difficult to
distinguish from forms of D. undulatum or D. sabuletorum. The clustered setae, woolly
stems, and wide-spreading leaves with points contorted and incurved when dry aid in
recognition.

7. Dicranum fuscescens Turn.—Dull, green or brownish plants in + tomentose
tufts 2-4 (rarely 10) cm. high. Leaves falcate-secund (or sometimes flexuose-spreading),
often + crisped when dry, 4-7 mm. long, narrowly lanceolate, gradually and slenderly
acuminate, acute, somewhat keeled above; margins serrulate about % down, bistratose
above (sometimes indistinctly so or only in patches); costa 100-200 u wide near the
base, shortly excurrent, usually serrulate at back about 4 way down; upper cells 8-12 u
wide, subquadrate or short-rectangular, smooth or occasionally papillose at back,
usually in + regular longitudinal rows; lower cells rectangular and usually + pitted; alar
cells yellow-brown, not extending to the costa. Male plants large. Setae 8-20 mm. long;
capsules 1.75-3 mm. long, inclined or horizontal, curved and asymmetric, + furrowed
and often somewhat strumose when dry; annulus of 1-2(3) irregular rows of large, pale
cells, + persistent; operculum 2-2.5 mm. long. Spores 16-32 u.—n = 9, 10, 10+1, 12,
24

Conard, How to Know the Mosses, (ed. 2) fig. 59a-d, 120a-d. Grout, Mosses with Hand-lens
and ren figs. 42b, b’, c, 44c, Moss Flora of North America 1: Pi. 46E. —Fig. 239-240.

Dicranum nee 246, Leaf. 247, Cells at apex of leaf.

forest species, on rotten logs or stumps and less commonly on bark of exposed roots o
tree pase Circumpolar; Greenland to Alaska, south to California, Colorado, Michigan, ee
Tenn

CHEBOYGAN CO.—Reeses Bog, Hermits Bog, Iron Bridge, Livingston Bog. EMME
Arnott Lake, Pleasantview Swamp, Stutsmanville, Big Stone Bay. MACKINAC CO. er oan ak
Prentiss Bay, Cut River, Epoufette. PRESQUE ISLE CO. —Evergreen Beach.

Although it belongs to a rae complex, this species is surprisingly uniform
in eastern North America where it can be recognized by keeled leaves with more or
less thickened margins. Its crisped a secund leaves, when dry, and its occurrence on
rotten logs are useful in identification.

8. Dicranum spurium Hedw.—Rather coarse plants in loosely cohering, dull,
yellow-green or yellow-brown tufts (0.5)2-5(7.5) cm. high, moderately tomentose,
usually interruptedly foliate. Leaves crowded, erect-spreading when moist, arched and
loosely imbricate when dry and only slightly contorted, often + undulate, 3-5 mm.
long, 0.8-1.2 mm. wide, broadly acuminate from a broad, ovate base (narrowed toward
the insertion), acute, serrulate in the we ee or more; costa 90-150 uw wide just
above the alar region, percurrent or nearly so, + ridged at back and slightly (or
sometimes strongly) toothed above, ste aie spurred; upper cells 8-14 u wide,

88

usually irregular in shape and arrangement, transversely elongate, subquadrate, or
triangular, with unevenly thickened walls, strongly papillose at back (or very rarely
nearly smooth), sometimes projecting at back as scattered teeth; lower cells elongate
and porose; alar cells brownish, not extending to the costa. Male plants dwarfed. Setae
12-24 mm. long; capsules 1.75-2 mm. long, inclined to horizontal, curved and asym-
metric, furrowed and + strumose when dry; annulus of about 2 rows of large, pale
cells, + revoluble; operculum 1.7-2.5 mm. long. Spores 20-24 u.—n = 12, 24

Conard, How to Know the Mosses, (ed. 2) fig. 58e-h. Grout, Mosses with Hand-lens and
acc "tig, 45a, d, e, Moss Flora of North America 1: Pl. 46A.—Fig. 241-243.

On light, often sandy soil or on rock, apparently limited to dry, open places, such as
scrubby oak, aspen, or pine woods, or rocky heaths or barrens. Circumpolar; Newfoundland to
Michigan, south to Arkansas and North Carolina

CHEBOYGAN CO.—Devereaux Lake. EMMET CO.—Wycamp Lake, Cecil Bay. PRESQUE
ISLE CO.—Hammond Bay.

The shape of the leaves (broadly tapered from the base) and their arched-
imbricate position when dry give a turgid appearance to the individual plants. The
species approaches D. sabuletorum through intergrading forms.

9. Dicranum sabuletorum Ren. & Card.—Dull, yellow-brown plants in rather
jae tomentose tufts 1-3(5) cm. high, + comose. Leaves erect-spreading when moist,

ved or + crisped when dry, indistinctly undulate, 3-6 mm. long, rather broadly
eens broadly short-acuminate to slenderly long-acuminate, distinctly keeled,
serrulate in the upper 1/3 or more; costa (75)90-140(160)u wide near the base,
shortly excurrent (or sometimes merely percurrent), often densely serrulate and
papillose at back, sometimes nearly smooth, low-ridged; upper cells (9)11-15(17) wu
wide, usually irregular in shape and size, mostly + subquadrate with walls usually +
unequally thickened but arranged in well-marked longitudinal rows separated by fairly
conspicuous pellucid lines, rather obscure, moderately papillose at back or nearly
smooth, rarely projecting at back as scattered, very inconspicuous teeth at the
undulations; lower cells narrow and elongate and slightly to strongly porose; alar cells
brownish, not extending to the costa. Male plants dwarfed. Setae 13-22 mm. long;
capsules 1.5-2 mm. long, curved, inclined to horizontal, + strumose, somewhat fur-
rowed when dry; annulus well developed, of 2 rows of cells, fragmenting; operculum
1.5-1.7 mm. long. Spores 21.23 u.—n = 12

Conard, How to Know the Mosses, (ed. 2) fig. 59e-f, 120e-f. Grout, Mosses with Hand-lens
and Microscope, fig. 45b, c, c’ (as D. pallidum). Welch, Mosses of Indiana, fig. 59 (as D
condensatum) .—Fig. 244-245.

As the name indicates, on sand or other light soil, often on thin soil over rocks, rarely
humus, in dry, as coniferous forests. Endemic to North America; Nova Scotia to Michigan, ah
to Florida and Texa

EMMET CO.—Wycamp Lake, Big Stone Bay. PRESQUE ISLE CO.—Huron Beach, Evergreen
Beach, Clinton Lake.

Although closely related to D. spurium, D. sabuletorum is more likely to be
confused with other, less nearly allied species. The leaves of D. sabuletorum vary from
short, broad, and subacute, with a percurrent or shortly excurrent costa, to long,
narrow, and slenderly acuminate, with a rather long-excurrent costa. The plants may
resemble D. drummondii, but the single setae and less controrted leaves with a more
regular arrangement of cells in longitudinal rows help to distinguish D. sabuletorum.

89

10. Dicranum undulatum Brid. (D. bergeri Bland.)—Rather robust plants in very
compact, yellow-brown or green, moderately tomentose tufts 5-9(12) cm. high. Leaves
erect-curved, not much altered on drying or somewhat contorted, often + secund at
stem tips, 5.5-9 mm. long, oblong-lanceolate, gradually narrowed to a broad acumen,
broadly acute or obtuse, keeled (usually rather broadly so), undulate, serrulate in the
upper % or more, erect at the margins (or occasionally + revolute); costa 100-125 yu
wide near the base, ending near the apex, rather indistinctly ridged, serrate at back
above or sometimes nearly smooth; upper cells 11-16 w wide, irregularly short-rectangu-
lar, in irregular longitudinal rows, + pellucid, very incrassate, smooth or occasionally +
papillose at back, rarely a few cells projecting at back as scattered teeth. Male plants
dwarfed. Setae single, 15-30 mm. long; capsules 2.5-4mm. long, curved and asym-
metric, inclined or horizontal, + furrowed when dry, not strumose; annulus of 2(3)
rows of larger, pale cells, fragmenting; operculum 2.5-3 mm. or more long. Spores
16-23 u

Conard, How to Know the Mosses, (ed. 2) fig. 60d-e (as D. bergeri). Darlington, Mosses of
chigan, fig. 44 (as D. bergeri). Grout, Moss Flora of North America 1: Pl. 46B (as D.
sees _Fig. 246-247.

A characteristic species of peat bogs and boggy heaths, often growing in Sphagnum
hummocks (most frequently under clumps of black spruce). Circumpolar

CHEBOYGAN CO.—Gleasons Bog, Gates Bog, Maloneys Bog, Mud Lake, Little Lake 16.
EMMET CO.—Conway Bog, Bear Creek Township, Stutsmanville, Galloway Bog, Cecil Bay, French
Farm Lake. MACKINAC CO.—Hendricks, Little Dollar Lake, Peggley Lake. PRESQUE ISLE
CO.—Hammond Bay, Evergreen Beach, Loon Lake, Clinton Lake.

Best recognized by its occurrence in peat bogs, D. undulatum usually grows in
tall, compact tufts, with erect, broadly pointed, undulate leaves.

In typical form D. undulatum and D. drummondii are easily distinguished, but
intergrades can be difficult. D. drummondii usually grows in dry coniferous woods in
dull, loosely cohering tufts, whereas D. undulatum is characteristic of open bogs where
it grows in compact, often rather shiny tufts in the shelter of black spruce and
tamarack. The tomentum, abundant in both species, is more obvious in D. drummondii
because the leaves are more distantly spaced and more contorted when dry. The leaves
of D. drummondii are slenderly long-acuminate, narrowly acute, and sharply serrate in
the upper half and especially near the apex. In D. undulatum the leaves are broadly
acuminate and broadly acute or obtuse at the apex. The upper cells of D. drummondii,
small (8-11 ys), obscure, green, relatively thin-walled, subquadrate and arranged in
regular longitudinal rows, are clearly papillose at back and often project as scattered
dorsal teeth. In D. undulatum, the cells are larger (11-16 wu), thick-walled, irregular in
shape and arrangement, usually smooth, and only rarely projecting as teeth. Perhaps
the most reliable character for distinguishing difficult specimens is the areolation of the
leaf.

5. Paraleucobryum (Lindb.) Loeske .

Plants rather robust, in gray-green or yellow, somewhat shiny tufts, scarcely to
moderately radiculose below. Leaves erect-spreading to falcate-secund, lanceolate,
gradually acuminate, subtubulose above; costa broad (less than 1/3 to about 9/10 the
leaf base), filling most of the subula, without stereids but consisting of small,
thick-walled green cells in a median layer and thick-walled, hyaline cells dorsally and
ventrally (the green cells in P. longifolium also interspersed in rows among the hyaline

90

cells dorsally); cells of the lamina firm, smooth, hyaline, rectangular above, elongate
below, the alar cells well differentiated. Dioicous; inner perichaetial leaves sheathing.
Setae elongate, erect; capsules erect and symmetric, sometimes slightly curved, +
cylindric; operculum slenderly long-rostrate; peristome teeth 16, inserted below the
mouth, red-brown, lanceolate, forked 4% way down or more, obliquely or + vertically
striolate. Spores spherical. Calyptra smooth, naked, entire at base.—The name refers to
the costa which is similar in structure to that of Leucobryum.

Paraleucobryum longifolium (Hedw.) Loeske.—Plants green or yellowish, with a
slight grayish sheen when dry, + radiculose, 1-3.5 cm. high. Leaves usually falcate-
secund, sometimes stiffly erect and fragile, 4-7.5 mm. long, lance-subulate, serrulate
1/2-2/3 down, roughened at back above; costa more than 1/2 the width of the base
(sometimes as much as 2/3), filling all of the subula, consisting of 3-4 layers of
thick-walled cells of nearly uniform size, the middle and dorsal layers with inter-
mingled green cells (alternating rows of green and hyaline cells at back appearing as
fine, longitudinal striations at low magnifications); inflated alar cells extending to the
costa, hyaline to brownish; cells of lamina long-rectangular to linear, sometimes +
porose. Seta 9-20 mm. long, straw-colored; urn 1,5-3 mm. long, oblong: -cylindric,
straw-colored to brownish, tough-walled, usually irregularly furrowed when dry and
empty; annulus none; operculum slightly shorter than the urn; peristome teeth
obliquely or + vertically striolate to the apex. Spores 23-37 u, densely and minutely
roughened, obscurely green.—n = 12.

Con How to Know the ee ee 2) fig. 122a-d. Grout, Mosses with Hand-lens and
Mice Pl. 17 (as Dicranum), Mos ora of North America 1: Pl. 44H (as Dicranum).
Jennings, Mosses of Western sista pee i 1) Pl. 9 & (ed. 2) Pl. 13 (as Dicranum).—Fig.
248-250.

t commonly on rocks in relatively moist woods (but also on bases of trees and rotten
logs, or rarely humus or soil). Circumpolar; Alaska to Nova Scotia, south to Arizona, Colorado,
Michigan, and North Carolina.

CHEBOYGAN CO. a Point. EMMET CO.—Middle Village, Cecil Bay. MACKINAC
CO.—Mackinac Island, Bush

Resembling a Dicranum but recognized by a grayish sheen when dry and (with
the aid of a lens) the apparent striolation of the leaves caused by longitudinal strips of
green cells enclosed by hyaline cells. The leaves consist mainly of costa and in their
curious structure indicate a transition to the Leucobryaceae.

Other members of the family found in the Upper Peninsula:

Rhabdoweisia crispata (With.) Lindb. occurs at the Upper Falls of the
Tahquamenon River in Luce Co. The plants are very small. The leaves are narrow,
acute, serrulate hee with smooth, subquadrate upper cells; the alar cells are scarcely
differentiated. The inflorescence is autoicous, The capsules are erect and symmetric,
obconic and 8-furrowed when dry and empty, and the peristome teeth are linear and
smooth or faintly and irregularly striolate.

Dichodontium pellucidum (Hedw.) Schimp. occurs at the Lower Falls of the
Tahquamenon in Chippewa County. The plants are rather small. The leaves are rather
broad, oblong-lanceolate to lingulate, bluntly acute to rounded-obtuse, irregularly
dentate, with upper cells subquadrate and mammillose or conic-papillose; the alar cells
are not differentiated. The inflorescences are dioicous, the capsules suberect to curved
and nodding, smooth, with well-developed peristome teeth which are vertically

9]

| | \ \ a .
248 y \ | & |

Paraleucobryum longifolium. 248. Habit. 249. Leaves. 25. Cross-section of leaf.
Leucobryum glaucum, 251. Habit. 252. Leaves. 253. Cross-section at base of leaf.

pitted-striolate.North American records of Oreoweisia serrulata (Funck) De Not.
should be referred here.

Cynodontium schisti (Wahl.) Lindb., C. strumiferum (Hedw.) Lindb.,
torquescens Limpr. are small, autoicous mosses with lanceolate, acute, keeled leaves
with margins bistratose and recurved and cells somewhat roughened by coarse,
papillose bulges; the capsules are furrowed and the peristome teeth vertically pitted-
striolate. C. strumiferum has capsules curved, inclined, and strumose at base; the
peristome teeth are forked. C. schisti has erect, non-strumose capsules and undivided
peristome teeth. C. torquescens has erect, non-strumose capsules and forked peristome
teeth. All three occur in crevices of granitic rock.

o2

Dicranoweisia crispula (Hedw.) Lindb. ex Milde grows on granitic rock ledges and
crevices of cliffs, especially near streams, in the Upper Peninsula. The plants are small,
dull, densely tufted, with leaves crisped and curled when dry; the cells appear papillose
in section because of cuticular ridges running lengthwise; the perichaetial leaves are
convolute-sheathing, the innermost ending abruptly in a broadly pointed or truncate
apex; the capsules are erect and smooth; the peristome teeth are inserted below the
mouth, undivided, and papillose throughout.

LEUCOBRYACEAE

Rather coarse, whitish, grayish, or bluish-green or somewhat iridescent plants in
dense, often large cushions or hummocks. Stems erect. Leaves thick and fleshy, linear,
ligulate, or lanceolate from a somewhat broader base; costa filling the leaf point and
most of the base, consisting of 2 or more layers of large, empty, hyaline, porose cells
(leucocysts) enclosing a single + median layer of small, green cells (chlorocysts);
laminae very narrow, restricted to the base or sometimes extending above the shoulders
as an inconspicuous border, consisting of delicate, hyaline, oblong to linear cells. Setae
terminal, mostly elongate, erect; capsules erect and symmetric to inclined and
asymmetric, often strumose at base; peristome single, consisting of 8-16 lanceolate,
entire or bifid, smooth, papillose, or vertically pitted-striolate teeth.

Leucobryum Brid.

Plants medium-sized to large, in dense, whitish, grayish or bluish-green cushions
or hummocks. Stems forked. Leaves crowded, thick and fleshy, lanceolate or subulate-
lanceolate and subtubulose from an oblong-obovate or elliptic base; the laminae very
narrow, hyaline and delicate, essentially restricted to the base (extending somewhat
to well above the shoulders as an inconspicuous border), the remainder of the leaf
filled by the costa which consists of 2-8 layers of leucocysts enclosing a + median
layer of chlorocysts. Dioicous; male plants dwarfed, growing on or among the leaves
of the female plants. Setae elongate, erect; capsules inclined, asymmetric, 8-ribbed, often
strumose at base; annulus usually none; operculum long-rostrate; peristome teeth 16,
lanceolate, bifid to the middle, vertically pitted-striolate below, papillose above. Spores
spherical. Calyptra large, cucullate.-The generic name means white moss.

Leucobryum glaucum (Hedw.) Schimp.—Fairly robust plants in deep, hemispheri-
cal cushions usually 2-9 cm. high (but often much higher and forming very large
hummocks), sometimes bearing clusters of reduced leaves at the stem tips. Leaves erect
to erect-spreading or sometimes subsecund, 3-8 mm. long, lanceolate and concave or
subtubulose from an erect, oblong-obovate base 1.5-3 mm. long, acute or apiculate,
usually + serrulate at the tip; costa in cross-section near the insertion showing 5-7
irregular layers of leucocysts on either side of the middle, with 3-4 layers below and
2-3 above the chlorocysts; lamina 5-11 cells wide. Setae 9-17 mm. long, red-brown;
capsules 1.5-2 mm. long, curved and strongly inclined, strumose at base; annulus none;
operculum 1.5-2 mm. long. Spores 13-18 m4, nearly smooth or very finely papillose.
Calyptrae often not split completely to the base which clasps the tip of the seta until
maturity of the capsule.—n = £11, 11,

en, Mosses of Florida, Pl. 24 (figs. 1, o Conard, How to Know the Mosses, (ed. fig.
29. Darlington, Mosses of Michigan, fig. 46. Grout, Moss Flora of North America 1: $2.
Jennings, Mosses of Western Pennsylvania, (ed. 1) Pl. 10, (ed. 2) Pl. 14. Welch, Mosses of ee
fig. 62.—Fig. 251-253.

93

ommon in moist forests on soil or humus (usually associated with very decomposed wood
hum Aaa occasional on rotten logs or stumps or bark at base of trees. Europe, Azores, Madeira,
Canary Islands; Caucasus; Japan; Newfoundland to Minnesota, south to Florida and Louisiana.

CHEBOYGAN CO.—Douglas Lake, Reeses Bog, Gorge of Carp Creek, Iron Bridge, Colonial
Point, Bryants Bog, Vincent Lake, Grass Bay. EMMET CO.—Arnott Lake, Galloway Bog, Carp
Lake, Cross Village, Middle Village, Five Mile Creek, Cecil Bay. MACKINAC CO.-—Bois Blanc
Island, Hendricks. PRESQUE ISLE CO.—Hammond Bay, Evergreen Beach, Presque Isle Lighthouse,
Clinton Lake

Sometimes called WHITE MOSS or PIN CUSHION MOSS, Leucobryum glaucum
is especially associated with wood humus, sometimes growing in large mounds
representing logs or stumps in an advanced stage of decomposition. Occasionally one
sees colonies up to several feet in diameter.

Sporophytes are rare, but small brood-leaves are sometimes produced in dense
axillary clusters resembling loose cabbage heads, and the leaves often proliferate
radicles at their tips and no doubt form new plants by regeneration following
accidental breakage of the leaves. I would guess, however, that it is a dwarf male plant
rooted on a leaf that Burrell (Bryol. 10: 107-111. Pl. 12, fig. 1. 1907) illustrated,
rather than a small plant produced by regeneration, as he thought

The localized dispersal of the species as a result of accidental disturbance is
neatly illustrated by Stormer’s observation that Leuwcobryum had spread along numer-
ous paths that developed between 1935 and 1964 and in a wooded area made available
for public camping (Mosses with a Western and Southern Distribution in Norway, p.
113. 1969).

Plants scratched loose by game birds or in some other accidental way may
continue growth as rounded and unattached “moss balls.’’ Burrell (Bryol. 10: 107-111.
1907) attributed such growths of Leucobryum glaucum to the scratching of pheasants.
Other reports, of the same species, include papers by Gilbert (Bryol. 9: 72. 1906) and
Dixon, as “Fawley Buns” (Bryol. 15: 31-32. 1912). Dixon, in the same article,
reported similar growths of Echinodium hispidum in New Zealand resulting from the
scratching of wood hens. In The Student’s Handbook of British Mosses, he mentioned
moss balls of Porotrichum alopecurum. Further information on this phenomenon in
New Zealand was provided by Martin (Bryol. 55: 65-70. 1952). Shacklette reported a
similar development of Grimmia apocarpa on Amchitka Island, Alaska, where plants
become detached from their normal rock substrate by wind and assume a spherical
growth form as a result of continual rolling on dune sands. These resemble “glacier
mice” developed in a similar way on glaciers in Iceland and Alaska and moss balls on
sand in Norway and Jan Mayen (Lid, Nytt Mag. Naturw. 78: 101-104. 1938). Heusser
(Bull. Torrey Bot. Club 99: 34. 1972) reported, with a photograph, hundreds of
rounded polsters of Drepanocladus berggrenii (C. Jens.) Roth on glacial ice in Alaska.

Moss balls call to mind other vagabonds, such as the tumbleweeds of the
American prairies or the manna which fell from Heaven as food for the Children of
Israel. Manna is believed to have been, in part, a lichen, Lecanora esculenta, which
occurs in arid wastes of North Africa and the Near East. It becomes detached and
blown great distances by the wind. Sometimes eaten by men and his animals, it has
food value, although it is abrasive to the teeth!

Leucobryum albidum (Brid.) Lindb., found once in Washtenaw County in the
extreme southern part of the state, and common farther south, is much smaller with
leaves only 2-4 mm. long; the leaf point is about the same length as the base and the
costa consists of 2 (or rarely here and there, 3) hyaline cells above and below the
green cells on either side of the median region at leaf base.

94
ENCALYPTACEAE

Plants rather small to moderately robust, loosely to densely tufted. Stems erect,
forked. Leaves elliptic to oblong-ovate or oblong-obovate, rounded-obtuse to
acuminate, sometimes apiculate or awned, + concave or keeled; margins evenly
papillose-crenulate or crenate-serrulate above, entire and sometimes revolute below;
costa single, strong, ending below the apex to shortly excurrent or ending in the awn,
prominent and often papillose or + toothed at back, consisting of a dorsal stereid band
and large ventral cells; upper cells hexagonal, firm-walled, bulging, and densely
forked-papillose on both surfaces and obscure (or, in Bryobrittonia, mammillose and
pellucid); basal cells pale, lax, oblong (with thickened, brown cross-walls in eee
the basal marginal cells smooth, pale, and linear in several rows. Setae termin
elongate, erect or somewhat flexuose, usually twisted above when dry; capsules ae

d symmetric, cylindric, with a short and usually inconspicuous neck, smooth or
furrowed; annulus present; operculum erect, long-rostrate; peristome lacking, single, or
double, the 16 teeth lanceolate to linear or filiform, the endostome segments rising
from a low membrane, pale and delicate, often + adherent to the teeth. Calyptra
mitrate, large, conic-cylindric and long-rostrate, fringed at base.

Encalypta Hedw.

Rather small to fairly robust plants in dense tufts, dull-green or brownish-green
and often glaucous above, brown and radiculose below. Leaves elliptic or ovate to
oblong-obovate, rounded-obtuse to acute or acuminate, often apiculate or hyaline-
awned, usually concave or somewhat keeled, often folded and contorted when dry;
costa ending well below the apex to percurrent or + excurrent, sometimes disappearing
in the awn; upper cells green and obscure, densely forked-papillose on both surfaces;
basal cells large, lax, smooth, oblong, with brown, thickened cross-walls, the basal
marginal cells pale and linear in several rows. Setae usually twisted above when dry,
erect or flexuose; capsules smooth or furrowed; peristome lacking, single, or double,
the teeth 16, lanceolate to filiform, entire or variously divided, papillose, the segments
pale and delicate, papillose, usually + adherent to the teeth, rising from a low
membrane adherent to the teeth._The name means covered with a veil, in reference to
the large calyptra resembling a candle-snuffer, hence the common name of EXTINGUISH-
ER MOSS. The lax basal cells with thickened and colored cross-walls are useful in
recognition of the genus from sterile material. Other interesting features of the genus
include the variability of the peristome, single, double, or lacking, from one species to
another and the very distinctive and recognizable spores of the various species.

1. Plants relatively pee ickiond sterile and _commonly bearing abundant, brown, filifor
brood bodies on stems; uppermost and perichaetial leaves of fruiting plants piliferous, the
others muticous; ae ually furrowed; ie double; spores 16- 6 wu, very finely
p 3. E. procera

de she smaller, usually sess without brood ieee leaves ae ee or awned; capsules

t spirally furrowed; peristom clas spores 27 uw or more, not papi
eee acute and hyaline-awned; capsules eae Haenebe panes ae HK,
warty; calyptra slightly erose at oe E, rhaptocarpa

2. Leaves rounded-obtuse to broadly acute, abruptly narrowed to a oi ‘apiculus;
capsules smooth; spores 28-36 uw, wrinkled-reticulate; calyptra deeply fringed at base
1. &. ciliata

1. Encalypta ciliata Hedw.—Plants 5-40 mm. high, dull-green or yellow-brown
above, brown below. Leaves + folded and contorted when dry, 2.5-4 mm. long, elliptic
to oblong-obovate, rounded-obtuse to broadly acute, ending abruptly in a smooth,

95

yellow apiculus; margins plane or erect above, or + incurved at the apex, revolute
below the middle; costa ending near the apex to shortly excurrent, smooth at back or
sometimes slightly toothed near the apex; upper cells 11-18. Autoicous. Setae
4-10 mm. long, yellow or brown; capsules 2-3 mm. long, light- to dark-brown, smooth
(but usually very finely wrinkled-striolate when old), thin-walled and often becoming
torn or shredded wtih age; annulus poorly developed, of 1 row of cells; operculum
1-1.6 mm. long; peristome single, the teeth lanceolate, finely papillose and often +
striolate as well, brown or rarely reddish-brown. Spores spherical, 28-36 uy, smooth (or
very rarely faintly roughened) but irregularly wrinkled-reticulate within. Calyptra
smooth, fringed at base.—n = 13

Conard, How to Know the Mosses, (ed. 2) fig. 77. Darlington, Mosses of Michigan, fig. 47.
Grout, Mosses with Hand-lens and Microscope, Pl. 37, Moss Flora of North America, 1: Pl.
71B.—Fig. 254-257

On ledges or in crevices of limey cliffs, also on soil of banks, especially in the shelter of
overhanging sod. Europe, Africa, and Asia; Alaska to Labrador, south to California, Arizona, New
Mexico, Iowa, and Pennsylvania; Mexico and Ecuador

Encalypta ciliata, 254. Leaves. 255. Capsules. 256. Calyptra. 257. Spor
Ecalypta procera. 258. Leaf. 259. Apex of perichaetial leaf. 260. " Brood-bodies. 261.
ihe

calypta rhaptocarpa. 262. Habit. nee Leaves. 264. Upper cells of leaf. 265. Basal cells of
leaf. ves Capsule. 267, Calyptra. 268. Spor

96

CHEBOYGAN CO.-—Fairy Island (Douglas Lake), also “Cheboygan.” EMMET CO.—Cross
Village

Recognized by smooth capsules with a single, red-brown peristome. The spores
smooth on the outside but irregularly wrinkled-reticulate within, are oddly distinctive.

2. Encalypta rhaptocarpa Schwaegr.—Plants 5-30 mm. high, dull-green to yellow-
ish or brownish above, brown below. Leaves somewhat keeled and incurved but usually
not much contorted when dry, 2.2-3 mm. long, ovate, broadly acuminate, acute and
hyaline-awned; margins plane, or erect near the apex; costa ending below the apex to
percurrent or disappearing in the awn, smooth or somewhat papillose at back,
especially below; upper cells 11-16. Autoicous. Setae 6-10 mm. long, brown or
red-brown; capsules 2-3 mm. long, dark- or red-brown, abruptly narrowed at the base,
strongly straight-ribbed, wrinkled at the short neck when dry; annulus of 2 rows of
small, usually persistent cells; operculum 1.1-2 mm. long; peristome single, the teeth
lanceolate, brown or reddish, rather coarsely papillose (sometimes with pre-peristome
fragments adhering to their bases). Spores (27)31-45 uw, spherical or subreniform,
coarsely warty-papillose, the papillae about 5-8 u in diameter. Calyptra smooth or +
scabrous above, slightly erose at base.—n = 26

Conard, How to Know the Mosses, (ed. 2) fig. 76e-f. Grout, Moss Flora of North America 1:
Pl. 69B.—Fig. 262-268.

On soil, often in crevices of dry, calcareous cliffs or in the shelter of turf overhanging steep
banks. Circumpolar; Greenland to Alaska and south to New York, Michigan, South Dakota, New
Mexico, Arizona, and California.

CHEBOYGAN CO. are Island (Douglas Lake). EMMET CO.—Cross Village. MACKINAC
CO.—Mackinac Island, Bush B

Distinctive features include the capsules with straight ribs, a single peristome, and
large, warty spores. In most books the specific epithet is spelled rhabdocarpa, but the
original spelling in Schwaegrichen’s herbarium and in his publication was rhaptocarpa.
A western species, E. vulgaris Hedw., collected in Ontario by Macoun (if label
information can be trusted), differs mainly in the absence of a peristome.

3. Encalypta procera Bruch—Plants 15-20 mm. high, dull-green or brownish-green
above, brown below, often bearing abundant, brown, filiform brood-bodies on simple
or branched radicles on the stems. Leaves + folded and contorted when dry, 2-4 mm.
long, oblong-ovate or oblong-elliptic, obtuse or broadly acute (or occasionally
rounded-obtuse), the upper and perichaetial leaves hyaline-awned; margins plane or
erect above, often reflexed or revolute in the lower 1/2-2/3; costa ending well below
the apex to percurrent or disappearing at the base of the awn, roughly papillose at
back toward the base and often papillose nearly or quite to the apex; upper cells
10-16 uw. Autoicous. Setae 11-17 mm. long, red-brown; capsules 2.5-3 mm. long, spirally
ribbed; annulus revoluble in fragments; operculum 1.5-1.8 mm. long; peristome double,
the teeth long-filiform, red, finely papillose, perforate near the base and joined
together by trabeculae, the segments linear, about 2/3 as long as the teeth and +
adherent to them below, yellow or orange-brown. Spores spherical, 16-26 u, very finely
papillose. Calyptra densely papillose above and occasionally + papillose nearly to the
base, lacerate-fringed. (Usually sterile and bearing abundant brood-bodies in eastern
North America.)—n = 26+1,

OF

Grout, Moss Flora of North America 1: Pl. 72 (figs. 1-14). Jennings, Mosses of Pennsylvania,
(ed. 2) Pl. 62 (as £. streptocarpa).—Fig. 258-261.

A calciphile, growing on or in crevices of damp rocks and cliffs, also on soil of cliff ledges
and road banks. Circumpolar; Greenland to Alaska, south to Idaho, Missouri, Ohio, and North
Carolina.

CHEBOYGAN CO.-—Biological Station campus. EMMET CO.-—Levering, Good Hart, Cross
Village, Cecil Bay, Big Stone Bay. MACKINAC CO. vee Corner, pest Island, Castle Rock,
McKays Creek, Bush Bay, Ozark, Rexton. PRESQUE ISLE CO.—Ocqueoc F

The essentials of this species include the autoicous inflorescence, long, spirally
ribbed capsules, and long, double peristome (with the endostome adhering to the
exostome and, therefore, easily overlooked). Sporophytes are rare in eastern North
America but common in the West. Our eastern plants become rather large because of
continued vegetative growth and are best recognized by masses of brown, filiform
brood-bodies borne on simple or branched radicles on the stems, often crowding the
leaves and causing them to be incurved-contorted when dry. All records of EF.
streptocarpa Hedw. from North America are based on such sterile forms, which do,
however, fruit on occasion and are indeed autoicous, proving them to be E. procera.
There is no positive evidence (such as a supposed central strand) for calling them EF.
streptocarpa, and whenever fruiting plants have been found there has been no question
of their identity with E. procera.

The leaves of the sterile form are muticous. The perichaetial leaves and upper
leaves of fruiting plants are hyaline-awned.

POTTIACEAE

Small to robust plants, usually in dense, dull, sometimes radiculose tufts. Stems
erect, simple or forked. Leaves in several rows, mostly narrowly lanceolate, usually
contorted when dry; costa single, well developed, mostly with 1 or 2 stereid bands;
upper cells generally small and obscure, subquadrate or hexagonal, firm- or thick-
walled, and mostly papillose; lower cells mostly pale and oblong, not differentiated at
the basal angles. Sporophytes mostly terminal; setae usually elongate, erect; capsules
mostly erect and symmetric, usually subcylindric, smooth; annulus sometimes differ-
entiated; operculum generally differentiated and usually conic-rostrate; peristom
usually present, single, the 16 teeth erect, oblique, or spirally wound together, entire
or deeply divided to a low or high basal membrane into 32 filiform, papillose divisions.
Spores usually spherical. Calyptrae cucullate.

Our genera fall into two large and heterogeneous subfamilies: The TRICHO-
STOMOIDEAE (Gymnostomum, Weissia, Aes Bryoerythrophyllum, Didymodon,
and Barbula) generally have a costa with t stereid bands of thick-walled cells
separated by a median row of large empty tis cells and leaf cells with papillae
usually blunt and simple (or none). The POTTIOIDEAE (Phascum, Pottia, Desmatodon,
and Tortula) generally have only a dorsal stereid band, and the leaf cells most often
have C-shaped papillae. (The Germans aptly refer to them as hufeisenférmig, or
horse-shoe shaped.)

1. Basal cells of leaves large, lax, and hyaline, extending up the margins above the shoulders
3.

as a V-shaped border Tortella
1. Basal cells various, not extending upward as a V-shaped border
2. Leaf margins involute 2. Weissia
2. Leaf margins plane or recurved
3. Capsules immersed, without an operculum, dehiscing irregularly 7. Phascum
3. Capsules exserted, dehiscing by means of an operculum

98

4. Leaf margins plane or narrowly recurved on 1 side below the middle; peristome none

5. Plants caespitose on wet, calcareous rocks; leaves with essentially parallel sides, blunt
or acute to apiculate, not awned; costa with 2 stereid bands; peristome oe
1. Gymnostomum
5. Plants scattered or gregarious, on disturbed soil; ae without parallel sides,
oblong-lanceolate, oblong-ovate, or obovate, en stout apiculus or awn:
margins usually plane (rarely somewhat recurved on both sides at the ss costa
with 1 stereid band; peristome none (or rarely very rudimentary 8. Pottia, p.p.
4. Leaf margins recurved on both sides; peristome mostly present and well developed
6. Basal cells subquadrate to oblong, not much enlarged, + firm-walled

7. Plants scattered or gregarious; leaves ending in a stout apieulis or short awn; costa

with 1 stereid band; peristome none (or rarely very rudimentar 8. Pottia, p.p.
7. Plants tufted; leaves muticous or moderately apiculate; peristome present and well
developed
8. Peristome teeth split nearly to the base into 32 filiform divisions and spirally
wound together 6. Barbula
8. Peristome teeth deeply but often irregularly bifid into filiform divisions, erect or
somewhat inclined, not spirally wound together a Didymodon

6. Basal cells + enlarged, lax, pale, thin-walled
9. Leaves lanceolate, with a few irregular, pellucid teeth near the apex; costa with 2

stereid bands 4. ee diel as
9. Leaves broader, oblong-lanceolate to oblong-ovate or elliptic, not toothed;
with 1 stereid band
10. Leaves acute or subulate to hair-pointed; basal leaf cells enlarged, usually in
rather abruptly differentiated groups; peristome consisting of 32 filiform
divisions spirally wound together above a high, tesselated basal membrane 10. Tortula

entiated; peristome teeth split to a low basal membrane into 2 (or . slender
divisions, somewhat spiraled but not wound together 9. Desmatodon

1. Gymnostomum Nees, Hornsch. & Sturm

Plants small to fairly robust, growing on wet, usually calcareous rocks in dense,
dull, green, glaucous, or yellowish tufts, rusty-brown below. Stems erect, forked. Leaves
a curved or incurved-contorted when dry, erect-spreading or rarely recurved when
moist, linear-lanceolate to broadly ligulate, acute and sometimes apiculate or bluntly
pointed to obtuse, concave or somewhat keeled; margins entire, plane or recurved on 1
side below; costa ending near the apex, homogeneous in structure or sometimes with 1
or 2 stereid bands; upper cells rounded or rounded-hexagonal, firm-walled, pluri-
papillose on both surfaces and obscure (or rarely nearly smooth and pellucid); lower
cells short-rectangular to irregularly oblong, smooth, pale, firm- or thick-walled.
Dioicous; perichaetial leaves scarcely differentiated. Setae terminal, elongate, erect or
somewhat flexuose; capsules erect and symmetric, ovoid, oblong-cylindric, or obovoid,
smooth; operculum obliquely long-rostrate, sometimes remaining attached to the
columella after dehiscence; annulus and peristome none. Calyptra smooth, naked.—The
name means naked mouth, referring to the absence of a peristome.

Leaves acute and typically ending in a sharp, smooth, usually pellucid cell; margins recurved

ide below; capsules dull, Wee: obovoid, systyliou recurviro strum
Leaves acute or bluntly pointed, no ing i mooth, pellucid cell; margins plane
capsules ovoid or oblong-cylindric, ae the operculum readily ae 1. G. geruginosum

1. Gymnostomum aeruginosum Sm.—Small plants in dirty-green tufts usually
2-7 mm. high (or rarely 40 mm.). Leaves loosely erect-incurved and moderately crisped
when dry, spreading when moist, often with recurved tips, about 1-1.3 mm. long,
ligulate or linear-lanceolate, bluntly acute to narrowly rounded-obtuse (with an
occasional leaf ending in a sharp, pellucid cell), concave but not particularly keeled;

99

margins plane, often sinuate, papillose-crenulate; costa disappearing near the apex, with
2 stereid bands and a median row of guide cells; upper cells 7-11 yu; rounded-hexagonal,
obscure and pluripapillose; lower cells short-rectangular, moderately firm-walled. Setae
3-5 mm. long, yellow; capsules 0.6-0.8 mm. long, oblong-cylindric, shiny, pale-brown,
thin-walled, finely shriveled-striolate when dry and empty; aes 0.5-0.7 mm
long, readily deciduous. Spores 10-13 yw, very slightly roughened.—n = 13

Breen, ee of Florida, Pl. 36 (figs. 5-8), as G. caleareum. Darlington, Mosses of Michigan,
fig. 50. Grout, Mosses with Hand -lens and Microscope, Pl. 25 (as G. ney Jennings, Mosses of
Western pee aes (ed. 1) Pl. 12 & (ed. 2) Pl. 16 (as G. calcareum).—Fig. 269-271.

n moist or wet, shaded, generally calcareous cliffs or boulders, rarely on soil. Circumpolar;
southeastern Alaska and British Columbia to Newfoundland, south to Arizona, Colorado, Arkansas,
and North Carolina; Mexico and Guatemala

CHEBOYGAN CO.—Mill Creek. MACKINAC CO.—Mackinac Island, Ozark.

The plants are quite small, and a confusion with the still smaller Seligerias is
possible, Sporophytes are common and aid in distinguishing from the following species
which has broader, systylious capsules. Sterile material is distinguished by the plane
leaf margins and blunter leaf tips of G. aeruginosum.

Because outcropping limestone is rare on the south side of the Straits,
Gymnostomum aeruginosum figures as a rarity at Mill Creek, in company with other
calciphiles of local interest, including Seligeria campylopoda and Platydictya con-
fervoides. Nichols (Trans. Conn. Acad. Arts & Sci. 22: 249-467. 1918) reported a
pronounced calciphilous element on gypsum outcrops in Nova Scotia, featuring such
mosses as Distichium inclinatum, Gymnostomum aeruginosum, Encalypta procera,
Myurella sibirica, Thuidium abietinum, and Tortula mucronifolia. This is a typically
calcicolous assemblage similar to that found on the minuscule cliffs at Mill Creek and
in Mackinac and Presque Isle Counties.

Gyroweisia tenuis (Hedw.) Schimp., a rare moss found on soft, red Cambrian
sandstone in Houghton County in the Upper Peninsula, is much like G. aeruginosum
but has longer, more cylindric capsules, well-developed annulus, differentiated peri-
chaetium, and blunt, rounded leaf apex with the costa ending well below it. The
differences are well illustrated by Steere (Bryol. 42: 16-23. 1939), although he erred in
using the name Gymnostomum calcareum Nees, Hornsch. & Sturm for the Michigan
material of G. aeruginosum which he actually illustrated. (Gymnostomum calcareum is
a Mediterranean species which has been found in California but not the eastern United
States.)

2. Gymnostomum recurvirostrum Hedw.—Plants small to moderately robust,
5-30 mm. high (or rarely 50mm. or more), in dark-green or yellow-brown tufts,

frequently interruptedly foliate, sometimes + papillose. Leaves loosely erect and
somewhat contorted or occasionally appressed in 5 spiral rows when dry, erect-spread-
ing, often with + recurved tips when moist, 0.8-2 mm. long, linear-lanceolate to
oblong-lanceolate, acute (or rarely some leaves blunt), usually ending in a sharp,
smooth, pellucid cell, keeled; margins papillose-crenulate above and occasionally
papillose-serrate somewhat above the base, recurved on 1 or very rarely both sides
below; costa vanishing near the apex, with a small dorsal stereid band; upper cells
8-11 y, irregularly hexagonal, subquadrate, or short-rectangular, obscure and densely
papillose to nearly smooth and pellucid; lower cells + enlarged, irregularly oblong,
thick-walled and often porose. Setae 5-11 mm. long, yellowish, becoming red-brown

100

with age; capsules 1-1.5 mm. long, obovoid or sometimes subcylindric, usually dark-
brown, dull, firm-walled, smooth; operculum 1-1.2 mm. long, remaining attached to the
columella for some time after dehiscence. Spores 16-20 yu, very finely roughened.—n =
eg oe

Conard, How to Know the Mosses, (ed. 2) fig. 97a-c. Darlington, Mosses of Michigan, fig. 52
(as Hymenostylium). Grout, Mosses with Hand-lens and Microscope, Pl. 24 (as G. curvirostre).
Jennings, Mosses of Western Pennsylvania, (ed. 1) Pl. 12 (as Hymenostylium curvirostre), (ed. 2) P
16 (as H. recurvirostrum). Welch, Mosses of Indiana, fig. 69.—Fig. 272-273.

On wet, ripen cliffs (and locally on low ridges of dune sand around moist depressions
along the south shore of the Straits). Europe, Asia, Philippine Islands; New Guinea; Mexico,
Guatemala, West ee Alaska to Greenland and south to Texas and Arizona, widespread in the
East, south to Tennessee and Arkansas.

EMMET CO.—Cecil Bay.

The leaf margins narrowly recurved on one side offer a convenience in
determination. The systylious capsules, though curious and very different in appearance
from those of G. aeruginosum, scarcely justify the segregation of the genus Hymeno-
stylium. | expect a good deal of variation in plants growing in aquatic or subaquatic
habitats and particularly those found in limey places where alkalinity and pH vary with
seasonal fluctuations in the water supply. For this reason, I am not much impressed
with forms with papillose stems, referable to var. latifolium (Zett.) Flow. ex Crum
[var. scabrum (Lindb.) Grout], or those with cells less papillose than usual, or nearly
smooth, and therefore pellucid, called the var. commutatum (Mitt.) Grout. Neither of

anomalous habitat. The species is almost certain to be found in Mackinac County or
Presque Isle County where wet, outcropping limestone rocks are occasional.

Eucladium verticillatum (Brid.) BSG (fig. 274-275) has been found on Summer
Island in Lake Michigan (Delta Co.). It is a tufa-forming species similar to Gymno-
stomum recurvirostrum in both habitat and appearance, but recognized by plane leaf
margins which are irregularly serrate near the junction of the green, upper cells and the
pale, enlarged basal cells. The chromosome number reported for EF. verticillatum is n =
13

2. Weissia Hedw.

Small plants in dull, green, yellowish or brownish, loose or dense tufts. Leaves
becoming somewhat larger toward the stem tips, strongly curled and contorted when
dry, erect-spreading or spreading when moist, narrowly lanceolate or linear-lanceolate
from a rather narrow, oblong base, acute and apiculate, + subtubulose above, especially
when dry; margins incurved or inrolled, usually from the shoulders to the apex; costa
shortly excurrent as a smooth, pellucid apiculus, with 2 stereid bands; upper cells
small, hexagonal, firm-walled, densely papillose on both surfaces and obscure; lower
cells short-rectangular, smooth, pale. Perichaetial leaves not much differentiated. Setae
terminal, mostly elongate, straight; capsules erect and symmetric, ellipsoidal to
oblong-cylindric, annulus none or narrow and persistent; operculum rostrate; stomata
at extreme base of the capsule; peristome lacking (sometimes with the mouth + closed
by an evanescent membrane) or consisting of 16 short (and sometimes rudimentary),
entire or somewhat cleft and perforate, erect, papillose teeth. Calyptrae smooth,
naked._Named by Hedwig for Friederich Wilhelm Weiss, a cryptogamic botanist of

i re
JOUE
Soy )(¢

Gymnostomum aeruginosum, 269. Habit. 270. Leaves. 271. Cells at leaf tip.
Gymnostomum recurvirostrum. 272. Leaf. 273. Capsule.
Eucladium verticillatum, 274. Leaves. 275. Cells at ein of leaf shoulder.
Weissia controversa. DG: Habit. 277. Leaves. 278. Cross-section of upper portion of leaf
(diagrammatic). 279. Cells at leaf tip.

G6ttingen. (The name in older books was often spelled Weis and the genus Weisia. |
prefer to adapt the generic name to the usual spelling of the surname. Anyway, the
spelling used by Hedwig in the Species Muscorum, 1801, was Weissia.)

Weissia controversa Hedw.—Plants about 2-5 (or very rarely 10) mm. high. Leaves
1-2.5, sometimes 3 mm. long, linear-lanceolate from an oblong base; costa 45-60 uw wide
near the base; upper cells 6-8 y. Autoicous (but often appearing to be dioicous). Setae
3-15 mm. long (but usually only 4-8 mm.), yellow; capsules 0.7-1.5 mm. long, oblong-
cylindric, + plicate when dry and empty, yellow-brown, becoming chestnut-brown with
age; annulus poorly differentiated; operculum obliquely long-rostrate, 0.5-0.8 mm. long;
peristome teeth short, lanceolate, sometimes slightly and ae cleft or perforate,
red-brown. Spores 15-20 yp, finely papillose, brown.—n = 13,

Breen, Mosses of Florida, Pl. 38 (figs. ae Conard, How to oY the Mosses, (ed. 2) fig. 84
(as W. aioe sar i n, ton, Mosses oF Michigan, fig. 49. Grout, Mosses with Hand-lens and
Microscope, Pl. 23 (as W. viridula). Jennings, Mosses of Western Pee eee (ed. 1) Pl. 12 & (ed.
2) Pl. 16 (as W. Sei Welch, Mosses ae Indiana, fig. 67 (as W. viridula).—Fig. 276-279.

, calcareous soil in exposed and pte habitats, such as old fields and roadsides.
Widely pith in Europe, Asia, Africa, New Zealand; Mexico, Central and South America, West
Indies; British Columbia to Nova Scotia and throughout the United States

CHEBOYGAN CO.-—Biological Station campus, Mill Creek. EMMET CO.—Brutus, Pellston
cau aes Lake, Sturgeon Bay Point. MACKINAC CO.—Allenville. PRESQUE ISLE CO.—
Pose

102

Recognized by its small size and crisped leaves with involute upper margins, this
species seems rare locally, perhaps because like most denizens of old fields it escapes
competition by completing its life cycle quickly in spring or fall, when bryological
activity is locally at a minimum. Similarly, at some off-season, one might find
Astomum, which is a Weissia with immersed, cleistocarpous sporophytes. Astomum
muhlenbergianum (Sw.) Grout has been found in Michigan.

Claire Williams (Bryol. 69: 361-365. 1966) found large numbers of sporophytes
that were hybrids between Astomum muhlenbergianum and Weissia controversa, in
Ontario. All the hybrids were produced on Astomum gametophytes and in the
presence of both parent species. Similarly, Reese and Lemmon (Bryol. 68: 277-283.
1965) reported hybridization between Weissia controversa and Astomum ludovicianum,
in Louisiana. The poorly formed spores produced by some of the intermediates gave
supporting evidence of hybrid origin. Because of such hybridization Reese and
Lemmon transferred all the North American species of Astomum to Weissia. Crundwell
& Nyholm (Jour. Bryol. 7(1): 7-19. 1972) cited several other generic crosses linking
Astomum, Hymenostomum, and even Tortella. | feel that here, as in the Funariaceae,
intergeneric crosses may occur occasionally and under unusual circumstances, but in
practice there is almost never any difficulty in recognizing them as distinct genera.
After all, closely related genera are branches of a phylogenetic tree rather than separate
creations. It is convenient and right to retain such recognizable branches (or evolution-
ary trends) as Funaria and Entosthodon, for example, as genera. One should expect
occasional cases of cytological compatibility. In nature, spatial, temporal, and ecologi-
cal barriers make hybridization unlikely.

Khanna (Bryol. 63: 1-16. 1960) made an interesting study of variability in
Weissia exserta (Broth.) Chen, which may also be regarded as Hymenostomum
exsertum (Broth.) Broth., in the Punjab and concluded that it is a young and vigorous
species which originated as a result of hybridization between Astomum crispum
(Hedw.) Hampe and Weissia controversa Hedw., followed by chromosome doubling.

3. Tortella (Lindb.) Limpr., nom. cons.

Plants small to medium-sized, in loose or dense, green, yellowish, or brown tufts.
Stems erect, simple or forked, + radiculose below. Leaves usually crisped and contorted
when dry, erect-spreading or spreading from an erect, oblong base when moist,
long-lanceolate, linear-lanceolate, or subulate, acute or obtuse, often apiculate; margins
plane or erect, often incurved near the apex, often slightly and irregularly notched,
somewhat undulate; costa strong, percurrent to excurrent, smooth at back, shiny and +
conspicuous when dry, with 2 stereid bands; upper cells rounded-hexagonal, green,
obscure, densely papillose on both surfaces; lower cells laxly long-rectangular, delicate,
hyaline, smooth, sharply set off from the upper cells in a V-shaped region extending
above the shoulders as a short or fairly elongate border. Perichaetial leaves usually not
much differentiated. Setae terminal, single, elongate, erect, becoming reddish with age;
capsules erect and symmetric or somewhat curved, cylindric or ovoid-cylindric, smooth,
greenish-yellow or yellow-brown; annulus present or lacking; operculum high-conic, 4%
as long as the urn or more; stomata at extreme base of the capsule; peristome teeth
inserted somewhat below the mouth, 16, split to a low basal membrane into 32
filiform, reddish, densely spiculose-papillose divisions which are wound together in 1-3
spirals (or rarely rudimentary or merely obliquely inclined). Calyptrae smooth,
naked.—The name is a diminutive meaning twisted, in allusion to the twisted peristome
teeth. The abrupt differentiation of the basal cells along a V-shaped line is the most
obvious and useful character of the genus.

103

1. Leaves rigidly erect and not much contorted when dry, with long, linear, fragile tips
Let: ae
1. Leaves cee or incurved and strongly contorted when dry, not ending in a linear

fragile
2. ee Paden long-subulate, spirally contorted when dr 2. T. tortuosa
2. Leaves acu oe e or obtuse and apiculate, irregularly incurved and contorted when dry
3. Autoicous; leaves only slightly keeled when moist, not cucullate at the apex 3. T. humilis
3% Dison jae keeled and cucullate even when moist 4. T. inclinata

1. Tortella fragilis (Hook. ex Drumm.) Limpr.—Plants in dense, rigid, green to
brown or yellowish tufts up to 4.5 cm. high. Leaves erect-incurved and somewhat twisted
around the stem when dry, erect-spreading when moist, about 4 or 5mm. long,
linear-lanceolate and long-subulate from a broader base, the subula thickened and
mostly broken off; costa filling the subula; upper cells 9-11 ux. Dioicous; perichaetial
leaves not much differentiated. Setae 15-20 mm. long; capsules 1.8-3 mm. long; annulus
none; peristome teeth long, spirally wound together. Spores about 9 yu, minutely
roughened or nearly smooth.

Grout, Mosses with Hand-lens and Microscope, Pl. 31.—Fig. 280.

On calcareous soil or rock (often in the crevice of cliffs), often on sand, logs, or other
substrata near lakes and streams. Europe, North Africa, Asia; New Zealand (Campbell Island);
Greenland to Alaska, south to New Jersey, the Great Lakes, Iowa, Colorado, and Orego

CHEBOYGAN ie —Biological Station campus, Carp Creek, Iron Bridge. EMMET CO.—
Arnott Lake, Cecil Bay, Big Stone Creek, Ramona Beach. MACKINAC CO.-—St. Martins Point, Bush
Bay. PRESQUE ISLE Co. ee Beach, Hammond Bay, Mast Point.

The rigid, fragile leaves which are not much contorted when dry are unmistakable.

2. Tortella tortuosa (Hedw.) Limpr.—Plants about 1-5.5cm. high, in dense,
green, yellow-green, or yellow-brown tufts. Leaves strongly contorted with spirally
curled tips when dry, flexuose-spreading when moist, 4-6.5 mm. long, linear-lanceolate
and gradually subulate-acuminate from a somewhat broader base, concave; costa
excurrent as a long apiculus, yellowish (or rarely brownish); upper cells about 7-10 yu.
Dioicous; perichaetial leaves slender and erect at base, slenderly subulate. Setae 9-27
(rarely 35) mm. long; capsules 1.5-3.3 mm. long; annulus none; operculum 1.5-2 mm.
long; peristome teeth long and spirally wound together. Spores 9-11 yw, finely papillose
or nearly smooth.—n = 13.

Conard, How to Know the Mosses, (ed. 2) fig. 94d. Darlington, Mosses of Michigan, fig. 53.
Grout, Mosses with Hand-lens and Microscope, fig. 73.—Fig. 281-283.

On eae rock or soil, common on cliffs and boulders, also locally sand near the
Great Lakes. Circumpolar; Greenland ie Alaska, south to Connecticut, Det iania: et fren
Lakes region, Iowa, Mates and Oreg

CHEBOYGAN CO.—Hermits Bog, Camp Knight and Fairy Island (Douglas Lake), Mill Creek,
Grass Bay, Indian River. EMMET CO.—Arnott Lake, Ramona Beach, Five Mile Creek, Middle
3 : ckina

Island, St. Martins Point, McKays Creek, Bush Bay, Prentiss Bay, Ozark, Caffey Corner, Rexton.
PRESQUE ISLE CO.—Mast Point, Evergreen Beach, Presque Isle lighthouse, Rainy River Falls,
Ocqueoc Falls.

Robust for the genus. The long, subulate leaves spirally twisted and also curled at
the tips when dry are highly distinctive. Tortella tortuosa sometimes has broken leaves,
but it is only rarely confused with the much more rigid 7. fragilis, in which the leaves
are more or less erect, not much contorted when dry, and regularly broken.

104

3. Tortella humilis (Hedw.) Jenn.—Rather small plants (about 5, rarely 10 mm.
), green, or yellow-green. Leaves curled and contorted when dry, spreading when
moist, 2.2-3.5 mm. long, oblong-lanceolate from a broader base, narrowly to broadly
acute, short-apiculate, concave, not cucullate; costa short-excurrent; upper cells 6-7 yu.
Autoicous; perigonia (often several) small and delicate, stalked, in axils of upper leaves;
perichaetial leaves not much differentiated. Setae 7-18 mm. long; capsules 1.5-3 mm.
long; annulus deciduous in fragments; operculum 1.2-1.5 mm. long; peristome teeth
long, spirally wound together. Spores 7-11 u, finely roughened to nearly smooth.—n =
1526

nard, How to Know the Mosses, (ed. 2) fig. 94a-c. Grout, Mosses with Hand-lens and
Microscope, Pl. 32 (as T. caespitosa). Jennings, Mosses of Western Pennsylvania, (ed. 1) Pl. 13, (ed.
2) Pl. 17. Welch, Mosses of Indiana, fig. 71.—Fig. 284.

On bark at base of trees, rocks, soil, logs, and stumps in rather dry, partially shaded
places; probably limited to calcareous habitats. Europe and North Africa; Haiti, Jamaica, Brazil, and
the Galapagos Islands; British Columbia; reported from Arizona; ia read in eastern North
America from Vermont and Ontario to Minnesota and the Gulf State

CHEBOYGAN CO.-—Fairy Island (Douglas Lake), Colonial Point, near Mud Lake. MACKI-
NAC CO.—Ozar

Rare locally. Best recognized by relatively broad (but not cucullate) acumina and
autoicous inflorescences.

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Tortella ee 280. Leaf.

Tortella tortuosa. 281. pla 282. Leaves. 283. Cells at margin of leaf shoulder.
Torte lla h ae 284.

Tortella inclinata. a ae

105

4. Tortella inclinata (R. Hedw.) Limpr.—Rather small plants in yellowish or
brown, usually dense tufts up to about 1 cm. high. Leaves incurved, twisted and
somewhat curled when dry, rigidly erect or erect-spreading when moist, 2.2-3 mm.
long, oblong-lanceolate and scarcely tapered from a somewhat broader base, broadly
acute or obtuse and minutely apiculate, concave and cucullate; costa very shortly
excurrent, brown; upper cells 7-104. Dioicous; perichaetial leaves conspicuously
differentiated, yellow, elongate and erect at the base, curled at the tips when dry,
narrowly lance-subulate with a long-excurrent costa. Setae 15-27 mm. long; capsules
1.5-2 mm. long; annulus none; operculum 1-2 mm. long; peristome teeth long, spirally
wound together. Spores 7-11 u, smooth or finely roughened.

Grout, Moss Flora of North America 1: Pl. 84A (as T. inclinatula).—Fig. 285.

A calciphile growing on rock or soil, locally on sand in exposed places along the Great
eae “Europe, Caucasus, and Persia; Alaska, British Columbia, Alberta, Michigan, Ontario, and
Verm

T CO.—Cross Village, Sturgeon Bay, Cecil Bay. MACKINAC CO.—12 mi. W. of St.
Ignace. PRESQUE ISLE CO.—Mast Point, Lamprey Experiment Station, Evergreen Beach.

A rare species recognized by its occurrence on stabilized dune sands and by
short, broad, cucullate leaf tips.

Trichostomum tenuirostre (Hook. & Tayl.) Lindb. (fig. 286-287), a rock-growing
moss known from the Upper Peninsula, is similar to a Tortella, but the hyaline basal
cells do not extend upward along the margins, and the peristome teeth are erect rather
than spirally wound together. The leaf margins are usually somewhat wavy and often
sinuate and irregularly notched. The chromosome number is recorded as n = 13.

4. Bryoerythrophyllum Chen

Plants small to rather robust, in loose or dense, green to yellowish, brownish, or
red-brown tufts, usually brick-red below. Stems simple or forked. Leaves contorted
when dry, erect-or wide-spreading from an erect base when moist, narrowly lanceolate
to oblong-ligulate, broadly acute to apiculate, + keeled; margins revolute in the lower
% or more, often with a few irregular, pellucid teeth above; costa strong, subpercurrent
or, more often, ending as a pellucid mucro, with 2 stereid bands; upper cells
subquadrate or rounded-hexagonal, obscure, densely papillose on both surfaces, the
papillae often C-shaped (rarely pale, smooth, and incrassate along the upper margins);
lower cells rather conspicuously enlarged, oblong, lax, thin-walled, hyaline or red-
brown and pellucid. Setae terminal, elongate, erect; capsules erect and symmetric or
curved and inclined, cylindric or ovoid, smooth; annulus revoluble (at least in B.
recurvirostrum), operculum conic to conic-rostrate; peristome teeth 16, erect, joined at

ase aS a very low membrane, perforate along the median line or + bifid, reddish,
finely papillose. Calyptrae smooth, naked._The name means red-leaved moss and refers
to the brick-red color developed in many species of the genus. It was chosen to replace
the name Erythrophyllum, which had been used previously for an alga. The genus is a
segregate from Didymodon which, like Barbula, is the kind of catch-all that has been
called a portemanteau (or coat-rack) genus. The large, lax, and thin-walled basal cells,
the C-shaped papillae of upper cells, and the large annulus seem to justify segregation.

Bryoerythrophyllum recurvirostrum (Hedw.) Chen—Rather small plants (5-20,
rarely 30 mm. high), in loose or dense tufts, green, yellow-green, or brown above, rusty
or brick-red below. Leaves crisped and curled when dry, erect- or wide-spreading when

106

moist, 2-2.8 mm. long, oblong- to linear-lanceolate from an oblong, erect base, bluntly
acute to apiculate; margins revolute in the lower 2/3 or nearly to the apex, often
irregularly dentate near the apex; costa subpercurrent or (more often) shortly
excurrent as a pellucid mucro; upper cells about 8 uw wide, subquadrate and hexagonal,
firm-walled, with C-shaped papillae; lower cells hyaline to pale red-brown. Paroicous
and synoicous. Setae 6-16 mm. long, red; capsules 1.3-2.5 or rarely 3 mm. long, erect
and symmetric or occasionally slightly curved, cylindric, red-brown; annulus revoluble;
operculum 0.5-1 mm. long, obliquely long-rostrate (or sometimes merely conic to
short-rostrate); peristome teeth pale-brown or yellowish, linear-lanceolate, perforate or
+ bifid, minutely papillose to nearly smooth. Spores 15-16, smooth or finely
roughened.—n = 12+1, 13, 13+1.

Conard, How to Know the Mosses, (ed. 2) fig. 91a-<d (as Didymodon). Darlington, Mosses of
Michigan, fig. 57. Grout, Mosses with Hand-lens and Microscope, PI. 27 (as Didymodon rubellus).—
Fig. 288-291.

common calciphile, usually on moist soil or rocks, often in disturbed, open situations
(such as roadbanks), also flourishing in deep, undisturbed woods on wet logs, rotten stumps, ¢
Europe, Asia, Africa, New Guinea, New Zealand; Greenland to Alaska, south to California, Arizona,
Texas, Arkansas, and New Jersey; Mexico.

CHEBOYGAN CO.-—Gorge of Carp Creek, Iron Bridge, Bryants Bog, Passino School, near

Levering, Mill Creek. EMMET CO.—Middle Village, Cecil Bay, Big Stone Creek. MACKINAC

O.—Bois Blanc Island, Bush Bay, Detour, Ozark, Rexton. PRESQUE ISLE CO.—Hammond Bay,
Evergreen Beach, Clinton Lake.

The lax, thin-walled basal cells and irregularly dentate margins near the leaf tips
are distinctive. In the field, one can recognize the plants from the brick-red color
sometimes demonstrable only at the base of the tufts, sometimes throughout.

5. Didymodon Hedw.

Plants slender to fairly robust, in loose or dense, dull, olive-green, brown, or
red-brown tufts. Stems forked. Leaves + erect and often contorted when dry,
erect-spreading when moist, narrowly lanceolate and often subulate from a broader
base; margins entire, generally revolute; costa strong, ending near the apex, often filling
the subula and excurrent as a fleshy continuation of it, with 2 stereid bands; cells
small, rounded-quadrate, usually obscure, smooth or bluntly papillose; lower cells short

d firm, rectangular or somewhat elongate near the costa, yellowish or hyaline.
Perichaetial leaves scarcely differentiated. Setae terminal, elongate, erect; capsules erect
and symmetric, oblong-cylindric, smooth; annulus none or narrow and variously
differentiated; operculum conic-rostrate; peristome teeth 16, erect or slightly oblique,
joined at base, undivided or variously perforate, sometimes split nearly to the base into
2(or 3) filiform divisions, + papillose. Calyptrae smooth, naked.—The name refers to
twin divisions of the peristome teeth.

Didymodon rigidulus Hedw.—Rather small, dirty-green or yellow-brown, usually
densely tufted plants up to 15 mm. high, commonly producing clusters of very small,
brown, globose or rarely ovoid gemmae of 3-4 cells on branched stalks in upper leaf
axils (especially when sterile). Leaves somewhat contorted when dry, erect- to
wide-spreading when moist, 1-1.5 mm. long, lanceolate and gradually acuminate or
subulate-acuminate to a slender, blunt, fleshy tip; margins recurved in the lower
1/2-2/3, entire or obscurely sinuate-notched and thickened above, the tip of the lamina
also thickened; costa percurrent or disappearing in the fleshy apex, covered by short,

Ns -—
—aS
_—— y

cee

vas,
ae
FEN

Y ff

|

LA

I

Trichostomum tenuirostre. 286. Leaves. 287. Cells at margin of leaf shoulder.

Bryoerythrophyllum recurvirostrum. 288. Habit. 289. Leaves. 290. Cells at leaf tip. 291.
Cells of leaf base.

Didymodon rigidulus, 292. Habit. 293. Leaves. 294. Cells at leaf tip. 295. Brood-bodies. 296.
Portion of peristome.

green cells in the upper half; upper cells about 8-10 yu, irregularly rounded-quadrate,
thick-walled, slightly and often obscurely papillose. Dioicous. Setae 5-13 mm. long, red;
capsules 1-2 (rarely 2.5) mm. long, red-brown; operculum 0.6-0.9 mm. long, its cells in
oblique rows; peristome teeth somewhat inclined, deeply and often irregularly bifid,
yellow-brown, densely papillose. Spores 8-16 yu, very finely roughened.—n = 12, 13.

Breen, Mosses of Florida, Pl. 40 (as D. fuscoviridis). Grout, Moss Flora of North America 1:
Pl. 90B.—Fig. 292-296.

108

damp or wet, calcareous rocks and soil, often on rocks in streams. Circumpolar; New
Brunswick to Michigan and southward to Ohio, Tennessee, and Florida.

MACKINAC CO.—Mackinac Island, Bush Bay, Ozark. PRESQUE ISLE CO.—Rainy River
Falls.

Didymodon rigidulus is a species of little character, best recognized by its
axillary gemmae. Growing on calcareous rocks in and along streams, it often occurs
with Hyophila involuta and Fissidens obtusifolius (neither known from our area).
Didymodon rigidulus is similar in gametophytic appearance to Barbula fallax, which has
elongate cells covering the costa in the upper part of the leaf.

Didymodon tophaceus (Brid.) Lisa (fig. 297-299) is a widely distributed species
of wet, limey habitats. It has considerable variability but can be recognized by its
decurrent leaves, a most unusual feature in the Pottiaceae. It is active in tufa
formation, sometimes depositing around itself limey accretions which have been called
didymodontoliths! Tufa is a soft limestone which on hardening becomes a porous,
brownish stone called travertine. The Roman Coliseum was built of travertine (a word
derived from the name of an ancient Roman town, now known as Tivoli). Not many
years ago the more elegant American banks often had floors and_ staircases of
travertine. (This was an elegance associated with the gleaming brass cuspidors once
found in all public buildings.) Tufa is formed by photosynthetic plants removing
carbon dioxide from waters charged with soluble calcium bicarbonate, causing the
precipitation around them of insoluble calcium carbonate. Didymodon tophaceus has
been found in Michigan but not locally, owing no doubt to a dearth of wet, calcareous
habitats. Travertine formation by mosses in Oklahoma is described by Wilson & Guest
(Okla. Geol. Notes 21: 310-316. 1961) and Emig (Bryol. 21: 25-27. 1918) and in Utah
and Idaho by Flowers (Bryol. 36: 26-27. 1933). See also Berner (Rev. Bryol. et Lichén.
18: 59-65. 1949).

The chromosome number recorded for Didymodon tophaceus is n = 12.

Hyophila involuta (Hook.) Jaeg. & Sauerb. (fig. 300-304) has been collected on
boulders along Au Train Bay, Lake Superior, in Alger Co. H. involuta is recognized
with ease because of its habit, as well as its calcareous habitat. The oblong-spatulate
leaves, crowded in a somewhat rosulate fashion at the stem tips, are rolled, incurved,
and blackish when dry. The leaf margins are usually unevenly serrulate, and the upper
cells are small, arranged in transverse rows, and bulging on the upper surface.
Polymorphous brood bodies are commonly produced on branched stalks in leaf axils.

6. Barbula Hedw., nom. cons.

Plants small to medium-sized, gregarious or in dull, dark-green, brownish, or
reddish tufts, simple or forked. Leaves usually larger toward the stem tips, erect and
somewhat contorted when dry, narrowly lanceolate or oblong-ovate to lingulate, acute,
obtuse, or rounded-obtuse, often apiculate; margins entire, nearly always revolute,
sometimes to the apex or nearly so; costa subpercurrent to shortly excurrent, mostly
with 2 stereid bands; upper cells rounded-quadrate, obscure, sometimes smooth but
usually papillose on both surfaces (the papillae sometimes C-shaped); basal cells
subquadrate to short-rectangular, hyaline or yellowish, and usually smooth. Dioicous,
perichaetial leaves scarcely or strongly differentiated and convolute-sheathing. Setae
terminal, erect, elongate, red or rarely yellow; capsules erect and symmetric or nearly
so, oblong- or ovoid-cylindric; annulus none or, more often, present and persistent,
rarely revoluble; operculum high-conic to long-rostrate; stomata few, at extreme base

109

of the urn; peristome (rarely lacking or rudimentary) inserted below the mouth,
consisting of 16 teeth split to a low basal membrane into 32 filiform, papillose, reddish
divisions which are spirally wound together. Calyptrae smooth, naked—The generic
name means little beard, referring to the tufted peristome of 32 hair-like divisions.
Barbula and Didymodon are separated for convenience on the basis of the peristome—
long and twisted above a low basal membrane in Barbula, shorter and erect or merely
obliquely inclined in Didymodon. Associated with twisted peristomes are a spiral
arrangement of the cells of the operculum.

1. Leaves obtuse or broadly acute and minutely mucronate; margins slightly revolute near the
base; inner perichaetial leaves enlarged, pale, sheathing, truncate or rounded to short-
pointed; setae ye 1. B. er
1. Leaves acute, not at all or stoutly apiculate; margins revolute in the lower 1/2-2/3 o
more; peal leaves scarcely differentiated ; setae reddis
2. Leaves broadly or bluntly acute, stoutly ae culate 2. B. unguiculata
2. Leaves acuminate and acute, not apiculat 3. B. fallax

1. Barbula convoluta Hedw.—Small plants in dense, green or yellowish tufts up
to 5, rarely 15mm. high. Leaves incurved-contorted and somewhat spirally twisted
around the stems when dry, erect-spreading when moist, about 0.8-1.2 mm. long,
oblong-lanceolate or oblong-ligulate from a broader base, rounded-obtuse to broadly
acute, minutely apiculate (at least in upper leaves); margins slightly revolute at base,
plane above; costa ending at or just below the apex, with 2 stereid bands; upper cells
densely papillose and obscure, rounded-quadrate or rounded-hexagonal, about 7-9 u
wide; lower cells shortly oblong, firm-walled, pale, smooth. Inner perichaetial leaves
conspicuously enlarged, pale-yellow, convolute-sheathing, truncate to rounded or
broadly pointed. Setae 10-20 mm. long, slender, bright-yellow to straw-colored, twisted
when dry; capsules 1.2-1.8mm. long, erect or slightly curved and inclined, sub-
cylindric, slightly tapered from a broader base, very short-necked, light-brown, +
furrowed when dry and empty; annulus revoluble; operculum narrowly high-conic,
slightly inclined, about 1mm. long, its cells in oblique rows; peristome teeth
dusty-pinkish. Spores 7-9 u, smooth.—n = 11, 13, 14

Darlington, Mosses of Michigan, fig. 56. Grout, Mosses with Hand-lens and Microscope, Pl.
28. Jennings, Mosses of Western Pennsylvania, (ed. 1) Pl. 13, (ed. 2) Pl. 17.—Fig. 305-307.

bare, calcareous soil, often in open, disturbed places, common on roadbanks, along paths,
and in old fields. Circumpolar; oa and British Columbia to New Brunswick, south to Baja
California, Oklahoma, and Tenness

CHEBOYGAN CO.—Douglas Lake, Riggsville, Duncan Bay, Mill Creek, Afton Tower.
EMMET Ae —Brutus, Alanson, Pellston Hills, Van, Levering, Sturgeon Bay, Cecil Bay, Big Stone
Bay. MACKINAC CO.-—Bois Blanc Island, Mackinac Island, Garnet, McKays Creek, Detour,
Allenville, Ozark, Rexton. PRESQUE ISLE CO.—Ocqueoc Falls, Posen

The yellow setae and distinctly differentiated perichaetial leaves are taxonomical-
ly useful.

2. Barbula unguiculata Hedw.—Plants in light- or dirty-green tufts 5-25 mm. high.
Leaves erect and contorted when dry, erect-spreading when moist, 1-2 or 2.5 mm. long,
oblong-lanceolate from an ovate base, blunt or broadly acute, abruptly apiculate;
margins revolute in the lower 1/2-2/3; costa shortly excurrent, papillose at back, with
2 stereid bands; upper cells 8-10 u, rounded-hexagonal, + thick-walled, pluripapillose on
both surfaces; lower cells pale, smooth, rectangular, thin-walled. Dioicous. Setae
5-13 mm. long, reddish; capsules 1-2 mm. long; annulus none; operculum conic-

»

aU a\a
WA\ r) ¥

\ we
I Deer Cc

i

M\
Vie [i (|
NU | I \ Ls A.
VA ie
40305 \a ee

yop involuta, 300. Habit. 301. Leaves. 302.

Cross- o of leaf. 304. Brood-bodies

rbula convoluta. 305. Habit. 306. Leaves. 307. Upper cells of leaf.
eet unguiculata, 308. Habit. 309. Leaves. 310. Cells at leaf tip.
Barbula fallax. 311. Leaves. 312. Upper cells of lea

Didymodon tophaceus. 297. Leaves, 298. Cells at leaf tip. 299. Cells of leaf decurrency.
Hyophila Up

subulate; peristome reddish. Spores 9-13 u, smooth.—n = 1 P41, T241, 132 1341
13+2, 14, 14+2, 24.

Conard, How to Know the Mosses, (ed. 2) fig. 92. Grout, Mosses with Hand-lens and
Microscope, fig. 72. Jennings, Mosses of Western Pennsylvania, (ed. 1) Pl. 13, (ed. 2) Pl. 17. Welch,
Mosses of Indiana, fig. 73.—Fig. 308-310

n calcareous soil in disturbed places, particularly at roadsides and in old fields. Circum-
polar; riper widespread in the East from Nova Scotia to Minnesota, south to Florida and
Louisia

Eth
302 >

per cells of two leaves. 303.

AB /\

G / | ‘ Fs
ee s 5 | | ] /
BS Si0\ A
SSeeO\ ;
(OSELO)

111

CHEBOYGAN CO.—Biological Station Campus, Riggsville, Afton, Mill Creek, near Mackinaw
City. EMMET CO.—Maple River, Pellston Hills, Conway Bog. MACKINAC CO.—Allenville, Hessel,
Bois Blanc Island. PRESQUE ISLE CO.—Mast Point, Ocqueoc River, Posen.

The specific name refers to the apiculus, supposedly claw-like, which is particu-
larly strong on upper and perichaetial leaves.

3. Barbula fallax Hedw.—Plants in brownish tufts 5-20 (rarely 40) mm. high.
Leaves erect and + incurved or contorted when dry, erect-spreading when moist,
1-2 mm. long, lanceolate or ovate-lanceolate, acuminate, acute; margins revolute in the
lower half or nearly to the apex, with 2 stereid bands, not covered by short cells;
upper cells 7-11 yu, irregularly rounded-quadrate, oblate, and angular, thick-walled, with
a single low, often bluntly forked papilla on either surface; lower cells pale, smooth,
subquadrate to short-rectangular, moderately thick-walled. Setae 8-25 mm. long, red;
capsules 1-2 mm. long; annulus none; operculum long, conic-subulate; peristome teeth
dark- to orange-red. Spores 10-11 uw, smooth.—n = 9, 10, 11, 13, 13+1.

Grout, Mosses with Hand-lens and Microscope, PI. 29. Welch, Mosses of Indiana, fig.
74,—-Fig. 311-312.
On moist calcareous soil, particularly on roadbanks. Circumpolar; Greenland; British
Columbia; Nova Scotia to Minnesota and south to New York, Ohio, Missouri, and Louisiana.
CHEBOYGAN CO.-Biological Station campus, Camp Knight, Cheboygan, Afton, Mill Creek.
EMMET CO.—Pellston Hills, Conway Bog. MACKINAC CO.—Bois Blanc Island, Mackinac Island,
McKays Creek, Rexton. PRESQUE ISLE CO.—Ocequeoc Falls, Mast Point, Posen.

The specific epithet means false or deceitful and seems appropriate because the
species is a fooler, having little character and thus easily misidentified. The elongate
cells covering the upper surface of the costa help in distinguishing this from
Didymodon rigidulus.

Barbula michiganensis Steere was described from the Upper Peninsula, where it
grows on sandstone cliffs above Lake Superior at Pictured Rocks in Alger County:
Leaves ovate, acuminate, and acute; costa ending near the leaf apex, covered on the
upper side by elongate cells; very small, spherical gemmae (20-30 w in diameter and
consisting of 2-6 cells) abundant in leaf axils. It can be identified readily on
comparison with the illustrations in Grout’s Moss Flora, vol. 1, plate 89A.

Similar to B. fallax in appearance, and also having elongate cells covering the
costa, is B. reflexa (Brid.) Brid. It has leaves squarrose-recurved when moist and upper
cells sharply papillose on both surfaces. It has been found in Ontario and elsewhere
farther to the east.

7. Phascum Hedw.

Plants very small, simple or forked, scattered to gregarious. Leaves few, crowded,
larger toward the stem tips, oblong-lanceolate to ovate or obovate, acute to acuminate
and generally piliferous; margins usually entire, revolute; costa ending below the apex
to excurrent, with a single, dorsal stereid band; upper cells subquadrate to hexagonal,
pluripapillose (with C-shaped papillae) on 1 or both surfaces, or rarely smooth; lower
cells lax, thin-walled, pale, smooth, rectangular. Perichaetial leaves not much differ-
entiated. Setae very short, often curved; capsules immersed or shortly exserted, single

112

or sometimes paired, ellipsoidal to globose, bluntly apiculate; operculum and peristome
none; stomata sometimes present. Calyptra small or minute, cucullate or rarely
mitrate.-A name used by Theophrastus for a lichen, supposedly Usnea barbata, and
applied to mosses by Linnaeus in a sense which also included modern genera as diverse
as Cryphaea and Splachnum. Schreber, in 1770, in restricting the name to a more
modern sense appropriately quoted the ancients as saying that nature is never more
perfect than in small things—a statement rendered more meaningful to his contem-
poraries by the increasing use of lenses. (In those times a lens was a Flohglas used in
searching for fleas!) Speaking of things small, Stark (in A Popular History of British
Mosses, 1860) brought out of the Edinburgh Encyclopaedia an apt remark: “It has
been observed by a writer equally elegant and profound (Pascal) that ‘Man is placed in
the middle between two infinities— the infinitely great and the infinitely little—both of
which are incomprehensible to him.”

Phascum cuspidatum Hedw.—Small, green to yellow-brown plants 2-4 mm. high.
Leaves curved and contorted when dry, erect or erect-spreading when moist, keeled,
about 2-3.5 mm. long, oblong-lanceolate to oblong-obovate, acute to acuminate and
ending in a pale-yellowish, smooth awn of varying length; margins revolute along the
middle, sometimes nearly to the apex, entire; costa excurrent, brownish, smooth; upper
cells subquadrate to hexagonal, sometimes slightly elongate, about 13-18u wide,
smooth or more commonly with numerous C-shaped papillae on both surfaces
(occasionally less papillose at the margins in an indistinct border). Autoicous. Setae
very short, + curved; capsules immersed, pale-brown, shiny, 1-1.5 mm. long, ellipsoidal
or sometimes globose, stoutly and bluntly apiculate; stomata in 1 row at base of the
capsule. Spores spherical and slightly elongate, 22-44 u, brown, densely papillose to
tuberculate. Calyptra rather small to medium-sized, cucullate, smooth, naked.—n = 21,
26, 28, 29, 42, 52 (var. americanum, n = 26, 27+1-2, 28+1-2, 29-30).

ard, How to Know the Mosses, (ed. 1) fig. 48c & (ed. 2) fig. 75¢ (as var. americanum).
Dethetor, Mosses of Michigan, fig. 59. Grout, Mosses with Hand-lens and Microscope, fig. 68,
Moss Flora of North America 1: Pl. 91C (as var. americanum).—Fig. 313-315.

On limey soil in old fields, pastures, and other grassy places, at roadsides, for example, in
winter and spring (November-May). Not yet found in the Straits counties, no doubt because of its
late appearance, which is out of phase with the botanical activity associated with the Biological
Station. Europe, North Africa, and the Caucasus; British Columbia ” California, and Utah, across
the continent to Quebec, and widespread in the eastern United State

Phascum floerkeanum Web. & Mohr, found in several localities in southern
Ontario, may well occur in Michigan. The plants, though minute, stand out because of
their red-brown color. The leaves are only 1 mm. long, or less, concave, ovate, and
reddish-awned, and the upper cells of the leaf are 11-14 u in diameter. The inflores-
cences are paroicous, the spores yellowish and nearly smooth, and the calyptrae
mitrate. The species grows on bare patches of calcareous soil in old fields and other
disturbed places and matures its fruits between October and April.

8. Pottia (Reich.) Furnr.

Very small, scattered or gregarious plants. Leaves generally larger and more
crowded at the stem tips, erect to spreading, + concave or keeled, oblong-ovate,
oblong-lanceolate, or lingulate, apiculate to awned, rarely rounded at the apex; margins
plane or recurved, sometimes crenulate or serrulate above; costa mostly percurrent to
excurrent, with dorsal stereids only; upper cells hexagonal, pluripapillose on both
surfaces or rarely smooth; lower cells pale, smooth, oblong. Perichaetial leaves generally

113

not differentiated. Setae mostly + elongate; capsules usually exserted, erect and
symmetric, sometimes wide-mouthed, usually with a short neck; annulus present or
lacking; operculum low-convex to conic, short-pointed to rostrate; peristome often
none or rudimentary, sometimes consisting of 16 irregularly 2-3-forked or perforate (or
sometimes entire), erect teeth from a low basal membrane inserted at the mouth.
Calyptrae cucullate, smooth or papillose, naked._Named for Johann Friedrich Pott, a
professor of botany in Braunschweig.

Plants red-brown, up to 2mm. high, not rosulate; leaves 1-1.5 mm. long, lanceolate to
ovate- eee ending in re eddish apiculus or awn; upper cells of leaves 12-15 yu
dense y papillose; annulus none; operculum short-pointed; spores spinulose; aaa finely
papi . P. davalliana
Plants eee th or yellow-brown, up to 5mm. high, + rosulate; leaves 1.5- nee long,
blong-lanceolate to obovate, ending in a yellow-brown apiculus or awn; upper cells
15-22 wu, smooth (or occasionally indistinctly papillose); annulus of 1 row of cells,
persistent; operculum rostrate; spores finely papillose; calyptra smooth 2. P. truncata

1. Pottia davalliana (Sm. ex Drake) C. Jens.—Plants up to about 2 mm. high,
red-brown. Leaves erect and not much contorted when dry, erect-spreading when
moist, 1-1.6mm. long, ovate-lanceolate or lanceolate, gradually acuminate, acute,
ending in a strong, red or red-brown apiculus or short awn; margins entire, revolute
below, sometimes nearly to the apex; costa excurrent; upper cells hexagonal, 12-15 u
wide, densely papillose (with C-shaped papillae). Paroicous. Setae 1.5-3 mm. long,
yellowish, slender and + flexuose; capsules 0.4-1 mm. long, short-oval to hemispheric,
usually wide-mouthed when dry and empty, brown, thin-walled, vertically striate-
wrinkled when dry; annulus none; operculum low-conic, shortly and obliquely pointed;
peristome none. Spores 29-33 yx, somewhat ellipsoidal, densely spinulose, yellow-brown.
Calyptrae delicately papillose throughout.—n = 27+1, 28, 30.

Williams, Bryol. 64: 49, figs. 1-8.—Fig. 316-318.

oneer species found on bare spots of calcareous soil of old fields (especially second-year
sue fields), pastures, lawns, and roadsides; a fall ephemeral, sometimes also appearing in the

der conditions of unusual moisture. Europe, North Africa, Madeira; Ontario and
Michio

EMMET CO.-—Brutus. PRESQUE ISLE CO.—Posen.

The sterile plants can be spotted in the field by their red-brown color and
resemblance to scattered plants of Ceratodon purpureus in miniature. (It is coincidental
that they grow in places where one also finds such a weed as Ceratodon.

2. Pottia truncata (Hedw.) Ftirnr. ex BSG.—Plants about 3-5 mm. high, green to
yellowish or brownish. Leaves larger and more crowded toward the stem tips, 1.5-2.5,
rarely 3mm. long, erect or erect-spreading and somewhat contorted when dry,
erect-spreading when moist, oblong-lanceolate to oblong-ovate or obovate, acute or
acuminate, ending in a smooth, yellow-brown, stout apiculus or awn; margins plane (or
sometimes + recurved at the middle), entire or slightly crenulate near the apex; costa
short- to long-excurrent; upper cells irregularly hexagonal to short-rhombic, 15-22 u
wide, firm-walled, smooth or occasionally indistinctly papillose at back. Autoicous (or
paroicous, the antheridia naked in axils of upper leaves). Setae 2-5, rarely 8 mm. long,
orange- to red-brown; capsules 0.5-1.5 mm. long, obovoid or obconic and becoming
wide-mouthed when dry and empty, brown, thin-walled, finely wrinkled-striate when
dry; annulus of 1 row of cells, persistent; operculum obliquely long-rostrate from a
convex-conic base; peristome lacking (or occasionally rudimentary, delicate, and
papillose). Spores 22-32 u, spherical or nearly so, finely papillose.—n = 20, 25, 26, 52.

114

raee os ax =¢ a r

Phascum cuspidatum. 313. Habit . 314. Leaves. 315. Upper cells of leaf

Desmatodon obtusifolius. 322, Habit. 323. ‘Leaves. 324. Upper cells at leaf margin. 325.
Portion ee on and annulus. 326. Stomate.

Conard, How to Know the Mosses, (ed. 2) fig. 142. Darlington, Mosses of Michigan, fig. 60.
Grout, Mosses with Hand-lens and Microscope, Pl. 33 (as P. truncatula), Moss Flora of North
America 1: Pl. 95 (figs. 20-21). Jennings, Mosses of Western Pennsylvania, (ed. 1) Pl. 13, (ed. 2) Pl.
17.—Fig. 319-321.

A fall ephemeral (sometimes appearing in the summer under ‘unusually moist conditions), on
bare sot of calcareous soil in fallow fields (especially second- thes clover, alfalfa, or hay fields),
pastures, meadows, roadside ditches, etc. Europe, North Africa, Asia, Canary Islands; Uruguay;
Australia; SW aciasta and British Columbia; Nova Scotia to Ontario and Michigan, south to
Maryland.

115

CHEBOYGAN CO.—Tower. EMMET CO.-—Galloway Lake, Brutus. MACKINAC CO.—
Allenville.

Much smaller but similar in aspect, both gametophytically and sporophytically,
to Physcomitrium pyriforme. The excurrent costa of Pottia truncata will help in
distinguishing the two. Also, Physcomitrium pyriforme fruits in the spring, Pottia
truncata in the fall.

The var. major (Web. & Mohr) BSG (n = 52), also known as P. intermedia
(Turn.) Fiirnr., has been found in Ontario. The plants are somewhat larger, with leaf
margins usually more strongly recurved. The capsules are somewhat longer (1-1.2,
rarely 1.5mm. long), oblong-cylindric, not wide at the mouth or tapered to the
base._The differences are discussed and illustrated by Saito (Bull. Nat. Sci. Mus.
Tokyo 16: 60-91. 1973—see especially fig. 1, 2).

9. Desmatodon Brid.

Small plants in usually dense, green to yellow tufts. Stems simple or forked.
Leaves crowded, erect and often flexuose or contorted when dry, erect-spreading when
moist, concave or keeled-concave, oblong-ovate or oblong-lanceolate to obovate, acute
or obtuse, usually apiculate or hair-pointed; margins usually revolute below, sometimes
nearly to the apex, often serrulate toward the apex, frequently bordered by linear cells
below, often by short, yellowish, incrassate cells above; costa subpercurrent to
excurrent, with a dorsal stereid band and covered ventrally by a layer of large, often
papillose cells; upper cells hexagonal or oblong-hexagonal, thin-walled, smooth or more
commonly pluripapillose (with C-shaped papillae); lower cells lax, thin-walled, rectan-
gular, smooth, hyaline. Perichaetial leaves not much differentiated. Setae elongate, erect,
usually twisted when dry, smooth; capsules mostly erect, symmetric and oblong-
cylindric to narrowly cylindric, sometimes broadly oblong, asymmetric and strongly
inclined, smooth or finely wrinkled-striate; annulus present, usually persistent;
operculum conic, short- to long-rostrate, often obliquely so, its cells mostly in oblique
rows; stomata in 1 row at extreme base of the capsule; peristome inserted somewhat
below the mouth, split nearly to a low membrane into 2-3 filiform, papillose divisions
which are sometimes irregularly joined above, often + obliquely inclined or slightly
wound together, rarely none or rudimentary. Spores small, spherical, and smooth or
large, shortly ellipsoidal, and roughened. Calyptrae smooth, naked.—Bridel applied the
name, in an ininspired way, in reference to a narrow band formed by the fusion of
peristome teeth into a low basal membrane. (The basal membrane of Tortula is so
much more striking, that any emphasis on that of Desmatodon seems unfortunate!)

Desmatodon Aarau (Schwaegr.) Schimp.—Plants dull-green, gregarious or
caespitose, about 3-6mm. high. Leaves erect-spreading when moist, + incurved-
contorted when dry, 12 2 mm. long, oblong, oblong-lanceolate, or ovate-lanceolate,
rounded-obtuse, often apiculate (uppermost leaves rarely short-awned); margins entire,
revolute from near the base almost to the apex, sometimes strongly so; costa terete,
subpercurrent, sometimes disappearing in the pale apiculus; upper cells 8-11 wide,
irregularly hexagonal, obscure, densely papillose, usually not differentiated at the
margins but sometimes somewhat paler and more incrassate. Autoicous (but usually
appearing dioicous). Setae 6-12 mm. long; capsules (1)1.5-2(2.5) mm. long, narrowly
(or rarely broadly) cylindric, erect or sometimes slightly curved, finely wrinkled-striate
when dry and empty; operculum 0.5-0.6 mm. high, stoutly conic, sometimes conic-
rostrate and + inclined, its cells in oblique rows; peristome teeth whitish to pale

116

orange-red, rather weakly developed, about 140-275 uw high, irregularly divided into 2
slender prongs of unequal length. Spores 8-11(16), greenish-yellow, smooth,
spherical.—n = 26.

Conard, How to Know the Mosses, (ed. 2) fig. 82a-c. ae Mosses of Michigan, fig. 61.
Grout, Mosses with Hand-lens and Microscope, Pl. 34 (as D. arenaceus), Jennings, Mosses of
Western Pennsylvania, (ed. 1) Pl. 13 (as D. arenaceus), (ed. %) Pl. 17. Welch, Mosses of Indiana, fig.
80.—Fig. 322-326.

On damp, shaded rock, on cliffs or boulders, stone walls, bridge supports, etc., rarely on soil;
mostly commonly on sandstone, but probably preferring limey substrates. Europe; Caucasus and
Asia Minor; Greenland; Alaska to New Brunswick, south to California, Colorado, Texas, and
Tennessee.

CHEBOYGAN CO.—Mill Creek. MACKINAC CO.—Mackinac Island.

10. Tortula Hedw., nom. cons.

Plants small to large, in dull, green, glaucous-green, or brownish tufts, rarely
scattered, often tinged with red, simple or forked, usually radiculose below. Leaves
often larger and more crowded at the stem tips, sometimes in rosulate clusters, erect-
to wide-spreading or squarrose to recurved when moist, usually erect, folded lengthwise
and twisted around the stem when dry, lingulate, obovate or elliptic (usually widest
above the middle), broadly rounded or obtuse, rarely acute, often apiculate or awned;
margins usually entire, occasionally bordered by short, incrassate, pale or colored cells
or by linear cells, nearly always revolute throughout or in part; costa usually strong,
often terete, mostly percurrent to excurrent, with dorsal stereids only; upper cells
hexagonal, thick-walled, usually strongly pluripapillose and obscure (rarely smooth or
weakly papillose), the papillae mostly circular or C-shaped; lower cells usually large,
rectangular, lax, thin-walled, smooth and hyaline, yellow-brown or reddish, usually in
rather abruptly differentiated groups. Perichaetial leaves mostly not or slightly dif-
ferentiated. Setae terminal, rt elongate, erect; capsules cylindric to ovoid-cylindric,
erect and symmetric or somewhat curved; annulus persistent or revoluble; operculum
narrowly conic or conic- oe peristome teeth 16 (rarely lacking), inserted below
the mouth, usually long, divided to a basal membrane of varying length (generally a
high, tesselated tube) into 32 filiform, papillose teeth typically wound together in
l-several spirals. Calyptrae smooth, naked.—The name (like Tortella) is a diminutive
meaning twisted, in reference to the long, twisted peristome teeth. In England, Tortula
has been called, for similar reasons, the SCREW MOSS. The basal membrane of the
peristome is, in most species, very high and marked off in a curious pattern of squares.

Leaves erect-spreading when moist, acute to subulate-acuminate; cells smooth or weakly
1. T. mucronifolia

Leaves squarrose-recurved when moist, ending in a hyaline hair-point; cells densely papillose
2. T. ruralis

1. Tortula mucronifolia Schwaegr.—Plants loosely tufted or mixed with other
mosses, dull, dark-green, sometimes tinged with yellow or brown, 5-15 mm. high.
Leaves keeled and contorted when dry, erect-spreading when moist, 2.5-3.5, rarely
5mm. long, oblong or oblong-obovate, acute or acuminate; margins revolute in the
lower 1/2-2/3; costa excurrent as a stout, yellow mucro or hair-point; upper cells
(13)16-21(28) u wide, irregularly subquadrate and rounded-hexagonal, with firm,
yellowish walls + thickened at the corners, smooth or nearly so, smaller and more
incrassate near the margins, often forming a + distinct border of short, yellowish cells;
basal cells elongate-rectangular, not sharply delimited. Autoicous. Setae 8-22 mm. long;

iat,

capsules (2)3.5-4.5 mm. long, erect but usually somewhat curved, narrowly cylindric;
annulus persistent; operculum 1.8-2.4 mm. long, erect, conic-subulate; peristome teeth
long, the basal membrane pale, pinkish or orange, becoming white with age, about as
long as the teeth which are closely wound together and orange or rarely pink. Spores
11-15 yu, very finely papillose.—n = 12, 24, 26, 30.

Conard, How to Know the Mosses, (ed. 2) fig. 146. Grout, Mosses with Hand-lens and
Microscope, Pl. 36. Welch, Mosses of Indiana, fig. 81. Fig. 327-329.

On calcareous soil in crevices ie cliffs and boulders and frequently in sheltered niches on
banks or on and among rocks. Cir ee Greenland to Alaska, south to New York, the Great
Lakes, Iowa, New Mexico, and Californi

CHEBOYGAN CO.—Fairy Island Cees Lake), Colonial Point, Mill Creek. EMMET
CO.—Middle Village, Cross Village, Cecil Bay

The nearly smooth upper cells and the poorly delimited basal cells are distinctive,
in a negative way, as compared with most species of Tortula.

2. Tortula ruralis (Hedw.) Gartn., Meyer & Scherb.—Rather small to moderately
robust plants in loose or dense, hoary tufts, often blackish when dry, typically reddish
below and bright-green above when moist. Leaves loosely appressed, folded, and
somewhat twisted around the stem when dry, wide-spreading to squarrose-recurved
when moist, 3-4 mm. long (including the awn), keeled, oblong, obtuse or rounded-
obtuse and ending abruptly in a long, white, spinulose awn (+ reddish at base); margins
revolute nearly to the apex; costa red or brown, densely spiculose at back; upper cells
9-13, green and often opaque, densely papillose on both surfaces, with C-shaped
papillae. Dioicous. Setae 10-18 mm. long; capsules 2.5-3 mm. long, long-cylindric,
sometimes slightly curved; annulus persistent; operculum high-conic; peristome teeth
long, twisted above a high basal membrane, yellowish to salmon-pinkish. Spores 7-11 y,
very finely roughened.—n = 12 (and, as T. intermedia, n = 12, 13, 13+1, 23+2, 2842.

, How to Know the Mosses, (ed. 2) fig. 83d-e. Darlington, Mosses of Michigan, apes
62. Grout, Mosses with Hand-lens and Microscope, Pl. 35. Welch, Mosses of Indiana, fig. 83.—Fig.
330-332.

On soil or rocks in dry, sunny, calcareous habitats, locally common on sand along the shores
of the Great Lakes. Circumpolar; throughout much of North Ameri

CHEBOYGAN CO.-—Biological Station campus, Colonial Point, Mill Creek, Indian River.
EMMET CO.-—Cross Village, Middle Village, Five Mile Creek, Ramona Beach, Cecil Bay, Big Stone
Bay, Waugoshance Point. MACKINAC CO.—Bois Blanc Island, Meee Island, St. Martins Bay,
McKays Creek, Bush Bay, Epoufette, Ozark, Caffey Corner. PRESQUE ISLE CO.—Hammond Bay,
Hoeft State Park, Bells Landing, Mast Point, Rainy River Falls.

Locally common as a sand-binding species, especially along shores of the Great
Lakes. Tortula ruralis is a pronounced calciphile, easily recognized by the brick-red
color of the plants, at least in the older parts, and by the leaves which are
squarrose-recurved when moist. The high, checkered membrane of the peristome, the
laxly oblong basal cells in rather well-marked groups rounded above, and the circular
and C- avy papillae are characteristic of most species of Tortula.

So our material in eastern North America may -be T. intermedia (Brid.)
Berk. (= Tm ralis var. crinita De Not.). A decision requires a re-evaluation of a circle
of confusion eee around 7. ruralis. As Steere, in Grout’s Moss Flora, said: ‘‘This
species is most apt to be taken for an underdeveloped form of T. ruralis, from which it
may be distinguished by the darker color, the spreading, but not squarrose leaves, , the
leaf margins plane above, the smaller leaf cells, and the more emarginate leaf apex

118

aes _ a
ee Seo
y

Tortula mucronifolia, 327. Habit. 328. Leaves. 329. Upper cells of leaf.
Tortula ruralis, 330. Habit. 331. Leaves. 332. Upper cells of leaf.
Tortula papillosa, 333. Habit. 334. Leaves. 335. Upper cells of leaf.

Tortula norvegica (Web.) Wahl. ex Lindb. is a western, montane species much
like 7. ruralis. It is characterized by a lightly roughened costa which projects as a
slender, shiny, red, nearly smooth awn. It has been found once in Michigan, on
Summer Island in Delta County. It is a calciphile.

Tortula papillosa Wils. ex Spruce (fig. 333-335) is a small moss growing scattered
on trunks of trees (and especially on bark at base of roadside elms and maples). It has
oblong-obovate, rounded-obtuse, abruptly piliferous leaves bearing an abundance of
globose, multicellular gemmae on the upper surface of the costa. Its cells are smooth
or unipapillose. It has been found in more southerly parts of the state.

Tortula pagorum (Milde) De Not., also small and scattered on bark of trees, has
not yet been found in Michigan. It is widespread throughout much of the eastern
United States, where it is associated with the activities of man, being found most

119

commonly in towns and elsewhere near habitations. It is no doubt an introduced
species, as only female plants have been found in eastern North America; only male
plants occur in Europe. (Fruiting plants have been found in Australia.) Anderson’s
systematic study of the American distribution (Bryol. 46: 47-66. 1943) is worth noting
as an example of biologically interesting information gleaned from floristic study.
Tortula pagorum has hyaline awns on its leaves, densely papillose cells, and leaf-like
propagula in the leaf axils.

caulon muticum (Hedw.) C. M. var. rufescens (Jaeg.) Crum is a minute moss
which is widespread in eastern North America and might be discovered locally in
ephemeral situations, such as sandy or clayey soil in old fields, vacant lots, or mowed
grass at roadsides, in winter and spring. The plants are only 1-2 mm. high, bulbiform,
yellowish to reddish-brown, not trigonous; leaves 1-2 mm. long, concave but not
keeled, broad and oblong-ovate to elliptic, cuspidate-apiculate and recurved at the tip;
margins recurved near apex of leaf; costa narrow, brown, subpercurrent or shortly
excurrent; upper cells 2-3:1, smooth. Capsules immersed, spherical, without
operculum or peristome. Spores spherical, 30-52 u, yellowish, smooth.

Aloina rigida (Hedw.) Limpr. and A. brevirostris (Hook. & Grev.) Kindb. (fig.
336-337) may yet be discovered in Michigan. They are pioneers of bare, limey silt,
especially on road banks and river bluffs. They resemble an Aloé in miniature because
of fleshy, concave leaves. The costa is hidden below a mass of green, branched
filaments. The two species may be distinguished, as follows: A. rigida—leaves
1.5-2.5mm. long, oblong-lingulate; upper cells 34:1; capsules 1.8-2.5 mm. long,
narrowly cylindric; operculum long and narrow and usually erect. A. brevirostris—leaves
1 or rarely as much as 1.8mm. long, suborbicular to broadly lingulate; upper cells
1-3:1; capsules 1-2 mm. long, broadly cylindric to ovoid-cylindric; operculum short-
rostrate and oblique.

Pterygoneurum ovatum (Hedw.) Dix. (fig. 338-339) should be sought for on bare
patches of calcareous soil in lawns, pastures, and clover fields. It is best known by the
lamellae borne on the upper surface of the costa.

Scopelophila ligulata (Spruce) Spruce (fig. 340-343), a rock-growing moss some-
what resembling a Tortula (also known as Merceya), has been found once in Michigan’s
Upper Peninsula. It has usually been found on copper-bearing rocks and is sometimes
called the COPPER MOSS for that reason. It has been suggested that it is actually linked
to the sulphur often associated with copper and possibly betraying chemosynthetic
ability. Interesting references to copper mosses include Brooks (New Zeal. Jour. Bot.
9: 674-677, 1972), Persson (Rev. Bryol. et Lichén. 17: 76-78. 1948; Jour. Hattori Bot.
Lab. 17: 1-18. 1956), Schatz (Bryol. 58: 113-120. 1955), and Shacklette (U.S. Geol.
Surv. Bull. 1198-G, 1967). The moss, not fruiting in North America, is rather small
and has lingulate leaves obtuse or rounded at the apex, becoming somewhat bordered
because of thickened cell walls with age; the upper cells are small, isodiametric,
smooth; the basal cells are laxly rectangular within but noticeably narrower and firmer
at the margins.

Whitehead and Brooks (Bryol. 72: 501-507. 1970) showed that aquatic mosses
can accumulate uranium.

GRIMMIACEAE

Plants small to relatively robust, perennial, in dull, mostly dark, rigid tufts or
mats, mostly on dry, exposed rocks, rarely on wet rocks or in water. Stems

120

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342

JO
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Aloina brevirostris. 336. Habit. 337. Leaf.

Pterygoneurum ovatum, 338. Habit. 339. Leaf.

Scopelophila ligulata. 340. Habit. 341. Leaves. 342. Upper cells at margin of leaf, 343. Basal
marginal cells of leaf.

erect-ascending or prostrate, forked or fasciculately branched, rarely with short,
tuft-like branches in a pinnate arrangement. Leaves in many rows, not at all
complanate, rarely secund, usually very hygroscopic, rarely crisped when dry, mostly
lanceolate and acute, sometimes oblong-ovate and acute to rounded-obtuse, often
piliferous; costa single, well developed; upper cells generally small and obscure,
sometimes papillose, often bistratose toward the apex or at the margins (sometimes
rounded-quadrate and sinuose-nodose or rectangular to linear and very strongly
sinuose); lower cells elongate, often somewhat to strongly sinuose-nodose. Setae
terminal or lateral, erect or curved, short or elongate; capsules subglobose to cylindric,
immersed or exserted, mostly erect and symmetric; annulus sometimes differentiated;
operculum mostly rostrate; peristome usually present, single, the teeth 16, long and
slender to short and broad, entire or variously cleft or perforate, mostly papillose.
Calyptrae generally small, mitrate or cucullate, naked, sometimes campanulate, occa-
sionally plicate—A family of rock-growing mosses.

Basal leaf cells short, quadrate or oblong, with walls straight or only slightly nodose 1. Grimmia
Basal cells long-linear, with strongly nodose lateral walls 2. Rhacomitrium

1. Grimmia Hedw.

Plants small or medium-sized, in dull, rigid, typically dark-green, brown, or
blackish tufts or mats, sometimes reddish, often hoary. Stems erect or ascending,
forked or fasciculately branched. Leaves in many rows, often larger at the stem or
branch tips in comose tufts, appressed when dry and rarely crisped or spiraled, usually
strongly hygroscopic (on wetting usually suddenly bending backward before becoming
erect and finally erect- or wide-spreading), mostly lanceolate and gradually tapered to
an acute apex, sometimes oblong or ovate, occasionally blunt or rounded-obtuse,
mostly piliferous, generally keeled; margins plane or revolute on | or both sides below,
mostly entire; costa strong, subpercurrent, frequently entering the base of the awn;
ee cells generally small and obscure, rounded-quadrate with walls frequently sinuose

nodose, mostly smooth; lower cells mostly elongate-rectangular, frequently +
sinuose- aoe near the costa, frequently pale and subquadrate at the basal margins,
sometimes with thickened cross-walls. Perichaetial leaves not much differentiated or
sometimes conspicuously enlarged. Setae terminal, very short to elongate, erect to
arcuate; capsules immersed to exserted, obovoid to subglobose to oblong-cylindric,
mostly symmetric, smooth or ribbed; annulus present or lacking; operculum often
rostrate, occasionally falling with the columella; peristome teeth usually present, the
teeth flat, broad or lanceolate, sometimes subulate, often blunt or truncate, sometimes
perforate or cleft above, + papillose. Calyptrae small, mitrate or cucullate, naked, not
or indistinctly plicate-—Named in honor of J. F.C. Grimm, a physician and botanist of
Gotha, Germany. A genus of rock-growing mosses in small, rigid, often blackish or
dark-brown tufts or cushions, often confused with Orthotrichum. Orthotrichum grows
on bark more often than rocks, has a rather large, campanulate-mitrate, plicate, and
often hairy calyptra, a fairly well-marked sterile neck, 8 or 16 peristome teeth, and
often an inner peristome as well. (The single peristome of Grimmia is homologous to
the inner peristome of Orthotrichum, having originated from a similar primordial cell
layer.) In Orthotrichum, the texture of the plants is softer, and the walls of leaf cells
are never sinuose or nodose-thickened.

1. Leaves plane at the Balsa perichaetial leaves similar to upper vegetative leaves; seta
excentrically attached to the capsule, curved; capsule bulging at 1 side at base; columella
not falling with the operculum 3. G. plagiopodia
1. Leaves revolute at the margins; perichaetial leaves noticeably enlarged; seta centrally
attached, straight; capsule symmetric; columella falling with the operculum

3. Plants in small, dense tufts about 1 (rarely 3.5) cm. high; leaves 1.3-2 mm. long, usually
1

entire (except for the awn), smooth at back . G. apocarpa
3.Plants in rather large, loose tufts 2-7 cm. high; leaves 1.8-2.5 mm. long, repand-
denticulate above, coarsely papillose at back of the costa la. G. apocarpa var. ge

2. Plants broadly cae to ovate, muticous or ete any hyaline-apiculate; spore
9-17 yw; calyptra m
4, Plants rather eal 1-2(rarely 4) cm. high; leaves broadly lanceolate to ovate-lanceolate,
usually entire 2. G. alpicola
4, Plants ie robust, 2-5 cm. high; leaves ovate-lanceolate, repand-denticulate above
2a. G. alpicola var. rivularis

Geneva Sayre’s key to North American species of Grimmia (Bryol. 55: 251-260.
1952) is useful in determining sterile material.

