¡

t
I
240 MOSSES [MEM. NEw YORK BOT. GARD. vo

of
Mosses a1c
Steven P. Churchill and Noris Salazar Allen Ar
:

68:
Illustrations :

M. Gloria Mora-Gonzales and Alberto Castillo

Mosses (Musci) :

Plants i= gametophyte) foliose, growing from a single tetrahedral (although a bifacial cell can be found in t
few forms, e.g., Ftssidens) apical cell. Stem prostrate or erect, outer surface, with or without a cuticle, in few
forms, with paraphyllia (Cyrtohypnum, Thuidiurz), pseudoparaphyltia present (many pleurocarps) or absent, I

in cross section with @olytrichales) or without (most mosses) internal conducting tissues, hyalodermis present
or absent. Leaves entire, never dissected, spirally arranged on the stem (sometimes in two rows, distichous), t\
with or without a definite arrangement on the stem, mostly isophyllous but some taxa with anisophylly (one or
two rows of leaves different in size and arrangement on the stem), leaves mostly unistratose (all cells chloro- :
:

phyllous or witlr chlorophyllous and hyaline cells in different areas of the leaf or multistratose (with various
layers of cellÉ, all chlorophyllous or with a central chlorophyllous layer srirrounded by 1 (rarely more) layers of
hyaline cells), with or without midrib; midrib one or two (rarely more), sometimes the number varying within
the same gametophyte. Leaf ceils chlorophyllous, with discoid chloroplasts or hyaline (devoid of protoplast
and usually porate), smooth or papillose; celi walls thin or thick, smooth or pitted, without trigones. Rhizoids
multicellular, branched, hyaline or colored, cell walls thin or thick, smooth or papillose. A¡theridia on the main
shoot, on short shoots or on lateral branches, oval to + sphericai, pedicellate, surrounded by paraphyses,
perigonial bracts differentiated or not from vegetative ieaves; archegonia apical (acrocarpous mosses) or on
lateral branches (pleurocarpous mosses), vessel shaped with a venter and a narrow neck, surrounded by para-
physes, perichaetial leaves differentiated or not from vegetative leaves. Sporophye mostly exerled or inmersed,
capsule spherical to oval or elliptical, with urn and operculum or without an operculum; operculum apiculate
to conic or long rostrate united to the urn by a ring of cells, the annulus; urn multistratose, with or without
stomata, outer cells smooth, papillose, mamillose or spinose, neck differentiated or not, columella present,
its distal part in some taxa (Polytrichales) expanded into a membrane that closes the mouth of the urn (the
epiphragm); mouth of the urn surrounded by one (single) or two (double) rows of teeth, the peristome or
peristome absent. Capsule dehiscent by vertical slits, an opercuium or indehiscent (cleistocarpous). Calyptra
mitrate or cuculate, smooth or pleated, naked or hairy. Seta short or long, yellowish-green to green when
I
young, straw-yeilow to red-brown when mature, smooth, papillose, sometimes spiculose to spinose or with
multicellular filaments (Calyptrochaeta), with or without a central strand. Protonema long persistent, fila-
mentous and ramified or less frequent thallose, usually producing more than one foliose gametophyte.
Worldwide, the mosses (Musci) are estimated to include 12,800 species (a more realistic figure would be
8000 species), in 900 genera. The diversity in the Neotropics is estimated at 76 families, 402 genera and
2600 (2300-2950) species. For those desiring a taxonomic sequence of families, the following list is pro-
vided. This reflects some of the current ideas about the general patterns of evolutionary history with regards
to mosses. It must be noted, however, that this is not necessarily a phylogenetic sequence. The monophyly of
most families (or higher levels) has not been demonskated; rather, it is a traditional-based classification i
fotrnded on a priori assumptions resulting in polythetic groups. Currentiy there is a healthy debate with regard t
to the definition of family limits and relationships within and between families. For that reason, the families
of the mosses are treated here in a singie alphabetical sequence and are not subdivided into orders as in the l

treatrnent of the Hepaticae. Since this manuscript was completed (ca. 1997) a substantial number of phyloge- :

netic studies have appeared (at least in abstracts), both morphologic,al and molecular, very few combining both;
one can anticipate major changes in our understanding of relationships and resulting classifications at all leveis.

