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MOSSES OF GUATEMALA

EDWIN B.« BARTRAM

FIELDIANA: BOTANY

VOLUME 25

Published by

CHICAGO NATURAL HISTORY MUSEUM

JANUARY 31, 1949

MOSSES OF GUATEMALA

MOSSES OF GUATEMALA

EDWIN B. BARTRAM

FIELDIANA: BOTANY

VOLUME 25

Published by

CHICAGO NATURAL HISTORY MUSEUM

JANUARY 31, 1949

THE LIBRARY OF THE

ra 2 3 1949

PRINTED IN THE UNITED STATES OF AMERICA

BY CHICAGO NATURAL HISTORY MUSEUM PRESS

CONTENTS

Families Included in Volume 25

Sphagnaceae 4

Andreaeaceae 7

Fissidentaceae 8

Ditrichaceae 24

Seligeriaceae 31

Dicranaceae 32

Leucobryaceae 67

Calymperaceae 73

Encalyptaceae 81

Pottiaceae 82

Grimmiaceae 140

Funariaceae 146

Splachnaceae 151

Bryaceae 153

Mniaceae 182

Drepanophyllaceae 184

Eustichiaceae 185

Rhizogoniaceae 186

Meeseaceae 187

Bartramiaceae 189

Erpodiaceae 204

Orthotrichaceae 206

Helicophyllaceae 230

Rhacopilaceae 231

Hedwigiaceae 232

Cryphaeaceae 236

Leucodontaceae * 242

Prionodontaceae 245

Trachypodaceae 247

Pterobryaceae 249

Meteoriaceae 260

Phyllogoniaceae 273

Neckeraceae 275

Lembophyllaceae 288

Pilotrichaceae 290

Hookeriaceae 293

Leucomiaceae 322

Hypopterygiaceae 323

Fabroniaceae 325

Leskeaceae 330

Amblystegiaceae 341

Brachytheciaceae 350

Entodontaceae 362

Plagiotheciaceae 371

Sematophyllaceae 383

Hypnaceae 397

Hylocomiaceae 415

Diphysciaceae 419

Polytrichaceae 420

INTRODUCTION

In the course of three botanical expeditions to Guatemala,

sponsored by Chicago Natural History Museum, Drs. Paul C.

Standley and Julian A. Steyermark accumulated extensive collec-

tions of mosses from nearly every department. These collections

have been beautifully supplemented by the later explorations of

Dr. Aaron J. Sharp during 1945 and 1946 in the highlands of Guate-

mala, principally at altitudes between 1,250 and 3,500 yieters, where

the bryophyte flora is exceedingly rich and diversified.

It has been my privilege to study all of these collections and the

results, supplemented by whatever other information I have been

able to gather, are presented in the following pages. Although Dr.

Sharp's collections have not been listed in full detail, all of his im-

portant discoveries, representing an addition of more than seventy

species to the previously known flora, are included in the present

work. As a result, the moss flora of Guatemala and its affinities

are now far better known than those of any other Central American

country.

As this is the first attempt at a realistic analysis of the mosses

of any restricted tropical American country, the task is, to some

extent, explorative. For, until many of the tropical American genera

are disencumbered of numerous dubious species by careful revisional

studies, the specific entities involved are open to a wide variety of

interpretations.

Being astride the Cordilleran axis, extending from the Atlantic

to the Pacific and possessing a highly diversified terrain, Guatemala

should and does support a rich moss flora broadly representative

of the American tropics. Since the literature relating to tropical

American mosses is widely scattered and often difficult of access,

it seems desirable to describe briefly, key out and illustrate the species

known to occur in this limited area.

One of the more interesting problems involves the relationship

of the Guatemalan flora with that of the surrounding and contribu-

tory regions. As might be logically expected, the Mexican types are

abundantly represented. However, the most significant feature is

2 FIELDIANA: BOTANY, VOLUME 25

the occurrence of numerous species typical of the northern United

States and Canada, including Fissidens taxifolius, Ditrichum gigan-

teum, Distichium capillaceum, Dicranella varia, Blindia acuta,

Dicranum flagellare, Barbula reflexa, Barbula icmadophila, Tortella

tortuosa, Grimmia apocarpa, Mnium serratum, Bryum cuspidatum,

Meesea longiseta, Orthotrichum anomalum, Cratoneuron filicinum,

Campylium stellatum, Campylium chrysophyllum, Hygrohypnum pa-

lustre, Drepanocladus aduncus, Brachythecium rutabulum, Eurhyn-

chium pulchellum, Plagiothecium denticulatum, Hypnum cupressiforme,

Rhytidium rugosum, Hylocomium brevirostre, Diphyscium foliosum

and many others of similar affinities. It is probable that this group

is made up of species forced southward during glacial times. These

species persisted in isolated communities in the highlands of Guate-

mala even after the retrograde migration had taken place. Many

of these records mark the extreme southern limit of distribution of

these species in North America.

Evidently the Cordillera served as a main highway of north-

south migration. It is interesting, therefore, to record as immigrants

from the opposite direction such typical Andean species as Ditrichum

gracile, Dicranella vaginata, Campylopus Jamesoni, Amphidium

cyathicarpum, Holomitrium pulchellum, Hymenostomum Jamesoni,

Streptopogon erythrodontus, Streptopogon rigidus, Entosthodon acidotus,

Actinodontium megalocarpum, Eustichia Spruceana, Rhizogonium

Lindigii, Eucatagonium politum and Pseudodimerodontium bolivianum,

all of which find their northern limits in Guatemala or adjacent

Mexico.

A small group of tropical Brazilian mosses, comprising Coleo-

chaetium Standleyi, Philophyllum tenuifolium and Puiggariella auri-

folia, all three genera new to North America, suggests that in the

past the tropical American flora may have been more closely in-

tegrated than it is now.

Among the well-known Caribbean species, found principally in

the eastern lowlands, are Sphagnum meridense, Fissidens pellucidus,

Dicranella subinclinata, Syrrhopodon ligulatus, Syrrhopodon incom-

pletus, Syrrhopodon lycopodioides, Calymperes lonchophyllum, Tortula

mniifolia, Philonotis glaucescens, Micromitrium mucronifolium, Pseu-

docryphaea flagellifera, Jaegerinopsis squarrosa, Or,thostichopsis tetra-

gona, Pireella cymbifolia, Papillaria nigrescens, Meteoriopsis patula,

Neckeropsis undulata, Neckeropsis disticha, Helicodontium capillare,

Entodon macropodus, Sematophyllum caespitosum and Taxithelium

planum. Many of these species reach Florida on the north and range

widely into northern South America in the opposite direction.

BARTRAM: MOSSES OF GUATEMALA 3

The following local mosses extend north through Mexico to

Arizona and New Mexico: Anoectangium arizonicum, Anoectangium

obtusifolium, Merceya ligulata, Husnotiella revoluta, Symblepharis

helicophylla, Dicranum rhabdocarpum, Tortula fragilis, Ptychomitrium

Leibergii, Ptychomitrium serratum, Brachymenium mexicanum, Bryum

truncorum, Orthotrichum Bartramii, Anacolia laevisphaera, Bartramia

microstoma, Braunia secunda, Fabronia ciliaris, Fabronia Wrightii

and Pleuropus Bonplandii. A smaller group, comprising Ditrichum

ambiguum, Timmiella anomala, Barbula vinealis, Grimmia tricho-

phylla, Orthodontium pellucens and Eurhynchium praelongum, extends

into California. Evidently with increasing distance from the Con-

tinental Divide, the number of species common to both areas

decreases.

The Guatemalan moss flora may be roughly divided into three

zones. The lowland mosses up to altitudes of about 1,500 meters

are broadly representative of the Caribbean regions. Here such

typical families as Fissidentaceae, Pterobryaceae, Meteoriaceae,

Hookeriaceae and Sematophyllaceae are abundantly distributed.

The second zone, representing the interior highlands from 1,500 or

2,000 meters up to 3,500 meters, supports a much more diversified

and highly intriguing flora, including many surprising vagrants from

far distant northern and southern latitudes. Such families as

Dicranaceae, Pottiaceae, Bryaceae, Bartramiaceae and Orthotri-

chaceae are developed to an amazing extent. Pottiaceae alone

account for 71 species distributed in 25 genera. The rocky summits

of the higher mountains from 3,600 to 4,600 meters above sea level

are truly alpine in character. Typical of these bleak, rugged domes

are the following rupestrine species: Andreaea rupestris, Distichium

capillaceum, Encalypta vulgaris, Grimmia ovalis, Rhacomitrium

crispulum and Hedwigidium imberbe. In more sheltered places and

a little below the bare rocks the upper fringes of the rich highland

flora are encountered.

The strange mixtures present a puzzling problem in phytogeog-

raphy, especially when two species of the same genus such as

Ditrichum giganteum of northern United States and Yukon and

Ditrichum gracile of the South American Andes are found growing

in close proximity. It is hardly possible to appraise the full signi-

ficance of these facts now, but the evidence surely indicates that

Guatemala is one of the principal focal points of geographical dis-

tribution in tropical North America.

While probably far from complete, the total of approximately

519 species and 205 genera is broadly representative of the local

4 FIELDIANA: BOTANY, VOLUME 25

mosses and may cover at least 80 per cent of the actual flora. The

percentage of endemic species is relatively small. Including the

new species, approximately 58, or about 11 per cent of the total

known flora, are not known outside of Guatemala. As the adjacent

regions are more thoroughly explored, many of these endemics may

prove to have a wider distribution than our present knowledge

indicates.

No one can realize the shortcomings of the task in hand more

keenly than I do. Yet I am hopeful that the work may prove useful

to students of the local moss flora and in future investigations relating

to Mexico and the other Central American countries.

The types of all new species are in the author's herbarium. A

complete series of Standley's and Steyermark's collections, including

duplicate types of the new species collected by them, can be found

in the herbarium of Chicago Natural History Museum. The full

series collected by Sharp is deposited in the herbarium of the Uni-

versity of Tennessee. As many species were collected in large

quantities, a wide distribution of representative specimens will no

doubt be made among the principal North American herbaria.

ORDER 1. SPHAGNALES

Typical peat mosses forming dense tufts or cushions in bogs or

on wet mountain slopes. Stems elongate, pale green tinged with

brown or red, without radicles, composed of a central core of lax

cells enclosed in a woody cylinder and surrounded by one to several

layers of large, hyaline cells. Branches in fascicles, crowded at tips

of stems in dense heads. Branch leaves composed of two kinds of

cells in one layer; a network of linear chlorophyllose cells in the

meshes of which are large rhomboidal hyaline cells usually porose

and reinforced by spiral fibers. Stem leaves similar in structure

but dissimilar in shape. Capsules dark brown, subglobose, with a

convex lid, on a short fleshy stalk or pseudopodium. Peristome

lacking.

A very distinct order comprising a single genus, Sphagnum, of

cosmopolitan distribution.

1. SPHAGNACEAE

1. SPHAGNUM Linn., Sp. PI. 1106. 1753.

1. Cortical cells of stems and branches not fibrillose 2

Cortical cells of stems and branches fibrillose . . .3

BARTRAM: MOSSES OF GUATEMALA 5

2. Chlorophyllose cells of branch leaves exposed on both surfaces, pigment brown

4. S. subsecundum

Chlorophyllose cells of branch leaves exposed on inner surface of leaf, pigment

red 3. S. meridense

3. Chlorophyllose cells of branch leaves central and included ... 1. S. magellanicum

Chlorophyllose cells of branch leaves exposed on inner surface of leaf

2. S. imbricatum

1. SPHAGNUM MAGELLANICUM Brid., Muse. Rec. 2, (1): 24. 1798.

Coarse plants tinged with brown. Cortical cells of stems and

branches fibrillose. Stem leaves Ungulate; branch leaves broadly

ovate, deeply concave, Chlorophyllose cells central and enclosed on

both surfaces of leaf by the hyaline cells. (Fig. 1, A-C.)

Dept. Huehuetenango: Steyermark 49918, 49921, 49922.

Distribution: Labrador to Alaska south to Florida and California,

Bermuda, Europe, Asia, South America.

Rare locally and confined to alpine regions.

2. SPHAGNUM IMBRICATUM Hornsch., Russow, Beitr. Torfm. 21.

1865.

Plants pale greenish brown. Cortical cells of stems and branches

fibrillose. Stem leaves Ungulate; branch leaves imbricated, ovate,

Chlorophyllose cells exposed on inner surface of leaf, lower hyaline

cells ridged or fringed on inner walls where overlying the chloro-

phyllose cells. (Fig. 1, D-G.)

Dept. Huehuetenango: Steyermark 49912.

Distribution: Alaska, eastern North America south to Gulf

States, Cuba, British Honduras, Europe, Asia.

Alpine meadow. Like the preceding, this species is known locally

only from alpine regions. Dr. LeRoy Andrews informs me that he

has a collection from British Honduras. The peculiar fringed fibrils

on the inner walls of the hyaline leaf cells are a marked feature.

3'. SPHAGNUM MERIDENSE (Hampe) C. M., Syn. 1: 95. 1848.

Sphagnum acutifolium meridense Hampe, Linnaea 20: 66. 1847.

Rather slender pale plants usually tinged with red. Stem leaves

triangular-ovate, bordered, toothed at apex. Branch leaves laxly

imbricated, oblong-ovate, concave, abruptly short pointed, apex

involute, truncate, toothed. Chlorophyllose cells exposed on inner

surface of leaf. (Fig. 1, H-J.)

FIELDIANA: BOTANY, VOLUME 25

FIGURE 1

A-C, Sphagnum magellanicum: A, stem leaf, X14; B, branch leaf, X14; C,

cross section of leaf cells, X270.

D-G, Sphagnum imbricatum: D, stem leaf, X14; E, branch leaf, X14; F, basal

cells of branch leaf, XllO; G, cross section of leaf cells, X270.

H-J, Sphagnum meridense: H, stem leaf, X14; I, branch leaf, X14; J, cross

section of leaf cells, X270.

Dept. Alta Verapaz: Standley 92577, 92590, 92620; Steyermark 1*3955. Dept.

Huehuetenango: Steyermark 1*971*6, 50183, 51898, 51950, 51951. Dept. El Pro-

gresso: Steyermark 1*3097, 1*31*55. Dept. Zacapa: Steyermark 1*2631, 1*3203, 1*3257,

1*3287. Dept. Chiquimula: Steyermark 30989. Dept. Jutiapa: Steyermark 31935.

Distribution: Florida, West Indies, Mexico, Central and South

America.

By far the most frequent species of Sphagnum and apparently

broadly distributed at altitudes above 1,200-1,500 meters. The

characteristic ruddy tinge is usually a dependable indicator of this

species.

4. SPHAGNUM SUBSECUNDUM Nees, Sturm, Deuts. Fl. Crypt. 17.

1819.

Rather robust, brittle plants, tinged with brown. Cortical cells

of stem in one layer. Stem leaves lingulate, entire. Divergent

branches cuspidate pointed, branch leaves broadly ovate, apex

truncate, toothed; hyaline cells strongly fibrillose with numerous

small pores on the dorsal face; chlorophyllose cells subrectangular

in section, exposed on both surfaces of leaf. (Fig. 2, A-D.)

Dept. Huehuetenango: Sharp 1*968. Dept. Baja Verapaz: Sharp 511*3.

BARTRAM: MOSSES OF GUATEMALA 7

Distribution: Greenland and Labrador south to Gulf states and

Mexico; California to Washington.

Wet rocks and swamp area at low to moderately high altitudes.

These seem to be the first records of the species in Central America.

The brown pigment, cortical cells of the stem in one layer and the

chlorophyll cells exposed on both surfaces of the branch leaves are

good diagnostic features.

ORDER 2. ANDREAEALES

Small dark colored, brittle rupestrial mosses of high altitudes.

Stems slender, simple or branched. Leaves crowded, cells small,

incrassate, in one layer; costa lacking or single. Capsules terminal,

exserted on a short pseudopodium when ripe, without lid or peristome,

splitting vertically into 4 (6-8) valves at maturity; columella

persistent, spores smooth or papillose.

2. ANDREAEACEAE

One family and one genus only represented in North America.

FIGURE 2

A--D, Sphagnum subsecundum: A, stem leaf, X14, B, branch leaf, X14; C, part

of cross section of stem, XllO; D, cross section of leaf cells, X270.

E-H, Fissidens Svihlae: E, plant, XI; F, leaf, X30; G, upper leaf cells and

margin, X270; H, cells of duplicate blade, X270.

8 FIELDIANA: BOTANY, VOLUME 25

1. ANDREAEA Hedw., Sp. Muse. 47. 1801.

1. ANDREAEA RUPESTRIS Hedw., Sp. Muse. 47. 1801.

Brittle reddish brown plants in dense tufts or cushions. Stems

1-3 cm. long, simple or forked. Leaves crowded, imbricated when

dry, ovate-lanceolate, concave, ecostate, muticous or slightly

pointed, usually papillose on back, to 1.5 mm. long; cells incrassate,

linear below, rounded-quadrate above. Capsules small, finally

exserted on a short pseudopodium. (Fig. 3, A-D.)

Dept. San Marcos: Steyermark 35542, 3551*60,, 3554.7, 35548a, 35549. Dept.

Solola: Steyermark 47494.

Distribution: Greenland to Alaska, south along mountains to

Georgia and California, Europe, Tasmania, New Zealand, southern

South America.

Exposed rock faces and crevices; locally confined to highest

elevations, 3,800-4,600 m.

var. ALPESTRIS Thed., Bot. Not. 1849: 79. 1849.

Differs from the typical form in the smaller, blunter leaves.

Dept. San Marcos: Steyermark 35786, 36089. Dept. Quezaltenango : Steyer-

mark 34209.

Distribution: Greenland south to New York and Alaska south

to Montana and Washington; also Europe.

ORDER 3. BRYALES

The great majority of mosses are comprised in this large order.

Variable in detail, they seem to grow from a filamentose protonema.

The spores and columella are developed from the endothecium.

Capsules borne on a definite seta of variable length, indehiscent or

opening by a lid; peristome present or lacking.

3. FISSIDENTACEAE

Small to medium sized plants with distichous, equitant leaves

flattened in one plane, split to the costa on the inner side of the

basal part into two blades clasping the stem. Lamina cells uniform,

hexagonal or rounded, usually in one layer; costa ending in or below

apex; seta terminal or lateral. Capsule erect or inclined; peristome

simple, of 16 teeth, entire or split to or below middle into two

subulate forks; spores small.

BARTRAM: MOSSES OF GUATEMALA 9

1. FISSIDENS Hedw., Sp. Muse. 152. 1801.

A large and very distinct group with the characters of the family.

I have profited largely by Dr. Grout's recent revision of the

North American species of Fissidens (Grout 20), which has clarified

many problems in this intricate genus.

1. Leaves bordered entirely or in part with elongated cells 2

Leaves not bordered with elongated cells 12

2. Leaves flaccid, cells large and lax 3

Leaves firm, cells small and dense 5

3. Costa ending far below apex, capsules horizontal, asymmetrical

3. F. reticulosus

Costa longer, capsules suberect, symmetrical f 4

4. Border distinct to apex 1. F. mollis

Border ending well below apex 2. F. dissitifolius

5. Leaf cells unipapillate 6. F. Svihlae

Leaf cells smooth or pluripapillate 6

6. Dorsal blade of leaf long decurrent 4. F. longidecurrew

Dorsal blade not decurrent 7

7. Stem leaves unbordered, peristome teeth undivided 10. F. muriculatus

Stem leaves bordered in part, peristome teeth forked 8

8. Leaf cells papillose, dense and obscure 9

Leaf cells smooth, distinct 11

9. Border weak, confined to basal part of duplicate blades 7. F. leptopodus

Border strong, extending to or beyond apex of duplicate blades 10

10. Apical and dorsal blades often variously bordered 8. F. Weiri

Apical and dorsal blades not bordered 11. F. elegans

11. Border narrow, of one layer of cells 5. F. repandus

Border strong, bistratose 9. F. Steyermarkii

12. Cells of leaf blade in 2 or more layers 24. F. grandifrons

Cells of leaf blade in one layer 13

13. Aquatic, slender floating plants 25. F. debilis

Not aquatic, plants tufted or gregarious 14

14. Marginal leaf cells incrassate, forming a definite border

17. F. austro-adiantoides

Marginal leaf cells not differentiated 15

15. Small plants, leaf margins crenulate 16

Robust plants, leaf margins entire below, apex rounded 20

16. Leaf cells large, smooth, distinct 12. F. pellucidus

Leaf cells small, papillose or mammillose, obscure 17

17. Leaf cells pluripapillate, minute plants, leaves rounded. . . 16. F. pusillissimus

Leaf cells unipapillate 18

10

FIELDIANA: BOTANY, VOLUME 25

18. Leaves subacute 13. F. Donnellii

Leaves rounded or obtuse 19

19. Leaves less than 1 mm. long, ovate 15. F. Steerei

Leaves 1-1.5 mm. long, oblong, abruptly contracted to an obtuse apex

14. F. radicans

20. Setae lateral 21

Setae terminal 23

21. Leaves subentire 22. F. polypodioides

Leaf margins crenulate or serrate 22

22. Leaves acute, margins serrulate, costa excurrent 21. F. taxifolius

Leaves obtuse, serrate near apex, costa percurrent 23. F. Oerstedianus

23. Leaves broadly acute 20. F. similiretis

Leaves broadly rounded 24

24. Leaves lingulate, apex rounded 19. F. lingulatus

Leaves oblong-lingulate, apex obtuse 18. F. asplenioides

1. FISSIDENS MOLLIS Mitt., Journ. Linn. Soc. 12: 600. 1869.

Dioicous; stems 1-1.5 cm. long, often branched. Leaves strongly

contorted when dry, flaccid and laxly spreading when moist, to 4 mm.

long, linear-lanceolate, short acuminate, strongly bordered all

around, border confluent at apex; costa ending below apex; cells

FIGURE 3

A-D, Andreaea rupestris: A, plant, XI; B, leaf, X20; C, apex of leaf, X270;

D, leaf of var. rupestris, X20.

E-F, Fissidens mollis: E, leaf, X14; F, apex of leaf, XllO.

G-H, Fissidens dissitifolius: G, leaf, X14; H, apex of leaf, XllO.

BARTRAM: MOSSES OF GUATEMALA 11

lax, hexagonal, thin-walled, to 40 M long above. Capsule small,

inclined. (Fig. 3, E-F).

Dept. Izabal: Steyermark ^1781 a.

Distribution: Mexico, Costa Rica, West Indies, South America.

On damp rocks at low altitude. More robust than F. dissitifolius

and distinguished at once by the leaf border continuous to the apex.

2. FISSIDENS DISSITIFOLIUS Sull., Proc. Am. Acad. 5: 274. 1861.

Smaller than F. mollis. Stem less than 5 mm. long. Leaves

oblong-ovate, short acuminate, about 1.5 mm. long, border narrow

and indistinct, ending below apex about opposite tip of costa; costa

ending above middle of apical blade; cells laxly hexagonal, to 25-30

fj. long in the apical blade. Capsule small, inclined. (Fig. 3, G-H.)

Dept. Chiquimula: Steyermark 30260.

Distribution: Mexico, Cuba, Porto Rico.

On damp rocks at low altitude. Uncomfortably near the follow-

ing species to which it is closely allied.

3. FISSIDENS RETICULOSUS (C.M.) Mitt, Journ. Linn. Soc. 12: 603.

1869.

Conomilrium reticulosum C. M., Syn. 2: 525. 1851.

Conomitrium hookeriaceum C. M., Bull. Herb. Boiss. 5: 173. 1897.

Plants small, about 2 mm. high. Leaves 4-9 pairs, contorted

when dry, larger upward, to 3 mm. long, lanceolate, acuminate,

bordered to near apex; costa ending near middle of apical blade;

cells lax, 35-40 M long, 10-14 n wide, thin-walled. Capsule nodding

or horizontal, asymmetrical; urn less than 1 mm. long.

Distribution: Mexico, West Indies.

No Guatemalan collections have been seen but C. hookeriaceum

C.M. is cited as a synonym of F. reticulosus (Grout 16, p. 171).

The shorter costa and asymmetrical capsules should distinguish it

from F. dissitifolius.

4. FISSIDENS LONGIDECURRENS The>., Smithson. Misc. Coll. 782: 10.

1926.

Slender, tufted, brownish yellow plants. Stems to 12-15 mm.

long, laxly foliate. Leaves strongly crispate when dry, oblong-

lanceolate, broadly acuminate, bordered all around, to 2 mm. long,

12

FIELDIANA: BOTANY, VOLUME 25

dorsal blade long decurrent; costa ending just below apex; cells

irregularly hexagonal, 6-8 M- Capsule small, suberect. (Fig. 4,

A-B.)

Dept. Quezaltenango : Standley 671*70.

Distribution: Mexico.

Shaded bank at high altitude. Readily identified by the long

decurrent dorsal blade often extending nearly to the next leaf below.

5. FISSIDENS REPANDUS Wils., Kew Journ. Bot. 3: 52. 1851.

Fissidens tortilis Hampe & C. M., Bot. Zeit. 22: 340. 1864.

Fissidens Carionis C. M., Bull. Herb. Boiss. 5: 171. 1897.

Fissidens fasciculato-bryoides C. M., Bull. Herb. Boiss. 5: 172. 1897.

Small greenish yellow plants in lax colonies. Stems 6-8 mm. long.

Leaves decurved and much crisped when dry, lanceolate, bordered

all around, to 2 mm. long, short and broadly acuminate, dorsal

lamina narrowed toward base; costa ending just below apex; cells

small, rounded, hexagonal, 8-10 n- Seta 6-7 mm. long; capsule

horizontal to suberect; urn oblong, 1 mm. long. (Fig. 4, C-E.)

Dept. Alta Verapaz: Standley 701+69, 701+73. Dept. Huehuetenango: Steyer-

mark 501+57. Dept. Quezaltenango: Steyermark 31+086a, 31+092a; Standley 85199.

Dept. Sacatepequez: Standley 58650. Dept. Zacapa: Steyermark 291+52.

B

T

D

FIGURE 4

A-B, Fissidens longidecurrens: A, part of stem and leaf, X14; B, apex of leaf,

X110.

C-E, Fissidens repandus: C, plant, XI; D, leaf, X14; E, apex of leaf, XllO.

F-I, Fissidens leptopodus: F, plant, XI; G, leaf, X14; H, apex of leaf, XllO;

I, basal margin of duplicate blade, X270.

BARTRAM: MOSSES OF GUATEMALA 13

Distribution: Florida, Mexico, West Indies, South America.

On soil, trees and walls at moderate altitudes. This seems to

be a rather frequent species locally. The small size and strongly

crisped leaves bordered throughout will serve to identify it easily.

6. FISSIDENS SviHLAE Bartr., Bryol. 50: 202. 1947.

Small, gregarious plants, yellowish green. Stems 1-2 mm. high.

Leaves erect-spreading, slightly flexuous when dry, about 1.5 mm.

long, narrowly lanceolate, entire, acuminate, bordered all around with

a narrow cartilaginous band of linear cells; costa percurrent; upper

leaf cells hexagonal, diam. 8-10 n, strongly unipapillate, cells of

duplicate blades lax, hyaline and smooth, to 40 or 50 /* long. Seta

5 mm. long; capsule erect, minute; peristome teeth deeply cleft.

(Fig. 2, E-H.)

Dept. Suchiate: near Chicacoa, Svihla 2871.

Endemic.

Nearest F. yucatanensis Steere but leaf cells less than half as

large. The papillae are about 5 /x high and are best seen in profile

on the upper leaves which are slightly twisted when the plant is

mounted entire. In many respects and especially the lax areolation

of the duplicate blades this species closely resembles F. Kegelianus

C. M. but the unipapillate leaf cells are at once distinctive.

7. FISSIDENS LEPTOPODUS Card., Rev. Bryol. 37: 120. 1910.

Fissidens michoacanus Th6r., Smithson. Misc. Coll. 782: 12. 1926.

Small sordid green plants. Stems 2-3 mm. high. Leaves 5-20

pairs, 1-1.25 mm. long, oblong-lanceolate, acute; costa ending in

or near apex; upper leaves indistinctly bordered toward base of

duplicate blades with several elongated cells; margins crenulate;

cells small, obscure, densely papillose. Capsule ovoid, suberect.

(Fig. 4, F-I.)

Dept. Peten: Bartlett 12155, 121*85, 1251*5, 12553; Lundell 2100a.

Distribution: Mexico, Trinidad.

On disintegrated limestone at low altitudes. F. Garberi L. & J.

in which the border is confined to the perichaetial leaves should

eventually be found in Guatemala but so far I have seen no collec-

tion that could be definitely referred to this species.

14 FIELDIANA: BOTANY, VOLUME 25

8. FISSIDENS WEIRI Mitt., Journ. Linn. Soc. 12: 602. 1869.

Fissidens Howelli Bartr., Proc. Cal. Acad. Sci. IV. 21 : 78. 1933.

Small yellowish green plants, closely gregarious. Stems 4-5 mm.

long with 6-9 pairs of leaves. Leaves erect-spreading, little altered

when dry, about 2 mm. long, oblong-lanceolate, acute, border strong

and pellucid below, distantly denticulate, ending far below apex of

apical blade and often spurred on the inner side above; costa pellucid,

ending just below apex; margins of apical blade minutely crenulate

where unbordered; cells minute, very obscure, densely and minutely

papillose. Seta terminal, 2-3.5 mm. long; capsule suberect, cylin-

drical, urn 1 mm. long. (Fig. 5, A-C.)

Dept. El Progresso: Steyermark 4.3697.

Distribution: West Indies, Galapagos Islands, Brazil.

On earth at high altitude. A plastic species but easily recognized

by the variable leaf border, often lacking on the apical and dorsal

blades but when well developed extending half way or more up the

apical blade and frequently spurred on the inner edge.

9. FISSIDENS STEYERMARKII Bartr., No. Amer. Flora 153: 177. 1943.

Dioicous. Rather robust, dull green terrestrial plants, densely

gregarious. Stems to 1.5 cm. long and 3-4 mm. wide with leaves,

sparsely radiculose below. Leaves in numerous pairs, the lower

minute, gradually larger upward, the upper to 3.5 mm. long and 0.6

mm. wide, lightly contorted when dry, erect-spreading and often

falcate when moist, oblong-lanceolate, short acuminate, bordered

all around, the border strong, cartilaginous, bistratose and confluent

with the percurrent costa at apex; cells distinct, hexagonal, with

firm pellucid walls, 8-10 M in diam., smooth or very faintly papillose.

Seta terminal, solitary, about 8 mm. long; capsule inclined, urn

1 mm. long; lid short, conical, deep red; peristome teeth about 375 n

high, deeply bifid, the forks erect, coarsely and densely papillose;

spores pale, papillose, diam. 10-18 M- (Fig. 5, D-F.)

Dept. San Marcos: Steyermark 36576 TYPE. Dept. Quezaltenango: Steyer-

mark 33889. Dept. Suchitepequez : Steyermark 35310.

Endemic. On moist rocks and slopes at medium to high altitudes.

Although evidently near F. plurisetus Bartr. of Panama the

distinctions are sharply defined and well maintained. In F. Steyer-

markii the setae are constantly solitary in all three collections

representing over a hundred fruiting plants; in all parts these plants

BARTRAM: MOSSES OF GUATEMALA

15

are about twice the size of F. plurisetus and the leaf cells by contrast

are distinct and smooth or very faintly papillose.

10. FISSIDENS MURICULATUS Spruce, Journ. Linn. Soc. 12: 593.

1869.

Small slender plants with numerous pairs of leaves. Stems 4-5

mm. long. Leaves curved when dry, 1-1.5 mm. long, oblong, broadly

acute, not bordered, dorsal blade ending in a rounded lobe at base;

margins crenulate all around; costa nearly percurrent; cells obscure,

6-8 ju, rather bluntly papillose. Perichaetial leaves indistinctly

bordered at base of duplicate blades; seta terminal, 1.5 mm. long;

capsules suberect, peristome teeth entire. (Fig. 5, G-J.)

Dept. Sacatepequez: Standley 88961 a.

Distribution: West Indies, Brazil.

On tree at moderate altitude. The undivided peristome teeth

and faintly bordered perichaetial leaves are clearly diagnostic. Dr.

Grout has confirmed the determination with the comment that the

papillae of the leaf cells are shorter and blunter than in the type.

If Moenkemeyera is recognized as a valid genus this species would

belong there.

B

H

FIGURE 5

A-C, Fissidens Weiri: A, plant, Xl; B, leaf, X14; C, part of margin of apical

blade, X270.

D-F, Fissidens Steyermarkii: D, plant, Xl; E, leaf, X14; F, apex of leaf, X 110.

G-J, Fissidens muriculatus: G, plant, Xl; H, leaf, X14; I, apex of leaf, XllO;

J, part of peristome, XllO.

16

FIELD IANA: BOTANY, VOLUME 25

FIGURE 6

A-B, Fissidens pellucidus: A, leaf, X30; B, upper leaf cells and margin, X320.

C-D, Fissidens pusillissimus: C, leaf, X 68; D, upper leaf cells and margin, X270.

E-G, Fissidens taxifolius: E, plant, XI; F, leaf, X20; G, apex of leaf, XllO.

11. FISSIDENS ELEGANS Brid., Bryol. Univ. 2: 691. 1827.

Small dull green plants, laxly gregarious. Stems 4-5 mm. long.

Leaves numerous, curved with deflexed points when dry, oblong-

lanceolate, acute, about 1 mm. long, strongly bordered on the

duplicate blades only; margins of apical and dorsal blades minutely

crenulate; costa pellucid, percurrent; cells minute, about 5 /*, obscure,

papillose. Seta terminal, about 4 mm. long; capsule suberect. (Fig.

7, A-D.)

Dept. Quezaltenango : Standley 86644 (distributed as F. radicans).

Distribution: Mexico, West Indies, South America.

On wet rock at moderate altitude. The strongly bordered dupli-

cate blades and unbordered apical and dorsal blades distinguish this

species from any of its local allies.

12. FISSIDENS PELLUCIDUS Hornsch., Linnaea 15: 146. 1841.

F. subcrenatua Schp., in C. M. Syn. 2: 531. 1851.

F. rufulus Sull., Proc. Am. Acad. 5: 275. 1861.

F. Wrightii Jaeg., Enum. Fissid. 12. 1869.

Small gregarious plants, green tinged with reddish brown. Stems

about 5 mm. long, with 8 or 10 pairs of leaves, 1.5 mm. wide with

BARTRAM: MOSSES OF GUATEMALA

17

leaves. Leaves slightly curved when dry, well spaced, not over-

lapping, 1 mm. long, oblong-ovate, bluntly acute, unbordered, dorsal

blade ending abruptly at base of costa; margins crenulate; costa

strong, brownish, ending below apex; leaf cells hexagonal, smooth,

large and pellucid, to 15 n in diam. Seta slender, 3 mm. long;

capsule small, ovoid, erect. (Fig. 6, A-B.)

Dept. Baja Verapaz: Sharp 5181.

Distribution: Georgia, Mexico, West Indies to Brazil.

Moist bank at moderate altitude. The small size, unbordered

leaves with relatively large, smooth, pellucid cells are distinctive as

compared with all the other Guatemalan species.

13. FISSIDENS DONNELLII Aust., Bot. Gaz. 4: 151. 1879.

Autoicous; minute plants. Stems short. Leaves numerous,

narrowly oblong, usually broadly subacute, unbordered, crenulate

all around; costa ending 3-5 cells below apex; cells unipapillate.

Seta terminal, 2-4 mm. long; capsule small, erect. (Fig. 7, E-F.)

Dept. Alta Verapaz: Steyermark M086, 44957, 44968. Dept. Quezaltenango :

Standley 84S17.

Distribution: Florida, Mexico, West Indies.

B

FIGURE 7

A-D, Fissidens elegans: A, plant, Xl; B, leaf, X14; C, apex of leaf, XllO;

D, apex of duplicate blades, XllO.

E-F, Fissidens Donnellii: E, leaf, X14; F, apex of leaf, X270.

G-I, Fissidens radicans: G, leaf, X14; H, apex of leaf, XllO; I, apex of leaf,

X270.

18 FIELDIANA: BOTANY, VOLUME 25

On wet banks and calcareous rocks at medium to high altitudes.

The toothed margin of the duplicate blades may be a distinctive

feature of this species.

14. FISSIDENS RADICANS Mont., Ann. Sci. Nat. 11, 14: 345. 1840.

Slightly larger than F. Donnellii. Stems often innovating.

Leaves oblong, 1-1.5 mm. long, scarcely tapering above until just

below apex where the blade is contracted to a short, obtuse point;

costa ending below apex; margins of duplicate blades finely crenulate

and similar to apical margins.

Dept. Peten: Lundell 3387 in part. (Fig. 1, G-I.)

Distribution: Florida, Mexico, Panama, Porto Rico, British and

French Guiana.

A lowland species. The larger size and different shaped leaves

will help to distinguish this species from F. Donnellii.

15. FISSIDENS STEEREI Grout, No. Amer. Flora 153: 191. 1943.

Minute plants similar to F. Donnellii. Leaves less than 1 mm.

long, broadly ovate, rounded at apex, unbordered, crenulate-serrate

all around; costa ending 5-8 cells below apex; cells hexagonal, 8-10 n,

coarsely unipapillate, the marginal row somewhat transversely

elongate and pellucid. Sporophyte unknown. (Fig. 8, A-C.)

Dept. Jalapa: Steyermark 32913.

Distribution: Porto Rico.

Base of waterfall at medium altitude. The relatively broader,

shorter leaves, rounded at apex and with the costa ending slightly

lower will distinguish this species from F. Donnellii.

16. FISSIDENS PUSILLISSIMUS Steere, Ann. Bryol. 10: 116. 1938.

Minute, laxly gregarious plants, yellowish green. Stems less

than 2 mm. long and less than 1 mm. wide with leaves. Leaves

in 4 to 6 pairs, 0.3-0.5 mm. long, oblong, obtusely rounded, un-

bordered; costa strong, brownish, ending well below apex; margins

papillose-crenulate all around; dorsal blade ending at leaf insertion

or in the reduced lower leaves ending some distance above base of

costa; cells strongly pluripapillose and obscure. (Fig. 6, C-D.)

Dept. Peten: Lundell 29^8.

Distribution: British Honduras.

BARTRAM: MOSSES OF GUATEMALA

19

Bark of tree at low altitude. A rare lowland species so small

and inconspicuous that it is likely to be collected only by chance.

The obscure pluripapillose leaf cells should distinguish this species

from its near allies in the Crenularia Section without much trouble.

17. FISSIDENS AUSTRO-ADIANTOIDES C. M., Bull. Herb. Boiss. 5:

547. 1897.

Fissidens incrassatolimbatus Card., Rev. Bryol. 37: 119. 1910.

Robust plants, 6-7 cm. high, branched. Stems to 8 mm. wide

with leaves, densely foliate. Leaves flexuous with contorted points

when dry 5-6 mm. long, 1 mm. wide, oblong-lanceolate, short

acuminate, bordered all around with a distinct band of incrassate

cells in 4-5 rows and in 2 layers in spots; margins coarsely and ir-

regularly toothed toward apex; costa percurrent; cells 12-15 M in

diam., irregularly rounded, with thick, pellucid walls. Sporophyte

not seen. (Fig. 8, D-E.)

Dept. Alta Verapaz: Turckheim 7722 (as F. incrassatolimbatus Card.); Standley

7160k, 89819. Dept. Huehuetenango : Steyermark 48865, 48575, 48866a.

Distribution: Jamaica.

On rocks and trees at medium altitudes. I have a single plant

from the type collection of F. incrassatolimbatus Card. (Turckheim

FIGURE 8

A-C, Fissidens Steerei: A, leaf, X14; B, leaf, X54; C, apex of leaf, X270.

D-E, Fissidens austro-adiantoides: D, leaf, X14; E, apex of leaf, XllO.

F-H, Fissidens asplenioides: F, leaf, X14; G, apex of leaf, XllO; H, upper leaf

cells and margin, X270.

20 FIELDIANA: BOTANY, VOLUME 25

7722} which seems to be inseparable from the collections cited above

which Dr. Grout has referred to F. austro-adiantoides C. M. In all

of these collections the border is bistratose in spots here and there

but never continuously. The apical teeth are irregular and vary

considerably on the same plant. I have not seen any authentic

material of F. Bourgaeanus Besch., but the distinctions seem rather

vague and I should not be surprised if eventually both F. austro-

adiantoides and F. incrassatolimbatus will have to be included in

F. Bourgaeanus.

18. FISSIDENS ASPLENIOIDES Hedw. Sp. Muse. 156. 1801.

Rather robust yellowish green plants growing in dense colonies.

Stems usually simple, 1.5-5 cm. long, densely foliate. Leaves

erect-spreading with strongly circinate tips when dry, about 3 mm.

long, 0.5 mm. wide, ligulate, unbordere'd, obtusely rounded at apex;

margins minutely crenulate all around; costa ending well below apex;

cells irregularly hexagonal, dense, not incrassate, 8-10 M- Seta

terminal, 4-6 mm. long; capsule oblong, inclined. (Fig. 8, F-H.)

Dept. Alta Verapaz: Standley 69651, 70502, 71151. Dept. San Marcos:

Steyermark 35988, 36010, 36488, 36494, 36647; Standley 68483, 864.58. Dept.

Totonicapan: Standley 65942. Dept. Quezaltenango : Standley 65328, 83267,

83279, 83504, 83685, 84607, 85121, 85529, 85939, 85667, 85951, 85998; Steyermark

34723, 35111, 35139, 35153. Dept. Suchitepequez : Steyermark 46852. Dept.

Sacatepequez: Standley 58957. Dept. Solola: Standley 62350, 62749. Dept.

Chimaltenango: Standley 61529, 62013, 79906. Dept. Zacapa: Steyermark 42221,

42672, 43204. Dept. Chiquimula: Steyermark 30922.

Distribution: Wide in tropical regions throughout the world.

On damp banks, rocks and trees at medium to high altitudes.

Easily recognized by the slender fronds with the leaf points neatly

coiled backwards when dry. It is often richly colored and is by far

the commonest species of the genus throughout Central America.

19. FISSIDENS LINGULATUS C. M., Bull. Herb. Boiss. 5: 172. 1897.

IFissidens gracilifrondeus C. M., Bull. Herb. Boiss. 5: 172. 1897.

Moderately large plants. Stems to 1.5 cm. long. Leaves to 20

pairs, broadly rounded and slightly crenulate at apex, about 2 mm.

long; costa ending below apex; duplicate blades to % the length of

the leaf; cells rounded, 7-10 n, mammillose, smaller toward margins.

(Fig. 9, A-B.)

Cuesta de Atitlan: Bernoulli & Cario 115.

Distribution: Mexico.

BARTRAM: MOSSES OF GUATEMALA

21

The only plants I have seen are from Mexico and these seem to

differ from F. asplenioides in the broader leaves with the apices

uniformly rounded without any suggestion of an apiculus. The

duplicate blades are also relatively longer but the distinctions are

neither sharp nor very convincing.

20. FISSIDENS SIMILIRETIS Sull. var. GUADALUPENSIS (Schimp.)

Grout, No. Amer. Flora 153: 193. 1943.

Very similar in every way to F. asplenioides except that the leaf

apex is broadly acute instead of obtusely rounded. (Fig. 9, C-D.)

Dept. San Marcos: Steyermark 35701*, 361*92 (both distrib. as F. asplenioides).

Distribution: With the species, West Indies.

On shaded banks at high altitudes. The distinctions between

these plants and F. asplenioides are slight. In fact the series of

F. asplenioides from Guatemala shows considerable variation in the

form of the leaf apex from broadly rounded to obtuse and minutely

apiculate so that both F. lingulatus and F. similiretis guadalupensis

as represented here might be included in the form circle of F. as-

plenioides without much violence to conservative judgment.

FIGURE 9

A-B, Fissidens lingulatus: A, leaf, X14; B, apex of leaf, XI 10.

C-D, Fissidens similiretis var. guadalupensis: C, leaf, X14; D, apex of leaf,

X110.

E-G, Fissidens polypodioides: E, leaf, X14; F, apex of leaf, X54; G, upper leaf

cells and margin, X270.

22 FIELDIANA: BOTANY, VOLUME 25

21. FISSIDENS TAXIFOLIUS Hedw., Sp. Muse. 155. 1801.

Medium sized plants, laxly gregarious, dark green. Stems 6-7

mm. high, about 3 mm. wide with leaves. Leaves numerous,

crowded, with strongly circinate tips when dry, widely spreading

when moist, 2 mm. long, oblong-lanceolate, acute or short acuminate,

unbordered; margins serrulate all around; costa pale, short excurrent;

cells small, dense, opaque, rounded-hexagonal, diam. 8-10 n, convex

on free surfaces, one or two rows at margins paler but not forming a

distinct border. Setae lateral from near base of stem, , to 15 mm.

long; capsule inclined, asymmetrical, peristome teeth bright red.

(Fig. 6, E-G.)

Dept. Alta Verapaz: Sharp 2987. Dept. Baja Verapaz: Sharp 2860.

Distribution : Canada and eastern United States south to Florida,

Missouri and Arizona.

Shaded banks and slopes at moderate altitudes. Here the leaves

are more sharply pointed than in most of the United States collec-

tions but otherwise the agreement is close. This is a significant

discovery in line with the occurrence of so many north temperate

types in the highlands of Guatemala. The species has not been

recorded before in North America south of the Mexican border.

22. FISSIDENS POLYPODIOIDES Hedw., Sp. Muse. 154. 1801.

Dioicous; robust, frondose yellowish green plants. Stems simple

or sparingly branched, 2-8 cm. long, 5-7 mm. wide. Leaves numer-

ous, not crowded, curved at tips when dry, oblong-lingulate, abruptly

rounded and bluntly apiculate, entire, not bordered, 4-5 mm. long,

1 mm. wide above; costa percurrent; cells hexagonal, to 20 n long,

smaller toward margins. Setae lateral, about 1 cm. long; capsule

inclined, narrowly pyriform. (Fig. 9, E-G.)

Dept. Huehuetenango : Steyermark 49748. Dept. San Marcos: Steyermark

37261. Dept. Zacapa: Steyermark 29962, 42658, 43306. Dept. Chiquimula:

Steyermark 31020.

Distribution: Southeastern United States, Mexico, West Indies,

Central and South America.

On damp banks, rocks and trees at medium to high altitudes.

Although the apical margins are usually repand and the apex varies

considerably in outline the teeth are not quite as sharp and pro-

nounced as in F. Oerstedianus.

BARTRAM: MOSSES OF GUATEMALA

23

23. FISSIDENS OERSTEDIANUS C. M., Syn. 2: 529. 1851.

Slightly more robust than the preceding species, fronds 8-10

mm. wide. Leaves serrate near apex with sharp, irregular teeth.

Capsule horizontal, subcylindric. (Fig. 10, A-B.)

Dept. Zacapa: Steyermark $318.

Distribution: Costa Rica, Panama.

On moist banks at high altitude. The distinctions between this

species and F. polypodioides are not always as clear as they might

be and I am doubtful if they can be specifically separated.

24. FISSIDENS GRANDIFRONS Brid., Muse. Rec. Suppl. 1 : 170. 1806.

Dull brownish green plants in dense mats. Stems 2-4 cm. long

or longer, rigid, often branched below, densely foliate, fronds 2-3 mm.

wide. Leaves rigidly erect-spreading, about 3 mm. long, unbordered,

linear-lanceolate, bluntly acute, entire, opaque; costa ending in

apex; cells hexagonal, incrassate, in 2 or more layers except at

margins. Setae lateral, capsules erect, fruit rare. (Fig. 10, C-D.)

Dept. Huehuetenango : Steyermark 5000,4, 50005. Dept. San Marcos: Steyer-

mark 36897.

Distribution: Southern Canada, United States, Mexico, Europe,

Asia.

FIGURE 10

A-B, Fissidens Oerstedianus: A, leaf, X8; B, apex of leaf, X54.

C-D, Fissidens grandifrons: C, leaf, X14; D, apex of leaf, X54.

E-G, Fissidens debilis: E, leaf, X8; F, apex of leaf, X54; G, upper leaf cells,

X270.

24 FIELDIANA: BOTANY, VOLUME 25

On wet rocks or submerged in streams in calcareous regions at

high altitudes. These collections seem to represent the southern

limit of distribution in North America.

25. FISSIDENS DEBILIS Schwaegr., Suppl. I2: 11. 1816.

Fissidens julianus Schimp., Flora 21: 271. 1838.

Conomitrium Turckheimi C. M., Bull. Herb. Boiss. 5: 173. 1897.

Slender floating plants, yellowish at tips, dark brown below.

Stems branching, 3 cm. long or often much longer. Leaves distant,

spreading, flexuous when dry, to 5-6 mm. long, linear-lanceolate,

bluntly acute; costa ending well below apex; cells irregularly hexa-

gonal, to 25 p. long, smaller toward margins. Fruit rare, terminal on

short lateral branches, seta shorter than capsule. (Fig. 10, E-G.)

Dept. Jutiapa: Standley 75512.

Distribution: United States, Mexico, South America, Europe,

Africa.

On rocks in stream at moderate altitude. This sterile collection

is one of the smaller forms with stems only 2-3 cm. long. Muller

also cites two collections from Alta Verapaz.

4. DITRICHACEAE

Slender densely caespitose plants. Stems erect, sparingly

branched. Leaves narrowly lanceolate, entire or slightly toothed

near apex; costa percurrent; cells smooth, not differentiated at basal

angles. Seta erect; capsules erect or nodding; peristome simple,

of 16 slender teeth, entire or split nearly to base into 2 filiform forks;

lid conical or beaked; annulus broad; spores small.

1. Capsules immersed, peristome lacking 1. Bryomanginia

Capsules exserted, peristome present 2

2. Leaves 2 ranked, peristome teeth obliquely striolate 2. Distichium

Leaves not 2 ranked, peristome teeth papillose 3

3. Upper leaf cells elongate, capsules smooth 4. Ditrichum

Upper leaf cells quadrate, capsule furrowed when dry 3. Ceratodon

1. BRYOMANGINIA TheY., Rec. de Trav. Crypt. 1. 1931.

Small, densely tufted alpine plants. Leaves linear, concave,

obtuse, entire; costa faint, short; cells oval, smooth, elongate below.

Seta short; capsule small, ovoid, immersed; peristome lacking;

annulus large; lid convex.

BARTRAM: MOSSES OF GUATEMALA

25

1. BRYOMANGINIA SAINT PIERREI Th£r., Rec. de Trav. Crypt. 2. 1931.

Autoicous; small, brittle, reddish brown plants growing in dense,

compact cushions. Stems erect, to 1.5 cm. high. Leaves erect,

1.5 mm. long, linear, deeply concave, obtuse; margins erect, entire;

costa about 50 n wide below, narrower upward, poorly defined, ending

near or above mid-leaf; upper cells oval, incrassate, smooth, oblique,

to 15 n long, 6-8 /* wide, inner basal cells rectangular, pellucid, 12-

15 n wide, to 75 M long, narrower toward margins. Capsule ovoid,

smooth, 0.5-0.6 mm. long, immersed or emergent, on a short, fleshy

seta about 0.5 mm. long; peristome none; annulus large and persis-

tent, about 65 M high, of 2-3 rows of cells; lid convex, mammillate;

spores pale brown, minutely papillose, diam. 25 /x- (Fig. 11, A-E.)

Dept. San Marcos: Sharp 5^23.

Distribution: Mexico.

Non-calcareous boulder near summit of Volcan Tajumulco. A

rare, alpine species previously known only from the type locality on

Nevada de Toluca, Mexico. Superficially the plants are suggestive

of Andreaea but the structural details are of course distinctive.

2. DISTICHIUM Bry. Eur. fasc. 29-30. 1846.

Slender, silky plants in dense tufts. Stems dichotomously

branched, densely tomentose below. Leaves distichous, abruptly

D

FIGURE 11

A-E, Bryomanginia Saint Pierrei: A, plant, XI; B, capsule, X20; C, leaf,

X20; D, upper leaf cells and margin, X270; E, basal leaf cells, X270.

F-H, Ditrichum rufescens: F, plant, Xl; G, leaf, X12; H, capsule, XlO.

26 FIELDIANA: BOTANY, VOLUME 25

narrowed to a spreading, subulate point from an oblong, sheathing

base; costa long excurrent. Seta elongate; capsule suberect; peri-

stome teeth irregularly divided.

1. DISTICHIUM CAPILLACEUM (Hedw.) Bry. Eur. fasc. 29-30. 1846.

Cynontodium capillaceum Hedw., Sp. Muse. 57. 1801.

Plants densely tufted or mixed with other mosses. Stems to

3 cm. or more high. Leaves 4-5 mm. long, in 2 ranks, the slender,

spreading, coarsely papillose point longer than the erect, clasping

base. Costa long excurrent; basal cells linear, gradually becoming

subquadrate above shoulders. Seta slender, to 2 cm. long; capsule

erect, oblong-cylindric; peristome teeth 16, obliquely striolate.

(Fig. 12, A-C.)

Dept. Huehuetenango: Standley 81627, 81672, 83090a.

Distribution: Cosmopolitan in temperate, arctic and antarctic

regions and at high altitudes in the tropics.

On rocks in alpine regions. The widely spreading, papillose leaf

points readily separate this species from any of the local species

of Ditrichum.

3. CERATODON Brid., Bryol. Univ. 1:480. 1826.

Plants densely tufted. Stems erect, closely foliate. Leaves

ovate-lanceolate, contorted when dry; margins recurved; costa short

excurrent; cells smooth, subquadrate, elongate below. Seta erect,

elongate; capsule suberect; peristome teeth split nearly to base.

1. CERATODON STENOCARPUS Bry. Eur. fasc. 29-30. 1846.

Ceratodon vulcanicus C. M., Bull. Herb. Boiss. 5: 191. 1897.

Tufts yellowish above, brown below. Stems to 2 cm. or more long.

Leaves crowded, curved and contorted when dry, 1.5-2 mm. long,

ovate-lanceolate, acuminate; margins recurved nearly to apex,

coarsely toothed near tip; costa subpercurrent; upper cells quadrate,

incrassate, basal cells rectangular. Seta about 2 cm. long, pale

yellow; capsules suberect or often arcuate and inclined, brown, urn

2 mm. long, sulcate when dry; peristome teeth brown, sharply

papillose, divided nearly to base. (Fig. 12, D-F.)

Dept. Huehuetenango: Standley 83092. Dept. San Marcos: Steyermark 35787,

36118. Dept. Totonicapan : Standley 62699a, 8^63, 8^551 . Dept. Quezaltenango :

Steyermark 3^212, 3^625; Standley 67620, 85751 a, 85752. Dept. Chimaltenango :

Standley 61909, 80602.

BARTRAM: MOSSES OF GUATEMALA

27

Distribution: Southern Arizona, Mexico, south along Andes to

Bolivia, also southern Europe, tropical Asia, Africa.

On ledges, rocks and dry banks at high altitudes. These col-

lections are uniformly different from the cosmopolitan C. purpureus

in the pale setae and more erect, paler capsules. Occasionally a

capsule will be nearly horizontal but the great majority are only

slightly inclined to suberect.

4. DITRICHUM (Timm.) Hampe, Flora 50: 181. 1867.

Small tufted plants. Stems erect. Leaves narrowly lanceolate,

subulate-acuminate; costa strong; upper cells linear or oval, basal

cells rectangular, alar cells not differentiated. Seta slender, elongate;

capsules erect or slightly curved; peristome teeth 16, papillose,

divided to base or cleft above.