1. Grimmia apocarpa Hedw.—Plants in small, dense, dark-green or brownish tufts
1-3.5 cm. high. Leaves erect when dry, erect-spreading when moist, 1.3-2 mm. long, +
keeled, lance-acuminate, ending in a rather shbrt, hyaline, denticulate awn but often
muticous; margins entire, revolute nearly to the apex; costa not papillose at back,

U22

disappearing in the base of the awn; upper cells + bistratose toward the apex and at
the margins, 5-7 u, subquadrate, with thick, sinuose walls; basal cells subquadrate and
short-rectangular. Autoicous; perichaetial leaves erect, conspicuously enlarged, broadly
ovate. Setae about 0.5mm. long, straight; capsules immersed, symmetric, broadly
ellipsoidal, wide-mouthed, about 1 mm. long, smooth; annulus none; columella falling
with the operculum which is obliquely short- to long-rostrate from a convex base;
stomata few, at extreme base of the urn; peristome teeth lanceolate, red, + perforate
above, finely papillose. Spores 7-12 u, finely roughened. Calyptrae small, cucullate.—n
= 12, 13, 14, 26 (as Schistidium confertum).

ard, How to Know the Mosses, (ed. 2) fig. ae f. Pa Mosses of Michigan, fig.
63. Gr ae Mosses with Hand-lens and Microscope, PI. ings, Mosses of Western Pennsyl-
vania, (ed. 1) Pl. 14, (ed. 2) Pl. 18. Welch, Mosses of aa fig. a —Fig. 344-346.

both siliceous and calcareous rocks, especially in dry, exposed places. Nearly cosmopoli-
tan; Alaska to Greenland, south to California, Arizona, New Mexico, Texas, Arkansas, and Georgia.

This widespread, supposedly ubiquitous species has been collected in the Upper
Peninsula but not in our area of Michigan. It has been confused and confounded, here
and abroad, with G. alpicola. The latter, common on the limited rocky habitats of the
Straits region, has broader, muticous or minutely hyaline-tipped leaves, mitrate calyptrae,
and somewhat larger spores.

la. Var. stricta (Turn.) Hook. & Tayl.—Fairly robust plants in loose, dark-green
to red-brown tufts 2-7cm. high. Leaves erect-spreading and sometimes + secund,
1.8-2.5 mm. long; margins usually + notched-denticulate near the apex; costa coarsely
papillose at back toward the apex; cells usually coarsely unipapillose on both
surfaces—n = 13, 26.

Fig. 347-349.

On granitic or calcareous rocks in shady places. Circumpolar; across aa south to
Arizona oa Mexico in the West, Georgia, eee and the Great Lakes in the Eas

MACKINAC CO.—Ozark, McKays Creek.

The relatively robust plants in loose, red-brown tufts and the button-like papillae
at back of the costa are good characters of this well-marked variety. The leaves are
usually shortly piliferous, but not always. The reddish color and larger, looser tufts will
distinguish such a form from G. alpicola and the narrower leaves from the large tufts.
of G. alpicola var. rivularis.

. Grimmia yates Hedw.—Plants in dark-green to brown, loose or dense, rigid
tufts 1S rarely 4c high. Leaves erect when dry, erect- spreading when moist,
1.5-2.5 mm. long, ele’, ovate-lanceolate to ovate, muticous or sometimes hyaline-
tipped; margins revolute below, thickened and entire or sometimes + notched above;
costa percurrent or nearly so, smooth or occasionally + papillose at back above; upper
cells 7-9 uw, irregularly subquadrate, thick-walled. Autoicous; perichaetial leaves erect,
longer and broader, 3-3.5mm. long, often hyaline-tipped. Setae 0.8-1 mm. long,
straight; capsules immersed, erect and symmetric, wide-mouthed, 1-1.5 mm. long;
columella falling with the obliquely short- to long-rostrate operculum; peristome teeth
red, + perforate above, finely papillose. Spores 9-18 uw. Calyptrae small, mitrate.—n =
13, 14.

Fig. 350-353.

123

On both calcareous and acidic rocks, usually in dry, exposed habitats. Circumpolar; across
Canada to Alaska, south to California and Texas, widespread in eastern North America, south to
Arkansas and North Carolina; Mexico and Ecuador; Australia.

CHEBOYGAN CO.-—Biological Station campus, Colonial Point, Mill Creek. EMMET CO.—
Pellston Hills, Cross Village, Middle Village, Wilderness State Park, Cecil Bay. MACKINAC
CO.—Bois Blanc Island, Mackinac Island, St. Martins Point, McKays Creek, Bush Bay, Epoufette,
Ozark, Caffey Corner. PRESQUE ISLE CO.—Grand Lake, Ocqueoc Falls, Rainy River Falls, Posen.

InQ O03
peace

Grimmia apocarpa. 344. Leaves. 345. Upper cells of leaf. 346. Inner basal cells of leaf.
rimmia apocarpa vat. stricta, 347. Leaves. 348. Tip of leaf in profile. 349. Upper cells of

leaf at margin.
immia alpicola, 350. Habit. 351. Leaves. 352. Upper cells at margin of two leaves. 353.

Capsule and perichaetial leaves.

immia alpicola var. rivularis. 354. Leaf. 355, Cells at leaf tip.
i ] i . 358. Upper cells of leaf. 359. Capsule with

rimmia plagiopodia. 356. Habit. 357. Leaves
perichaetial leaf and dry capsule (at upper right).

124

Distinguished from G. apocarpa by somewhat broader leaves which are bluntly
muticous or at most hyaline-apiculate, spores of a greater size range, and mitrate
calyptrae, this species grades into the var. rivularis, which occupies wet habitats and
has broad, notched-denticulate leaves.

2a. Var. rivularis (Brid.) Wahl.—Robust, dark-green or brown plants, 2-5 cm.
high. Leaves 2-2.5mm. long, broadly ovate, muticous; margins revolute below,
thickened and repand-denticulate above; costa smooth or sometimes + papillose at
back. Autoicous; perichaetial leaves 3.5-4 mm. long. Setae 0.6-0.8 mm. long; capsules
1-1.5 mm. long.—n = 13.

Fig. 354-355.

On both calcareous and non-calcareous rocks in stream beds, often in places periodically
inundated. Europe and Asia; Greenland; Newfoundland to Pennsylvania and Michigan; Alaska and
the Yukon to Arizona, New Mexico, and northern Mexico; Costa Rica

CHEBOYGAN CO.—Mill Creek.

The habitat (on wet rocks) and broad, muticous leaves with upper margins
thickened and repand-denticulate are marks by which this variety may be recognized,
at least in its more extreme manifestations.

One might expect the var. dupretii (Thér.) Crum in calcareous areas of the state.
It is a small form (3-5 mm. high), with leaves only 1-1.3 mm. long and entire and with
spores only 9-11 uw in diameter. The leaves are sometimes, but not always rounded-
obtuse.

Sayre (Bryol. 55: 251-259. 1952) made some pertinent remarks on the
taxonomy of the genus which “is an extremely variable one and hard and fast lines
between species are not always easy to recognize. This factor is the source of much of
the synonymy, and especially of the periodic rise and fall of varietal names. Some
authors have noted the variations with such care that the typical form of the species is
lost sight of.... The author accepts a reasonable amount of responsibility if the key

anomalas.” It is my opinion that much of the trouble associated with Grimmia
apocarpa has resulted from a misunderstanding of the extreme variability of G. alpicola
and a confusion of the two, with a resulting tendency to segregate unimportant
expressions of G. apocarpa on the basis of meaningless comparisons with what is
actually G. alpicola! The history of difficulties presented by G. alpicola is summarized
by Sayre (Bryol. 49: 3-7. 1946), although she goes farther than I and considers G.
alpicola a mere variety of G. apocarpa.

Grimmia agassizii (Sull. & Lesq. ex Sull.) Jaeg. & Sauerb. occurs on wet rocks in
or near streams or waterfalls in the Upper Peninsula. It is related to G. alpicola but has
nearly flat, narrow leaves, with nearly parallel sides, and margins plane and not
thickened.

3. Grimmia plagiopodia Hedw.—Small plants (up to 10mm. high), in dense,
dark-green to brown, somewhat hoary tufts. Leaves crowded and imbricate, scarcely
altered when dry, keeled-concave, the lower leaves about 1-1.5 mm. long, oblong-ovate
and obtuse to acute and muticous or shortly hyaline-pointed, the upper and _peri-

125
chaetial leaves about 2-2.5 mm. long, broadly oblong or oblong-lanceolate, ending in a
long, hyaline, denticulate hair-point which is usually broad at the base; margins erect
and entire; costa channeled, subpercurrent or disappearing in the awn; upper cells
unistratose, green, 7-12 4, rounded-quadrate or somewhat longer than broad, with
thick, smooth or slightly irregular walls; basal cells pale or hyaline, subquadrate in
lower leaves, shortly rectangular in upper leaves, with rather thin but firm, straight
walls. Autoicous. Setae shorter than the capsules (about 0.2-0.3 mm. long), slender,
curved or curled; capsules immersed, erect or somewhat inclined, nearly hemispheric or
broadly oblong, wide-mouthed, bulging on 1 side at the base, about 1-1.5 mm. long;
annulus none; operculum low-convex, mammillate to stoutly and bluntly apiculate;
exothecial cells large, rather thin-walled; stomata large, at extreme base of the capsule;
peristome teeth 16, inserted below the mouth, spreading to reflexed when dry,
lanceolate, freely cleft and variously perforate (but not cribrose) at the apex,
red-brown, finely papillose. Spores 9-14, smooth. Calyptrae covering only the
operculum, mitrate, rostrate, deeply lobed, not plicate, smooth.—n = 26.

Grout, Moss Flora of North America 2: Pl. 5 (below).—Fig. 356-359.

ically on sandstone, sometimes on limestone or concrete of bridge abutments, culverts,
or old Peer Europe; Caucasus; central Asia; Alaska to California and New Mexico and, in
the East, rare and scattered from Ontario to the Dakotas, south to Wisconsin, Ohio, and Maryland.

CHEBOYGAN CO.-—Mill Creek, Camp Knight (Douglas Lake).

A rare moss resembling G. apocarpa because of brownish leaves ending in hyaline
hair-points. On closer inspection, however, it proves to be very distinctive owing to the
wide-mouthed capsules bulging on one side at base and thus causing the short, curved
(or even curled) setae to be excentrically attached. The specific epithet refers to the
curved seta.

Grimmia anodon BSG, broadly distributed from South Dakota westward, has a
narrow annulus and no peristome. It has been found in southern Ontario.

The role of lichens as pioneers on bare rock has no doubt been over-emphasized.
Bryophytes such as Andreaea, Rhacomitrium, and Grimmia may precede lichens.
Catherine Keever (Ecol. 38: 422-429. 1957) found that she could easily grow such a
primary invader, Grimmia laevigata, on crushed granite in petri dishes by regeneration
from shoots or plant fragments. Such regenerants grew better on granite if lichens were
absent. She found that most herbarium specimens of Grimmia laevigata retain viability
as long as three years, and some 10-year old specimens will grow. The fact that she
was eee to germinate the spores suggests that this (and perhaps other) rock-growing
mo may depend less on spore formation than on regeneration of fragments
Sere er eile broken off during dry periods.

Also occurring in the Upper Peninsula of Michigan are the following:

mmia affinis Hoppe & Hornsch. ex Hornsch. Autoicous; leaves keeled,
piliferous, revolute on 1 side below; setae straight, 2.5-3.5 mm. long, longer than the
emergent or slightly exserted capsules.

Grimmia hartmanii var. anomala (Hampe ex Schimp:) Monk. Leaves keeled, the
uppermost leaves of occasional branches slenderly acute and hyaline-tipped or short-
awned, the others blunt or rounded and muticous, often bearing at their tips large,
pale-green or orange-brown, globose brood-bodies; upper cells usually covered by
longitudinal cuticular ridges.

126

Grimmia pilifera P.-Beauv. Distinguished from G. affinis by dioicous inflores-
cences; leaf margins revolute on | or both sides below; setae only 1 mm. long, not
longer than the immersed capsules.

Grimmia pulvinata (Hedw.) Sm. ex Sm. & Sowerby, a western species found
disjunctively in several midwestern localities, grows in small, hoary cushions. The leaves
are keeled and long-awned, with margins recurved on one or both sides, and the basal
cells have walls rather thin and not or only slightly sinuose. The setae, erect and
twisted when dry, are curved when moist; and the capsules, immersed to exeouied. are
brown and wrinkled-striate when dry.

Grimmia torquata Hornsch. ex Grev., known from the granitic rocks of Ontario’s
Lake Superior region, has awned leaves which are twisted, curled, and spirally ranked
when dry; the costa is shiny when dry; and red, globose gemmae are produced in
chains from the back of the costa on the lower part of leaves terminating each year’s
rowth

Grimmia unicolor Hook. ex Grev. Leaves rigidly erect or imbricate when dry,
very concave, oblong-ligulate from a broader base, rounded or rounded-obtuse and
somewhat cucullate at the apex, muticous; seta 3-4 mm. long, + flexuose-curved when
dry, erect when moist; capsules exserted. It somewhat resembles G. agassizii but has a
dioicous (rather than autoicous) inflorescence and leaves concave and more or less
cucullate at the tip, with cells from the shoulders upward in two or more layers.

2. Rhacomitrium Brid.

Fairly robust plants in dull, often rigid, extensive mats. Stems creeping or
crowded and ascending, freely branched, sometimes with short, tuft-like branches in a
pinnate arrangement. Leaves erect or spreading, oblong-lanceolate, acute or acuminate
and often piliferous, or rarely blunt or rounded at the apex; costa ending below the
apex to excurrent or disappearing in the awn; cells sometimes bistratose toward the
apex or at the margins above, incrassate with nodose lateral walls at least in the leaf
base, sometimes throughout, linear below, short, pale, and straight-walled at the basal
angles, subquadrate or short-rectangular to linear above, smooth or sometimes
papillose. Dioicous; perichaetia lateral. Setae elongate, usually straight; capsules erect
and symmetric, oblong-ovoid to oblong-cylindric, smooth, wrinkled-striate, or obscurely
plicate; annulus well developed; operculum long-rostrate; peristome teeth erect, fused
at base and + completely split to or below the middle into 2 terete or filiform,
papillose divisions; pre-peristome present, consisting of a low and inconspicuous,
smooth, hyaline membrane adhering to the base of the teeth. Spores spherical.
Calyptrae conic-mitrate, often rough (at least at the apex), naked, not plicate.—The
name indicates a torn cap, referring rather inappropriately to the mitrate calyptra
which is somewhat lobed at base. The nodose lateral walls of the cells, at the base of
the leaf and sometimes throughout, are unique features of the genus.

Leaves ending in a toothed but not papillose hair-point; cells oe 1. R. heterostichum
Leaves ending in a toothed, papillose hair-point; cells papillos 2. R. canescens

1. Rhacomitrium heterostichum (Hedw.) Brid.—Tufts green to brown, sometimes
+ hoary, freely branched. Leaves 2-2.5 mm. long, lanceolate, acuminate, ending in a
flat, denticulate, hyaline awn of various lengths, sometimes merely hyaline-tipped or
even muticous; margins entire, revolute on 1 or both sides, bistratose above; costa

127

ending near the apex, not ridged at back; upper cells nodose, smooth (or in section
sometimes appearing papillose because of thick walls), 1-1.5:1. Setae twisted when dry,
slightly curved or flexuose when moist, 3-8mm. long; capsules oblong-cylindric,
1.2-2 mm. long, smooth; peristome teeth unequally bifid nearly to the base, brown,
rather coarsely papillose. Spores 9-12 u, papillose. Calyptrae smooth or + scabrous
above.—n = 13, 14; 2n = 28

Conard, How to Know the Mosses, (ed. 2) fig. oe Grout, Moss Flora of North America
SRI 22s pentaee Mosses of Western Pennsylvania, (ed. 2) Pl. 52 (as var. gracilescens).

siliceous rocks. Circumpolar; Alaska to Greenland and south ee California, the Great
akes, and North Car vast No ete ay ene in our area where it has been collected once on an
erratic boulder at Cecil Bay b ; Katz, July 25, 1 . (The species occurs naturally in ae
ke Superior region of Nichicah Be Ontario, and it has been collected on Sugar Islan
Chippewa County.)

2. Rhacomitrium canescens (Hedw.) Brid.—Plants in loose or dense, green,
yellowish, or brownish mats. Stems curved-ascending, sparsely fasciculate and often
with few to many tuft-like horizontal branches. Leaves erect and often + curved with
flexuose-spreading awns when dry, erect to wide-spreading when moist, 1.5-2.5 mm.
long, lanceolate to ovate, slenderly to broadly acuminate, ending in a short to long,
hyaline, serrulate, papillose awn; margins unistratose, revolute in the lower 2/3 or

10-25 mm. long, smooth; capsules 1.2-2.5 mm. long, ovoid-cylindric, irregularly plicate
when dry and empty; annulus tardily deciduous; operculum up to 2 mm. long,
long-subulate; peristome teeth red-brown, very long, split nearly to the base into 2
filiform, densely papillose divisions. Spores 7-9 u, smooth or nearly so. Calyptrae
papillose above.—n = 12, 13.

Conard, How to Know the Mosses (ed. 2), fig. 62. Grout, Mosses with Hand-lens and
Microscope, fig. 62.—Fig. 360-363.

A pioneer growing on sand or gravel at shores or in soil among rocks. Circumpolar;
Greenland to Alaska and south = California, Colorado, Michigan, and New York.

und once in our area, near Bryants Bog (at the southwest side of Douglas Lake),
Ghebone Co., on a sand mound next to a ditch in open oak-aspen woods (D. Caporello, 1971).

Unmistakable because of flat, toothed, papillose awns and papillose leaf cells.
The fo, ericoides (Brid.) Ménk. (fig. 364) is evenly pinnate because of an abundance of
short, horizontal, tuft-like branchlets. Though distinctive in appearance, this seems to
be only an ecological expression. Sand-binding plants seem to have few or no

(Hedw.) Brid. at the specific level for the following reasons: In northwestern Europe, it
sometimes grows mixed with R. canescens and then can be recognized by branching
alone. It also has somewhat narrower leaves with a rather longer costa and cells with
papillae only about half as high. It tends to be somewhat more oceanic in distribution
(though otherwise similar in ecology).

Rhacomitrium lanuginosum (Hedw.) Brid., not yet found in Michigan, has leaves
ending in long-decurrent, hyaline hairpoints with coarse knot-like teeth at their
margins. The leaf cells are linear throughout and (except in the awn) smooth. It is a

128

re
IOC

DD OAD OSS OS
CS On SR =~
SNaetn = —s

i
3

Rhacomitrium canescens. 360. Habit. 361. Leaf. 362. Upper cells of leaf. 363. Cells near
base of leaf.
Rhacomitrium canescens, fo. ericoides. 364. Habit.

common species of arctic and alpine tundra, especially among rocks in talus or fell
fields; it may occur somewhere in the Lake Superior region.

In Keweenaw and Marquette Counties, we have R. patens (Hedw.) Hub.,
distinctive species with the costa conspicuously ridged at back and the upper i
short, not nodulose. The long-standing dispute over placing this ae in oe or
Rhacomitrium is neatly resolved by referring it to a monotypic , as Drypto
patens (Hedw.) Brid., as proposed recently by Crundwell (Trans. Brit, ta Soc. 6(2).
323-324. 1971).

Other species of the Upper Peninsula, both with muticous leaves, are R. aciculare
(Hedw.) Brid., with broad, elliptic leaves which are obtuse to rounded and dentate at
the apex, and R. fasciculare (Hedw.) Brid., with narrowly lanceolate, entire leaves
which are slenderly tapered to a narrow, blunt tip.

129
HEDWIGIACEAE

Fairly robust, rigid plants in loose or dense mats, freely and irregularly to
subpinnately branched, crowded and erect-ascending or creeping (or pendent with long,
2-3-pinnate branches), sometimes stoloniferous. Leaves crowded in many rows, ap-
pressed when dry, + spreading when moist, broad and concave, ovate-lanceolate,
oblong-ovate, or panduriform, acute, short-acuminate, or piliferous, sometimes plicate,
entire (or rarely abruptly toothed at the base of the awn), bordered or unbordered;
costa none; cells finely to coarsely pluripapillose on both surfaces, with firm or
incrassate walls, subquadrate or oblong to linear above, linear at the middle of the base
and variously differentiated at the basal angles. Perichaetia mostly at the ends of main
stems or branches; perichaetial leaves + elongate, erect; paraphyses filiform. Setae short
to elongate, erect; capsules immersed to exserted, erect and symmetric, subglobose to
ovoid or obovoid, with a short, thick neck; annulus none; operculum low-convex and
sometimes umbonate to conic and apiculate or rostrate; stomata in the neck; peristome
none (or in Cleistostoma, consisting of 16 short ‘teeth deeply inserted on a low

accepted as the starting point for the nomenclature of mosses (exclusive of Sphagnum).
Professor of botany at Leipzig, he was a skilled microscopist and the first to
understand the sexual function of antheridia and archegonia of mosses. (Goethe,
himself a botanist, recorded in his diary, in 1797, a visit to Hedwig’s laboratory where
he saw elegant preparations and drawings.)

Hedwigia ciliata (Hedw.) P.-Beauv.—Coarse and usually rather robust plants in
dull mats, green, yellow, or brown and usually grayish or glaucous when dry, often
hoary. Stems spreading or curved-ascending, irregularly branched, not stoloniferous.
Leaves erect or appressed (or sometimes subsecund) with apices sometimes wide-
spreading when dry, spreading when moist, 1.3-2.3mm. long, sometimes faintly
striolate when dry, broadly oblong-ovate, acute to acuminate, nearly always ending in a
hyaline tip or a short to long, broad, hyaline acumen; margins revolute in the lower
2/3 or nearly to the hyaline tip or acumen, irregularly spinulose-serrate at the apex,
not bordered; upper cells subquadrate to short-oblong or rhombic with + thickened

noticeably poroce toward the base; median basal cells linear, smooth, yellow, in-
crassate, and porose, the basal marginal cells green and quadrate in many rows.
Autoicous; perichaetial leaves conspicuously long-ciliate along the upper margins, the
cilia hyaline and nodose or toothed. Setae very short; capsules deeply immersed,
subhemispheric with a short and inconspicuous neck which is longitudinally wrinkled
when dry, wide-mouthed, the rim usually incurved or inrolled when dry, smooth,
pale-brown below, red-brown toward the mouth, shiny when dry; peristome none;
operculum plano-convex and sometimes umbonate. Spores 20-30 u, obscurely angled,
finely papillose. Calyptra very small, conic-mitrate, hairy.—n = 10, 10+1, 11, 21, 22.

Breen, Mosses of Florida, Pl. 42 (figs. 14). Conard, How to Know the Mosses, (ed. 2) fig.
63. Darlington, Mosses of soe fig. 95. Grout, Mosses with Hand-lens and Microscope, ae ar
& 51 (as H. albicans). Jennings, Mosses of Western Pennsylvania, (ed. 1) Pl. 30, (ed. 2) P
Welch, Mosses of Indiana, fig. 3» 89. —Fig. 365-369.

On dry, granitic boulders in the open or in partial shade (in open woods and often in
scrubby pastures or at roadsides), rarely on calcareous rocks. Nearly cosmopolitan; widespread in
North America.

130

we >) ©)
{aC

JEQXUE IAS

( 368

Hedwigia ciliata, 365. Habit. 366. Leaves. 367. Upper cells of leaf. 368. Inner basal cells of
leaf. 369. Perichaetial leaves.

CHEBOYGAN CO.—Reeses Bog, Douglas Lake, Colonial Point. EMMET CO.—Pellston Hills,
Good Hart, Middle Village, Cross Village, Mackinaw City Hardwoods, Cecil Bay. MACKINAC
CO.—Mackinac Island, Ozark.

The plants are rigid and gray when dry but become surprisingly soft, green, and
otherwise altered in appearance because of spreading leaves when moist. The length of
the hyaline leaf points is highly variable. The specific name refers to long, jointed,
hyaline cilia fringing the perichaetial leaves.

Although generally placed near Orthotrichum in a group of more or less
pleurocarpous families, Hedwigia seems to me enough like Grimmia to be accom-
modated in the same family. Since it lacks a peristome one can only guess at its true
relationship, whether to the Haplolepideae (as I suspect) or the Diplolepideae.

FUNARIACEAE

Plants small to medium-sized, light-green or yellowish, gregarious to loosely
tufted. Stems erect, simple or forked, radiculose only at base. Leaves small below,
larger and more crowded above, usually concave, elliptic to oblong-lanceolate, oblong-
ovate, or obovate, usually acute or acuminate; costa single, ending near the apex to
excurrent; cells smooth, pale, large, lax and thin-walled, oblong-hexagonal above,
sometimes narrower at the margins, rectangular below. Mostly autoicous or sometimes
polygamous, rarely synoicous or paroicous; perigonia usually at the ends of | or rarely
2 short basal branches, antheridia often in the perichaetia as well; perigonia with
short, usually clavate paraphyses; perichaetia terminal, with few, small, filiform

131

paraphyses, or none. Setae mostly elongate; capsules sometimes immersed, usually
long-exserted, erect and symmetric or inclined and asymmetric, usually with a short
but distinct neck; annulus often present; stomata usually differentiated, restricted to
the neck, consisting of a central slit in a single, rounded guard cell; operculum usually
differentiated; peristome lacking, single or double, the teeth 16, lanceolate, usually
papillose-striate, the endostome consisting only of segments opposite the teeth.
Calyptrae smooth, small and mitrate or large, cucullate, and long-rostrate, often
becoming deeply lobed at base.
Capsules erect, symmetric, wide-mouthed (especially when dry and empty), smooth
1. Physcomitrium
Capsules inclined, strongly asymmetric, small-mouthed, sulcate 2. Funaria

1. Physcomitrium (Brid.) Flirnr.

Small, gregarious or loosely tufted, light-green or yellowish plants with simple or
forked stems. Leaves erect or erect-spreading, contorted when dry, oblong-lanceolate to
oblong-ovate or obovate, bluntly pointed to acute or acuminate; costa ending near the
apex to short-excurrent; cells large and lax, oblong-hexagonal above, rectangular below,
sometimes + swollen at the basal angles. Autoicous or occasionally polygamous;
perigonial branch basal, with clavate paraphyses, antheridia sometimes in the peri-
chaetium as well; perichaetia with few, small, filiform paraphyses. Setae mostly
elongate, erect; capsules mostly exserted, erect and symmetric, hemispheric to
urceolate or short-pyriform, wide at the mouth, with a short, usually distinct neck;
annulus narrow and persistent or broader and revoluble in fragments; operculum
convex, sometimes apiculate or rostellate; peristome none. Calyptra large, long-beaked,
inflated at base and deeply split into 3 (or less often 2 or 4) lobes.—The name refers
to the inflated calyptra. By the time of maturity, however, the calyptra becomes
deeply lobed and loses its bladder-like nature (and the name seems more suitable to
Funaria, in which the calyptra retains its inflated character).

Physcomitrium pyriforme (Hedw.) Hampe—Plants 3-10, rarely 25 mm. high.
3-5 mm. long, oblong-lanceolate to oblong-ovate, acuminate; margins plane,
subentire to bluntly serrulate or serrate above the middle; costa ending near the apex
or very rarely short-excurrent; upper cells oblong-hexagonal, paler and narrower at the
margins or sometimes + inflated in 1-2 rows as an indistinct border; lower cells large
and oblong, often somewhat inflated at the basal angles. Autoicous or polygamous.
Setae 4-14, rarely 30 mm. long; capsules 1-2 mm. long, globose-pyriform,,variable when
dry and empty but usually constricted below the flaring mouth and at the short neck;
annulus narrow, persistent; exothecial cells transversely oblong in 8-17 suboral rows.
Spores 24-42, rarely 50 yu, papillose.—n = 9, 26, 27, 36, 52, 54 (and, in experimental
material, 18, 24, 69, 71, 72 and 2n = 48+2, 128, 173).

een, Mosses of Florida, Pl. 50. Conard, How to Know the Mosses, (ed. 2) fig. 162a-d.
Bae eg Mosses of Michigan, fig. 67. Grout, Mosses with Hand-lens and nae ele fig. 96 (as
P. turbinatum), Moss Flora of North America 2: Pl. 30 (as P. turbinatum). Jennings, Mosses of
Western Pennsylvania, (ed. 1) Pl. 16 & ae 2) Pl. 20 (as P. turbinatum). Welch, Gee of Indiana,
fig. 95 (as P. turbinatum).—Fig. 370-374

n disturbed soil, frequently along roads and in old fields, in spring. nae re Islands,
North Africa, Europe a and the Caucasus; widespread in eastern North America; Mex

CHEBOYGAN CO.—Colonial Point, Inverness Township, Passino School, Duncan Bay.
EMMET CO.—Brutus. PRESQUE ISLE CO.—Posen

Physcomitrium pyriforme. 370. Habit. 371. Leaves. 372. Cells at apex of leaf. 373. Capsule
(dry). 374. Calyptra.
Physcomitrium immersum. 375. Habit.
Physcomitrella patens, 376, Habit. 377. Leaf. 378. Upper cells of leaf. 379. Exothecial cells.
380. Stomata at base of capsule.
anorrhegma serratum. 381. Habit. 382. Leaves. 383. Cells of upper part of leaf. 384.
Exothecial cells.

An interesting feature is the apical inflorescence in which one can demonstrate
the whole developmental sequence from archegonial initial to fertilized egg and young
embryo stages. (Antheridia found in basal buds and sometimes also mingled with
archegonia in the apical inflorescence are not so easily observed.) This spring weed is
sometimes called the TOP MOSS or the URN MOSS, both names referring to the shape
of the capsules on drying (contracted below the wide mouth and at the short neck).
The specific name means pear-shaped and is not as appropriate as turbinatum (or
top-shaped), an American synonym formerly in common use.

133

Physcomitrium hookeri Hampe is a prairie species with a large revoluble annulus.
(It has not been found in Michigan.)

Physcomitrium immersum Sull., (fig. 375; n = 54), not yet found in Michigan,
has immersed capsules. The large, beaked, and inflated calyptra, as well as the
structural features of urn and operculum, make the generic position immediately clear.
It is similar in some ways to two other members of the family which occur in
ephemeral situations farther south in Michigan: Physcomitrella patens (Hedw.) BSG
(fig. 376-380; n = ca. 16) has capsules rupturing irregularly, with exothecial cells very
lax and thin-walled. Aphanorrhegma serratum (J. Hook. & Wils. ex Drumm.) Sull. (fig.
381-384) has operculate capsules with collenchymatous exothecial cells. Both genera
have small calyptrae.

The genera of Funariaceae are well marked and easily distinguished, and yet
numerous intergeneric hybrids, some actually produced in the laboratory, indicate close
cytological and genetic ties. Crosses of Funaria with Physcomitrium and of Physco-
mitrium with Physcomitrella indicate that the absence of peristome or operculum does
not necessarily evidence a great degree of evolutionary divergence. Hybrids in the
Pottiaceae, between Weissia and Astomum, and the Ditrichaceae, between Ditrichum
and Pleuridium, further illustrate the close relationship of inoperculate and gymno-
stomous genera or species to operculate and peristomate ancestors. Hybrids between
Physcomitrium and Aphanorrhegma provide other evidence of relationship within the
Funariaceae, and Funaria-Entosthodon hybrids de-emphasize the importance of asym-
metry of capsules as a generic distinction. References to the literature on supposed
hybrids occurring in nature may be found in papers by Timm (Hedwigia 67: 1-44.
1927) and Wettstein (Bibl. Gen. 1: 1-38. 1925). Allen (Bot. Rev. 1: 269-291. 1935)
lists numerous hybrids induced between species of Funariaceae. Other information on
hybridization is given above, under Weissia. Osada (Misc. Bryol. et Lichen. 4: 118.
1967) illustrated a possible hybrid having the capsule of a Polytrichum (such as P.
formosum) but the distinctive leaf lamellae of Pogonatum alpinum.

Natural hybrids are unknown in liverworts, although they have been induced
between various species of Sphaerocarpos (see Allen, Bot. Rev. 1: 269-291. 1935).

Apogamy—the production of sporophytes directly from gametophytic tissue
rather than a normal process of fertilization—has been induced in a number of species,
for example, in Physcomitrium coorgense, an Indian species, by Lal (Phytomorph. 11:
263-269, 1961; also in Maheshwari & Swamy, Plant Tissue and Organ Culture, pp.
363-383. 1963). A wounded seta may regenerate as a diploid protonema eventually
forming leafy shoots which may reproduce sexually or sometimes vegetatively, forming
capsules on stalk-like extensions of the shoot. This type of apogamy, which begins as
apospory, has been reported in Phascum cuspidatum, Physcomitrium pyriforme, P.
pyriforme X Funaria hygrometrica and the reciprocal, Desmatodon ucrainicus, D.
randii, Grimmia pulvinata (incompletely), and Tetraphis pellucida. Additional data on
apogamy are provided by Bauer (Planta 46:604-618. 1956; Handbuch der Pflanzen-
physiologie 18: 235-256. 1967), Chopra & Rashid (Bryol. 70: 206-208. 1967), Hughes
(A Study of Apogamy, 1958; New Phytol. 68: 883-900. 1969), Lazarenko (Dokl. Bot.
Sci. 134: 177-179. 1961 and 162: 962-964. 1965; Cytol. y Genet. 1965: 158-173.
1965; Dopov. Akad. Nauk Ukraine SSR 11: 1524-1525. 1963 and ser. B, 5: 459-461.
1969), Lazarenko et al. (Dopov. Akad. Nauk Ukraine SSR 10: 1381-1384. 1961),
Rashid & Chopra (Phytomorph. 19: 170-178. 1969), Springer (Zeitschr. Indukt. Abst.
Vererb. 69: 249-262. 1935), and Wettstein (Ber. Deutsch. Bot. Ges. 60: 399405.
1942). Other species in which apogamy has been observed are Funaria hygrometrica,
Pottia lanceolata, Pottia intermedia (= P. truncata var. major), Amblystegium
juratzkanum (A. serpens var. juratzkanum), Leptodictyum riparium, and Brachy-

134

thecium campestre. Viable spores have been produced in some apogamously produced
sporophytes (see Lazarenko, 1965). It should be noted that sporophytes have been
produced apogamously only on diploid gametophytes.

The production of diploid gametophytes by the regeneration of sporophytic
tissue (apospory), especially from segments of setae, is easily induced in the laboratory
and may also occur in nature, thus accounting for pairs of species differing most
notably in sex and chromosome number, as in Mnium, for example. It was Pringsheim
who first induced apospory in mosses (Monatschr. K. Akad. Wiss. Berlin 1486:
425-429. 1876). Reference should be made to the Marchals’ early work on the
regeneration of setae (Bull. Acad. Roy. Belg., Sci. 1907: 765-789. 1907; 1909:
1249-1288. 1909; 1911: 750-756. 1911) and Wettstein’s work on hybridization and
genetics of aposporously produced polyploids (Zeitschr. Indukt. Abst. Vererb. 33:
1-236, 253-257. 1924; Bibl. Genet. no. 10, 1928; pp. 233-272 in Verdoorn’s Manual of
Bryology, 1932). LaRue (Papers Mich. Acad. Sci., Arts & Lett. 11: 225-241. 1929)
demonstrated regeneration from the setae of 14 species of Michigan mosses. Lowry
(Mem. Torrey Bot. Club 20, part 2, pp. 142. 1948) provided cytological evidence that
some species of Mnium could have originated through some such process of regenera-
tion in nature. Apospory probably occurs commonly in nature, providing for polytopic
origins of some species, but observations are few indeed: Bower (Science, N.S. 20:
524-536. 1904) reported that Brizi in 1892 had observed regeneration from a capsule
of Funaria which had been accidentally buried. Other literature on apospory includes
Allen (Bot. Rev. 1: 269-296. 1935), Bauer (Ber. Deutsch. Bot. Ges. 70: 424-432.
1957), Bornhagen (Beih. Bot. Centralbl. 46: 407-434. 1930), Lazarenko (Dokl. Bot.
Sci. 162: 962-964. 1965; Cytol. y Genet. 1: 15-26. 1967), Lowry (Bryol. 67: 147-149.
1954; Amer. Jour. Bot. ae 410-414. 1954), Moutschen (Lejeunea 15: 41-50. 1951;
Congr. Intern. Bot. Rapp. et Comm., sect. 16: 114-121. 1955), Redfearn & Meyers
(Bryol. 52: 197-201. 1949), pea (Bot. Zeit. 34: 690. 1876), Schratz (Biol. Zentralbl.
44: 593-623. 1924), Schweitzer (Flora 116: 1-72. 1923, and Stahl (Bot. Zeit. 34: 690.
1876)

The fact that apogamy can be induced by altering culture conditions (see
especially Lal, Phytomorph. 11: 263-269. 1961, and Wettstein, Ber. Deutsch. Bot. Ges.
60: 399-405. 1942) and apospory can result from wounding proves that gametophytes
and sporophytes have the same genetic potential and are indeed homologous. Hence
the antithetic theory of the origin of the sporophyte which assumes a lack of
homology loses strength.

Apospory is probably not an important factor in the evolution of mosses but
more significant surely than in liverworts where the sporophytes are smaller and
simpler and the setae short-lived. Apospory has been induced in a few liverworts by
Bornhagen (Biol. Zentralbl. 46: 578-586. 1926), Lang (Ann. Bot. 15: 503-510. 1901),
Raudzens & Matzke (Amer. Jour. Bot. 55: 1190-1196. 1968), Matzke & Raudzens
(Proc. Nat. Acad. Sci. 59: 752-755. 1968; in Gunckel, Current Topics in Plant Science,
pp. 117-119. 1969), and Schwarzenbach (Arch. Julius Klaus-Stift. Vererb. Soz.-anthr.
Rassenhyg. 2: 91-141. 1926).

The special techniques of counting moss chromosomes are given by Snider (Mich.
Bot. 9: 67-71. 1970) and Lewis (Trans. Brit. Bryol. Soc. 3(2): 279-284. 1957).

Heald’s study of regeneration of gametophytic structures (Bot. Gaz. 26: 169-210.
1898) is worthy of reference.

2. Funaria Hedw.

Small to medium-sized plants, gregarious or in bright-green or yellowish, loose
tufts. Stems usually simple. Leaves small below, becoming larger and more crowde

135

above, erect, concave, oblong-ovate to obovate, usually acute or acuminate; margins
erect, entire or bluntly serrulate above the middle; costa usually ending near the apex,
sometimes excurrent; cells large, lax and thin-walled, oblong-hexagonal above, scarcely
differentiated at the margins, oblong below. Autoicous; perigonia at the ends of 1 or
sometimes 2 short, basal branches; perigonia with clavate paraphyses; perichaetia
apparently lacking paraphyses. Setae elongate, capsules inclined to pendulous (rarely
suberect), asymmetric, usually curved, broadly pyriform, small-mouthed, mostly plicate
or sulcate when dry and empty; annulus large and revoluble or none; operculum
convex or rarely conic, its cells in oblique rows; peristome double, the 16 teeth
obliquely directed, lance-acuminate, papillose-striate, often strongly trabeculate, fre-
quently appendiculate, particularly above and united in a latticed disc at the apex;
segments lanceolate, opposite the teeth and 1/3-2/3 or sometimes nearly as long as the
teeth. Calyptra large, cucullate, long-rostrate, inflated at base —The name refers to the
cord-like twisting of the dry seta of Funaria hygrometrica, commonly known as the
CORD MOSS.

Entosthodon differs from Funaria only in having erect capsules and a lesser
development of the peristome. Bopp (Zeitschr. Bot. 42: 331-352. 1954; Ber. Deutsch.
Bot. Ges. 69: 455-468. 1956; Zeitschr. Indukt. Abst. Vererb. 88: 600-607. 1957) has
found that phototropic curvature of such capsules is controlled by the calyptra (and in
a purely mechanical way, since a dead calyptra has the same effect as a living one).
This fact, plus the induction of intergeneric hybrids, presents a challenge to the
traditional separation of Entosthodon from Funaria (which is still a considerable
convenience). Entosthodon does not occur as far north as Michigan.

Funaria hygrometrica Hedw.—Plants 4-10 mm. or more high, tall and laxly foliate
throughout or, more often, shorter, nearly naked below and densely foliate and +
bulbiform above. Upper leaves 1-3 mm. long, usually crowded and erect, very concave,
oblong-ovate to broadly obovate, acute or apiculate to short-acuminate, entire or
somewhat serrulate above; costa subpercurrent to short-excurrent; cells somewhat
inflated and oblong-hexagonal (1-2:1) above, longer and oblong below. Setae 12-45,
rarely 80mm. long, slender and flexuose, usually strongly hygroscopic; capsules
2-3 mm. long, broadly pyriform, curved and very asymmetric, horizontal or pendulous,
deeply sulcate, strongly oblique at the small mouth (usually bent just below the mouth
when dry and empty); annulus revoluble; operculum low-convex; peristome teeth
brown and papillose-striate below, pale and papillose above, strongly trabeculate,
appendiculate above, joined as a latticed-disc at the tips; segments about 2/3 as long as
the teeth, pale-yellow, finely papillose-striate. Spores 11-24 y, spherical, smooth.—n =
14, 20, 21, 28, 56 (and, experimentally, 12, 28, 30, 60, 240).

Conard, How to Know the Mosses, (ed. 2) fig. 163a-d. Darlington, Mosses of Michigan, fig.
68. Grout, Mosses with Hand-lens and Microscope, fig. 97, Moss Flora of North America 2: Pl. 38.
Jennings, Mosses of Western Pennsylvania, (ed. 1) Pl. 27, (ed. 2) Pl. 21. Welch, Mosses of Indiana,
fig. 97.—Fig. 385-389.

A weed, very common on wet soil in disturbed habitats, often in burned-over places,
sometimes on old wo rock, common in lawns and greenhouses. Cosmopolitan (but largely
replaced by var. calvescens in tropical latitudes).

CHEBOYGAN CO.—Reeses Bog, Gorge of Carp Creek, Colonial Point, Bryants Bog, Inverness
Township, Mill Creek, Cheboygan, Duncan Bay, Weber Lake. EMMET CO.—Brutus, Maple River,
Cross Village, Wycamp Creek, Good Hart, Cecil Bay. MACKINAC CO.-—Bois Blanc Island, Mackinac
Island, Nunns Creek, Ozark. PRESQUE ISLE CO.—Ocqueoc River, Mast Point, Lamprey Experi-
ment Station, Evergreen Beach.

The structure and life cycle of Funaria hygrometrica are well known to anyone
who has had a course in botany. It is called the CORD MOSS, as explained above, and in

Vi nit
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388
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| i ; \ 391 393
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ath

392
Funaria se ea ce 385. Habit. 386. Leaf. 387. a at leaf apex. 388. Peristome as
viewed from open end of capsule, Snel: 389. Calypt
ea sae crassinervium, 390. Habit. 391.
Ephemerum cohaerens, 392. Lea

Ephemerum spinulosum. 393. Leaf.

Leaf.

Marshall’s Mosses and Lichens at least, the WATER MEASURING CORD MOSS! The
French call it LA CHARBONNIERE (a charcoal or coal peddler); and Mrs, Britton aptly
referred to it as CINDERELLA as it loves ashes and is often found on wet soil where
there A been a camp fire.
mouth of the capsule is remarkably oblique, and the large annulus is
Scene to watch as it slowly unrolls in a drop of water. The peristome teeth are
curved-sigmoid and attached at their tips to a small sieve-like disc. The endostome
segments are unusual in being opposite the teeth rather than alternating with them.
Solomon “spake of trees, from the cedar tree that is in Lebanon even unto the
hyssop that springeth out of the wall.” Stark, in A Popular History of British Mosses
stated that a moss, Entosthodon fascicularis (Hedw.) C. M., related to Funaria, is this
Hyssopus Salamonis, citing the far-fetched authority of Sir J. E. Smith and the dubious
evidence of a specimen in Linnaeus’ herbarium collected in the Holy Land by
Hasselqvist, who was one of Linnaeus’ students. The Moldenkes, in Plants of the Bible,
made no mention of this collection or of any moss which had been called a hyssop.
The word was used a number of times in the Bible, and numerous suggestions have
been made as to its identity, the most likely contenders apparently being the caper or
the Syrian marjoram.

In England, as summarized by Watson (Structure and Life History of Bryo-

137,

phytes), Funaria hygrometrica is among the first colonists following a fire. It is quickly
succeeded by Ceratodon purpureus and then by Polytrichum juniperinum and P.
piliferum in association with various Cladonias. The last phase commonly endures for
many years. The abundance of Funaria in the first phase corresponds to a short period
when the substrate has a high pH and a high nutrient content (especially of potash),
soon lost by leaching. This information, based in part on a study of a burned-over
English heath (Fritsch & Salisbury, New Phytol. 14: 116. 1915) is just as applicable in
North America, although Skutch (Ecol. 10(2): 177-189. 1929) found in both Maine
and Jamaica that Marchantia polymorpha was a common pioneer, followed by
Polytrichum, Peltigera, and eventually Pteridium. Skutch provided a good review of
literature on the early stages of plant succession following forest fires (including
abundant reference to Funaria and Ceratodon in early stages). Graff (Bull. Torrey Bot.
Club 63: 67-74. 1936) has provided further information on the invasion of burns by
Marchantia in the American West.

Marshall & Averill (Ecol. 9: 533. 1928) provided data on the alkalinity of
burned-over forest soils in northern Idaho where some 125,000 acres of forest were
swept by fire in 1926. One year later 58 of 60 scattered tests showed a pH of 7-8, in
sharp contrast to the acid reaction normal to forest soils. (Two tests in the acid range
could be related to incomplete destruction of duff and humus.) The explanation of
such pH values lies in the high potash content of wood ashes. It may be recalled that
lye obtained incidental to clearing and burning hardwood forests was important in the
economy of early American settlers. Lye was produced by soaking ashes in water. It
was used with animal fat, such as lard, in making soap. It was also used in removing
the hulls from corn in making hominy. It should be noted, however, that O’Toole and
Synnott (Jour. Ecol. 59: 121-125. 1971) reported that Marchantia polymorpha and
Funaria hygrometrica invaded Irish blanket peat after fertilization with calcium carbonate
and phosphorus, but response to increased nitrogen and potassium was poor.

At Douglas Lake both Funaria and Marchantia are uncommon, though no doubt
normally successional to fire in wet places.

It may be that the spores of some weeds remain viable for long periods of time
and help in that way to explain some widespread distributions. Malta (Acta Univ. Latv.
4: 235-246. 1922) was able to germinate spores of Funaria hygrometrica 13 years after
their maturity and those of Ceratodon purpureus (an even more ubiquitous weed) after
16 years. Meyer (Bryol. 44: 69-75. 1941) reported growing Funaria hygrometrica
spores which were nine years old, and Hoffman (Bryol. 73: 634-635. 1970) grew some
which were 11 years old. Meyer found, however, that spores of the weedy Physco-
mitrium turbinatum (= P. pyriforme), which is closely related to Funaria hygrometrica,
could be grown only from the current and preceding year’s crop. Other literature on
longevity of spores: Bernstein (Acta Hort. Bot. Univ. Latv. 3: 33-36. 1928); Sussman
in Ruhland et al, Handbuch der Pflanzenphysiologie 15: 1086-1093. 1965).

Schimper (in Recherche anatomiques et morphologiques sur les mousses. Stras-
bourg, 1848) is said to have reported spore germination after 50 years, but Malta
(1922) mentioned that Correns cast doubt on the record. Malta (Acta Univ. Latv. 1:
125-129. 1921) reported germinating spores of Grimmia pulvinata from a specimen
nearly 70 years old. But later (Acta Univ. Latv. 4: 235-246. 1922) he confessed that
he had been unable to duplicate the results and therefore considered them based on

error. He also observed protonematal growth in a 52-year-old specimen of Grimmia
ovalis but was not sure whether the growth resulted from spore germination or
regeneration.

Spore production may actually be superfluous in some mosses where germination
is particularly chancy. Keever (Ecol. 38: 422-429. 1957) could easily grow Grimmia
laevigata, an invader of bare rock, by regeneration from shoots or fragments but not

138

from spores, even though spores are produced commonly and abundantly. This suggests
that rigid and brittle plants of dry places, likely to be crumbled by accident anyway,
may reproduce more efficiently and more certainly by regeneration than by spores.

Hughes (New Phytol. 68: 883-900. 1969) reported that spores of Grimmia
pulvinata germinate readily to form protonematal growth but no leafy shoots result.
Cuttings of setae, however, regenerate by a protonematal growth which sometimes
produces leafy shoots. Such diploid shoots can apparently produce capsules vege-
tatively (by apogamy), but no complete development was observed. Again, it would
seem that normal reproduction from spores may be dispensed with in favor of
accidental breakage, even though spores are regularly produced in G. pulvinata, as in G.
laevigata.

The ability of mosses to occupy extremely xerophytic habitats or withstand
extreme fluctuations in temperature and humidity is well known. Malta (Acta Univ.
Latv. 1: 125-129, 1921) reported reviving herbarium specimens, after long periods of
storage, as follows: Anoectangium compactum, 19 years; Dicranoweisia cirrata, 9 years;
Grimmia elatior, 70 months; Anomodon longifolius, 29 months; Bryum argenteum, 23
months; Orthotrichum rupestre, 22 months; Grimmia muehlenbeckii, 18 months;
Rhynchostegiella compacta, 8 months (growth of brood bodies). Patterson (Bryol. 46:
1-13. 1943), using a respiration method found greater viability in xeric mosses than in
hydric and mesic types (after one year of desiccation).

Glime (Bryol. 74: 383-386. 1971) found that a species of Fontinalis died after
only 55 hours of laboratory desiccation, but two species, F. novae-angliae and F.
dalecarlica, removed from a stream and laid on the bank survived stranding for one
year. She reported that Polytrichum ohioense lived as long as seven months under
laboratory conditions of desiccation.

Xerophytism in bryophytes, including the relation of osmotic pressures and
resistance to drought and cold, have been studied by Abel (Oesterr. Akad. Wiss. Wien
Math. Naturw. KI., Abt. 1, 165: 619-707. 1956), Hinshiri & Proctor (New Phytol. 70:
527-538. 1971), Hofleur (Ber. Deutsch. Bot. Ges. 15: 94-107. 1942), Hosokawa &
Kubota (Jour. Ecol. 45: 579-591. 1957), Irmscher (Jahrb. Wiss. Bot. 50: 387-449.
1912, reviewed in Bryol. 15: 67-69. 1912), Johnson & Kokila (Bryol. 73: 682-686.
1971), Ochi (Bot. Mag. Tokyo 65: 10-12, 112-118. 1952), and Patterson (Amer. Jour.
Bot. 33: 604-611. 1946; Bryol. 67: 390-396. 1964). Ochi (Bull. Soc. Plant Ecol.
Sendai 1: 182-187. 1952), on investigating xeromorphism in nine Japanese mosses,
concluded that drought resistance is more dependent on osmotic pressure and
permeability rather than structural modification. Apparent adaptations for water
conservation are listed by Grout (Bryol. 11: 97-100. 1908) and Smith (Univ. Kans. Sci.
Bull. 46: 433-474. 1966).

A review of recent research on bryophytes and water relations, edaphic factors,
light, radiation, air pollution, and growth form was presented by Miller (Plant Sci.
Bull., Dec. 1971: 34-37). An earlier review on water relation was provided by
Magdefrau (Ann. Bryol. 10: 141-150. 1937).

Funaria flavicans Mx., a weed in the southern United States, less common
northward but extending as far as southern Ontario, differs from F. hygrometrica in
having scarcely hygroscopic setae, slightly asymmetric capsules which become moder-
ately plicate on drying, and emarginate or bluntly bilobed endostome segments about
one-fourth the length of the peristome teeth.

Funaria americana Lindb., a rare species of scattered distribution in the East, has
smooth capsules, no annulus, segments about three-fourths the length of the peristome
teeth, and subentire leaves.

159

Pyramidula tetragona (Brid.) Brid. is a prairie species which may yet be found in
Michigan. It is something like Physcomitrium, but the seta is short and the
eperistomate capsules are enclosed, even at maturity, in a large 4-angled calyptra which
allows spore dispersal through a lengthwise tear.

EPHEMERACEAE

Minute and delicate ephemerals occurring usually as scattered buds on a sparse or
abundant protonema. Leaves, few, the lower small, the upper linear-setaceous to
lanceolate or ovate-lanceolate, broadly to narrowly acuminate, entire to serrate; costa
single and often weak to obsolete or lacking; cells oblong-rhomboidal, mostly lax and
transparent, sometimes papillose because of projecting upper ends. Capsules sessile,
ovoid or globose, sometimes apiculate, without a neck, indehiscent or opening along a
line of weakness near the middle; exothecial cells lax and thin-walled; stomata
superficial, with 2 guard cells, or sometimes lacking; columella mostly resorbed at
maturity; annulus and peristome none. Spores large, reniform, brown, papillose.
Calyptrae small and mitrate or minute and consisting of little more than the unaltered
remains of the archegonium.

Ephemerum Hampe, nom. cons.

Protonema typically abundant. Leaves not or only slightly shriveled or contorted
when dry, linear-setaceous to lanceolate or ovate-lanceolate, acuminate, entire to
strongly serrate; costa well developed or rarely lacking; upper median cells oblong-
rhomboidal, mostly lax and rather thin-walled, smooth or, more often, papillose
because of projecting upper ends. Capsules ovoid, apiculate, indehiscent. Calyptrae
small, mitrate, smooth or papillose, naked.—The name refers to the short-lived nature of
the plants. Elizabeth G. Britton referred to them appropriately enough as EMERALD
DEWDROPS. Although the Ephemera have a reputation for rarity, they are not hard
to find if one knows what they look like, where they grow, and when to find them. They
are pioneers of disturbed soils, able to grow in temporarily available habitats (especially
fallow fields in the second year of succession) at times when competition is at a
minimum, from fall to spring. Like more weedy and ubiquitous species of such
habitats they are favored by a short life cycle. The protonema, though abundant and
no doubt physiologically active, is probably not as persistent as generally thought, in
view of the short life cycle.

Leaves broadly oblong or oblong-lanceolate, rather abruptly narrowed to a recurved acumen
an

distinct, asymmetric, irregularly and coarsely serrate shoulders; cells smooth, in

oblique rows at the shoulders 2. E. cohaerens
Leaves much narrower, more gradually acuminate from the base or from poorly marked
shoulders; cells papillose, not in oblique rows 1. E. crassinervium

1. Ephemerum crassinervium (Schwaegr.) Hampe—Plants green or yellowish.
Leaves 0.8-1.5 (rarely 2.5) mm. long, about 0.2 mm. wide, linear-setaceous, scarcely to
strongly toothed in the upper 2/3; costa filling the acumen and distinct to the base,
papillose at back; upper median cells oblong-rhomboidal, about 4-6:1, usually papillose,
often strongly so. Dioicous. Capsules shortly and usually obliquely apiculate; stomata
few, scattered throughout the exothecium. Spores 50-75 (rarely 95) X 35-504,
coarsely papillose.—n = 27.

Bryan & Anderson, Bryol. 60: 97, figs. 11-19, & 102, fig. 54. Conard, How to Know the
Mosses, (ed. 2) fig. 40e. Grout, Mosses with Hand-lens and Microscope, fig. 66.—Fig. 390-391.

140

On moist soil es disturbed places, particularly in old fields, also on river banks etc., in re
summer to early spring, i a on latitude. Connecticut to Ontario, Michigan, and Kansa
south to Florida and ian

EMMET CO.—Brutus. PRESQUE ISLE CO.—Posen.

Easily recognized by the slender, linear-setaceous leaves with the costa nearly
filling the acumen and with cells papillose.

Z hemerum cohaerens (Hedw.) Hampe—Plants green or typically brownish
when mature. Largest leaves up to 1.5 mm. long, oblong or oblong-lanceolate, rather
abruptly narrowed (sometimes conspicuously so) to a recurved acumen from fairly
distinct, asymmetric, irregularly serrate to spinose and + incised shoulders about 2/3 up
the leaf, otherwise serrulate to serrate in the upper 1/2 or more; costa obsolete at base,
becoming stronger and thicker above, percurrent or excurrent, + papillose at back;
lower cells large, lax, and oblong or oblong-hexagonal, gradually becoming smaller and
rhombic to oblong and arranged in + diagonal rows in the middle 1/3 (near the base of
the acumen), smooth or sometimes + papillose. Polyoicous (dioicous or autoicous).
Stomata distributed over the entire capsule. Spores rather finely to coarsely papillose,
45-54 X 60-95 uw. Calyptrae usually smooth.—n = 27.

n & Anderson, Bryol. 60: 99, figs. 28-35 & 102, fig. 56. Conard, How to Know the
Mosses ee 2) fig. 40d. D arlington, Mosses of Michigan, fig. 65. Grout, Mosses with Hand-lens and
Microscope, fig. 65.—Fig. 392.

re soil in wet places, along streams and in pastures, marshes, or meadows, both spring
and fall re locally, at least, also in mid-summer). Europe; ot n Canada, south to Florida,
west to Iowa and Texas. Not yet found in our area, this speci hus been collected in nearby
Alpena and Montmorency Counties and in numerous more uy counties

The asymmetric leaves with irregularly toothed or jagged shoulders and obliquely
arranged cells are unmistakable. The only other species known from Michigan
(somewhat resembling £. crassinervium) is E. spinulosum Bruch & Schimp. ex Schimp.
(fig. 393), which has linear-lanceolate leaves, spinose-serrate because of spreading or
recurved teeth; the costa is percurrent but does not fill the apex; and the cells are
papillose. (The chromosome number of E£. spinulosum is n = 27.

The photomicrographs published by Bryan and Anderson (Bryol. 60: 67-102.
1957) provide the easiest means of identifying North American species.

SPLACHNACEAE

Plants small to fairly robust, gregarious or, more often, loosely to densely
caespitose, usually + tomentose, mostly growing on bones, dung or other decaying
organic matter. Stems erect, forked, often bearing minute, claviform-cylindric filaments
in axils of upper leaves. Leaves soft, mostly broad, ovate-lanceolate to oblong or
spatulate, entire to dentate or rarely laciniate-toothed, sometimes bordered; costa
single, strong, generally ending below the apex; cells smooth, lax and pale, relatively
large, oblong or oblong-hexagonal above, the lower cells oblong, not differentiated at
the angles. Perichaetial leaves scarcely differentiated; perigonia often large and con-
spicuous, with filiform paraphyses. Setae mostly elongate, erect; capsules mostly
exserted, erect and symmetric, usually with a conspicuously differentiated neck (the
hypophysis), sometimes merely elongate, often swollen or expanded; annulus mostly
lacking; operculum convex to conic (very rarely undifferentiated); stomata abundant,

141

with 2 guard cells; peristome nearly always present, single, the 16 teeth various in
form, frequently in groups of 2 or 4 and ofter + fused, entire or rarely deeply forked.
Calyptra mitrate or rarely cucullate, naked or sometimes hairy.

Splachnum Hedw.

Plants in soft, loose, pale- or yellow-green, somewhat shiny tufts, radiculose
below. Stems loosely foliate, simple or forked. Leaves large, usually more crowded
toward the stem tips, erect or erect-spreading, soft, somewhat flexuose-contorted when
dry, broadly obovate, obtuse to acute or serrate above; costa ending in or below the
apex; cells large, lax, thin-walled, pale, smooth, oblong-hexagonal above, rectangular
below. Autoicous or dioicous; perichaetia scarcely differentiated; perigonia large and
cupulate, the bracts with long, narrow, spreading acumina. Setae smooth, slender and
mostly elongate, sometimes very long because of continued growth after maturity of
spores; capsules erect, consisting of a small, subcylindric, brownish, firm-walled urn and
a broader, more delicate, globose, pyriform, or turbinate hypophysis strikingly
different from the urn in color and texture and generally wrinkled on drying, usually
continuing growth after spore maturation and sometimes becoming greatly expanded
and skirt- or umbrella-like; annulus none; operculum convex or hemispheric, often
somewhat umbonate, sometimes stoutly and bluntly apiculate; exothecial cells irregu-
larly hexagonal, + incrassate, often somewhat collenchymatous, smaller and usually +
oblate in numerous suboral rows; cells of the hypophysis, large, lax, thin-walled,
hexagonal; stomata in the upper part of the hypophysis; columella shortly exserted,
peristome teeth 16, inserted at or below the mouth, sometimes in pairs, reflexed when
dry, inflexed when moist, lanceolate, densely and minutely papillose on the outer
surface, derived from 3 peristomial layers of cells and chambered. Spores spherical or
ellipsoidal, small, smooth. Calyptrae small, conic-mitrate, often becoming split on 1
side, not constricted at the base, smooth, naked.—Splachnum was a name used by
Dioscorides presumably for the lichen Lobaria pulmonaria, which has a lung-like
appearance roughly similar to the wrinkled hypophysis of some species of Splachnum.
In spite of a predilection for dung as a substrate, the species of Splachnum are among
the most elegant and colorful of all mosses.

Splachnum ampullaceum Hedw.—Plants 1-4 cm. high. Leaves 3.54 mm. long,
long-lanceolate to narrowly oblong-ovate, slenderly acuminate; margins deeply spinose-
dentate in the upper half or less; costa disappearing in the acumen somewhat below
the apex. Dioicous (male and female plants invariably mixed). Setae 15-65 mm. long,
slender, flexuose, red or red-brown; urn 1-1.2 mm. long, yellow-brown; hypophysis
much broader, 2-6 mm. long, top-shaped, rugose, pinkish to pale-purple, becoming
dark-brown with age; operculum hemispheric, blunt; peristome teeth inserted near the
mouth, united in pairs, pale-brown or orange-brown. Spores 7-9.5 yu, subspherical
(slightly longer than wide), smooth.—n = 8, 9, 10 (and, in experimental material, n =
8, 16, 32)

Darlington, Mosses of Michigan, fig. 70. Grout, Mosses with Hand-lens and Microscope, fig.
94. Jennings, Mosses of Western Pennsylvania, (ed. 2) Pl. 52.—Fig. 394-397.

On old dung of cows (or Greer mea other herbivores, such as moose, deer, or horses) in
peat bogs and bog forests. ate polar; British ,Columbia and Alberta, Nova Scotia to New
England, West Virginia, and Mic

CHEBOYGAN CO.-Little Lake 16. EMMET CO.-—Stutsmanville Bog. PRESQUE ISLE
CO.—Hammond Bay

142

The large teeth on the margins of the leaves, often consisting of several cells,
distinguish S. ampullaceum from other members of the genus and, indeed, the entire
family.

Splachnum rubrum Hedw. (fig. 398-399), with a red, skirt-like hypophysis, occurs
on moose dung on Isle Royale in Keweenaw County. S. luteum Hedw. has a lovely
yellow skirt. Of them, Mrs. Britton (The Observer 7: 637-645. 1896) said, “Fairy
parasols, red and yellow, that is what they seem to be, not ordinary umbrellas meant
to be taken out in the rain, but bright, gay, silky, showy things, only used by Titania
when in holiday attire. But the books will tell you that they are called ‘Umbrella
Mosses.’” (She failed to comment on the name PETTICOAT MOSS.)

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pease ampullaceum. 394. Habit. 395. Capsule (dry). 396. Leaves. 397. Upper cells of

leaf at m
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Tetraplodon munioides. 400. Habit. 401. Dry capsule. 402, Operculum and calyptra. 403.

Leaf

143

The odd habitats of Splachnum and clas genera have excited considerable
interest. Kerner von Marilaun’s account is pleasa

“In mountain districts, where cattle ote pass to and from the meadows
and alps, one notices on their halting grounds, and along their tracks, moss of a
conspicuous green colour growing on circumscribed spots. On closer examination we
find that we have here an example of the remarkable group of the Splachnaceae, and
that it has selected the cow-dung to be its nutrient substratum. Each growth of
emerald green, Splachnum ampullaceum, is strictly limited to the area of a lump of
dung... hundreds of little green stems, thickly clothed with leaves, emerge, and the
spore cases, which resemble tiny antique jars, and are amongst the prettiest exhibited
by the world of mosses, become visible as well.”

He further reported that Splachnum rubrum and S. luteum grow only on
reindeer dung in arctic latitudes of Europe. In North America, they are found on
moose dung. He recorded that Tayloria serrata is seen only near cow chalets on
decomposing human feces. I have not observed that species in the field, but I have
found Haplodon wormskjolfii on what seemed to be human dung near Eskimo
dwellings in Alaska.

Tayloria rudolphiana, in Europe, apparently grows most often on mossy trunks
and branches of Fagus silvatica and Acer pseudo-platanus, where it is associated with
the excrements of birds of prey.

Hooker and Taylor, in the Muscologia Britannica reported that they once saw
Tetraplodon angustatus, which normally grows on the dung of carnivores, on the foot
of an old stocking, and the same species was found by a friend of their covering the
half decayed hat of a traveler who had perished on ne mountain of St. Bernard in
Switzerland.

n northernmost Alaska, on Point Barrow, where Eskimos have slaughtered
whales since time immemorial, are vast swards of Splachnum vasculosum, which also
grows on

The genus offers a number of adaptive features of interest. The seta continues
growth even after the maturation of spores, suggesting an evolutionary attempt, similar
to that of Anthoceros, to develop a long-lived and independent sporophyte. Coupled
with this is a high degree of sterilization of sporophytic tissue, a trend continued so
successfully by higher plants. The greatly expanded hypophysis has numerous stomata
and a spongy internal structure similar to a mesophyll. The sporophyte is, in fact, a
functional approach to a sporophyll. (The sexual dimorphism of some other mosses is
associated with chromosomal dimorphism and sometimes spore size differences and
further illustrates an evolutionary experiment with sporophyll development which was
carried to a more successful conclusion by vascular plants.)

Vaizey (Ann. Bot. 5: 1-10. 1890) came to the conclusion that the highest
development of the moss sporophyte is reached in Splachnum in which the hypophysis
serves the same functions as the leaves of higher plants. He even went so far as to
suggest homology. Further speculations on such homologies (or analogies) are pre-
sented by Crum (Bryol. 69: 205-207. 1966).

Splachnum has many adaptations for the dispersal of spores by insects (and
presumably such as also visit fresh dung for breeding purposes). The hypophysis may
be spherical and wrinkled when dry or enlarged and expanded, in both cases brightly
colored like a flower and, one would think, attractive to insects and providing a
landing space for them (although insects are not sensitive to the same spectrum of
color as we are). The dung on which the plants grow is thoroughly humified and, to a
man at least, smells only like fresh earth, but the hypophysis emits a faint, unpleasant
odor apparently through the stomata; this is especially noticeable under conditions of
humidity. I have noticed sour or musty odors in Splachnum and Tetraplodon; the
smell is obvious when plants are taken from the dampness of a vasculum at the end of

144

a collecting day, but it is difficult to demonstrate even moments afterward. A final
feature conducive to insect dispersal is that the spores adhere in clumps and are not
likely to be blown free in the air. The columella is thickened at the apex and acts as a
plug when the capsule is moist, but it is exposed, together with the mass of spores, by
shrinkage of the dry capsule.

Information on dispersal of spores of Splachnaceae is provided by Bequaert
(Bryol. 24: 1-4. 1921), Bryhn (Biol. Centralbl. 17: 48-55. 1897), Erlanson (Bryol. 33:
13-14. 1930), Walsh (Trans. Brit. Bryol. Soc. 1(5): 487. 1951), and Wettstein (Osterr.
Bot. Zeit. 70: 65-77. 1921).

Steere (Amer. Nat. 92: 5-20. 1958) suggested an interesting correlation of the
odors emitted by capsules of various Splachnaceae to the evolution of the family: ‘‘So
far as I know, these are the only mosses whose capsules have specific odors. The
capsule of Tetraplodon has the strong sharp odor of an acetic ester. Splachnum ovatum
smells strongly of lactic acid, and S. luteum is reported to have the odor of a butyl
compound. ... A correlation that may have some significance is that the more highly
modified and specialized species, morphologically, also have the odor of the more
complex organic compound. The biochemical evolution in Splachnum ...would seem
to promise a fertile field for investigation.”

We have, in the Upper Peninsula, two species of Tetraplodon which grow on the
dung of carnivores (such as wolf), bones, or owl pellets. The hypophysis is elongate-
pyriform and narrower than the spore-case; the 16 teeth of the peristome are at first
more or less joined in 4’s, later in 2’s. Tetraplodon angustatus (Hedw.) BSG is small,
with leaves oblong-lanceolate, and gradually subulate, irregularly serrate, and _ its
capsules are brown and not or slightly exserted; the hypophysis is also brown.
Tetraplodon mnioides (Hedw.) BSG (fig. 400403) is more robust and has oblong-
ovate, rather abruptly subulate, entire leaves and capsules which are red (becoming
dark- red or blackish) and long-exserted; the hypophysis is area darker than the
capsule. Chromosome numbers reported for T. mnioides are n = 11,

Several other species of Splachnaceae might conceivably occur in the northern
reaches of the Great Lakes area, including Splachnum sphaericum Hedw. (also known
as S. ovatum Hedw.), Tayloria acuminata Hornsch. (fig. 404-409), T. lingulata (Dicks.)
Lindb., T. serrata (Hedw.) BSG (fig. 414-418), and T. splachnoides (Schleich. ex
Schwaegr.) Hook. (fig. 410-413). Splachnum sphaericum has a purplish to blackish
hypophysis only slightly broader than the urn (and resembles in capsule shape a
Tetraplodon). It occurs on moose dung. Tayloria lingulata grows scattered on wet soil
especially at the margins of dried-up pools no doubt enriched by dung of wading birds.
It takes the name from its tongue-shaped leaves which are broadly rounded at the
apex.

SCHISTOSTEGACEAE

A family of a single genus, Schistostega Mohr, and one species: Schistostega
pennata (Hedw.) Web. & Mohr. LUMINOUS MOSS. CAVE MOSS. GOBLIN GOLD.—
Plants small and delicate, 4-7 mm. high, gregarious or loosely tufted, radiculose only at
the base, growing from a persistent protonema which is luminous because of light
reflected from vesiculose cells in conidia-ltke chains. Plants dimorphous: Sterile stems
erect-flexuose, naked below, pinnately foliate and frondiform above; leaves 0.7-1.2 mm.
long, distichous-complanate, little altered when dry, oblong or oblong-lanceolate,
slightly asymmetric, acute, confluent at base because of decurrencies, ecostate; margins
plane, entire (or sinuolate), indistinctly bordered); cells laxly oblong-rhomboidal,
thin-walled, smooth, pellucid, 16-20 u wide and about 5: 1 above, somewhat narrower

{hy

410 | | ce

ayloria acuminata. 404. Leaves. 405. Leaf cells. 406. Axillary club hairs. 407. Capsules,

moist a left) and dry. 408. Two moist capsules showing inrolled peristome. 409. Exothecial cells.
Tayloria splachnoides. 410. Leaves. 411. Leaf cells. 412. Capsules, wet and dry. 413.
Exothecial cells.

in a marginal row, rectangular below. Fertile plants (both male and female) mixed with
the more conspicuous sterile plants, more slender, with smaller, erect-spreading leaves
inserted all around the stem in a crowded tuft at its tip. Dioicous. Setae 2-5 mm. long,
pale, slender, flexuose, smooth; capsules erect and symmetric, very small (0.4-0.5 mm.
high), subglobose or ovoid (i.e., wide at the base and small at the mouth), smooth,
annulus and peristome none; exothecial cells irregularly hexagonal, thickened at the
corners, with 1-2 rows at the mouth darker, smaller, and subquadrate, often resembling

146

a persistent annulus; stomata none (or few and difficult to observe); operculum
plano-convex. Spores spherical, 10-13 u. smooth. Calyptra very small, conic-mitrate.—n
1, 14.

d, How to Know the Mosses, (ed. 2) fig. 31. Darlington, Mosses of Michigan, fig. 71.

Grou osse

including ap ah of the protonema, are provid :
Inoue (Dare Nimo Wakaru Koke No Subete, p. 60, 1971), Kobayabi & Watari (Jour. Jap. Bot. 11:
433-437. 1935), al Kanda (Hikobia 6: 60- 75. 1972).—Fig. 419-422.

A rare species not likely to be found in the limey habitats of the Straits Counties but

tring in Scotts Cave west of Au Train in Alger County. It is characteristically found in caves
associated with sandstone rocks but also occurs in other dark recesses, under old barns or in
cavities under upturned roots of trees, for example. Europe; Japan; British Columbia to Washington
and Alberta; rare and scattered from Wisconsin to Newfoundland and south to New York and

Rhode Is

The genus is defined by its dimorphous growth habit and luminous protonema
which makes possible its growth in dark habitats. The epithet pennata, meaning
feathery, is aptly descriptive, as was an epithet formerly in use, osmundacea, referring
to the fern-like appearance of sterile plants. The generic name, meaning split
operculum, resulted from an erroneous observation that the operculum becomes

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chistostega pennata, 419. ‘Hab bit of sterile and fruiting plants. 420. Cells at leaf tip. 421.
Portion of protonema in diffuse light. 422. Protonematal cell lighted from one side.

147

radially split into tooth-like segments. William Wilson, unable to find radiating fissures
on examination of living material, remarked that “the operculum is the very thickest
and the most sturdy that I ever met with... filling the mouth exactly like a bung,
composed strictly of cells of an hexagonal form pervading the thickness of the lid, and
not infrequently disposed so as to stand in rows from the centre to the circumference,
so that when any part of the lid is obliquely placed, with respect to the eye, the
partitions of the cells, in perspective, represent dark lines resembling radii” (in Smith’s
English Flora, vol. 5, part 1, p. 16. 1844).

The luminous appearance of the moss is caused by reflection from lens-like cells
of the protonema. The account given by Kerner von Marilaun* (in his Pflanzenleben,
translated by F. W. Oliver as The Natural History of Plants) has appeal: ‘“‘On looking
into the interior of the cave, the background appears quite dark, and an ill-defined
twilight only appears to fall from the centre on to the side walls; but on the level floor
of the cave innumerable golden-green points of light sparkle and gleam, so that it
might be imagined that small emeralds had been scattered over the ground. If we reach
curiously into the depth of the grotto to snatch a specimen of the shining objects, and
examine the prize in our hand under a bright light, we can scarcely believe our eyes,
for there is nothing else but dull lustreless earth and damp, mouldering bits of stone of
yellowish-grey colour! Only on looking closer will it be noticed that the soil and stones
are studded and spun over with dull green dots and delicate threads, and that,
moreover, there appears a delicate filigree of tiny moss-plants, resembling a small
arched feather stuck in the ground. This phenomenon, that an object should only shine
in dark rocky clefts, and immediately lose its brilliance when it is brought into the
bright daylight, is so surprising that one can easily understand how the legends have
arisen of fantastic gnomes and cave-inhabiting goblins who allow the covetous sons of
earth to gaze on the gold and precious stones, but prepare a bitter disappointment for
the seeker of the enchanted treasure; that, when he empties out the treasure which he
hastily raked together in the cave, he sees roll out of the sacks, not glittering jewels,
but only common earth.... On the floor of rocky caves one may discern by careful
examination two kinds of insignificant-looking plant-structures, one a web of threads
studded with small crumbling bodies, and the other bluish-green moss-plants resembling
tiny feathers. The threads form the so-called protonema, and the green moss-plants
grow up as a second generation from this protonema ... the gleams do not issue from
the green moss-plants, but only from their protonema.

“From the much branched threads ...numerous twigs rise up vertically, bearing
groups of spherical cells arranged like bunches of grapes. All the cells of a group lie in
one plane, and each of these plants is at right angles to the rays of light entering
through the aperture of the rocky cleft. Each of the spherical cells contains
chlorophyll-granules, but in small number ...and they are always collected together on
those sides of the cells which are turned towards the dark background of the cave....
Taken together, these chlorophyll-granules form a layer which under low power of the
microscope appears as a round green spot...the light which falls on such cells
through the opening of a rocky cleft behaves like the light which reaches a glass globe
at the further end of a dark room. The parallel incident rays which arrive at the globe
are so refracted that they form a cone of light, and since the hinder surface of the
globe is within this cone, a bright disc appears on it. If this disc, in which the refracted
rays of light fall, is furnished with a lining, this also will be comparatively strongly
illuminated by the light concentrated, on it and will stand out from the darker
surroundings as a bright, circular patch. ... It is well worthy of notice that the patch
of green chlorophyll-granules on the hinder side of the spherical cell extends exactly so
far as it is illumined by the refractive rays, while beyond this region, where there is no

*The grandfather of Fritz von Wettstein, famed for his work on the genetics of mosses.

148

illumination, no chlorophyll granules are to be seen. The refracted rays which fall on
the round green spot are, moreover, only partially absorbed; in part they are reflected
back as from a concave mirror, and these reflected rays give ...a luminous appearance.
This phenomenon, therefore, has the greatest resemblance to the appearance of light
which the eyes of cats and other animals display in half-dark places, only illumined
from one side, and so does not depend upon a chemical process, an oxidation, as
perhaps does the light from a glow-worm or of the mycelium of fungi which grows on
decaying wood. Since the reflected light-rays take the same path as the incident rays
had taken, it is clear that the gleams of the Schistostega can only be seen when the
eye is in the line of the incident rays of light. In consequence of the small extent of
the aperture through which the light penetrates into the rock cleft, it is not always
easy to get a good view.... If we hold the head close to the opening, we thereby
prevent the entrance of the light, and obviously in that case no light can be reflected.
It is, therefore, better when looking into the cave to place one’s self so that some light
at any rate may reach its depth. Then the spectacle has indeed an indescribable
charm.”

And thus the gold of the Nibelungen is reduced to scientific fact!

The spherical cells of the protonema of this “‘singularly brilliant vegetable” were
associated with its luminous nature by Bowman as long ago as 1829 (Mag. Nat. Hist.
2: 407) and illustrated by him in 1830 (Mag. Nat. Hist. 3: 462-463. Fig. 115).

Other references of interest: Albrecht-Rohner (Ber. Schweiz. Bot. Ges. 61:
428-460. 1951), Gaisberg & Finck (Flora 120: 143-175. 1926), Johnson (Bryol. 29:
17-19. 1926), Lye (Lindbergia 1: 205-213. 1973), Toda (Jour. Coll. Sci. Tokyo 40(5):
1-30. 1918), Webster (Nat. Hist. 58: 309-311. 1949

A similar protonema occurs in an Australian luminous moss, Mittenia plumula
(Mitt.) Lindb., described in full by Stone (Proc. Roy. Soc. Victoria 74(2): 119-124.
1951). Mrs. Stone also presented the developmental morphology in a complete and
excellent ied including the development of the peristome (Austral. Jour. Bot. 9(2):
124-151. 1).

ae thallose liverworts of the genus Cyathodium (Targioniaceae) grow in caves
and other poorly lighted places and emit a yellowish luminescence from their thalli.

Schistostega and Mittenia illustrate so well that mosses have fairly definite
ecological requirements or, at least, preferences. The species are both structural and
biological entities. This biological role is explored by Crum (Jour. Hattori Bot. Lab.
35: 269-298. 1972). Some information on habitat preferences is given in Glenn &
Welch (Proc. Indiana Acad. Sci. 40: 87-101. 1931) and Welch (Bryol. 36: 11-13. 1933;
Proc. Indiana Acad. Sci. 82: 105-107. 1955). Other references are given throughout
this book, under Sphagnum, Scopelophila, Mielichhoferia, Funaria, Splachnum,
Ephemerum, etc. One can get further information from Richard’s summaries of the
literature on ecology of bryophytes (in Verdoorn’s Manual of Bryology, pp. 367-395.
1932; Jour. Ecol. 28: 245-248. 1940).

BRYACEAE

Plants mostly in loose or dense, green or yellowish tufts. Stems erect (or,
Rhodobryum, erect from a rhizome-like primary stem), simple or forked, often densely
radiculose. Leaves in many rows (rarely somewhat complanate with + dimorphous
leaves), usually small below, larger and more crowded above, often in comose tufts,
usually erect or erect- spreading, rarely much altered on drying, lanceolate to ovate,
usually acute, sometimes awned or hair-pointed, frequently bordered; costa single, well
developed, often excurrent; upper cells elongate, typically pale, relatively large,
smooth, rhomboid-hexagonal to rhomboidal or linear above, rectangular or sometimes

149

subquadrate at base. Perichaetia terminal (in Mielichhoferia at the ends of short, basal
branches); perichaetial leaves not much differentiated; perigonia with filiform
paraphyses. Setae + elongate, erect or + curved near the tip; capsules mostly inclined
to pendulous, sometimes curved, usually symmetric or nearly so, ovoid, pyriform, or
oblong-cylindric, with a fairly well-developed, narrow, tapered neck, nearly always
smooth; annulus usually present, usually large and revoluble; operculum convex to
short-conic, umbonate or apiculate, rarely short-rostrate; stomata numerous, in the
neck; peristome nearly always present, rarely single and consisting of only exostome or
endostome, the teeth 16, mostly lanceolate, slenderly pointed, often bordered,
papillose on the outer surface, prominently trabeculate at back; endostome typically
consisting of 16 delicate, pale, keeled segments from a generally well-developed basal
membrane, nearly always alternating with the teeth and usually separated by 1-3
delicate cilia. Spores spherical. Calyptrae cucullate, smooth, naked.
1. Leaves linear to lanceolate, not distinctly bordered; leaf cells elongate (4:1 or longer)

2. Leaves linear-subulate, the costa filling about 4% the base and most of the subula 1. Leptobryum
2. Leaves broader, with a narrower costa 2. Pohlia
1. Leaves broader, ovate or ovate-lanceolate, usually distinctly bordered; leaf cells less than

4:1

3. Stems rosulate, Sera erect from an underground runner; leaves serrate above
4.R

sporophytes often clust eee
3. Stems not particularly are not growing from an underground runner; leaves entire
or nearly so; sporophytes not clustered ryum

1. Leptobryum (BSG) Wils.

Slender plants in loose or dense, shiny, light- or yellow-green tufts, radiculose at
base. Stems simple, erect. Leaves small and remote below, abruptly larger and crowded
at the stem tips, erect- or wide-spreading, flexuose when moist, long-subulate or
setaceous from a broader, oblong base; margins plane, entire or somewhat denticulate
at the apex; costa very broad, filling about % the leaf base and most of the subula,
subpercurrent to shortly excurrent; cells long-linear, broader and long-rectangular
below. Setae terminal, smooth, slender, flexuose-curved; capsules strongly inclined to
pendulous, pyriform from a long, narrow neck, small at the mouth, smooth, shiny;
annulus revoluble; operculum convex-conic, mammillate; peristome teeth 16, lance-
acuminate, yellowish, densely papillose, trabeculate; endostome hyaline, finely
papillose, about as long as the teeth, with a high basal membrane, keeled and perforate
segments, and well-developed, appendiculate cilia normally in 3’s.-The name means
slender or delicate Bryum.

Leptobryum pyriforme (Hedw.) Wils.—Plants about 0.5-1.5 cm. high (rarely
more). Stems black, flexuose, rarely producing red, globose brood-bodies in leaf axils
or on radicles and protonemata, regularly producing minute, reddish club-hairs in axils
of upper leaves. Upper leaves 4-5 mm. long, sometimes denticulate at the tip.
Synoicous or sometimes dioicous. Setae 10-45 mm. long, red-yellow or orange-brown;
capsules 1.7-2.5 mm. long, brown or yellow-brown, horizontal to pendulous; cilia of
endostome in 3’s or occasionally in 4’s. Spores 9-13, smooth or very finely
papillose.—n = 20, 21, 22, 24.

Breen, Mosses of Florida, Pl. 64. Cgnard, How to Know the Mosses, (ed. 2) fig. 123.

yaar Mosses of Michigan, fig. 73. Grout, Mosses with Hand-lens and Microscope, fig. 105,

Moss Flora of North America 2: 76C. Jennings, Mosses of Western Pennsylvania, (ed. 1) Pl. 17, (ed.
2) Pl. 21. Welch, Mosses of Indiana, fig. 116.—Fig. 423-425.

On 1, rotten wood, humus, or occasionally rock in wet places, particularly in swamps,
often in tee ar or otherwise disturbed habitats, common in greenhouses. Circumpolar; South

150

432

423

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Pohlia nutans, 426. . 427, Leaves. 428. Upper cells of leaf. 429. Portion of peristome
and annulus. 430. Poition pent enlarged.

Pohlia cruda, 431, Leaves. 432. Upper cells of leaf.

merica, New Zealand, Tasmania, and New Guinea; Greenland to ig south to California,
Arizona, and New Mexico in the West, eee eastern North Amer

CHEBOYGAN CO.—Reeses Bog, Hermits Bog, Gorge of Carp Creek, ne sige alee
Station campus, Colonial Point, Mill Creek, Weber Lake. EMMET CO. _~Gallow y Bog s Lake,
Wycamp Lake, Cecil Bay. MACKINAC CO. —Mackinac Island, Prentiss “a percotn ISLE
CO.—Lamprey Experiment Station, Evergreen Beach.

Dixon described this as a “very elegant and beautiful plant, both in the leaf and
in the fruit.” The narrow, almost setaceous leaves, suggestive of Dicranella, seem
anomalous in the Bryaceae, but the pear-shaped capsules leave no question as to
relationship. The production of brood bodies on the rhizoids enable Leptobryum to
colonize bare soil readily and explain its common occurrence on flower pots in

151

greenhouses. It is interesting to observe that Marchantia polymorpha and Lunularia,
also common in greenhouses, bear gemmae in large numbers, and Ceratodon purpureus,
another invader of disturbed soils, produces gemmae on its rhizoids. (The brood-bodies
of Leptobryum are difficult to find in dried specimens, though apparently common in
living material, judging by published accounts.)

2. Pohlia Hedw.

Plants small to fairly robust, in loose or dense, green, yellowish, or rarely
reddish, sometimes glossy tufts, sometimes merely gregarious. Stems simple or forked.
Leaves generally crowded at the stem tips, erect or nearly so, not much altered on
drying, narrow and elongate, lanceolate and acute or rarely obtuse, + toothed near the
apex, not bordered; costa ending at or below the apex; upper cells usually long and
narrow, oblong-rhomboidal to linear; lower cells somewhat shorter and laxer, rectang-
ular. Setae elongate, flexuose and twisted when dry, usually curved or hooked at the
tip; capsules inclined to pendulous (or rarely suberect), symmetric or nearly so, clavate
or pyriform, rarely shorter and ovoid or subcylindric, with a short or long neck,
smooth; annulus usually present; operculum mostly convex-conic, mammillate or
apiculate to short-rostrate; stomata sometimes + immersed; peristome double, the 16
teeth lanceolate, yellowish or brownish, often bordered, finely papillose; endostome
with a well-developed basal membrane, segments keeled and often perforate, about as
long as the teeth, rarely rudimentary, and cilia rarely appendiculate (sometimes short
or lacking)._Named for Johann Ehrenfried Pohl, a physician of Dresden. The genus is
often confused with Bryum, which generally has broader leaves which are often
bordered and often awned and also has much shorter leaf cells.

1. Plants producing vermicular brood-bodies in the leaf axils

2. Plants glossy, with abundant brood bodie 4. P. proligera
2. Plants dull or only slightly shiny, with -several brood bodies per leaf axil, sometimes
rather abundantly produced 3. P. annotina var. decipiens

1. Plants not producing brood bodies
3. Capsules short; exothecial cells irregularly hexagonal, with em walls

4. Plants green or yellow; annulus present; stomata superficial 3.2: lescuriana
4, Plants with een leaves and reddish stems (particularly noticeable in contrast in
the field); annulus none; stomata immersed 6. P. wahlenbersgii

3. aoe cere exothecial cells oblong, with straight walls
shiny, whitish, yellowish, or bluish-green; upper leaf cells long-linear,
irae alled 1. P. ‘cruda
5. Plants dull or slightly shiny, green or yellowish; upper cells rhomboidal, thick-walled
2. P. nutans

1. Pohlia cruda (Hedw.) Lindb.—Plants in loose, soft, very shiny, light- or
whitish-green, yellowish, or bluish tufts 1-5 cm. high. Stems simple or forked, red.
Leaves gradually larger above, loosely erect or erect-spreading, not much altered on
drying, 2-3.5(5)mm. long, lance-acuminate, somewhat decurrent; margins plane or
narrowly reflexed, distinctly serrulate in the upper 1/2-2/3; costa ending near or
considerably below the apex, usually red below, flexuose above; upper cells very long
and narrow, thin-walled. Dioicous, paroicous, or occasionally synoicous. Setae
10-35 mm. long, red-yellow; capsules horizontal to pendulous (suberect when dry and
empty), pale-brown, 2.5-4 mm. long, oblong-cylindric or somewhat clavate, the neck
1/3-1/2 the capsule length but not clearly differentiated when dry; annulus revoluble;
operculum convex or conic, blunt or umbonate, rarely apiculate; endostome segments
perforate, the cilia well-developed, + nodulose, in 2’s or sometimes 3’s. Spores 13-24 uy,
finely papillose.—n = 10, 10+4, 11, 14, 22, 40.

132

Darlington, Mosses of Michigan, fig. 74. Grout, Moss Flora of North America 2: Pl.
74A.—Fig. 431-432.

soil or humus in moist, shaded places, particularly in crevices of cliffs and on banks,
often in the protection of overhanging turf (common in calcareous ane Very widespread in
both Northern and Southern Hemispheres (but in uplands or mountains); Greenland ie a
south to North Carolina, the Great Lakes region, eee Colorado, Arizona, and Califor

CHEBOYGAN CO.—Douglas Lake. EMMET CO.—Maple River, Cross Village.

n Dixon’s words, this is “‘a very fine and beautiful species known at once by the
metallic, almost opalescent sheen on the leaves.” (Other species of Pohlia with a
notable sheen are P. proligera and P. bulbifera.) The specific epithet means raw,
unripened, or immature, bearing a connotation of a fresh or youthful appearance.

2. Pohlia nutans (Hedw.) Lindb.—Plants in loose, green or yellowish, not or
somewhat shiny tufts 1-4 cm. high. Stems usually simple, red. Leaves longer and more
crowded above, 2-4 mm. long, erect, not much altered when dry (but often somewhat
twisted around the stem), lance-acuminate; margins + reflexed when dry, denticulate
toward the apex; costa subpercurrent; upper cells narrowly rhomboidal or linear,
thick-walled. Paroicous. Setae 14-33 (rarely 50) mm. long, orange-yellow; capsules
2.5-4 mm. long, orange-brown, horizontal to pendulous, the neck about 1/3 its length,
sometimes contracted when dry; annulus revoluble; operculum convex-conic, acute or
bluntly apiculate; endostome segments perforate or gaping, the cilia well-developed,
nodose or somewhat appendiculate, in 2’s or sometimes 3’s. Spores 16-21 uy, finely
papillose.—n = 11, 14, 21, 22, 23,

Conard, How to Know the Mosses, (ed. 2) fig. 145a-d. Grout, Mosses with Hand-lens and
Microscope, fig. 107, Moss Flora of North America 2: Pl. 75A. Jennings, Mosses of Western
Pennsylvania, (ed. 1) Pl. 17, 18 & (ed. 2) Pl. 21, 22 (as Webera nutans & var. triciliata).—Fig.
426-430.

Common on turfy soil, decaying logs and especially tops of rotten stumps, old Sphagnum
hummocks, and soil in rock crevices. Widespread in both Northern and Southern Hemispheres;
Greenland to Alaska, south to Georgia, Arkansas, Colorado, Arizona, and California.

CHEBOYGAN CO.-—Douglas Lake, Reeses Bog, Hermits Bog, Gorge of Carp Creek, Iron
Bridge, Colonial Point, Vincent Lake, Gates Bog, Livingston Bog, Bryants Bog, Mud Lake, Grass
Bay, Little Lake 16, Weber Lake. EMMET CO.—Arnott Lake, Galloway Bog, Larks Lake, Carp
Lake, Pleasantview Swamp, Stutsmanville, Wycamp Lake, Five Mile Creek. MACKINAC CO.—Bois
Blanc Island, St. Martins Bay, McKays Creek, Prentiss Bay, Hog Island Point, Cut River, Epoufette,
Ozark, Hendricks, Little Dollar Lake. PRESQUE ISLE CO.—Lamprey Experiment Station, Clinton
Lake, Loon Lake, Shoepac Sink Holes.

The orange setae and capsules help in field identification. | recently identified
Pohlia nutans sporophytes as a nesting material. The collector, Mrs. Nicholas L
a of Mount Pleasant, Michigan, wrote: “The nest was about two feet above the

round in a small Cornus racemosa shrub and was very striking looking because of the
(inated) hundred or so moss sporophytes in the lining. It was a wet day so that the
moss appeared gold-colored and was thus quite noticeable.”
larke & Greene (Trans. Brit. Bryol. Soc. 6(1): 114-128. 1970, 6(2): 278-295.
1971) compared the reproductive behavior of Pohlia cruda and P. nutans from arctic,
temperate, and subantarctic localities. They found that the paroicous P. nutans fruits
in all three areas but sporophytes mature much faster in the arctic. The dioicous
P. cruda fruits rather commonly in arctic, less commonly in temperate, and very rarely
n subantarctic sites. A comparison of plants grown together in the laboratory showed
ene adaptation of reproductive processes to local environments. For example,

153

British plants were able to complete gametangial maturation under short or long-day
conditions, but those of South Georgia matured gametangia only under long days. The
length of the maturation cycle varied greatly with temperature in P. nutans but not in
P. cruda. P. nutans could complete its maturation in warmer temperatures than P.
cruda. South Georgia plants had a lower optimum temperature for maturation than
British plants which had a lesser tolerance to frost.

The relationship of photoperiod to sporophyte development (in Polytrichum) was
studied by Hughes (New Phytol. 61: 266-273. 1969).

Pohlia elongata Hedw., which grows on ledges or in crevices of cliffs in the
Upper Peninsula, is somewhat similar. It is paroicous (or sometimes autoicous); its
capsules have a very long neck (about as long as the urn); and the cilia of the
endostome are rudimentary or lacking.

3. Pohlia annotina (Hedw.) Lindb.—Small plants in loose, yellow to dark-green,
not or only slightly shiny tufts. Stems reddish below, bearing in leaf axils (especially
toward the apex of sterile stems) 1-several small, yellowish, ovoid brood-bodies with
3-4 leaf points. Leaves not crowded, erect-spreading wet or dry, 1.2-2 mm. long,
narrowly lanceolate, decurrent; margins plane (or slightly reflexed when dry), +
serrulate at the apex; costa yellowish, ending somewhat below the apex; cells
rhomboidal, thin-walled. Dioicous. Setae slender and flexuose, reddish, 18-35 mm. long;
capsules 2.5-3 mm. long, long-pyriform, with a distinct, narrow neck about as long as
the urn, horizontal to pendulous, brown when mature; annulus revoluble; operculum
conic or convex and mammillate; exothecial cells rounded-hexagonal, with slightly
wavy walls; endostome segments perforate, the cilia in groups of 24. Spores 18-22 yn,
finely papillose.

Grout, Moss Flora of North America 2: Pl. 78 (fig. 2).—Fig. 433-435.

Not found in our area, the typical variety has been found in nearby Chippewa County (on
Sugar Island) and also in Alger County. Europe and the Azores; Asia Minor and Siberia; Greenland;
British Columbia to California and Idaho; Michigan and Ontario to New York and New Hampshire.

Var. decipiens Loeske (var. Joeskei Crum, Steere & Anders.)—Brood-bodies small,
yellow or pale-rusty, shortly vermicular, twisted, 1-several in the leaf axils.

Conard, How to Know the Mosses (ed. 2) fig. 145e. Grout, Moss Flora of North America 2:

Pl. 78 (fig. 1L).—Fig. 436.
n soil in moist, open places, especially near lakes and ponds or in ditches. Europe;
southeastern Alaska to Washington and Idaho; New Brunswick to Wisconsin, south to Georgia and
owa.

CHEBOYGAN CO.-—Vincent Lake.

Wilczek & Demaret (Bull. Jard. Bot. Nat. Belg. 40: 405-422. 1970) found that in
plants grown in culture slender, twisted propagula (as in P. annotina var. decipiens)
represent a young stage of growth, longer, cylindric and obovoid ones (as in P.
proligera) a somewhat older stage, and large, ovoid or obovoid ones (as in P. annotina
var. annotina) a mature stage. Consequently they recognized only one species. I reserve
judgment, at least as regards P. proligera, as have never had trouble recognizing it by
the size, shape, and luster of its leaves. They may be right in considering P. annotina
var. decipiens a mere developmental form. However, its range in eastern North America
is significantly different from that of the typical variety. From P. proligera, the var.
decipiens is distinguished by a relatively dull appearance and twisted brood bodies.

154

WWI YN)
Al {| | II )
(\/ }| iH | Vv

439

Pohlia annotina. 433. Leaves. 434. Upper ie a leaf. 435. Brood-bodies.
an annotina var. decipiens, 436. Brood-b

Pohlia proligera. 437, Leaves. 438. Upper as of leaf. 439. Brood-bodies.
Pohlia bulbifera, 440. one 441. Upper cells of leaf. 442. Brood-bodies.
Pohlia rothii. 443. Hab

4. Pohlia proligera (Kindb. ex Limpr.) Lindb. ex Arn.—Plants small and slender,
gregarious or loosely tufted, glossy, green, yellowish, or occasionally red-tinged. Stems
simple, red; brood-bodies very numerous in axils of upper leaves, small, vermicular,
with 1-3 leaf primordia, green to yellow or rusty. Leaves erect or spreading and often

L55

+ flexuose or twisted when dry, erect- to wide-spreading when moist, soft, 1-1.5 mm.
long, rather broadly oblong-lanceolate, acute, + decurrent; margins plane or somewhat
reflexed, slightly serrulate toward the apex; costa subpercurrent or ending somewhat
below the apex; cells linear-rhomboidal, thin-walled. Dioicous. Setae 20-40 mm. long,
slender and flexuose; capsules horizontal to pendulous, 2.5-3 mm. long, shortly
oblong-cylindric and short-necked; annulus revoluble; operculum convex-conic; exo-
thecial cells short with wavy outlines; endostome segments perforate, the cilia in 2’s
ane 3’s, nodulose. Spores 16-18 u, finely papillose. —n = 11

, Mosses with Hand-lens and Microscope, fig. 109, Moss Flora of North America 2: Pl.
718 fe L. AK). —Fig. 437-439.

n moist disturbed soil of roadbanks, also in crevices of cliffs. Europe; Greenland to Alaska
and south to California, Colorado, Michigan, and New York.

MACKINAC CO.—Cut River.

The conspicuous sheen and the small size provide an easy means of identifica-
tion. The brood-bodies have been aptly compared to an empty glove

Pohlia bulbifera (Warnst.) Warnst. (fig. 440-442), which occurs on Sugar Island
(Chippewa Co.), is a glossy plant with 1-3 obovoid or top-shaped, yellowish brood-
bodies per leaf axil. P. annotina has leaves less glossy and brood-bodies ovoid and not
tapered to the base.

Not yet discovered in Michigan is Pohlia rothii (Corr. ex Limpr.) Broth. (fig.
443), which has glossy leaves and produces singly in the leaf axils large, red, ovoid
gemmae with rather well-developed leaf points.

5. Pohlia lescuriana (Sull.) Grout—Small, gregarious or loosely tufted, dull or
somewhat shiny, green or yellow plants about 4-7 mm. high. Leaves 1.5-3 mm. long,
narrowly lance-acuminate; margins narrowly recurved, slightly and remotely serrulate
near the apex; costa ending near the apex; cells long, linear-rhomboidal. Dioicous.
Setae 9-17 mm. long, orange-brown or red; capsules about 1mm. long, shortly

yriform from a short, distinct neck, horizontal to pendulous, brownish; annulus well
nif ented in 1 row, deciduous; operculum convex, apiculate; stomata superficial (or
slightly sunken, but scarcely immersed); exothecial cells irregularly hexagonal, with
walls somewhat wavy and thickened at the corners; endostome segments perforate, the
cilia well-developed, slightly nodulose, in groups of 2-3. Spores 15-20u, faintly
roughened

Grout, Mosses with Hand-lens and Microscope, Pl. 45. Jennings, Mosses of Western
Pennsylvania, (ed. 1) Pl. 18 (as Webera), (ed. 2) Pl. 22 (as Pohlia ct —Fig. 444,

O mp coil of roadbanks or on banks of streams oe ditches. ge Nova Scotia to
Michigan os Iowa, south to Ohio and the District of Colum

MACKINAC CO.—Mackinac Island.

The very small, shortly pyriform capsules and the short exothecial cells with
wavy walls thickened at the corners are recognizable features of this rare species.

6. Pohlia wahlenbergii (Web. & Mohr) Andr.—Plants in extensive, loose or dense,
soft tufts 1.5-7 cm. high, whitish- or glaucous-green when moist, pale green, yellowish,
or brownish when dry, not shiny. Stems reddish, forked. Leaves rather distant, erect or

156

erect-spreading, slightly twisted when dry, 1.2-2.6mm. long, lanceolate or ovate-
lanceolate, acute, slightly to distinctly decurrent; margins plane or somewhat reflexed
below, especially when dry, serrulate toward the apex; costa yellowish or reddish,
ending near the apex; upper cells rather lax and thin-walled, oblong-rhomboidal, 3-7:1,
somewhat narrower toward the margins. Dioicous; perigonia large and conspicuous,
cupulate, the bracts with spreading points. Setae 25-40 mm. long, slender and flexuose;
capsules pendulous, 1.5-2 mm. long, shortly oblong-ovoid from a short neck; annulus
none; operculum low-conic, apiculate; exothecial cells hexagonal, with slightly irregular
outlines, not or slightly collenchymatous; stomata immersed; endostome segments
perforate, the cilia nodulose, in 2’s and 3’s. Spores 15-20 yu, minutely papillose.—n =
10, 10+1, 11

now the Mosses, (ed. 2) fig. 144. Grout, Mosses with Hand-lens oy
Tod a see (as Searches albicans). Jennings, ae of Western Pennsylvania, (ed. 1) P
18 & (ed. 2) Pl. 22 (as Mniobryum). Welch, Mosses of Indiana, fig. 118.—Fig. 445-448.

On wet soil or rock in seepage around springs, also in ditches and in the overflow of brooks.
Circumpolar; also North Africa, South America, and Kerguelen; Greenland to Alaska, south to
North Carolina, Arkansas, and Guatemala.

CHEBOYGAN CO.—Reeses Bog, Hermits Bog, Burt Lake. EMMET CO.—Maple River, Big
Stone Bay. MACKINAC CO.-—Bois Blanc Island, St. Martins Point, Epoufette. PRESQUE ISLE
CO.—Lamprey Experiment Station, Evergreen Beach, Ocqueoc River.

Easily recognized in nature by the seepy habitat and soft, whitish leaves
contrasting with red stems. Dried plants lose this whiteness and contrast and may be
mistaken for Philonotis fontana, which is larger, coarser, densely tufted, and at
white (differing more in texture than in color).

Pohlia filiformis (Dicks.) Andr., a rare montane species occurring in the Lake
Superior region, differs from all our other Pohlias in having small, julaceous stems with
shiny leaves which are erect or appressed, concave, broad (ovate-lanceolate to
subelliptic) and broadly pointed to obtuse and apiculate.

3. Bryum Hedw.

Small to robust plants, gregarious to densely tufted. Stems mostly forked, often
radiculose. Leaves usually small and remote below, larger and more crowded above,
sometimes in + rosulate tufts, erect-appressed to erect-spreading, usually not much
altered on drying, + concave, mostly rather broad, ovate to elliptic or lanceolate,
sometimes decurrent, usually acute or acuminate, sometimes awned, rarely obtuse or
rounded, often bordered by narrow cells; costa strong, rarely ending below the apex,
often excurrent; upper cells usually rather broad and transparent, mostly rhomboid-
hexagonal; lower cells rectangular. Setae terminal, elongate, curved or hooked at the
tip; capsules mostly nodding or pendent, subcylindric, clavate, or pyriform, rarely
ovoid or subglobose, symmetric or somewhat curved, especially in the neck which is
well developed and usually evident, smooth; annulus revoluble; operculum convex or
conic, umbonate or rarely apiculate; peristome double, the 16 teeth lanceolate,
slenderly acuminate, yellow-brown, mostly hyaline at the tips, papillose on the outer
surface, often bordered, trabeculate at back; endostome with a well-developed basal
membrane, keeled and usually perforate segments (very rarely lacking), and slender,
often nodose or appendiculate cilia (rarely rudimentary or lacking).—A Greek word for
moss, used by Dioscorides for some cryptogam and applied by Dillenius to this genus
and its relatives. A large and difficult genus—the Crataegus of bryology. Specific

Loy

a
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Se eee =

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SN
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ee ij 4o\dV |
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ff \
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We

“449 le

Pohlia lescuriana, 444. Hab
Pohlia wahlenbergii. 445.

and dry.
ryum algovicum. 449. Leaves. 450. Upper cells of leaf. 451. Capsule. 452. Portion of

it.
Habit. 446. Leaves. 447. Upper cells of leaf. 448. Capsules, wet

peristome.

characters are based primarily on sexual conditions and details of the peristome.
Fortunately, the seven species in our area are easily separated, and three of them can
be recognized in a sterile condition and in the field.

1. Endostome with cilia rudimentary or lacking
2. Leaf margins revolute; polygamous; capsules rarely 4mm. long; endostome adherent to
exostome in the lower % or more; lamellae at back of exostome teeth extensively joined
1. B. algovicum

i
2. Leaf margins not or slightly revolute; autoicous; capsules unusually long and slender,
46 mm. long; endostome free; lamellae at back of exostome teeth not extensively join
by vertical walls 2. B. uliginosum
1. Endostome with cilia well developed
3. Small, silvery, julaceous plants; upper leaf cells hyaline, conspicuously differentiated from
lower cells 3. B. argenteum

158

3. Larger plants, not silvery or julaceous; upper cells not hyaline or conspicuously
differentiated from lower cells
4. Costa not or rarely somewhat excurrent
5. Leaves noticeably decurrent (especially on sterile stems); leaf margins revolute
5.

throughout; upper cells thick-walled fiak ele
5. Leaves not particularly decurrent; leaf margins + reflexed below; upper cells nae
thin-walled 4.B. ain
4, Costa regularly and distinctly excurrent :
6. Synoicous; upper cells 3-4: 6. B. lisae var. cuspidatum
6. Dioicous; upper cells about 7:1 7. B. caespiticium

1. Bryum algovicum Sendtn. ex C. M.—Plants variable in size but usually about
5mm. high, densely tufted, often radiculose below, green or yellowish-green. Leaves
erect and slightly contorted when dry, erect-spreading when moist, somewhat larger,
more crowded, and spreading at the stem tips, ovate-lanceolate, slenderly long-
acuminate, somewhat decurrent; margins revolute; costa reddish at base, long-excurrent,
remotely toothed near the tip; upper median cells oblong-hexagonal, about 2-3:1, fairly
thick-walled, gradually narrowed toward the margins forming an ill-defined border,
longer and narrower in the acumen. Polygamous (usually synoicous, but often
appearing autoicous or dioicous; innovations on fruiting plants often ending in a
perigonium). Setae (4)15-40 (rarely 60) mm. long, red or red-brown; capsules
pendulous, 1.5-3.5, rarely 4mm. long, pyriform to clavate-cylindric, with a well-
developed neck; operculum small, convex-conic, mammillate to apiculate; peristome
teeth brown and opaque below, hyaline above, the lamellae at back extensively joined
by vertical walls; endostome well developed but usually adhering to the exostome in
the lower half or more, the segments perforate, the cilia none or 1-2 and irregularly
developed. Spores 18-37 u (somewhat variable in size even in the same capsules), finely
papillose to nearly smooth.—n = 10, 27, 30.

Conard, How to Know the Mosses, (ed. 2) fig. 148e (as B. ae Grout, Moss Flora of
ee America 2: Pl. 84 & 97B (as B. pendulum). Jennings, Mosses of West n Pennsylvania, (ed. 2)
22 (as B. angustirete), Welch, Mosses of Indiana, fig. 119 (as B. pendilhirn), “Fig. 449-452.

n sand or gravel, often in seepage; locally in depressions in low dunes near Lake Michigan
and Lake Huron. Circumpolar; Greenland to Alaska and south to the District of Columbia, Ohio,
Illinois, Colorado, and Arizona

EMMET CO.—Cecil Bay, Big Stone Bay, Waugoshance Point. PRESQUE ISLE CO.—Grace
Harbor, Evergreen Beach.

Bryum algovicum grows in the sands along the shores of the Great Lakes, usually
in depressions which are wet or seepy during part of the year. It has the ability to
continue growth through successive layers of sand. Its unmistakable feature, the
chambered appearance of the peristome caused by the adherence of endostome to
exostome around air spaces, is often best observed microscopically before complete
soaking. With a handlens, the species can be recognized by the apparent absence of the
endostome and the tendency for the teeth to stand erect rather than incurved, as in
mosses with endostome and exostome separate.

2. Bryum uliginosum (Brid.) BSG—Plants in loose or dense, green, yellowish, or
brownish tufts, rarely more than 10 mm. high. Leaves equally spaced, not particularly
crowded, erect and contorted when dry, erect-spreading when moist, up to 5mm
long, ovate-lanceolate, long-acuminate, not decurrent; margins not or slightly revolute,
serrulate near the apex; costa yellowish, percurrent or short-excurrent; cells relatively
lax and thin-walled, oblong-hexagonal or short-rhomboidal, about 3-3.5:1 above, linear
in several marginal rows forming a well-marked border. Autoicous; perigonia small and

159

inconspicuous, at the ends of short branches. Setae usually very long, 25-75 mm. long,
red or red-yellow; capsules suberect to horizontal or + pendulous, 4-6 mm. long,
clavate or clavate-cylindric (typically very long and slender), oblique at the mouth, the
well-defined neck usually about as long as the urn, + curved; operculum convex-conic,
sometimes apiculate; peristome teeth yellow-brown; endostome segments rather broad,
cilia none or rudimentary (or occasionally long and nodulose but weak and coherent,
in 3’s). Spores 18-29 y, finely papillose.—n = 10.

Conard, How to Know the Mosses, (ed. 2) fig. 148a-d. Grout, Moss Flora of North America
2: Pl. 87 below (as B. cernuum). Jennings, Mosses of Western Pennsylvania, (ed. 1) Pl. 18 (as B.
cernuum).—Fig. 453-455.

On peaty or sandy banks of streams or on rotten wood in wet places, probably a calciphile.
Circumpolar; Labrador to Manitoba, south to New a and Illinois; in the West, from British
Columbia and Alberta to Oregon; Texas and New Mex

CHEBOYGAN CO.~—Iron Bridge (over Carp Creek).

A great rarity in eastern North America, known locally from a single locality;
best recognized by the long setae and long, slender capsules.