Lrenerunr. Buck, W. R. & D. H. Vitt. 1986. Suggestions for a new familial classiñcation of pleurocarpous mosses.
Taxon 35: 21-60. De Luna, E., A. Withey,D. Gonzalez & B. D. Mishler. 1999. The transition to'pleurocarpy: A
-
phylogenetic analysis of the main diplolepidous lineages based on rácl sequences and morphology. The Bryologist
102: 634450.
- Edwards, S. R. 1984. Homologies
New Manual of Bryologlt 2: 658495.
and inter-relationships of moss peristome. In: R. M. Schuster (ed.),
Mishler, B. D. & S. P Chu¡chill. 1984. A cladistic approach to the phylogeny
-
) vol-. 86,20011 KEYS TO SUBCLASSES & FAMILIES 241

of tlre "bryophytes." Brittonia 36: 406-424. Mishier, B. D. et al. 1994. Phylogenetic relationships of the "green
-
algae" aad "bryophytes". Annals of the Missouri Botanical Garden B 1: 451-483. Newton, A. E. & E. De Luna. 1999.
-
A survey of morphological characters for phylogenetic study of the transition to pleurocarpy. The Bryologist 102: 65 1-
682. D. H. 1984. Classification of the Bryopsida. In: R. M. Schuster{ed.), New Manual of Bryology2:697-759.
-Vift, I

Class Sphagnopsida Family Orthotrichaceae Family Leskeaceae

Order Sphagnales Family Rhachitheciaceae Family Myriniaceae
Farnily Sphagnaceae Order Leucodontales Family Plagiotheciaceae
Family Anomodontaceae Family Pterig¡mandmceae
Class Andreaeopsida Family Catagoniaceae Family Regmatodontaceae
Order Andreaeales Family Cryphaeaceae Family fugodiaceae
Family And¡eaeaceae Family Fo¡tinalaceae Family Sematophyllaceae
Family Leptodontaceae Family Stereophyllaceae
Class Polytrichopsida Family Lepyrodontaceae Family Syraphyodootaceae
Order Polytrichales Family Leucodontaceae Family Thamnobryaceae
Family Polytrichaceae Family Meteoriaceae Family Thuidiaceae
Family Neckeraceae Order Fissidentales
Class Bryopsida Family Phyllogoniaceae Family Fissidentaceae
(Subclass Bryidae) Family Prionodontaceae Order Bryoxiphiales
Order Funariales Family Pterobryaceae Farnily Bryoxiphiaceae
Family Ephemeraceae Family Racopilaceae O¡der Dicranales
Family Funariaceae Family Rhacocarpaceae Family Archidiaceae
Family GigaspeÍnaceae Family Sorapillaceae Family Bruchiaceae
Family Splachnobryaceae Family Trachypodaceae Family Dicranaceae
) Order §plachnales Order Hookeriales Family Ditrichaceae
Family Sptachnaceae Family Adelotheciaceae Famiiy Eustichiaceae
Order Bryales Family Daltoniaeeae Family Leucobryaceae
Family Aulacomniaceae Family Hookeriaceae Family Seligeriaceae
Family Bartramiaceae Family Leucomiaceae Order Pottiales
Family Bryaceae Family Pilotrichaceae Family Calymperaceae
Family Meesiaceae Order Hypnales Family Leucophanaceae
Family Mniaceae Family Ambiystegiaceae Family Pottiaceae
Family Phyllodrepaniaceae Family Brachytheciaceae Order Buxbaumiales
Family Rhizogoniaceae Family Entodontaceae Family Diphysciaceae
Order Orthotr.ichales Family Fabroniaceae Order Encalyptales
Family Erpodiaceae Family Hydropo goaaceae Family Encalyptaceae
Family Hedwigiaceae Family Hylocomiáceae Order Grirnniales
Family Helicophyllaceae Family Hypnaceae Family Grimmiaceae
Family Macromitriaceae Family llypopterygiaceae