1. Leaf base oblong, abruptly narrowed at shoulders 1. D. gracile

Leaf base ovate, gradually narrowed upward 2

2. Stems 6-10 cm. long 3

Stems 2-3.5 cm. long 4

3. Leaves 5-8 mm. long 3. D. giganteum

Leaves 3-4 mm. long 2. D. longicaule

U(

c

X8.

H

FIGURE 12

A-C, Distichium capillaceum: A, leaf, X8; B, apex of leaf, XllO; C, capsule,

D-F, Ceratodon stenocarpus: D, plant, XI; E, leaf, X20; F, capsule, X8.

G-I, Ditrichum gracile: G, leaf, X8; H, apex of leaf, XllO; I, upper leaf cells

and margin, X270.

28 FIELD IANA: BOTANY, VOLUME 25

4. Seta 5 mm. long, peristome teeth irregularly cleft, not bifid. 4. D. Steyermarkii

Seta 1-2 cm. long, peristome teeth split to base 5

5. Dioicous, seta to 1 cm. long 5. D. ambiguum

Paroicous, seta to 2 cm. long 6. D. rufescens

1. DITRICHUM GRACILE (Mitt.) Par., Ind. Bryol. ed. 1 : 393. 1895.

Cynontodium gracile Mitt., Jour. Linn. Soc. 12: 43. 1869.

Dioicous; slender, glossy, tawny plants. Stems 2-4 cm. long.

Leaves erect with points often spirally twisted when dry, to 4 mm.

long, abruptly linear-subulate from an oblong, clasping base, toothed

at extreme apex; costa long excurrent; basal cells linear, incrassate,

often very narrow and hyaline toward margins, quickly shorter

toward leaf shoulders, irregularly oval above. Seta about 12 mm.

long; capsule suberect. (Fig. 12, G-I.)

Dept. Solola: Steyermark -47.437, .47503.

Distribution: Mexico, Ecuador.

On exposed rocky summit of Volcan Atitlan. The spiral twisting

of the leaf points is quite obvious. Several species of the southern

hemisphere show the same character but it is not shared by any

other North American species I know of.

2. DITRICHUM LONGICAULE Bartr., Bryol. 49: 109. 1946.

Tall, slender plants in yellowish green tufts. Stems to 10 cm.

long, sparsely radiculose below. Leaves suberect, to 4.25 mm. long,

not crowded, curved and flexuous when dry, rather quickly con-

tracted from an oblong-ovate, concave base to a long, narrowly

linear subula, flat above and coarsely toothed at apex; margins

erect; costa broad below, long excurrent; basal cells linear with

thickened, pellucid walls, much shorter and irregularly oval above.

Sporophyte unknown. (Fig. 13, A-C.)

Dept. San Marcos: Between San Sebastian and summit of Volcan Tajumulco,

alt. 3,800-4,600 m., Steyermark 355 H, TYPE.

Similar in general appearance to robust forms of D. flexicaule

but distinct in the flat, linear leaf subula which is coarsely toothed

at and near the extreme apex. D. crinale (Tayl.) Par. of Ecuador

has longer leaves (6-7 mm.) with longer, finer, setaceous points.

3. DITRICHUM GIGANTEUM Williams, Bull. N. Y. Bot. Card. 2: 113.

1901.

Dioicous; plants in deep, dense tufts, yellowish green above,

brown below. Stems branched, 10 cm. or more long. Leaves

BARTRAM: MOSSES OF GUATEMALA

29

laxly spreading, often falcate, to 7-8 mm. long, narrowly lanceolate,

gradually long acuminate, slightly toothed near apex; costa long

excurrent; basal cells linear with thick, pitted walls, upper cells

oval. (Fig. 13, D-F.)

Dept. Huehuetenango: Standley 83085, 83086, 83087, 83090c, 83091 a.

Distribution: Alaska, northern United States.

In shade of juniperus forest at high altitudes. Although lacking

fruit the identity of these collections is reasonably sure. The gap

in distribution is wide but not without precedent when the alpine

moss flora of Guatemala is considered as a whole.

4. DITRICHUM STEYERMARKII Bartr., Bryol. 49: 110. 1946.

Slender, dull yellowish green plants, densely tufted. Stems 1.5-

2 cm. high, simple or branched above, sparingly radiculose below.

Leaves erect-appressed when dry, erect-spreading when moist, 3.5-

4 mm. long, gradually subulate-acuminate from an oblong, concave

base, sharply serrate at extreme apex; costa broad and indistinct

below, short excurrent; upper leaf cells subquadrate, diam. 8-10 n,

basal cells narrowly rectangular, hyaline. Perichaetial leaves

]

B

Li

H

J

FIGURE 13

A-C, Ditrichum longicaule: A, leaf, X14; B, apex of leaf, XllO; C, upper leaf

cells and margin, X270.

D-F, Ditrichum giganteum: D, leaf, X8; E, apex of leaf, XllO; F, upper leaf

cells and margin, X270.

G-J, Ditrichum Steyermarkii: G, plant, Xl; H, leaf, X14; I, apex of leaf,

XllO; J, part of peristome, XllO.

30 FIELDIANA: BOTANY, VOLUME 25

abruptly narrowed from a clasping base to a linear-subulate point.

Seta short, 5 mm. long; capsule erect, ovoid-cylindrical, barely

exceeding the tips of the perichaetial leaves; urn scarcely 2 mm.

long; operculum red, conic-rostrate, 1 mm. long; peristome teeth 16,

pale, densely papillose, irregularly cleft, not bifid; spores smooth,

diam. 10 M- (Fig. 13, G-J.)

Dept. San Marcos: Along road between San Sebastian at km. 21 and km. 8,

8-18 miles northwest of San Marcos, alt. 2,700-3,800 m., Steyermark 35657, TYPE.

Crevices of banks of dry slope. A highly individual species

characterized by the short setae and irregularly cleft peristome teeth.

5. DITRICHUM AMBIGUUM Best, Bull. Torr. Bot. Club 20 : 117. 1893.

Dioicous; dull yellowish green plants. Stems to 1.5 cm. high.

Leaves crowded, erect with spreading points when dry, 1.5-2 mm.

long, ovate-lanceolate, subulate-acuminate; margins slightly re-

curved, entire or weakly toothed; costa percurrent; cells rectangular,

incrassate. Seta 8-9 mm. long; capsule erect, cylindric; lid conic-

rostrate, nearly 1 mm. long; peristome teeth divided to the short

basal membrane, densely and sharply papillose. (Fig. 14, A-D.)

Dept. Sacatepequez: Standley 6024.5.

Distribution: British Columbia to California.

On dry open bank at moderate altitude. The differences between

this collection and authentic material of D. ambiguum are negligible

and I have little hesitation in referring them here.

6. DITRICHUM RUFESCENS (Hampe) Broth., E. & P. Nat. Pflanzenf.

I3: 300. 1901.

Leptotrichum rufescens Hampe, Linnaea 31: 521. 1862.

Paroicous; antheridia in a bud-like cluster just below the peri-

chaetium. Slender, silky plants, densely tufted, green above, reddish

brown below. Stems erect, to 1 cm. high, sparsely radiculose. Leaves

erect, flexuous, the uppermost to 3.5 mm. long, slenderly subulate-

acuminate from a short, ovate base; margins erect, entire; costa

broad and poorly defined below, excurrent with a few blunt teeth

at apex; cells linear, smooth. Seta to 2 cm. long, pale, becoming

reddish with age; capsule slightly curved, urn to 3 mm. long, small-

mouthed; lid conic-rostrate, 1 mm. long; annulus broad; peristome

teeth erect, pale red, cleft to base into 2 filiform, densely papillose

forks; spores smooth, pale, diam. 10-12 /*. (Fig. 11, F-H.)

BARTRAM: MOSSES OF GUATEMALA

31

Dept. El Quiche: Sharp 5307. Dept. Huehuetenango : Sharp 4879,

4950.

Distribution: Mexico to Colombia, West Indies, Venezuela.

Moist, shaded banks at moderately high altitudes. This species

fruits freely and will be easily recognized by the paroicous inflores-

cence.

5. SELIGERIACEAE

Slender or small mostly rupestral plants. Leaves subulate-

acuminate from a broader base; costa strong, excurrent; cells smooth,

alar group strongly differentiated in Blindia. Seta erect or curved;

capsule pyriform, wide-mouthed; peristome single, of 16 undivided

teeth.

1. BLINDIA Bry. Eur. fasc. 33-36. 1846.

Plants medium sized. Stems branched. Leaves lanceolate; cells

narrow, smooth, incrassate, inflated and colored at basal angles in

a conspicuous group. Seta elongate; capsule turbinate when dry,

peristome teeth 16, smooth, entire; annulus lacking.

1. BLINDIA ACUTA (Hedw.) Bry. Eur. fasc. 33-36. 1846.

Weisia acuta Hedw., Sp. Muse. 71. 1801.

B

H

FIGURE 14

A-D, Ditrichum ambiguum: A, plant, Xl; B, leaf, X14; C, apex of leaf, XllO;

D, part of peristome, XllO.

E-H, Blindia acuta: E, plant, Xl; F, leaf, X14; G, apex of leaf, XllO; H, basal

leaf cells, X270.

I-K, Trematodon longicollis: I, plant, Xl; J, leaf, X14; K, capsule, X8.

32 FIELDIANA: BOTANY, VOLUME 25

Dioicous; plants tufted; stems slender, branched. Leaves sub-

secund, long subulate from a triangular-lanceolate, concave base,

3-3.5 mm. long, entire; costa strong, long excurrent, obscurely

toothed at tip; cells linear, smooth, very incrassate, shorter at ex-

treme base and colored across insertion, alar group large and con-

spicuous, subquadrate, deep brown. Seta 3-10 mm. long; capsule

turbinate when dry and empty. (Fig. 14, E-H.)

Dept. Quezaltenango : Standley 83684.

Distribution: Greenland to Alaska south to northern United

States, Europe, Asia.

On wet sand at high altitude. This collection is sterile but as

far as the vegetative characters are concerned it is certainly good

Blindia acuta.

6. DICRANACEAE

Plants often robust, closely tufted. Stems simple or forked,

densely foliate, tomentose below. Leaves erect or secund, often

crispate, lanceolate; costa single, usually well developed; basal cells

rectangular, smaller and usually subquadrate above, alar group

usually strongly differentiated. Seta mostly elongate, straight or

cygneous; capsules erect or curved, cylindric or ovoid, often plicate;

peristome single, of 16 teeth, cleft to or below middle, usually strio-

late below, papillose above; lid conic-rostrate; calyptra cucullate,

entire or fringed at base.

1. Capsule neck slender and spongy, longer than urn 1. Trematodon

Capsule neck inconspicuous, shorter than urn 2

2. Alar cells clearly differentiated 3

Alar cells not or scarcely differentiated 9

3. Costa broad, }/$ the width of leaf base or more 4

Costa narrow, less than ^ the width of leaf base 7

4. Seta strongly cygneous or flexuous when moist 5

Seta erect and straight 6

5. Upper leaf cells oval or rhomboidal, calyptra usually fringed . . 5. Campylopus

Upper leaf cells linear, calyptra not fringed 8. Dicranodontium

6. Calyptra fringed, peristome teeth undivided 7. Pilopogon

Calyptra not fringed, peristome teeth bifid 6. Atradylocarpus

7. Leaves with a hyaline border 17. Leucoloma

Leaves not bordered 8

8. Peristome teeth papillose, perichaetium conspicuous 15. Holomitrium

Peristome teeth striolate, perichaetium inconspicuous 16. Dicranum

BARTRAM: MOSSES OF GUATEMALA 33

9. Leaf cells mammillose or papillose 10

Leaf cells smooth 11

10. Peristome lacking, leaf cells papillose 10. Amphidium

Peristome present, leaf cells mammillose 12. Oncophorus

11. Costa broad, occupying more than Yi of leaf base 9. Brothera

Costa narrow, less than % the width of leaf base 12

12. Seta stout, strongly curved when moist 4. Campylopodium

Seta slender, erect 13

13. Leaves appressed, male flower conspicuous, discoid 2. Aongstroemia

Leaves spreading, male flower inconspicuous, gemmiform 14

14. Leaves erect-spreading, not crispate 3. Dicranella,

Leaves crispate when dry 15

15. Small, delicate plants, capsule 8 ribbed, seta short 11. Rhabdoweisia

Larger plants, capsule smooth, seta elongate 16

16. Leaf base obovate, sheathing, abruptly narrowed to blade. .13. Symblepharis

Leaf base oblong, gradually narrowed upward 14. Dicrarioweisia

1. TREMATODON Michx., Fl. Bor. Amer. 2: 289. 1803.

Small gregarious plants. Leaves slenderly pointed from an ovate,

concave base; costa ending below apex; cells smooth. Seta elongate;

capsule curved, with a neck about twice as long as urn; peristome

single, of 16 teeth.

1. TREMATODON LONGICOLLIS Michx., Fl. Bor. Amer. 2: 289. 1803.

Trematodon reflexus C. M., Syn. 1 : 459. 1848.

Autoicous: stems 2-3 mm. high. Leaves spreading, flexuous,

abruptly linear from an ovate, concave base, blunt and toothed at

apex; costa stout, ending under apex; cells rectangular, more elon-

gate below. Seta yellow, 1-3 cm. long; capsule curved, cylindric,

urn 2 mm. long, neck often longer than urn and strumose at base;

peristome teeth reddish brown from a low basal membrane, verti-

cally striolate; annulus broad; lid long beaked; spores about 20/z in

diam. (Fig. 14, I-K.)

Distribution: Eastern United States, Mexico, Cuba, South

America, Europe, Philippines, New Guinea, Ceylon.

This widely distributed species has been recorded from Guate-

mala but I have seen no local collections and it does not appear in

any of Standley's or Steyermark's collections.

34

FIELDIANA: BOTANY, VOLUME 25

u

D

t

H

FIGURE 15

A-C, Aongstroemia jamaicensis: A, fertile plant, Xl; B, male plant, Xl;

C, leaf, X 14.

D-F, Aongstroemia orientalis: D, plants, Xl; E and F, leaves, X27.

G-I, Dicranella vaginata: G, plant, Xl; H, leaf, X14; I, capsule, X8.

2. AONGSTROEMIA Bry. Eur. fasc. 33-36. 1846.

Stems short to slender and elongate, julaceous. Leaves ap-

pressed, ovate; costa strong; cells rather elongate and smooth. Seta

smooth, erect; capsule erect, •ovoid-cylindric; peristome teeth when

present inserted below rim, often split or perforate.

Plants 4-6 cm. high, leaves acuminate 1. A. jamaicensis

Plants less than 1 cm. high, leaves obtuse 2. A. orientalis

1. AONGSTROEMIA JAMAICENSIS C. M., Bull. Herb. Boiss. 5: 554.

1897.

Dioicous; antheridial buds conspicuous. Plants glossy; stems

mostly simple, laxly tufted, to 6-7 cm. high. Leaves appressed,

abruptly subulate from an oblong-ovate, clasping base, to 4-5 mm.

long; margins erect; costa long excurrent in a slender, smooth awn;

lowest cells rectangular, becoming linear- vermicular upward. Seta

3-4 mm. long; capsule cylindric, urn 1.5-2 mm. long, brown, exceeded

by tips of perichaetial leaves; peristome teeth reddish, narrow,

papillose, forked about half way down, perforate below. (Fig. 15,

A-C.)

Dept. San Marcos: Steyermark 35951, 3 61*1*1 a, 36^2; Standley 864.20. Dept.

Totonicapan: Standley 65920, 659^1. Dept. Quezaltenango : Standley 6807 It,

8^926.

BARTRAM: MOSSES OF GUATEMALA 35

Distribution: Jamaica, Mexico, Costa Rica.

Damp banks at high altitudes. The delicate, filiform, glossy

stems of this species are distinctive and not likely to be confused

with anything else.

2. AONGSTROEMIA ORIENTALIS Mitt., Trans. Linn. Soc. 2: 154.

1891.

Small, slender, yellowish green plants, closely gregarious. Stems

simple or little branched, less than 1 cm. long. Leaves minute,

appressed with secund points, 0.5-0.8 mm. long, ovate, obtuse;

margins erose-denticulate nearly to base; costa ending below apex;

cells oval-rhomboidal, smooth, incrassate, elongate below. Seta

8-10 mm. long; capsule erect; peristome lacking. (Fig. 15, D-F.)

Dept. Huehuetenango : Steyermark 50228. Dept. San Marcos: Steyermark

36090, 361 18a.

Distribution: Mexico, Himalayas, Burma, Yunnan, Philippine

Islands.

Dry slopes at high altitudes. This curious little moss has been

found in fruit in Mexico but the local collections are sterile.

3. DICRANELLA Schimp., Coroll. Bry. Eur. fasc. 13. 1855.

Small terrestrial plants growing in tufts or mats. Leaves spread-

ing, narrowly lanceolate; costa stout; cells smooth, alar group not

differentiated. Seta slender, erect; capsules erect or inclined;

peristome of 16 reddish teeth usually cleft to about middle; lid conic-

rostrate, oblique.

1. Leaves squarrose-spreading, from an erect, obovate, sheathing base

1. D. vaginata

Leaves erect-spreading from insertion 2

2. Capsules cernuous, asymmetrical 4. D. varia

Capsules erect, symmetrical 3

3. Peristome teeth irregularly papillose on outer surface 4

Peristome teeth striolate on outer surface 5

4. Peristome teeth 225-250 n high, leaves abruptly narrowed from an oblong

base 6. D. Sharpii

Peristome teeth 100 n high, leaves ovate-lanceolate 5. D. alpina

5. Seta 10-15 mm. long, peristome 200-250 M high 6

Seta 5-6 mm. long, peristome under 125 /* high 7

6. Seta reddish, capsules contracted under mouth when dry. . .3. D. subinclinata

Seta yellowish, capsules not contracted under mouth 2. D. Hilariana

7. Blade of inner perichaetial leaves as long as basal part 7. D. lagunaria

Blade of inner perichaetial leaves twice as long as basal part

8. D. brachyblepharis

36 FIELDIANA: BOTANY, VOLUME 25

1. DICRANELLA VAGINATA (Hook.) Card., La Flore Bryol. d. Ter.

Mag. etc. 60. 1908.

Dicranum vaginatum Hook., Muse. Exot. pi. 141. 1820.

Dicranella Standleyi Bartr., Contrib. U. S. Nat. Herb. 263: 57. 1928.

Plants laxly gregarious, slender, 1-4 cm. high. Upper leaves

3-4 mm. long from an obovate, erect, clasping base abruptly nar-

rowed to a spreading setaceous point; costa percurrent; basal cells

rectangular, shorter and irregular at shoulders, subquadrate and

slightly incrassate above. Seta erect, 10-12 mm. long; capsules

erect or nodding; peristome teeth reddish brown, densely papillose,

divided to below middle; lid long subulate-rostrate. (Fig. 15, G-I.)

Dept. San Marcos: Standley 66247. Dept. Jutiapa: Steyermark 31938.

Distribution: Mexico, Costa Rica, Colombia, Ecuador.

On damp banks at high altitudes. It seems more logical to

confine Aongstroemia to the species with erect, appressed leaves and

include Aongstroemia vaginata (Hook.) Card, in Dicranella where

it belongs from every point of view.

2. DICRANELLA HILARIANA (Mont.) Mitt., Journ. Linn. Soc. 12:

31. 1869.

Dicranum Hilarianum Mont., Ann. Sci. Nat. II. 12: 52. 1839.

Small, laxly tufted, pale green plants. Leaves spreading, to

2 mm. long, narrowly lanceolate, gradually narrowed to a blunt,

toothed apex; costa stout, ending below apex; upper cells short

rectangular, more elongate below. Seta about 10 mm. long; capsule

erect; peristome teeth vertically striolate below, to 200 ^ long,

divided to below middle. (Fig. 17, A-D.)

Dept. Quezaltenango : Standley 653^8.

Distribution: Southern United States, Mexico, West Indies,

Central and South America.

The above collection is sterile and doubtful but the species

should eventually be found in Guatemala, which is well within its

geographical range.

3. DICRANELLA SUBINCLINATA Lor., Moosst. 160. 1864.

Slender, yellowish green plants, densely tufted. Stems about

1 cm. high, sparsely branched. Leaves erect with slightly contorted

points when dry, more rigid when moist, gradually linear-lanceolate

from an ovate base, to 2 mm. long, bluntly rounded and toothed

BARTRAM: MOSSES OF GUATEMALA

37

at apex; costa ending just below apex; upper cells rectangular with

firm, pellucid walls, gradually becoming linear toward base. Seta

to 8 or 10 mm. long, reddish; capsule dark brown, contracted below

mouth when dry; peristome as in D. Hilariana. (Fig. 16, A-C.)

Dept. Baja Verapaz: Sharp 2948.

Distribution: Mexico, Central America, West Indies.

On bank at moderate altitude. Readily separated from D.

Hilariana by the stouter, reddish setae and the capsules contracted

below the mouth when dry.

4. DICRANELLA VARIA (Hedw.) Schimp., Coroll. Bry. Eur. 13.

1855.

Dicranum varium Hedw., Sp. Muse. 133. 1801.

Small, slender, densely tufted plants, brownish green. Stems to

4 or 5 mm. high. Leaves erect when dry, erect-spreading or slightly

secund when moist, the upper to 1.5 mm. long, smaller below, tri-

angular-lanceolate, short acuminate; margins recurved below, den-

ticulate near apex; costa percurrent; cells linear. Seta 7-8 mm.

long, reddish; capsule nodding, curved and asymmetrical, urn about

1 mm. long, wide-mouthed; peristome relatively large, teeth reddish,

f\

G

FIGURE 16

A-C, Dicranella subinclinata: A, plant, Xl; B, leaf, X14; C, apex of leaf,

X110.

D-F, Dicranella varia: D, plant, Xl; E, leaf, X20; F, capsule, X12.

Cr-J, Dicranella Sharpii: G, plant, Xl; H, leaf, X40; I, capsule, X8; J, part

of peristome, XllO.

38 FIELDIANA: BOTANY, VOLUME 25

250-300 M high, cleft half way down, striolate; annulus lacking; lid

short rostrate. (Fig. 16, D-F.)

Dept. Baja Verapaz: Sharp 2948.

Distribution: New Brunswick to Alaska south to Mexico,

Florida and Cuba.

On moist bank at moderate altitude. Several collections have

been recorded from Mexico and Cuba but the Guatemalan speci-

mens extend the range appreciably to the southward.

5. DICRANELLA ALPINA (C. M.) Par., Ind. Bryol. Suppl. 115. 1900.

Angstroemia alpina C. M., Bull. Herb. Boiss. 5: 186. 1897.

Plants similar to the following species. Seta elongate; capsule

erect; peristome teeth about 0.1 mm. high, papillose (not striolate).

Nagula, alt. 9,000 ft., Bernoulli & Cario 6^. Dept. Alta Verapaz: Turckheim

6653.

Distribution: Mexico.

6. DICRANELLA SHARPII Bartr., Bryol. 50: 202. 1947.

Slender, tufted, yellowish plants. Stems erect, 6-7 mm. high.

Leaves erect, minute below, the upper to 4 mm. long, abruptly

lanceolate-subulate from an oblong base about 1 mm. long, sharply

acute; margins entire; costa well defined excurrent; upper leaf cells

very narrow, gradually becoming rectangular below with firm lateral

walls. Seta 7 mm. long, yellowish; capsule erect, oblong, brownish,

urn 1.5 mm. long, slightly angulate when dry; peristome teeth 225-

250 n high, irregularly cleft to about middle, papillose, -not striolate;

lid obliquely rostrate; annulus broad; spores papillose, diam. 16-18 p..

(Fig. 16, G-J.)

Dept. El Quiche: Sharp 2^8. Dept. Baja Verapaz: Sharp 5188.

Endemic.

On banks at moderate altitudes. This species seems to be clearly

distinct from both D. barbensis Ren. & Card, and D. alpina C. M.

in the much longer peristome teeth and the leaves abruptly narrowed

above the oblong base to an almost setaceous point with the costa

plainly excurrent. The length of the peristome varies somewhat

with the size of the capsules but even in the smaller forms the teeth

are over 200 M high.

BARTRAM: MOSSES OF GUATEMALA 39

7. DICRANELLA LAGUNARIA (C. M.) Broth., E. & P. Pflanzenf. I3:

309. 1901.

Aongstroemia lagunaria C. M., Bull. Herb. Boiss. 5: 187. 1897.

Small plants. Stems to 8 mm. high. Leaves crowded, erect-

spreading, ovate-lanceolate, bluntly acute; costa ending below apex;

cells slightly incrassate, rectangular, more elongate below. Seta

5-6 mm. long; capsule erect; peristome teeth about 0.1 mm. long,

striolate below, entire or irregularly cleft.

Laguna del Pino, Bernoulli & Cario 116.

Endemic.

Known only from the type collection.

8. DICRANELLA BRACHYBLEPHARIS (C. M.) Mitt., Journ. Linn. Soc.

12: 34. 1869.

Aongstroemia brachyblepharis C. M., Syn. 1 : 435. 1848.

Small plants to 1 cm. high. Leaves distant, ovate-lanceolate;

costa subpercurrent; cells rectangular. Perichaetial leaves to 4 mm.

long, gradually narrowed to a point twice as long as the ovate base;

seta 5 mm. long; capsule erect; peristome as in preceding species.

(Fig. 17, E-H.)

Distribution: Mexico, Jamaica.

I have seen no local specimens of either this species or D. lagunaria.

4. CAMPYLOPODIUM (C. M.) Besch., Ann.

Sci. Nat. V. 18:189. 1873.

Angstroemia Sect. Campylopodium C. M., Syn. 1 : 429. 1848.

Small plants similar in habit and appearance to Dicranella but

differing markedly in the stout, curved or cygneous setae. Capsules

ribbed when dry; peristome teeth striolate, cleft about half way down.

1. CAMPYLOPODIUM PUSILLUM (Schimp.) Williams, No. Amer. Flora

152:94. 1913.

Campylopus pusillus Schimp., Mem. Soc. Sci. Nat. Cherb. 16: 165. 1872.

Dicranum magnirete C. M., Bull. Herb. Boiss. 5: 186. 1897.

Dicranum Turckheimii C. M., Bull. Herb. Boiss. 5: 186. 1897.

Laxly gregarious, yellowish green plants. Stems to 1 cm. high.

Leaves spreading, flexuous, to 4 mm. long, abruptly narrowed from

40

FIELDIANA: BOTANY, VOLUME 25

FIGURE 17

A-D, Dicranella Hilariana: A, plant, Xl; B, leaf, X14; C, apex of leaf, XllO;

D, part of peristome, X134.

E-H, Dicranella brachyblepharis: E, plant, Xl; F, part of peristome, X134;

G, stem leaf, X14; H, perichaetial leaf, X14.

I-K, Campylopodium pusillum: I, plant, Xl; J, leaf, X14; K, capsule, X14.

a short, ovate, clasping base to a long subulate point; costa stout,

excurrent, basal cells rectangular, alar cells not differentiated, shorter

and irregular at shoulders, linear above in the. narrow blade. Seta

stout, brown, to 6 mm. long, flexuous when dry, cygneous when

moist; capsule oval; lid obliquely rostrate; spores coarsely papillose,

diam. 20-24 /x. (Fig. 17, I-K.)

Dept. San Marcos: Standley 8651 5a.

Distribution: Mexico, Jamaica, South America.

On tree in wet forest at rather high altitude. The short, broad

leaf base without any differentiated alar cells, abruptly narrowed to

a long, subulate point will distinguish this species from Campylopus.

5. CAMPYLOPUS Brid., Muse. Rec. Suppl. 4: 71. 1819.

Dioicous; small to robust plants, densely tufted. Stems often

branching, radiculose below. Leaves erect or curved, ovate-lanceo-

late, slenderly acuminate, channelled above; margins usually toothed

above; costa very broad below, percurrent or excurrent, often ribbed

on back; basal cells narrow, alar group enlarged, hyaline or colored,

usually conspicuous, upper cells mostly rhomboidal to short rec-

tangular. Seta usually strongly cygneous when moist; capsules

BARTRAM: MOSSES OF GUATEMALA 41

ovoid, usually ribbed when dry; peristome teeth divided about half

way down, striolate below; lid rostrate; calyptra cucullate, generally

fringed at base.

The species of this difficult genus may be separated into three

groups based on the structure of the costa in cross section.

Costa without stereid bands (Pseudocampylopus) : C. guate-

malensis.

Costa with stereid band on dorsal side only (Eucampylopus) :

C. Chrismari, C. flexuosus, C. fragilis, C. concolor, C. Jamesoni,

C. introflexus.

Costa with stereid bands on both sides of median guide row

(Palinocraspis): C. savannarum, C. filifolius, C. arctocarpus,

C. Richardi.

1. Marginal cells of leaf base short, quadrate, chlorophyllose . .8. C. savannarum

Marginal cells of leaf base elongate, hyaline 2

2. Leaves generally with hyaline tips 3

Leaves with concolorous tips 4

3. Costa strongly ridged on back, basal cells hyaline, thin walled . . 7. C. introflexus

Costa smooth or faintly ridged on back, basal cells incrassate, porose

11. C. Richardi

4. Cells of leaf base with firm, pellucid walls 5

Cells of leaf base lax, with thin, delicate walls 7

5. Basal cells more or less pitted, costa with stereid bands on both sides of

median guide row 6

Basal leaf cells not pitted, costa with dorsal stereid band only, ventral cells

large 3. C. flexuosus

6. Leaves in interrupted tufts, apex slender 9. C. filifolius

Stems equally foliate, apex of leaf short and stout 10. C. arctocarpus

7. Leaf base distinctly bordered to shoulders with 6-10 rows of linear, hyaline

cells 2. C. Chrismari

Leaf base not distinctly bordered 8

8. Alar cells not differentiated 4. C. fragilis

Alar cells strongly differentiated 9

9. Leaves 4-4.5 mm. long, entire except at extreme apex, costa without stereids

1. C. guatemalensis

Leaves 10 mm. or more long, costa with dorsal stereid band 10

10. Costa less than 1 mm. wide 5. C. concolor

Costa over 1 mm. wide 6. C. Jamesoni

1. CAMPYLOPUS GUATEMALENSIS Bartr., Bryol. 49: 110. 1946.

Slender, compactly tufted, pale green plants, slightly glossy.

Stems branched, tomentose nearly to tips, 2-6 or 7 cm. high. Leaves

42

FIELDIANA: BOTANY, VOLUME 25

FIGURE 18

A-E, Campylopus guatemalensis: A, part of plant, Xl; B, leaf, X14; C, apex

of leaf, XllO; D, calyptra, X14; E, part of cross section of costa, X270.

F-J, Campylopus Chrismari: F, part of plant, Xl; G, leaf, X8; H, apex of

leaf, XllO; I, calyptra, X14; J, part of cross section of costa, X270.

erect-spreading, 4-6 mm. long, narrowly lanceolate, tubulose above,

entire except for a few teeth at extreme apex; costa at least % the

width of leaf base, long excurrent, without stereids; basal cells

rectangular, thin walled, alar group conspicuous, hyaline or brownish,

inflated and auriculate, upper cells small, rhomboidal. Seta 5 mm.

long, bent near middle or strongly sinuous; capsule elliptic, urn

1.5 mm. long; calyptra fringed at base. (Fig. 18, A-E.)

Dept. Totonicapan: Standley 83 101 a. Dept. Quezaltenango : Standley 86121,

86125, 86137, 86159 TYPE, 86182, 86186; Steyermark 3^656.

Endemic.

On rocks in alpine regions. Although near C. Schimperi Milde

even in structural details, I feel that these plants are best treated

as a distinct species. The longer leaves, in some cases reaching 5-6

mm., more spreading both moist and dry, are distinctive features.

It will be separated from C. Chrismari by the unbordered leaf base,

the more conspicuous alar cells and the calyptra fringed at the base.

2. CAMPYLOPUS CHRISMARI (C. M.) Mitt., Journ. Linn. Soc. 12:

88. 1869.

Dicranum Chrismari C. M., Bot. Zeit. 13: 761. 1855.

BARTRAM: MOSSES OF GUATEMALA 43

Slender, silky, glossy, yellowish green plants growing in compact

tufts. Stems branched, to 6-7 cm. high. Leaves spreading, often

secund, to 6-7 mm. long, from a short, ovate base tapering gradually

to a long, tubulose, setaceous point, denticulate only at extreme

apex; costa long excurrent, with a weak stereid band on the dorsal

side only; basal cells rectangular, thin walled, very narrow and

elongate toward margins forming a wide, distinct hyaline border,

enlarged alar cells few, inconspicuous, upper cells irregularly rhom-

boidal, longer than wide. Seta 12-15 mm. long, cygneous; capsule

narrowly ovoid; calyptra not fringed at base. (Fig. 18, F-J.)

Dept. San Marcos: Steyermark 36101*. Dept. Totonicapan: Standley 62688,

62706a, 62707, 62729, 65869, 81*1*17, 81*502, 84516, 81*562a. Dept. Quezaltenango :

Standley 67691, 67771*, 671*1*9, 85868, 85890; Steyermark 31*182, 31*81*6. Dept.

Chimaltenango: Standley 6181*l*a, 6181*7.

Distribution: Mexico, Costa Rica.

On banks, rocks and trees in alpine regions. The scattered stereid

cells on the dorsal side of the costa indicate that this species should

be included in the Sec. Eucampylopus. In well developed plants the

wide border of narrow cells extending nearly to the top of the leaf

base is a reliable diagnostic character.

3. CAMPYLOPUS FLEXUOSUS (Hedw.) Brid., Muse. Rec. Suppl. 4:

71. 1819.

Dicranum flexuosum Hedw., Sp. Muse. 145. 1801.

Campylopus gracilicaulis Mitt., Journ. Linn. Soc. 12: 83. 1869.

Dicranum Hellerianus Hampe, Verb. Zool.-Bot. Ges. Wien 19: 507. 1869.

Campylopus tallulensis S. & L. Sull., Ic. Muse. 27. 1872.

Dicranum Donnellii Aust., Bot. Gaz. 4: 150. 1879.

Dicranum subleucogaster C. M., Bull. Torr. Bot. Club 5: 49. 1879.

Campylopus Sargii Roll., Bull. Soc. Bot. Belg. 38': 8. 1900.

Campylopus Roellii Ren. & Card., Bull. Soc. Bot. Belg. 381: 9. 1900.

Campylopus straminifolius Bartr., Contr. U. S. Nat. Mus. 263: 63. 1928.

Campylopus hondurensis Bartr., Field Mus. Nat. Hist. Bot. Ser. 49: 351.

1929.

Variable plants; stems 1-6 or 7 cm. high, often with clusters of

microphyllous branchlets near tips. Leaves rigid or flexuous when

dry, oblong-lanceolate, gradually narrowed to a channeled, subulate

point, serrulate toward apex; costa mostly excurrent; basal cells

short rectangular toward costa, narrower toward margins, gradually

or quickly becoming smaller and subquadrate upward, with firm,

pellucid, unpitted walls, alar cells usually inflated and auriculate,

44

FIELDIANA: BOTANY, VOLUME 25

FIGURE 19

A-D, Campylopus flexuosus: A, plant, Xl; B, leaf, X8; C, apex of leaf, X134;

D, one side of leaf base, X134.

E-F, Campylopus fragilis: E, leaf, XlO; F, one side of leaf base, X134.

hyaline or colored, upper cells short rhomboidal. Seta 8-10 mm.

long, strongly curved or cygneous; capsule ovoid, ribbed; calyptra

fringed at base. (Fig. 19, A-D.)

Dept. Alta Verapaz: Standley 71051, 9062, 90784, 92068. Dept. San Marcos:

Standley 68592, 68613. Dept. Totonicapan: Standley 6266 4. Dept. Quezalten-

ango: Steyermark 33471; Standley 65334, 67371, 67417 (distr. as C. Chrismari),

67^27 (as C. Chrismari), 67429 (as C. Chrismari), 67460 (as C. Chrismari), 83321,

83376, 83386, 85913, 85916, 85964, 86013, 86033, 86044 (as C. Chrismari), 86048,

86051. Dept. Chimaltenango : Standley 61910 (as C. Chrismari). Dept. Guate-

mala: Standley 58424, 80620, 80728. Dept. Zacapa: Steyermark 42660. Dept.

Chiquimula: Steyermark 30608,

Distribution: Southern United States, Mexico, Central America,

Europe.

On damp banks, trees and logs at medium to high altitudes.

C. flexuosus is well marked in a broad way by the rectangular cells

of the leaf base with firm unpitted, pellucid lateral walls, appreciably

larger toward costa and gradually narrower toward margins. It is

an exceedingly variable species and many closely related forms have

been described from tropical and subtropical North America based

on more or less trivial and inconstant characters which to my mind

are not amenable to any orderly or practical classification. These

rectangular basal cells change gradually to the small, rhomboidal

cells of the upper leaf blade but the gradation is so irregular in plants

of the same tuft or even on the same stem that I doubt if it can be

BARTRAM: MOSSES OF GUATEMALA 45

used as a specific indicator. For this reason it seems as though

C. gracilicaulis Mitt, naturally falls into the same concept. ,The

robust forms with tall stems and broader costa include C. Roellii

and C. Hellerianus while at the other extreme small plants about

1 cm. high with the costa only 150 M wide or less seem to be insepar-

able from C. Sargii.

4. CAMPYLOPUS FRAGILIS (Turn.) Bry. Eur. fasc. 41. 1847.

Dicranum flexuosum fragile Turn., Muse. Hib. 74. 1804.

Rather small, densely tufted, yellowish green plants. Stems 1-4

cm. high, densely foliate. Leaves suberect and slightly flexuous

when dry, narrowly lanceolate from a pale, oblong base, serrulate

toward apex; basal cells rectangular, thin walled, hyaline, narrower

toward margins and shorter and subrhomboidal toward leaf shoulders,

upper cells short rhomboidal, differentiated alar cells few or none,

never auriculate. Seta 5-8 mm. long; calyptra fringed. (Fig. 19,

E-F.)

Dept. Alta Verapaz: Standley 69112 (as C. Chrismari). Dept. Huehuetenango :

Standley 65 8M (as C. Chrismari); Steyermark 50188. Dept. San Marcos: Steyer-

mark 36391. Dept. Totonicapan: Standley 8^107. Dept. Chimaltenango: Standley

58745a.

Distribution: Florida, Jamaica, Europe, Asia, Africa.

On damp banks and trees at medium to high altitudes. These

collections are all sterile but the typical leaf base with few or no

enlarged alar cells confirms the determination with reasonable

certainty.

5. CAMPYLOPUS CONCOLOR (Hook.) Brid., Bryol. Univ. 1 : 476. 1826.

Dicranum concolor Hook., Muse. Exot. tab. 138. 1820.

Robust pale green plants, slightly lustrous. Stems to 6 cm. high,

uniformly and densely foliate. Leaves spreading, with long flexuous

or secund setaceous points, gradually narrowed from a slender, con-

cave base, 10-14 mm. long, serrulate for some distance below apex;

costa excurrent, to 0.8 mm. wide below, stereids on dorsal side only;

basal cells rectangular, thin walled, quickly changing to the small,

irregular, subquadrate cells of the upper lamina which is only 1 or 2

cells wide for some distance down, enlarged alar cells auriculate, pale

or hyaline. Seta short; capsules asymmetrical; calyptra fringed.

(Fig. 20, A-D.)

Dept. San Marcos: Standley 86229, 86300, 86391; Steyermark 36799. Dept.

Quezaltenango : Steyermark 34325; Standley 85675.

46

FIELDIANA: BOTANY, VOLUME 25

D

H

FIGURE 20

A-D, Campylopus concolor: A, leaf, X6; B, apex of leaf, XllO; C, basal leaf

cells next costa, X134; D, upper leaf cells and margin, X270.

E, Campylopus Jamesoni: E, leaf, X6.

F-I, Campylopus introflexus: F, plant, XI; G, leaf, X8; H, upper leaf cells and

margin, X270; I, part of cross section of costa, X270.

Distribution: Venezuela, Colombia, Peru.

On damp banks at high altitudes. Although uniformly sterile

these notable collections agree in all essential particulars with

authentic material of C. concolor from northern South America.

This seems to be the first record of the species in North America.

6. CAMPYLOPUS JAMESONI (Hook.) Jaeg., Adumb. 1: 126. 1874.

Dicranum Jamesoni Hook., Ic. PI. r. t. 179. 1841.

Campylopus Standleyi Bartr., Contr. U. S. Nat. Mus. 263: 57. 1928.

More robust than C. concolor. Leaves 10-20 mm. long; costa

1-1.6 mm. wide below; auriculate alar cells smaller, more numerous

and more deeply colored. Seta 12-14 mm. long; capsules asymmet-

rical, curved. (Fig. 20, E.)

Dept. Huehuetenango : Steyermark 501 88a.

Distribution: Costa Rica, Colombia.

Limestone bluff of Caxin, summit Sierra de las Cuchumatanes,

3,700 m. The local record is based on a fragmentary stem in poor

condition but enough to establish the species in the local flora. When

TheYiot's notes on C. concolor and C. Jamesoni were published

BARTRAM: MOSSES OF GUATEMALA 47

(Archiv. d. Bot. 2: 185. 1928) neither species had been recorded

from North America.

7. CAMPYLOPUS INTROFLEXUS (Hedw.) Brid., Bryol. Univ. 1: 472.

1826.

Dicranum introflexum Hedw., Sp. Muse. 147. 1801.

Densely tufted plants; stems to 4 cm. or more high. Leaves

5 mm. or more long, laxly appressed when dry, oblong-lanceolate,

subulate pointed, subtubulose above, ending in a hyaline, toothed

point; costa excurrent, broad, with numerous serrated ridges 2-6

cells high on back, stereid band on dorsal side only; basal cells

narrowly rectangular, alar group inconspicuous, upper cells obliquely

rhomboidal. Setae often aggregated, 6-9 mm. long, scabrous near

tips; capsule ovoid, rugose at base; calyptra fringed. (Fig. 20, F-I.)

Dept. Huehuetenango: Siandley 65838, 6581*6, 82295, 82308, 821*32. Dept.

San Marcos: Standley 68535. Dept. Totonicapan: Standley 81*103. Dept. Quezal-

tenango: Steyermark 33188, 31*831*; Standley 65563, 661*75, 81*232, 81*735, 81*71*0.

Dept. Sacatepequez: Siandley 58816, 61229. Dept. Solola: Steyermark 1*6911,

1*71*1*7, 1*71*58 (as C. Richardi), 1*71*78, 1*71*59 (as C. Richardi), 1*71*95; Standley

6231*9. Dept. Guatemala: Standley 58360, 80600, 80603, 80732. Dept. Jutiapa:

Standley 75591*. Dept. Jalapa: Steyermark 32163, 32602.

Distribution: Wide in Europe, North America, South America,

New Zealand, Pacific Islands.

On banks and rocks at medium to high altitudes. An abundant

species of broad distribution and exceedingly variable. The hyaline

basal cells, inconspicuous alar cells, the typical Eucampylopus

costal structure and the high dorsal ridges will help to separate it

from C. Richardi.

8. CAMPYLOPUS SAVANNARUM (C. M.) Mitt., Journ. Linn. Soc.

12: 85. 1869.

Dicranum savannarum C. M., Syn. 2: 596. 1851.

Robust yellowish green plants; stems to 5 cm. long or longer,

often branched, densely tomentose. Leaves crowded, 4-6 mm. long,

tubulose above, oblong-lanceolate, acuminate, sharply serrate toward

apex; costa with two stereid bands, excurrent, concolorous or hyaline

at tip; basal cells short rectangular, incrassate, smaller and sub-

quadrate at margins, upper cells oval-rhomboidal. Fruit not seen.

(Fig. 21, A-F.)

Dept. San Marcos: Steyermark 37135. Dept. Quezaltenango : Steyermark

33672, SS673a, 31*308. Dept. Jalapa: Standley 76731*. Dept. Chiquimula: Steyer-

mark 30599.

48

FIELDIANA: BOTANY, VOLUME 25

H

n

FIGURE 21

A-F, Campylopus savannarum: A and B, plants, Xl; C, leaf, X8; D, basal

margin of leaf, X270; E, apex of leaf, X54; F, part of cross section of costa, X270.

G-I, Campylopus filifolius: G, plant, Xl; H, leaf, X14; I, apex of leaf, X270.

Distribution: Costa Rica, British Guiana, Dutch Guiana.

On trees, rocks and banks at medium altitudes. The quadrate

or even transversely elongate marginal cells of the leaf base are very

distinctive. When I described C. Bartletti from British Honduras

C. savannarum was not known from North America, but I am very

doubtful if the Honduran plant can be maintained as a distinct

species. The hyaline leaf tip is variable, often short or lacking and

again well developed.

9. CAMPYLOPUS FILIFOLIUS (Hornsch.) Mitt., Journ. Linn. Soc. 12:

76. 1869.

Dicranum filifolium Hornsch., Fl. Bras. I2: 12. 1840.

Slender plants to 4 or 5 cm. long. Leaves in interrupted tufts,

curved when dry, 4-6 mm. long, from a short, narrowly ovate base

gradually narrowed to a long setaceous point, serrulate for some

distance below apex; costa excurrent, with two stereid bands, lamina

very narrow above; basal cells rectangular, incrassate, alar cells

conspicuous, reddish, slightly auriculate, upper cells rhomboidal.

Seta 10-15 mm. long, cygneous when moist; capsule oblong, furrowed;

calyptra fringed. (Fig. 21, G-I.)

Dept. Huehuetenango : Steyermark 49734.

Distribution: Costa Rica, Brazil.

BARTRAM: MOSSES OF GUATEMALA

49

On log at medium altitude. The interruptedly foliate stems and

the slender setaceous pointed leaves with concolorous tips will

separate this species from any of the local Palinocraspis group.

10. CAMPYLOPUS ARCTOCARPUS (Hornsch.) Mitt., Journ. Linn. Soc.

12: 75. 1869.

Dicranum arctocarpum Hornsch., Fl. Bras. 1 : 12. 1840.

Densely tufted plants; stems uniformly foliate, radiculose nearly

to apex. Leaves crowded, curved or flexuous when dry, 4 mm. or

more long, oblong-lanceolate, gradually narrowed to a stout, denticu-

late point, channelled above; costa short excurrent, with two stereid

bands; basal cells rectangular, incrassate, pitted, alar group reddish,

conspicuous, extending to costa, upper cells rhomboidal. Seta 6-7

mm. long; capsule oblong; calyptra fringed. (Fig. 22, A-C.)

Dept. Alta Verapaz: Standley 71298, 92300. Dept. Totonicapan: Slandley

65887 (as C. Richardi). Dept. Quezaltenango : Standley 83^21. Dept. Jalapa:

Steyermark 321^89, 32J^93a.

Distribution: Jamaica, South America.

On damp banks, trees and logs at moderate altitudes. These

collections are sterile and not well developed. The costal structure

is definitely of the Palinocraspis type and the leaves all with stout,

concolorous tips so it seems fairly certain that they belong here.

H

A-C, Campylopus arctocarpus: A, plant, Xl;>B, leaf, X14; C, apex of leaf,

X110.

D-H, Campylopus Richardi: D, part of plant, Xl; E, leaf, X8; F, basal leaf

cells, X270; G, median leaf cells, X270; H, part of cross section of costa, X270.

50 FIELDIANA: BOTANY, VOLUME 25

11. CAMPYLOPUS RICHARDI Brid., Muse. Rec. Suppl. 4: 73. 1819.

Medium sized to robust plants, yellowish green at tips, dark brown

below. Leaves erect, nearly straight when dry, crowded, in comal

tufts on the fertile stems, to 6 mm. or more long, oblong-lanceolate,

usually with a distinct hyaline, toothed point; costa lightly ribbed

on back, with two stereid bands, excurrent; basal cells linear or

rectangular with pitted, incrassate lateral walls, alar group reddish,

conspicuous, median and upper cells obliquely linear-rhomboidal,

incrassate. Seta 6-8 mm. long, scabrous near tip; capsules elliptic,

rough at base; calyptra fringed. (Fig. 22, D-H.)

Dept. Quiche: Standley 62461. Dept. Huehuetenango : Standley 62586. Dept.

Quezaltenango : Standley 66393 in part, 664.06 in part. Dept. Chimaltenango :

Standley 61676, 6^361.

Distribution: Mexico, Costa Rica, West Indies, South America.

On damp, shaded banks at medium to high altitudes. This is

a plastic species. The basal cells in the local plants are often shorter

than in typical plants from Guadeloupe and the leaves of the sterile

stems frequently concolorous at the tips although some plants in-

variably show the characteristic hyaline hair points.

6. ATRACTYLOCARPUS Mitt., Journ. Linn. Soc. 12: 71. 1869.

Autoicous; stems radiculose, densely tufted. Leaves crowded,

curved or secund, setaceous pointed from a lanceolate base, serrulate

above; costa long excurrent, with two stereid bands; basal cells

rectangular, alar group pale, fragile, not auriculate. Seta straight

or slightly flexuous, elongate; capsule erect; peristome teeth divided

nearly to base, striolate below; lid long beaked; calyptra long, cucul-

late, not fringed.

Annulus present, capsules cylindric 2. A. costaricensis

Annulus lacking, capsules oblong 1. A. longisetus

1. ATRACTYLOCARPUS LONGISETUS (Hook.) Bartr., Bryol. 49: 110.

1946.

Dicranum longisetum Hook., Muse. Exot. tab. 139. 1820.

Dicranum sublongisetum C. M., Bull. Herb. Boiss. 5: 185. 1897.

Plants brownish to yellow, not glossy. Stems 2-3 cm. long or

longer, often branched. . Leaves erect or slightly falcate-secund,

6-11 mm. long, from a short ovate base, long setaceous pointed,

serrulate far below apex; costa wide below, long excurrent; basal

BARTRAM: MOSSES OF GUATEMALA

51

cells rectangular, alar group fugacious, upper cells linear. Seta

15-25 mm. long, straight or slightly flexuous; capsule oblong, urn

2 mm. long, lightly ribbed when dry; annulus lacking; peristome teeth

reddish, divided more than half way down and perforate below; lid

long and slenderly beaked; calyptra entire at base. (Fig. 23, A-C.)

Chemal: Bernoulli & Carlo 99.

Distribution: Costa Rica, Panama, northern South America.

I have seen no plants of this species from Guatemala but D. sub-

longisetum C. M. is evidently the same thing.

2. ATRACTYLOCARPUS COSTARICENSIS (C. M.) Bartr., Bryol. 49: 110.

1946.

Leptotrichum costaricense C. M., Bot. Zeit. 16: 161. 1858.

Atractylocarpus mexicanus Mitt., Journ. Linn. Soc. 12: 71. 1869.

Plants similar to A. longisetus but smaller. Leaves 5-7 mm. long,

pale yellow, slightly secund. Capsule narrowly cylindric, smooth,

urn 2-3 mm. long; annulus wide; peristome teeth divided nearly to

base, forks slender, papillose; lid dark red, 1.5 mm. long; calyptra

often reaching nearly to base of capsule, entire at base. (Fig. 23, D-F.)

B

H

X8.

FIGURE 23

A-C, Atractylocarpus longisetus: A, plant, XI; B, leaf, X6; C, capsule, X8.

D-F, Atractylocarpus costaricensis: D, plant, Xl; E, leaf, X6; F, capsule, X8.

G-J, Pilopogon gracilis: G, plant, Xl; H, leaf, X6; I, capsule, X8; J, calyptra,

52 FIELDIANA: BOTANY, VOLUME 25

Dept. Baja Verapaz: Standley 6981+1 in part. Dept. Quiche: Standley 62368.

Dept. Huehuetenango : Standley 81365. Dept. Totonicapan: Standley 81+535.

Dept. Sacatepequez: Standley 65223a. Dept. Chimaltenango: Standley 57827a.

Dept. El Progresso: Steyermark 1+31+36. Dept. Jalapa: Steyermark 321+93.

Distribution: Mexico, Honduras, Costa Rica.

On logs and banks mostly at high altitudes. A frequent species

confined to Mexico and Central America as far as known.

7. PILOPOGON Brid., Bryol. Univ. 1: 519. 1826.

Dioicous; slender plants in dense tufts. Stems branched. Leaves

erect, subulate-acuminate; costa broad with dorsal and ventral

stereid bands; basal cells rectangular, lacking a distinct alar group.

Perichaetial leaves long, convolute; seta elongate, erect; capsule

cylindric, curved; peristome teeth not divided, papillose; annulus

lacking; calyptra fringed at base.

1. PILOPOGON GRACILIS (Hook.) Brid., Bryol. Univ. 1: 519. 1826.

Didymodon gracile Hook., Muse. Exot. tab. 5. 1818.

Pilopogon gracilis var. Bernoullii C. M., Bull. Herb. Boiss. 5: 185. 1897.

Pilopogon glabrisetus C. M., Bull. Herb. Boiss. 5: 551. 1897.

Yellowish green, glossy plants. Stems to 6 cm. or more long,

radiculose, uniformly foliate. Leaves 6 mm. long, gradually subulate

from a narrowly oblong base, denticulate near apex; costa short

excurrent; basal cells thin walled, rectangular, gradually becoming

shorter and irregular above. Perichaetial leaves with long, setaceous

points often reaching the capsule; seta 1.5-2 cm. long, slender,

slightly rough above; capsule smooth, dark brown, urn 2-2.5 mm.

long; peristome teeth slender, papillose, entire; lid conic-rostrate;

annulus lacking; calyptra long, fringed at base. (Fig. 23, G-J.)

Dept. San Marcos: Standley 86195, 861+1+1+, 86511, 86515. Dept. Quezalten-

ango: Standley 85686.

Distribution: Mexico, Costa Rica, West Indies, South America.

On damp banks and trees mostly at high altitudes. Readily

distinguished from the allied genera by the conspicuous perichaetial

leaves, the curved, cylindrical capsules and the undivided peristome

teeth.

8. DICRANODONTIUM Bry. Eur. fasc. 41. 1847.

Dioicous; stems slender, tomentose, simple or branched. Leaves

slightly falcate-secund, long setaceous pointed from an ovate base;

BARTRAM: MOSSES OF GUATEMALA

53

costa broad below, long excurrent, with dorsal and ventral stereid

bands; basal cells rectangular, slightly pitted, narrower toward

margins, upper cells linear. Seta curved when moist; capsules erect,

oblong; annulus lacking; peristome teeth divided more than half

way down, vertically striolate below; lid rostrate; calyptra entire

at base.

1. DICRANODONTIUM DENUDATUM (Brid.) E. G. Britt., No. Amer.

Flora 152: 151. 1913.

Dicranum denudatum Brid., Muse. Rec. Suppl. 1 : 184. 1806.

Rather robust, pale green plants. Stems 5 cm. or more long,

uniformly foliate. Leaves falcate-secund, 6-10 mm. long, from a

short, ovate, concave base gradually long setaceous pointed, serrulate

above; costa excurrent; basal cells laxly rectangular toward costa,

much narrower toward margins, upper cells linear. Seta 1 cm. long,

curved or cygneous when moist; capsule smooth; peristome teeth

divided nearly to base; calyptra long. (Fig. 24, A-C.)

Dept. Chiquimula: Steyermark 31000.

Distribution: Eastern United States, Alaska, Mexico, Europe.

FIGURE 24

A-C, Dicranodontium denudatum: A, plant, Xl; B, leaf, X6; C, upper leaf

cells and margin, X270.

D-H, Oncophorus guatemalensis: D, plant, Xl; E, leaf, X8; F, apex of leaf,

XllO; G, upper leaf cells and margin, X270; H, part of cross section of leaf margin,

X270.

I-J, Symblepharis helicophylla: I, plant, Xl; J, leaf, X8.