3. Bryum argenteum Hedw.—Small plants in loose or dense, whitish or silvery
tufts 3-12 mm. high. Stems reddish, forked. Stem leaves few, ovate-lanceolate. Branch
leaves imbricate wet or dry, concave, 0.5-1.2 mm. long, broadly ovate, apiculate to
filiform-acuminate, not bordered; margins entire, plane or slightly reflexed below; costa
slender, reddish at base, ending well below the apex or disappearing in the acumen;
upper cells rhomboidal, hyaline and rather thin-walled toward the apex, thicker-walled
and yellowish below, and subquadrate to short-rectangular and reddish in an extensive
basal area. Dioicous. Setae 8-20 mm. long, red; capsules 1-2 mm. long, pendulous to
pendent, oblong-cylindric from a short neck, usually red; operculum low-conic, blunt
to acute or apiculate; peristome teeth brownish below, hyaline above; endostome
segments keeled and perforate, the cilia nodose to appendiculate, in 2’s and 3’s. Spores
10-15 uw, smooth or faintly roughened.—n = 9+1, 10, 10+1, 11, 12 (as var. lanatum),
20.

Con How to Know the Mosses, (ed. TOR 149. Jennings, Mosses of Western
Rom ahr (ed. 1) Pi. 20, ek 2) Pl. 24.—Fig. 456-4

On bare soil in disturbed places, in old fields, along roads, in paths, in cracks of sidewalks,
etc. Cosmopolitan; widespread in North America.

CHEBOYGAN CO.-Biological Station campus, Iron Bridge, Colonial Point, Riggsville,
Duncan Bay, Weber Lake. EMMET CO.—Alanson, Cross Village, Five Mile Creek, Ramona Beach,
Cecil Bay. MACKINAC CO.—Mackinac Island, Nunns Creek, Bush Bay

The SILVERY BRYUM is easily recognized by the color of the leaves which are
crowded together on small, catkin-like innovations. The pink or red capsules are
attractive in contrast with the silvery gametophytes. The species is a ubiquitous weed,
thriving even in the atmospheric pollution of large cities. It is a pronounced nitrophile,
occurring in tell-tale abundance in urine-soaked soils around Eskimo habitations in the
Arctic, for example, or around dog kennels.

4. Bryum capillare Hedw.—Dark-green or brownish plants in dense tufts usually
less than 10 mm. high. Filiform, brown, papillose brood-bodies often clustered in leaf
axils. Leaves erect and usually + spirally contorted when dry, erect-spreading when

160

dS

VIZ SLIVYYOR

\ f \ ‘ ‘
\ Vase ) SL \ SAS
\\ TOS
7 NATO

Bryum uliginosum. 453. Leaves. 454. Upper cells. 455. Capsule.

Bryum argenteum, 456. Habit. 457. Leaves of julaceous branches. 458. Upper cells of leaf.
459. Cells at basal margin of leaf. 460. Capsule.

Bryum capillare. 461. Habit. 462. Leaves. 463. Upper cells of leaf. 464. Brood-body. 465.
Capsule.

=

moist, elliptic to obovate, abruptly cuspidate to awned; margins plane or variously
reflexed, serrulate toward the apex; costa ending well below the apex to long-
excurrent; upper cells oblong-hexagonal, 1.5-3:1, green, thin-walled, with 1-2 (rarely 4)
rows of linear cells in a pale, narrow, unistratose border. Dioicous. Setae about
20-35 mm. long; capsules inclined to pendulous, clavate-cylindric, 2.5-4.5mm. long;
operculum convex-conic, apiculate; cilia 14, appendiculate. Spores 8-15 uy, nearly
smooth.—n = 9, 10, 10+1, 10+2, 10+2-3, 12, 15, 20, 24.—Most, if not all collections
from eastern North America belong to the var. flaccidum (Brid.) BSG, a lax expression
of the species with remote leaves only moderately spiral-contorted when dry and
typically provided with axillary, filiform brood-bodies. The range of the variety is
probably much like that of the species.

161

Breen, Mosses of Florida, Pl. 62 (figs. 14). Conard, How to Know the ‘Mosses, (ed. 2) fig.
5 Grout, Mosses with Hand-lens and Microscope, Pl. 48. Jennings, Mosses of Western
Pennsylvania, (ed. 1) Pl. 20, (ed. 2) Pl. 24.—Fig. 461-465.

On rock, soil, or humus, especially on banks, commonly on bark at base of trees or in
crotches or drainage channels on tree trunks. Circumpolar; Aleutian Islands and across the
continent of North America south to California, Arizona, Texas, and Florida; Mexico, Central
America, Chile; West Indies.

CHEBOYGAN CO.-—Riggsville, Little Lake 16, Colonial Point, south end of Burt Lake, Mill
Creek, Weber Lake. EMMET CO.—Pellston Hills, Cros s Village, Five Mile Creek, Cecil Bay, Big
Stone Bay. MACKINAC CO.—Cut River, Ozark, Caffey as Little Dollar Lake.

The spirally contorted, dark-green leaves with thin-walled cells are distinctive, and
with a little practice, Bryum capillare can be recognized in the field by an aspect
rather like that of a Mnium. The brown, filiform brood-bodies, looking like stout
rhizoids, also help.

5. Bryum pseudotriquetrum (Hedw.) Gartn., Meyer & Scherb.—Relatively Sa
plants, in dense, dark-green or reddish tufts, radiculose below, 1.5-5, sometimes
high. Stems simple or forked, red, rarely felted above with filamentous, ae
brood-bodies, + tomentose below. Leaves usually rather remote below, somewhat larger
and more crowded above, twisted and contorted when dry, erect-spreading when moist,
2.5-4mm. long, oblong-lanceolate to ovate-lanceolate, acute (and often cuspidate) to
acuminate, narrow at the insertion and narrowly but distinctly decurrent; margins
entire or serrulate near the apex, strongly revolute throughout; costa red below, red or
brownish above, percurrent or short-excurrent; upper cells oblong-hexagonal, 2-3:1,
with firm walls, linear and very thick-walled in several marginal rows forming a strong,
often colored border. Dioicous or synoicous. Setae 25-60 mm. long, dark-red below,
red-brown above; capsules 3-4mm. long, inclined to pendulous, usually straight,
clavate, the neck sometimes as long as the umm; operculum large, convex-conic,
apiculate; peristome teeth brownish-yellow; cilia of endostome well-developed, ap-
pendiculate, in 3’s (or sometimes also in 2’s and 4’s). Spores 11 -20 w, smooth or
faintly roughened.—n = 9, 10, 10+1, 11, 20, 22.

Conard,
75. Grout, Mosses with Hand-lens and Microscope, Pl. 47 (as B. binum). Jennings, Mosse
Western Pennsylvania, (ed. 1) PI. 19 & (ed. 2) Pl. 23 AnD binum). Welch, Mosses of Indiana, i
122.—Fig. 466-470.

nard, How to Know the ae (ed. 2) fig. i 7 Darlington, Mosses of Michigan, tye
of

On wet soil or humus, sometimes on rocks or decayed wood, common in swamps and near
brooks md onds. Circumpolar; South America; Greenland to Alaska, south to California,
Colorado, Arkansas, and North Carolina.

CHEBOYGAN CO.—Reeses Bog, Hermits Bog, Gorge of Carp Creek, fetes Bridge, Vincent
Lake, Livingston Bog, Inverness Township, Mud Lake, Duncan Bay, Grass B , Mill Creek, Trout
Creek. EMMET CO.—Pleasantview Swamp, Carp Lake, Arnott Lake, Saiees aoe Wycamp Lake,
Cecil Bay, Big Stone Bay, Conway Bog. MACKINAC CO.—Bois Blanc Island, Mackinac Island, St.
Martins Bay, McKays Creek, Cut River, Ozark. PRESQUE ISLE CO.—Evergreen Beach, Hammond
Bay, Mast Point, Grace Harbor, Rainy River Falls, Clinton Lake.

Very common in wet habitats and easy to recognize in a sterile state because of
red stems and obviously decurrent leaves with rather broad points. It has often been
confused with B. weigelii Spreng., which occurs in more northerly parts of Michigan
and has soft leaves, shriveled when dry, with lax, thin-walled cells and an indistinct
border.

162

Bryum tortifolium Funck ex Brid., which occurs on Sugar Island in nearby
Chippewa County, can be characterized as follows: weak, slender plants in soft, loose,
green to yellow-brown tufts; leaves oblong-ovate to elliptic, rounded at the apex,
inconspicuously decurrent, plane and entire at the margins; costa ending somewhat
below the apex; cells 2-5:1, rather thin-walled, longer and narrower in a few marginal
rows in an indistinct border. Dioicous; peristome complete, with appendiculate cilia. It
has been confused with red plants of the arctic properly referred to B. cryophilum
Mart. It is larger and has leaves with more distinctly differentiated borders which are
very narrowiy revolute below.

6. Bryum lisae De Not. var. cuspidatum (BSG) Marg.—Plants in dense, green or
yellowish tufts usually scarcely more than | cm. high. Leaves small and distant below,
larger and crowded above, moderately contorted when dry, oblong-lanceolate to
ovate-lanceolate, acuminate, ending in a smooth or + toothed point formed of the +
long-excurrent costa; margins revolute nearly to the + serrulate apex, eee
bordered; upper cells rhomboidal (about 3-4:1), usually longer in the acumen, linear
2-5 marginal rows. Synoicous. Setae up to about 2.5 cm. long; capsules horizontal
subpendulous, up to about 3 mm. long, the neck generally somewhat shorter than the
rest of the capsule; annulus revoluble; operculum convex and apiculate; cilia of
endostome in 2’s and 3’s, appendiculate. Spores 11-18 yu, finely papillose.—n = 10, 20,
also 22 and 30 (as B. affine)

Grout, Moss Flora of 8 America 2: Pl. 92B (as B. cuspidatum). Jennings, Mosses of
Western Makes (ed. 1) Pl. 19 (as B. affine ao intermedium), (ed, 2) Pl. 23. Welch, Hae
of Indiana, fig. 120 (as B. By tee —Fig. 471-473.

weed, on rocky or sandy, often wet soild in disturbed places, especially on roadbanks or
sides 2 ditches, sometimes also on rock, humus, or rotten wood. Circumpolar; widespread in North
Americ

CHEBOYGAN CO.—Reeses Bog, Gorge of Carp Creek, Douglas 7 Smiths Bog, Ellis
Township, Long Lake, Mill Creek, Little Lake 16, Weber Lake. EMM T CO, —Conway Bog,
Wycamp Lake. PRESQUE ISLE CO.—Presque Isle Lighthouse, Hammond ai Evergreen Beach.

Recognized by its weedy habitat, perfect peristome, and synoicous inflorescence,
it is most likely to be confounded with B. caespiticium.

The name Bryum cuspidatum (BSG) Schimp., so long in use for this common
and widespread species, is nomenclaturally unacceptable (being a later homonym), and
B. creberrimum Tayl., recently in use as a replacement, has proved to be something
else. Until various synonyms at the specific level can be tested and verified, one can
use the name B. lisae var. cuspidatum.

7. Bryum caespiticium Hedw.—Plants densely caespitose or gregarious and mixed
with other mosses, light- or yellow-green, rarely more than 1 cm. high. Leaves small
and distant below, larger and crowded above, ovate-lanceolate, long-acuminate; margins
revolute, entire or slightly serrulate in the apex; costa red at base, yellowish above,
long-excurrent as a smooth or slightly serrulate point; upper cells thomboidal, about
7:1, somewhat longer toward the apex and the margin but not forming a distinct
border; basal cells broader and shorter, the alar cells subquadrate. Dioicous. Setae up
to 3cm. long; capsules inclined to pendulous, up to 4mm. long, clavate to oblong-
obovoid, sometimes curved, the neck shorter than the rest of the capsule; annulus
revoluble; operculum short-conic; stomata in the neck; cilia of endostome in groups of
3-4, appendiculate. Spores 12-15 wu, or nearly smooth.—n - 9+1, 10, 10+1, 10+5, 11,
20, 20+1 (and, experimentally, 20, 40, etc.).

163.

\
i

Bryum pseudotriquetrum. 466. Habit. 467. Portion of sterile stem showing decurrent leaves.
468. Leaves. 469. Upper cells of leaf. 470. Capsule.
m lisae var. cuspidatum, 471. Leaves. 472. Upper cells of leaf. 473. Capsule.
Bryum caespiticium. 474. Leaf. 475. Upper cells of leaf. 476. Capsule.
Rhodobryum roseum, 477. Habit. 478. Leaf. 479. Upper cells at margin of leaf.

Conard, How to Know the Mosses, (ed. 2) fig. 151a-d. Grout, Mosses with Hand-lens and
Microscope, fig. 113. Jennings, Mosses of Western Pennsylvania, (ed. 1) Pl. 20, (ed. 2) Pl. 24.
Welch, Mosses of Indiana, fig. 121.—Fig. 474-476.

On soil in disturbed places. Circumpolar; Tasmania and New Zealand; widespread in North
America.

CHEBOYGAN CO.—Reeses Bog, Colonial Point. EMMET CO.—Pellston, Big Stone Bay.

164

Like many species of Bryum, B. caespiticium is based on highly technical
characters. I recognize it by its weedy habitat, perfect peristome, and dioicous
inflorescence. (It has narrower leaf points with longer cells and a silkier over-all
appearance than B, lisae var. cuspidatum, but I am unable to recognize it with certainty
by those characters alone.)

Other species of Bryum known from the Upper Peninsula:

Bryum muehlenbeckii BSG grows on wet rocks near streams in dense, dull,
reddish or dark-green tufts tinged with red or brown. It looks rather like a Pohlia. The
leaves are concave and often cucullate, broad and broadly pointed, with erect margins
and a red, percurrent costa. The cells are rhomboidal (3-4:1), with firm, reddish walls;
they are narrower in a single marginal row. The plants are dioicous, and the peristome
is complete, with well-developed segments and cilia.

Bryum pallens (Brid.) Sw. ex Roehl. grows in wet places, but not always on
rock. It is somewhat like B. creberrimum but is dioicous, with spores larger (19-24 y),
leaves more abruptly narrowed to a shorter tip, costa ending in a shorter excurrency,
and cells shorter (2-3:1) and laxer.

4. Rhodobryum (Schimp.) Hampe

Plants fairly robust, dark-green and often red-tinged, gregarious or loosely tufted.
Primary stems subterranean, rhizome-like; secondary stems erect, rosulate-foliate,
innovating through the rosettes or below them. Leaves remote, small, and scale-like
below, much larger and crowded at the stem tips, contorted when dry, wide-spreading
at least when moist, oblong-obovate, broadly acute and often abruptly pointed,
generally bordered and toothed above; costa strong, very broad below, tapered above,
ending slightly below the apex to excurrent as a sharp point; upper cells large and
broad, oblong-hexagonal, firm-walled; lower cells elongate-rectangular. Perichaetial
leaves smaller, lance-acuminate. Setae 1-3 or more per perichaetium, long, + hooked or
curved at the tip; capsules horizontal to pendent, curved, oblong-cylindric with a short
neck, smooth; annulus present; operculum convex and apiculate; peristome double, the
16 teeth lanceolate, yellow- to red-brown below, hyaline at the tips. finely papillose, +
bordered, trabeculate at back; endostome consisting of a high basal membrane, keeled
and perforate segments, and nodose to appendiculate cilia._The generic name translates
into the common name, ROSE MOSS, and it refers to a flower-like arrangement of
leaves at the tips of the upright stems, and perhaps to red tinges, as well.

Rhodobryum roseum (Hedw.) Limpr.—Plants in loose tufts 1-5 cm. high. Upper
leaves crowded in a rosette, erect and variously twisted and contorted when dry,
wide-spreading when moist, 5-10mm. long, oblong-obovate, abruptly cuspidate;
margins revolute in the lower 2/3, serrate in the upper 1/3-1/2; costa ending in the
cuspidate apex or shortly excurrent; upper cells green, oblong-rhomboidal, about 4-6:1,
thin- or moderately thick-walled, considerably pitted, 1-2 rows at the margins narrow
and yellowish in a + distinct border. Dioicous. Setae 1-8 from the same perichaetium,
25-40 mm. long; capsules 4-7 mm. long, oblong-cylindric from a short neck, somewhat
curved, horizontal to pendent; annulus revoluble; operculum conic, acute; cilia of
endostome in 3’s and 4’s, appendiculate. Spores 15-22 yu, very minutely roughened to
finely papillose.—n = 10, 10+1, 11 (and, experimentally, with colchisine, 20).

165

Conard, Sees ae Mosses, (ed. 2) fig. 147. Darlington, Lae of Michigan, fig. 76.
Grout, Mosses ith Ha nd-len d Microscope, fig. 116. Jennings, Moss Western Pennsylvania,
(ed. 1) Pl. 20 (as R. oN (ed. 2) Pl. 24. Welch, Mosses of Indiana, "fie, 126.—Fig. 477-479.

On thin humus or soil over boulders in woods, or sometimes on the forest floor, on logs, or
bark at the base of trees. Circumpolar; Newfoundland oe Manitoba, south to North Carolina and
Akransas; British Columbia and Alberta; Arizona and Mexi

CHEBOYGAN CO.—Grapevine Point, Reeses Bog, Gorge of Carp Creek, Fontinalis Run, Mud
Lake, Ellis Township, Little Lake 16. EMMET CO.— W. Branch of Maple River, Harbor Springs,
Five Mile Creek, Middle Village, Big Stone Bay, Cecil Bay. MACKINAC CO.—Bois Blanc Island,
Bush Bay, McKays Creek, Ozark. PRESQUE ISLE CO.—Presque Isle lighthouse, Rainy River Falls,
Clinton

A very attractive moss, in many ways similar to a Mnium, but actually more
nearly related to Bryum as evidenced by the elongate leaf cells and rather indistinct
border of narrower, yellowish cells. Sporophytes are rare.

It has been suggested by numerous authors (and most recently Iwatsuki &
Koponen, Acta Bot. Fenn. 96: 1-22. 1972) that all our material in eastern North
America should be referred to R. ontariense (Kindb.) Kindb. (although the differences
are slight and the ranges greatly overlapping).

Mielichhoferia mielichhoferi (Funck ex Hook.) Loeske (fig. 480-483), one of the
rare copper mosses, has been found at Pictured Rocks along the south shore of Lake
Superior. The plants are dioicous, with both male and female inflorescences lateral (at
or near the base of stems or at the dichotomies formed by branches); the capsule is
erect and symmetric and the peristome single, consisting of endostome only. Informa-
tion on the biology of the copper mosses is given above in a discussion of
Scopelophila.

By analyzing rocks and mineral uptake, Coker (Trans. Brit. Bryol. Soc. 6(2):
317-322. 1971) was able to determine the probable origin of Scottish specimens of
Mielichhoferia, (a copper moss) and Saelania (a calciphile) with inadequate collection
data.

MNIACEAE
Plants small or, more often, cane Sea gregarious to loosely tufted, dull,
dark-green or yellow-green, often + entose. Stems erect, simple or forked, rarely

small and remote below, larger and more crowded above, often forming a terminal tuft
or rosette, usually not greatly contorted when dry, mostly broad, elliptic to obovate,
usually broadly pointed and often apiculate, frequently bordered by linear cells and
often strongly toothed, the teeth sometimes in pairs; costa single, strong, broad at the
base, tapered above, generally ending somewhat below the apex; cells mostly rounded-
quadrate or hexagonal, mostly firm- or thick-walled, smooth (or very rarely mam-
millose), becoming longer and mostly rectangular below. Perigonia discoid, usually with
clavate paraphyses; perichaetia bud-like, with small, lanceolate inner bracts and few,
filiform paraphyses. Setae terminal, single or clustered, mostly elongate, usually +
hooked at the tips; capsules inclined to pendent, very rarely erect, symmetric or nearly
so, oblong-ovoid to cylindric, rarely subglobose, short-necked, smooth; annulus usually
present; operculum convex to obliquely rostrate; stomata nearly always immersed,
mostly restricted to the neck; peristome double, with 16 lanceolate teeth; endostome
typically with a high basal membrane, keeled segments alternating with the teeth, and
usually well-developed cilia (in Orthomnium the endostome is reduced to a rather low
membrane; in Cinclidium the endostome is longer than the teeth, and the segments are
coalesced at their tips). Spores spherical. Calyptrae mostly cucullate.

166

Endostome about as long as the exostome, consisting of a high basal membrane, see

segments, and well-developed cilia; leaves mostly toothed but, if entire, not ending in a
long, cuspidate point 1. Mnium

Endostome considerably longer than the exostome, consisting of a pleated dome pees

by keeled columns alternating with the exostome teeth; leaves entire, abruptly cuspidate
2. Cinclidium

1. Mnium Hedw., nom. cons.

Medium-sized to fairly robust plants in light- to dark-green, yellowish, brown, or

reddish tufts, often + tomentose below. Stems simple or forked by innovations, erect
(or sometimes sterile stems spreading or prostrate, + complanate-foliate, and stoloni-
form-attenuate). Leaves erect- to wide-spreading when moist, generally crisped and
contorted when dry, elliptic or oblong-ovate to obovate or nearly rounded, broadly
pointed to rounded at the apex and frequently mucronate to cuspidate, usually
bordered by linear cells in 1-several layers, entire to strongly toothed, sometimes in
pairs; costa ending well below the apex or more often subpercurrent to shortly
excurrent; upper cells hexagonal to rounded- or oblong-hexagonal, often thickened at
the corners and minutely pitted, smooth, frequently in oblique rows, longer and
oblong near the base. Inner perichaetial leaves small and lanceolate. Setae single or
clustered, elongate, smooth; capsules suberect to inclined or pendent, oblong-cylindric
from a short, usually inconspicuous neck; annulus present; operculum convex-conic,
sometimes apiculate to rostrate; stomata mostly immersed and nearly always restricted
to the neck; peristome teeth inserted near the mouth, lance-acuminate, yellow or
ellow-brown, densely and finely papillose, bordered and trabeculate; endostome
yellowish to brown, finely papillose, with a high basal membrane, broad segments
perforate and finally gaping along the keels, often with irregular margins, cilia well
developed, nodulose, in groups of 2-4. Calyptrae cucullate, smooth, naked.—An ancient
Greek name for a moss. Mnium is often confused with Bryum, perhaps because of
similarly nodding capsules. The leaves in Mnium are generally broader and variously
elliptic to obovate and strongly bordered, the leaf cells are more or less isodiametric
rather than oblong-hexagonal in a kind of fish-net pattern, and the capsules are
cylindric rather than tapered to the base by way of a well-defined neck.

The genus is a traditional one, and I prefer to retain it as a convenience, even
though it can be subdivided on the basis of good structural and cytological differences.
Following Koponen (Ann. Bot. Fenn. 5: 117-151. 1968), one could assign our species
to the following genera: Mnium—with double-toothed leaves (MM. lycopodioides, M.
marginatum, M. orthorrhynchum, M. spinulosum, and M. stellare); Plagiomnium—with
single-toothed leaves (M. cuspidatum, M. drummondii, M. affine, M. medium, and M.
longirostrum), Rhizomnium—with entire leaves and a thick cartilaginous border (M.
punctatum and M. pseudopunctatum); and Pseudobryum—with entire, scarcely
bordered leaves and diagonally elongate cells (M. cinclidioides). Koponen’s study is one
of the few papers on mosses attempting to analyze species relationships on the basis of
primitive vs. advanced characters.

Lowry (Mem. Torrey Bot. Club 20, part 2, pp. 142. 1948) laid the cytological
basis for such a generic separation (using, in the main, Michigan materials) and also
showed a haploid-diploid relationship between M. orthorrhynchum and M. marginatum
and also between M. affine and M. medium. In both pairs of species, the differences
are essentially quantitative and sexual, the haploids being smaller in various ways and
dioicous, the diploids monoicous (paroicous or synoicous). This raises the interesting
possibility of speciation through natural apospory (the production of polyploid,
bisexual gametophytes by the regeneration of sporophytic tissue). It is known, of
course, that apospory can be induced in many mosses, as demonstrated especially by
the well publicized research of the Marchals and Wettstein. (See references following
Physcomitrium, above.

167

1. Leaves distinctly toothed
2. Leaf border none or obsolete 1. M. stellare
2. Leaf border ee ee
3. Leaves doubly se
4. Dioicous
5. Upper cells firm-walled, only slightly thickened at the corners 4. M. orthorrhynchum
5. Upper cells with delicate, pellucid walls clearly thickened at the corners
2. M. lycopodioides
4. Paroicous (and sometimes synoicous)
6. ee leaf cells hexagonal, with thin walls not thickened at the corners; setae often
. Sspinulosum
6. Upper eu rounded-hexagonal or subquadrate, with firm walls thickened at the
orners; setae single 3. M. marginatum
3. Te singly serrate
7. Leaves toothed only in the upper hal

f
8. Cells 14-26 u, irregularly rounded-hexagonal, with rather thick walls thickened at the

corners; setae sing cuspidatum
8. Cells 25-50 w, hexagonal, with thin or firm walls not thickened at the corners; ae
generally in groups of 7. M. drummondii
7. pr aa nearly to the base
9. Dio 8. M. affine
9: Seen (or sometimes paroicous)
10. Opercul + apiculate; ieee restricted to the neck; leaves cuspidate from a
rounded-obtuse apex, sharply toothed; upper leaf cells 28-55 (77) pw . M. medium
10. Operculum long-rostrate; panes scattered over much o e capsule; aa
cuspidat

a rounded-obtuse to truncate or ee apex, bluntl
10. M.

Sree
1. Leaves entire (or very indistinctly toothed)

11. Leaves obscurely bordered by sue narrower cells in 3-4 marginal rows, obscurely
serrulate; upper cells elongate, in oblique rows 1. ; eu cuaiiors

11. Leaves distinctly bordered by linear an entire; upper cells not elongate or in obliqu
IOWS

12. Dioicous 12. M. punctatum

12. Synoicous 13. M. pseudopunctatum

1. Mnium stellare Hedw.—Rather small plants in loose, dark-green or red-brown
tufts up to 2 or 3, sometimes 5 cm. high, somewhat radiculose below. Stems erect or
ascending, dark-red at least below. Leaves loosely erect or spreading, somewhat wavy
or irregularly curved when dry, 1.5-2.5 mm. long, elliptic or ovate-elliptic, obtuse and
often abruptly short-pointed, decurrent, not or very indistinctly bordered, the upper
leaves irregularly denticulate toward the apex because of broad, unicellular teeth
mostly projecting at an angle on both surfaces, sometimes nearly paired; costa ending
well below the apex, brown or reddish; upper cells irregular in size and shape, 15-30 u,
hexagonal or rounded- to quadrate-hexagonal, with thin or slightly thickened walls
somewhat thickened at the corners, 1-2 marginal rows sometimes slightly longer and
not or somewhat darker. Dioicous. Setae 10-20 mm. long, single, reddish; capsules
2-3 mm. long, oblong-cylindric from a short neck, pendulous to pendent, pale- to
dark-brown, darker at the neck; peristome teeth yellow-brown; endostome golden-
brown, with cilia in groups of 2-4. Spores 20-29 y, very finely papillose.—n = 6+1, 7.

Conard, How to Know the Mosses, (ed. 2) fig. 157. Grout, Mosses with Hand-lens and
Mico ene fig. 21. Jennings, Mosses of Western Bees. (ed. 1) Pl. 22, (ed. 2) Pl. 26.
Welch, Mosses of Indiana, fig. 127.—Fig. 484-485.

On soil, humus, logs, and rock but most ’often on the bases of trees and ee. in wet
woods. Erne and Asia; Nova Scotia to Saskatchewan, south to North Carolina and Arkansa

CHEBOYGAN CO.—Reeses Bog, Gorge . Carp Creek, Iron Bridge, Mill Creek, Mud Lake.
EMMET CO.—Larks Lake, Pleasantview Swamp, Middle Village, Wycamp Lake, Sturgeon Bay.
MACKINAC CO.—Mackinac Island, Castle Rock, feyae PRESQUE ISLE CO. ie River Falls.

168

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Mnium marginatum, 487. Upper cells of leaf. 488. Capsule.
Mnium orthorrhynchum. 489. Leaves. 490. Upper cells of leaf.
Mnium spinulosum. 491. Leaf. 492. Upper cells of leaf.

Mnium stellare is easily recognized with a hand-lens by the virtual absence ofa
leaf border and the costa ending well below the apex.

2. Mnium lycopodioides Schwaegr.—Relatively slender plants in loose, light- or
dark-green tufts 1-4.5 cm. high, somewhat radiculose below. Stems reddish. Leaves
twisted and contorted when dry, erect-spreading when moist, 3-4, rarely 6.5 mm. long,
lanceolate or ovate-lanceolate, acute and + apiculate, decurrent, with a thickened
border of linear, pellucid cells, doubly serrate, costa reddish, percurrent or confluent

169

with the border to form an apiculus, generally toothed at back above; cells irregularly
rounded-hexagonal, with thin, delicate, and usually hyaline walls somewhat thickened
at the corners, 17-24 (occasionally 30) u. Dioicous. Setae single, orange-red, 15-40 mm.
long, capsules nearly horizontal or somewhat pendulous, 2.5-5.5 mm. long, oblong-
cylindric from a short neck, yellow-brown; operculum stoutly and obliquely rostrate;
stomata immersed, in the neck; peristome teeth greenish-yellow; endostome yellow-
brown, with perforate ee and cilia in 3’s. Spores finely papillose, 19-29 pw.
Calyptrae brown, smooth.—

Grout, Moss Flora of North America 2: Pl. 104A, 105 (fig. 3).—Fig. 486.

On damp, shaded soil or rock, often on cliffs, also on humus and rotten logs. Circumpolar;
British Columbia and Alberta to Oregon and Montana; Nova Scotia to Manitoba, south to
Kentucky and Michigan

CHEBOYGAN CO.—Douglas Lake.

The delicate leaf cells with hyaline walls somewhat thickened at the corners are
distinctive.

2. Mnium marginatum (With.) Brid. ex P.-Beauv.—Relatively slender plants in
loose, green or brownish tufts 1-3, sometimes 4 cm. high, somewhat radiculose below.
Stems reddish. Leaves crisped and contorted when dry, erect-spreading when moist,
2-4.5 mm. long, elliptic or oblong-ovate, acute and long-apiculate, decurrent, with a
thickened border of linear, usually reddish cells, doubly serrate at least near the apex
(the teeth often reduced, sometimes nearly lacking); costa subpercurrent or, more
often, confluent with the border to form the apiculus, reddish, not toothed at back;
upper cells 24-30, rarely 38, hexagonal or shortly oblong-hexagonal, with firm,
usually pale-brown walls thickened at the corners, often in longitudinal rows. Paroicous
or occasionally synoicous. Setae single, orange-yellow or reddish at least below,
13-25 mm. long; capsules 1.5-3.5mm. long, oblong-cylindric from a short neck,
yellow-brown, nearly horizontal to pendulous; operculum stoutly and obliquely
rostrate; stomata immersed, in the neck; exostome teeth brownish or golden-brown;
endostome golden-brown or brown, with segments perforate and cilia mostly in 3’s but
sometimes in 2’s or 4’s. Spores 22-32 py, finely papillose. Calyptrae brown, smooth.—n

arlington, Mosses of Michigan, fig. 77 (as M. serratum). Grout, Mosses with Hand-lens and
Microscope, Pl. 51 above (as M. serratum), Moss Flora of North America 2: 105 (fig. 1). Jennings,
Mosses of Western Pennsylvania, (ed. 1) Pl. 21 and (ed. 2) Pl. 25 (as M. serratum). Welch, Mosses
of Indiana, fig. 128 (as M. serratum).—Fig. 487-488

On soil or rock in moist, shady places, particularly on banks of streams or on cliffs.
Circumpolar; Mexico and Guatemala; Falkland Islands; Alaska to Arizona and New Mexico;
Manitoba and New Brunswick to Arkansas and North Carolina

CHEBOYGAN CO.—Reeses Bog, Mill Creek. EMMET CO.—Maple River, Wilderness State
Park. MACKINAC CO.-—Bois Blanc Island, Ozark.

This species and the next are distinguished by sex, cell size, and chromosome
number, M. marginatum being a diploid, M.,orthorrhynchum a haploid. M. marginatum
is paroicous (or rarely synoicous), and its leaf cells, 24-30 u, or more, are generally
arranged in longitudinal rows.

4. Mnium orthorrhynchum Brid.—Plants rather small and slender, in loose tufts
about 10-20, occasionally as much as 70cm. high, green with reddish tinges. Leaves

170

erect-spreading, + wavy and twisted when dry, 2-3, sometimes 4.5 mm. long, oblong-
ovate, acute to short-apiculate, decurrent, bordered by linear cells in more than 1
layer, doubly serrate; costa nearly always confluent with the border to form the
apiculus, not or slightly toothed at back near the tip, reddish; upper cells 15-22 y,
hexagonal, firm-walled, somewhat thickened at the corners. Dioicous. Setae single,
13-18 mm. long, reddish; capsules nearly horizontal to pendulous, relatively long and
slender, cylindric, 2-4mm. long, abruptly contracted to a short neck; operculum
obliquely rostrate, 1-1.5 mm. long; stomata immersed, in the neck; exostome teeth
greenish-yellow; endostome brown, with perforate segments and cilia in 2’s. Spores
20-31 wu, very finely roughened. Calyptrae brown, smooth.—n =

, Mosses with Hand-lens and Microscope, Pl. 51 below, Moss Flora of North America
2: Pl. CG (Ee. 2).—Fig. 489-490.

On damp, shaded rock, soil, humus, and bark at base of trees, common on stream banks or
cliffs. oe Greenland to Alaska, south to North Carolina, Arkansas, New Mexico, Arizona,
and Californ

CHEBOYGAN CO.—Douglas Lake, Reeses Bog, Hermits Bog, Iron Bridge, Burt Lake, Mill
Creek. EMMET CO.—Maple River, Cross Village, Wycamp Creek, Big Stone Bay. M INAC
CO.—Mackinac Island, McKays Creek, Ozark, Caffey Corner. PRESQUE ISLE CO.—Ocqueoc River.

Best recognized by the small cells (less than 24 uw) and dioicous inflorescence.

5. Mnium spinulosum BSG—Plants in loose, deep-green tufts tinged with red,
0.5-1 or sometimes 2.5 cm. high, radiculose only at base. Stems becoming red. Leaves
erect, only slightly contorted when dry, 2.5-3.5 (or sometimes 4.5) cm. long, broadly
obovate, cuspidate, decurrent, strongly bordered, the border thick and rounded in
section, red or brown, sharply double-serrate in the upper half or somewhat more;
costa shortly excurrent, brown or red, not toothed at back; cells 21-25, rarely 35 y,
irregularly hexagonal, with thin walls not thickened at the corners. Paroicous. Setae
single, paired, or sometimes in 3’s, 8-35 mm. long, red-yellow; capsules 1.5-3 mm. long,
oblong-cylindric, short-necked, horizontal to pendulous, brownish, dark red-brown at
the mouth; operculum obliquely rostrate from a convex-conic base; stomata immersed,
in the neck; peristome teeth dark, red-brown; endostome light- or golden-brown, with
cilia sometimes single but usually in 2’s and 3’s. Spores 18-21 u, finely papillose.—n =
8

Grout, Mosses with Hand-lens and Microscope, fig. 118f-h. Jennings, Mosses of Western
Pennsylvania, (ed. 2) Pl. 65.—Fig. 491-492.

On humus, soil, logs, and rocks in coniferous ee Circumpolar; Nova Scotia to Alaska,
south to Maryland, the Great Lakes, Colorado, and Ore

HEBOYGAN CO.—Reeses Bog, Hermits Bog, Gorge of Carp Creek, Douglas Lake,
eres Bog, Mud Lake, Grass Bay. EMMET CO.-—Arnott ie, Sherett ek Pleasantv view
Swamp, Pellston Hardwoods, Carp Lake, Cross Village, Big Stone Bay, Conway Bog, Ramona
Beach. MACKINAC CO. —~Mackinac Island, Bois Blanc Island, Cadi Rock, M oKay s Creek, Cut
River, Ozark, Little Dollar Lake, McNamara Pond. PRESQUE ISLE CO.—Mast Poin. Kelsey Pond.

Grout called this the RED-MOUTHED MNIUM, and it has been called the
FLAPPER MOSS in allusion to the same feature. The species is recognized by its
dark- cael leaves which are broad for the double-toothed group, as well as its red
peristom

171

. Mnium cuspidatum Hedw.—Plants in dark- or yellow-green, loose, often
extensive tufts up to 2 or 3.5 cm. high, somewhat radiculose below. Stems green, the
sterile ones spreading or horizontal, often alee attenuate, t+ complanate-foliate.
Leaves erect or erect-spreading, strongly crisped and contorted when dry, about
3-4.5mm. long, obovate, acute to cuspidate-acuminate, decurrent, bordered by 24
rows of linear cells in 1 layer, with single, sharp, unicellular teeth in the upper half;
costa percurrent or shortly excurrent, greenish; upper cells irregularly rounded-
hexagonal, rather thick-walled and + thickened at the corners, 14-26 uw. Synoicous.
Setae single, 10-30 mm. long, yellowish or brownish; capsules 2-3 mm. long, pendent,
yellowish or brownish; operculum convex-conic; stomata immersed, in the neck;
peristome teeth yellow; endostome brown, the basal membrane fenestrate, cilia in
groups of 2-4. Spores 24-31 y, finely papillose. —n = 6, 12.
nes with Lge lens and Microscope, fig. 117e-f (as M. Dac Moss Flora a

. Jennings, Mosses of Western Pennsylvania, (ed. 1) Pl. 21, (ed. 2) P
Ss.

Gro
North America 2:
25. Welch, fe of ae fig. 129.—Fig. 493

soil and humus, particularly on wooded banks, also on logs, stumps, bases of trees, or
n nee woods. Circumpolar; Labrador to Alaska, south to Oregon and Arizona, throughout

rock, i
eastern North America

CHEBOYGAN CO.—Reeses Bog, Pine Point, near Smiths Bog, Ingleside, Colonial Point,
Livingston Bog, Mud Lake, Duncan Bay. EMMET CO.-—Pellston Hills, Levering, Carp Lake, Wycamp

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Mnium cuspidatum. 493. Habit. 494. Leaf. 495. Upper cells at margin of leaf.
nium drummondii. 496. Habit. 497. Leaf. 498. Upper cells at margin of leaf.

ie

Lake, Middle paves! Five Mile Creek. MACKINAC CO.—Bois Blanc Island, Mackinac Island,
McKays Creek, Prentiss Bay, Gros Cap, Cut River, Ozark. PRESQUE ISLE CO.—Hammond Bay,
Mast Point, Ocqueoc River Falls.

Mnium cuspidatum is one of the most common mosses in eastern North America,
an. of moist hardwood forests but also found in many other habitats, in

r lawns and in drainage channels on roadside maple trees or in their crotches, for
ae. It is often called the WOODSY MNIUM (translating a synonym formerly in
usage, M. sylvaticum). It is recognized by obovate leaves, crisped when dry, and
toothed only above the middle. In a sterile condition it consists of spreading stolons
with leaves loosely arranged in two rows. The relatively small, rounded-hexagonal leaf
cells are rather thick-walled and more or less thickened at the comers.

7. Mnium drummondii Bruch & Schimp.—Plants in loose, light-green or yellowish
tufts 2-3 or sometimes 5 cm. high, radiculose below. Stems green to brownish, the
sterile ones spreading, somewhat stoloniform-attenuate and + complanate-foliate.
Leaves loosely erect or erect-spreading, not much contorted when dry, 4-6 mm. long,
broadly obovate, cuspidate-acuminate, decurrent, bordered by 2-3 rows of linear cells
in | layer, with single, sharp, mostly unicellular teeth in the upper half; costa greenish,
subpercurrent to shortly excurrent; upper cells 25-50 u, hexagonal with straight, thin
or rather firm walls, not at all thickened at the corners. Synoicous (rarely also
producing a large terminal perigonium with differentiated bracts). Setae 1-3 from the
same perichaetium, 12-28 mm. long, yellow or orange-yellow; capsules 2-3 mm. long,
pale-yellow, pendulous to pendent; operculum convex-conic; stomata superficial, in the
neck; exostome yellow or greenish-yellow; endostome brown, the cilia in groups of 2-3.
Spores 18-24 yw, finely papillose.—n = 6

Grout, Moss Flora of North America 2: Pl. 100 right, 101C.—Fig. 496-498.

On soil, humus, bark at base of trees, and rocks in woods. arate British Columbia to
Nova Scotia and south to Maryland, Michigan, Minnesota, and Washingt

CHEBOYGAN CO.—Reeses Bog, Iron Bridge (Carp Creek). EMMET CO.-—Levering.
MACKINAC CO.—Mackinac Island.

A rare species rather similar to the M. cuspidatum, recognized by clustered
sporophytes, leaves scarcely contorted when dry, and cells larger and more regularly
hexagonal, with thin walls not thickened at the corners.

. Mnium affine Bland. ex Funck, sensu l/ato—Plants in green or yellow-green,
loose or dense tufts about 3-9 cm. high, usually brown-tomentose below. Sterile stems
spreading to deflexed, often elongate. Leaves erect or spreading and strongly contorted
when dry, spreading when moist, 5-8 mm. long, elliptic to oblong-obovate, broadly
acute to rounded-obtuse, abruptly cuspidate-pointed, decurrent, bordered by 24 rows
of linear cells in 1 layer, toothed nearly to the base by teeth of 1-3 cells (or the teeth
often short and blunt and sometimes nearly lacking); costa excurrent as a cuspidate
point; upper median cells nearly hexagonal or more often oblong-hexagonal in oblique
rows, 22-35 X 33-65 u, thin- or moderately thick-walled, inconspicuously pitted and
somewhat thickened at the corners. Dioicous. Setae single (or rarelysin 2’s or 3’s),
18-50 mm. long, yellowish above, reddish below; capsules pendulous, oblong-cylindric,
3-5 mm. long, brown or brownish-yellow; operculum apiculate; stomata somewhat
immersed, in the short and inconspicuous neck; exostome teeth yellowish; endostome
brown, the basal membrane not or slightly perforate, segments broadly fenestrate,

173

appendiculate at the apex, cilia in groups of 2-4. Spores 26-29 4, minutely papillose.—n
= 6 (also as var. ciliare, n = 6, and as var. rugicum, n =

en, Mosses of Florida, ie 67 oe 5 a Grout, Mosses with Hand-lens and Microscope,
Pl. 49 a fig. 117a-c (as var. re), g, M Flora of North America 2: Pl. 101A. Jennings,
Mosses of Western Tete a 1) Pl. 2 oe (ed. 2) Pl. 26 (as var. ciliare and var. rugicum).
Welch, Mosses of Indiana, fig. 131.—Fig. 499-501 (var. ciliare).

On wet soil, humus, and rotten wood, in swampy woods, frequently among willows and other
shrubs alon; small streams. Circumpolar; South America; Greenland to Alaska and south to
California, Wyoming, Texas, and Florida.

CHEBOYGAN ne —Reeses Bog, Hermits Bog, Gorge of Carp Creek, Iron Bridge, Livingston
Bog, Colonial Point, Mud Lake, Grass Bay, Little Lake 16. EMMET CO.-—Pellston Hills, Pleasant-
view Swamp, ar Village, Five Mile Creek, Conway Bog. MACKINAC CO.—Mackinac Island, Cut
River, Hendricks, McNamara Pond. PRESQUE ISLE CO.—Lamprey Experiment Station, Evergreen
Beach, Presque Isle lighthouse, Clinton Lake.

Mnium affine is an Old World species represented in North America by the var. ciliare
C. M. (leaves broadly decurrent; marginal teeth distinct and often very long, consisting of
1-4 cells; setae single) and var. rugicum (Laur.) BSG (leaves not decurrent; marginal teeth
short, of 1-2 cells but usually reduced and often nearly lacking; setae clustered). The
var. rugicum is usually sterile and poorly developed and even at best difficult to
separate from depauperate expressions of var. ciliare. The differences between the var.
ciliare and the var. affine are small indeed. In the var. affine, the setae are clustered

and the leaves narrowly decurrent. Such differences assume some importance, however,
because of geographic separation: The var. affine occurs in Europe and the Caucasus;
the var. ciliare is limited to North America; and the var. rugicum is circumpolar.

We have both var. ciliare and var. rugicum in the Douglas Lake region. They are
somewhat similar to M. cuspidatum, which is synoicous and has marginal teeth only in
the upper half of leaves. Mnium medium is even closer in appearance and relationship,
differing mainly in the synoicous inflorescence and larger size. The leaves are broadly
decurrent and cee toothed all around, but the teeth are much shorter than those
of M. affine var. ciliar

The var es like the var. affine, grows in wet, eutrophic habitats, especially
in thickets along streams, the var. ciliare in mesic situations in deciduous or mixed
woods.

9. Mnium medium BSG—Plants in loose, light- or yellow-green tufts about
2.5-7 cm. high, tomentose. Stems greenish; sterile stems few, spreading or deflexed, not
much elongate. Leaves erect-spreading or spreading when moist, wide-spreading and
strongly crisped and contorted when dry, 5-10 mm. long, elliptic to oblong-obovate,
rounded-obtuse and abruptly short-cuspidate, decurrent, bordered by 24 rows of linear
cells in 1 layer, with single, rather sharp, unicellular teeth nearly to the base; costa
shortly excurrent; upper cells 28-55 (and sometimes up to 77) 4, irregularly rounded-
and oblong-hexagonal, with firm or rather thick walls, thickened at the corners, pitted.
Synoicous and paroicous. Setae 1-3 (sometimes 7) from the same perichaetum,
20-45 mm. long, pale-yellow, becoming red or orange-red with age; capsules 3-4 mm.
long, brown, pendulous or pendent; operculum convex-conic, + apiculate; stomata
immersed, in the neck; peristome teeth yellow or brownish; endostome brown, the cilia
in groups of 2-4. Spores 24-29 y, finely papillose—n = 12.

Grout, Moss Flora of North America 2: Pl. 101 above, 101B. Jennings, Mosses of Western
an ae (ed. 1) Pl. 21, (ed. 2) Pl. 25. Welch, Mosses of Indiana, fig. 130.—Fig. 502-505.

On , humus, rock, and bases of trees in wet woods. Circumpolar; Greenland to Alaska,
south to enn Arizona, Colorado, Arkansas, and Maryland.

174

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Mnium oS var. eaee 499. me ou Upper cells of leaf. 501. bap alae om
Mnium medium, 502. Habit. 503. 504. Upper cells of leaf. 505. Capsu
Mnium eclecae 506. Leaves. rir Leaf cells. 508. Capsule.

CHEBOYGAN CO.—Reeses Bog, Iron 7 (Carp Creek), Colonial Point, Ellis Township,
Douglas Lake, Mud Lake. EMMET CO.—Wycamp Lake, Cross Village. MACKINAC CO.—Mackinac
Island, McKays Creek, Cut River, Epoufette

Mnium medium has a somewhat distinctive appearance based largely on size and
defying description, although the clustered sporophytes generally give a clue to its
identity. The species is related to M. affine, differing essentially in size, sex, and
chromosome number (being a diploid).

10. Mnium longirostrum Brid.—Plants in light or dark-green tufts about 2-4 cm.
high, brown-radiculose below. Stems greenish; sterile stems numerous, deflexed to
horizontal, up to 5 or 6 cm. long, with rather distant, + 2-ranked leaves. Leaves larger
and + crowded at the stem tips, spreading, recurved and strongly contorted when dry,
4-7 mm. long, elliptic, rounded-obtuse or emarginate and abruptly mucronate to

~

175

cuspidate, slightly decurrent, strongly bordered by 3-5 rows of linear cells in | layer,
serrulate all around by small and inconspicuous, blunt teeth of 1 cell; costa
subpercurrent to shortly excurrent; upper cells 22-24 y, irregularly oblong -hexagonal, in
oblique rows, firm- or fairly thick-walled, pitted, thickened at the comers. Synoicous.
Setae 1-4 per perichaetium (often 5-7 in tropical America), yellow, becoming orange or
red, 15-35 mm. long; capsules 2.5-3mm. long, oblong, pendulous, yellow-brown;
operculum obliquely long-rostrate; stomata immersed, scattered over much of the
capsule; exostome teeth yellowish; endostome golden-brown or brown, with perforate
segments and cilia in groups of 2-4. Spores 22-33 y, very finely roughened. Calyptrae
pale-brownish, smooth.—n = 7, 7(6+x/y), 12, 14, 21.

, Mosses of Florida, Pl. 66 (fig. 1-3). Grout, Mosses with Hand-lens and Microscope, Pl,
50 ee eee —Fig. 506 -508.

On damp, shaded rock and soil. Nearly cosmopolitan; widely scattered but uncommon
North America, from Hers Columbia and Alberta to Oregon and Colorado, also southeastern
Canada to the Gulf of Mex

MACKINAC CO.—Mackinac Island.

Mnium longirostrum can be recognized by its elliptic leaves which are more or
less rounded to truncate or emarginate at the apex and abruptly short-apiculate. Other
useful taxonomic features include the collenchymatous leaf cells, rostrate opercula, and
stomata distributed over most of the capsule. (The single record from our area may be
a result of mislabeling. I would be more inclined to accept a record from the montane
habitats of the Lake Superior region.)

11. Mnium cinclidioides Htib.—Plants in loose, green, or yellowish tufts about
5-7, sometimes 15 cm. high, somewhat shiny when dry, moderately radiculose below.
Leaves rather distant, erect or erect-spreading, somewhat contorted when dry, usually
about 6-7 mm. long (rarely only 3mm.), elliptic, rounded or rounded-obtuse and
shortly and bluntly apiculate, not decurrent; margins slightly and bluntly serrulate all
around; costa ending somewhat below the apex; upper cells large, elongate in oblique
rows, about 3-4:1, with walls thin or moderately thickened, pitted, not thickened at

Dioicous. Setae single (or rarely paired), slender and flexuose, orange-yellow,
20-55 mm. long; capsules 2-3 mm. long, yellow, becoming brown with age, dark and
brownish at the mouth, pendent; operculum convex-conic, apiculate; stomata com-
pletely immersed, in the neck; peristome teeth dark-brown; endostome yellow-brown,
with + coalesced cilia in groups of 2-4. Spores 31-40 y, finely papillose.—n = 6, 7.

Grout, Mosses with Hand-lens and Microscope, fig. 120. Jennings, Mosses of Western
Pennsylvania (ed. 1) Pl. 23, (ed. 2) Pl. 27.—Fig. 509-510.

On wet soil or humus, often in swampy depressions or dried-up pools in mixed woods or
ree swamps. Circumpolar; Greenland to Alaska and south to New Jersey, Michigan, and
Monta

CHEBOYGAN CO.—Burt Lake (at mouth of Carp Creek), Iron Bridge (Carp Creek). EMMET
CO —Larks Lake.

This rare species of wet habitats is somewhat similar to M. punctatum but
distinguished by the blackish color of the interior of the tufts, large, broad leaves
without a distinct border, and leaf cells elongate in diagonal rows.

176

12. Mnium punctatum Hedw.—Plants in loose, pale- to dark-green, often red-
tinged tufts, usually less than 5 but sometimes as much as 12cm. high, often
tomentose. Leaves erect or spreading and moderately contorted when dry, spreading
when moist, 3.5-9 mm. long (but usually about 6 or 7 mm. long), elliptic or obovate
from a narrow base, broadly rounded to emarginate and usually ending in a short,
blunt mucro, entire, strongly bordered by linear, often reddish cells usually in several
layers; costa usually confluent with the border to form the mucro but often ending
somewhat below the apex; upper cells irregularly hexagonal to oblong-hexagonal,
sometimes elongate in oblique rows, 1-3:1, 50-100 (rarely 150) X 35-50 u, thin-walled
to fairly thick-walled, pitted. Dioicous. Setae single (or rarely paired), becoming
reddish, with a very short neck, yellowish or brownish, horizontal to pendulous;
annulus persistent; operculum stoutly rostrate; stomata completely immersed, in the
neck; peristome teeth yellowish or brownish-yellow; endostome brown, with cilia
cohering, in 2’s or 3’s. Spores 29-41 yu, finely papillose.—n = 6+1, 6+x/y, 7; 2n = 14.

rout, Mosses with Hand-lens and Microscope, fig. 118a-e (including var. elatum). Jennings,
osses of ‘We stern Pennsylvania, (ed. 1) Pl. 23 & (ed. 2) Pl. 27 (including var. elatum). Welch,
Sasi of Indiana, fig. 132.—Fig. 511-513.

On wet soil, humus, rotten logs, and rocks in swampy woods, especially along brooks and
near springs. Circumpolar; Greenland to Alaska, south to Georgia, Arkansas, Colorado, and
California

CHEBOYGAN CO.—Reeses Bog, Hermits Bog, Iron Bridge, Gorge of ee Creek, Livingston
Bog, Grass Bay. EMMET CO.-—Levering, Maple River, Pleasantview Swamp, Wycamp Lake, Conway
Bog. MACKINAC CO.—Bois Blanc Island, Cut River, Epoufette. PRESQUE ISLE CO. ~Hammo nd
Bay, Evergreen Beach, Kelsey Pond, Hendricks.

A common species of wet habitats (especially Thuja swamps), characterized by
broad, emarginate leaves ending in a short, stout apiculus formed by the confluence of
costa and heavy, cartilaginous border. (It is likely to be confused with M. pseudo-
punctatum, which is synoicous and has a shorter costa.)

The male inflorescence has a daisy-like beauty. The specific epithet means dotted
and refers to a dark cluster of antheridia surrounded by petal-like leaves. Such a
discoid perigonium forms a splash cup conferring possible advantage in the dispersal of
sperms.

The var. elatum Schimp. is a luxuriant form of particularly wet habitats. In its
best expression it is large and handsome with large leaves, without an apiculus and
with the costa ending below the apex. It is locally common.—n = 7.

13. Mnium pseudopunctatum Bruch & Schimp.—Dark-green plants in rather
loose, tomentose tufts about 4-6cm. high. Leaves erect or erect-spreading and
moderately contorted when dry, spreading when moist, 4.5-6 mm. long, nearly rounded
to broadly obovate, rounded or slightly retuse at the apex; margins entire, bordered by
1-3 rows of cells, the border unistratose above, somewhat thickened but scarcely
cartilaginous below; costa usually ending well below the apex (by about 2-14 cells),
often indistinctly forked at the tip; upper cells oblong-hexagonal, elongate in oblique
rows, about 45-90 X 27-38 u, mostly about 1.5-2:1, with relatively thick, pitted walls.
Synoicous, Setae single, 13-45 mm. long, orange-brown or reddish; capsules 2-2.3 mm.
long, pendent, narrowly ellipsoidal, pale-brown, darker at the short neck and at the
mouth; annulus of 2 rows of cells; operculum stoutly and obliquely rostrate; stomata
immersed, in the neck; peristome teeth dark-brown, densely papillose below, vertically

177

Mnium cinclidioides. 509. Leaves. 510. Upper cells of leaf.
Mnium punctatum. 511. Habit. 512. Leaf. 513. Cells at apex of leaf.
Mnium pseudopunctatum., 514. Leaf. 515. Upper cells of leaf.

striate and papillose above; endostome brownish, with cilia in 3’s. Spores 28-48 y,
densely and finely papillose.—n = 13, 14.
Fig. 514-515.

On soil and humus in wet places, particularly in fens and wooded swamps, such as Thuja
bogs, probably a calciphile. Circumpolar; Greenland to Alaska, south to Colorado, the Great Lakes,
and Maine.

CHEBOYGAN CO.—Reeses Bog, Mud Lake. EMMET CO.—Maple River, Arnott Lake,
Wycamp Lake, Cecil Bay, Conway Bog. MACKINAC CO.—Macckinac Island. PRESQUE ISLE
CO.—Lamprey Experiment Station, Evergreen Beach, Clinton Lake.

178

Distinguished from the much more common M. punctatum by the synoicous
inflorescence and costa usually ending well below the leaf apex. M. andrewsianum
Steere, a rare and local species distributed across Siberia and northern North America,
has been found in the Lake Superior region of Ontario (Red Rock) and Michigan
(Alger Co.). It grows in boggy habitats and is similar to M. punctatum and M.
pseudopunctatum. It is dioicous with leaves only 1-2 mm. long, not mucronate, with
an unistratose border and with the costa ending far below the apex.

2. Cinclidium Sw.

Fairly robust plants in bright-green to red-brown tufts, shiny and translucent (or
dewey-looking) when growing, often brown or blackish and matted with brown radicles
below. Stems simple or forked. Leaves erect, spreading, or recurved, oval or broadly
obovate, acute to obtuse or rounded and + apiculate, entire, strongly bordered by
linear cells; costa percurrent or nearly so or sometimes ending in the apiculus; upper
cells rounded-hexagonal or somewhat elongate in a transverse or oblique direction,
smooth, firm- or rather thick-walled, often pitted; lower cells rectangular. Setae single,
elongate, erect; capsules pendent, oblong-cylindric or broadly pyriform, with a short
neck; annulus deciduous in fragments; operculum hemispheric, sometimes mammillate;
stomata immersed, restricted to the neck; peristome deeply inserted, the teeth short
and blunt or emarginate, often irregular at the margins, pale- or greenish-yellow, finely
papillose; endostome much longer, dark- or yellow-brown, finely papillose, originally a
continuous, covered cylinder but becoming ruptured at maturity because of adherence
to the teeth, resulting in a pleated dome supported by irregular, keeled columns
alternating with the teeth. Spores spherical. Calyptra cucullate, smooth, naked.—The
name refers to spaces of a lattice, in this case to spaces of the endostome which is like
a cupola supported by pillars alternating with the teeth of the exostome.

Cinclidium stygium Sw.—Plants in loose tufts 3-8, rarely 13 cm. high, red-brown
or green with red tinges above, dark-red or blackish below (light-green and dewey-wet
in appearance when growing). Stems reddish, becoming blackish with age. Leaves
erect-spreading, contorted when dry, 4-6 mm. long, broadly obovate, rounded-obtuse
and abruptly narrowed to a long, red apiculus; margins often + revolute below; costa
reddish, shortly excurrent; cells obliquely or transversely elongate-hexagonal, with
rather thick, pitted walls, not thickened at the corners, 3-4 marginal rows narrow and
thick-walled forming a strong, red or red-brown border. Synoicous. Setae 40-65 mm.
long, slender, flexuose, rarely in pairs; capsules 2-3 mm. long, shortly pyriform; annulus

of 1 row of cells; operculum sometimes mammillate; peristome teeth pale-yellowish;
endostome dark-brown. Spores extremely variable even in the same capsules, 31-68 y,
very ea roughened.—n = 14.
ington, Mosses of Michigan, fig. 78. Grout, Mosses with Hand-lens and Microscope, Pl. 5
(fig. 14), rae Flora of North America 2: Pl. 106. _Fig. §16-519.

en, calcareous fens, near trickling springs or vee — near pools. Circumpolar;
Gon oe Alaska and south to Quebec, New York, and Mich

CHEBOYGAN CO.—Grass Bay. EMMET CO.—Cecil Bay. MACKINAC CO.-—Bois Blanc Island,
Mud Lake. PRESQUE ISLE CO.—Hammond Bay, Evergreen Beach, Clinton Lake

Rare in our area. Similar to Mnium punctatum in many ways. The younger
leaves have a lovely dewey-green freshness that is lost on drying. The older parts are
red or purplish-black (a gloomy, stygian color). The strong, red border and the strong
apiculus, red even in young leaves, are good characters for recognition.

179

\
y

pe
ote AY

Cinclidium stygium. 516. Habit. 517. Leaves. 518. Upper cells at margin of leaf. 519.
Portion of peristome

immia megapolitana. 520. Habit. 521. Leaves. 522. Upper cells at leaf margin. 523.
Cross-section of costa.

TIMMIACEAE -

Rather coarse plants in dull, dark-green or yellowish tufts, brown below, rigid
when dry. Stems erect, simple or forked. Leaves in many rows, sometimes somewhat
larger and + tufted at the stem tips, spreading to wide-spreading from an erect,
half-sheathing base, incurved-contorted when dry, keeled or subtubulose, unbordered, +
serrate; costa single, ending at or near the apex, sometimes toothed at back above; cells
of the limb small and green, rounded-quadrate or rounded-hexagonal, mammillose on
the upper surface, those of the sheathing base long-rectangular to linear, narrower
toward the margins, pale and hyaline, orange, or red, sometimes papillose at back.
Perichaetial leaves not much differentiated; paraphyses of perigonia filiform. Setae
terminal, elongate, erect; capsules inclined to pendulous, oblong-ovoid or oblong-
cylindric from a short neck, brown, firm, often furrowed when dry and empty;
annulus broad, sometimes revoluble; operculum hemispheric, usually + apiculate;

180

stomata in the neck or sometimes also in the lower half of the urn; peristome double,
inserted well in from the mouth and attached to it by a membrane, the 16 teeth
abruptly incurved when dry, lanceolate, rarely perforate along the middle or split at
the apex, yellowish and papillose or transversely punctate-striate below, whitish,
papillose, and vertically barred above; endostome as long as the teeth, consisting of a
high, keeled, cross-striolate basal membrane and 64 papillose cilia mostly + grouped or
attached in 2’s or 4’s (especially at the tips), sometimes appendiculate on the inner
side. Spores spherical. Calyptrae long and narrow, cucullate, smooth and naked, often
adhering to the tip of the seta at maturity.—Consisting of a single genus, Timmia
Hedw., named for Joachim Christian Timm of Malchin in the former German state of
Mecklenburg, author of the Florae Megapolitanae Prodromus. The endostome consist-
ing of 64 cilia is an unusual and attractive feature of the genus. Mount Timmia on
Ellesmere Island was recently named for this genus, at the suggestion of a Canadian
bryologist, Guy Brassard.

Timmia megapolitana Hedw.—Plants in loose, dirty-green tufts 1.5-6, rarely 8 cm.
high, + radiculose below. Leaves 5-8, rarely 10 mm. long, subtubulose and curled when
dry, wide-spreading from an erect base and concave when moist, gradually oblong-
lanceolate from an oblong, sheathing base, broadly and bluntly acute; margins coarsely
serrate above, serrulate to the shoulders or somewhat below, incurved when dry, plane
or erect when moist; costa subpercurrent, mammillose on the upper surface (and
sometimes somewhat mammillose-roughened near the apex at back); upper cells
8-13(16) u wide, rounded-hexagonal; basal cells oblong-linear, yellow-brown or brown,
those of the upper median region of the sheath often bearing several papillae in a row
over the lumina at back. Autoicous (with 1-several perigonial buds at the base of the
perichaetium). Setae single or occasionally paired, 10-30mm. long, becoming red;
capsules 2.5-3 mm. long, broadly oblong-cylindric, curved and asymmetric, not or only
slightly furrowed when dry; annulus persistent; operculum hemispheric, not apiculate;
stomata in the lower half of the urn or nearly throughout; peristome teeth yellow-
brown and finely papillose below, pale-yellow or whitish and coarsely papillose and
vertically barred above; endostome with 64 evenly spaced, yellowish or whitish,
papillose filaments bearing fine, inwardly directed appendages at the nodes. Spores
11-16 u, very a4 papillose. Calyptra clasping the tip of the seta after falling from
the urn.—n = 12,

Conard, How to Know the Mosses, (ed. 2) fig. 55. Grout, Mosses with Hand-lens and
Microscope, PL 43. Jennings, Mosses of Western Deets (ed, 1) Pl. 25 & (ed. 2) Pl. 29 (as T.
cucullata). Welch, Mosses of Indiana, fig. 108.—Fig. 520-523.

On moist or wet, shaded soil or humus, sometimes on rotten logs, typically along streams,
sometimes on banks of ravines. oe Alaska to Quebec, south to Nebraska, Arkansas, and
Virginia (more common in the East.)

CHEBOYGAN CO.—Reeses Bog, Douglas Lake, Mill Creek, Black Lake. EMMET CO.—Maple
River, Wycamp Lake, Cross Village. MACKINAC CO.—Mackinac Island. PRESQUE ISLE CO.-—
Ocqueoc River.

Resembling an Asrichum or a Polytrichum in aspect. The leaves rolled and
incurved when dry, with ventrally bulging cells, and the details of the peristome are
highly distinctive. The way in which the calyptra remains attached to the tip of the
seta after slipping off the mature capsule aids in recognition. The var. bavarica (Hessl.)
Brid., a western expression which has been, found in the Thunder Bay District of
Ontario, has smaller, more angular cells often somewhat longer than wide (6-7 u wide
and isodiametric or sometimes about 11 long). In montane habitats of the Upper
Peninsula, we have Timmia austriaca Hedw. It has cells of the leaf base orange and

181

smooth, filaments of the endostome somewhat nodose and in groups of four, and
dioicous inflorescences.

Lazarenko (Bryol. 60: 14-17. 1957) noted a curious behavior in the double
peristome of Timmia megapolitana var. bavarica: The teeth are joined at their tips in
pairs thus forming eight triangles, the broad apices of which cannot penetrate between
the cilia of the endostome, as in double bryoid or hypnoid peristomes. Because of the
joined tips, the teeth spread outward in their lower parts and inward at the tips. When
hygroscopic movements begin, the teeth squat down and bulge outward at the middle,
while the incurved tips skim up and down ploughing the spores and jumping over
nodes of the cilia. Because of such disturbance the cilia throw out spores rather than
merely allow their dispersal in a salt-shaker fashion.

AULACOMNIACEAE
Plants small to rather robust, in dull, green, yellow-green, or eas ee
usually tomentose tufts. Stems erect, simple or forked. Leaves in many r not o

rarely somewhat complanate, erect to erect-spreading, scarcely altered = an
when dry, sometimes + larger at the stem tips, lanceolate to oblong-ovate or elliptic,
acute to rounded at the apex, sometimes apiculate, concave or keeled, unbordered,
generally toothed above; costa single, strong, ending somewhat below the apex (or
sometimes considerably below it), with 2 stereid bands; upper cells small, rounded,
rounded-hexagonal, or elliptic, thick-walled, smooth or unipapillose on both surfaces;
lower cells mostly somewhat longer or rectangular, sometimes swollen at the insertion.
Setae terminal, + elongate, erect; capsules usually inclined to horizontal, symmetric or
+ curved and asymmetric, oblong-ovoid to cylindric from a short neck, plicate when
dry; annulus broad; operculum conic to rostrate, sometimes obliquely so; stomata
restricted to the neck; peristome double, the 16 teeth lanceolate, papillose; endostome
hyaline, with a low or high basal membrane, narrow segments sometimes slit or gaping
along the median line, and delicate to stout cilia. Calyptrae cucullate.

Aulacomnium Schwaegr.

Plants small to robust, in dull, yellow-green, yellow-brown, or green, radiculose
tufts. Stems erect, simple, or forked. Leaves crowded, erect or erect-spreading, scarcely
altered on drying to strongly contorted, oblong-lanceolate to ovate or elliptic, acute to
obtuse or rounded, sometimes apiculate, not or somewhat decurrent, concave or
keeled; margins entire or toothed above, usually + revolute; costa tapered, usually
flexuose above, typically ending somewhat below the apex; cells rounded or shortly
oblong to elliptic, + incrassate and sometimes collenchymatous or stellate, smooth or
unipapillose on both surfaces, short and scarcely differentiated to somewhat elongate
below, sometimes colored and somewhat swollen at the insertion. Setae elongate,
smooth; capsules suberect to horizontal, subcylindric, asymmetric and usually some-
what curved, contracted below the mouth when dry; annulus revoluble; operculum
bluntly conic to stoutly and obliquely short-rostrate; peristome teeth pale, yellow-
brown; endostome hyaline, with a high basal membrane, keeled and usually gaping
segments, and long, slender, nodulose cilia in groups of 24 (or rarely 5). Spores
spherical. Calyptrae smooth, naked._The name means furrowed Mnium, referring to a
moss with ribbed capsules.

182

1. Plants autoicous; leaves concave, broadly oblong or oblong-ovate, coarsely serrate in the
upper half, erect and + homomallous, scarcely contorted when dry; cells smoo
1. A. heterostichum
1. Plants dioicous; leaves + keeled, narrower, oblong-lanceolate, serrulate near ie apex,
loosely erect and curved or contorted when dry, not homomallous; cells unipapillose on

2. Fairly robust plants; leaves contorted when dry; upper cells collenchymatous and +
stellate; cells at leaf insertion + swollen and usually brownish in sees Ss sterile

extension of the stem and also, sparsely, along the extension 3. A. palusine
2. Rather small plants; leaves + curved but usually not particularly contorted when dry
upper cells irregularly rounded or rounded-hexagonal; cells at insertion not or an

slightly differentiated; sterile stems usually bearing a dense cluster of very small, fusiform
brood bodies at the end of a seta-like extension of the stem (and not along its length)
2. A. androgynum

Aulacomnium heterostichum (Hedw.) BSG—Plants in rather loose, green or
sometimes yellow-brown tufts 1.5-4 cm. high. Leaves rather soft, concave and slightly
undulate, erect and somewhat homomallous, not altered on drying, 2-3.5 mm. long,
oblong or oblong-ovate, not decurrent, subacute or obtuse and apiculate; margins plane
or narrowly recurved below the middle, coarsely serrate in the upper 4 or more; costa
not flexuose, ending somewhat below the apex; upper cells small, green, smooth,
thick-walled, irregularly rounded-quadrate, rounded-hexagonal, and transversely short-
oblong, 9-11, often 15 wide; basal cells similar, those at the insertion somewhat
longer, not colored or swollen. Autoicous; perigonia small, axillary. Setae 8-17 mm.
long; capsules 2.5-3.5 mm. long, suberect or inclined, somewhat curved; operculum
sr ae ie rostrate; endostome segments not perforate. Spores 9-15 u, very finely
papillose.—n = 12.

een, Mosses of Florida, Pl. 57 (fig. 5- as Conard, How to eae the Mosses, (ed. 2) fig.
3, sie Mosses with Hand -lens and Microscope, fig. 100. Jennings, Mosses of Western
Pennsylvania, (ed. 1) Pl. 24 & (ed. 2) Pl. 28. Welch, Mosses iv Indiana, fie. 109.—Fig. 524-526.

On soil or the bases of trees and stumps in rather dry to moist woods. Widespread in eastern
North America

CHEBOYGAN CO.—Douglas Lake.

An attractive moss common farther south, in oak-hickory woods, where it is often
associated with Bartramia pomiformis on shaded ravine banks. The broad, coarsely
toothed leaves and smooth cells are diagnostic.

The development of the peristome is described by Blomquist and Robertson
(Bull. Torrey Bot. Club 68: 569-584. 1941). Other references concerning peristome
development include Cavers (New Phytol. Reprint 4, pp. 177-189. 1910-11), Evans &
Hooker (Bull. Torrey Bot. Club 40: 97-109. 1913, on Ceratodon purpureus), Kreulen (Lind-
bergia 1: 153-160. 1973), Proskauer (Amer. Jour. Bot. 45: 560-563. 1958, on Funaria
hygrometrica), Saito (Bot. Mag. Tokyo 69: 53-58. 1956, on Dicranum japonicum),
Saito & Shimozu (Bot. Mag. Tokyo 68: 55-60. 1955, on Bartramia crispata), Stone
(Austral. Jour. Bot. 9: 124-151. 1961, on Mittenia plumula), van der Wijk (Rec. Trav.
Bot. Neerl. 26: 289-395. 1929; pp. 1-40 in Verdoorn’s Manual of Bryology. 1932).

For detailed structure and evolutionary significance of peristomes: Philibert (Rev.
Bryol. 11: 49-52, 65-72, 81-87. 1884; 12: 67-77, 81-85. 1885; 13: 17-27, 81-86. 1886;
14: 9-11, 81-90. 1887; 15: 6-12, 24-28, 37-44, 50-60, 65-69, 90-93. 1888; 16: 1-9,
38-44, 67-77. 1889; 17: 8-12, 25-29, 38-42. 1890; B: 36-38, 41-56. 1896; 28: 56-59,
127-130. 1901; 29: 10-13. 1902) or, for an abridged translation, Taylor [Bryol. 65(3):
175-212. (1962)1963]; van der Wijk (Buxbaumia 5: 10-17. 1951). Grout’s notes on

.

t tye °
; soe :
“Ne ife yoy
Ie 5
}

Af
<ceoe =
PSEIS

CESSES.

ws

525

Aulacomnium heterostichum. 524. Habit. 525. Leaf. 526. Upper cells of leaf.
Aulacomnium androgynum. 527. Habit of sterile plant. 528. Leaf. 529. Cells at insertion of

leaf.
Aulacomnium palustre. 530. Habit. 531. Leaf. 532. Upper cells of leaf. 533. Basal cells of

leaf.

peristome structures (Bryol. 4: 31-32. 1901; 5: 53-56, 73-74, 94-95. 1902; 6: 63-65.
1903; 7: 37-39. 1904) are commendable because of illustrations. Details of peristome
structures as revealed by the scanning electron microscope are beautifully portrayed by
Robinson (Bryol. 74: 473-473. 1972).

Aulacomnium heterostichum is quite unlike other members of the genus in
aspect, and efforts have been made to set it apart as the genus Arrhenopterum, on the
basis of broad, toothed, secund leaves and an apparent lack of “‘pseudopodia” and
propagula. Small deciduous leaves were found, however, in a colony of plants in
southern Michigan by Lowry and Steere (Bryol. 49: 30-32. Fig. 1-6. 1946). Though
probably induced by some unusual feature of the environment, such brood leaves
provide some evidence of relationship to A. palustre and A. androgynum.

184

2. Aulacomnium androgynum (Hedw.) Schwaegr.—Small plants in rather dense,
dark-green or sometimes yellow-brown tufts 1-2.5 (rarely 4.5) cm. high; sterile plants
commonly bearing very small, + spindle-shaped propagula in dense, globose clusters of
the end of a smooth, seta-like extension of the stem. Lower leaves small and distant,
upper larger and + crowded, loosely erect and somewhat curved but not much
contorted when dry, 1-3.5 mm. long, narrowly to broadly oblong-lanceolate, acute,
decurrent, somewhat keeled; margins narrowly revolute below and often nearly to the
apex (or in leaves of innovations sometimes plane), distinctly serrulate near the apex or
in the upper % and often crenate-serrulate nearly to the base; costa flexuose above,
ending somewhat below the apex; cells irregularly rounded or rounded-hexagonal,
moderately thick-walled, 5-11 u wide (mostly 7-9 yw), bluntly unipapillose on both
surfaces; basal cells similar, those at the insertion sometimes + elongate, not brownish
or swollen. Dioicous; male plants repeatedly forked by innovations, the perigonia large,
terminal. Setae 11-25 mm. long; capsules 2-2.3 mm. long, suberect to strongly inclined,
not curved; operculum bluntly conic; segments of endostome gaping. Spores 9-11 y,
smooth.—n = 10, 11-12, 12.

ard, How to Know the Mosses, (ed. 2) fig. 85e-. Grout, Moss Flora of North America 2:
Pl. see Fig. 527-529.

A woodland species, on soil or humus of banks, sometimes on rotten wood. Europe, Canar
aoe Sapa Alaska to Newfoundland, south to California, Wyoming, the Great Lakes, and West

CHEBOYGAN CO.-—Gorge of Carp Creek, Hee Bridge, Burt Lake. EMMET CO.—Stutsman-
ville, Cross Village. MACKINAC CO.—McNamara Pond.

The specific epithet, indicating a hermaphroditic or monoicous sexual situation,
was inappropriately applied by Linnaeus. The species is much less common than A
palustre, with which it is likely to be confused.

3. Aulacomnium palustre (Hedw.) Schwaegr.—Plants fairly robust, in loose or
dense, yellow, yellow-green or yellow-brown, often conspicuously tomentose tufts
3-9 cm. high (rarely less); sterile stems often bearing clusters of + triangular propagula
at the end of a warty, seta-like extension of the stem and also, sparsely, along the
extension. Leaves erect-spreading, twisted and contorted when dry, 24mm. long,
keeled, oblong-lanceolate, acute or acuminate, somewhat decurrent; margins serrulate
or serrulate-papillose toward the apex, revolute in the lower %4; costa flexuose above,
ending somewhat below the apex; upper cells 9-13 u wide, isodiametric, thick-walled,
collenchymatous and + stellate, unipapillose on both surfaces; basal cells similar except
for several rows of pale to brownish, somewhat swollen cells with moderately
thickened, straight walls at the insertion. Dioicous; perigonia large, terminal. Setae
25-45 mm. long; capsules 2.5-4 mm. long, strongly inclined to horizontal, curved;
operculum bluntly conic to stoutly and obliquely short-rostrate; segments of endo-
stome gaping. Spores 11-13 uw, smooth.—n = 9, 10,

een, Mosses of Florida, Pl. 57 (figs. 14). Conard, How to Know the Mosses, (ed. 2) fig.
ie aaa Mosses of Michigan, fig. 79. Grout, Mosses a Hand-lens and Microscope, fig.
, 102, Moss Flora of North America 2: Pl. 66A. Jennings, Mosses of Western Pennsylvania, (ed.

7 aT 24, (ed. 2) Pl. 28. Welch, Mosses of Indiana, fig. 110.—Fig. 530-533.

Very common on various substrata, particularly soil or humus, in swamps, fens, and
burned-over bogs. Nearly le iat Greenland to Alaska, south to the Gulf of Mexico, cee
Mexico, Arizona, and Califor

185

CHEBOYGAN CO.—Reeses Bog, Hermits Bog, Gorge of Carp Creek, Iron Bridge, Douglas
Lake, Gleasons Bog, Wolffs Bog, Maloneys Bog, Livingston Bog, Bryants Bog, Gates Bog, Vincent
Lake, Smiths Bog, Mud Lake, Grass Bay, Little Lake 16, Weber Lake. EMMET CO.—Pellston,
Levering, Galloway Bog, Maple River, Pleasantview Swamp, Stutsmanville, Conway Bog, Wycamp
Lake, Wilderness State Park, Cecil Bay. MACKINAC CO.—Bois Blanc Island, Rexton, St. Martins
Bay, Nunns Creek, St. Martins Point, McKays Creek, Hog Island Point, Epoufette, Ozark, Peggley
Lake, Hendricks. PRESQUE ISLE CO.—Evergreen Beach, Hammond Bay, Mast Point, Clinton Lake.

This exceedingly common moss is larger than A. androgynum and has brownish,
swollen cells near the leaf insertion. The gemmae are triangular (and thus more or less
leaf-like) and borne along a seta-like extension of the stem, in a leaf-like arrangement,
and also crowded in an apical cluster. A. androgynum has tiny, spindle-shaped gemmae
borne only at the apex of a similar extension of the stem (sometimes called a
pseudopodium).

Aulacomnium palustre may be green or brown, but it always has a sickly yellow
cast as well.

Aulacomnium acuminatum (Lindb. & H. Arn.) Par., a rare arctic and boreal
species not included in Grout’s Moss Flora, has been found in the Thunder Bay
District of Ontario. Compared with A. palustre, the plants are quite robust, lack brood
bodies, and have leaves which are erect or imbricate and not at all contorted when dry;
the margins are entire and strongly revolute nearly the length of the leaf.

Common northward is A. turgidum (Wahl.) Schwaegr., distinguished from A.
palustre especially by rounded-obtuse leaves with smooth cells. The leaves are imbricate
and scarcely altered on drying. When well developed, the stout, julaceous, essentially
simple stems look like large yellowish caterpillars.

MEESIACEAE

Plants slender to rather robust, gregarious to tufted, dull, dirty-green or yellowish
to light-green and rather shiny, radiculose or tomentose below. Stems erect, forked.
Leaves in 3-8 rows, erect or erect-spreading to squarrose-recurved from an erect base,
lance-ligulate to ovate-lanceolate or obovate, acute to rounded-obtuse, unbordered,
sometimes toothed above, rarely to the base; costa single, strong, mostly ending below
the apex; cells rounded- or oblong-hexagonal or rectangular, mostly with firm- or
thick-walls, smooth (or rarely mammillose); lower cells rectangular, often delicate and
hyaline. Paraphyses of perigonia clavate, those of the perichaetia (if present) filiform.
Setae terminal, elongate, erect-flexuose; capsules suberect or inclined, strongly curved
and asymmetric, oblong-pyriform from a rather short or fairly long neck, small- and
oblique-mouthed, smooth; annulus narrow, deciduous in fragments; operculum small,
convex or bluntly short-conic; stomata numerous, in the neck; peristome double, in
Paludella consisting of 16 well-developed teeth and segments of a length equal to them,
rising from a rather low basal membrane, otherwise the 16 teeth much shorter than the
endostome, fused at base or nearly throughout, blunt or truncate at the apex, the
endostome consisting of a very low basal membrane adhering to the teeth, long, linear,
keeled segments slit along the middle and mostly irregular at the margins, and cilia
rudimentary or lacking (the inner peristomial layer sometimes incompletely resorbed,
leaving a + fragmentary membrane between the segments with cilia sometimes
represented by irregular chains of nodose deposits on the membrane). Spores generally
large. Calyptrae cucullate, smooth.

186

1. Leaves strongly squarrose-recurved from an erect base, in 5 distinct rows; cells mammillose
both surfaces; segments about as long as the tee 3. Paludella
1. Leaves erect, spreading, or squarrose but not recurved, not in ae rows, or if so
longe

2. Soft, delicate plants with leaves oblong-lanceolate, rather ae narrowed to a short,
sharp point, shrunken when dry; leaf cells lax and thin-walled; a a leaves not
ie

differentiated Amblyodon
2. Coarser, firmer plants; leaves not shrunken when dry, ieee ras to lance-ligulate,
rounded-obtuse or gradually acute; cells firm; perichaetial leaves elongat 2. Meesia

1. Amblyodon BSG, nom. cons.
A monotypic genus, so named because of blunt peristome teeth.

Amblyodon dealbatus (Hedw.) BSG—Plants small, gregarious or tufted, pale- or
bright-green with red tinges, shiny, soft, somewhat radiculose at base, up to 1 (or
rarely 3) cm. high, considerably forked by short branches. Leaves crowded, erect and
not or slightly contorted when dry, loosely erect or erect-spreading when moist,
2-4 mm. long, keeled (especially when dry), oblong-lanceolate, gradually or abruptly
acute, margins plane, + serrulate near the apex; costa very thick and about 4% or more
the width of the leaf at base, narrower above, ending somewhat below the apex,
becoming reddish to blackish; cells smooth, pale, lax and thin-walled, oblong-hexagonal
and shortly oblong-rhomboidal, narrower in a single marginal row but not forming a
noticeable border, not much different at base. Polygamous (perichaetia and perigonia
on the same plants but archegonia occasionally found in the perigonia). Seta
12-40 mm. long, orange-brown, smooth; capsules 2-3mm. long, elongate-pyriform,
strongly curved and asymmetric from an erect, well-marked neck, yellow-brown;
operculum bluntly conic; peristome teeth inserted near the mouth, short, lanceolate,
blunt or truncate, finely papillose, yellow- to dark-brown, trabeculate at back;
endostome up to twice as long as the teeth, distinctly papillose to nearly smooth,
hyaline or yellowish, consisting of a low basal membrane, narrow, keeled, and
perforate segments, and cilia lacking or reduced to single, short, stout stubs. Spores
29-40 uw, spherical, brown or orange-brown, finely to rather coarsely papillose.—n = 14,

?

arlington, Mosses of Michigan, fig. 81. Grout, Moss Flora of North America 2: Pl.
63A. -. $34-535,

n peaty soil or aes on logs in wet or seepy places such as bogs and river banks.
Europe, Caucasus, Pers and Central Asia; Yukon a —— Columbia and Colorado, also
Manitoba to Nova Scotia a south to the Great Lakes regi

MACKINAC CO.—Mackinac Island. PRESQUE ISLE CO.—Hammond Bay.

Thy lax, thin-walled leaf cells suggest the Funariaceae, but the capsule gives
indisputable evidence of a relationship to Meesia and Paludella

2. Meesia Hedw., nom. cons.

Plants medium-sized to fairly robust, in dense, sordid-green or yellowish tufts,
brown or blackish and densely radiculose below. Stems bearing small club-hairs with
red bases in axils of uppermost leaves and perichaetial and perigonial bracts. Leaves
erect to wide-spreading or squarrose, sometimes clearly 3-ranked, not or distinctly
decurrent, oblong-lanceolate or oblong-ovate to lance-ligulate, acute to rounded-obtuse;
margins entire or serrulate, plane or revolute; costa strong, broader at the base

187

(sometimes as much as % or more the width of the leaf base), ending near the apex;
upper cells short-rectangular, smooth, with thin to firm walls; lower cells longer,
rectangular, pale. Perichaetial leaves similar, not or somewhat elongate. Setae usually
very long, yellow-brown; capsules elongate-pyriform, curved and asymmetric from an
erect, elongate, well-defined neck, inclined to horizontal, brown, shiny; annulus rather
poorly differentiated; operculum hemispheric to bluntly conic; peristome inserted
below the mouth, the teeth short, blunt or truncate, brown or yellow-brown, minutely
papillose and indistinctly cross-striolate but usually appearing nearly smooth; endo-
stome 2-4 times as long as the teeth, yellowish-brown, smooth, consisting of a low
basal membrane, narrow, keeled, and slightly perforate segments with irregular,
crenate-bulging margins, and 2-4 rudimentary or irregularly developed cilia, sometimes
+ continuous and about as long as the segments but more often represented by a chain
of fragments adhering to a delicate, incomplete, hyaline membrane resulting from
incomplete resorption of the inner peristomial layer. Spores large, irregularly spherical.
—The genus was named for David Meese, a Dutch gardener. The pear-shaped capsule
curved above a long, erect neck gives a distinctive look to the genus and the family.

Plants 1-2.5 cm. high; leaves erect, not 3-ranked, keeled, or decurrent, narrowly lance-
linear-ligulate; margins strongly recurved, entire 1... Pies

Plants 3-12cm. high; leaves spreading to squarrose from an erect base, ovate-lanceolate,
3-ranked, keeled and decurrent; margins not or sometimes indistinctly revolute, serrulate
above the shoulders 2. M. triquetra

1. Meesia uliginosa Hedw.—Plants 10-25 mm. high. Leaves not 3-ranked, erect
and often + contorted when dry, erect when moist, 2-4mm. long, narrowly lance-
ligulate to linear-ligulate, obtuse or rounded-obtuse, not keeled or decurrent; margins
entire, revolute in the lower 1/2 or nearly to the apex; costa about 1/2-2/3 the width of
the leaf base. Polygamous (usually dioicous, but sometimes autoicous or with both
synoicous and antheridial inflorescences on the same plants). Setae (10)20-70 mm.
long; capsules 1.5-4 mm. long; operculum hemispheric and flat-topped. Spores 40-51 y,
brown, densely and finely papillose.—n = 13, 13+1, 14, 14+1.

Con How to Know the Mosses, (ed. 2) fig. 159a,e-f. Grout, Moss Flora of North
America, a Pl. 64A.—Fig. 536-538.

ee or peaty humus, rarely on logs, especially in marly fens or swamp forests,
ev of cliffs; a eons: Circumpolar; Greenland to Alaska and south to New
aa Mine Cnn and Califor

CHEBOYGAN CO.—Reeses Bog, Iron Bridge, Nichols Bog, Duncan Bay, Grass Bay. EMMET
CO.—Cecil Bay. MACKINAC CO.—Cedarville. PRESQUE ISLE CO.—Evergreen Beach, Hammond
Bay, Mast Point.

The erect, ligulate, blunt leaves with the costa very broad, especially at the base,
are unmistakable

2. Meesia triquetra (L. ex Richt.) Angstr.—Plants 3-12cm. high. Leaves 3-
ranked, somewhat to strongly contorted when dry, spreading to squarrose-spreading
from an erect base, (1.5)2.5-4 mm. long, ovate-lanceolate, keeled above the shoulders,
gradually narrowed to a slenderly acute apex, distinctly decurrent; margins plane or
sometimes + reflexed or revolute near the. shoulders, distinctly serrulate from the
shoulders to the apex; costa somewhat broader at base but considerably less than 1/3
the width of the leaf. base. Dioicous; perichaetial leaves not very elongate, about
4-4.5 mm. long. Setae 30-100 mm. long; capsules 3-4.5 mm. long; operculum bluntly
conic. Spores 33-38 y, yellow-brown, very minutely roughened or nearly smooth.—n =
10

188

Satan

Ce ecsacis 5

) oy
1
—
te
__

Wy
SHY
j |

ye

Amblyodon dealbatus. 534. Leaf. 535. Cells at apex of leaf.

Meesia uliginosa, 536. Habit. 537. Leaf. 538. Upper cells of leaf.
Meesia triquetra. 539. Habit. 540. Leaf. 541. Upper cells of leaf.
Paludella squarrosa, 542. Portion of plant. 543. Leaves. 544. Capsule.

Conard, How to Know the Mosses, (ed. 2) fig. 159b-<d. Darlington, Mosses of Michigan, fig.
82 (as M. tristicha). Grout, Mosses with Hand-lens and Microscope, Pl. 42.—Fig. 539-541.

A calciphile, on wet soil or humus in marly bogs or swamp forests. Circumpolar; Alaska to
Greenland, south to California, Ohio, and Pennsylvania.

CHEBOYGAN CO.—Mud Lake, Little Lake 16. EMMET CO.—Conway Bog.

Extremely rare locally but easily recognized by the spreading leaves in three
obvious ranks. A similar species, M. longiseta Hedw., has been found in a few localities
in other glaciated portions of the eastern United States and adjacent Canada. It has
entire leaves (or rarely somewhat serrulate), elongate perichaetial leaves, synoicous
inflorescences, and brown spores about 36-44 win diameter.

189

3. Paludella Brid.

Rather robust plants in dense tufts, dull, pale-green to yellowish- or brownish-
een above, brown or rarely blackish and tomentose below; small, purplish, clavate
hairs (‘‘club-hairs’”) in leaf axils. Leaves in 5 distinct rows, strongly squarrose-recurved
from an erect base, scarcely altered on drying, obovate, broadly acuminate and acute,
narrowly long-decurrent, strongly keeled; margins narrowly recurved below and broadly
reflexed at the middle, serrulate above; costa slender, serrulate at back above, vanishing
below the apex; upper cells rounded-hexagonal, rather thick-walled, mammillose on
both surfaces; basal cells smooth, rectangular. Dioicous; perigonia rosulate; perichaetial
leaves elongate, lance-acuminate. Setae long, brown or reddish; capsules suberect or
inclined, subcylindric, tapered from a short neck, asymmetric, + curved, brown;
operculum convex or bluntly low-conic; peristome teeth lanceolate, pale-yellow,
densely and finely papillose; segments of endostome about as long as the teeth,
narrowly perforate along the keel, lightly papillose to nearly smooth, rising from a
rather low basal membrane, the cilia lacking or reduced to mere stubs. Spores
spherical, relatively small, finely papillose.—The name is a diminutive referring to a
swampy habitat.

Paludella squarrosa (Hedw.) Brid.—Tufts 4-7, rarely 15cm. high. Leaves
1.5-2 mm. long; upper cells 10-13 u wide. Setae 2.5-4.5 cm. long; capsules 2-3 mm.
long. Spores 15-21 wu, brownish.

Darlington, Mosses of Michigan, fig. 80. Grout, Moss Flora of North America 2: Pl.
63B.—Fig. 542-544,

A species of fe ae Ri not ae found in the Straits counties, although known from a
few localities in both Upper d Low t Peninsulas of Michigan (Alger, Keweenaw and Montcalm
Counties). It should be Soe espec ially at the sedgey margins of beach pools fringing the Great

Lakes. Circumpolar; Alaska to Greenland, south to New York, Michigan, Wyoming, and Montana

Certainly one of our most attractive and curious mosses, unlike anything else in
its tomentose stems and extremely squarrose-recurved leaves (and yet in structure not
too unlike Meesia triquetra).

CATOSCOPIACEAE

Slender plants in very dense, often deep tufts, dark-green to yellowish-brown
above, dark-brown or blackish and radiculose below. Stems erect, forked. Leaves in
many rows, incurved and somewhat crisped when dry, erect-spreading when moist,
narrowly lance-acuminate, slenderly acute, keeled; margins entire, narrowly recurved
below; costa single, narrow but well developed, disappearing in the apex; upper cells
small, quadrate to short-rectangular, green, smooth, firm-walled; lower cells larger,
oblong to sublinear, quadrate at the basal margins. Dioicous; perichaetial leaves larger,
lance-subulate from a half-sheathing base, plane at the margins; perigonia bud-like, with
filiform paraphyses. Setae terminal, erect, slender and elongate, dark-red; capsules
inclined or horizontal, very small, shortly ovoid from a short neck, very asymmetric
and curved, with the mouth directed downward at maturity, smooth, dark red-brown
to nearly black, becoming glossy and brittle; annulus and stomata none; operculum
convex, bluntly pointed; peristome teeth 16, lanceolate, short, irregular, and fragile,
pale, finely papillose, incurved; endostome a very low membrane, sometimes lacking.
Spores large. Calyptra cucullate, smooth, naked, sometimes adhering to the tip of the
seta at maturity.—Consisting of a single monotypic genus, Catoscopium Brid., named in

190

reference to the capsule which is curved so that the mouth is directed downward (and
hence “looks” down).

Catoscopium nigritum (Hedw.) Brid.—Plants 1.5-6 cm. high. Leaves 0.8-1.5 mm.
long. Setae 8-25 mm. long; capsules 0.6-1 mm. long. Spores 33-48 yw, spherical and
ellipsoidal, finely papillose.—n = 13, 14.

Darlington, Mosses of Michigan, fig. 83. Grout, Moss Flora of North America 2: PL.
67E.—Fig. 545-547,

A calciphile of wet eae typically on peaty humus at the margins of pools in fens,
sometimes on limey mud or limestone cliffs, at edges of beach pools or on wet sand in estes
along the Great Lakes. enact Alaska to Greenland, south to New Brunswick, Michigan
Iowa, and British Columbia.

CHEBOYGAN CO.—Duncan Bay. EMMET CO,—Wilderness State Park, Cecil Bay. MACKINAC
CO.— Bois Blanc Island. PRESQUE ISLE CO.—Mast Point, Hammond Bay, Evergreen Beach.

The leafy plants are similar to Ceratodon in aspect and structure; they grow in
very compact, wet, blackish tufts recognizable even in a sterile state and especially in
ecological context. The sporophytes, often produced in great abundance, resemble
minute golf clubs, justifying the name GOLF CLUB MOSS. The tiny capsules are a
beautiful, fresh green until fully formed but later become black and shiny. They
remind me of someone looking over his shoulder, calling to mind Satchel Paige’s
advice, in How to Keep Young: Don’t look back. Something may be gaining on you.

BARTRAMIACEAE

Small to robust plants in loose or dense tufts, generally tomentose below. Stems
erect, with forked, fastigiate, or whorled branching. Leaves in many rows, ovate-
lanceolate to linear- or lance-subulate, usually acute or acuminate, sometimes abruptly
enlarged and clasping at the base, unbordered, toothed at the margins above and often
at the back of the costa; costa single, ending below the apex to excurrent as a toothed
point or awn; upper cells rounded-quadrate to oblong or linear, mostly firm-walled,
papillose at or near | or both ends or sometimes centrally, rarely smooth; lower cells
usually larger, lax, smooth, and hyaline, not or slightly differentiated at the basal
angles. Paraphyses of perigonia filiform or clavate; perichaetia! leaves usually longer but
not much differentiated. Setae terminal (but often becoming lateral because of
innovations), mostly + elongate, straight or sometimes curved; capsules usually inclined
or horizontal (rarely erect or pendulous), subglobose or ovoid, mostly asymmetric and
oblique at the mouth, usually furrowed; annulus usually none; operculum convex or
short-conic, rarely rostrate (usually flat and umbonate when dry); stomata restricted to
the base of the capsule; peristome double, single, rudimentary, or lacking, inserted well
inside the mouth and generally attached to it by a membrane, the 16 teeth lanceolate,
yellow-brown or red-yellow, smooth or papillose, mostly unbordered, usually trabecu-
late and often with intermediate thickenings at back; endostome occasionally adhering
to the teeth as fragments or somewhat shorter, consisting of a well-developed basal
membrane, segments perforate and finally split along the keels, and cilia in groups of
1-3 (usually short, sometimes lacking). Calyptrae small, cucullate, smooth.

1. Leaf cells finely desig striolate but not papillose 1. Plagiopus
1. Leaf cells papillose a t both ends
2. Leaves linear- oe or subulate from an erect and + sheathing base; upper cells in 2-3
layers (at least at the margins); peristome teeth without intermediate thickenings at back

2. Bartramia
2. Leaves ovate or lanceolate, not erect or sheathing at a upper cells unistratose;
peristome ok usually with intermediate thickenings at bac 3. Philonotis

191
1. Plagiopus Brid.

Plants in dense, radiculose, dull, olive-green or brownish tufts. Stems erect or
ascending, triangular in section with a cortical layer of lax, thin-walled, hyaline cells.
Leaves loosely erect or erect-spreading and somewhat curved when dry, erect-spreading
when moist, narrowly lanceolate, acuminate, keeled above, not sheathing at base;
margins revolute in the lower half or more, bistratose and doubly serrate toward the
apex; costa percurrent or shortly excurrent, prominent at back and + toothed toward
the apex; cells subquadrate and short-rectangular, thick-walled, and minutely verrucu-
lose-striolate above, longer and thinner-walled below, especially toward the costa at
base. Perichaetial leaves scarcely differentiated. Setae elongate, erect or flexuose-curved;
capsules when moist suberect, subglobose, small-mouthed, and indistinctly striate, when
dry somewhat curved and inclined to horizontal, + ovoid, flattened at both ends,
strongly furrowed; operculum small, short-conic or convex; peristome inserted some-
what below the mouth, double, the teeth smooth or very minutely papillose,
lanceolate, with intermediate thickenings at back (the thickened cross-walls of the
ventral plates not coinciding with the cross-walls of the dorsal plates); endostome
consisting of a well-developed basal membrane and broad segments, the cilia none or
very short and rudimentary._The name means oblique or curved foot, making a not
very apt reference to the more or less flexuose seta.

Plagiopus Ses (Sw.) Limpr.—Plants 2-10 cm. high. Leaves 2-3.5 mm. long;
upper cells about 9y wide, 1-2:1. Synoicous. Setae 7-18 mm. long; annulus none;
operculum low-convex; peristome teeth smooth or very finely papillose, golden-brown
or red-brown; endostome pale-brown, about 2/3 the length of the teeth, smooth or
faintly papillose. Spores ellipsoidal, 20-26 u, coarsely papillose.—n = 7.

Conard, How to Know the Mosses, (ed. 2) fig. 111 (as P. oederi). Darlington, Mosses of
Michigan, fig. 84 (as P. oederi). Grout, Mosses with Hand-lens and Microscope, fig. 104b, d, g (as
Bartramia oederi), Moss Flora of North America 2: Pl. 67G (as P. oederi).—Fig. 548-549.

damp, shaded cliffs or in crevices, sometimes on humus over boulders, generally
calciphilous. Circumpolar; Hawaii; Greenland to Alaska and south to Virginia, lowa, Colorado, and
Oregon.

MACKINAC CO.—Mackinac Island.

Similar to Bartramia pomiformis but with leaves somewhat curved though
scarcely contorted when dry and cells slightly roughened by verruculose striations but
not papillose.

2. Bartramia Hedw.

Plants small to rather robust, in loose to dense, often rigid tufts, dull, green,
yellowish, or glaucous above, yellow-brown and radiculose or tomentose below. Stems
not triangular in section, with a cortex of small, firm cells. Leaves sometimes crisped
when dry, gradually or abruptly narrowed from an erect, sometimes sheathing base to
a linear-lanceolate or subulate point, toothed at the margins above and often at the
back of the costa; costa prominent at back, vanishing below the apex to excurrent;
upper cells in 2-3 layers (sometimes only at the margins), small, subquadrate to
oblong-linear, firm-walled, papillose at the ends on both surfaces; lower cells long-
rectangular to linear, pale, smooth, thin-walled. Setae usually elongate, straight or
sometimes curved; capsules mostly inclined and asymmetric, subglobose, oblique at the
mouth, ribbed (becoming deeply furrowed, contracted at the middle, flattened at the

192

——-
ee

La
———-

1 \
h \
N \\
Vy

—
Boras
——

i) il ! \\ o il }

aa nigritum, 545. Habit. 546. Leaves. 547. Cells at leaf tip.
Plagiopus oederiana, 548. Leaf. 549.

Ba aia pomiformis. 550. Habit
Capsule, moist.

pper cells at margin of lea
S51.
dry

Leaf. 5

52. Upper cells at margin of
Philonotis fontana, 554. Leaves. 555. Upper cells at margin of leaf. 556. Capsule, wet and

ends, and curved when dry), rarely wrinkled rather than ribbed; operculum convex or
convex-conic; peristome single or dcuble, rarely lacking, the teeth without intermediate
thickenings at back; endostome with a basal membrane, segments well developed or

often reduced or lacking, and cilia none or poorly developed. Spores spherical and
reniform._Named for the noted plant collector and explorer,
Bartram wa

John Bartram of

ee an who corresponded with Linnaeus and sent mosses to Dillenius. John
so an ancestor of a recent, great American bryologist,
Bartram.

Edwin B.
artramia pomiformis Hedw.—Plants in rather dense

, soft tufts 2-6cm. high
light- or yellow-green or yellow-brown, sometimes + glaucous, tomentose below. Leaves

ae

leaf. 553.

550

193

+ spreading and flexuose or crisped from an erect base when dry, loosely erect or
erect-spreading when moist, 4-5.5mm. long, linear-lanceolate, acuminate; margins
revolute almost to the apex, bistratose and doubly serrate; costa shortly excurrent,
serrate at back toward the apex; upper cells subquadrate and short-rectangular, about
1-2:1, thick-walled, unipapillose on both surfaces; basal cells pale, long-rectangular,
rather thin-walled. Synoicous and autoicous (the perigonia next to the perichaetia).
Setae 8-20 mm. long, erect-flexuose; capsules 1.5-2 mm. long, inclined to horizontal,
globose to ovoid, deeply furrowed when dry; annulus none; operculum convex;
peristome teeth inserted somewhat below the mouth, lanceolate, red-brown, very finely
papillose, strongly barred at back; endostome pale-brown, shorter than the teeth,
consisting of a well-developed basal membrane and broad segments. Spores spherical
and + ellipsoidal, brown, coarsely papillose, 20-24 u.—n = 8, 8+1; 2n = 16

Conard, How to Know the Mosses, (ed. 2) fig. 89. Darlington, Mosses of Michigan, fig. 85.
Grout Mosses with Hand-lens and Microscope, Pl. 44 and fig. 104a, c, ie f, Moss Flora of North

i : . 68A-B (as B. glaucoviridis & B. circinatula). Jennings, Mosses of Western
Pennsylvania, (ed. 1) Pl. 24, (ed. 2) Pl. 28. Welch, Mosses of Indiana, fig. ILL —Fig. 550-553.

n moist, ee soil of steep banks in woods and often on ledges, crevices, or faces of
cliffs. Europe, Asia, Madeira, New Zealand; Greenland; Alaska to Oregon and Montana; Labrador to
Manitoba, south to the Gulf States.

CHEBOYGAN CO.—Douglas Lake (at Polypody Bluff and on the campus of the aaa
Station). EMMET CO.—Maple River dam. PRESQUE ISLE CO.—Lamprey Experiment Statio

This is the APPLE MOSS, rare here but common throughout most of eastern
North America and often associated with Aulacomnium heterostichum. In addition to
the apple-like capsules (characteristic of the family), the slender, crisped, and contorted
leaves with papillose cells are distinctive.

tane parts of the Upper Peninsula we have Bartramia ithyphylla Brid.:
Leaves abruptly subulate from a broad, sheathing base, rigidly erect when dry; upper
cells 2-3-stratose except at the margins; costa indistinct above and appearing to fill
most of the subula; margins merely serrulate.

3. Philonotis Brid.

Plants small to moderately robust, usually in dense tufts, dull- or bright-, bluish-
or whitish-green or yellowish, radiculose to tomentose. Stems forked or frequently
whorled (below inflorescences). Leaves erect to spreading and sometimes secund,
usually not much altered on drying, ovate to lanceolate, occasionally blunt but usually
acuminate and narrowly acute; costa percurrent to excurrent; upper cells unistratose,
oblong to linear or shortly rhomboidal, papillose at 1 or both ends or rarely smooth;
lower cells usually larger and thinner-walled. Perigonia bud-like or larger and discoid
with bracts abruptly narrowed from a broad, sheathing base, the paraphyses filiform-
clavate; perichaetia with filiform paraphyses. Setae elongate, erect; capsules strongly
inclined to horizontal, subglobose with a short neck, asymmetric, ribbed (furrowed,
usually contracted at the middle, and flattened at the base when dry); operculum
convex-conic, blunt or mammillate; peristome teeth dark- or red-brown, densely
papillose; endostome consisting of keeled, vertically papillose-striate segments from a
high basal membrane and cilia rudimentary to fairly well-developed. Spores reniform,
orange or brown, densely papillose.-The name means loving moisture.

Leaves rather abruptly acuminate from a + broad base; upper leaf cells papillose at the Be

ends on the outer surface, at the upper ends on the inner surface P. fontana
aerial rae ee gradually acuminate from a narrow base; upper cells papillose only ms the

u ends 1. P. marchica

194

1. Philonotis marchica (Hedw.) Brid.—Plants in loose or dense, bright-green tufts

2-5 (sometimes 8) cm. high. Leaves erect-spreading to spreading, not or slightly curved,

1.3-2.3 mm. long, lanceolate, acuminate, acute; margins serrulate nearly to the base,

plane or nearly so; costa excurrent; upper cells linear, papillose at the upper ends.

Dioicous; perigonia discoid, Setae 24cm. long; capsules 2-2.5mm. long. Spores
26 u

Grout, Moss Flora of North America 2: Pl. 69G. Welch, Mosses of Indiana, fig. 112.

1 in wet habitats, roadbanks, near springs, etc. Circumpolar; British Columbia to
Arizona and widespread in eastern North America from southern Canada to the Gulf of Mexico.

CHEBOYGAN CO.—On a roadbank northeast of Burt Lake (near Livingston Bog).

e leaf margins plane or nearly so and the leaf cells papillose only at the upper
ends are sufficient to distinguish this uncommon species locally. On Sugar Island in
nearby Chippewa County, however, Philonotis capillaris Lindb. ex C. J. Hartm. is
found. It has shorter leaves (0.5-1.1 mm. long), with oblong cells (24:1).

2. Philonotis fontana (Hedw.) Brid.—Plants in dense, bright-green or yellowish
tufts 3-10(16) cm. high (pale or whitish and noticeably red-tinged when growing).
Stems red. Leaves erect, erect-spreading, or + falcate-secund, |.1-2 mm. long, ovate-
lanceolate, + abruptly acuminate; margins serrulate, revolute in the lower % or more;
costa short- to long-excurrent; upper cells oblong-linear or linear, papillose at the lower
ends on the outer surface and at the upper ends on the inner surface. Dioicous;
perigonia discoid. Setae 2-4 cm. long; capsules 2-3 mm. long. Spores 24-30 u.—

Conard, How to Know the Mosses, (ed. 2) fig. 87. Darlington, Mosses of Michigan, fig. 86.
mee Mosses wi ith Ha nd-le ens and Microscope, fig. 103, Moss Flora of North America 2: Pl. 69A
, 7OA-E (including varieties). Jennings, Mosses of Western Pennsylvania, (ed. 1) Pl. 25, (ed. e Pl.
aA Welch, Mosses of ane fig. 113, 114 (as var. pumila), 115 (as var. falcata),—Fig. 554-556

On soil or rock in wet, often seepy places, on banks of ditches, around springs, along
brooks, etc., usually in the open. Circumpolar; widespread in North America south to northern
Mexico.

CHEBOYGAN CO.—Gorge of Carp Creek, Iron Bridge, Reeses ee Burt Lake, oe
Point Nipigon. EMMET CO.—Wilderness State Park, Cecil Bay, Conway Bog, Ramona Beach.
MACKINAC CO.-Bois Blanc Island, Mackinac Island, St. Martins Point, Epoufette. PRESQUE
ISLE CO.—Mast Point, Hammond Bay, Evergreen Beac

This is a species of great variability and, as Dixon said, “No good purpose seems
served by giving varietal names to the numberless variations which occur.” Flowers,
after “‘spending a lifetime with the genus,” wrote me shortly before his death that he
would no longer recognize any of the pas varieties included in his revision of the
genus in aren s Moss Flora of North Amer.

white or bluish-glaucous color of ite leaves, when fresh, contrasts with the
red oad (Pohlia wahlenbergii, of similar habitats, is softer and more delicate and
does not form compact tufts.) The leaf cells have papillae on both ends (at the lower
end on the outer surface and the upper end on the inner), a feature that will usually
suffice to distinguish this, the most common species of a bad genus.

ORTHOTRICHACEAE

Plants small to fairly robust, dull, green, brown, yellow-brown, or reddish, in
small cushions or extensive tufts or mats. Stems erect or erect-ascending and generally

195

forked or long and creeping with numerous erect-ascending and often forked branches,
radiculose below or along the creeping stems. Leaves crowded in many rows, often
curled or spirally contorted when dry, ovate- to ovate-lanceolate or oblong-lanceolate,
mostly acute, often apiculate, rarely awned, usually keeled, mostly entire; costa single,
ending somewhat below the apex to excurrent; upper cells mostly rounded-hexagonal,
occasionally elongate, firm- or thick-walled, green, usually uni- or pluripapillose on
both surfaces, sometimes bulging to mammillose, rarely smooth; lower cells mostl

elongate-rectangular to linear, often incrassate, hyaline or yellowish. Perichaetial leaves
rarely much differentiated. Setae mostly terminal, mostly erect, very short to elongate,
sometimes roughened; capsules immersed to exserted, erect and symmetric, broadly
ovoid to oblong-cylindric, sometimes pyriform, often ribbed; annulus usually present,
operculum rostrate; stomata present, sometimes immersed; peristome usually double,
sometimes single, rarely lacking, the teeth 16 but often united or in pairs, sometimes
completely united into an irregular membrane, lanceolate, usually papillose or variously
striolate, sometimes with a fragmentary pre-peristome; endostome often reduced or
lacking, the basal membrane (sometimes lacking) low, the segments linear to lanceolate,
8 or 16, alternating with the teeth, cilia none. Calyptrae cucullate or mitrate, naked or
hairy, smooth or plicate.