Keys to the sutrclasses and families of mosses of tropical America

The lollowing keys are i¡ltended to assist in the identification of mossss Íiom the Neotropics. The keys
are designed to stress gametophytic features whenéver possible. Sporophytes are often not present in col-
lected material, and, in some cases, not known in the Neotropics or elsewhere. For this reason, and because
a number of families exhibit considerable variation, families are often keyed out in more than one place. The
keys often emphasize salient features of families with primary emphasis on gametophytic characters, sec-
ondariiy on sporophytic characters, and lastly on habitat or geography. In using the keys, one normally
follows the iead that b'est fits the two altemative choices given. When one reaches a point in the keys in
which neither lead appears appropriate, one then should follow both, hopefully ending with two aiternative
farnilies. Both should then be compared carefully with the descriptions, keys to genera, and illuskations. If
neither altemative appears appropriate, then one must backtrack in the keys to find where the problem of
intrrpreting a particular character exists. In any case, one should read carefuliy the family description, keep-
ing in mind the variation exhibited by that family (this applies equaily to genera).
 I
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242 MOSSES IMEM. NEW YORK BOT. GARD. i

i'
{
Key to the subclasses 'i
' ,

Plants typical of boggy ormarshy sites, whitish; branches spirally arranged in fascicles; laminal cells
of stem and branch leaves altemating between leucocyst (hyaline cells) and chlorocyst (green) cells; :

capsulesglobose,supportedbyapseudopodium....'......... i
1. Piants mostly of drier sites, mostly.green, yellow or brown; laminal cells uniformly similar with chloro- i,
phyll, or chlorophyll cells layered between hyaline cells above and below; not altemating between hyaline !
and green ceils; capsules variously shaped, supported by a pseudopodium or seta 2 I
2. Plants small, dark red or blackish, largely restricted to open high elevations, in páramo and puna;
l

k
capsules supported by a pseudopodium, opening by 4 (division tbroughout) or 8 (distal tip) slits or
i.
i
2. Plants small to medium sized or robust, present in all environments; capsules stegocarpic or cieistocarpic, I
supported by a seta, generally with a peristome and operculum I
j

3. Leaves bearing rows of lamellae cin distal upper surface; capsules with a single series of peristome teeth, !
l
teeth 16, 32, or 64, distally attached to a circular membrane (epiphragm)........... Polytrichopsida (Polytrichaceae) 'i,,
3. Leaves lacking rows of lamellae, orvery rarelypresent (see Grimmiaceae, Pottiaceae); capsules either .t
with a peristome in a single series with 16 teeth, or a double series with the outer series of 16 teeth
(exostome), and inner series with a basal membra:re bearing 16 segments and cilia (endostome), in a t
number of cases the peristome is reduced ....... BryQsida I
i
-:i.
General key to the families
Plants whitish, laminal cells alternating or iayered between leucocyst and chlorocyst cel1s ,.................-... Section 1
Plants variou§ly green, yellow to golden, or broWn to blackish; laminal cells uniform, neither layered or
altemating between leucocyst and chlorocyst ceil walls...................-.........................--......i,........................................2 j
t'
2. Upper distal surface of leaf with discontinuous or continuous rows of lamellae Section 2 Y
2. Upper distal surface iacking lamellae !

3. Plants acrocarpous; stems erect or occasionally spreading, solitary or in short to long tufts or cushions: i
sporoph¡es terminai on stems or branched innovations; peristome single (a single series of 16 teeth or !
.':
divided into 32 n¿urow segments) or double (two series, outer series of 16 teeth, inne¡ series often with i.
16 segments and often cilia atop a short to tall membrane)
3..'Plants pleurocarporis, stems creeping, spreading, or pendent, frondose or dendroid from conspicuous or
I
I
inconspicuous creeping primary stems; sporophytes iateral on stems; peristome double or variously reduced ............ 6 I
i¡
4. Leaves arranged in 2 ranlís (distichous) or 4 ranks (but appearing 2-ranked) Section 3 I
1 I
a.
5. Laminal ceils papillose o¡ mammillose Section 4
t
¡.
5. .i.
6. Leafcosta absent, short and forked; or elongate and double '1 'l i

ult
6. o t,
t\
Leaves distinctly 2-ranked, usually strongly folded Section 6 t
7. Leaves in 3 or more raoks, when complanate often with lateral asymmetric leaves (occasionally partially t
i
folded on one side) and median symmetric leaves 8
.t