54 FIELDIANA: BOTANY, VOLUME 25

On damp ground in cloud forest. These plants are more robust

than usual but this may be due to the excess of moisture. In other

respects the shape and structure of the leaves are typical.

9. BROTHERA C. M., Gen. Muse. 258. 1901.

Small yellowish green plants, slightly glossy, in dense mats.

Stems short, sparingly radiculose. Leaves crowded, subulate-acu-

minate from a lanceolate base, subtubulose; costa broad, excurrent,

without stereids; lamina cells rectangular, hyaline. Seta erect,

slender; capsule erect, oblong-ovoid; peristome single, teeth cleft

to base into two subulate, papillose forks; lid rostrate; calyptra

cucullate, fringed at base.

1. BROTHERA LEANA (Sull.) C. M., Gen. Muse. 259. 1901.

Campylopus Leana Sull., A. Gray Man. Ed. 2: 619. 1856.

Stems short, branched, less than 5 mm. high, usually with apical

clusters of rudimentary leaves which serve the purpose of vegetative

reproduction. Leaves flexuous when dry, to 2.5 mm. long, entire

or minutely toothed at apex; margins erect or inflexed; costa 60 M

wide below, poorly defined, excurrent, in cross section showing a

median row of chlorophyllose cells with bands of lax, hyaline cells

on both sides; cells of leaf base rectangular, at basal angles lax

and delicate forming small, poorly defined auricles. The local plants

are sterile. (Fig. 25, A-C.)

Dept. Huehuetenango : Sharp 493 9a. Dept. Quezaltenango : Sharp 1995.

Distribution : Pennsylvania, Ohio, Tennessee, Minnesota, Mexico,

also Asia.

On banks and peaty soil at moderately high altitudes. The plants

in both of these collections consist almost entirely of the deciduous

brood leaves and are so deformed that it is difficult to find a normal

leaf.

10. AMPHIDIUM (Nees) Schimp. emend. Bry. Eur.

Coroll. 39. 1856.

Densely tufted plants, olive green above, brown below. Stems

slender, sparingly radiculose. Leaves strongly crisped when dry,

linear-lanceolate, costate to apex; upper leaf cells rounded-quadrate,

papillose, basal cells narrowly rectangular, pellucid. Seta short;

capsule barely exserted, strongly 8 ribbed, contracted below mouth

BARTRAM: MOSSES OF GUATEMALA

55

and urceolate when dry; peristome none; lid obliquely rostellate

from a convex base; calyptra cucullate, naked.

1. AMPHIDIUM CYATHICARPUM (Mont.) Broth., E. & P. Nat.

Pflanzenf. I3: 460. 1902.

Zygodon cyathicarpus Mont., Ann. Sci. Nat. 106. 1845.

Autoicous; stems under 1 cm. high, sparingly branched. Leaves

crispate when dry, flexuous-spreading when moist, linear-lanceolate,

acuminate; margins narrowly recurved near shoulders of leaf, plane

above, distantly and shallowly toothed or notched in upper half;

costa pale, distinct, ending in or just below the sharp apex; upper

cells rounded-quadrate, diam. 8-10 n, with firm pale walls, papillose,

basal cells narrow, smooth, pellucid. Seta about 1 mm. long, often

slightly curved; capsule oblong, wide-mouthed, with 8 brownish

longitudinal ribs when dry; peristome none. (Fig. 25, D-E.)

Dept. Quezaltenango : Sharp 2290.

Distribution: Western South America, Hawaii, Australia, New

Zealand, Africa.

On boulder at high altitude. The appearance of this austral

species in Guatemala is exceedingly interesting. The only other

locality I am aware of in the northern hemisphere is on the summit

H

FIGURE 25

A-C, Brothera Leana: A, plant, XI; B, leaf, X22; C, abnormal'leaf, X22.

D-E, Amphidium cyathicarpum: D, plant, Xl; E, capsule and upper leaves,

X14.

F-I, Rhabdoweisia fugax var. tenerrima: F, plant, Xl; G, capsule, X26;

H, leaf, X20; I, basal leaf cells, X368.

56 FIELDIANA: BOTANY, VOLUME 25

of Haleakala, Maui, Hawaii. As far as I can see the Guatemalan

plants are typical in every respect. The inflorescence is autoicous

and the narrow leaves show the upper margins shallowly toothed

or sinuate. As the plants are well fruited I think there can be

little doubt regarding the accuracy of the determination.

11. RHABDOWEISIA Bry. Eur. fasc. 33-36. 1846.

Small plants growing in extensive green mats. Leaves strongly

curled when dry, narrowly linear-lanceolate, acuminate; costa ending

near apex; upper leaf cells rounded-quadrate, chlorophyllose, smooth,

basal cells rectangular, hyaline; Seta erect; capsule exserted, ovoid,

8 ribbed when dry; peristome well developed but fragile; lid rostrate.

1. RHABDOWEISIA FUGAX (Hedw.) Bry. Eur. var. TENERRIMA

Bartr., Bryol. 50: 203. 1947.

Low, delicate plants, laxly tufted. Stems 2-3 mm. high. Leaves

crispate when dry, to 2 mm. long, linear-lanceolate, acuminate;

margins plane, minutely crenulate above; costa ending just below

apex; upper cells rounded-quadrate, diam. 8-10 M, often transversely

elongate, smooth, basal cells rectangular, delicate, thin-walled and

hyaline, to 60 n long, 12 /j. wide. Seta erect, yellow, 2 mm. long;

capsule erect, ovoid, 0.5 mm. long, 8 ribbed and urceolate when dry;

peristome teeth very fragile, to 150 n long, filiform from a short,

broad base; spores pale, diam. 10-12 M- (Fig. 25, F-I.)

Dept. San Marcos: Sharp 5413a. Dept. Quezaltenango : Sharp 5085.

Endemic.

Moist bank and decaying log at high altitudes. These plants

are appreciably smaller and more delicate than any European speci-

mens I have seen. The stems are only 2-3 mm. high, the leaves up

to 2 mm. long and the basal leaf cells thin-walled hyaline and about

60 M long changing abruptly to the short chlorophyllose cells above.

In most cases only the expanded bases of the peristome teeth are

evident, the fragile, filiform tips being almost invariably broken off

in operculate capsules, but these vestiges will at once distinguish

the plants from Amphidium cyathicarpum to which they bear a close

resemblance.

12. ONCOPHORUS Brid., Bryol. Univ. 1: 389. 1826.

Autoicous; dull yellowish green plants in dense tufts. Stems often

branched. Leaves crisped when dry, lanceolate, usually serrate

BARTRAM: MOSSES OF GUATEMALA 57

above; margins recurved, bistratose; costa subpercurrent, with dorsal

and ventral stereid bands; upper cells small, basal cells rectangular.

Seta elongate, curved when moist; capsule nodding, sulcate; peris-

tome teeth divided to middle; lid obliquely rostrate; calyptra cucul-

late.

1. ONCOPHORUS GUATEMALENSIS Bartr., Bryol. 49: 111. 1946.

Densely tufted plants, dull yellowish green above, brown below.

Stems 2-3 cm. long, simple or branched, sparingly radiculose below.

Leaves crisped when dry, erect-spreading when moist, 5-6 mm. long,

narrowly lanceolate, acute, keeled above; margins recurved below,

irregularly serrate above, usually bistratose; costa stout, ending

just below the acute apex; upper cells rounded-quadrate, lightly

mammillose on both sides; inner basal cells narrowly rectangular,

wider and more lax toward margins at the basal angles. Seta 3-4

mm. long, stout, yellowish, erect-flexuous when dry, strongly curved

or cygneous when moist; capsule oblong-cylindrical, suberect, urn

1.5 mm. long, strongly ribbed when dry; annulus lacking; lid obliquely

conic-rostrate, 0.75 mm. long; peristome teeth reddish, vertically

striolate on outer plates below, cleft about half way down, forks

papillose above; calyptra cucullate; spores papillose, diam. 15-18 /*.

(Fig. 24, D-H.)

Dept. Quezaltenango : Uppermost ridge to summit of Volcan Zunil, alt.

3,000-3,800 m., Steyermark 3J^869c, S!t872.

Endemic.

On dry slopes below summit. A clean cut species characterized

by the short, cygneous setae. The genus has not been recorded

before in North America south of the United States; hence these

noteworthy collections have an added interest.

13. SYMBLEPHARIS Mont, Ann. Sci. Nat. II. 8:252. 1837.

Autoicous; medium sized plants in compact tufts. Stems erect,

densely foliate. Leaves abruptly linear-lanceolate from an obovate,

clasping base, the points widely spreading and crispate when dry;

costa excurrent; basal cells rectangular, upper cells small and dense.

Setae single or aggregated, elongate; capsule cylindrical, erect;

peristome teeth divided to below middle; lid bbliquely rostrate;

calyptra entire at base.

58 FIELDIANA: BOTANY, VOLUME 25

1. SYMBLEPHARIS HELICOPHYLLA Mont., Ann. Sci. Nat. II. 8: 252.

1837.

Stems about 3 cm. high. Leaves 6-7 mm. long, from a strongly

clasping obovate base about 2 mm. high quickly narrowed to a

spreading, grooved, subulate point, distantly denticulate above;

margins erect; costa slender, excurrent; cells smooth, narrowly

rectangular, thin walled and hyaline in base, irregularly subquadrate

with firm walls above. Setae 10-12 mm. long, straight; capsule

smooth, erect, 3-4 mm. long; peristome teeth red, deeply divided,

vertically striolate; lid about 0.8 mm. long; calyptra extending half

way down urn. (Fig. 24, I-J.)

Dept. Huehuetenango: Standley 81653, 8180^, 81813. Dept. Quezaltenango :

Steyermark 3^116, 3472^a, 3^725a, 34869a; Standley 67656, 67679. Dept. Chimal-

tenango: Standley 5877^, 609^9a, 60960.

Distribution: New Mexico, southern Arizona, Mexico, Costa

Rica, Panama, Asia.

On trees, logs and moist banks at high altitudes. The abruptly

spreading, strongly curled leaf points and the slender, erect, cylin-

drical capsules, deep red at mouth, are very characteristic. The

plants fruit abundantly throughout their range.

14. DICRANOWEISIA Lindb., Oefv. Sv. Vet.-Akad.

Forh. 21 : 230. 1864.

Tufted plants with erect, branching stems. Leaves flexuous or

crispate, subulate pointed from an ovate base, entire; costa short

excurrent; cells smooth, small and subquadrate above, rectangular

below. Seta erect, elongate; capsules suberect; peristome teeth

papillose, entire or cleft at apex.

1. DICRANOWEISIA CALCAREA Bartr., Bryol. 49: 111. 1946.

Dioicous; plants densely tufted, dull brownish yellow. Stems

2-3 cm. high, encrusted with a calcareous deposit, laxly foliate.

Leaves erect-flexuous, 2.5-3 mm. long, entire, rather abruptly subu-

late-acuminate from a short, ovate, concave, decurrent base; margins

erect or slightly recurved on one side below; costa strong, short

excurrent; cells smooth, incrassate, rectangular below and linear

toward margins, irregularly subquadrate to elongate above, 10 /*

wide, 12-25 /* long, the marginal rows smaller. Seta erect, smooth,

8-10 mm. long; capsules inclined, urn dark brown, 1.5 mm. long,

BARTRAM: MOSSES OF GUATEMALA

59

asymmetrical with a short neck; peristome none as seen (capsules

all old and overripe). (Fig. 26, A-C.)

Dept. Huehuetenango : Above San Juan Ixcoy, Sierra de los Cuchumatanes,

alt. 2,400 m., Steyermark 50001, TYPE.

Endemic. .

On dripping rock at base of waterfall on encrusted limestone in

forested ravine. This species is included in Dicranoweisia with

considerable reservation. It has some affinities with Hymenolopsis

tolucensis The"r. of Mexico but appears to be quite distinct in the

decurrent leaf angles and the lack of differentiated alar cells. Until

the peristome characters are known it seems wiser to follow the

more conservative plan.

15. HOLOMITRIUM Brid., Bryol. Univ. .1: 226. 1826.

Plants medium sized, tufted. Stems branched, tomentose, often

with terminal clusters of short, microphyllous branchlets. Leaves

crowded, narrowly lanceolate from a broader base, crisped when

dry; basal cells linear, alar group conspicuous, upper cells rectangular

to subquadrate, incrassate; costa percurrent. Inner perichaetial

leaves convolute, with long, setaceous points, often reaching the

H

FIGURE 26

A-C, Dicranoweisia calcarea: A, plant, XI; B, leaf, X14; C, upper leaf cells

and margin, X270.

D-E, Holomitrium flexuosum: D, plant, XI; E, leaf, X8.

F-H, Holomitrium falcatum: F, plant, Xl; G, leaf, X8; H, upper leaf cells,

X270.

60 FIELDIANA: BOTANY, VOLUME 25

capsule. Seta erect; capsules erect, cylindrical; peristome teeth red,

papillose, usually split along median line.

1. Leaves subentire, seta 5 mm. long 6. H. pulchellum

Leaves serrate or serrulate, seta 1 cm. or more long 2

2. Leaves spreading from insertion, serrulate above 3

Leaves with an erect, clasping base, serrate half way down 5

3. Leaves strongly falcate-secund, 3-4 mm. long 2. H. falcatum

Leaves not falcate-secund, 5 mm. or more long 4

4. Alar cells conspicuous, forming a group about 200 n high 1. H.flexuosum

Alar cells inconspicuous, forming a band less than 100 ^ high . . 3. H. terebellatum

5. Upper leaf cells quadrate 4. H. arboreum

Upper leaf cells elongate, 1 : 4 or 5 5. H. Standleyi

1. HOLOMITRIUM F.LEXUOSUM Mitt., Journ. Linn. Soc. 12: 57. 1869.

Stems 4 cm. or more high, yellowish green above, brown below.

Leaves spreading on all sides, 5 mm. or more long, narrowly lanceo-

late from an oblong base, grooved above, serrulate toward apex;

costa excurrent, toothed on back above; upper cells rectangular,

shorter at margins, basal cells narrowly rectangular, pitted, all

smooth and incrassate, alar group prominent, extending nearly to

costa. Sporophyte not seen. (Fig. 26, D-E.)

Dept. Huehuetenango: Standley 81822.

Distribution: Mexico, Costa Rica, Ecuador.

On tree in Juniperus forest at high altitude. These plants are

undersize but in other respects agree with the species.

2. HOLOMITRIUM FALCATUM Bartr., Bryol. 49: 111. 1946.

Near H. flexuosum Mitt, but apparently distinct in the shorter

stems and smaller leaves, 3-4 mm. long (5-10 mm. long in H. flex-

uosum), which are conspicuously falcate-secund both moist and dry

giving the plant a very characteristic appearance. (Fig. 26, F-H.)

Dept. Totonicapan: Near Cumbre del Aire, on road between Huehuetenango

and Sija, alt. 3,000-3,450 m., Standley 65906.

Endemic.

The distinctions outlined above are not very impressive and un-

less they can be correlated with some sharper differences in the

sporophyte it may be desirable to reduce H. falcatum to a variety

of H. flexuosum.

BARTRAM: MOSSES OF GUATEMALA

61

3. HOLOMITRIUM TEREBELLATUM C. M., in Ren. & Card. Bull.

Soc. Bot. Belg. 311: 151. 1893.

Robust plants forming deep tufts, yellowish green above, brown

below. Stems to 5 cm. or more high, branched, densely foliate,

clothed with reddish tomentum. Leaves widely spreading from the

insertion, flexuous with strongly curled points when dry, 5-6 mm.

long, gradually narrowed from an oblong, concave, entire base to

a narrow, lanceolate, grooved point, acuminate; margins erect,

undulate above, distantly and irregularly toothed down to leaf

shoulders; costa excurrent. Toothed on back near apex; upper cells

subquadrate to short rectangular, smooth, incrassate, gradually

becoming narrowly linear and porose below, alar group very fragile

and inconspicuous, forming a poorly defined band across base of

leaf less than 100 M high. (Fig. 27, A-C.)

Dept. Baja Verapaz: Sharp 2759.

Distribution: Costa Rica.

On tree trunk at moderate altitude. Previously considered a

Costa Rican endemic where it is decidedly uncommon. The Guate-

malan plants lack fruit and are undersized but the essential characters

leave little doubt as to its identity.

FIGURE 27

A-C, Holomitrium terebellatum: A, plant, Xl; B, leaf, X8; C, one side of leaf

base, X68.

D-F, Holomitrium pulchellum: D, plant, Xl; E, leaf, X20; F, apex of leaf,

X66.

62

FIELDIANA: BOTANY, VOLUME 25

sss

B D iXXXn,

FIGURE 28

A-D, Holomitrium arboreum: A, plant, Xl; B, leaf, X8; C, apex of leaf,

XllO; D, upper leaf cells and margin, X270.

E-F, Holomitrium Standleyi: E, leaf, X8; F, upper leaf cells and margin, X270.

G-I, Dicranum flagellare: G, plant, Xl; H, leaf, X8; I, upper leaf cells and

margin, X270.

4. HOLOMITRIUM ARBOREUM Mitt., Journ. Linn. Soc. 12: 58. 1869.

Stems 2-3 cm. high, yellowish green, brown below. Leaves

crowded, strongly crisped when dry, 4-6 mm. long, linear-lanceolate

from an erect, ovate, clasping base, strongly serrate above middle;

costa percurrent; upper cells subquadrate, basal cells linear, incras-

sate, pitted, alar group distinct. Tips of perichaetial leaves often

reaching the capsule. Seta 1-1.5 cm. long; capsule cylindric, urn

4-5 mm. long; lid subulate-rostrate. (Fig. 28, A-D.)

Dept. Peten: Lundell 2679 (as H. calycinum). Dept. Jalapa: Steyermark

32552 (as H. calycinum), 33 Ilk (as H. calycinum).

Distribution: Mexico to Bolivia.

On trees and rocks at moderate altitudes. The short upper leaf

cells often wider than long and in one layer throughout will separate

this species from any of its local allies.

5. HOLOMITRIUM STANDLEYI Bartr., Contr. U. S. Nat. Mus. 263:

66. 1928.

Plants similar in habit and appearance to H. arboreum but with

the upper and median leaf cells elongate especially toward costa

BARTRAM: MOSSES OF GUATEMALA 63

where they are often 4-5 times as long as wide with sinuous lateral

walls. (Fig. 28, E-F.)

Dept. Alta Verapaz: Standley 91596.

Distribution: Costa Rica.

On limestone at moderate altitude. In this species the leaf cells

are not all elongate but the areolation is quite different from that

of H. arboreum. The distinction is none too marked however and

a broader series of specimens might show that they are not wide

enough apart for practical segregation.

6. HOLOMITRIUM PULCHELLUM Mitt., Journ. Linn. Soc. 12: 60. 1869.

Forming dense, compact cushions, yellowish green above, brown

below. Stems to 2 cm. high, branched, tomentose. Leaves crowded,

very crispate when dry, flexuous-spreading when moist, 2-2.5 mm.

long, gradually lanceolate from an ovate, slightly clasping base,

acuminate, canaliculate; margins erect, entire or minutely toothed

at extreme apex; costa short-excurrent; upper leaf cells rounded,

strongly incrassate, smooth, diam. 5-7 n, basal cells narrowly rec-

tangular with firm, pale walls. Perichaetium about half as long as

the seta, leaves convolute with spreading points; seta erect, yellow,

5 mm. long; capsule erect, oblong, urn 1.5 mm. long. (Fig. 27, D-F.)

Dept. Quezaltenango : Sharp 2199a.

Distribution: Ecuador.

On tree at moderately high altitude. This is evidently a very rare

species. It is represented in the Mitten Herbarium by only one

collection from the type locality in Ecuador.

Compact cushions of tightly curled leaves studded with attrac-

tively colored, short-stalked capsules standing well above the con-

spicuous perichaetia give these plants an especially neat and trim

look. As Mitten aptly remarks it is "a very pretty moss." The

short setae and nearly entire leaves are unique characters among

the American species of Holomitrium. It is a striking addition to

the North American moss flora.

16. DICRANUM Hedw., Sp. Muse. 126. 1801.

Dioicous; medium sized to robust plants. Leaves lanceolate,

often falcate-secund ; costa narrow but strong, with dorsal and

ventral stereid bands, often toothed on back; leaf cells mostly smooth,

alar group inflated, usually colored and conspicuous. Seta erect;

64 FIELDI ANA: BOTANY, VOLUME 25

capsules cylindrical, erect or curved; peristome teeth red, vertically

striolate, cleft about half way down; lid long beaked; calyptra

cucullate, entire at base.

1. Upper leaf cells small, subquadrate 1. D. flagellare

Upper leaf cells elongate, pitted 2

2. Leaves short pointed, costa not winged on back 2. D. rhabdocarpum

Leaves long acuminate, costa winged on back 3. D. frigidum

1. DlCRANUM FLAGELLARE Hedw., Sp. Muse. 130. 1801.

Plants yellowish green, tufted; stems 1-3 cm. high, often with

fragile, microphyllous branchlets in axils of upper leaves. Leaves

3-3.5 mm. long, crispate when dry, lanceolate; margins erect, in-

flexed above, toothed near apex; costa percurrent; upper cells

quadrate or short rectangular, basal cells rectangular with firm,

pellucid walls, alar group colored, conspicuous. Seta 1-2 cm. long;

capsules cylindric, erect. (Fig. 28, G-I.)

Dept. Huehuetenango: Standley 81705, 8180J^a.

Distribution: Southern Canada, United States, Mexico, Europe,

Asia.

On logs and limestone at high altitudes. The characteristic

flagellate branchlets are very scarce in these two local collections

but in other particulars the plants are typical.

2. DICRANUM RHABDOCARPUM Sull., Mem. Am. Acad. II. 4: 172.

1849.

Pale or yellowish green glossy plants. Stems 1-2 cm. or more

long. Leaves crowded, suberect and nearly straight when dry, 3-5

mm. long, ovate-lanceolate, serrulate above, acute; costa ending

below apex, toothed on back above; basal cells rectangular, alar

group inflated and colored, upper cells elongate, incrassate, pitted.

Seta 1.5-2 cm. long; capsules cylindric, erect. (Fig. 29, A-D.)

Dept. Quezaltenango: Standley 67694a, 677 ±3, 677Ma, 67749a.

Distribution: Mountains of Colorado, new Mexico and Arizona,

Mexico.

On rocks and banks at high altitudes. These collections are

small, underdeveloped and sterile but there is little doubt concerning

their identity.

BARTRAM: MOSSES OF GUATEMALA

65

3. DICRANUM FRIGIDUM C. M., Bot. Zeit. 17: 219. 1859.

Robust plants in extensive deep mats, yellowish green and

lustrous above, brown below. Stems to 10 cm. or more long, tomen-

tose. Leaves spreading, flexuous or falcate-secund, scarcely undulate,

linear-lanceolate from an ovate base, serrate in upper half, 10-13

mm. long; costa ending below apex, with two sharply serrated

wings on back; cells all elongate with thickened, pitted walls, alar

group brown. Setae aggregated, 1-3, red, to 5 cm. long; capsule

cylindric, curved, urn 4-5 mm. long. (Fig. 29, E-H.)

Dept. Quiche: Aguilar 1115. Dept. Huehuetenango : Steyermark 50173, 51901.

Dept. Zacapa: Steyermark 43290.

Distribution: Mexico to northern South America.

On damp ground at high altitudes. Near D. rugosum (Hoffm.)

Brid. but uniformly distinct in the longer, scarcely undulate leaves,

longer setae and longer capsules.

17. LEUCOLOMA Brid., Bryol. Univ. 2:218. 1827.

Dioicous; pale green, silky plants in soft, loose tufts. Stems

branched, sparsely radiculose. Leaves flexuous or secund, gradually

subulate-lanceolate from an ovate base; costa narrow; chlorophyllose

FIGURE 29

A-D, Dicranum rhabdocarpum: A, plant, Xl; B, leaf, X8; C, apex of leaf,

XllO; D, upper leaf cells and margin, X270.

E-H, Dicranum frigidum: E, part of plant, Xl; F, leaf, X6; G, apex of leaf,

XllO; H, upper leaf cells and margin, X270.

66

FIELDIANA: BOTANY, VOLUME 25

H

D

FIGURE 30

A-D, Leucoloma serrulatum: A, plant, XI; B, leaf, X8; C, apex of leaf, XllO;

D, upper leaf cells and margin, X270.

E-H, Leucoloma Crugerianum: E, plant, Xl; F, leaf, X8; G, apex of leaf,

XllO; H, cells and margin near mid-leaf, X270.

cells small, papillose, marginal cells smooth, narrow and hyaline

forming a distinct border, alar group large, auriculate. Seta erect;

capsules cylindric; peristome teeth divided to or below middle.

Leaves with a distinct median band of short, papillose cells extending to or near

base 1. L. serrulatum

Leaves without a median band of short cells in basal part 2. L. Crugerianum

1. LEUCOLOMA SERRULATUM Brid., Bryol. Univ. 2: 752. 1827.

Stems to 6 cm. or more long. Leaves 4-7 mm. long, straight or

curved, narrowly subulate-acuminate, grooved above, serrulate to-

ward apex; costa excurrent; marginal cells linear, hyaline, forming

a distinct border merging with the basal cells below, chlorophyllose

cells small, oblong, papillose on back, extending nearly to insertion

in a broad, sharply defined median band. Sporophyte not seen.

(Fig. 30, A-D.)

Dept. Izabal: Steyermark 38805a, 89203, 41765. Dept. Alta Verapaz: Steyer-

mark 45561, 45614, 45615, 46376; Standley 90639. Dept. Quezaltenango : Steyer-

mark 34350. Dept. Zacapa: Steyermark 29827, 42789. Dept. Chiquimula:

Steyermark 31030.

Distribution: Mexico, Costa Rica, West Indies, British Guiana.

On logs and trees at low and medium altitudes. These collections

are sterile as are all the numerous specimens in my herbarium but

BARTRAM: MOSSES OF GUATEMALA 67

the broad median band of green cells reaching nearly to the insertion

is a distinctive feature.

2. LEUCOLOMA CRUGERIANUM (C. M.) Jaeg., Adumb. 1: 116.

1872-73.

Dicranum Crugerianum C. M., Syn. 2: 588. 1851.

Stems short, fragile. Leaves narrowly lanceolate, subulate

pointed, flexuous and widely spreading when dry, 3-5 mm. long,

tubulose above, serrulate near apex; costa excurrent; border of linear

cells one row wide above, gradually wider below and merging with

the basal cells, green cells subquadrate, sharply papillose on back

above, irregularly longer below where they merge with the basal

cells. Sporophyte not seen. (Fig. 30, E-H.)

Dept. Peten: Lundell 28^0. Dept. Quezaltenango : Standley 8^880.

Distribution: Mexico, British Honduras, Costa Rica, West Indies,

Venezuela.

On banks at low to medium altitudes. The narrower leaves,

indistinctly bordered and without a well defined median band of

green cells will distinguish this species from L. serrulatum.

7. LEUCOBRYACEAE

Compactly tufted, whitish green plants, leaves fragile, consist-

ing mostly of a broad, thick costa showing in cross section a central

row of small chlorophyllose cells (chlorocysts) covered on both sides

by one or more layers of large hyaline cells (leucocysts), porose

on the inner walls. Sporophyte as in Dicranaceae.

1. Capsules immersed, calyptra fringed at base 1. Ochrobryum

Capsules exserted, calyptra entire at base 2

2. Capsules ovoid, inclined, asymmetrical 3. Leucobryum

Capsules cylindrical, erect, symmetrical 2. Octoblepharum

1. OCHROBRYUM Mitt., Journ. Linn. Soc. 12: 108. 1869.

Low, dull, whitish green plants forming dense tufts. Leaves

erect to slightly spreading, crowded, rigid when moist, linear-lanceo-

late from a narrowly ovate base, subtubulose above, bluntly pointed,

leucocysts in one layer on each side of the median row of chlorocysts;

lamina cells narrowly rectangular, confined to basal part. Seta

short, terminal; capsules immersed; peristome lacking; calyptra

conic-rostrate, slender, fringed at base.

68

FIELDIANA: BOTANY, VOLUME 25

H

FIGURE 31

A-D, Ochrobryum obtusifoUum: A, plant, Xl; B, leaf, X18; C, apex of leaf,

X68; D, part of cross section of leaf, X68.

E-F, Syrrhopodon ligulatus: E, plant, Xl; F, leaf, X16.

G-H, Syrrhopodon lycopodioides: G, plant, Xl; H, leaf, X6.

1. OCHROBRYUM OBTUSIFOLIUM (C. M.) Mitt., Journ. Linn. Soc. 12:

108. 1869.

Schistomitrium obtusifoUum C. M., Bot. Zeit. 577. 1857.

Leaves 3-3.5 mm. long, obtuse or bluntly acute, often with brush-

like clusters of brownish filaments on the dorsal face near apex which

varies considerably from broadly rounded to acute but always with

a minute apiculate point; margins erect or inflexed so that the blade

is deeply grooved or subtubulose above; lamina cells thin- walled,

hyaline, in 4-6 rows, evident only on the basal margins. (Fig. 31,

A-D.)

Dept. Baja Verapaz: Sharp 2660a.

Distribution: Costa Rica, Colombia.

On decaying log at rather low altitude. The only other North

American records of this rare little species are from Costa Rica where

it was collected by Standley. Here as in Costa Rica the plants are

characterized by conspicuous clusters of brownish septate filaments

on the dorsal faces of the leaf tips.

Although quite similar in appearance to some of the smaller

forms of Leucobryum albidum the lack of any expanded leaf base

coupled with the leucocysts in two layers, one above and one below

the median row of chlorocysts, cannot fail to distinguish it upon

careful examination.

BARTRAM: MOSSES OF GUATEMALA 69

2. OCTOBLEPHARUM Hedw., Sp. Muse. 50. 1801.

Autoicous; whitish plants tinged with brown or red, growing in

tufts or cushions. Leaves spreading, lingulate from a hyaline base,

plane above, apiculate and serrulate at apex; costa with a median

row of small, triangular chlorocysts and 3-4 layers of leucocysts

on both sides; lamina small and narrow, confined to base. Seta

erect; capsules erect, cylindrical; peristome teeth 8 or 16; calyptra

cucullate, entire at base.

1. Leaves less than 10 mm. long 2

Leaves 15-20 mm. long 3

2. Peristome teeth 8, leaves usually not fragile 1. 0. albidum

Peristome teeth 16, leaves very fragile 2. 0. pulvinatum

3. Plants yellowish, leaf base oblong, tapering above 3. O. erectifolium

Plants purplish, leaf base obovate, rounded above 4. 0. Mittenii

1. OCTOBLEPHARUM ALBIDUM Hedw., Sp. Muse. 50. 1801.

Plants in dense cushions. Stems to 3 cm. high, branched.

Leaves widely spreading or recurved, to 6 mm. or more long, oblong-

lingulate from a slightly broader erect base, rounded, apiculate and

serrulate at apex. Seta 4-7 mm. long; capsule oblong, 1-1.5 mm.

long; peristome teeth 8, brown, faintly striolate; lid obliquely

rostrate. (Fig. 32, A-D.)

Dept. Izabal: H. Johnson 10^6; Standley 728 J^l; Steyermark 1^1830, 418S1,

Dept. Baja Verapaz: Standley 69770. Dept. Retalhuleu: Standley 885^6. Dept.

Chiquimula: Steyermark 81226. Dept. Jalapa: Standley 77^11; Steyermark 32091.

Dept. Santa Rosa: Standley 78030.

Distribution: Pantropical, southern Florida.

On tree trunks. Mostly in the lowlands. The rather short,

fleshy, strap-shaped leaves, not or rarely fragile will readily identify

this common, widely distributed species.

2. OCTOBLEPHARUM PULVINATUM (Doz. & Molk.) Mitt., Journ.

Linn. Soc. 12: 109. 1869.

Arthrocormus pulvinatus Doz. & Molk., Fl. Bryol. Surinam 6. 1854.

Plants similar to 0. albidum but with more erect, less fleshy and

very fragile leaves. Seta 10 mm. or more long; capsules about 2 mm.

long; peristome teeth 16, in 8 pairs, nearly smooth. (Fig. 32, E-G.)

Dept. Peten: Lundell 2700, 27S6a, 2769. Dept. Izabal: Steyermark 38738.

Distribution: Costa Rica, British Honduras, West Indies, north-

ern South America.

70

FIELDIANA: BOTANY, VOLUME 25

D

FIGURE 32

A-D, Octoblepharum albidum: A, plant, Xl; B, leaf, X8; C, capsule, X8;

D, part of cross section of costa, XllO.

E-G, Octoblepharum pulvinatum: E, plant, Xl; F, leaf, X8; G, part of cross

section of costa, XllO.

H-I, Octoblepharum erectifolium: H, plant, Xl; I, leaf base, XlO.

J-K, Octoblepharum Mittenii: J, plant, Xl; K, leaf base, XlO.

On tree trunks at low altitudes. Although sterile these collec-

tions seem to meet all the requirements of 0. pulvinatum.

3. OCTOBLEPHARUM ERECTIFOLIUM Mitt., No. Amer. Flora 152: 162.

1913.

Plants glossy, tinged with brown. Leaves erect, rigid, fragile,

1.5-2 cm. long, narrowly ligulate from a narrowly oblong base

tapering at shoulders; apex acute, irregularly sinuate; lamina at leaf

base narrow, tapering upward, 150-160 ^ wide on each side of costa,

inner cells 3-4 times as long as wide. Sporophyte unknown. (Fig.

32, H-I.)

Dept. Alta Verapaz: Standley 91692.

Distribution: Costa Rica, Jamaica, Trinidad.

On trees at moderate altitude. The coloring alone should separate

this species from 0. Mittenii but in addition there seem to be tangible

differentiating characters in the shape and areolation of the leaf base.

4. OCTOBLEPHARUM MITTENII Jaeg., Adumb. 1 : 169. 1871-72.

Octoblepharum longifolium Mitt., Journ. Linn. Soc. 12: 110. 1869.

BARTRAM: MOSSES OF GUATEMALA 71

Plants deeply tinged with purple, glossy with an iridescent sheen.

Leaves fragile, 1.5 cm. or more long, narrowly ligulate from an

obovate base, broadly rounded at shoulders, apex obtuse, apiculate;

interior lamina cells short rectangular, about 40 M wide and 1-2

times as long, thin walled, in two layers, narrowly linear-rhomboidal

and in one layer toward margins. Sporophyte unknown. (Fig. 32,

J-K.)

Dept. Alta Verapaz: Standley 91700.

Distribution: British Honduras, Costa Rica, Brazil.

On log at moderate altitude.

3. LEUCOBRYUM Hampe, Flora 20: 282. 1837.

Whitish green plants in dense cushions. Leaves crowded, spread-

ing or flexuous, from an ovate base narrowed to a subtubulose point,

composed almost entirely of the costa, lamina reduced to a narrow

hyaline margin below; in cross section showing a central row of

small, angular chlorocysts with 1-4 layers of leucocysts on both sides.

Seta elongate; capsules inclined; peristome dicranoid.

1. Leucocysts in 2 layers throughout the leaf 4. L. Martianum

Leucocysts in 4 or more layers in thickest part of leaf base 2

2. Leaves 5-10 mm. long, tip longer than base 3. L. antillarum

Leaves less than 5 mm. long, tubulose tip about equaling basal part 3

3. Leucocysts in 4 layers in thickest part of leaf 1. L. albidum

Leucocysts in 5-6 layers in thickest part of leaf 2. L. Polakowskyi

1. LEUCOBRYUM ALBIDUM (Brid.) Lindb., Oefv. Sv. Vet.-Akad.

Forh. 20: 403. 1863.

Dicranum albidum Brid., Muse. Rec. 21: 167. 1798 and Sp. Muse. 205. 1806.

Leucobryum incurvifolium C. M., Bull. Herb. Boiss. 5: 174. 1897.

Relatively small plants; stems 1-3 cm. high. Leaves crowded,

imbricated when dry, to 4.5 mm. long, the subtubulose point shorter

than or equal in length to the broad base, in cross section near base

showing 2-3 layers of leucocysts on each side of the median row of

chlorocysts in the thicker parts of leaf. Seta elongate, slender, red;

capsule nodding, curved, ribbed when dry; lid beaked, about as long

as urn. (Fig. 33, A-C.)

Dept. Alta Verapaz: Standley 69121, 90630, 92207; Steyermark Ml>19, 45630.

Dept. Zacapa: Steyermark 42230.

Distribution: Eastern United States, Mexico, Costa Rica, Ba-

hamas, West Indies.

72 FIELDIANA: BOTANY, VOLUME 25

On logs and soil at low to medium altitudes. Rather variable

in development and scarcely distinct from the following species

which will probably have to be included with it.

2. LEUCOBRYUM POLAKOWSKYI (C. M.) Card., Mem. Soc. Sci. Nat.

Cherb. 32: 82. 1900.

Ochrobryum Polakowskyi C. M., Besch. Journ. de Bot. 11 : 151. 1897.

The distinctions between this species and L. albidum are not

impressive. Here the subtubulose leaf points are a little longer and

the leucocysts in the thicker parts of the leaf base in about 3 layers

on each side of the chlorocyst row but these differences are not

always clearly correlated or maintained. (Fig. 33, D-E.)

Dept. Alta Verapaz: Standley 90779, 91456. Dept. San Marcos: Steyermark

36675. Dept. Solola: Steyermark 47985.

Distribution: Mexico, Costa Rica, Jamaica.

On logs and soil, mostly at medium altitudes.

3. LEUCOBRYUM ANTILLARUM Schimp., Besch. Ann. Sci. Nat. VI.

3: 190. 1876.

Stems longer than in L. albidum. Leaves spreading, flexuous,

5-10 mm. long, the subtubulose part from slightly longer to 2 or 3

times as long as the leaf base, in cross section showing 2-3 layers

of leucocysts on each side of the chlorocysts in the thicker parts of

the base; hyaline lamina about 8 cells wide. (Fig. 33, F-G.)

Dept. Alta Verapaz: Standley 92316, 92321 . Dept. Quezaltenango : Steyermark

Distribution: Florida, West Indies, Central and South America.

On logs and hummocks in swamps at moderate altitudes. This

is not a very convincing species. The Florida plants as described

and illustrated in Grout's Moss Flora of North America are certainly

not typical and it is often difficult to see how L. antillarum differs

from L. glaucum (Hedw.) Schimp.

4. LEUCOBRYUM MARTIANUM (Hornsch.) Hampe, Linnaea 17: 317.

1843.

Dicranum Martianum Hornsch., Fl. Bras. 1 : 11. 1840.

Plants in lax mats; stems 1-2 cm. high. Leaves crowded, falcate-

secund, 5-6 mm. long, gradually narrowed from an ovate base to

slender, subtubulose point; leaf base not thickened, leucocysts in

BARTRAM: MOSSES OF GUATEMALA

73

a single layer on each side of the chlorocysts throughout the leaf,

chlorocysts nearer the dorsal surface in upper part of leaf; hyaline

lamina about 6 cells wide. Seta to 2 cm. long, red; capsules curved,

ribbed when dry, strumose. (Fig. 34, A-C.)

Dept. Izabal: Standley 72767.

Distribution: Costa Rica, West Indies, northern South America.

On log at low altitude. The leaf structure showing the leucocysts

in only 2 layers throughout is sharply distinctive.

8. CALYMPERACEAE

Small to fairly robust plants growing in tufts, mostly on trees.

Leaves lanceolate from a pale sheathing base, often with a narrow

hyaline border, more rarely unbordered or with thickened, concolor-

ous margins; costa strong; upper cells small, usually papillose; inner

basal cells abruptly large and hyaline (cancellinae). Seta erect,

usually elongate; capsules erect, cylindrical; peristome of 16 papillose

teeth or lacking; calyptra cucullate or campanulate.

Peristome present, calyptra cucullate 1. Syrrhopodon

Peristome lacking, calyptra campanulate and persistent, leaves usually with an

intramarginal border of elongated cells 2. Calymperes

B

FIGURE 33

A-C, Leucobryum albidum: A, plant, Xl; B, leaf, X12; C, part of cross section

of leaf near base, X68.

D-E, Leucobryum Polakowskyi: D, leaf, X12; E, part of cross section of leaf

near base, X68.

F-G, Leucobryum antillarum: F, leaf, X6; G, part of cross section of leaf near

base, X68.

74 FIELDIANA: BOTANY, VOLUME 25 '

1. SYRRHOPODON Schwaegr., Suppl. 22: 110. 1824.

Plants green or brownish; stems branched. Leaves crowded, the

whitish, imbricated bases often conspicuous, lanceolate or ligulate,

with either a thickened or hyaline border; costa stout, subpercurrent,

often spinose on one or both sides; upper cells small, changing

abruptly to the large, hyaline cancellinae cells of the leaf base.

Capsules cylindrical; peristome teeth 16, papillose; calyptra cucullate.

1 . Leaves bordered with narrow, elongated cells 2

Leaf border thickened, doubly serrate, marginal cells not differentiated 5

2. Robust plants, leaf margins spinose-serrate with paired teeth . 6. S. lycopodioides

Small plants, leaves entire or nearly so 3

3. Leaves ligulate, apex rounded 2. S. ligulatus

Leaves lanceolate or linear, acute 4

4. Leaves long-pointed, bordered nearly to apex 1. S. prolifer

Leaves acute, unbordered in upper ^ 5. S. parasiticus

5. Basal leaf cells reddish 4. S. Bernoullii

Basal leaf cells hyaline S. S. incompletus

1. SYRRHOPODON PROLIFER Schwaegr., Suppl. 22: pt. 2, 99. 1827.

Plants fragile, pale or yellowish green; stems 1-2 cm. long,

branched. Leaves crowded, flexuous when dry, 3-6 mm. long or

longer, linear from a pale, oblong base, acute, sharply toothed near

apex, otherwise entire, with a narrow hyaline border of elongated

cells extending nearly to apex; costa ending below apex, spinose

near tip; upper cells small, dense, papillose, obscure, cancellinae

filling nearly all of the leaf base, in 2 layers, usually acutely angled

above. Seta 5-8 mm. long; capsule cylindric, lid beaked; calyptra

covering more than half the urn. (Fig. 34, D-F.)

Dept. Solola: Steyermark ^7986a.

Distribution: Mexico, West Indies, Central and South America.

On rock at medium altitude. Just a few stems segregated from

other mosses but enough for identification. It seems reasonably

sure that S. flavescens C. M., including the list of synonyms given by

Williams (33, p. 376), may be included here. The leaves vary con-

siderably in length but little in structural details.

2. SYRRHOPODON LIGULATUS Mont., Syll. 47. 1856.

Small, brownish, brittle plants, densely tufted. Stems to 1 cm.

high, branched, densely foliate. Leaves strongly curled when dry,

BARTRAM: MOSSES OF GUATEMALA

75

to 3 mm. long, ligulate from a scarcely wider oblong, pale, entire

base, apex broadly rounded, often minutely apiculate, unbordered

or with a single row of very narrow, hyaline, marginal cells here and

there in the blade; costa ending below apex; upper cells papillose,

obscure, diam. 6-8 n, cancellinae in 4 or 5 rows, broadly rounded

above, bordered by about 5 rows of linear, pellucid cells. Seta red,

3-4 mm. long; capsule cylindrical, dark brown, urn 1 mm. long;

lid subulate-rostrate (Fig. 31, E-F.)

Dept. Baja Verapaz: Sharp 270^.

Distribution: Florida, West Indies, northern South America.

On tree at rather low altitude. Like S. lycopodioides this species

is a representative Caribbean type extending from Florida through

the West Indies to the Guianas but not recorded before from Central

America.

3. SYRRHOPODON INCOMPLETUS Schwaegr., Suppl. 21: 119. 1824.

Syrrhopodon decolorans C. M., Bull. Herb. Boiss. 5: 188. 1897.

Fairly robust, brownish green plants in deep tufts. Leaves

crowded, 4-5 mm. long, abruptly ' broadly linear from a hyaline,

FIGURE 34

A-C, Leucobryum Martianum: A, plant, Xl; B, leaf, X8; C, part of cross

section of leaf near base, X68.

D-F, Syrrhopodon prolifer: D, plant, Xl; E, leaf, X14; F, upper leaf base and

margin, X110.

G-I, Syrrhopodon incomplete: G, plant, X 1 ; H, leaf, X 10; I, upper leaf margin,

X134.

76 FIELDIANA: BOTANY, VOLUME 25

obovate base, basal margins serrulate, margins of blade concolorous,

thickened and doubly serrate, apex rounded and occasionally bearing

clusters of propagula; costa nearly percurrent; upper cells subquad-

rate or longer than wide, cancellinae in 10-12 rows, rounded above.

Seta 6-7 mm. long; capsule oblong, narrowed at mouth; peristome a

short, pale cylinder not exceeding the rim. (Fig. 34, G-I.)

Dept. Peten: Lundell 2035; Bartlett 12250, 12^88. Dept. Izabal: Standley

72788.

Distribution: Florida, Mexico, Honduras, British Honduras,

Costa Rica, Panama, West Indies, northern South America.

On trees at low altitudes. A common Caribbean species easily

known by the obovate, whitish leaf base and the thickened, winged

margins of the leaf blade serrate on the edges.

4. SYRRHOPODON BERNOULLII C. M., Bull. Herb. Boiss. 5: 189.

1897.

Brownish green plants growing in deep tufts; stems to 4-5 cm.

high, branched. Leaves to 6-7 mm. long, linear-lanceolate from an

oblong-ovate golden brown base; margins serrulate at base, thickened,

concolorous and doubly serrate above; costa nearly percurrent;

upper cells slightly elongate; cancellinae in numerous rows, not

sharply defined, rounded above. Seta to 18 mm. long; capsule

oblong, small mouthed; peristome teeth pale brown, coarsely papil-

lose. (Fig. 35, A-C.)

Distribution: Nicaragua, Costa Rica, Panama, Guadeloupe,

Cocos Island.

I have seen no Guatemalan collection and the species is evidently

infrequent locally although widely but sparingly distributed else-

where.

5. SYRRHOPODON PARASITICUS (Sw.) Besch., Ann. Sci. Nat. VIII.

1 : 298. 1895.

Encalypta parasitica Sw., Fl. Ind. Occ. 3: 1759. 1806.

Plants laxly gregarious or mixed with other mosses; stems to 2 cm.

high. Stem leaves linear-lanceolate from a slightly broader base,

narrowly and irregularly bordered in the median part with elongated

cells in several rows, sharply pointed, entire or minutely serrulate

above; costa percurrent; upper cells irregularly hexagonal, cancellinae

in 15-20 rows, acutely angled above. Comal leaves shorter and

broader, often bearing on the inner face near the costa conspicuous

BARTRAM: MOSSES OF GUATEMALA

77

filiform propagula. Seta short; capsule erect, cylindrical; peristome

teeth short and irregular, barely exceeding the rim. (Fig. 35, D-G.)

Dept. Peten: Lundell 301>7. Dept. Alta Verapaz: Standley 90775a.

Distribution: Florida, Mexico, Panama, West Indies, Galapagos

Islands.

On trees and logs at low altitudes. The irregular, narrow, yellow-

ish border, ending far below the apex and the characteristic propagula

of the terminal leaves easily separate this species from its congeners.

6. SYRRHOPODON LYCOPODIOIDES (Sw.) C. M., Syn. 1 : 538. 1849.

?Dicranum lycopodioides Sw., Prod. Fl. Ind. Occid. 3: 1066. 1806.

Robust plants forming dense, deep tufts, yellowish at tips,

brown below. Stems to 6 cm. high, branched, clothed with brown

tomentum. Leaves widely spreading, to 1 cm. or more long, gradually

lanceolate from a pale, erect, scarcely wider base, acuminate, keeled

below, blade strongly bordered with narrow cells, border thickened,

brownish, spinose-serrate with paired teeth, cancellinae cells short

rectangular, gradually merging with the upper leaf cells which are

rounded -quadrate, smooth and incrassate. Seta elongate; capsule

oblong-cylindrical. (Fig. 31, G-H.)

Dept. Baja Verapaz: Sharp 2760.

FIGURE 35

A-C, Syrrhopodon Bernoulli!: A, plant, Xl; B, leaf, X8; C, apex of leaf, X134.

D-G, Syrrhopodon parasiticus: D, plant, Xl; E, comal leaf, X8; F, stem leaf,

X8; G, apex of comal leaf, X134.

78 FIELDIANA: BOTANY, VOLUME 25

Distribution: Costa Rica, West Indies, northern and western

South America to Bolivia.

On tree trunk at moderate altitude. A frequent species in the

West Indies but uncommon in Central America where it has been

collected before only in Costa Rica.

2. CALYMPERES Sw., in Schwaegr. Suppl. I2: 333. 1816.

Small to moderately robust, mostly corticolous plants growing

in tufts. Leaves crispate when dry, lanceolate or ligulate from a

broader, whitish base; costa stout, often thickened at apex and bear-

ing apical clusters of propagula; lamina cells small, changing abruptly

to the lax, hyaline cancellinae cells of the leaf base, frequently with

narrow, intramarginal bands of elongate cells (teniolae) extending

through the shoulders upward into the blade; margins usually

thickened and serrate. Seta short; capsule subcylindric; peristome

lacking; calyptra campanulate, plicate, persistent.

1. Leaves 10 mm. or more long, upper cells transversely elongate

5. C. lonchophyllum

Leaves less than 5 mm. long, upper cells rounded 2

2. Upper leaf cells about 4 /* in diam 1. C. Donnellii

Upper leaf cells 6-8 /* in diam 3

3. Teniolae short, leaf border very thick 4. C. nicaraguense

Teniolae extending above midleaf , leaf border slightly thickened 4

4. Teniolae 2-3 cells in from margin at shoulders, leaves entire above

2. C. Richardi

Teniolae 4-8 cells in from margin at shoulders, leaves serrate above

3. C. emersum

1. CALYMPERES DONNELLII Aust., Bot. Gaz. 4: 151. 1879.

Plants less than 1 cm. high, often forming green mats. Leaves

incurved and crisped when dry, 2.5-5 mm. long, broadly linear from

an oblong base, tubulose above; margins thickened, irregularly

doubly serrate above, serrulate below; costa stout; upper cells

minute, diam. 4-5 /x, papillose; teniolae distinct, 8-12 cells in from

margin at shoulders, extending about half way up the blade; cancel-

linae in 12-14 rows, acutely angled above. Seta 5 mm. long; capsule

about 2 mm. long. (Fig. 36, A-E.)

Dept. Peten: Lundell 2072a. Dept. Izabal: Steyermark J^lSlSa.

Distribution: Florida, Honduras, Panama, West Indies, northern

South America, Cocos Island.

On rocks and tree trunks at low altitudes.

BARTRAM: MOSSES OF GUATEMALA 79

2. CALYMPERES RICHARDI C. M., Syn. 1 : 524. 1849.

Stems to 1 cm. high, rarely higher. Leaves incurved and crisped

when dry, small below, the upper 3-4 mm. long, oblong-ovate from

a slightly wider variable base, broadly acute or obtuse; margins

entire above, serrulate at shoulders; costa stout, scabrous on both

sides above, ending below apex; upper cells rounded, distinct, 6-8 n\

teniolae 1-5 cells in from margins at shoulders, extending into the

thickened border toward apex; cancellinae rounded above. Ab-

normal leaves narrower, club-shaped, bearing numerous propagula.

Seta 3 mm. long; capsule narrowly oval. (Fig. 36, F-H.)

Distribution: Florida, Mexico, British Honduras, West Indies,

Brazil.

On trees at low altitudes. A frequent species, widely distributed

in Caribbean regions but apparently rare or overlooked in the local

area.

3. CALYMPERES EMERSUM C. M., Bull. Herb. Boiss. 5: 189. 1897.

Similar in appearance to C. Richardi. Leaves to 4 mm. long,

oblong-linear from a narrowly obovate, serrulate base, acute;

FIGURE 36

A-E, Calymperes Donnellii: A, plant, Xl; B, abnormal leaf, X14; C, normal

leaf, X14; D, apex of normal leaf, X134; E, cells and margin near leaf shoulder,

X270.

F-H, Calymperes Richardi: F, normal leaf, X14; G, abnormal leaf, X14;

H, cells and margin near leaf shoulder, X270.

I-K, Calymperes emersum: I, leaf, X14; J, cells and margin near leaf shoulder,

X270; K, upper leaf cells and margin, X270.

80

FIELD IANA: BOTANY, VOLUME 25

H

FIGURE 37

A-C, Calymperes nicaraguense: A, plant, Xl; B, leaf, X12; C, apex

X134.

D-G, Calymperes lonchophyllum: D, moist plant, Xl; E, leaf, X4; F,

leaf, X18; G, leaf cells near shoulder, X270.

H-J, Encalypta vulgaris: H, plant, Xl; I, leaf, X14; J, calyptra, X8.

of leaf,

base of

margins of blade serrulate; costa ending just below apex; upper

cells papillose, angular, 7-8 juJ cancellinae acutely angled above;

teniolae extending nearly to apex and 4-8 cells in from margins at

shoulders. Sporophyte not seen. (Fig. 36, I-K.)

Distribution: Florida, Honduras, Panama.

Distinguished from C. Richardi by the narrower leaf blade serru-

late on the margins and the longer teniolae. No Guatemalan speci-

mens have been seen.

4. CALYMPERES NICARAGUENSE Ren. & Card., Bull. Soc. Bot. Belg.

332: 117. 1894.

Calymperes Carionis C. M., Bull. Herb. Boiss. 5: 189. 1897.

Rather robust, dark green plants. Stems densely foliate, to 2.5

cm. high. Leaves crispate when dry, to 4.5 mm. long, linear from

an obovate, serrulate base, bluntly pointed ; margins of blade strongly

thickened, serrulate; costa nearly percurrent, scabrous on both sides

above; upper cells rounded or angular, mammillose on ventral face,

nearly smooth on back; teniolae 8-10 cells in from margins at

shoulders, soon merging with the thickened border above and

disappearing below shoulders; cancellinae rounded or broadly angled

BARTRAM: MOSSES OF GUATEMALA 81

above. Seta 5-6 mm. long; capsule cylindrical, about 2.5 mm. long.

(Fig. 37, A-C.)

Distribution : Nicaragua, British Honduras, Costa Rica, Panama,

Guadeloupe.

I have a part of the type collection from Nicaragua but have

not seen the material described as C. Carionis from Guatemala.

Like its associates it is a lowland species.

5. CALYMPERES LONCHOPHYLLUM Schwaegr., Suppl. I2: 333. 1816.

Plants with short stems and very long leaves, growing in exten-

sive yellowish green mats. Stems under 5 mm. long. Leaves

crowded, 10-15 mm. or more long, flexuous or strongly curled when

dry, narrowly linear from a very short, ovate, serrulate base, acute;

margins of blade thickened and distantly doubly serrulate; costa

nearly percurrent; cells of blade often in two layers, smooth, trans-

versely oval, incrassate, longer diameter 8-10 /*; teniolae lacking

or very indistinct; cancellinae in two layers toward costa, in 6-7

rows, rounded or truncate above. Seta 10-12 mm. long, often

slightly scabrous above; capsule 2 mm. long. (Fig. 37, D-G.)

Dept. Peten: Lundell 2051; Bartletl 12441, 12472. Dept. Izabal: Steyermark

38891, 39200. Dept. Alta Verapaz: Steyermark 44465, 44995.

Distribution: British Honduras, Costa Rica, West Indies, north-

ern South America.

On trees and logs at low altitudes. Readily known by the long,

narrow leaves, much longer than the stems.