1. Calyptrae cucullate; brood-bodies common, on radicles or in leaf axils; leaf margins plane;
leaf cells short nearly to the insertion . Zygodon
1. Calyptrae mitrate; brood-bodies, if present, on leaves; leaf margins mostly revolute; leaf
cells elongate below
2; ae bordered at base by 1-several rows of pale, short cells wie aren poner
he

inner basal cells linear, yellowish, thick-walled, usually in what radiating r
eee with a rather long, tapered neck, + spindle-shaped; ae superficial, Pee
to the neck . Ulota

. Leaves not bo ed at base, the inner basal cells not as above; capsules short-necked,
not spindle-shaped; stomata superficial or immersed, not restricted to the neck 1. Orthotrichum

1. Orthotrichum Hedw.

Plants small to fairly robust, typically in small tufts, mostly dark-green or
brownish, dull, radiculose at base, erect or ascending, forked or fasciculately branched.
Leaves crowded, rarely much contorted when dry, ovate-lanceolate to oblong-
lanceolate, acute or rarely blunt or rounded, very rarely piliferous, mostly + keeled,
nearly always revolute at the margins, generally from the base eae! to the apex; costa
rather strong, usually ending near the apex; upper cells green, + rounded, thick-walled,
incrassate, usually pluripapillose on both surfaces; lower cells rectangular (or sub-
quadrate) to linear, not or somewhat incrassate, hyaline or pellucid, sometimes shorter
at the margins but not forming a hyaline border. Autoicous or rarely dioicous;
perichaetial leaves ae slightly differentiated. Setae terminal, straight, very short to
fairly long; capsules immersed to exserted, oblong-ovoid to subcylindric from a +
tapered neck, smooth or 8- or 16-ribbed (on drying often becoming deeply furrowed

and sometimes strongly contracted below the mouth), the ribs corresponding to
colored bands of exothecial cells; annulus none or poorly differentiated and persistent;
operculum stoutly rostrate from a convex or conic base; stomata superficial or
immersed, not restricted to the neck; peristome mostly double but sometimes single
and rarely lacking, usually with 16 broadly lanceolate teeth, sometimes approximate or
joined in pairs, occasionally with a pre-peristome, and 8 or 16 filiform segments
alternating with the teeth. Spores spherical, brown, papillose, Calyptrae campanulate-
mitrate, covering most of the urn, usually + plicate, naked or hairy, smooth or rarely
papillose.—The generic name refers to the straight (upright) hairs on the calyptrae
of many of the species (but sometimes sparse). The stomata have taxonomic im-
portance in this genus. Species with superficial stomata have guard cells at the

196

surface, but those with immersed stomata have guard cells sunken in chambers partially
or almost completely roofed over by projections of surrounding cells. Those wi
superficial stomata have a haploid chromosome number of six; those with immersed
stomata have (with one known exception) 11. Confusion with Grimmia can be avoided
by referring to the discussion of that genus, which belongs to the Haplolepideae rather
than Pa esr eae
the few attempts to analyze phylogenetic relationships and the direction
of Hele, was made by Vitt (The Family Orthotrichaceae (Musci) in — America,
1970) using the “Wagner Tree’? method. Koponen’s work on Mnium (Ann. Bot. Fenn.
5: 117-151. 1968) and Seki’s on the Sematophyllaceae (Jour. Sci. Hiroshima aot er.
B., div. 2, 12: 1-80. 1968) also attempt objective analysis of evolutionary relationships.
Techniques for the evaluation of characters distinguishing species are exemplified
by Briggs (New Phytol. 64: 366-386. 1965), Greig-Smith (Trans. Brit. Bryol. Soc. 2:
458-469. 1954), Iwatsuki (Bryol. 61: 68-78. 1958), Lefebvre & Lennes (Taxon 18:
291-299. 1969), Koponen (Ann. Bot. Fenn. 4: 67-73. 1967), Rahman (Jour. Bryol. 7:
169-179. 1973), Redfearn (Bryol. 59: 256-262. 1956), Smith (Trans. Brit. Bryol. Soc.
6(1): 56-68. 1970), and Suzuki (Jap. Jour. Bot. 16: 227-268. 1958).

1. Leaves with erect margins and broad, blunt to rounded t
gin

tips 1. O. obtusifolium
s and narrow, blunt or acute tips

h
when dry; segments rather spe smooth o . sordidum
3. Capsules narrowly cylindric, smooth or rately obscurely plicate, not contracted below
the mouth; segments vey ioe densely papillo 3. O. speciosum var. elegans
2. Stomata immersed

3. Capsules oblong-cylindric, strongly ribbed oe moderately contracted below the mout
nearly so

4. ae (eee on rock; capsules shortly exserted; peristome teeth erect or + spreading

. anomalum
4. Pant ae bark; capsules immersed to emergent; peristome teeth mostly recurved when

5; Capsules small and delicate, straw-colored, wrinkled or sometimes lightly ribbed and
at all contracted below the mouth when dry; exothecial cells obscurely or
peice rather clearly differentiated in bands 4 cells wide; peristome teeth erect (or
occasionally becoming + reflexed with age), soon splitting into 16 small and irregular
divisions; endostome none . O, pusillum
5. Capsules darker, not delicate, distinctly ribbed and usually + contracted below the
mouth when dry; exothecial cells well differentiated; peristome teeth 8 or eventually
becoming split into 16, but the divisions not particularly small or irregular; endostome
segments present
6. Leaves apiculate and sometimes sparsely oes at the tip; cells surrounding stomata
not notably y jutting or otherwise differentia . O. pumilum
6. Leaves not apiculate or toothed at the pee cells surrounding stomata clearly
differentiated

7. Capsules strongly is and very strongly contracted below the mouth when dry;
tibs about c wid ll

pically dark-brown; cells surrounding stomata
projecting eee ie peristome teeth 8 and usually not splitting into 16 ee

. O. stellatum

7. Capsules rather narrowly and ae crn and not or slightly ea below

the mouth when dry; ribs abou wide and pale-brown; cells surrounding

stomata Hae _steply or aa ie cies peristome teeth 8, regularly splitting
into 16 divisio

O. ohioense

1. Orthotrichum obtusifolium Brid.—Plants in dense, = dark-green to brown
or yellowish tufts 5-10(25) mm. high. Leaves about 1.5- long, erect-appressed,
scarcely altered when dry, ovate (or rarely lanceolate), se or rounded at apex,
concave, with margins erect and entire, usually bearing abundant small, green or
red- brown, shortly cylindric brood bodies 45 cells long on the upper surface of the
lamina (and sometimes on both surfaces); costa slender, ending well below the apex,
narrowly channeled; upper cells isodiametric, 13-18 u wide, strongly thickened at the

197

corners, green, unipapillose on both surfaces; lower cells rectangular, 2-4:1, with firm
walls often + pitted, pale and smooth, Dioicous. Seta less than 0.5 mm. long; urn
immersed to emergent, about 2 mm. long including the neck (about as long as the
spore sac), elongate-pyriform, cylindric and + strangulate when dry and empty, strongly
8-plicate, dark-brown; exothecial cells clearly differentiated along the ribs; stomata
superficial; peristome double, the teeth fused together in 8 pairs, red or orange-red,
densely papillose or irregularly papillose-striolate, reflexed when dry, the segments 8,
brownish-yellow, smooth or striolate, well developed, of 2 rows of cells, erect. Spores
16-18 yp. Calyptra not plicate, naked, papillose throughout.—n = 11.

Conard, How to Know the Mosses, (ed. 2) fig. 67. Grout, Mosses with Hand-lens and
Microscope, fig. 91. Jennings, Mosses of Western Pennsylvania, (ed. 2) Pl. 63. Welch, Mosses of
Indiana, fig. 100.—Fig. 557-562.

n the bark of trees (commonly on poplars), rarely on old fence rails. Europe; Japan;
Yukon to Labrador and Newfoundland, south to California, Arizona, the Great Lakes, and North
olina.

CHEBOYGAN CO.-—Reeses Bog, Carp Creek, Smiths Bog, Burt Township Hall, Riggsville,
Passino School, Mud Lake. EMMET CO.—Alanson, Levering, Larks Lake, Pellston Dunes, Pellston,
west branch of Maple River, Pellston Hills, Stutsmanville, Middle Village, Wycamp Lake. MACK-
INAC CO.—Bois Blanc Island, Mackinac Island, Gros Cap, Caffey Corner, Hendricks. PRESQUE
ISLE CO.—Lamprey Experiment Station, Loon Lake, Onaway.

Perhaps our most common Orthotrichum, unlike all the others because of the
leaves with rounded-obtuse apices and erect margins.

9. Orthotrichum sordidum Sull. & Lesg. ex Aust.—Plants in small, dense,
dark-green to yellow-brown cushions about 6-10 mm. high. Leaves erect and slightly
contorted when dry, erect-spreading when moist, 1.5-3 mm. long, oblong- or ovate-
lanceolate, acute; margins entire, revolute nearly to the apex; costa ending slightly
below the apex; upper cells 8-13 yu, irregularly rounded and rounded-hexagonal,
incrassate, pluripapillose on both surfaces; basal cells rectangular, moderately incrassate
and pitted. Autoicous. Setae up to 2.5 mm. long; capsules emergent to shortly
exserted, 1-1.8 mm. long, yellow-brown, oblong-cylindric and tapered at the neck,
moderately constricted below the mouth and strongly 8-ribbed when dry and empty;
exothecial cells moderately differentiated along the ribs; stomata superficial; peristome
teeth orange-brown, strongly reflexed when dry, fused in 8 pairs (occasionally
becoming + split), densely and finely papillose, cancellate at the apex; segments 8,
brownish-yellow, relatively stout (consisting of 2 rows of cells), smooth or + obscurely
papillose. Spores 17-24 yw. Calyptra plicate, hairy, smooth.—n = 6.

Grout, Mosses with Hand-lens and Microscope, PI. 39. Welch, Mosses of Indiana, fig.
99.—Fig. 563-566.

n the bark of trees. Newfoundland and Nova Scotia to Ontario and south to New England
and the Great Lakes; Japan.

CHEBOYGAN CO.—Hermits Bog, Gorge of Carp Creek, Colonial Point, Livingston Bog,
Riggsville, Mud Lake, Cheboygan, Mill Creek, Mud Creek, Black Lake. EMMET CO.—Brutus,
Pellston Dunes, Pellston, Pellston Hills, Middle Village, Good Hart, Wycamp Lake. MACKINAC
CO.—Bois Blanc Island, Mackinac Island, Gros Cap, Epoufette, Hendricks. PRESQUE ISLE
CO.—Onaway, Rainy River Falls, Posen.

The capsules are strongly ribbed and somewhat contracted below the mouth
when dry, and the peristome teeth are orange and somewhat latticed at their tips. This
is our only species of the superficial stomate group with distinctly ribbed or
strangulate capsules.

ee
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Orthotrichum obtusifolium. 557. Habit. 558. Leaves.
Brood-bodies. $61. Capsule, dry. 562. Portion of peristome

rthotrichum sordidum, 563. Leaves. 564. Upper cells of leaf. 565. Capsule, dry. 566.
Stomate.

559. Upper cells of leaf. 560.

Orthotrichum speciosum var. elegans. 567. Leaves. 568, Upper cells of leaf. 569. Capsules,

3. Orthotrichum speciosum Nees ex Sturm var. elegans (Schwaegr. ex Hook. &
Grev.) Warnst.—Plants 5-10, rarely 20 mm. high, dark-green or sometimes brownish.
Leaves loosely erect when dry, erect-spreading to spreading with recurved tips when
moist, lanceolate, acuminate, acute; margins revolute nearly to the apex; costa ending
at or near the apex; upper cells about 7-11 wide, irregularly rounded and shortly
oval, incrassate, pluripapillose. Autoicous. Setae short: capsules emergent to shortly
exserted, about 1.2-1.7 mm. long, narrowly cylindric, smooth (or rarely faintly plicate
when dry and empty); exothecial cells not differentiated in longitudinal bands; stomata

199

superficial; peristome teeth 8, recurved when dry, pale orange-brown, densely papillose;
endostome segments 8, incurved when dry, stout and about as long as the teeth,
densely papillose. Spores 13-26 py. Calyptrae hairy, smooth.—n = 6, 12, 12+1, 13 (all in
O. speciosum vat. speciosum).

Grout, Moss Flora of North America 2: Pl. SOD (as O. elegans).—Fig. 567-569.

On trees, rarely on old logs, fence rails, or rock. Alaska to Newfoundland, south to Oregon,
Michigan, and New York (common in the East but usually replaced by O. speciosum in its robust
typical form in the West).

CHEBOYGAN CO.—Reeses Bog, Carp Creek, Douglas Lake, Riggsville, Cheboygan, Grass
Bay, Mullett Lake. EMMET CO.—Levering, Brutus, Pellston Dunes, Pellston, Pellston Hills, Five
Mile Creek, Wycamp Lake, Cecil Bay, Wilderness State Park, Big Stone Bay. MACKINAC CO.-—Bois
Blanc Island, Mackinac Island, McKays Creek, McNamara Pond. PRESQUE ISLE CO.—Lamprey
Experiment Station, Rainy River Falls, Clinton Lake, Loon Lake.

The var. elegans is recognized by its narrowly cylindric, smooth capsules, with
yellow-brown peristome teeth strongly recurved (like the handle of a mug) when dry
and pale, stout, papillose segments incurved when dry. (Orthotrichum speciosum has
been found in its typical form in Keweenaw Co. and also on Sugar Island in Chippewa
Co. It is larger, with broader, obscurely ribbed capsules.)

4. Orthotrichum anomalum Hedw.—Plants in rigid, dark- or blackish-green to
brownish cushions about 10-20 mm. high. Leaves erect and scarcely contorted when
dry, erect-spreading when moist, 2-3 mm. long, ovate- or oblong-lanceolate, broadly (or
rarely slenderly) acute and blunt; margins entire, revolute nearly to the apex; costa
ending slightly below the apex; upper cells about 10-11 u wide, irregularly rounded-
hexagonal, + incrassate, pluripapillose on both surfaces; lower cells rectangular, smooth,
pale, somewhat incrassate. Autoicous. Setae 2.5-3 mm. long; capsules 1.5-2 mm. long,
barely to shortly exserted, oblong-cylindric, becoming tapered at the neck, constricted
above the middle and strongly furrowed when dry (with 8 strong ribs usually
alternating with 8 less prominent ones), light- to chestnut-brown, annulus none;
operculum stoutly and bluntly rostrate; exothecial cells differentiated along the ribs;
stomata immersed; peristome teeth at first fused in pairs, later 16, erect or spreading
when dry, yellow-brown, faintly to distinctly sinuose-striolate, often with fragmentary
pre-peristome thickenings at the base; endostome consisting of 8 irregular, smooth cilia,
often rudimentary or lacking. Spores 10-16 uy, papillose. Calyptrae smooth, sparsely to
moderately hairy.—n = 10+1, 11.

Conard, How to Know the Mosses, (ed. 2) fig. 71a-c. Darlington, Mosses of Michigan, fig. 88.
Grout, Mosses with Hand-lens and Microscope, fig. 86.—Fig. §70-572.

On rocks, usually if not always calcareous. Europe; Japan; Mexico and Haiti; Alberta to
Nova Scotia, south to Oregon and Arizona, Colorado, Iowa, and Virginia.

EMMET CO.-—Pellston Hills, Big Stone Bay. MACKINAC CO.—Mackinac Island, McKays
Creek, Bush Bay.

Unmistakable by its habitat, on rock, and its exserted capsules with immersed
stomata. Orthotrichum anomalum presents no problems locally, but some characters
useful in cases of doubt are the faint, sinuous markings on the peristome teeth and the
pre-peristome fragments attached to the outer surface near the base of the teeth. The
intermediate ribs on the capsule are distinctive, though not always present. Our only
species growing on rock, Orthotrichum anomalum is superficially similar to Ulota
hutchinsiae (Sm.) Hamm., which prefers acid, siliceous rocks over limestone. (It has

200

been found once in our area, perhaps on an erratic boulder, and it occurs on Sugar
Island in nearby Chippewa Co. It is nearly black except at the growing tips and grows
in extensive, loose colonies. Its capsules are gradually tapered to a long neck, the
sieeti teeth are reflexed, and the stomata are superficial. Also the basal leaf cells

, to some extent, the radiating pattern typical of Ulota, with cells in rows fanning
aa ee the base toward the shoulders.

5. Orthotrichum stellatum Brid.—Small, yellow-brown plants about 3-5 mm. high.
Leaves rather rigid, erect and scarcely contorted when dry, spreading or erect-spreading
when moist, 1.8-2.3mm. long, oblong-lanceolate, blunt or bluntly acute; margins
revolute nearly to the apex, entire; costa ending at or near the apex; upper cells 9-13 py
wide, irregularly rounded, moderately incrassate, low-papillose. Brood-bodies pale-
brown, cylindric or shortly filamentous, occasional on leaves. Autoicous. Capsules
emergent, 1.5-2 mm. long, oblong-cylindric when moist, narrower, deeply 8-ribbed,
strongly strangulate, and tapered to the seta when dry, chestnut-brown; exothecial cells
clearly differentiated in bands + 8 cells wide; stomata immersed, with surrounding cells
clearly differentiated and projecting nearly horizontally; peristome teeth 8 (rarely
split), reflexed when dry, densely and finely papillose; segments 8, linear, of 2 rows of
cells, smooth. Spores 11-18 y. Calyptrae plicate, naked or sometimes + hairy, smooth.

, Mosses with Hand-lens and Microscope, fig. 88 (as O. strangulatum). Jennings, Mosse
of Western’ aa (ed. 1) Pl. 15 (as O. braunii), (ed. 2) Pl. 19. Welch, Mosses of Indiana, ‘is,
102.—Fig. 573-575.

On trees, rarely on logs, stumps, or fence rails. ey and Scotland: Newfoundland to
Wisconsin and Iowa, south to Arkansas and North Carolin

EBOYGAN CO.—Reeses Bog, Carp ah ee EMMET CO.—Alanson, Wycamp
Creek, Middle Village. PRESQUE ISLE CO.—Belkna

The specific epithet refers to the peristome teeth which may spread like the rays
of a star when dry. The species has been nomenclaturally confused with O.
strangulatum P.-Beauv. (fig. 576-578), a limestone species with strangulate capsules and
16, rather than eight peristome teeth.

The conspicuously strangulate capsules with straight, heavy, chestnut-brown
ridges which become contiguous on complete contraction are unmistakable. In QO.
pumilum, the species most likely to be confused with O. stellatum, the capsules are
fairly strangulate and the ribs dark-brown and prominent, but on contraction of the
suboral region they do not become contiguous, and they often seem slightly spiraled.
Of course, the apiculate and even denticulate apices of many leaves in O. pumilum are
distinctive. In O. ohioense, the capsules are at most only slightly strangulate, and the
tibs, though distinct, are pale, yellow-brown, narrow, and widely spaced.

Orthotrichum pallens Bruch ex Brid. (fig. 579-581) is a western species found
disjunctively in nearby Alpena County and in the Upper Peninsula in Keweenaw
ounty. It is autoicous, as most Orthotricha are, but has the perigonia at the ends of
short branches rather than in leaf axils near the perichaetia. It has strongly ribbed,
strangulate capsules with slightly to distinctly immersed stomata, peristome teeth
united in eight pairs, 16 slender endostome segments, and a naked calyptra. A related
species, also disjunct from a broad western range in Keweenaw County, O. alpestre
Hornsch. ex BSG, has axillary perigonia below the perichaetia, eight stout segments,
and a somewhat hairy calyptra (with papillae on the hairs).

\ | | ! . \ | | | f HI ty
Vs) LY i sal

Orthotrichum anomalum. 570. Leaves. 571. Upper cells of leaf. 572. Capsules, dry.
Orthotrichum stellatum. 573. Leaves. 574. Upper cells of leaf. $75. Capsule, dry.
Orthotrichum strangulatum. 576. Leaves. 577. Upper cells at leaf margin. 578. Capsules, dry.
Orthotrichum pallens. 579, Leaves. 580. Upper cells of leaf. 581. Capsules, dry

6. Orthotrichum ohioense Sull. & Lesq. ex Aust.—Small, brownish plants up to
10 mm. high. Leaves rigid, erect when dry, erect-spreading when moist, 2-2.5 mm. long,
oblong-lanceolate, bluntly acute; margins revolute nearly to the apex, entire; costa
ending slightly below the apex; upper cells 9-12 wide, irregularly rounded or
transversely oval, rather evenly incrassate, rather obscurely papillose on both surfaces.
Brood-bodies apparently lacking. Autoicous. Setae about 0.7-0.8 mm. long; capsules
emergent, oblong-cylindric, pale-brown, narrowly and distantly ribbed and not or only
slightly constricted below the mouth when dry and empty; exothecial cells differ-
entiated in longitudinal bands 4 cells wide; stomata at middle of the capsule, immersed
because of greatly differentiated, thick-walled cells jutting steeply or almost vertically

202

over the stomatal chamber, peristome single, the teeth reflexed when dry, 8, soon
splitting into 16 small, lanceolate divisions, densely and finely papillose, reflexed when
dry, sometimes with inconspicuous pre-peristome thickenings at base; segments 8,
smooth; Spores 15-21 ys. Calyptrae plicate, hairy, smooth.—n = 10+1, 11

Conard, How to Know the Mosses, (ed. 2) fig. 73f-¢. Grout, Mosses with Hand-lens and
icroscope, fig. 87. Jennings, Mosses of Western Pennsylvania, (ed. 1) Pl. 15, (ed. 2) Pl. 19. Welch,
Mosses of Indiana, fig. 103.—Fig. 582-585.

On bark of trees. Nova Scotia to Michigan, south to South Carolina, Mississippi, and
Arkansas,

CHEBOYGAN CO.-—Reeses Bog, Gorge of Carp Creek. EMMET Co.—Middle Village. MACK-
INAC CO.—Hendricks.

A species commonly misunderstood but distinguished by pale capsules with
narrow and remote but fairly well-marked ribs, 8 peristome teeth becoming split into
16 small, pale divisions, and stomatal chambers covered by greatly differentiated cells
jutting almost vertically outward.

7. Orthotrichum pumilum Sw.—Small, dark- or blackish-green plants. Leaves
erect when dry, spreading when moist, typically rather soft, about 2 mm. long,
ovate-lanceolate, obtuse or acute, apiculate or slightly toothed at the extreme apex;
margins revolute nearly to the apex; costa ending near the apex; upper cells 13-20 y,
usually rather pellucid, rounded because of + thickened corners, sometimes + broader
than long. Brood-bodies occasional on the leaves. Autoicous. Capsules emergent, about
1.5 mm. long, oblong-ovoid, strongly 8-ribbed and strangulate (sometimes strongly so)
when dry, dark-brown; exothecial cells clearly differentiated in bands + 8 cells wide
(cells of ribs not greatly incrassate below the suboral zone as in O. stellatum); stomata
immersed, the cells surrounding the stomatal cavities usually not jutting or notably
differentiated: peristome teeth 8, reflexed, brownish, densely papillose; segments 8,
yellow, smooth. Spores 13-17 pw. Calyptrae plicate, naked or + hairy.—n = 10+1, 1

, How to Know the Mosses, (ed. 2) fig. 73a-e. Grout, Mosses with Hand-lens and
mses es 89 (as O. schimperi). Welch, Mosses of Indiana, fig. 104. —Fig. 586-589.

On trees. Germany and Austria; New England to Ontario, South Dakota, and Nebraska,
south to North Carolina and Louisiana

CHEBOYGAN CO.—Riggsville, Mud Lake, Passino School. EMMET CO.—Brutus, Pellston,
Maple River, Pellston Hills, Five Mile Creek. MACKINAC CO.—Hendricks.

Known by apiculate upper and perichaetial leaves and strongly ribbed capsules
which may be more or less strangulate (but the ribs are not completely drawn together
as in O. stellatum).

8. Orthotrichum pusillum Mitt.—Small, dark- or blackish-green plants about 2 or
3mm. high, scattered or loosely tufted. Leaves loosely erect when dry, erect- -spreading
when moist, 1.5-2.5 mm. long, oblong-lanceolate, bluntly acute; margins revolute nearly
to the apex, entire (except for irregular denticulations at the apex of perichaetial
leaves); costa ending slightly below the apex; upper cells about 11-13 y wide, rounded
to transversely oval, incrassate, pluripapillose on both surfaces. Brood-bodies sometimes
produced on the leaves. Autoicous. Setae about 0.8-1 mm. long; capsules immersed to
emergent, 1-1.75 mm. long, oblong-cylindric, delicate and thin-walled, somewhat
wrinkled to lightly 8-plicate and not at all constricted below the mouth when dry and

203

i

Orthotrichum ohioense. 582. Leaves. 583. Upper cells of leaf. 584. Capsule, dry. 585.
Stomate.

Orthotrichum pumilum. 586. Leaves. 587. Cells at leaf tip. 588. Upper cells of leaf. 589.
Capsule, dry.

Orthotrichum pusillum. 590. Leaves. 591. Upper cells of leaf. 592. Capsule, dry.

empty, straw-colored; exothecial cells obscurely to rather distinctly differentiated in
bands 4 cells wide; stomata immersed, the surrounding cells not particularly jutting or
greatly differentiated; peristome teeth erect or sometimes becoming reflexed with age,
soon becoming split into 16 small, irregular divisions, pale-brown, densely and finely
papillose, often opaque; endostome none. Spores 13-18 yp. Calyptra plicate, smooth,
naked.—n = 11.

Fig. 590-592.
On bark of trees. Throughout the eastern United States.

CHEBOYGAN CO.-Riggsville, Passino School. EMMET CO.—Levering, Cross Village, Middle
Village, Ramona Beach. MACKINAC CO.—Hendricks. PRESQUE ISLE CO.—Clinton Lake.

Among the species with immersed stomata, this one is distinctive because of
small, delicate, pale, somewhat wrinkled to obscurely plicate capsules with the
peristome becoming split into 16 small and irregular divisions which are recurved rather
than reflexed.

2. Ulota Mohr ex Web.

Relatively small plants usually in small tufts, green, yellowish, or brownish above,
dark-brown to blackish and + radiculose below. Stems erect, forked. Leaves crowded,

204

mostly crisped and contorted when dry, lanceolate or linear-lanceolate from a broader,
+ concave base, sharply to bluntly acute, + keeled; margins usually + revolute along the
middle (especially when dry); costa percurrent or nearly so; upper cells rounded or
elliptic, thick-walled, mostly papillose; cells of the sheathing base linear, yellowish, very
thick-walled and sometimes porose, usually smooth, | or more marginal rows short and
pale with thickened cross-walls. Perichaetial leaves not or slightly differentiated. Setae
terminal, straight, + elongate; capsules exserted, erect and symmetric, cylindric,
fusiform, or narrowly urceolate, with a relatively long, tapering neck, 8-ribbed (or
merely puckered at the mouth): annulus none; operculum stoutly rostrate from a
convex or convex-conic base; exothecial cells differentiated along the ribs; stomata
superficial, restricted to the neck; peristome mostly double, the teeth 16, often united
in pairs, lanceolate; segments of endostome 8 or rarely 16 (sometimes lacking), linear.
Calyptrae mitrate, deeply lobed, + hairy.—-Ulota means something curled and refers in
this case to the crisped leaves of some species of the genus.

1. Capsules pyriform, smooth except at the puckered mouth 1. U. coarctata

1. Capsules oblong-cylindric to fusiform, plicate
saxicolous; leaves slightly curved but aren crisped when dry, 1.7-2 lon

neck of capsules not clearly delimited; spores 9-1 3. a aie
2. Plants corticolous; leaves strongly crisped when ae 2-3 mm. long; capsules with a long,
tapered neck; spores 18-24 u 2. U. crispa

1. Ulota coarctata (P.-Beauv.) Hamm.—Plants yellow-green to brown. Leaves
erect and not or slightly contorted when dry, erect-spreading when moist, 1.5-2.5 mm.
long, lance-acuminate from a short, concave base, acute; Margins entire, narrowly
revolute along the middle; costa ending somewhat below the apex; upper cells smooth
or indistinctly papillose on both surfaces, rounded or elliptic, about 7 X 794.
Autoicous. Setae 3-4mm. long; capsules 1.5-2.2 mm. long, pyriform, long-necked,
strongly puckered by 8 short ribs and nearly closed at the mouth when dry; peristome
teeth at first fused in 8 pairs, later 16, not recurved when dry, densely papillose;
segments 8, very short and rudimentary. Spores 17-22 u, ellipsoidal, papillose.
Calyptrae densely pilose.—n = 9+1.

Conard, tae to Know the Mosses, (ed. 2) fig. 65f (as U. ludwigii). Grout, Mosses with
Hand-lens and M croscope, fig. 84 (as U. ludwigii). Jennings, Mosses of Western Pennsylvania, (ed.
1) Pl. 15, (ed. 2) PL 19 (as U. ludwigii).-Fig. 593-595.

On bark of trees in damp woods, rather rare. Europe; Newfoundland to Michigan and south
to North ponen and Tennessee.

EMMET CO.—Cecil Bay. MACKINAC CO.—Mackinac Island, Epoufette, Hendricks Tower.

A curious little moss with pear-shaped capsules abruptly narrowed to a small,
puckered mouth. (My impression is that it grows most often on yellow birch trees in
beech-maple-hemlock forests and probably near streams.

2. Ulota crispa (Hedw.) Brid.—Plants green, yellow-green, or yellow-brown.
Leaves erect and strongly crisped when dry, erect-spreading when moist, 2-3 mm. long,
narrowly lanceolate from a broader, concave base, acute or narrowly obtuse; margins
entire, plane or sometimes slightly and irregularly revolute along the middle; costa
vanishing near the apex; upper cells rounded-quadrate to elliptic, unipapillose on either
surface, 8-10. Autoicous, Setae 2-4mm. long; capsules 1.5-2 mm. long, oblong-
cylindric from a long, tapered neck, becoming contracted below the mouth and (rather
abruptly) to the neck when dry, strongly 8-ribbed; peristome teeth united in 8 pairs,
finely papillose, often perforate near the tips, reflexed when dry; segments 8, well

205

developed, smooth or rarely very finely papillose. Spores 18-24 u, spherical, papillose.
Calyptrae densely pilose.—n = 10+1, 10+2, 19+1, 19+2, 20+2, 21.

eed How to Know the Mosses, a 2) fig. 65c-e. Grout, Mosses with Hand-lens and
Microscope, Pl. 38. Jennings, Mosses of Western Pennsylvania, (ed. 1) Pl. 16, (ed. 2) Pl. 20. Welch,
Mosses of Indiana, fig. 106.—Fig. 596-599.

k of trees, usually in moist woods. Europe, North Africa, ra Islands, Asia;
southeastern Alaska; Newfoundland to Eanes south to Tennessee and Georg

CHEBOYGAN CO.-—Reeses Bog, Hermits Bog, Gorge of Carp Creek, Iron Bridge, Fontinalis
Run, Livingston Bog, Cheboygan, Grass Bay, Mill Creek, Mud Creek (at Black Lake). EMMET
CO.—Pellston, Larks Lake, Cecil ce y, Five Mile Creek, Ramona Beach. MACKINAC CO.—Bois
Blanc Island, Mackinac Island, St. ins Point, Gros Cap, Cut River, Epoufette, Ozark, Hendricks.
PRESQUE ISLE CO.—Hammond soe ee Bea Clinton Lake.

In small tufts on bark in moist, coniferous forests, contrasting with the
Orthotricha which occur most commonly on roadside hardwoods. The pale, ribbed
capsules with a conspicuous neck and the strongly crisped leaves are the recognizable
features of the species.

. Ulota hutchinsiae (Sm.) Hamm.—Small plants in rigid tufts (about 1-1.5 cm.
high), aa yellow-brown to dark-green above, blackish below. Leaves erect and slightly
curved but scarcely contorted when dry, erect-spreading when moist, 1.7-2 mm. long,
lanceolate, gradually narrowed to a slender, bluntly acute apex; margins + revolute on
1 or both sides in the lower % or sometimes more; costa ending near the apex, upper
cells nearly isodiametric, irregularly rounded or elliptic, 8-12 X 7-8, very thick-
walled, pellucid, nearly smooth to bluntly and usually obscurely unipapillose on both
surfaces. Autoicous. Setae 2.5-3.5 mm. long; capsules about 1.5-2 mm. long, oblong-
cylindric (the neck not clearly delimited, usually no more than about % as long as the
urn), 8-plicate when dry and empty; operculum stoutly long-rostrate from a convex
base; exothecial cells differentiated along the ribs; peristome teeth united in 8 pairs
(rarely separating with age), erect on dehiscence but generally becoming reflexed,
cancellate and often somewhat striolate at the tips, densely papillose below, sometimes
with irregular, rather conspicuous, papillose pre-peristome thickenings at the base;
segments 8, fairly well developed (1/2- 2/3 the length of the teeth), weakly striolate-
roughened (or rarely rather aga so). Spores spherical, papillose, 9-14 yu. Calyptra
copiously hairy.—n = 1

Conard, How to ae the Mosses, Pcs 2) fig. 65a-b (as U. americana). oped Mosses
of Michigan, fig. 89. Grout, Moss Flora of North America 2: Pl. as U. Caner ennings,
Mosses of Western Sen (ed. 1) Pl. ae (ed. 2) Pl. 19 (as U. americana).—Fig. 600-602.

Usually (and perhaps always) on acid, siliceous rocks. Europe; Japan; neveuery Alaska;
widespread in eastern North America from Newfoundland and Labrador to Minnesota and scuth to
Oklahoma, Arkansas, and Georgia. The only specimen from the Douglas Lake es is one at the
New York Botanical Garden ares collected on granite at Cheboygan by E. Clark in 1941.
The record is accepted with reservation, as it is possible, though unlikely, that the species grew
there on a glacial erratic. (It fen (ee collected on Sugar Island in Chippewa County.)

This species is named for Ellen Hutchins (1785-1815) of County Cork, Ireland,
who collected bryophytes and algae, several of which bear her name. Best recognized
by its saxicolous habitat, rigid leaves scarcely contorted when dry, and plicate capsules
with a poorly marked neck. Although not likely to be confused with any other Ulota,
it is superficially similar to Orthotrichum anomalum, which grows on calcareous rather
than siliceous rocks. From that species, Ulota hutchinsiae can be distinguished, as

—— See Nese Oc es

= >

S
~

NE
LY P

Ulota coarctata, 593. Habit.
Ulota crispa. 596. Habit. 59

594. Leaves. 595. Upper cells at margin of leaf.
7, Leaves. 598. Upper cells at margin of leaf. 599. Cells of alar

region.
Ulota hutchinsiae, 600. Habit. 601. Leaves. 602. Upper cells at margin of leaf.
Zygodon virridissimus var. rupestris. 603. Habit. 604. Leaves. 605. Cells at leaf tip. 606.

Brood-bodies.

follows: Its capsules are gradually tapered to a longer neck, the peristome teeth
reflexed, the stomata superficial, and the cells at the base of the leaves arranged, at

least to a degree, in a radiating pattern.

3. Zygodon Hook. & Tayl.

Rather small plants, scattered or in loose or dense, green, yellowish, brownish, or
reddish tufts. Stems erect or ascending, simple or forked, mostly matted with radicles,

207

usually bearing small, green or brown, ellipsoidal to cylindric brood-bodies on radicles
and in leaf axils. Leaves erect and often contorted when dry, spreading to squarrose-
recurved when moist, sometimes clearly S-ranked, mostly oblong to linear-lanceolate
and acute or + acuminate (frequently ending in a pellucid apiculus), rarely lingulate
and rounded at the apex; margins mostly plane, rarely recurved below, entire or +
dentate toward the apex (or rarely nearly to the base); costa prominent at back,
ending in or below the apex, sometimes excurrent; cells usually uniform nearly to the
base, rounded-quadrate or rounded-hexagonal and usually thick-walled, pluripapillose
(or less often smooth or unipapillose) on both surfaces, rectangular or rarely linear and
usually hyaline near the insertion. Perichaetial leaves scarcely differentiated. Setae
terminal, elongate, erect, usually yellowish; capsules erect and symmetric, subcylindric
or narrowly pyriform, usually with a fairly long neck, mostly 8-plicate or furrowed;
annulus very poorly differentiated, persistent; exothecial cells sometimes differentiated
in longitudinal bands; operculum nearly always obliquely long-rostrate from a low,
convex-conic base; stomata superficial, in the neck; peristome lacking to rudimentary
or well developed and double or single (and then consisting of the endostome), the 16
lanceolate teeth + joined in pairs at first, papillose, the 8 or 16 segments linear, usually
weakly papillose. Spores spherical. Calyptrae cucullate, mostly smooth, naked or rarely
hairy, not plicate.—The name signifies yoked or paired teeth, referring to the fact that
the 16 teeth of the peristome are at first joined in pairs.

Zygodon Nabari (Dicks.) Brid.—Plants scattered or in small, loose tufts,
bright- to dark-green or brownish, not or somewhat tomentose. Brood-bodies 3-6-

with + recurved tips when moist, 1-2 mm. long, keeled below, oblong-lanceolate, acute,
rather gradually apiculate, ending in a rather long, clear cell; margins plane (but
somewhat broadly reflexed when dry), entire, somewhat wavy; costa ending well below
the apex; upper cells 7-10, rounded or rounded-hexagonal, thick-walled, rather
coarsely pluripapillose on both surfaces; basal cells pale, subquadrate and short-
rectangular in a few rows = the insertion. Dioicous. Setae 4-7 mm. long, yellow;
capsules 1-1.5 mm. lon arrowly pyriform, furrowed, yellow-brown; operculum
slenderly and obliquely ae rostrate; exothecial cells not or slightly differentiated in
bands; peristome none or rarely represented by a very low, somewhat Epes
membrane. Spores brown, papillose, 11-14 wu. Calyptrae smooth, naked.—n = 12

Grout, Moss Flora of North America 2: Pl. 58A, 59C.—Fig. 603-606 (var. rupestris).

On bark of trees. New England to North Carolina; Michigan and Sree aca tia to
California; Mexico; Canary Islands; Europe, Caucasus, Asia Minor, and North Afric

EMMET CO.-—Larks Lake. PRESQUE ISLE CO.—Clinton Lake.

A species of little character, most often found scattered and sterile. The acute,
gradually apiculate leaves ending in one or more pellucid cells give a hint of its
identity. Our material, like most North American collections, belongs to the var.
rupestris Lindb. ex C. J. Hartm., which is characterized by narrower leaves, more
gradually narrowed to a longer apiculus of 1-3 pale cells in one row and by brood
bodies lacking vertical septations.

Other Orthotrichaceae known from Michigan:
Drummondia prorepens (Hedw.) Schimp. Stems elongate and creeping, with

numerous erect-ascending branches. Leaves imbricate when dry, not contorted, oblong-
lanceolate, blunt or acute, channeled at the costa; cells rounded and green throughout,

208

smooth. Capsules exserted, smooth; peristome single, consisting of 16 very short,
truncate teeth. Spores multicellular, 60-100 uw. Calyptrae cucullate.—On bark of trees.
Widespread in eastern North America but not likely to be found in the Straits region.

Amphidium lapponicum (Hedw.) Schimp.—Plants in dense cushions. Stems erect,
repeatedly forked. Leaves curled and contorted when dry, oblong-lanceolate; upper
cells rounded-quadrate, densely warty-papillose; basal cells differentiated, pale.
Autoicous; perichaetial leaves pale, sheathing, broad, short-pointed. Capsules half-
emergent, urceolate and 8-ribbed when dry and empty; peristome none. Calyptrae
cucullate (n = 13, 16).—On cliffs, especially on undersides of projecting ledges. A
northerly and montane species likely to be found on the limestone outcrops of
Mackinac County, for example.

Amphidium mougeotii (BSG) Schimp., of montane distribution farther east, is
dioicous, with long, narrow, pointed, and scarcely sheathing perichaetial leaves and
capsules just barely exserted. The leaves are moderately flexuose when dry and
linear-lanceolate; the upper cells, only moderately opaque, are both papillose and
longitudinally striolate, and the basal cells are small and thick-walled.

CRYPHAEACEAE

Plants rather slender to fairly robust, generally rigid, in loose, dull or somewhat
shiny tufts. Primary stems creeping and stoloniform, sparsely radiculose; secondary
stems usually + erect-ascending (rarely pendent or floating), irregularly to subpinnately
branched; paraphyllia none or sometimes abundant. Leaves crowded in many rows,
mostly imbricate when dry, + spreading when moist, concave, smooth or sometimes
plicate, acute, acuminate, or rarely long-acuminate from an ovate or lanceolate base,
unbordered; costa single, ending near the middle to subpercurrent or rarely short-
excurrent (rarely short and double in some leaves); cells mostly short, oval to narrowly
elliptic, usually thick-walled, smooth or rarely papillose, elongate toward the base and
quadrate to rounded- or oblate-quadrate often in many oblique rows at the basal
angles. Perichaetia at the ends of very short or sometimes undifferentiated branches;
perichaetial leaves often greatly differentiated, elongate, erect and sheathing, with or
without a costa, with elongate cells. Setae mostly very short; capsules immersed to +
exserted, erect and symmetric, oblong-ovoid or rarely subglobose or subcylindric,
thin-walled, smooth or sometimes + furrowed when dry; annulus mostly differentiated,
operculum conic and acute, rarely nearly flat and apiculate to short-rostrate; peristome
inserted below the mouth, mostly double, the teeth narrowly lanceolate or lance-
subulate, pale or yellowish, + papillose; segments from a low basal membrane, narrowly
lance-subulate to linear, rarely keeled and perforate, cilia none or sometimes rudi-
mentary. Calyptrae small, conic-mitrate, or rarely split on 1 side and + cucullate,
usually + rough.

Forsstroemia Lindb.

Slender to moderately robust plants in loose, dull or somewhat shiny, green,
yellowish, or brownish tufts. Secondary stems erect, sparsely to freely branched, the
branches sometimes curved. or subcircinate when dry; paraphyllia none. Leaves loosely
erect to imbricate, smooth or somewhat plicate when dry, erect-spreading when moist,
ovate-lanceolate to broadly oblong-ovate, gradually or abruptly short- to rather
long-acuminate; margins reflexed or + revolute, entire or serrulate at the apex; costa

209

single (sometimes weak) and %-%4 the leaf length, sometimes short and double or
lacking in some leaves; cells smooth, thick-walled, rhombic to oblong-fusiform above,
rounded-quadrate to oblate-oblong in many rows at the basal margins. Perichaetial
leaves long and sheathing, short- to long-acuminate. Setae short (less than 5 mm. long),
straight, smooth; capsules immersed to exserted, oblong-ovoid or oblong-cylindric,
smooth; annulus narrow and fragmenting or none; operculum conic, acuminate to
rostrate; stomata none; peristome teeth narrowly lanceolate, transparent, white to
yellowish or reddish, smooth or finely papillose above, sometimes perforate along the
middle; endostome lacking to rudimentary. Spores spherical. Calyptrae cucullate,
usually + hairy.—Named for Johan Eric Forsstr6m, a Swedish pastor who traveled in
Fennoscandia with Wahlenberg and later sent mosses to Swartz from St. Barthélemy
and elsewhere in the West Indes.

Forsstroemia trichomitria (Hedw.) Lindb.—Relatively robust plants in soft,
whitish-green to pale-yellowish or brownish, slightly shiny tufts. Secondary stems
subpinnately branched; branches straight, frequently stoloniform-attenuate at their tips.
Leaves erect and somewhat plicate when dry, erect-spreading when moist, 1.2-2 mm.
long, concave, oblong-ovate to oblong-lanceolate, acute or short-acuminate; margins
entire to slightly serrulate at the apex, reflexed; costa very slender and extending above
the leaf middle or shorter and often double or nearly lacking; upper cells oblong-
fusiform (about 3.5:1), somewhat shorter at the apex. Autoicous; perichaetial leaves
long-sheathing, usually reaching the urn, sometimes surpassing it. Setae 1-3.5 mm. long;
capsules immersed to shortly exserted, 1.5-1.8 mm. long, oblong-ovoid; annulus none,
operculum stoutly and obliquely rostrate; peristome teeth whitish, becoming yellow
with age, faintly roughened or nearly smooth; endostome adhering to the teeth,
consisting of a very delicate and irregular, pale membrane with segments sometimes
visible as fragments. Spores 21-33 yu, very finely papillose. Calyptrae sparsely pilose.

reen, Mosses of Florida, Pl. 113. Darlington, Mosses of Michigan, fig. 96. Grout, Mosses
with Hand-lens and Microscope, Pl. 86. Jennings, Mosses of Western Pennsylvania, (ed. 1) Pl. 32,
(ed. 2) Pl. 36 (as Leptodon). Welch, Mosses of Indiana, fig. 241 (as Leptodon).—Fig. 607-610.

On bark of trees (rarely on rock). Eastern Asia; widespread in eastern North ee south
to northeastern Mexico. Known from a single collection from our area “Mackinaw” (C. F. Wheeler,
1890, in herb. Michigan State University) aaa cluded here wit reservation, as the Straits area is
far north of the expected range of the speci

Somewhat resembling a Leucodon, but (among other differences) softer and
paler, with secondary stems straight, spreading from the substrate, and subpinnately
branched.

LEUCODONTACEAE

Plants mostly rather robust and rigid in loose mats, with creeping, apie:
sparsely radiculose or rarely tomentose primary stems and numerous, erect, curv
ascending, or rarely pendent, simple or branched secondary stems without are
paraphyllia none. Leaves crowded in many rows, ovate or ovate-lanceolate, +
acuminate, somewhat decurrent, often concave, unbordered; costa single, double, or
sometimes lacking; cells thick-walled, mostly smooth, rarely unipapillose on 1 or both
surfaces, oval or rhombic to linear-rhomboidal above, elongate below, with subquadrate
and often oblate-oblong cells in many oblique rows at the basal margins. Dioicous;
paraphyses filiform, few; perichaetia terminating short branches; inner perichaetial
leaves elongate and sheathing. Setae very short to elongate; capsules immersed to

210

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Forsstroemia trichomitria. 607. Habit. 608. Leaves. 609. Upper cells of leaf. 610. Pseudo-
paraphyllia.

exserted, mostly erect, symmetric, subglobose to ovoid or subcylindric; annulus mostly
differentiated; operculum conic or obliquely rostrate from a conic base; stomata
mostly lacking; peristome double or the endostome sometimes rudimentary or ap-
parently lacking, the 16 teeth lanceolate or lance-subulate, mostly papillose, rarely +
cross-striolate, unbordered; basal membrane low, segments lacking or rudimentary, or
rarely well developed, narrow and sometimes as long as the teeth, cilia none. Calyptrae

cucullate, naked or sparsely hairy.

Leucodon Schwaegr.

Coarse and often robust plants in dull, dark-green or yellow-brown, rigid tufts.
Secondary stems simple or sparsely branched, usually terete, often solonifem
attenuate. Leaves erect or appressed to subsecund when dry, erect- to wide-spreading
when moist, ovate or ovate-lanceolate, acute or acuminate, mostly plicate; margins
plane to reflexed or revolute, entire or serrulate at the apex; costa lacking; cells
smooth, thick-walled, shortly oblong-rhomboidal above, becoming longer below, yellow
at the insertion, rounded-quadrate and transversely oblong in many rows at the margins
in the lower half or more. Perichaetial leaves very long-sheathing, acuminate, with all
cells elongate. Setae very short to elongate, smooth; capsules mostly exserted, erect
and symmetric, oblong-cylindric or ellipsoidal, narrow at the mouth, smooth; annulus
none; operculum conic or obliquely rostrate; stomata none; peristome teeth inserted
below the mouth, whitish, becoming yellowish with age, narrowly lanceolate, mostly
perforate, papillose; endostome rudimentary, sometimes apparently lacking. Spores
spherical. Calyptrae often split on 1 side nearly to the base which envelopes the tip of
the seta until the capsule is mature, smooth (or frequently plicate at base), naked.—
Named for the pale or whitish teeth of the peristome.

Slender, flexuose branchlets regularly produced in clusters in the axils of some leaves; leaves

ovate-lanceolate, gradually acuminate, plicate 2. L. brachypus var. andrewsianus
Slender branchlets not produced in leaf axils; leaves broadly oblong-ovate, rather abruptly
short-acuminate, not or obscurely plicate 1. i. julaceus

1. Leucodon julaceus (Hedw.) Sull.—Plants rather small to moderately robust,
dark-green to yellow-brown. Secondary stems short, simple or forked, julaceous and
straight or curved when dry, frequently stoloniform-attenuate; brood-branchlets absent.
Leaves particularly crowded, erect or appressed when dry, wide-spreading when moist,
not at all secund, 1-1.3mm. long, broadly oblong-ovate, rather abruptly short-
acuminate, concave, not or only obscurely plicate when dry; margins plane or
somewhat reflexed, serrulate at the extreme apex; upper cells linear-fusiform, those at
the apex with somewhat bulging walls at the back. Dioicous; perichaetial leaves usually
ending well below the urn, sometimes as long as the seta or slightly longer. Setae
2.5-8mm. long, straight or somewhat curved, dark-red; capsules shortly exserted,
1.1-1.5 mm. long, oblong-ellipsoidal, brown or yellow-brown, red at the mouth, +
constricted below the mouth when dry; operculum obliquely rostrate; peristome teeth
finely perforate; endostome a low, pale, papillose membrane. Spores 33-47 py, varying
in size in the same capsule, densely and finely papillose. Calyptrae tardily deciduous,
split nearly to the base on 1 side and clasping the seta below the urn.—n = 10

en, Mosses of Florida, Pl. 111. Grout, Mosses with Hand-lens and Microscope, fig. 212a,
13d- ae nnings, Mosses of Western Pennsylvania, a. 1) Pl. 32, (ed. 2) Pl. 36. Welch, Mosses of
ea fig. 240.—Fig. 611-612

Typically on the trunks of trees (but sometimes on the bases or on exposed roots),
occasionally on decayed logs and mps or rocks. Wides ee eens in eastern North America; northern
exico. Known from a single local collection, from the Gorge of Carp Creek, Cheboygan Co. ( C.
B. ee 1). The origin of the specimen may be questioned, ee as the normal range of the
species is far southward.

The plants are smaller than those of L. brachypus var. andrewsianus and differ
also in the absence of axillary branchlets and in abruptly short-acuminate leaves with
upper cells somewhat bulging at back.

NO
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Leucodon julaceus. 611. Leaves. 612. Upper cells of leaf.

Leucodon brachypus var. andrewsianus. 613. Habit. 614. Leaves. 615. Upper cells. 616. Basal
calls at margin.

2. Leucodon brachypus Brid. var. andrewsianus Crum & Anders. (L. sciuroides
sensu Amer. authors)—Plants variable in size but usually moderately coarse and robust,
dull, dirty-green to brownish. Secondary stems terete and usually curved when dry;
stoloniform branches often present; brood-branchlets few to numerous in leaf axils.
Leaves crowded, appressed, and sometimes slightly secund when dry, erect-spreading
when moist, plicate wet or dry, 1.4-2.2mm. long, ovate-lanceolate, acuminate,
subentire or slightly serrulate at the extreme apex; upper cells narrowly oblong-linear,
+ flexuose, thick-walled and somewhat pitted, without irregularly thickened walls
bulging at the back of the leaf tip. Inflorescences and sporophytes unknown.—n = 11.

Grout, Mosses with Hand-lens and Microscope, fig. 212b, g-i (as L. sciuroides).—Fig. 613-616.

On the bark of trees in open woods (locally in both hardwood and coniferous swamps),
rarely also on limestone rocks. Widespread in eastern North America from Nova Scotia through the
Great Lakes and south to North Carolina.

CHEBOYGAN CO.~—Reeses Bog, Hermits Bog, Gorge of Carp Creek, Iron Bridge, Smiths
Bog, Livingston Bog, Colonial Point, Riggsville, Mud Lake, Mill Creek, Mud Creek (at Black Lake).
EMMET CO.-—Levering, Larks Lake, Maple River, Middle Village, Five Mile Creek. MACKINAC
CO.-—Bois Blanc Island, Mackinac Island, St. Martins Point, Gros Cap, Peggley Lake, Caffey Corner,
Hendricks, Ozark. PRESQUE ISLE CO.—Rainy River Falls, Rainy Lake.

213

The clusters of branchlets in the leaf axils can always be found somewhere in a
collection of this common moss so long referred to L. sciuroides (Hedw.) Schwaegr.,
an Old World species with similar branchlets but otherwise quite unlike the American
plants. The branches curve upward when dry in the manner of a Pylaisiella.

FONTINALACEAE

Plants elongate, slender to robust, aquatic or submerged at least at high water,
mostly dull and dark, irregularly to subpinnately branched, prostrate and trailing from
a single point of attachment, radiculose only at base and usually denuded of leaves and
branches for some distance above the base. Leaves in 3 rows and sometimes clearly
ranked, erect, erect-spreading, or sometimes falcate-secund, little changed on drying,
broadly ovate to narrowly lanceolate and blunt or acute to acuminate or long-
subulate, keeled or concave and often cucullate at the apex or nearly flat, entire or +
toothed at the apex, often + decurrent, sometimes auriculate; costa lacking or single
and percurrent to long-excurrent; upper cells oblong-hexagonal to linear, smooth; lower
cells shorter, broader and often pitted and colored, the alar cells sometimes clearly
differentiated. Perichaetia at the end of short branches; perichaetial leaves greatly
differentiated, sheathing. Setae very short to elongate, straight; capsules immersed to
exserted, erect and symmetric, ovoid to cylindric, smooth; annulus and stomata none;
operculum bluntly conic to rostrate; peristome double, the teeth sometimes shorter
than the endostome, 16, narrowly lance-acuminate, entire or perforate to gaping along
the middle, sometimes united in pairs at the tips, mostly coarsely papillose; endostome
without a basal membrane, the segments filiform, mostly united at least above by
lateral projections to form a keeled, latticed cone open at the apex, rarely free and
nodose to appendiculate. Calyptrae conic-mitrate or cucullate.

Leaves long and slender, acuminate to setaceous, + falcate-secund, keeled, costate; a cells

scarcely differentiated Dichely ma
Leaves broader, blunt or acute, not secund, flat, concave or keeled, ecostate; “ee cells
somewhat differentiated 1. Fontinalis

1. Fontinalis Hedw.

Plants submerged and trailing, freely and usually irregularly branched above a
denuded base, slender and filiform to robust, variously colored, usually dull and dark.
Stem and branch leaves similar (except sometimes in size), in 3 rows (sometimes
obviously ranked), erect or erect-spreading, not much altered on drying, broadly ovate ~
to lanceolate, acute or sometimes blunt at the apex, sometimes + toothed at the tip,
keeled, concave, or nearly flat, sometimes decurrent and often + auriculate; costa
lacking: cells oblong- to linear- rhomboidal, smooth; alar cells + differentiated. Mostly
dioicous; perichaetial leaves large, broadly oblong-ovate, blunt or rounded at apex or
sometimes broadly short-pointed. Setae very short; capsules immersed to emergent;
operculum conic; peristome double, the 16 teeth about as long as the endostome,
narrowly lanceolate, flat, often joined in pairs at the apex, sometimes perforate or cleft
along the median line, red, smooth or papillose, the segments filiform and usually
joined (at least above) by lateral appendages into a + plicate, latticed cone open at the
top, rarely free and appendiculate. Spores spherical. Calyptrae mitrate, long-rostrate,
scarcely reaching below the operculum, smooth, naked.—The name refers to springs or
“fountains.” (Our local stream, Fontinalis Run, is named not for the moss but for the
Brook Trout, Salvelinus fontinalis.) Surely the most difficult genus of mosses,
Fontinalis, sometimes called WATER MOSS or BROOK MOSS, is usually found in
running water. The leaves are sometimes keeled and arranged in three distinct rows. In

214

a pre-Linnaean publication (but not the famous Historia Muscorum), Dillenius gave one
of these, now known as F. antipyretica Hedw., the name Selago aquatica because of a
resemblance to our modern concept of Lycopodium. | have seen F. antipyretica fo.
robusta (Card.) Card. (F. gigantea Sull. ex Sull. & Lesq.) in dry, stranded masses with
dark-green, rigid, keeled leaves in obvious rows and looking much like a clump of the
Crow Berry, Empetrum nigrum, in dry arctic tundra. We have none of the species with
keeled and ranked leaves locally. The most common of them, F. antipyretica, was
so-named by Linnaeus because of use by Swedish peasants in caulking between the
chimney and the walls of houses in a vain hope of preventing fires.

It is commonly assumed that bryophytes are simple plants, evolved from aquatic
green algae and barely eking out an existence on land. Some of them may be derived
directly from the algae, but the division Bryophyta is no doubt polyphyletic in origin,
and the Musci at least seem to be derived by reduction from primitive vascular plants
already adapted to terrestrial living and the aquatic species secondarily derived from
terrestrial origins.

quatic mosses are actually few and exceptional and they are highly evolved
along widely divergent phylogenetic lines. All of them are related to species or genera
obviously well adapted to life on land. The well-developed peristomes of Fontinalis and
most other truly aquatic mosses, together with the vestigial stomata prevalent
throughout the class, give evidence of reduction from a terrestrial stock. The vascular
tissue of Polytrichum gametophytes, obviously xylem and phloem in form, arrange-
ment, and function, indicates a genetic potential, possessed but not used by the diploid
sporophyte. I believe the mosses and the most primitive pteridophytes were derived
from the same source, from a terrestrial plant with vascular tissue, swimming sperms,
well-defined gametophytes, attached, unbranched sporophytes, no roots or leaves, and
homospory

Leaves somewhat concave, narrowly lanceolate and narrowly acuminate 1. F. disticha
Leaves nearly flat, ovate and broadly pointed to ovate-lanceolate and narrowly acuminate
2. F. hypnoides

1. Fontinalis disticha Hook. & Wils. ex Drumm.—Slender plants about 15 cm.
long, in rather shiny, yellowish or brownish masses. Stem leaves loosely erect or
erect-spreading, slightly concave, 4-5 mm. long, narrowly lanceolate, gradually long-
acuminate, acute, slightly decurrent; cells linear-rhomboidal. Branch leaves similar,
loosely erect, imbricate at branch tips. Dioicous. Setae virtually lacking; capsules
immersed or slightly emergent, cylindric or oblong-cylindric, 1.7-3 mm. long;
operculum large, high-conic, acute; peristome inserted somewhat below the mouth,
granulose, the teeth not or only occasionally perforate at base, the segments strongly
appendiculate throughout but joined in a trellis only above. Spores 13-22 y, finely
papillose.

en, Mosses of Florida, Pl. 118 a7 n Grout, Moss Flora of North America 3: Pl. 75
(figs. ic 4) Welch, Mosses of Indiana, fig. 2

Attached to rocks or bases and roots of trees and a and submerged in pools or streams
(probably in stagnant or sluggish water), eastern United Sta

CHEBOYGAN CO.-~—Pigeon River.

2. Fontinalis hypnoides C. J. Hartm.—Rather slender to medium-sized plants in
loose, soft, usually somewhat shiny, yellow-brown, or (less often) bright- to sordid-
green, irregularly branched masses. Stem leaves usually remote, loosely erect or
erect-spreading, slightly concave or nearly flat, somewhat clasping at the insertion,

215

3-4.5 mm. long, ovate and rather broadly acuminate or ovate-lanceolate and longer-
pointed, acute, entire or + serrulate at the apex, decurrent; cells long-rhomboidal or
linear-rhomboidal; alar cells somewhat enlarged, short-oblong, sometimes brownish.
Branch leaves similar, 2-3, sometimes 4 mm. long. Dioicous; perichaetial leaves broadly
rounded at apex. Setae about 0.25 mm. long; capsules immersed, about 1.5-2 mm.
long; operculum large, bluntly conic; peristome granulose (the teeth sometimes +
perforate at base), the endostome latticed throughout. [Spores 14-20 u, smooth or
finely papillose, according to Welch.]

Conard, How to Know the Mosses, (ed. 2) fig. 253a-d (as F. duriaei). Grout, Moss Flora of
North America 3: Pl. 77 (as F. nitida, F. tenella, F. duriaei, & F. hypnoides). Welch, Mosses of
Indiana, fig. 250 (as F. duriaei).—Fig. 617-620 (as var. duriaei).

Attached to roots, logs, twigs, or rocks and submerged in shallow, flowing water, less
commonly at the margins of lakes. Europe, Asia, Africa, Brazil; widespread in North America, from
Alaska to Nova Scotia and south to California, Arizona, Texas, Missouri, and Pennsylvania.

CHEBOYGAN CO.—Beavertail Creek, Bessey Creek, Gorge of Carp Creek, Iron Bridge, E.
branch of Maple River, Lancaster Lake, Fontinalis Run, Nigger Creek, Mill Creek, 3 miles north of
Cheboygan, “limestone creek” south of Mackinaw City, Duncan Bay, Myers Creek and Mud Creek
(at Black Lake). EMMET CO.—Maple River, Arnott Lake, between Levering and Carp Lake, Carp
Lake. MACKINAC CO.-—Prentiss Bay. PRESQUE ISLE CO.—Rainy River Falls.

Our material can be referred with no hesitation to F. duriaei Schimp., but I see
no way of sorting it from the variability of F. hypnoides, the differences being at best

Fe See
OS
ae:
@

Ga

=

RS
Sep
===
St

Frontinalis hypnoides var. duriaei, 617. Habit. 618. Leaf. 619. Upper cells of leaf. 620. Alar
cells.

216

rather insignificant. Larger, laxer, softer forms can be referred to var. duriaei (Schimp.)
Husn., especially small forms to var. hypnoides.

ecause surface waters rapidly percolate into our loose sands or into the porous
limey bedrock, we have few permanent brooks and therefore only a poor showing of
Fontinalis. We have at least eight species in the state: F. antipyretica Hedw., F.
disticha Hook. & Wils. ex Drumm., F. dalecarlica BSG, F. hypnoides var. duriaei, F.
neo-mexicana Sull. & Lesq., F. paige ee Sull., F. patula Card., and F. sphagnifolia
(C. M.) Wijk & Marg. Records of two southern species, F. filiformis Sull. & Lesq. ex
Aust. and F. flaccida Ren & Card., should be viewed as phytogeographically improb-
able. It is also unlikely that the western Ff. neo-mexicana actually came from Manistee
County, although the specimen is correctly named and the collector, E. J. Hill, did
indeed gather plants in the county

e following key may aid in identification of species collected elsewhere in

ichigan. For further aid in identification, see Welch’s monograph in Grout’s Moss

Flora of North America (vol. 3) or her world-wide Monograph of the Fontinalaceae.

1. Leaves nae obviously 3-ranked
igh

2. Keel strai F. patula
2. Keel c
3. Leaves oblong-lanceolate, acute; cells linear; perichaetial leaves narrow, a tly
apiculate F, neo-mexicana

3. Leaves ovate-lanceolate or broader, obtuse; cells linear-rhomboidal; perichaetial ete
very broad, obtuse F. antipyretica
1. Leaves not keeled or ranked

. Leaves Keer flat F. hypnoides var. duriaei
4. Leaves + con

5. Branch tips pada because of appressed leaves
6. Margins of leaves reflexed when dry; leaves + erect; branches conspicuously terete
perichaetial leaves broadly acute or site F. dalecarlica
6. Margins not reflexed when dry; leaves generally looser, more spreading except at
branch tips which are less es terete; perichactial leaves obtuse or rounded
F. novae-angliae
5. Branch tips not particularly terete; leaves loosely erect-spreadin
7. Leaf margins broadly involute above, sometimes nearly throughout
F.

. Sphagnifolia (summer form)
7. Leaf margins not involute

8. Leaves narrowly lanceolate, acuminate; upper cells linear-rhomboidal F. disticha
8. Leaves broader, lanceolate to ovate-lanceolate, obtuse; upper cells shorter, rhomboidal
F. sphagnifolia (spring form)

2. Dichelyma Myr.

Plants rather slender but elongate, in loose, rather glossy, yellow-brown,
golden-brown, or sordid-green mats submerged at high water. Stems prostrate and
trailing, irregularly branched. Stem and branch leaves similar, in 3 ranks, mostly secund
and sometimes falcate or circinate, keeled and + conduplicate, lanceolate to linear-
lanceolate, gradually acuminate and sometimes filiform- anes not or slightly decur-
rent; margins serrulate toward the apex; costa subpercu to long-excurrent; cells
long, linear-rhomboidal, smooth, only slightly Enea at the basal angles.
Dioicous; perichaetial leaves long, oblong, acuminate, convolute and + twisted,

emerging from the side of the perichaetium) to exserted, erect and symmetric,
oblong-cylindric, brown or yellow-brown; operculum conic, acute to obliquely rostrate;
exothecial cells hexagonal; peristome teeth inserted somewhat below the mouth,
shorter than the endostome, narrowly lanceolate, perforate along the middle, brown,
papillose, with pre-peristome thickenings at base; endostome consisting of long, flat,

2h]

linear segments joined by lateral appendages only at the apex or throughout to form a
trellis-like cone, brown, papillose. Spores spherical. Calyptrae long, cucullate, eae
the capsule and clasping the tip of the seta until maturity.—This genus was name
reference to a cleft veil, or calyptra.

Leaves lanceolate, acute or obtuse, rather clearly 3-ranked, strongly keeled and conduplicate;
2.

costa subpercurrent Dz pallescens
Leaves linear-lanceolate, filiform-acuminate, only obscurely 3-ranked, keeled but no
obviously conduplicate; costa long-excurrent 1. D. capillaceum

1. Dichelyma capillaceum (With.) Myr.—Plants of moderate size, in yellow-brown
or dark-green, not or somewhat shiny masses. Leaves loosely erect and usually +
secund, indistinctly 3-ranked, keeled but not noticeably conduplicate, 3-7 mm. long,
linear-lanceolate, filiform-acuminate; costa long-excurrent. Perichaetial leaves up to
7.5mm. long. Setae 2-4 mm. long; capsules 1.5-2.2 mm. long, immersed but becoming
laterally emergent, oblong-cylindric; operculum obliquely long-rostrate,; exostome teeth
orange-brown, densely spiculose-papillose; endostome similarly orange-brown and
spiculose-papillose, with long, perforate segments joined only at the tips in a small
trellis. Spores 13-17 wu, very minutely papillose to nearly smooth.

een, Mosses of Florida, Pl. 119 (fig. 5-7). Conard, How to Know the Mosses, (ed. 2) fig.
ut, Mosses with Hand-lens and Microscope, fig. 219, Moe Flora of North America 3: Pl.
79 (figs. °38-42), Welch, Mosses of Indiana, fig. 251. Rig, 621.

oots or bases of trees, shrubs, or stumps, sometimes on fallen twigs, branches, or rocks,
in swamps, submerged at high water. Europe; eastern North America.

CHEBOYGAN CO.-—Carp Creek.

The leaves are long-setaceous and not obviously ranked or strongly keeled. The
plants can be taken, in carelessness, for a Dicranum or a Drepanocladus!

2. Dichelyma pallescens BSG—Plants relatively robust, yellow-brown or green,
somewhat shiny. Leaves rather clearly 3-ranked, falcate-secund, strongly keeled and
conduplicate, 3-4 mm. long, lanceolate and rather broadly acuminate, acute or obtuse;
costa subpercurrent or ending somewhat below the apex. Perichaetial leaves up to
7.5 mm. long. Setae 3-5 mm. long; capsules immersed but becoming laterally emergent,
1.2-2 mm. long, oblong-cylindric; operculum obliquely rostrate; exostome teeth brown,
finely papillose; endostome segments darker-brown, finely papillose, perforate, joined
only at the tips as a small trellis. Spores 11-15, minutely roughened or nearly
smooth.

out, Mosses with Hand-lens and Microscope, fig. 220, Moss Flora of North America 3: PI.
79 (Gee 43-47). Jennings, Mosses of Western Pennsylvania, (ed. 1) Pl. 31, (ed. 2) Pl. 35.—Fig.
6 4.
O ots and bases of trees or sticks and stones in swampy woods, submerged at high water.
Northeastern United States and adjacent parts of Canada.
CHEBOYGAN CO.-—Colonial Point, Black Lake.

s compared with D. capillaceum, the leaves are paler, somewhat glossy, clearly
3-ranked, strongly keeled, and not as slenderly pointed.

Dichelyma falcatum (Hedw.) Myr. occurs elsewhere in the state. Like D.
pallescens it has lanceolate leaves which are strongly keeled-conduplicate and clearly

624

Dichelyma capillaceum. 621. Leaf.
Dichelyma pallescens, 622. Habit. 623. Leaf. 624. Upper cells.
Neckera pennata. 625. Habit. 626. Leaves, 627. Upper cells of leaf.

3-ranked. But the leaves are slenderly acute or even setaceous-pointed with a
percurrent to excurrent costa, and the endostome segments are joined throughout their
lengths to form a conspicuous conical trellis.

NECKERACEAE

Mostly rather robust, attractive plants in loose, often glossy tufts, with filiform,
creeping primary stems and erect to pendent, simple to pinnate secondary stems,
usually without paraphyllia. Leaves crowded, mostly complanate, often undulate, of
many forms, mostly oblong-ovate or elliptic, sometimes cordate, acute to rounded or

219

truncate at apex, frequently + asymmetric; costa mostly single, sometimes weak and
ending well below the apex, rarely double or lacking; cells mostly smooth, rounded-
quadrate, rhombic, or rhomboidal above, elongate or linear below. Perichaetia lateral;
perichaetial leaves differentiated. Setae very short to elongate; capsules immersed or
sometimes exserted, mostly erect, ovoid to cylindric; annulus mostly undifferentiated;
operculum conic, mostly rostrate, often obliquely so; stomata present or lacking;
peristome double, the teeth 16, lanceolate, sometimes with a pre-peristome; basal
membrane + well-developed, segments linear, sometimes variously perforate or cleft,
occasionally rudimentary, cilia usually none. Calyptrae mitrate or cucullate, often
hairy.

Leaves oblong-ovate or ovate-lanceolate, acute or acuminate, undulate wet or dry; capsules

immersed . Neckera
Leaves broadly oblong-lingulate, rounded at apex or occasionally rounded-obtuse and
apiculate, smooth; capsules exserted 2. Homalia

1. Neckera Hedw.

Plants medium-sized or + robust, in loose, green, brownish-green, or yellowish,
somewhat glossy mats, often stoloniferous. Secondary stems erect or spreading to
horizontal, simple to pinnate or bipinnate; branches blunt or attenuate; paraphyllia
usually lacking. Leaves inserted in 8 rows but spreading and complanate, asymmetric,
oblong-lingulate to oblong-ovate, acute to acuminate or rounded to truncate or
emarginate, sometimes blunt and apiculate, mostly undulate, shortly decurrent; margins
entire or serrulate, inflexed on 1 side below; costa single and extending %-% the leaf
length or short and double or lacking; cells smooth, mostly + pitted, rhombic to
rhomboidal at the apex, mostly oblong-linear below, + differentiated at the basal angles
(small and quadrate). Perichaetial leaves long-sheathing, rather narrowly acuminate.
Setae short or elongate; capsules immersed to long-exserted, ellipsoidal to oblong-
cylindric, erect and symmetric; operculum obliquely rostrate from a conic base;
peristome teeth narrowly lanceolate, smooth or papillose, sometimes variously striolate
below; basal membrane of endostome rather low (and sometimes lacking); segments
linear, sometimes slit along the keel, sometimes short and fugacious, cilia none. Spores
spherical. Calyptrae cucullate or rarely mitrate, sometimes hairy._Named for Noel
Martin Joseph de Necker (1730-1793), a French-born botanist of Mannheim who
studied both sexual and asexual reproduction in bryophytes.

Neckera pennata Hedw.—Plants medium-sized to fairly large, in soft, shiny,
light-green to yellow-green mats. Secondary stems horizontal or somewhat pendent,
subpinnately branched; attenuate branches few or none; paraphyllia none. Leaves
undulate, 2-2.5 mm. long, oblong-lanceolate to oblong-ovate, acute or acuminate;
margins serrulate at the apex and indistinctly sinuose-serrulate to the middle or below;
costa short and double or lacking; upper cells oblong-linear, flexuose + thick-walled,
somewhat shorter and oblong-rhombic at the apex, subquadrate and shortly rectangular
in small alar groups. Autoicous; perichaetial leaves usually considerably longer than the
capsules, rarely slightly exceeding the mouth. Setae shorter than the capsules (about
1mm. long or less); capsules immersed, about 1.5mm. long, oblong-ovoid, brown;
annulus and stomata none; operculum obliquely apiculate to short-rostrate from a
conic base; peristome teeth inserted below the mouth, pale, sometimes irregularly
perforate and often cohering at the tips, obscurely cross-striolate at base, smooth or
nearly so above; endostome consisting of short, delicate, irregular, linear segments
mostly adhering to the teeth, the basal membrane apparently lacking. Spores 18-32 y,
densely papillose. Calyptrae small (covering the operculum), cucullate, smooth,
naked.—n = 10.

Conard, How to Know the Mosses, (ed. 2) fig. 300. Darlington, Mosses of Michigan, fig. 98.
rout, Mosses with Hand-lens and Microscope, fig. 215, 216 (as var. oligocarpa). Welch, Mosses of
Indiana, fig. 238.—Fig. 625-627

Typically on the trunks of trees in moist, coniferous woods, occasionally on logs or rocks.
Europe, Asia, Africa, Australia, and New Zealand: Alaska to Labrador, south to Arizona, New
Mexico, the Great Lakes, and Tennessee.

CHEBOYGAN CO.—Reeses Bog, Hermits Bog, Gorge of Carp Creek, Iron Bridge, Grass Bay.
EMMET CO. ~Levering, Pleasantview Swamp, Larks Lake, Cross Village, Good Hart, Cecil Bay, Five

Martins Point, McKays Creek, Prentiss Bay, Gros Ca ap, Cut er, Epoufette, ape Pond.
PRESQUE ISLE CO.—Presque Isle Lighthouse, eee ier Station, Clinton L

Although the plants become faded and shaggy, they can be remarkably beautiful,
recognizable at a glance by the flat branches spreading outward from trunks of trees,
with soft, shiny, complanate, wavy leaves and an abundance of immersed capsules.

2. Homalia (Brid.) BSG

Plants slender or moderately robust in strongly flattened, green, or yellow, shiny
mats. Stems creeping, simple to irregularly or subpinnately branched, often stolon-
iferous; paraphyllia none. Leaves in 8 rows, strongly complanate and appearing
distichous, smooth, oblong-cultiform to nearly circular, blunt or rounded-obtuse to
rounded, shortly decurrent; margins entire or somewhat serrulate (or rarely coarsely
toothed) at apex, inflexed on 1 or both sides at base; costa single, extending to or
beyond the leaf middle, or rarely lacking or very short and double; cells smooth, small
and rounded-hexagonal to rhombic or rarely linear above, becoming + elongate and
usually linear below. Perichaetial leaves short-sheathing, lance-acuminate. Setae
elongate; capsules mostly erect, sometimes inclined and + curved with age, rarely
pendulous, oblong or subcylindric from a narrow base, red-brown, smooth; annulus
well developed; operculum obliquely rostrate from a conic base; peristome teeth
lance-subulate, yellow or brownish and cross-striolate below, hyaline and papillose at
the tips, with well-developed trabeculae; endostome pale-yellowish, smooth or finely
papillose, the basal membrane high, the segments linear-lanceolate, keeled, perforate,
cilia none, rudimentary and fugacious, or rarely well-developed and appendiculate.
Spores spherical. Calyptrae cucullate, mostly naked.—_The name refers to the flattened
stems and branches owing to strongly complanate leaves.

Homalia trichomanoides (Hedw.) BSG—Plants in shiny, green or yellowish, very
flat mats. Stems freely and irregularly branched (rarely producing terete-foliate,
stoloniform branches with small, pointed leaves). Stem and branch leaves similar,
wide-spreading and very strongly complanate, smooth or faintly striolate and often
decurved at the tips when dry, 1-2 mm. long, asymmetric, oblong-lingulate, rounded at
the apex or less often obtusely pointed to apiculate; margins inflexed on 1 side at
base, irregularly serrulate in the upper 1/3 or more, faintly serrulate nearly to the base
on the convex side; costa slender, usually single and extending %-% the leaf length,
occasionally short and double; cells firm-walled, rhombic (1-1.5:1) near the apex,
gradually becoming oblong-linear below, porose near the insertion. Autoicous. Setae
9-11 mm. long, red, smooth; capsules 1.5-2 mm. long, suberect and slightly asymmetric,
narrowly oblong-cylindric, tapered at the neck; annulus large, fragmenting; operculum
long-rostrate; stomata few, in the neck; peristome teeth yellowish, bordered below;
endostome pale and smooth, as long as the teeth, with cilia none or occasionally mie
and oa Spores 14-16 yw, very finely papillose. Calyptrae smooth and naked.—

LT Et

pA

Conard, How to Know the Mosses, (ed. 2) fig. 223 (as H. jamesii). Darlington, Mosses of Michi-
gan, fig. 99. Grout, Mosses with Hand-lens and Microscope, fig. 214.—Fig. 628-630.

A calciphile, on cliffs and boulders (especially at their bases), less commonly on bark at base
of trees. Circumpolar; British Se to Oregon; Arizona; Mexico; Newfoundland to Wisconson,
south to Tennessee and Arkan

CHEBOYGAN CO.-—Grapevine Point (Douglas Lake), Carp Creek. EMMET CO.—Levering,
Larks Lake, Wilderness State Park. MACKINAC CO.—Ozark, Caffey Corner.

A pretty, flat, shiny moss somewhat resembling a leafy liverwort. Many
pleurocarpous mosses, like this one, are complanate-foliate (and often conspicuously
flattened), but the leaves are actually inserted in many rows. In a few genera of
acrocarps, such as Fissidens, Bryoxiphium, Distichium, and Schistostega, the leaves
result from the activities of a 2-sided apical cell and are indeed in two rows. Such
plants have a 1/2 phyllotaxy, ie., only two leaves arise from the circumference of the
stem and are thus spaced at 180°. This arrangement is repeated in a spiral in which
every Other leaf is aligned in the same row. When leaves of each cycle are more
numerous, the spiral arrangement is more obvious, at times conspicuous. Since the
apical cell is normally 3-sided, it might be expected that the leaves would be in three
vertical rows, but in most mosses the segments of the apical cell are asymmetric, the
right-hand side being much thicker then the left, so that as growth proceeds the apical
cell gradually turns counter-clockwise, in some genera more strongly than in others.
This torsion growth tends to shift leaves spirally and distort the original 1/3
arrangement. Kerner von Marilaun (The Natural History of Plants, pp. 396-407. 1904)
pointed out that the position of leaves is usually related to their shape. If the leaves
are obovate and short-stalked, as in most oaks, they are arranged in five rows in a 2/5
phyllotaxy. If they are lanceolate or oval, as in almond or peach trees, they usually
have a 3/8 phyllotaxy. The long, narrow leaves of many willows have a 5/13
phyllotaxy. Similarly, in mosses, the narrower the leaves the more numerous the rows.
The broad leaves of Mniums display a 1/3 arrangement; the linear leaves of Poly-
trichums have 3/8, 5/13, or even more complex phyllotaxies. It is my impression that
the 2/5 phyllotaxy is the most common among mosses. In this arrangement, the leaves
are inserted at distances equal to 2/5 the circumference of the stem. Any five leaves
joined in succession according to age are arranged in a spiral of two revolutions, and
every fifth leaf will occupy corresponding positions (first, sixth, eleventh, etc., or
second, seventh, twelfth, etc.).

Charles Darwin once wrote to Asa Gray that he could spare him from an uneasy
grave by explaining why different plants have leaves spaced in varying spiral arrange-
ments.

A thoughtful discussion of phyllotaxy in evolution is provided by Wagner
(Florida State Univ. Studies 13: 48-71. 1954). He (following Fleischer) divided the
Neckeraceae into two subfamilies: the Neckeroideae, with leaves inserted in eight rows,
and the Homalioideae, with leaves inserted in four rows. However, Touw (Blumea 11:
373-425. 1962) showed that all the Neckeraceae could be interpreted as having an
8-ranked phyllotaxy, sometimes apparently 4-ranked because of complanation, asym-
metry, and differential curvature of leaves. Information on distichous and pseudo-
distichous leaf arrangements provided earlier by van der Wijk (Acta Bot. Neerl. 6:
386-391. 1957) support Touw’s interpretations.

FABRONIACEAE

Small or very small plants in green or yellowish, dull or shiny. mats. Stems
creeping, radiculose in scattered tufts, freely branched; branches ascending; paraphyllia

pee)

usually none. Leaves crowded, appressed when dry, spreading on all sides or rarely
secund when moist, + concave, ovate or lanceolate, mostly long-acuminate, usually not
decurrent, neither bordered nor plicate; costa single and slender, rather short (never
percurrent), rarely lacking; cells mostly prosenchymatous, mostly thin-walled, smooth,
quadrate or oblate-oblong at the basal angles. Perichaetia lateral; perichaetial leaves
rarely sheathing. Setae elongate; capsules erect and symmetric, oblong-cylindric, with a
short, thick neck, often longitudinally wrinkled and sometimes constricted below the
mouth when dry; annulus persistent and apparently lacking or rarely deciduous;
operculum broad and umbonate or conic, sometimes rostrate; exothecial cells lax and
thin-walled, sometimes with very irregular walls; peristome single (either the endostome
or rarely the exostome lacking) or double, the teeth 16 (often united in pairs before
dehiscence), flat, without trabeculae, rarely bordered; endostome, if present, consisting
of subulate segments only or sometimes of a basal membrane and keeled segments.
Calyptra cucullate, usually naked.

Anacamptodon Brid.

Small plants in dense, dark-green or yellowish mats. Stems creeping, somewhat
radiculose, freely and irregularly branched; branches + ascending. Leaves erect and +
homomallous when dry, spreading or subsecund when moist, moderately concave,
gradually acuminate from an ovate or oblong-ovate base; margins erect and entire;
costa single, ending at or above the leaf middle; cells green, oblong-rhombic to shortly
oblong-rhomboidal above, rectangular at base, not much differentiated at the basal
margins. Autoicous; perichaetial leaves somewhat elongate, loosely erect, lance-
acuminate, with a slender costa and elongate cells. Setae elongate, straight, rather
stout, smooth; capsules erect and symmetric, oblong-cylindric, strongly contracted
below the mouth and at the distinct neck when dry; annulus narrow; operculum
convex, stoutly and obliquely apiculate to short-rostrate; stomata superficial, in the
neck; exothecial cells hexagonal, with straight wells; peristome teeth inserted somewhat
below the mouth, strongly recurved when dry, yellow-brown, lanceolate, densely and
finely papillose; endostome consisting of linear, smooth, red-brown segments about
1/2-2/3 as long as the teeth and alternating with them. Spores spherical. Calyptrae
smooth, naked.-The name means that the teeth of the peristome are bent backward.

Anacamptodon splachnoides (Froel. ex Brid.) Brid.—Leaves about 1 mm. long;
costa 1/2-2/3 the leaf length (or sometimes shorter); upper cells oblong-rhombic to
shortly oblong-rhomboidal, about 13 wide and 2.5-4:1. Setae 8-12 mm. long,
yellow-brown or orange-red; capsules 1-2 mm. long, yellow-brown, shiny-orange at the
wide mouth when dry. Spores 9-11 yw, faintly roughened to very finely papillose.

en, Mosses of Florida, Pl. 116. Darlington, Mosses of Michigan, fig. 102. Grout, 7
with Hand: lens and Microscope, fig. 210. Jennings, Mosses of Western Pennsylvania, (ed. 1) P
(ed. 2) Pl. 37. Welch, Mosses of Indiana, fig. 242.—Fig. 631-634

the bark of trees, particularly in the protection of crotches, fissures, or eee rarely
on ne or stumps. Central Europe; widespread (but uncommon) in eastern North Americ

CHEBOYGAN CO.—Colonial Point. EMMET CO.—Pellston Hills, Cross Village. MACKINAC
CO.—Mackinac Island.

The gametophytes are dark-green and nondescript in appearance, somewhat
resembling an Amblystegium on a larger scale. The capsules are erect, and a deep
constriction just below the mouth is made the more conspicuous by closely recurved

Homalia trichomanoides, 628. Habit. 629. Leaves. 630. Cells at leaf apex.

Anacamptodon splachnoides, 631. Habit. 632. Leaves. 633. Cells at leaf tip. 634. Capsules,
wet and dry.

Fabronia ciliaris. 635. Habit. 636. Leaves. 637. Cells of upper portion of leaf.

peristome teeth. The specific name no doubt refers to the recurved peristome, as any
other resemblance to the genus Splachnum seems remote. The “primordial utricle” is a
conspicuous feature of the leaf cells. (On drying the plasma membrane becomes
withdrawn from the cell walls, and even on soaking the cell contents remain collapsed
and give the leaves a microscopic effect of blurred opacity, reminding me of soapy
water.) The KNOTHOLE MOSS grows especially on moist, rotten bark, often indeed in
knotholes.

Other members of the family found as rarities in the southern part of the state
are Fabronia ciliaris (Brid.) Brid. (fig. 635-637) and Schwetschkeopsis fabronia
(Schwaegr.) Broth. (fig. 638-640). Both are very small and grow on bark of trees.

224

LESKEACEAE

Plants slender to fairly robust, in loose or dense, mostly dull and rigid, dark,
green or brownish mats. Stems creeping or sometimes arched or ascending, irregularly
to pinnately branched (sometimes 2-3-pinnate in flat fronds); paraphyllia very
abundant or sparse to lacking. Stem and branch leaves often differentiated, ovate or
ovate-lanceolate, blunt or acute to acuminate; costa single and strong, rarely short and
double or lacking; cells subquadrate or rounded-hexagonal to rhombic, rarely
thomboidal, smooth or variously papillose, rarely mammillose. Perichaetia lateral;
perichaetial leaves differentiated. Setae elongate, mostly smooth; capsules erect and
symmetric or inclined to horizontal and asymmetric, cylindric or oblong-cylindric;
annulus generally present; operculum bluntly conic to rostrate; stomata usually present;
peristome double, various in form, inserted far below the mouth with the endostome
much longer than the exostome (in Rhegmatodon) or inserted mostly at or near the
mouth with the endostome nearly as long as the exostome, the 16 teeth lanceolate,
blunt to subulate, smooth, papillose, or variously striolate, sometimes bordered, mostly
trabeculate; basal membrane of the shee stome low to high, segments well developed
and often keeled or sometimes narrow, irregular, or lacking, cilia rudimentary or
lacking to well developed. Spores one Calyptrae mostly cucullate, usually
naked.—For convenience, the family is presented here in a most inclusive sense,
corresponding to the Theliaceae, Leskeaceae, and Thuidiaceae of some authors. Either
treatment is artificial, arbitrary, and unsatisfactory as a phyletic grouping. A better
arrangement is suggested, as follows:

My tentative concept of the Leskeaceae calls for capsules erect and symmetric

usually well-developed cilia—from a high basal membrane). I would group the North
American genera as follows: LESKEACEAE (Leskea, Leskeella, Lescuraea, Pteri-
gynandrum, Lindbergia, Thelia, Haplohymenium, Anomodon, Herpetineurum) and
THUIDIACEAE § (Pseudoleskeella, Pseudoleskea, Myurella, Heterocladium, Haplo-
cladium, Thuidium, Helodium). | would dispense with the Theliaceae. Whether
Pterigynandrum and Myurella are properly placed is debatable. Rhegmatodon is
excluded, as probably worth familial distinction.

1. Costa short and usually double or none
2. Plants subpinnately branched; stem and branch leaves differentiated; capsules inclined
and asymmetric 5. Heterocladium
2. Plants irregularly branched; stem and branch leaves similar; capsules erect and symmetric
3. Plants freely branched; stems and branches slenderly terete or filiform; peristome teeth

variously striolate below, smooth above; endostome consisting of linear, unequal
segments 4, Pterigynandrum
3. Plants simple or sparsely branched; stems and Siac ate or terete but not
filiform; peristo teeth Senet tbet below, papillos ove; ostome consisting of
a well-developed basal membrane, keeled segments, and short or conan cilia 3. Myurella
1. Costa well developed, single
4. Leaf cells smooth 2. Leskeella

4. Leaf cells obscurely bulging-papillose to conspicuously uni- to pluripapillose
5. Paraphyllia none or very few and inconspicuous; plants irregularly branched
6.Stems differentiated from branches, creeping, filiform, with reduced leaves; leaf
cells pluripapillose 6. Anomodon
6.Stems and branches not much differentiated, with similar leaves; cells obscurely
bulging-papillose to distinctly unipapillose
7. Plants scattered or in loose, thin mats; brood branchlets common in axillary clusters;
leaves abruptly narrowed to a short, pale or yellowish awn-like acumen; cells
distinctly unipapillose 8. Lindbergia

225

7. Plants in dense mats; brood branches none; leaves gradually narrowed to an acute o

T
obtuse apex; cells obscurely bulging on both surfaces (especially at back) 1. Leskea
5. Paraphyllia numerous or abundant; plants pinnate (and usually very obviously so)
8. Paraphyllia filiform, very numerous and felted 11. Helodium

8. Paraphyllia polymorphous, not filiform, not felted
9. Apical cell of branch leaves with 2 or more papillae and truncate; plants regularly
1-3-pinnate and frondose; paraphyllia very a idium
9. Apical cell of branch leaves paneer and sharply pointed; plants 1- -pinnate, not
frondose; paraphyllia fairly num
10, Plants loosely subpinnate; pees not julaceous; leaves oe crowded; leaf cells
with a single, low papilla at back; capsules inclined and asymmetr 9. Haplocladium
10. Plants closely and regularly pinnate; foe eae: leaves eae leaf cells
stoutly unipapillose; capsules erect and symm elia

1. Leskea Hedw.

Rather small plants in dull, dark-green or brownish mats. Stems creeping, sparsely
radiculose, freely to subpinnately branched; branches horizontal or ascending; para-
phyllia few, mae lanceolate. Stem and branch leaves not much differentiated. Leaves
erect or somewhat incurved when dry, erect-spreading or spreading when moist,
somewhat are oblong-ovate to ovate-lanceolate, sometimes biplicate at base;
margins plane or + recurved below, entire or subserrulate above; costa subpercurrent or
ending somewhat below the apex, often flexuose-curved above; upper cells small,
irregularly quadrate-hexagonal, thick-walled, obscure, + unipapillose at back or on both
surfaces; lower cells somewhat longer, those toward the margins aa amee
Autoicous; perichaetial leaves elongate, erect, pale-brown, acuminate. Setae smooth;
capsules erect and symmetric or somewhat curved, cylindric or oblong- Ane

Schwetschkeopsis fabronia. 638. Habit. 639. Leaf. 640. Upper cells of leaf.
skea gracilescens. 641. Habit. 642. Leaves. 643. Upper cells of leaf.

226

annulus narrow; operculum bluntly conic; stomata few, at base of capsule; peristome
teeth incurved when dry, linear-lanceolate, whitish or yellow, densely papillose
throughout or, more often, cross-striolate near the base; endostome erect wet or dry,
pale, papillose, consisting of a low basal membrane, long, linear, somewhat keeled,
ee segments (sometimes rudimentary), and cilia none or rudimentary. Calyptra
cucullate, smooth, naked.-Named for Professor Gottfried Leske of Leipzig.

Leskea gracilescens Hedw.—Branches spreading. Stem leaves 0.6-0.8 mm. long.
Branch leaves usually + crowded, rather rigid and erect or somewhat incurved-erect
when dry, erect-spreading when moist, 0.4-0.5 mm. long, ovate or ovate-lanceolate,
acute or sometimes bluntly pointed, + biplicate at base; margins irregularly revolute in
the lower % or more (especially when dry); costa ending near the apex; upper cells
7-11 u, obscurely bulging on both surfaces, but especially at back. Setae 5-8 mm. long,
yellow-brown, becoming orange-brown to reddish with age; capsules 1.5-2.2 mm. long,
oblong-cylindric, straight (or sometimes somewhat curved before dehiscence); peristome
teeth pale, yellowish or brownish-yellow, cross-striolate at base; segments somewhat
shorter than the teeth. Spores 12-14 yw, very finely papillose.—n = 10+1, 11.

Conard, How to Know the Mosses, 1) fig. 129a-c, (ed. 2) fig. 188a-c. Grout, Moss Flora
of North America 3: Pl. 71B.—Fig. 641-643

Found only once in the Straits area, on a roadside maple four miles west of Van in Emmet
County. It is common in more southern parts of Michigan and occurs in nearby Alpena County.
Widespread in eastern North America.

The species of Leskea are not well marked, and their names have been much
misused: L. polycarpa Hedw., not known from Michigan (though wide-ranging and to
be expected here) has leaves somewhat longer than wide and somewhat oblique and
subsecund at the tips, and the capsules are subcylindric and curved. Leskea obscura
Hedw., which has been found in southernmost Michigan, differs from L. gracilescens in
having broad, concave, non-plicate leaves, rounded or rounded-obtuse at the apex, and
erect at the margins.

Pseudoleskea is somewhat similar to Leskea in appearance but occurs on rock in
upland habitats in montane areas. The paraphyllia are numerous, the inflorescences
dioicous, and the peristomes hypnaceous in structure and ornamentation. In Keweenaw
Co., we have P. patens (Lindb.) Kindb. It has leaf cells isodiametric, with a central
papilla on either surface. In Keweenaw and Ontonagon Counties, we have P, radicosa
(Mitt.) Macoun & Kindb. Its leaf cells are somewhat elongate (2-3:1) with those
toward the leaf tips papillose on the upper surface because of projecting upper ends.
(It is most unusual for such papillae to occur on the upper surface.

2. Leskeella (Limpr.) Loeske

Slender plants in loose or dense, dull, dark-green or brown, rigid mats. Stems
creeping, subpinnately branched; paraphyllia none. Stem and branch leaves somewhat
differentiated. Stem leaves erect or appressed when dry, erect-spreading when moist,
rather abruptly subulate from an ovate-lanceolate base; margins narrowly recurved
below, entire or occasionally slightly serrulate near the tip; costa nearly filling the
subula and percurrent or nearly so; cells small and dark, smooth, thick-walled, rounded
or slightly elongate in the upper median region, usually somewhat elongate in the
subula, oblate-quadrate in numerous rows at the basal angles. Branch leaves shorter,
gradually acuminate or sometimes acute or rarely bluntly acute, with the costa shorter,

2a

ending somewhat below the apex and not filling most of the subula. Dioicous;
perichaetial leaves pale, erect, elongate, slenderly acuminate from a sheathing base.
Setae elongate, smooth; capsules erect and symmetric or slightly curved, oblong-
cylindric, brownish; annulus narrow; operculum stoutly and obliquely short-rostrate
from a convex-conic base; stomata at base of the capsule; peristome teeth inserted near
the mouth, erect, yellowish, lanceolate, acuminate, transversely and obliquely striolate
below, smooth or papillose above; endostome pale, finely papillose, the basal mem-
brane moderately high, segments irregular, fragmentary or filiform, rarely lanceolate
and keeled, cilia none or rudimentary. Calyptrae cucullate, smooth, naked.—A
diminutive of Leskea.

Leskeella nervosa (Brid.) Loeske—Plants nearly always producing dense, axillary
clusters of brood-branchlets (or mere buds). Stem leaves 0.8-1 mm. long, entire or
rarely somewhat serrulate at the apex; costa smooth at back; upper median cells 7-9 uw
wide, rounded or somewhat elongate, those in the acumen about 1.5-2:1. Branch leaves
0.4-0.6(0.8) mm. long, subulate to acute (rarely bluntly acute). Setae 9-15 mm. long;
capsules 1.5-2.5 mm. long; annulus of 2 rows of cells; peristome teeth smooth above;
segments of endostome irregular or rudimentary, the cilia none or delicate and
irregular. Spores 13-17 yu, very finely papillose.

t, Mosses with Hand-lens and Microscope, Pl. 57 (figs. 41-54), as Leskea. Jennings, Mosses
of Weert Pennsylvania, (ed. 1) Pl. 35, (ed. 2) Pl. 39.—Fig. 644-646.

n bark of trees (especially at base) and on rock, often on brush or old logs in dry or fairly
moist, shay places. Circumpolar; Alaska to Labrador, south to Pennsylvania, Michigan, Colorado,
and Arizo

CHEBOYGAN CO.-—South Fishtail Bay and Polypody Bluff (both at Douglas Lake), Gorge
of Carp Creek, Iron Bridge, Beavertail Creek, Colonial Point, Mill Creek. EMMET CO.—Levering,
Maple River, Cross Village, Middle Village, Five Mile Creek, Cecil Bay, Big Stone Creek.
MACKINAC CO.—Mackinac Island, McKays Creek, Gros Cap, Caffey Corner, Ozark. PRESQUE
ISLE CO.—Rainy River Falls, Hoeft State Park, Evergreen Beach

A very common moss resembling Platygyrium repens in the production of ~
branchlets in axillary clusters but distinguished by smaller size, dull, dark-green or
blackish color, and leaves tapered to narrow tips more or less filled by the single costa.

The single local record of Pseudoleskeella tectorum (Funck ex Brid.) Kindb. ex
Broth. (South Fishtail Bay, Douglas Lake, C. B. Arzeni) is geographically and
ecologically unlikely. It is a saxicolous moss of the Rocky Mountains and a few
outlying localities in the East, in the Lake Superior region of Michigan (Keweenaw

fe)

and sometimes extending as much as two-thirds the leaf length. Also, the peristome is
hypnaceous in character.

3. Myurella BSG

Small plants in loose or dense, light- or yellow-green, often grayish or bluish mats
or cushions, rigid when dry. Stems crowded and erect-ascending or looser and creeping,
forked or irregularly branched, often + stoloniform-attenuate, somewhat radiculose in
scattered tufts; branches terete, sometimes julaceous, blunt or tapered; paraphyllia
none. Leaves crowded and imbricate or less crowded and erect- to wide-spreading,
concave, rounded-ovate or broadly ovate, blunt or rounded-obtuse to abruptly

228

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Leskeella nervosa. 644. Habit. 645. Leaves. 646. Upper cells of leaf.

es Julacea, 647. Habit. 648. Leaves. 649. Upper cells at margin of lea

Myurella sibirica, 650. Habit of plant and one branch at a slightly higher eee: 651.
Leaves. 652. Cells near apex of leaf.
apiculate or acuminate; margins erect, + serrulate to strongly dentate; costa short and
usually double, often indistinct or lacking; cells small and firm-or thick-walled, rhombic
or ellipticrhomboidal, smooth or papillose at back, centrally or because of projecting
upper ends, shortly oblong near the insertion and irregularly and inconspicuously
subquadrate toward the basal margins. Perichaetial leaves somewhat elongate, erect
lance-acuminate, toothed, red-brown, ecostate, with linear cells. Setae elongate, slender
and flexuose, smooth; capsules erect and symmetric, oblong-obovoid from a distinct
smooth; annulus well developed; operculum conic or conic-

in the neck; peristome teeth lance-subulate, pale, yellow or

yellow-brown and cross-striolate below, hyaline and papillose above, narrowly
bordered, trabeculate; endostome pale, finely papillose, with a well-developed basal
membrane, keeled segments, and short or elongate, single and paired cilia. Calyptrae
cucullate, smooth, naked.—Myurella suggests a little mouse tail, alluding to the form of
the branches, especially in M. julacea. A small genus of pretty little plants distinctive
because of the terete or even julaceous, fragile stems and branches and their glaucous

neck, yellow-brown,

color.

229

Branches julaceous, blunt; leaves crowded and appressed, rounded-ovate, rounded or
rounded-obtuse, occasionally short-apiculate, serrulate, papillose at back because of
1. M. julacea
Branches terete, tapered; leaves less crowded, erect- or wide-spreading, broadly ovate,
abruptly apiculate to acuminate, dentate, stoutly unipapillose at back of the lumina
. M. sibirica

1. Myurella julacea (Schwaegr.) BSG—Plants light-green, pale, yellowish or green-
ish and glaucous. Branches julaceous, blunt. Leaves crowded, imbricate wet or dry,
faintly striolate when dry, 0.3-0.5 mm. long, rounded-ovate, rounded or rounded-
obtuse and occasionally short-apiculate; margins serrulate all around; costa faint or
lacking; upper cells irregularly rhombic, papillose at back because of projecting upper
ends (or sometimes smooth). Dioicous. Setae 6-15 mm. long, orange-yellow, capsules
1-1.2mm. long; endostome segments perforate; cilia short, stout, single or paired.
Spores 9-11 u, very finely roughened.

Darlington, Mosses of Michigan, fig. 100. Grout, Moss Flora of North America 3: Pl. 50 left.—
Fig. 647-649.

Generally in crevices of shaded, moist cliffs, sometimes on soil or humus, especially at base
of trees or on banks of brooks, apparently preferring calcareous habitats. Circumpolar; Greenland
to Alaska, south to New York, Wisconsin, Colorado, and Oregon.

CHEBOYGAN CO.-—East of Levering, Pine Point (Douglas Lake). Bryants Bog, Mill Creek,
Duncan Bay, Grass Bay, Black Lake. EMMET CO.—Cecil Bay, Big Stone Creek. MACKINAC
CO.—Bois Blanc Island, Bush Bay. PRESQUE ISLE CO.-Hammond Bay, Evergreen Beach, Clinton
Lake.

2. Myurella sibirica (C. M.) Reim.—Plants light- or yellow-green to glaucous.
Branches terete, often somewhat flagellate-attenuate. Leaves usually not crowded,
loosely erect- to wide-spreading, 0.35-0.6 mm. long, broadly ovate, abruptly narrowed
to a long apiculus or a short, slender, flexuose acumen; margins irregularly dentate to
spinulose-dentate all around; costa short and double, often indistinct, frequently 1 fork
longer and extending about 1/3 the leaf length; cells rhombic, each bearing a single
large papilla over the lumen at back. Dioicous. Setae about 10-12 mm. long, becoming
red; capsules about 0.8-1 mm. long; endostome segments not perforate; cilia well
developed, + nodulose, single or paired. Spores 11, smooth or very finely papillose.

Conard, How to Know the Mosses, (ed. 2) fig. 176 (as M. careyana). Grout, Moss Flora of
North America 3: Pl. 51B (as M. careyana). Welch, Mosses of Indiana, fig. 229 (as M. careyana).—
Fig. 650-65 2.

Typically in deeply shaded, moist niches and crevices of cliffs, sometimes on protected soil
of banks or sides of damp logs, often in calcareous habitats. Circumpolar (but apparently rare in
northern Eurasia; Nova Scotia to Minnesota, south to North Carolina and Arkansas.

CHEBOYGAN CO.-—Mill Creek, Black Lake. MACKINAC CO.—Mackinac Island. PRESQUE
ISLE CO.—Evergreen Beach.

4. Pterigynandrum Hedw.

Plants very slender to small or medium-sized, in loose or dense, sometimes shiny,
green, yellowish, or brownish mats. Stems creeping or curved-ascending, freely and
irregularly branched; branches terete and often nearly filiform, frequently curved;
paraphyllia none or very few at the branch insertions. Leaves erect or imbricate and
sometimes + homomallous, concave, elliptic or oblong-ovate, sharply or bluntly acute,
shortly decurrent; margins narrowly reflexed in the lower half, serrulate above; costa

230

short and double; cells mostly rather short, oblong-rhombic to oblong-rhomboidal,
flexuose, generally thick-walled, papillose at back because of thickened and projecting
ends; alar cells small and subquadrate in few to several rows, forming small
inconspicuous groups. Dioicous; perichaetial leaves pale and erect, broadly lanceolate,
gradually acuminate, entire. Setae elongate, smooth; capsules erect and symmetric,
cylindric; annulus well developed; operculum conic-rostrate; stomata in the neck;
peristome teeth rather short, lanceolate, pale, yellow-brown and variously striolate
(obliquely, transversely, and vertically) below, hyaline and smooth above, not trabecu-
late; endostome hyaline, smooth, consisting of stoutly linear segments, unequal, short
or sometimes nearly as long as the teeth. Calyptrae cucullate, smooth, naked._Named
by Hedwig presumably in reference to lateral inflorescences (at the side or in the
“wings’’).

Pterigynandrum filiforme Hedw.—Small, slender plants in low, dull, dark-green,
yellowish, or brownish mats; branches terete or julaceous, often curved, frequently
slender and nearly filiform; brood-bodies common in leaf axils, small, stalked, ovoid,
golden-brown, 2-celled. Stem leaves crowded, erect or imbricate, about 0.6-1 mm. long,
broadly ovate or elliptic, gradually or rather abruptly narrowed to a sharp or rather
blunt, acute point; costae rarely % or more the leaf length; cells oblong-rhombic to
oblong-rhomboidal, thick-walled, coarsely papillose at back; alar cells subquadrate in
few to several rows. Branch leaves similar but smaller. Setae 5-15 mm. long, yellow-
brown to orange; capsules 1-2.5 mm. long, yellow-brown; operculum stoutly long-
rostrate. Spores 11-13 yw, very finely papillose.

Darlington, Mosses of Michigan, fig. 106. Grout, Mosses with Hand-lens and Microscope, fig.
134.—Fig. 653-656.

On dry, shaded rocks or sometimes on bark of exposed roots or bases of trees. Circumpolar;
Alaska to California and Montana, also Quebec and New Hampshire to North Carolina and the
Great Lakes region.

CHEBOYGAN CO.~—Lake bluff near Bryants Bog.

The brood-bodies, so commonly produced in leaf axils, were described and
illustrated by Crum (Bryol. 56: 98-100. 1953). They had been reported earlier from
Switzerland and formed the basis for Wheldon’s var, montanense (Rev. Bryol. 32: 7-8.
1905)

5. Heterocladium BSG

Plants small to moderate in size, in flat, mostly dull, dark-green to yellow-brown
mats. Stems creeping, subpinnately branched, stoloniform at the ends; paraphyllia very
few; pseudoparaphyllia present. Leaves dimorphous, not plicate. Stem leaves erect-
spreading to squarrose, gradually or abruptly narrowed to a slender acumen from a
sheathing, cordate-ovate, narrowly decurrent base; margins plane, serrulate all around:
costa short and indistinct, forked or double, sometimes single; cells elongate through-
out or oblong-linear in the middle of the leaf base and small and hexagonal to rhombic
toward the margins and above, smooth or papillose with 1-several small papillae. Branch
leaves small, broadly ovate, acute to blunt or rounded-obtuse, often imbricate or
secund, with short upper cells. Dioicous; perichaetial leaves pale, erect at base, with
spreading to squarrose acumina. Setae elongate; capsules strongly inclined to hori-
zontal, curved, oblong-cylindric, short-necked, constricted below the mouth when dry;
annulus differentiated; operculum convex-conic, blunt or rostrate; peristome teeth
fused at base, lance-subulate, yellow-brown and cross-striolate below, pale and papillose

231

Pterygynandrum filiforme. 653. Habit. 654. Leaves. 655. Upper cells at margin of leaf. 656.
Brood- ees
rocladium dimorphum. 657. Habit. 658. Stem leaves. 659. Branch leaves. 660. Upper
lea

Het
cells os branch
n0modon eae 661. Habit. 662. Branch leaves. 663. Cells at tip of branch leaf.

above, bordered, trabeculate; endostome pale and finely papillose, with a high basal
membrane, keeled and narrowly perforate segments, and well-developed, nodulose cilia
in groups of 1-3. Calyptrae cucullate, smooth, naked._The name translates as different
branch, referring to the differentiated stem and branch leaves.

Heterocladium dimorphum (Brid.) BSG—Rather small plants in loose or dense,
rigid, dark-green to yellow-brown mats. Stems loosely pinnate. Stem leaves about
0.7-1 mm. long, pale, rather abruptly narrowed to a slender, loosely spreading to
recurved acumen from an erect, cordate-ovate base; costa double, very slender and
often indistinct but usually 1/3-1/2 or slightly more the length of the leaf; cells
thombic to oblong-rhombic, thick-walled, and unipapillose at back because of project-
ing upper ends; cells of the base oblong or oblong-linear, thick-walled, pale,
smooth, those toward the margins small, subquadrate to short-rhombic, and papillose.
Branch leaves smaller, imbricate when dry, wide-spreading when moist, rounded- or
oblong-ovate, acute or (especially toward the branch tips) rounded at the apex,
serrulate nearly all around; costa short and indistinct or up to % the leaf length,
sometimes single; upper and basal marginal cells small, hexagonal to short-rhombic,
thick-walled, unipapillose at back because of projecting upper ends. Setae 13-16 mm.

222

long, flexuose, red-yellow to orange-red, smooth; capsules ].2-2 mm. long; annulus of 2
rows of cells, deciduous in fragments; operculum blunt; stomata few and small, at base
of capsule; cilia of endostome 1-3. Spores 13-15 y, very finely papillose.

Darlington, Mosses of Michigan, fig. 109 (as H. squarrosulum). Grout, Moss Flora of North
America 3: Pl. 48 (figs. 18-40), as H. squarrosulum. Fig. 657660.

On soil, humus, rocks, or bases of trees and stumps in moist woods, particularly on steep
banks of streams. Europe and the Caucasus; British Columbia to Oregon and Montana; Newfound-
land to Michigan and New York; also reported from Greenland.

CHEBOYGAN CO.-—Grapevine Point (Douglas Lake). EMMET CO.—Big Stone Bay.