8. Leaves ecostate (most leaves lacking a costa, a few leaves may have a very weak costa) Section 7 i

8. Leaves with costa, short¡nd forked or elongate and double Section I

9. Leaves skongly dimorphic, upper or lower leaves on stems smaller and differing in shape than larger I
,)
laterai leaves Section 9 I

9, LeavesÁondmorphic, usuallydiffering if at all between smaller branch leaves, or lateral leaves asJ¡m- .. i
f.
metric and median leaves slmmetric 10 i
t.
I 0. Laminal cells mammillose oi papillose ...................,. §ection 10
j
10. Laminal cells smooth Section 11
-f 1.

Section 1. Plants whitish, occasionally reddish or purplish tinged; laminal cells differentiated, I

alternating or layered between hyaline cells (leucocysts) and green cells (chlorocysts).
. Ir

,I
1 Plants §pically of boggy or seepy sites; forming a compact head of clustered short branches (capitulum),
below capitulum, stems with short to elongate fasciculate branches; leaves of stem and branch drmorphic; .t'
in cross section lamina unistratose, cells alternating between chlorocysts and leucocysts §phagnaceae l1

1. Plants of moist to semidry sites, often lignicolous or epiphytic; stem and branch leaves monomorphic,
1

mostly evenly distributed along stem§; in cross section lamira bi- to multistratose, chlorocysts layered
tretween I or more layers of leucocysts above and below...... ........_....... 2
t
f
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voi-. 86, 20011 KEYS TO SUBCLASSES & FAMILIES

2. Leaves fai*ly singly costate, chlorocyst cells clustered near center on abaxial side (back) of leaf ...... Leucophanaceae
2. Leaves appearing ecostate; chiorocyst cells evenly distributed in a single row befween leucocyst cells,
not forming a cluster of cells toward the center............. . Leucobrvaceae

Section 2. Upper distal surface of leaves with rows of lamellae or short filaments
1 . Lamellaé in long continuous rows, occasionaliy rows few or discontinuous; peristome teeth 16, 32, or
64, e¡tt¡e, usually with tips ofteeth attached to an epiphragrn Polytrichaceae
i. Lamellae forming short ¡ows or occurring as shoí fiiaments scattered over surface; peristome of 16 teeth,

2. Distal upper surface covered by filaments, often papillose, if wi& lamellae then leaves ovate with margins

2. Distal upper surface covered by rows of lamellae, with Ieaf rnargins erect to incurved; calypka campan-

PLANTS ACROCARPOUS
Plants acrocarpous, stems mostly erect or occasionaliy spreading, solitary or in short to tall loose or dense
tufts; sporophytes terminai on stems or branched innovations;peristome single or double, rarfi absent.

Section 3. Plants acrocarpous. Leaves arranged in 2 ranks (distichous)

or in 4 ranks but appearing 2-ranked
1 . Lea,f costa long excurrent, 2-3 times longer than lamina length; lower and basaL cells linear-re ctangular;
confined to páramo and puna Ditrichaceae p"p.
1. Leaf costa subpercurrent to short excuüent (rarely iong excurrent), iower a¡rd basal cells quadrate to
rectaugular; lowiand to hjgh montane, rarely in 2
2. Leaves exhibiting an extended dorsal and ventral lamina a sheath-iike leaf base (vaginant lami.nae) ............ 3
2. Leaves lacking an extended dorsai and vbntral lamina &om a sheath-like base.... 4
3. Piants soiitary or formiog iufts, erect to suberect; marginal border cells of leaf linear or oblong-lineaE

3. Plants forming mats, procumbent (pleruocarpous); marginal border cells very iarge, subrectangulal thin-
rvalled; capsules ir¡mersed ..
4. Leaves skongiy asymmétric; ;"; ;i;;;; ;; ;;; ;;" ;;,",*";; r"**"" ,"**'"r" ;*;;;;;;;,::fP*iaceae
stem tips; wet lowiands .:............. Phyllodrepaniaceae
4. Leaves + symmetric or weakly asymmetric; costa longitudinally centered on leaf; gerrrmae if present irr
leaf axils; nostly montane or subalpine 5
5 Leaves folded, conduplicate 6
5 Leaves flat, not iolded; lamiral cells smooth '7

6 Laminal cells pluripapiliose; leaves short oblong-ovate, less than I mm iong ........ Eustichiaceae
6 Laminal cells smooth; ieaves oblong-lanceolate, mostly 2-3 mm long........ Bryoxiphiaceae
,7
Stems spreading or creeping; leaves elliptical, bordered Mniaceae p.p
7 Stems erect; leaves ovate, lacking border........"... .. Rhizogoniaceae p-p.