9. ENCALYPTACEAE

Coarse, rather robust, densely tufted plants. Leaves crisped

when dry, ligulate, obtuse or apiculate; costa strong, ending below

apex or excurrent; upper cells hexagonal, densely papillose, basal

cells rectangular, smooth. Seta erect; capsule erect, cylindrical;

peristome lacking or variable; lid beaked; calyptra large and con-

spicuous, campanulate, erose or fringed at base.

1. ENCALYPTA Hedw., Sp. Muse. 60. 1801.

A single genus with the characters of the family.

1. ENCALYPTA VULGARIS Hedw., Sp. Muse. 60. 1801.

Autoicous; stems 0.5 cm. or more high, tomentose below. Leaves

lightly crisped when dry, 2-3 mm. long, oblong-ligulate from a

82 FIELD IANA: BOTANY, VOLUME 25

slightly broader, yellowish base, broadly acute or obtuse; margins

erect or slightly recurved below, papillose-crenulate in blade; costa

stout, reddish, ending below apex; upper cells hexagonal, very ob-

scure, densely papillose, basal cells rectangular, often with thickened

end walls, linear toward margins. Seta 5 mm. or more long, red;

capsule furrowed and wrinkled when dry and empty; peristome

lacking; calyptra straw colored, covering capsule, ragged at base,

scabrous above. (Fig. 37, H-J.)

Dept. Huehuetenango : Standley 83090b.

Distribution: Western United States; Europe, Asia, Africa.

On soil at high altitude. This species is frequent in the mountains

of 'Arizona and New Mexico but I know of no other record south

of the border.

10. POTTIACEAE

Small to moderately robust, densely tufted plants, mostly rupe-

strine or terrestrial and partial to calcareous substrata. Stems erect,

usually branched. Leaves often crisped when dry; costa strong;

upper cells small, usually papillose, often obscure, basal cells rec-

tangular, often hyaline. Seta erect, elongate, smooth; capsules

erect, subcylindric, symmetrical, smooth; lid beaked; peristome

lacking or of 16 erect or spirally twisted teeth from a basal mem-

brane, entire or divided into 2 filiform, papillose forks; calyptra

cucullate.

1. Costa with dorsal and ventral stereid bands 2

Costa with dorsal stereid band only 19

2. Sporophyte lateral 1. Anoectangium

Sporophyte terminal (except Pleurochaete) 3

3. Leaf margin involute 4

Leaf margin plane or revolute 5

4. Peristome lacking 3. Hymenostomum

Peristome present 4. Weisia

5. Leaves with a thickened border 21. Morinia

Leaf border not thickened 6

6. Hyaline basal cells extending upward along leaf margin 7

Hyaline basal cells not as above 8

7. Leaf margins entire, sporophyte terminal 13. Tortella

Leaf margins toothed, sporophyte lateral 14. Pleurochaete

8. Peristome lacking 9

Peristome present 12

BARTRAM: MOSSES OF GUATEMALA 83

9. Leaves broadly Ungulate 17. Hyophila

Leaves narrower, usually lanceolate 10

10. Lid remaining attached to columella after dehiscence 7. Hymenostylium

Lid not persistent, free from columella 11

11. Leaves long and slenderly acuminate 9. Turckheimia

Leaves acute or obtuse 5. Gymnostomum

12. Lamina cells in 2 layers 12. Timmiella

Lamina cells in one layer 13

13. Peristome teeth spirally twisted 20. Barbula

Peristome teeth erect or nearly so 14

14. Leaf base obovate, strongly sheathing 15

Leaf base ovate, not sheathing the stem 16

15. Leaves toothed near apex, short pointed 8. Rhamphidium

Leaves entire, slenderly acuminate 11. Pseudosymblepharis

16. Leaves strongly toothed above 16. Leptodontium

Leaves entire or nearly so 17

17. Leaf margins revolute 19. Didymodon

Leaf margins plane 18

18. Leaves Ungulate, apex rounded 18. Weisiopsis

Leaves lanceolate, sharply pointed 10. Trichostomum

19. Costa broad, ventral surface densely filamentose 24. Aloinella

Costa narrow, without filaments 20

20. Upper lamina cells and margins bistratose 15. Trichostomopsis

Leaf cells in one layer 21

21. Leaf cells smooth 22

Leaf cells papillose 23

22. Peristome lacking, leaves bordered with several rows of paler incrassate

cells 2. Merceya

Peristome present, leaves unbordered or bordered with elongated cells

22. Streptopogon

23. Leaf apex rounded, peristome very rudimentary 6. Husnotiella

Leaf apex pointed, peristome well developed 24

24. Peristome teeth spirally twisted 25. Tortula

Peristome teeth not or scarcely twisted 23. Desmatodon

1. ANOECTANGIUM Schwaegr., Suppl. I1: 33. 1811.

Slender plants in bright green tufts or cushions. Leaves lanceo-

late or oblong, crispate when dry; costa subpercurrent; cells small,

papillose, more elongate and pellucid below. Setae lateral, elongate,

slender; capsule oblong; lid obliquely rostrate; peristome none;

calyptra cucullate.

84 FIELD IANA: BOTANY, VOLUME 25

1. Leaves lanceolate, gradually acuminate 2

Leaves oblong, broadly pointed 3

2. Plants bright or yellowish green above, upper leaf cells obscure, densely papillose

2. A. compactum

Plants glaucous green above, upper leaf cells distinct, less papillose

3. A. incur vans

3. Leaves apiculate 1. A. euchloron

Leaf apex obtusely rounded 4

4. Lamina cells often in 2 layers 5. A. obtusifolium

Lamina cells in one layer 4. A. arizonicum

1. ANOECTANGIUM EUCHLORON (Schwaegr.) Mitt., Journ. Linn.

Soc. 12: 176. 1869.

Gymnostomum euchloron Schwaegr., Suppl. 22: 83. 1827.

Stems about 1 cm. high, radiculose below, branched. Leaves

spirally contorted and somewhat crispate when dry, to 1.25 mm.

long, narrowly oblong, keeled, obtuse, mucronate; margins plane,

papillose-crenulate; costa subpercurrent, scabrous on back above;

upper cells about 5 /z, densely papillose, obscure, more pellucid and

distinct at base and rectangular toward costa. Setae lateral, to

6 mm. long; capsule ovoid-cylindric, barely 1 mm. long. (Fig. 38,

A-D.)

Dept. Huehuetenango : Standley 821+3 4. Dept. San Marcos: Standley 68696.

Dept. Quezaltenango: Standley 84569, 84597, 85142; Steyermark 33619. Dept.

Retalhuleu: Standley 88158. Dept. Sacatepequez : Standley 58637. Dept. Chimal-

tenango: Standley 57918, 6163 4, 62050. Dept. Guatemala: Standley 59689.

Dept. Zacapa: Steyermark 42459. Dept. Jalapa: Steyermark 32190. Dept. Santa

Rosa: Standley 78361.

Distribution: Wide in tropical America, Hawaii, Malaysia, Africa.

On banks and rocks at moderate altitudes. The broadly pointed,

apiculate leaves are distinctive in comparison with the other local

species.

2. ANOECTANGIUM COMPACTUM Schwaegr., Suppl. I1: 36. 1811.

Anoectangium condensatum Schimp., Besch. Prodr. Bryol. Mex. 16. 1871.

Compactly tufted plants, bright or yellowish green above, brown

below. Stems to 4 cm. or more long. Leaves strongly contorted

when dry, 1-1.75 mm. long, linear-lanceolate, keeled, short acumi-

nate; margins erect, papillose-crenulate; costa subpercurrent, papil-

lose on back above; upper cells 6-8 M, densely papillose, obscure,

inner basal cells short rectangular, pellucid, incrassate. Seta about

BARTRAM: MOSSES OF GUATEMALA

85

10 mm. long; capsule ovoid-cylindric, 1-1.5 mm. long. (Fig. 38,

E-G.)

Dept. Huehuetenango : Standley 82880. Dept. San Marcos: Steyermark 85696,

35823, 35824; Standley 85392. Dept. Quezaltenango : Standley 84182, 84751,

85263; Steyermark 34?65a, 34?66a. Dept. Suchitepequez: Steyermark 4683 la.

Dept. Sacatepequez: Standley 58931. Dept. Solola: Steyermark 46930. Dept.

Chimaltenango: Standley 61540. Dept. El Progresso: Steyermark 43058, 43339,

43660, 43661. Dept. Jalapa: Steyermark 32438.

Distribution: Greenland, northern United States, Mexico, South

America, Africa, Asia, New Zealand.

On banks and rocks mostly at high altitudes. The distinctions

between A. condensatum and A. compactum are so tenuous that I

doubt if they can be separated in practice. I agree with The"riot

(29, p. 94) that they are conspecific.

3. ANOECTANGIUM INCURVANS (Schimp.) Bartr., Bryol. 49: 111.

1946.

Gymnostomum incurvans Schimp. in Besch., Prodr. Bryol. Mex. 15. 1871.

Stems 2-3 cm. high, densely reddish tomentose below, dark green

and often glaucous above. Leaves strongly incurved when dry,

B

G

FIGURE 38

A-D, Anoectangium euchloron: A, plant, X 1 ; B, leaf, X20; C, apex of leaf, X 134;

D, upper leaf cells and margin, X270.

E-G, Anoectangium compactum: E, leaf, X20; F, apex of leaf, X134; G, upper

leaf cells and margin, X270.

H-J, Anoectangium incurvans: H, leaf, X20; I, apex of leaf, X134; J, upper

leaf cells and margin, X270.

86 FIELDIANA: BOTANY, VOLUME 25

2-2.4 mm. long, narrowly linear-lanceolate, acuminate; margins

erect, minutely papillose-crenulate; costa percurrent; upper cells

distinct, rounded, incrassate, lightly papillose, diameter 8-10 n,

inner basals rectangular, smooth, pellucid. Inflorescence lateral

(TheYiot 27, pt. 3, p. 8); sporophyte not seen. (Fig. 38, H-J.)

Dept. Huehuetenango : Standley 81205 (as Hymenostylium recurvirostrum) .

Dept. Zacapa: Steyermark 1*3169.

Distribution: Mexico.

On rock and bank at moderate altitude. The leaf structure of

this species is quite similar to that of Hymenostylium recurvirostrum

'but the plane leaf margins and lateral inflorescence suggest that it

may be more aptly included in Anoectangium.

4. ANOECTANGIUM ARIZONICUM Bartr., Moss Fl. No. Amer. I3: 192.

1938.

Slender plants in dense cushions, bright green above, brown

below. Stems to 2.5 cm. long, branched. Leaves crispate when

dry, to 1 mm. long, oblong-ligulate, carinate-concave, obtusely

rounded, not apiculate; margins erect, papillose-crenulate; costa

ending below apex, rough on back; upper cells small, papillose,

incrassate, elongate and short rectangular near costa at base. Fruit

unknown. (Fig. 39, A-C.)

Dept. El Quiche: Sharp 21*91*. Dept. El Progresso: Sharp 5099.

Distribution: Arizona, Mexico.

Moist boulders and bark of trees at medium to rather high alti-

tudes. A wide range extension of a species previously known only

from Arizona and northern Mexico.

5. ANOECTANGIUM OBTUSIFOLIUM (Broth. & Par.) Grout, Moss Fl.

of No. Amer. I3: 150. 1938.

Molendoa obtusifolia Broth. & Par., Rev. Bryol. 40: 36. 1913.

Forming dense tufts or mats, glaucous green above, brown

below. Stems branched, brittle, sparingly radiculose below, to

1.5 cm. high. Leaves incurved when dry, 1.5-2 mm. long, oblong-

ligulate, obtuse; margins plane, papillose-crenulate; costa ending

below apex; upper cells obscure, papillose, diam. 6-8 n, often in

two layers, basal cells short rectangular with firm, pale, pellucid

walls. (Fig. 39, D-F.)

Dept. Baja Verapaz: Sharp 2805.

BARTRAM: MOSSES OF GUATEMALA

87

Distribution: Arizona, Mexico.

On tree at moderately low altitude. As these plants lack fruit

the determination remains questionable but the bluntly rounded,

plane margined leaves with the lamina cells here and there in two

layers suggest this species.

2. MERCEYA Schimp., Syn. Ed. 2. 852. 1876.

Rather robust plants with branched stems. Leaves ligulate,

obtuse, bordered with 3-8 rows of incrassate, often colored cells;

upper leaf cells irregularly rounded, smooth, basal cells rectangular;

costa ending below apex, in cross section showing a stereid core

with 1 or 2 layers of large, thin walled cells on the ventral surface.

Seta terminal, slender; capsule erect, ovoid-cylindric; lid conic-

rostrate, peristome none.

1. MERCEYA LIGULATA (Spruce) Schimp., Syn. Ed. 2. 852. 1876.

Encalypta? ligulata Spruce, Muse. Pyren. No. 331. 1845; Trans. Bot. Soc.

3s: 187. 1850.

Weisia agoyanensis Mitt., Journ. Linn. Soc. 12: 135. 1869,

Weisia cataracta Mitt., Journ. Linn. Soc. 12: 135. 1869.

Merceyopsis mexicana Bartr., Journ. Wash. Acad. Sci. 18, no. 21: 577. 1928.

B

\

FIGURE 39

A-C, Anoectangium arizonicum: A, plant, Xl; B, leaf, X30; C, apex of leaf,

X110.

D-F, Anoectangium obtusifolium: D, plant, Xl; E, leaf, X28; F, part of cross

section of leaf, X270.

G-H, Tortella tortuosa: G, plant, Xl; H, leaf, XlO.

88 FIELD IANA: BOTANY, VOLUME 25

Stems 1-5 cm. high, densely tufted. Leaves crowded, contorted

when dry, 1.5-5 mm. long, ligulate or oblong-spatulate, rounded,

obtuse or acute, often bordered with 38 rows of thick walled, colored

cells; costa ending below apex or nearly percurrent; upper cells

quadrate or slightly elongate, with firm walls, 10-15 n, more in-

crassate and thicker toward margins, basal cells oblong, thin walled ;

margins entire, slightly recurved below. Seta yellowish, erect,

2-5 mm. long; capsule ovoid-cylindric, lightly furrowed when dry

and empty, urn 1-1.5 mm. long; peristome lacking; lid erect, short

conic-rostrate. (Fig. 40, A-E.)

Dept. Quezaltenango: Standley 67426, 674.28, 83336, 83385, 83389, 86049,

86053; Steyermark 34663, 34665, 34657, 34977. Dept. Chiquimula: Steyermark

30977.

Distribution: Tennessee, Arizona, Mexico, Costa Rica, Ecuador,

Europe.

On wet banks and rocks at high altitudes. As far as I can see

there is but one species in North America extending south to Ecuador.

The leaves vary considerably in outline from spatulate and broadly

rounded to oblong and acute. The leaf border of incrassate cells

is variable, often very prominent and again hardly noticable.

3. HYMENOSTOMUM R. Brown, Trans. Linn.

Soc. 122:573. 1819.

Small plants with crisped leaves; partial to calcareous habitats.

Differing from Weisia only in the lack of a peristome; the mouth

of capsule covered with a fugacious membrane developed from the

top of the columella.

1. HYMENOSTOMUM JAMESONI (W. Arn.) Broth., E. & P. Nat.

Pflanzenf. I3: 386. 1902.

Gymnostomum Jamesoni W. Arn., in Wern. Transact. 5: 200(7).

Small, laxly gregarious plants. Stems 2-3 mm. high. Leaves

strongly contorted when dry, erect-spreading when moist, to 2 mm.

long, linear-lanceolate from a pale, ovate base, acute, mucronate,

entire; margins narrowly involute above base; costa pale, 50-60 n

wide below, excurrent in a sharp, pale mucro; upper cells small,

opaque, obscure, papillose, basal cells rectangular, pellucid, smooth.

Seta 4-5 mm. long, pale yellow; capsule erect, ovoid, urn 1-1.1 mm.

long;' peristome lacking; lid conic-rostrate, 0.5 mm. long. (Fig. 40,

F-H.)

BARTRAM: MOSSES OF GUATEMALA

89

Dept. Quezaltenango : Sharp 1947. Dept. Zacapa: Steyermark

Distribution: Brazil.

On soil at moderate to high altitudes. A critical study of the

tropical American species of this genus is essential before they can

be named with any certainty. As far as I can see there are no im-

portant differences between Sharp's 1947 and plants from Brazil,

determined as H. Jamesoni. Steyermark's 42746 is sterile and there-

fore questionable, but it may be remarked that the leaves here

match very closely the type material of H. mexicanum Card, and

also those of H. tortile Bry. Eur.

4. WEISIA Hedw., Sp. Muse. 64. 1801.

Stems short, branched. Lower leaves small, upper much longer,

crisped when dry, narrowly lanceolate from a broader base; upper

margins involute; costa short excurrent; basal cells hyaline, upper

cells small, opaque, papillose. Seta elongate; capsule erect; peristome

teeth variable, entire or divided.

Costa rarely over 50/* wide below 1. W. viridula

Costa 70-75 n wide below 2. W. jamaicensis

FIGURE 40

A-E, Merceya ligulaia: A, plant, Xl; B, leaf, X16; C, apex of leaf, X134;

D, upper leaf cells and margin, X270; E, capsule, X14.

F-H, Hymenostomum Jamesoni: F, plant, Xl; G, leaf, X14; H, capsule, X8.

I-L, Weisia viridula: I, leaf, X14; J, apex of leaf, X134; K, capsule, X8;

L, part of peristome, X134.

90 FIELDIANA: BOTANY, VOLUME 25

1. WEISIA VIRIDULA Hedw., Sp. Muse. 68. 1801.

Plants bright green or yellowish green in low tufts. Stems to

5 mm. or more high. Upper leaves lanceolate, sharply pointed,

2.5-3 mm. long; margins strongly involute and entire above; costa

narrow, excurrent, 35-45 /x wide at base; upper cells 6-7 //, densely

papillose, opaque, basal cells rectangular, hyaline, smooth. Seta

3-7 mm. high; capsule oblong-cylindric; peristome teeth variable,

short and truncate or lanceolate. (Fig. 40, I-L.)

Dept. Jutiapa: Standley 75228a.

Distribution : Cosmopolitan but apparently infrequent in Central

America.

On soil at moderate altitude.

2. WEISIA JAMAICENSIS (Mitt.) Grout, Moss Fl. of No. Amer. 1:

157. 1938.

Tortula jamaicensis Mitt., Journ. Linn. Soc. 12: 147. 1869.

Trichostomum Purpusi Card., Rev. Bryol. 36: 73. 1909.

Trichostomum involvens Card., Rev. Bryol. 40: 34. 1913.

Stems to 1 cm. or more high. Leaves 3-4 mm. long, linear from

a broader oblong-ovate, hyaline base, sharply pointed; margins

strongly involute; costa strong, 75-80 n wide below, excurrent;

basal cells rectangular, upper cells small, dense, opaque, papillose.

Seta 8-15 mm. long; capsule cylindric, urn 2-2.5 mm. long; peristome

teeth divided nearly to base into 32 filiform, papillose forks. (Fig. 41,

A-D.)

Dept. Alta Verapaz: Standley 89664. Dept. Sacatepequez: Standley 58985a.

Distribution: Southern United States, Mexico, West Indies.

On banks and rocks at moderate altitude. In no. 89664 the leaves

are typically narrow and sharply pointed while in no. 58985a they

are broader and bluntly pointed, but I believe both collections may

safely be referred here.

5. GYMNOSTOMUM Hedw., Sp. Muse. 30. 1801.

Dioicous; small, slender plants usually occurring in limestone

regions. Leaves narrowly lanceolate, contorted when dry; basal

cells rectangular, hyaline, upper cells small, papillose, obscure. Seta

elongate; capsule erect, ovoid; peristome lacking; lid not persistent.

Stems short, leaves Ungulate, often obtuse 2. G. calcareum

Stems elongate, leaves lanceolate, acute 1. G. aeruginosum

BARTRAM: MOSSES OF GUATEMALA

91

1. GYMNOSTOMUM AERUGINOSUM Sm., Fl. Brit. 3: 1163. 1804.

Gymnostomum rupestre Schleich., Cat. 29. 1807.

Slender plants in compact cushions, green above, pale brown

below, incrusted with a calcareous deposit. Stems about 3 cm.

high. Leaves to 1.6 mm. long, narrowly linear-lanceolate, broadly

acute; margins plane; costa stout, about 60 p. wide below, ending

below apex; upper cells 10-12 n, obscure, densely papillose, rec-

tangular and hyaline at base. Seta short; capsules short oblong.

(Fig. 41, E-G.)

Dept. Huehuetenango : Standley 82781 (sterile).

Distribution: Southern Canada to Texas and Arizona, Europe,

Asia, China, Japan.

On damp, calcareous bank at moderate altitude. Not recorded

before from below the United States border.

2. GYMNOSTOMUM CALCAREUM Nees & Hornsch., Bry. Germ. 1:

53. 1823.

Stems short, 4-8 mm. high. Leaves less than 1 mm. long, similar

to those of G. aeruginosum but often obtusely rounded at apex.

FIGURE 41

A-D, Weisia jamaicensis: A, plant, XI; B, leaf, X14; C, apex of leaf, X134;

D, capsule, X8.

E-G, Gymnostomum aeruginosum: E, plant, Xl; F, leaf, X26; G, apex of leaf,

X134.

H-L, Gymnostomum calcareum: H, plant, Xl; I and J, leaves, X26; K, apex

of leaf, X134; L, upper leaf cells and margin, X270.

92 FIELDIANA: BOTANY, VOLUME 25

Seta 4-5 mm. long; capsule short ovoid, urn about 0.7 mm. long.

(Fig. 41, H-L.)

Dept. Huehuetenango: Standley 656^.1, 65661, 6580^. Dept. Sacatepequez :

Standley 58936. Dept. Guatemala: Standley 62866.

Distribution: Newfoundland to California south to West Virginia

and Arizona, Europe, Asia, Africa.

On damp banks and cliffs at moderate to rather high altitudes.

These collections are typical and several of them well fruited.

6. HUSNOTIELLA Card., Rev. Bryol. 36:71. 1909.

Plants small, closely tufted. Stems short, erect. Leaves ovate-

lingulate, rounded at apex; margins entire, revolute; costa ending

below apex, with stereids on dorsal side only; cells small, papillose,

obscure, rectangular and smooth below. S^ta elongate; capsules

erect; peristome very rudimentary or none.

1. HUSNOTIELLA REVOLUTA Card., Rev. Bryol. 36: 71. 1909.

Stems to 1 cm. high, olive green above, brown below. Leaves

strongly contorted when dry, about 1 mm. long, lingulate, rounded

at apex; margins revolute except near base; costa strong, ending

below apex, wider and spurred above; cells rounded, mammillose,

8-10 n, basal cells short rectangular, hyaline. Seta 6-8 mm. long;

capsules erect, subcylindric. (Fig. 42, A-C.)

Dept. Jalapa: Steyermark 32156.

Distribution: Southwestern United States, Mexico.

On shaded bricks of path at moderate altitude. The short,

lingulate, rounded leaves with revolute margins and thick, spurred

costa are distinctive characters.

7. HYMENOSTYLIUM Brid., Bryol. Univ. 2: 81. 1827.

Dioicous; slender, compactly tufted plants, green above, brown

below. Stems long, fragile, tomentose. Leaves curved when dry,

narrowly lanceolate, acuminate; margin recurved below; costa end-

ing below apex; cells rounded, papillose, rectangular below. Seta

elongate; capsules ovoid, wide mouthed; peristome lacking; lid

remaining attached to columella after dehiscence.

BARTRAM: MOSSES OF GUATEMALA

93

1. HYMENOSTYLIUM RECURVIROSTRUM (Hedw.) Dix., Rev. Bryol.

etLich. 6:96. 1933.

Gymnostomum recurvirostrum Hedw., Sp. Muse. 33. 1801.

Stems to 3-4 cm. long or longer. Leaves crowded, to 2 mm. or

more long, keeled; margins recurved on one or both sides below;

upper cells distinct, rounded or angular, diam. 8-10 n, papillose, in-

crassate, rectangular basal cells few. Seta 8-10 mm. long; capsule

erect and slenderly beaked, lid persistent. (Fig. 42, D-F.)

Dept. Alta Verapaz: Steyermark 44575; Standley 70882a. Dept. Huehue-

tenango: Standley 81621, 81672a, 81688; Steyermark SOOOOa, 50146a, 50152, 50282a,

51235. Dept. San Marcos: Steyermark 35701, 36888a. Dept. Zacapa: Steyermark

42934.

Distribution: Labrador to Alaska south to South Carolina, Cali-

fornia and West Indies, wide in Europe, Asia, New Zealand.

On calcareous rocks and bluffs at medium to high altitudes.

These collections are all sterile and although variable have in com-

mon narrowly lanceolate leaves with at least one margin recurved

below and distinct upper cells. Some of the forms with longer,

crispate leaves are quite similar to Amphidium Mougeotii (Bry.

Eur.) but it seems more conservative to include them here until

the problem can be clarified by fertile collections.

B

FIGURE 42

A-C, Husnotiella revoluta: A, plant, Xl; B, leaf, X26; C, apex of leaf, X120.

D-F, Hymenostylium recurvirostrum: D, leaf, X26; E, apex of leaf, X120;

F, upper leaf cells and margin, X270.

G-K, Rhamphidium dicranoides: G, plant, Xl; H and I, leaves, X26; J, apex

of leaf, X120; K, leaf cells and margin near apex, X270.

94 FIELDIANA: BOTANY, VOLUME 25

8. RHAMPHIDIUM Mitt., Journ. Linn. Soc. 12: 45. 1869.

Small, slender plants. Leaves lanceolate from a sheathing base;

margins erect; costa ending below apex; basal cells linear, upper

cells subquadrate, smooth. Seta elongate; capsule nodding, oblong-

cylindric; peristome teeth divided nearly to base into 2 filiform,

papillose forks; lid long beaked.

r-

1. RHAMPHIDIUM DICRANOIDES (C. M.) Bartr., Bryol. 49: 112. 1946.

Leptotrichum dicranoides C. M., Syn. 2: 612. 1851.

ITrichostomum hyophilaceum C. M., Bull. Herb. Boiss. 5: 191. 1897.

Small, slender, yellowish plants. Stems erect, to 10 or 12 mm.

high, reddish. Leaves crispate when dry, spreading when moist,

scarcely 1 mm. long, lanceolate from a short, broad, clasping base,

concave, obtuse; margins erect, distantly denticulate above middle;

costa ending below apex, toothed on back above; upper cells sub-

quadrate to slightly elongate, smooth, distinct, diam. about 10 //,

basal cells narrowly rectangular. Seta slender, reddish, 1 cm. long;

capsule suberect to nodding, urn 1.5-1.8 mm. long. (Fig. 42, G-K.)

Dept. Retalhuleu: Standley 87201, 87206. Dept. Suchiate: Svihla 2837,

281t5, 2856.

Distribution: Alabama, Louisiana, Mexico, Central America,

West Indies, South America.

On wet banks at moderate altitudes. These collections differ

in no way from Liebman's original gathering from Mexico. There

is nothing very distinctive in the description of Trichostomum

hyophilaceum C. M. and none of the type material is available. In

all probability it will prove to be a synonym of R. dicranoides.

9. TURCKHEIMIA Broth., Ofv. F. Vet-

Soc. Forh. 52, no. 7: 2. 1909-1910.

Dioicous; very small, pale green plants. Leaves linear-lanceo-

late, subulate-acuminate, entire; costa percurrent; upper cells small,

rounded, basal cells oblong, hyaline. Seta elongate; capsule erect,

cylindric; peristome in our species' none or rudimentary.

1. TURCKHEIMIA GUATEMALENSIS Broth., Ofv. F. Vet.-Soc. Forh.

52, no. 7: 2. 1909-1910.

Stems 1-2 mm. high, branched. Leaves few, flexuous-spreading

when dry, about 2 mm. long, linear-lanceolate; costa ending in the

BARTRAM: MOSSES OF GUATEMALA 95

subula; upper cells rounded-hexagonal, 8 M, obscurely mammillose,

occasionally in 2 layers at margins above, basal cells lax, oblong,

hyaline. Seta 5-8 mm. long, slender; capsule cylindric, urn 0.5-1

mm. long, tapering to seta. Lid and peristome not seen. (Fig. 43,

A-D.)

West Guatemala: Livingston, H. J. Turckheim 1908.

Endemic.

This species is described as having no peristome but Mrs. Britton's

notes accompanying part of the type collection in the New York

Botanical Garden Herbarium indicate that traces of a peristome

remain. It is evidently rare and has never been collected again to

my knowledge.

10. TRICHOSTOMUM Hedw. emend. Bruch, Flora 2: 393. 1829.

Dioicous; small to medium sized, tufted plants. Leaves narrow,

crisped when dry; margins plane; costa percurrent or excurrent,

with dorsal and ventral stereid bands; upper cells small, papillose,

basal cells rectangular, pellucid. Seta elongate, erect; capsule

cylindric; peristome teeth 16, erect, papillose, bifid or rudimentary;

lid conic-rostrate.

1. Leaves linear-lanceolate, acute 1. T. cylindricum

Leaves wider, apex obtuse, mucronate 2

2. Stems 1-2 mm. high, leaves less than 1.5 mm. long 3. T. pygmaeum

Stems 5-10 mm. high, leaves 3-4 mm. long 2. T. brachydontium

1. TRICHOSTOMUM CYLINDRICUM (Bruch) C. M., Syn. 1: 586.

1849.

Weisia cylindrica Bruch, in Brid. Bryol. Univ. 1 : 806. 1826.

Plants loosely tufted, yellowish above, brown below. Stems to

2.5 cm. high. Leaves crispate when dry, brittle, linear-lanceolate

from an erect, pale base, 3-4 mm. long, acute; margins plane, often

sinuate or notched above; costa excurrent in a short, pellucid

apiculus; upper cells rounded, papillose, obscure, basal cells rec-

tangular. Seta 1-1.5 cm. long, slender, yellowish; capsules cylindric,

erect or slightly curved; peristome teeth short, erect, irregular.

(Fig. 43, E-G.)

Dept. Alta Verapaz: Standley 69650a. Dept. San Marcos: Standley 66063;

Steyermark 36115a, S6120. Dept. Sacatepequez : Standley 58832. Dept. Chimal-

tenango: Standley 58?40a, 60962a. Dept. Guatemala: Standley 58J^85a, 80601.

Distribution: Greenland to Manitoba south to North Carolina

and Arizona, South America, Europe, Asia, Africa, Japan.

96

FIELDIANA: BOTANY, VOLUME 25

I

FIGURE 43

A-D, Turckheimia guatemalensis: A, plant, XI; B, leaf, X14; C,

X120; D, upper leaf cells and margin, X270.

E-G, Trichostomum cylindricum: E, plant, Xl; F, leaf, X14; G,

X120.

H-J, Trichostomum brachydontium: H, plant, Xl; I, leaf, X14; J,

X120.

apex of

apex of

apex of

leaf,

leaf,

leaf,

On banks,, rocks and logs at medium to high altitudes. These

collections vary considerably but no more so than in the north where

the species is proverbially plastic. Some of the collections show

the pellucid basal cells extending up the margins as in Tortella but

this anomalous feature is neither constant nor well marked.

2. TRICHOSTOMUM BRACHYDONTIUM Bruch, in Flora P. 2: 393.

1829.

Rather coarse plants in lax, brownish green tufts. Stems 1-2 cm.

high. Leaves crowded, contorted and incurved when dry, 3-4 mm.

long, narrowly oblong-lanceolate, short pointed, mucronate by the

short excurrent costa, not fragile; upper cells small, dense, very

opaque, basal cells rectangular, pellucid. Seta 1-1.2 cm. long;

capsule ovoid-cylindric; peristome short or rudimentary. (Fig. 43,

H-J.)

Dept. Huehuetenango : Standley 81181; Steyermark 50282. Dept. Zacapa:

Steyermark ^2205. Dept. Jalapa: Standley 76^8, 77310.

Distribution: Mexico, Jamaica, Europe, Asia, Africa, Japan.

On damp banks and rocks at moderate altitudes. These collec-

tions are without fruit but the vegetative characters are in every

way similar to T. brachydontium.

BARTRAM: MOSSES OF GUATEMALA 97

3. TRICHOSTOMUM PYGMAEUM Bartr., Bryol. 49: 112. 1946.

Small, dull green, densely tufted plants. Stems very short,

1-2 mm. high. Lower leaves minute, the upper to 1.4 mm. long,

crisped when dry; oblong-lanceolate, concave, obtuse, mucronate;

margins erect, papillose-crenulate; costa ending in or just below

apex, papillose on back above; upper cells densely papillose, obscure,

diam. 6-8 M, basal cells rectangular, smooth, pellucid. (Fig. 44, A-C.)

Dept. Alta Verapaz: along Rio Icvolay, north and northwest of Finca Cubil-

guitz to Quebrada Diabalo, alt. 300-350 m., Steyermark M770, TYPE.

Endemic.

While lacking any marked characters, this species seems to be

clearly different from any of its local associates in the short stems

and small, bluntly pointed, concave leaves.

11. PSEUDOSYMBLEPHARIS Broth., E. & P. Pflanzenf.

Ed. 2, 10:261. 1924.

Plants fairly robust, growing in deep tufts. Stems branched.

Leaves strongly curled when dry, lanceolate from an obovate, sheath-

ing base, acuminate; margins erect; costa excurrent; basal cells

linear, hyaline, upper cells small, papillose. Seta elongate; capsule

cylindric; peristome teeth erect, irregularly cleft.

1. PSEUDOSYMBLEPHARIS CIRCINATA (Schimp.) Broth., E. & P.

Pflanzenf. Ed. 2, 10: 261. 1924.

Syrrhopodon rircinatus Schimp., in Besch., Prodr. Bryol. Mex. 28. 1871.

Stems to 3 cm. or more high, yellowish green, brown below.

Leaves crowded, the long points widely spreading and circinate when

dry, rather brittle, to 10-12 mm. long, narrowly linear-lanceolate

from an erect, obovate, strongly clasping base, gradually subulate

acuminate; upper margins minutely papillose-crenulate; costa ex-

current in a smooth, sharp point; upper cells subquadrate, densely

papillose, obscure, basal cells linear, hyaline, becoming incrassate

toward shoulders with sinuose lateral walls, long and narrow toward

margins and often forming an indistinct border to above leaf

shoulders. Fruit unknown. (Fig. 44, D-G.)

Dept. Alta Verapaz: Standley 71601. Dept. Huehuetenango: Steyermark

48575 (as P. subulata sp. nov.), 1*8928(1, SOltfb. Dept. Chimaltenango: Standley

58781 c. Dept. Chiquimula: Steyermark 31693.

Distribution: Mexico, Costa Rica.

98

FIELDIANA: BOTANY, VOLUME 25

f

B

/

H

FIGURE 44

A-C, Trichostomum pygmaeum: A, plant, Xl; B, leaf, X14; C, apex of leaf,

X120.

D-G, Pseudosymblepharis circinata: D, plant, Xl; E, leaf, X6; F, apex of

leaf, X120; G, upper leaf cells and margin, X270.

H-J, Timmiella anomala: H, plant, Xl; I, leaf, X12; J, apex of leaf, X120.

On trees and rocks at moderate altitudes. Distinguished from

Trichostomum principally by the obovate, clasping leaf base. The

narrow marginal cells of the leaf base and the slender, subulate leaf

point have a suggestive parallel in Trichostomum angustatum (Mitt.)

Fleisch. of Malaysia.

12. TIMMIELLA (DeNot.) Limpr., Laubm. 1: 590. 1888.

Timmiella Sec. of Trichostomum DeNot., Cron. Briol.

Ital. 1 : 14. 1886.

Plants fairly robust, laxly tufted. Leaves strongly contorted

with incurved margins when dry, oblong-lanceolate, toothed above;

costa strong; upper cells rounded, in 2 layers except near margins,

mammillose on ventral face, basal cells rectangular, hyaline. Seta

elongate; capsule cylindric; peristome teeth from a low basal mem-

brane, divided into 32 filiform, papillose forks, erect or slightly

twisted; lid conic-rostrate.

1. TIMMIELLA ANOMALA (Bry. Eur.) Limp., Laubm. 1: 592. 1888.

Barbula anomala Bry. Eur. fasc. 13-15. 1842.

Autoicous or synoicous; stems 1 cm. or more high. Upper

leaves broadly linear from an ovate base, 4-5 mm. long, concave,

BARTRAM: MOSSES OF GUATEMALA 99

acute; margins erect or inflexed, serrate toward apex; costa broad

below, ending near apeX; basal cells hyaline, upper cells 7-10 n,

the dorsal layer smooth. Seta 15-20 mm. long, slender; capsules

narrowly cylindric, curved, urn 4 mm. long; lid 1.5 mm. long;

peristome teeth about 1 mm. long, slightly twisted. (Fig. 44, H-J.)

Dept. Sacatepequez: Standley 65266.

Distribution: California, Arizona, Mexico, Europe, Asia.

On damp bank at high altitude. The narrowly pointed leaves

suggest that this may be the form described as T. subanomala

Besch. from Mexico, but I doubt that it is specifically distinct.

13. TORTELLA (C. M.) Limp., Laubm. 1: 599. 1890.

Barbula Sec. Tortella C. M., Syn. 1 : 599. 1849.

Medium sized plants growing in mats. Stems erect, radiculose.

Leaves usually strongly crisped when dry, widely spreading when

moist, linear-lanceolate; margins erect or inflexed; costa strong,

excurrent, glossy on back when dry; upper cells small, rounded,

papillose, basal cells rectangular, hyaline, extending up margins in

a more or less distinct border. Seta elongate; capsules erect or nearly

so; peristome teeth split to base into 32 filiform, papillose forks,

spirally twisted; lid beaked.

1. Hyaline border of broad, rectangular cells extending more than half way up

leaf 1. T. Richardsii

Hyaline border of narrow cells confined to basal part of leaf 2

2. Leaf apex abruptly rounded and mucronate 2. T. guatemalensis

Leaf apex slenderly subulate-acuminate 3

3. Robust plants, stems to 4 cm. or more high 3. T. tortuosa

Small plants, stems under 1 cm. high 4. T. mollissima

1. TORTELLA RICHARDSII Bartr., Bryol. 49: 112. 1946.

Dull, olive green plants in dense mats. Stems to 2 cm. high,

simple or branched, densely foliate, slightly radiculose below.

Leaves flexuous-spreading moist and dry, slightly contorted, 4-7

mm. long, rather quickly linear-lanceolate from an ovate base,

sharply acute at apex, channelled, bordered to above middle with

3-4 rows of rectangular, hyaline cells; margins erect, minutely

papillose-crenulate above; costa brown, stout, short excurrent in a

pellucid apiculus; upper cells rounded, obscure, diameter about 6 /z>

basal cells rectangular, smooth, pellucid. Fruit unknown. (Fig. 45,

A-D.)

100

FIELDIANA: BOTANY, VOLUME 25

B

FIGURE 45

A-D, Tortella Richardsii: A, moist plant, XI; B, leaf, X8; C, one side of leaf

about half way up, X120; D, apex of leaf, X120.

E-G, Tortella guatemalensis: E, moist plant, Xl; F, leaf, X8; G, apex of leaf,

X120.

H-J, Tortella mollissima: H, moist plant, Xl; I, leaf, X8; J, apex of leaf, X120.

Dept. Alta Verapaz: Vicinity of Cubilguitz, 1^-2 miles south of Cubilguitz,

alt. 300-350 m., Steyermark M389, TYPE.

Distribution: British Honduras.

A very distinct and handsome species which may well prove to

be generically distinct when the fruit is known. The rectangular,

hyaline border cells average about 15 n wide and up to 60 /j. long

and merge gradually with the basal areolation instead of con-

tinuing in a distinct marginal band below as in Pleurochaete.

I take pleasure in naming this unique plant for Mr. Donald

Richards whose labors in packeting and labeling the extensive

Guatemalan collections have greatly facilitated the study of the

material.

2. TORTELLA GUATEMALENSIS Bartr., Bryol. 49: 113. 1946.

Plants with the habit of T. tortuosa (Turn.) Limp. Stems

scarcely 5 mm. high, laxly tufted, pale green. Leaves crowded,

strongly contorted with circinate points when dry, laxly spreading

when moist, to 5.5 mm. long, linear-lanceolate from a scarcely wider

base, abruptly rounded and mucronate at apex, channelled above,

lamina fragile and often broken; margins plane, lightly undulate

BARTRAM: MOSSES OF GUATEMALA 101

above, minutely papillose-crenulate; costa very strong below, about

90 M wide, excurrent in a sharp, pellucid mucro; upper leaf cells

densely papillose, obscure, diam. 8-10 M, basal cells laxly rectangular,

about 15 M wide, extending up margins but much less conspicuously

than in T. tortuosa. Sporophyte unknown. (Fig. 45, E-G.)

Dept. Alta Verapaz: Standley 8987 ^ (as Trichostomum angustinerve Card.).

Dept. Huehuetenango : Standley 82556a (as T. tortuosa). Dept. Retalhuleu:

Standley 88397 TYPE (as T. tortuosa). Dept. Chimaltenango: Standley. Dept.

Guatemala: Standley 80361 (as T. tortuosa).

Endemic.

On trees and damp banks at moderate altitudes. I thought at

first that these plants could be included in T. tortuosa but a more

careful comparison shows that this is not practicable. The differences

are not marked but consistent. In the Guatemalan plants the leaves

are abruptly rounded at the apex, the basal cells are wider and more

lax and less conspicuously extended up the margins.

3. TORTELLA TORTUOSA (Hedw.) Limp., Laubm. 1 : 604. 1890.

Tortula tortuosa Hedw., Sp. Muse. 124. 1801.

Robust brownish plants in dense tufts, yellow at tips, dark brown

below. Stems to 4 cm. high, branched. Leaves widely spreading,

strongly contorted when dry, to 6 mm. long, gradually linear-lanceo-

late from a short, ovate base, entire, subulate-acuminate, points

fragile and usually broken off; margins erect, undulate; costa strong,

excurrent; basal cells linear, porose, pellucid, extending well up

margins to above leaf-shoulders; lamina cells small, obscure, densely

papillose. Seta reddish, to 3 cm. long; capsule cylindrical; peristome

teeth red, twisted in several turns. (Fig. 39, G-H.)

Dept. El Quiche: Sharp 5342. Dept. Huehuetenango: Sharp 4811.

Distribution: Northern United States and Canada south to the

Gulf of Mexico.

On calcareous bluffs and boulders at moderately high altitudes.

Although sterile, these collections are thoroughly typical of the species

which has not been recorded before in North America south of the

Mexican border.

4. TORTELLA MOLLISSIMA Broth., Bryol. 50: 203. 1947.

Plants densely tufted forming extensive, low mats, yellowish or

brownish green. Stems rarely over 1 cm. high. Leaves crowded,

102 FIELDIANA: BOTANY, VOLUME 25

strongly curled toward tips when dry, 4-8 or 9 mm. long, narrowly

linear from a short, pale, shining, oblong-ovate base, tapering

gradually to slender, subulate-acuminate tips; margins erect, not

undulate; costa short-excurrent; upper cells minute, obscure, densely

papillose, basal cells rectangular, hyaline, thin-walled, extending

upward along margins higher than toward costa but not forming a

conspicuous border. Seta 13 mm. long, reddish, slender; capsule

erect, cylindrical, urn 2 mm. long; peristome not seen. (Fig. 45,

H-J.)

Dept. Peten: Lundell 2863. Dept. Alta Verapaz: Sharp 29 W, 3002. Dept.

Baja Verapaz: Sharp 2696. Dept. Jalapa: Steyermark 32553.

Distribution: Mexico, Porto Rico, Jamaica, Trinidad.

On boulders at relatively low altitudes. This seems to be a

typical Caribbean type extending into the lowlands of Mexico and

Guatemala in conformity with the usual distributional picture. The

only fertile plant I have seen came from Sharp's 3002, north of

Coban, Alta Verapaz, alt. 4,400 ft. The capsule is too old to show

the peristome structure but the sporophyte appears to be relatively

smaller and more slender than in T. tortuosa. In T. mollissima the

shorter stems, very narrow blades rarely over 0.22 mm. wide and

not undulate on the edges, coupled with the less conspicuous border

of narrow cells extending only a short way up the basal leaf margins,

seem to be good diagnostic features as compared with T. tortuosa.

14. PLEUROCHAETE Lindb., de Tort. 253. 1864.

Plants laxly erect, in loose tufts. Stems flexuous, branched, not

radiculose. Leaves lanceolate from an erect, sheathing base,

bordered in lower half; margins toothed above base; costa percurrent;

upper cells subquadrate, papillose, median basal cells chlorophyllose,

marginal rows rectangular, pellucid, extending up margins in a

narrow border. Seta lateral; peristome long, slightly twisted.

1. PLEUROCHAETE LUTEOLA (Besch.) TheY., Smiths. Misc. Coll. 782:

14. 1926.

Trichostomum luteolum Besch., Prod. Bryol. Mex. 34. 1871.

Robust plants forming deep, lax tufts, yellowish green above,

brown below. Stems to 8 cm. long, branched, densely foliate. Leaves

erect-spreading with tightly curled points when dry, squarrose-

spreading when moist, 4-5 mm. long, linear-lanceolate from an

erect, concave, clasping, oblong-ovate base, acuminate, with a dis-

BARTRAM: MOSSES OF GUATEMALA

103

tinct border of elongate, pellucid cells extending more than half

way up the leaf margins; margins erect, undulate, denticulate with

irregular teeth to below the leaf shoulders; costa percurrent; inner

basal cells linear with firm, pellucid lateral walls, laxly rectan-

gular and pellucid in 6 or 7 rows at margins, gradually merging

above with the small, subquadrate, densely papillose lamina cells,

border near mid-leaf 3-4 rows wide of narrowly rectangular, pellucid

cells with slightly porose, pale, incrassate walls, sharply defined

from the small, obscure lamina cells. (Fig. 46, A-D.)

Dept. Huehuetenango: Sharp 48^8, 4864, 4970, 5361.

Distribution: Mexico, Ecuador.

On calcareous boulders and soil at moderately high altitudes.

Distinguished from P. squarrosa (Brid.) Lindb. by the more robust

habit, elongated inner basal cells and the hyaline border extending

farther up the margins. No. 4864 in the above series approaches

P. squarrosa in the shorter interior basal cells but the hyaline border

extends higher up the leaf as in P. luteola.

15. TRICHOSTOMOPSIS Card., Rev. Bryol. 36: 73. 1909.

Small, tufted, green plants; stems short. Leaves crowded, crisped

when dry, lanceolate, acute; margins lightly reflexed; costa per-

FIGURE 46

A-D, Pleurochaete luteola: A, plant, X%; B, leaf, X12; C, upper leaf cells

and margin above mid-leaf, X340; D, basal leaf cells, X340.

E-G, Leptodontium excelsum: E, leaf, X12; F, cells and margin near mid-leaf,

X400; G, basal leaf cells, X400.

104 FIELDIANA: BOTANY, VOLUME 25

current, with stereid band on dorsal side only; upper cells papillose,

often bistratose, basal cells rectangular, hyaline. Seta terminal,

elongate; capsules erect; peristome teeth divided nearly to base into

2 filiform, papillose forks, slightly twisted.

1. TRICHOSTOMOPSIS DIAPHANOBASIS (Card.) Grout, Moss Fl. of

No. Amer. 1 : 228. 1939.

Didymodon diaphanobasis Card., Rev. Bryol. 37: 125. 1910.

Plants densely tufted, brownish green above, paler below. Stems

about 1 cm. high. Leaves contorted when dry, 1.5-2 mm. long,

lanceolate, acute; costa percurrent; upper cells rounded, papillose,

10-12 IJL, bistratose at margins, basal cells lax, rectangular, hyaline.

Seta 7-8 mm. long; capsule ovoid-cylindric, urn about 2 mm. long;

peristome teeth slightly twisted, about 0.5 mm. long; lid conic-

rostrate. (Fig. 47, A-D.)

Dept. Quezaltenango : Standley 664.89a in part, 66502.

Distribution: Arizona, Texas, Mexico.

On rocks at high altitude. The lax, thin walled, hyaline cells

of the leaf base and the leaf blade broader above may assist in separat-

ing this species from Didymodon Godmanianus.

16. LEPTODONTIUM Hampe, Linnaea 20: 70. 1847.

Usually dioicous; slender to robust plants in lax tufts or mats.

Stems short to very long, mostly laxly foliate. Leaves contorted

or crispate when dry, lanceolate or lingulate, usually coarsely toothed

above; costa strong, ending in or near apex, with a thick dorsal

stereid band and a thinner band on the ventral face; lamina cells

rounded, papillose, basal cells rectangular, hyaline. Perichaetial

leaves sheathing; seta erect; capsules cylindric; peristome teeth 16,

irregularly cleft; lid beaked.

1. Stems slender, often short, leaves less than 2.5 mm. long 2

Stems robust, elongate, leaves over 2.5 mm. long 4

2. Leaves minutely denticulate 1. L. filescens

Leaves coarsely serrate 3

3. Leaves oblong-lingulate 3. L. Orcutti

Leaves linear-lanceolate 2. L. Valerianum

4. Papillae of leaf cells multifid 5

Papillae of leaf cells simple or forked 6

5. Inner basal leaf cells thin-walled, hyaline 4. L. gracile

Inner basal leaf cells firm, pellucid 5. L. subgracile

BARTRAM: MOSSES OF GUATEMALA

105

6. Leaf base obovate 6. L. acutifolium

Leaf base ovate 7

7. Lamina cells dense, basal cells with straight lateral walls 7. L. excelsum

Lamina cells incrassate, basal cells with sinuose lateral walls 8

8. Leaves erect-spreading when moist, long and slenderly acuminate

10. L. ulocalyx

Leaves squarrose-recurved when moist, short acuminate 9

9. Papillae of leaf cells simple, low 9. L. sulphureum

Papillae of leaf cells high, often forked 8. L. exasperatum

1. LEPTODONTIUM FILESCENS (Hampe) Mitt., Journ. Linn. Soc. 12:

50. 1869.

Trichostomum filescens Hampe, Linnaea 32: 128. 1863.

Plants yellowish green. Stems slender, red, laxly foliate, to 1.5

cm. long. Leaves appressed and slightly contorted when dry, to

1.2 mm. long, oblong-ovate, acute, apiculate, keeled; margins re-

curved near mid-leaf, papillose-crenulate, slightly denticulate near

apex; costa ending below apex, papillose on back; inner basal cells

rectangular, incrassate, upper cells irregularly rounded, densely

papillose, obscure. Seta 8-9 mm. long; capsule erect. (Fig. 47, E-I.)

Dept. Quezaltenango : Standley 67663a.

M

FIGURE 47

A-D, Trichostomopsis diaphanobasis: A, moist plant, Xl; B, leaf, X14; C,

upper leaf cells and margin, X270; D, basal leaf cells, X270.

E-I, Leptodontium filescens: E, moist plant, X 1 ; F and G, leaves, X 14; H, apex

of leaf, X120; I, upper leaf cells and margin, X270.

J-M, Leptodontium Valerianum: J, plant, Xl; K, leaf, X14; L, apex of leaf,

X120; M, upper leaf cells and margin, X270.

106 FIELDIANA: BOTANY, VOLUME 25

Distribution: Mexico to Colombia.

On moist bank at high altitude. Rarely collected and usually

sterile. The slender stems and small, subentire leaves are distinctive.

2. LEPTODONTIUM VALERIANUM Bartr., Journ. Wash. Acad. Sci.

19: 18. 1929.

Slender, yellowish green plants; stems 1 cm. long or longer,

densely tomentose below. Leaves incurved and contorted when dry,

about 2 mm. long, linear-lanceolate, short acuminate; margins erect,

coarsely and irregularly serrate above; costa strong, subpercurrent;

upper cells 7-8 y., densely papillose, occasionally in 2 layers near

margins, basal cells oblong, pellucid. Sporophyte unknown. (Fig.

47, J-M.)

Dept. San Marcos: Steyermark 356 96a.

Distribution: Costa Rica.

On moist, shaded bluffs at high altitude. This species has some

anomalous characters but until the sporOphyte is known it may

be retained here.

3. LEPTODONTIUM ORCUTTI Bartr., Journ. Wash. Acad. Sci. 21:

289. 1931.

Zygodon gracilis var. americana Grout, Moss Fl. No. Amer. 2: 141. 1935.

Leptodontium flexifolium var. americanum Grout, Moss Fl. No. Amer. 1: 171.

1938.

Slender plants, yellowish green above, brown below. Stems to

4 cm. long but usually shorter. Leaves contorted and incurved

when dry, to 2 mm. long, oblong-lingulate, abruptly acute, keeled;

margins slightly recurved, irregularly serrate above; costa ending

below apex; upper cells rounded, 7-10 n, densely papillose, 4-6 rows

at margins often incrassate forming a pellucid border, basal cells

rectangular. Seta about 12 mm. long, pale yellow; capsule suberect,

cylindric, urn 2 mm. long; peristome teeth divided to base, forks

lightly granulose; lid conic-rostrate. (Fig. 48, A-D.)

Dept. Huehuetenango : Steyermark 50172. Dept. San Marcos: Steyermark

35523. Dept. Totonicapan: Standley 62667, 62685, 84512a, 84562. Dept. Que-

zaltenango: Standby 67687, 67715b, 67717, 67727a, 67731, 67739, 67759a, 86136,

86137; Steyermark 34164, 34165a. Dept. Solola: Steyermark 47496. Dept. Chimal-

tenango: Standley 58745, 60980, 61078.

Distribution: North Carolina, Mexico, Costa Rica.

On trees, logs, banks and rocks at high altitudes. Until this

genus is more critically studied I feel that it is more practical to

BARTRAM: MOSSES OF GUATEMALA

107

use the less cumbersome name for the North American plants which

are obviously closely allied to L. flexifolium (Sm.) Hampe. In our

plants the leaves are often conspicuously bordered, the spores average

a little larger and there are slight differences in the sporophyte.

4. LEPTODONTIUM GRACILE C. M., Bull. Torr. Bot. Club 23: 474.

1896.

•Stems 3-4 cm. long, yellowish green at tips, brown below. Leaves

crowded, strongly contorted when dry, about 4 mm. long, ovate,

short acuminate; margins recurved about half way up, plane and

irregularly serrate above; costa ending below apex; inner basal cells

rectangular, lax, thin walled, hyaline, shorter and strongly papillose

toward margins, changing abruptly above to the rounded upper cells

which are strongly papillose with multifid papillae. Sporophyte

unknown. (Fig. 48, E-H.)

Dept. Totonicapan: Standley 62686, 62687, 8US2, 84U1, WW, 8M6.

Distribution: Mexico, Costa Rica, Bolivia.

On damp banks at high altitudes. The large area of delicate,

hyaline basal cells changing quickly to the small, chlorophyllose

cells of the margins and to the similar lamina cells above is a note-

worthy feature of this species.

H

FIGURE 48

A-D, Leptodontium Orcutti: A, plant, Xl; B, leaf, X14; C, apex of leaf, X120;

D, upper leaf cells and margin, X270.

E-H, Leptodontium gracile: E, plant, Xl; F, leaf, X14; G, upper leaf cells and

margin, X270; H, inner basal leaf cells, X270.

108

FIELDIANA: BOTANY, VOLUME 25

H

FIGURE 49

A-D, Leptodontium subgracile: A, plant, XI; B, leaf, X14; C, apex of leaf,

X120; D, inner basal leaf cells, X270.

E-H, Leptodontium acutifolium: E, plant, XI; F, leaf, X14; G, upper leaf

cells and margin, X270; H, inner basal leaf cells, X270.