6. Anomodon Hook. & Tayl.

numerous rows, erect or imbricate or sometimes incurved-contorted when dry, erect to
wide-spreading when moist, rarely subsecund, lingulate, lanceolate, or lance-acuminate
from a broad, oblong or ovate, generally broadly decurrent base, obtuse or rounded to
acute or acuminate; margins mostly plane and entire or nearly so; costa strong,
flexuose, yellow, mostly ending well below the apex; cells mostly small, hexagonal,
thin-walled, green, obscure, and densely pluripapillose to the base (but rarely coarsely
unipapillose or longer, rhombic, and smooth throughout); cells at middle of insertion
oblong, incrassate, smooth, and pellucid. Dioicous; perichaetial leaves moderately

mentary or none. Calyptrae cucullate, smooth (or rarely papillose above), naked.—The
name Anomodon signifies an abnormal peristome. It was based on a faulty interpreta-
tion of the segments of the endostome as arising between the teeth of the exostome,
rather than from an inner peristomial layer as in Bridel’s concept of Neckera (and,
indeed, in all mosses with a double peristome).

1. Leaves ending in a short or long, hyaline hair-point; margins revolute 1. A. rostratus
1. Leaves not ending in a hair-point; margins plane
2. Secondary stems spreading, freely branched, with many branches conspicuously tapered
to stoloniform-attenuate; leaves acute and mostly apiculate 2. A. attenuatus
2. Secondary stems ascending, less freely branched, with branches often slender but not
particularly attenuate; leaves broadly pointed, obtuse, rounded-obtuse, or broadly
rounded

3. Leaves erect or imbricate and scarcely contorted when dry 3. A. minor
3. Leaves erect and incurved-contorted when dry
4. Leaves scarcely tapered from the shoulders to the rounded apex, rounded-auriculate at
Ss _ A, rugelii
4. Leaves tapered from the shoulders to the narrow, obtuse or rounded-obtuse apex,
broadly decurrent 5. A. viticulosus

233

1. Anomodon rostratus (Hedw.) Schimp.—Plants in yellowish, brownish-yellow,
or sometimes dark-green, dense mats. Secondary stems and branches crowded and
imbricate when dry, erect or erect-spreading when moist, ovate, acuminate, ending in a
hyaline hair-point of varying length. margins usually revolute to the base of the
acumen, papillose-crenulate; costa ending well below the apex. Setae (5)10-13 mm.
long, red-brown, becoming blackish; capsules 1-1.5 mm. long, oval, oblong-cylindric, or
occasionally cylindric, smooth, becoming dark-brown with age; annulus of 2 rows of
cells, deciduous in fragments; operculum 0.8-1.1 mm. long, obliquely rostrate; stomata
at the extreme base of the urn; peristome teeth pale yellowish or yellow-brown,
becoming white, cross-striolate below; endostome segments not perforate, the cilia
none or rudimentary. Spores green, very finely papillose or nearly smooth, 11-16 yu.
Calyptrae smooth.—n = 10+1, 11.

Breen, Mosses of Florida, Pl. 94 (figs. 3-4). Grout, Mosses with Hand-lens and Microscope,
fig. 138. Jennings, Mosses of Western Pennsylvania, (ed. 1) Pl. 34, (ed. 2) Pl. 38. Welch, Mosses of
Indiana, fig. 235.—Fig. 661-663.

On rock, soil, and bark at base of trees or sometimes stumps, in rather moist, shady places;
probably preferring calcareous habitats. Europe; Bermuda, Haiti, Jamaica, Mexico, and Guatemala;
widespread in eastern North America; Arizona.

CHEBOYGAN CO.—Colonial Point, Cheboygan, Mill Creek. EMMET CO.—Cecil Bay, Big
Stone Bay. MACKINAC CO.—Mackinac Island, Hessel, McKays Creek, Bush Bay, Prentiss Bay, Ozark,
Caffey Corner, PRESQUE ISLE CO.—Rainy River Falls, Clinton Lake.

Clearly distinguished from our other species by crowded, terete branches and
ovate-acuminate leaves with hyaline hair-points.

2. Anomodon attenuatus (Hedw.) Htib.—Plants in rather loose, dark-green or
yellowish to brown mats. Secondary stems loosely spreading; branches usually decurved
when dry, tapered or flagelliform-attenuate. Leaves curved and homomallous, loosely
erect when dry, erect-spreading when moist, broadly oblong-lanceolate from an ovate,
broadly decurrent base, acute and usually ending in a pale apiculus; margins plane,
papillose-crenulate all around, often sparsely serrulate near the apiculus; costa ending
well below the apex. Setae 13-27 mm. long; capsules 2-3 mm. long, cylindric, smooth,
light-brown; annulus none; operculum 1.2-1.3 mm. long, obliquely rostrate; stomata at
extreme base of urn; peristome teeth pale-yellow, becoming white, often faintly
cross-striolate below, bordered; endostome segments often narrowly perforate along the
keel, cilia rudimentary or none. Spores very finely papillose, 10-15 u. Calyptrae
smooth.

Breen, Mosses of Florida, Pl. 97 (figs. 1-2). Grout, Mosses with Hand-lens and Microscope,
figs. 135d, 137. Jennings, Mosses of Western Pennsylvania, (ed. 1) Pl. 34, (ed. 2) Pl. 38. Welch,
Mosses of Indiana, fig. 234.—Fig. 664-666.

bark, especially at the base of trees, also on logs, stumps, rocks, and sometimes soil;
probably preferring calcareous habitats. Europe; Asia; Mexico, Guatemala, Cuba, and Jamaica;
Colorado, New Mexico, and Arizona; widespread in eastern North America.

CHEBOYGAN CO.-—Gorge of Carp Creek, Douglas Lake, Fontinalis Run, Bryants Bog, Mud
Lake, Mill Creek, Mud Creek (at Black Lake). EMMET CO.—Maple River, Carp Lake, Larks Lake,
Good Hart, Middle Village, Five Mile Creek, Cecil Bay, Big Stone Bay. MACKINAC CO.-—Bois
Blanc Island, Mackinac Island, McKays Creek, Bush Bay, Prentiss Bay, Gros Cap, Ozark, Caffey
Corner. PRESQUE ISLE CO.—Rainy River Falls, Clinton Lake.

Our most common member of the genus, best recognized by drooping, tapered
branches and broadly short-pointed leaves.

3. Anomodon minor (Hedw.) Fuirnr.—Rather coarse plants in loose, dark- or
glaucous-green to yellowish- or brownish-green mats. Primary stems and stolons with
small leaves with wide-spreading or squarrose tips; secondary stems and branches
erect-ascending. Leaves erect or imbricate and not or only slightly contorted when dry,
somewhat spreading when moist, broadly oblong from an ovate, broadly decurrent base,
rounded or rounded-obtuse at the apex; margins plane, papillose-crenulate; costa ending
well below the apex. Setae 6-12 mm. long, pale-yellow; capsules 1.5-2.3 mm. long, oblong-
cylindric or cylindric, brown, smooth or + furrowed when dry; annulus revoluble; oper-
culum (0.3)0.6-0.9 mm. long, obliquely rostrate; stomata none; peristome teeth white or
pale and yellowish to brownish, smooth at the base; endostome adhering to the teeth,
consisting of a very low, scarcely discernible membrane and rudimentary segments.
Spores brownish, finely papillose or smooth, 11-15 yu. Calyptrae smooth.—n = 11.

een, Mosses of Florida, Pl. 97 (figs. A 6). Conard, How to Know the Mosses, (ed. 2) fig.
185. oan Mosses with Hand-len d Microscope, PI. . Jennings, Mosses of Western
Pennsylvania, (ed. 1) Pl. 34, (ed. 2) Pl. 38. ae Mosses of Indiana, fig. 233.—Fig. 667-668.

n trunks or bases of trees, sometimes on logs, stumps, or rocks; probably preferring
calcareous habitats. Eastern Asia; Mexico; western Texas and Arizona; widespread in eastern North
America

CHEBOYGAN CO.-—Fontinalis Run, Colonial Point, Mud Lake. EMMET CO.—Burt Lake,
Maple River, Carp Lake, ee Pes Five Mile Creek, Cecil Bay, Wilderness State Park.
MACKINAC CO.—Bois Blanc Is]

Most likely to be confused with A. rugelii, this species is distinguished from it
(and all the other species) by broadly lingulate, rounded-obtuse leaf points which are
erect and scarcely incurved or contorted when dry. As Grout said, “Any Anomodon
with any of the leaves acute, apiculate or serrulate at the apex is pretty surely not this
species.’

4. Anomodon rugelii (C. M.) Keissl—Rather coarse plants in dark-green, yellow-
ish or brownish, dense mats; primary stems often producing stolons with small,
wide-spreading or squarrose leaves; secondary stems and branches erect-ascending.
Leaves incurved-contorted when dry, erect-spreading when moist, broadly oblong-
ligulate and scarcely tapered from an ovate, broadly auricled, non-decurrent base,
rounded at the apex; margins plane, papillose-crenulate all around, often irregularly and
sparsely denticulate at the apex; costa ending well below the apex. Perichaetial leaves
erect, pale-green, ligulate-pointed and obtuse from a broader, sheathing base, strongly
costate, cells green and papillose to below the shoulders, margins crenulate-papillose
above, crenate-serrulate toward the insertion: Setae 8-12 mm. long, yellow, becoming
red-yellow with age; capsules 2-2.5mm. long, cylindric, dark-brown, smooth or
irregularly furrowed when dry; annulus of 2(3) rows of cells, remaining attached to the
operculum after dehiscence; operculum 0.7-0.8 mm. long, obliquely rostrate; stomata at
extreme base of the urn; peristome teeth white, smooth at base; endostome segments
adhering to the teeth, irregular and unequal or rudimentary (sometimes 1/3 as long as
the teeth), cilia none. Spores brownish, finely papillose or nearly smooth, 11-16 yu.
Calyptrae smooth.—n = 10+1, 11.

Cc ae = Know the Mosses, (ed. 2) fig. 186d. Grout, Mosses with Hand-lens and
ees fig. 135a-c (as A. apiculatus). Jennings, Mosses of Western Pennsylvania, (ed. 1) Pl. 34
(as A, a. el 9) Pl. 38.—Fig. 669-670.

On the bark of trees and stumps (often at the base) and sometimes on rocks; probably
preferring eee habitats. Europe; Asia; Maine to Ontario and Michigan, south to fen
and Georgia

235

‘ A ye "
NY \(
Anomodon eee se Habit. oe eae? leaves. 666. Cells at tip of branch leaf.

Anomodon minor, 667. Habit. 668. L
Anomodon nen 669. Leaf. 670. ae at basal auricles.
Anomodon viticulosus. 671. Leaf

HEBOYGAN CO.-—Gorge of Carp Creek, Fontinalis Run. EMMET CO.—Pellston, Five Mile
Creek, Levering, Larks Lake, Carp Lake, Big Stone Bay. MACKINAC CO.-—Prentiss Bay, Gros Cap.

Similar to A. minor but with leaves incurved-contorted when dry, often
irregularly denticulate at the apex and somewhat fimbriate-auriculate at the base. A.
viticulosus has longer, more tapered leaf points.

5. Anomodon viticulosus (Hedw.) Hook. & Tayl.—Plants coarse and robust, in
dark-green, yellowish, or brownish, usually dense mats. Primary stems and stolons with
all, wide-spreading to squarrose leaves; secondary stems and branches generally
ascending. Leaves erect and + incurved-contorted when dry, erect-spreading when
moist, long, lance-ligulate from an ovate, broadly decurrent base, tapered to a rather
narrow, obtuse apex; margins plane, crenulate-papillose; costa ending somewhat below
the apex. Setae 8-16mm. long, yellow, becoming + red-yellow with age; capsules
2.4-3 mm. long, cylindric, dark-brown, smooth or occasionally indistinctly furrowed
when dry and empty; annulus of 3 rows of cells, deciduous in fragments; operculum

236

0.8-1 mm. long, obliquely rostrate; stomata none; peristome teeth pale-yellow or white,
smooth at the base; endostome segments 1/3-2/3 as long as the teeth, often unequal,
linear, keeled, not perforate, cilia none or single, stout stubs. Spores brown, finely
papillose, 11-19 uw. Calyptrae smooth.—n = 10+1, 11.

Darlington, Mosses of Michigan, fig. 110. Grout, Mosses with Hand-lens and Microscope, fig.
136.—Fig. 671.

n limestone cliffs, less commonly on trunks, bases, and exposed roots of trees, in mois
shady places, Europe, Asia, northern Africa; Michigan to Nova Scotia, south to Arkansas aa
Tennesse

EMMET CO.—Levering. MACKINAC CO.—Mackinac Island, McKays Creek. (Common in the
sinkholes of Alpena County.

Larger than A. rugelii, with long-tapered, ligulate-lanceolate leaf points.

Not yet found in this part of Michigan, though widespread in much of eastern
North America, is Haplohymenium triste (Ces. ex De Not.) Sull. (fig. 672-675).
Typically on trunks of trees, the plants are Anomodon-like in appearance and
structure. They are slender, irregularly branched, with stem and branch leaves similar,
imbricate when dry, wide-spreading when moist, abruptly narrowed from a broad,
ovate base to a lingulate or lanceolate point which is usually broken off.

7. Thelia Sull.

Plants of medium size, in dense, rigid, dull, green, yellow-green, yellow-brown,
grayish, or bluish mats. Stems elongate and creeping or sometimes more crowded and
ascending, usually radiculose, sometimes densely so, irregularly branched or more often
regularly 1-pinnate; branches ascending, short and blunt, slender and terete or stout
and julaceous; paraphyllia very few to fairly abundant (at least on the stems),
polymorphous, laciniate or ciliate at the margins. Stem and branch leaves similar or
somewhat differentiated. Branch leaves imbricate wet or dry, very concave, deltoid-
ovate, abruptly short- to long-apiculate, + decurrent; margins erect, dentate or
irregularly spinose-dentate above and + ciliate below; costa single, slender, ending at or
above the leaf middle (or very rarely and atypically short and double); cells
firm-walled, rhombic, coarsely unipapillose on the lumina at back nearly to the
insertion (the papillae simple or elaborately branched), somewhat elongate near the
costa at base, subquadrate at the basal angles. Phyllodioicous; the male plants minute,
resting on leaves or tomentum of female plants; perichaetial leaves pale-yellow,
enlarged, erect and sheathing, gradually subulate from an oblong base (with long,
flexuose cilia with spreading to recurved teeth at their margins), with costa weak or
obsolete and cells elongate, those at the apex + papillose. Setae rather short to
elongate, yellow to red-yellow, smooth, flexuose; capsules erect and symmetric,
cylindric to ovoid-cylindric, yellow or brown, smooth or somewhat wrinkled-striate
when dry and empty; annulus none or poorly developed and persistent or sometimes
fragmenting; operculum long, conic-subulate, slightly oblique; stomata at extreme base
of urn; peristome teeth inserted at or somewhat below the mouth on a very low
membrane, white or whitish-yellow, long, narrowly lanceolate, smooth near the base,
densely papillose above, not bordered or trabeculate; endostome pale and _papillose,
consisting of a rather low basal membrane, short, irregular or rudimentary segments,
and no cilia. Calyptrae cucullate, smooth, naked._The name refers to the strong
papillae on the leaf cells.

23),

Thelia hirtella (Hedw.) Sull.—Plants green, yellow-brown, or grayish-yellow. Stems
creeping, densely radiculose, closely 1-pinnate; branches slender and terete or some-
times stouter and julaceous; paraphyllia fairly numerous (especially on the stems).
Stem leaves about 1-1.3 mm. long, strongly ciliate at the margins. Branch leaves about
1 mm. long, usually not particularly crowded, rather abruptly narrowed to a short to
fairly long apiculus or piliform acumen which is usually flexuose or spreading when
dry; margins irregularly dentate all around and usually + spinose-ciliate, especially
toward the base; costa single and %-% the leaf length (or occasionally short and
double); upper cells about 8 wide, about 1.5:1, each bearing a simple, high, curved
papilla at back (rarely an exceptional papilla bluntly bifid). Setae 5-12 mm. long;
capsules 1.5-2.5 mm. long; annulus none or occasionally slightly differentiated in about
2 rows and fragmenting. Spores 12-18 u, nearly smooth to finely roughened.

, Mosses of Florida, fe 98 (figs. 1-6). Conard, How to Know the Mosses, (ed. 2) fig.

179. ea "Moss ses with Hand-lens and Microscope, fig. 139, Moss Flora of North America 3: Pl.

Jennings, Mosses of Were Pennsylvania, (ed. 1) Pl. 34, (ed. 2) Pl. 38. Welch, Mosses of
Indiana, fig. 230.—Fig. 676-680.

Haplohymenium triste. 672. Habit, dry. 673. Portion of plant, moist. 674. Leaves. 675.
Upper cells of leaf.
Thelia hirtella, 676. Habit. 677. Paraphyllia. 678. Stem leaf. 679. Branch leaf. 680. Upper
cells oe branch leaf.
fee brachyptera. 681. Habit, dry. 682. Portion of plant, moist. 683. Leaves. 684.
Cells Ns eee

238

Usually on the bark of hardwoods (occasionally —elsewhere—on junipers), especially on the
bases of trees or on their exposed roots, sometimes on decayed logs or stumps, rarely on soil or
rocks, in rather dry, open woods. Widespread in eastern North America; northeastern Mexico.

CHEBOYGAN CO.—Fontinalis Run.

The high, curved, conic papillae at the back of the leaves scarcely compare in
shock value with the elaborately branched papillae of 7. asprella Sull. and T. lescurii
Sull. The dwarf male plants are not only very small but also few and hard to find.

Thelia asprella has been found in southern Michigan. It has much the same range
from the Great Lakes southward as Thelia lescurii and has been greatly confused
with it. Thelia asprella nearly always grows on bark at base of trees. It has creeping,
radiculose, pinnately branched stems; its branches are slender, terete, and simple; its
leaves, usually not particularly crowded, are ciliate-papillose above, long-ciliate below,
acute or obtuse, and long-apiculate to piliform-apiculate. Thelia lescurii, by contrast,
grows on sandy soil. Its stems are crowded and ascending, not or slightly radiculose,
irregularly branched; its branches short and julaceous, often forked; its leaves crowded,
not or somewhat ciliate-papillose at the margins, obtuse, shortly apiculate.

8. Lindbergia Kindb.

Rather slender plants in loose, dull, rigid, dark-green, yellowish, or brownish
mats. Stems creeping, freely and mostly irregularly branched; paraphyllia few or
lacking. Leaves crowded, appressed when dry, wide-spreading to squarrose when moist,
somewhat concave, ovate or ovate-lanceolate, slenderly short-acuminate, + decurrent,
not plicate; margins erect, entire or rarely indistinctly toothed near the apex; costa
fairly strong, ending well below the leaf apex; cells rounded-hexagonal or rhombic,
with firm or thickened walls, smooth or unipapillose over the lumina, smaller and
uadrate or oblate-rectangular in many rows at the basal margins. Autoicous; peri-
chaetial leaves enlarged, pale, erect, lanceolate- to subulate-acuminate from a sheathing
base, entire or finely toothed, with a short costa and elongate, smooth cells. Setae
elongate; capsules erect and symmetric, sometimes weakly curved, oblong-cylindric,
smaller at the mouth, brown; annulus sometimes differentiated; operculum conic,
blunt; peristome inserted well below the mouth, the teeth lanceolate, blunt, fused at
base, pale or yellow, + papillose, not striolate, with low trabeculae at back below;
endostome a low, finely papillose membrane. Calyptrae cucullate, smooth, naked.—
Named for the great Scandinavian bryologist, Sextus Otto Lindberg.

Lindbergia brachyptera (Mitt.) Kindb.—Paraphyllia none; brood-branchlets com-
mon in axillary clusters. Leaves 0.9-1.4mm. long, ovate or ovate-lanceolate, gradually
or abruptly narrowed to a pale or yellowish acumen; margins entire or faintly serrulate
above; costa 1/2-2/3 the leaf length; upper median cells thick-walled, rounded to
thombic (8-104 wide and 1-2:1, bluntly unipapillose on both — surfaces,
rounded-quadrate and often oblate in many oblique rows at the basal margins; apical
cells oblong, smooth. Setae 6-8 mm. long, yellow-brown, smooth; capsules 1-1.5 mm
long, yellow-brown, smooth; annulus none; operculum convex-conic; stomata in the
neck; peristome teeth yellow, densely papillose; endostome yellowish. Spores 18-23 yp,
very minutely roughened.

gton, Mosses of Michigan, fig. 105. Grout, Moss Flora of North America 3: Pl. 51A (as
L. mn. Welch, Mosses of Indiana, fig. 228.—Fig. 681-684.

239

On trunks of trees, particularly on roadside elms and maples, occasionally on old logs or
fence rails, very rarely on rocks. Caucasus and the Himalayas; Japan; widespread in eastern North
America; Arizona and western Texas.

CHEBOYGAN CO.-Gorge of Carp Creek, Burt Lake, Riggsville, Black Lake. EMMET
CO.—Brutus, Pellston Hills, Larks Lake, between Levering and Cross Village, Cross Village, Sturgeon
Bay. PRESQUE ISLE CO.—Hammond B

The plants generally occur as scattered strands. The dark-green leaves, squarrose
when moist and tipped with short hair-points are distinctive, as are the dense clusters
of short, axillary branchlets and the rounded to rhombic, bluntly unipapillose cells.

9. Haplocladium (C. M.) C. M.

Plants rather small to medium-sized, in loose, dull, yellowish, brownish, or
light-green mats. Stems prostrate, subpinnately branched; branches spreading or ascend-
ing; paraphyllia few to abundant, especially on the stems, polymorphous. Stem and
branch leaves somewhat differentiated, erect to somewhat incurved and sometimes
subsecund when dry, erect-spreading when moist: Stem leaves often biplicate, ovate,
short- to long-acuminate; margins plane or, more often, + revolute below; costa
extending well into the acumen to excurrent, often flexuose above; cells irregularly
thombic, firm-walled, bearing a single, low papilla at back. Branch leaves smaller and
narrower, ending in a unipapillose cell. Autoicous; perichaetial leaves pale, erect,
elongate, narrowly long-acuminate from a lanceolate base. Setae elongate, smooth,
becoming reddish; capsules strongly inclined to horizontal, curved and constricted
below the mouth when dry, oblong-cylindric; annulus well developed; operculum conic,
acute to short-rostrate; stomata at base of the capsule; peristome teeth lance-subulate,
yellow or yellow-brown and cross-striolate below, pale and papillose above, bordered,
trabeculate; endostome pale-yellowish, finely papillose, with a high basal membrane,
keeled and sometimes narrowly perforate segments, and well-developed, nodose or
appendiculate cilia in groups of 2-4. Calyptrae cucullate, smooth, naked.—The name
refers to the branching system, once-pinnate as compared with the bipinnate or even
tripinnate systems of many species of Thuidium.

Haplocladium microphyllum (Hedw.) Broth.—Plants of medium size. Stems and
branches loosely foliate; paraphyllia few to fairly abundant. Stem leaves loosely erect,
frequently flexuose or subsecund at the tips when dry, about 0.55-1.3 mm. long, ovate
or ovate-lanceolate, gradually or abruptly long-acuminate, biplicate at base; margins
revolute below, sinuate-serrulate above; costa subpercurrent. Branch leaves erect and
often + incurved when dry, ovate-lanceolate, acuminate. Setae often flexuose,
15-30 mm. long; capsules 1.5-2 mm. long; operculum apiculate to stoutly short-rostrate;
cilia of endostome appendiculate, in groups of 3-4. Spores 9-11 u, nearly smooth.—n =
10, 10+1, 11

Gro Mosses with Hand-lens and Microscope. Pl. 55 (as Thuidium). Jennings, Mosses of
Western sae area (ed. 1) Pl. 35, (ed. 2) Pl. 39. Welch, Mosses of Indiana, fig. 223 (as
Thuidium).—Fig. 685-690.

On soil, humus, ae other substrata in the shade, often in rather dry, disturbed, brushy
habitats. nee and Asia; Mexico, Central and South America, West Indies; British Columbia to
Nova Scotia, south to Florida and Texas; Arizona

CHEBOYGAN CO.—Fontinalis Run, Colonial Point, Smiths Bog, Douglas Lake. EMMET
CO.—Larks Lake, Middle Village, Five Mile Creek.

240

Haplocladium virginianum (Brid.) Lindb. (fig. 691-692), to be expected in our
area, looks rather more like a Thuidium and can be confused with T. minutulum
(which has pluripapillose leaf cells). As compared with Haplocladium microphyllum, it
has rather rigid, julaceous branches with crowded leaves, more numerous paraphyllia,
and stem leaves not plicate, broadly short-acuminate, with margins plane or nearly so
and irregularly dentate above.

10. Thuidium BSG

Small to fairly robust plants in dull, green, yellowish, or brownish, loose, often
rigid mats. Stems creeping to curved-ascending, regularly 1-3-pinnate and often
frondose; paraphyllia abundant, papillose, often polymorphous. Stem and branch leaves
differentiated: Stem leaves ovate or subcordate, acuminate, narrowed to the base and
somewhat decurrent, generally + biplicate; margins + revolute below, not or somewhat
toothed above; costa ending below the apex or rarely excurrent; cells mostly uniform,
rounded- to oblong-hexagonal, firm- or thick-walled, minutely pluripapillose on 1 or
both surfaces or coarsely unipapillose at back. Leaves of primary branches often rather
similar to stem leaves, those of secondary and tertiary branches much smaller, concave,
with erect margins, mostly ovate, acute or sometimes obtuse or rounded, usuall

Perichaetial leaves + elongate, erect, pale, lanceolate or ovate-lanceolate, long-
acuminate, sometimes ciliate at the margins, the costa ending in the acumen or
subpercurrent, the cells elongate, smooth or somewhat papillose. Setae elongate,

ow
oblong-cylindric, smooth; annulus of 2-3(4) irregular rows of cells, often tardily
deciduous; operculum conic or convex-conic, often obliquely rostrate; stomata in the
neck; peristome teeth yellow or yellow-brown and cross-striolate below, pale and
papillose above; endostome pale, yellowish or brownish, finely papillose, consisting of a
high basal membrane, keeled and narrowly perforate segments, and + nodulose cilia in
groups of 2-4 (rarely rudimentary or lacking). Calyptrae cucullate, smooth, naked (or
rarely sparsely hairy)._Named because of a resemblance in feathery, frondose branch-

ing (of some of the species) to Thuja, the arbor vitae or northern white cedar.
Commonly called the FERN MOSSES.

if peas a dioicous plants; leaf cells stoutly unipapillose
2 s subere -pinnate 3. T. abietinum
2; a oe or curved-ascending, 2-3-pinna
3. Stem leaves incurved at base, wide- bette Hs or recurved at the tips, with margins plane
or nearly so and costa almost filling the acumen; paraphyllia with papillae at the upper
ends of the cells; operculum hehe aie 4. es aul
3. Stem leaves erect, with margins revolute and costa ending below the apex and n
filling the acumen; paraphyllia with papillae not restricted to cell ends; se te
rostrate
4. Stem | ie ending in a capillary point; inner perichaetial leaves ciliate 5.7. delicatulum
. Stem lea ending in a capillary point formed by 2-7 or more uniseriate cells;
pericha oan oe not ciliate (or only rarely + ce Sa. T. delicatulum var. radicans
1. Small, autoicous plants; leaf cells minutely pluripapillos
5. Plants 1-pinnate; branches terete when dry; leaves cabetas when dry, erect-spreading
when moist; leaf cells papillose at bac . Scitum
5. Plants 1- or more commonly 2-pinnate; branches not terete; leaves incurved- catenulate
when dry, wide-spreading when moist; leaf cells papillose on both surfaces 1. 7. minutulum

1. Thuidium minutulum (Hedw.) BSG—Small, dark-green plants. Stems 2-4 cm.
long, creeping, 1-2-pinnate; paraphyllia numerous on stems and primary branches, few
on secondary branches, simple, 3(or rarely as many as 6) cells long. Stem leaves erect

oe }J 696 ss 607-4 Min ~ 700

Haplocl
tip of eae leaf. 689. Upper cells of eee Tee 69 araphyllia.
Haplocladium tite 691. Stem leaves. 692. Branch leaves

Thuidium minutulum, 693. Stem leaves. 694. Branch leaves. 695. Cells at apex of branch
leaf. a Paraphyllia

yu
nuidium scitum. 697. Stem leaves. 698. Branch leaves. 699. Upper cells at margin of leaf.
700. Serene

or somewhat incurved when dry, wide-spreading or squarrose when moist,
0.3-0.55 mm. long, usually gradually acuminate from an oblong-ovate, obscurely

firm-walled, pluripapillose on both surfaces; costa 2/3-4/5 the leaf length, sometimes
vanishing near the apex. Leaves of primary branches incurved-catenulate when dry,
wide-spreading when moist, ovate, short-acuminate with 1-2 cells often projecting at
iis apex as a hyaline, pluripapillose apiculus; costa 2/3-4/5 the leaf length. Leaves of

econdary branches about 0.2-0.25 mm. long, ovate or oblong-ovate, broadly acute or
a ater blunt, particularly at the ae of branches; costa 1/2-7/ 10 the leaf length;

cells 6-8 u, the terminal cells pluripapillose, not projecting. Autoicous; perichaetial

242

leaves up to 1.4mm. long, entire or somewhat irregular at the margins above. Setae
7-21 mm. long, red, smooth; capsules 0.7-1.5 mm. long, oblong or oblong-ovoid, curved
and asymmetric, inclined to horizontal; operculum long-rostrate, usually about | mm.
long; cilia of endostome nodulose, in 1’s-3’s. Spores 9-12“, smooth or indistinctly
roughened.—n = 9+2, 11.

een, Mosses of Florida, Pl. 101 (figs. 5-8). Grout, Mosses with Hand-lens and Microscope,
fig. ions 123a, 128, Moss Flora of North America 3: Pl. 37 upper left. Jennings, Mosses of
Western Pennsylvania, (ed. 1) Pl. 35, (ed. 2) Pl. 39. Welch, Mosses of Indiana, fig. 221.—Fig
693-696.

On decayed wood or on rocks and soil, often on exposed roots or bases of trees in moist
woods. Europe, oo Bermuda; widespread in eastern North America; Middle and South
America; West Indie

CHEBOYGAN CO.~—Reeses Bog, Hermits Bog, Burt Lake, Gorge of Carp Creek, Carp Creek,
Black Lake. PRESQUE ISLE CO.—Clinton Lake.

The distinguishing features of 7. minutulum include small size, 1-2-pinnate
branching, leaves incurved when dry, and pluripapillose leaf cells.

A similar but much smaller moss not yet found in Michigan but to be expected
on moist, shaded limestone rocks and pebbles especially in stream valleys is T.
pygmaeum BSG, which is best determined by papillae on stems and_ branches.
Haplocladium virginianum (Brid.) Broth., not yet found in our range, can be mistaken
for Thuidium minutulum. In case of doubt, the number of papillae on leaf cells is
diagnostic, single in Haplocladium, pluripapillose in Thuidium minutulum. (The branch
leaves ending in more than one point is a generic character of Thuidium which should
also be helpful.)

2. Thuidium scitum (P.-Beauv.) Aust.—Rather small, rigid, dark-green, yellow-
brown, or brown plants. Stems creeping, 1-pinnate; branches short, terete; paraphyllia
abundant on stems and branches, linear to ovate, often branched. Stem leaves erect
when dry, erect-spreading when moist, broadly ovate, gradually or abruptly narrowed
to a slender or filiform acumen, concave, not or moderately biplicate; margins reflexed
or loosely revolute below; costa vanishing in the base of the acumen; upper cells
irregularly rounded-hexagonal, firm-walled, about 6 u wide, pluripapillose at back; cells
near costa at base oblong. Branch leaves appressed when dry, wide-spreading when
moist, small, ovate, acute. Autoicous. Setae 9-19 mm. long, yellow or red-yellow
smooth, flexuose; capsules 1.2-2 mm. long, suberect to horizontal, oblong- cylindric,
asymmetric, constricted below the mouth and at the short neck when dry; operculum
stoutly and obliquely rostrate; cilia of endostome in 2’s and 3’s. Spores 10-13 y, finely
papillose.—n = 11.

Conard, How to Know the Mosses, (ed. 2) fig. 192e. Grout, Mosses with Hand-lens and
Microscope, figs. 126a, 133b, c, Moss Flora of North America 3: Pl. 44B. Jennings, Mosses of
Western Pennsylvania, (ed. 1) Pl. 35 & (ed. 2) Pl. 39 (as Rauia).—Fig. 698-700.

On bark, especially at the base of trees, occasionally on logs or stumps or rocks. Widespread
in eastern North America, New Brunswick to Michigan, south to Missouri and North Carolina.

CHEBOYGAN CO.—Colonial Point. EMMET CO.—Pellston Hills, Carp Lake, Cross Village,
Cecil Bay, Wilderness State Park. MACKINAC CO.—Gros Cap. PRESQUE ISLE CO.—Hoeft State
Park.

The small, rigid, once-pinnate plants with branch leaves appressed when dry are
suggestive of 7. abietinum. Size alone will distinguish them, but a more basic

243

difference is the pluripapillose leaf cells of 7. scituwm contrasting with the unipapillose
cells of 7. abietinum. Thuidium minutulum is 1-2-pinnate, with leaves incurved-
catenulate when dry.

3. Thuidium abietinum (Hedw.) BSG—Relatively robust, rigid, dark-green,
yellowish, or dark-brown plants. Stems up to 12 cm. high, erect-ascending, 1-pinnate;
branches short, + unequal, terete, tapered; paraphyllia abundant on stem and branches,
linear and simple or branched or lanceolate. Stem leaves erect when dry, erect-spread-
ing when moist, 1.2-1.8 mm. long, ovate, broadly acuminate, plicate, orange-yellow at
the insertion; margins irregularly revolute below the middle and sometimes above it as
well; costa 3/4 or more the leaf length; cells rhombic or irregularly oblong-hexagonal,
thick-walled, stoutly unipapillose at back (the papillae occasionally bluntly forked).
Branch leaves erect when dry, erect-spreading when moist, 0.6-0.7 mm. long, ovate,
acute; cells rounded-quadrate or rhombic, about 7-11, the terminal cell with 2 or
more papillae. Dioicous; perichaetial leaves up to about 4mm. long, serrulate above.
Setae about 25 mm. long, reddish, flexuose; capsules 2-3 mm. long, cylindric, strongly
curved, inclined; operculum obliquely very short-rostrate; cilia of endostome single and
paired; Spores 9-11 us, very finely roughened.—n = 11.

Conard, How to Know the Mosses, (ed. 2) fig. 192a-d. Grout, Mosses with Hand-lens and
Microscope, figs. 126b, 133d, Moss Flora of North America 3: Pl. 37 lower left. Welch, Mosses of
Indiana, fig. 220.—Fig. 701- 705.

On rock and sandy, gravelly, or turfy soil in dry, usually calcareous habitats, frequently on
old dunes or dry hillsides, in open, coniferous stands, on ledges, at bases of cliffs, or among talus
rocks. eae Greenland to Alaska and south to Arizona, Colorado, Iowa, Indiana, and
Virgin

CHEBOYGAN CO.-—Biological Station Campus, Pine Point and Fairy Island (Douglas Lake),
Mill Creek, EMMET CO.—Cross Village, Cecil Bay, Big Stone Bay. MACKINAC CO.-—Bois Blanc
Island, Mackinac Island, Bush Bay, Prentiss Bay, St. Martins Point, McKays Creek. PRESQUE ISLE
CO.—Mast Point, south of Hoeft State Park

The large, wiry, loosely once-pinnate aspect of the plants is unmistakable in the
field, as is the xerophytic habitat in exposed places, such as dune sands.

4. Thuidium recognitum (Hedw.) Lindb.—Rather robust plants in rigid, light- or
yellow-green to yellow-brown mats. Stems 4-9 cm. long, spreading or arched-ascending,
bipinnate and frondose; paraphyllia very abundant (especially on stems and primary
branches), mostly filiform and branched, also lanceolate, the cells strongly papillose
(mostly at or near their upper ends), the terminal cell usually with 2-3 large, divergent
papillae. Stem leaves + incurved at base with wide-spreading tips when dry, spreading
with reflexed tips when moist, about 1 mm. long, broadly ovate, abruptly acuminate,
distinctly plicate; margins papillose-serrulate, more strongly serrulate in the acumen,
plane (at least when the leaf is removed from the stem) or rarely + revolute below;
costa broadened above, nearly filling the acumen, and disappearing near the apex;
upper cells 8-10 wide (up to 244 long), rhombic or oblong-rhombic, thick-walled,
stoutly unipapillose. Leaves of primary branches smaller, with a shorter costa. Leaves
of secondary branches about 0.2 mm. long, erect-spreading wet or dry, ovate, acute;
costa 1/3-2/3 the leaf length; upper cells rhombic, 8-10 X 8-12, incrassate, stoutly
unipapillose, the papillae simple, curved-spinose (usually 8-12 u high), the terminal cell
truncate, pluripapillose. Dioicous; perichaetial leaves up to 4.2 mm. long, with margins
denticulate (and sometimes dentate or notched near the base of the acumen). Setae
20-42 mm. long, reddish, smooth; capsules 2-3.5 mm. long, cylindric, curved and

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aca ee 701. Habit. 702. Stem leaves. 703. Branch leaves. 704. Upper cells of
leaf. 705. Para

Hata ae 706. Stem leaves. 707. Paraphyllia. 708. Perichaetial leaf. 709.
Capsule

horizontal; operculum high-conic or bluntly short-rostrate, about 0.7-1 mm. long; cilia
of endostome nodulose, in 2’s or 3’s. Spores 11-16 u, smooth or nearly so.—n = 11.

Conard, How to Know the Mosses, (ed. 2) fig. 194a-b. Grout, Mosses with Hand-lens and
Microscope, - 127. Jennings, Mosses of Western Pennsylvania, (ed. 1) Pl. 36, (ed. 2) Pl. 40.
elch, Mosses of Indiana, fig. 219.—Fig. 706-709.

On soil, humus or rocks and sometimes also on logs, bases of trees, or exposed roots, in wet
or dry woods, open Se or meadows. Circumpolar; Alaska to Newfoundland, south to Georgia,
Arkansas, and Oklahom

245

EBOYGAN CO.—Reeses Bog, Hermits Bog, Gorge of ais Creek, ae eee Fairy
Island ies Lake), Wolffs Bog, Mud Lake, Grass Bay. EMMET CO.—Leve , Galloway Bog,
Pleasantview Swamp, Stutsmanville, Cross Village, Cecil Bay, Wiens State Park, Co ae Bog.
MACKINAC CO.—Mackinac Island, St. Martins Creek, McKays Creek, Prentiss Bay, Hog Island
Point, Epoufette, Ozark, Caffey Corner, Hendricks. PRESQUE ISLE CO.—Presque Isle Lighthouse,
Evergreen Beach, Hammond Bay, Mast Point, Rainy River Falls, Rainy Lake, Clinton Lake.

See T. delicatulum, below, for means of distinguishing this common species, so
commonly misunderstood.

5. Thuidium delicatulum (Hedw.) BSG—Rather robust, green or yellowish plants.
Stems 3-8cm. long, spreading or arched-ascending, 2-3-pinnate and + frondose;
paraphyllia very abundant (especially on stems and primary branches), polymorphous
but mostly filiform and branched, the papillae not restricted to cell ends, the terminal
cell with several rather small papillae. Stem leaves appressed when dry, erect-spreading
when moist, 0.6-1.4 mm. long, triangular-ovate, gradually or abruptly narrowed to a
broad acumen, plicate; margins papillose-serrulate, revolute from the base to the
acumen; costa ending in the acumen well below the apex; upper cells irregularly
oblong-hexagonal, 6-10 X 8-12 u, moderately aE, coarsely unipapillose at back.
Leaves of primary branches erect-spreading, up to 0.5 mm. long, ovate, acute; costa
1/2-2/3 the leaf length; cells 6-8 X 8-12 y, cee Danese stoutly unipapillose, the
papillae 4-12 u high, curved and often + forked, the apical cell truncate, pluripapillose.
Leaves of secondary branches similar but smaller. Dioicous; perichaetial leaves up to
5mm. long, ciliate below, denticulate above. Setae 15-45 mm. long, reddish, smooth;
capsules 1.8-4mm. long, suberect and moderately curved to horizontal and arcuate,
cylindric, asymmetric; operculum long-rostrate, 0.75-2 mm. long; cilia of endostome in
2’s and 3’s. Spores 12-24 uw, smooth.—n =

Conard, How to Know the Mosses, (ed. 2) figs. 193, 194c. Darlington, Mosses of Michigan,
fig. 107. Grout, Mosses with Hand-lens and Microscope, figs. 122b, 123b, 128, 133a. Jennings,
Mosses of Western Pennsylvania, (ed. 1) Pl. 36, (ed. 2) Pl. 40. Welch, Mosses of Indiana, fig.
217.—Fig. 710-717.

On soil or humus, decayed wood, and rocks, often on exposed roots or bases of trees, in
moist or wet places. Europe and Asia; Alaska and British Columbia; ane Labrador; econatee
in eastern North America; Mexico to South America and the West Indie

CHEBOYGAN CO.—Reeses Bog, Hermits Bog, Gorge of wae Creek, Iron ee oe
Creek), Colonial Point, Vincent Lake, Livingston Bog, Mi reek, Grass Bay. EMM
Pellston, Pleasantview Swamp, Wycamp Lake. MACKINAC CO.—Bois ees Island, See ee
Epoufette. PRESQUE ISLE CO.—Lamprey Experiment Station, Evergreen Beach, Clinton Lake.

Often known as the COMMON FERN MOSS.

Sa. Thuidium delicatulum var. radicans (Kindb.) Crum, Steere & Anders.—Inter-
grading with 7. delicatulum in most respects but distinguished as follows: Stem leaves
1-1.5 mm. long (averaging 1.2 rather than 0.9 mm.), ending in a slender, hyaline point
consisting of 2-8 cells in a single row; perichaetial leaves usually not ciliate (sometimes
a few are present, and rarely they are quite well developed); annulus less well
developed, of 2-3 rows of cells scarcely differentiated from suboral cells.—n = 11

Grout, Moss Flora of North America 3: Pl. 44A (as T. philibertii),—Fig. 718-719.

On soil, humus, and logs in moist or wet woods. Circumpolar; Alaska to British Columbia
and ee Quebec, northern Ontario, and Manitoba south to Iowa, Michigan, and Virginia

Thuidium delicatulum. 710. Habit. 711. Stem leaf. 712. Leaves of primary branches. 713.
Leaves of apna branches. 714. Upper cells of branch leaves. 715. Paraphyllia. 716. Perichaetial
leaf. 717. Cap

CHEBOYGAN CO.—Reeses Bog. EMMET CO.—Levering, Wycamp Lake. MACKINAC CO,—
Mackinac Island. PRESQUE ISLE CO.—Kelsey Pond.

Few species as common, widespread, and distinctive as T. delicatulum have been
greatly confused. A summary of the distinctive features of 7. delicatulum, T.
Baie var. radicans (T. philibertii Limpr.), and T. recognitum follows:

T. delicatulum has stem leaves averaging 0.9mm. long, appressed when dry,
erect-spreading when moist, not plicate, rather gradually acuminate, with the costa
disappearing in the acumen well below the apex and with margins recurved from the
base to the acumen. The paraphyllia bear small papillae along the sides of the cells
(rather than at the ends). The perichaetial leaves are abundantly ciliate. The operculum
is long-rostrate from a conic base and usually 1-2 mm. long.

The var. radicans cannot always be segregated satisfactorily. It has stem leaves
longer peak about 1.2mm. long), more finely acuminate, ending in a slender

247

point consisting of 2-8 hyaline cells in a single row. The perichaetial leaves, usually
thought to be eciliate, actually have a few rather short cilia in most instances and
sometimes have rather numerous and well-developed ones.

Thuidium recognitum has stem leaves which are shortly, broadly, and abruptly
acuminate. They are plicate, with margins plane or nearly so (when removed from the
stems), and the costa appears to be excurrent but actually spreads out and disappears
among elongated cells of the acumen. The tips of the stem leaves spread widely from a
concave and incurved base. The perichaetial leaves lack cilia. The paraphyllia are large
and usually spinose-papillose at or near the ends of cells. The operculum is not rostrate
(or only shortly and bluntly so) and is only about 1 mm. in length or less.

11. Helodium (Sull.) Warnst., nom. cons.

Relatively robust plants in soft tufts or mats, green or yellow-green or sometimes
brownish, sometimes slightly shiny. Stems erect or spreading, freely branched and

en
narrowed at the base, plicate, keeled; margins irregularly revolute nearly to the apex,
entire or minutely toothed near the apex; costa strong, disappearing below the apex or
sometimes subpercurrent; cells pellucid, thin-walled or incrassate, smooth or unipapil-
lose at back (centrally or near the upper end), oblong-hexagonal to sublinear above,
laxly oblong below. Perichaetial leaves erect, pale, plicate, lanceolate, slenderly
long-acuminate. Setae elongate, smooth; capsules inclined to horizontal, + curved,
oblong-cylindric, brown; annulus readily deciduous; operculum conic; peristome broad-
ly lance-subulate, yellow-brown and cross-striolate below, nearly smooth at the tips,
bordered, trabeculate; endostome as long as the teeth, yellowish, nearly smooth, the
basal membrane high, segments lance-subulate, keeled, not or very narrowly perforate,
cilia in 3’s, delicate but well developed. Calyptrae cucullate, smooth, naked.—_The name
refers to the marshy habitats in which the various species are found.

Helodium blandowii (Web. & Mohr) Warnst.—Plants in dense, yellow or light-
green masses 5-12 cm. high. Stems red-brown, erect, closely 1-pinnate; branches nearly
equal, straight, somewhat tapered, wide-spreading. Stem leaves erect or somewhat
erect-spreading, 1.2-1.5 mm. long, broadly ovate, gradually short-acuminate, abruptly
narrowed to the insertion and ciliate near the base; margins serrulate or sinuate-
serrulate above; costa about 4/5 the leaf length; upper cells oblong-fusiform or
oblong-rhomboidal, 3-5:1, firm-walled, strongly unipapillose at back near the upper
ends or nearly centrally. Branch leaves erect or + erect-incurved and moderately
contorted when dry, + erect-spreading when moist, 0.5-0.8 mm. long, narrowly ovate,
acute, ending in a sharp, unipapillose cell. Autoicous. Setae 40-60 mm. long, reddish;
capsules 3-4 mm. long; annulus well developed, of about 2 rows of cells; operculum
conic, blunt to apiculate; stomata in the neck; cilia of endostome in 3’s (or
occasionally paired), nodulose. Spores 11-15 ut, very minutely papillose.—n = 12

arlington, Mosses of Michigan, fig. 108. Grout, Mosses with Hand-lens and Microscope, figs.
123c, 124, 132 (as Thuidium).—Fig. 720-723.

In wooded or open bogs or swamps, frequently in brushy eres Hite streams or ditches.
Grane: Alaska to Greenland and south to Colorado and Ariz Illinois, Ohio, and New
York.

248

719

sly

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4) WW

SS.

= - ar
Se Oh

Thuidium delicatulum vat.
Helodium blandowii. 720.

Paraphy llium
Helodun blandowii var.

branch leaf.
Helodium paludosum. 726. Stem and branch leaves. 727. Upper cells of branch leaf

radicans. 718. Stem leaves. 719. Perichaetial leaf,
Habit. 721. Stem and branch leaves. 722. Upper cells of leaf. 723.

helodioides, 724. Stem and branch leaves. 725. Upper cells of

CHEBOYGAN CO.—Reeses Bog, Hermits Bog, Gorge of Carp Creek, Iron Bridge (Carp
Creek), Douglas Lake, Mud Lake, Little Lake 16. EMMET CO.—Wilderness State Park, Gladys a
Bear Creek Township. MACKINAC CO.—Peggley Lake. PRESQUE

Galloway Bog, Conway Bog,
ISLE CO.—Evergreen Beach, Kelsey Pond.

Helodium blandowii has somewhat the look of a Thuidium. The erect, once-
pinnate stems felted with filiform paraphyllia, the softness of the leaves, and the
swampy habitat aid in recognition. In this, and other species of Helodium, the

249

paraphyllia are found not only on stems and branches but also on the bases of the
costa.

The var. helodioides (Ren. & Card. ex Roll) Crum, Steere & Anders. (fig.
724-725) may be expected in hardwood or tamarack swamps. The plants are somewhat
smaller and somewhat more rigid, with stems spreading and loosely branched or
irregularly pinnate and with leaves costate nearly to the apex.

Helodium paludosum (Sull.) Aust. (fig. 726-727), known from southern Michi-

gan, has longer cells (5-6:1), smooth or weakly papillose at back because of projecting
upper angles.

AMBLYSTEGIACEAE

Plants small to robust, in loose to dense tufts or mats, mostly in wet places,
sometimes submerged. Stems creeping to suberect, irregularly to pinnately branched;
paraphyllia sometimes abundant but mostly lacking. Stem and branch leaves similar, in
many rows, rarely somewhat flattened, imbricate or erect to wide-spreading or
sometimes squarrose-recurved, frequently secund, symmetric, mostly oblong-lanceolate
and acuminate, rarely ovate or elliptic and blunt or even rounded at the apex; costa
single and sometimes very long or short and often forked, double, or lacking; cells
oblong-hexagonal to long-linear, usually smooth, often differentiated at the basal angles
and sometimes abruptly inflated in well-marked alar groups. Perichaetia lateral;
perichaetial leaves differentiated. Setae elongate, red or red-yellow, smooth; capsules
inclined to horizontal, oblong-cylindric, asymmetric and usually strongly curved,
constricted below the mouth when dry and empty, smooth; annulus mostly differ-
entiated; operculum conic or convex-conic, acute or mammillate to apiculate, rarely
short-rostrate; stomata in the neck; peristome double, the 16 teeth lanceolate,
cross-striolate below, papillose above, bordered, trabeculate; endostome with a high
basal membrane, broad, keeled and often perforate segments, and usually well-
developed cilia. Spores spherical. Calyptrae cucullate, naked.—A family characteristic of
wet habitats. The arcuate capsules and yellow-brown peristome teeth contrast with the
shorter capsules and red-brown teeth of the Brachytheciaceae typical of dryer habitats.

1. Paraphyllia present, generally numerous, not restricted to branch bases 1. Cratoneuron
1. Paraphyllia none or few and restricted to branch bases
und
3. Leaves very concave, short-acuminate, imbricate and usually + secund, sometimes
strongly so; costa well developed, often single and double on the same plants 7. Hygrohypnum
3. poe not particularly concave, long-acuminate, not imbricate, falcate-secund through-
ut or occasionally only at ee ai stems and branches; costa single 6. ee
2: Taye erect or spreading, not se
4. Leaves lanceolate or ovate- eee eee variously acuminate
5. Leaves gradually or abruptly narrowed from an ovate base to a channeled acumen,
mostly wide-spreading or squarrose above the base; costa single or short and double

. Campylium
5. ae gradually narrowed from a yes Or ener base to a rather flat
, erect or + spreading from the insertion; costa single
a Plants rather small to fairly ree cells | ineatthompeidal to a aha
3. Leptodictyum

6. Plants small; leaf cells short, oblong-rhombic to shortly rhomboidal
7. Costa not particularly strong, usually ending near the leaf middle, rarely disappear-
ing in the acumen or subpercurrent . Amblystegium
7. Costa very stout, subpercurrent to excurr 4. Hygroamblystegium
4. Leaves broad and elliptic or nearly ae broadly rounded at the apex and
sometimes apiculate

250

8. Costa single and well developed 8. Calliergon
8. Costa short and double or lacki
9. Alar cells clearly differentiated in conspicuous groups 9. Calliergonella
9. Alar cells not differentiated or only a few somewhat inflated in small and
inconspicuous groups 10. Scorpidium

1. Cratoneuron (Sull.) Spruce

Plants medium-sized to fairly robust, in dense, often deep cushions, green,
bluish-green or yellow to brown, usually somewhat radiculose below. Stems creeping
or, more typically, crowded and erect or ascending, regularly or sometimes irregularly
pinnate; paraphyllia present, usually rather abundant, sometimes very few and difficult
to find. Leaves striate or plicate, sometimes erect or erect-spreading but mostly
falcate-secund, slenderly acuminate from a broadly ovate or ovate-lanceolate, some-
times decurrent base; margins plane or slightly recurved near the insertion, serrulate all
around or at the base and sometimes also at the apex; costa single, sometimes very
strong, disappearing in the subula to percurrent; cells + incrassate, smooth or rarely
papillose at back because of thickened cell angles, short and oblong or oblong-
thomboidal to rather long and linear, somewhat larger and inflated in a few rows
across the insertion or abruptly inflated and thin- or thick-walled in well-developed,
concave alar groups. Dioicous; perichaetial leaves elongate, erect, lance-acuminate.
Capsules curved-cylindric; annulus of 2-3 rows of small cells; operculum acute, stoutly
apiculate, or short-rostrate; endostome with well-developed, nodulose cilia._The name
refers to a strong costa.

Cratoneuron filicinum (Hedw.) Spruce—Plants medium-sized to fairly robust,
often rigid, in extensive cushions up to about 12cm. high but usually considerably
less, green, yellowish, or brownish, somewhat radiculose below, radiculose in tufts
above. Stems spreading or more often crowded and erect-ascending, irregularly or more
typically pinnately branched; paraphyllia usually none or few but sometimes fairly
numerous, variously lanceolate, unbranched. Stem leaves erect or erect-spreading,
sometimes secund to falcate-secund, + striate, 1-1.8 mm. long, ovate-lanceolate to
broadly ovate, rather abruptly acuminate, the acumen usually not flexuose and twisted;
margins often somewhat recurved at the extreme base, serrulate nearly all around (the
acumen nearly entire); costa rather strong, percurrent or ending in the acumen
somewhat below the apex, rarely filling the apex; cells smooth, firm-walled, usually
shortly oblong, sometimes oblong-rhomboidal, about 3-6:1, abruptly inflated in
concave, hyaline or yellow-brown, thin- or thick-walled alar groups. Branch leaves
shorter and narrower, usually + falcate-secund. Setae 25-35 mm. long; capsules
2-2.5 mm. long; operculum apiculate to short-rostrate; cilia in 3’s and 4’s. Spores
15-18 u, very finely roughened.—n = 10, 11, 20, 30.

arlington, Mosses of Michigan, fig. 114. Grout, Mosses with Hand-lens and Microscope, Pl.
75. res Mosses of Indiana, fig. 174.—Fig. 728-732.

wet soil or rocks in seepage or overflow from springs or backwater of brooks; apparently
limited to ue ane habitats. Europe, Asia, Africa, Madeira; New Zealand; Guatemala; — a and
British Columbia to Newfoundland, south to California, New Mexico, Texas, and Alabam

CHEBOYGAN CO.—Reeses Bog, Gorge of Carp Sats Iron Bridge, Mill Creek, Grass Lake,
Grass Bay. EMMET CO.—Maple River, Sturgeon Bay Point, Cecil Bay, Alanson, Ramona Beach.
MACKINAC CO.-—Bois Blanc Island, Mackinac Island. PeESOUE ISLE CO.—Hammond Bay,
Evergreen Beach.

Cratoneuron filicinum is recognized by its seepy, calcareous habitat and upright
or decumbent, pinnately branched stems ending in an unbranched extension pointing

251

upward like a long, skinny finger curved at the tip. The paraphyllia are few and usually
difficult to find, but the rhizoids in tufts along the stems are distinctive. The plants are
often harsh and rigid owing to the deposition of calcium carbonate around the older
portions. The other species known from the state, C. commutatum (Hedw.) Roth, is
much more like a Drepanocladus in appearance and structure but has an abundance of
paraphyllia. The epithet commutatum refers to changes effected by deposition of limey
material from the water resulting in a degree of petrification. (Chromosome numbers
reported for C. commutatum are n = 7

2. Campylium (Sull.) Mitt.

Plants small and slender to moderately robust, in green to yellow or golden-
brown, often shiny tufts or mats. Stems creeping or sometimes crowded and
erect-ascending, irregularly to subpinnately branched; paraphyllia none or sometimes
few, at the branch bases. Leaves suberect, ovate or cordate, shortly decurrent at the
base and gradually to rather abruptly narrowed to a long, channeled acumen which is
usually wide-spreading to squarrose-recurved; margins erect, mostly entire; costa single,
slender, and + elongate or short, double, or lacking; cells narrowly prosenchymatous to
linear, smooth, + differentiated in somewhat excavate alar groups (small, quadrate, and
firm-walled or pale, lax, and + inflated). Perichaetial leaves sheathing at base,
lance-subulate, striate. Capsules curved, subcylindric; annulus present; operculum acute
or apiculate; endostome with usually perforate segments and 1-3 well-developed,
nodulose cilia.-The name, signifying curved, may refer to the leaves which are
commonly curved and spreading or possibly to the curved capsules.

1. Costa none or very short and double
2 ae enoee with creeping oe leaves smooth, finely serrulate all around; alar

cells
mall, qua . hispidulum
2 Pint usally eee robust, with erect- "a stems; leaves generally + ‘striolate
n dry,e ; alar cells inflated, subrectan

3. ieee: ee striolate when dry, erect Or a spreading from an erect base, oe

lly
ust . stellatum
3. Leaves not crowd t or eae striolate when dry, wide-spreading from the

a. C. stellatum var. protensum

dle
. Leaves erect-spreading, gradually acuminate; alar cells enlarged and inflate 4.C. polygamum

d
4. Leaves wide-spreading to squarrose, abruptly acuminate; alar cells not or only slightly
inflate

5. Leaves crowded, often ae secund at tips; upper cells linear; alar cells sub
quadrate, firm-walled; we Pu Oy chrysophyilum

5. Leaves remote, not eens acre rt cells gy thomboidal (4-6:1); alar eile shor
oblong, not eve pan Bae autoico 3, a radicale

1. Campylium hispidulum (Brid.) Mitt.—Very slender plants in bright-green or
yellowish mats. Stems creeping, irregularly branched. Leaves squarrose, 0.5-0.6 mm
long (or stem leaves rarely 1mm. long), very concave, ovate, rather abruptly

: i

none; upper cells oblong-rhomboidal; alar cells numerous, subquadrate and shortly
oblong. Autoicous. Setae 8-15 mm. long; capsules 1-2 mm. long, constricted below the
mouth when dry and empty; operculum stoutly and bluntly oblique-apiculate;
exostome teeth brown; endostome pale-brownish, with non-perforate segments and cilia
in groups of 2 and 3. Spores 9-13 uw, very finely roughened.—n = 14.

Conard, How to Know the oe (ed, 2) fig. 256. Grout, Mosses with Hand-lens ae
Microscope, fig. 168. Jennings, Mosses of Western Pena varis, (ed. 1) Pl. 41, (ed. 2) Pl.
Welch, Mosses of Indiana, fig. 175. “Fig. 733-735.

728

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ee 2
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J

Cratoneuron filicinum, 728, eee 729. Stem and branch leaves. 730. Upper cells at margin
of leaf. 731. Alar cells. 732. Paraphy

Campylium aan ee 733. Habit Leaves. 735. Upper cells of leaf.

Campylium chrysophyllum. 736.

Campylium radicale. 737. Leaf. ey yor cells.

On soil, rocks, and logs, usually in rather moist forests; probably a calciphile. Circumpolar;
Saskatchewan to Labrador, south to the Gulf of Mexico; Mexico; reported from northern South
America.

CHEBOYGAN CO.—Douglas Lake, Reeses Bog, Carp Creek, Colonial Point, Mud Lake.
EMMET CO.—Arnott Lake, Galloway Bog, Pellston or Middle Village, Wilderness State Park.
MACKINAC CO.—Bois Blanc Island, Mackinac Island, McKays Creek, Ozark. ea E ISLE
CO.—Hammond Bay, Evergreen Beach, Hoeft State Park, Ocqueoc Falls, Rainy River Falls.

253

Campylium hispidulum is our smallest representative of the genus, recognized by
crowded, cordate leaves abruptly narrowed to a wide-spreading or squarrose, channeled
acumen. The costa is very short and double (or none), in contrast to the single costa
of C. chrysophyllum.

2. Campylium chrysophyllum (Brid.) J. Lange—Plants variable in size but rather
small, in green, yellow-green, or yellow-brown, sometimes + shiny mats. Stems
creeping, freely and irregularly branched. Leaves typically wide-spreading to squarrose,
often somewhat secund at the tips, ovate or lanceolate, rather abruptly narrowed to a
channeled acumen, rounded to the base, usually faintly striolate when dry; margins
entire above, not or slightly sinuate at the base; costa slender, single, 1/2-2/3 the leaf
length; upper cells oblong-linear; alar cells quadrate to short-oblong, not inflated,
firm-walled. Dioicous. Setae 10-27 mm. long, becoming red; capsules 2-3 mm. long;
operculum stoutly oblique-apiculate; exostome teeth yellow-brown; endostome pale-
brownish, finely papillose, with non-perforate segments and cilia in groups of 2-3.
Spores 13-15 y, faintly roughened.—n = 9, 20

Breen, Mosses of Florida, Pl. 69 (figs. 1-3). Darlington, Mosses of Michigan, fig. 115. Grout,
Mosses with Hand-lens and Microscope, fig. 169. Jennings, Mosses of Western Pennsylvania, (ed. 1)
Pl. 41, (ed. 2) Pl. 45. Welch, Mosses of Indiana, fig. 177.—Fig. 736

On rocks, soil, rotten wood, and bases of trees in moist to wet woods. Circumpolar; Mexico,
Guat emer (Gey ter Antilles, northern South America; British Columbia to Newfoundland, south to
Oregon, Arizona, and Colorado; throughout eastern North America.

CHEBOYGAN CO.—Reeses Bog, Hermits Bog, Douglas Lake, Iron Bridge, Lancaster Lake,
Mud Lake, Cheboygan, Mill Creek, Grass Bay. EMMET CO.—Arnott Lake, Maple River, Stutsman-
ville, Cecil Bay, Big Stone Bay, Five Mile Creek. MACKINAC CO.—Mackinac Island, St. Martins
Point, Hessel, Ozark. PRESQUE ISLE CO.—Mast Point, Hammond Bay, Evergreen Beach, Presque
Isle Lighthouse, Ocqueoc Falls, Rainy River Falls, Grand Lake, Clinton Lake.

Campylium chrysophyllum is rather small but considerably stouter than C.
hispidulum, from which it is easily distinguished by its leaves with a single costa. The
leaves are crowded and typically squarrose from a clasping base, but occasionally they
are slightly to strongly secund.

3. Campylium radicale (P.-Beauv.) Grout— ee plants in soft, loose, light-
green or yellowish mats, resembling an Amblystegium in habit. Leaves remote,
1-1.3 mm. long, wide- spreading from the insertion, an abruptly narrowed to a
slender, channeled acumen, decurrent; margins plane, entire or nearly so; costa single,
slender, 1/2-2/3 the leaf length; upper cells rather shortly rhomboidal, 4-6: 1; alar cells
somewhat enlarged, oblong, hyaline. Autoicous. Setae 3-4 cm. long; capsules about
2mm. long; peristome teeth brown; cilia of endostome 1-3. Spores ca. 9-11 yu, finely
roughened,

Breen, Mosses of Florida, Pl. 69 (figs. 8-10). Grout, ane ee ess lens and Microscope,
fig. 175 (8-10), as Hypnum radicale, H. bergenense, & Amblystegium hygrophilum. Jennings,
Mosses of Western Pennsylvania, (ed. 1) Pl. 38 & (ed. 2) Pl. 42 ee abba radicale).—Fig.
737-738.

On decaying leaves, twigs, humus, and mucky soil in oy habitats. Europe; probably
common but usually overlooked in much of eastern North Americ

CHEBOYGAN CO.—Douglas Lake, Livingston Bog, Cheboygan.

254

Campylium radicale is somewhat like an Amblystegium in aspect, because of its
growth in thin, loose mats and its remote leaves widely spreading from the insertion,
but it has channeled leaf tips, a characteristic of Campylium never found in
Amblystegium. Among our species of Campylium, it is rather easily recognized by
remote leaves spreading from the insertion, broad at the base, abruptly acuminate, and
singly costate.

4. Campylium polygamum (BSG) C. Jens.—Plants moderately robust, in loose or
dense, yellow-green to golden-brown, slightly shiny mats. Stems prostrate or ascending,
hcl branched. Leaves erect-spreading or somewhat spreading, 2-2.5 mm. long,
ovate-lanceolate, gradually narrowed to a long, slender, channeled acumen, entire; costa
slender, single and ending at or somewhat above the midleaf (rarely shorter and +
forked); upper cells linear, becoming + thick-walled and sometimes porose with age;
cells at middle of the insertion incrassate and porose, those in the alar regions
short-rectangular, inflated and often golden-brown, in distinct groups. Polygamous.
Setae 2-4 cm. long; capsules about 2.5 mm. long. Spores ca. 16 uy, finely papillose.—n =
11, 20

en, Mosses of Florida, Pl. 69 (figs. 4-7). Grout, Mosses with Hand-lens and Microscope,
fig. an. Een nings, Mosses of Western Pennsylvania, (ed. 1) Pl. 42, (ed. 2) Pl. 46. Welch, Mosses of
Indiana, fig. 178.—Fig. 739.

soil or humus in meadows, swamps, or wooded bogs. Europe (and presumably nearly
cosmopolitan); ae in North America, but scattered from British Columbia to Quebec and
south to the Gul

CHEBOYGAN CO.—Reeses Bog, Burt Lake, Douglas Lake. EMMET CO.—Pellston Hills, Cecil
Bay. MACKINAC CO.—Bois Blanc Island, Peggley Lake, Rexton, PRESQUE ISLE CO.—Mast Point,
Evergreen Beach.

Campylium polygamum resembles C. stellatum, especially in size, but has leaves
less spreading, narrower, and singly costate. As in C. stellatum, the alar cells are rather
well differentiated.

5. Campylium stellatum (Hedw.) C. Jens.—Plants moderately robust, in loose or
dense, yellow, golden, or brownish, + shiny mats. Stems irregularly branched,
ascending. nee crowded, 1.5-3 mm. long, striolate when dry, broad and cordate-ovate
at the base, gradually narrowed to a long, channeled, suberect or more typically widely
spreading acumen, subentire; costa lacking or short, slender, and single or forked;
upper cells linear, + thick-walled, becoming porose with age; alar cells rather
conspicuously differentiated, enlarged, thin-walled, hyaline, short-oblong (becoming
brownish and thicker-walled with age). Dioicous. Setae 2-3.5 cm. long; capsules about
3mm. long. Spores about 16 y, finely papillose.—n = 10, 18+2, 20, 22.

Grout, Mosses with Hand-lens and Microscope, Pl. 77. Jennings, Mosses of Western
Pennsylvania, (ed. 1) Pl. 42, (ed. 2) Pl. 46. Welch, Mosses of Indiana, fig. 176.—Fig. 740-742.

On soil or humus in wet, calcareous habitats, such as rich fens, meadows, roadside ditches,
and seepage around springs. ie Greenland to Alaska and south to Colorado, Ohio, and
Pennsylvania.

CHEBOYGAN CO.—Reeses Bog, Mill Creek, Grass Bay, Indian River. EMMET CO. = Scan
Lake, Wycamp Lake, Cecil Bay, Wilderness State Park, Waugoshance Point, Conway Bog, Ramona
Beach. MACKINAC CO.-—St. Martins Bay, Nunns Creek, St. Martins Point, Epoufette, en
PRESQUE ISLE CO.—Hammond Bay, Evergreen Beach, Clinton Lake.

Campylium polygamum. 739, Lea
Campylium stellatum, 740. Betas of branch. 741. Leaf. 742. Alar cells.

Campylium stellatum is highly pnaapere in of calcareous fens. It is typically
robust, with a compact and tufted manner of growth and a rich, golden color. The
leaves are more or less squarrose-spreading and, Saah dry, longitudinally striolate. As
in C. polygamum, which can be confused with it, C. stellatum has somewhat inflated
alar cells, but it lacks a costa. (The name refers to spreading leaf tips giving the stems
and branches something of a star-like appearance at their tips.

5a. Var. protensum (Brid.) Bryhn ex Grout—Plants rather slender. Leaves not
crowded, not or slightly striolate when dry, wide-spreading to squarrose, mostly from
the insertion, abruptly narrowed from a broad base to a long, slender acumen.—n = 10.

Grout, Moss Flora of North America 3: Pl. 17A (fig. 3).
CHEBOYGAN CO.—Reeses Bog, Grass Bay.

In open, boggy habitats, such as sedge meadows or margins of beach cee Europe, Asia,
and North America, presumably ranging with the species but relatively uncomm

Smaller than the typical variety and somewhat resembling Campylium chryso-
phyllum. When well developed, it also bears a similarity to Rhytidiadelphus loreus,
though constructed on a smaller scale.

256
3. Leptodictyum (Schimp.) Warnst.

Plants slender to fairly robust, in thin, usually dull, dirty-green or yellowish,
sometimes shiny mats, usually in very wet places, sometimes submerged. Stems
creeping or ascending, freely and irregularly or sometimes subpinnately branched. Leaves
soft, mostly widespreading and occasionally loosely complanate, oblong-lanceolate,
acute or acuminate, not channeled above, smooth; margins plane and entire or nearly
so; costa single, slender, ending at or above the leaf middle, rarely subpercurrent; cells
+ elongate, oblong-hexagonal to rhomboidal or long-linear, smooth, mostly thin-walled,
broader and oblong at base, age differentiated at the basal angles. Autoicous;
perichaetial leaves lance-acuminate, + plicate. Capsules curved; annulus differentiated;

filiform, + nodose cilia in 2’s-4’s.-The name refers to the fine network formed by the
outlines of the leaf cells (and is not particularly appropriate).

1. Leaves 2.5-4 mm. long; leaf cells linear-rhomboidal, 8-15:1 1. L. riparium
1. Leaves 2 mm. or less long; leaf cells oblong-rhomboidal, 2-6: 1

2. Leaves broadly short-acuminate 2. L. brevipes
2. Leaves slenderly acuminate 3. L. trichopodium

1. Leptodictyum riparium (Hedw.) Warnst.—Medium-sized to relatively robust,
green, yellowish, or golden to brownish plants in loose, flat mats. Stems creeping;
branches usually horizontal and wide-spreading. Leaves rather distant, wide-spreading
(and often appearing somewhat 2-ranked), 2.5-4.5 mm. long, lanceolate, gradually
long-acuminate; costa %4-% the leaf length; upper cells long, linear-rhomboidal (about
8-15:1), those at the basal angles laxly rectangular. Setae 13-25 mm. long; capsules
2.2-2.5 mm. long. Spores 11-13 yu, finely papillose—n = 9+1, 10, 12, 20, 24, 36, 40.
48, 96 (and experimentally 12, 48, 96).

Conard, How to Know the Mosses, (ed. 2) fig. 225. Darlington, Mosses of Michigan, fig. 111.
Grout, Mosses with Hand-lens and Microscope, fig. 176 (as Amblystegium), Moss Flora of North
America 3: Pl. 15A (figs. 1-6). Jennings, Mosses of Western Pennsylvania, (ed. 1) Pl. 38 & (ed. 2)
Pl. 42 (as ee Welch, Mosses of Indiana, fig. 155, 156 (as fo. longifolium ), 158 (as fo.
obtusum),—Fig. 7

On humus, leaf litter, and twigs and branches on the ground in wet places, commonly in
wooded swamps in shallow depressions eo in spring. Circumpolar; widespread in North
America; Mexico, Guatemala, Cuba, and Hai

CHEBOYGAN CO.-—Burt Lake, Douglas Lake, Reeses Bog, Gorge of Carp Creek, Iron Bridge,
Fontinalis Run, Smiths Bog, Black River, Cheboygan, Mill Creek. EMMET CO.—Pleasantview
Swamp, Carp Lake, Levering. PRESQUE ISLE CO.-Hammond Bay

Like other mosses of wet habitats (and most members of this “wet family’’),
Leptodictyum riparium is extremely variable. It grows most commonly on the floor of
hardwood swamps where it is submerged in the spring but becomes stranded as the
water table lowers. Typically it has straight, wide-spreading, rather complanate, entire,
long-pointed leaves with long cells.

We have both the fo. bai a (Schultz) Grout, a robust, usually submerged
form with leaves 3.5-4.5 mm. long, ending in a long, filiform acumen, and the
fluitans (Lesq. & James) Grout, a robust, submerged form with leaves about 3 mm.
long and shorter-acuminate than those of the fo. longifolium. Neither seems very
distinctive when compared with other expressions of this plastic species.

Zo7

=o! + / y K ees
aaa | N iy

ee he ele 743. Habit. 744. Three leaves. 745. Upper cells of leaf at margin.
746. Cells of alar re

2. Leptodictyum brevipes (Card. & Thér. ex Holz.) Broth.—Rather small plants.
Stems creeping. Leaves stiffly erect-spreading and concave when dry, more spreading
when moist, about 1.2-2mm. long, ovate or ovate-lanceolate, gradually short-
acuminate; costa extending about 2/3 the leaf length; upper cells oblong-rhomboidal,
4-6:1; basal and alar cells subquadrate to short-rectangular, not conspicuously dif-
ferentiated. Setae 1-2.5cm. long; capsule 2-2.5mm. long. Spores 9-13 y, faintly
roughened.

ow to Know the Mosses, (ed. 2) fig. 226. Grout, Moss Flora of North America 3:
1SB, ie “fig. os -h). Welch, Mosses of Indiana, fig. 160.

n moist, disturbed places, especially among grasses in partially shaded lawns, sometimes o
bark at base of trees, decayed wood, and other See Scattered in eastern North America ae
Minnesota and Missouri to Sabet and West Virg

The broad, rather stiffly erect-spreading, concave leaves with shortly acuminate
tips and rather short cells are distinguishing features. Leptodictvum brevipes has not

258

been found in Michigan, although reported earlier from the Douglas Lake region.
Because of its weedy occurrence across a broad range from Minnesota to Michigan, and
because it has been generally misunderstood, it seems likely to occur in the state and
no doubt commonly.

3. Leptodictyum trichopodium (Schultz) Warnst.—Plants in flat, loose, light-green
or yellowish mats. Branches spreading or ascending. Leaves not crowded, erect- to
wide-spreading, 1.3-1.9 mm. long, ovate-lanceolate, rather gradually acuminate; margins
subentire to + distinctly serrulate, especially at base; costa 0.6-0.7 the leaf length;
upper cells rhomboidal, 3-6:1, cells at base short-oblong in several marginal rows. Setae
10-30 mm. long; capsules 2-2.5 mm. long. Spores 11-15 yw, finely papillose.—n = 20 (as
L. kochii).

Conard, How to Know the Mosses, (ed. 2) fig. 217a-e & f (as var. kochii). Grout, Mosser
with Hand-lens a ee ce fig. 173 (as Amblystegium kochii), Moss Flora of North h America
3: Pl. 16 (fig. 12). Jennings, Mosses of Western Pennsylvania, (ed. 1) Pl. 38 & (ed. 2) Pl. 42 (as
Amblystegium kochii). Welch, ecu of Indiana, fig. 161, 162 (as var. Kochii).—Fig. 747-749.

On various substrata in ey places, particularly in swamps. Circumpolar; widespread in
North America, including Mexi

CHEBOYGAN CO.—Douglas, Burt, and Mud Lakes, Mill Creek. EMMET CO.—Maple River.

Smaller than L. riparium and somewhat resembling a large expression of
Amblystegium varium but with softer leaves, a shorter costa, and longer cells with
thinner walls. I make no attempt to distinguish L. trichopodium from its var. kochii
(BSG) Broth.

4. Hygroamblystegium Loeske, nom. cons.

Plants rather small to large and coarse, in soft or rigid, dull, dark-green, blackish,
or yellowish-brown mats in wet places, often submerged. Stems prostrate or crowded
and ascending, freely and irregularly branched, often denuded of leaves in older parts;
paraphyllia limited to branch bases. Leaves erect or somewhat incurved or rarely
subsecund, oblong-ovate or oblong-lanceolate, gradually acuminate, usually bluntly
acute, not or only slightly decurrent; margins plane, entire or nearly so; costa broad,
ending at or near the apex or nearly filling the acumen to excurrent; cells firm,
smooth, oblong-rhombic to shortly oblong-rhomboidal (about 2-6:1), generally some-
what enlarged and shortly oblong and thick-walled in 1 or more rows across the
insertion, the alar cells not differentiated. Autoicous; perichaetial leaves erect, lance-
acuminate. Capsules curved-cylindric; annulus well developed, of 2 rows of cells;
operculum acute to apiculate; endostome with not or narrowly perforate segments and
well-developed, nodose cilia in 2’s and 3’s._The name signifies an Amblystegium of
aquatic habitats. The genus is scarcely more than a large edition of Amblystegium with
a notably stout costa.

1. Plants large, coarse, and usually rigid; es ] broad (110-140 p wide), terete, Paes the
apex of the leaf and stoutly and shortly excurrent 2. A. i
1. Plants rather small, softer; costa much narrower, not terete, ending near the leaf apex oO
excurrent as a slender poi
2. Leaves gradually acuminate and bluntly acute; costa ending near the apex 1. H. tenax
2. Leaves rather abruptly subulate; costa excurrent as a slender poi
la. H. tenax var. spinifolium

259

1. Hygroamblystegium tenax (Hedw.) Jenn.—Plants rather small, dark-or yellow-
green, usually rather soft. Stem leaves not crowded, erect or sometimes wide-spreading,
somewhat incurved when dry, (0.7)1-1.5 mm. long, ovate or ovate-lanceolate, gradually
short-acuminate; costa yellowish, fairly strong, disappearing near the apex; upper cells
oblong-rhombic, 2-3:1, at the insertion 2-3 rows somewhat-enlarged, usually becoming
+ yellowish. Branch leaves oblong-lanceolate (or occasionally ovate), bluntly acuminate,
0.45-0.8 mm. long, erect-incurved. Setae 10-25mm. long; capsules 2-2.5 (rarely
3.5)mm. long. Spores 13-18 yw, faintly roughened or minutely papillose.—n = 12, 20,
30

, Mosses of Florida, Pl. se (figs. 5-8). Conard, How to Know the Mosses, (ed. 2) fig.

San Gro, eee ith Hand-lens and pee xe 174 (1a-d), i eal irriguum.
nnings, M of Ween eer ee (ed. 1) P , (ed. 2) Pl. 43 (as A. irriguum). Welch,
Mosses of ene fig. 169 (as H. wriguum). ee oe 752, 753 (H. HR iatie), 754 (4.

orthocladon

Hygroamblystegium tenax var. spinifolium. Be “Leaves
Hygroamblystegium noterophilum. 756. Leav

260

On wet rocks in and beside brooks, not limited to calcareous habitats. Europe, North Africa,
Madeira, oi — Caucasus; New Brunswick to Minnesota, south to Arkansas and Florida; Arizona
and Mexico.

CHEBOYGAN CO.—Reeses Bog, Hermits Bog, Carp Creek, Gorge of Carp Creek, Iron Bridge,
Burt Lake, Grass Bay, Duncan Bay, Mill Creek. EMMET CO.—Maple River, Wycamp Lake,
Wilderness State Park. MACKINAC CO.-—Mackinac Island, Cut River. PRESQUE ISLE CO.—
Evergreen Beach, Rainy River Falls.

la. var. spinifolium (Schimp.) Jenn.—Slender, yellowish-brown or brown plants
with leaves slightly incurved or subsecund when dry, shortly ovate and rather abruptly
narrowed to a slender subula largely filled by the excurrent costa.

Grout, Mosses with Hand-lens and Microscope, fig. 174 (1a’), as sede irriguum
eneten, pe Flora of North America 3: Pl. 16 (fig. 4), as H. irriguum var. spinifolium. Welch,
Mosses of Indiana, fig. 170 (as H. irriguum var. spinifolium).—Fig. 155.

Europe; Ontario, Michigan, British Columbia, and Colorado.

CHEBOYGAN CO.—Carp Creek and Gorge of Carp Creek. MACKINAC CO.—Mackinac
Island.

Hygroamblystegium tenax is relatively soft and slender, with leaves somewhat
incurved when dry. Its leaves are broader than those of Amblystegium varium, with
costa stouter. The var. spinifolium has leaves abruptly acuminate, with the costa nearly
filling the subula and excurrent.

Hygroamblystegium fluviatile (Hedw.) Loeske (fig. 753) is a poor thing scarcely
worth segregating from H. tenax. At best, it differs in being somewhat larger and in
having a somewhat broader costa (about 50-100 wide) which is blackish-green,
becoming pees brown with age, and upper cells 3-6:1, those at the insertion
greenish. H. tenax, by contrast, has the costa somewhat narrower, the upper cells
2-3:1, those at “ insertion + yellowish. These differences, by no means absolute,
should be viewed in terms of the variability commonly exhibited by aquatic calciphiles.

Hygroamblystegium orthocladon (P.-Beauv.) Loeske (fig. 754) is an unimportant
form of H. tenax with smaller, bluntly pointed leaves.

2. Hygroamblystegium noterophilum (Sull. & Lesq. ex Sull.) Warnst.—Large,
coarse, typically rigid plants in dark-green or brownish mats, often encrusted with lime
and often bristly because of persistent costae from which leaf laminae have eroded.
Stem and branch leaves similar, rigidly erect (only atypically soft and + incurved),
(1.5)2-3 mm. long, oblong-lanceolate, acuminate; costa very strong (110-140 wide
near the base), terete above, excurrent as a stout but mostly short point; upper cells
shortly oblong-rhomboidal, 3-5:1, not much differentiated at the insertion. [Only
imperfect sporophytes were seen. |

Grout, Moss Flora of North America 3: Pl. 17B.—Fig. 756.

Attached to wet, calcareous rocks, usually submerged in running water. New York and
Ontario to Minnesota, south to Missouri and Pennsylvania; Chile.

CHEBOYGAN CO. a Creek. EMMET CO.—Crooked Lake, Ramona Beach. PRESQUE
ISLE CO.—Rainy River Fa

The plants are much larger, coarser, and stiffer than those of H. tenax var.
spinifolium, and the costa is much stouter

5. Amblystegium BSG

Plants slender, green, yellowish, or brownish, in dull mats, usually in wet places.
Stems creeping, irregularly or subpinnately branched. Leaves small, somewhat concave,
erect- to wide-spreading when moist, ovate or lanceolate, gradually or rather abruptly
acuminate; margins plane, entire or + toothed; costa rather slender, ending at or above
the leaf middle, sometimes disappearing in the acumen; cells green, thin- or firm-
walled, smooth, oblong-rhombic or oblong-hexagonal above, somewhat longer and
broader at the base and subquadrate or shortly rectangular in poorly differentiated alar
groups. Autoicous; perichaetial leaves erect, pale, lanceolate, acuminate, weakly plicate.
Setae elongate, usually reddish; capsules cylindric or oblong-cylindric, curved and
inclined, becoming horizontal, arcuate, and constricted below the mouth when dry and
empty, smooth; annulus differentiated; operculum conic, acute or apiculate; endostome
with cilia nodulose, in groups of 1-3. Spores spherical.—With a blunt lid (or “roof”),
referring apparently to the fact that the operculum does not end in a beak. A swampy
habitat, slender, arched capsules, lanceolate leaves with a slender costa, and short cells
characterize the genus.

Costa disappearing in the acumen; upper cells 2-3:1 2. A. varium
Costa 1/2-2/3 the leaf length; upper cells 3-5:1 1. A. serpens

1. Amblystegium serpens (Hedw.) BSG—Small plants in loose, soft, green or
yellowish mats. Leaves crowded or remote, erect to wide-spreading, 0.5-0.9 mm. long,
lanceolate, gradually acuminate, + serrulate; costa 1/2-2/3 the leaf length; upper cells
oblong-rhomboidal, 3-5:1; cells at base broader and subrectangular, a few at the basal
margins subquadrate and short-oblong. Setae 9-23 mm. long; capsules 1.5-2 mm. long.
Spores 11-18 yu, finely papillose—n = 7, 10, 10+1, 11, 12, 14.2197 1941; 1942, 20, 22:
AO (and, from experimental material, 24, 28); 2n = 40.

Breen, Mosses of Florida, Pl. 71 (figs. 1-5). Darlington, Mosses of Michigan, fig. 112. Grout,
Mosses with Hand-lens and Microscope, fig. 171, Moss Flora of North America 3: Pl. 16 (figs.
1la-b), as A, juratzkanum. Jennings, Mosses of Western Pennsylvania, (ed. 1) Pl. 37 & (ed. 2) Pl. 41
(as A. serpens & A. juratzkanum). Welch, Mosses of Indiana, fig. 163.—Fig. 757-761.

Vv common, on wet soil, humus, rotten wood, and other substrata in swampy places.
Nearly cosmopolitan; widespread in North America.

CHEBOYGAN CO.—Douglas Lake, Smiths Bog, Reeses Bog, Colonial Point, Riggsville, Grass
Bay. EMMET CO.—Arnott Lake, Pellston Hills, Carp Lake, Ellis Township, Cross Village, Middle
Village, Five Mile Creek, Big Stone Bay. MACKINAC CO.-—Bois Blanc Island, Mackinac Island.
PRESQUE ISLE CO.—Rainy Lake, Ocqueoc River, Clinton Lake.

The small leaves with short, firm cells and short midrib are distinctive characters.
In its more typical form the leaves are more or less erect when dry. In the so-called
var. juratzkanum (Schimp.) Rau & Herv., which some authors consider a troublesome
species (but which I generally ignore), the leaves are wide-spreading wet or dry.
(Chromosome counts for the latter include n = 11+1, 12, 12+1, 13, 20.)

2. Amblystegium varium (Hedw.) Lindb.—Small plants in loose, yellowish or
green, often + rigid mats. Stem leaves often considerably longer and broader than the
branch leaves, about 0.6-1.2 mm. long, ovate-lanceolate, somewhat decurrent, with
costa stronger and more nearly percurrent, about 22-33 u wide near the base. Branch
leaves loosely erect and usually slightly incurved when dry, spreading or erect-spreading
when moist, very variable in size and shape, about 0.35-0.7 mm. long, lanceolate,
gradually and slenderly acuminate; margins entire or slightly serrulate; costa disappear-

262

759 |, x \ | —_

\ if
aC . 761
if] 762
/ ' |

J / VS / /\ \ if

ie [ NN I ) /
‘Ve iN I)
758 “ ¥
y ve we oe gs
wor gees ss
; 757 763

mblystegium serpens, 757. Habit. 758. Leaves. 759. Cells of branch leaf. 760. Upper cells
of branch leaf (shorter and more typical). 761. Capsules
Amblystegium varium. 762. Leaves. 763. Alar cells.

ing in the acumen, often flexuose above, about 17-26 u wide near the base; upper cells
oblong-rhombic, about 2-3:1; lower cells shortly oblong-hexagonal, a few at the basal
angles subquadrate. Setae 12-22 mm. long; capsules 1.2-2 mm. long; cilia 1-3. Spores
9-14 2, minutely papillose.—n = 10, 12+1, 19+1, 20, 22.

Grout, Mosses with Hand-lens and Microscope, Pl. 78 & fig. 175(2). Jennings, Mosses of
Western nates eae (ed. 1) Pl. 37, (ed. 2) Pl. 41. Welch, Mosses of Indiana, fig. 165.—Fig
762-763.

On soil, humus, logs, rocks, and other substrata, in wet, shady places. Circumpolar;
widespread in North America; Mexico; Haiti

CHEBOYGAN CO.—Polypody Bluff (Douglas Lake), Reeses Bog, Hermits wee Gorge of
Carp Creek, Iron Bridge, Smiths Bog, Colonial Point, Ingleside, Black Lake. EMMET CO.—Five
Mile Creek, Pellston Hills) MACKINAC CO.—Gros Cap, Caffey Corner. PRESQUE aie CO.—Near
Belknap, Clinton Lake.

As the name suggests, this is a variable and troublesome species. The plants are
somewhat larger than those of A. serpens. The main character is the long costa which
disappears in the narrow acumen. The basal cells of the leaf are typically dark and
somewhat enlarged. The species approaches Hygroamblystegium tenax, but the leaves
are softer and narrower and the costa considerably narrower. The habitat of H. tenax
on wet rocks (and commonly submerged) is, of course, distinctive.

6. Drepanocladus (C. M.) Roth

Plants generally rather robust, in extensive mats or deep tufts, green, yellow-
green, golden or brownish, sometimes red, often + glossy, mostly growing in wet
places, sometimes submerged. Stems creeping or erect-ascending, freely and irregularly
to pinnately branched, or rarely nearly simple; paraphyllia few, at branch bases. Leaves
nearly always falcate-secund, + concave, sometimes striate or plicate, lanceolate or
ovate-lanceolate, slenderly long-acuminate, frequently + auriculate at base; margins
erect, entire or + serrulate; costa single, ending near the leaf middle or in the acumen,
sometimes excurrent; cells mostly long-linear, smooth, sometimes thick-walled and
porose at base or throughout, usually differentiated at the basal angles, sometimes in
well-marked, excavate groups extending nearly to the costa, hyaline, thin-walled, and
inflated or sometimes colored and thick-walled. Perichaetial leaves erect, lanceolate,
slenderly long-acuminate, mostly plicate. Capsules + curved, usually strongly so;
annulus mostly differentiated; operculum apiculate; endostome with often perforate
segments and nodulose cilia in groups of 1-4.-The name refers to the falcate-secund
leaves which give the branch tips a hooked, sickle-like appearance. This is a genus of
mostly aquatic and subaquatic species, varying chaotically with seasonal fluctuations in
water level and associated changes in salinity and hydrogen-ion concentration.

1. Leaves plicate or striolate when dry
2. Leaves plicate, with a few enlarged, hyaline alar cells; cortical cells large, hyaline,
Ned 1.

thin-walle . uncinatus
_ Leaves merely striolate when dry, without enlarged alar or cortical cells 3. D. vernicosus
1. Leaves smooth or (if striolate when dry, the alar cells conspicuously inflated)
3. Leaves entire
4, Alar cells not or very slightly differentiated 2. D. revolvens
4. Alar cells inflated, in large and conspicuous groups
5. Cells of leaf base clearly elongate, oblong or oblong-linear 4. D. aduncus
5. Cells of leaf base shortly oblong or oblong-hexagonal 4a. D. aduncus var. polycarpus

3. Leaves serrulate, at least at the apex ;
_ Leaves relatively flat, often striolate when dry; costa rather broad, usually extending
well into the acumen; alar cells abruptly inflated, in large, decurrent auricles usuall
reaching the costa; dioicous 5. D. exannulatus
_ Leaves concave, smooth; costa narrow, ending well above the midleaf; alar cells often
somewhat enlarged but not abruptly differentiated, not in decurrent auricles; autoicous
6. D. fluitans

1. Drepanocladus uncinatus (Hedw.) Warnst.—Plants rather slender to fairly
robust, in loose, yellow, brownish, or green, somewhat shiny mats or tufts. Stems
creeping to more crowded and erect or ascending, irregularly to subpinnately branched;
cortical cells large and hyaline. Leaves strongly falcate-secund, 2.5-4 mm. long,
gradually long-subulate from a lanceolate base, not decurrent, slightly to strongly
plicate (especially when dry), remotely serrulate toward the apex; costa ending above
the middle or in the acumen; upper cells linear-flexuose, those at the insertion oblong,
thick-walled and porose, the alar cells oblong-hexagonal, hyaline, and thin-walled in
small, inconspicuous alar groups. Autoicous; perichaetial leaves greatly elongate
(4-8 mm. long), + plicate. Setae 15-32 mm. long; capsules 1.7-3 mm. long; annulus
present; cilia of endostome 1-3. Spores 10-16 y, very finely papillose.—n = 9+1, 10, 11,
15.203

Conard, How to Know the Mosses, (ed. 2) fig. 210a-d. Grout, Mosses with Hand-lens and
Microscope, Pl. 72.—Fig. 764-766.

On rocks, decaying wood, humus, or bark at base of trees in moist or dry, coniferous
forests. Nearly cosmopolitan; Greenland to Alaska, south to California, Arizona, New Mexico, Ohio,
and Pennsylvania.

264

CHEBOYGAN CO.—Reeses Bog, Hermits Bog, Gorge of Carp Creek, Iron Bridge, Burt Lake,
Grass Bay. EMMET CO.—Arnott Lake, Pleasantview Swamp, Cecil Bay. MACKINAC CO.-—Bois
Blanc Island, Mackinac Island, St. Martins Point, Cut River, Epoufette, Ozark. PRESQUE ISLE
CO.—Mast Point, Lamprey Experiment Station, Evergreen Beach.

Unlike other members of the genus, D. uncinatus grows in relatively dry niches
(such as on logs) in dry or moist, coniferous forests. It can be quite beautiful, with
golden or green, glossy leaves which are circinately secund and deeply plicate, not
unlike those of Prilium. It is variable, however, and often scraggly in appearance. A
common expression is the so-called var. plumulosus (BSG) Warnst., which is more
slender, with leaves not or only slightly striate and nearly entire. The var. symmetricus
(Ren. & Card.) Grout has capsules small, erect, symmetric, and cylindric; its leaves are
less strongly plicate than in the typical variety and entire or slightly denticulate. (It has
been collected in the Lake Superior region of Ontario.)

2. Drepanocladus revolvens (Sw.) Warnst.—Rather robust plants in dense, glossy,
typically reddish to blackish-purple tufts, sometimes green, yellow, or brownish (with
or without red tinges). Stems erect or ascending, irregularly to subpinnately branched;
central strand present but very small and inconspicuous, consisting of 3-4 small cells;
outer cells of stems and branches hyaline and thin-walled (usually remaining attached
to the bases of leaves stripped off for examination and obviously broader than leaf
cells). Leaves usually crowded, strongly falcate-secund with flexuose tips, smooth (or
very rarely and very slightly striolate) when dry, 3-4 mm. long, concave, lanceolate or
ovate-lanceolate, gradually narrowed to a long, filiform, flexuose point; margins erect,
entire; costa slender, about %4 the leaf length; cells long-linear, flexuose, with thick
walls, a few rows at the insertion oblong-linear and somewhat darker with thick, +
porose walls, at the basal angles a few, hyaline, slightly enlarged cells. Autoicous. Setae
30-55 mm. long; capsules 2-3 mm. long; annulus present; cilia of endostome in groups
of 1-3. Spores 13-18 yu, finely papillose—n = 10+1, 20, 22+1, 23.

Darlington, Mosses of Michigan, fig. 120. Grout, Mosses with Hand-lens and Microscope, Pl.
71, Moss Flora of North America 3: Pl. 28B.—Fig. 767-768.

On wet soil in swampy places, near springs, margins of lakes, etc., apparently most common
in alkaline bogs. Circumpolar; Colombia; New Guinea; Alaska and British Columbia, across the
continent and south to Michigan, Quebec, and Nova Scotia.

CHEBOYGAN CO.-—Little Lake 16. EMMET CO.—Cecil Bay, Big Stone Bay. MACKINAC
CO.—Bois Blanc Island.

The plants are a beautiful purplish-red, with long, flexuose, circinate leaf tips.
The basal cells are incrassate and porose and contrast with the large, hyaline cortical
cells which generally strip off with the leaves on dissection. A few alar cells are often
hyaline and somewhat enlarged.—Locally rare, perhaps limited to more acid situations.

Var. intermedius (Lindb. ex C. J. Hartm.) Rich. & Wall.—Plants yellow-brown.
Leaves with shorter acumina, not or rarely somewhat striolate. Cortical cells of stem
enlarged; cells across insertion of leaf not much differentiated, a few alar cells hyaline
and somewhat inflated (though inconspicuous). Locally common in rich (calcareous)
fens. It occupies the same habitats as D. vernicosus and looks like it too. I have
trouble finding any characters separating them. I believe that D. revolvens var.
intermedius is only an expression of D. vernicosus, but without further study I lack
the courage to flout tradition. (The fact that D. revolvens is acid-loving also makes me
wonder if such a calciphile as the so-called var. intermedius is genetically allied to it.)
Circumpolar; common in more northerly latitudes of North America.

265

\\

\

—S 4 Z {fit
2
as
WG

Ss
a
wy

HAS ‘Se f
Sg yy

WS Y /
a

——

Drepanocladus uncinatus, 764. Habit. 765. Leaf. 766. Alar cells.
Drepanocladus revolvens. 767. Leaf. 768. Alar cells.
Drepanocladus vernicosus, 769. Leaves. 770. Alar cells.

3. Drepanocladus vernicosus (Lindb. ex C. J. Hartm.) Warnst.—Plants in green or
yellowish, sometimes brownish, rarely reddish tufts. Stems erect-ascending, + pinnately
branched; central strand none; cortical cells not large (not often remaining attached to
leaves stripped from the stems and not obviously broader than leaf cells). Leaves
usually not crowded, + striolate when dry, falcate-secund, 1.5-2 mm. long, lanceolate
to ovate-lanceolate, gradually narrowed to a relatively short or sometimes long, slender,
flexuose-twisted acumen, entire; costa ending somewhat above the leaf middle; upper
cells linear-flexuose; basal cells shorter, more incrassate, porose, and colored in about 2
ows at the leaf insertion, a few at the basal angles occasionally pale and slightly
differentiated. Dioicous. Setae 35-65 mm. long; capsules 2-3.5 mm. long; annulus of 2-3
rows of cells; cilia in 2’s and 3’s. Spores 12-22 u, nearly smooth to finely papillose.

266

Mosses with Hand-lens and Microscope, fig. 160 and Pl. 71. Welch, Mosses of Indiana,
fig. re “Fig. 769-770.

On wet soil or humus in calcareous fens. Circumpolar; Greenland to Alaska, south to
Oregon, Iowa, Ohi, and Pennsylvania.

CHEBOYGAN CO.—Reeses He Douglas Lake, Grass Bay. EMMET CO.-—Cecil Bay, Big
Stone Bay, Lawrence Lake, Conway, Bear Creek Township. MACKINAC CO.—Mackinac Islan d,
Bois Blanc Island, Mud Lake, Rexton. PRESQUE ISLE CO.—Mast Point, Hammond Bay, Evergreen
Beach, Ocqueoc Lake, Kelsey Pond, Clinton Lake.

The leaves are finely striate when dry. The cortical cells are not large and
hyaline, as in D. revolvens var. intermedius, but the extreme basal cells may be hyaline
in a single row. Above them are one or sometimes two rows of thick-walled cells
forming a darker line across the leaf. D. revolvens var. intermedius is similar, but the
cells across the insertion may be less clearly differentiated, the cortical cells enlarged,
and a few alar cells hyaline and somewhat inflated. As stated above, I have grave
doubts of any genetic difference, regardless of characters which in some specimens
allow pigeonholing.

4, Drepanocladus aduncus (Hedw.) Warnst.—Rather robust plants in soft, loose
or dense, dark- to yellow-green or brownish mats or tufts. Stems spreading or more
crowded and suberect, irregularly to pinnately branched. Leaves falcate-secund, not
plicate or striolate when dry, ovate-lanceolate and gradually narrowed to a short or
long, channeled acumen, entire; costa narrow or stout, ending at or above the leaf
middle; cells linear-flexuose above, those at base long-rhomboidal, not or somewhat
porose, the alar cells clearly differentiated, yellow-brown or hy aline, thin- or rather
thick-walled, and inflated in large, non- decurrent, auriculate groups scarcely extending
to the costa. Dioicous. Setae about 2.5 cm. long; capsules 2-2.5 mm. long; annulus
present. Spores about 164, papillose—n = 12, 20 (as D. kneiffii).

, How to Know the Mosses, (ed. 2) fig. 202a, d. Grout, Mosses yes Hand-lens and
Microscope, PL 74 (fig. 1), also fig. 163b, Moss Flora of North America 3: Pl. 24D, 27A (figs. an
Jennings, Mosses of Western Pennsylvania, (ed. 2) Pl. 67. Welch, Mosses of soil fig. 183.—
771-772.

On soil in wet, often calcareous habitats (swamps, meadows, boggy woods), eae
submerged or emergent, at least during high water, in the shallow water of pools, sprin
ditches. Circumpolar; Mexico, ee nea New Zealand, Kerguelen; eee in ne th
America, especially common in the North.

CHEBOYGAN CO,—Hermits Bog, Reeses Bog, Gorge of Carp Creek, Burt Lake, Douglas
Lake, Ingleside, Bryants Bog, Smiths Bog, Passino School, Lancaster Lake, near Mackinaw City,
Duncan Bay, Black Lake, Indian River. EMMET CO.—Levering, Arnott Lake, Crooked River, Carp

Pegeley Lake. PRESQUE ISLE CO.—Mast Point, Grace Harbor.

4a. Var. polycarpus (Bland. ex Voit) Roth—Leaves slightly to strongly falcate-
secund, broadly oblong-ovate, usually rather shortly as cells of the leaf base
shortly oblong or oblong- hexagonal, the alar cells in + decurrent auricles extending
nearly to the costa.
Mosses with Hand-lens and Microscope, Pl. 74 (fig. 2), Pl. 74 (figs. 10-11), as fo.

Gro
gracilescens, neta Flora of North America 3: Pl. 26D & ee oe i), Pl. 27A (figs. k-l) & 28A (as
fo. gracilescens). Welch, Mosses of Indiana, fig. 186. —Fig. 77

A plant of wet, calcareous habitats, sometimes submer rged. Nearly eae throughout
arctic America and south to Pennsylvania, Missouri, New Mexico, and Californ

267

y, oN mae \
Kf

{| N

iA . t \

’ J | \ ia
772 : | 774
fe” ra
77\ Gees

Drepanocladus aduncus vat. aduncus. 771. Leaf. 772. Cells at alar region.
Drepanocladus aduncus vat. polycarpus, 773. Leaves. 774. Alar cells.

CHEBOYGAN CO.—Reeses Bog, Gorge of Carp Creek, Burt Lake, Linné Bog, Mud Lake,
Munro Lake, Lancaster Lake, Black Lake, Black River, Duncan Bay. EMMET CO.—Maple River,
Carp Lake, Arnott Lake, Alanson. MACKINAC CO.—Bois Blanc Island. PRESQUE ISLE CO.—Grace
Harbor, Ocqueoc Falls.

Drepanocladus aduncus is very common and exceedingly variable. It is usually
soft and green to yellow or brown, with leaves not crowded, moderately falcate, and
not plicate, with alar cells clearly differentiated in large groups. The entire leaf margins
provide a quick means of separation from D. fluitans and D. exannulatus. The typical
variety has long basal cells, whereas those of the var. polycarpus are short. The var.
polycarpus fo. gracilescens (BSG) Ménk. has been collected at Duncan Bay in
Cheboygan County; it is an unusually slender form of little significance. The var.
kneiffii (BSG) Monk. is a common form with leaves more or less remote and not or
only slightly falcate; I see no good reason to bother with it. Finally, D. sendtneri
(Schimp.) Warnst. has alar cells usually orange-colored and rather thick-walled in fairly
distinct groups and with a stronger costa; I include it here as a probable ecological
expression, although I am not sure of its taxonomic limits or ecological preferences. (It
has been collected in Cheboygan and Delta Counties and is probably common
elsewhere in especially calcareous habitats.) A number of local collections qualify as
var. aduncus fo. capillifolius (Warnst.) Monk., a robust, generally submerged form with
straight or + curved leaves with long, slender acumina and long, often excurrent costa;
both var. aduncus and var. polycarpus apparently produce such capillifolius forms.

Calliergidium pseudostramineum (C. M.) Grout, which bears some resemblance to
Drepanocladus and Calliergon, has been found in Delta County. The leaves, crowded

268

and more or less imbricate, are striolate when dry, ovate or ovate-lanceolate and
broadly tapered to a narrow, rounded point; the margins are entire and the alar cells
abruptly inflated.

5. Drepanocladus exannulatus (BSG) Warnst.—Plants green or yellow-brown,
sometimes red-tinged to purple-red, up to 25 cm. long. Stems + pinnately branched.
Leaves twisted and striate when dry, somewhat to strongly falcate-secund, 2-5 mm.
long, lanceolate and gradually long- to filiform-acuminate, the apex sometimes sub-
obtuse; margins plane, subentire or more often + serrulate, sometimes nearly to the
base; costa rather broad, extending well into the acumen, sometimes ending near the
apex; upper cells linear; alar cells hyaline, thin-walled, and conspicuously inflated in
auriculate groups extending nearly to the costa. Dioicous. Setae 30-75 mm. long;
capsules 2.2-3 mm. long; annulus none; cilia in groups of 2 or 3. Spores 15-24 u, very
finely papillose.—n = 11, 12.

nard, How to Know the Mosses, (ed. 2) fig. 203. Grout, Mosses with Hand-lens and
Microscope, Pl. 74 (figs. 7, 8, 16), Moss Flora of North America 3: Pl. 29B, C. Jennings, Mosses of
Western Pennsylvania, (ed. 1) Pl. 40, (ed. 2) Pl. 44. Welch, Mosses of Indiana, fig. 189.—Fig.
775-777.

In wet places, sometimes submerged. Circumpolar; Colombia; Greenland to Alaska and south
to California, Colorado, Michigan, and Pennsylvania.

CHEBOYGAN CO,.—Livingston Bog, Smiths Bog, Vincent Lake, Mud Lake, Little Lake 16.
EMMET CO.—Sherett Lake, Stutsmanville. MACKINAC CO.-—Little Dollar Lake.

As compared with D. fluitans, this species is more pinnately branched and often
purplish, with leaves more or less flat and often finely striate when dry and alar cells
in large, abruptly differentiated, excavate groups. The serrulate leaf margins alone will
distinguish it from the myriad forms of D, aduncus.

Drepanocladus exannulatus, like other mosses of wet or aquatic habitats, varies
endlessly with conditions of environment. The taxonomic implications of suc
variations are beautifully illustrated by Smirnova’s transplant studies, summarized in
English by Lodge (Trans. Brit. Bryol. Soc. 2(4): 575-577. 1955). According to her
findings, the var. rotae (De Not.) Loeske is merely a submerged form of the species. In
dryer than normal conditions, a short-leaved and short-celled var. alpinus (Grav.) Wijk
& Marg., also known as the fo. brachydictyus (Ren.) Smirn., develops. With good
aeration and high light intensity, this form becomes purple and may be called the var.
purpurascens Schimp. On extreme exposure, the fo. tundrae (Arn.) Monk., often
recognized at the specific level as D. tundrae (Arn.) Loeske, results.

Lodge (Jour. Linn. Soc. London, Bot. 56: 218-224. 1959) found by varying
environmental conditions in culture that growth forms of D. exannulatus and D.
fluitans could be greatly altered, whereas certain structural details, such as the nature
of the alar cells, remained relatively unchanged.

Sonesson (Bot. Notis. 119: 379-400. 1966) grew Drepanocladus trichophyllus
(Warnst.) Podp. (a relative of D. exannulatus with a long excurrent costa) from spores
and reported that “four different morphological categories are possible to distinguish in
the progeny.”

6. Drepanocladus fluitans (Hedw.) Warnst.—Plants in loose, green, yellow-green,
or brown, shiny masses. Stems sparsely branched. Leaves rather distant, falcate-secund,
at least at the tips of stems and branches, 3-4 mm. long, oblong-lanceolate, gradually
narrowed to a long, flexuose acumen, somewhat narrowed to the base, not or only

269

slightly decurrent, serrulate at the apex; costa slender, ending well above the midleaf;
cells linear-flexuose; alar cells oblong, hyaline or colored, usually somewhat enlarged,
occasionally inflated but not forming auricles. Autoicous. Setae 50-75 mm. long;
capsules about 2-3 mm. long; annulus none. Spores 17-27 y, finely papillose.—n = 20,
22, 2242, 24; also 20 (as var. falcatus).

Grout, Mosses with Hand-lens and Microscope, Pl. 73, also fig. 163a, Moss Flora of North
America 3: Pl. 29A. Jennings, Mosses of Western Pennsylvania, (ed. 2) Pl. 69. Welch, Mosses of
Indiana, fig. 188.—Fig. 778-780.

Drepanocladus exannulatus. 775, Leaves. 776. Cells at leaf tip. 777. Alar cells.
Drepanocladus fluitans. 778. Leaf. 779. Cells at leaf tip. 780. Alar cells.
Hygrohypnum luridum. 781. Habit. 782. Leaves.

t soil of ditches, swamps, bogs, meadows, swales, or lake margins, often submerged.
Nearly cosmopolitan; widespread in cae latitudes of North America, south to California,
Colorado, Michigan, and West Virginia.

CHEBOYGAN CO.—Reeses Bog, Gleasons Bog, Bryants Bog, ean Bog, Mud Lake.
EMMET CO.-—Sherett Lake, Gladys Bog, Stutsmanville. MACKINAC CO.—Rex

In contrast to D. exannulatus, D. fluitans is softer, shinier, and often brownish or
reddish but never wine-red or purple. The leaves, not striate, are scarcely narrowed to
the base; the alar cells are sometimes slightly inflated but usually not abruptly
differentiated and not occupying excavate groups. Williams (Bryol. 33: 32. 1930)
reported this species from a depth of 62 feet in a lake in northern Wisconsin.

Drepanocladus fluitans var. uncatus Crum, Steere & Anders., generally known as
the var. falcatus (C. Jens.) Roth and sometimes as D. schulzei Roth, is a rather
common form of the species with alar cells slightly more differentiated than usual.