Section 4. Plants acrocarpous. Leaves in 3 or more rows. Laminal cells

papillose or mammillose {cells often isodiametric or shortiy elongate)
Basal portion ofieafexhibiting a cancellinae (enlarged clear cells), strolgly differentiated &om basal
margin and distal cells; marginal or intramarginal border often presenl; laminal cells above isodiametric,
gemmae nearly always present on leaves, particularly leaf tips; plants mostly epíphytic Calymperaceae
Basal portion of leaf lacking a cancellinae (see Kingiobryutn etc. in páramo), cells either similar to distal
cells or gradually differentiated toward base, if differentiated then leaf border lacking; gemmae absent
frcm leaves 2
2. Leaves ecostate; mostly on rocks 3
2. Leaves costate; on various subskates including rocks 4
l- Leaf margins plaae; laminal cells unipapillose on back; plants small, dark red or blackish, in paramo and
puna; capsules opening by 4 (división throughout) or 8 (distal tip) slits or valves Andreaeaceae p.p.
3. Leaf margins recurved; laminal ceils I to several papillose, papillae in a row; plants medium sized dark
green to brown or graytsh; mid-montane to páramo or puria; capsules operculate, Iacking a peristome .... Hedwigiaceae
4. Leaves compietely to partially bistratose distatly ........... 5
4. Leaves unistratose or only partially biskatose, particularly restricted to margins B
 j.
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244 MOSSES IMEM. NEW YORK BOT. GARD. i vot
.i
5. Peristome of 16 teeth in 8 pairs; leaves spathulate, apex acute to apiculate Rhachitheciaceae p.p. 5.
5. Peristome of 16 unpaired teeth; leaves mostly ovate-lanceolate to lanceolate, if spathulate then costa
1

6.
i
6. Laminal cells subquadrate to short rectangular, appearing bipapi11ose..... Ditrichaceae p. p.
i
i 6.
6. Laminal cells rounded, mammillose 7 l

7. Leaf margins uni-, bi-, or tristratose; perichaetial leaf costa long excurrent with distal lamina erose or '7.
I
fimbriate; sporophytes immersed; seta short; capsules strongly asymmetric, obliquely and broadly ovoid ¡

below, narrow toward mouth; peristome double; plants usually found on soil or sandstone ............. Diphysciaceae tt 7.
7. Leaf margins bistratose, sometimes incomplete below; perichaetial leaf costae subpercurrent; sporophytes i
i
exserted; seta elongate; capsules s¡rrnmetric; peristome single; ptants exclusively found on rocks .... Grimmiaceae p.p. ¿. 8.
8. Laminal cells stellate, unipa¡iillose, papillae over cell lumen; basal cells inflated; leaf margins recurved 'i
{.
nearly throughout; distal stems naked with a few highly reduced leaves; plants ofmarshes and bogs at
high elevations Aulacomniaceae
I
, !
:
8. Laminal cells not stellate, uni- to pluripapillose; basal cells not conspicuously inflated; lacking special- 9.
ized distal branches or stems (sometimes with deciduous leaves); plants mostly of semi-wet or dry sites ..,...:,..... 9 9.
9. Laminal cells narrowly to somewhat broadly rectangular with projecting papillae at cell ends (rarely
.í :

10. l
I
single over cell lumen); leaf margins often coarsely toothed, teeth single or more often double; capsules 10.1
i
subglobose'inclinedoriferectthenimmersed....,................, i t
9. Laminal cells mostly isodiametric or sliortiectangular; papillae mostly over cell lumina or cells mam- t
.¡" 11. (

millose; leaf margins entire, crenulate or a few teeth at apex; capsules ovoid to cylindrical ,;........ l0 I
:7
10. Laminal cells often bipapillose; capsules immersed to shortly exserted; peristome double or reduced; (
t
calyptra campanuláte, plicate or not, hairy or naked; epiphytic or rarely on rocks
.¿
Orthotrichaceae p.p. 11. (