5. LEPTODONTIUM SUBGRACILE Ren. & Card., Bull. Soc. Bot. Belg.

31:153. 1892.

Leptodontium brachyphyllum Broth. & Ther., Bull. Acad. Int. de Geog. Bot.

40. 1906.

Stems slender, flexuous, 3-8 cm. long or longer, yellowish green

above. Leaves not crowded, appressed with contorted points when

dry, spreading when moist, 2.5-5 mm. long, ovate-lanceolate, short

acuminate; margins recurved more than half way up, irregularly

serrate above; costa ending just below apex; basal cells linear with

firm lateral walls, papillose nearly to insertion, upper cells densely

papillose with multifid papillae. Sporophyte unknown. (Fig. 49,

A-D.)

Dept. San Marcos: Steyermark 35789. Dept. Totonicapan: Standley 83135.

Dept. Quezaltenango : Steyermark 34163a. Dept. Jalapa: Steyermark 322M, 33119.

Distribution: Mexico, Costa Rica, Colombia, Peru, Bolivia.

On dry and moist banks and trees at medium to high altitudes.

The only noticeable difference between L. subgracile and L. brachy-

phyllum is in the length of the stems. Apparently the plants growing

in moist habitats have longer stems while those found on dry or

rocky banks have shorter stems. As the structural details are

identical, I feel that they can safely be combined in one species.

BARTRAM: MOSSES OF GUATEMALA 109

6. LEPTODONTIUM ACUTIFOLIUM Mitt., Journ. Linn. Soc. 12: 51.

1869.

Stems slender, to 10 cm. long, yellowish above, brown below.

Leaves appressed and contorted when dry, widely spreading with

decurved points when moist, 2.5-3 mm. long, lanceolate from an

erect, obovate, clasping base, acuminate; margins recurved below,

sharply serrate above; costa percurrent; basal cells linear with

sinuose, incrassate lateral walls, upper cells 6-9 n, papillose. Seta

1.25 cm. long; capsule ovoid-cylindric; lid short beaked. (Fig. 49,

E-H.)

Dept. San Marcos: Steyermark 35523a, 3552!+. Volcan del Fuego, Godman &

Salvin (type).

Distribution: Ecuador, Bolivia.

On dry ridges in pine woods at high altitudes. Clearly dis-

tinguished by the upwardly dilated leaf base, the narrow basal cells

and the small, dense, obscure lamina cells.

7. LEPTODONTIUM EXCELSUM (Sull.) E. G. Britt, Bryol. 11: 66.

1908.

Syrrhopodon excelsus Sull., Muse. Allegh. 170. 1848.

Holomitrium serratum (Schp.) C. M., Syn. 2: 587. 1851.

Leptodontium brevisetum Mitt., Journ. Linn. Soc. 12: 50. 1869.

Growing in intricate mats or masses, yellowish green at tips.

Stems elongate, flexuous, profusely branched. Leaves strongly

contorted when dry, widely spreading when moist, often radiculose

at tips, about 3 mm. long, ovate-lanceolate, acuminate, deeply

carinate, slightly decurrent; margins recurved below, erect and

coarsely spinose-serrate at least half way down; costa short-excurrent;

lamina cells small, dense, rounded-quadrate, slightly incrassate,

finely papillose, diam. 5-6 /x near mid-leaf, larger toward apex,

inner basal cells narrowly rectangular with straight lateral walls,

smooth, pellucid, smaller toward margins. (Pig. 46, E-G.)

Dept. Quezaltenango : Sharp 231^.

Distribution: Southern Alleghenies, Mexico.

On tree at rather high altitude. Leptodontium is so broadly

represented here that Guatemala may well be considered as the

center of distribution for the genus. This is a noteworthy collection

extending the range of the species well to the southward.

110

FIELDIANA: BOTANY, VOLUME 25

FIGURE 50

A-C, Leptodontium exasperatum: A, plant, Xl; B, leaf, X14; C, upper leaf

cells and margin, X270.

D-F, Leptodontium sulphureum: D, plant, Xl; E, leaf, X14; F, upper leaf

cells and margin, X270.

8. LEPTODONTIUM EXASPERATUM Card., Rev. Bryol. 36: 74. 1909.

Plants yellow above, brown below. Stems branched, to 6-8 cm.

Ipng. Leaves crowded, contorted when dry, squarrose-recurved

when moist, to 4 mm. long, ovate, acuminate; margins broadly re-

flexed below, serrate above; costa percurrent, papillose on back;

basal cells linear, sinuose, upper cells rounded, distinct, very in-

crassate, coarsely papillose with strong, simple or forked papillae

about 10-12 fj. high. Sporophyte unknown. (Fig. 50, A-C.)

Dept. Huehuetenango : Steyermark 49956. Dept. Totonicapan: Standley

6266 1+a (as L. Sulphureum').

Distribution: Mexico.

On trees at high altitudes. The more broadly reflexed leaf

margins and especially the high, spine-like papillae of the leaf cells

seem to be good diagnostic characters as compared with L. sul-

phureum.

9. LEPTODONTIUM SULPHUREUM (C. M.) Mitt., Journ. Linn. Soc.

12: 51. 1869.

Trichostomum sulphureum C. M., Syn. 2: 626. 1851.

Leptodontium helicoides Card., Rev. Bryol. 36: 75. 1909.

BARTRAM: MOSSES OF GUATEMALA 111

Plants pale yellow; stems to 10 cm. long, branched. Leaves

contorted when dry, squarrose-recurved when moist, 3-3.5 mm. long,

sometimes indistinctly 3 ranked, slightly undulate, ovate-lanceolate,

short acuminate; margins recurved below, serrate above; costa sub-

percurrent, minutely papillose on back; basal cells linear, incrassate,

with sinuose lateral walls, upper cells rounded, incrassate, densely

papillose with low, rounded papillae scarcely 3 M high. Perichaetial

leaves 7-8 mm. long, sheathing, laxly areolate, cells elongate, lateral

walls straight or only slightly sinuose; seta 7 mm. long, pale yellow;

capsule ovoid-cylindric, urn 3.5 mm. long; lid 1 mm. long; peristome

teeth about 0.1 long, pale, nearly smooth, indistinctly articulate;

spores 20-30 M- (Fig. 50, D-F.)

Dept. Quiche: Standley 62471, 62539. Dept. Huehuetenango : Standley 62629a,

82720, 83084a; Steyermark 50508. Dept. San Marcos: Standley 68589, 86314 (in

prime fruit). Dept. Totonicapan: Standley 84514- Dept. Quezaltenango : Steyer-

mark 33942; Standley 67840, 83678 (as L. ulocalyx), 85653. Dept. Chimaltenango :

Standley 57827 (as L. ulocalyx). Dept. Guatemala: Standley 80570 (as L. ulocalyx).

Dept. Jutiapa: Standley 75933 (as L. ulocalyx), 76423 (as L. ulocalyx). Dept.

Zacapa: Steyermark 4231 9a. Dept. Jalapa: Standley 75583, 76849, 77303.

Distribution: Mexico, Nicaragua, Costa Rica, Bolivia.

On damp, shaded banks and trees at medium to high altitudes.

The pale yellow or yellowish green color, the shorter, broader leaf

points and the leaves more decurved when moist will help to separate

this species from L. ulocalyx. Occasional collections with more

slenderly pointed leaves are puzzling but whether these are hybrids

or intermediate forms I do not know.

10. LEPTODONTIUM ULOCALYX (C. M.) Mitt, Journ. Linn. Soc. 12:

51. 1869.

Trichostomum ulocalyx C. M., Syn. 1: 578. 1849.

Trichostomum Sartorii C. M., Linnaea 38: 637. 1874.

Leptodontium perannulatum Williams, Bull. Torr. Bot. Club 34: 572. 1907.

Plants usually more slender than L. sulphureum and with a more

decided brownish cast. Leaves slenderly acuminate, less strongly

decurved when moist. Perichaetial leaves similar to L. sulphureum

but with firmer cells and more thickened, sinuose lateral walls;

capsules to 4 mm. long; annulus broad and persistent. (Fig. 51,

A-D.)

Dept. Alta Verapaz: Standley 92603. Dept. Huehuetenango: Standley 81117,

81695, 81698. Dept. San Marcos: Steyermark 36094, 36096. Dept. Totonicapan:

Standley 62729a, 84512, 84535a. Dept. Quezaltenango: Steyermark 34122, 34869b,

34851,34914a; Standley 67660a, 6766Sb, 67675, 67756, 83741. Dept. Sacatepequez :

112 FIELDIANA: BOTANY, VOLUME 25

Standley 65275. Dept. Chimaltenango : Standley 609J^9b, 618Mb, 61858, 61869,

61870. Dept. Guatemala: Standley 80608, 80732a. Dept. El Progresso: Steyer-

mark ^3114. Dept. Jalapa: Steyermark 321*61, 38051*.

Distribution: Mexico, Costa Rica, Venezuela.

On banks, trees and rocks at medium to high altitudes. This

species and L. sulphur eum are closely allied. The distinctions are

relative only and not sharp. I am tempted to keep them separate

but am doubtful if the distinctions will hold. An extreme variant

is represented by the following variety.

LEPTODONTIUM ULOCALYX var. CIRRIFOLIUM (Mitt.) Bartr., comb. nov.

Leptodontium cirrifoUum Mitt., Journ. Linn. Soc. 12: 52. 1869.

Stems longer, to 15-20 cm. long. Leaves with strongly contorted,

crispate points when dry, long and slenderly acuminate.

Dept. Totonicapan: Standley 62708, 62722a. Dept. Solola: Steyermark -47-455,

Distribution: Panama, Ecuador.

On banks at high altitudes. The longer stems and the relatively

longer more slender, crispate leaf points seem to be the only differ-

ences between this form and typical L. ulocalyx.

17. HYOPHILA Brid., Bryol. Univ. 1:760. 1826.

Dioicous; small, tufted, green or brownish plants. Stems

branched. Leaves crowded, when dry contorted with incurved

margins, Ungulate or spatulate, acute or obtuse, entire or weakly

toothed above; costa ending in or near apex; upper cells subquadrate,

basal cells rectangular. Seta terminal, erect; capsule exserted, erect;

peristome lacking; lid beaked.

Upper leaf cells distinct, slightly papillose, seta 5-10 mm. long ...... 1. H. tortula

Upper leaf cells obscure, densely papillose, seta 2-3 mm. long. . .2. H. microcarpa

1. HYOPHILA TORTULA (Schwaegr.) Hampe, Bot. Zeit. 1846: 267.

1846.

Gymnostomum tortula Schwaegr., Suppl. 22: 78. 1827.

Pottia riparia Aust., Muse. Appl. 112. 1870.

Pottia denticulata C. M., Bull. Herb. Boiss. 5: 190. 1897.

Pottia subcrenulata C. M., Bull. Herb. Boiss. 5: 190. 1897.

Pottia reflexifolia C. M., Bull. Herb. Boiss. 5: 190. 1897.

Stems 2 or 3 cm. high, frequently with clusters of stalked, multi-

cellular brood bodies in axils of comal leaves. Leaves oblong, lingu-

BARTRAM: MOSSES OF GUATEMALA

113

late or spatulate, short pointed, 2-3 mm. long; margins inflexed

below, plane above, often with several coarse, distant teeth near

apex; costa usually percurrent in a short, concolorous apiculus;

upper cells rounded, distinct, 8-12 ju, with firm walls, slightly papil-

lose, basal cells rectangular. Seta about 1 cm. long, reddish; capsule

cylindric, urn 1.5-2 mm. long; lid 0.5 mm. long; annulus wide.

(Fig. 51, E-G.)

Dept. Peten: Steyermark 45910, 1+5911; Lundell 2020. Dept. Izabal: Standley

73010; Steyermark 39922. Dept. Alta Verapaz: Steyermark 44762. Dept. Huehue-

tenango: Standley 82217, 82902. Dept. Quezaltenango : Steyermark 33929a. Dept.

Retalhuleu: Standley 88544. Dept. Sacatepequez: Standley 62234. Dept. Chimal-

tenango: Standley 64514. Dept. Chiquimula: Steyermark 31122. Dept. Jalapa:

Standley 76802, 76816, 77210. Dept. Santa Rosa: Standley 77966.

Distribution: Eastern United States, Arizona, Mexico, West

Indies, Central America, Brazil.

On damp rocks at low altitudes. A common and variable species

in the American tropics where it fruits freely. The synonymy is

quite extensive.

2. HYOPHILA MICROCARPA (Schimp.) Broth., E. & P. Pflanzenf. I3:

403. 1902.

Trichostomum microcarpum Schimp., Ann. Sci. Nat. Ser. 6, 3: 198. 1876.

FIGURE 51

A-D, Leptodontium ulocalyx: A, plant, Xl; B, leaf, X14; C, inner basal leaf

cells, X270; D, leaf of var. cirrifolium, X6.

E-G, Hyophila tortula: E, plant, XI; F, leaf, X12; G, apex of leaf, X120.

H-J, Hyophila microcarpa: H, plant, Xl; I, leaf, X14; J, capsule, X14.

114 FIELDIANA: BOTANY, VOLUME 25

Dioicous; stems less than 5 mm. high. Leaves crowded, incurved

and contorted when dry, to 2 mm. long, oblong-lanceolate, acute;

margins erect or slightly inflexed; costa percurrent; upper cells small,

dense, papillose, obscure and opaque, basal cells oblong, smooth,

pellucid. Seta 2-3 mm. long; capsules short ovoid to subglobose,

urn scarcely 1 mm. long; lid obliquely rostrate, 0.5 mm. long. (Fig.

51, H-J.)

Dept. Jutiapa: Standley 75228.

Distribution: Guadeloupe, Martinique.

On damp bank at moderate altitude. These plants are apparently

dioicous. No antheridia were found but the buds seen are all purely

archegonial. The leaves are rather more bluntly pointed than in

the Martinique plants but the sporophyte characters agree perfectly.

18. WEISIOPSIS Broth., Ofv. Finska Vet.-Soc. Forh. 62,

Avd. A, No. 9:7. 1920.

Autoicous; small terrestrial plants in extensive colonies. Leaves

contorted when dry, long Ungulate, broadly rounded; margins plane;

costa ending below apex; lamina cells small, papillose, basal cells

rectangular, pellucid, smooth. Seta erect, slender; capsule ovoid-

cylindric, erect; peristome teeth linear-subulate, erect, finely papil-

lose; lid conic-rostrate.

1. WEISIOPSIS OBLONGA TheY., Rev. Bryol. et Lich. 5: 95. 1932.

Stems to 3 mm. high, simple or branched, yellowish green above,

paler below, sparsely radiculose. Leaves with incurved, strongly

contorted points when dry, erect-spreading when moist, 1.5-1.8 mm.

long, oblong-lingulate, broadly rounded or truncate, carinate-con-

cave; margins erect, papillose-crenulate; costa pale, ending below

apex; upper cells rounded-hexagonal, diam. 10 M, scarcely incrassate,

highly convex, smooth, inner basal cells lax, rectangular, thin-walled,

pellucid, to 20 n wide, narrower toward margins. Seta very slender,

pale yellow, to 6 mm. long; capsule erect, urn 0.6-1 mm. long;

peristome teeth slender, red, well spaced, 0.2 mm. long, 15 M wide at

base, papillose; lid rostrate, oblique, 0.5 mm. long. (Fig. 52, A-C.)

Dept. Huehuetenango: Sharp 4981.

Distribution: Mexico.

On bank at moderately high altitude. The lingulate, plane-

margined, broadly rounded leaves in combination with the relatively

long, widely spaced peristome teeth should simplify the recognition

BARTRAM: MOSSES OF GUATEMALA

115

of this attractive and rare little moss previously known only from

the type locality in Mexico.

19. DIDYMODON Hedw., Sp. Muse. 104. 1801.

Small to moderately robust, tufted plants; stems branched,

radiculose below. Leaves crowded, erect-spreading, lanceolate;

margins recurved; costa strong, with dorsal and ventral stereid bands;

upper cells small, papillose, elongated and smooth below. Seta

elongate; capsules oblong to cylindrical; peristome teeth entire or

divided, not twisted; lid conic-rostrate.

1. Capsules curved 2. D. campylocarpus

Capsules erect 2

2. Leaves apiculate, usually toothed near apex 1. D. recurvirostris

Leaves not apiculate, entire 3

3. Leaf margins erect, basal cells hyaline, delicate 3. D. alticaulis

Leaf margins recurved, basal cells with firm walls 4

4. Leaves lanceolate or Ungulate, rounded-obtuse, costa ending below apex

6. D. tophaceus

Leaves ovate-lanceolate, costa percurrent or excurrent 5

5. Leaf apex blunt, costa percurrent 5. D. fusco-viridis

Leaf apex sharp, costa ending in subula or excurrent 4. D. Godmanianus

FIGURE 52

A-C, Weisiopsis oblonga: A, plant, XI; B, leaf, X20; C, part of peristome, X54.

D-F, Didymodon fusco-viridis: D, plant, XI; E, leaf, X22; F, apex of leaf, X 120.

G-I, Didymodon alticaulis: G, plant, Xl; H, leaf, X12; I, basal leaf cells, X400.

116 FIELDIANA: BOTANY, VOLUME 25

1. DIDYMODON RECURVIROSTRIS (Hedw.) Jennings, Man. Mosses

West. Pa. 97. 1913.

Weissia recurvirostra Hedw., Sp. Muse. 71. 1801.

Trichostomum aeneum C. M., Syn. 2: 628. 1851.

Trichostomum leucodon C. M., Bull. Herb. Boiss. 5: 192. 1897.

Synoicous; slender, tufted plants, yellowish green at tips, reddish

brown below. Stems to 4-5 cm. high. Leaves crisped when dry, to

3 mm. long, linear-lanceolate from an erect, pale, clasping base,

apiculate; margins revolute nearly to apex, strongly to obscurely

toothed near the point; costa ending near apex; basal cells narrowly

rectangular, hyaline, upper cells small, papillose, obscure. Seta

15-18 mm. long, slender, red; capsule suberect, cylindrical, urn to

3 mm. long; peristome teeth linear, reddish, rarely split; lid about

1 mm. long. (Fig. 53, A-C.)

Dept. Huehuetenango: Steyermark 50268a. Dept. San Marcos: Steyermark

35732, 36092. Dept. Totonicapan: Standley 81>029a. Dept. Quezaltenango :

Standley 67627a, 67665, 67705a, 84274, 84318, 84340, 86004, 86145; Steyermark

34055, 34092, 34093, 34147, 34249, 34598, 34611, 35151. Dept. Sacatepequez :

Standley 65244, 65246a. Volcan de Agua, Godman & Salvin (as D. aeneum).

Distribution: Greenland to Alaska south to New Jersey, Iowa,

New Mexico and Arizona, Mexico, wide in Europe, Asia, Africa,

New Guinea, Tasmania, New Zealand.

On soil, rocks and trees in limestone regions at medium to high

altitudes. This species may usually be recognized by the rusty-

red coloring, the sharply apiculate leaves and toothed apical margins.

D. aeneus (C. M.) Besch. has been credited to Guatemala but as

far as I can see it is only a robust form of D. recurvirostris. The

inflorescence of D. aeneus is monoicous but this alone is hardly

a specific character. From the description T. leucodon seems to

belong here also.

2. DIDYMODON CAMPYLOCARPUS (C. M.) Broth., E. & P. Pflanzenf.

I3: 405. 1902.

Trichostomum campylocarpum C. M., Syn. 2: 628. 1851.

?Barbula Jamesoni Tayl., Lond. Journ. Bot. 1846: 48. 1846.

?Syrrhopodon Jamesoni Tayl., Lond. Journ. Bot. 1847: 331. 1847.

Tortula arcuata Mitt., Journ. Linn. Soc. 12: 163. 1869.

Plants similar in appearance and coloring to the smaller forms

of D. recurvirostris. Stems about 1 cm. high. Leaves strongly con-

torted when dry, 1.5-2 mm. long, broadly linear from a slightly wider

base, acute, apiculate; costa ending below apex; margins plane above,

BARTRAM: MOSSES OF GUATEMALA 117

toothed near apex, recurved on one side below; inner basal cells

laxly rectangular, thin walled, hyaline or slightly colored, changing

abruptly to the chlorophyllose cells of the basal margins and the

small, subquadrate, papillose cells of the blade. Seta slender, red,

12-15 mm. long; capsules short, cylindrical, curved, wide-mouthed,

urn 1.5 mm. long; peristome teeth red, from a short basal membrane,

irregularly cleft. (Fig. 53, D-G.)

Dept. Quezaltenango : Standley 67765.

Distribution: Mexico, Costa Rica, Colombia, Ecuador.

On damp bank at rather high altitude. In addition to the

shorter, plainly arcuate capsules this species differs from D. recur-

virostris in the relatively broader leaf blade with the margins plane

above the calymperoid base. I have not seen the type of Barbula

Jamesoni Tayl. but if this and Syrrhopodon Jamesoni Tayl. prove

to be the same as D. campylocarpus, as I suspect, there is no apparent

reason why D. Jamesoni (Tayl.) should not be the acceptable name.

3. DIDYMODON ALTICAULIS Bartr., Bryol. 50: 204. 1947.

Relatively robust, bright green plants in dense tufts. Stems to

1.5 cm. high. Leaves crowded, strongly contorted with circinate

tips when dry, 3-4 mm. long, linear-lanceolate from a pale, clasping

base, blunt at apex; margins erect, entire; costa percurrent; basal

cells narrowly rectangular, delicate, hyaline, upper cells very obscure,

densely papillose, diameter 10 M, in one layer. Seta slender, flexuous,

reddish below, paler above; capsule oblong-cylindrical, urn 2 mm.

long; peristome teeth erect, to 0.5 mm. long, irregularly cleft; lid

conic-rostrate, 1.5 mm. long. (Fig. 52, G-I.)

Dept. El Quiche: Sharpe 5286.

Endemic.

Boulder in river bed at moderate altitude. Suggestive of Tri-

chostomopsis diaphanobasis (Card.) Grout in the delicate, hyaline

areolation of the leaf base but widely different in the costal structure

with thick stereid bands on both sides of the median guide row and

also in the unistratose lamina cells.

4. DIDYMODON GODMANIANUS (C. M.) Bartr., Bryol. 49:113. 1946.

Barbula Godmaniana C. M., Bull. Herb. Boiss. 5: 193. 1897.

Tortula campylocarpa Mitt, (nee Tayl.), Journ. Linn. Soc. 12: 159. 1869.

?Barbula strictidens C. M., Bull. Herb. Boiss. 5: 193. 1897.

118

FIELDIANA: BOTANY, VOLUME 25

K

FIGURE 53

A-C, Didymodon recur virostris: A, plant, Xl; B, leaf, X14; C, apex of leaf,

X120.

D-G, Didymodon campylocarpus: D, plant, Xl; E, leaf, X14; F, apex of leaf,

X120; G, capsule, X8.

H-K, Didymodon Godmanianus: H, plant, Xl; I, leaf, X14; J, apex of leaf,

X120; K, capsule, X8.

Small, pale, slender plants; stems 5-6 mm. high. Leaves crispate

when dry, about 2 mm. long, linear-lanceolate from a broader, ovate

base, acuminate; margins entire, recurved below; costa ending in

the fleshy, subulate point or excurrent; cells distinct, rounded, in-

crassate, papillose, 8-10 /z, more elongate near costa at base. Seta

6-8 mm. long, reddish; capsule erect, urn cylindric, narrowed at

mouth, 1-2 mm. long; peristome teeth erect, deeply divided, about

0.3 mm. long; lid 1 mm. long. (Fig. 53, H-K.)

Volcan de Fuego, Godman & Salvin. Dept. Sacatepequez: Standley 58531 in

part (as Trichostomopsis diaphanobasis) .

Endemic.

On old bricks at moderate altitude. Standley's 58531 differs in

no way from the type collection. The peristome teeth are erect

and the species should therefore be included in Didymodon, I have

not seen the specimens from Ecuador which Mitten cites under

T. campylocarpa Tayl. but have examined a part of the Godman

& Salvin collection upon which Miiller bases his species. It is

possible that B. strictidens belongs here but no material is available

for comparison.

BARTRAM: MOSSES OF GUATEMALA 119

5. DIDYMODON FUSCO-VIRIDIS Card., Rev. Bryol. 36: 83. 1909.

Small, densely tufted, dark olive green plants. Stems 5-8 mm.

high, densely foliate. Leaves erect and slightly contorted when

dry, widely spreading when moist, 1-1.5 mm. long, ovate-lanceolate,

blunt at apex, carinate-concave; margins entire, narrowly recurved

near mid-leaf; costa strong, percurrent; leaf cells chlorophyllose, the

upper small, rounded, faintly papillose, basal cells short-rectangular

with firm, pale walls. Seta red, 8 mm. long; capsule ovoid-cylindric,

dark brown; lid conic-rostrate, 0.6 mm. long; peristome teeth pale

brown, erect, 0.3 mm. long. (Fig. 52, D-F.)

Dept. Solola: Svihla 2888.

Distribution: Mexico.

On rock at moderate altitude. The shorter, bluntish leaf points

will hardly fail to separate this species from D. Godmanianus (C. M.).

Here as in the type collection from Mexico spherical, brown, axillary

propagula are often abundant.

6. DIDYMODON TOPHACEUS (Brid.) Jur., Laubm. 100. 1882.

Trichostomum tophaceum Brid., Method. Muse. 84. 1822.

Dioicous; plants in dense, dull, olive green tufts, brown below.

Stems to 3 cm. or more long. Leaves incurved when dry, ovate-

lanceolate or lingulate, apex obtuse or rounded, entire; margins

revolute below; costa strong, ending below apex; upper cells distinct,

rounded, incrassate, slightly papillose, rectangular below. Seta

8-12 mm. long, red; capsules cylindric, glossy; peristome teeth ir-

regularly divided, variable; lid conic-rostrate. (Fig. 54, A-D.)

Dept. Huehuetenango : Standley 656 40, 81586 (as Gyroweisia obtusifolia),

82407 (as Gyroweisia obtusifolia). Dept. Quezaltenango : Steyermark 8^985 (as

Gyroweisia obtusifolia). Dept. Sacatepequez: Standley 58979 in part (as Gyroweisia

obtusifolia).

Distribution: New York to British Columbia south to Tennessee

and Arizona, Mexico, Bolivia, Europe, Asia, Africa.

On damp banks in limestone regions at moderate altitudes. A

variable species but usually easily recognized by the rounded or

bluntly pointed leaves with the costa ending below the apex. Dr.

Andrews has a significant note on this species in The Bryologist,

44, p. 105. 1941.

20. BARBULA Hedw., Sp. Muse. 115. 1801.

Dioicous; small or medium sized, tufted plants partial to cal-

careous soil or rocks, yellowish or brownish green. Stems erect.

120 FIELDIANA: BOTANY, VOLUME 25

Leaves lanceolate, usually contorted when dry; margins entire,

mostly revolute; costa strong, percurrent or excurrent with dorsal

and ventral stereid bands; upper cells small, usually papillose and

obscure, basal cells more elongate and smooth. Seta elongate, erect;

capsules cylindric; peristome teeth divided to base into 32 filiform,

reddish forks, spirally twisted; lid long beaked; calyptra cucullate;

spores small.

1. Upper leaf cells subquadrate, smooth, pellucid 2

Upper leaf cells rounded or angular, mostly papillose 4

2. Leaves widest near middle, mucronate 13. B. agraria

Leaves widest at base, apex rounded or obtuse 3

3. Leaves narrowed from a wider base to a slender subulate point

1. B. subulifolia

Leaves gradually narrowed from a slightly wider base to a broadly ligulate

point 2. B. stillicidiorum

4. Leaves lanceolate, tapering to an acute apex (except B. brunneola) 5

Leaves oblong or lingulate, apex obtuse, usually mucronate 11

5. Costa long excurrent 6

Costa percurrent or nearly so 7

6. Leaves abruptly contracted to the subula, margins erect. .3. B. icmadophila

Leaves gradually tapering, margins revolute 4. B. Bescherellei

7. Leaves 4-6 mm. long, spirally contorted when dry, cells smooth

5. B. crassicostata

Leaves 2.5 mm. or less long, slightly curved when dry, cells papillose 8

8. Cells of ventral face of costa linear, distinct from lamina cells 9

Cells of ventral face of costa similar to lamina cells 10

9. Leaves squarrose when moist, acuminate, costa percurrent 8. B. reflexa

Leaves erect-spreading when moist, rounded, costa ending below apex

9. B. brunneola

10. Leaves erect-imbricated when dry, margins slightly recurved below

6. B. teretiuscula

Leaves curved with spreading points when dry, margins revolute to above

middle 7. B. vinealis

11. Perichaetial leaves convolute 12

Perichaetial leaves not convolute 13

12. Stems 1-3 cm. high, perichaetial leaves acute 10. B. Pringlei

Stems under 1 cm. high, perichaetial leaves blunt 11. B. hypselostegia

13. Leaves rounded, costa ending below apex 12. B. linguaefolia

Leaves mucronate, costa percurrent or excurrent 14

14. Leaf margin plane in upper half, recurved below 14. B. Cruegeri

Leaf margin strongly recurved from base nearly or quite to apex 15

15. Leaf margins spirally revolute to apex, basal cells smooth 15. B. spiralis

Leaf margins recurved to just below apex, cells papillose nearly to insertion

16. B. orizabensis

BARTRAM: MOSSES OF GUATEMALA

121

1. BARBULA SUBULIFOLIA Sull., Proc. Am. Acad. 1861: 277. 1861.

Plants densely tufted, pale or olive green above, brown below.

Stems red, to 2 cm. long, slender. Leaves contorted when dry,

1.5-2 mm. long, subulate-lanceolate from a broader base, bluntly

pointed; margins recurved below, denticulate at extreme apex; costa

ending in or just below apex; upper cells irregularly quadrate, 6-10 M,

pellucid, smooth, basal cells rectangular. Seta red, 12-18 mm. long;

capsule cylindric; peristome teeth red, spirally twisted in several

turns, about as long as urn; lid long beaked, about as long as urn.

(Fig. 54, E-H.)

Dept. Huehuetenango: Steyermark 50791. Dept. Chimaltenango: Standley

808W, 80842.

Distribution: Costa Rica, West Indies, Ecuador.

On wet banks and damp rocks at moderately high altitudes. A

variable species with respect to the shape of the leaves and the form

of the apex but readily distinguished by the smooth, quadrate, dis-

tinct upper leaf cells and the long, tightly twisted peristome.

2. BARBULA STILLICIDIORUM Card., Rev. Bryol. 37: 126. 1910.

Plants similar to B. subulifolia in habit, coloring and structural

details. Leaves broadly ligulate from a scarcely wider base, obtusely

rounded and crenulate at apex. (Fig. 54, I-K.)

B

FIGURE 54

A-D, Didymodon tophaceus: A, plant, Xl; B and C, leaves, X14; D, apex of

leaf, X120.

E-H, Barbula subulifolia: E, plant, Xl; F, leaf, X14; G, upper leaf cells and

margin, X270; H, capsule, X8.

I-K, Barbula stillicidiorum: I and J, leaves, X14; K, apex of leaf, X120.

122 FIELDIANA: BOTANY, VOLUME 25

Dept. Izabal: Steyermark 38628, 398^9. Dept. Alta Verapaz: Steyermark

^6317; Standley 92003. Dept. Retalhuleu: Steyermark 34569. Dept. Solola:

Standley 62769. Dept. Chimaltenango : Standley 64483, 64494, 80844, 8087 It,

80943. Dept. Chiquimula: Steyermark 30661. Dept. Jalapa: Steyermark 32235,

32276, 32915, 32917, 32920; Standley 76945.

Distribution: Mexico, Porto Rico.

On wet banks and damp rocks at low to medium altitudes. Al-

though distinct in the extreme this species is closely allied to B. subuli-

folia and may eventually have to be reduced to synonymy.

3. BARBULA ICMADOPHILA Schimp., Bry. Eur. fasc. 43. Suppl. II.

1850.

Slender, brownish plants in lax tufts; stems 1-1.5 cm. long.

Leaves appressed with spreading points when dry, not contorted,

1-1.5 mm. long, abruptly linear-subulate from an ovate base;

margins erect, entire; costa long excurrent; cells rounded, incrassate,

nearly smooth, slightly elongate near costa at base. Sporophyte

not seen. (Fig. 55, E-G.)

Dept. San Marcos: Steyermark 35816.

Distribution: Canada, Montana, Europe, Asia.

On tree trunks at high altitude. The habitat is a peculiar one

for this species and it is far out of its known range, but a careful

comparison. with authentic material leaves little doubt that it belongs

here.

4. BARBULA BESCHERELLEI Sauerb. in Jaeg., Adumb. 2: 673. 1878.

Plants green or often tinged with brown, laxly tufted. Stems

slender, 1-4 cm. long. Leaves slightly contorted when dry, ap-

pressed or flexuous-spreading, 1.5-2 mm. long, ovate-lanceolate,

subulate-acuminate; margins revolute; costa long excurrent; cells

rounded, incrassate, lightly papillose, 7-9 n, slightly larger and

oblong near costa at base. Seta red, 1-2 cm. long; capsule cylindric;

peristome teeth spirally twisted; lid long beaked. (Fig. 55, A-D.)

. Dept. Huehuetenango : Steyermark 50152a, 50456 (as B. vinealis), 82106.

Dept. Totonicapan: Standley 83127 (as B. teretiuscula). Dept. Quezaltenango :

Standley 67599, 83378, 83399, 85802, 85975. Dept. Sacatepequez: Standley

59017, 65211. Dept. Baja Verapaz: Standley 69735. Volcan de Fuego, Godman

& Salvin. Dept. Santa Rosa: Bernoulli 657.

Distribution: Arizona, New Mexico, Mexico, Costa Rica.

On shaded banks and bases of trees mostly at high altitudes.

Mitten's description of Barbula rectifolia Tayl. does not apply to

BARTRAM: MOSSES OF GUATEMALA

123

the collections of Godman & Salvin cited under this heading which

show the leaves with a long, excurrent costa and structurally different

in no important details that I can see from B. Bescherellei.

5. BARBULA CRASSICOSTATA Bartr., Bryol. 49: 114. 1946.

Robust plants in deep tufts, dull yellowish green above, brown

below. Stems 3-4 cm. high, sparingly radiculose below. Leaves

spreading, strongly contorted with circinate points when dry, widely

spreading when moist and somewhat falcate-secund at the tips, 5-6

mm. long, lamina fragile, gradually linear-lanceolate from a short,

triangular-ovate base; margins entire, recurved below, erect above;

costa stout, brown, 150 n wide below, percurrent or excurrent in a

short, fleshy point; leaf cells smooth and incrassate, short and sub-

quadrate below, irregularly rounded above, diameter about 10 M-

Fruit unknown. (Fig. 55, H-K.)

Dept. San Marcos: Along road between San Sebastian at km. 21 and km. 8,

8-18 miles northwest of San Marcos, alt. 2,700-3,800 m., Steyermark 35695 TYPE,

S5715.

Endemic.

Suggestive of Pseudosymblepharis circinata in general appearance

but widely different in the shape and structure of the leaves. The

FIGURE 55

A-D, Barbula Bescherellei: A, plant, Xl; B, leaf, X14; C, apex of leaf, X120;

D, capsule, X8.

E-G, Barbula icmadophila: E, plant, Xl; F, leaf, X14; G, apex of leaf, X120.

H-K, Barbula crassicostata: H, plant, Xl; I, leaf, X12; J, upper leaf cells and

margin, X270; K, basal leaf cells, X270.

124 FIELDIANA: BOTANY, VOLUME 25

short basal areolation and recurved margins ^re indicative of Barbula

but without fruit one can only guess^at its generic position.

6. BARBULA TERETIUSCULA Schimp., in C. M., Syn. 1: 614. 1849.

Plants slender, laxly tufted, brownish; stems branched, sub terete.

Leaves rigidly erect, appressed, not contorted, 1-1.5 mm. long, ovate-

lanceolate, acuminate; margins slightly recurved below; costa per-

current; upper cells rounded, 8-10 n, papillose, oblong and pellucid

near costa at base. Seta red, 6-8 mm. long; capsule ovoid-cylindric,

urn 1.5 mm. long; peristome teeth reddish, twisted; lid conic-rostrate, ,

0.5 mm. long. (Fig. 57, A-D.) \

Dept. Sacatepequez: Standley 65262a.

Distribution: Mexico.

At high altitude. The erect, closely imbricated leaves give the

stems a characteristic look as compared with the other local species

but apart from this the species has no particularly distinctive

features.

7. BARBULA VINEALIS Brid., Bryol. Univ. 1 : 830. 1826.

Plants tinged with reddish brown, tufted; stems 1-3 cm. long.

Leaves appressed and lightly twisted with spreading points when

dry, about 2 mm. long, narrowly lanceolate from an ovate base,

acuminate; costa stout, percurrent; margins recurved to above

middle; upper cells small, dense and incrassate, larger and short

rectangular below. Seta red, 10-12 mm. long; capsule cylindric,

urn 2 mm. long; peristome teeth laxly twisted; lid conic-rostrate,

to 0.7 mm. long. (Fig. 57, E-H.)

Dept. Huehuetenango : Standley 81892 (c. fr.). Dept. Quezaltenango : Standley

85251. Dept. Jalapa: Steyermark 32815?

Distribution: Alaska to Mexico, east to Idaho and Montana,

Europe, Asia, Africa.

On rocks at high altitudes. These are the first records of B.

vinealis in Central America. The species is notoriously variable but

the Guatemalan plants deviate in no essential way from the specific

concept.

8. BARBULA REFLEXA (Brid.) Brid., Method. Muse. 93. 1822.

Tortula reflexa Brid., Muse. Recent Suppl. 1 : 255. 1806.

Slender, reddish-brown plants in dense, depressed tufts. Stems

2.5-3 cm. long, branched, decumbent, julaceous when dry. Leaves

BARTRAM: MOSSES OF GUATEMALA

125

crowded, imbricated when dry, squarrose-spreading when moist,

1.5 mm. long, ovate-lanceolate, short acuminate, strongly keeled,

slightly decurrent; margins recurved to above mid-leaf; costa per-

current, showing linear cells on the ventral face in contrast to the

small, papillose lamina cells, basal cells short, oval, incrassate, with

pellucid walls, elongate only near insertion. Sporophyte rare, as

in B. fallax Hedw. (Fig. 56, A-B.)

Dept. Huehuetenango: Sharp £9 %0, 5020.

Distribution: Northern United States and Canada south to

Virginia, New Mexico and Colorado.

On calcareous bluffs and outcrops at high altitudes. These

collections seem to be thoroughly typical of the species as it occurs

in temperate regions and extend the known range appreciably to the

southward.

9. BARBULA BRUNNEOLA C. M., Bull. Herb. Boiss. 5: 196. 1897.

Very dark, rich brown plants, densely tufted but easily separated.

Stems erect, to 1.5 cm. high, brittle, usually branched. Leaves

appressed and imbricated when dry, spreading when moist, 1.5-2

mm. long, linear-lanceolate from an ovate base, bluntly rounded at

the apex; margins recurved to above mid-leaf; costa strong, dark-

brown, ending below apex; basal cells short rectangular with incras-

FIGURE 56

A-B, Barbula reflexa: A, plant, Xl; B, leaf, X32.

C-G, Barbula brunneola: C, plant, Xl; D, leaf, X28; E, apex of leaf, X68;

F, basal leaf cells, X400; G, capsule, X12.

126

FIELDIANA: BOTANY, VOLUME 25

D

H

FIGURE 57

A-D, Barbula teretiuscula: A, plant, Xl; B, leaf, X14; C, apex of leaf, X120;

D, capsule, X8.

E-H, Barbula vinealis: E, plant, Xl; F, leaf, X14; G, apex of leaf, X120;

H, capsule, X8.

I-L, Barbula agraria: I, plant, Xl; J, leaf, X14; K, apex of leaf, X120; L,

capsule, X8.

sate, brownish walls, smooth, upper cells small, rounded, papillose.

Perichaetial leaves erect, acuminate; seta dark brown, 6-7 mm.

long; capsule cylindrical, urn 2 mm. long, dark brown; peristome

0.45 mm. high, teeth pale brown, twisted in about one turn; lid

conic-rostrate, 1 mm. long. (Fig. 56, C-G.)

Dept. Huehuetenango: Sharp 4.966.

Endemic.

On limestone boulders at high altitudes. No authentic material

of this species is available for comparison, but as the above collection

agrees perfectly with the original description and is likewise sharply

distinct from any of the other local species, I feel reasonably confident

in referring it here. The costa shows linear cells on the ventral face

as in B. reflexa and 5. fallax so the species is evidently closely allied

to this group.

10. BARBULA PRINGLEI Card., Rev. Bryol. 36: 85. 1909.

Plants to 3 cm. high, in dense tufts, yellowish green above,

brown below. Stems branched, the sterile shoots often with axillary

bulbils. Leaves contorted with incurved points when dry, 1.5 mm.

long, 0.5 mm. wide, lingulate from an ovate base, obtuse, apiculate;

BARTRAM: MOSSES OF GUATEMALA 127

margins plane; costa very stout, brownish, excurrent in a minute

apiculus; basal cells short, subquadrate, with pellucid, incrassate

walls, slightly elongate near costa at extreme base, upper cells minute,

opaque, obscure, papillose. Perichaetial leaves erect, convolute,

acute; seta slender, 15 mm. long, reddish; peristome teeth 1 mm. long,

twisted in several turns. (Fig. 58, A-D.)

Dept. Huehuetenango: Sharp 4814, 4915.

Distribution: Mexico.

On limestone bluff and outcrops at moderately high altitudes.

11. BARBULA HYPSELOSTEGIA Card., Rev. Bryol. 36: 84. 1909.

Small, densely tufted plants similar to B. Pringlei but with shorter

stems, rarely over 5-6 mm. high. Perichaetial leaves bluntly pointed.

Dept. Huehuetenango: Sharp 4984.

Distribution: Mexico.

On calcareous soil at moderately high altitude. In structural

details these plants resemble B. Pringlei too closely for comfort.

The sterile stems show the same ovoid, axillary bulbils, the leaves

are relatively shorter and more broadly pointed, but the distinctions

are far from sharp and the degree to which the perichaetial leaves

are pointed varies considerably. On the whole I doubt if they can

be maintained as separate species.

i

12. BARBULA LINGUAEFOLIA Bartr., Bryol. 50: 204. 1947.

Laxly tufted, reddish brown plants. Stems erect, to 1 cm. high,

laxly foliate. Leaves spreading, not contorted when dry, 1.5-2 mm.

long, 0.5 mm. wide, lingulate, rounded-obtuse; margins recurved to

above mid-leaf, plane and papillose-crenulate above; costa ending

below apex; basal cells rectangular, thin- walled, smooth, upper cells

rounded-quadrate, not incrassate, papillose. Seta slender, reddish,

10-12 mm. high; capsules curved when dry, erect when moist,

narrowly cylindrical, urn 2 mm. long; lid 1 mm. long, subulate-

rostrate; peristome teeth reddish, twisted in several turns. (Fig. 58,

E-I.)

Dept. Suchiate: Strihla 2879a.

Endemic.

A unique and distinctive species comparable to no other North

American Barbula that I am familiar with. The perfectly lingulate

leaves with the costa ending below the broadly rounded apex are

128

FIELDIANA: BOTANY, VOLUME 25

D

FIGURE 58

A-D, Barbula Pringlei: A, plant, Xl; B, leaf, X14; C, apex of leaf, X68;

D, perichaetial leaf, X14.

E-I, Barbula linguaefolia: E, plant, Xl; F, leaf, X14; G, upper leaf cells and

margin, X400; H, basal leaf cells, X270; I, capsule, X14.

suggestive of Tortula but the costa in cross section shows both dorsal

and ventral stereid bands.

13. BARBULA AGRARIA Hedw., Sp. Muse. 116. 1801.

?Barbula subagraria C. M., Bull. Herb. Boiss. 5: 195. 1897.

Small, gregarious, almost stemless plants. Leaves crowded in a

terminal rosette, slightly contorted when dry, ovate-lanceolate,

widest near middle, acute, concave, about 2 mm. long; margins

erect; costa ending in the mucronate point; upper cells subquadrate,

mammillose on ventral face, basal cells oblong, pellucid. Seta reddish,

about 1 cm. long; capsule cylindric; peristome teeth long, twisted;

lid long rostrate. (Fig. 57, I-L.)

Mazatenango: Bernoulli 127 in part (as B. subagraria C. M.).

Distribution: Florida, Louisiana, Texas, Mexico, West Indies.

On calcareous soil and rocks. I have seen no material of this well-

known species from Guatemala but Muller's description of B. sub-

agraria leaves little doubt that it belongs here.

14. BARBULA CRUEGERI Sond. in C. M., Syn. 1 : 618. 1849.

Plants small, yellowish; stems red, less than 1 cm. long, rarely

longer, usually with obovate, stalked propagula in the leaf axils.

BARTRAM: MOSSES OF GUATEMALA

129

Leaves contorted when dry, 1.5-2 mm. long, oblong-lanceolate,

obtuse, mucronate; costa percurrent; margins narrowly recurved

below, plane above; upper cells small, opaque, densely papillose,

basal cells rectangular, incrassate, pellucid. Seta red; capsule

oblong-cylindric; lid long beaked; peristome teeth red, closely

twisted. (Fig. 59, A-D.)

Dept. Peten: Bartlett 125^8. Dept. Jalapa: Steyermark 32950.

Distribution: New Jersey to Texas, Mexico, Costa Rica, West

Indies, northern South America.

On damp banks at moderate altitudes. This species is apparently

less common through Mexico and Central America than in the West

Indies where it is widely distributed.

15. BARBULA SPIRALIS Schimp. in C. M., Syn. 1 : 622. 1849.

Barbula perlinealis C. M., Bull. Herb. Boiss. 5: 195. 1897.

Medium sized plants, laxly tufted, dull yellowish green; stems

to 2 or 3 cm. long, branched. Leaves spirally contorted when dry,

1.5-2.5 mm. long, oblong-lanceolate from a broader, pale base, obtuse,

mucronate; margins strongly recoiled from near base to apex;

costa very strong, broader above, excurrent in a short, pale mucro,

FIGURE 59

A-D, Barbula Crugeri: A, plant, Xl; B, leaf, X14; C, apex of leaf, X120;

D, propagula, X120.

E-G, Barbula spiralis: E, plant, Xl; F, leaf, X14; G, apex of leaf, X120.

H-K, Barbula orizabensis: H, plant, Xl; I, leaf, X14; J, apex of leaf, X120;

K, basal leaf cells, X270.

130 FIELDIANA: BOTANY, VOLUME 25

papillose on back; upper cells small, densely papillose, very obscure,

basal cells linear, smooth, pellucid toward costa, shorter and chloro-

phyllose toward margins. Seta reddish, 12-15 mm. long; capsule

narrowly ovoid-cylindric, urn about 3 mm. long; peristome teeth

tightly twisted; lid conic-rostrate, 1-1.5 mm. long. (Fig. 59, E-G.)

Dept. Totonicapan: Standky 83157. Dept. Quezaltenango : Standley 664.17,

66490, 66494, 66497a, 83888. Dept. Zacapa: Standley 74401. Dept. Chiquimula:

Steyermark 30662. Dept. Jalapa: Steyermark 32816a; Standley 76707, 76777.

Distribution: Arizona, New Mexico, west Texas, Mexico.

Dry, shaded banks and rocks at medium to high altitudes. The

broadly revolute leaf margins extending from the apex nearly to the

base and the large area of elongated, pellucid basal cells make this

species easy of recognition.

16. BARBULA ORIZABENSIS C. M., Linnaea 40: 638. 1876.

Plants similar in size and appearance to B. spiralis. Leaves

lanceolate from an ovate base, obtuse, mucronate; margins recurved

from just above base nearly to apex, plane for a short distance

below point; cbsta strong but not broader above, short excurrent;

upper cells dense and opaque, basal cells rectangular, papillose almost

to insertion. Small, globose, multicellular propagula are frequent

if not constant in the upper leaf axils. Sporophyte not seen. (Fig.

59, H-K.)

Dept. Huehuetenango: Standley 81224, 82428a, 82591, 82780, 82904, 83029a.

Dept. Guatemala: Standley 62865.

Distribution: Mexico.

On damp, shaded banks at moderately high altitudes. The

distinctions between this species and B. spiralis are narrow but

decisive. The papillose basal leaf cells and recurved but not revolute

margins becoming flat just below the apex are constant characters.

EXCLUDED SPECIES

The following species are not available for comparison and can-

not be accurately placed from the descriptions.

Barbula pellata Schimp., Bull. Herb. Boiss. 5: 192. 1897.

Barbula lagunicola C. M., Bull. Herb. Boiss. 5: 194. 1897.

Barbula suberythropoda C. M., Bull. Herb. Boiss. 5: 194. 1897.

Barbula lonchostega C. M., Bull. Herb. Boiss. 5: 195. 1897.

BARTRAM: MOSSES OF GUATEMALA 131

21. MORINIA Card., Rev. Bryol. 37: 124. 1910.

Rather robust, tufted plants, resembling Tortula. Leaves erect

and slightly contorted when dry, squarrose-recurved when moist,

lanceolate, acute, carinate; margins revolute, thickened above,

denticulate toward apex; costa strong, with stereid bands on both

sides of the median guide row; upper cells small, densely papillose,

basal cells rectangular, hyaline. Seta elongate; capsules cylindric;

peristome teeth from a low basal membrane, divided to base, forks

spirally twisted.

1. MORINIA EHRENBERGIANA (C. M.) The'r., Smiths. Misc. Coll.

854:22. 1931.

Barbula Ehrenbergiana C. M., Syn. 1 : 636. 1849.

Barbula trichostomoides Besch., Prodr. Bryol. Mex. 38. 1871.

Morinia trichostomoides (Besch.) Card., Rev. Bryol. 37: 124. 1910.

Dioicous; plants brown; stems to 3 or 4 cm. high. Leaves

crowded, about 3 mm. long, strongly keeled, with a thickened

border; margins revolute more than % up, irregularly denticulate

for some distance below apex; costa brown, percurrent or short

excurrent, smooth on back; upper cells rounded, about 8 n, very

obscure, 2-3 stratose in several rows at margins forming a distinct,

thickened border, basal cells narrowly rectangular, thin walled,

hyaline. Seta 8-10 mm. long, thick, reddish ; capsules erect or slightly

curved, urn 4 mm. long; lid conic-rostrate, 1.5 mm. long; peristome

teeth reddish, tightly spiraled. (Fig. 60, A-D.)

Dept. San Marcos: Steyermark 3556 J^b, 85900. Dept. Quezaltenango: Standley

6 7601 a.

Distribution: Mexico.

On trees at high altitudes. The costal structure and the leaves

with a thickened border, toothed toward apex, are distinctive

characters in comparison with Tortula. Previously known only

from Mexico.

22. STREPTOPOGON Wils. mss.; Mitt, in

Lond. Journ. Bot. 51. 1869.

Medium sized, corticolous plants tinged with brown, in lax tufts;

stems branched, laxly foliate. Leaves contorted when dry, often

bordered; margins recurved below, entire or toothed; costa with a

dorsal stereid band only, ending near apex or long excurrent; cells

132 FIELDIANA: BOTANY, VOLUME 25

lax, smooth and rectangular below, oval-hexagonal above. Seta

short; capsules oblong-cylindric, exserted; peristome teeth from a

low basal membrane, divided to base into 32 spirally twisted forks;

lid long conic; calyptra conical, barely covering lid, scabrous with

short, bristly hairs.

1. Leaves obtusely rounded, cucullate 3. S. cavifolius

Leaves acuminate 2

2. Leaves lanceolate, bordered 1. S. erythrodontus

Leaves oblong, unbordered 2. S. rigidus

1. STREPTOPOGON ERYTHRODONTUS (Tayl.) Wils., in Lond. Journ.

Bot. 51. 1851.

Barbula erythrodonta Tayl., Lond. Journ. Bot. 50. 1846.

Autoicous; stems to 3 cm. long. Leaves flexuous-spreading and

spirally contorted when dry, 5-6 mm. long, lanceolate from an oblong,

clasping base; margins recurved below, denticulate toward apex;

costa excurrent in a denticulate arista; upper cells oval-hexagonal,

to 50 M long, more elongate below, basal cells narrowly rectangular;

bordered all around with a narrow yellowish band of elongated cells

clearly differentiated from the cells within. Seta 2-3 mm. long;

capsule pale brown, urn 2-2.5 mm. long; lid rostrate, 1.5 mm. long;

calyptra mitriform, hispid; peristome teeth red, twisted in several

turns. (Fig. 60, E-G.)

Dept. Quezaltenango : Standley 66350a.

Distribution: Colombia, Ecuador, Peru, Bolivia, Hawaii, Mada-

gascar, Africa.

On tree at high altitude. This is the first North American

record for this interesting and attractive species.

2. STREPTOPOGON RIGIDUS Mitt., Bryol. 50: 205. 1947.

Dioicous; plants brownish green. Stems simple, about 15 mm.

high, densely foliate, slightly radiculose. Leaves slightly contorted

when dry, rigidly erect-spreading when moist, 3 mm. long, 0.8 mm.

wide, ovate-oblong, acuminate, concave, unbordered ; margins entire,

narrowly recurved almost to apex; costa stout, excurrent in a short,

clavate point, crowned with dense, sphaerical clusters of elliptical,

articulated propagula; upper cells hexagonal, smooth, thin- walled,

to 16 n wide, 32 // long, smaller at margins, interior basal cells laxly

rectangular, to 110 p. long, shorter toward margins. Fruit unknown.

(Fig. 61, A-B.)

BARTRAM: MOSSES OF GUATEMALA

133

Dept. El Quiche: Sharp 2425. Dept. Baja Verapaz: Sharp 2792, 2894, 2902.

Distribution: Costa Rica, Colombia, Ecuador, Brazil.

On bark of trees at moderate altitudes. The name is apparently

unpublished as no trace of it can be found in either Mitten's Musci

Austro-Americani or in the Paris Index. It has become familiar

through usage so it has seemed advisable to validate the combina-

tion. S. peruvianus Broth, may be the same thing but no authentic

material is available for comparison.

In habit and coloring the plants are suggestive of Tortula but

the sphaerical clusters of propagula at the tips of the probos-

coid leaf apices are unique. Sharp's collections are uniformly

sterile but abundant and in splendid condition. They will be a

welcome addition to American herbaria as the species is known in

North America only from a few meager collections from Costa Rica.

3. STREPTOPOGON CAVIFOLIUS Mitt., Journ. Linn. Soc. 12: 180.

1869.

Stems 1 cm. or more long. Leaves erect and slightly contorted

with incurved tips when dry, 2.5-3 mm. long, oblong, concave,

unbordered, obtuse and cucullate at apex, often bearing numerous

w

FIGURE 60

A-D, Morinia Ehrenbergiana: A, plant, Xl; B, leaf, X14; C, apex of leaf,

X120; D, upper leaf cells and margin, X270.

E-G, Streptopogon erythrodontus: E, plant, Xl; F, leaf, X8; G, upper leaf

cells and margin, X120.

H-J, Streptopogon cavifolius: H, plant, Xl; I, leaf, XlO; J, upper leaf cells and

margin, X120.

134 FIELDIANA: BOTANY, VOLUME 25

clavate or cylindrical, septate propagula on both faces near apex;

upper cells oblong-hexagonal, thin walled, to 50 n long and 20 /z

wide, smaller toward margins, basal cells more elongate, rectangular,

all with a persistent primordial utricle. Sporophyte unknown.

(Fig. 60, H-J.)

Dept. Alta Verapaz: Standley 69Wa, 69566c.