7. Hygrohypnum Lindb.

Plants slender to moderately robust, in green, yellow, or red-gold, + shiny, dense
mats. Stems creeping, irregularly branched; paraphyllia few, at the branch bases;
paraphysis-like hairs often in leaf axils. Leaves imbricate to spreading, often secund,
concave (sometimes strongly so and + cucullate at the apex), smooth or weakly plicate,
oblong-lanceolate and acuminate or broadly oblong-ovate to subcircular and blunt or
rounded at the apex; margins erect, sometimes incurved toward the apex, entire or
serrulate; costa often variable on the same plants, generally unequally forked and
usually short and weak, rarely single and nearly percurrent; cells smooth, oblong-linear
or linear-flexuose, often short and oblong-rhombic at the extreme apex, generally
yellow or orange at the middle of the insertion, the alar cells subquadrate or oblong,
hyaline or colored, sometimes clearly inflated in small, well-marked alar groups.
Perichaetial leaves erect, elongate, generally plicate. Capsules ovoid or oblong-cylindric,
+ curved; annulus present; operculum sometimes rostrate; endostome with perforate
segments and + nodose cilia (rarely lacking or rudimentary).—_A “Hypnum” of wet
habitats. The very concave leaves and variable costa are characteristic features of the
genus, as well as the occurrence on wet rocks

Hygrohypnum luridum (Hedw.) Jenn.—Plants of medium size, in dark- or
yellow-green to brownish, loose or dense mats. Leaves crowded, erect or, more often,
cund, 1-1.5 mm. long, concave, broadly to narrowly oblong-ovate, obtuse or acute,
sometimes apiculate; margins erect or incurved above, entire; costa variable even on the
same plants, yellowish, single, and 1/2-2/3 the leaf length, sometimes short and double
or forked; upper cells linear-flexuose; alar cells rather numerous, small, subquadrate,
opaque. Autoicous. Setae 13-20 mm. long; capsules about 2.5mm. long; cilia 1-3.
Spores 15-18 y, finely papillose so nearly smooth.—n = 10, 10+1, 11, 12+1

Conard, How to Know the Mosses, (ed. 2) fig. 273e. Darlington, Mosses of Michigan, fig.
116. Grout, Moss Flora of North America 3: Pl. 23 upper & lower right, lower left (as H. palustre,
var. subsphaericum, & var. julaceum).—Fig. 781-782.

wet rocks in and beside streams (especially common in cascading brooks and near
waterfalls) probably a calciphile. Cir cumpo olar; widespread in North America, across the continent
and south in the mountains to Colorado, in the East to Pennsylvania

CHEBOYGAN CO.—South of Burt Lake, Mill Creek. PRESQUE ISLE CO.—Ocqueoc Falls.

271

he leaves may be distinctly secund or erect and imbricate, but in spite of
considerable variability, the deeply concave, entire leaves with alar cells few and small
d costa variable, coupled with the subaquatic habitat, are useful differentia.
Braithwaite expressed the variability well when he said, “So polymorphous is this
species that a long series of varieties might be set up, without much benefit, as they do
not appear to maintain their stability, and even two forms of nerve may be found in
leaves from the same plant.” The species is locally rare enough not to cause too much
trouble.

n the Upper Peninsula we have two other species: H. ochraceum (Turn. ex Wils.)
Loeske has loosely falcate-secund leaves and large hyaline alar cells and similarly
enlarged cortical cells. It resembles Hypnum lindbergii but has a much stronger costa,
one-half or more the leaf length. The chromosome number has been recorded as n =
10, 11. H. molle (Hedw.) Loeske has leaves spreading from an erect base.

8. Calliergon (Sull.) Kindb.

Plants relatively robust, in deep, loose tufts, green or yellowish to red-brown or
purplish, sometimes + shiny. Stems erect or ascending, sparsely to freely and
sometimes pinnately branched; branches cuspidate at the tips because of crowded,
erect leaves; paraphyllia few, at branch bases. Stem leaves large, imbricate to
erect-spreading, + concave, often cucullate at the apex, rarely somewhat striate, oblong-
to rounded-ovate or suborbicular, broadly rounded and sometimes apiculate at the
apex, often producing radicles at back near the apex from scattered, pellucid cells;
costa mostly rather strong, single, extending somewhat beyond the leaf middle or
nearly to the apex, sometimes forked at the tip; cells smooth, linear-rhomboidal,
shorter toward the base, subquadrate or hexagonal and inflated in usually well-marked,
concave alar groups, rarely slightly differentiated at the basal angles. Branch leaves
smaller, narrower, and often subtubulose above. Perichaetial leaves erect, + long-

curved; annulus present or lacking; operculum blunt or acute; endostome with
sometimes perforate segments and delicate, sometimes nodose cilia in 2’s to 4’s.—The
name, meaning pretty work, refers to good workmanship or elegance in construction or
appearance.

1. Plants usually dark and brownish, terete when dry, turgid when moist; alar cells not ages
differentiated 2. C. trifarium

1. Plants usually pale and yellowish or greenish, + cuspidate at tips of stems and branches
but not pie ines or turgid; alar cells inflated in large, gradually or abruptly
differentiated gro

2. Plants relatively eraee leaves oblong or oblong-ovate

_ 2. Plants coarser and fairly robust; leaves broad, cordate-ova
3. Plants sparsely and irregularly branched; inflated alar ail gradually differentiated

. C. cordifolium

1. C. stramineum

3. Plants pinnately eee ee me cells abruptly differentiated
4. Dioicous; costa e ar the

m
4. Autoicous; costa ne a oe ees

3. C. giganteu

4. C. richardsonii

1. Calliergon stramineum (Brid.) Kindb.—Relatively slender plants in deep, loose or
dense, somewhat shiny, green or more often yellowish to straw-colored masses. Stems
5-12 cm. high (or rarely as much as 30cm.), sparsely branched, erect-ascending,
slenderly terete toward the apex and cuspidate-tipped because of crowded, imbricate
leaves. Stem leaves somewhat crowded, erect-spreading below, erect or imbricate above
when dry and erect or erect-spreading when moist, 1.2-2 mm. long, oblong or
oblong-ovate, rounded and cucullate at the apex, often radiculose near the tips,

‘S784

Calliergon stramineum, 783. Habit. 784. Leaves. 785. Alar cells.
Calliergon trifarium. 786. Habit. 787. Leaves. 788. Alar cells.

decurrent, entire; costa 3/4-5/6 the leaf length; cells at extreme apex shortly oblong or
thombic; upper median cells linear, becoming shorter and broader near the base,
inflated in fairly large but not sharply delimited, decurrent, hyaline alar groups (often
orange-yellow within). Dioicous. Setae 35-45 mm. long, red-yellow or red; capsules
about 2mm. long; annulus none; operculum apiculate: ee of endostome
narrowly perforate; cilia 1-2. Spores 13-15 yw, finely roughened.—n =

arlington, Mosses of Michigan, fig. 118. Grout, Moss Flora of North America 3: Pl. 20
above. a 783-785.

hallow pools or damp depressions in fens, bogs, or ae woods. Circumpolar;
ésni ener to Oregon, Michigan, Ohio, and renters Australia

CHEBOYGAN CO.-—Bryants Bog, Gleasons Bog, Nichols Bog, Livingston Bog, M e,
Little Lake 16. EMMET CO.-—Galloway Bog, Wilderness State Park. MACKINAC CO. come
Lake, Little Dollar Lake.

Calliergon stramineum, as the name indicates, is somewhat straw-colored. The
plants are pretty and easily recognized by their slender, nearly simple stems ending in

23

narrow, terete, shiny points. As Braithwaite so quaintly put it, ‘““The leaves often throw
out radicles from the tips.’ Even when such radicles cannot be found, large,
differentiated cells from which they are initiated may be demonstrated easily.

Much like Calliergon stramineum, and likewise often bearing radicles at the leaf
tips, is Calliergidium pseudostramineum (C. M.) Grout. It can be distinguished by the
fact that the leaves are not crowded and appressed in cuspidate points at the tips of
stems and branches, and in shape they are distinctly tapered to a narrowly rounded
apex. A rare species, it has been found once in Michigan, in Delta County. It may
occur elsewhere in the Upper Peninsula, in wet places, especially in the crevices of wet
rocks in stream gorges and near waterfalls. (It is probably an acidophile.) The
occurrence of radicles at leaf tips is not worth too much emphasis. They also occur on
occasion at leaf tips of Drepanocladus fluitans and D. exannulatus.

2. Calliergon trifarium (Web. & Mohr) Kindb.—Plants in loose mats 4-16 cm.
high, frequently mixed with other mosses, green or more often yellow- or golden-
brown, dark-brown below. Stems elongate, erect-ascending, simple or sparsely forked,
soft and flexible when moist, brittle and usually smoothly terete when dry, more
turgid when moist. Leaves crowded, erect or imbricate when dry, sometimes loosely
erect-spreading when moist, cucullate-concave, smooth or + striolate when dry,
1-1.6 mm. long, elliptic to oblong-ovate, not decurrent, rounded at the apex; margins
erect or frequently revolute at the extreme base, usually crenulate at the extreme apex;
costa slender, single (or occasionally double or forked), about 1/3-2/3 the leaf length,
rarely e; upper cells oblong-linear (short at the extreme apex); alar cells not
particularly differentiated, somewhat enlarged and oblong with firm, often yellowish
walls at the basal angles or almost across the insertion. Dioicous. Setae 30-50 mm.
long; capsules 2.5-3.5 mm. long; annulus broad; operculum blunt; endostome segments
perforate; cilia nodose, in 2’s and 3’s. Spores 13-15 y, very finely papillose.

Grout, Moss Flora of North America 3: Pl. 21 lower left.—Fig. 786-788.

Emergent from shallow water of pools or among sedges in rich fens (around beach pools in
the Great Lakes area); probably restricted to calcareous habitats. mae Greenland; British
Columbia to Labrador, south to Oregon, Michigan, Ohio, and Connectic

EMMET CO.-Cecil Bay. MACKINAC CO.—Bois Blanc Island. PRESQUE ISLE CO.-
Hammond Bay, Evergreen Beach, Clinton Lake.

A rare species of richly calcareous fens, usually growing as separate strands in
tufts of Drepanocladus or Scorpidium (and easily overlooked). In the field, the terete,
scarcely branched plants have a dark, varnished appearance, and the leaves have a spiral
arrangement which is nearly lost on drying.

3. Calliergon giganteum (Schimp.) Kindb.—Rather robust plants in deep, loose,
green, yellow-green, or brownish masses 7-20, rarely 35 cm. long. Stems ascending,
pinnately branched; stems and branches cuspidate at the tips. Stem leaves not crowded,
erect or erect-spreading except at the cuspidate tips, about 1.3-3 mm. long, broadly
cordate-ovate, rounded and cucullate at the apex, broadly decurrent, entire, often
indistinctly striate when dry; costa ending near the apex; upper cells linear-rhomboidal;
alar cells large and hyaline, abruptly differentiated, in large, concave, decurrent
auricles. Branch leaves narrower, usually wide-spreading and rolled when dry. Dioicous.
Setae 45-70 mm. long, orange-brown; capsules 2.5-3 mm. long; annulus none; oper-

274

*

culum apiculate; endostome segments very narrowly perforate; cilia well developed,

nodulose, in 2’s-4’s. Spores 13-17 yw, finely papillose.
Grout, Moss Flora of North America 3: Pl. 19A.—Fig. 789-791.

Emergent (or sometimes submerged) in shallow, usually temporary pools or ditches or at the
edge of springs or brooks, in bogs, fens, and wooded swamps. Circumpolar; Greenland to Alaska,

south to Washington, Colorado, Michigan, and Pennsylvania.

PRESQUE ISLE CO.—Hammond Bay, Evergreen Beach, Presque Isle Corporation, south of Onaway
k

Calliergon giganteum, 789. Habit. 790. Leaf. 791. Alar cells.

215

A common species of considerable variability and yet easily recognized by the
irregularly pinnate arrangement of branches, dioicous inflorescences, and abruptly
differentiated alar cells. (The branches are generally remote and unequal so that the
effect is by no means frondose.) Many recent authors have erroneously included C.
giganteum in the synonymy of C. cordifolium, which has sparse and irregular
branching, autoicous inflorescences, and gradually differentiated alar cells (not
bordered above by short cells).

4. Calliergon richardsonii (Mitt.) Kindb. ex Warnst.—Rather robust plants in soft,
loose, green or yellowish masses up to 15 cm. high. Stems ascending, irregularly or
occasionally subpinnately branched; stems and branches + cuspidate at the tips. Stem
leaves erect- or wide-spreading, concave and often somewhat rolled toward the apex
when dry, about 1.7-2 mm. long, oblong- or cordate-ovate, rounded-obtuse, sometimes
radiculose at back near the apex, somewhat decurrent, entire; costa about %-% the leaf
length, usually forked at the apex; alar cells abruptly enlarged, inflated, and hyaline in
large, concave groups. Autoicous. Setae 45-60 mm. long, orange-red; capsules 2.5-3 mm.
long; annulus none; operculum apiculate; endostome segments narrowly perforate, the
cilia well developed, nodulose, in groups of 2-3. Spores 15-22 u, very finely papillose.
—n = ca.

Grout, Moss Flora of North America 3: Pl. 17A (figs. 10-12).—Fig. 792.

On soil in wet depressions or shallow pools in muskeg and, in our area, spruce-white cedar
bogs. Circumpolar; pie se but widely scattered from Greenland to Alaska and south to
Meanebe: Michigan, and Verm

CHEBOYGAN CO.—Hebron Swamp.

Intermediate in many ways between C. giganteum and C. cordifolium, this
species, rare and local in occurrence, is distinguished by an autoicous inflorescence,
irregular branching, relatively short costa, and abruptly differentiated alar cells.

5. Calliergon cordifolium (Hedw.) Kindb.—Rather robust plants in soft, loose,
green or yellow-green masses about 7-15 cm. high. Stems ascending, sparsely and
irregularly branched; stems and branches cuspidate at the tips. Stem leaves not
crowded, erect-spreading (somewhat imbricate near the stem and branch tips),
1.8-3 mm. long, cordate-ovate, rounded and somewhat cucullate at the apex, sometimes
radiculose at back near the apex, broadly decurrent, entire; costa ending near the apex;
cells at the apex shortly oval or rhombic; upper median cells linear-rhomboidal; alar
cells large, inflated, and hyaline in large, decurrent groups but not abruptly differ-
entiated. Autoicous (or rarely polygamous, the perigonial buds rarely including a few
archegonia). Setae 35-65 mm. long, orange-brown; capsules 2-3 mm. long; annulus
none; operculum apiculate; segments of endostome narrowly perforate; cilia well
developed, nodulose, in 2’s and 3’s. Spores 13-18 yw, very finely papillose.—n = 10, 21,
20-22

Conard, How to Know the Mosses, (ed. 2) fig. 205. Grout, Mosses with Hand-lens and
Microscope, Pl. 76. Jennings, Mosses of Western Pennsylvania, (ed. 1) Pl. 40, (ed. 2) Pl. 44. Welch,
Mosses of Indiana, fig. 180.—Fig. 793-795.

On various substrata in or at the margins of temporary pools and along streams, frequently
in ditches or wet depressions, often emergent or temporarily submerged, in swamps, swales, or peat
bogs. Circumpolar; Greenland to Alaska and south to Washington, Montana, Illinois, and Pennsy]-
vania.

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Calliergon richard soni.

V2
792.
Calliergon cordifolium. 793. ae 794. Leaf. 795. Alar cells.

CHEBOYGAN CO.—Reeses Bog, Carp Creek, Douglas Lake, Burt Lake, Livingston Bog, Mud
Lake. EMMET CO.—Maple River, Pleasantview Sw wamp, Amott Lake, Galloway Bog, rie
State Park. MACKINAC CO.—Bois Blanc Island, re ers Lake, Koski Pond, McNamar
Pond. PRESQUE ISLE CO.—South of Onaway State Par

This common species has often he confused with C. giganteum, from which it
differs iene in its loose and irregular, sparse habit of branching, autoicous
inflorescences, and gradually differentiated alar cells.

277
9. Calliergonella Loeske

Rather robust plants in loose, shiny, green, yellow-green, or yellow-brown tufts
or masses. Stems greenish or yellowish, erect or ascending, irregularly or subpinnately
branched; cortical cells large and hyaline; stems and branches cuspidate at the tips;
paraphyllia few, at branch bases. Stem leaves erect or erect-spreading (crowded and
imbricate at tips of stems), concave, broadly oblong-ovate to elliptic, rounded and
usually bluntly and inconspicuously apiculate at the apex, entire; costa none or short
and double; upper cells smooth, linear-flexuose; cells at insertion thicker-walled and
porose, those at the basal angles subquadrate, inflated, hyaline, and thin-walled in
conspicuous auricles. Branch leaves shorter, narrower, oblong or oblong-lanceolate,
blunt or obtuse. Dioicous; perichaetial leaves erect, long, lance-acuminate. Capsules
curved-cylindric; annulus large, of about 4 rows of cells; operculum + apiculate;
endostome with entire or narrowly perforate segments and appendiculate cilia in 3’s
and 4’s—The name is a diminutive of Calliergon, and the plants have an aspect
suggestive of that genus.

Calliergonella cuspidata (Hedw.) Loeske—Plants up to about 10cm. high. Stem
leaves 1.8-2.2 mm. long. Setae 35-50 mm. long; capsules 3-4 mm. long. Spores 17-22 uy,
finely roughened.—n = 9, 10, 11.

arlington, Mosses of Michigan, fig. 119. Grout, Mosses with Hand-lens and Microscope, fig.
167 as Cillicmon). ‘Jennings, Mosses of Western Pennsylvania, (ed. 1) Pl. 40 (as Acrocladium), (ed.
2) Pl. 44.—Fig. 796-798.

In rich fens and sedge meadows; a calciphile. Europe, North Africa, Macaronesia, Asia,
Jamaica, Argentina, and New Zealand; British Columbia to California, Nova Scotia to Wisconsin,
south to Iowa and Tennessee.

CHEBOYGAN CO.-—Reeses Bog, Mud Lake, Little Lake 16. EMMET CO.—Alanson, Cecil
Bay, Wilderness State Park, Waugoshance Point, Conway, Bear Creek Township, Lawrence Lake.
MACKINAC CO.-—St. Martins Bay, Peggley Lake. PRESQUE ISLE CO.—Evergreen Beach, Clinton
Lake, Kelsey Lake.

The leaves are crowded and convolute at the cuspidate tips of stems and
branches. The species somewhat resembles Pleurozium schreberi but grows in much
wetter habitats and has much more cuspidate stem and branch tips, more strikingly
differentiated basal leaf auricles, and large cortical cells. When growing submerged, it is
more robust and more plumose, with spreading leaves.

10. Scorpidium (Schimp.) Limpr.

Plants robust, turgid or julaceous, in loose mats in very wet places, often
submerged in shallow, stagnant water, yellow-green or red-gold, sometimes brownish or
blackish, often slimy or lime-encrusted. Stems creeping or ascending, scarcely to
irregularly or subpinnately branched. Leaves large, crowded, very concave and often
cucullate, erect or + falcate-secund, smooth or striolate and sometimes + rugose when

margins erect or incurved above, entire; costa weak, short and double, or lacking; cells
smooth, linear, those at the insertion shorter, wider, incrassate and porose, a few cells
at the basal angles + differentiated (small and subquadrate, or somewhat inflated).
Dioicous; perichaetial leaves erect, lanceolate, striate. Capsules curved-cylindric; annulus
present; endostome cilia in 2’s and 3’s.-_Named for a fancied resemblance to a scorpion
with its tail curved upward, in reference to the curved-secund leaves of S. scorpioides.

Calliergonella cuspidata. 796. Habit. 797. Leaves. 798. Alar cells.

gsi + falcate-secund, at least at the tips of stems and branches; alar = hyaline

Hewes and inflated in small and inconspicuous groups .S. scorpioides
ee loosely imbricate to spreading, not at all secund or slightly so at branch a alar
cells nett oblong and subquadrate, not hyaline, thin-walled, or inflated . 5, turgescens

1. Scorpidium scorpioides (Hedw.) Limpr.—Plants robust and _ turgid, to

nequ:

encrusted with a slimy deposit. Leaves + secund, at least at the tips of stems and
branches, often falcate-secund, + striolate and a pecans and obscurely rugose-
wrinkled when dry, concave, 2-3, rarely 4.5 mm. long, elliptic to oblong-ovate, obtuse
or acute, often apiculate; costa weak, short and double, or none; upper cells long,

209

linear-flexuose; cells at insertion oblong-linear, thick-walled, and porose, a few cells at
the extreme angles hyaline, thin-walled, + inflated. Setae 30-45 mm. long; capsules
2-3 mm. long. Spores 13-15 uw, very finely roughened.—n = 8, 10+1; 2n = 16.

Conard, How to Know the Mosses, (ed. 2) fig. 267. Darlington, Mosses of Michigan, fig. 117.
Grout, Moss Flora of North America 3: Pl. 27C.—Fig. 799-800.

A calciphile, in shallow water and often submerged among sedges at the margins of pools in
rich fens or swales. Circumpolar; Andes of South America; Greenland to Alaska, south to Montana,
Wisconsin, Ohio, and New Jersey.

CHEBOYGAN CO.—Douglas Lake, Blanchards Bog, Mud Lake, Grass Bay, Black Lake.
EMMET CO.—Arnott Lake, Ramona Beach, Sturgeon Bay Point, Cecil Bay, Wilderness State Park.
MACKINAC CO.-—Bois Blanc Island, Mackinac Island, Mud Lake. PRESQUE ISLE CO.—Hammond
Bay, Evergreen Beach, Clinton Lake.

Cx
oe

“a
oe
ay

MEGIFL-

hide

bee |
We)
No)

Scorpidium scorpioides, 799. Habit. 800. Leaves.
Scorpidium turgescens. 801. Habit. 802. Leaf.

280

Scorpidium scorpioides can be quite beautiful with its robust, fatly julaceous
stems and branches and deep, red-gold color and attractive sheen. In Great Britain, it is
said to grow sometimes in acid habitats (with Drepanocladus revolvens), but in North
America it is limited to shallow water of highly calcareous fens where it is usually
submerged, covered with a limey deposit, and, as often as not, slippery, slimy, and
partially decomposed. The very broad, secund, and somewhat rugose leaves with short
points are distinctive. The species is suggestive of a Drepanocladus, differing most
clearly, of course, in the absence of a costa and also by short leaf points. As compared
with S. turgescens, its secund leaves and absence of differentiated alar cells are helpful
characteristics.

2. Scorpidium turgescens (T. Jens.) Loeske—Rather stout or robust plants in
dense, soft, yellow-brown, + shiny masses about 5-20 cm. high. Stems erect, sparsely
branched, usually rather turgid. Leaves crowded and loosely imbricate or sometimes
rather remote and erect-spreading to spreading, not at all secund or slightly so at
branch tips, concave or sometimes cucullate, not or slightly striolate when dry,
2-3.2 mm. long, broadly oblong-ovate, obtuse or rounded-obtuse and (at least in
ounger leaves) narrowed to a small, recurved apiculus, somewhat rounded to the
insertion, costa slender, double, about 1/3 the leaf length; upper cells linear; cells at
insertion yellow, oblong-linear, incrassate, and porose, those toward the angles shortly
oblong and subquadrate, firm- or thick-walled (not hyaline, thin-walled, and inflated).
[Sporophytes very rare; not seen.]

Grout, Moss Flora of North America 3: Pl. 26B (as Calliergon).—Fig. 801-802.
A merge ee in rich fens, often emergent or temporarily submerged. (It is said to
ccur in Great B n also in acid habitats.) Circumpolar; Ecuador; Greenland to Alaska, south to
British Columbia, ia. Michigan, Ontario, and Newfoundland.
MACKINAC CO.—Bois Blanc Island.
e plants are stoutly julaceous with broad, concave leaves erect or + spreading,

Th
rounded-obtuse and often abruptly apiculate. It has the look of a Calliergon, but the
costa is double, and the alar cells are scarcely inflated.

BRACHY THECIACEAE

Plants slender to robust, in loose or dense mats or tufts, often + shiny. Stems
creeping to erect-ascending, freely and irregularly to pinnately branched; paraphyllia
mostly lacking. Leaves in many rows, appressed os erect-spreading, occasionally +
secund, mostly plicate, broadly ovate to lanceolate, + long-acuminate (or rarely blunt
or rounded at the apex); costa single and well developed, mostly ending below the
apex, frequently projecting at the tip as a dorsal spine; cells prosenchymatous, mostly
elongate, rhomboidal to linear-flexuose, smooth or rarely papillose at back because of
projecting upper angles, laxer and often porose at base, mostly subquadrate at the
basal angles, sometimes forming well-marked alar groups. Perichaetia lateral; peri-
chaetial leaves differentiated. Setae + elongate, frequently roughened; capsules inclined
to horizontal or rarely erect and symmetric, mostly rather short, ovoid or oblong-
cylindric and asymmetric, + curved when dry and empty, the neck short and
inconspicuous, generally stomatose; annulus usually poorly differentiated; operculum
conic, blunt, acute, or rostrate; peristome double, the 16 teeth lance-subulate, generally
red-brown, mostly cross-striolate below, trabeculate at back; endostome generally free

281

and about as long as the teeth, the basal membrane usually high, the segments keeled,
usually lance-subulate, the cilia well developed or rarely rudimentary or lacking. Spores
spherical. Calyptra cucullate, usually naked.—_As compared with the Amblystegiaceae,
which generally slender, curved capsules with yellow-brown exostomes, the

Brachytheciaceae characteristically have short capsules with dark, red-brown peristome
teeth

The generic concepts have recently been revised by Robinson (Bryol. 65: 73-146.
1962), but I take a more traditional view, more or less that presented in Die
naturlichen Pflanzenfamilien.

1. Leaves broadly oblong-ovate and rounded at the apex, often appearing apiculate becau
of abruptly incurved upper margins; costa short and double
1. Leaves ovate to lanceolate, blunt to acuminate; costa sin
2. Stems erect, pinnately branched, conspicuously tomentose nearly to the apex; leaves
lanceolate, gradually acuminate, strongly plica , GENEL) RTE
: + ascending, irregularly ss subpinnately branched, occasionally
dendroid, not tomentose; leaves ovate to eae bluntly acute or Bee
acuminate, mostly striate or moderately plicate when
eaves acute or obtuse; ae cells or branch ence short and rhomboid-hexagonal,
“ differentiated from median c Eurhynchium
3. Leaves acuminate; apical in of branch leaves sionenic’ not much different from
median cells
4. Leaf cells eta at back because of projecting cell ends 3. Bryhnia
4. Leaf cells smooth
5. Leaves a concave, oes -ovate, abruptly filiform-acuminate 4. Cirriphyllum
5. Leaves not conspicuously concave, variously acuminate but not abruptly filiform
6. Operculum obliquely rere leaves smooth, ovate or ovate-lanceolate, usua
somewhat complanate
6. Operculum blunt or acute to apiculate; leaves mostly +
not or rarely obscurely complanate
7. Plants small and slender; leaves smooth, serrulate all around, especially at the oa
angles; costa subpercurrent, indistinct at the midleaf, often bearing rhizoids
subcylindric brood bodies at back above he Rhynchostegielt
7. Plants small or mostly medium-sized; leaves + plicate, entire or serrulate but
conspicuously toothed at the basal angles; costa ending above the leaf eles or
rarely subpercurrent, distinct at the middle, not bearing rhizoids or brood bodies
at back above

. Pleurozium

6. panenee
plicate, mostly lanceolate,

2. Brachythecium
1. Tomenthypnum Loeske

Rather robust plants in loose or dense, yellow-green to golden-brown tufts, rigid
and often glossy when dry. Stems erect, pinnately branched, conspicuously brown-
tomentose nearly throughout; branches horizontal and straight. Stem and branch leaves
similar, stiffly erect-spreading or occasionally + secund, strongly plicate, long-lanceolate

3/4-4/5 up the leaf; upper cells long-linear, rather thick-walled, smooth; cells at
insertion shorter, broader, and porose, the alar cells scarcely differentiated. Dioicous,
perichaetial leaves erect, long, lance-subulate, plicate or striate. Setae long, slender,
reddish, smooth; capsules orange-brown, strongly inclined to horizontal, oblong-
cylindric from a short neck, curved and asymmetric, constricted below the mouth
when dry; annulus of 3 layers of cells; operculum conic, acute; stomata in the neck;
peristome teeth lanceolate, dark, red-brown, cross-striolate below, papillose above
bordered, trabeculate; endostome yellow-brown, finely papillose, as long as the teeth,
with a high basal membrane, keeled, perforate segments, and well-developed, nodulose
ilia in 3’s and 4’s. Calyptrae naked._The name refers to the tomentum of felted hairs
characteristic of a moss otherwise suggestive of Hypnum (in a broad sense).

Tomenthypnum nitens (Hedw.) Loeske—Plants 5-15 cm. high. Stem leaves
3-4 mm. long, radiculose at the back of the costa at base; costa very slender and

202

807

Tomenthypnum nitens. 803. Habit. 804. Leaves. 805. Upper cells of le
Brachythecium velutinum. 806. Habit. 807. Branch leaves. 808. Upper ok of branch leaf.

difficult to see because of plications. Setae 27-47 mm. long; capsules 2-3 mm. long.
Spores 13-17 w, very finely papillose.—n = 10, 12
Darlington, Mosses of Michigan, fig. 127 (as Cae reenaa), Grout, Mosses with Hand-lens
and eeu. fig. 157 (as Camptothecium).—Fig. 8 5.
common in the North and characteristic of black spruce muskegs and fens, frequent in

Vv
the Great Lakes area in swampy, calcareous habitats such as Thuja swamps and open, marly bogs.
Circumpolar; Greenland to Alaska, south to Colorado, Wisconsin, and Pennsylvania.

CHEBOYGAN CO.—Reeses Bog, Gorge of Carp Creek, Iron Bridge, Mud Lake, Grass B

EMMET CO.-Cecil Bay, Conway Bog, Lawrence Lake. MACKINAC CO.—Mackinac Island.
Hendricks. PRESQUE ISLE CO.—Evergreen Beach, Hammond Bay, Mast Point, Kelsey Pond,
Clinton Lake.

A characteristic moss of rich fens where it is associated with Sphagnum
warnstorfii, Campylium stellatum, and other pronounced calciphiles, recognized by

acid habitats than the var. nitens, at the tops of Sphagnum hummocks with S. fuscum,
for example.

283
2. Brachythecium BSG

Plants slender to moderately robust, in green, yellowish, golden, or whitish-green
mats, often somewhat shiny. Stems creeping or curved and decumbent, sometimes
erect-ascending, irregularly to subpinnately branched, often stoloniform at the ends.
Stem and branch leaves usually somewhat differentiated: Stem leaves crowded
erect-spreading or spreading, occasionally subsecund, + concave, mostly + plicate; ovate
to lanceolate, usually long-acuminate, rarely short-pointed, short- to rather long-
decurrent; margins mostly plane, finely toothed all around or only toward the apex, or
sometimes entire; costa single, usually extending well up the leaf but rarely subpercur-
rent; cells smooth, mostly elongate, rhomboidal to oblong-linear, shorter and laxer at
the base and usually subquadrate or oblong-hexagonal at the basal margins, rarely
well-differentiated in + conspicuous alar groups. Branch leaves mostly shorter and
narrower, with costa weaker and usually shorter. Perichaetial leaves erect or squarrose
rom a sheathing base, slenderly long-acuminate. Setae + elongate, smooth or
roughened (sometimes only in the upper half); capsules inclined to horizontal or rarely
erect, typically rather short and broad, oblong-ovoid and asymmetric, sometimes longer
and cylindric, curved (especially when dry and empty), smooth; annulus usually
differentiated; operculum conic or convex-conic, blunt or acute; stomata in the short
neck; peristome teeth brown to red-brown, cross-striolate below, papillose at the tips,
trabeculate; endostome free, the basal membrane high, segments lance-acuminate,
narrowly perforate or gaping along the keel, cilia well developed and nodulose to
appendiculate (or rarely rudimentary or lacking). Calyptrae naked.—_The name means
short capsule; in most species of Brachythecium the capsule is relatively short and
thick as compared with many other Hypnum-like mosses, and Ambiystegium especially.

1. Costa disappearing in the acumen to percurrent
2. Plants small; branch leaves 0.6-0.9 mm. long; costa ending in the acumen; upper cells
3-5:1, see aa to shortly oblong-rhomboidal; setae 7-12 mm. long, rough
throughou 28 B. reflexum
2. Plants oe branch leaves 1-1.6 mm. long; costa percurrent or nearly so; upper cells
longer, about 5-8:1, oblong-rhomboidal; setae 8-22 mm. long, lightly papillose above
3. B. populeum
1. Costa ending well below the apex
3. Branch leaves entire or essentially so 8. B. turgidum
3. Branch leaves serrulate
4. Stems + dendroid; leaves (especially of stems) broadly and noticeably decurrent
. B. rivulare
4. Stems spreading and irregularly to subpinnately branched; leaves not or only slightly
decurrent
5. Plants small and slender
6. Plants soft and silky; branches not terete; leaves falcate-secund; quadrate alar cells
few; autoicous; setae rough throughout; capsules asymmetric and horizontal to
pendulous 1. B. velutinum
6. Plants coarser; branches terete or subjulaceous when dry, not falcate- sie ss eerie
alar cells numerous; dioicous; setae smooth; capsules nearly erect and sym

Ps a acuminatum
5. Plants medium-sized to rather robust

8. Dioicous; alar cells small, quadrate, rather opaqu 6. B. oxycladon
8. Autoicous; alar cells subquadrate or hot eSbIOne not particularly small, rather lax
and + pellucid 7. B. salebrosum
7. Setae rough, at least above; leaves smooth or sometimes indistinctly plica
9. Setae papillose only above a rarely smooth throughout); branch ae concave,
often homomallous, 1.3-2 mm. long 4. B. plumosum
9. Setae papillose Fea res Guar leaves not particularly concave, not homo-
mallous, 2-2.5 m

&
10. Branch leaves spreading, loosely complanate when dry, twisted at the apex; costa

284

sees te and usually projecting as a ape spine at back; margins + serrate
early all around; cilia of endostome appendiculate 9. B. curtum

10. Branch leaves erect, only se ea ae at the apex; costa smooth, rarely

projecting as a minute dorsal spine; margins less strongly toothed, serrulate onl
above or sometimes nearly to the base; cilia nodulose 10. B. rutabulum

Brachythecium velutinum (Hedw.) BSG—Small plants in dark- or yellow-
green, rather soft and silky mats, irregularly to subpinnately branched. Stem and
branch leaves similar. Branch leaves usually rather widely spaced, loosely erect- to
wide-spreading and + falcate-secund (especially at branch tips) when dry, erect-spread-
ing when moist, about 1-1.5 mm. long, narrowly to broadly lanceolate, gradually
acuminate, usually twisted at the apex, shortly decurrent, not or only faintly plicate;
margins plane, serrulate; costa extending well above the middle, serrulate at back
above; median cells linear, 8-12:1; quadrate alar cells few. Autoicous. Setae 7-15 mm.
long, brown or red-brown, rough throughout; capsules about 1.5-2 mm. long, oblong-
cylindric, horizontal or somewhat pendulous, curved and asymmetric, + contracted
below the mouth when dry; annulus of 2 rows of cells; operculum bluntly conic; cilia
of endostome nodose, in groups of 2 or 3. Spores 10-13 4, smooth or very minutely
roughened.—n = 10, HH, Te

Conard, How to Know the Mosses, (ed. 2) fig. 249. Grout, Mosses with Hand-lens and
essa PI. 66. Jennings, Mosses of Western Sens (ed. 1) Pl. 51, (ed. 2) Pl. 56.—Fig.
806-808.

n soil, sometimes on rock or bark at base of trees, in dry or rather moist woods, often on
sandy forest trails. Circumpolar; British Columbia to Labrador, south to Oregon, Montana,
Michigan, and New York.

CHEBOYGAN CO.~—Mullett Lake, Little Lake 16, Weber Lake. EMMET CO.—Pellston Hills,
Wycamp Lake, Big Stone Bay. MACKINAC CO.—Caffey Corner. PRESQUE ISLE CO.—Loon Lake.

This is our smallest Brachythecium. It is scarcely velvety, as the name suggests,
but as compared with B. reflexum, it is soft and silky, with loosely falcate-secund
leaves. It is characteristically found on light soil of forest trails, especially in dry, aspen
stands or in moister hardwood forests, such as beech-maple communities.

2. Brachythecium reflexum (Starke ex Web. & Mohr) BSG—Small, dark- or
yellow-green plants in loose or dense mats. Stems pinnately branched; branches terete.
Stem leaves about 1-1.2 mm. long, deltoid-ovate, abruptly narrowed to a flexuose-
spreading acumen, rounded to the insertion, excavate at the basal angles and broadly
decurrent. Branch leaves loosely erect to imbricate (and sometimes slightly homo-
mallous) when dry, erect-spreading when moist, about 0.6-0.9 mm. long, lanceolate,
gradually acuminate, sometimes + twisted at the apex, decurrent; margins serrate
above, serrulate nearly to the base, sometimes slightly reflexed; costa ending in the
acumen, serrulate at back at or near the tip; upper cells short, 3-5:1 or rarely
somewhat longer, oblong-hexagonal or shortly oblong-rhomboidal; alar cells short-
sedan ng or subquadrate. Autoicous. Seta 7-13 mm. long, rough throughout, orange-

own to dark-red; capsules 1.5-1.8 mm. long, strongly inclined to horizontal, shortly
ae asymmetric, not constricted below the wide mouth, brownish, becoming
blackish with age; annulus of 2 rows of cells, fragmenting: operculum conic or
convex-conic, mammillate to stoutly apiculate; cilia of endostome appendiculate, in 2’s
and 3’s. Spores 11-16 uy, finely papillose.—n = 10, 11, 15, 20

arlington, Mosses of Michigan, fig. 126. Grout, Mosses with Hand-lens and Microscope, fig.
148. a 809-812.

285

On rocks, soil, logs, stumps, exposed roots, and bases of trees in aes dry to moist woods.
Grccnate Alaska to Labrador, south to Oregon, Michigan, and Virgin

CHEBOYGAN CO.—Douglas Lake, Carp Creek, Colonial Point, Mud Lake, on Bay, Little
Lake 16. EMMET CO.—Maple River Township, Peliston Dunes, Pellston Hills, Wycamp Lake, Cross
Village, Five Mile Creek. MACKINAC CO.—Mackinac Island, Gros Cap, Ozark, Little Dollar Lake.
PRESQUE ISLE CO.—Mast Point, Loon Lake.

Most common on bark at base of trees (often with Hypnum pallescens,
eee adnatum, and Platydictya subtile), the plants are small and dark-green,
with stem leaves abruptly acuminate, rounded to the insertion, and broadly decurrent;
the costa of branch leaves is serrulate at back at or near the tip.

3. Brachythecium populeum (Hedw.) BSG—Plants relatively slender, in dense,
green or yellow-green mats. Branches slender and terete. Branch leaves erect and not or
only slightly plicate when dry, erect-spreading when moist, 1-1.6 mm. long, lanceolate,
slenderly acuminate; margins serrulate in the upper 4, revolute below; costa disappear-
ing at or near the apex; upper cells firm-walled, oblong-rhomboidal; alar cells
subquadrate in numerous rows. Stem leaves ovate-lanceolate, about 1.4-2 mm. long.
Autoicous. Setae 8-22 mm. long, lightly roughened above, reddish; capsules 1-2 mm.
long, strongly inclined to horizontal, asymmetric, oblong-ovoid; annulus of 1-2 rows of
cells; operculum conic, acute or apiculate; cilia of endostome in groups of 2 and 3.
Spores 12-14 yw, very finely papillose —n = 9, 10, 10+1, 11.

Conard, How to Know the Mosses, (ed. 2) fig. 246e. Grout, Mosses with Hand-lens and
Microscope, fig. 150.—Fig. 813-814.

n rock or on over rocks. Europe; Azores; Asia; British Columbia; Nova Scotia to Manitoba
and me to Virgini

CHEBOYGAN CO.—Colonial Point. EMMET CO.—Five Mile Creek. MACKINAC CO.-—Little
Dollar Lake.

Relatively slender plants, usually on rocks, with terete branches and erect-spread-
ing leaves tapered to a long, narrow acumen in which the costa disappears.

4. Brachythecium plumosum (Hedw.) BSG—Plants of medium size, green or,
more often, yellow to brownish, shiny. Leaves often homomallous, concave, smooth or
occasionally faintly striolate, 1 2: 2 mm. long, lanceolate or ovate Janceolate, acuminate,
serrulate above; costa about 2/3 the leaf length; upper cells linear; alar cells
subquadrate in small groups. Autoicous. Setae 7-16 mm. long, dark-red, papillose above
(or occasionally smooth throughout); capsules 1.3-2 mm. long, oblong-ovoid, asym-
metric, inclined to horizontal, dark-brown; annulus of 2 rows of cells; operculum conic
and acute to apiculate or short-rostrate; cilia of endostome appendiculate, in groups of
1-3. Spores 13-18 wy, nearly smooth.—n = 10, 10+1, 11

Conard, How to Know the Mosses, (ed. 2) fig..246 (as B. flagellare). Grout, Mosses with
Hand-lens and pues fig. 149. Jennings, Mosses of Western Pennsylvania, (ed. 1) Pl. 52 & (ed.
2) Pl. 57 (as B. flagellare and var. homomallum). Welch, Mosses of Indiana, fig. 153 (as B
flagellare).—Fig. 815.

On moist rocks (usually in or near streams), rarely on soil. Nearly aaa British
Columbia to Oregon and Arizona, Labrador to Manitoba and south to Georgia and Arkan

CHEBOYGAN CO.—Gorge of Carp Creek, Mill Creek. MACKINAC CO.—Ozark.

286

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Brachythecium reflexum, 809. Habit. 810. Stem leaves. 811. Branch leaves. 812. Upper cells
h

of branch leaf.
Brachythecium populem. 813. Stem leaf. a Branch leaves.

oe plumosum, 815. Branch
chythecium acuminatum. 816. ce ee 817. Upper cells of branch leaf. 818. Alar
cells of sent leaf. 819. Capsule.

The plants are characteristic of moist rocks along streams. Useful characteristics
include the yellowish or brownish color, and concave, smooth, and often homomallous
leaves.

5. Brachythecium gran (Hedw.) Aust.—Slender, green or yellowish, shiny

ats. reeping, closely branched; branches usually erect,

plants in extensive m ems
short, terete and usually an when dry. Stem and branch leaves similar,

287

usually crowded, erect when dry, erect-spreading or spreading when moist, somewhat
biplicate or sometimes nearly smooth, 1-1.5 mm. long, ovate or ovate-lanceolate,
gradually or abruptly acuminate, slenderly acute, often twisted at the apex; margins
teflexed below, serrulate in the upper half; costa about 2/3 to 3/4 the leaf length, often
ending in a minute tooth at back; upper cells linear-rhomboidal (shorter at the apex of
the leaf); alar cells subquadrate in several to numerous rows. Dioicous. Setae 7-20 mm.
long, dark-red, smooth; capsules erect and symmetric or slightly curved, cylindric,
1.5-2.2, rarely 3mm. long; annulus none; operculum conic, often obliquely apiculate;
stomata in the short neck; cilia of endostome none or rudimentary. Spores 15-17 yp,
very finely papillose or nearly smooth.

arlington, Mosses of Michigan, fig. 125 (as Chamberlainia cyrtophylla). Grout, Mosses with
Hand-lens fe Microscope, Pl. 65, ae fig. 15 os B. cyrtophyllum). Jennings, Mosses of Western

Pennsylvania, (ed. oe ee 48, 49 (as B. acuminatum & B. cyrtophyllum), (ed. 2) Pl. 53, 54 (as
Chamberlainia acuminata & B. cyrtophylla). Welch, Mees of Indiana, fig. 144 (as Chamberlainia),
145 (as C. ren Fig. 816-819.

On bark at base of trees and less frequently on rocks or soil. Saskatchewan to New
Brunswick and south to Texas and Florida.

CHEBOYGAN CO.—Reeses Bog, Polypody Bluff (near Douglas Lake). EMMET CO.—Pellston
Hills.

The plants are a small version of B. oxycladon, with branches terete when dry
and with rather numerous, small, quadrate alar cells. The capsules are nearly erect and
symmetric but not too different from the longish, somewhat inclined capsules of B.
oxycladon and scarcely justify a generic separation as Chamberlainia, in Grout’s
original concept. The genus Brachythecium in the inclusive and classical sense of the
Bryologia Europaea seems to me natural, recognizable, and “‘Linnaean,” and I see no
reason to re-apportion some of the species as Chamberlainia and Bryhnia in the
emended sense proposed by Robinson (Bryol. 65: 73-146. 1962).

Brachythecium acuminatum var. cyrtophyllum (Kindb.) Redf. ex Crum n. by
[B. cyrtophyllum Kindb., Ottawa Nat. 4: 63. 1890. B. acuminatum ssp. seer ie
(Kindb.) Kindb. ex Macoun & Kindb., Cat. Canad. Pl. 1: 191. 1892. B. acuminatum
var. cyrtophyllum (Kindb.) Redf., Ann. Mo. Bot. Gard. 59: 79.1972, comb. illeg.| Often
hidden in synonymy, this variety is recognizable by leaves shorter, broader, and more
concave, with shorter tips (acute to short-acuminate) and shorter cells. Redfearn
erroneously attributed the combination to Welch and provided no reference to a
basionym which would have validated the transfer in this name. The variety is
widespread, probably ranging with the species proper.

6. Brachythecium oxycladon (Brid.) Jaeg. & Sauerb.—Plants in extensive, some-
what shiny, yellow-green or yellow-brown mats. Stems creeping, freely branched;
branches usually closely foliate and + terete. Stem leaves up to 2.2 mm. long. Branch
leaves erect or erect-spreading, concave, plicate, 1.5-2 mm. long, lanceolate, gradually
acuminate, serrulate nearly all around; costa 2/3-3/4 the leaf length; upper cells
linear-rhomboidal; alar cells subquadrate, relatively small and somewhat grayish-opaque.
Dioicous. Setae 8-27 mm. long, usually reddish, smooth; capsules suberect or inclined,
curved, oblong-cylindric, about 2-3 mm. long; annulus none; operculum conic, acute or
+ acuminate; cilia paired. Spores 11-16 y, finely papillose.

ard, How to Know the Mosses, (ed. 2) fig. 241a-c,e. Grout, Mosses with Hand-lens and

Con
Microscope, PL 63. Jennings, Mosses of Western Pennsylvania, (ed. 2) Pl. 69. Welch, Mosses of
Indiana, fig. 150.—Fig. 820-822.

288

On shaded soil, rocks, rotten wood, and bases of trees, often in rather dry or disturbed
habitats. Europe; eastern North America from southeastern Canada to Minnesota and Nebraska,
south to North Carolina and Missouri.

CHEBOYGAN CO.—Wolffs Bog, Mud Lake. EMMET CO.—Maple River, Larks Lake, Cross
Village, Middle Village, Five Mile Creek, MACKINAC CO.—Mackinac Island, Sand Lake, Ozark,
Caffey Corner, McKays Creek.

Very common and most likely to be confused with B. salebrosum, which is
autoicous and has larger, laxer, more transparent alar cells. (The Bryologia Europaea
pictures which Grout reproduced in his Mosses with Hand-lens and Microscope describe
the areolation more eloquently than I could hope to do in words.)

Brachythecium digastrum C. M. & Kindb. ex Macoun & Kindb. is a smallish
version of B. oxycladon. It has broadly decurrent stem leaves, especially in forms with
leaves widely spaced. Its branch leaves are narrowed to broad, acute or shortly
acuminate, blunt, serrate points. The cells near the leaf tip are short, and the costa is
somewhat toothed at back. Unlike B. oxycladon, it appears to be autoicous. It is
variable and poorly understood, though seemingly widespread in eastern North
America.

7. Brachythecium salebrosum (Web. & Mohr) BSG—Rather robust plants in
loose, somewhat shiny, green, yellowish or brownish mats. Stems creeping or +
ascending, freely branched. Stem leaves 2-2.5 mm. long; branch leaves loosely erect or
erect-spreading, sometimes loosely falcate, usually strongly plicate, 1.8-2 mm. long,
lanceolate or ovate-lanceolate, gradually long-acuminate; margins irregularly recurved
below, serrulate above; costa about 2/3 the leaf length; upper cells linear, those at the
insertion shorter and broader with thicker, pitted walls, the alar cells subquadrate and
rather thin-walled, not opaque. Autoicous. Setae 10-27 mm. long, yellowish or reddish,
smooth; capsules rather shortly oblong-cylindric, curved and asymmetric, inclined to
nearly horizontal, 2-2.5 mm. long; annulus poorly developed in 1 row; operculum
conic, acute or apiculate; cilia of endostome (1)2. Spores 13-18 w, finely papillose.—n =
10+1, 13, 15, 20

sana How to Know the Mosses, (ed. 2) fig. 243. Grout, Mosses with Hand-lens and
Microscope, Pl. 62. Jennings, Mosses of Western ne (ed. 1) Pl. 49, (ed. 2) Pl. 54. Welch,
Mosses of Indiana, fig. 146.—Fig. 823-827.

Re shaded soil, stones, bases of trees, and logs, often in rather dry and disturbed places
(com in lawns). Circumpolar; Tasmania and Kerguelen; Alberta and eastward across the
ee widespread in eastern North America.

CHEBOYGAN CO.—Douglas Lake, Reeses Bog, Grapevine Point, Colonial Point, Maple River,
Ellis Township, Mill Creek. EMMET CO. —Carp Lake, Good Hart, Cecil Bay, ee Stone Bay.
MACKINAC CO.—Mackinac Island, McKays Crock. PRESQUE ISLE Co, —Hammond Bay

Very common in disturbed places and most likely to be confused with B.
oxycladon. The important differences are well illustrated by the Bryologia Europaea
illustrations reproduced in Grout’s Mosses with Hand-lens and Microscope (pp. 272 and
275). B. oxycladon is dioicous and has small, rather opaque, quadrate alar cells. B.
salebrosum is autoicous and has a more open areolation. of larger, laxer, subquadrate
cells at the basal angles.

Brachythecium oxycladon fo. falcatum Grout, known from several local areas,
including the type locality at Douglas Lake, seems to me a small form of this species
(somewhat resembling B. velutinum because of distinctly falcate-secund leaves).

289

AE
—
aa

Se
) ; ) F —_ =
NS .
ip

SOX

nines
Male
wae

Brachythecium oxycladon, 820. Branch leaf. 821. Alar cells of branch leaf. 822. Capsule.
Brachythecium iin 823. Habit. 824. Branch leaf. 825. Upper cells of branch leaf.

826. Alar cells of branch leaf. 827. Capsule.
Brachythecium sea 828. Branch leaf.

Brachythecium campestre (C. M.) BSG is much like B. salebrosum (although it
may be more closely related to B. curtum). The branch leaves are erect-spreading and
usually somewhat falcate-secund, not at all complanate, serrate above and serrulate
nearly to the base; the alar cells are subquadrate and shortly oblong; the costa is
toothed at back above; the inflorescences are autoicous and the setae lightly and
inconspicuously papillose. It has not been found in this part of Michigan.

8. Brachythecium turgidum (C. J. Hartm.) Kindb.—Stout plants in loose, shiny,
golden-green or golden-brown tufts. Stems erect-ascending, irregularly and sparsely

290

branched; branches stout and + turgid or subjulaceous, pointed. Stem and branch
leaves similar, loosely erect or erect-spreading, moderately plicate, 2.5-3 mm. long,
ovate-lanceolate, rather abruptly narrowed to a long, slender acumen, somewhat
decurrent; margins narrowly reflexed below, entire or occasionally slightly serrulate
above; costa ending above the leaf middle; upper cells linear; alar cells laxly
subquadrate or short-oblong. Autoicous. Setae 18-25 mm. long, reddish, smooth,
capsules about 2 mm. long, oblong-cylindric, asymmetric, nearly horizontal; annulus of
1 row of cells; operculum conic, apiculate; cilia of endostome 2(3). Spores 11-20 y,
finely papillose.—n = 10

Fig. 828.

On t soil and rocks near sere: pools, or waterfalls; probably limited to calcareous
habitats. Circumpolar; scattered and rare n the estern Hemisphere, from Greenland and Labrador
to Alaska and south to Quebec and Michigan .

MACKINAC CO.-—Bois Blanc Island. PRESQUE ISLE CO.—Mast Point.

Similar to B. salebrosum but larger, somewhat turgid, and growing in wet,
calcareous places.

9. Brachythecium curtum (Lindb.) Limpr.—Plants of moderate size, in loose,
yellow or light-green, shiny mats. Stems ascending or arched, irregularly branched.
Stem leaves ovate-lanceolate, acuminate, somewhat decurrent, with fairly numerous,
lax, oblong cells at the basal angles. Branch leaves not crowded, spreading and loosely
complanate wet or dry, not plicate, concave, 2-2.5mm. long, oblong-lanceolate,
gradually acuminate, twisted at the apex, slightly decurrent; margins plane, sharply
serrulate or, more often, serrate nearly all around; costa slender, about 5/6 the leaf
length, minutely toothed at back above and usually ending in a small dorsal tooth;
cells linear-flexuose, a few at the basal angles lax and oblong to subquadrate.
Autoicous; perichaetial leaves sheathing at base, with long, wide-spreading acumina.
Setae 10-30 mm. long, brownish or orange, becoming red, papillose-roughened through-
out; capsules 1.5-2 mm. long, horizontal or subpendulous, asymmetric, oblong, brown
or blackish-brown with age; annulus of 2 rows of cells; operculum conic, acute; cilia of
endostome appendiculate, in groups of 2 and 3. Spores 11-15 uw, smooth or minutely
roughened.—n =

Fig. 829.

On various substrata, often in bogs. Europe and eastern North America from Nova Scotia to
Minnesota and North Carolina

CHEBOYGAN CO.—Reeses Bog, Gorge of Carp Creek, eas Bog, Mud Lake. EMMET
CO.—Pellston Hills, eae MACKINAC CO.—Hendricks, Hesse

If my concept is correct, all American reports of B. starkei (Brid.) BSG can be
referred here. Though related to that species, B. curtum has a distinctive appearance
because of loosely spreading, more or less complanate, and more gradually tapered
leaves. The leaves are twisted at their tips.

10. Brachythecium rutabulum (Hedw.) BSG—Plants moderately robust, glossy,
green or yellow. Stems arched, freely and irregularly to subpinnately branched;
branches ascending. Stem leaves 2-3 mm. long, ovate, acuminate, broadly decurrent,
with numerous, large, oblong cells at the basal angles. Branch leaves loosely erect,

291

smooth or only slightly plicate when dry, 2-2.5 (rarely 3)mm. long, oblong- to
ovate-lanceolate, slenderly acuminate, with the apex occasionally twisted, not or only
slightly decurrent; margins distinctly serrulate in the upper half or less, sometimes
nearly to the base; costa very slender, about 2/3-3/4 the leaf length, smooth at back
and only rarely projecting as a minute, inconspicuous dorsal spine; cells long-linear,
flexuose, a few at the basal angles enlarged and oblong. Autoicous; perichaetial leaves
elongate, sheathing at base, with long, slender, somewhat serrulate, wide-spreading
acumina. Setae 13-30mm. long, red-brown or red, papillose-roughened throughout;
capsules 2.5-3 mm. long, horizontal, somewhat curved, asymmetric, oblong, dark- or
red-brown; annulus of 2 rows of cells; operculum conic, acute; cilia of endostome
nodulose, in groups of 2-4. Spores 13-20 yw, very finely roughened or nearly smooth.—n
= 5,6, 10, 10+1, 11, 12, 13, 20, 22; 2n = 20.

Conard, How to Know the Mosses, (ed. 2) fig. 247. Grout, ae with Hand-lens and
Microscope, Pl. 64, Moss Flora of North America 3: Pl. 8 (fig. 3), var. flavescens. he
Mosses of Western Pennsylvania, (ed. 1) Pl. 50, (ed. 2) Pl. 55. Welch, Mosses of Indiana, fig.
151.—Fig. 830.

On soil, rocks, logs, and bark at base of trees in moist, shady places. Nearly cosmopolitan;
eastern United States from New England to Michigan, south to Missouri, Kentucky, and Ne
Jersey.

CHEBOYGAN CO.—Reeses Bog, Burt Lake, Gorge of Carp Creek, Smiths Bog, Wolffs Bog,

Livingston Bog, Mill Creek, Mullett Lake. EMMET CO.—Five Mile Creek, Cecil Bay. MACKINAC

O.—Bois Blanc Island, Mackinac Island, St. Martins Point, Caffey Corner. PRESQUE ISLE
CO.—Presque Isle Lighthouse.

A troublesome species often resembling B. salebrosum, and like it autoicous, but
having a roughened seta and leaves less plicate and usually more glossy-green. It grows
in wetter habitats, too. It apes B. rivulare because of rather decurrent leaves but differs
in its autoicous inflorescences and non-dendroid form of growth.

11. Brachythecium rivulare BSG—Plants fairly robust, in extensive, pale- o
whitish- to yellow-green, + shiny mats. Primary stems creeping; secondary stems
typically erect-ascending and irregularly dendroid. Stem leaves 1.8-2.5 mm. long,
broadly oblong-ovate or gradually or rather abruptly short-acuminate, with alar cells
conspicuously enlarged and short-oblong in broadly decurrent groups. Branch leaves
erect or erect-spreading, concave, often slightly plicate, 1-1.5, sometimes 2 mm. long,
rounded-ovate to ovate-lanceolate, sharply or bluntly acute to short-acuminate, some-
what decurrent; margins serrulate above; costa about 2/3 the leaf length; upper cells
linear; alar cells lax or + enlarged. Autoicous (and apparently also dioicous). Setae
15-25 mm. long, red-brown, rough throughout; capsules asymmetric, inclined to
horizontal, 2-2.5 mm. long; annulus of about 2 rows of cells; operculum conic,
apiculate; cilia 2-3. Spores 15-22 yw, finely papillose—n = 6, 10+1, 11, 12, 13, 16

Conard, How to Know the Mosses, (ed. 2) fig. 248. Grout, Mosses with Hand-lens and
Microscope, fig. 147. Jennings, Mosses of Western Pennsylvania, (ed. 1) Pl. 51, (ed. 2) Pl. 56.
Welch, Mosses of Indiana, fig. 152.—Fig. 831-833.

n shaded soil, rocks, and other substrata in seepy places around springs and near brooks.
Circumpolar; Kerguelen; across Canada and the northern United States, south to Virginia and
Missouri in the East.

CHEBOYGAN CO.—Reeses Bog, Hermits Bog, Gorge of Carp Creek, Iron Bridge, Burt Lake,
Mill Creek, Grass Bay. EMMET CO.-—Sherett Lake, Wycamp Lake, Pellston Hills, Pleasantview
Swamp, Conway Bog. MACKINAC CO.—Mackinac Island, Cut River, Epoufette. PRESQUE ISLE
CO.—Hammond Bay, Evergreen Beach, Rainy River Falls.

Zo2

833

832

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Brachythecium curtum, 829. Branch leaves.
Brachythecium rutabulum, 830. Branch leaf.
Brachythecium rivulare, 831. Habit. 832. Stem leaf. 833. Branch leaves.

Brachythecium rivulare is recognized by its seepy habitat and pale, soft,
somewhat dendroid habit of growth. The alar cells are greatly differentiated in broad
decurrencies, especially on stem leaves. Being dioicous, it can be distinguished sexually
from occasional puzzling forms of B. rutabulum.

€ species was reported, by Aravilla Taylor, to be involved in the deposit of
bog iron ore in Indiana (Bryol. 22: 38-39. 1919). I am dubious of this, as I can think
of no obvious physiological method comparable to the carbon dioxide exchange
resulting in tufa deposition. Flowers (Bryol. 36: 26-27. 1933) reported a calcareous
tufa in Utah which had been deposited by a moss suggesting Brachythecium rivulare.

3. Bryhnia Kaur.

Plants slender to medium-sized, in loose or dense, green, yellowish, or brownish,
somewhat shiny mats. Stems ascending from a creeping base, subpinnately branched or

293

somewhat dendroid. Stem leaves somewhat larger than branch leaves, broadly ovate to
ovate-lanceolate, gradually or abruptly short- or fairly long-acuminate, decurrent.
Branch leaves ovate-lanceolate to lanceolate, acute or acuminate, decurrent, + striate
when dry; margins serrulate nearly all around; costa ending below the apex; upper cells
shortly oblong-rhomboidal to linear-rhomboidal, papillose at back because of projecting
upper ends, longer and broader at base, not much differentiated at basal angles.
Dioicous; perichaetial leaves clasping at a broader base and ending in a flexuose,
wide-spreading subula. Setae elongate, red, rough throughout, capsules strongly inclined
to horizontal, oblong-cylindric, somewhat curved, asymmetric, smooth; annulus narrow
but well differentiated; operculum conic and apiculate to obliquely short-rostrate;
stomata at base of the capsule; peristome teeth brown or red-brown and cross-striolate
below, pale and papillose at the tips, bordered, trabeculate; endostome pale yellow-
brown, papillose, with a high basal membrane, keeled, perforate segments, and
nodulose cilia in groups of (1)2-3. Calyptrae naked or with a few scattered hairs.—
Named for Nils Bryhn, a Norwegian bryologist.
Leaves + spreading, ovate or ovate-lanceolate, acute or broadly acuminate, twisted at the

apex (especially at branch tips), plane-margined; costa smooth at back, about 3/4-5/6 the

leaf length; upper cells shortly oblong-rhomboidal, 3-4:1, minutely papillose at back

. B. novae-angliae

Leaves loosely erect, lanceolate, acuminate, + flexuose but not twisted at the apex; margins

very narrowly recurved; costa toothed at back and ending in a spine; upper cells

oblong-linear, strongly papillose at back 2. B. graminicolor

1. Bryhnia novae-angliae (Sull. & Lesq. ex Sull.) Grout—Medium-sized plants in
soft, rather dense, bright- or yellow-green mats. Stems often somewhat dendroid. Stem
leaves about 1-1.3mm. long, broadly ovate or rounded-ovate, gradually or abruptly
narrowed to a short or moderately long acumen. Branch leaves usually rather remote,
spreading or erect-spreading and somewhat flexuose-contorted when dry, spreading
when moist, 0.6-1.2 mm. long, ovate to ovate-lanceolate, acute or broadly acuminate, +
twisted at the apex (especially at branch tips); margins plane; costa 3/4-5/6 the leaf
length, smooth at back; upper cells shortly oblong-rhomboidal, 3-4:1, minutely
papillose at back because of projecting ends. Setae 10-20 mm. long; capsules 1.5-2 mm.
long, dark red-brown; operculum apiculate to obliquely and stoutly short-rostrate; cilia
of endostome 1-3. Spores 13-18, minutely papillose or nearly smooth. Caly ptrae
apparently naked.—n = 11.

onard, How to Know the Mosses, (ed. 2) fig. 166d-e. Darlington, Mosses of Michigan, fig.
121. Grout, Mosses with Hand-lens an icroscope, fig. 152. Jennings, Mosses of Western
Pennsylvania, (ed. 2) Pl. 69. Welch, Mosses of Indiana, fig. 136.—Fig. 834-836.

On soil, humus, logs, or rocks in wet, shady places, particularly in seepage near brooks.
Europe and Asia; southeastern Alaska; Newfoundland to Ontario and the prairies, south to North
Carolina and Tennessee.

CHEBOYGAN CO.-—Reeses Bog, Douglas Lake, Burt Lake, Mill Creek. EMMET CO. —Pellston
Hills. MACKINAC CO.—Cut River.

Bryhnia novae-angliae grows in seepy habitats where Brachythecium rivulare
might also occur and is not unlike it in aspect. It can be distinguished, however,
because of its leaves with twisted points and papillae formed by cell ends projecting at
back. (B. graminicolor is much smaller and otherwise different, as indicated in the
key.

2. Bryhnia graminicolor (Brid.) Grout—Slender plants in dense, yellow-brown or
sometimes green mats. Stem leaves 1-1.1 mm. long, ovate-lanceolate, acuminate. Branch

294

leaves loosely erect (or occasionally erect-spreading) and somewhat flexuose when dry,
erect-spreading when moist, 0.4-0.6 mm. long, lanceolate, acuminate, not twisted at the
apex, margins very narrowly and indistinctly recurved nearly all around; costa ending
somewhat below the apex, toothed at back and ending in a small spine; upper cells
oblong-linear, 6-8:1 (or in occasional broader-leaved forms only about 4:1), strongly
papillose at back because of projecting ends. Setae 7-10 mm. long; capsules 1.5-2 mm.
long, brown; operculum narrowed to a rather short, oblique beak; cilia of endostome
1-2. Spores 13-15, minutely papillose or nearly smooth. Calyptrae with a few
scattered hairs.

Conard, How to Know the Mosses, (ed. 2), fig. 166a-c. Grout, Mosses with Hand-lens and
Microscope, fig. 153a. Jennings, Mosses of Western Pennsylvania (ed. 1) Pl. 54, (ed. 2) Pl. 59.
Welch, Mosses of Indiana, fig. 137.—Fig. 837-838.

n moist soil or rock (often calcareous), on banks of roads or streams, also on cliffs. New
England and Ontario to Minnesota, south to North Carolina and Arkansas.

CHEBOYGAN CO,—Fairy Island (Douglas Lake), Mill Creek. MACKINAC CO.—Caffey
Corner.

The plants are small, as compared with B. novae-angliae, with narrow, erect
leaves not twisted at the tip and leaf cells narrow and strongly papillose at the upper
ends

4. Cirriphyllum Grout

Plants slender to moderately robust, pale- or yellow-, rarely dark-green, mostly
shiny. Stems creeping to ascending, irregularly to subpinnately branched; branches
erect or ascending, terete and sometimes loosely julaceous. Leaves erect or erect-
spreading, often imbricate when dry, very concave and often cucullate, not or weakly
plicate, oblong or oblong-ovate or sometimes lanceolate, gradually or abruptly lance-
acuminate to subulate, + decurrent; margins erect, entire or serrulate; costa single,
about 1/2-2/3 the leaf length; upper cells smooth, linear; alar cells short-oblong to
subquadrate in somewhat differentiated groups. Dioicous: perichaetial leaves abruptly
or gradually narrowed to a long, slender acumen from a sheathing base. Setae elongate,
mostly rough; capsules inclined to horizontal, oblong-ovoid and asymmetric, rarely
suberect and subcylindric; annulus differentiated: operculum conic, acute to long-
rostrate; peristome teeth lanceolate, cross-striolate below, papillose above, bordered,
trabeculate at back; endostome free, about as long as the teeth, basal membrane high,
segments lance-acuminate, keeled and perforate, cilia well-developed, nodose. Calyptrae
naked.—The name refers to hair-pointed leaves. Like Rhynchostegium, also with a
long-beaked operculum, this genus is not as distinct from Brachythecium as one might
like.

Cirriphyllum piliferum (Hedw.) Grout—Medium-sized plants in soft, loose, shiny,
bright- or yellow-green mats. Leaves usually not particularly crowded, erect to
erect-spreading, 1.3-2.5 mm. long, very concave, not or only slightly and irregularly
plicate when dry, broadly oblong-ovate, + rounded at the apex and abruptly filiform,
broadly decurrent; margins erect, serrate at the apex, serrulate in the upper half or
more; costa extending about 2/3 the leaf length; cells of the decurrencies enlarged and
shortly oblong. Setae 10-30mm. long, rough throughout; capsules + curved and
oblong-cylindric, about 2.5mm. long; annulus of 2 rows of cells; operculum long-
rostrate (up to 1.8mm. long); stomata few, near the junction of urn and seta;
peristome teeth dark, red-brown; endostome pale-brown, with cilia in 2’s and 3’s.
Spores 11-15 y, finely papillose to nearly smooth.—n = 11, 12-14.

295

Darlington, Mosses of Michigan, fig. 124. Grout, Moss Flora of North America 3: Pl. 6 (figs.
1-23).—Fig. 839-841

s in wet, shady places (locally in Thuja swamps). Circumpolar; ae Victoria
Island ae Territories), Washington, Newfoundland to South Carolina and Michi

CHEBOYGAN CO.-—Gorge of Carp Creek, Reeses Bog. EMMET CO.—Sherett Lake.

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Bryhnia-novae-angliae. 834. Habit. 835. Leaves. 836. Upper cells of leaf.
Bryhnia graminicolor. 837. Leaves. 838. Upper cells of lea

Coripheyihim piliferum. 839. Habit. 840. Leaf. 841. Cells at leaf tip.
Bryoandersonia illecebra, 842. Habit. 843. Leaf.

296

The plants, suggestive of Brachythecium, are recognized by the soft, glossy
appearance of the tumid branches. The leaves, very concave and softly plicate when
dry, are abruptly narrowed to a slender, flexuose point (not really a hair-point as the
generic name suggests). Not yet found in Michigan but no doubt occurring in the
southern part is a typical denizen of the oak-hickory forest—Bryoandersonia illecebra
(Hedw.) Robins. (fig. 842-843), also known as Cirriphyllum boscii (Schwaegr.) Grout.
It is stoutly julaceous with broad, smoothly concave, cucullate leaves which end
abruptly in a short, twisted point. The leaves are clasping at base and have small,
thick-walled alar cells. The smooth setae and long-beaked opercula also aid in
recognition.

5. Eurhynchium BSG

Plants rather small to fairly robust, dark, sordid-green and dull or light-green,
yellowish, or brownish and often shiny when dry, in loose or dense mats or tufts.
Stems creeping, arched and decumbent, or ascending, irregularly to pinnately (or rarely
bipinnately) branched, sometimes + dendroid, frequently stoloniform; paraphyllia none
or few, at branch bases. Stem and branch leaves somewhat differentiated: Stem leaves
erect and sometimes appressed or more commonly spreading to squarrose, sometimes
loosely complanate, + concave, often plicate, broadly ovate or subcordate, shortly and
broadly acuminate or rather slenderly long-acuminate, somewhat to decidedly decur-
rent, costa ending at or above the leaf middle, often projecting at the tip as a dorsal
spine; cells smooth, linear or oblong-linear, much shorter at the apex, shorter and
broader at base, + differentiated at the basal angles (small and subquadrate or rarely
oblong and enlarged). Branch leaves narrower, broadly acute, obtuse, or rounded-
obtuse (or rarely gradually acuminate), sometimes twisted at the apex, the apical cells
generally shorter than the median, the terminal cell commonly short-rhombic. Peri-
chaetial leaves ending in a mostly squarrose-recurved subula. Setae elongate, smooth or
rough; capsules strongly inclined to horizontal, ovoid-cylindric to subcylindric, +
asymmetric, annulus differentiated; operculum slenderly and obliquely long-rostrate
from a convex-conic base; stomata in the short neck; peristome teeth cross-striolate
below, papillose above, bordered, trabeculate; endostome free, about as long as the
teeth, the basal membrane high, segments lance-acuminate, keeled and perforate, the
cilia mostly well developed, nodose or appendiculate. Calyptrae naked.—The name
alludes to the well-developed beak of the operculum which, together with the short
cells at the leaf tips, is the essential character of the genus.

1. Rather robust, autoicous plants of aquatic habitats, often submerged; branch leaves
1.2-1.5 mm. long or more; setae smoo . £. riparioides
1. Rather small, dioicous or phyllodioicous plants of moist but not aquatic habitats; branch

. Dioicous; branch leaves 0.8-1.2 mm. long, broadly ovate, acute; setae rough 2. FE. hians
2. Phyllodioicous; branch leaves 0.5-1 mm. long, narrowly oblong-ovate, bluntly acute to
rounded-obtuse; setae smooth 1. E. pulchellum

1. Eurhynchium pulchellum (Hedw.) Jenn.—Relatively slender, usually shiny,
green, yellowish, or brownish plants in extensive mats. Stems creeping, irregularly to
pinnately branched; branches horizontal or erect-ascending, + flattened or terete. Stem
leaves erect, about 1 mm. long, ovate, acute. Branch leaves crowded and erect or, more
typically, less crowded and erect to widely spreading; often slightly complanate,
0.5-1 mm. long, narrowly oblong-ovate, bluntly acute to rounded-obtuse, indistinctly
striate, often twisted at the apex; margins serrulate all around; costa extending 5/6 or
more the leaf length, + toothed at back near the tip and projecting as a dorsal spine;

297

cells rhomboidal, short at the apex. Phyllodioicous. Setae 10-20 mm. long, becoming
reddish, smooth; capsules 1.5-2 mm. long, oblong-cylindric, becoming somewhat con-
stricted below the mouth and at the short neck; peristome teeth brown; endostome
pale-brownish; cilia in groups of 2-3, well developed, nodulose. Spores 11-13 yu, smooth
or nearly so.—n = 10, 12 (as E. praecox).

Darlington, Mosses of Michigan, fig. 122. Grout, Mosses with Hand-lens and Microscope, Pl.

67 (as E. strigosum), Moss Flora of North America 3: Pl. 4 (figs. 1-17 & a-c), as E. diversifolium &

E. strigosum, Jennings, Mosses of Western Pennsylvania, (ed, 1) Pl. 53 & (ed. 2) Pl. 58 (as var.

robustum). Welch, Mosses of Indiana, fig. 139 (as var. robustum), 140 (as var. praecox).—Fig.
Ds

On soil or humus, particularly on low mounds in woods, or on rotten logs or stumps, bark
at base of trees, or rocks. Circumpolar; Mexico and Guatemala; northern oe America; Alaska to
Newfoundland, south to Arizona in the West, Georgia and Texas in the Eas

CHEBOYGAN CO.-—Reeses Bog, Hermits Bog, Gorge of Carp Creek, Iron Bridge, Fairy
Island (Douglas Lake), Colonial Point, aes Bog, Livingston ie Ellis Township, Mud Lake, Mill
Creek, Grass Bay, Little Lake 16. EMMET CO.—Levering, Arnott Lake, Pleasantview Swamp, Good
Hart, Middle Village, Wycamp Lake, ea Bay, Big Stone Bay, Conway Bog. MACKINAC CO.—Bois
cae Island, Mackinac Island, St. Martins Point, McKays Creek, ae Bay, Gros Cap, Epoufette,
Ozark, McNamara Pond. PRESQUE ISLE CO.—Ocqueoc River, Rainy River Falls, Bells Landing,
Mast Point, Lamprey Experiment Station, Evergreen Beach.

This is one of our most common mosses. It characteristically grows on old
stumps rotted nearly to the level of the forest floor or on low mounds of humic soil

ALI
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Eurhynchium pe eee 844. Habit. 845. Dwarf male plants growing on large female plant.
846. Leaves. 847. Cells at leaf t
Eurhynchium hians, 348. Leaves. 849. Cells at leaf tip.

298

(perhaps associated with windthrow of trees and kept in a disturbed condition by
extended exposure during the snow season). The plants are often pinnately branched
and frequently brownish. The small, ovate, bluntly pointed leaves, typically spreading
when moist are distinctive in comparison with other pleurocarps of woodland habitats.

The dwarf male plants are mere buds perched on the leaves of the female plants.
Lewis Anderson, investigating local collections, was able to show that a distinctive
chromosome recognizable at meiosis is transmitted to the male plants and is indeed a
sex chromosome. (For further information on sexual dimorphism, see the discussion of
Dicranum polysetum, above.)

2. Eurhynchium hians (Hedw.) Sande-Lac.—Dull or shiny, green or yellowish
plants with irregular to subpinnate branching. Stem leaves 1-1.2 mm. long, ovate or
triangular-ovate, slenderly short-acuminate. Branch leaves erect to erect-spreading,
0.8-1.2 mm. long, ovate (or sometimes ovate-lanceolate), acute, somewhat plicate when
dry, serrulate all around; costa ending in a dorsal spine about 3/4 up the leaf: cells
linear-rhomboidal, short and rhombic at the apex. Dioicous. Setae 10-27 mm. long,
reddish or orange-brown, rough; capsules oblong-cylindric, 1.5-2.5 mm. long; operculum
I-1.4 mm. long; peristome teeth brown; cilia of endostome nodulose, in pairs. Spores
9-13 m, very finely papillose to nearly smooth.—n = 7, 8, 10.

Conard, How to Know the Mosses, (ed. 2) fig. 233a-e. Grout, Mosses with Hand-lens and
Microscope, fig. 154. Jennings, Mosses of Western Pennsylvania, (ed. 1) Pl. 53 & (ed. 2) Pl. 58 (as
Oxyrrhynchium).—Fig. 848-849.

On soil and occasionally other substrata in damp, shady places. Europe; eastern Asia;
widespread in eastern North America; Arizona and New Mexico; Haiti.

CHEBOYGAN CO.—Gorge of Carp Creek, Iron Bridge (Carp Creek), Mill Creek, Stewart
Creek (south of Black Lake).

A plant of relatively wet habitats much resembling &. pulchellum but known by
its broader leaves and rough setae.

3. Eurhynchium riparioides (Hedw.) Rich.—Relatively robust plants in wide,
loose, somewhat rigid mats, dark-green or blackish below, often somewhat shiny and
green to yellowish or brownish above, irregularly branched; branches elongate, curved-
ascending. Stem leaves 1.5-2.5mm. long, erect to wide-spreading and often + homo-
mallous, only slightly concave, broadly ovate or oblong-ovate, ,acute, obtuse, or
rounded-obtuse; margins plane, serrulate nearly to the base; costa extending 4/5 or
more the leaf length, often projecting as a spine at back; cells green, linear-rhomboidal
above, short and broad at the apex. Branch leaves usually only about 1.5 mm. long or
less, often + pinched and twisted at the apex. Autoicous. Setae 12-23 mm. long,
orange, becoming reddish with age, smooth; capsules 1.5-2 mm. long, + curved,
oblong-ovoid, becoming constricted below the mouth and at the short neck; peristome
teeth dark-brown; endostome pale-brown, the cilia 2-3, well developed, nodose. Spores
13-16 uw, smooth.—n = 6, 8, 10, 11, 12, 20.

Conard, How to Know the Mosses, (ed. 2) fig. 232 (as E. rusciforme). Grout, Mosses with
Hand-lens and Microscope, fig. 155 (as E. rusciforme). Jennings, Mosses of Western Pennsylvania,
(ed. 1) Pl. S3 & (ed. 2) Pl. 58 (as Oxyrrhynchium). Welch, Mosses of Indiana, fig. 141.—Fig
850-852.

On wet rocks in or near brooks, often submerged in swiftly moving water or continually
splashed by waterfalls, probably preferring calcareous habitats. Europe, Macaronesia, North Africa,

299

Asia; British Columbia to California; Arizona, Mexico, and Guatemala; Jamaica and Haiti; northern
South America; Newfoundland to Minnesota and Oklahoma, south to Arkansas and North Carolina.

CHEBOYGAN CO.—Mill Creek.

Because of its aquatic habitat, this species has often been treated as a member of
the ‘“‘wet” Amblystegiaceae in the genus Platyhypnidium. Although it has an appear-
ance suggestive of Hygrohypnum, its true relationship as indicated by structure of
gametophyte and sporophyte is clearly with Eurhynchium pulchellum and £. hians.
The plants are larger, coarser, and darker than those of £. hians and grow in wetter,
often submerged habitats.

6. Rhynchostegium BSG

Plants rather small to moderately robust, in soft, pale, green or yellowish, rarely
dark-green mats, + shiny when dry. Stems creeping, irregularly to subpinnately
branched. Leaves erect or more often spreading, often somewhat complanate, some-
what concave, smooth, ovate or ovate-lanceolate, short- or long-acuminate; margins
somewhat reflexed below, mostly serrulate; costa rarely forked at the tip, 2-% the leaf
length; cells smooth, linear or oblong-linear, shorter and broader at the base, quadrate
to short-rectangular or not differentiated at the basal angles. Perichaetial leaves
sheathing at base, abruptly narrowed to a recurved subula. Setae elongate, smooth;
capsules strongly inclined to horizontal, ovoid- or oblong-cylindric, + asymmetric, often
constricted below the mouth when dry and empty; annulus differentiated; operculum
obliquely long-rostrate from a convex-conic base; stomata in the neck; peristome teeth
lance-subulate, cross-striolate below, papillose at the tips, bordered, trabeculate at back;
endostome about as long as the teeth, finely papillose, basal membrane high, segments
lance-acuminate, perforate along the keel, cilia mostly well developed. Calyptrae
naked.—The name refers to the beaked lid of the capsule, really the only reason, and
perhaps not a good one, for segregating the genus from Brachythecium.

Rhynchostegium serrulatum (Hedw.) Jaeg. & Sauerb.—Plants of moderate size, in
flat, shiny, bright- or yellow-green mats. Stem and branch leaves similar, usually not
crowded, scarcely altered on drying, wide-spreading and somewhat complanate (or
occasionally loosely erect), about 2mm. long (occasionally only 1.5 mm.), oblong-
ovate, slenderly acuminate, twisted at the apex; margins plane or erect, distinctly
serrulate in the upper 2/3 or nearly to the base; costa slender, ending about 3/4 up the
leaf, often ending in a minute and inconspicuous dorsal tooth and.often sparsely and
minutely toothed near the apex; cells linear-flexuose, not differentiated at the basal
angles. Autoicous. Setae flexuose, yellowish or orange-yellow, becoming orange-brown
or reddish with age, 10-30 mm. long; capsules oblong-cylindric but becoming narrow,
strongly contracted below the mouth, and strongly curved when dry and ee
1.5-2 mm. long, brown or red-brown; annulus of 2 rows of cells; operculum 1-1.5 m
long; et teeth dark- or red-brown, bordered, trabeculate; endostome pale ee
with nodulose cilia in groups of 2-3. Spores 9-11 wu, smooth.—n = 9, 10, 11

Breen, Mosses of Florida, Pl. 79. Conard, How to Know the oes (ed. 2) fig. 224 (as
Eurhynchium). Darlington, Mosses os Michigan, fig. 123. Grout, Mosses with Hand-lens and
Microscope, Pl. 68 (as Eurhynchium). Jennings, Mosses of Western Seat ce (ed. 1) Pl. 54, (ed.
2) Pl. 59. Welch, Mosses of Indiana, fig. 42 (as Eurhynchium).-—Fig. 853-857.

On soil, humus, rotten wood, and other substrata in moist or rather dry eae ga forests.
Widespread in eastern North America and also in uplands throughout tropical Ameri

300

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Rhynchostegium serrulatum, 853. Leaves. 854. Cells at leaf tip. 855, "Capsule. 856. Stomate.
857. Portion of peristome

Rhynchostegiella compacta. 858. Habit. 859. Leaves. 860. Cells at apex of leaf. 861. Cells at
margin of leaf base.

CHEBOYGAN CO.—Gorge of Carp Creek, Colonial Point. EMMET CO.—Pellston Hills, Five
Mile Creek

Best recognized by long-beaked opercula and smooth setae, R. serrulatum also
differs from various species of Brachythecium, most notably forms of B. rutabulum, by
a combination of characters: pale, wide-spreading, and loosely complanate leaves,
serrulate to the base or nearly so and ending in a slender acumen twisted at the tip.

7. Rhynchostegiella (BSG) Limpr.
mall plants in soft, green or yellowish, dull or shiny mats. Stems creeping,

S
irregularly to subpinnately branched; paraphyllia few, at branch bases. Leaves erect or
spreading, rarely somewhat secund, concave or nearly plane, narrowly lanceolate to

301

ovate-lanceolate, + long-acuminate; margins plane, entire or + serrulate; costa ending
near the leaf middle to subpercurrent, sometimes projecting at the tip as a dorsal spine;
cells smooth, linear or rhomboidal, scarcely differentiated at base, sometimes small and
subquadrate at the basal angles. Autoicous; perichaetial leaves + long-acuminate, not
squarrose. Setae elongate, slender and flexuose, red, smooth or papillose; capsules erect
to horizontal, oblong-ovoid, nearly symmetric, constricted below the mouth when dry
and empty; annulus differentiated; operculum apiculate to long-rostrate from a
convex-conic base; exothecial cells mostly + collenchymatous; peristome teeth lance-
subulate, papillose throughout or + cross-striolate below, papillose at the tips,
trabeculate at back; endostome free, about as long as the teeth, basal membrane high,
segments lance-acuminate, keeled and perforate, cilia mostly well developed. Calyptrae
naked.—A diminutive of Rhynchostegium.

da hagas compacta (C. M.) Loeske—Plants in dense, shiny, green or, more
typically, yellow mats. Stems usually crowded and ascending (but sometimes loose and
spreading), sa branched; branches short, ascending, usually + curved. Leaves
crowded, erect or erect-spreading, often homomallous when dry, 0.6-1.2 mm. long,
lanceolate, gradually acuminate, + decurrent; margins plane, serrulate all around,
especially at the base where the teeth are + spreading or recurved; costa percurrent or
nearly so, often indistinct at the midleaf, re-appearing above and frequently bearing
subcylindric brood-bodies or rhizoids at back; cells linear-rhomboidal, 5-9:1, sub-
quadrate at the basal angles. Setae 8-14mm. long, smooth, orange-yellow or red;
capsules 1.2-1.5mm. long, suberect or somewhat inclined, somewhat asymmetric,
oblong-cylindric from a short neck; operculum obliquely apiculate; stomata in the
neck; peristome teeth brown, papillose throughout or somewhat cross-striolate at base;
endostome pale, brownish-yellow, the cilia none or short and single. Spores 11-15 yu,
very finely papillose.

Conard, How to Know the Mosses, (ed. 2) fig. 222 (as sree Grout, Mosses with
Hand-lens and Microscope, fig. 172 (as Amblystegium).—Fig. 858

On wet, calcareous cliffs or bark at base of trees in wet woods. Europe; Greenland; Yukon
to California, Arizona, and Colorado; Quebec to South Dakota and south to Missouri and
Pennsylvania; Mexico

CHEBOYGAN CO.—Reeses Bog, Iron Bridge, Colonial Point, Grass Bay. MACKINAC
CO.—Bois Blanc Island.

Somewhat like an Amblystegium in size and appearance but softer and with
longer leaf cells (clearly like the Brachytheciaceae). The long costa disappears in the
acumen near the leaf apex, and the margins are serrulate in an interesting way, with
abrupt, recurved, and paired teeth near the leaf base. The common occurrence of
brood-bodies at the back of the leaf tips is an additional aid in identification.

8. Pleurozium Mitt.

Rather robust plants in loose, light-green or yellowish, + shiny mats. Stems
erect-ascending from a decumbent base, pinnately branched; branches loosely julaceous,
blunt or tapered; paraphyllia none. Leaves loosely imbricate, very concave, obscurely
wrinkled-plicate when dry, broadly oblong-ovate, rounded at the apex and generally
appearing recurved-apiculate because of abruptly incurved margins, somewhat decur-
rent; margins plane or reflexed near the base, broadly incurved above, crenulate to
serrulate at the apex; costa short and double; upper cells linear, smooth; cells at the
insertion thick-walled, porose, and orange, the alar cells shortly oblong, with thick,

302

orange walls, in rather small, triangular, often concave groups. Branch leaves shorter
and narrower. Dioicous; perichaetial leaves erect, elongate, oblong-lanceolate,
acuminate. Setae elongate, red or red-yellow, flexuose, smooth; capsules strongly
inclined to horizontal, curved, oblong-cylindric from a tapered neck, smooth; annulus
none; operculum conic or convex-conic, acute; stomata small, near the junction of seta
and urn; peristome teeth brown or brownish-yellow, finely papillose throughout (or
only indistinctly transversely papillose-striolate below), bordered; endostome pale-
brownish, finely papillose, with a high basal membrane, keeled and gaping segments,
and well-developed cilia in groups of 2-3. Calyptrae naked.-The name presumably
refers to the branches produced on either side of a suberect stem (in a loosely pinnate
arrangement). Because the costa is short and double and the alar cells noticeably
differentiated, Pleurozium is often placed in the Entodontaceae. However, the aspect—
so like Pseudoscleropodium— and the nature of the capsule suggest a closer tie to the
Brachy theciaceae.

Pleurozium schreberi (Brid.) Mitt.—Plants about 7-16 cm. high. Setae 20-43 mm.
long; capsules 2-2.5 mm. long; cilia of endostome nodulose or rarely + appendiculate.
Spores 11-20 u, very finely papillose.—n = 5, 10

nard, How to ners Bes snes (ed. 2) fig. 269 (as Calliergonella). Grout, Mosses with
Hand- ie and Microscope, fi 66 (as Callier, es Jennings, Mosses of Western Pennsylvania, (ed.
1) Pl. 44, (ed. 2) Pl. 48 (as pean "Fig, 862-

On humus, soil, and other substrata, usually in dry, open woods, but also in bogs or wet,
coniferous woods. Circumpolar; ta Rica; northern South America; across Canada, Alaska, and
the northern United States, south through the Great Lakes area and Virgi

CHEBOYGAN CO.—Reeses Bog, Hermits mee Gorge of Carp Creek, Iron Bridge, Bryants
en oe Point, Vincent Lake, Livingston Bog, Mud Lake, Grass Bay, Little Lake 16, Weber

E O.—Arnott Lake, Galloway Bog, Maple River Township, ne Swamp,
Stutsmanville Carp Lake, Wycamp Lake, Cecil Bay, Big yee Bay, Conway Bog. MACKINAC
CO.—Bois Blanc Island, Mackinac Island, St. Martins Point, McKays Creek, Bush a Epoufette,
— ide Dollar Lake, Caffey Corner, Hendricks, Prentiss Bay. PRESQUE ISLE CO.—Evergreen
Beach, Hammond Bay, Huron Beach, Mast Point, Ocqueoc River, Clinton Lake, Kelsey Pond, Loon

Linnaeus used the name Hypnum parietinum for this common northern moss, in
reference to its use in stopping chinks in the walls of houses in Scandinavia.
Braithwaite reported that it had been used in a similar way in shelters on ancient
island refuges in lakes or bogs in Great Britain. The plants can be recognized in the
field by red stems visible through semi-transparent leaves. (The leaves are light-green or
yellowish with something of a grayish overcast when dry). The species has been placed
in Hylocomium, and poorly branched forms of H. splendens can indeed be confused
with Pleurozium. The abundant presence of paraphyllia in Hylocomium is the best
means of establishing the identity of doubtful specimens.

e widespread occurrence of Pleurozium schreberi in the Northern Hemisphere,
and its disjunction in Central and South America are difficult to understand, as it
seldom fruits and has no obvious mode of vegetative reproduction. The mystery is
heightened, as noted by Watson (in Structure and Life History of Bryophytes), by the
fact that one rarely sees young growth stages as evidence of recent establishment or
active dispersal.

Newton (Trans. Brit. Bryol. Soc. 6(2): 230-243. 1971) found a measurable
difference in the size of the interphase heterochromatin in male and female plants of
Mnium undulatum, enabling him to sort out sterile plants as to sex potential. By
determining sex in this way, he showed that male and female plants are intermingled
more often than not (in a ratio of 1.3:1), but male plants are decidedly fewer (1:5.6 or

4
H

i

Pleurozium schreberi. 862. Habit. 863. Leaves. 864. Cells of alar region. 865. Capsule.

6.5). Fertilization is limited by the fact that not all male plants produce inflorescences
and distance is also a factor: The distance of a fertilized female plant from the nearest
male was normally only 2.5-12 cm. (in one instance 20 cm.

Hague and Welch (Bryol. 54: 214-215. 1951) attributed scarcity of fruits in
Bryoxiphium norvegicum to the fact that the sexes are nearly always found segregated
and inflorescences often lacking anyway. In a single fruiting specimen seen (from the
Wisconsin Dells), they found only archegonia.

Bedford’s study of sex distribution in colonies of Climacium dendroides (Northw.

304

Nat. 13: 213-221. 1938) is also worthy of reference, as is a recent study of
Rhytidiadelphus triquetrus by Reimann (Lindbergia 1: 219-224. 1973).

Pleurozium schreberi is one of the few mosses recorded from every vice-county
in the British Isles. There, the scarcity of sporophytes has been correlated with a rarity
of male plants by Longton & Greene (Ann. Bot. 33: 107-126. 1969). Fruiting
specimens were usually found where male and female plants grew intermixed. Fruiting
was induced in nature by transplanting from one area to another, in one instance,
between male and female colonies growing only 10 meters apart. In another study,
Longton & Greene (Ann. Bot. 33: 83-105. 1969) reported details of the life cycle in
Great Britain, where spores are shed nine to 12 months after fertilization.

Additional use of sex-ratio differences in Mnium are recorded by Newton (Ann.
Bot. 36: 163-178. 1922).

Uggla (Acta Phytogeogr. Suec. 41: 1-116. 1958) found that Pleurozium schrebert
and Hylocomium splendens often lack connections with the substrate and are therefore
usually completely consumed by forest fires. In northern Finland, they colonize burns
only after 30-40 years and then compete well with low-growing Cladonias and sods of
Ceratodon purpureus, Polytrichum juniperinum, and P. piliferum. The Polytricha
survive fires better because of penetration of mineral soil by rhizoids.

ENTODONTACEAE

Plants in loose, + shiny mats or tufts. Stems creeping or sometimes erect-
ascending (very rarely pendent), irregularly to pinnately branched; branches generally
julaceous, sometimes + flattened. Leaves crowded in many rows, usually appressed at
least when dry, sometimes + complanate, mostly symmetric, of various shapes; costa
single and + elongate (but never percurrent), very short and double, or lacking; cells
linear or oblong-linear, rarely narrowly elliptic, smooth, mostly quadrate and oblate-
rectangular at the basal angles, sometimes in many oblique rows, or rarely undiffer-
entiated. Inflorescences lateral; perichaetial leaves differentiated. Setae elongate;
capsules mostly erect and symmetric, mostly cylindric; annulus present or lacking;
operculum conic and often rostrate; peristome usually double (rarely the endostome
lacking), the teeth 16, papillose or variously striolate, rarely smooth or irregularly
thickened, the trabeculae lacking to well developed; endostome consisting of a mostly
low, keeled basal membrane and keeled, often narrowly perforate segments, cilia
lacking or rudimentary. Spores spherical. Calyptrae cucullate, mostly naked.

Entodon C. M.

Slender to moderately robust plants in green, yellow, or golden-brown, flat mats
or loose tufts. Stems creeping to ascending, irregularly to subpinnately branched;
branches short, blunt or acute, spreading, terete or + flattened. Leaves crowded,
concave, often + complanate, oblong-ovate or oblong-lanceolate, blunt or acute to
acuminate; margins erect or somewhat reflexed near the base, often serrulate at the
apex; costa very short and double or lacking; cells linear above, shorter and broader at
the base, hyaline and laxly quadrate in rather well-marked alar groups. Perichaetial
leaves elongate, acuminate from a + sheathing base, erect or spreading. Setae rather
short to elongate, smooth, reddish or yellow; capsules erect and symmetric, cylindric,
brown; annulus present or lacking; operculum usually obliquely rostrate; stomata few,
at extreme base of the urn; peristome teeth inserted somewhat below the mouth,
brown or red-brown, lanceolate, papillose or variously striolate or rarely smooth, not
or narrowly bordered; endostome usually about as long as the teeth, with a very low

305

basal membrane, linear, + keeled segments, the cilia none (or exceedingly rudimentary).
Calyptrae naked._The name refers to the teeth of the peristome which are within the
capsule, ie., inserted below the mouth. Generic characteristics include virtually
ecostate leaves with laxly quadrate alar cells and long, narrow, erect capsules with a
double, non-hypnaceous peristome. Color of seta (whether reddish or yellowish) and
surface markings on the peristome are of fundamental importance in classification of
the genus

Stem and branches distinctly flattened; leaves complanate; peristome teeth papillose
1. E. cladorrhizans

Stems and branches terete; leaves not complanate; peristome teeth smooth or nearly so
2. £. seductrix

1. Entodon cladorrhizans (Hedw.) C. M.—Bright- or yellow-green plants, some-
times tinged with golden-brown. Stems prostrate, subpinnately branched; both stems
and branches strongly flattened. Stem leaves erect or erect-spreading and complanate,
about 2 mm. long, broadly oblong-ovate, acute; margins serrulate (and often notched)
at the apex. Branch leaves smaller. Autoicous. Setae 7-20 mm. long, orange-brown;
capsules 2-3 mm. long, dark-brown; annulus of 2-3 rows of cells, persistent; operculum
high-conic or obliquely rostrate; peristome teeth finely papillose (or occasionally +
striolate), slightly bordered, often perforate above; segments smooth or slightly and
obscurely roughened. Spores 13-20 y, faintly roughened to finely papillose.

Breen, Mosses of Florida, Pl. onard, How to Know the Mosses, (ed. 2) fig. 290.
Jennings, Mosses of Western isles Nr Re 1) PI. 32, (ed. 2) Pl. 36.—Fig. 866-869.

On rotten wood, leaf ee og bases of trees in shady places. British Columbia and Alberta;
a ees in eastern North Amer

CHEBOYGAN CO.-—Colonial Point, Trout Creek, Cheboygan, Black River, Black Lake.
EMMET CO.—Harbor Springs.

Recognized by shiny, erect-spreading, complanate leaves.

2. Entodon seductrix (Hedw.) C. M.—Plants in green, yellowish, or brownish-
tinged mats. Stems creeping, subpinnately branched; stems and branches terete and
sometimes slightly flattened. Leaves concave, imbricate, 1-2 mm. long, oblong-ovate or
elliptic, abruptly acute to apiculate; margins entire or slightly serrulate at the apex.
Autoicous. Setae 5-16 mm. long, red or red-brown; capsules 2-3.5 mm. long, dark-
brown, usually somewhat wrinkled-striate when dry and empty; annulus of 2-3 rows
of cells, persistent; operculum stoutly and obliquely rostrate; peristome teeth smooth or
faintly and irregularly roughened, distinctly bordered; endostome segments smooth.
Spores 13-21 yw, finely papillose.—n = 11.

Mosses of Florida, Pl. 81. Conard, How to Know the Mosses, Sek 2) fig. 286a-g.
ae Grae of Michigan, fig. 128. Grout, Mosses with Hand-lens and Microscope, fig. 20S.
Jennings, Mosses of Western Pennsylvania, (ed. 1) Pl. 33, (ed. 2) Pl. 37. Welch, ee of ae
fig. 215.—Fig. 870-871.

On. rotten wood and bases of trees in shady, often rather dry places. Widespread in eastern
North America.

CHEBOYGAN CO.—Camp Knight (Douglas Lake), Riggsville, Black Lake. EMMET CO.—
Pellston Dunes, Larks Lake.

Shiny and attractive (though scarcely seductive), this species is distinguished from
E. cladorrhizans by its julaceous stems and branches.

Entodon cladorrhizans. 866. Habit of branch. 867. Leaves. 868. Upper cells of leaf. 869.
Cells in alar region.
Entodon seductrix. 870. Habit. 871. Leaves.

PLAGIOTHECIACEAE

Plants rather small to moderately robust, in + shiny, loose or dense mats. Stems
mostly creeping, irregularly or rarely pinnately branched; branches + flattened;
paraphyllia none. Stem and branch leaves similar, lanceolate to broadly ovate or
elliptic, obtuse or acute to acuminate, sometimes clearly decurrent, + concave, +
complanate, obliquely inserted, the dorsal and ventral leaves mostly symmetric and
erect or appressed, the lateral leaves often larger, + asymmetric and inflexed at the
base on alternating sides; costa single and + elongate, short and double, or lacking; cells
linear, oblong-linear, or sometimes oblong-rhombic, mostly smooth and chlorophyllose,
sometimes clearly differentiated at the basal angles and quadrate or shortly oblate-
oblong. Perichaetia lateral; perichaetial leaves rather small, erect, acuminate from a +
sheathing base. Setae + elongate, smooth; capsules erect or inclined to horizontal,
symmetric or nearly so; annulus mostly present; operculum convex-conic, sometimes
obliquely rostrate; peristome double, the 16 teeth lance-acuminate, mostly cross-
striolate below, bordered, trabeculate; endostome free, the basal membrane high,

307

segments broad and keeled, cilia present or lacking. Spores spherical. Calyptrae
cucullate, smooth, naked.

Plagiothecium BSG

Plants slender to medium-sized, in loose or dense, usually flat, green or yellowish,
shiny mats. Stems creeping or crowded and ascending, freely branched; cortical cells of
stems and branches large; pseudoparaphyllia none. Stem and branch leaves similar,
mostly strongly complanate, often asymmetric, oblong-ovate to ovate-lanceolate, acute
or acuminate, decurrent; margins often revolute, sometimes nearly to the apex, entire
or sparsely serrulate at the apex; costae double, unequal, frequently reaching 1/3-1/2
the leaf length, sometimes very short or lacking; cells smooth, linear-flexuose or
long-rhomboidal, shorter and broader at the base, laxly oblong in a few rows at the
basal margins in rather poorly differentiated, decurrent alar groups. Perichaetial leaves
moderately enlarged, sheathing, with somewhat spreading tips. Setae elongate, becom-
ing reddish with age; capsules suberect and symmetric to strongly eer or
horizontal and asymmetric, oblong-cylindric from a short neck, smooth or + longi-
tudinally wrinkled-plicate when dry and empty; annulus present; cee conic,
stoutly and obliquely apiculate to rostrate; stomata in the neck; peristome teeth
pale-yellowish and cross-striolate below, hyaline and papillose above, bordered,
trabeculate; endostome hyaline, papillose, with a high basal membrane, keeled segments
(not or very narrowly perforate), and nodulose cilia in groups of 2-3 (or occasionally
single, rarely lacking or rudimentary)._Named for the obliquely inclined capsules. The
outstanding feature of the genus is the alar cells which are somewhat differentiated in
narrow, decurrent groups. The leaves are generally complanate.

1. Leaves symmetric or nearly so, typically concave, not or only loosely and irregularly
complanate (and if so, shrunken, + contorted, and scarcely overlapping when dr
1. P. cavifolium
1. Leaves asymmetric, flat, distinctly complanate
2. Leaf cells about 5-7 » wide, linear-flexuose; capsules suberect, symmetric or nearly os
smooth, and not contracted below the mouth when dr P. laetum
2. Leaf cells broader, about 10-13 u wide, long-rhomboidal; capsules strongly inclined to
horizontal, curved and asymmetric, contracted below the mouth and usually + wrinkled-
plicate when dry 3. P. denticulatum

1. Plagiothecium cavifolium (Brid.) Iwats.—Plants of medium size, in + shin
yellowish to golden or sometimes bright-green mats. Stems and branches crowded,
ascending, and terete-foliate or prostrate and loosely and irregularly oomplanate-foliate
(often tapered and + complanate-foliate at the branch tips. Small, cylindric gemmae
sometimes produced in leaf axils or on leaves. Leaves crowded and erect or sometimes
fairly remote, irregularly complanate, and somewhat contorted when dry, usually
concave, 1.5-3mm. long, symmetric, ovate-lanceolate to oblong-ovate, abruptly nar-
rowed to a short, acute, often recurved apex or gradually narrowed to a short or rather
long acumen; margins often narrowly revolute near the base, sometimes nearly to the
apex, entire or occasionally sparsely serrulate at the tip; costa with 1 branch usually
1/3-1/2 the leaf length; cells long-rhomboidal, about 10-13 u wide. Dioicous. Setae
9-20 mm. long; capsules suberect and symmetric or somewhat inclined and + asym-
metric, 1.5-2.5 mm. long, smooth; annulus revoluble in fragments; operculum stoutly
rostrate; cilia of endostome 2-3. Spores 9-13 u, minutely roughened or nearly
smooth.—n = 9+1, 10, 11+1, 20.

Mosses of Florida, Pl. 89 (figs. 6-10), as P. sylvaticum. Grout, Moss Flora North
ees nN "PL 39C (as P. roeseanum). Jennings, Mosses of Western Pennsylvania, (ed. 1) Pl. 47 &

308

(ed. 2) Pl. 51 (as P. sylvaticum), Welch, Mosses of Indiana, fig. 209 (as P. roeseanum).—Fig.
872-874.

On shaded soil or humus (often overlying rocks or cliffs), sometimes on rotten wood or
bases of trees. Europe and Asia; Newfoundland to Minnesota and south to Georgia and Arkansas;
Alaska and the Aleutians.

CHEBOYGAN CO.—Burt Lake, Colonial Point, Polypody Bluff and Fairy Island (both at
Douglas ee Harare Bog, Mud Lake, nena Lake, Black Lake. EMMET CO. oo Arnott
Lake, Maple River Dam, Pleasantview Swamp, Middle Village, Cross Village, Wycamp Creek.
MACKINAC CO.-—Cut River, Ozark, Prentiss i PRESQUE ISLE CO.—Loon Lake, Rainy Lake.

Plagiothecium cavifolium is a common species of great variability. The branches
may be terete-foliate in a form commonly called P. roeseanum BSG or, in the form
generally referred in this country, through error, to P. sylvaticum (Brid.) BSG,
somewhat to strongly flattened, with leaves loosely complanate and somewhat shrunk-
en and scarcely overlapping when dry. The plants, characteristically yellowish, are not
as glossy as those of P. denticulatum or P. laetum

Plagiothecium latebricola BSG, a rare species of swamps and meadows, might
show up in our area in swampy habitats (on bases of rotten stumps or on old Carex or
fern tussocks, for example). The plants are very small and delicate, with erect-spread-
ing, not or slightly and loosely complanate, nearly symmetric leaves about 0.8-1.2 mm.
long, gradually narrowed to a slender, acute tip which is not recurved. The species has
been collected in Muskegon and Washtenaw Counties

Such names as Plagiothecium latebricola and Campylostelium saxicola ee
troublesome to most users of botanical Latin, as the specific epithets seem to be ou
of agreement with the neuter generic names. But the epithets are not adjectives, but
nouns in apposition.

2. Plagiothecium laetum BSG—Rather small plants in very glossy, ae or
yellow-green to golden mats. Stems and branches prostrate and very flat. Leaves
overlapping, spreading and strongly complanate, sometimes subsecund at the tips,
1.2-1.5 (rarely 2) mm. long, asymmetric, oblong, oblong-ovate, or oblong-lanceolate,
acute or acuminate; margins plane or narrowly revolute, sometimes nearly to the apex;
costa with 1 branch often 1/3-1/2 the leaf length; upper cells linear-flexuose, about
5-7 w wide. Autoicous. Setae 8-16 (rarely 20) mm. long; capsules suberect (or rarely
strongly inclined), symmetric or nearly so, 1.5-2 mm. long, pale- or yellow-brown,
smooth; annulus revoluble in fragments; operculum stoutly rostrate; cilia of endostome
1-3. Spores 9-12 x, smooth or nearly so.—n = 10, 10+1, 11.

Grout, Moss Flora of North America 3: Pl. 40 above.—Fig. 875-876.

Usually in coniferous woods, particularly on humus or soil of steep banks or overlying rocks
and cliffs, also on rotten wood and bases of trees. Europe; Alaska to California and New Mexico;
Newfoundland to Manitoba, south to Iowa and North Carolina

CHEBOYGAN CO.—Reeses Bog, Ingleside, Bryants Bog, Vincent Lake, Mud Lake, Little
Lake 16. EMMET CO.—Pellston Hills, Arnott Lake, Carp Lake, Wycamp Lake, Cross Village, Middle
Village, Five Mile Creek, Big Stone Bay. MACKINAC CO.—Bois Blanc Island, Mackinac Island, St.
Martins Point, Prentiss Bay, Cut River, Hendricks. PRESQUE ISLE CO.—Lamprey Experiment
Station, Mast Point.

Plagiothecium laetum is common in flat, very glossy mats on steep, wooded
banks and often on soil, humus, or bark at the bases of trees. As in P. denticulatum,
the leaves are crowded and very complanate; indeed, the species is in many ways

309

Plagiothecium cavifolium, 872. Leaves. 873. Cells at leaf tip. 874. Brood-bodies.

Plagiothecium laetum. 875. Leaves. 876. Cells at leaf tip.

Plagiothecium denticulatum. 877. Habit. 878. Leaf. 879. Cells of leaf tip. 880. Decurrent
alar cells.

merely a small edition of P. denticulatum, one of its synonyms being P. denticulatum
var. tenellum BSG. It is distinct, however, in having upper leaf cells narrower (5-7 u
wide) and linear-flexuose and capsules suberect, nearly symmetric, and not contracted
below the mouth when dry, and it grows in drier niches.

3. Plagiothecium denticulatum (Hedw.) BSG—Plants light- or, more often, dark-
green, shiny. Stems and branches flattened or somewhat ascending. Leaves usually
crowded and overlapping, wide-spreading and complanate, rarely somewhat contorted
when dry, 1.5-3mm. long, asymmetric, oblong-ovate, broadly acute to broadly
acuminate; margins sometimes plane but usually clearly revolute nearly to the apex,
sparsely serrulate at the tip; costa with 1 branch usually 1/3-1/2 the leaf length; upper
cells long-rhomboidal, about 10-13 wide. Small, cylindric, 3-7-celled brood-bodies
sometimes produced near the leaf tips. Autoicous (or, sometimes dioicous). Setae
15-30mm. long; capsules strongly inclined to horizontal, narrowly oblong-cylindric,
curved and asymmetric, 1.5-3mm. long, brown, smooth or + wrinkled-plicate, con-
tracted below the mouth when dry; annulus revoluble in fragments; operculum stoutly
rostrate; cilia of endostome 2-3. Spores 9-11 uw, smooth.—n = 10, 11, 20, 25.

310

‘onard, How to Know the Mosses, (ed. 2) fig. 294. Grout, Mosses with Hand-lens and
Microscope, Pl. 84. Jennings, Mosses of Western Pennsylvania, (ed. 1) Pl. 48, (ed. 2) Pl. 53. Welch,
Mosses of Indiana, fig. 207.—Fig. 877-880.

In wet woods, often in swamps, on rotten wood, bases of trees, soil, or humus Aa are
on soil or humus overlying rocks or cliffs). Europe, Macaronesia, Asia, Australia, and New Zealand;
Alaska to oo and New Mexico; Newfoundland to Manitoba and south to Illinois ead North
Carolina; Mexic

CHEBOYGAN CO.-—Iron Bridge, Ingleside, Smiths Bog, Colonial Point, Mud Lake. EMMET
CO.—Carp Lake, Stutsmanville, Wycamp Lake. MACKINAC CO.-—Bois Blanc Island, McNamara
Pond.

Plagiothecium denticulatum has shiny, crowded, complanate leaves with a few
small and irregular teeth at the very tips. The leaf margins are usually distinctly
recurved. As compared with flattened forms of P. cavifolium, the leaves are more
crowded, more complanate, more shiny, and more broadly pointed. In contrast to P.
laetum (which is much more common locally), it has broader, long-rhomboidal upper
leaf cells (about 10-13 rather than 5-7 u wide), and the capsules are curved-asymmetric
and contracted below the mouth when dry. It also grows in wetter habitats, on humus
at the swampy margins of pools, for example, whereas P. laetum characteristically
occurs on soil of steep banks or on bark at base of trees.