10. Laminal cells uni orpluripapillose, ormammillose; capsules mostly long, rarely short exserted; calyptra (
cucullate or if campanulate then long cylindricai, smooth; mostly terrestrial, occasionally epiphytic 11 : 12. t
;
I 1. Leaves bordered by few to several rows of hyaline cells, border extending from base to 1/3 ieaf length, I 12. t
or to near apex; laminal cells uni-, bi- or pluripapillose; alar cells well-differentiated Dicranaceae p. p. 13. t
I 1. Leaves iacking a border; cells variously omamented; alar cell undifferentiated 12 (
i
12. Leaves differentiated, base clasping, iimb spreading; plants of wet or moist sites ........................... 13 73. t
¡
12. Leaves undifferentiate{ or if so, then base not clasping stem or if clasping upper laminal cells pluripapillose .......... 14
13. Laminal cells subquadrate, mammillose; costa percu¡rent ................. Ditrichaceae p.p.
jt' 14. I
)
13. Laminal celis short-rectangular, rounded, papillae projecting at cell ends; costa short excurent.... Dicranaceae p.p. I

14. Leafcosta in cross section little differentiated, well-developed stereid band lacking 15 14. I
ir
14. Leaf costa in cross section usually well-developed, stereid band present, only below guide cells or both I .(
h 15. I
15. Laminal cells mammillose or weakly unipapillose, cells crowded, little space between celi lumens.... Dicranaceae p.p. l.1.
ia
_f
1 5 . Laminal cells piuripapillose, cells widely spaced, or if crowded, papillae extending between celi lumens .... .. .... I 6 §

16. Leaves spathulate or ifnarrowly oblong-lanceolate then hyaline basal cells extending upward along :
15. I
,
margin a sbort distance; peristome single, 16 teeth in 8 pairs Rhachitheciaceae p.p. ,,!, s

16. Leaves narrowly lanceolate to oblong- or ovate-lanceolate, basal cells uniform across; peristome double 16. I
or variously reduced, i6 teeth, separated or in 8 pairs Orthotrichaeeae p.p. i
t
1 I
17. Calyptrae long cylindrical-campanulate; leaves oblong-elliptical to -obovate, laminai cells coarsely pluri- t
16.
papillose, papillae usually branched; restricted to very high elevations (zacatonal, páramo or puna).... Encalyptaceae t
"-f(
17. Callptra cucullate; leaves of various shapes; laminal cells uni- to pluripapillose, simple or branched; 7
U

Section 5. Plants acrocarpous. Leaves in 3 or more rows, ralked or not. Laminal óells smooth 7 (
1. Plants minute, ephemeral or not; leaves costate or ecostate, capsuies cleistocarpous o¡ gymnostomous, É

mostly immersed 2 (
1. Plants conspicuous, mostly medium sized to iarge, not ephemeral; leaves costate; capsules stegocarpous 8. (
or rarely gymnostomous, mostly long exserted 't' c

2. Laminal cell walls firm, mostly rectanguiar to subquadrate 3 8. (

2. Laminal cell walls lax, short- to long-rhomboidal or hexagonal-rhomboidal 6 h

3. Leaves oblong to obovate or orbicular, margins entire; annulus well-developed Rhachitheciaceae p.p. 9 .t
3. Leaves mostly lanceolate, if obovate then distal leaf margins usually crenulate or weakly semrlate; annulus t e

.t
absent or well-developed 4 9
4. Spores very large, few Archidiaceae v
4- 20 ,l
5. Capsules with a¡ elongate neck, often haif or more the length of capsule, stomata oÍten numerous in l'
20. L

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t. vol-. 86, 20011 KEYS TO SUBCLASSES & FAMILIES 245

5. Capsules with an inconspicuous neck or neck lacking, stomata few or absent ........ Dit¡ichaceae p.p.
6. Stems fleshy, partially subterranean; capsules wrinkled or warfy, if smooth then exserted and leaves
6
l. 6. Stems Íiom a persistent protoÍema, not subterranean; capsules smooth or weakly bulging-mammillose,
7 immersed §phen[praceae
7. Lamirial celis ¡arrowly rectangular and strongly sinuose throughout except at basal margin; plants found