Distribution: Mexico, Colombia, Ecuador.

On trees in small tufts mixed with other mosses, at moderate

altitude. This curious and very individual species is evidently quite

rare. Mitten rather inaptly compares it to S. erythrodontus from

which it differs widely in appearance and detail.

23. DESMATODON Brid., Muse. Rec. Suppl. 4:86. 1819.

Plants small or medium sized, usually tufted, terrestrial. Leaves

incurved or slightly contorted when dry, oblong-lanceolate, broadly

pointed, subentire; costa strong, ending in or near apex, with a

stereid band on dorsal side only; upper cells small, papillose, basal

cells elongate, smooth. Seta elongate; capsules usually erect; lid

beaked; peristome teeth erect, divided nearly to base; calyptra

cucullate.

Small plants, stems under 5 mm. high, leaves oblong-lanceolate, 1-1.5 mm. long

1. D. Sprengelii

Robust plants, stems 2 cm. high, leaves Ungulate or spathulate, 2 mm. long

2. D. spathulifolius

1. DESMATODON SPRENGELII (Schwaegr.) Williams, Bull. Torr.

Bot. Club 46: 217. 1919.

Barbula Sprengelii Schwaegr., Suppl. 21: 64. 1824.

Desmatodon Garberi Lesq. & James, Man. 112. 1884.

Hyophila fragilis Card., Rev. Bryol. 36: 75. 1909.

Dioicous; small, densely tufted, green plants; stems 3-5 mm.

high, radiculose below. Upper leaves 1-1.5 mm. long, crowded, in-

curved when dry, broadly Ungulate, obtuse, often apiculate, entire

or toothed near apex; margins strongly inflexed; costa strong,

percurrent; upper cells small, dense, obscure, 5-8 n, mammillose on

upper face, basal cells oblong, smooth. Seta pale, to 5 mm. long;

capsules erect, ovoid-cylindric, urn 1.5 mm. long; lid conic-rostrate;

peristome teeth divided nearly to base, the forks strongly articulated,

erect. (Fig. 62, A-D.)

Dept. Peten: Bartlett 12541. Dept. Chiquimula: Steyermark 3171>3.

BARTRAM: MOSSES OF GUATEMALA

135

Distribution: Florida, Mexico, British Honduras, West Indies.

Moist cliff at moderate altitude. These collections represent the

wider leaved plants with entire apical margins previously referred

to D. Garberi. I have followed Grout in reducing this form to

D. Sprengelii but am not sure that they are conspecific.

2. DESMATODON (?) SPATHULIFOLIUS Bartr., Bryol. 50: 205. 1947.

Rather robust, densely tufted plants, green above, pale brown

below. Stems 2 cm. high, freely branched above from a stipe-like

base. Leaves crowded, strongly contorted when dry, widely spread-

ing when moist, to 2 mm. long, 0.6 mm. wide, lingulate from a nar-

rowed base, widest about middle, carinate-concave, broadly rounded,

mucronulate; margins entire or with a few blunt teeth near apex,

narrowly involute in upper half; costa strong, brownish, percurrent,

stereids lacking on ventral face; upper cells rounded, moderately

incrassate, papillose, interior basal cells short rectangular, becoming

quadrate toward margins. (Fig. 61, C-E.)

Dept. Huehuetenango : Sharp 52^5, 5247.

Endemic.

On moist calcareous rocks at moderate altitudes. Suggestive of

Hyophila in everything but the costal structure which shows a

H

A-B, Streptopogon rigidus: A, plant, Xl; B, leaf, X14.

C-E, Desmatodon spathulifolius: C, plant, Xl; D, leaf, X14; E, apex of leaf,

X54.

F-H, Tortula mniifolia: F, plant, Xl; G, leaf, X12; H, upper leaf cells and

margin, XllO.

136 FIELDIANA: BOTANY, VOLUME 25

stereid band on the dorsal side only. I have tentatively referred

the species to Desmatodon but its true affinity must wait upon the

discovery of fruit. Hyophila (?) lingulata Card, has narrower leaves

less broadly rounded at the apex and not mucronate.

24. ALOINELLA Card., Rev. Bryol. 36: 76. 1909.

Small, brownish, gregarious, gemmiform plants; stems very short.

Leaves fleshy, rigid, incurved when dry; margins broadly inflexed;

costa broad, densely clothed on the ventral face with septate,

chlorophyllose filaments; upper lamina cells incrassate, often broader

than long, basal cells rectangular, thin walled, hyaline. Seta erect,

elongate; capsules subcylindrical, peristome teeth divided to base

into 32 papillose, erect forks; lid beaked; calyptra cucullate.

1. ALOINELLA HAMULUS (C. M.) Bartr., Bryol. 49: 114. 1946.

Barbula hamulus C. M., Bull. Herb. Boiss. 5: 192. 1897.

Dioicous; stems 1-2 mm. high. Upper leaves crowded, brown,

1.5 mm. long, oblong-lingulate from a pale, laxly areolate base,

concave, strongly cucullate at apex; upper margins erose-denticulate,

broadly inflexed with the edges often overlapping; costa clothed on

the ventral face with filaments 2-3 cells high, excurrent in a short,

blunt mucro; upper lamina cells mostly transversely elongate, to

20 M long, 5-10 M wide, incrassate, basal cells rectangular, thin

walled, hyaline. Seta erect, reddish, 11 mm. long; capsule erect,

ovoid-cylindric, urn brown, 2 mm. long; peristome teeth rather short,

pale, from a low basal membrane, irregularly divided nearly to base,

the forks papillose, erect, about 0.25 mm. long. (Fig. 62, E-H.)

Dept. Quezaltenango: Bernoulli & Carlo 118.

Distribution: Mexico?

I feel reasonably sure that further collections from Guatemala

will prove that this species and A. catenula Card, of Mexico are

conspecific. One fruiting plant and some fragments of Barbula

hamulus have been seen through the courtesy of the New York

Botanical Garden. The seta is longer than mentioned by Cardot

in his description of A. catenula but numerous subsequent collections

of this species from Mexico show the setae varying from 6 to 14 mm.

long so this character evidently has little diagnostic value.

BARTRAM: MOSSES OF GUATEMALA 137

[ 25. TORTULA Hedw., Sp. Muse. 122. 1801. (in part).

Mostly medium sized to robust plants, often tinged with reddish

brown; stems, simple or branched. Leaves broad, ovate-lanceolate

or spatulate, erect and twisted when dry; margins entire, usually

revolute; costa strong, percurrent to long excurrent, with a thick

dorsal stereid band and a ventral layer of large cells; upper leaf

cells small, usually coarsely papillose, basal cells much larger,

hyaline or colored, usually sharply differentiated.

1. Leaves with a narrow, thickened border of elongated cells 3. T. mniifolia

Leaves unbordered 2

2. Costa long excurrent 4. T. guatemalensis

Costa percurrent or short excurrent 3

3. Leaves fragile, abruptly mucronate, without propagula 2. T. fragilis

Leaves not fragile, costa percurrent, with propagula 1. T. caroliniana

1. TORTULA CAROLINIANA Andrews, Bryol. 23: 72. 1920.

Plants green or reddish brown, in small tufts, often associated

with other mosses; stems to 1.5 cm. long. Leaves incurved when

dry, to 4 mm. long, oblong-lanceolate, carinate, short apiculate,

usually bearing numerous cylindrical, brownish septate propagula

on the ventral face of the upper lamina; margins recurved more than

half way up; costa brown, percurrent; upper cells rounded, 12-15 /x,

papillose, often smaller and more incrassate in several rows toward

margins, basal cells rectangular, smooth, thin walled.

Seta 7-8 mm. long, red; capsule cylindrical, to 3.5 mm. long,

exothecial cells short rectangular with thickened, brownish walls,

becoming smaller and rounded-hexagonal toward rim; annulus per-

sistent, 50 n wide, mostly of a single row of cells; peristome pale

red, 0.6-0.7 mm. high, teeth slightly twisted, from a pale basal

membrane projecting about 75 ^ above rim of capsule and slightly

higher than the annulus; lid conic-rostrate, 1 mm. long; spores pale

brown, minutely papillose, diameter 12-15 M- (Fig. 62, I-K.)

Dept. Alta Verapaz: Standley 69556b. Dept. Huehuetenango : Steyermark

50599b; Sharp 4809 in fruit. Dept. Totonicapan: Standley 84441a. Dept. Quezal-

tenango: Standley 886440, 841 93 in part. Dept. Sacatepequez: Standley 58825a,

63 71 Id.

Distribution: North Carolina, Tennessee, Mexico, Costa Rica.

On trees and banks at medium to high altitudes. The Guate-

malan plants are often more robust than those from the southern

Appalachians but otherwise are typical. The percurrent costa and

138

FIELDIANA: BOTANY, VOLUME 25

FIGURE 62

A-D, Desmatodon Sprengelii: A, plant, Xl; B, leaf, X14; C, apex of leaf,

X120; D, capsule, X8.

E-H, Aloinella hamulus: E, plant, Xl; F, leaf, X22; G, apex of leaf in profile,

X80; H, lamina cells, X270.

I-K, Tortula caroliniana: I, plant, Xl; J, leaf, X8; K, apex of leaf, X54.

the characteristic propagula on the upper leaf face are distinctive

features.

2. TORTULA FRAGILIS Tayl., Lond. Journ. Bot. 6: 333. 1847.

Tortula confusa Card., Rev. Bryol. 36: 87. 1909.

Tortula Pringlei Card., Rev. Bryol. 36: 87. 1909.

Moderately robust plants, usually reddish brown; stems from

3-4 mm. to 3-4 cm. high, matted with radicles below. Leaves

conduplicate and incurved when dry, very brittle, to 3 mm. long,

oblong-ovate, obtuse, mucronate; margins narrowly recurved below,

often lobed in the smaller, rounded, comal leaves; costa short

excurrent, brown; upper cells 10-15 n, densely papillose, inner basal

cells rectangular, thin walled, hyaline or colored, smaller and shorter

toward margins. Seta 8-14 mm. long, red; capsule cylindric;

peristome teeth from a distinct basal tube, spirally twisted. (Fig.

63, A-C.)

Dept. Quezaltenango: Standley 66^9^a, 67682. Dept. Sacatepequez: Standley

58825 (as T, parva), 588S2a (as T. parva). Dept. Jutiapa: Standley 75262.

Distribution: Virginia, West Virginia, southwestern United States,

Mexico, Colombia, Ecuador, Bolivia.

BARTRAM: MOSSES OF GUATEMALA 139

On trees and rocks at medium to high altitudes. The exceedingly

brittle lamina and the abruptly short mucronate apex distinguish

this species with little trouble. Some of the corticolous forms are

very reduced in stature.

3. TORTULA MNIIFOLIA (Sull.) Mitt., Journ. Linn. Soc. 12: 167.

1869.

Barbula mniifolia Sull., Proc. Am. Acad. 277. 1861.

Rather robust, brownish green, terrestrial plants in low, dense

tufts. Stems 6-7 mm. high, densely foliate, sparsely radiculose.

Leaves strongly curled and twisted when dry, spreading when

moist, to 5 mm. long, 1.7 mm. wide, oblong-lingulate from a

narrowed base, entire, obtuse, strongly bordered all around with a

narrow, thickened band of brownish, elongate cells; costa rather

slender, brown, merging with the border in the short, blunt point;

upper cells hexagonal, thin-walled, smooth, diameter 25-28 M,

gradually becoming more lax and rectangular below. Seta red,

10-12 mm. long; capsule oblong-cylindric, urn 2.25 mm. long;

peristome teeth from a short basal membrane. (Fig. 61, F-H.)

Dept. Baja Verapaz: Sharp 2900.

Distribution: Mexico, Costa Rica, West Indies, western South

America to Bolivia.

On moist bank at moderate altitude. The plants are suggestive

of Atrichum in a superficial way when dry and the brown bordered

leaves bear some resemblance to those of Mnium punctatum. It is

apparently uncommon and local in Central America.

4. TORTULA GUATEMALENSIS Bartr., Bryol. 49: 114. 1946.

Robust reddish brown plants with the habit and appearance of

T. norvegica (W. & M.). Stems to 5 cm. long, dichotomously

branched. Leaves squarrose-recurved when moist, ovate-lanceolate,

acute; margins plane throughout, often toothed at extreme apex;

costa long excurrent in a reddish, sparingly spinose awn; upper leaf

cells very opaque, densely papillose, 12 14 M, basal cells rectangular,

hyaline, shorter and narrower toward margins. Seta 8-10 mm. long;

capsule narrowly cylindrical, urn 4 mm. long; peristome 1 mm. long,

teeth slightly twisted from a short basal tube extending about 75 n

above the rim. (Fig. 63, D-G.)

Dept. San Marcos: Between San Sebastian and summit of Volcan Tajumulco,

alt. 3,800-4,600 m., Steyermark SS56Sa, S556b TYPE. Dept. Totonicapan: Standley

140

FIELDIANA: BOTANY, VOLUME 25

D

B

\S ~~^J 3

FIGURE 63

A-C, Tortula fragilis: A, plant, Xl; B, leaf, XlO; C, apex of leaf, X26.

D-G, Tortula guatemalensis: D, plant, Xl; E, leaf, X8; F, apex of leaf, X54;

G, capsule, X6.

Endemic.

Very similar to T. norvegica in the acutely pointed leaves and the

reddish awn but the plane margined leaves and short basal tube of

the peristome leave no doubt that it is a distinct species.

11. GRIMMIACEAE

Small to medium sized plants, mostly rupestrine, growing in

dense tufts or cushions. Leaves hygroscopic, often hyaline tipped;

upper cells small, usually opaque, often in 2 or 3 layers, basal cells

elongate, with straight or sinuous lateral walls; costa single, strong.

Seta terminal, usually elongate; capsules ovoid or cylindrical;

peristome single, teeth 16, entire or cleft above; calyptra mitriform

or cucullate.

1. Calyptra plicate, leaves muticous 3. Ptychomitrium

Calyptra not plicate, leaves hyaline tipped 2

2. Leaf cells sinuose, short above 1. Grimmia

Leaf cells nodulose, narrowly linear 2. Rhacomitrium

1. GRIMMIA Hedw., Sp. Muse. 73. 1801.

Densely tufted green plants; stems branched, radiculose below.

Leaves crowded, usually hyaline pointed; upper cells small, in several

layers especially toward margins, elongate and sinuose below. Seta

BARTRAM: MOSSES OF GUATEMALA

141

straight or curved; capsules ovoid, smooth or ribbed when dry;

peristome teeth 2-3 cleft above; lid short, conical; calyptra mitriform.

1. Capsules immersed 1. G. apocarpa

Capsules exserted 2

2. Seta erect, straight 2. G. ovalis

Seta arcuate or flexuous 3. G. trichophylla

1. GRIMMIA APOCARPA Hedw. var. GRACILIS (Schleich.) Web. &

Mohr., Bot. Taschb. 131. 1807.

Grimmia gracilis Schleich., Catal. Helv. (Ed. 2) 29. 1807.

Plants very dark brown or blackish, glossy, forming low, dense

mats. Stems to 2.5-3 cm. long, decumbent, densely foliate. Leaves

erect, curved or slightly secund when dry, 1-1.5 mm. long, ovate,

tipped with short, hyaline, denticulate hair-points; margins recurved,

often sinuate-dentate toward apex; costa percurrent, slightly toothed

on back near apex; basal cells short rectangular with firm, pale,

sinuous lateral walls, becoming shorter and rounded above. Peri-

chaetial leaves conspicuously larger, to 3 mm. long; capsule small,

oblong, immersed; lid rostrate from a conical base; peristome teeth

0.5 mm. long, entire, red, lanceolate, filiform pointed, nodulose

toward tips. (Fig. 64, A-C.)

Dept. Huehuetenango: Sharp 5000.

FIGURE 64

A-C, Grimmia apofarpa var. gracilis: A, plant, XI; B, stem leaf, X14; C,

sporophyte, X14.

D-G, Funaria obtusala: D, plant, Xl; E, leaf, X14; F, upper leaf cells and

margin, XllO; G, capsule, XlO.

142 FIELDIANA: BOTANY, VOLUME 25

Distribution: Northern United States and Canada south to

West Virginia, Tennessee, Arizona.

On limestone boulder at high altitude. This is an unusual record,

far to the south of the known range in North America. The plants

are in perfect fruit and are typical in all respects excepting the

peristome teeth which here instead of being cuneiform are narrowly

lanceolate with long, filiform, nodulose tips. Although this is an

anomalous feature I doubt if it has any real taxonomic value.

2. GRIMMIA OVALIS (Hedw.) Lindb., Acta Soc. Sci. Fenn. 10: 75.

1871.

Dicranum ovale Hedw., Sp. Muse. 140. 1801.

?Grimmia breri-exserta C. M., Bull. Herb. Boiss. 5: 200. 1897.

?Grimmia Bernoullii C. M., Bull. Herb. Boiss. 5: 200. 1897.

Grimmia praetermissa Card., Rev. Bryol. 36: 105. 1909.

Plants densely tufted, green or yellowish at tips, brown below;

stems branched, to 3 cm. high. Leaves 2.5-3 mm. long, imbricated

when dry, lanceolate, hyaline tipped; margin recurved on one side

below; costa ending below apex; upper cells rounded, sinuose, bistra- j

tose, elongated at base with straight or slightly sinuose lateral walls.

Seta erect, 2-3 mm. long; capsule erect, exserted, ovoid; lid conical;

peristome teeth 2-3 cleft to middle; calyptra mi triform. (Fig. 65,

A-C.)

Dept. San Marcos: Steyermark 3551+7a (as G. ovata), 3551+8 (as G. ovata),

3551+90, (as G. ovata). Dept. Quezaltenango : Standley 65526a (as G. praetermissa),

65530 (as G. praetermissa). Dept. Sacatepequez: Standley 65262 (as G. praeter-

missa).

Distribution: Greenland, British Columbia to California and

Arizona, Mexico, Europe, Asia.

On rocks and banks at high altitudes. The hyaline leaf tips vary

from very short or none to quite long but in a broad sense I can see

no advantage in separating these forms. The types of G. brevi-

exserta and G. Bernoullii have not been seen but the descriptions

strongly suggest that they belong here.

3. GRIMMIA TRICHOPHYLLA Grev., Fl. Edin. 235. 1824.

Yellowish green plants in lax tufts; stems to 3 cm. long. Leaves

erect and twisted when dry, 2-2.5 mm. long, linear-lanceolate from |

an ovate base, hyaline tip subentire; margins recurved below; costa

prominent at back; upper cells rounded, bistratose at margins, basal

BARTRAM: MOSSES OF GUATEMALA

143

cells linear, incrassate, slightly sinuose, shorter toward margins.

Seta 3-5 mm. long, strongly curved; capsule ovoid, yellowish, ribbed

with age; lid conic-rostrate; peristome teeth reddish, papillose, 2-3

cleft; calyptra mitriform. (Fig. 65, D-F.)

Dept. San Marcos: Steyermark S5683 in part, 36091, 36096a; Standley 851*10.

Dept. Totonicapan: Standley 8M51, 8^62, SWk-

Distribution: Western North America, Hawaii, Europe, Asia,

Africa, New Zealand.

On rocks at high altitudes. A widely distributed, variable species

but usually easily recognized by the leaves with a distinctly thickened

border and a long, nearly entire hyaline tip. As the plants fruit

freely the curved setae are noteworthy.

2. RHACOMITRIUM Brid., Mant. 78. 1819.

Robust rupestrine plants in loose, extensive mats; stems often

with numerous short lateral branchlets. Leaves lanceolate, usually

hyaline tipped; costa ending in or near apex; leaf cells elongate,

strongly sinuose or nodulose. Seta terminal, elongate; capsules

erect, ovoid-cylindric; lid long beaked; peristome teeth deeply 2-3

cleft into narrow forks; calyptra mitriform.

FIGURE 65

A-C, Grimmia ovalis: A, plant, XI; B, leaf, X22; C, capsule, X8.

D-F, Grimmia trichophylla: D, plant, Xl; E, leaf, X22; F, capsule, X8.

G-I, Rhacomitrium crispulum: G, part of plant, Xl; H, leaf, X14; I, upper

leaf cells and margin, X270.

144 FIELDIANA: BOTANY, VOLUME 25

1. RHACOMITRIUM CRISPULUM (H. f. & W.) H. f. & W., Fl. Tasm.

181. 1867.

Dryptodon crispulus H. f . & W., Fl. Ant. 1 : 57. 1843.

Trichostomum crispipilum Tayl., Lond. Journ. Bot. 5: 47. 1846.

Grimmia contermina C. M., Syn. 2: 655. 1851.

?Rhacomitrium fragile Ren. & Card., Rev. Bryol. 36: 106. 1909.

Plants hoary, yellowish green above, brown below; stems decum-

bent, to 8 cm. or more long, with numerous short lateral branchlets.

Leaves imbricated when dry, flexuous or secund, 3-3.5 mm. long,

ovate-lanceolate, acuminate, carinate, hyaline tip very variable,

from nearly obsolete to very long and strongly crisped; margin

recurved on one side; costa prominent at back; cells linear with

thickened, strongly nodulose lateral walls. Seta about 10 mm. long;

capsule cylindric, urn about 3 mm. long, erect or slightly curved;

lid subulate-rostrate. (Fig. 65, G-I.)

Dept. San Marcos: Steyermark 3551*0, 36101, 36102. Dept. Totonicapan:

Standley 81*l*35a, 81*1*1*5, 81*1*1*9, 81*1*61, 81*51*6. Dept. Quezaltenango : Standley

67709, 67710a, 67711, 6771k, 67715a, 67723; Steyermark 31*210, 31*211, 31+839.

Dept. Solola: Steyermark l*71*l*7a, 1*71*53, 1*71*57.

Distribution: Costa Rica to Fuegia, Africa, New Zealand, New

Guinea, Java, Sumatra, Borneo, Hawaii.

On rocks and rocky banks at high altitudes. Apart from the

hyaline tip, which is too variable to be used as a specific indicator,

the Cordilleran plants ranging north to Guatemala differ in no

essential way from those of other regions. It is evidently a plastic

and widely distributed species.

3. PTYCHOMITRIUM Furnr., Flora 2: 19. 1829.

Autoicous; medium sized tufted plants. Stems erect. Leaves

crisped when dry, lanceolate, entire or toothed above; cells smooth,

rounded above, narrower and elongate below. Setae erect, often

aggregated; capsules ovoid;* lid long beaked; peristome teeth 16,

divided nearly to base into narrow, papillose, erect forks; calyptra

mitriform, plicate, covering half the urn.

1. Leaves entire I. P. Leibergii

Leaves sharply serrate above 2

2. Capsule ovoid-cylindric 2. P. serratum

Capsule narrowly cylindrical 3. P. cylindrothecium

1. PTYCHOMITRIUM LEIBERGII Best, Bryol. 9: 80. 1906.

Plants 1-2 cm. high, brownish green, compactly tufted. Leaves

crowded, crispate with incurved points when dry, 3-5 mm. long,

BARTRAM: MOSSES OF GUATEMALA

145

linear-lanceolate from an ovate base, broadly acute; margins plane,

entire; costa strong, percurrent; upper cells rounded, 8-10 n, obscure,

often in 2 layers, basal cells oblong, hyaline. Seta 4-5 mm. long;

capsule ovoid, urn 1-1.4 mm. long; peristome teeth brown, densely

papillose, irregularly cleft; annulus broad; lid about 1 mm. long;

calyptra plicate, lobed at base; spores 15-25 M- (Fig. 66, A-D.)

Dept. Jalapa: Slandley 76792.

Distribution: Arizona.

On dry, shaded banks at moderate altitude. , These plants are

more robust and the spores average larger than in the Arizona col-

lections but these seem to be only trivial differences.

2. PTYCHOMITRIUM SERRATUM Bry. Eur. fasc. 2-3, Mon. 4. 1837

(name only).

Brachysteleum serratum C. M., Syn. 1 : 768. 1849.

Glyphomitrium serratum Mitt., Journ. Linn. Soc. 12: 106. 1869.

Robust, tufted plants, yellowish green above, brown below;

stems erect, 3-4 cm. high. Leaves crowded, strongly crisped when

dry, 5-6 mm. long, lanceolate, acuminate, apex acute, plicate near

FIGURE 66

A-D, Ptychomitrium Leibergii: A, plant, XI; B, leaf, X14; C, apex of leaf,

XI 20; D, calyptra, X8.

E-H, Ptychomitrium serratum: E, plant, Xl; F, leaf, X8; G, apex of leaf, X54;

H, capsule, X8.

I-K, Ptychomitrium cylindrothecium: I, leaf, X8; J, apex of leaf, X54; K,

capsule, X8.

146 FIELDIANA: BOTANY, VOLUME 25

base; margins narrowly recurved below, coarsely serrate toward

apex; costa percurrent; upper cells quadrate, incrassate, 8-10 n, often

in 2 layers near margins, basal cells linear, sinuose, hyaline. Setae

3-7 from one perichaetium, 3-5 mm. long; capsules ovoid-cylindric,

urn 2.5 mm. long; lid 1 mm. long; peristome teeth bifid to near base,

reddish; calyptra 2.5 mm. long, plicate, deeply lobed at base, serrate

on the plaits above. (Fig. 66, E-H.)

Dept. Huehuetenango : Standley 6591^3.

Distribution: Western Texas, Mexico.

On damp bank at high altitude. A handsome plant and one that

fruits freely throughout its range. The coarsely toothed leaves and

clustered, short setae are conspicuous features.

3. PTYCHOMITRIUM CYLINDROTHECIUM (C. M.) Par., Ind. Bryol.

1056. 1897.

Brachysteleum cylindrothecium C. M., Bull. Herb. Boiss. 5: 199. 1897.

Plants similar to P. serratum but leaves more slenderly acuminate,

usually strongly undulate in the upper half. Setae slender, clustered,

about 5 mm. long, yellowish; capsules narrowly cylindrical, 2-2.5 mm.

long; peristome teeth reddish, bifid, forks filiform, papillose; calyptra

as in P. serratum. (Fig. 66, I-K.)

Dept. San Marcos: Standley 85^15.

Endemic.

At high altitude. Although near P. serratum this species seems

to be distinct in the more slenderly pointed leaves and the narrowly

cylindrical capsules. Until a careful study of these and the closely

allied species from Mexico and Colombia is made the group cannot

be clearly resolved.

12. FUNARIACEAE

Small terrestrial plants with broad, soft, laxly areolate leaves

crowded in a comal tuft. Costa slender, usually ending below apex;

cells large, smooth, rhomboidal above, rectangular below. Seta

terminal, erect; capsules erect or curved, smooth or ribbed; peristome

single or double with segments opposite teeth or lacking; lid plano-

convex; calyptra smooth, long beaked.

1. Capsules strongly curved and asymmetrical 3. Funaria

Capsules erect and symmetrical 2

2. Lid conical, calyptra mi triform 1. Physcomitrium

Lid nearly flat, calyptra cucullate, inflated 2. Entosthodon

BARTRAM: MOSSES OF GUATEMALA 147

1. PHYSCOMITRIUM Brid., Bryol. Univ. 2:815. 1827.

Small, gregarious plants. Leaves contorted when dry, obovate

or spatulate; costa subpercurrent. Seta slender, elongate; capsules

erect, subglobose, wide mouthed, without peristome; lid plano-

convex, apiculate; calyptra mitriform, long beaked, lobed at base;

spores large.

1. PHYSCOMITRIUM OLLULA C. M., Bull. Herb. Boiss. 5: 174. 1897.

Small plants; stems simple. Leaves few, crispate, undulate-con-

cave when moist, rather broadly oblong from a long, very narrow

base, folded together, short acuminate, acumen distinctly serrulate,

recurved, narrowly bordered; costa reddish, slender, ending below

apex; cells lax, pellucid. Seta short, red, erect; capsule "tumescenti-

amphoroidea" ; lid minute, flat, apiculate.

Alta Verapaz: Pansamala, alt. 3,800 ft., H. v. Turckheim, Dec.

1887.

The above is a free transcription of the original description. None

of the original collection is available for comparison but I suspect

P. ollula will prove to be identical with P. subsphaericum Schimp.

of Mexico. The only noteworthy difference seems to be in the short-

ness of the setae which is a notoriously variable character in this

group.

2. ENTOSTHODON Schwaegr., Suppl. 2':44. 1823.

Small, autoicous plants with laxly areolate leaves. Seta slender,

elongate; capsules erect, symmetrical; peristome single, often rudi-

mentary, rarely lacking or double; lid convex, apiculate; calyptra

long beaked, inflated below. Distinguished from Funaria only by

the erect, symmetrical capsules.

1. Leaves obtuse, peristome well developed 3. E. longisetus

Leaves acuminate, peristome lacking or rudimentary 2

2. Leaves obovate, bordered, serrulate 1. E. Bonplandii

Leaves lanceolate, not bordered, entire 2. E. acidotus

1. ENTOSTHODON BONPLANDII (Brid.) Mitt., Journ. Linn. Soc. 12:

245. 1869.

Gymnostomum Bonplandii Brid., Bryol. Univ. 1: 101. 1826.

?Entosthodon microcarpus C. M., Bull. Herb. Boiss. 5: 174. 1897.

Small, green, gregarious plants; stems 2 mm. high. Upper leaves

few, crowded, 2 mm. long and a scant 1 mm. wide, obovate, concave,

148

FIELDIANA: BOTANY, VOLUME 25

H

FIGURE 67

A-C, Entosthodon Bonplandii: A, plant, XI; B, leaf, X18; C, upper leaf cells

and margin, X120.

D-F, Entosthodon acidotus: D, plant, Xl; E, leaf, X18; F, upper leaf cells

and margin, X120.

G-J, Entosthodon longisetus: G, plant, Xl; H, leaf, X18; I, upper leaf cells

and margin, X120; J, capsule, X14.

short acuminate, bluntly denticulate in upper half; costa ending

below apex; cells thin walled, oblong-hexagonal, about 25 p. x 65 /x,

narrower in several rows at margins. Seta 6-10 mm. long, reddish;

capsule erect, oblong-pyriform, urceolate and wide mouthed when

dry; peristome teeth very rudimentary, short, hyaline and truncate

or lacking. (Fig. 67, A-C.)

Dept. Alta Verapaz: Standley 9001 1*.

Distribution: Costa Rica, West Indies.

On damp bank at moderate altitude. No original material of

E. Bonplandii is available but the above collection corresponds in

every way with the description and memoranda of Mrs. Britton's

taken from the type. The description of E. microcarpus C. M.

suggests nothing very different and it is probably the same thing.

2. ENTOSTHODON ACIDOTUS (Tayl.) C. M., Syn. 2: 547. 1851.

Gymnostomum acidotum Tayl., Lond. Journ. Bot. 7: 279. 1848.

Stems slender, 2-4 mm. high. Leaves erect, appressed, 1-1.5 mm.

long, oblong-lanceolate, subulate-acuminate, entire; costa strong but

thin, ending below apex; cells oblong with firm, yellowish pellucid

BARTRAM: MOSSES OF GUATEMALA 149

walls, laxer below. Seta slender, red, 5-10 mm. long; capsules erect

or nodding, oblong-pyriform, tapering to a distinct neck, urn brown,

1-1.5 mm. long; peristome lacking. (Fig. 67, D-F.)

Dept. Chimaltenango: Standley 61015 in part (as Funaria microcarpa) .

Distribution: Ecuador, Bolivia.

On wet bank at high altitude. This is an interesting addition

to the North American moss flora. The erect, imbricated, entire,

unbordered leaves are sharply distinct from those of E. Bonplandii

and seem to be similar in all ways to Spruce's No. 444 from Mt.

Pichincha, Ecuador.

3. ENTOSTHODON LONGISETUS Schp., in Besch. Prodr. Bryol. Mex.

48. 1871.

Funaria epipedostegia Card., Rev. Bryol. 36: 109. 1909.

Small, yellowish green plants, densely gregarious. Stems 4-5 mm.

high. Upper leaves in a terminal tuft, decreasing in size below, to

2.8 mm. long, 1.2 mm. wide, spathulate-ovate from a narrow base,

widest about middle, obtuse, minutely apiculate, unbordered, entire;

costa ending below apex; upper cells irregularly hexagonal, thin-

walled, 25-30 ju wide, gradually more elongate and rectangular below.

Seta very variable in length, to 4 cm. long, slender, reddish; capsules

suberect, pyriform, narrowed to a distinct neck; peristome simple,

teeth red, well separated, linear-lanceolate. (Fig. 67, G-J.)

D.ept. Alta Verapaz: Sharp 29^1. Dept. El Quiche: Sharp 2356. Dept.

Totonicapan: Standley 8^021.

Distribution: Mexico.

On banks at moderately high altitudes. The broad, entire, un-

bordered leaves coupled with the peristomate capsules are good

diagnostic characters.

3. FUNARIA Hedw., Sp. Muse. 172. 1801.

Autoicous; plants gregarious; stems short. Lower leaves small,

the upper much larger and crowded in a terminal rosette, obovate,

acuminate; cells lax; costa strong, ending in or near apex. Seta

elongate; capsules pyriform with a long neck, usually unsymmetrical

and arcuate, sulcate, mouth oblique; annulus large; lid nearly flat;

peristome double, teeth 16, curved, segments opposite teeth.

Capsules smooth, annulus lacking 1. F. obtusata

Capsules sulcate, annulus present 2. F. hygrometrica

150

FIELDIANA: BOTANY, VOLUME 25

FIGURE 68

A-C, Funaria hygrometrica: A, plant, Xl; B, leaf, X14; C, capsule, X8.

D-F, Splachnobryum Bernoulli!: D, plant, Xl; E, leaf, X26; F, cells and

margin near leaf apex, X120.

G-I, Tayloria mexicana: G, plant, Xl; H, leaf, XlO; I, upper leaf cells and

margin, X54.

1. FUNARIA OBTUSATA Schimp., in C. M. Syn. 2: 540. 1851.

Small, densely tufted plants, dull yellowish green. Leaves

crowded in a terminal tuft, to 2.5 mm. long, 1.2 mm. wide, Ungulate,

obtuse, entire or sinuate toward apex by the projecting marginal

cells; costa slender, ending well below apex; upper cells irregularly

hexagonal, 25-30 n wide, elongate and rectangular toward base.

Seta yellow, 1 cm. long; capsule strongly arcuate and asymmetrical

when dry so that the mouth is vertical, faintly ribbed in lower half,

nodding, pyriform when moist; peristome double; annulus lacking.

(Fig. 64, D-G.)

Dept. Baja Verapaz: Sharp 279^.

Distribution : Mexico.

On soil at moderate altitude. Easily recognized by the short,

curved asymmetrical capsules with the mouth nearly vertical.

Previously known only from Mexico.

2. FUNARIA HYGROMETRICA Hedw., Sp. Muse. 172. 1801.

Funaria megapoda C. M., Bull. Herb. Boiss. 5: 175. 1897.

Plants in extensive colonies, pale green; stems to 1 cm. high.

Upper leaves contorted when dry, oblong-ovate, concave, short

BARTRAM: MOSSES OF GUATEMALA 151

acuminate, 2-4 mm. long, entire or weakly toothed; costa sub-

percurrent; upper cells hexagonal, elongated below. Seta 1-5 cm.

long, flexuous; capsule pyriform, unsymmetrical, sulcate; peristome

teeth spirally curved, dark red, united at tips, segments shorter

than teeth; calyptra long beaked, inflated below. (Fig. 68, A-C.)

Dept. Alta Verapaz: Standley 90790. Dept. Huehuetenango: Standley 65931,

82119, 81261, 82730. Dept. San Marcos: Standley 86265, 85SW, 85373, 86^36.

Dept. Totonicapan: Standley 83989, 8^1+75. Dept. Quezaltenango: Standley

66019, 66U8, SSltOS. Dept. Sacatepequez: Standley 58031, 586^2, 61175, 63052,

6S707. Dept. Chimaltenango: Standley 61112, 80316, 80325. Dept. Guatemala:

Standley 8050^. Dept. Zacapa: Steyermark ^2^60. Dept. Chiquimula: Steyer-

mark 31640. Dept. Jalapa: Steyermark 32680. Dept. Santa Rosa: Standley

77819.

Distribution: Cosmopolitan.

On bare soil, burned ground, banks etc. Many of the above

collections, especially from lower altitudes, represent the variety

calvescens (Schwaegr.) Bry. Eur. but at high altitudes the typical

form is not uncommon.

13. SPLACHNACEAE

Small to medium sized plants with erect stems and relatively

broad leaves, laxly areolate as in Funaria. Costa ending below apex

or excurrent. Seta elongate; capsules cylindrical, usually with a

distinct hypophysis; peristome single, teeth 16, often in 8 pairs.

Leaves entire, apex rounded, not bordered 1. Splachnobryum

Leaves serrate, apiculate, bordered 2. Tayloria

\

1. SPLACHNOBRYUM C. M., Verh. z. b. Ges. Wien 503. 1869.

Dioicous; small, slender, gregarious plants. Leaves not crowded,

Ungulate, obtuse, subentire; costa weak, ending below apex; cells

smooth and lax. Seta slender, elongate; capsule cylindric, erect;

peristome teeth papillose, well spaced, irregularly cleft; lid conical;

calyptra short, cucullate.

1. SPLACHNOBRYUM BERNOULLII C. M., Verh. z. b. Ges. Wien 505.

1869.

Splachnobryum crenulatulum Card., Rev. Bry. 36: 86. 1909.

Plants green; stems 5-8 mm. high. Leaves oblong-ovate, 1-1.5

mm. long, rounded and crenulate at apex; margins recurved toward

base; costa ending below apex; upper cells irregularly hexagonal, to

152 FIELDIANA: BOTANY, VOLUME 25

10-12 /i wide, smooth, more elongate below. Seta 4-5 mm. long,

slender; capsule cylindric, urn 1 mm. long, pale yellow, reddish at

mouth; peristome teeth 16, linear, coarsely papillose, deeply inserted;

lid conical, 0.25 mm. long; spores 13-17 n, smooth. (Fig. 68, D-F.)

Dept. Alta Verapaz: Standley 70961 (as S. obtusum C. M.?). Dept. Zacapa:

Standley 73880 (as S. obtusum?).

Distribution: Mexico, Honduras.

On wet rocks and banks at low altitudes. Until the tropical

American species of this genus are restudied it seems advisable to

refer the local collections here. The Arizona collections referred to

S. Bernoullii are apparently not the same thing and as far as I

know S. Bernoullii has not been found north of Mexico. The

sporophyte characters in the above description are from Standley's

No. 53516 from Honduras.

2. TAYLORIA Hook., Journ. Sci. and Arts 3: 144. 1816.

Medium sized plants; stems erect. Leaves not crowded, con-

torted when dry, lingulate or' spatulate, entire or serrate, often

bordered; cells lax; costa strong. Seta elongate; capsule erect with

a tapering neck; peristome teeth 16, single or paired; lid conical;

calyptra inflated below, smooth or pilose.

1. TAYLORIA MEXICANA (The>.) Bartr., Bryol. 49: 115. 1946.

Orthomnium mexicanum Ther., Rev. Bryol. et Lich. 5 : 103. 1932.

Moderately robust, pale green plants; stems about 1.5 cm. high,

densely reddish tomentose below, laxly foliate. Leaves strongly

contorted when dry, widely spreading when moist, 4 mm. long, 2 mm.

wide, broadly spatulate, obtuse, short apiculate, narrowly bordered;

margins recurved at extreme base, plane above, irregularly dentate

with short teeth; costa ending below apex; leaf cells lax, oblong-

hexagonal, gradually becoming rectangular below, 2-3 rows at

margins broadly rectangular, hyaline, forming an indistinct border

above middle. Seta short, smooth, 2-3 mm. long; capsule erect,

narrowly cylindrical, to 4.5 mm. long; peristome teeth evenly

spaced, brown, entire, to 300 M long, minutely vertically papillose-

striolate on the outer plates; spores brown, 15 M- (Fig. 68, G-I.)

Dept. Quezaltenango : Steyermark 34093a. Dept. San Marcos: Sharp 54.77.

Dept. Chimaltenango : Slopes of Volcan de Acatenango above Las Calderas, alt.

2,400-2,700 m., in dense, wet, Chiranthodendron forest, Standley 61932e.

Distribution: Mexico.

BARTRAM: MOSSES. OF GUATEMALA 153

This interesting and attractive species has some affinities with

both T. Jamesoni (Tayl.) and T. Moritziana C. M. From the

former it differs in the shorter setae, narrower capsule and narrower

leaf border and from T. Moritziana in the short seta, evenly spaced

peristome teeth and bordered leaves with short, blunt marginal

teeth. Unfortunately the calyptrae are not available.

14. BRYACEAE

Small to very large plants, usually tufted. Stems radiculose

below, simple or with subfloral innovations. Lower leaves small,

the upper larger, lanceolate to obovate; costa strong, usually per-

current or excurrent; cells smooth, prosenchymatous, linear to

rhomboidal, often narrower toward margins. Seta elongate; capsules

mostly inclined to pendulous, usually tapering to a distinct neck;

lid convex, apiculate; peristome normally double; calyptra cucullate;

spores small.

1. Peristome teeth lacking 1. Mielichhoferia

Peristome teeth present 2

2. Inner peristome without basal membrane 2. Orthodontium

Inner peristome with basal membrane 3

3. Capsule suberect, segments of inner peristome none or rudimentary

7. Brachymenium

Capsule inclined or pendulous, segments well developed 4

4. Upper leaf cells narrow, linear or narrowly rhomboidal 5

Upper leaf cells broad, rhomboidal 7

5. Stems julaceous, leaves closely imbricated 8. Anomobryum

Stems not julaceous, leaves spreading 6

6. Leaves narrow, costa broad 3. Leptobryum

Leaves broader, costa narrow 4. Pohlia

7. Upper leaf cells lax, wide, 15 n or more broad 8

Upper leaf cells firm, less than 15 /x wide 9

8. Leaves dimorphous, lateral rows larger than the dorsal rows . 6. Epipterygium

Leaves uniform 5. Mniobryum

9. Segments of inner peristome split into widely divergent forks

9. Acidodontium

Segments not forked 10

10. Seta usually solitary, stems not stoloniferous 10. Bryum

Setae aggregated, stems stoloniferous 11. Rhodobryum

1. MIELICHHOFERIA Hornsch., Bryol. Germ. 22: 179. 1831.

Slender, tufted plants. Leaves ovate-lanceolate, toothed above;

costa strong, ending near apex; cells linear-rhomboidal. Seta slender;

154

FIELDIANA: BOTANY, VOLUME 25

FIGURE 69

A-C, Mielichhoferia praticola: A, plant, XI; B, leaf, X26; C, upper leaf cells

and margin, X270.

D-F, Orthodontium pellucens: D, plant, Xl; E, leaf, X20; F, upper leaf cells

and margin, X270.

G-I, Leptobryum pyriforme: G, plant, Xl; H, leaf, X18; I, capsule, X8.

capsules pyriform, erect to horizontal; outer peristome lacking,

segments of endostome linear from a short basal membrane; lid

convex.

1. MIELICHHOFERIA PRATICOLA Card., Rev. Bryol. 38: 3. 1911.

Synoicous; plants in close, fragile tufts, yellowish green above,

brown below; stems 2-4 cm. high. Leaves suberect, 1.5 mm. long,

narrowly lanceolate, acuminate; margins plane, denticulate toward

apex; costa strong, ending below apex; cells linear-rhomboidal,

with firm walls, shorter below. Seta about 15 mm. long; capsule

cylindric, curved, erect or nodding, urn to 3 mm. long; segments of

endostome filiform, smooth. (Fig. 69, A-C.)

Dept. San Marcos: Steyermark 855 Wa, 355^6. Dept. Quezaltenango : Steyer-

mark

Distribution: Mexico, Costa Rica.

Rock crevices at high altitudes. These collections appear to be

identical with M . praticola but until a review of the tropical North

American species is made the specific distinctions must remain

uncertain.

BARTRAM: MOSSES OF GUATEMALA 155

2. ORTHODONTIUM Schwaegr., Suppl. 22: 123. 1827.

Small, delicate, yellowish green plants. Leaves numerous, long

and narrow, subentire; costa nearly percurrent; cells linear-rhom-

boidal. Seta elongate; capsule cylindric, sulcate when dry; peristome

double, teeth 16, slender, fragile, segments of endostome 16, not

united below, about as long as teeth.

1. ORTHODONTIUM PELLUCENS (Hook.) Bry. Eur. fasc. 23-24. 1844.

Bryum pellucens Hook., Ic. PI. 1: 34. 1837.

Autoicous; plants densely tufted; stems 3-8 mm. high. Leaves

flexuous when dry, linear-lanceolate, minutely denticulate near apex,

about 3 mm. long; costa slender, ending just below apex; cells linear-

rhomboidal, incrassate, shorter, laxer, thin walled and brownish at

base. Seta slender, to 1 cm. long; capsule nodding, pyriform with

a short neck, lightly sulcate when dry; peristome teeth and segments

about equal in length. (Fig. 69, D-F.)

Dept. Quezaltenango : Steyermark 31^6^2.

Distribution: California, Costa Rica, West Indies, South

America.

On rocks near sulfur terraces at high altitude. This collection

is sterile but is well within the known range of the species and

evidently belongs here.

3. LEPTOBRYUM (Bry. Eur.) Schimp., Coroll. 64. 1855.

Bryum subg. Leptobryum Bry. Eur. fasc. 46-47. 1851.

Slender, pale green, closely tufted plants. Leaves narrow, setace-

ous, costa broad ; cells linear above, shorter and broader below. Seta

elongate; capsule pyriform, glossy; peristome double, segments with

median slits, cilia appendiculate.

1. LEPTOBRYUM PYRIFORME (Hedw.) Schimp., Coroll. 64. 1855.

Webera pyriformis Hedw., Sp. Muse. 169. 1801.

Synoicous; stems about 1 cm. high, laxly foliate. Leaves flexuous

when dry, linear-setaceous, 2-3 mm. long; margins plane, denticulate

above; costa broad below, excurrent; cells linear. Seta terminal,

slender, to 3 cm. long; capsule pendulous, glossy, narrowed to a rather

long, wrinkled neck; lid hemispherical; peristome complete, teeth

156 FIELDIANA: BOTANY, VOLUME 25

yellowish, segments of endostome from a high basal membrane;

cilia 3, strongly appendiculate. (Fig. 69, G-I.)

Dept. San Marcos: Slandley 66232.

Distribution: Wide and nearly cosmopolitan.

On damp bank at high altitude. Infrequent throughout Mexico

and Central America and apparently confined to moderately high

altitudes.

4. POHLIA Hedw., Sp. Muse. 171. 1801.

Plants tufted or associated with other mosses. Stems erect,

simple or innovating. Leaves lanceolate, not distinctly bordered,

denticulate above; costa usually ending below apex; cells narrow,

usually linear, shorter and rectangular below. Seta elongate;

capsules clavate or pyriform, inclined or pendulous; peristome double,

segments of endostome with median slits; cilia nodose.

1. Capsule slender, with a long neck 1. P. spectabilis

Capsule clavate, with a short neck 2

2. Leaves pale green with metallic luster 2. P. cruda

Leaves yellowish green, without luster 3

3. Setae aggregated, short 3. P. polycarpa

Seta solitary, elongate 4

4. Capsules suberect, endostome rudimentary 6. P. tenuiseta

Capsules nodding, endostome well developed 5

5. Capsules ovoid, leaves short acuminate 4. P. papillosa

Capsules oblong-cylindric, leaves slenderly acuminate 5. P. peracuminata

1. POHLIA SPECTABILIS (C. M.) Broth., E. & P. Pflanzenfam. I3:

547. 1903.

Bryum spedabile C. M., Syn. 2: 583. 1851.

Webera cylindrica (Mont.) Schimp., in Besch., Prodr. Bryol. Mex. 52. 1871.

Paroicous; antheridia in axils of comal leaves. Plants slender,

yellowish green; stems about 1 cm. high, nearly bare below. Comal

leaves crowded, erect, 3-3.5 mm. long, lanceolate, acuminate;

margins recurved below, denticulate toward apex; costa strong,

ending just below apex or percurrent; cells linear, incrassate, shorter

and rectangular at base. Seta slender, 2.5-3 cm. long; capsules

nodding or horizontal, to 6-7 mm. long, cylindric, often curved,

neck not much narrowed, shorter than the rest of the capsule; lid

conical, apiculate; peristome teeth yellow, minutely papillose, seg-

ments of endostome narrow, nearly as long as teeth, scarcely slit, cilia

2-3, short and rudimentary, nodose; spores 12-15 M- (Fig. 70, A-C.)

BARTRAM: MOSSES OF GUATEMALA 157

Dept. San Marcos: Steyermark S5900a. Dept. Totonicapan: Standley 62712,

62697, 62699, 65909. Dept. Quezaltenango : Standley 676 14a, 67680, 67686,

67690, 67694, 67695, 67716, 67755, 67759, 83404, 83407, 86115, 86121a; Steyer-

mark 34186, 34165.

Distribution: Mexico, Costa Rica, Colombia.

On damp banks, logs and trees at high altitudes. This frequent

species fruits freely and will command attention at once by the rela-

tively large and conspicuous, elongated capsules. When well

developed the fruit is longer than in any of the nearly related species

north of the Mexican border but the structural features are very

similar.

2. POHLIA CRUDA (Hedw.) Lindb., Muse. Scand. 18. 1879.

Mnium crudum Hedw., Sp. Muse. 189. 1801.

Plants slender, loosely tufted, pale green with a pronounced

metallic luster; stems to 4 cm. high, radiculose below. Lower

leaves small and distant, above about 3 mm. long, ovate-lanceolate,

short acuminate, denticulate toward apex; costa reddish below, end-

ing below apex; upper cells linear, to 85 n long, comal leaves often

longer, narrower and slenderly acuminate. Seta 1.5-2 cm. long,

reddish; capsule nodding or horizontal, oblong-cylindric, neck short;

lid short, conical; peristome complete, teeth yellow, segments of

endostome widely split, cilia 2-3, nodose. (Fig. 70, D-F.)

Dept. San Marcos: Steyermark 35482, 36085. Dept. Totonicapan: Standley

62693, 62724, 84516a. Dept. Quezaltenango: Steyermark 34193a, 34251, 34254.

Distribution: Wide in North America, south along Cordillera to

Antarctica, Europe, Asia, Australia, New Zealand.

On rocks and trees at high altitudes. Several of these collections

are more robust than the average but otherwise typical.

3. POHLIA POLYCARPA (Mitt.) Broth., E. & P. Pflanzenfam. I3: 548.

1903.

Bryum polycarpum Mitt., Journ. Linn. Soc. 12: 293. 1869.

Webera Costesii Card. & The>., Rev. Chil. Hist. Nat. 1917: 13. 1917.

?Bryum aggregatum Hampe, Bull. Herb. Boiss. 5: 181. 1897.

Dioicous? Small, dull green plants; stems slender, about 1 cm.

high, nearly naked below. Upper leaves crowded in a comal tuft,

1.5-2 mm. long, ovate-lanceolate, acuminate, denticulate toward

apex; costa ending below apex; margins plane or narrowly recurved

below; cells linear, shorter and broader at base. Setae aggregated,

158

FIELDIANA: BOTANY, VOLUME 25

FIGURE 70

A-C, Pohlia spectabilis: A, plant, Xl; B, leaf, X12; C, capsule, X4.

D-F, Pohlia cruda: D, plant, Xl; E, leaf, X14; F, upper leaf cells and margin,

X270.

G-I, Pohlia polycarpa: G, plant, Xl; H, leaf, X14; I, apex of leaf, X54.

2-4 from one perichaetium, 6-8 mm. long; capsules brown, oblong-

cylindric, 2.5-3 mm. long, neck about equaling the rest of the

capsule; peristome short, segments of endostome narrow from a short

basal membrane, cilia none or very rudimentary; lid conical, apicu-

late. (Fig. 70, G-I.)

Volcan de Fuego, Godman & Salvin.

Distribution: Chile.

The clustered, short setae should readily identify this species

which is known locally only from the type locality.

4. POHLIA PAPILLOSA (C. M.) Broth., E. & P. Pflanzenfam. I3: 552.

1903.

Bryum papillosum C. M., Syn. 1 : 326. 1849.

?Bryum didymodontium Mitt., Journ. Linn. Soc. 12: 289. 1869.

Dioicous; plants slender, dull green; stems to 1.5 cm. high.

Lower leaves minute and distant, the upper 1.5-2 mm. long, narrowly

lanceolate, decurrent, acuminate, denticulate toward apex; margins

plane or narrowly recurved below; costa strong, ending just below

apex; cells linear. Seta slender, variable in length, from 3-8 cm.

long; capsules nodding, ovoid, small mouthed, short necked, 2-3

mm. long, appearing papillose by the strongly convex exothecial

BARTRAM: MOSSES OF GUATEMALA

159

cells; lid convex, apiculate; peristome short, teeth densely but

minutely papillose, segments of endostome narrow from a high basal

membrane, about as long as teeth, cilia none or rudimentary.

Sterile stems often with linear, vermicular propagula in the leaf

axils. (Fig. 71, A-C.)

Dept. San Marcos: Standley 66268, 66269, 86511a, 86515b; Steyermark 35631.

Dept. Quezaltenango: Standley 66375, 8SS08, 836^5, 85986, 86025. Dept.

Suchitepequez: Steyermark S5S2\, S5S62.

Distribution: Mexico, Costa Rica, West Indies, northern South

America.

On damp banks and trees at rather high altitudes. The long

setae and short, small, wide mouthed capsules are very distinctive.

Mitten's description of Bryum didymodontium is not very convincing

as compared with Muller's species, and I strongly suspect they are

conspecific.

Sterile plants may be distinguished from Mniobryum Wahlen-

bergii by the presence of gemmae in the upper leaf axils.

5. POHLIA PERACUMINATA Bartr., Bryol. 50: 206. 1947.

Dioicous; tufts yellowish green and slightly glossy above, reddish

tomentose below. Stems slender, laxly foliate, 1-1.5 cm. high.

FIGURE 71

A-C, Pohlia papillosa: A, plant, XI; B, leaf, X14; C, capsule, X8.

D-F, Mniobryum Wahlenbergti: D, plant, Xl; E, leaf, X14; F, upper leaf

cells and margin, X270.

G-I, Epipterygium immarginatum: G, plant, Xl; H, lateral leaf, X14; I,

dorsal leaf, X14.

160

FIELDIANA: BOTANY, VOLUME 25

H

D

FIGURE 72

A-C, Pohlia peracuminata: A, plant, Xl; B, leaf, X26; C, capsule, XlO.

D-G, Pohlia tenuiseta: D, plant, Xl; E, leaf, X14; F, capsule, XlO; G, part

of peristome, XllO.

H-I, Anomobryum semiovatum: H, leaf, X14; I, apex of leaf, XllO.

Leaves erect-spreading, 1.5 mm. long, ovate-lanceolate, long and

slenderly acuminate; margins recurved, minutely denticulate toward

apex; costa ending in acumen; upper cells linear-rhomboidal, laxer

and short rectangular toward base, not incrassate. Seta red, flexu-

ous, to 2 cm. long; capsule pendulous, urn 3 mm. long with neck,

oblong-cylindrical, neck about half the length of urn; lid conical;

peristome teeth yellow, endostome from a high basal membrane,

cilia 2, short, nodose. (Fig. 72, A-C.)

Dept. Huehuetenango : Sharp ^780a. Dept. San Marcos: Sharp 5451a.

Endemic.