SEMATOPHYLLACEAE

Plants slender to moderately robust, in soft or rather rigid mats, green or
yellowish to yellow-brown or golden, often shiny. Stems creeping or sometimes
erect-ascending, irregularly to subpinnately branched; paraphyllia mostly lacking. Stem
and branch leaves mostly uniform, crowded in many rows but often + complanate,
often homomallous or distinctly secund, mostly symmetric, variously shaped; costa
very short and double or lacking; cells mostly long-rhomboidal or linear, smooth o
variously papillose, nearly always clearly differentiated at the basal angles (hich
abruptly and conspicuously inflated at the extreme angles in | or more transverse
rows). Perichaetia lateral; perichaetial leaves differentiated. Setae elongate, smooth or
sometimes + papillose; capsules inclined to pendulous and + asymmetric or rarely erect
and symmetric, ovoid- or oblong-cylindric, smooth; annulus usually lacking; operculum
convex or conic, usually rostrate; exothecial cells frequently collenchymatous;
peristome mostly double, the teeth 16, mostly lance-subulate, cross-striolate below (or
only rarely smooth or papillose), sometimes furrowed along the median line, usually
strongly trabeculate at back; endostome (rarely lacking) nearly always free, about as
long as the teeth or rarely much longer, basal membrane mostly high, segments
lance-subulate and keeled (or sometimes filiform, rarely short and broad), cilia usually
present. Spores spherical. Calyptrae cucullate or rarely mitrate, naked.

Leaves erect-spreading and somewhat complanate, not or sometimes very slightly falcate a
. Het

the tips; alar cells subquadrate in several rows ne ee ee
Leaves spreading and loosely complanate, falcate-secund at the tips; alar cells inflated an
oblong in 1 row 2. Brotherella

1. Heterophyllium (Schimp.) Kindb.

Plants medium-sized or moderately robust, in green or yellowish to brownish,
somewhat shiny, extensive, flat mats. Stems creeping, subpinnately branched; branches
horizontal, acute, straight or curved, obscurely flattened; paraphyllia few, at the bases
of branches. Stem and branch leaves sometimes + differentiated, erect or erect-spread-

311

ing, often + secund, concave, lanceolate to ovate, short- to long-acuminate; costa very
short and double or none; cells smooth, linear, yellowish at the insertion, subquadrate
and somewhat inflated with moderately thickened walls in several rows at the basal
angles forming small but well-marked, concave, hyaline or yellowish alar groups.
Perichaetial leaves clearly differentiated, elongate. pela elongate, smooth; capsules

apiculate or rostrate; exothecial cells not elena peristome teeth lance-
acuminate, cross-striolate and bordered below, papillose above, trabeculate at back,
with a zig-zag median line; endostome hyaline or yellowish, papillose, the basal
membrane high, segments broad and keeled, not or narrowly perforate, cilia + well
developed. Calyptrae cucullate.-The name alludes to the somewhat dimorphous leaves
of stems and branches and is not appropriate to our species which is more properly
placed in the Hypnaceae as Callicladium haldanianum (Grev.) Crum, Bryol. 74: 167.
1972

Heterophyllium haldanianum (Grev.) Kindb.—Plants bright-green, yellowish, or
brownish. Branches rather short, unequal, tapered, straight or somewhat curved,
especially at the tips, somewhat flattened. Stem and branch leaves similar, scarcely
altered on drying, erect or erect-spreading, often slightly secund (particularly near the
ends of branches), somewhat flattened but not distinctly complanate, about 1-1.7 mm.
long, ovate or ovate-lanceolate, gradually narrowed to a short or fairly long acumen;
margins plane, entire; cells long-linear, flexuose, somewhat shorter and broader at the
insertion; subquadrate alar cells well marked, often yellow-brown. Autoicous. Setae
16-32 mm. long, brown to red, slender, flexuose; capsules suberect to inclined,
narrowly cylindric, somewhat curved (especially below the mouth), 1.7-3 mm. long;
operculum stoutly and obliquely rostrate from a conic base; peristome teeth yellow-
brown; cilia of endostome single, short to elongate, fragile. Spores 12-16 yu, finely
papillose.—n = 11, 20+2, 22

oo ae How to Know the Mosses, (ed. 2) fig. 275. Darlington, Mosses of Michigan, fig. 132.
Grout, Mosses with Hand-lens and Microscope, fig. 191 (as Hypnum). Jennings, Mosses of Western
ac ne (ed. 1) Pl. 46, ee 2) Pl. 50 (as Sieroaden haldanianus). Welch, Mosses of Indiana,
fig. 199.—Fig. 881-884.

ually on logs or stumps, in both wet and dry ane ae in partial shade (at the edges
of woo Pid or in brushy thickets), also occasionally on bark at of trees. Europe; seu ger biea
in eastern North America, from Nova Scotia to Manitoba, a ue Missouri and Virg

CHEBOYGAN CO.—Reeses Bog, Hermits Bog, Gorge of Carp Creek, Iron Bridge, Sedge
Point, Colonial Point, Vincent Lake, Smiths Bog, Wolffs Bog, Bryants Bog; Livingston Bog, Mud
Lake, Duncan Bay, Grass Bay, Little Lake 16, Weber Lake. EMMET CO.—Maple River, Pleasantview
Swamp, Levering, Arnott Lake, Pellston Hills, Wycamp Lake, Big Stone Bay. MACKINAC
CO.-—Bois Blanc Island, St. Martins Point, McKays Creek, Prentiss Bay, Cut River, Gros Cap,
Epoufette, Ozark, Hendricks, Little Dollar Lake. PRESQUE ISLE CO.—Ocqueoc River, Lamprey
Experiment Station, Clinton Lake

This species has considerable character because of the short, tapered, and
somewhat flattened branches (resembling small swords) and also because of the curved
but nearly erect capsules.

The hygroscopic movements of peristome teeth result from the unequal deposi-
tion of material on adjacent walls of two peristomial layers. Each tooth reacts to
moisture as a bimetallic strip reacts to temperature. The endostome consists of a single
layer of material and is therefore not hygroscopic. It is a common oversimplification to
say that peristome teeth spread outward when dry and thus allow spores to be
dispersed when atmospheric conditions are suitably dry. As Patterson (Bryol. 56:
157-159. 1953) pointed out, in many (but not all) epiphytic mosses, the peristomes

Heterophyllium haldanianum. 881. Habit. 882. Leaves. 883. Alar cells. 884. Capsule.

open when wet. According to Lazarenko (Bryol. 60: 14-17. 1957), it is only in a small
group of epiphytic mosses with reduced double peristomes, such as Orthotrichum, that
the teeth spread outward under dry conditions. In a much larger group of epiphytes,
those with perfect double peristomes, the teeth bend inward both wet and dry, and the
peristome is closed except at a time of change from wet to dry or vice versa, i.e., at
the beginning and end of rain or heavy dew. Dispersal of spores under such conditions
insures adherence to a substrate and sufficient moisture for germination. These mosses
are thus protected from the hazards of long-range dispersal. (Of course, some spores
washed down to lower parts of the same tree may be redistributed by air currents
during dry periods.) As shown by Lazarenko (loc. cit.), the double peristome of
Heterophyllium actually prevents the escape of spores under both dry and wet
conditions. However, the teeth regularly break off in response to hygroscopic changes
(after only 10-20 movements), and the spores are then salted into the air currents as
though from a gymnostomous capsule.

Other references on spore dispersal in mosses: Goebel (Flora 80: 459486.
1895), Pfaehler (Bull. Soc. Vaudoise Sci. Nat. 40: 41-132. 1904), Sayre (Gard. Jour. 2:
51-53. 1952), Steinbrinck (Flora 84: 131-158. 1897), Ingold (Spore Discharge in Land
Plants, 1939; Trans. Bot. Soc. Edinb. 38: 76-88. 1959; Spore Liberation, 1965).

313

Other information on dispersal mechanisms is given in discussions on Splachnum,
Buxbaumia, and Tetraphis. The airgun method of explosively discharging spores from
dried capsules of Sphagnum also deserves mention. The effectiveness of such mech-
anisms are discussed by Crum (Jour. Hattori Bot. Lab. 35: 269-298. 1972.)

2. Brotherella Loeske ex FI.

Plants slender to medium-sized, in green, yellowish, or rarely brownish-green
mats, + glossy. Stems creeping, freely branched. Stem and branch leaves similar,
crowded, + complanate and secund to falcate-secund, concave, oblong-ovate or
oblong-lanceolate, slenderly acuminate; margins recurved below, serrulate above,
ecostate; cells smooth, long-rhomboidal or linear, yellow at the insertion, rather large,
oblong, inflated, and hyaline or yellow in a single transverse row at the basal angles
(with a few small, isodiametric cells above them). Perichaetial leaves plicate, subulate-
acuminate, toothed above. Setae elongate; capsules mostly + inclined, oblong-ovoid and
somewhat asymmetric to subcylindric; annulus persistent; exothecial cells not collen-
chymatous; operculum short- to long-rostrate; peristome teeth subulate-acuminate, with
a zig-zag median line, cross-striolate, trabeculate; endostome about as long as the teeth,
free, yellow, basal membrane rather high, segments keeled and often perforate, cilia
+rudimentary. Calyptrae cucullate Named for the great Finnish bryologist, V. F.
Brotherus, author of the Musci in Engler and Prantl’s Die natiirlichen Pflanzenfamilien.

Brotherella recurvans (Mx.) Fl.—Very shiny, green, yellow, or golden plants.
Stems subpinnately branched. Leaves 1-1.4mm. long, erect-spreading and loosely
complanate with falcate-secund tips or more clearly falcate-secund, oblong-ovate or
sometimes oblong-lanceolate, slenderly acuminate; margins sharply serrulate above; cells
linear above, shorter, broader, and yellow at the insertion with about 4-8 cells at the
basal angles conspicuously inflated and yellowish in a single transverse row, with 1-2 or
more rows of moderately inflated, usually hyaline cells above them. Dioicous. Setae
7-17 mm. long, orange-brown; capsules 1-1.5 mm. long, suberect to strongly inclined,
asymmetric, oblong-cylindric, smooth, brown; annulus of about 2 rows of small cells;
operculum obliquely rostrate, the beak often nearly as long as the urn; peristome teeth
yellowish to brownish; endostome segments not perforate, cilia 1-2, long and delicate.
Spores 13-18 yp, very faintly roughened.—n = 6

arlington, Mosses of Michigan, fig. 131. Grout, Moss Flora of North America 3: Pl. 35A
Jennings, Mosses of Western Pennsylvania, (ed. 1) Pl. 47 & (ed. 2) Pl. 51 (as Stereodon). Welch,
Mosses of Indiana, fig. 197.—Fig. 885-889.

On , humus, and bases of trees in moist woods. From Nova Scotia through the Great
Lakes and ane to North Carolina and Iowa; reported from Japan

CHEBOYGAN CO.—Reeses eee Hermits Bog, Gorge of Carp Creek, Burt Lake, Bryants Bog,
Ingleside, ees Bog, Grass Bay, Mud Lake. EMMET CO.—Galloway Bog, Arnott Lake, Carp
, Pleasantview Swamp, Stutsmanville, Good Hart, Five Mile Creek, Wycamp Lake, Conway

Bog. “MACK INAC CO.-—Bois Blanc Island, Mackinac Island, Cut River, Epoufette, pees
cN a Pond, St. Martins Point, McKays Creek, Prentiss Bay. PRESQUE ISLE CO.—Clinto
Lake, Kelsey Pon

Recognizable because of an extraordinary golden sheen. Only Plagiothecium
laetum, among local mosses, is so shiny. Brotherella is easily distinguished because of
soft, crowded, complanate leaves with secund tips. The few abruptly inflated cells at
the extreme angles are distinctive among our local Hypnum-like mosses, but they are
only a poor imitation of the beautiful “bubble cells” of Sematophyllum. The latter

886 | : dy UY oe

Brotherella recurvans. 885. Habit. 886. Leaves. 887. Cells at leaf tip. 888. Alar cells. 889.
Capsule.
Sematophyllum demissum. 890. Leaves. 891. Upper cells of leaf. 892. Alar cells.

genus, represented in a single locality in southern Michigan by S. demissum (Wils.)
Mitt. (fig. 890-892), is also distinguished by collenchymatous exothecial cells and
suberect, homomallous leaves.

Brotherella tenuirostris (Bruch & Schimp. ex Sull.) Fl. often resembles forms of
om

complanate, 0.5-1 mm. long, moderately serrulate above; the capsules are erect and
symmetric. Brown, papillose, filiform brood-bodies are often found in the leaf axils.
The species has been found in several southern counties (Eaton, Ingham, and Van
Buren).

HYPNACEAE

Plants small to robust, in loose or dense, often shiny mats. Stems mostly
creeping, sometimes ascending, irregularly branched to very regularly pinnate and
sometimes frondose; paraphyllia generally lacking. Leaves of stems and branches mostly
uniform, often homomallous to secund or falcate-secund, occasionally somewhat

315

complanate, rarely plicate, of various shapes, mostly +

long-acuminate; costa short and
double or none;

cells prosenchymatous (mostly linear), smooth or occasionally
papillose at back because of projecting upper angles, very rarely with several papillae in
a row over the lumen at back, mostly differentiated a the basal angles, small and

quadrate or sometimes moderately enlarged and inflated in rather small but usually
distinct alar groups. Perichaetia lateral; perichaetial ees diticientinted! Setae elongate,
mostly smooth; capsules erect or more commonly inclined to horizontal, ovoid to
cylindric, straight or curved and asymmetric; annulus usually differentiated, operculum
conic or sometimes convex, often rostrate; peristome nearly always double, the 16
teeth lance-subulate, usually cross-striolate below, usually bordered and strongly
trabeculate; endostome (rarely lacking) usually consisting of a high basal membrane,
broad, keeled segments, and well-developed, nodose cilia (but the teeth may be
papillose, basal membrane low, segments filiform, and cilia rudimentary or lacking).
Spores spherical. Calyptrae cucullate, smooth, naked or rarely + hairy.

1. Stem and branch ee aa ee stem leaves broadly decurrent; leaf cells papillose at

back because of projec 9. Ctenidium
1. Stem and branch ea not particularly differentiated, not or somewhat decurrent; cells

not papillose
2. Plants rather large, handsome, pinnately branched to form triangular fronds; leaves

strongly plicate . Ptilium
2. Plants small to medium-sized, not particularly striking in appearance, variously branched
eck ee | plicate

mm. 4. Platydictya
3. Plants small to fairly robust; leaves more are 0.5 mm. long
4. Capsules erect and symmetric
5. Plants dark-green with a golden, oily sheen; branches + straight; leaves erect when
dry; clusters of brood branchle el in leaf axils; margins reflexed; ce basal
membrane of endostome low; cilia

one 1. Platygyrium
5. Plants light-green or yellowish, a eelt shiny; branches curved when dry; leaves

appressed and somewhat homomallous when dry; margins erect; brood branchlets
lacking; autoicous; basal membrane well cate a cilia rudimentary 2. Pylaisiella

4. Capsules variously inclined, curved, and asymmetr

6. Leaves 0.5-0.8 mm. long, erect or appressed, not e all complanate or secund

3. Homomallium
6. Leaves longer, mostly + complanate or secund
7. Leaves secund or falcate-secund; alar cells differentiated, small and quadrate or
larger and + infla 7. Hypnum
tate not or somewhat secund at the tips; alar cells scarcely
ed
8. Leaves strongly ae apical cells shorter than the median
8. pe not or loosely complanate; apical cells not differentiated
9, Leaves 0.7-1.2 m ee entire; capsules oblong-cylindric, smooth
9. ene 1.2-1.5 mm. long, serrulate; capsules curved-cylindric, plicate

6. Taxiphyllum

5. Isopterygium
8. Herzogiella

1. Platygyrium BSG, nom. cons.

Rather small plants in flat, dark-, yellowish-, golden-, or brownish-green, glossy
mats. Stems creeping, freely branched; branches ascending, often somewhat curved,
short, usually bearing clusters of minute brood-branches in axils of upper leaves. Stem
and branch leaves similar, erect or appressed when dry, erect-spreading when moist,
concave, ovate- or oblong-lanceolate, acuminate, slightly decurrent; margins entire or

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triangular alar groups. Dioicous; perichaetial leaves somewhat elongate, with +
ing tips. Setae reddish, smooth; capsules erect and symmetric, cylindric; annulus of 2-4
rows of cells, persistent; operculum obliquely conic-rostrate; stomata few, at extreme
base of the urn; peristome teeth inserted near the mouth, lanceolate, yellow-brown,

316

transversely striolate-papillose at base, papillose above; endostome yellowish, smooth,
about 2/3 as long as the teeth, consisting of a very low basal membrane, nearly linear,
non-perforate segments, and no cilia. Calyptrae naked._Named in reference to the
broad annulus.

Platygyrium repens (Brid.) BSG—Leaves 0.8-1.3 mm. long; upper median cells
about 6-10:1. Setae 10-25 mm. long; capsules 1.5-2.2 mm. long. Spores 11-15 y, finely
papillose.

en, Mosses of Florida, Pl. 88 (figs. 7-10). Conard, How to Know the Mosses, (ed. 2) fig.
261. Erne Mosses of Michigan, fig. 137. Grout, Mosses with Hand-lens and Microscope, fig.
208. Jennings, Mosses of Western Pennsylvania, (ed. 1) Pl. 33, (ed. 2) Pl. 37. Welch, Mosses of
Indiana, fig. 206.—Fig. 893-897.

logs and stumps, bark of trees, and occasionally on rocks, often in rather dry, brushy,
yee places, such as roadsides, pastures, and bag of woodlots. Circumpolar; British Columbia
o New Brunswick, throughout eastern North Am

CHEBOYGAN CO.—Reeses Bog, Hermits Bog, Gorge of Carp Creek, Iron Bridge, Pine Point,
Livingston Bog, Grass Bay, Weber Lake. EMMET CO.—Pellston Dunes, Maple see Pleasantview
Swamp, Arnott Lake, Cross Village, Good Hart, Middle Village, Wilderness State Park, Alanson,
Conway Bog. MACKINAC CO.-—Bois Blanc Island, Mackinac Island, St. Martins Point, McKays
Creek, Gros Cap, Cut River, Epoufette, Ozark, Hendricks. PRESQUE ISLE CO. _Evergreen Beach,
Lamprey Oe Station, Mast Point, Rainy River Falls.

Platygyrium repens is common on trunks of trees, especially in open places in
woods or in brushy ecotones, as blackish-green mats with a curious oily sheen. The
axillary clusters of brood-branchlets (actually little buds) are invariably present and
provide the easiest means of recognition. Leskeella nervosa has similar branchlets, but
the plants are a dull dark-green, the leaves are narrower and long-pointed, and the
costa is single and nearly fills the leaf apex. Homomallium adnatum, which grows on
rock and also on bark at base of trees, is distinguished from Platygyrium by erect leaf
margins.

2. Pylaisiella Kindb.

come rather slender or medium-sized, in flat, yellowish, light- to dark-green, or
brownish, + shiny mats. Stems creeping, irregularly or subpinnately branched; branches
short, erect-ascending, usually curved. Stem and branch leaves similar, crowded, usually
homomallous, loosely appressed when dry, erect-spreading when moist, concave, ovate-
or oblong-lanceolate, short- to long-acuminate; margins mostly erect and entire; cells
linear or linear-rhomboidal, small and quadrate in several rows at the basal angles.
Autoicous; perichaetial leaves oblong- or ovate-lanceolate, + long-acuminate, serrulate at
the apex. Setae smooth; capsules erect and symmetric (or rarely somewhat curved),
ovoid- or oblong-cylindric; annulus present or lacking; operculum conic, sometimes +
rostrate; peristome deeply inserted, the teeth cross-striolate (or sometimes smooth or
indistinctly striolate), bordered, trabeculate; endostome free or + adherent to the teeth,
the basal membrane well developed, segments lance-subulate, keeled and variously
perforate or split, cilia mostly rudimentary. Calyptrae naked.—A diminutive of Pylaisia,
the name given the genus in honor of an early French bryologist, Bachelot de la Pylaie.

better-developed annulus, endostome with a very low basal membrane (not exserted
above the mouth), narrower segments, and cilia completely lacking.

CE ANZ : “J
aaa

Platygyrium repens. 893. Branch and brood-branchlets. 894. Leaves. 895. Cells at margin of
leaf near apex. 896. Cells at basal angle of leaf. 897. oe ule.

Pylaisiella polyantha. 898. Leaves. 899. Alar c
Pylaisiella selwynii. 900. Habit. 901. Leaves. 902. Alar cells.

Quadrate ae ae relatively few (only 3-9 along the margins); endostome free . P. polyantha
Sees many (about 15-20 along the margins); endostome partially Pete to
cen _ 2. P. selwynii

1. Pylaisiella polyantha (Hedw.) Grout—Plants in glossy, green or yellowish mats.
Leaves 1-1.3 mm. long, ovate or ovate-lanceolate, gradually or rather abruptly long-
acuminate, entire or slightly serrulate above; quadrate alar cells relatively few (3-9 at
the margins). Setae 6-10 mm. long; capsules 1.3-2 mm. long, oblong-cylindric; annulus
narrow; operculum conic or conic-apiculate; segments of endostome free. Spores 9-15 u
finely papillose.—n = 11

Conard, How to Know the Mosses, (ed. 2) fig. 288a-e (as Pylaisia). Darlington, Mosses of
Michigan, fig. 136 fm pare Grout, Moss Flora of North America 3: Pl. 35G (as Pylaisia).—Fig.
898-899.

On bark of trees. Circumpolar; wiht Canada and Alaska, presumably south to Arizona in
the West and the Gulf of Mexico in the E

318

eer oa CO.—Reeses Bog, Gorge of Carp ae Iron Bridge, Mullet Lake. EMMET
CO.—Wycamp Creek, Mackinaw City Hardwoods, Cecil Bay, Big Stone Bay. MACKINAC CO.—Bois
Blanc ae. Mackinac Island. PRESQUE ISLE CO, ar eet Isle Lighthouse, Hammond Bay.

The plants are looser and the branches less tightly curled than in P. selwynii. The
segments of the endostome are completely free from the exostome, and so both parts
of the peristome can be seen with a hand-lens; in our other species, P. selwynii, the
endostome is at least partly adherent to the exostome and cannot be detected in a dry
condition under low magnifications.

2. Pylaisiella selwynii (Kindb.) Crum, Steere & Anders.—Plants in rather glossy,
green or yellow-green mats. Branch leaves crowded, 0.5-1.3 mm. long, ovate-lanceolate
and acuminate, entire or nearly so; alar cells numerous (15-20 at the margins and
extending about 1/3 up the leaf). Setae 6-14mm. long; capsules 1-2 mm. long,
oblong-cylindric; annulus of 2-3 rows of cells; operculum conic, obliquely blunt-tipped;
segments of endostome adhering to the teeth in the lower 2/3. Spores 13-24 u, densely
papillose.—n = 10+1, 11.

Conard, How to Know the ae (ed. 2) fig. 287 (as Pylaisia). Grout, Mosses with
Hand-lens ar Microscope, Pl. 85 (as Pylaisia schimperi). Jennings, Mosses of Western Pennsylvania,
(ed. 1) Pl. 33 (as ae aie (ed. 2) Pl. 37 (as Pylaisia). Welch, Mosses of Indiana, fig. 204
(as Pylaisia).—Fig. 900-902

bark of trees. Apparently rare in northern Europe, Siberia, and Japan; southern Canada
to the Gulf of Mexico; reported from Arizona; Sonora.

CHEBOYGAN CO. es Bog, Gorge of Carp Creek, Douglas Lake, Smiths Bog, Colonial
Point, Mill Creek. EMMET CO.-—Larks Lake, Cross Village, Wilderness State Park. MACKINAC

Mackinac Island, Gros Cap, Ozark, Caffey Corner, Hendricks. PRESQUE ISLE CO.—Evergreen
Beach, Clinton Lake.

The neatly curved branches are smaller and more compact than those of P.
polyantha.

Pylaisiella intricata (Hedw.) Grout, widespread in the Northeast but not known
to occur in Michigan, has segments wholly adherent to the teeth, alar cells less than 10
at the margins, and spores 20-30 yu.

3. Homomallium (Schimp.) Loeske

Plants small to medium-sized, in flat, dark- or brownish-green to yellowish, dull
or + shiny mats. Stems creeping, freely and irregularly to subpinnately branched;
branches short, straight or somewhat curved; paraphyllia few. Stem and branch leaves
similar, erect to erect-spreading or homomallous, concave, oblong-lanceolate or oblong-
ovate, acuminate; margins erect, entire or + serrulate near the apex; upper cells
thomboidal, smooth or papillose at back because of projecting cell ends; alar cells
quadrate in numerous rows. Autoicous; perichaetial leaves elongate, loosely sheathing.
Setae smooth; capsules inclined to horizontal, curved and constricted below the mouth
when dry and empty; annulus differentiated; operculum convex-conic, blunt to
apiculate or rostellate; peristome teeth cross-striolate below, papillose above, bordered
and trabeculate; endostome papillose, with a high basal membrane, keeled and
sometimes perforate segments, and well-developed, nodulose cilia in groups of 1-3.
Calyptrae naked.—Named in reference to a tuft (or woolly mat) in which the leaves are
curved or directed in the same way (characteristic of some of the species).

319

Homomallium adnatum (Hedw.) Broth.—Small plants in dull, dark-green or
blackish to brownish mats. Stems freely but irregularly branched; branches short and
straight. Leaves erect (or very slightly secund) when dry, erect-spreading when moist,
0.5-0.8 mm. long, oblong-ovate, rather abruptly short-acuminate; margins entire or
nearly so; upper cells smooth, thick-walled, shortly oblong-rhomboidal, about 5-6:1;
alar cells quadrate in triangular groups extending about 1/3 up the leaf. Setae
6-12 mm. long, orange-yellow; capsules 1-1.5 mm. long, yellowish; operculum obliquely
and stoutly apiculate or rostellate; stomata at extreme base of the urn; segments not
perforate; cilia in groups of 1-2. Spores 11-12 u, smooth or very finely papillose.

Conard, How to Know nee Mosses, (ed. 2) fig. 259. Darlington, Mosses of Michigan, fig. 134.
Grout, Mosses with Hand-lens and eee Hie figs. 202(5) & 203 (as Amblystegiella). Jennings,
Mosses of Western Beesyianis (ed. 1) Pl. 39, (ed. 2) Pl. 43.—Fig. 903-905.

On rocks (especially limestone), rarely on bases of hardwood trees. Eastern North America;
Japan.

CHEBOYGAN CO.-—Pine Point (Douglas Lake).

Rather similar to Platygyrium repens in appearance but differs most noticeably in
having leaves with erect, rather than reflexed margins. The habitats, of course, are
normally quite different.

4. Platydictya Berk.

Plants small and slender, in dark-, light-green or brownish, dull or slightly shiny
mats. Stems mostly creeping, irregularly branched; paraphyllia none. Leaves very small,
slightly concave, erect-spreading or weakly secund when moist, lanceolate or lance-
subulate, not or shortly and indistinctly decurrent; margins plane and entire or
serrulate; cells rhomboidal or oblong-hexagonal, mostly short (2-4, rarely 8 times as
long as broad), shorter and somewhat laxer at base, subquadrate and poorly dif-
ferentiated at the basal angles. Perichaetial leaves erect, sheathing at base, oblong-
lanceolate, long-acuminate. Setae slender, reddish, smooth; capsules mostly erect and
symmetric or weakly curved (sometimes inclined and more distinctly curved), oblong-
cylindric, mostly constricted below the mouth when dry and empty, smooth; annulus
differentiated; operculum convex, blunt or. obliquely saa peristome teeth cross-
striolate below, bordered, + trabeculate; basal membrane of endostome low to
moderately high, segments well developed, cilia single and Spee to lacking or
rarely in 2’s and 3’s and well developed._The name apparently refers to a broad-
meshed network formed by the outlines of the short leaf cells and is not obviously
appropriate. (The genus has been called Amblystegiella, because of some resemblance
to Amblystegium, but that name is nomenclaturally unavailable.)

1. Silky, green or yellowish, soft plants with serrulate leaves; alar cells ae differentiated ;
perichaetial leaves spinulose-denticulate . jungermannioides

1. Dull, dark-green, + rigid plants with subentire leaves; quadrate alar cells more numerous;
perichaetial leaves entire or slightly denticulate

2.On_ bark _ base of trees; leaves 0.25-0.5 mm. long; cells 3-5:1; capsules erect a

symmetric or nearly so; operculum conic, obliquely pointed . P. subtile
2. On Geren rocks; leaves 0.15-0.3 mm. long;,cells 2-3:1. capsules curved and strongly
inclined; operculum convex-conic, not ‘obligate pointed 3. P. confervoides

1. Platydictya jungermannioides (Brid.) Crum—Soft, silky, green or yellowish
plants with creeping stems and ascending branches. Short-cylindric or clavate brood-
bodies sometimes produced in leaf axils. Leaves erect-spreading, 0.17-0.3, rarely

Homomaltium adnatum. 903. Habit. 904. Leaves. 905. Cells of upper part of leaf.
tydictya jungermannioides, 906. Habit. 907. ee 908. Cells at apex of two leaves. 909.
Cells leaf base.
Platydictya subtile. 910. Leaves. 911. Cells at tip of two leaves. 912. Alar cells.
Platydictya confervoides. 913. Leaves. 914. Areolation of entire leaf.

0.5 mm. long, lanceolate, acuminate, not or somewhat narrowed to the insertion,
margins serrulate throughout (but especially at the base); cells elongate throughout,
oblong- pega and about 3-4:1 above, a few at the basal angles subquadrate

usuall more than 3, sometimes as many as 7 at the margins). Dioicous;
ead leaves denticulate. Setae i 11mm. long; capsules erect and symmetric or
nearly so, contracted below the mouth when dry, 0.6-1 mm. long; operculum
convex-conic and mammillate to oes cilia of endostome none or 1-2. Spores
11-13 py, finely papillose.

Grout, Mosses with Hand-lens and Microscope, fig. 202(2), as Amblystegiella sprucei. —Fig.
906-909.

In damp, shaded places, especially in crevices of cliffs, also found in sheltered habitats on
logs, bases of trees, and soil of banks or roots of upturned trees. Circumpolar; Canada and Alaska,
south to New Mexico, the Great Lakes, and New England.

321

CHEBOYGAN CO.—Reeses Bog, Colonial Point, Grass Bay. EMMET CO.—Arnott Lake, Cecil
Bay, Wilderness State Park. MACKINAC CO. ~Mackinac Island, McKays Creek.

Extremely small, delicate, pale-green plants resembling a minute Amblystegium
but lacking a costa; this species is most likely to be confused with Jsopterygium
pulchellum, which is larger, very glossy, and has entire leaves which are in two
apparent rows but more or less curved and homomallous.

2. Platydictya subtile (Hedw.) Crum—Rather dull, green to brownish plants with
creeping stems and erect branches. Leaves erect-spreading or subsecund, 0.25-0.5 mm.
long, lanceolate, acuminate, + narrowed to the insertion, entire or nearly so; upper
cells oblong-rhomboidal, about 3-5:1; subquadrate alar cells rather numerous (about
7-16 along the margins). Autoicous; perichaetial leaves entire or nearly so. Setae
7-12 mm. long; capsules erect and symmetric or slightly inclined, oblong-cylindric, not
or slightly contracted below the mouth when dry, about 1-1.5 mm. long; operculum
conic, obliquely apiculate to rostellate; cilia of endostome none or rudimentary. Spores
9-13 uw, finely papillose.—n = 10

Conard, o Know the Mosses, (ed. 2) fig. 260a-d (as AA leds Grout, Mosses
with Hand-lens oe ere figs. 201, 202(3), as Amblystegiella, —Fig. 910-912

n bark at base of hardwood trees. Europe; Asia; southeastern Canada to Minnesota, south
to Pennsylvania and Illinois.

CHEBOYGAN CO.—Gorge of Carp Creek, Fairy Island, Colonial Point, Mud Lake, Mill
Creek. EMMET CO.—Larks Lake, Pellston Hills, Big Stone Bay. MACKINAC CO.—Mackinac Island,
Prentiss Bay, Gros Cap, Ozark, Hendri cks.

Though small, the largest of the genus. The habit of growth at the base of
hardwood trees is the easiest character to use in recognition.

3. Platydictya confervoides (Brid.) Crum—Dark-green, rather rigid plants with
creeping stems and erect-ascending branches. Leaves erect or rarely subsecund when
dry, erect-spreading when moist, 0.2-0.3 mm. long, lanceolate or ovate-lanceolate,
acute, not or somewhat narrowed to the insertion, entire or nearly so; upper cells
short, 2 (rarely 4):1, irregularly oblong-rhombic; basal cells short, those at the basal
angles inconspicuously small and subquadrate (3-6, sometimes 12 along the margins).
Autoicous; perichaetial leaves subentire. Setae 6-8 mm. long; capsules 0.8-1.2 mm. long,
suberect to nearly horizontal; operculum convex-conic, apiculate; cilia of endostome
1-3, rather well developed. Spores 11-13 y, finely papillose.—n =

Conard, How to Know the Mosses, (ed. 2) fig. 260e-f (as Amblystegiella). Darlington, Mosses
of Michigan, fig. 135 (as Amblystegiella). Grout, Mosses with Hand-lens and Microscope, fig.
202(1), as Amblystegiella. Welch, Mosses of Indiana, fig. 202 (as Ambiystegiella).—Fig. 913-914.

limestone rocks in moist, shady places, especially in stream valleys. Europe and the
anaes southeastern Canada and northeastern United States (and presumably in the Rocky
Mountains).

CHEBOYGAN CO.—Mill Creek. MACKINA€ CO.—Mackinac Island, Ozark.
Very small, dark-green plants characteristically found in the valleys of streams on

moist, shaded, calcareous rocks. The subquadrate alar cells are helpful in determina-
tion.

B22
5. Isopterygium Mitt.

mall or medium-sized plants in soft, light- or yellowish-green, sometimes whitish
mats, often somewhat glossy. Stems creeping or sometimes ascending, mostly irregu-
larly branched. Stem and branch leaves similar, oblong- or ovate-lanceolate, acuminate,
not or only slightly decurrent, the dorsal and ventral leaves obliquely appressed and
symmetric, the lateral leaves usually spreading, + complanate, sometimes + asymmetric
and frequently somewhat secund; margins plane, entire or serrulate; cells smooth (or
rarely papillose at back because of projecting angles), narrow and elongate (mostly
linear) above, shorter and thicker-walled or sometimes lax at the insertion, not or
slightly differentiated at the basal angles. Perichaetial leaves erect, sheathing at base,
gradually or abruptly subulate. Setae smooth; capsules small, suberect to strongly
inclined, + asymmetric, oblong-ovoid to subcylindric from a short, distinct neck;
annulus sometimes differentiated; operculum conic or convex-conic, sometimes
rostrate; peristome teeth cross-striolate, bordered, trabeculate; basal membrane of
endostome high, segments keeled and sometimes perforate, cilia 1-3, delicate, some-
times nodose or + appendiculate. Calyptrae naked._The name means equal wings or
sides and probably refers to the fact that the lateral leaves are often somewhat
spreading and complanate, thus forming indistinct rows.

Isopterygium pulchellum (Hedw.) Jaeg. & Sauerb.—Plants small and slender, in
shiny, bright-green or yellow mats. Stems creeping, irregularly branched; branches often
ascending. Leaves usually not crowded, erect- to wide-spreading and somewhat to
distinctly complanate, often + secund at the tips, 0.7-1.2mm. long, narrowly
lanceolate, slenderly acuminate, entire; cells linear-flexuose, scarcely differentiated at
the basal angles. Autoicous. Setae 8-l16mm. long, orange or al becoming
orange-red; capsules suberect and nearly symmetric to inclined and + asymmetric,
oblong-cylindric, smooth, 1-1.6 mm. long; annulus of 2 rows of cells; operculum
bluntly low-conic; stomata at extreme base of the urn; endostome segments non-
perforate, the cilia single (or occasionally paired), nodulose. Spores 9-13 yu, minutely
roughened.—n = 10, 11,

Darlington, Mosses of Michigan, fig. 130.—Fig. 915-917.

n damp soil, eae rock, or bark at base of wees in deep shade, often under overhanging
turf, in crevices, or undersides of logs. Circumpolar; New Zealand; Greenland to Alaska, south
to California, oa Colorado, Wisconsin, and New York.

CHEBOYGAN CO.-—Hermits Bog. EMMET CO.—Arnott Lake, Pleasantview Swamp.
PRESQUE ISLE CO.—Rainy Lake.

Indeed pretty and small, as the name indicates, the plants are usually interlaced
with other mosses in moist, protected niches. The branches are erect or ascending, and
the leaves, shiny when dry, are flattened into two apparent rows and also somewhat
curved and homomallous.

Isopterygium distichaceum (Mitt.) Jaeg. & Sauerb. has been found at the Lower
Falls of the Tahquamenon River in nearby Chippewa County (and also in Alger Co.).
is larger, with glossy leaves wide-spreading and complanate, often somewhat

(Brid.) Lindb., the brood branches are distributed throughout the length of the stems
and branches.)

323

Isopterygiopsis muelleriana (Schimp.) Iwats., a rare species on non-calcareous
rock in the Upper Peninsula (Ontonagon Co.), differs from Jsopterygium in its rigidly
distichous-complanate leaves, usually crowded and pectinate in arrangement, and large,
hyaline, relatively thin-walled cortical cells. Somewhat similar forms of Jsopterygium
pulchellum with rigidly complanate leaves have thick-walled cortical cells and narrower
leaves.

6. Taxiphyllum FI.

Plants small to medium-sized, in flat, green or yellow, often glossy mats. Stems
prostrate, sparsely to irregularly or subpinnately branched; pseudoparaphyllia generally
present and lanceolate. Stem and branch leaves similar, often crowded, spreading and
complanate in 2 apparent rows, oblong-ovate or oblong-lanceolate and short- to

916

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= =

ma

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FAAS

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Isopterygium ioaciaas 915. Habit. 916. Leaves. 917, Cells of alar reg
Taxiphyllum deplanatum. 918. Habit. 919. Leaves. 920. Cells at leaf ea "921. Cells at basal
margin of leaf. 922. Paraphyllia.

324

long-acuminate (or rarely oblong-lingulate and blunt), mostly concave, not or some-
what asymmetric; margins plane or sometimes narrowly recurved, at least below,

and thus appearing papillose, shorter at the apex and at the insertion,
subquadrate in small alar groups. Dioicous; perichaetial leaves erect at base, with
slender, somewhat spreading acumina. Setae smooth; capsules erect or inclined,
oblong-ovoid with a distinct neck; annulus persistent, of 2 rows of cells; operculum
obliquely rostrate; peristome teeth cross-striolate below, papillose above, bordered,
trabeculate; endostome hyaline, papillose, with a high basal membrane, keeled seg-
ments, and cilia in groups of 2 or 3 (rarely — Calyptrae naked. _The name refers
to the apparent arrangement of leaves in two r

Taxiphyllum deplanatum (Bruch & Schimp. ex Sull.) Fl.—Plants in loose or
dense, light-green or yellow, shiny mats. Pseudoparaphyllia lanceolate. Leaves crowded
and overlapping, spreading and strongly complanate, not plicate, 1.5-2 mm. long,
ovate-lanceolate or occasionally oblong-lanceolate, slenderly acuminate, narrowly acute;
margins plane, or narrowly recurved near the base, serrulate nearly all around (more
strongly so near the apex). Setae 6-16 mm. long; capsules 1.3-2 mm. long, erect or
somewhat inclined, not or only somewhat asymmetric, constricted below the mouth
and at the neck when dry; stomata in the neck; exostome teeth very pale and
brownish or yellowish; endostome with non-perforate segments and cilia in 2’s. Spores
about 11-13 4, nearly smooth.

Conard, How to Know the Mosses, (ed. 2) fig. 297 (as sail enear it Grout, Mosses with
Hand-lens and Microscope, fig. 199 (as Plagiothecium), Moss Flora of North America 3: Pl. 42B (as
Plagiothecium). Jennings, Mosses of Western Pennsylvania, (ed. 2) P . 69 (as Isopterygium). Welch,
Mosses of Indiana, fig. 210 (as Plagiothecium).—Fig. 918-922,

On calcareous rock or soil. Widespread in eastern North pu from Saskatchewan to
Quebec, south to North Carolina and Louisiana; Arizona and New Mexi

CHEBOYGAN CO,—Douglas Lake. MACKINAC CO.—Mackinac Island, Caffey Corner.

Recognized by the crowded and overlapping, spreading and complanate leaves
with narrow points. A related species, not found in the state, is T. taxirameum (Mitt.
l., known in most books as T. geophilum (Aust.) Fl.; it has leaves relatively distant
and more broadly pointed.

7. Hypnum Hedw.

Plants slender to fairly robust, in soft, green, yellowish, brownish-, or golden-
green mats or tufts. Stems mostly creeping, irregularly to pinnately branched; stems
and branches usually hooked at the tips; paraphyllia none or few. Stem and branch
leaves similar, crowded, secund or falcate-secund, often in 2 apparent rows, concave,
symmetric, ovate- or oblong-lanceolate, acuminate, not or somewhat decurrent; margins
erect or reflexed below, rarely revolute, entire or serrulate above; cells smooth, linear
or rhomboidal, mostly incrassate and porose at the insertion, usually clearly dif-
ferentiated at the basal angles (small and quadrate or laxly oblong, inflated, pale and
thin-walled to thick-walled and colored). Perichaetial leaves slenderly long-acuminate,
generally + striate. Setae smooth; capsules inclined to horizontal (or rarely suberect),
usually + curved and asymmetric, oblong-cylindric; annulus usually present; operculum
conic or convex-conic, mammillate to apiculate or + rostrate; stomata at base of the
capsule; peristome teeth cross-striolate and bordered below, papillose above, trabeculate
at back; endostome with a well-developed basal membrane, not or narrowly perforate,

B25

keeled segments, and cilia in 2’s and 3’s (rarely single). Calyptrae naked.—The generic
name, derived from a Greek word for sleep (as is the word hypnosis), was applied in
antiquity to some epiphytic mosses or lichens used as medicinal ingredients. Dillenius,
in 1741, used the name for pleurocarps of the sort sometimes called FEATHER
MOSSES. Such mosses would seem suitable for stuffing pillows and thus inducing
sleep, but Dillenius tried, in the spirit of scholasticism, to justify ancient usage by
imagining a soporific fragrance emanating from them. The genus has continued until
recent times to be used in an inclusive sense, meaning most of the pleurocarps, and
with good justification as most of them have the “perfect double” peristome (also
known as the “hypnaceous” peristome described above), giving obvious evidence of
common origin. In the interest of practicality, the genus has been subdivided into
many genera in several families. These families, based mainly on gametophytic
characters, show virtual uniformity in sporophytic features, especially in details of
peristome structure. For those who question the value of families so poorly dif-
ferentiated and so difficult to define, I would point out that in Lesquereux and James’
Manual of the Mosses of North America, of 1884, there are 195 species of
Hypnum—making a truly pesky and unmanageable genus! Even in Dixon’s admirable
Students Handbook of British Mosses, last revised in 1924 and still used as a standard
reference, the genus embraced at least 15 genera in common usage today, 11 of them
distributed in three families in this work. (It is interesting to note that in his many
publications Dixon exhibited Dr. Jekyll-Mr. Hyde characteristics in lumping genera
shamelessly at home but splitting them with equal abandon abroad!)

1. Alar cells small, quadrate, not thin-walled or inflated; capsules suberect
2. Rather robust plants with opaque, quadrate alar cells in rather conspicuous groups (about
4-6 along the leaf margins, below them a few somewhat larger, hyaline or orange-brown
a H. imponens
2. Rather small plants with numerous opaque, quadrate alar cells in triangular grou
3. Dioicous; leaves crowded and appressed, giving stems and branches a terete Bae ade
leaf margins erect, entire or serrulate above; alar cells many aA. ce ee
3. Autoicous; leaves less crowded, not appressed; leaf margins sae reflexed poe
serrulate ne ay all around, sharply so above; alar cells rather numerou H. pallescens
1. Alar cells hyaline, + enlarged or inflated, thin-walled; capsules inclined
4. Stem eee closely and regularly pinnate; cortical cells not se somewhat enlarged;
ee serrulate to well below the apex, usually in the upper 1/3-1/
5. Cortical cells not enlarged; pseudoparaphyllia none; leaves broad, ingular -ovate, rather
broadly acuminate, abruptly rounded above a few inflated alar cells; dioico
. H. curvifolium
5. Cortical cells somewhat enlarged; pseudoparaphyllia at branch insertions; leaves narrow
oblong-lanceolate, slenderly long-acuminate, somewhat rounded to the insertion;
autoicous 1. A. fertile
4. Stems usually ascending, loosely and irregularly branched; cortical cells Den Gere

ea

6. Leaves somewhat complanate, usually strongly concave and conspicuously falcate-
secund, often + furrowed when dry, not or slightly rounded to the insertion; alar cells
abruptly inflated in large decurrencies delimited above by small cells 1. H. lindbergii

6. Leaves loosely complanate, meuerae tely concave, curved-secund at the tips, not longi-
tudinally furrowed but often wrinkled transversely when dry, rounded to the
insertion; alar cells enlarged a not particularly inflated, in gradually differentiated
groups 2. H. pratense

1. Hypnum lindbergii Mitt. Fairly robust plants in loose, light-green to yellowish
or brownish, shiny mats. Stems ascending, irregularly or subpinnately branched,
cortical cells of stems and branches large and hyaline (stripping off with leaves on
dissection and obviously broader than the leaf cells). Leaves erect-spreading and +
complanate with falcate-secund tips (especially so at tips of stems and branches),
smooth or sometimes obscurely plicate when dry, concave, about 1-2.5 mm. long,
oblong-ovate, gradually narrowed to a fairly broad or slender, acute acumen, decurrent;

326

margins plane, entire or sometimes serrulate at the apex; costa short and double; upper
cells linear; basal cells shorter, broader, thick-walled and pitted, becoming yellowish;
alar cells abruptly inflated, hyaline, and thin-walled, in large, decurrent groups
delimited above by small, + quadrate cells. Dioicous. Setae 25-37 mm. long; capsules
inclined to horizontal, curved and asymmetric, 2-2.5 mm. long, + furrowed when dry

d em annulus large; operculum convex-conic and apiculate; cilia 2-4. Spores
13-22 p, very finely papillose.—n = 9+1, 10, 11.

Mosses of Florida, Pl. 84 (figs. 14). Conard, How to Know the Mosses, (ed. 2) fig.
oie a W. pratense) Grout, Mosses with Hand-lens and Microscope, fig. 188 (as H. patientiae).
ings, Mosses of Western Pennsylvania, (ed. 1) Pl. 46 (as Stereodon arcuatus), (ed. 2) Pl. 50 (as

. Dae Welch, Mosses of Indiana, fig. 194 (as H. arcuatum).—Fig. 923-924.

wet soil, humus, rotten wood, etc., in swampy places, particularly near oo and ponds.
Gaeae: fee North America to Washington, Colorado, and the Gulf of Mex

CHEBOYGAN CO.-—Reeses Bog, Hermits Bog, Gorge of Carp Creek, Iron Bridge, Burt |
North Fishtail Bay (Douglas Lake), Smiths Bog, Levering— enh an road, Mill Creek, Grass B
oe tview Swamp, Levering, Larks Lake, Wycamp Lake, Cecil Bay, Big Stone a

mona Beach. MACKINAC CO.-—Bois Blanc ane ven Island, McKays Creek, Cut River,
Ora PRESQUE ISLE CO.-—Evergreen Beach, Hammond Bay, Mast Point, Clinton Lake, Kelsey
Pond, Rainy Lake.

Hypnum lindbergii, very common in swampy habitats, is pale, whitish-green, and
glossy when fresh. The falcate leaves are irregularly plicate when dry, scarcely rounded
to the insertion, with abruptly differentiated alar cells. Like 1. pratense, it has large
cortical cells. This character, found in several species of Hypnum in a more restricted sense,
scarcely justifies the segregation of H. lindbergii and H. pratense as Breidleria.

2. Hypnum pratense Koch ex Spruce—Plants of medium size, in yellowish to
golden, shiny mats. Stems ascending, subpinnately branched; cortical cells of stems and
branches large and hyaline. Leaves erect-spreading and loosely complanate, with tips
slightly to moderately curved-secund (especially so at tips of stems and branches),
concave, often obscurely wrinkled-rugose when dry, about 2 mm. long, oblong-ovate,
gradually acuminate, acute, somewhat decurrent; margins plane, usually serrulate at the
tips; cells linear; basal cells shorter, wider, thick-walled, pitted, and yellowish, a few
alar cells somewhat enlarged or inflated, not sharply delimited by obviously smaller
cells. Dioicous. Setae 20-35 mm. long; capsules inclined to horizontal, curved and
asymmetric, about 2.5 mm. long, not furrowed; annulus broad; operculum convex-
conic, apiculate; cilia 2-3. Spores ca. 10-16 yw, very finely papillose.—n = 9+1, 10

Grout, Moss Flora of North America 3: Pl. 33 below. Jennings, Mosses of Western
Pennsylvania, (ed. 2) Pl. 70 (as Stereodon).—Fig. 925-927.

A calciphile growing on wet soil or humus in open swampy places. Circumpolar; arctic
America to British Columbia, Colorado, Michigan, and Pennsylvania.

CHEBOYGAN CO.—Douglas Lake, Reeses Bog, Vincent Lake. EMMET CO.—Arnott Lake.
MACKINAC CO.—McKays Creek, Rexton Lake. PRESQUE ISLE CO.—Hammond Bay.

Hypnum pratense has been greatly misunderstood in this country, perhaps as a
result of an unfortunate comparison with Plagiothecium given in Grout’s Moss Flora. It
is not likely to be confused with any species of Plagiothecium, although it is true that
it has leaves somewhat more complanate and less hamate than those of H. lindbergii,
from which it differs further in having leaves rounded to the insertion and alar cells
smaller and less inflated in gradually differentiated groups.

327.

Hypnum curvifolium has more regularly pinnate branching and more falcate-
secund leaves with margins serrulate to well below the apex; also, its cortical cells are
not noticeably large and hyaline.

. Hypnum cupressiforme Hedw.—Plants in + shiny, green, yellowish, or brown-
ish mats. Stems creeping or ascending, freely and often pinnately branched; paraphyllia
few, small, lanceolate. Leaves crowded, loosely imbricate, homomallous or falcate-
secund, concave, oblong-ovate, or ovate-lanceolate, gradually or rather abruptly
acuminate; margins erect, entire or serrulate above: upper cells rhomboidal to
linear-rhomboidal, often thick-walled; alar cells many, small, quadrate, grayish-opaque.
Dioicous. Setae up to 30 or 40mm. long; capsules suberect and curved-cylindric;
annulus present; operculum rostrate from a convex-conic base; cilia of endostome
usually 1-2. Spores 18-20 yu, finely papillose.—n = 10, 11, 16.

Conard, How to Know the Mosses, (ed. 2) fig. 283a-e. Jennings, Mosses of Western
Pennsylvania, “ed. 1) Pl. 46 & (ed. 2) Pl. 50 (Stereodon cupressiformis var. filiformis).—Fig.
28-929,

On calcareous soil or rock, often in dry, exposed places. eee cosmopolitan; widespread in
seve: an montane regions in North America but not common in the East. Known from our
a fro single collection on Mackinac Island, made by C. F. Wheeler, September 5, 1892 (in
ip Michigan coe University). The record may be viewed with suspicion.

——-——

Se
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Oe aX)
SS 9245 St

Hypnum lindbergii. 923. apes 924. Alar c
Hypnum pratense. 925. Leaves. 926. Cells Be ‘leaf tip. 927. Alar cells.
Hypnum cupressiforme. oe Leaf. 929. Alar cells.

328

Notoriously polymorphous in Europe but not causing much trouble in North
America. It can be recognized by numerous small, quadrate, opaque alar cells. Plants
may be green or brown but often have a characteristic grayish overcast. The crowded,
imbricate, smoothly concave leaves with secund tips are recognizable to the practiced
eye.

H. A. Miller has told me that Hypnum cupressiforme and allied species are often
sold as “sheet moss” at Christmas time, even in modern supermarkets.

4. Hypnum pallescens (Hedw.) P.-Beauv.—Small plants in dark- or yellow-green,
slightly shiny mats. Stems creeping, pinnately branched; paraphyllia few, small,
lanceolate. Leaves crowded, erect-spreading and + falcate-secund, 0.6-1 mm. long,
concave, gradually or abruptly acuminate from an oblong-ovate base, acute; margins
reflexed below, serrulate nearly throughout, more strongly so toward the apex; upper
cells linear-rhomboidal (about 6-8:1), thick-walled; cells at the insertion shorter, wider,
and yellowish; alar cells rather numerous, small, quadrate, thick-walled, + opaque.
Autoicous. Setae 6-13 mm. long; capsules suberect, curved-cylindric, about 1.5-2 mm.
long; annulus present; operculum obliquely rostrate from a convex base; cilia usually
paired, nodulose. Spores 12-13 u, finely papillose.—n = 10+1, 11; 2n = 2

Conard, How to Know the Mosses, (ed. 2) fig. 285 (as H. reptile). Grout, Mosses with
Hand-lens ant Microscope, fig. 190 (as H. reptile), Moss Flora of North America 3:
Jennings, Mosses of Western Pennsylvania, (ed. 1) Pl. 45 & (ed. 2) Pl. 49 (as Stereodon reptilis).
Welch, Mosses of Indiana, fig. 195 (as H. reptile).—Fig. 930-932.

On bark at base of trees, less commonly on logs, rocks, and other substrata, in woods.
ee British Columbia to Nova Scotia, south to Arizona in the West, North Carolina in the

CHEBOYGAN CO.—Reeses Bog, Hermits Bog, Colonial Point, Douglas Lake, Bryants Bog,
Livingston Bog, Mud Lake, Mullett Lake, Grass Bay, Little Lake 16, Weber Lake. EMMET
CO.—Pellston Hills, Five Mile Creek, Wycamp Lake, Cecil Bay, Wilderness State Park, Conway Bog.
MACKINAC CO.-—Bois Blanc Island, Mackinac Island, St. Martins Point, McKays Creek, Bush Bay,
Cut River, Ozark, Hendricks, Little Dollar Lake. PRESQUE ISLE CO.—Presque Isle Lighthouse,
Evergreen Beach, Hammond Bay, Mast Point, Rainy River Falls, Clinton Lake, Loon Lake.

Very common at the base of trees, Meee hardwoods, where Brachythecium
reflexum, Platydictya subtile, and H allium adnatum also grow, this species is
recognized by its small size, falcate ee vith serrulate acumina, and margins recurved
below. Grout tried to distinguish H. pallescens and H. reptile Mx. (largely on the basis
of altitude and distribution), but the type specimens are essentially similar (and even if
one wishes to recognize a specific difference, the names would need to be reversed in
usage to conform to the type concepts).

5. Hypnum imponens Hedw.—Plants fairly robust and handsome, in yellow-
brown to yellow-green mats. Stems creeping, regularly pinnate, red-brown; paraphyllia
relatively numerous, polymorphous, often lobed or dentate. Stem leaves about 2 mm.

linear, incrassate and porose, usually orange; alar cells quadrate, incrassate and
somewhat opaque in rather conspicuous groups (usually about 4-6 cells along the
margin), below them a few somewhat inflated, quadrate, clear or orange-brown cells.
Branch leaves smaller with fewer quadrate alar cells. Dioicous. Setae 10-35 mm. long,

329

red; capsules 2.5-3 mm. long, cylindric, suberect, symmetric or slightly curved, smooth
red-brown; annulus none or narrow and poorly differentiated; operculum high-conic
and sometimes shortly and stoutly rostrate; cilia of endostome single, variable in
length, often well developed but fragile. Spores 15-22 y, very finely papillose.—n = 6-7
fim ale

arlington, Mosses of Michigan, fig. 133. Grout, Mosses with Hand-lens and Microscope, fig.
b, 185. Jennings, Mosses of Western Pennsylvania, (ed. 1) Pl. 45, (ed. 2) Pl. 49 (as
panera Welch, Mosses of Indiana, fig. 191.—Fig. 933-937.

Typically on rotten logs but often on rocks, humus, or bark at base of trees in moist woods
Europe and Asia; Yukon; Newfoundland to Wisconsin and south to the Gulf of Mexico.

HEBOYGAN CO.-—Reeses Bog, Gorge of Carp Creek, Iron Bridge, Livingston Bog, Mud
Lake, Weber Lake. EMMET CO.—Carp Lake, Five Mile Creek, Mackinaw City Hardwoods, Big

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Hypnum pallescens. 930. Habit. 931. Leaves. 932. Alar cells.
Hypnum imponens, 933. Habit. 934. Leaf. 935. Alar eile 936. Paraphyllia. 937. Capsule.

330

Stone Creek. MACKINAC CO.-—Prentiss Bay, Cut River, McNamara Pond. PRESQUE ISLE
CO.—Lamprey Experiment Station, Evergreen Beach, Hoeft State Park.

Hypnum imponens has red-brown stems and, for the genus, relatively numerous,
often branched paraphyllia. The leaves are not much narrowed to the insertion (as
compared with H. curvifolium), and the rather thick-walled, orange-brown, quadrate
alar cells are distinctive, as are the suberect capsules.

Mat-forming mosses, such as Hypnum imponens, are important in forest seeding.
In the Great Lakes region, sugar maples ordinarily spring up in countless numbers in
upland forests that have been clean cut, occupying the surface to the exclusion of all
other tree species and thus completely controlling succession. But sugar maple seedlings
will not grow on logs. If logs are left behind after cutting, they provide habitats
favorable to the establishment of hemlock and yellow birch in mixture with maple.
(See also the discussion of Polytrichum juniperinum, which may serve in the
re-forestation of pasture lands by providing a seed bed.)

6. Hypnum curvifolium Hedw.—Plants relatively robust, in yellow-green to
brownish aie light-green), + shiny, flat mats. Stems prostrate, orange-brown or
reddish, regularly pinnate; cortical cells not enlarged; pseudoparaphyllia none. Stem
leaves smooth, falcate-secund, triangular-ovate, gradually narrowed to a relatively broad
acumen, abruptly rounded at the insertion and somewhat decurrent; margins plane or
erect, serrulate at the apex and sometimes sinuate-serrulate in the upper third or more,
sinuolate near the base; cells linear-flexuose above, shorter, broader, porose, and often
yellow or orange at the insertion, at the basal angles a few hyaline, thin-walled,
inflated cells forming short decurrencies, above them a few, small and inconspicuous,
short-oblong cells. Apparently dioicous. Setae 20-47 mm. long, orange-yellow; capsules
2.5-3mm. long, strongly curved, obovoid-cylindric from a tapered neck, furrowed
when dry and empty; annulus broad, persistent; operculum convex-conic, apiculate;
cilia in 2’s and 3’s, well developed, nodulose. Spores 18-24 u, very finely papillose.

Conard, How to Know the Mosses, (ed. 2) fig. 281. Grout, Mosses with Hand-lens and
Microscope, fig. 187. Jennings, Mosses of Western hak: (ed. 1) Pl. 46 & (ed. 2) Pl. 50 (as
Stereodon). Welch, Mosses of Indiana, fig. 193.—Fig. 938-940.

On rocks, soil, logs, or bark at base of trees in moist or wet, shaded places. Japan;
widespread in eastern North America

MACKINAC CO.—Mackinac Island.

The broadly acuminate leaves are abruptly narrowed to a shortly decurrent base.
The few cells of the decurrencies are somewhat enlarged, thin-walled, and hyaline. The
curved and strongly inclined capsules are plicate when dry and empty

7. Hypnum fertile Sendtn.—Plants relatively slender, in soft, yellow-green or
yellow-brown, flat mats. Stems prostrate, orange or brownish, closely and evenly
pinnate; cortical cells somewhat enlarged; pseudoparaphyllia at branch insertions. Stem
leaves smooth, falcate-secund, about 1.5-2 mm. long, concave, oblong-lanceolate, very
slenderly long-acuminate, somewhat rounded’ to the insertion and decurrent; margins
erect, entire or somewhat serrulate near the apex; cells long-linear, flexuose, somewhat
thicker-walled and + porose at the insertion; alar cells oblong, thin-walled, inflated, and
hyaline in small decurrencies bordered above by a small group of small, obscure cells.
Branch leaves similar but smaller, with broader acumina, serrulate in the upper half.
Autoicous. Setae 13-25 mm. long, reddish, flexuose; capsules 2-2.2 mm. long, hori-

331
zontal, curved and asymmetric, oblong-cylindric, smooth or somewhat furrowed when
dry and empty; annulus revoluble; operculum conic, blunt or acute; cilia of endostome

in 2’s, well developed, nodose or appendiculate. Spores 11-13, very finely papil-
lose.—n = 11.

Grout, Mosses with Hand-lens and Microscope, fig. 189. Jennings, Mosses of Western
Pennsylvania, (ed. 1) Pl. 45 & (ed. 2) Pl. 49 (as Stereodon).—Fig. 941-942.
Tennessee.

On rotten logs in woods. Europe and Asia; Labrador to Manitoba, south to Michigan and
CHEBOYGAN CO,—Colonial Point, Little Lake 16.
Smaller than H. curvifolium, with more slenderly acuminate leaves which are not

rounded to the insertion. A few cells at the extreme basal angels of the leaves are
somewhat inflated, hyaline, and thin-walled, and the cortical cells are rather large.

8. Herzogiella Broth.
Plants of rather small size in loose or dense, shiny, green, yellowish, or brownish

mats. Stems creeping, freely and irregularly branched; branches spreading or ascending;
cortical cells large and thin-walled; pseudoparaphyllia none; brood-bodies none. Leaves

ee
SS
Se

aie
S Gres rae a
NE :

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Se

=
SSS
SSS
ISS.
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Fp BS =
a gag is

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a
a

Ke

Hypnum curvifolium, 938. Leaf. 939. Alar cells. 940. Capsule, dry.
Hypnum fertile. 941, Leaf. 942. Alar cells.

Herzogiella turfacea. 943. Habit. 944. Leaves. 945. Cells of alar region. 946. Capsule, dry.

332

of stems and branches similar, wide-spreading and not or loosely complanate, ovate or
oblong-lanceolate, slenderly acuminate, not or strongly decurrent; margins plane,
serrulate in the upper 1/2-2/3; cells re scarcely differentiated at the basal angles or
hyaline, thin-walled, and inflated in small but distinct, decurrent alar groups.
Autoicous. Setae smooth; capsules suberect to strongly inclined, oblong-cylindric,
becoming curved-cylindric, constricted below the mouth, and plicate when dry and
empty; annulus well developed; operculum bluntly low-conic; stomata at base of the
um; peristome yellowish or yellow-brown and cross-striolate below, hyaline and
papillose above, bordered, trabeculate; endostome hyaline, finely papillose, with a high
basal membrane, keeled, non-perforate segments, and nodulose, single or paired cilia.
Calyptrae naked._Named for Theodor Herzog, distinguished bryologist and author of
Die Geographie der Moose.

Herzogiella turfacea (Lindb.) Iwats.—Rather small, yellow- or bright-green plants
in loose mats. Branches spreading. Leaves wide-spreading and loosely and indistinctly
complanate, often + secund at the tips, sometimes somewhat plicate when dry,
1.2-1.5 mm. long, oblong-lanceolate, gradually narrowed to a long, flexuose acumen,
not decurrent; cells linear-flexuose, scarcely differentiated at the basal angles. Peri-
chaetial leaves flexuose or spreading at the tips. Setae 8-18 mm. long, red; capsules
strongly inclined and horizontal, 1.5-2 mm. long; peristome teeth yellowish; cilia of
endostome paired. Spores 9-11 u, minutely roughened.—n =

ut, Mosses with Hand-lens and Microscope, Pl. 83 (as near Jennings, Mosses of
Western. an (ed. 1) Pl. 47 & (ed. 2) Pl. 51 (as /sopterygium).—Fig. 943-946.

n rotten logs and stumps, less commonly on humus or bark at base of trees, in moist,
coniferous Woods Europe and Asia; Newfoundland to South Dakota, south to Michigan, Ohio, and
South Carolin

CHEBOYGAN CO.—Reeses Bog, Hermits Bog, Gorge of Carp Creek, Iron Bridge, Burt Lake,
Bryants Bog, Livingston Bog, Mud Lake, Mullett Lake, Duncan Bay, Grass Bay, Little Lake ie

Cecil Bay, Conway Bog, Five Mile Creek, Ramona Beach. MACKINAC CO.—Bois Blanc Island,
Mackinac Island, Epoufette, Little Dollar Lake. PRESQUE ISLE CO.—Ocequeoc River, Lamprey
Experiment Station, Evergreen Beach, Clinton Lake.

The plants are easily recognized but difficult to characterize. The narrow,
wide-spreading leaves give an impression of being complanate with somewhat decurved
tips, but they are really loosely arranged and on inspection seem scarcely complanate
at all

Herzogiella striatella (Brid.) Iwats., a species of more northern or montane
distribution, occurs at the Upper Falls of the Tahquamenon River in Luce County and
also at the Lower Falls in Chippewa County (and elsewhere in the Lake Superior
region). It has leaves spreading from an erect base, not or obscurely flattened and not
at all secund, with abruptly inflated, decurrent alar cells.

9. Ctenidium (Schimp.) Mitt.

Plants rather small or medium-sized, in dense, soft, green, yellowish, or golden-
brown, shiny mats. Stems creeping or more crowded and ascending, pinnately
branched; -branches short; paraphyllia few, at branch insertions. Stem and branch leaves
differentiated: Stem leaves crowded, spreading to squarrose or squarrose-recurved and
usually + secund at the tips, smooth or somewhat plicate, symmetric, abruptly
acuminate from an ovate-cordate, broadly decurrent base, distinctly serrate all around;

335

cells linear above, subquadrate in the decurrencies. Branch leaves loosely falcate-secund,
smaller, lanceolate, gradually acuminate, only slightly decurrent, with upper cells
papillose at back because of projecting upper angles. Dioicous; perichaetial pee erect,
oblong, with abruptly acuminate tips. Setae smooth or + papillose; capsules strongly
inclined to horizontal, oblong-cylindric, not or somewhat narrowed below the mouth
when dry, smooth; annulus present; operculum conic, sometimes apiculate; peristome
teeth cross-striolate below, papillose above, bordered and trabeculate; endostome
densely and finely papillose, with a high basal membrane, keeled and not or somewhat
perforate segments, and nodose cilia in groups of 2-3. Calyptrae usually + hairy.—The
name means comb-like, branched in a pectinate manner.

Ctenidium molluscum (Hedw.) Mitt.-Stem leaves 1.2-2 mm. long. Branch leaves
about 1-1.4mm. long. Phyllodioicous. Setae 13-20mm. long, smooth, red-yellow,
becoming red with age; capsules about 2mm. long; operculum apiculate; stomata in
the neck. Spores 11-15 y, very finely papillose. Calyptrae very sparsely hairy.—n = 7, 8,
10

Conard, How to eee the Mosses, (ed. 2) fig. 280 (as Hypnum). Darlington, Mosses of
Michigan, fig. 139. Grout, Mosses with Hand-lens and Microscope, fig. 183 (as Hypnum). Jennings,
Mosses of Western ate (ed. 1) Pl. 42, (ed. 2) Pl. 46. Welch, Mosses of Indiana, fig. 196
(as Hypnum).—Fig. 947-950.

On soil or humus and rock, sometimes on decayed wood, locally on the floor of Thuja
swamps. Europe, North Africa, Macaronesia, and Asia; Newfoundland to Michigan, south to South
Carolina and Louisiana

CHEBOYGAN CO.-Iron Bridge. EMMET CO.—E. of Woodland School, Arnott Lake.

SS a { J Ee VY) : Les AY ( | ‘ al :
Pee \e as

Cte a ge 947. Habit. 948. Stem leaves. 949. Branch leaves. 950. Cells of upper
part of branch 1

334

Rather small mosses of a soft, feathery appearance, notable for the cordate-
triangular, broadly decurrent, and abruptly acuminate stem leaves and leaf cells
minutely papillose at the upper angles at back.

10. Ptilium De Not.

Robust, strikingly handsome plants in extensive, bright-green, yellow, or golden,
shiny mats. Stems erect-ascending, at least toward the tips, simple or forked, regularly
l-pinnate, forming flat, oblong or oblong-triangular fronds whcih are hooked or rolled
at the tips; branches crowded, horizontally spreading, hooked at the tips; paraphyllia
few, at branch bases. Stem leaves considerably larger than branch leaves, loosely
falcate-secund, broadly triangular-ovate, gradually long-acuminate, plicate; margins
plane, serrulate above; cells smooth, long-linear, flexuose, those at the insertion shorter,
broader, incrassate, and porose, a few cells at the basal angles short-rectangular, pale
and thin-walled in inconspicuous groups. Branch leaves circinate and secund, plicate,
oblong-lanceolate, slenderly long-acuminate; margins irregularly revolute below,
serrulate above; costa short and double or lacking; cells at insertion and at basal angles
only poorly differentiated. Dioicous; perichaetial leaves very long, erect, lance-subulate.
Setae smooth; annulus narrow; operculum conic, apiculate; peristome teeth cross-
striolate and bordered below, papillose above, trabeculate; endostome pale, papillose,
the basal membrane high, segments broad, keeled, not or narrowly perforate, cilia well
developed, nodulose. Calyptrae naked.—_The name refers to a plume-like appearance.

Ptilium crista-castrensis (Hedw.) De Not.—Plants 3.5-11 cm. high, beautifully
plumose, green to golden, shiny. Stem leaves 2-3 mm. long. Branch leaves 1.2-2 mm.
long. Setae 25-45 mm. long, reddish; capsules nearly horizontal, 2-3 mm. long, chest-
nut-brown; peristome teeth dark, reddish-brown; endostome pale-yellow; cilia in groups
of 2 or 3, rarely single. Spores 12. 16 uw, smooth or very finely papillose.—n = 9+1, 10,
10+1, 11, 20.

Conard, How S Know the Mosses, Se fig. 279 (as Hypnum). Darlington, Mosses of
Michigan, Ss 138. Grout, Mosses with Han and Microscope, figs. 181, 182, 184a,c,e (all as
Hypnum). Jennings, aes of Western aa (ed. 1) Pl. 45, (ed. 2) Pl. 49.—Fig. 951-953.

On humus and old mossy logs in moist forests. Sapicatea Alaska to Newfoundland, south
to Oregon, Montana, Iowa, Michigan and North Carolina

CHEBOYGAN CO.—Reeses Bog, Hermits Bog, Gorge of Carp Creek. EMMET CO.—Levering,
Galloway Bog, Maple River, Stutsmanville, Middle Village, Wycamp Lake, Cecil Bay, Big Stone Bay,
Conway Bog. MACKINAC CO.-—Bois Blanc Island, Mackinac Island, St. Martins aaeee ee
Rexton Lake, Ozark, McKays Creek. PRESQUE ISLE CO.—Lamprey Experiment Statio

A truly splendid moss, appropriately called the KNIGHT’S PLUME. The shiny,
feathery fronds curved at the tips, with clearly plicate leaves, are distinctive, although
the species is similar in many ways, including the elongate perichaetium, to Drepano-
cladus uncinatus. In rare cases of doubt, the single costa of the Drepanocladus will
decide the matter.

In the Popular History of British Mosses (1860), Stark referred to Ptilium
crista-castrensis as “one of the prizes which botanical tourists should endeavour to
carry south, as a remembrance of the land of brown heath and shaggy wood, for its
graceful feathery branches suit admirably either for the scientific cabinet or the
drawing-room scrapbook.

The ssecihie epithet would seem to have an ending grammatically out of
agreement with the generic name. Crista (a crest) is actually a noun in apposition

335

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Ptilium crista-castrensis. 951. Habit. 952. Stem leaf. 953. Branch leaf

rather than a simple epithet and castrensis (castle or fort) agrees with it rather than

with Prilium.
It has been erroneously stated that this moss is figured in the coat of arms of the
Dukes of Lancaster. Similar looking plumes are featured there and in numerous other

devices of heraldry, but not mosses!
CLIMACIACEAE

Rather large, gregarious mosses on swampy soil. Primary stems subterranean and
rhizome-like, tomentose; secondary stems typically erect and dendroid; branches simple

336

or 1-2-pinnate, densely foliate, terete, + attenuate; paraphyllia abundant, filiform,

cordate, blunt or acute, + auriculate at base, + plicate, toothed at least above; costa
single, strong, broad at base, ending below the apex; cells smooth, rhomboidal to linear
above, with firm or thick walls, colored and + porose at base, scarcely to clearly
differentiated at the basal angles. Dioicous; perichaetia on secondary stems and bases
of branches; perichaetial leaves long-sheathing. Setae elongate, sometimes clustered;
capsules cylindric, smooth, erect and symmetric or curved and strongly inclined;
annulus none; operculum rostrate from a convex base; stomata in the neck; peristome
double, the 16 teeth lanceolate, red-brown, papillose or cross-striolate, bordered,
trabeculate; endostome about as long as the teeth, the basal membrane very low to
fairly high, segments keeled and narrowly perforate along the middle to gaping and
finally split to the apex, cilia none. Calyptrae cucullate.

Climacium Web. & Mohr

Plants dull, dark-green or yellow-brown, typically erect and dendroid. Branch
leaves oblong-ovate to subcordate, bluntly acute to short-acuminate, coarsely toothed
above, + decurrent and auriculate at base; costa toothed at back above; cells
rhomboidal to linear, colored and pitted at the insertion, not much differentiated at
the basal angles. Setae single or clustered; capsules erect and symmetric; peristome
teeth dark-brown, papillose; endostome yellowish, with a very low basal membrane and
linear segments papillose in vertical lines, gaping and finally split along the median line.
Spores spherical. Calyptrae covering the entire capsule, somewhat twisted.—_The name
refers to the broad perforations of the endostome segments which suggest a ladder.

Climacium dendroides (Hedw.) Web. & Mohr—Plants shiny when dry, yellow or
yellow-green, rather soft, 3-9, rarely 13 cm. high. Branch leaves erect, nearly smooth to
sulcate, 2-3 mm. long, ovate or ovate-lanceolate, + rounded above the insertion but not
or only slightly auriculate; upper median cells 5-9:1, oblong-rhomboidal; alar cells pale,
thin-walled, and slightly inflated in small and inconspicuous groups. Setae 18-45 mm
long; capsules 1.5-3 mm. 7 one cylindric; operculum + rostrate. Spores 13-22 u,
minutely papillose.—n = 9+2,

Conard, How to Know the Mosses, (ed. 2) fig. 195d-e. Darlington, Mosses of Michigan, fig.
94, Grout, Mosses with Hand-lens and Microscope, fig. 158d-e, 159a. Welch, Mosses of Indiana, fig.
134.—Fig. 954-958

On soil or humus in shady, swampy places. Circumpolar; Alaska to northern Quebec, south
to pains Wisconsin, New Mexico, Arizona, and California.

CHEBOYGAN CO.—Reeses Bog, Hermits Bog, Carp Creek, Iron Bridge, Smiths Bog, Vincent
Lake, Fontinalis Run, Bessey Creek, Douglas Lake, Colonial Point, Mud Lake, Grass Bay, Weber
Lake. EMMET CO.—Maple River, Pleasantview Swamp, Levering, Galloway mae Cross Village, ne
Bay, Big Stone Bay, Conway Bog. MACKINAC CO.—Bois Blanc toland. St. Martins Point, Ho
Island Point, Epoufette, McNamara Pond, Hendricks, Peggley Lake, Round sand, eee Case
PRESQUE ISLE CO. —Mast Point, Lamprey Experiment Station, Clinton Lake, Rainy Lake.

Climacium dendroides, commonly known as the TREE MOSS, has a broad,
northern distribution overlapping in eastern North America with the more southerly C.
americanum Brid., which differs as follows: Plants dull and dirty-green, with auriculate
branch leaves with shorter upper cells (2-5, rarely 7:1) and longer, cylindric capsules
(3.5-6 mm. long). Both species occur in southern Michigan. Sporophytes are rare in
either species.

337

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Climacium dendroides. 954. Habit. 955. Leaves. 956. Upper cells of leaf. 957. Paraphyllia.
958. Capsule.

Climacium kindbergii Hg & Card.) Grout is an irregularly branched form of C.
americanum induced by floodin

Thamnobryum alleghaniense (C. M.) Nieuwl. resembles a stepped-on Climacium,
but the branches are drooping and swept to one side, and there are no paraphyllia. It
is characteristic of sandstone areas and may be discovered in the Lake Superior region.

Climacium and other mosses have been used in decorating women’s hats, but as
Frances Tripp (British Mosses ...1888) observed, not very well: “And a lady’s cap
which we saw in a window was in no wise adorned by having sprays of artificial moss
tacked all over it...Nor is a bunch of moss which has died of thirst suitable for
trimming a bonnet. The chief beauty bonnets at present possess is their being fresh and
clean; dry moss is particularly fusty looking, and it is not improved by being dyed of a
leather color(!!), or a violent blue-green (!!!), the latter being the worst, as a bad
match, for moss is never a blue-green.”

The economic uses of bryophytes, decorative and otherwise, are recorded by
Ando (Jour. Biol. Soc. Hiroshima Univ. 7: 23-26. 1957), Nelson & Carpenter (Econ.
Bot. 19: 70. 1965), Stark (British Mosses, ed. 2, 1860), Thieret (Bull. Chicago Nat.
Hist. Mus., Sept. 1954, pp. 4, 6; Econ. Bot. 10: 75-91. 1956), and Welch (Proc.

338

Indiana Acad. Sci. 58: 31-46. 1948). There is no dearth of information on uses of
Sphagnum (which see).

The studied simplicity of Japanese moss gardens approaches the Zen Buddhist
ideal of beauty—nothingness. Avery’s essay (Plants & Gardens 22: 44-48. 1967)
provides insight into the meaning of Japanese gardens in general. Iwatsuki & Kodama
(Econ. Bot. 15: 264-269. 1961) give an informative bryological review of moss
gardens. Other information is provided by Ando (Plantes de Mont. 5: 290-294. 1971),
Pullar (Plants & Gardens 22: 32-33. 1967), Schofield (Davidsonia 1: 18. 1970), and
Steere (Garden Jour. 18: 2-11. 1968).

In Sunset Ideas for Japanese Gardens (edited by Jack McDowell), directions are
given for starting moss cultures (p. 69). Partially dried moss is crumbled over a layer of
cheesecloth overlying a sand-peatmoss mixture (or sawdust) in a flat. A second piece of
cheesecloth is laid on top and kept moist. In about 4% months the double layer of
cloth with attached moss can be lifted out as an interwoven turf and cut into pieces
which can be plastered with mud or wired in place for growth. I have found that
vermiculite is better, because organic substrates so quickly mold. | recommend using
ecologically undemanding perennials, such as Mnium cuspidatum or Brachythecium
salebrosum. New growth formed by regeneration works better than mats transplanted
from nature. Constant humidity and indirect lighting (under a greenhouse bench, for
example) favor growth of cultures.

Forman (Bryol. 71: 344-347. 1968) reported that caloric values of 13 mosses
from Mt. Washington, New Hampshire, are among the lowest in the ecosystem. Such
values may be related to a low metabolic rate, evidenced by slow growth to a small
size, and also to a virtual lack of tissue differentiation providing for food storage. On
the other hand, Sugawa (Hikobia 2: 119-124. 1960) found some food value in Barbella
pendula (Sull.) Fi. He found it rich in vitamin B>, in dry and living states. On adding
powdered material to food given to puppies and chickens, he noted no distaste for the
mixtures and no ill effects. In fact, animals so fed gained more weight than those in a
control series. (The sample was, however, very small.)

Robert Pegau has told me, in correspondence, that reindeer herded in Alaska
scarcely digest mosses at all, although lichens (which have high food values) are readily
digested.

In nature animals rarely eat bryophytes, possibly because of their taste! Mizutani
(Misc. Bryol. et Lichen. 2: 100. 1961) reported that 12 out of 95 dried specimens of
the liverwort Jamesoniella autumnalis were so bitter that gargling was necessary to
wash the taste from the mouth. Other specimens were less bitter, and some not at all,
there being some apparent geographic pattern in the phenomenon in Japan. Another
liverwort, Porella vernicosa, had a terrible taste, difficult to describe, in both dry and
fresh conditions. A moss, Rhodobryum giganteum, had a sickening sweetness in fresh
but not dry material; this taste too had to be rinsed from the mouth.

Most mosses in eastern North America have an unpleasant taste, something like
raw green beans and, when strong, rather peppery and distinctly unpleasant, rarely

medicinal flavor, something like witch hazel or camphor. I have not sampled enough of
the thallose liverworts to generalize. Preissia quadrata is hot enough to tingle the tip of
the tongue and burn the back of the mouth. Conocephalum conicum has a spicy-sweet
smell, not registering in the mouth as a taste, however. Sphagna generally have a slight
fishy-oily taste. I find some variation between specimens, here and there, but no
patterns of geographic differentiation. Nor do I taste anything other than straw-flavors
in dry material!

Skin rashes sometimes affecting forest workers, known as cedar, spruce, or wood
poisoning, have been traced to such lichen substances as usnic acid. A similar rash may

339

be caused by contact with certain liverworts, especially species of Frullania and
apparently other bryophytes, including mosses, as well (Mitchell et al., Arch. Derm.
100: 46-49, 1969),

Edible or not, bryophytes do have importance in nature, in providing shelter to
small animals or seed beds for other plants, in controlling erosion and flooding, and in
altering drainage and preserving permafrosted conditions of arctic soils.

Water bears (tardigrades) may be associated with mosses or lichens on the bark
of trees and elsewhere. Such an association may only express a similar ability to
withstand frequent drying. It was reported by Felféldy and Iharos (Borbasia 7: 31-38.
1947) that tardigrades are associated with Didymodon tophaceus but only in tufts
above the wet zone at lake shores in Hungary; they believed that it was not the species
of moss but the uniformity of conditions offered by the tufts that favored the
tardigrades. Richards (Trans. Brit. Bryol. Soc. 1: 16. 1947) reported that Fontinalis
antipyretica, introduced into South Africa to harbor small crustaceans as food for
trout, had an adverse effect on the larvae of aquatic insects. The associations of
arthropods and bryophytes are reviewed by Gerson (Bryol. 72: 495-500. (1969) 1970).

But all uses aside, who can measure the comfort that mosses and liverworts add
to a cheerless planet, just by being?

“The Mosses are humble plants, but they have no insignificant part to play

either in the economy of Noe or in the physiognomy of the landscape. Trees, walls,

I and ruins assume miling or picturesque aspect under their covering in its

varied colours.”

Fiquier, The Vegetable World, 1872.

I like Frances Tripp’s observation (in British Mosses ... 1888): “It is in winter,
and in earth’s barren spots, that the moss best flourishes. Is it not a type of the
countless little mercies, which. in times of trial, are often sent to us?” But her prose
got purple: ‘““Mosses are useful to the insect tribe, countless numbers of which find
homes among their branches, and roam about in their shades as in mighty forests,
looking with their thousand eyes upon the wonders of their leaves, and sunning their
wings of purple and of gold, and burnishing their shining armour upon the polished
columns of their urns.”

HYLOCOMIACEAE

Rather robust plants in loose mats or tufts. Stems creeping or erect-ascending,
sparsely to pinnately branched, sometimes frondose; paraphyllia none, few, or man
Leaves crowded in several rows, + concave, symmetric, broadly ovate to oblong-
lanceolate, broadly acute to gradually or abruptly + long-acuminate, sometimes plicate
or rugose; costa single or double and fairly elongate, often slender or weak, sometimes
papillose at back because of projecting angles, not or + differentiated at the basal
angles. Dioicous; perichaetia lateral; perichaetial leaves erect or sheathing at base,
long-acuminate, the subulae usually wide-spreading. Setae elongate, smooth; capsules
horizontal to pendulous (or rarely erect), oblong-ovoid to oblong-cylindric, mostly
asymmetric; annulus generally present; operculum conic or convex, acute, mammillate,
or shortly and obliquely rostrate; stomata in the neck; peristome teeth lance-subulate,
cross-striolate and bordered below, papillose above, trabeculate; endostome with a high
basal membrane, keeled and perforate or gaping segments, and often nodose or
appendiculate cilia in 2’s and 4’s. Spores spherical. Calyptrae cucullate, smooth, naked.

Paraphyllia none or very few and restricted to the branch bases; stem and branch leaves not
much differentiated 1. Rhytidiadelphus
Paraphyllia very abundant, on stems and branches; stem and branch leaves differentiated
2. Hylocomium

340
1. Rhytidiadelphus (Lindb. ex Limpr.) Warnst.

Rather coarse and robust plants in loose, green or yellowish, dull or + shiny mats or
tufts. Stems decumbent or erect-ascending, sparsely branched to subpinnate or closely
branched in various directions; branches short and blunt or long and tapering, often
decurved and sometimes radiculose at the tips; paraphyllia and pseudoparaphyllia none.
Stem and branch leaves similar, crowded, erect and oblong-ovate or cordate at the base
and gradually or abruptly acuminate, generally squarrose, squarrose-recurved, or
falcate-secund at the acumen, mostly + striate or plicate; margins plane, serrulate to
serrate; costa double, short or fairly long and reaching the leaf middle or more, or
sometimes lacking; cells linear, smooth or spinose-papillose at back because of
projecting angles, broader and shorter below, + incrassate, porose, and colored at the
insertion, not or somewhat differentiated at the basal angles. Perichaetial leaves with
squarrose-recurved subulae. Capsules horizontal, oblong-ovoid, short-necked, asym-
metric, somewhat furrowed when dry; annulus of 2-3 rows of cells; operculum conic
and apiculate; cilia of endostome nodulose, in groups of 1-3.—Although the name has
been interpreted as meaning twin wrinkles, it was intended to refer to a brotherly
relationship to the genus Rhytidium (which has wrinkled or undulate leaves).

Leaves gradually tapered > a noe: flat, erect- to wide-spreading acumen, irregularly plicate;

leaf cells dentate-papillose at bac L. R. triquetrus
se ae abruptly cone to a slender, channeled, squarrose or squarrose-recurve
cumen, not plicate; cells smooth 2. R. squarro sus

1. Rhytidiadelphus triquetrus (Hedw.) Warnst.—Plants dark- to bright- or yellow-
green. Stems decumbent or erect, up to 20cm. long, irregularly and unequally
branched to closely and equally branched on all sides; stems and branches orange-red;
branches horizontal, mostly tapered and decurved at the tips. Stem leaves erect- to
wide-spreading from a clasping base wet or dry, 3.5-5 mm. long, ovate-cordate, broadly
acuminate, irregularly plicate; margins densely serrate in the upper half, rather
remotely serrulate nearly to the base; costae slender, nearly parallel, extending about
2/3 the leaf length; upper cells oblong-linear to linear, thin- or rather firm-walled, +
porose, coarsely dentate-papillose at back because of projecting cell ends (and
sometimes also rather finely papillose here and there on the upper surface because of
similar projections); cells at insertion thick-walled, porose and yellow, the alar cells
scarcely differentiated. Branch leaves erect or erect-spreading from a clasping base,
1.5-3mm. long (or sometimes more), oblong-ovate and acuminate, usually distantly
serrulate all around but sometimes densely serrate near the apex. Setae 15-45 mm.
long; capsules 1.5-3 mm. long, red-brown; cilia of endostome in groups of 1 or 2.
Spores 20-24 yp, faintly papillose to nearly smooth.—n = 5, 6, 10

Grout, Mosses with Hand-lens and Microscope, fig. 141 (as Hylocomium), Moss Flora “4
North America 3: Pl. 30B. Jennings, Mosses of Western Pennsylvania, (ed. 1) Pl. 43, (ed. 2) P
47,—Fig. 959-961

On soil, humus, and logs in ee (or moist forests. Circumpolar; Alaska to Labrador, south to
California, Means Arkansas, and Flor

CHEBOYGAN CO.—Reeses Bog, Hermits Bog, Gorge of Carp Creek, Iron Bridge, ae el
EMMET CO.—Levering, Pleasantview Swamp, Maple River, Cecil Bay, Big Ston e Bay, Con Bog.
MACKINAC CO.-—Bois Blanc Island, Mackinac Taand, St. Martins Point, McKays Creek, Bush Bay,
Prentiss Bay, Cut River, Epoufette, Ozark, Caffey Corner. PRESQUE ISLE CO.—Mast Point,
Lamprey Experiment Station, Clinton Lake.

A coarse, shaggy moss in loose tufts in coniferous woods, R. triquetrus has
branches wide-spreading and somewhat 3-ranked. The leaves are crowded and both

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Rhytidiadelphus triquetrus. 959. Habit. 960. Leaf. 961. Upper cells of leaf.

plicate and rugose. The orange leaf insertion, strong, double costa, and leaf cells
stoutly papillose at the ends at back are distinctive features helpful in identification
T ommon name of SHAGGY MOSS seems appropriate, as does R

S, in reference to a coarse and loutish appearance. (I like less Nina Marshall’s
me on TRIANGULAR WOODREVELER!)

342

2. Rhytidiadelphus squarrosus (Hedw.) Warnst.—Plants bright- or yellow-green,
somewhat shiny (often grayish-green when dry), up to 15 cm. high. Stems decumbent
or loosely ascending, remotely and irregularly to subpinnately branched; stems and
branches orange-red; paraphyllia none. Stem leaves usually not particularly crowded,
2.5-3.2 mm. long, cordate-ovate, rounded to the insertion, and strongly sheathing at
base, rather abruptly narrowed to a long, slender, channeled, squarrose or squarrose-
recurved acumen, not plicate; margins serrulate at the tip and indistinctly sinuate-
serrulate to the middle or more; costae ending about 4 up the leaf; cells linear, smooth
at back; alar cells somewhat enlarged, pale, and oblong in fairly well-marked groups.
Branch leaves somewhat smaller, usually more crowded and less strongly spreading.
Setae 17-33 mm. long; capsules 1.8-2.5 mm. long; cilia in 3’s. Spores 11-17 u, very
finely papillose.—n = 6-8 (as estimated by Heitz), 10.

Conard, How to Know the Mosses, (ed. 2) fig. 289. Darlington, Mosses of Michigan, fig. 141.
Grout, Mosses with Hand-lens and Microscope, fig. 142.—Fig. 966-967.

Very rare in our area. On soil, humus, logs, or rocks in wet, shady places. ore and Asia;
Alaska to oe and Montana; Greenland: Newfoundland to Michigan and Tennessee.

CHEBOYGAN CO.—Reeses Bog and “Douglas Lake.”

Known by its pale color, rather tall and sparsely branched stems, and squarrose-
recurved stem leaves.

Rhytidiadelphus loreus (Hedw.) Warnst. is similar to R. squarrosus. It has leaves
irregularly plicate and alar cells scarcely differentiated. Occurring in the East and also
in the West, it may occur as a disjunct in the Lake Superior region.

2. Hylocomium BSG

Fairly robust plants in loose, yellow or olive-green, dull or shiny mats or tufts.
Stems spreading or ascending, arcuate, irregularly pinnate to regularly 2- to 3-pinnate
and frondose from a stipitate base, sometimes layered because of repeated annual
growth from arched-ascending, stoloniform innovations; paraphyllia abundant, filiform,
branched. Stem and branch leaves differentiated: Stem leaves loosely erect or
erect-spreading, irregularly plicate, broadly oblong-ovate and rather abruptly narrowed
to a short or long, often + rugose acumen; margins plane or reflexed at base, serrate or
serrulate all around; costa double, short or sometimes reaching the leaf middle, or
single; cells linear, smooth or spiculose here and there at back because of projecting
ends, oblong, incrassate, porose, and yellowish at the insertion, not differentiated at

L

convex base; cilia of endostome 2-4, rarely 1-2, nodulose.—The name means woods-
inhabiting

Stem regularly 2-3-pinnate and frondose in a layered arrangement resulting from successive
years’ growth from a curved-ascending stoloniform innovation, costa double; a a

ot twisted at the apex H. splendens
— irregularly pinnate, not la eee or ee from a stoloniform shoot; eee single;
anch leaves mostly twisted at the 2. H. pyrenaicum

. Hylocomium splendens (Hedw.) BSG—Plants in dull, green or yellowish mats.
Stems stiff and wiry, regularly 2-3-pinnate in horizontal fronds in a step-wise

343

comium splendens. 962. Habit. 963. Stem leaves. 964. Branch leaves. 965. Cells from
upper part i: branch leaves.

arrangement, each frond produced by an arched and ascending, stoloniform innovation
arising from the middle of the previous year’s growth; stems and branches reddish.
Stem leaves loosely erect, 2-3 mm. long, lightly plicate, broadly oblong-ovate, abruptly
narrowed to short or long, often rugose acumen; margins recurved below, serrulate
above, + inflexed at the base of the acumen; costa double, less than 1/3 to nearly 1/2
the leaf length; upper cells linear-flexuose, spiculose-papillose here and there at back
because of projecting upper angles. Leaves of primary branches 1.2-1.5 mm. long,
oblong-ovate, acute or acuminate. Leaves of branchlets 0.4-0.8 mm. long, oblong-
lanceolate, sharply or bluntly acute, with margins incurved above, serrulate in the
upper half, often sinuate-serrulate nearly to the base. Setae 12-30 mm. long, orange-
brown or reddish, + flexuose; capsules 1.5-2.7 mm. long, brown, somewhat constricted
below the mouth when dry; operculum stoutly and obliquely rostrate, 1.6-2 mm. long.
Spores 13-18 pw, very finely papillose—n = 10, 10+2, 11, 12

Conard, How to Know the Mosses, (ed. 2) fig. 172. ange Mosses of Michigan, fig. 140.
Grout, Mosses with Hand-lens and Microscope, fig. i s H. proliferum), Moss Flora of North
America 3: Pl. 31 above. Jennings, Mosses of Western Pen scuteaia. (ed. 1) Pl. 43, (ed. 2) Pl.
47,—Fig. 962-965.

n humus, soil, and rocks in oe or wet tp haan end in deeply shaded ravines,
Ren eat also on rotten logs or bases trees. Circumpolar; orted from New Zealand; in
North America south to Oregon, Colorado, owe and Nor th Caro atin: na

344

CHEBOYGAN CO.—Reeses Bog, Hermits Bog, Gorge of Carp Creek, Iron Bridge, Grass Bay.
EMMET CO.—Pleasantview Swamp, Five Mile Creek, Wycamp Lake, Cecil Bay, Big Stone Creek,
Conway Bog. MACKINAC CO.—Mackinac Island, St. Martins Point, McKays Creek, Prentiss Bay,
Hog Island Point, Cut River, Epoufette, Ozark. PRESQUE ISLE CO. —Presque Isle Lighthouse,
Evergreen Beach, Hammond Bay, Mast Point, Clinton Lake.

Hylocomium splendens is attractive and, indeed, splendid. It is sometimes called
the MOUNTAIN FERN MOSS because of a resemblance to Thuidium delicatulum, the
Fern Moss. I like the name STAIR-STEP MOSS because the branching is neatly
2-3-pinnate in successive sprays corresponding to annual layers of growth. (Poorly
developed plants may be confused with Pleurozium schreberi, but the absence of
paraphyllia in Pleurozium is a decisive difference.)

2. Hylocomium pyrenaicum (Spruce) Lindb.—Plants rather robust, in light- or
yellow-green to yellow-brown, moderately shiny mats. Stems ascending, remotely and
irregularly pinnate, not stipitate or layered, without stoloniform innovations; branches
often cuspidate-tipped. Stem leaves loosely erect or erect-spreading when dry, erect-
spreading when moist, 2.2-2.5 mm. long, broadly oblong-ovate, gradually or abruptly
acuminate, narrowed to a non-decurrent base, irregularly plicate; margins + revolute
below (especially when dry), spinulose-serrate in the upper 1/3-1/2; costa single, ending
2/3-3/4 up the leaf, usually in a dorsal spine; cells oblong-linear, flexuose, with thin or
rather firm walls, smooth at back. Branch leaves somewhat differentiated, 1.5-2.5 mm.
long, broadly oblong-ovate and abruptly short-acuminate to oblong-lanceolate and more
gradually acuminate, with mostly twisted apices. Setae slender, flexuose, reddish,
10-22 mm. long; capsules red-brown, shortly ovoid-cylindric, 1-1.8 mm. long; annulus
none; operculum conic, bluntly apiculate, 1 mm. long; cilia in 2’s or occasionally
single. Spores 15-20 u, very finely papillose.—n = 9, 11+1, 12

Grout, Moss Flora of North America 3: Pl. 31B.—Fig. 968-969.

On logs, humus, or rocks in woods. Europe and Asia; ie and the Aleutians, British
Columbia and Colorado; Newfoundland to New York and Michig

CHEBOYGAN CO.—Reeses Bog, Iron Bridge (Carp Creek), Little Lake 16.

Only indistinctly pinnate and not forming layered innovations, the plants have
rather stout, tumid branches tapered to the ends and in the field resemble a
Brachy thecium.

Hylocomium umbratum (Hedw.) BSG (fig. 970-973) occurs not too far from the
Straits, at the Lower Falls of the Tahquamenon River (Chippewa County). The plants
are relatively slender. They branch in a bushy fashion from an inconspicuous stipe and
do not form flat fronds corresponding to the growth of successive years (as in H.
splendens). The stem leaves are decurrent, spinulose-serrate all around, and bicostate
1/2-2/3 their lengths. The leaf cells are smooth. (The chromosome number is recorded
as n= 6+1,7

Not yet found in Michigan though widely distributed from Ontario and Ohio
eastward and southward is Hylocomium brevirostre (Brid.) BSG (fig. 974-975). It has
abundant paraphyllia, allying it to Hylocomium, and yet its aspect is something like
that of Rhytidiadelphus loreus or R. squarrosus. The plants do not form flat fronds,
the costa is double, and the leaf cells are smooth. It differs from H. umbratum in
being larger and coarser and having leaves of stipes and innovations with squarrose tips,
and its stem leaves are pinched and rugose at the base of the acumen, serrate in the
upper half or more, serrulate below, and not decurrent.

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Hylocomium pyre . 968. Stem leaf. 969. Branch lea
Hylocomium Lene he Stem leaf. 971. Branch ae 972. Parpahyllia. 973. Cells at

margin of upper part of branch
Hylocomium brevirostre. a Stem leaf. 975. Branch leaf.

Rhytidium rugosum (Sull.) Kindb., which occurs in the Upper Peninsula, is an
attractive golden-brown moss, erect-ascending and often regularly pinnate, with leaves
falcate-secund and both plicate and rugose. The costa is single. It grows on rocky soil
in dry, exposed places and is most often associated with limestone.

BUXBAUMIACEAE

Small, annual plants, scattered or gregarious, growing from a + persistent
protonema. Stems very short, simple. Leaves few, very small, broadly ovate or

346

ovate-lanceolate, green at base, hyaline above, ciliate-margined, ecostate; cells laxly
oblong-hexagonal, the marginal cilia continuing growth to form a tomentum over the
stem and the base of the seta. Dioicous; male plants minute, consisting of a single
bract resembling a clam shell and enclosing a single antheridium on a cygneous stalk,
resting on the protonema; female plants producing terminally usually 1 archegonium
and no paraphyses, the perichaetial leaves not differentiated. Setae elongate, stout and
rigid, reddish, warty-papillose; capsules oblique to horizontal from a short, stout, erect
neck, ovate or narrowly oblong-ovate, narrow at the mouth, flat or concave on the
upper side, swollen on the lower, the upper and lower surfaces sometimes separated by
a distinct or indistinct rim; annulus none, dehiscence aided by a mass of enlarged,
ovoid or globose cells between the epidermis and the peristome; operculum bluntly
conic, falling with the top of the columella; stomata at junction of neck and seta,
superficial or immersed; exostome consisting of 1-4 concentric series of irregular and
sometimes rudimentary, smooth or papillose teeth, sometimes partially joined as a
membrane, increasing in size toward the endostome; endostome a pale, 16-plicate
membrane forming a truncate cone (open at the apex). Spores spherical. Calyptrae
shorter than the operculum, cucullate, conic or thimble-shaped, naked.—The family
consists of a single genus, Buxbaumia Hedw. Johann Christian Buxbaum, a German
physician-botanist, discovered the genus in 1712 near the mouth of the Volga River
and described it in 1728, without giving it a name. He wished to follow the example
of Marchant and make it into a new genus named for his father but called to mind the
fox who was derided because he begged grapes, not for himself, but for his sick
mother. Haller in 1744 named the genus for the modest Buxbaum. (Fabricius
re-christened the genus Hippopodium in reference to the horseshoe-shaped ridge around
the capsule of B. aphylla, and Palisot de Beauvois called it Saccophorus because of the
belly-like swelling at the base of the capsule.)

Buxbaumia aphylla Hedw.—Setae erect or nearly so, about 4-11 mm. long,
capsules strongly inclined to nearly horizontal from a stout, erect neck, chestnut-
brown, broadly ovoid, 3-5 mm. long, nearly as broad as long, flattened on the upper
side, angled between the upper and lower surfaces, the epidermis peeling back a short
distance from the mouth after dehiscence; operculum about 1 mm. long; stomata

ersed, consisting of 2 guard cells or a single rounded cell with a central pore;
exostome relatively well developed, consisting of a single, brown, smooth membrane
dissected at the apex into 16 short, unequal teeth; euiasions well developed,
pale-tawny, papillose. Spores smooth, 6.5-8 u.—n = 8; 2n =

Conard, How to Know the Mosses, (ed. 2) fig. 30a. Darlington, Mosses of Michigan, fig. 143.
Grout, Mosses with Hand-lens and Microscope, fig. 19 & Pl. 8 left, Moss Flora of North America |:
Pl. 73A (fig. a). Jennings, Mosses of Western Pennsylvania, (ed. 1) Pl. 26, (ed. 2) Pl. 30. Welch,
Mosses of Indiana, fig. 63.—Fig

neer of disturbed, sandy or clayey soil, sometimes on logs or stumps, in partial shade
in moist sores nd also in dry, open woods, frequently on banks of trails or roads; locally
common on sandy soil and old wood in aspen stands (successional to ae Europe and Asia;
Greenland; Yukon to Colorado; Michigan to Nova Scotia, south to Delaw

oo Co. sa the Gorge of Carp Creek, Iron Bridge, Hogback Road, Biological
Station campus, near Bryants Bog, Grapevine Point, Colonial Point, Mud Lake, Little Lake 16.
EMMET CO. “Peliston Hills, pet Bay, Wilderness State Park. MACKINAC CO.—Ozark. PRESQUE
ISLE CO.—Huron Beach, Evergreen Beach.

“This is as strange a thing as e’er I looked on,
As disproportionate in manners as in shape.”
It was Elizabeth G. Britton, who thus quoted Shakespeare to describe these
HUMP-BACKED ELVES (The Observer 7: 105-113. 1896), Dillenius, some 200 years

347

ago, referred to Buxbaumia aphylla as the queen of mosses, and Grout called it a
BUG-ON-A-STICK. The capsule (which might be described as elegantly tear-shaped) is
as large as that of a Polytrichum and quite oversized in proportion to the seta and the
extremely reduced gametophyte.

It is said that the great W. J. Hooker became interested in mosses as a result of
discovering a colony of this curious moss. Braithwaite said, “This strange plant has the
annoying habit of disappearing from the stations it occupies, probably due to some
change in the constituents of the substratum on which it is produced, and thus we can
never rely upon finding it a second time in the same locality.” The reason for its
ephemeral character has to do with a pioneer role in succession. Occupying barren,
disturbed habitats, it is quickly crowded out by larger, more vigorous competitors.
However, it has been produced in abundance year after year on dry, sandy soil and old
wood in the aspens near the entrance to the Biological Station, where succession has
been kept at a standstill by inhospitable edaphic conditions. (From my own memory, I
can say that it was there in 1958 and every year from 1965 through 1973.) It is easily
spotted by a blackish alga-like crust which may be partly protonema but almost
invariably includes a growth of a minute leafy liverwort (Cephaloziella). It grows in
burned-over areas not yet occupied by an understory of Pteridium, in open Cladonia
barrens where soils are virtually lacking in humus, except for occasional bits of wood.
The Cladonia barrens, occupied especially by reindeer lichens, mark places of excessive
burning. It was common practice to cut the pines once abundant in the area in such a
way that their crowns fell together for convenience in trimming, thus leaving great
slash piles which burned more intensely than surrounding areas.

Uggla (Acta Phytogeogr. Suec. 41: 94. 1958), studying the effects of fire in
northern Scandinavia, found Buxbaumia aphyila in 1947 on sites burned in 1933 and
fully exposed to the sun. He was unable to locate the species in 1949, 1953, or 1957
or find it on sites burned in 1920 or 1941 and concluded that sporophytes were
produced only at a certain stage in the succession following fire. (Uggla’s findings on
Buxbaumia are summarized by Jones, Trans. Brit. Bryol. Soc. 6(1): 56-68. 1970.)
Locally, in Michigan, the calendar of events has been quite different. According to
Kilburn (Papers Mich. Acad. Sci., Arts & Lett. 45: 77-81. 1960), the best quality of
pine and hemlock were removed from Station property between 1870 and 1880. Poor
quality conifers were cut between 1880 and 1900 and hardwoods from 1900 to 1920.
For 40 years after the original cutting was completed, in 1880, fires occurred here and
there almost constantly, with major burns in 1892, 1901, 1911, and 1923. An area
along the power strip north of Reeses Bog and east of Carp Creek, more or less the
area where Buxbaumia is now so abundant, was burned in 1880, 1890, 1896, 1899,
and 1907. It presumably escaped the fires of 1901, 1911 and 1923 which damaged
nearby areas closer to Douglas Lake (according to unpublished information given me
by Edward Voss). Thus the area has apparently been available to Buxbaumia at least
since 1923 (50 years) and probably since 1907 (66 years).

Buxbaumia is often associated with old logs or bits of wood humus. It has been
considered saprophytic in nutrition more because of morphological reduction than any
physiological evidence. Haberlandt (Jahrb. Wiss. Bot. 17: 359-598. 1866) considered
the extreme reduction of the gametophyte an evolutionary response to saprophytism.
He considered the rhizoids similar to the hyphae of fungi in form and probably also in
function and thought that the protonema could support the small and evanescent leafy
gametophyte but not the large sporophyte. Actually the sporophyte has abundant
chlorophyll until a late stage of maturity and looks to be capable of self-support. It
has been demonstrated in other mosses that the sporophyte is nutritionally inde-
pendent (Bold, Amer. Jour. Bot. 27: 318-322. 1940; Rastorfer, Phyton 19: 169-177.
1962; Wolf, Nature 181: 579-580. 1958).

It is interesting to note that Buxbaum, 250 years ago, considered B. aphylla a

348

moss, while noting its fungus-like appearance, although Haller, Dillenius, and other
early authors indeed considered it a fungus!

Capsules are held aloft in a torch-like manner when young but later become
tilted owing to the influence of light on the distribution of auxin. Evidence of such
phototropism was provided by a study in the open aspen stands at Douglas Lake,
where it was found that the pointed ends of capsules are generally directed toward the
greatest incidence of light, 45% to the southwest and 30% to the south (Taylor, Mich.
Bot. 11: 70-73. 1972). Diphyscium foliosum, so similar in capsule form, shows a
similar orientation toward the light source. Phototropism has been investigated in the
setae of Splachnum; see Walsh, Naturalist, no. 823: 137-139. 1947.

An interesting feature of Buxbaumia is its sexual dimorphism.

Buxbaumia minakatae Okam. (B. subcylindrica Grout)—fig. 977—should be
sought on rotten logs and stumps in lowland woods, in such places as Tetraphis and
Lophocolea grow. It has setae only 3-5 mm. long, capsules only slightly inclined,
lance-cylindric, not flattened dorsally or angled between the upper and lower surfaces.
It has been found near Galesburg in southern Michigan.

DIPHYSCIACEAE

Small perennials in dense sods, dull, dark-green, becoming brown or blackish with
age. Protonema persistent, bearing peltate appendages. Stems mostly very short, erect,
usually simple. Lower leaves small, crowded, crisped and sometimes inrolled when dry,
+ spreading when moist, lingulate or lingulate-subulate, rounded-obtuse to acute,
mostly entire; costa single, strong, ending below the apex or filling it; upper cells
2-3-stratose from the costa nearly to the margins, obscure, rounded-quadrate or
rounded-hexagonal, often oblate, incrassate, plane or mammillose on both surfaces;
basal cells unistratose, pale, smooth, rectangular or oblong-hexagonal. Upper and
perichaetial leaves longer, erect, scarious, ovate-lanceolate, lance-subulate, or linear,
generally fringed at the apex and awned by the excurrent costa. Setae terminal, very
short; capsules immersed, obliquely ovoid-conic, very narrow at the mouth, rounded at
the base and swollen on 1 side, thin-walled, not angled or differentiated in color
between the upper and lower surfaces; annulus present; operculum conic, acute;
peristome teeth 16, very short and irregular, + separate; endostome a conspicuous,
whitish, membranous, densely papillose, open cone with 16 keels projecting between
the teeth. Calyptrae scarcely covering the operculum, conic or thimble-shaped, smooth
and naked.—Represented in our area only by Diphyscium Mohr, named because the
capsule wall is fairly distinct from the spore sac, being connected with it by short
filaments, resulting in a double “bladder.” (There is no really good reason to separate
the Diphysciaceae from the Buxbaumiaceae, other than tradition.)

Diphyscium foliosum (Hedw.) Mohr—Small plants in rigid, dark-green or more
often brown to blackish, extensive tufts. Stems very short. Leaves strongly curled and
contorted when dry, erect-spreading when moist, 1.3-4 mm. long (becoming longer
toward the stem tips), broadly ligulate or lingulate, obtuse or rounded-obtuse and
usually + cucullate at the apex; margins erect, crenate above the base; costa vanishing
near the apex; upper cells 6.5-7.5 4 wide, irregularly rounded and often somewhat
oblate, incrassate, mammillose-bulging on both surfaces. Dioicous; perichaetial leaves up
to 8mm. long, with costa long-excurrent asa brown, spinulose awn and cells delicate
with notably thickened cross-walls. Capsules nearly sessile, greenish-yellow, becoming
brown, about 3mm. long; annulus well developed; operculum 1.1-2 mm. long; stomata
numerous, at base of urn; peristome teeth very short and irregular (usually difficult to

Buxbaumia aphylla. 976. Habit (sporophyte).
Buxbaumia minakatae. 977. Habit (sporophyte).