On bank and tree trunk at high altitudes. Bryum Seleri C. M.

is described as having the leaves "breviter acuminata" which should

preclude any confusion with the new species. The dioicous inflores-

cence and sporophyte characters suggest faintly some affinity with

P. Drummondii (C. M.) Andrews but the leaves of the Guatemalan

plants are much more slenderly acuminate and the basal leaf cells

lax and delicate.

6. POHLIA TENUISETA Bartr., Bryol. 50: 206. 1947.

Dioicous; slender plants in dense, pale green tufts. Stems erect,

5-10 mm. high, radiculose at base. Upper leaves erect-spreading,

BARTRAM: MOSSES OF GUATEMALA 161

2-2.3 mm. long, narrowly ovate-lanceolate, acuminate; margins

recurved below, minutely denticulate above; costa ending below

apex; upper cells linear becoming short rectangular at base. Seta

very slender, to 5 cm. long, flexuous, red; capsules suberect, oblong-

cylindric from a short neck, small-mouthed, to 3.5 mm. long; lid

convex, apiculate; peristome teeth yellow, about 0.2 mm. high,

minutely papillose, endostome imperfect, consisting of a low, hyaline

basal membrane with irregularly cleft segments about as long as

teeth, cilia none or rudimentary; spores minutely papillose, diameter

20-25 M. (Fig. 72, D-G.)

Dept. El Quiche: Sharp 5336. Dept. Huehuetenango : Sharp 4909.

Endemic.

On banks at moderate altitudes. This is an unusual species in

several respects. The long, slender, bright red setae, suberect

capsules and imperfect inner peristome are characters in the aggre-

gate widely different from those attributed to any other member

of the genus.

EXCLUDED SPECIES

BRYUM SELERI C. M., Bull. Herb. Boiss. 5: 181. 1897.

This species is evidently referable to Pohlia but it cannot be

accurately determined from the description.

5. MNIOBRYUM (Schimp. ex p.) Limpr., Laubm. 2: 272. 1892.

Mniobryum Schimp. in Bry. Eur. fasc. 46-47 et Consp. ad Vol. 4. 1851.

Plants in lax pale green tufts; stems elongate, radiculose below,

laxly foliate. Leaves slightly contorted when dry, ovate-lanceolate;

costa ending below apex; cells lax, rhomboidal. Seta elongate;

capsule ovoid, nodding or pendulous; peristome complete, cilia

nodose; lid convex, apiculate.

1. MNIOBRYUM WAHLENBERGII (Web. & Mohr.) Bartr., comb. nov.

Hypnum Wahlenbergii Web. & Mohr., Bot. Taschenbuch 280. 1807.

Mnium albicans Wahlenb., Fl. Lapp. 353. 1812.

Dioicous; stems to 5 cm. long, often branched, slender. Leaves

1.5-2 mm. long, slightly decurrent, short acuminate; margins slightly

recurved below; plane and denticulate above; costa reddish toward

base, ending below apex; cells thin walled, to 15 or 20 n wide and

162 FIELDIANA: BOTANY, VOLUME 25

100 n long, narrower toward margins. Seta to 2 cm. or more long;

capsule ovoid, wide mouthed, neck short; peristome teeth slender,

yellowish, segments of endostome split, cilia 2 or 3, nodose. (Fig. 71,

D-F.)

Dept. San Marcos: Steyermark 36523.

Distribution: Nearly cosmopolitan.

Wet banks and rocks at medium to high altitudes. No fertile

plants have been seen from the local area. Sterile stems are fre-

quently quite slender with reduced leaves.

6. EPIPTERYGIUM Lindb., Oefv. K. Vet.-Akad.

Forh. 1863:599. 1863.

Dioicous; plants rather small, pale and dull green tinged with

red ; stems simple, radiculose below. Lower leaves small and distant,

the upper not crowded, complanate, dimorphous, the lateral rows

ovate, larger and broader than the dorsal rows; costa ending above

mid-leaf; cells very lax and thin walled, narrower toward margins

often forming a distinct, colored border. Seta elongate; capsule

pendulous, small, ovoid; peristome complete, segments of endostome

from a high basal membrane, cilia well developed, nodose.

Leaves with a reddish border 2. E. lepidopiloides

Leaves not bordered 1. E. immarginatum

1. EPIPTERYGIUM IMMARGINATUM Mitt., Journ. Linn. Soc. 12: 319.

1869.

Densely gregarious, medium sized plants; stems about 2 cm. high.

Lateral leaves broadly ovate, short apiculate, 2.5 mm. long; margins

plane, minutely denticulate above; costa slender, ending well above

mid-leaf; cells rhomboidal-hexagonal, to 20 M x 120 n, gradually nar-

rower and longer toward margins but not forming a distinct border;

dorsal leaves about 1 mm. long, lanceolate, acuminate. Seta 12-15

mm. long; capsule small, pendulous. (Fig. 71, G-I.)

Dept. Quezaltenango : Steyermark 3^736; Standley 65321 . Dept. Sacatepequez:

Standley 59^6 in part.

Distribution: Costa Rica.

On wet banks and rocks at high altitudes. The longer costa

and the indistinct, concolorous leaf border will aid in separating

this species from the following.

BARTRAM: MOSSES OF GUATEMALA

163

2. EPIPTERYGIUM LEPIDOPILOIDES (C. M.) Par., Suppl. Ind. Bryol.

143. 1900.

Bryum lepidopiloides C. M., Bull. Herb. Boiss. 5: 185. 1897.

Similar to E. immarginatum but smaller and more deeply tinged

with red. Stems under 1 cm. high. Lateral leaves to 2.5 mm. long,

oblong-ovate, short apiculate, entire; costa ending about mid-leaf

or below; cells as in E. immarginatum but colored toward margins

forming a rather indefinite reddish border; dorsal leaves much

smaller, narrowly lanceolate, slenderly acuminate. Sporophyte not

seen. (Fig. 73, A-D.)

Dept. Suchitepequez: Steyermark 1*6850.

Endemic. •

On shaded cliff face at high altitude. Very distinct from E.

Wrightii (Sull.) Lindb. of the West Indies in the oblong, not obovate,

lateral leaves, less abruptly apiculate and the narrower, more

elongated leaf cells.

7. BRACHYMENIUM Schwaegr., Suppl. 21: 131. 1824.

Small to medium sized plants, densely tufted. Leaves imbricated

or contorted when dry; costa strong, often long excurrent; cells

FIGURE 73

A-D, Epipterygium lepidopiloides: A, lateral leaf, X14; B, dorsal leaf, X14;

C, upper leaf cells and margin, X120; D, apex of leaf, X54.

E-G, Brachymenium systylium: E, plant, XI; F, leaf, X14; G, apex of leaf,

X54.

H-J, Brachymenium barbe-montis: H, plant, Xl; I, leaf, X14; J, upper leaf

cells and margin, X270.

164 FIELDIANA: BOTANY, VOLUME 25

rhomboidal above, short rectangular below. Seta elongate; capsule

suberect; peristome double, teeth normally developed, endostome

imperfect, consisting of an irregular basal membrane without

segments or cilia.

1. Costa ex current in a long, hyaline hair point 1. B. systylium

Costa not as above 2

2. Leaves spirally twisted when dry, narrowly bordered 4. B. macrocarpum

Leaves erect-imbricated when dry, unbordered 3

3. Small plants, leaves less than 1 mm. long, cells not incrassate

2. B. barbe-montis

Large plants, leaves 1-1.5 mm. long, cells very incrassate. . . .3. B. mexicanum

1. BRACHYMENIUM SYSTYLIUM (C. M.) Jaeg., Adumb. 2: 117.

1874-75.

Bryum systylium C. M., Syn. 1 : 320. 1849.

Bryum Carionis C. M., Bull. Herb. Boiss. 5: 180. 1897.

Plants 1-3 cm. high, densely tufted, matted with reddish brown

radicles below; stems branched. Leaves crowded, often in inter-

rupted tufts, closely imbricated, unbordered, oblong-ovate, concave,

with long, hyaline hair points; costa strong, excurrent in a long,

subentire hair point; margins erect, denticulate above; cells rhom-

boidal-hexagonal becoming linear toward margins, quadrate toward

base. Seta 1.5 mm. long or longer; capsule suberect, ovoid-cylindric,

3.5 mm. long; lid bluntly conical; peristome teeth slender, brown,

papillose; endostome a low, yellowish membrane slightly exceeding

the rim. (Fig. 73, E-G.)

Dept. Alta Verapaz: Standley 69650, 71730, 717J*3a, 90775. Dept. Huehuete-

nango: Standley 81690, 82867; Steyermark 50598, 50599a. Dept. Quezaltenango :

Standley 81*261, 85738; Steyermark 31*101*, 31*106. Dept. Sacatepequez : Standley

59011. Dept. Solola: Steyermark 1*6959. Dept. Chimaltenango: Standley 61856.

Dept. Jalapa: Standley 75581*.

Distribution: Arizona, New Mexico, Mexico, Central and South

America.

On trees, logs and damp, shaded banks at medium to high alti-

tudes. Variable and frequent but easily recognized by the closely

imbricated, hair-pointed leaves.

2. BRACHYMENIUM BARBE-MONTIS C. M., Bull. Soc. Roy. Bot. Belg.

31 : 165. 1892.

Small, densely tufted, terrestrial plants; stems to 7 mm. high,

slender. Leaves erect and closely imbricated when dry, scarcely

BARTRAM: MOSSES OF GUATEMALA 165

1 mm. long, ovate, concave, aristate; margins erect, entire; costa

strong, excurrent in a stout, concolorous arista; cells narrowly

rhomboidal, not incrassate, narrower toward margins, short rec-

tangular below. Seta slender, reddish, about 15 mm. long; capsule

erect, ovoid with a short neck, wrinkled when dry, 1.5 mm. long; lid

bluntly conical; peristome teeth linear, densely papillose, endostome

a high membrane more than half the height of the teeth; annulus

compound. Sterile stems frequently with small, foliose gemmae in

upper leaf axils. (Fig. 73, H-J.)

Dept. Guatemala: Standley 891*38. Dept. Jalapa: Standley 76785, 77527.

Distribution: Costa Rica.

On shaded banks at moderate altitudes. The small size and

minute, unbordered leaves will distinguish this species without

much trouble.

3. BRACHYMENIUM MEXICANUM Mont., Ann. Sci. Nat. Bot. Ser. II,

9: 54. 1838.

Dull, yellowish green plants, closely tufted; stems to 1 cm. high,

often branched. Lower leaves small, upper crowded in a comal tuft,

closely imbricated, broadly ovate, about 1.5 mm. long, concave,

cuspidate, entire, unbordered; costa excurrent in a short, concolorous

point; upper cells rhomboidal-hexagonal, incrassate, basal cells

quadrate. Seta 12-25 mm. long; capsule erect, ovoid-cylindric,

tapering to a slender neck; lid bluntly conical; annulus broad;

peristome teeth brownish, papillose, endostome about half the

height of the teeth, irregularly laciniate on the edge. (Fig. 74, A-D.)

Dept. Huehuetenango: Standley 82522a. Dept. Quezaltenango : Standley

83261 a. Dept. Jutiapa: Standley

Distribution: Texas, Mexico.

On banks and rocks mostly at rather high altitudes. The short

pointed, unbordered leaves in compact comal tufts give this species

a characteristic look.

4. BRACHYMENIUM MACROCARPUM Card., Rev. Bryol. 38: 6. 1911.

Rather robust plants in green mats or tufts; stems about 1 cm.

high, densely radiculose below, simple or branched. Leaves crowded,

spirally twisted when dry, oblong-ovate, obtuse, cuspidate, to 2 mm.

long, 1 mm. wide; margins strongly recurved, denticulate near apex;

costa stout, excurrent in a toothed, concolorous point; upper cells

166

FIELDIANA: BOTANY, VOLUME 25

vxi

D F

FIGURE 74

A-D, Brachymenium mexicanum: A, plant, Xl; B, leaf, X14; C, upper leaf

cells and margin, X270; D, part of peristome, X120.

E-G, Brachymenium macrocarpum: E, plant, Xl; F, leaf, X14; G, upper leaf

cells and margin, X270.

H-J, Anomobryum filiforme: H, plant, Xl; I, leaf, X14; J, upper leaf cells

and margin, X270.

hexagonal, thin walled, densely chlorophyllose, elongate in 1 or 2

rows at margins forming a narrow border, basal cells quadrate. Seta

red, 1.5-2 cm. long; capsule erect, ovoid-cylindric, small mouthed,

3-4 mm. long; lid conical, curved, a scant 1 mm. long; peristome

teeth brown, densely papillose, endostome a high yellowish, papillose

cylinder, laciniate on the edge. (Fig. 74, E-G.)

Dept. Peten: Lundell 2056.

Distribution: Florida, Mexico.

On rocks, logs and trees at low altitude. Sharply distinct from

the other local species in that the leaves are spirally twisted when

dry with the margins recurved nearly to apex.

8. ANOMOBRYUM Schimp., Syn. Ed. I, 382. 1860.

Dioicous; slender, pale green, glossy plants; stems evenly foliate

and terete. Leaves erect, closely imbricated, ovate, concave, entire;

costa ending below apex; upper cells narrow, more lax and rhomboidal

below. Seta elongate; capsule pendulous; peristome double and

complete.

BARTRAM: MOSSES OF GUATEMALA 167

1. Leaves plicate, upper cells to 50 n long 2. A. plicatum

Leaves not plicate, upper cells to 100 /i long 2

2. Leaves bluntly pointed, costa ending below apex 1. A. filiforme

Leaves acute, costa percurrent 3. A. semiovatum

1. ANOMOBRYUM FILIFORME (Dicks) Husn., Muse. Gall. 2?2. 1888.

Bryum filiforme Dicks., PI. Crypt, fasc. 4: 16. 1801.

?Bryum perappresum C. M., Bull. Herb. Boiss. 5: 182. 1897.

Plants tufted, to 2 cm. or more high, glossy; stems julaceous,

with subfloral innovations. Leaves numerous, erect and rigidly

imbricated, 1-1.5 mm. long, obtuse to broadly acute; margins erect,

denticulate toward apex; costa ending below apex; upper cells linear,

to 100 M or more long, incrassate, vermicular, broader, shorter and

thin walled below. Seta slender, flexuous, to 2 cm. long; capsule

oblong-cylindric, neck distinct, 3-4 mm. long; peristome teeth

yellow, segments of endostome from a high basal membrane, split,

cilia appendiculate. (Fig. 74, H-J.)

Dept. Huehuetenango : Standley 82^28. Dept. San Marcos: Standley 66271;

Steyermark 85675, 35993. Dept. Totonicapan: Standley 65921, 8M25. Dept.

Quezaltenango: Standley 67829, 67861, 83313, 83430, 83377, 852^5, 85987, 8601>5;

Steyermark 33929, 8^596, 8^618, 8^916, 3^920. Dept. Sacatepequez: Standley

65273. Dept. Chimaltenango: Standley 62322, 6^80.

Distribution: Greenland to Alaska south to New York, Wisconsin

and Minnesota, Mexico, Costa Rica, South America, Europe, Africa.

On damp banks and rocks at medium to high altitudes. The

species is quite variable and I cannot satisfactorily segregate the

var. mexicanum (Schimp.) Par. from the type concept. It fruits

freely in Mexico and Costa Rica and is apparently broadly dis-

tributed.

2. ANOMOBRYUM PLICATUM Card., Rev. Bryol. 36: 112. 1909.

More robust than A. filiforme; stems to 5 cm. or more high.

Leaves broadly ovate, about as broad as long, rounded at apex,

crenulate-denticulate about half way down, often with 1 or 2 notice-

able plicae on either side of costa; costa stout, ending below apex;

upper cells narrowly rhomboidal, 8-10 M wide, 25-50 n long, in-

crassate, much narrower and linear in several rows toward margins,

basal cells shorter and broader. (Fig. 75, A-C.)

Dept. Huehuetenango: Standley 81087, 81179 in part.

Distribution: Mexico.

168 FIELDIANA: BOTANY, VOLUME 25

Alpine meadow. Distinct from A. filiforme in the orbicular-

ovate leaves and shorter, broader upper leaf cells.

3. ANOMOBRYUM SEMIOVATUM (Brid.) Jaeg., Adumb. 1 : 602. 1874-

75.

Bryum semiovatum Brid., Bryol. Univ. 1 : 846. 1826.

Slender yellowish green plants forming low, dense mats. Stems

decumbent, to 2 cm. long, julaceous when moist. Leaves laxly

appressed when dry, tightly imbricated when moist, to 1.5 mm.

long, ovate, concave, acute; margins erect, entire; costa percurrent;

upper cells rather lax, linear-rhomboidal, 12-15 /JL wide, 60-100 /z

long, laxer toward base. Fruit not seen. (Fig. 72, H-I.)

Dept. Solola: Svihla 2890b.

Distribution: Costa Rica, Ecuador, Peru.

At moderate altitudes. Unless I am much mistaken this collec-

tion represents a robust form of this species. It is well distinguished

from A. filiforme by the laxer upper leaf cells, percurrent costa and

acute leaf points. Furthermore the leaf points when dry are spread-

ing, not closely appressed, so that the stems lack the characteristic

julaceous appearance of A. filiforme.

9. ACIDODONTIUM Schwaegr., Suppl. 22: 152. 1827.

Dioicous; plants medium sized growing in dense, green tufts

matted together with reddish brown radicles below; stems with

numerous subfloral innovations. Leaves lanceolate, piliform acumi-

nate, bordered; margins recurved below; costa ending below apex

or excurrent; cells hexagonal above, rectangular below. Seta

elongate; capsule large, suberect or nodding, ovoid, long necked;

peristome double, teeth papillose, segments of endostome split into

2 divergent forks, cilia rudimentary; lid conical, short.

1. ACIDODONTIUM MEGALOCARPUM (Hook.) Ren. & Card., Bull. Soc.

Bot. Belg. 31 : 162. 1892.

Bryum megalocarpum Hook, in Kunth., Syn. PI. Aeq. 59: 1822-28.

Addodontium floresianum C. M., Bull. Soc. Bot. Belg. 31 : 162. 1892.

Stems about 2 cm. high. Leaves numerous, spirally contorted

when dry, 3 mm. long, ovate-lanceolate, narrowed to a slender,

toothed, hair-like point; margins narrowly recurved near base, plane

above, denticulate toward apex; costa slender, ending near base of

BARTRAM: MOSSES OF GUATEMALA

169

acumen; upper cells hexagonal, thin walled, 1-2 rows at margins

narrowly linear, not forming a distinct border, basal cells laxly

rectangular. Seta 2.5-3.5 cm. long; capsule ovoid-cylindric, abruptly

contracted to a slender, rugose neck, small mouthed ; peristome teeth

broad below, abruptly contracted to a long subulate point, segments

of endostome from a high basal membrane, forks widely divergent.

(Fig. 75, D-G.)

Dept; Alta Verapaz: Standley 90878.

Distribution: Costa Rica, Ecuador, Colombia.

On trees at moderate altitude. Allowing for a reasonable varia-

tion in leaf outline there seems to be no appreciable difference

between A. floresianum and A. megalocarpum. The conspicuous

capsules and the curiously forked segments are distinctive characters.

10. BRYUM Hedw., Sp. Muse. 178. 1801 in part.

Small to very large, tufted plants; stems with subfloral innova-

tions, radiculose below. Leaves usually ovate-lanceolate, often

bordered with narrower cells, entire or nearly so; costa excurrent

or ending in or near apex; upper cells rhomboidal. Seta terminal,

elongate; capsules clavate or pyriform, mostly horizontal or pen-

FIGURB 75

A-C, Anomobryum plicatum: A, plant, Xl; B, leaf, X14; C, upper leaf cells

and margin, X270.

D-G, Acidodontium megalocarpum: D, plant, Xl; E, leaf, X14; F, part of

peristome, X54; G, capsule, X4.

170 FIELDIANA: BOTANY, VOLUME 25

dulous; peristome double, usually complete, teeth 16, lanceolate,

endostome with a high basal membrane bearing 16 keeled, split

segments and appendiculate cilia; lid convex, apiculate; calyptra

small, fugacious.

1. Leaves silvery white or yellow 2

Plants green, often with a reddish or brownish tinge 3

2. Plants silvery white, capsule oblong with a short neck 3. B. argenteum

Plants yellow, capsule narrowly cylindrical, with a long neck. 4. B. chryseum

3. Capsule short, with a thick, spongy neck, abruptly contracted to seta 4

Capsule elongate, with a slender, tapering neck 5

4. Capsule cylindrical, 2 mm. or more long, leaves 2-3 mm. long

5. B. coronatum

Capsule subglobose, 1 mm. or less long, leaves about 1.5 mm. long

6. B. microbalanum

5. Leaves bluntly pointed 6

Leaves acuminate 7

6. Leaves not bordered, ovate-lanceolate 1. B. Crugeri

Leaves strongly bordered, orbicular-oval 2. B. mnioides

7. Very robust plants, stems to 12 or 15 cm. long, leaves long decurrent

13. B. procerum

Stems shorter, seldom over 3-4 cm. long, leaves scarcely decurrent 8

8. Leaf cells lax, thin-walled 8. B. capillare

Leaf cells firm, thick-walled 9

9. Costa long excurrent, synoicous 9. B. cuspidatum

Costa short excurrent, dioicous 10

10. Stems evenly foliate, leaves contorted and appressed when dry

7. B. pseudotriquetrum

Stems with the leaves imbricated, crowded above, often in rosulate tufts. . . 11

11. Upper leaves in conspicuous rosulate tufts, leaf cells elongate. 10. B. truncorum

Upper leaves crowded but not in rosulate tufts, leaf cells short, 1:2 or less. . 12

12. Leaf margins with short, single teeth 11. B. Mangini

Leaf margins spinose-serrate, teeth often in pairs 12. B. geminidens

1. BRYUM CRUGERI Hampe, in C. M., Syn. 1 : 300. 1849.

Dioicous; rather small, pale green, slightly glossy plants, densely

tufted; stems red, branched. Lower leaves widely spreading, upper

more erect, 1.5-2 mm. long, oblong-ovate, concave, bluntly acute;

margins plane, subentire; costa slender, percurrent; cells narrowly

rhomboidal, linear near margins, lax and broad near base. Seta

1.5-2 cm. long, slender; capsule pendulous, clavate with a tapering

neck; peristome teeth dark brown, segments widely perforate along

keel, cilia appendiculate. (Fig. 76, A-C.)

BARTRAM: MOSSES OF GUATEMALA

171

Dept. Izabal: Standley 721*62. Dept. Zacapa: Steyermark 29389. Dept.

Chiquimula: Steyermark 30878.

Distribution: Florida, Costa Rica, West Indies, South America.

On damp ground at low altitudes. The deeply concave, short

pointed unbordered leaves will readily distinguish this species.

2. BRYUM MNIOIDES (Schimp.) Broth., E. & P. Pflanzenfam. I3:

574. 1904.

Webera mnioides Schimp., Ann. Sci. Nat. 6, Ser. 3: 204. 1876.

Fragile, brownish green plants in dense tufts or cushions. Stems

to 2 cm. high, laxly foliate, with clusters of brownish, septate fila-

ments in the upper leaf axils. Leaves contorted when dry, widely

spreading when moist, orbicular-oval, slightly concave, decurrent,

obtuse or broadly rounded, bordered, subentire to faintly sinuate,

1.5-1.8 mm. long, 1.2 mm. wide; margins erect; costa strong, brown,

ending below apex; upper cells short hexagonal, 3-5 rows at margins

linear with brown, incrassate walls, forming a strong border, basal

cells rectangular. Fruit not seen. (Fig. 78, A-C.)

Dept. El Quiche: Sharp 21*51*. Dept. Baja Verapaz: Sharp 28 11*, 2818.

Distribution: Guadeloupe.

D

FIGURE 76

A-C, Bryum Crugeri: A, plant, Xl; B, leaf, X14; C, upper leaf cells and

margin, X270.

D-G, Bryum argenteum: D, plant, Xl; E, leaf, X14; F, leaf of var. lanatum,

X14; G, capsule, X8.

H-L, Bryum chryseum: H, plant, Xl; I and J, leaves, X14; K, capsule, X8; L,

apex of leaf, X54.

172 FIELDIANA: BOTANY, VOLUME 25

On moist rocks at moderate altitudes. As these collections are

sterile the determination is questionable but the broadly ovate

leaves with the costa ending below the short, blunt apex, compare

favorably with the original description and with specimens from

Guadeloupe.

3. BRYUM ARGENTEUM Hedw., Sp. Muse. 181. 1801.

?Bryum subcorrugatum C. M., Bull. Herb. Boiss. 5: 182. 1897.

?Bryum lagunicolum C. M., Bull. Herb. Boiss. 5: 183. 1897.

Dioicous; small, silvery white, often densely tufted plants; stems

red, fragile. Leaves crowded, imbricated, broadly ovate, acuminate,

entire; costa ending below apex or excurrent in forms; upper cells

hyaline, narrow, basal cells quadrate, chlorophyllose. Seta slender,

red, about 1 cm. high; capsule pendulous, oblong, short necked;

peristome complete. (Fig. 76, D-G.)

Dept. Alta Verapaz: Standley 7174.3. Dept. Huehuetenango : Steyermark

48928; Standley 81111, 81714, 81721. Dept. Totonicapan: Standley 83156, 84447.

Dept. Quezaltenango: Standley 66497, 83400, 83459, 84155; Steyermark 34625a,

34626, 34835. Dept. Sacatepequez : Standley 58644, 58646. Dept. Guatemala:

Standley 80716. Dept. Chiquimula: Steyermark 30611. Dept. Jutiapa: Standley

75711, 78450. Dept. Jalapa: Standley 76713, 77499. Dept. Santa Rosa: Standley

78248.

Distribution: Cosmopolitan.

On banks, rocks, trees, etc., at medium to high altitudes. A

cosmopolitan species with an extensive synonymy. Many of the

above collections represent the var. lanatum (P. B.) Bry. Eur. with

the costa excurrent but there are closely intergrading forms.

4. BRYUM CHRYSEUM Mitt., Journ. Linn. Soc. 12: 304. 1869.

?Bryum guatemalense Hampe, Bull. Herb. Boiss. 5: 182. 1897.

Dioicous; plants small, yellowish, slightly glossy, densely tufted;

stems to 5-6 mm. high, julaceous. Leaves closely imbricated with

spreading points, about 1 mm. long, broadly ovate, concave, slenderly

acuminate, minutely denticulate toward apex; costa excurrent in a

concolorous, denticulate point; upper cells linear-rhomboidal, basal

cells quadrate, chlorophyllose. Seta to 20 mm. long; capsule hori-

zontal, cylindrical with a tapering neck; peristome complete. (Fig.

76, H-L.)

Dept. Quezaltenango: Standley 84815.

Distribution: Mexico.

BARTRAM: MOSSES OF GUATEMALA

173

On dry bank at rather high altitude. The yellowish, terete stems

with the leaf points widely spreading on all sides and the narrow

capsule wij:h a tapering neck clearly distinguish this species from

any form of B. argenteum.

5. BRYUM CORONATUM Schwaegr., Suppl. I2: 103. 1816.

Dioicous; plants green, tufted, radiculose below; stems short,

slender, rarely over 1 cm. high. Leaves numerous, erect and slightly

contorted when dry, to 2-3.5 mm. long, ovate-lanceolate, acuminate,

concave, entire; costa usually excurrent; cells narrowly rhomboidal,

narrower toward margins but not forming a distinct border, broader

and shorter below. Seta slender, red, to 2 cm. or more long; capsule

red, pendulous, oblong, 2-2.5 mm. long, with a short, spongy,

rounded neck; peristome complete. (Fig. 77, A-C.)

Dept. Jutiapa: Standley 76102.

Distribution: Pantropical, north to Florida.

On damp banks, rocks, etc., mostly at low altitudes. Readily

known by the peculiar short necked capsules.

6. BRYUM MICROBALANUM Card., Rev. Bryol. 36: 112. 1909.

Similar to B. coronatum but smaller. Plants yellowish; stems

less than 5 mm. high. Leaves erect, imbricated when dry, about

FIGURE 77

A-C, Bryum coronatum: A, plant, Xl; B, leaf, X14; C, capsule, X8.

D-F, Bryum microbalanum: D, plant, Xl; E, leaf, X14; F, capsule, X8.

G-J, Bryum pseudotriquelrum: G, plant, Xl; H, leaf, X14; I, apex of leaf,

X54; J, upper leaf cells, X270.

174 FIELDIANA: BOTANY, VOLUME 25

1.5 mm. long, lanceolate, acuminate; margins recurved below; costa

excurrent. Seta 1-2 cm. long, slender, reddish; capsule minute,

pendulous, subglobose with a short, thick, rounded neck, about 1 mm.

long. (Fig. 77, D-F.)

Dept. Huehuetenango: Steyermark ^9662.

Distribution: Mexico.

On sandy river flat at rather low altitude. These plants seem to

be identical with the type collection from Mexico and are distin-

guished from B. coronatum by the smaller leaves and shorter, sub-

globose capsules.

7. BRYUM PSEUDOTRIQUETRUM (Hedw.) Schwaegr., Suppl. I2: 110.

1816.

Mnium pseudotriquetrum Hedw., Sp. Muse. 109. 1801.

Synoicous or dioicous; plants rather robust, green toward tips,

brown and radiculose below. Stems erect, to 5 cm. or more high.

Leaves not crowded, contorted when dry, about 3 mm. long, ovate-

lanceolate, short pointed; margins entire or slightly toothed toward

apex, recurved; costa strong, reddish, percurrent or short excurrent;

cells rhomboidal-hexagonal, moderately incrassate, several rows at

margins long and narrow forming a distinct border, shorter, broader

and reddish at base. Seta to 4 cm. or more long; capsule sub-

pendulous, clavate, with a long neck, to 5-6 mm. long; peristome

large, complete. (Fig. 77, G-J.)

Dept. Huehuetenango: Standley 81180 (as B. bimum). Dept. San Marcos:

Steyermark 35971.

Distribution: United States and northward, Europe, Asia.

Wet alpine meadows. These collections are sterile but the vegeta-

tive characters differ in no way from typical plants of farther north.

8. BRYUM CAPILLARE Hedw., Sp. Muse. 182. 1801.

?Bryum Bernoullii C. M., Bull. Herb. Boiss. 5: 183. 1897.

Bryum vulcanicolum C. M., Bull. Herb. Boiss. 5: 184. 1897.

Rather small tufted plants, green or brownish; stems rarely over

1 cm. high. Lower leaves small, distant, upper leaves larger, crowded

in a comal tuft, strongly contorted when dry, obovate, cuspidate,

often serrulate above; margins recurved below; costa ending below

apex to long excurrent; cells broadly hexagonal, thin walled, 2 or 3

rows at margins long and narrow forming a distinct border. Seta

BARTRAM: MOSSES 6F GUATEMALA

175

to 3 cm. long; capsule large, often curved, cylindric with a tapering

neck; peristome large, complete. (Fig. 79, A-D.)

Dept. Izabal: Steyermark 39807. Dept. Huehuetenango: Standley 81087d;

Steyermark ^9130, 50268b. Dept. San Marcos: Steyermark 36086, 36097, 36103.

Dept. Quezaltenango : Steyermark 34139, 34630 (as Brachymenium spirifolium?) ;

Standley 67627 (as Bryum erythroneurori), 8^225, 86005. Dept. Chimaltenango:

Standley 587^0 (as Brachymenium spirifolium?) . Dept. Guatemala: Standley

80601a. Dept. Jutiapa: Standley 75163, 75671.

Distribution: Nearly cosmopolitan.

On damp banks, meadows, walls, etc., at medium to high altitudes.

The strongly contorted leaves, often spirally twisted when dry, with

broad, thin walled cells may usually be relied upon to distinguish

this widespread, variable species. No. 36103 cited above is synoicous

and represents the form usually referred to as var. torquescens (Bry.

Eur.) which, as Andrews remarks in his recent treatment of the

species north of Mexico, differs only in the synoicous inflorescence.

9. BRYUM CUSPIDATUM (Bry. Eur.) Schimp., Syn. (Ed. 2) 430.

1876.

Bryum bimum var. cuspidatum Bry. Eur. fasc. 6/9: 50. 1839.

Synoicous; small, tufted plants. Stems to 5 mm. high, sparingly

radiculose. Leaves erect, flexuous and slightly spreading when

FIGURE 78

A-C, Bryum mnioides: A, plant, XI; B, leaf, X18; C, upper leaf cells and

margin, XllO.

D-F, Bryum cuspidatum: D, plant, XI; E, leaf, X14; F, apex of leaf, X66.

176 FIELDIANA: BOTANY, VOLUME 25

moist, to 4 mm. long, oblong-lanceolate, acuminate; margins re-

curved below, denticulate toward apex; costa long excurrent in a

slightly denticulate arista; upper cells linear-rhomboidal, narrower

in several rows at margins forming an indistinct border, basal cells

oblong, lax, often tinged with red. Seta slender, red, to 3 cm. long;

capsules pendulous, ovoid-cylindrical, reddish brown. (Fig. 78,

D-F.)

Dept. Baja Verapaz: Sharp 5156, 5158.

Distribution : United States and Canada.

On soil at low altitudes. These records are far to the south of

the range previously credited to the species in North America but

they seem to be typical in all essential particulars.

10. BRYUM TRUNCORUM Brid., Sp. Muse. 3: 50. 1817.

Bryum andicola Hook, in Kunth, Syn, PI. Aeq. 1 : 58. 1822.

Bryum streptorhodon C. M., Bull. Herb. Boiss. 5: 179. 1897.

Dioicous; plants usually in dense tufts, green or yellowish above;

stems to 3 cm. high, radiculose below. Leaves contorted when dry,

distant below, the upper crowded in a dense, rosulate tuft, 3-3.5 mm.

long, 1.5 mm. wide, obovate, short acuminate, usually with numerous

brown, septate, cylindrical, papillose propagula in the axils; margins

recurved below, toothed above; costa short excurrent; cells rhom-

boidal-hexagonal, 2-3 rows at margins linear and incrassate forming

a distinct border. Seta 1 or 2 from the same perichaetium, 2 cm.

or more long; capsule large, subpendulous; peristome complete.

(Fig. 79, E-H.)

Dept. Huehuetenango : Standley 81717, 82553. Dept. San Marcos: Standley

66247b. Dept. Totonicapan: Standley 65886a, 65888, 83103. Dept. Quezal-

tenango: Standley 65558, 83396, 83805, 85736, 86065; Steyermark 34241. Dept.

Suchitepequez : Steyermark 46712. Dept. Sacatepequez : Standley 59499, 63077.

Dept. Chimaltenango: Standley 60068, 8003 8a, 80152, 80323. Dept. El Progresso:

Steyermark 43698. Dept. Zacapa: Steyermark 42640. Dept. Chiquimula: Steyer-

mark 30592. Dept. Jutiapa: Standley 76320. Dept. Jalapa: Steyermark 32263,

32482, 32495, 32816; Standley 76571. Dept. Santa Rosa: Standley 78097, 78101.

Distribution: Texas and Arizona, Mexico, West Indies, South

America, also wide in southern hemisphere.

On banks, trees, logs, etc., mostly at medium altitudes. I have

followed Andrews' interpretation of this species, which seems a very

happy solution of a complex problem. The complete synonymy is

evidently very extensive.

BARTRAM: MOSSES OF GUATEMALA

177

11. BRYUM MANGINI Ren. & Card., Rev. Bryol. 36: 115. 1909.

Robust densely tufted plants; stems to 5 cm. high, evenly and

densely foliate, matted together with reddish brown tomentum.

Leaves contorted when dry, obovate, short pointed, to 3.5 mm. long,

1.5 mm. wide; margins strongly revolute nearly to apex, bluntly

serrulate above; costa percurrent; cells short oval-hexagonal, in-

crassate, rarely more than twice as long as wide, narrower toward

margins but not forming a distinct border, larger and yellowish at

base. Seta solitary, slender, 2-3 cm. long; capsule nodding or hori-

zontal, cylindrical with a tapering neck, 4-5 mm. long; peristome

complete, endostome with a high basal membrane, cilia 2-3, short,

nodose or weakly appendiculate. (Fig. 80, A-C.)

Dept. Huehuetenango: Steyermark 50185, 50268. Dept. Chimaltenango:

Standley 58739, 60962, 61857.

Distribution: Mexico.

On trees, logs and limestone bluffs at high altitudes. This seems

to be a well marked species differing from B. truncorum in the

evenly foliate stems and unbordered leaves with shorter, more

incrassate cells.

12. BRYUM GEMINIDENS Bartr., Bryol. 49: 115. 1946.

Robust, densely tufted, yellowish green plants; stems to 5 cm.

high, evenly foliate, densely tomentose. Leaves strongly contorted

H

FIGURE 79

A-D, Bryum capillare: A, plant, XI; B, leaf, X8; C, apex of leaf, X54; D,

upper leaf cells and margin, X120.

E-H, Bryum truncorum: E, plant, XI; F, leaf, X8; G, apex of leaf, X54;

H, upper leaf cells and margin, X120.

178

FIELDIANA: BOTANY, VOLUME 25

F

A-C, Bryum Mangini: A, plant, Xl; B, leaf, X8; C, upper leaf

margin, X270.

D-F, Bryum geminidens: D, plant, Xl; E, leaf, X8; F, upper leaf

margin, X270.

cells and

cells and

when dry, 3.5-4 mm. long, 1.5-2 mm. wide, obovate, abruptly

apiculate; margins recurved below, plane and strongly spinose-

serrate above with the teeth often paired; cells short, incrassate,

oval-hexagonal, about 1:2, several rows at margins long and narrow

forming a distinct border; costa excurrent. Sporophyte unknown.

(Fig. 80, D-F.)

Dept. San Marcos: Between San Sebastian and summit of Volcan Tajumulco,

alt. 3,800-4,600 m., among rocks on top of ridge leading to rocky dome, Steyer-

mark 35519 TYPE.

Endemic.

Distinct from B. Mangini in the plainly bordered leaves, spinose-

serrate above with the teeth often in pairs. From B. procerum it

differs in the leaves, which are not decurrent, and in the shorter,

incrassate areolation.

13. BRYUM PROCERUM Schimp., in Besch., Prodr. Bryol. Mex. 55.

1871.

Rhodobryum elatissimum Bartr. in herbaria.

Plants large to very robust, yellowish green; stems to 15 cm. long,

evenly foliate, densely tomentose nearly to tips. Leaves not crowded,

contorted when dry, to 10 mm. long, ovate-lanceolate, acuminate,

decurrent, strongly bordered; margins narrowly recurved below,

BARTRAM: MOSSES OF GUATEMALA 179

plane and spinose-serrate in upper half with the teeth often in pairs;

costa ending below apex; cells rhomboid-hexagonal, thin walled, to

100 M long, very narrow in 3-4 rows at margins forming a distinct,

pale border, gradually laxer below. . Setae 1-3 from the same peri-

chaetium, red, to 5 cm. long; capsule horizontal, oblong-cylindric

with a tapering neck, to 5 mm. long; lid conical; annulus broad,

compound; peristome teeth brownish, segments of endostome from

a high basal membrane, widely split, cilia 2-3, strongly appendicu-

late; spores 10-12 M. (Fig. 81, A-B.)

Dept. Huehuetenango: Standley 81626, 81768; Steyermark 1*8391, 5061*3.

Dept. San Marcos: Standley 85398, 851*820.; Steyermark 35628. Dept. Totoni-

capan: Standley 62711*a, 62727, 83101, 81*006, 81*1*85. Dept. Quezaltenango :

Steyermark 31*071*, 31*102. Dept. Solola: Steyermark 1*7513, 1*7562. Dept. Chimal-

tenango: Standley 5876 1*, 60967, 61109a, 61831.

Distribution: Mexico.

On damp banks, rocks, trees etc. at medium to high altitudes.

In addition to the robust habit these plants are distinguished by

the ovate-lanceolate, decurrent and strongly bordered leaves with

the margins spinose-serrate above and the elongate, rhomboidal,

thin walled cells.

EXCLUDED SPECIES

Bryum pergracilescens C. M., Bull. Herb. Boiss. 5: 184. 1897.

Bryum perminutum C. M., Bull. Herb. Boiss. 5: 184. 1897.

No material relating to either of these species is available for

comparison.

11. RHODOBRYUM (Schimp.) Limpr., Laubm. 2: 444. 1892.

Bryum subg. Rhodobryum Schimp., Syn. 381. 1860.

Robust terrestrial plants, stoloniferous, in lax mats; stems erect,

often interruptedly foliate. Lower leaves small and distant, upper

leaves often crowded in rosette-like tufts, bordered, serrate above;

costa strong; upper cells rhomboidal, basal cells rectangular. Seta

single or aggregated, elongate; capsules large, pendulous; peristome

complete.

1. Leaf border 5-6 cells wide above 2. R. confluens

Leaf border about 2 cells wide above 2

2. Plants yellowish, leaves oblong, little narrowed below 3. R. utriculosum

Plants dark green tinged with red, leaves spatulate 1. R. Beyrichianum

180

FIELDIANA: BOTANY, VOLUME 25

D

FIGURE 81

A-B, Bryum procerum: A, leaf, X8; B, upper leaf cells and margin, X120.

C-D, Rhodobryum Beyrichianum: C, leaf, X4; D, upper leaf cells and margin,

X120.

1. RHODOBRYUM BEYRICHIANUM (Hornsch.) Par., Ind. Bryol. 1115.

1894-98.

Mnium Beyrichianum, Hornsch., Fl. Bras. 1 : 45. 1840.

Plants large, dark green tinged with red; stems 2-6 cm. or more

high, radiculose below. Lower leaves small, upper much larger,

crowded or in rosulate tufts, contorted when dry, to 12 mm. long,

6 mm. wide, broadly spatulate, short acuminate; margins slightly

recurved at extreme base, plane and sharply spinose-serrate above;

costa strong, percurrent; cells oval-hexagonal, thin walled, 2-3 rows

at margins elongate forming a narrow distinct border. Seta 4 cm.

or more long; capsule large, curved, cylindric with a tapering neck.

(Fig. 81, C-D.)

Dept. Peten: Lundell 2730; Bartlett 12604, 12636. Dept. El Progresso;

Steyermark 43530, 43780. Dept. Chiquimula: Steyermark 30843.

Distribution: Mexico, Central America, South America.

On ground at medium altitudes. A critical study of the tropical

American species is essential before the species and their respective

ranges can be limited with any satisfaction.

2. RHODOBRYUM CONFLUENS (C. M.) Par., Ind. Bryol. 1115. 1894-

98.

Bryum confluens C. M., Bull. Herb. Boiss. 5: 179. 1897.

BARTRAM: MOSSES OF GUATEMALA

181

Rather small yellowish green plants, laxly tufted; stems to 6-7 cm.

high, proliferous from the comal tufts. Upper leaves in small rosu-

late tufts, contorted when dry, to 8 mm. long, 4 mm. wide, obovate,

short acuminate, strongly bordered; margins recurved more than

half way up, plane and serrulate above; costa short excurrent; upper

cells oval-hexagonal, thin walled, 25 n wide, 50 n long, linear and

incrassate in 5-6 rows at margins forming a wide, distinct border,

lax and rectangular below. Sporophyte unknown. (Fig. 82, A-B.)

Dept. Huehuetenango: Steyermark 50076 (as Bryum truncorum). Dept.

Chimaltenango: Standley 57819a (as R. Beyrichianum).

Endemic.

Wet, shaded bank at moderately high altitude. I have not seen

the original collection but these specimens agree closely with the

description and seem to be well distinguished by the widely bordered

leaves with short upper cells.

3. RHODOBRYUM UTRICULOSUM (C. M.) Par., Ind. Bryol. 1122.

1894-98.

Bryum utriculosum C. M., Bull. Herb. Boiss. 5: 180. 1897.

Plants robust, yellowish green; stems to 5 cm. high, nearly naked

below. Upper leaves in large rosulate tufts, to 11 mm. long, 2.5 mm.

B

D

X120.

FIGURE 82

A-B, Rhodobryum confluens: A, leaf, X8; B, upper leaf cells and margin, X120.

C-D, Rhodobryum utriculosum: C, leaf, X4; D, upper leaf cells and margin,

182 FIELDIANA: BOTANY, VOLUME 25

wide, oblong-lanceolate, acuminate; margins recurved about half way

up, undulate, plane and sharply serrate above; costa percurrent;

upper cells rhomboidal-hexagonal, thin walled, 1 : 3 or 4, one or two

rows at margins elongate forming a narrow, indistinct border, basal

cells laxly rectangular. (Fig. 82, C-D.)

Dept. San Marcos: Steyermark 87278.

Endemic.

Moist pine slopes at moderate altitude. The oblong leaves from

a scarcely narrower base with the margins recurved seem clearly to

separate this species from R. Beyrichianum. I have not seen the

type and the determination is based on the description.

EXCLUDED SPECIES

BRYUM LATO-CUSPIDATUM C. M., Bull. Herb. Boiss. 5: 180. 1897.

The type is not available and the species cannot be placed from

the description.

15. MNIACEAE

Medium sized, broad leaved plants in tufts or mats. Leaves

large, the upper often in rosulate tufts, short pointed, bordered,

serrate with single or paired teeth; costa strong; cells broadly hexa-

gonal. Seta elongate, single or aggregated; capsules usually pendu-

lous, oblong, short necked; peristome double, complete, bryoid in

structure.

1. MNIUM Hedw., Sp. Muse. 188. 1801.

Plants with the characters of the family. Setae often aggregated ;

capsules oblong-ovoid, subpendulous; peristome double, complete,

segments of endostome from a high basal membrane, cilia nodose.

Leaves oblong or obovate, apiculate, marginal teeth single. . . .1. M. longirostrum

Leaves ovate, acuminate, marginal teeth in pairs 2. M. serratum

1. MNIUM LONGIROSTRUM Brid., Muse. Recent. 23: 106. 1803.

Mnium orbifolium C. M., Bull. Herb. Boiss. 5: 176. 1897.

Synoicous; plants in loose mats; fertile stems about 2 cm. high,

sterile stems longer. Leaves large, to 7 mm. long, obovate, rounded

above, short apiculate, bordered with 3-4 rows of narrow cells;

BARTRAM: MOSSES OF GUATEMALA

183

margins bluntly serrate with short, single teeth well toward base;

costa percurrent; cells rounded-hexagonal with thickened corners.

Seta solitary or aggregated, about 2 cm. long; capsule pendulous,

oblong, urn to 3 mm. long; lid long rostrate; annulus broad; peristome

large and well developed. (Fig. 83, A-D.)

Dept. Alta Verapaz: Standley 70050, 71629, 91392. Dept. Huehuetenango :

Steyermark 1*8756. Dept. San Marcos: Steyermark 35731, 37518. Dept. Chimal-

tenango: Standley 61821. Dept. El Progresso: Steyermark 1*31*51. Dept. Zacapa:

Steyermark 1*2388. Dept. Chiquimula: Steyermark 3081*2. Dept. Jalapa: Steyer-

mark 32839.

Distribution: Cosmopolitan, mostly in tropical and subtropical

regions.

On banks, logs, trees, etc., at medium to high altitudes. Frequent

and usually fruiting.

2. MNIUM SERRATUM Brid., Muse. Rec. 22: 84. 1803.

Synoicous; rather slender, greenish plants, laxly tufted; stems

erect, simple, laxly foliate, to 2 cm. high. Leaves crisped when dry,

about 3 mm. long, oblong-ovate, short acuminate, with a strong

reddish border, serrate with short, usually paired teeth; costa strong,

smooth on back, percurrent; upper cells rounded-quadrate with

noticeably thickened corners, basal cells more elongate. Seta solitary,

FIGURE 83

A-D, Mnium longirostrum: A, plant, XI; B, leaf, X4; C, upper leaf cells and

margin, X120; D, capsule, X4.

E-F, Mnium serratum: E, leaf, X8; F, upper leaf cells and margin, X120.

184 FIELD IANA: BOTANY, VOLUME 25

to 2 cm. long; capsule large, oblong-cylindric; lid rostrate; peristome

complete. (Fig. 83, E-F.)

Dept. San Marcos: Steyermark S5711a, 35713, 35832. Dept. Huehuetenango :

Steyermark 48399.

Distribution: Wide in temperate North America, Europe, Asia.

On wet rocks, banks and in meadows at higk altitudes. The

local collections are all sterile and often poorly developed but they

surely belong here.

16. DREPANOPHYLLACEAE

Small to medium sized laxly tufted plants. Leaves in 4 rows,

laterally spreading on both sides, very inequilateral, broad and

convex on one side of costa, narrow and concave on the other side;

costa percurrent; cells short. Seta terminal, elongate; capsule erect;

peristome single.

1. MNIOMALLIA C. M., in Journ. Mus. Godeffr. 5:60. 1873-74.

Small, slender, gregarious plants; stems branched, complanate-

foliate. Leaves very unequally divided by the costa, obliquely ovate,

short pointed; costa strong; cells rounded, smooth or papillose.

Sporophyte not seen.

1. MNIOMALLIA VIRIDIS (Mitt.) C. M., Journ. Mus. Godeffr. 5: 61.

1873-74.

Drepanophyllum viride Mitt., Journ. Linn. Soc. 12: 318. 1869.

Mniomallia Bernoullii C. M., Bull. Herb. Boiss. 5: 176. 1897.

Plants to 1 cm. high, dull green; stems branched, often with

clustered brood filaments at tips, radiculose below, 1.5 mm. wide

with leaves. Leaves numerous, obliquely inserted, to 0.7 mm. long,

oblong-ovate, acute, arcuate when moist, overlapping, very asym-

metrical; margins narrowly inflexed above and minutely serrulate

in upper half; costa strong, nearer the concave side, percurrent;

cells rounded-quadrate with firm, pellucid walls, coarsely and dis-

tinctly papillose. Sporophyte unknown. (Fig. 84, A-C.)

Dept. Alta Verapaz: Steyermark M995a.

Distribution: Brazil, Ecuador.

On log at medium altitude. This interesting and highly individual

species has much the appearance of a small Fissidens to the naked

BARTRAM: MOSSES OF GUATEMALA 185

eye but under a microscope the oddly shaped leaves are unmistak-

able. I have not seen the original of M. Bernoullii but certainly the

description suggests nothing different.

17. EUSTICHIACEAE

Slender, bright green plants in dense tufts, interwoven with

brownish radicles below; stems branched. Leaves numerous, dis-

tichous, deeply carinate, ovate, cuspidate; margins erose-denticulate;

costa strong, excurrent; cells small, papillose. Seta slender, elongate;

capsules suberect; peristome teeth lacking, endostome of 16 vertically

striolate segments, slightly perforate, united at base; lid long and

slenderly beaked; calyptra cucullate.

1. EUSTICHIA (Brid.) Mitt., Journ. Linn. Soc. 12: 603. 1869.

Phyllogonium sect. Eustichia Brid., Bryol. Univ. 2: 674. 1827.

Plants with the characters of the family.

1. EUSTICHIA SPRUCEANA (C. M.) Par., Ind. Bryol. Suppl. 153. 1900.

Diplostichum Spruceanum C. M., Hedwigia 36: 85. 1897.

?Diplostichum miradoricum C. M., Hedwigia 36: 85. 1897.

Stems to 4 cm. high, considerably branched, about 1 mm. wide

with leaves. Leaves spreading, with erect or incurved points, under

1 mm. long, deeply carinate-concave, cuspidate by the strong,

pellucid, excurrent costa; margins erect, finely and irregularly

denticulate; cells 8-10 n, rather obscure, papillose, several rows at

margins often slightly elongate but not forming a distinct border.

Seta 1.5-2 cm. long; capsule ovoid, wide mouthed, erect or curved,

sulcate when dry. (Fig. 84, D-F.)

Dept. San Marcos: Steyermark 35700, 35703, 36^8. Dept. Quezaltenango :

Standley 837 1*5.

Distribution: Mexico, Costa Rica, South America.

Moist cliffs and bluffs at high altitudes. The various species

proposed by Muller in Hedwigia 1897 appear to be based on very

trivial distinctions. My numerous specimens from various parts of

South America look much alike and I suspect they are merely forms

of one broadly distributed species possibly referable to E. longirostris

(Brid.). Dr. Reimers advised me some years ago that the original

of E. miradorica (C. M.) from Mexico could not be located in

186

FIELDIANA: BOTANY, VOLUME 25

FIGURE 84

A-C, Mniomallia viridis: A, plant, Xl; B, leaf, X54; C, upper leaf cells and

margin, X270.

D-F, Eustichia Spruceana: D, plant, Xl; E, leaf, X28; F, upper leaf cells and

margin, X270.

G-I, Rhizogonium Lindigii: G, plant, Xl; H, leaf, X28; I, upper leaf cells

and margin, X270.

Berlin but the brief description leaves little doubt that it is the

same as E. Spruceana.

18. RHIZOGONIACEAE

Plants medium sized, densely tufted; stems erect, radiculose

below. Leaves narrow, spreading, strongly serrate with single or

paired teeth; costa strong; cells small, rounded, incrassate, smooth.

Setae elongate, lateral near base of stem; capsule nodding or hori-

zontal, curved; lid beaked; annulus present; peristome double,

complete.

1. RHIZOGONIUM Brid., Bryol. Univ. 2: 663. 1827.

Plants with the characters of the family.

Leaves in 2 rows, bifarious, serrate with single teeth 1. R. Lindigii

Leaves spreading on all sides, serrate with paired teeth 2. R. spiniforme

1. RHIZOGONIUM LINDIGII (Hampe) Mitt., Journ. Linn. Soc. 12:

328. 1869.

Mnium Lindigii Hampe, Ann. Sci. Nat. ser. V, 4: 345. 1865.

BARTRAM: MOSSES OF GUATEMALA 187

Dioicous; slender, yellowish brown, glossy plants; stems about

2 cm. high, nearly naked below, flexuous. Lower leaves small and

distant, the upper numerous in 2 opposite rows, 1-1.5 mm. long,

ovate, cuspidate by the excurrent costa, not bordered, coarsely

incised serrate with simple teeth. Inflorescence basal. Sporophyte

not seen. (Fig. 84, G-I.)

Dept. Zacapa: Steyermark 43289.

Distribution: Costa Rica, Colombia, British Guiana, Brazil.

On shaded bluffs at high altitude. The bifarious, unbordered

leaves distinguish this species at a glance. This is the northern

limit of the range as now known.

2. RHIZOGONIUM SPINIFORME (Hedw.) Bruch, Flora 29: 134. 1846.

Hypnum spiniforme Hedw., Sp. Muse. 236. 1801.

Normally synoicous; inflorescence basal. Plants yellowish green

in deep tufts, densely radiculose at base; stems to 3-5 cm. high,

flexuous. Leaves numerous, not crowded, linear-lanceolate, gradu-

ally acuminate, to 8 mm. long; margins thickened, spinose-serrate

with paired teeth from near base; costa strong, toothed on back

above; cells rounded, incrassate, several rows at margins in 2 layers

forming a thickened border. Seta slender, flexuous, to 7 cm. long;

capsule nodding or horizontal, curved; lid obliquely beaked; peris-

tome teeth brownish, close, segments of endostome narrow from a

high basal membrane, cilia nodose. (Fig. 85, A-C.)

Dept. Izabal: Steyermark 38895, 41895. Dept. Alta Verapaz: Standley 90412,

90^21, 90433, 91420, 91662, 91667, 91962. Dept. Huehuetenango: Steyermark

48798. Dept. San Marcos: Steyermark 37260. Dept. Zacapa: Steyermark 29829,

42556, 43225. Dept. Chiquimula: Steyermark 30815.

Distribution: Cosmopolitan in tropical and subtropical regions

reaching the southeastern United States.

On logs, trees and humus at low altitudes. Frequent and usually

fruiting.

19. MEESEACEAE

Plants of bogs and wet places; stems erect. Leaves spreading,

lanceolate, cells small, smooth; costa single, strong. Seta terminal,

long; capsules curved, with a long, prominent neck; peristome double,

the teeth usually blunt and shorter than the segments, cilia rudi-

mentary; lid short, conical.

188

FIELDIANA: BOTANY, VOLUME 25

E

FIGURE 85

A-C, Rhizogonium spiniforme: A, plant, Xl; B, leaf, X8; C, upper leaf cells

and margin, X270.

D-F, Meesea longiseta: D, leaf, X8; E, capsule, X8; F, apex of leaf, X120.

G-I, Anacolia laevisphaera: G, plant, Xl; H, leaf, X14; I, basal leaf cells, X270.

1. MEESEA Hedw., Sp. Muse. 173. 1801.

Plants with the characters of the family. Leaves decurrent; costa

ending below apex. Segments of endostome often transversely

connected.

1. MEESEA LONGISETA Hedw., Sp. Muse. 173. 1801.

Synoicous; plants rather densely tufted, yellowish green; stems

densely radiculose below. Leaves numerous, contorted when dry,

lanceolate from an ovate base, to 3-4 mm. long, acute or blunt at

apex; margins entire, plane or narrowly recurved toward base; costa

ending below apex; upper cells rhomboidal, 2-4:1, rectangular toward

base. Seta to 8 or 10 cm. long but often shorter; capsule curved,

nodding, pyriform with a long neck; peristome teeth short, obtuse,

segments of endostome much exceeding the teeth. (Fig. 85, D-F.)

Dept. Alta Verapaz: Standley 92739.

Distribution: Rare and local in North America, Honduras (as

M. Ulei C. M.), Europe, Asia.

Terrestrial in bogs at moderate altitude. The unexpected occur-

rence of this species in Guatemala naturally led to a comparison

with Standley's No. 56214a from Honduras, which I referred to

BARTRAM: MOSSES OF GUATEMALA 189

M. Ulei C. M. After comparing the two Central American collec-

tions with each other and with M. longiseta I am convinced that they

are identical. The leaves of M. longiseta are invariably described

as plane margined but I find the basal edges are often narrowly

recurved on one or both sides.

20. BARTRAMIACEAE

Small to robust tufted plants; stems branched or with whorled

subfloral innovations. Leaves usually narrow and acute; cells narrow,

papillose at or near end walls. Setae short or long; capsules mostly

globose and cernuous, ribbed when dry. Peristome usually double

or imperfect, teeth 16, segments of endostome shorter than teeth,

often poorly developed; lid convex or conical. .

1. Seta short, 2-3 mm. long 2

Seta elongate 3

2. Peristome lacking, dioicous 1. Anacolia

Peristome present, synoicous 2. Leiomela

3. Synoicous, very small plants 4. Bartramidula

Dioicous, plants larger 4

4. Leaves plicate, at least at base, alar cells differentiated 6. Breutelia

Leaves not plicate, alar cells not differentiated 5

5. Leaves linear-lanceolate from a sheathing base 3. Bartramia

Leaves lanceolate, not sheathing at base 5. Philonotis

1. ANACOLIA Schimp., Syn. Ed. 2, 513. 1876.

Dioicous; plants fairly large, tufted, densely felted with brown

tomentum below. Leaves appressed when dry, lanceolate; costa

stout, percurrent or excurrent; margins recurved below, plane and

serrate above; cells oblong, usually papillose. Seta terminal, short;

capsules nearly erect, subglobose; peristome lacking or of 16 fragile,

rudimentary teeth.

Basal cells elongate, lamina cells in 2 layers 1. A. laevisphaera

Basal cells short, lamina cells in 1 layer 2. A. intertexta

1. ANACOLIA LAEVISPHAERA (Tayl.) Flowers, Moss Fl. of No. Amer.

23: 155. 1935.

Glyphocarpus laevisphaera Tayl., Lond. Journ. Bot. 1846: 56. 1846.

Bartramia subsessilis Tayl., Lond. Journ. Bot. 1847: 334. 1847.

Plants yellowish green, closely tufted; stems 2-4 cm. long,

branched. Leaves crowded, erect with flexuous points when dry,

190 FIELDIANA: BOTANY, VOLUME 25

3-4 mm. long, narrowly lanceolate, long acuminate, sharply serrate

above; costa excurrent; upper cells in 2 layers, oblong, papillose,

basal cells rectangular, to 65 // long, shorter toward margins. Seta

2-3 mm. long; capsule erect, globose, 2 mm. in diameter, small

mouthed; peristome none as seen; spores papillose, about 25 n. (Fig.

85, G-I.)

Dept. Quezaltenango : Standley 84.170.

Distribution: Arizona, New Mexico, Mexico, wide in South

America.

On damp bank at high altitude. Uniformly sterile in North

America as far as I know but readily known by the elongated basal

leaf cells and the bistratose upper cells. The sporophyte characters

are described from fruiting plants collected in Ecuador.

2. ANACOLIA INTERTEXTA (Schimp.) Jaeg., Adumb. 2: 699. 1879.

Bartramia intertexta Schimp. in C. M., Syn. 1: 503. 1849.

Robust plants, yellowish, strongly tinged with brown, growing

in extensive mats; stems slender, decumbent, to 7 or 8 cm. long,

copiously branched, densely felted with red tomentum nearly to

tips. Leaves appressed when dry, 3 mm. long, lanceolate from an

ovate, plicate base, subulate-acuminate; margins strongly recurved

more than half way up, serrulate nearly to base; costa excurrent;

cells rounded, incrassate, in one layer, coarsely papillose, narrower

toward margins but not elongate below. Perichaetial leaves longer,

setaceous pointed; seta 2-3 mm. long, curved; capsule large, globose,

pale brown, glossy, diameter 3 mm.; peristome none. (Fig. 86,

A-C.)

Dept. Huehuetenango: Standley 81090, 81677, 81691, 81852, 83088a; Steyer-

mark 50231. Dept. San Marcos: Steyermark 35896 (c. fr.). Dept. Quezaltenango:

Steyermark 3^764, 3^765; Standley 8^180. Dept. Solola: Steyermark 1^6932.

Distribution: Mexico.

On limestone rocks, banks and trees at high altitudes. Mitten

confused this species with A. setifolia as explained by TheYiot. The

short, rounded distinct leaf cells of A. intertexta are very distinctive.

2. LEIOMELA (Mitt.) Broth., E. & P. Pflanzenf. I3: 634. 1904.

Bartramia subsec. Leiomela Mitt, in part, Journ. Linn. Soc. 12: 253. 1869.

Synoicous; robust, dull yellowish green plants, densely tufted,

felted with brown tomentum below; stems erect, branched. Leaves

narrow, setaceous, serrulate; costa long excurrent; cells narrowly

BARTRAM: MOSSES OF GUATEMALA

191

rectangular, papillose at apical angles. Perichaetial leaves longer

than stem leaves; seta terminal, short; capsule ovoid; lid plano-

convex; peristome teeth 16, deeply inserted, endostome rudimentary.

1. LEIOMELA BARTRAMIOIDES (Hook.) Par., Ind. Bryol. Ed. 2, 3:

132. 1905.

Leucodon bartramioides Hook., Ic. PI. Rar. 1: tab. 71. 1837.

Stems to 7 cm. high. Leaves crowded, erect-spreading when

dry, to 8 mm. long, linear-subulate from an erect, pale, oblong base;

margins plane, minutely serrulate nearly to base; costa excurrent,

toothed on back above; upper cells oblong, obscure, coarsely papillose,

basal cells linear, smooth, hyaline, brownish near insertion. Peri-

chaetial leaves 14-16 mm. long, with long, fragile, capillary, con-

colorous points; seta 1 mm. or less long; capsule immersed, 2.5 mm.

long, 1.5 mm. wide, pale and rather glossy, smooth; peristome teeth

irregularly cleft. (Fig. 86, D-F.)

Dept. Alta Verapaz: Standley 71682. Dept. Huehuetenango : Steyermark

1*875]*, 1*9790 (c. fr.). Dept. San Marcos: Standley 68552a. Dept. Quezaltenango :

Standley 85997, 8601Sa. Dept. Chimaltenango: Standley 61080.

Distribution: Costa Rica, Jamaica, South America.

On trees at medium to high altitudes. Easily recognized by the

very narrow, plane margined leaves and immersed capsules. Numer-

FIGURE 86

A-C, Anacolia interiexta: A, plant, Xl; B, leaf, X14; C, basal leaf cells, X270.

D-F, Leiomela bartramioides: D, plant, Xl; E, leaf, X8; F, upper leaf cells,

X270.

G-I, Bartramia microstoma: G, plant, Xl; H, leaf, X8; I, capsule, X8.

192 FIELDIANA: BOTANY, VOLUME 25

ous purplish or brown, naked, filiform shoots are often produced

near the tips of the sterile stems. These shoots bear clusters of

rhizoids from which grow minute plants.

3. BARTRAMIA Hedw., Sp. Muse. 164. 1801.

Plants erect, tufted, bright or yellowish green; stems branched.

Leaves narrow, serrulate; costa strong, often excurrent; upper cells

quadrate or elongate, papillose, obscure, basal cells linear, smooth.

Seta short to elongate; capsules exserted, subglbbose, furrowed

when dry; peristome double, single or none, endostome often rudi-

mentary. The local species all belong in the section Vaginella.

1. Perichaetial leaves to 10-12 mm. long, much longer than stem leaves

3. B. Mathewsii

Perichaetial leaves not differentiated 2

2. Leaves erect, rigid, appressed when dry 2. B. potosica

Leaves with spreading points when dry 1. B. microstoma

1. BARTRAMIA MICROSTOMA Mitt., Journ. Linn. Soc. 12: 272. 1869.

Synoicous; plants compactly tufted, green above, brown below;

stems 2-4 cm. high. Leaves crowded, suberect with spreading,

curved points when dry, 3-5 mm. long, linear-subulate from an erect,

oblong, hyaline, sheathing base, wider at shoulders than below,

subula opaque, serrulate; margins narrowly recurved just above

shoulders, plane above; costa obscure above, excurrent; upper cells

small, oblong, obscure, papillose. Seta slender, 6-8 mm. long,

straight or curved; capsule nodding, ovoid, 1.5 mm. long, small

mouthed; peristome teeth reddish brown, about 225 n long, endos-

tome rudimentary; lid convex with a blunt point; spores reniform,

to 28 M long. (Fig. 86, G-I.)

Dept. San Marcos: Steyermark 36116 (as B. Mathewsii). Dept Quezaltenango:

Standley 66^08, 664U, 67749a. Dept. Sacatepequez: Standley 65202.

Distribution: Arizona.

On shaded banks at rather high altitudes. The sporophyte is

very similar to that of B. potosica excepting the endostome, which

appears to be constantly more rudimentary.

2. BARTRAMIA POTOSICA Mont., Ann. Sci. Nat. Ser. II, 9: 56. 1838.

Dioicous; plants yellowish green; stems 2-3 cm. high, usually

simple, densely radiculose below. Leaves rigidly erect and appressed

BARTRAM: MOSSES OF GUATEMALA

193

when dry, fragile, points often broken off, 4-6 mm. long, abruptly

linear-subulate from an erect, oblong, hyaline base, broader at

shoulders than below, subula opaque, sharply serrulate; costa obscure

above, excurrent; upper cells narrowly oblong, obscure, papillose.

Seta 3-6 mm. long, usually curved; capsule suberect, ovoid, glossy,

1.5-2 mm. long; peristome double, teeth brown, segments of endos-

tome shorter than teeth; spores reniform, 22-28 M- (Fig. 87, A-D.)

Dept. Totonicapan: Standley 65936. Dept. Quezaltenango: Steyermark 34192,

34193.

Distribution: Mexico, Colombia to Chile.

•

On shaded banks and rocks at high altitudes. Noticeably distinct

from B. microstoma in the rigidly erect, fragile leaves. Brotherus

includes this species in the dioicous group while Mitten describes it

as synoicous. The plants I have examined are dioicous but the

inflorescence may be variable.

3. BARTRAMIA MATHEWSII Mitt., Journ. Linn. Soc. 12: 273. 1869.

Dioicous? Plants yellowish green above, brown below; stems

simple or sparingly branched, to 3 cm. high. Leaves crowded, erect,

appressed, brittle, the points often broken off, 4-6 mm. long, similar

FIGURE 87

A-D, Bartramia potosica: A, plant, Xl; B, leaf, X8; C, broken leaf, X8;

D, upper leaf cells and margin, X270.

E-G, Bartramia Mathewsii: E, plant, Xl; F, stem leaf, X8; G, comal leaf, X8.

H-J,tBartramidula patula: H, two plants, Xl; I, leaf, X22; J, upper leaf cells

and margin, X346.

194 FIELDIANA: BOTANY, VOLUME 25

in structure to those of B. potosica; margins narrowly recurved just

above shoulders. Perichaetial leaves much longer, 10-12 mm. long,

with long, capillary, concolorous points; capsule erect, oblong,

globose; peristome imperfect, teeth none? Segments well developed.

Sporophyte not seen. (Fig. 87, E-G.)

San Marcos: Steyermark 3551>6b, 355M, 36115.

Distribution: Ecuador, Peru.

Rock crevices and boulders at very high altitudes. The con-

spicuously long pointed perichaetial leaves are suggestive of Leiomela

but the stem leaves follow the pattern of Bartramia. These collec-

tions agree perfectly with the original collection by Mathews from

Peru.

Brotherus interprets Mitten's ambiguous description of the

sporophyte to mean that the inner peristome only is present. How-

ever a single capsule in a collection from Chile on the sheet bearing

the type specimen shows a short, curved seta about 4 mm. long and

the teeth of the outer peristome well developed. Until this problem

is clarified the peristome structure must remain in doubt.

4. BARTRAMIDULA Schimp., Bry. Eur. fasc. 29-30. 1846.

Small, slender plants with whorled, subfloral innovations. Leaves

small, lanceolate, erect-spreading; costa percurrent; cells oblong,

papillose. Seta slender, straight or curved; capsule small, sub-

globose; peristome usually lacking.

Peristome lacking 1. B. patula

Peristome present 2.. B. Turckheimi

1. BARTRAMIDULA PATULA (Mitt.) Jaeg., Adumb. 2: 698. 1877-78.

Bartramia patula Mitt., Journ. Linn. Soc. 12: 255. 1869.

Synoicous; plants less than 1 cm. high with several slender sub-

floral innovations. Leaves erect-spreading, 1-1.5 mm. long, lanceo-

late, acuminate; margins plane, serrate above middle; costa percur-

rent; cells narrowly oblong to linear, smooth to very faintly papillose.

Seta 5-8 mm. long, flexuous or arcuate, slender; capsule globose-

pyriform, brown, rugulose, about 1.5 mm. long, mouth small;

peristome lacking. (Fig. 87, H-J.)

Volcan de Agua; Godman & Salvin.

Endemic.

BARTRAM: MOSSES OF GUATEMALA

195

Evidently a rare, local species. The only plants I have seen are

from the Mitten Herbarium in New York.

2. BARTRAMIDULA TURCKHEIMI (C. M.) Par., Ind. Bryol. Suppl.

36. 1900.

Bartramia Turckheimi C. M., Bull. Herb. Boiss. 5: 187. 1897.

Synoicous; small, densely tufted plants, yellowish green, matted

together with brown tomentum below. Stems to 7 mm. high, tipped

with 4-6 short, whorled innovations. Leaves crowded, erect-spread-

ing, to 2 mm. long, narrowly lanceolate, slenderly acuminate;

margins recurved; costa excurrent in a long, denticulate hair-point;

leaf cells linear, papillose at upper ends. Seta curved, to 7 mm.

long; capsule globose, sulcate, 1.5 mm. in diameter; peristome

double, teeth about 110 M high, broad, brownish, truncate, smooth,

endostome rudimentary, fragments as long as teeth, pale yellow,

minutely papillose, segments and cilia lacking; spores brown,

diameter 40-45 M- (Fig. 88, A-C.)

Dept. El Quiche: Sharp 2568.

Endemic.

On bank at moderate altitude. This is a noteworthy collection

as the species is apparently known only from the type gathering by

FIGURE 88

A-C, Bartramidula Turckheimi: A, plant, Xl; B, leaf, X26; C, part of peri-

stome, X68.

D-G, Philonotis Bernoullii: D, plant, Xl; E, leaf, X28; F, apex of leaf, X68;

G, capsule, X8.

196 FIELDIANA: BOTANY, VOLUME 25

Turckheim from near Coban, Alta Verapaz, and no material is

available in the American herbaria. Muller describes the capsules

as gymnostomous but Brotherus notes (E. & P. Ed. 2, 10: 460) that

an examination of an unripe capsule from the type collection shows

a peristome structure but only fragments of the outer peristome were

seen. This observation is confirmed by Sharp's collection, which is

in good fruit and shows the peristome described above.

5. PHILONOTIS Brid., Bryol. Univ. 2: 15. 1827.

Plants of varying size, partial to wet places. Stems with whorled,

subfloral branches. Leaves appressed when dry, lanceolate; costa

strong, percurrent or excurrent; cells narrow, usually papillose. Seta

elongate; capsules subglobose, cernuous, furrowed when dry; peris-

tome double, teeth 16, segments of endostome from a high basal

membrane, cilia well developed.

1. Autoicous 1. P. longiseta

Dioicous 2

2. Stems hooked at tips .'6. P. uncinata

Stems not hooked 3

3. Leaves obtuse, costa ending below apex 3. P. gracillima

Leaves acute or acuminate, costa percurrent or excurrent 4

4. Costa percurrent 2. P. glaucescens

Costa long excurrent 5

5. Robust plants, seta erect, 2 cm. or more long 4. P. sphaericarpa

Very slender plants, setae flexuous or curved, less than 1.5 cm. long

5. P. Bernoullii

1. PHILONOTIS LONGISETA (Rich.) E. G. Britt., Bryol. 14: 44. 1911.

Bartramia longiseta Rich, in Michx., Fl. Am. Bor. 2: 301. 1803.

Bartramia graminicola C. M., Linnaea 38: 632. 1874.

Autoicous; plants green, tufted, tomentose below; stems about

2 cm. high. Leaves 1-1.5 mm. long, lanceolate, acuminate; costa

excurrent; margins revolute, serrulate; cells linear, papillose at upper

ends. Seta about 2.5 cm. long; capsule nodding, 2 mm. long. (Fig.

89, A-C.)

Dept. Quezaltenango : Standley 85907. Dept. Zacapa: Steyermark 43187.

Distribution: Eastern and southeastern United States, Mexico.

On damp banks at moderate to rather high altitudes. There

seems to be no appreciable difference between P. graminicola (C. M.)

and P. longiseta. If anything the setae are a little longer in the

BARTRAM: MOSSES OF GUATEMALA

197

Guatemalan plants, nearly 3 cm. at times, but this is an inconstant

character.

2. PHILONOTIS GLAUCESCENS (Hornsch.) Par., Ind. Bryol. 923. 1894.

Barlramia glaucescens Hornsch., Fl. Bras. 1: 40. 1840.

Bartramia tenella C. M., Syn. 1: 481. 1849.

?Bartramia scobinifolia C. M., Bull. Herb. Boiss. 5: 188. 1897.

Dioicous; small plants, pale green; stems slender. Leaves

crowded, often subfalcate, less than 1 mm. long, lanceolate, acumi-

nate; costa percurrent; margins recurved; cells linear-oblong, papil-

lose at upper ends. Seta 1-1.5 cm. long, erect; capsule inclined,

ovoid, furrowed when dry. (Fig. 89, D-G.)

Dept. Izabal: Steyermark 41782; H. Johnson 1123. Dept. Alta Verapaz:

Steyermark MS 92, 45815, 45817; Standley 71748. Dept. Huehuetenango : Standley

82307; Steyermark 51169, 51170. Dept. San Marcos: Standley 66247a; Steyermark

35710. Dept. Quezaltenango : Standley 65481, 67856, 84818, 87051, 87216; Steyer-

mark 35163. Dept. Sacatepequez: Standley 59357, 59957, 62177, 63285. Dept.

Chimaltenango: Standley 62053, 79724, 80852. Dept. Jutiapa: Standley 75584.

Dept. Jalapa: Steyermark 32135, 32916, 32990. Dept. Santa Rosa: Standley 77781 .

Distribution: Southern United States, Mexico, West Indies,

Central and South America.

o

T

i

A \

FIGURE 89

A-C, Philonotis longiseta: A, plant, XI; B, leaf, X26; C, apex of leaf, X120.

D-G, Philonotis glaucescens: D, plant, XI; E and F, leaves, X24; G, apex of

leaf, X120.

H-I, Philonotis gracillima: H, leaves, X24; I, apex of leaf, X120.

198 FIELDIANA: BOTANY, VOLUME 25

On damp banks and rocks at low and medium altitudes. Fre-

quent and widely distributed. The small leaves with the costa

percurrent and margins recurved simplify the recognition of this

rather variable species.

3. PHILONOTIS GRACILLIMA Aongstr., Oefv. Sv. K. Vet.-Akad. Forh.

33: 17. 1876.

Dioicous; small, slender, green plants; stems to 1.5 cm. long.

Leaves oblong-ovate, bluntly pointed, to 1 mm. long; margins plane

or recurved, bluntly serrulate; costa ending below apex; cells oblong

to rhomboidal, weakly papillose at upper ends. Sporophyte as in

P. glaucescens. (Fig. 89, H-I.)

Dept. San Marcos: Steyermark 35729, 36884- Dept. Chimaltenango: Standley

57909, 6155k. Dept. Guatemala: Standley 62961. Dept. Zacapa: Steyermark

291+13(1.

Distribution: Southern United States, Mexico, West Indies,

Central and South America.

On damp banks at medium altitudes. This species intergrades

with P. glaucescens but may usually be separated, since some of

the leaves are obtusely rounded with the costa ending below the tip,

and the upper cells are broader and more pellucid.

4. PHILONOTIS SPHAERICARPA (Hedw.) Brid., Bryol. Univ. 2: 25.

1827.

Mnium sphaericarpum Hedw., Sp. Muse. 197. 1801.

?Bartramia chrysoblasta C. M., Bull. Herb. Boiss. 5: 188. 1897.

Dioicous; plants rarely over 2-3 cm. high, yellowish green, tomen-

tose below. Leaves erect, closely imbricated, oblong-lanceolate,

slenderly acuminate, 1.5-2 mm. long; margins revolute, sharply

serrulate; costa long excurrent in a slender, spinulose point; cells

linear, papillose at upper ends, oblong below. Seta 2-2.5 cm. long;

capsule cernuous, subglobose, brown, furrowed when dry, 2 mm.

long. (Fig. 90, A-C.)

Dept. Huehuetenango : Steyermark 50083. Dept. San Marcos: Steyermark

37313, 37315. Dept. Quezaltenango : Standley 81+827, 81+830; Steyermark 33621.

Dept. Solola: Steyermark 1+71+25, 1+71+87.

Distribution: Florida, Mexico, West Indies, Central and South

America.

On damp banks at medium to high altitudes. Distinguished

from P. glaucescens by the long excurrent costa and from P. uncinata

by the erect leaves.

BARTRAM: MOSSES OF GUATEMALA 199

5. PHILONOTIS BERNOULLII (C. M.) Par., Ind. Bryol. Suppl. 265.

1900.

Bartramia Bernoullii C. M., Bull. Herb. Boiss. 5: 187. 1897.

Dioicous? No antheridial buds seen. Slender, yellowish green

plants, laxly gregarious. Stems 6-8 mm. high, erect, densely tomen-

tose, with 4-6 slender, whorled innovations about 5 mm. long.

Leaves appressed, under 1 mm. long, narrowly triangular-lanceolate,

slenderly acuminate; margins narrowly recurved; costa long ex-

current in a denticulate, concolorous point; cells narrowly rec-

tangular, papillose at upper ends, wider and laxer toward base.

Seta slender, red, 12-14 mm. long, flexuous or slightly curved;

capsules subglobose, about 2 mm. in diameter, sulcate when dry;

peristome double, teeth acuminate, 200 yu high, segments of endos-

tome nearly as long as teeth; spores reniform, diameter 20-25 M-

(Fig. 88, D-G.)

Dept. Suchiate: Svihla 2872. Dept. Guatemala: Svihla 2800, 2801.

Endemic.

On damp soil at moderate altitudes. A considerable risk is

assumed in naming a species without authentic material for com-

parison but in this instance the plants agree so closely with the

original description that I am reasonably confident the name is

correctly applied.

The extremely slender stems clothed with minute, narrow leaves,

gradually tapering to a long, setaceous point formed by the excurrent

costa, and the filiform, flexuous or even arcuate setae are widely

different from any form of P. sphaericarpa (Hedw.) Brid.

6. PHILONOTIS UNCINATA (Schwaegr.) Brid., Bryol. Univ. 2: 22.

1827.

Bartramia uncinata Schwaegr., Suppl. I2: 60. 1816.

Dioicous; plants very similar to P. sphaericarpa but with the

upper leaves falcate and hooked at the tips of the stems. Setae

to 2.5 cm. or more long. (Fig. 90, D-E.)

Dept. Alta Verapaz: H. Johnson 982.

Distribution: Southern United States, Mexico, West Indies,

Central and South America.

On clay bank. This is the only collection I have seen from

Guatemala but it may prove to be more widely distributed locally.

200

FIELDIANA: BOTANY, VOLUME 25

B

A-C,

D-E,

F-H,

X270.

FIGURE 90

Philonotis sphaericarpa: A, plant, Xl; B, leaf, X28; C, apex of leaf, X120.

Philonotis uncinata: D, plant, Xl; E, leaf, X28.

Breutelia tomentosa: F, plant, Xl; G, leaf, X14; H, basal angle of leaf,

6. BREUTELIA Schimp., Coroll. 85. 1856.

Dioicous; male flower discoid. Usually robust plants with erect,

branched stems, densely tufted. Leaves lanceolate, acuminate,

plicate at base; margins serrulate; cells linear, papillose, well differ-

entiated at basal angles. Seta usually elongate; capsules cernuous,

furrowed; peristome double, endostome with well developed seg-

ments, cilia rudimentary; lid short, convex.

1. Leaf base erect and sheathing 2

Leaves spreading from insertion, base not sheathing 3

2. Stems slender, leaf base sulcate, often with a pocket on either side of costa

near shoulders 5. B. Brittoniae

Stems robust, leaf base plicate, without pockets 6. B. deflexifolia

3. Seta short, arcuate 2. B. subarcuata

Seta elongate, straight 4

4. Basal cells quadrate across width of leaf 3. B. jamaicensis

Quadrate alar cells few or none 5

5. Basal angles of leaf laxly areolate, decurrent 4. B. auriculata

Basal angles of leaf not as above 1. B. tomentosa

1.

1894.

BREUTELIA TOMENTOSA (Sw.) Schimp., in Ind. Bryol. 155.

, Bryum tomentosum Sw., Fl. Ind. Occ. 3: 1837. 1806.

Plants yellowish green, laxly tufted; stems to 10 cm. or more

long but usually shorter, variously branched, densely felted with

BARTRAM: MOSSES OF GUATEMALA 201

reddish brown tomentum below, branches in subfloral whorls on

fertile stems. Leaves spreading from insertion, occasionally sub-

secund, 3-4 mm. long, narrowly lanceolate from an ovate base,

slenderly acuminate; margins narrowly recurved below, distantly

serrulate above; costa slender, excurrent; cells narrowly linear, in-

crassate, papillose above, smooth toward base, colored across

insertion, very few at basal angles irregularly oblong, pellucid. Setae

1-2 cm. long; capsules nodding, ovoid, 3 mm. long. (Fig. 90, F-H.)

Dept. Alta Verapaz: Standley 7107^. Dept. San Marcos: Steyermark 35981,

36M9a, 36799a; Standley 86194, 86205, 86296, 86^00, 86^69. Dept. Totonicapan:

Standley 65919. Dept. Quezaltenango : Standley 67710. Dept. Baja Verapaz:

Standley 69908.

Distribution: Mexico, West Indies, Central and South America.

On wet banks at moderate to high altitudes. Variable but readily

recognized by the spreading leaves with only a few differentiated cells

at the extreme basal angles.

2. BREUTELIA SUBARCUATA (C. M.) Schp. in Besch., Prodr. Bryol.

Mex. 60. 1871.

Bartramia subarcuata C. M., Syn. 2: 617. 1851.

Plants yellowish green; stems to 8 or 10 cm. long, copiously

branched. Leaves crowded, spreading or often subfalcate, 4 mm.

long, lanceolate from a short, broadly ovate, erect, slightly clasping,

plicate base, margins recurved to or above mid-leaf, sharply serrulate

above; costa excurrent; cells linear, sharply papillose, 4-5 rows at

basal margins lax, oblong, pellucid, extending well up the basal

margins. Setae 4-6 mm. long, curved, reddish; capsules subglobose,

3-3.5 mm. long, not furrowed. (Fig. 91, A-C.)

Dept. Huehuetenango: Standley 83086b. Dept. San Marcos: Steyermark

35492 (c. fr.), 36099. Dept. Quezaltenango: Standley 67685a (as B. deflexifolia),

67701 (as B. deflexifolia), 677Ub (as B. deflexifolia), 677^9 (as B. deflexifolia),

67753 (as B. deflexifolia); Steyermark 3^163, 3^854. Dept. Chimaltenango:

Standley 61010 (as B. deflexifolia). Dept. Guatemala: Standley 80699. Dept.

Jutiapa: Steyermark 31922.

Distribution: Mexico, Colombia.

On forested banks, trees and rocks at high altitudes. When in

fruit the short, arcuate setae are distinctive. Sterile plants may be

distinguished from B. deflexifolia by the more branched stems and

the leaves often secund from a less strongly clasping base.

202

FIELDIANA: BOTANY, VOLUME 25

D

FIGURE 91

A-C, Breutelia subarcuata: A, plant, Xl; B, leaf, X14; C, basal angle of leaf,

X120.

D-F, Breutelia jamaicensis: D, plant, Xl; E, leaf, X14; F, basal angle of leaf,

X120.

3. BREUTELIA JAMAICENSIS (Mitt.) Jaeg., Adumb. 1 : 558. 1873-74.

Bartramia jamaicensis Mitt., Journ. Linn. Soc. 12: 265. 1869.

?Bartramia erythrocaulis C. M., Syn. 1: 473. 1849.

Philonotis Schlumbergeri Schimp. in Besch., Prodr. Bryol. Mex. 59. 1871.

Philonotis haitensis Ren. & Card., Bull. Soc. Roy. Bot. Belg. 29: 177. 1890.

Plants medium sized, yellowish or green, densely tufted; stems

to 3 or 4 cm. high, densely tomentose below. Leaves closely im-

bricated, appressed when dry, 2-3 mm. long, ovate-lanceolate;

margins recurved below, serrulate above; costa excurrent; upper

cells linear, papillose, gradually shorter and broader below, basal

cells subquadrate clear across the leaf. Seta about 1.5 cm. long,

red; capsule ovoid, nodding, strongly furrowed, 2.5 mm. long.

(Fig. 91, D-F.)

Dept. San Marcos: Standley 68496, 86417, 86468; Steyermark 86493. Dept.

Totonicapan: Standley 65924. Dept. Quezaltenango : Standley 66798, 67076,

83383, 84576, 84579, 84642, 84761, 86765. Dept. Suchitepequez: Steyermark

35320. Dept. Chimaltenango: Standley 61059. Dept. Zacapa: Steyermark 42204,

42207. Dept. Chiquimula: Steyermark 31123. Dept. Jalapa: Steyermark 32858.

Distribution: Mexico, Costa Rica, Jamaica.

On damp banks, rocks^and trees at medium to high altitudes.

A variable plant but readily known by the large area of subquadrate

cells extending across the leaf base.

BARTRAM: MOSSES OF GUATEMALA

203

4. BREUTELIA AURICULATA Bartr., Bryol. 49: 115. 1946.

Robust plants, green above, brown below; stems 10-12 cm. long,

densely reddish tomentose below. Leaves erect-spreading, some-

what flexuous when dry, 4 mm. long, oblong-lanceolate, gradually

slenderly acuminate, faintly plicate at base, with conspicuous, laxly

areolate, decurrent auricles; margins plane, minutely serrulate nearly

to base; costa percurrent; cells narrowly linear, sharply papillose at

upper ends, very lax, smooth, hyaline or brownish at extreme base,

laxly rectangular and hyaline in the decurrent auricles. Setae

12-14 mm. long, flexuous, reddish; capsules subglobose, inclined,

sulcate. (Fig. 92, A-C.)

Dept. San Marcos: Finca El Porvenir along Rio Chopal, south-facing slopes

of Volcan Tajumulco, alt. 1,300-1,500 m., Steyermark 37^62 TYPE.

Endemic.

Sharply distinct from any other species of the genus familiar to

me in the lax basal cells and the laxly areolate, hyaline, decurrent

auricles.

5. BREUTELIA BRITTONIAE Ren. & Card., Bull. Soc. Bot. Belg. 31:

161. 1893.

Stems slender, branched, to 10 cm. long, densely reddish tomen-

tose below. Leaves crowded, 3-4 mm. long, plicate, abruptly lanceo-

FIGURE 92

A-C, Breutelia auriculata: A, plant, Xl; B, leaf, X14; C, basal angle of leaf,

X120.

D-F, Breutelia Britloniae: D, part of plant, Xl; E, leaf, X14; F, basal angle

of leaf, X120.

204 FIELDIANA: BOTANY, VOLUME 25

late from a short, erect, sulcate, closely clasping, obovate base,

acuminate, points squarrose-spreading, often with a small pocket of

lax cells on each side of the costa near the top of the leaf base;

margins slightly recurved at leaf shoulders, otherwise plane, serrulate

above; costa excurrent; cells linear, papillose above, smooth at base,

several rows at basal margins more lax, oblong and pellucid. Setae

1.5 cm. long, erect; capsules nodding, ovoid, 3 mm. long, furrowed.

(Fig. 92, D-F.)

Dept. Huehuetenango : Steyermark 50072. Dept. San Marcos: Steyermark

35790, 35982. Dept. Totonicapan: Standley 62731, 6273 h. Dept. Chimaltenango:

Standley 61081>a.

Distribution: Costa Rica, Colombia.

On banks and trees at high altitudes. The characteristic little

pockets or cavities of lax cells are not always evident but occur at

least in some leaves of every plant examined. The slender habit

and short leaf base, broader at the shoulders than below will separate

it from B. deflexifolia.

6. BREUTELIA DEFLEXIFOLIA Card., Rev. Bryol. 37: 4. 1910.

Stems to 10 cm. long, sparingly branched, densely tomentose

below. Leaves crowded, deflexed or widely spreading, 5-6 mm. long,

lanceolate from an erect, ovate, lightly plicate base, slenderly acumi-

nate; margins recurved to about mid-leaf, serrulate above; cells

narrowly linear, incrassate, papillose 5-6 rows at basal margins

rectangular, lax and hyaline, forming a border nearly to the leaf

shoulders. Seta erect, 7 mm. long, red, stout, curved at tip; capsule

pendulous, subglobose, nearly smooth; lid conical (Fig. 93, A-C.)

Dept. Huehuetenango: Steyermark 501 73a (c. fr.). Dept. El Progresso:

Steyermark J^3094.

Distribution: Mexico.

Terrestrial at high altitudes. More robust than B. Brittoniae,

the leaves larger and the margins strongly recurved below.

21. ERPODIACEAE

Autoicous; small, delicate, soft plants, usually corticolous, grow-

ing in mats. Stems lax in structure, prostrate, branched, usually

flattened. Leaves crowded, broad, ecostate, unbordered; cells

rounded-hexagonal, smooth or papillose. Sporophyte at ends of

short lateral branches; seta short, erect; capsule erect, thin walled,

persistent; peristome lacking or simple; calyptra mi triform, plicate.

BARTRAM: MOSSES OF GUATEMALA 205

1. ERPODIUM (Brid.) C. M., Bot. Zeit. 1:774. 1843.

Anoectangium subg. Erpodium Brid., Bryol. Univ. 2: 167. 1827.

Plants with the characters of the family. Leaves imbricated,

concave; cells rounded, smooth or papillose. Perichaetial leaves

erect; seta short; capsules erect, exserted (in our species); annulus

broad; lid nearly flat; peristome lacking; calyptra mitriform, plicate,

lobed at base.

Leaf apex acuminate, cells smooth 2. E. Pringlei

Leaf apex rounded, cells papillose 1. E. domingense

1. ERPODIUM DOMINGENSE (Brid.) C. M., Bot. Zeit. 1: 774. 1843.

Anoectangium Erpodium domingense Brid., Bryol. Univ. 2: 167. 1827.

Plants yellowish green in close mats; stems flattened, to 1.5 mm.

wide with leaves, radiculose. Leaves closely imbricated when dry,

oblong-lingulate, to 1 mm. long, entire, rounded at apex; cells large,

papillose, rather obscure. Seta less than 0.5 mm. long; capsule

exserted, cylindrical, pale, 1 mm. long; lid plano-convex; calyptra

plicate, scabrous on the plaits, lobed at base, fugacious; spores

25-30 M. (Fig. 93, D-F.)

Distribution: Texas, Mexico, West Indies.

Usually on trees. This well known tropical American species

is recorded by Steere from the department of Peten (Lundell 2325)

but I have seen no collection from the local area.

2. ERPODIUM PRINGLEI E. G. Britt, Bull. Torr. Bot. Club 32: 266.

1905.

Plants tinged with brown; stems creeping, radiculose, closely

applied to the substratum, branches numerous, short, blunt, sub-

terete. Leaves closely imbricated, 1-1.3 mm. long, broadly ovate,

concave, short acuminate, entire; cells rounded-hexagonal, about

20 n, smooth, distinct, becoming wider than long below mid-leaf

and smaller at margins. Perichaetial leaves erect, clasping; capsule

partly exserted, urn 1 mm. long; spores 25-35 M- (Fig. 93, G-I.)

Dept. Santa Rosa: Standley 791>27.

Distribution: Mexico.

On exposed rock at low altitude. The collection is sterile and

the habit on rock unusual but otherwise the plants are indistinguish-

able from the corticolous specimens from Mexico.

206

FIELDIANA: BOTANY, VOLUME 25

22. ORTHOTRICHACEAE

Plants growing on rocks or trees in tufts or mats. Stems erect

or creeping with erect branches. Leaves crowded, hygroscopic,

lanceolate; costa strong, usually percurrent; upper cells rounded,

usually papillose, elongate below. Seta terminal; capsules immersed

or exserted, smooth or plicate; peristome usually present and double,

the teeth often united in pairs, segments of endostome narrow; lid

generally beaked ; calyptra mostly mitriform or campanulate, smooth

or plicate, usually pilose.

1. Stems erect ". 2

Stems creeping, often with erect branches 3

2. Calyptra small, cucullate 1. Zygodon

Calyptra large, campanulate 2. Orthotrichum

3. Leaves long decurrent, spinose-tuberculate at basal angles, basal cells short

3. Coleochaetium

Leaves not as above 4

4. Calyptra large, campanulate, not plicate, lobed at base 6. Schlotheimia

Calyptra mitriform, usually plicate, laciniate at base 5

5. Leaves bordered below with several rows of elongated cells, basal cells short

5. Micromitrium

Leaf base not bordered, basal cells usually elongate 4. Macromitrium

H

FIGURE 93

A-C, Breutelia deflexifolia: A, part of plant, Xl; B, leaf, X12; C, basal angle

of leaf, X120.

D-F, Erpodium domingense: D, plant, Xl; E, leaf, X54; F, upper leaf cells

and margin, X270.

G-T, Erpodium Pringlei: G, plant, Xl; H, leaf, X30; I, upper leaf cells and

margin, X270.

BARTRAM: MOSSES OF GUATEMALA 207

1. ZYGODON Hook. & Tayl., Muse. Brit. 70. 1818.

Slender, tufted plants; stems erect, dichotomously branched.

Leaves lanceolate, contorted when dry, entire or serrate above;

costa strong; upper cells small, rounded, incrassate. Seta terminal,

elongate; capsules suberect, 8 ribbed; peristome single, double or

none; calyptra cucullate, fugacious.

1. Leaf apex rounded 1 . Z. obtusifolius

Leaf apex acuminate 2

2. Leaves strongly squarrose-recurved when moist 3. Z. campylophyllus

Leaves erect-spreading when moist 2. Z. Reinwardtii

1. ZYGODON OBTUSIFOLIUS Hook., Muse. Exot., tab. 159. 1819.

Zygodon spathulaefolius Besch., Prodr. Bryol. Mex. 43. 1871.

Autoicous; plants small, in compact reddish brown tufts; stems

6-12 mm. high, branched, densely reddish tomentose below. Leaves

lingulate, broadly rounded, to 1 mm. long; margins papillose-

crenulate, recurved below; costa ending below apex, scabrous on

back; cells small, rounded-quadrate, incrassate, coarsely papillose.

Seta 4-5 mm. long; capsule erect or slightly inclined, cylindrical,

strongly ribbed, urn 1.25 mm. long; peristome double, teeth blunt,

in 8 pairs, segments 8, as long as teeth. (Fig. 94, A-C.)

Dept. Baja Verapaz: O. F. Cook & C. B. Doyle 255.

Distribution: Mexico, South America, New Zealand, Asia.

A highly individual species clearly distinguished by the broadly

rounded lingulate leaves with the costa ending well below the apex.

2. ZYGODON REINWARDTII (Hornsch.) Al. Br., Bry. Eur. 4, mon. 9.

1838.

Syrrhopodon Reinwardtii Hornsch., Nov. Act. Acad. Leop. 142: 700.

Zygodon circinatus Schimp. in Besch., Prodr. Bryol. Mex. 43. 1871.

Synoicous or heteroicous; tufts dense, yellowish green; stems

about 2 cm. high, tomentose below. Leaves crispate when dry,

to 1.5 mm. long, oblong-lanceolate, short acuminate, carinate,

decurrent; margins erect, coarsely and irregularly serrate near apex;

costa ending in or near apex; basal cells rectangular, upper cells

rounded, incrassate, papillose. Seta 1-2 cm. high; capsule nearly

cylindrical, small mouthed; peristome single, teeth 16, short and

often rudimentary; spores 20-25 M- (Fig. 94, D-F.)

Dept. Huehuetenango : Standley 81121,. in part. Dept. El Progresso: Steyer-

mark ltS119. Dept. Jalapa: Steyermark S2758a.

208

FIELDIANA: BOTANY, VOLUME 25

FIGURE 94

A-C, Zygodon obtusifolius: A, plant, Xl; B, leaf, X26; C, apex of leaf, X120.

D-F, Zygodon Reinwardtii: D, plant, Xl; E, leaf, X26; F, apex of leaf, X120.

G-J, Zygodon campylophyllus: G, moist plant, Xl; H, leaf, X16; I, upper

leaf cells and margin, X270; J, capsule, X8.

Distribution: Alaska, Mexico, Costa Rica, West Indies, South

America, Pacific Islands, India, Africa.

On trees and banks at medium to "high altitudes. The sharply

toothed apical leaf margins can usually be relied upon as a good

diagnostic character for the typical form but the var. subintegri-

folius is not without difficulties. Fruiting plants in good condition

are essential for accurate determination in many of the species.

var. SUBINTEGRIFOLIUS Malta, Monog. 122. 1926.

Leaves entire or nearly so.

Dept. San Marcos: Standley 85285. Dept. Totonicapan: Standley 8453 J^a.

Dept. Quezaltenango : Steyermark 3^.9Hb.

Distribution: South America.

These collections all lack good fruit. I had determined them as

Z. Liebmannii Schimp. on account of the subentire leaves, but Dr.

Grout, who has kindly examined them in the course of his studies

in this group, thinks that they might better be referred here.

3. ZYGODON CAMPYLOPHYLLUS C. M., Syn. 1 : 680. 1849.

Dioicous; stems slender, to 10 cm. high, branched, tomentose

below. Leaves erect and slightly contorted when dry, squarrose-

BARTRAM: MOSSES OF GUATEMALA 209

recurved when moist, 2 mm. long, lanceolate, decurrent, carinate;

margins often slightly reflexed above, sharply serrate toward apex;

costa ending below apex; upper cells small, rounded or angular, in-

crassate, papillose, rectangular and smooth below. Seta 5-6 mm.

long; capsule cylindric, 2.5 mm. long; peristome double, teeth broad

and blunt, in 8 pairs, segments 8, narrow; lid slenderly beaked,

curved. (Fig. 94, G-J.)

Dept. Huehuetenango: Standley 81 621 a, 81732, 81743, 81749, 81801, 83088

(c. fr.), 83089 (c. fr.); Steyermark 48374. Dept. Totonicapan: Standley 84521b.

Dept. Chimaltenango: Standley 58781 a.

Distribution: Mexico.

On trees and shaded limestone rocks in alpine regions. The

long, slender, much branched stems with the leaves squarrose-

recurved when moist will identify this individual species with little

difficulty. It will almost surely be mistaken for a Leptodontium at

first glance. A pertinent query is how this species differs from

Z. gracilis Wils.

2. ORTHOTRICHUM Hedw., Sp. Muse. 162. 1801.

Plants tufted, growing on trees or rocks; stems erect. Leaves

hygroscopic, imbricated, lanceolate, mostly entire; costa strong;

upper cells small, incrassate, papillose, rectangular below. Seta

terminal, short; capsules immersed or emergent, often 8 ribbed;

peristome usually double, teeth 16, often in pairs, segments 8 or 16,

narrow; calyptra campanulate, plicate, often pilose.

1. Stomata superficial 1. O. pycnophyllum

Stomata immersed 2

2. Capsules exserted, rupestrine plants 2. O. anomalum

Capsules immersed or emergent, corticolous plants 3

3. Upper and perichaetial leaves toothed 3. O. Bartramii

Leaves entire 4. O. malacophyllum

1. ORTHOTRICHUM PYCNOPHYLLUM Schimp. in C. M., Syn. 1: 709.

1849.

Orthotrichum recurvans Schimp. in C. M., Syn. 1: 709. 1849.

Orthotrichum Lozani Card., Rev. Bryol. 36: 107. 1909.

Autoicous; plants to 3 or 4 cm. high. Leaves ovate-lanceolate,

3-4 mm. long, acuminate; costa percurrent; margins recurved to

just below apex; upper cells rounded, incrassate, papillose, basal

210 FIELDIANA: BOTANY, VOLUME 25

cells linear, nodulose, shorter and broader toward margins. Seta

variable, to 2.5 mm. long; capsules immersed or exserted, nearly

smooth or lightly ribbed in upper half, ovoid-cylindric, sulcate when

dry and empty, stomata superficial, near middle of urn; peristome

teeth in 8 pairs, papillose, segments 16, about as long as teeth, 2 cells

wide, papillose; spores 16-20 /z. (Fig. 96, A-C.)

Dept. Totonicapan: Standley 62666a.

Distribution: Mexico.

On tree at high altitude. Very near 0. speciosum Nees. The

only noticeable difference is in the segments of the endostome which

in 0. pycnophyllum are supposed to number 16 but in some capsules

I find only 8.

2. ORTHOTRICHUM ANOMALUM Hedw., Sp. Muse. 162. 1801.

Autoicous; densely tufted, dark green rupestrine plants. Stems

1 cm. or more high, simple or branched. Leaves imbricated when dry,

strongly hygroscopic, to 3 mm. long, oblong-lanceolate, broadly

acute; margins revolute, entire; costa brown, ending just below apex;

upper cells irregularly rounded, incrassate, papillose, basal cells

rectangular, thin-walled, smooth. Seta 1.5 mm. long; capsules

exserted, ovoid-cylindric, tapering below, urn 2 mm. long, stomata

immersed; peristome double, teeth erect when dry, faintly striolate,

segments of endostome rudimentary and fragile; calyptra pilose.

(Fig. 95, A-C.)

Dept. Huehuetenango : Sharp 5000a.

Distribution: Northern United States and Canada south to

New Mexico.

On limestone boulder at high altitude. A significant collection

marking another long range extension to the south. These plants

are associated with Grimmia apocarpa var. gracilis just as they

might be in northeastern United States.

3. ORTHOTRICHUM BARTRAMII Williams, Bryol. 28: 76. 1925.

Mostly autoicous; small compactly tufted plants, yellowish

green at tips, brown below. Stems to 1 cm. high, often branched.

Leaves appressed when dry, widely spreading when moist, about

2 mm. long, oblong-ovate, pinched at apex to a short, blunt, toothed

point; margins recurved nearly to apex; costa ending below apex;

upper cells rounded, incrassate, papillose, basal cells rectangular near

BARTRAM: MOSSES OF GUATEMALA

211

I

D

\A

FIGURE 95

A-C, Orthotrichum anomalum: A, plant, Xl; B, capsule, X8; C, stoma of

capsule, X270.

D-G, Orthotrichum Bartramii: D, plant, Xl; E, leaf, X16; F, apex of leaf,

X68; G, sporophyte, XlO.

H-J, Orthotrichum malacophyllum var. guatemalense: H, plant, X 1 ; I, leaf, X 12 ;

J, part of peristome, X68.

costa, subquadrate toward margins. Seta scarcely 1 mm. long;

capsules emergent, ovoid, urn to 1.5 mm. long, ribbed, stomata

immersed; peristome double, teeth papillose, paired, segments of

endostome 8; calyptra pilose. (Fig. 95, D-G.)

Dept. Quezaltenango: Sharp 2317.

Distribution: Arizona.

On tree trunk at moderately high altitude. An interesting range

extension of this species hitherto known only from the type locality

in Arizona. The Guatemalan plants are not exactly typical but the

toothed upper leaves suggest this species rather than 0. tenellum

Bruch.

4. ORTHOTRICHUM MALACOPHYLLUM Card. var. GUATEMALENSE

Bartr., Bryol. 50: 207. 1947.

Autoicous; small dull yellowish plants tinged with brown, densely

tufted. Stems to 1 cm. high, branched. Leaves contorted when

dry, spreading when moist, to 3 mm. long, narrowly oblong-lanceolate,

bluntly acute, carinate, decurrent; margins entire, recurved below;

costa ending below apex; upper cells rounded, incrassate, minutely

212 FIELDIANA: BOTANY, VOLUME 25

papillose, basal cells rectangular with firm, pellucid, sinuose lateral

walls. Seta very short; capsules immersed, oblong, urn 2 mm. long,

stomata immersed; peristome teeth paired, minutely papillose,

segments of endostome 8, vertically striolate; calyptra pilose. (Fig.

95, H-J,)

Dept. El Quiche: Sharp 2^32, 2^5, 2^63. Dept. Baja Verapaz: Sharp 2926 a.

Endemic.

On trees and shrubs at moderate altitudes. These collections

are an exact counterpart of the Mexican species except that the

segments of the endostome are uniformly vertically striolate instead

of papillose.

3. COLEOCHAETIUM (Besch.) Ren. & Card.,

Bull. Soc. Bot. Belg. 332: 120. 1894.

Orthotrichum subg. Coleochaetium Besch.,

Fl. Bryol. Reun. 1879: 66. 1879.

Medium sized brownish green plants in lax, intricate tufts; stems

creeping, branched. Leaves crowded, fragile, decurrent; costa

strong; cells uniform, rounded, small, papillose and pellucid, rec-

tangular and spinose in the decurrent portion. Seta short, erect;

capsules exserted, 8 ribbed; peristome double; calyptra campanulate,

smooth, pilose (sporophyte not seen).

1. COLEOCHAETIUM STANDLEYI Bartr., Bryol. 47: 21. 1944.

Probably dioicous. Plants wiry, laxly caespitose, dull sordid

green, brown below. Primary stems creeping, radiculose, irregularly

branched, branches up to 5 cm. long, laxly and irregularly rebranched,

branches obtuse or often attenuate and radiculose at the tips,

flexuous when moist, variously curved and contorted when dry.

Leaves crowded, 5 ranked, appressed when dry, squarrose-spreading

when moist, about 2 mm. long, 1 mm. wide, shortly ligujate-lanceo-

late from an ovate base, carinate above with the points fragile and

usually broken off, acute, apiculate, strongly decurrent; margins

narrowly recurved below, sharply and finely papillose-serrate above;

costa brownish, strong, ending below apex; leaf cells obscure, densely

papillose with sharp, salient papillae, rounded, about 10 n in diameter,

in the decurrent angles large, rectangular and pellucid, strongly

armed with high, spine-like tubercles up to 20 ^ long. Fruit unknown.

(Fig. 96, D-G.)

BARTRAM: MOSSES OF GUATEMALA

213

Dept. Huehuetenango : Rio Pucal, about 14 km. south of Huehuetenango,

alt. about 1,780 m., Standley 82293.

Endemic.

In this highly individual species the decurrent leaf auricles com-

posed of large, rectangular, pellucid cells strongly armed with spinose

tubercles is a striking character. The species is evidently near

C. scaberrimum (Broth.) Broth, of Brazil but the plants are wiry,

laxly branched and strongly contorted when dry whereas the

Brazilian plants are described as rigid, densely branched with strict

branchlets. No specimen of C. scaberrimum is available for com-

parison but it seems evident that the Guatemalan plant is distinct.

It is a privilege to associate Dr. Standley's name with this unique

* addition to the Central American moss flora.

4. MACROMITRIUM Brid., Muse. Rec. Suppl. 4: 132. 1819.

Plants slender to robust, in dense mats; stems elongate, creeping,

branches numerous, erect, densely foliate. Leaves lanceolate or

oblong; costa strong; upper cells small, smooth or papillose, basal

cells, usually elongated. Seta smooth or scabrous; capsules exserted,

erect, ovoid, smooth or ribbed; peristome single, double or lacking;

calyptra large, mitriform, naked or pilose, deeply laciniate below;

lid mostly long beaked.

FIGURE 96

A-C, Orlhotrichum pycnophyllum: A, plant, Xl; B, leaf, X8; C, capsule, X8.

D-G, Coleochaetium Standleyi: D, part of moist plant, Xl; E, leaf, X14;

F, basal angle of leaf, X120; G, upper leaf cells and margin, X270.

214 FIELDIANA: BOTANY, VOLUME 25

Grout's timely and important studies in this group (Grout 18)

have been followed closely in the treatment of the local species.

1. Stems slender, leaves rigid and appressed when dry, basal cells short 2

Stems more robust, leaves spreading, basal cells elongate 3

2. Calyptra pilose, peristome a rudimentary membrane. . . .1. M. hymenostomum

Calyptra naked or nearly so, peristome of 16 short teeth 2. M. filiforme

3. Capsule mouth small, puckered or plicate 4

Capsule mouth wide, not puckered 5

4. Leaves lanceolate, acuminate, cells incrassate 3. M. stratosum

Leaves Ungulate, short pointed, cells thin- walled ., 4. M. Richardi

5. Capsules ribbed 6

Capsules smooth 12

6. Seta scabrous 7

Seta smooth 8

7. Calyptra pilose, peristome teeth long, subulate-acuminate. .7. M. homalacron

Calyptra naked, peristome teeth short, truncate 6. M. longifolium

8. Leaves lingulate, obtuse, mucronate '. 5. M. altituberculosum

Leaves lanceolate, acute 9

9. Leaves entire 11. M. Podocarpi

Leaves toothed above 10

10. Leaf cells highly convex or mammillose 9. M. serrulatum

Leaf cells smooth or slightly convex 11

11. Seta 8-10 mm. long, leaves squarrose-recurved when moist. 8. M. punctatum

Seta 4-5 mm. long, leaves erect-spreading 10. M. guatemalense

12. Calyptra pilose, leaves spirally ranked 12. M. pentastichum

Calyptra naked, leaves not seriate 13

13. Leaves ending in a long, fragile point, mostly broken off at base of acumen