Habit.
s pellucida. 979. Habit of sterile plant. 980. Habit is eee plant. 981. Leaves.
982. ee leaf. 983. Cells at leaf apex. 984. Portion of peristo

discern, although the whitish endostome is ee Spores 9-10y, finely
papillose. Calyptrae conic, acute._n = 8, 9, 1 = 16.

Conard, How to Know the aoe (ed. 2) fig. 30b-d. Darlington, Mosses of Michigan, fig.
144, Grout, Mote: with Hand-lens and Microscope, fig. 20 (as Webera sessilis), Pl. 8 right.
Jennings, _Mosses of Western See nen (ed. 1) Pl. 25, (ed. 2) Pl. 29. Welch, Mosses of Indiana,
fig. 64.—Fig. 978.

In moist hardwood forests, on soil or humus of shaded banks in late stages of moss-lichen
succession, Pee es on rock walls of stream ees ma and Madeira; Japan; widespread in
ein North America; Mexico, Guatemala, and Jam

CHEBOYGAN CO.-—Near Bryants Bog, Colonial Point, Munro Township, Mullett Lake.
EMMET CO.-—Pellston Hills, Middle Village Road east of Good Hart, Cross Village. MACKINAC
CO.—Mackinac Island.

The broad, sloping upper surface of the capsules apparently serve a purpose in
spore dispersal. According to Goebel, pelting rain drops cause spores to be emitted in

i 35) 6)

little puffs to a distance of two inches. (Diphyscium foliosum has been called the
POWDER GUN MOSS.) As in Buxbaumia, the capsules are directed toward the light,
but the adaptive significance of this is lost on me.

William C. Steere has said that Diphyscium looks like scared rabbits in the grass.
It may be recognized when sterile by broadly ligulate, blunt leaves consisting of several
layers of cells, those at the upper and lower surfaces conspicuously bulging. When dry
the leaves are dark and crisped, looking like they were dipped in hot butter (a happy
comparison supplied, again, by Dr. Steere).

TETRAPHIDACEAE

Plants small and tufted or minute and scattered to gregarious. Stems erect,
mostly simple. Leaves erect-spreading in 3 ranks (sometimes obviously so), scarcely
altered on drying, ovate to oblong-lanceolate or lance-acuminate, acute, not bordered;
costa single, weak and sometimes absent or well-developed and ending well below the
apex to subpercurrent; upper cells irregularly rounded-hexagonal or oblong-rhomboidal,
smooth; lower cells elongate. Autoicous. Setae terminal, elongate; capsules cylindric or
ovoid-cylindric, erect and symmetric or somewhat curved, smooth; annulus none;
operculum conic or conic-apiculate; peristome single, the 4 teeth inserted below the
mouth, narrowly triangular (actually pyramidal, representing the entire tissue within
the operculum and consisting of whole cells elongated lengthwise of the teeth),
yellow-brown, smooth. Spores spherical. Calyptrae narrowly mitrate, plicate, naked.—A
curious feature of this family of two small genera is the presence of “‘frondiform
leaves” which appear on the protonema at the first development of the stem. They are
more or less ligulate to spatulate. In Tetradontium brownianum, they are often forked.
In that species they are abundant and persistent. In 7. pellucida, they disappear early
and are rarely seen, although they can be found easily on rotten wood where Tetraphis
abounds. (A more suitable name than “frondiform leaves” might be protonematal
flaps.)

Tetraphis Hedw.

Rather small plants in dense, dull, green or red-brown tufts. Stems erect-flexuose,
naked below, rather densely foliate above, radiculose only at base, simple or, if bearing
perigonia, sometimes forked; sterile stems frequently ending in a rosulate cluster of
stalked, discoid brood-bodies surrounded by rounded bracts of varying size. Leaves +
keeled and somewhat tristichous, somewhat decurrent, the lower leaves ovate and
acute, the upper and perichaetial leaves gradually longer and narrower, lanceolate and
gradually acuminate; margins entire, plane or broadly reflexed; costa ending well below
the apex in lower leaves, subpercurrent in upper and aera leaves; cells irregularly
rounded-hexagonal and thick-walled above, oblong near the insertion, or in upper
leaves oblong-linear in most of the base. Perigonia at ae ends of innovations from the
base of sterile plants or on separate, simple or forked male shoots freely mingled with
the others, with abundant paraphyses. Capsules erect and symmetric or slightly curved,
cylindric, red or brown at the mouth; operculum conic; exothecial cells elongate
below, small and thicker-walled in several suboral rows; stomata none; peristome teeth
narrowly long-triangular. Calyptrae whitish below, sometimes slightly roughened near
the apex._The name means bearing or producing four, referring to the four peristome
teeth. This name was used by Hedwig in 1801 and has legal priority over Georgia,
which Ehrhart dedicated to his patron George III of England, who was also Elector of
Hannover.

351

Tetraphis pellucida Hedw.—Plants about 8-15 mm. high. Leaves plane and entire
at the margins (or broadly reflexed, at least when attached to the stems), lower leaves
about 1-2 mm. long, the upper and perichaetial leaves up to 3 mm. long; upper cells
about 13-20yu. Setae 6-14mm. long, twisted when dry, erect or + flexuose, smooth;
capsules 2-3mm. long; operculum about 1mm. long; peristome teeth about
0.5-0.9 mm. long. Spores 10-13 wu, nearly smooth or finely roughened.—n = 7, 8; 2n =
14

Conard, How to Know the rs (ed. 2) fig. 51. Darlington, Mosses of Michigan, fig. 26.
Grout, Mosses with Hand-lens and oscope, fig. 13 (as Georgia), Moss Flora of North America
1: Pl. 3 right. Jennings, Mosses of western PennsyWania, (ed. 1) Pl. 26, (ed. 2) Pl. 30. Welch,
Mosses of Indiana, fig. 20.—Fig. 979-984.

Typically on very rotten stumps or logs, sometimes on wood humus, rarely on soil or rock.
Circumpolar; Labrador to Alaska, south to California, Arizona, Colorado, Arkansas, and South
a.

CHEBOYGAN CO.—Reeses Bog, Hermits ae Gorge of Carp Creek, Iron ee Colonial
Point, Gleasons Bog, Wolffs Bog, Livingston Bog, M ud Lake, Grass Bay, Weber Lake ET
Cc

Mackinac Island, St. Martins Point, McKays Creek, Prentiss Bay, Cut River, Epoufette, Ozark,
Hendricks, McNamara Pond. PRESQUE ISLE CO.—Lamprey Experiment Station, Evergreen Beach,
Clinton Lake, Loon Lake, Rainy Lake.

gemmae cups suggest those of Marchantia polymorpha. Dispersal by rain
drops oie on such splash-cup structures has been investigated by Brodie (Canad.
Jour. Bot. 29: 224-234. 1951; Proc. Indiana Acad. Sci. 66: 65-73. 1957). In related
investigations, Brodie and Gregory (Canad. Jour. Bot. 31: 402-410. 1953) showed that
spores and other reproductive bodies may be more readily blown from cupulate
structures (especially in some species of Cladonia) than from flat surfaces, owing to a
vortex of turbulence which sucks out spores and lifts them above the relatively still air
near the surface into the main airflow

Tetraphis geniculata Girg. ex Milde has setae abruptly bent midway (like a knee)
and rough above the bend. It occurs in the East and also in the West and should be
looked for as a disjunct in the Lake Superior region.

Also, in Michigan’s Upper Peninsula we have Tetradontium brownianum (Dicks.)
Schwaegr., a rare moss of dark recesses, such as crevices of cliffs or undersides of
siliceous or granitic ledges. The minute and bud-like plants are gregarious but not
caespitose. The protonematal flaps are forked (and they are numerous and persistent).
The costa is weak, ending well below the apex of the leaf or sometimes lacking. The
upper cells are oblong-rhomboidal, the capsules oblong-ovoid, and the operculum
obliquely apiculate.

POLYTRICHACEAE

Plants mostly robust and rigid, usually tufted (rarely scattered or gregarious and
growing from a persistent protonema). Stems erect, sometimes from a + decumbent
base, usually simple. Lower leaves small, appressed, remote and sometimes scale-like,
the upper larger and more crowded, generally differentiated into a pale, broad,
sheathing base and a narrow, + spreading limb which is often bistratose nearly to the
margins, with long, green lamellae on the upper side of the costa and often covering
most of the lamina as well (sometimes lamellose on the lower side of the leaf also);

352

margins mostly strongly toothed, sometimes bordered by elongate cells; costa single,
strong, ending near the apex to excurrent; upper cells isodiametric, smooth or rarely +
papillose (the cuticle often longitudinally striate); cells of leaf base mostly rectangular
to linear, narrower toward the margins. Mostly ees perigonia cupulate or discoid,
often conspicuously rosulate, often producing an innovating branch from the center,
with intermingled claviform and filiform eee perichaetial leaves not or some-
what differentiated. Setae elongate; capsules erect or nearly so to horizontal, cylindric
or 4-6-angled, + flattened, and prismatic, sometimes clearly differentiated at the neck;
annulus none or very narrow; operculum acute to rostrate from a convex-conic base;
peristome single, consisting of 32-64 (rarely 16) short, lingulate teeth attached at their
tips to a discoid expansion of the tip of the columella, consisting of whole cells which
are elongate and fiber-like and follow the contour of the teeth in several concurrent
series, mostly pale except for the midline. Spores spherical, smooth. Calyptrae
cucullate, sometimes roughened at the tip, rarely naked, usually densely pilose.

The peristome teeth of the Polytrichaceae consist of whole cells which become
greatly elongate and compressed. Thus, at maturity, the teeth consist of several layers
of fine threads extending from the apex of one tooth through the basal membrane and
up again to the apex of the adjacent tooth, those in the middle being even more
compressed and forming an apparent median line. The tips of the teeth are attached to
the expanded, tympanum-like top of the columella. When dry, the columella is
retracted, leaving space between the teeth for spores to be dispersed by a salt-shaker
mechanism.

Smith (Mem. N.Y. Bot. Gard. 21(3): 1-83. 1971) presented some refinements on
generic concepts in the Polytrichaceae (not followed in the present treatment),
together with a wealth of information on peristome structure and vascularization.

Leaves undulate, toothed at back of lamina and costa, bordered, doubly serrate; lamellae
restricted to the costa; capsules cylindric, not angled; calyptra naked or sparsely
short-bristled at the tip 1. Atrichum

Leaves not undulate, + toothed at back of costa, not bordered, entire or singly serrate;
lamellae covering the costa and most of the lamina as well; capsules cubic to prismatic, 4-
or sometimes 6-angled; calyptra completely covered by long hairs 2. Polytrichum

1. Atrichum P.-Beauv., nom. cons.

Plants medium-sized to fairly robust, in loose, dull, dark-green tufts, becoming
brown or red-brown with age. Leaves crisped and contorted when dry, erect or
erect-spreading when moist, usually undulate and toothed at back of lamina and costa,
lingulate to oblong-lanceolate, acute or sometimes rounded-obtuse, not sheathing at
base, nearly flat or somewhat concave or keeled toward the apex, bordered by elongate
cells and singly or doubly serrate; costa narrow, ending somewhat below the apex to
percurrent, covered on the upper side by rather few, long, usually somewhat wavy,
green lamellae with undifferentiated marginal cells; cells unistratose, green throughout,
smooth or rarely + papillose, hexagonal above, short-rectangular below. Perichaetial
leaves scarcely differentiated. Setae single or sometimes clustered; capsules cylindric
from a short, tapered neck, suberect or somewhat inclined, often somewhat curved,
smooth; annulus and stomata none; operculum long-rostrate from a convex or
hemispheric base; exothecial cells smooth; peristome teeth 32, pale with a yellow to
red-brown mid-line. Calyptrae smooth and naked or sometimes sparsely toothed or
bristly at the tip._The name refers to the absence of hairs on the calpytra, in marked
contrast to the usual condition in the family. A more interesting name, earlier but
taxonomically unacceptable, Catharinea, was provided by Ehrhart in honor of Russia’s
benevolent despot for reasons that Bridel guessed at in a snide way (an reverentia? an
gratitudine? an benevolentiae captatione?).

353

Plants pout 10-20 mm. high; leaves + concave toward the apex; lamellae + wavy, 5-14
obscuring %-% the leaf in the upper third; ee cells 11-19 yw, thick-walled,

eer isodiametric, only occasionally ia than lon . A. angustatum
one typically taller than on mm.; leaves + keeled ae the ; lamellae straight, 2-5
s high, obscuring less an % the a in the upper third: ees cells 18-30 uw wide,

hy tather thin-walled, ee than lon 2.A _ undulatum

1. Atrichum angustatum (Brid.) BSG—Plants medium-sized, 10-20 (rarely 45)
mm. high. Leaves undulate (especially when dry), toothed at back in oblique rows
corresponding to the undulations, 3.5-7, rarely 9mm. long, oblong-lanceolate or
narrowly oblong, acute, + concave above; margins with mostly double teeth; costa
subpercurrent, toothed at back above; lamellae 5-6 (rarely 9), wavy, 5-14 cells high,
obscuring %4-% the width of the upper 1/3 of the leaf; upper cells 9-19 u wide,
incrassate, irregularly isodiametric, sometimes + longer than wide, plane or nearly so on
the lower surface, bulging on the upper, smooth or + papillose (sometimes strongly so)
on the lower and sometimes also on the upper surface. Dioicous. Setae 10-30 mm.
long, occasionally in pairs; capsules 2.7-6.5 mm. long, erect or + inclined, slightly or
rarely strongly curved; operculum 2-3 mm. long; Bees teeth 32. Spores 9-14 yu.
Calyptrae + toothed to short-bristled at the apex.—n = 7,

Breen, Mosses ee Florida, Pl. 120. Conard, How to Know the Mosses, (ed. 2) fig. 43. Grout,
Mosses with Hand-lens and Microscope, fig. 14c (as Catharinea), Moss Flora of North America 1
1 includ ar. plurilamellatum, A. papillo ; A. macmillanii). Jennings, Mosses of

31 (as Atrichum angustatum, A. plurilamellatum, & A. papillosum). Welch, Mosses of
Indiana, Pl. 120.—Fig. 985-986.

On light, often sandy soil in dry, open woods and frequently in disturbed places, such as
poor lawns, roadbanks, bare mounds in woods. Europe and Asia; Newfoundland to Manitoba and
south to the Gulf of Mexico.

CHEBOYGAN CO.-—Gorge of Carp Creek, Colonial Point, Grapevine Point, Mud Lake, Little
Lake 16. EMMET CO.—Pellston, Pellston Hills, Maple River Township. MACKINAC CO.—Mackinac
Island, Gros Cap, Ozark.

trichum undulatum (Hedw.) P.-Beauv.—Relatively robust plants, 1.5-7 cm.
high. ate undulate (especially when dry), toothed at back in oblique rows
cients to he. undulations, 4.5-9.5mm. long, oblong-lanceolate or rarely
oblong-elliptic, acute, + keeled at the tip; margins bordered, with mostly double teeth,
costa subpercurrent, toothed at back above; lamellae 4-6, nearly straight, 2-5 cells high,
obscuring less than 1/4 the width of the upper 1/3 of the leaf; upper cells 18-30 u
wide, hexagonal and + transversely elongate, not or sometimes incrassate, usually +
thickened at the corners, smooth. Appearing dioicous but occasionally producing
antheridia the first year and archegonia the next. Setae 14-40 mm. long, occasionally in
2’s, very rarely in 3’s; capsules 3.5-6mm. long, suberect or somewhat inclined,
curved; operculum 1.5-3mm. long. Spores 12-16 yu. Calyptrae toothed and con
bristled at the tip.—n = 7, 14, 16, 20, 21, 22, 28, 42.

Conard, How now the Mosses, (ed. 2) fig. 44. Grout, Mosses with Hand-lens and

Microscope, fig. es ie Catharinea), Moss Flora of North America 1: Pl. 53B (figs. 1-12).

ennings, Mosses of Western Pennsylvania, (ed. 1) Pl. 26, 27 & (ed. 2) Pl. 30, a Ae Catharinea
undulata, var. alleghaniensis, & var. minus). Welch, Mosses of Indiana, fig. 21.—Fig. 9

On rich, humic soil in moist woods and shady ravines and also on clay or mud, especially
along streams. Europe, North Africa, Madeira and the Azores, Asia ea or, China, and Japan; Alaska
to California; Nova Scotia to Minnestoa, south to the Gulf of Mexi

354

Atrichum angustatum. 985. ah 986. Leaf.

Atrichum undulatum, 987

Polytrichum piliferum. re

Polytrichum juniperinum. ro cee 990. Cross-section of leaf. 991. Lamellae as seen in
section.

CHEBOYGAN CO.—Burt Lake, Mud Lake. EMMET CO. Seas Hills. MACKINAC CO,.—
Cut River, Ozark. PRESQUE ISLE CO.—Lamprey Experiment Statio

Atrichum undulatum has been called the WAVY CATHARINEA (or, irreverenily,
the UNDULATING CATHARINE). It grows in moist, hardwood forests on rich,
humified soil, whereas A. angustatum is more often found on poor, disturbed soils in
the open, on roadbanks, for example. As compared with A. angustatum, it is usually
robust and often quite attractive (at least in a moist, growing condition), with lamellae
few and low, covering only about 1/8-1/10 of the upper half of the leaves; in A.
angustatum, the lamellae are more numerous and higher and cover about 1/4-1/3 the
upper half of the leaves.

Atrichum undulatum is, in essence, a diploid (or triploid) expression of A.
angustatum. The sexual relationship usually expressed by such a haploid-diploid species
pair is obscured by the fact that in A. undulatum antheridia are produced one season

ED)

and archegonia me next (making it, more often than not, difficult to demonstrate a
monoicous pote

Our pee can be sorted (although not too successfully) as follows: Var.
undulatum, relatively small plants of rather dry, weedy habitats, usually appearing
dioicous but potentially autoicous, with lamellae 2-4 or rarely 6 cells high and capsules
strongly curved and inclined. Var. altecristatum Ren. & Card., medium-sized plants of
moist woodland habitats, usually appearing dioicous but potentially autoicous, with
lamellae 4-6 and sometimes 9 cells high and capsules slightly curved and inclined. Var.
oerstedianum (C. M.) Crum, large, dioicous plants (usually more than 3 cm. high), in
wet habitats, with lamellae 4-6 cells high and especially long, narrow, somewhat curved
and inclined capsules (often more than 5mm. long). The var. altecristatum, best
recognized by its well-developed lamellae approaching those of A. angustatum, appears
to have twice the chromosome number as A. undulatum var. undulatum.

Atrichum crispum (James) Sull. occurs in Michigan’s Upper Peninsula on wet
banks of ditches or woodland streams. It resembles Mnium in aspect and has smooth
leaves not or slightly toothed at back, few, low, and inconspicuous lamellae, large cells
(25-40 u wide), and short capsules (2-3 mm. long), tapered to the neck.

2. Polytrichum Hedw.

Coarse and robust, rigid plants in loose or dense, dark, green, brownish, or
bluish-green tufts. Leaves erect or erect-spreading when dry, erect- to wide-spreading
and + recurved when moist, long-lanceolate from an enlarged, sheathing base, usually +
awned, almost entirely covered on the upper surface from the shoulders to the apex by
crowded, green lamellae with mostly differentiated terminal cells; margins plane, erect,
or abruptly inflexed, entire or coarsely toothed above the shoulders, unbordered; costa
broader and thicker above the shoulders, often toothed at back toward the apex,
usually excurrent as a + elongate, toothed awn or hair-point; cells of the upper lamina
bistratose nearly to the margins, smooth and thick-walled, subquadrate to hexagonal,
those at the shoulders very incrassate and transversely oblong, the cells of the sheath
oblong or oblong-linear and yellow-brown, narrower and paler at the margins. Dioicous
(or very rarely autoicous); perichaetial leaves not much differentiated but sometimes
considerably elongate. Setae single; capsules suberect to horizontal, cubic or prismatic,
4-6-angled, gradually or abruptly narrowed to a short, thick apophysis; annulus none or
poorly differentiated in a single row; operculum rostrate from a flat or convex base;
stomata large and numerous, somewhat immersed, 1-2-celled, in the upper part of the
apophysis or in a constriction between urn and apophysis; peristome teeth 64.
Calyptrae densely pilose.—These are the HAIR CAP MOSSES; the generic name too refers
to many hairs felting the calyptra. The large capsules, frequently broken off and
apparently eaten by small animals (perhaps mice or rabbits), resemble grains of wheat.
The name BIRD WHEAT is sometimes used for various species of Polytrichum. Poly-
trichum is almost always an indicator of acid conditions.

1. Leaves ending ae a vhs nae -point 1. P. piliferum
1. Leaves not endin a hyaline h
2. Leaf margins Be ene eeeed? Hen the shoulders and covering the lame a
3. Plants of bogs, growing in compact tufts and matted with rhizoids well up the stems;
leaves not at all curved when dry, not conspicuously awne 2a. P. jun pee nes var. affine
3. ae gases in dry habitats, growing in loose tufts, only moderately radiculose at o 2
e of the eons leaves usually somewhat curved when dry, en ae in a shor
e ae Ge reddish a ae
Leaf margins toothed, ee or erect, not inflexed or covering the lamellae

356

. Terminal cells of lamellae clearly notched (as seen in section); capsules deeply
constricted between urn and apophysis; exothecial cells bulging, with a central thin spot
P. commune
4. Terminal cells not or very slightly notched; capsules not or only ee constricted
etween urn and apophysis; exothecial cells not bulging, without a central thin spot
5. Terminal cells of lamellae subquadrate or transversely oblong, flat-topped or sometimes
re 4. P. ohioense
5. Terminal cells rounded or narrowly elliptic, not much differentiated from other cells
of the lamellae
6. Terminal cells of lamellae narrowly elliptic; cells of leaf sheath oblong-linear; marg
of upper a of leaf very narrow (usually less than 4 cells wide), the cells seat
0-12 uw ormosum
6. Terminal ain rounded or shortly oval; cells of sheath shorter (3-5:1), oblong; margin
of upper part of leaf somewhat wider, the cells about 15-18 yu wide
3a. P. formosum var. aurantiacum

1. Polytrichum piliferum Hedw.—Relatively short, stout plants (1-4 cm. high) in
loose, brown or glaucous-green tufts. Stems comose above a naked and flexuose base.
Leaves erect when dry, erect-spreading when moist, 4.5-7 mm. long, lanceolate from a
broader base, stoutly needle-like above the shoulders because of broadly infolded,
subentire margins, ending in a long, hyaline, spinulose (or rarely smooth) hair point;
lamellae about 24-29, crenate-dentate in profile, 5-7 cells high, with marginal cells
conic or pyriform and mammillose in section. Setae 20-43 mm. long; capsules erect
when young, inclined when mature, 2-3.5 mm. long, cubic to short-prismatic, sharply
4-5-angled, strongly constricted between urn and apophysis; annulus a single row of
cells; stomata in the constriction between urn and capsule. Spores 8-13 u.—n = 6, 7; 2n

onard, How to Know the Mosses, (ed. 2) fig. 48g. arte: Mosses of Michigan, fig. 147.

Grout, ee with Hand-lens and Microscope, Pl. 17g-h, p, Moss Flora of North America 1: Pl.

as (figs. 1-13). Jennings, Mosses of Western Pennsylvania, (ed. 1) Pl. 28, (ed. 2) Pl. 32 & 64.
Welch, Mosses of Indiana, fig. 31. —Fig. 988.

On dry, often sandy or gravelly oe Piscine in exposed pe ee on thin soil over
rocks, bare roadbanks, sunny knolls or ail banks in forests, and often in old fields. Circumpolar;
southern South America a Australia; Clea: . Alaska, south California, Colorado, Iowa,
Ohio, and North Carolina

CHEBOYGAN CO.—Above the Gorge of Carp Creek, Pine ek Vincent Lake, Smiths Bog,
Mud Lake, Little Lake 16, Weber Lake. EMMET CO. —Stu nville, French Farm Lake.
MACKINAC CO.-—Bois Blanc Island, St. Martins sie Ozark, Hendricks. PRESQUE ISLE CO.—
Huron Beach, Rainy River Falls, Shoepac Sink Holes

Easily recognized by hyaline hair-points and inflexed leaf margins.

Hughes (New Phytol. 61: 266-273. 1962) has studied the effects on daylength on
the development of the sporophyte of this species. Marsh and Koerner (Ecology 53:
489-493. 1972) studied its role in stabilizing dune sands along the shores of Lake
Superior.

2. Polytrichum juniperinum Hedw.—Plants rather small to fairly robust, in loose,
green, bluish-green, or red-brown tufts 1-13 cm. high. Stems moderately radiculose at
or near the base. Leaves straight or + curved. and erect, erect-spreading, or rarely
wide-spreading when dry, spreading or wide-spreading when moist, 4-8 mm. long,
lanceolate and needle-like above the shoulders because of broadly infolded, entire
margins (the sheathing base 1-2.5 mm. long), acuminate, ending in a short to fairly
ong, red or brown, toothed awn; costa excurrent, usually toothed at back above;
lamellae 23-41, crenate in profile, 6-7 cells high, the marginal cells in section

cRyi

ovoid-conic, with thick, smooth, hyaline walls. Perichaetial leaves elongate, with long,
slender awns. Setae 20-60 mm. long; capsules suberect to horizontal, longer than broad,
2.5-5 mm. long, dark-brown and somewhat glaucous, sharply 4-angled, deeply con-
stricted to the short apophysis; annulus none; exothecial cells not or somewhat
bulging, with a rounded or elliptic, central thin spot; stomata with 2 guard cells, in the
constriction between urn and apophysis. Spores 8-9 u.—n = 6, 7, 14; 2n = 14.

Con How Know the Mosses, (ed. 2) fig. 48a-f. Grout, Mosses with Hand-lens and
Microscope fg, 170, o’, Moss Flora of North America 1: Pl. 64B (figs. 1-10). Jennings, Mosses of
Western ee, eer 1) Pl. 29, (ed. 2) Pl. 33. Welch, Mosses of Indiana, fig. 29.—
989-991.

On soil and rocks, usually in dry, exposed or partially shaded places, frequently on bare
roadbanks, in dry, open woods, or on sunny knolls in moist woods, dry heaths, pastures, etc.
Widespread in both Northern and Southern Hemispheres and wee deeiuted in North America.

CHEBOYGAN CO.—Reeses Bog, Gorge of Carp Creek, Be Bridge, Grapevine Point, Bryants
Bog, Vincent Lake, Long Lake, Mud Lake, Weber Lake. EMMET CO.—Maple River Township,
Pellston Hills, Carp Lake, Cross Meee French Farm ore MACKINAC CO.-Bois Blanc Island,
Mackinac Island, St. Martins Point, McKays Creek, Cut River, Ozark, meat Corner, PRESQUE
ISLE CO.—Lamprey Experiment Station, Evergreen Beach, Shoepac Sink Hole

Commonly known as JUNIPER MOSS. The leaves resemble the scales of
Juniperus, in shape and in bluish-green color. The inflexed leaf margins covering the
green lamellae protect the photosynthetic tissue from desiccation. This is our most
common member of the genus, related to P. piliferum and like it characteristic of xeric
habitats; it is distinguished by the red awns rather than hyaline hair points.

P. juniperinum and P. piliferum are common on sterile soil in areas altered
by fire. In northern Finland and the Swedish Lapland, Uggla (Acta Phytogeogr. Suec.
41: 1-116. 1958) found that they reach their maximum development some 10-20 years
after a fire and survive burning well because of rhizoids penetrating the soil.

In Nova Scotia, Nichols (Trans. Conn. Acad. Arts & Sci. 22: 249-467. 1918)
noted that the pioneer trees of pastures are generally white spruce which germinate
most prolifically in carpets of Polytrichum. In our area, P. juniperinum would be the
most likely Polytrichum invading upland pastures. However, Gates (Bot. Gaz. 90:
233-259. 1930) reported that moss-Cladonia mats in the aspen stands at Douglas Lake
are too dense for seed penetration, absorb water too quickly to allow the soaking of
seeds, and because of frequent wetting and drying heave seedlings from the soil. (The
role of mossy logs as a seed bed was mentioned, under Hypnum imponens, above )

2a. Var. affine (Funck) Brid.—Plants relatively slender but tall and rigid, in very
compact tufts 4-16 cm. high, usually dark-brown, nearly always matted with rhizoids
well up the stems. Leaves erect and straight when dry, erect-spreading when moist,
3.5-6 mm. long, not conspicuously awned. Setae 18-60 mm. long; capsules 2.5-3 mm.
long.—n = 7; 2n =
ut, Moss Flora of North America 1: Pl. 64B (figs. 12-15), as var. alpestre. Jennings,
s of Western Pennsylvania, (ed. 1) Pl. 29 & (ed. 2) Pl. 33 (as P. alpestre). Welch, Mosses of
Indiana, fig. 30 (as var. alpestre).

In vas and damp heaths, hie in Sphagnum hummocks, sometimes forming extensive turfy
mats in ned-over peatlands. Circumpolar; South .America; Greenland to Alaska, south to
Wee Gas Illinois, Ohio, ne North Carolina.

HEBOYGAN CO.—Reeses Bog, Iron Bridge, Bryants Bog, Gleasons Bog, Gates Bog, Mud
Lake, Gn Bay, wate re 16, Weber Lake. EMMET CO.—Maple River Township, Stutsmanville,
Galloway Bog Lake, Cecil Bay, Conway Bog. MACKINAC CO.—Epoufette, Ozark,
Hendricks. PRESQUE ISLE CO.—Hammond Bay, Evergreen Beach, Loon Lake, Clinton La ke.

358

The var. affine can be recognized by long stems compacted together by a woolly
growth of rhizoids and also by its occurrence in boggy habitats where it often serves as
an indicator of fire. It may also indicate dry, late stages of bog succession. The tops of
old Sphagnum hummocks often support a good growth of Polytrichum and species of
Cladonia. The Polytrichum may represent a peak development rather than a recent
invasion, however, as Sphagnum hummocks are often made possible in the first place
by support offered by Polytrichum.

3. Polytrichum formosum Hedw.—Moderately robust, dark-green or brownish
plants in loose tufts about 4-7 cm. high. Leaves loosely erect or erect-spreading when
dry, flexuose-spreading to wide-spreading and recurved when moist, 6.5-10 mm. long,
lance-acuminate from a sheathing base 1.5-2.5 mm. long, ending in a brown or
red-brown, toothed point; margins erect, sharply serrate nearly to the sheath, the
unistratose margin about 2/3 up the leaf 4-7, sometimes 10 cells wide; lamellae
covering most of the leaf width above the shoulders, 22-58, entire in profile, 3-7 cells
high, the marginal cells in section narrowly conic to elliptic, with somewhat thickened,
hyaline or pale-brown walls, smooth; costa rather shortly excurrent, toothed at back
near the tip; median cells of sheath long and narrow, oblong-linear; perichaetial leaves
not much differentiated. Setae 35-50mm. long; capsules 4-5 mm. long, 4-angled,
weakly constricted between urn and apophysis, pale- or yellow-brown, suberect to
horizontal; annulus none; exothecial cells not bulging, without a central thin spot;
stomata |- or 2-celled. Spores 11-13 u.—n = 7, 14; 2n = 14, 17

Conard, How to Know the Mosses, (ed. 2) fig. 49e-f. Grout, Moss Flora of North America
1: Pl. 61A. Jennings, Mosses of Western Pennsylvania, (ed. 1) Pl. 72 (as P. gracile).—Fig. 992-994,

On soil or rocks in damp, hardwood or coniferous woods, often in boggy places. Widespread
in both Northern and Southern Hemispheres; Alaska to Newfoundland and south to Oregon and
Colorado in the West and the Gulf of Mexico in the East.

CHEBOYGAN CO.—3 miles north of Cheboygan. EMMET CO.-—Pellston Hardwoods.
MACKINAC CO.—Hendricks, Little Dollar Lake.

3a. Var. aurantiacum (Brid.) C. J. Hartm.—Tufts 2-5, rarely 15 cm. high. Leaves
usually somewhat curved when dry, rarely strongly curved-contorted, 5-8 mm. long,
lance-acuminate from a broader, oblong base; margins incurved when dry, the
unistratose margin about 2/3 up the leaf 5-8, rarely 20 cells wide; lamellae 18-50, 3-6
cells high, subentire to somewhat crenulate in profile, the marginal cells not enlarged,
rounded to elliptic as seen in section, with scarcely thickened walls; median cells of
sheath rectangular, about 3-5:1. Basal membrane of peristome only slightly projecting
above the mouth of the capsule. Spores 17-22 u.—n = 7,

Conard, How to Know the Mosses, (ed. 2) fig. 49a-d. Grout, Mosses with Hand-lens and
Microscope, fig. 16b (as P. gracile), Moss Flora of North America 1: Pl. 61B (as P. gracile).—Fig.
995-996.

On soil and humus in hardwood or coniferous woods. Circumpolar; Labrador to Alaska and
south to Colorado, Michigan, and North Carolina.

CHEBOYGAN CO.-—Grapevine Point (Douglas Lake), Little Lake 16.

The variety is distinguished ae the typical form by broader leaf margins
(noticeably inflexed when dry) and short, rectangular cells of the leaf sheath. In
pical P. formosum the free ees of the lamina is not continuous to the leaf base,
whereas in the variety it is broadened at the shoulders and obviously continuous to the
base.

359

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Polytrichum formosum. 992. Leaf. 993. Nae as seen in section. is Cells of leaf base.
Polytrichum formosum var. aurantiacum, 995. Leaf. ee Cells or i bas
Polytrichum ohioense. 997. Lamellae as seen a section. 998. C
trichum commune. 999. Habit. 1000. Leaf. 1001. paeilae as seen in section. 1002.
Capsule. 003, Calyptra. 1004. Portion of peristome.

4. Polytrichum ohioense Ren. & Card.—Moderately robust plants in loose,
dark-green to brown tufts 1.5-6 cm. high. Leaves erect or erect-spreading when dry,
wide-spreading and recurved when moist, 6-10.5 mm. long, long-lanceolate from a
sheathing base about 2 mm. long, acuminate, ending in a short, reddish, toothed awn;
margins erect and coarsely toothed nearly to the shoulders; lamellae 30-41, covering
almost the width of the leaf above the shoulders, entire or nearly so in profile, 3-7
cells high, the marginal cells in section somewhat broader, flat-topped or somewhat
retuse, with thick, brown, smooth walls; costa rather shortly excurrent, sparsely
toothed at back near the apex. Dioicous or rarely autoicous; perichaetial leaves not

360

much differentiated. Setae 15-85 mm. long; capsules 2-5 mm. long, pale-brown, sub-
erect to strongly inclined, not or only slightly constricted between urn and apophysis,
4- or rarely 5-angled; annulus none; exothecial cells not bulging, without a central thin
spot; stomata numerous (but difficult to observe), with 2 guard cells or consisting of a
central slit in a single guard cell. Spores 11-13 u.—n = 14.

nard, How to Know the Mosses, (ed. 2) fig. 50e-g. Grout, Mosses with Hand-lens and
ia. fig. 16a (as P. smithiae), 17e, Moss Flora of North America 1: Pl. 62A. Jennings,
Mosses of Western Pennsylvania, (ed. 1) Pl. 28, (ed. 2) Pl. 32. Welch, Mosses of Indiana, fig.
26.—Fig. 997-998.

On soil or humus in woods. Circumpolar; widespread in eastern North America; New Mexico.

CHEBOYGAN CO.—Carp i Colonial Point, Mud Lake. EMMET CO.-—Pellston Hills,
Carp Lake. MACKINAC CO.—Cut Riv

Polytrichum ohioense resembles P. formosum in the appearance of gametophytes
and sporophytes, but the flattened marginal cells of the lamellae, as viewed in section,
are distinctive. They may be somewhat retuse, but those of P. commune are broader
and more conspicuously notched. The wet ‘habitat of P. commune is very different
from the mesic woodlands favored by P. ohioense. (Polytrichum ohioense can often be
spotted in the field by yellowish setae.)

5. Polytrichum commune Hedw.—Robust, dark-green or brownish plants in loose
or fairly dense tufts 4-45 cm. high. Stems usually erect from a flexuose and decumbent
base. Leaves erect or somewhat spreading at least at the tips and + rolled when dry,
spreading to recurved when moist, 6-10 mm. long, lance-acuminate from a sheathing
base about 2-3 mm. long (the yellow-brown base often shiny and conspicuous —
dry); margins plane or erect, sharply serrate nearly to the sheath; costa excurrent,
toothed at back; lamellae 21-55, covering nearly all the leaf above the shoulders, ne
cells high, entire or slightly crenulate in profile, the marginal cells in section somewhat
to considerably wider, notched or depressed in the middle (and occasionally divided
into 2 cells), with thick, brown, smooth walls. Perichaetial leaves long and narrow
(about 9-15 mm. long), scarious-margined, with costa long-excurrent. Setae 50-90 mm.
long; capsules 3-5 mm. long, red-brown and + glaucous, 4-angled, deeply constricted
above a distinct apophysis; annulus none; stomata with 2 guard cells, in the
constriction between urn and apophysis; exothecial cells bulging, with a circular to
slit-like thin spot. Spores 8 u.—n = 6, 7, 14

en, Mosses of Florida, Pl. 124. Conard, How to Know the Mosses, (ed. 2) fig. 50a-d.
Grout, ee with Hand-lens and Microscope, fig. ‘Wad, f f. Moss Flora of North America 1: Pl. 63.
Jennings, Mosses of Western Pennsylvania, (ed. 1) Pl. 29, (ed. 2) Pl. 33. Welch, Mosses of Indiana,
fig. 27, 28 (as var. perigoniale).—Fig. 999-1004.

On soil in wet, boggy woods. Nearly cosmopolitan; Greenland to Alaska and south to
California, eee Wyoming, Texas and Florida.

CHEBOYGAN CO.—Reeses Bog, Douglas Lake, ea Bog, Bryants Bog, Nichols Bog,
Vincent Lake, Mud Lake, Mullett Lake, Weber Lake. EMMET CO.—Pellston, Stutsmanville, Wycam
Lake. Wycamp Creek, French hia Lake, Big Stone MACKINAC CO. —Epoufette, Cranberry
Lake, Hendricks, McNamara Pon

The COMMON HAIR CAP MOSS is especially characteristic of wet habitats, at
the edge of coniferous swamps or in the moat surrounding bogs, for example. It is not
particularly common in our area.

361

Linnaeus wrote that this large and conspicuous moss was used in Lapland for
stuffing pillows and other bedding and that he had often slept on a bed of
Polytrichum during his travels in the North. (Withering reported that bears use it too!)
In Great Britain it was occasionally used in former times in making small brooms for
dusting curtains, carpets, and beds. Dillenius said that an oil expressed from it was used
by ladies of his time on their hair—a truly fine application of the Doctrine of
Signatures!

The leaf bases are usually exposed and shiny, giving a good clue to the identity
of the plants, but I depend entirely on the appearance of the terminal cells of the
lamellae in cross-sections of leaves. They are broad and notched in P. commune. In P.
ohioense, they are not much broader than the other cells of the lamellae and flat (or
occasionally slightly retuse). P. ohioense, of course, grows in moist hardwood forests,
not in boggy places.

Pogonatum is very similar to Polytrichum, especially in gametophytic structure
and appearance. The calyptra is hairy, as in Polytrichum, but the capsules are cylindric.
In northern Michigan we have Pogonatum alpinum (Hedw.) Rohl. (at the Lower Falls

exothecial cells and no stomata.

Similar in many ways to P. alpinum is P. urnigerum (Hedw.) Par. (a montane
species likely to occur in the Lake Superior region). When fresh, it has an attractive
bluish color quite different from the dark green of P. alpinum. It is further
distinguished by smaller size and growth in looser tufts. The leaves are flatter and
broader; the marginal cells of the lamellae are shorter (with the lumen resembling in
section the gabled end of a house); and the capsules are erect and narrowly cylindric,
with bulging exothecial cells and no stomata. The capsules of P. alpinum are nothing
like those of Pogonatum urnigerum or any other Pogonatum in a strict sense, see
Smith (Mem. N.Y. Bot. Gard. 21(3): 1-83. 1972) for a consideration of relationships.

We do not have Pogonatum pensilvanicum (Hedw.) Par., which grows as scattered
individuals from an abundant, persistent protenema on roadbanks farther to the east.

Tansley and Chick (Ann. Bot. 15: 1-33. 1901) demonstrated some startling
resemblances to the conductive tissues of Polytrichum and the so-called vascular plants.
The stems of Polytrichum have a central core of elongate cells (hydroids) similar in
shape to tracheids and an enveloping sheath of cells suggestive of sieve tubes (leptoids)
and companion cells. The vascular cylinder is enclosed by endodermis with Casparian
strips, and leaf traces join the midrib of the leaves to the conductive elements of the
stem. The radial arrangement of “xylem” and “phloem” in the decumbent base and
the concentric arrangement in the erect, leafy portion of the stem are similar to those
in the primary steles of roots and stems, respectively.

The seta has a central cylinder of less obviously differentiated tissues, owing
perhaps to overall simplication of the sporophyte. Spiral and annular thickenings have
been observed in comparable tissues in the columella of a hornwort, Dendroceros, by
Proskauer (Phytomorph. 10: 1-19. 1960). Finocchio (Bull. Torrey Bot. Club 94: 18-20.
1967) reported vessel-like cells in the gametophytic “cortex” of some pleurocarpous
mosses having only a small central strand or none. These cells, living at maturity, are

362

moderately elongate, with broad, oblique end walls and large, simple pits on the sides
and ends, sometimes in a scalariform pattern. Arrangement in short vertical series
suggests a conductive function. Such cells may be primitive or vestigial vascular
elements—or phylogenetically quite unrelated to vascular tissues!

In mosses, including Polyirichum, the movement of water is mainly external,
although some is internal and presumably by way of the xylem-like cells. [See the
discussion and references following Sphagnum recurvum above. Some further informa-
tion on water relations is provided by Anderson and Bordeau (Ecol. 36: 206-212.
1955); Buch (Soc. Sci. Fenn. Comm. Biol. 9(16): 1-44. 1945, 9(20): 1-60. 1947);
Klepper (Bryol. 66: 41-54. 1963; Nova Hedwigia 15: 13-20. 1968); Roberts and Haring
(Ann. Bryol. 10: 97-114. 1938.)] Callus plugs develop in the sieve tubes of
Polytrichum, and Eschrich and Steiner (Planta 74: 330-349. 1967) have demonstrated
that carbon-labeled assimilates indeed move in this “phloem.”

The stems of Polytrichum are hard and woody, befitting their size, but the
vascular tissue is not lignified. Lignin provides at the cellular level the strength and
stability mecessary for nearly all plants that successfully cope with a dry land
environment. It is a constant feature of vascular plants, large or small, from the fossil
Psilophytales upward. There is no obvious need for lignin in the low-growing
bryophytes. Among North American examples, lignin is absent from all structures of
the liverworts Marchantia, Riccia, and Jungermannia and from gametophytes of the
mosses Bryum, Rhodobryum, Mnium, Funaria, and Polytrichum, although present in
some peristomes (Siegel, The Plant Cell Wall, 1962; Amer. Jour. Bot. 56: 175-179.
1969). Lignin has been found in Sphagnum, of a different type from that in higher
plants (Black et al., Jour. Applied Chem. 5: 484-492. 1955; Bland et al., Phytochem.
7: 1373-1377. 1968). It occurs in two of the giant mosses of New Zealand, Dawsonia
and Dendroligotrichum, both somewhat related to Polytrichum (Siegel, loc. cit.), and
Scheirer (Bryol. 75: 305-314. 1972) has demonstrated that in Dendroligotrichum it is
indeed associated with vascular tissue, though not limited to the “xylem.

Lignin defies chemical definition, and therefore one can scarcely draw con-
clusions concerning evolutionary origins from the foregoing. However, it is worthy of
comment that both the Polytrichaceae and the Dawsoniaceae give morphological
evidence of a primitive nature and a relationship to the Psilophytales, and lignin
supplies a chemical link. The Dawsoniaceae, centered in the Antipodes, are phyto-
geographically associated with many other primitive organisms, plant and animal.

eferences on the structure and significance of conductive tissue in mosses,
especially the Polytrichaceae, include Berthier et al. (Compte Rendu Séanc. Acad. Sci.
Paris 272: 2868-2871. 1971), Eschrich & Steiner (Planta 82: 33-49. 1968), Hébant
(Natur. Monspel., Bot. 18: 293-298. 1967; Compte Rendu Séanc. Acad. Sci. Paris 226:
2190-2192. 1968; Rev. Bryol. et Lichén. 36: 721-731. 1970; Phytomorph. 20:
390-410. 1971), Kawai (Ann. Rept. Bot. Gard. Univ. Kanazawa 2: 1-25. 1969, 16:
21-60, 83-111. 1971), Kawai & Ikeda (Sci. Rept. Kanazawa Univ. 15: 71-98. 1970).
Lorch (in Linsbauer, Handbuch der Pflanzenanatomie 11, 2, 7(1): 1-358. 1931), Smith
(Mem. N.Y. Bot. Gard. 21(3): 1-83. 1971).

Vascular tissue is well developed in large, erect mosses, especially in the
Polytrichaceae and Dawsoniaceae, but progressively less evident in smaller acrocarps
and low-growing, matted pleurocarps. Because they have no provision for lignified
support or secondary growth and have no specialized food storage or other provision
for dormancy, mosses have remained small and adapted themselves to small niches not
suited to plants more liberally endowed in size and structure and have thus escaped the
evolutionary pressures acting on larger terrestrial plants. Reduction in size and
simplification of structure (often associated with a telescoping of the life cycle) has
been a major force in the evolution of bryophytes. The uncertainties of fertilization by

363

swimming sperms have been largely avoided by a trend toward monoecism and,
especially, asexual forms of propagation. [The phylogenetic meaning of Buxbaumia’s
highly simplified gametophyte is discussed by Goebel (Flora 76: 92-104. 1892; Ann.
Bot. 6: 355-360.-1892) and van der Wijk (Buxbaumia 1: 2-4. 1947).]

The mosses are not direct offshoots from the aquatic green algae but reduced
from primitive vascular plants already adapted to life on land. Swimming sperms, a
well-defined alternation of generations, and dependence of sporophyte on gametophyte
fit as well in a downgrade sequence from vascular plants as in an upgrade series from
algae. Other evidence of a terrestrial ancestry includes cytokinesis by cell plate
formation (rather than centripetal pinching) and spores developed in tetrads as a direct
result of meiosis, as well as presence of a cuticle, stomata in the capsule wall, and
peristome mechanisms for spore dispersal. Few mosses are truly aquatic and even those
that are, such as Fontinalis, are related to terrestrial groups and have structures like
peristomes and stomata obviously suited to aerial functions. Also the spores of mosses
have a size suited to wind transport and no obvious adaptations for aquatic dispersal.

It is convenient to treat bryophytes as a single division because of obvious
similarities in life cycle. Yet they are polyphyletic in origin. I recognize five lines of
evolution as classes: Hepaticopsida, Anthocerotopsida, Sphagnopsida, Andreaeopsida,
and Bryopsida. The liverwort-like Anthocerotopsida may be derived from more highly
organized vascular plants, like the mosses. Hepatics may have come more directly from
green algae; yet the oldest fossil liverworts show some evidence of vascularization not
exhibited by any of the algae.

The evolution of bryophytes is difficult to follow because of independent trends
of adaptative change in gametophyte and sporophyte and also because morphologists
have been loath to recognize the homology of the two generations—hence favoring
such terms as dioicous rather than dioecious, phyllid rather than leaf, and caulid rather
than stem. However, the genetic potential is there, whether the chromosome number is
haploid or diploid. Sporophytic tissue can be made to regenerate as leafy diploid
gametophytes. Gametophytes can be made to bypass fertilization and form sporangia
by an abnormal budding process. Such buds can be directed toward sporangial
development or gametophytic expression depending on conditions of culture. Thus the
chromosome level does not control the expression of characters and the two genera-
tions, gametophyte and sporophyte, must indeed be homologous. Because the sporo-
phyte normally remains attached to the gametophyte, it is subject to different
environmental pressures, expresses itself differently, and evolves independently within
the framework of identical genes. It is the conditions of growth that have kept the
sporophyte from expressing its potential as a leafy, sexual plant. It follows for similar
reasons that vascular tissue in the gametophyte and the sporophyte are homologous
and also homologous to xylem and phloem in the higher plants.

I see the common ancestor of mosses and Psilophytales (and all other vascular
plants) as terrestrial but low growing and suited for fertilization under conditions of
intermittent wetness. The sporophyte was attached to a small, simple gametophyte,
much as in Anthoceros; it was unbranched and leafless, ended in a sporangium, and
had the essential elements of vascularization. From that point, mosses arose as the
gametophyte made do with microhabitats and developed into a leafy form, first erect
and simple, later prostrate and branched; higher plants specialized in an elaboration of
the sporophyte, beginning with dichotomous branching, later developing leaves, sporo-
phylls, heterospory, seeds, roots, secondary growth, etc. Cooksonia, of the Upper
Silurian, is the earliest known vascular plant and close to the hypothetical ancestor of
mosses, but it has a forked sporophyte. The early development of mosses has not yet
been discovered in the fossil record. Cooksonia is older than the oldest “probable”
fossil moss, Muscites plumosus of the Lower Carboniferous (Thomas, Ann. Bot. 36:

364

155-161. 1972). The undoubted mosses of the Permian, much later, are highly evolved
and far removed from the level of gametophytic complexity presented by the
Polytrichaceae, which in terms of extant mosses seem primitive.

Reviews of bryophyte fossils are given by Jovet-Ast (Traité de Paléobotanique,
vol. 2, pp. 17-186. 1967) and Lacey (Biol. Rev. 44: 189-205. 1969).

ADDENDUM

Sphagnum quinquefarium (Lindb. ex Braithw.) Warnst. has been found once in
Michigan (in a Thuja swamp at Epoufette, Mackinac Co., H. Crum & N. G, Miller, July 28,
1966). It is light-green to pink and resembles S. warnstorfii when dry because the narrow
branch leaves are arranged in 5 spiral rows and spreading at the tips. However, the pores
on the outer surface of the branch leaves are larger and elliptic. The stem leaves are
rather small and oblong-triangular, ending in a short, acute, concave tip; the hyaline
cells are often 1-2-divided, not or somewhat fibrillose, and the border is conspicuously
differentiated at the base. Pores are often found in the stem cortex, but the small,
pointed stem leaves with commonly divided hyaline cells and the dry aspect of the
branch leaves separate this species from S. russowii.

A DICTIONARY OF TERMS RELATING TO MOSSES

“When I use a word,” Humpty moe said in rather a scornful tone, “it means just what I
choose it to mean—neither more nor less
Lewis Carroll, Through the Looking Glass

William C. Steere used to emphasize the importance of terminology by reading to
his ee in systematic botany this account of how horses are harnessed in Germany:
man stands up the horses on each side of the thing that projects from the
front ond of the wagon, throws the gear on top of the horses, and passes the thing
that goes forward through a ring, and hauls it aft, and passes the other thing through
the other ring and hauls it aft on the other side of the other horse, opposite to the
first one, after crossing them and bringing the loose end back and then buckles the
other thing underneath the horse, and takes another thing and wraps it around the
thing I spoke of before, and puts another thing over each horse’s head, and puts the
iron thing in his mouth, and brings the ends of these things aft over his back, after
buckling another one around under his neck, and hitching another thing on a thing
that goes over his shoulders, and then takes the slack of the thing which I mentioned a
while ago and fetches it aft and makes it fast to the thing that pulls the wagon, and
hands the other things up to the driver.
Mark Twain, A Tramp Abroad

ABAXIAL—away from the stem or axis, opposed to adaxial; (see also dorsal).
ACAULESCENT —stemless or apparent tly so.

ACIDOPHILE-—a plant of acid ee an oxylophile.

ACINACIFORM —scimitar-sha

ACROCARPOUS —producing re sporophyte at the end of a stem or main branch (opposed to
pleurocarpous). oe as mosses generally grow erect in tufts (rather than mats) and are
sparsely branc

ACROGENOUS— a an ois meristem

ACUMEN (pl. acumina)—a slender, tapering point; ACUMINATE -slenderly tapered.

ACUTE -sharply ee (less than 90°
ADAXIAL—toward stem or axis, oppo osed to abaxial; (see also ventral).
ADHERENT-sticking to another organ, as opposed to coherent, or sticking to an organ of the
nd.

ADNATE-—fused with or attached to another org.

AERUGINOSE~—bluish-green, the color of scene ae (verdigris).

AGGREGATE —clustere

ALAR CELLS~—cells at the basal angles of a leaf, often Sa ceata in pee size, or color.

ALVEOLATE—with depressions on ss surface, sometimes applied to

AMORPHOUS -without definite sha

ogee oa eee oe of en ee representing a third row, often reduced or even

ing; in a few mosses, smaller iia the upper surface of stems and branches, variously

peabareinen he the lateral leav

AMPHITHECIUM—an embryonic tissue oy a developing capsule, surrounding the central endo-
thecium.

AMPLEXICAUL-clasping the s

ANASTOMOSING— Seer ee sometimes applied to the joining and rejoining in irregularly
divided peristome teeth.

ANDROECIUM-the male inflorescence.

ANDROGYNOUS-—male and female sex organs in the same inflorescence.

ANISOSPOROUS —producing spores of two overlapping size classes in the same capsule (sometimes
associated with asexual dimorphism); differentiated from the heterosporous condition of

365

higher plants in which the microspores and megaspores are of two distinct sizes and are
produced in different structures.

ANNOTINOUS-~—in yearly growths (the junction with the vee year’s growth evident because of
crowded leaves or other signs of interrupted grow

ANNULUS~—a ring of differentiated cells between the mouth of the capsule and the operculum,
aiding in dehiscence; ANNULAR -ring-like.

sear nnn the male reproductive organ, a globose to broadly cylindric, stalked structure

a pa (antherozoids).

ANTHROPOCH se, ANTHROPOGENOUS ~associated with man or aided in dispersal by the
acti ae

ANTRORSE_ nee a earad or (in reference to papillae) toward the leaf tip.

APICULUS~—a ss abrupt point; APICULATE—abruptly short-pointed (mucronate is even shorter;
cuspidate is longer and stouter).

APOPHYSIS—a cai or expansion at the base of the capsule (also called an hypophysis).

APPENDICULATE—with short, transverse projections

APPRESSED— as — i te closely applied to the stem.

APPROXIMATE —clos oad

ARBOREAL— spa ont

ARCHEGONIUM —the female | as organ, a flask-shaped structure producing an egg.

ARCUATE -curved like a b

ARENICOLOUS-growing 0

AREOLAE-small, aon areas ee a network.

AREOLATION-—the cellular network of a leaf.

ARISTATE-—bristle-pointed, ending in an awn (arista).

ARTICULATE-—with thickened joints, jointed

ASCENDING —directed obliquely upward.

ASPERULOUS~—slightly roughened.

ASSURGENT —directed upward.

ASTOMOUS-—without an sea ae hence indehiscent; also called cleistocarpous.

ATTENUATE-—narrowly taper

AURICLE~—a small, ear-like ane or lobe at the basal margins of a leaf; AURICULATE—with basal

AUTOICOUS—with archegonia and antheridia in separate inflorescences on the same plant;
CLADAUTOICOUS-—with the male inflorescence on a separate branch; GONIOAUTOICOUS

inflorescence; PSEUDAUTOICOUS-—with dwarf male plants epiphytic on the female;
RHIZAUTOICOUS—with the male inflorescence on a very short mess attached to the

AWN -~—a bristle- or hair-point, usually formed by an excurrent costa.
AXIL—the angle between leaf and stem; AXILLARY ~—in the leaf axils.

BASAL MEMBRANE~—a delicate but often well-developed membrane at the base of the inner
peristome of many mosses, commonly terminating in s oe and cilia.
BEAK—the prolonged apex of an aster ai se called a

Bl-—a prefix, meaning 2 h 2 nerves or costae; BICUSPIDATE— with 2 short points;
OUS-—with 2 divisions (or legs”): BIFARIOUS-—in 2 rows; BIFID and BIFURCATE—
forked, divided into : ATE—doubly paired, in 4’s; = 2 lobes;
pels: —in pairs; BIPARTITE—in 2 parts; BIPINNATE —twice- -pinnately branched; BISERI-

T layers
BLOOM-—a ee of powdery or waxy material causing a whitish or bluish overcast; see also

BOG~—a waterlogged open type of vegetation poor in mineral nutrition because all water is derived
from precipitation, rather oak from the soil; often used bee this work and elsewhere for poor
fens (which see)). A bog is usually dominated by Sphagn

BORDERED -having margins differentiated from the rest of is leaf in shape, size, color, or
thickness of cells.

BOREAL-—northern, especially referring to the boreal coniferous forest which extends around the
Northern Hemisphere, from the treeless tundra southward to temperate latitudes (and

BRACTS —specialized leaves surrounding reproductive organs.

BROOD BODIES-—reduced buds, leaves, or branches (propagula) or small, globose, oe or
cylindric to filamentous, septate bodies (gemmae) serving in vegetative reproductio io

BULBIFORM —bulb-shaped.

BULBIL—a small, reduced bulb-shaped bud serving in vegetative reproduction.

BYSSACEOUS —divided into threads.

367

CADUCOUS—deciduous, regularly ae off.

CAESIOUS —bluish or bluish-glaucou

CAESPITOSE-tufted, growing in Bees or
OUS-—containing calcium Pe nee or dolomite r

te eee —a plant of calcareous habitats; opposed to Heike (catia, a plant not
cupying calcareous habitats; CALCICOLOUS —living in calcareous habita

CALLOSE_ hard and thick.

CALYPTRA—a membranous hood over the ob sporophyte, developed from the arche-

gonium, in true mosses ruptured near the base, carried upward on the elongation of the seta,

ontinuing growth to form a cap over the Rae

CAMPANULATE —bell-shaped.

CANALICULATE—channeled.

CANCELLATE -—lattice-like

CANCELLINAE-—the sharply defined, large, empty and usually hyaline basal leaf cells of the

Calymper
CANESCENT —hoary becau use of hyaline hair-points on the leave
CANOPY ena branches borne well above the Stas as in dendroid forms (cf.

Clim eee

CAPIL LEARY ir-like.

CAPITATE (pion ae one and compact forming a head-like capitulum.

CAPSULE-—the spore case, often differentiated into an upper spore-bearing urn and a sterile basal
portion called the neck (which may be considerably differentiated as a broad apophysis or

CARINATE keeled like the bottom of a boat.
CARNEOUS —flesh-colored; CARNOSE —fleshy.
CARTILAGINOUS-—firm and tough.
CASTANEOUS-—chestnut-brown.
CATENULATE-—chain-like.
CAULINE-referring to the stem; CAULID—haploid stem of bryophyte; CAULESCENT— aes
CENTRAL STRAND~—a small poe of elongate cells forming a central axis of some stem
CERNUOUS-—nodding or dro
CER VINE —dark-tawny eg aire ee a deer).
CHARTACEOUS —paper
CHLOROCYSTS-—small, inte. green cells forming a network enclosing the large, hyaline cells
ee of a Sphagnum leaf, also called green cells or chlorophyll cells; also applied to
mall, green cells enclosed by layers of hyaline cells in the leaves of Leucobryaceae or
ingled with them in Paraleucobryum.
CHLOROPHYLLOSE —green, containing chlorophyll.
CILIA —delicate, thread-like structures alternating with the segments of the inner peristome of man
mosses; also applied to hair-like appendages fringing leaves or calyptrae; CILIATE—fringed

CINEREOUS —ashy-gra

CIRCINATE-—curved ae

CIRCUMBOREAL— “videspread | in the Northern Hemisphere, circumpolar.

CIRRATE (CIRROSE)—curled

CLADAUTOICOUS— monoicous, with the male inflorescence on a separate branch.
CLADOCARPOUS-—producing the sporophyte at the end of a special, short branch.

CLAVATE (CLA VIFORM)-slender, elongate, and thickened upward, club-like.

CLEISTOCARPOUS —lacking an operculum and hence irregularly dehiscent (opposed to stego-

COALESCED —fused together.
ae —pressed eeoen squeezed in, constricted, sometimes applied to strangulate capsules
f Orthotrichum, for e
COCHLEARIFORM _ron nd Pa deeply concave, shaped like the bowl of a spoon (not snail-like or
as the derivation would sugges
COHERENT, COHERING-sticking together, sticking to another organ of the same
COLLENCHYMATOUS-with cell walls thickened at the comers; in liverworts such Hiickennae are
called trigones.
COLLUM-—the tapered base or neck of a ca

COMMISSURE-margin ranch leaves of oe , the ormed by the adjacent walls of
hyaline and green noel hence, pores cee? ne commissures are arranged along the lateral
margi

ins
COMOSE-hairy, but nee me bryologists to refer to leaves larger and more crowded, eee tufts
or comae at the stem tips; extremely comose stems may be rosulate in appearan

368

COMPLANATE-filattened together or compressed in 1 plan

COMPLICATE-folded lengthwise; COMPLICATE- CARINATE— sharply folded along a keel.

COMPRESSED —flattened.

CONCOLOROUS-—of the same color

CONDUPLICATE -strongly olde along the middle.

CONFERVOID— a filamen

CONFLUENT -—me

CONNATE-—fused, ene togethe

CONNIVENT —approaching or aes at the tips, directed to a single point, converging.

CONSTRICTED—abruptly narrowed.

CONTORTED~—bent into pone curves, eae twisted.

CONVOLUTE-rolled together and forming a shea

CORDATE -heart-shape

CORIACEOUS —tough, leath

CORMOPHYTE— ss Henrie ae (i.e€., eee ag hae and fern ate seed plants).

CORTEX-—differentiated outer layers of branch, surrounding a central cylinder;
CORTICAL-referring to the outer cells . a or poe. CORTICOLOUS -¢growing on

bark.

COSMOPOLITAN-occurring in all major floristic zones of the world.

COSTA-—nerve of a leaf, sometimes double, sometimes single ease a midrib; COSTATE—with a
costa; ECOSTATE—without a costa.

CRENATE-with rounded teeth; CRENULATE—with minute, rounded teeth.

CRIBROSE-finely perforated, sieve-like.

cae! (CRISPATE)—wavy (like crisp bacon); often used more loosely to mean variously curled,

isted and contorted.

CRISTATE- crested; bearing a crest-like ridge; with a pale margin resembling a

CRYPTOPORE—having immersed stomata, with the guard cells sunken ee ie level of the
exothecial cells and often + covered by them (opposed to phaneropor re).

CUCULLATE~—hooded or hood-shaped; a cucullate calyptra is conic and split up 1 side, resembling
am — hood; also used to describe leaves concave at the tips. A mitrate calyptra may be

base or several-lobed.

CULTRIFORM- shaped like a knife, in bryology meaning curved and asymmetric, like a short,
oe scimitar (apparent an incorrect form resulting from a confusion with cultiform, or
shaped like a p are

CUNEATE —wedge- fae cuneif

CUPULATE-—cup-shaped, pre re swollen.

ok —cupola-like.

CUSHION —wit h stems + erect but somewhat ee to form small tufts (as in Orthotrichum).

up
CUTICLE-—the non-cellular covering of such ae s leaves or capsules, often ee roughened;

es :
CYGNEOUS~—curved downward like a swan’s neck.
CY MBIFORM —concave a broadly bee -shaped.

DECIDUOUS —falling off.

DECLINATE —descending in an arch.

DECUMBENT —ascending ses a eens base, lying prostrate but with ascending ti
DECURRENT-—with the margins extending down the stem below the leaf ace as ridges or

arrow wings.

DECURVED-—curved downward

DEFLEXED~—bent or curved downward.

DEFOLIATE—denunded of leaves.

DEHISCENT-splitting open, referring to capsules opening by valves or by an operculum, or

DELTOID —equilaterally triangular

DENDROID~—branched above a trunk- like base and resembling a

DENTATE—with sharp teeth directed ela DENTICULATE “heey dentate.

DENUDED (DENUDATE)-—with leaves worn off.

DEOPERCULATE-—without the ope Shins aie to a capsule after the operculum has fallen.
veloped.

e
DESCENDING a gradually downward.
DEXTRORSE-—twisted to the right (opposed to sinistrorse).
DIAPHANOUS —transparent.

369

DICHOTOMOUS —equally forked, with paired branch

DIMIDIATE-split on : aa (cucullate calyptrae are NS and hooded).

DIMORPHOUS - of 2 f

DIOICOUS —with eons and antheridia on separate plants, dioecious.

DISCOID (DISCIFORM)-flattened and disc-like or plate-like; male inflorescences may be discoid
Caer small flowers), cupulate (rounded and swollen), or gemmiform (small and

DISPOSITIO —see phyllotaxy

DISTAL—away from the base or cae of attachment (opposed to proximal).

DISTICHOUS—in 2 opposite

DIVARICATE-widely eee fee each other.

DIVERGENT —spreading from each other.

DIVISURAL LINE-—the median line of a peristome tooth, usually zig-zag, sometimes furrowed.

DORSAL-the back, lower, or abaxial surface of a leaf, the outer surface of a peristome tooth, or
the upper surface of a flattened plant body; DORSAL LAMINA-in Fissidens the portion of
ca leaf blade at the back of the costa, opposite the sheathing base; DORSAL PLATES—the

uter surface of exostome teeth.
DORSIVENTRAL_ flattened, with distinct upper and lower surfaces.

ECHINATE-—roughened ae eee spiny projections; ECHINULATE—minutely spiny.

ECOSTATE—without a

EDENTATE-—without ae ne

ELLIPTIC— essentially oblong but convex at sides and ends; ELLIPSOIDAL-—a solid having an
elliptic outline.

EMARGINATE-broad at the apex and shallowly notched Gee than retuse).

EMERGENT— anepad el aes referring to capsules only partly exceeding the tips of the
perichaetial leav

ENDEMIC -limited ae a seis country or floristic area.

ENDOHYALOCYST— abies empty, hyaline cell of cancellinae occupying the leaf base in the
Calymper

ENDOSTOME-—the inner peristome (usually consisting of segments, often from a basal membrane,
frequently alternating wit

eke aaron the inner cell layer of lining of a capsule, forming a spore-sac; usually applied to

er ieee tissue of a young capsule (surrounded by the amphithecium).

sae lackin osta.

ENSIFORM~—sword- shaped.

ENTIRE —not at all toothed.

EPIDERMIS—a Reale ee of cells at the ere eaeimmred applied to ae liverworts).

ssa e pee —in Polytrichaceae, a circular formed by bee xpanded tip of the
lla, es ile the ar Ae ee Lae ee dehiscenc
EQUITANT ae addling like rider on rseback, referring to the aah ds icate and strongly
sheathing leaf bases = Fissidens
ERECT-SPREADING— ing at an angle of about 45° or less, erecto-patent.

EROSE -irregularly astre or ragged, as though gnawed.

EUTROPHIC-tich in minerals, hence + basic or alkaline.

EXANNULATE- reece an annulus

EXASPERATE-roughen

EXCAVATE~—abruptly Rollowed out.

EXCURRENT—extending beyond the ap

EXOSPORE-the outer wall or surface of a

EXOSTOME-—the outer peristome, eee ne oh teeth (only rarely undivided).

EXOTHECIUM-—the outermost layer of the capsule wall, made up of Seton cells.

EXSERTED— De emee and exposed, applied to capsules which project beyond the tips of the
perichaetial leave

EXSICCATI—dried pee ns; usually referring more specifically to widely distributed, published
sets of specimens, with printed labels, used as standards for comparison and often cited in
monographic works.

FALCATE—curved like the blade of a sickle; FALCATE-SECUND-strongly curved and turned to 1

FASCICULATE— eae together, in bundles or fascicles.

FAST ee branches nearly parallel and of + equal length; also erect and columnar,
because ne th type of branching (like a pated Poplar
FEN—an Bn boggy habitat receiving nutrients from Lace (soligenous), hence more

mineral-rich than a bog (ombrotrophic in nutrition, receiving water and nutrients only by

370

precipitation). BOGS are acid; FENS may be acid to neutral or alkaline. RICH FENS are
dominated by sedges, with Sphagnum a minor element. POOR FENS (often called bogs)
are Sphagnum dominated and are floristically similar to true bogs but richer in species

FENESTRATE mane ea window-like openings, a more extreme condition than perforate.

FERRUGINOUS—

ee es an fiber-like bos apres (fibrils); applied to hyaline cells of Sphagnum

h the fibrils may be s or

FILIFORM. slender and elongate, ae. thread- like.

FIMBRIATE-frin

FLABELLATE (FLABELLIFORM)-— fan-shaped.

FLACCID—soft, limp, flabby

FLAGELLUM (pl. FLAGELLA)~a slender branch, referring to minute, axillary brood- anche oo
sometimes to long, <a tapering stems of branches; FLAGELLATE or FLAGELL
FORM -—like the lash of a whip.

FLEXUOSE-—slightly and aenttn bent, twisted, or wavy.

FOLIOSE—leafy

FOOT —the aa absorbing organ of - ene

FORCIPATE-—curved inward like nip

‘—arched

rot)
ia

—pitted.
RONDOSE-closely and regularly branched in 1 plane and resembling a fern frond.
aaa capsule (or more loosely the sporophyte).
FUGACIOUS —vanishing or readily fallin

FUNDUS-—the base of a peristome tooth.

FURCATE-—forked.

FURFURACEOUS - scaly.

FUSCOUS—dull, dark-brown, often tinged with blac
FUSIFORM-—narrow and tapered at both ends, aap: -shaped.

GAMETOPHYTE-—the dominant, sexual ecipeina in mosses, the green, leafy plant; in liverworts,
the gametophyte may be leafy or thallos

GASTEROPODOUS —bulging on 1 side (like a ceerny see also gibbous and ventricose.

GEMINATE-in pairs.

GEMMA (pl. GEMMAE)~—a small, globose, elliptic, or cylindric body of a few cells serving in
tl reproduction (see also brood-body and propagulum); GEMMIFEROUS (GEMMI-

OUS)-—bearing gemmae; oes (GEMMACEOUS)~—bud-like.

eee Ge abruptly bent, like a

GIBBOUS— dace: 2 1 side above, senbine a hunched back.

GLABROUS-—smo

GLAUCOUS— aca a whitish, grayish, or bluish overcast (resembling the waxy bloom on a plum).

GLOBOSE -spherical.

GONIDIA-rarely used to refer to gemmae

GONIAUTOICOUS—monoicous, with the male inflorescence small, ona and axillary.

GRANULOSE-roughened with minute, blunt projections, minutely

GREGARIOUS ~growing close together but not in tufts or mats.

GUARD CELLS-the cells surrounding a stoma, usually paired and kidney-shaped, occasionally a
single cell with a central slit

GUIDE CELLS-—large, pas uae eed — cells in a median row as viewed in the
cross-section of the costa of m

GUTTULATE-—-with He haste noinde ee of strong wall thickening, having the appearance
of drops of liqu

GYMNOSTOMOUS— cam a perist

GY NOECIUM-—the female a ae (more often applied to hepatics).

GYRATE-circinate, coiled.

HABIT -—the aspect or eee appearance of a plant.

HASTATE _abmptly oe and auriculate at base, shaped like an arrow.

HELICOID-spirally twi

HELIOPHILOUS _prsterrin sunny places, photophilous.

HETEROGENEOUS —dissimilar

HETEROICOUS —with ae forms of inflorescence on the same plants (or various plants of the
same species); also called polyoicous and polygamous.

371

HETEROMALLOUS-pointing in all directions (opposed to homomallous, all pointing the same

ay).

HISPID- ae short, stiff ipa bristly.

HOARY-—whitish or gray

HOMOGENEOUS ler aiaee

HOMOMALLOUS— _pointing the same way.

HUMICOLOUS-~¢growing on the ground.

HYALINE-—colorless and transparent.

Me gees —in Sphagnum ees s, a large, empty, ares water-storage cell enclosed in a

work of green chlorocysts; zie called hyaline c

AYALODERM_ —in Sphagnum, a cortex of large, empty, ie cells.

HYDRIC—very wet or aquatic; HYDROPHYTE-an aquatic plant, growing submerged or floating;
HYDROPHILOUS —in water, submerged or floating.

HYDROIDS-tracheid-like conductive cells in the axial strand of many mosses, sometimes also in
the costa a not usually in contact with the central strand.

aoe eee TE-—a plant ee wet habits but not in water; HYGROPHILOUS—adapted to wet (but

aes abita
H¥OROSCOPIC (HYGROMETRIC)- changing position with a change in humidity, readily absorb-

HYPNOID "PERISTOME-a * ‘perfect”’ double peristome, with 16 lanceolate teeth, cross-striolate at
ase me trabeculate at back and a well-developed endostome consisting of a high basal

membrane and 16 lanceolate, keeled segments alternating with 1 or more cilia.
HYPOPHYSIS_a conspicuous swelling or expansion of the neck of the capsule (also called an
apophysis)

IMBRICATE-closely appressed and overlapping.
IMMARGINATE-—without a border
IMMERSED— SEE covered; immersed capsules are exceeded by the tips of the perichaetial

it mmersed stomata have guard cells + sunken below the level of the exothecial cells.
INAPERTURATE SPORES —lacking “apertures” or thin spots in the exine (the most common type
of moss spore).

INCISED— Be into sharp divisions separated by narrow sinuse

INCLINED CAPSULES-~—less than vertical, between erect and ee

INCLUDED-enclosed, not reaching the surface or extending beyond the surrounding organs.
INCOHERENT -rarely used to describe loose tufts (which become separated on collection).
INCRASSATE-—thickened.

INCURVED-—curved upward and inward, applied to leaf tips and margins.

INDEHISCENT—without an operculum

INFLATED-—swollen

INFLEXED— bent upward and inward, applied to leaf margins (opposed to reflexed, bent under and

INFLORESCENCE ~a cluster of sex organs and the leaves or bracts surrounding them
pees ens —a new ee a branch formed after the maturity of sex organs, usually at the base

of an inflor
INSERTION. the aes St ohare of a leaf, branch, or peristome.
ed.

—in

INVOLUTE-inrolled, applie od to leaf margins (opposed to revolute).

ISODIAMETRIC—about as broad as long (including square, rounded, or hexagonal).

ISOPHYLLOUS-—in Sphagnum, stem and branch leaves similar; HEMI- ISOPHYLLOUS —stem leaves
not greatly differentiated.

JUGUM (pl. JUGA)-—a pair of leaves, ee applied to leaves of Fissiden
JULA eon smoothly cylindric, like orm or a catkin, referring re stems or branches with
crowded and imbricate leaves,

KEELED-sharply folded along the middle, like the keel of a boat, carinate.

LACERATE-—deeply and irregularly slashed or t

LACINIATE—dissected into fine, deep, irregular Paivions (laciniae), ciliate-fringed.

LACUNOSE-—with irregular lumina because of deeply porose cell walls, or perforated, sometimes
referring to cells with very bees walls (the lumina resembling a blank space or cavity).

LAMELLAE- green ridges or plates on the costa or lamina of some moss leaves.

LAMINA (Pl. LAMINAE)-—the leaf blade (as distinguished from the costa

LANCEOLATE-—lance-shaped, narrow and tapered from the base pti than ovate).

LANUGINOSE-—woolly, tomentose.

O72

LATERAL-—at the oo opposed to terminal.
LAX-~—soft or loose, usually referring to large thin-walled c
LEAF TRACE~a pee uation of cells of the midrib rie roug igh the cortex to the axial stra

nd; if
aig at all, usually ending blindly in the cortex as false leaf traces, True leaf traces occur
in some species of Polytrichum.

LENTICULAR (LENTIFORM)-—doubly convex, lens-shaped.

LEPTODERMOUS-—thin-walled, referring to leaf cells and also to capsules with soft or delicate

opposed to pachydermous).

LEPTOID—conductive cells in their best development similar to sieve tubes in form and function,
found in the axial strand of stems and setae of polytrichaceous mosses and in setae of many
other mosses.

LEPTOME-—a phloem-like tissue consisting of leptoids and parenchymatous cells.

LEUCOCYST-—the large, empty, hyaline leaf cells of Sphagnum, Leucobryaceae, and Paraleu-

cobryum (also called hyalocysts).

LID—operculum

LIGULATE-— strap-sha aped.

LIMB-—the upper part of a leaf, contrasted with the base.

LIMBATE-bordered.

LINEAR-—very narrow and elongate, with nearly parallel sides (narrower than ligulate).

LINGULATE-—tongue-shaped, oblong with a broadened apex.

LOBE~a major division of a leaf; LOBULE~—a small, basal division of a bilobed leaf.

LORICATE-—equally narrow throughout (referring to the appearance given by a corset or cuirass).

LUMEN (pl. LUMINA)-—the cell cavity.

LUNATE (LUNULATE)-—crescent-shaped.

LURID —wan or gastly pale, sometimes referring to a dingy yellow-brown color.

MACRONEMATA (sing. MACRONEMA)-large, freely branched rhizoids produced around branch

imo :
MAMMILLA (pl. MAMMILLAE)—a bulging protuberance with a blunt, nipple-like tip;
MAMMILLOSE (MAMMILLATE)-—convex with a blunt central projection. (Note:
correctly spelled mamilla, mamillose, etc.)
MAT~—a densely interwoven, horizontal form of growth (cf. Hypnum).
MEDIAN—middle; MEDIAN LEAF CELLS-—cells in the upper middle of the leaf, in leaves with a
well-developed costa, those between costa and margin about 2/3 up the leaf.
MEMBRANE GAPS-—irregular openings in the outer pel = hyaline cells of Sphagnum stem leaves.
MERISTEM-—the growing point or a zone of continuing growth.
MESIC—moist, neither very wet nor very dry as Geen MESOPHYTIC—adapted to a
temperate climate or a ‘ey moist habit
apse eee —more or less neutral in eee neither acid nor basic (cf. eutrophic and
trophic).
smekoN (pl. MICRA)-—one-thousandth of a millimeter (represented by the Greek letter py).
MICRONEMATA (sing. MICRONEMA)—minute club-shaped hairs, resembling paraphyses, borne in
leaf axils in some Mniaceae, aig tlt and Splachnaceae; small rhizoids produced on stems,
leaf decurrencies, costae, or laminae, smaller and less branched than macronemata
ea

MIDRIB—a mid-vein or single costa.
MITRATE (MITRIFORM)-—conic and undivided or equally lobed at base, similar to a bishop’s
miter, referring to calyptrae Oe to cucullate or split on 1 side).
MONILIFORM— resembling a string of bea
MONOICOUS —with antheridia and cies! nia on the same plant, including autoicous, synoicous,
ous, and polyoicous (opposed to dioicous).
ee SPORES— Ba a single scar on the surface of the exine (as in Funaria hygrometrica
m caespitici

NONOPODIAL BRANCHING -differentiation into primary and secondary shoots (or stems and
branc

‘En form, structure

MUCRO~-a short, abrupt point; MUCRONATE— —ending abruptly in a short point usually caused by
a shortly excurrent costa; APICULATE indicates a somewhat longer tip and CUSPIDATE a
longer, sate tooth-like tip.

MULTI-—prefix meaning many, as in multicellular.

MURICULATE-— ican with minute, sharp points (diminutive of muricate).

MUTICOUS-—without a point or awn

373

Dae ees sexual dimorphism, with male plants dwarfed, see also phyllodioicous and pseudo-

NAVICULA R—boat-sh aped.
ee the sterile basal portion of a capsule, sometimes considerably differentiated.

NIDULANT —nesting, rarely mae to refer to the bud-like plants of Ephemerum nested on an
ant isaac

NITROPHII OUS_ prefe Panta ae rich in nitrogenous compounds.

NODOSE-with ot! like ‘hic kenings; NODULOSE—minutely knobbed.

NUTANT-—nodding, drooping

OB-—a ay indicating inversion, as in obovate Saas shaped, with the broader portion at the apex

an the base), obconic, obcordate, e

OBLATE-. ae than long.

OBLONG -—rectangular.

OBSCURE —dark, indistinct

OBSOLETE -scarcely evident, almost lacking (usually referring to the costa).

OBTUSE-broadly pointed (more than 90°)—used by some authors to mean blunt or rounded.

OCHRACEOUS —brownish-yellow.

OCREA (pl. OCREAE)—a thin, filmy sheath terminating the vaginula, rarely noticeable.

OLIGO-—a prefix denoting few; OLIGOPOROUS—with a few pores; OLIGOTROPHIC—poor in
minerals, hence acid in reaction (opposed to eutrophic).

OLIVACEOUS— olive-green.

OPAQUE -—dark, not transparent or translucent

Ep meee the lid covering the mouth of a moss capsule, falling at maturity to release the

Spor
ORAL_refersing to the mouth of a capsule.
ORBICULAR-—nearly circular.
ORIFICE—mouth of a capsule.
OVAL-shortly elliptic.
OVATE-—egg-shaped (with the base broader than the apex).
OXYLOPHILE (OXYLOPHYTE)-—a plant of acid habitats, an acidophile.

PACHYDERMOUS -thick-walled, referring to cell ee a to firm and tough or leathery capsules.
PALMATE-—with finger-like lobes radiating oa the
PANDURIFORM-shaped like the body of a vio Beebe with a sinus on either side).
ao ae —a minute protuberance of various forms; PAPILLOSE—roughened by 1 to many minute
tuberance
PARAPHYLLIA~ small, ey filiform, lanceolate, or leaf-like, sometimes branched scales often
pee ed t branches of pleurocarpous mosses; PSEUDOPARAPHYLLIA—
hy llia-like Seis fea to the br. ee insertions.
PARAPHYSES— hyaline or yellowish, septate hairs mingled with the antheridia (and often with
onia).
PARENCHYMA~a tissue of relatively undifferentiated cells, usually thin-walled and isodiametric
(hence with broad ends); PARENCHYMATOUS-—composed of + short cells joined end to
end (opposed to prosenchymatous
PAROICOUS —with antheridia and archego onia in the same inflorescence but not mixed, the
eridia in the axils of uppermost and perichaetial leaves.
PATENT spreading at 45° or more (used by some authors to mean spreading at 26-45°, patulous
meaning 46-90°
PECTINATE-—resembling ac
PEDICEL-—the stalk or ee on ene the capsule is borne.
PELLUCID-—clear, translucent or transparent.
PELTATE-shield-like, flat and attached by a stalk on the underside.
eae wees Ronee PENDULOUS— somewhat drooping, inclined more than horizontally; with
and branches hanging, a growth form common in tropical cloud forests.
PENICILLATE _like an artist’s brush
R P

e apex.
PERFECT PERISTOME~see hypnoid peristome.
PERICHAETIUM-—the female inflorescence, consisting as leaves or bracts surrounding the
archegonia, opposed to the male ves scence ee m).
PERIGONIUM-—the male inflorescence, a cluster of leaves or meee surrounding the antheridia.
PERISTOME~a es or double circle of Bath ins side the mouth of the capsule, the outer
peristome (or exostome) consisting of teeth, the inner peristome (or endostom) consisting
of segments, ane alternating with cilia, often rising from a basal membra

374

PHANEROPORE-—with superficial stomata, that is, the guard cells ae the same level as other
exothecial cells and not sunken in chambers (opposed to cryptopor

PHOTOPHILOL 1a et well-lighted habitats, heliophilous; opposed . sciaphilo ilou

PHY LLODIOICOUS~—nanandrous; with dwarf male plants which rest on leaves or aan of the
larger female plants.

PHYLLOTAXY-—the arrangement of leaves in spirals on a stem, formerly referred to as dispositio;
e.£., a 3/8 means that 8 leaves occupy 3 complete turns, and disp. 2/5 means that 5
leaves occupy 2 turns of the spiral.

PILIFEROUS— Bes ene

PILOSE—long-hair

PINNATE-—with numerous, spreading branches on 2 sides of the axis and thus resembling a eer

PITTED—with small openings or pores through the walls of ae cells, sometimes marked b
oo depressions in thick-walled cells, also called por

PLAN

Srrunoe «ous. producing the sporophytes sien (from a mere perichaetial bud or a short,
specialized branch) rather than at the stem ; PLEUROCARPOUS mosses are usually
prostrate and freely branched and grow in mats eee than tufts

PLICATE-—folded in longitudinal pleats (plicae).

PLUMOSE-closely and regularly pinnate, feather

PLURI-—a prefix denoting many, as in plripailose pluriseriate, ox pluristratose.

POLYEDAPHIC—occurring on many kinds of substrate

POLYGAMOUS-—with antheridia and Seon in separate inflorescences and also mixed together

e same plants (also called polyoicous and heteroicous).
POLYMORPHOUS ar ea a many forms, variable.

PORES~—small openings or “pits” in the walls of some cells, sometimes conspicuous as depressions
in thick walls between adjacent cells; in Sphagnum the pores are particularly large and
conspicuous in the exposed walls of the hyaline cells of branch leaves and also in outer stem
cells; porose cell walls may be sasanan or quite obviously perforated.

PREDOMINANT -—very conspicuous, promin

PRIMARY STEM-—the main stem, often ae or rhizome-like with reduced or scale-like leaves.

PRIMORDIAL UTRICLE-referring to the cytoplasmic membrane which is sometimes particularly
noticeable as a result of shrinkage from the cell wall on drying

PROCESSES —segments of the endostome.

PROCUMBENT -spreading, prostrate.

PROLIFEROUS-—continuing growth by the production of a new stem (or innovation).

sale Nee (pl. PROPAGULA)-reduced bud, branch, or leaf serving in vegetative reproduction

e also eae body).

PROSENCHYM TOUS—with narrow, elongate cells overlapping at the ends opposed to paren-
peering PROSENCHYMA ~a tissue made up of elongate cells with tapered ends.

PROSTRATE—creeping.

PROTEAN-—exceedingly variable.

PROTONEMA (pl. PROTONEMATA)— —green, branched filaments produced on germination of

a

PROTONEMATOUS BS aa macronemata capable of producing protonemata if detached from
stems (Mniac

PSEUDO. —a ee a ane false; PSEUDANNULUS~—an apparent annulus consisting of small,

non-vesiculose cells; PSEUDAUTOICOUS or PSEUDOMONOICOUS—with dwarf males grow-

ing on female plants; PSEUDOPARAPHYLLIA-—small, unistratose, lanceolate or filiform

structures sega ar ane? often found in pleuroc arpous mosses in small numbers

the gametophytic ne below the sporophyte in Sphagnum and re ee serving the

Sphagnum branch leaves, consisting of fibril rings not enclosing a perforation; PSEUDO-
STOMATA-~—in Sphagnum capsules, vestigial stomata, consisting of two guard cells but no

pore.
PUGIONIF ORM —dagger-shaped.
PULVINATE-cushion-like.
PUNCTATE-—dotted, usually referring to aes markings; PUNCTULATE—minutely dotted.
PUNGENT —ending in a stiff or hard poin
PYRIFORM-—pear-shaped.

375

QUADRATE-square.
QUINQUEFARIOUS—in 5 rows.

Lee ee sees eae non-chlorophyllose threads attaching the plants to the substrate,
ms, occasionally growing from leaf tips; the radicles of mosses have oblique
eae "RADICU LOSE-—covered with rhizoids.

RAMENTUM-a woolly covering or tomentum, made up of radicles

RAMULI (sing. RAMULUS)-—the smaller divisions of a much branched plant; RAMULOSE—with
small br

RANK -—r1ow.

RECEPTACLE—an expanded and + elevated structure bearing sex organs and eventually sporo-
phytes in the Marchantiaceae, rarely used in the Musci to refer to the tissue on which the
reproductive organs are

RECURVED-—curved downward or ‘backward, referring to leaf margins or tips, marginal teeth, or
peristome sree

REFLEXED—bent backward.

REGULAR— ae ne Hie

I ed

REPAND— slightly sinuose.

REPENT —creeping.

RESOR in a eence or erosion of parts of cell walls in Sphagnum leaves; RESORPTION
W-a furrow produced by the disappearance of the outer cell walls at the leaf
of some species of Sphagnum, best seen in cross-section.

RETICULATE~—in a network.

RETORT CELLS—cells with a pore at the upper end, i a short, projecting neck, found in

the cortex of branches in many species of Sphagnum.

RETUSE-slightly indented at a broad apex.

pein emia away, referring to an annulus which falls in a ring; REVOLUTE -rolled

ward and backward, eae to leaf margins.

RHIZAUTOICOUS- monoicous with the male inflorescence on a ie short branch attached to the

female stem by rhizoids and thus appearing to be a separate p

RHIZOIDS— ee or branched, septate filaments, dead at eee anchoring the plant and

sometimes + covering the stem (see we radicles).

RHIZOME-—a slender, horizontal, subterranean, root-like

RHOMBIC—diamond-shaped; RHOMBOIDAL_ ‘longer than honte oblong-hexagonal.

hinky.
ROSTRATE—beaked; ROSTELLATE-shortly or minutely beaked.
ROSULATE-like a sae aris to leaves enlarged and crowded at tips of stems and
resembling small flow

RUFOUS —reddish.
RUGOSE-transversely wrinkled or undulate; RUGULOSE-slightly wrinkled crosswise.
RUPESTRAL--growing on rocks (also called saxicolous).

SACCATE-—abruptly and deeply concave, forming a sack.
SAPROPHYTE-—living on dead organic matter, Bee to parasite.
SAXICOLOUS~growing on rocks (rupestral).
SCABROUS-—rough.
SCALARIFORM —ladder-like.
SCALPELLIFORM~—asymmetric and Sepealic a blade of a pen-knife.
SCARIOUS—dry and thin, scale-like or Hage
Naan —preferring shady habita
CLERO —the thick-walled ie ‘forming a woody cylinder in stems and branches of

Sp i a
SECUND— re to ls
SEGMENTS-—the tooth- like divisions of the inner peristome, sometimes called processes.
SEPTATE ee by cell walls, having partitions.
SERIATE-—in :
SERRATE— eee et with marginal teeth pointing forward; mre SERRATE —with teeth
tlapping and joined in pairs; SERRULATE-—minutely serra
SESSILE—without a stalk or seta, hence immerse
SETA-—the stalk supporting the capsule
SETACEOUS —bristle-like.
SHEATHING—+ surrounding and clasping the stem or the base of the seta.
SHOULDER-the area where a leaf base is abruptly narrowed to the limb.

376

SIGMOID—somewhat S-shaped

SILICEOUS—having a high silicon content, sandy (usually poor in bases); SILICICOLOUS growing
on sandstone or other oT

SINISTRORSE-—twisted to the

SINUATE (SINUOSE)—wavy; ROLE —minutely or slightly wavy.

SINUS—a depression between two lobes or prominences, the ‘entation between lobes of a leaf
(liverworts).

SORDID—of a dirty or muddy colo

ele ncieas tapered from a broad, rounded apex (similar to lingulate but more abruptly

rowed from a broad a :

SPICULOSE_ sharply and va toothed or papillose

SPINDLE-SHAPED —fusiform; narrow, elongate, and tapered at both ends.

SPINOSE-spiny, with sharp, slender teeth or projections; SPINULOSE—minutely spiny.

SPORANGIUM-—the spore-sac of a capsule or, more loosely, the entire capsule.

sae Birr mostly spherical, nearly always unicellular bodies produced in the capsule as a

of reduction division, giving rise on germination to protonemata.

sonore the spore-bearing generation; the spore-bearing plant, produced by the fertilization
of an egg, remaining attached to the gametophyte and partially dependent on it, typically
consisting of foot, seta, and capsu le.

ia ts at an angle of 45° or more; wide-spreading means nearly 90°, squarrose 90°,

uarrose- siecahecee 90° with recurved tips.

SQUALID sordid or dir

SQUARROSE— ae at ase angles; SQUARROSE-RECURVED-~—spreading at right angles, with
the tips curved downward.

STEGOCARPOUS— ee aoe cult differentiated (opposed to cleistocarpous or astomous).

STELLATE -star-sha

STEREIDS-—slender, ae thick-walled, fiber-like ae me in groups in the costa of some
mosses; also applied to linear, thick- sau as

STERILE— barren, not fertile, lacking sporophyt

STIPE—the unbranched base of a dendroid or ae plant; STIPITATE— . a trunk- ee base

STOLONIFORM-rteferring to slender, eee spreading branches or stems, with sm r and
— differentiated leaves and ae aaa or “rooting” near oe tips, abla

ers or stolons of a strawberry,

STOMA (pl. STOMATA)—opening in the cab vil usually in the neck, surrounded by guard
cells.

STRAMINEOUS -straw-colore

STRANGULATE-—deeply constricted below the mouth of a capsule

STRIATE—marked ee ri longitudinal ridges (striae); STRIOLATE— finely ridged.

STRICT —straight and r

STRUMOSE-with a oe -like swelling (or struma) at the base, applied to some capsules;

TRUMULOSE-—with a small or indistinct swelling at 1 side of the capsule base

SUB-—a prefix meaning nearly, almost, somewhat, as in subacute, subentire, oubparcurrent, or
subquadrate; also used to mean under, as in submedian, subinvolucral.

SUBULA~a long, slender point; SUBULATE-—slenderly long-acuminate, shaped like a needle or awl.

aan strongly plicate, with deep longitudinal folds (more extreme than striate).

SURCULUS~a sucker-like Serene deel used to refer to upright secondary stems arising from a
onan primary stem or “rhiz

UTURE-—a line formed by the aon ‘of 2 par
SYNOICOUS-—with antheridia and archegonia ae in the same inflores
SYSTYLIOUS—with the operculum remaining attached to the tip of the aienela after dehiscence.

TEETH Raley of a single peristome or of the exostome of a moss with a double peristome;
of the endostome are called segments or processes.

TENIOLAE the intramarginal border of linear cells in the leaves of Calymperes.

TERETE-—rounded in cross-section, cylindric.

TERRESTRIAL~growing on the ground, ferricolous.

TESSELLATED-~—checkered, in a pattern of squares, applied to the basal membrane of Tortula.

TETRAD~a group of 4 developing or rarely mature spores; because of poland act in tetrads some
mature spores are ellipsoidal, angular, or + kidney-shaped, sometimes tetrahed.

THALLOSE-—flat, not much differentiated, and cordate or ribbon-like in ee form.

TOMENTOSE-—woolly, densely radiculose, covered by a tomentum of rh s.

ie —a heater ti _ the outer peristome; also applied to ne ee or projections at the

TORTUOSE_ irregularly bent or twisted.

ie

TRABECULATE —with projecting cross-bars (trabeculae) at the back of some peristome teeth.
TRI-—a_ prefix meaning thrice; TRIGONOUS, TRIQUETROUS-—3-cornered; TRIFARIOUS,

TRIGONES-—thickened corners of cells.

TRILETE SPORES-~—having a 3-pronged scar on the wall (as in Sphagnum).

TRIRADIATE-—3-pronged, often used to refer to the scar of a trilete spore.

TRUNCATE~—abruptly cut off or squared off at the apex.

TUBULOSE-—tube-like, usually referring to leaves with strongly incurved mar

TUFA~—a porose limestone formed by deposition from calcareous Nica Gin formed because of
‘the CO, relationships—in photosynthesis and respiration—of mosse

TUMID —swollen, inflated.

TURBINATE-—top-shaped.

TURF —a tufted form of ees with stems erect and parallel, often in extensive clones.

TURGID—plump or swollen

UMBONATE-—convex with an ll rounded, central point.
UMBRACULIFORM —umbrella-like
UNCINATE—hooked.
UNDULATE-wavy; RUGOSE is a more extreme condition.
UNEQUAL-~—asymmetric.
Sel arene ier: in a point similar to an animal’s
UNI-—a prefix g 1, as in unipapillose, unistratose, ot uniseriate.
URCE POLATE —urmshaped, applied to capsules constricted nes a wide mouth and abruptly
wed at the base.
URN- te iene bearing portion of a capsule, as opposed to the sterile nec
RICLE~—a vesicle, rarely used to refer to the enlarged cortical ei ‘of Sphagnum (see also
primordial Pe

VAGINATE— sheathing; VAGINANT LAMINA-—in caer ae expanded lamina at base of the
he

leaf which clasps the stem and the base of the leaf a
VAGINULA-—the ring or sheath enveloping the base of a seta, ae from the base of the

ied o
VENTER-the swollen basal portion es a arc ae m, contai ainin

lower surface of a See pias or tha

VENTRICOSE~—bulging on 1 side below eae : omc:
VERMICULAR~-narrowly cylindric and c d, like a worm.
VERRUCULOSE —ir oe pas

VESICULOSE —inflated, bladder-like.

VILLI (sing. VILLUS)— ee
VITTATE-striped.

WEFTS-—a loosely interwoven, souls ome growth form (as in Thuidium).
WHORLED —arranged in a ring or
WIDE-SPREADING~spreading at a ome angle but less than 90° (which is squarrose).

XERIC—very dry; XEROMORPHIC-structurally adapted for dryness; XEROPHILOUS-—preferring
dry places; XEROPHYTE~—a plant of dry, often hot places.

ZYGOMORPHIC~—bilaterally symmetric.

An of
a _. : ——
Fy ww - F = ;

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INDEX

Subject Index

Allergies 338
Antibiosis 33
Arthropod association 339; tardigrades 339

Bog iron 292

Calyptra, function ie
Chemosynthesis 1
Chromosomes ee toe counting 134

Dispersal ere 18, $1, 52, 70, 82, 93, 97,
119, 137-138, 143- 144, 150-151, 181, 302,
ce 313, 349, 351; insect dispersal 143- 144;

gabo nds (moss balls) 93

Distribution 62, 71, 302; nunataks 62

Dwarf males 84, 298

Ecology and habitats 1, 18-19, 53-54, 66, 70, 99,

108, 119, 125, 127, 133, 136- 137, 139- 150,

152, 159, 165, 251, 304, 339, 347, 357-358;
d .

ers
7; sand-binders 127;
submerged mosses 53-54; weeds 137, 149-150,
159

Environmental conditions 18-19, 33; acidity 18,
38, 153; ion-ex

change
Evolution 133- 134, ra 4, 166, 214, 221, 362-

64; methods o y 196; reduction 133,

362-364; Bee 134, 166, 354

Fawley buns 93

Food: caloric and vitamin content 338; odor 143,
144, 338; taste 338

Fungal infection (Tilletia sphagni,
schimperi) 23

Helotium

Genetics oe 133-134, 298, 354
Glacier m
Goblin Gold 146-148

Limestone deposits: didymodontoliths 108; trav-
ertine 108; tufa 108, 251, 292

Leaves: Fissidens 52; isophylly 39, 46; phyllotaxy
221

pene? family concept 224
uminescence 146-148

Manna 93
Tee cycles 84-85, 304
Mineral uptake 119, 165
Moss balls 93

Park, Mungo 61-62

Peristomes 121, 181, 224, 311-312, 325; hypna-
ceous 224, 325; perfect 224, 325; structure
and ile ae 70, 148, 182-183

Photoperiod 85, 1

ES ate ee ae

Pygm

Regeneration 134, 137-138; apogamy 133; apo-
166

Reproduction: asexual 119, 125, 137, 138, 160;
hybrids 66, 102, 133; sexual 119, 129, 153,
298, 302-304: sexual dimorphism 84, 143,
298, 348; sterility 302-304

Saprophytes: Cryptothallus 42; Buxbaumia 347

Seeding 330, 357

Sphagnum, cacsification ae ecology 18-19; fun-
gal infection 23; ion-exchange 18; isophylly
39, 46; eee 19; ee res 23; spores 18;
surgical pads 33; uses ae variability 39, 46,
48, 60; water relations

Spores 18; germination a 125; dimorphism
143; microspores 23; numbers 70-71; paly-
nology 18; viability 137

Succession: bog 18-19, 358; fire 70, 136, 137,

"387: pioneers 70, 125, 136, 139,

347; rock 125; soil 150

Taxonomy: cytotaxonomy 166, 196; methods of
study 39, 166, 196, 298; variability 39, 46,
48, 60, 69, 100, 124-125, 267-268

Thuidiaceae, family concept 224

Uses 19, 328, 337-339; decoration 328, 337; nest-
ing material 152; gardens 33

Vascular tissue 214, 362; lignin 362

Taxonomic Index

Acaulon muticum var. rufescens 119
Aloina brevirostris 119

rigida 119
Amblyodon dealbatus 186
Am spender serpens 261
erpens var. juratzkanum 261

399

varium 261

Amphidium lapponicum 208

Ree anes
camptodon splachnoides 222

anes rothii 50

rupestris 50

400

Anomodon attenuatus 233

viticulosus 235
Aphanorrhegma serratum 133
Apple Moss 193
Astomum muhlenbergianum 102
Atrichum angustatum 353

crispum 355

undulatum 353

var. altecristatum 355

var. oerstedianum 355

var. undulatum 355
Aulacomnium acuminatum 185

androgynum 18

heterostichum 182

palustre 184

turgidum 185

Barbula convoluta 109

fallax 111
michiganensis 111
reflexa ey :
unguiculata 109

Bartramia ia phyla 193
pomiformis 192

Bird Wheat 355

Blindia acuta 65

Bog Moss

Brachythecium acuminatum 286
acuminatum var. crytophyllum 287

Broom Mos

Bryoandersonia illecebra 296
Bryoerythrophyllum recurvirostrum 105
Bryoxiphium norvegicum

var. flaccidum 160
cryophilum
lisae var. de ea 162
see A ii 164

pallen
sre aaneees 161

tortifolium 162

minakatae

Callicladium haldanianum 311
Calliergidium pseudostramineum 267, 273
Calliergon cordifolium

ge
+ me
qe
a
~

a
Calliergonella cuspidata 277
Campylium chrysophyllum 253
51

rotensum 255
Catoscopiim nigritum 190
Cave Mos
aes purpureus 69
Cinclidium stygium 178
Cinderella 136
Cirriphyllum piliferum 294
Climacium americanum 336

ss 135, 1
ip eh commutatum 251
filic
Gs ptsthalis saat 42
lus

Ctenidium mol m 333
Cynodontium sceist’ 91
strumiferu

torquescens 91
Desmatodon obtusifolius 115
Dichelyma capillaceum 217

atum 217

pallesc es s 217
Dichodontium eee 90
Dicra el cervic 76

villeana ao

Dicranum bonjeanii 85

coparium

spurium 87
undulatum 89
viride 80
per an Lae 106
tophac 08
DPR aan eee 348
Distichium capillaceum 67
inclinatum 67
Ditrichum flexicaule 72
flexicaule var, densum 72
ile 72

rhynchostegium 73
Drepanocladus aduncus 266

aduncus fo. capillifolius 267

var. kneiffii 267

var. polycarpus 266

o, gracilescens 267

exannulatus 268

fo. brachydictyus 268

fo. tun 2
var. alpinus 2
var. purpurascens 268
var. rotae 268

var, ea 264
sendtneri 267
rare ite 268
uncinatus 263
var. symmetricus 264
r. plumulosus 26
vernicosus
Drummondia prorepens 207
Dryptodon patens 128

Emerald Dewdrops 139
Fnalype ciliata 94
cera 9

ee a 96
streptocarpa 97

vulgaris 96

Entodon cladorrhizans 305
seductrix

Entosthodon 135, 136

Ephemerum cohaerens 139
crassinervium 139
spinulosum 4

Eucladium verticillatum 100

Eurhynchium hians 298
seanere 296
riparioides 298

Ete Moss 94

Fabronia ciliaris 223
Feather Mosses 325
Fern Mosses 240
Fissidens adiantoides 55
bryoides 60

cristatus 55
exiguus 60

exilis 59
fontanus 52
grandifrons 52
minutulus 60
obtusifolius 58

Flapper Mos

dalecarlica

aei 216
var. hypnoides 214, 216

neo-mexicana 216

novae-angliae 216

patula 216

sphagnifolia 216
Fork Moss 80
Forsstroemia trichomitria 209
Woes americana

flavicans 138
inaeoanss 135

Goblin Gold 144
Golf Club Moss 190
Grimmia affinis 125

var, dupretii 124
var, rivularis 124

unic
Gymnostomum aeruginosum 98
calcareu
recurvirostrum 99
ar, commutatum 100

var. |
Gyroweisia tenuis 9

Hair Cap Mosses 355

Haplocladium microphyllum 239
virginianum 240

Haplohymenium triste 236

Hedwigia ciliata 129

Helodium blandowii 247
blandowii var. helodioides 249
paludosum 249

Helotium schimperi 2

Herzogiella striatella 332
turfacea 332

Heterocladium dimorphum 231

Heterophyllium haldanianum 311

Homalia trichomanoides 220

402

Homomallium adnatum 319
Hump-backed Elves

Hygroamblystegium fluviatile 260
2

noterophilum
orthocladon 260
ae ee
inifolium 260
Hygrohypim luridum 270
sale

ie m 271
yin brevirostre 344

splendens 342
umbratum
Hyophila involuta 59, 108
Hypnum aera 327
curvifoliu

e 326
eee eer 136

Isopterygiopsis muelleriana 323
Isopterygium distichaceum 322
e

pulchellum 322
Juniper Moss 357

Knight’s Plume 334
Knothole Moss 223

La Charbonniére 136

Leptobryum pyriforme 149

Leptodicty um brevipes 257
25

Leskea gracilescens 226

polycarpa 226
Leskeella nervosa 227
Leucobryum albidum 93

glaucum 92

Leucodon brachypus var. andrewsianus 212

julaceus 211
Lindbergia a 238
Long-Neck Moss 7
Luminous ie: 144

Meesia — 188
i iq a 187
a 187
Mi sicher mielichhoferi 165
Mnium

ine
affine var. affine 173
var. ciliare 173

cuspidatum 171

drummondii 172

longirostrum 1
lycopodioides 168
marginatum
mediu

m 17
[ical ein 169
pseudopunctatum 176

Mountain Fern Moss 344
Myurella julacea 229
sibirica 229

Neckera pennata 219

Oncophotrus virens 79
wahlenbergii 79

Orthotrichum alpestre 200
anomalum 199

m 197
speciosum var. Boe 198
var. specio 199
stellatum
strangulatum 200

Paludella squarrosa 189

Paraleucobryum longifolium 89

Peat Moss 17

Petticoat Moss 142

Phascum ae 112
floerkeanum 112

Philonotis capillaris 194
fontana

Physcomitrella patens 133
Physcomitrium hookeri 133

nium ae
Plagiopus oederiana 191
Plagiothecium ayiliani 307
denticulatum 309
etu
latebricola 308
Plat eae confervoides 321
jungermannioides 319
subtile 321
Platygyrium repens 316

Pleuriditrichum marylandicum 66

Pleuridium palustre
subulatum 66

Pleurozium schreberi 302

Pogonatum alpinum 361
dentatum 361
pensilvanicum 361
urnigerum 361

Pohlia annotina 153
annotina var. annotina 153
var. decipiens 153
bulbifera 155
cruda 151
elongata 153
filiformis 156
lescuriana 155

wahlenbergii 155
Polytrichum commune 360
formosum 358
ar. aurantiacum 358
juniperinum 3

truncata 113

Pottioideae 97

Powder Gun Moss 350

Pseudobryum 166

Pseudoleskea patens 226
radicosa 226

Pseudoleskeella tectorum 227

Pterygoneurum ovatum 119
Ptilium crista-castrensis 334
Pylaisiella intricata 318
polyantha 317
selwynii
Pyramidula tetragona 139

Red-Mouthed Mnium 170
Rhabdoweisia crispata 90
Rhacomitrium aciculare 128
canescens 127
fo. ericoides 127
fasciculare 128
heterostichum 126

Rhodo seum 164
Rhynchostegiella compacta 301
Rhynchostegium serrulatum 299
Rhytidiadelphus loreus 342

3

Rhytidium rugosum 345
Rose Moss 164
Rough Neck Moss 341

Saelania ape tatiee 71

Schistostega pennata 144

Schwetschkeopsis cprens 223

Scopelophila ligulata 119

Scorpidium scorpioides 278
turgescens 280

Seligeria calcarea 63
campylopoda 63

donniana 63

Sematophyllum demissum 314
Shaggy Moss 341
Silvery Bryum 159
Sphagnum an eae! 18
sect, Cusp
oe Gen 18
sect, Polyclada 18
sect. Rigida 18
sect. Sphagnum 18
sect. Squarrosa 18
sect. Subsecunda 18
subg. Inophloea 17
ee Litophloea 18
nulatum var. porosum 36

girgensohnii 40

imbricatum 24

lindbergii 33

She har 21
6

pulchrum 33
quinquefarium 364
recurvum
var. amblyphyilum 33
2

var. platyphyllum 39
var, ru
var. subsecundum 38

tere
warnstorfii 43
wulfi
Splachnum ampullaceum 141
luteum 142
rubrum 142
sphaericum 144
Stair-Step Moss 344
Sword Moss 62

Taxiphyllum deplanatum 324
taxirameum
Tayloria acuminata 144

403

404

lingulata 144
serrata 144
splachnoides 144
Tetradontium brownianum 351
Tetraphis geniculata 351
pellucida 351
Tetraplodon angustatus 142
mnioides
Thamnobryum alleghaniense 337
Thelia asprella 238

ery 243
a. oe 23
Timmia Senn 180

— ana 180

I. aoe arica 180
Tae cae nitens 281
nitens var. falcifolium 282

Top Moss

Tortella fragilis 103
humilis 104
inclinata 105
tortuosa 103

Tortula ones 117
mucr nifo lia 116

Moss 336
Trematodon ambiguus 78
oe ollis

Trian ar Woodreveler 341

dricus 74
Trichostomum tenuirostre 105
Ulota coarctata 204
crispa 20
hutchinsiae 205
Umbrella Moss 142

Undulating Catharine 354
Urn Moss 132

Water- oe Cord Moss 136
Water s 213

Wavy eae 354

Uke fan 101

White Moss 93

ci S 5 Heads 80

Zygodon viridissimus 2

0
sacred var. rupestris 207
r, viridissimus 207

Composition and photo- ae printing by
, IN

BRAUN-BRUMFIE
Ann Arbor, an