:e '7
.Larrn:rrral cells not sinuose throughout, walls entire, or if sinuose not strongly so and reetricted in distri-
bution, not throughout; plants foünd on various substrates includi*g rock§.............. .......................... 8
l.
,¡
8. Leaf margins bordered, border extending to apex or ending somewhat belów, 4djacent inner laminal
cells large, often hexagonal-elongate üo broadly fusiform. or rhomboidal 9
i

i 8. Leaf margias lackiag a border, or if borde¡ed then confined to leaf base; marginal cells simila¡to or
only slightiy differentiated from inner laminal cells ..,........... .............. 12
9, Laminal cells su\uadrate aod rouoded to oblong-oval, distaliy obliquely arranged Mniaceae p.p.
9 9. Laminal eells rhombic to elongate-hexagonal, not obliguely aranged 10
10. Leaf border consisting of long linear cells; capsules often inclined to penduious ...,..... Bryaceae p.p. .t,'

10. Leaftrorder consisting oflarge. rectangular to fusiform cells; capsules erect, ifincli¡ed or subpendulous
e then seta hygroscopic 1l
1i. Capsules with a corspicuously módihed, enlarged neck, often i¡flated or larger ihan urn; peristome ?
0 1, siirgle, 16 teeth united in pairs, or B. teeth, oÍ}en recurved when dry; plants usually epiphyi'c, on dung
"jI or decaying plant material Splachnaceae p.p
). ii
I 1- Capsules rvith an inconspicuous or narrowed neck; peristome absent, or single and represented by an
¡
I exostorle of 16 teeth, or doub,e, teeth incwved when wet; plants mostly on soil ..........- Funaria¿eae
I
I 1-2. Alar cells differentiated, quadmte to obiong or oval, often dark eolored, yellow to red..........
n. klar
).
;l 13. Alar cells dark red; low, somewhat irregular membrane formed a¡ound and external to peristome; plants
2 i on rocks associated with streaias, mostly at high elevations §eligeriaeeae p.p
¿ 13. Alar cells often reddish-brown or golden; 1ow, external rnembrane lacking; plants on various substrates
1
4 {: and at low to high elevations D. icranaceae p.p.

). rl, 1 4. Leaves appearing 3-raaked; seta conspicuously elongate (to 10 cm long or rnore); exostome reduce'd,

). l endostome well-deveioped; plaats of mid to liigir eievations in boggy sites Meesiaceae

5 i
14. Leaves spirally arranged, or rnore than 3-ranked; seta usually shorter; peristome various; plants either
l
7 I5. Laminai cells distally linear-obiong or -fusiform and flexuose or not; capsdes inclined to more often
).
f pendulous; peristome double, endostonrial eiiia often present, or represented by only a hyaline endo-
6
I
i
i5. Laminal cells isgdiametric to short or long rectangular;capsules mostly erect to inclined; peristome
t. .1, single or double §-hizogoniaceae) i6
I i6. Laminal eells above base isodiametric, smail, thick-walled; plants often dark green, reddish-brorvn, or
t.
i walls thin- to
16. Laminal cells often short to long rectangular, or distally grading into subquadrate cells,
e
I .'ratherthick-walled;p1antmostly1ighttodarkgreen
I
i7. Costa in cross section generally well-developed, stereids above asd below guide cells or only below,
|. I upper and lower epidermal cells differentiated or not; capsules often long exserted; peristome te€th
i
't'
I
often divided into 32 filaments; plants on'soil o¡ rocks, dry or wet sites, at ali elevations ...........".... Pottiaceae p.p.
i7. Costa in crcss sectian little or not differentiated; capsules short exserted or immersed, 4-valved or with
I peristome teeth perforate or divided only distally; piants confined to rocks a*d at very high elevations
z ,i
18. Capsules 4-valved (peristome and operculum lacking); leaves strongly subulate from an ovate base or
t
7 I ovate with a very broad, cliffuse costa fo ca. i12lamina length Andreaeaceae p. p.
l i
I
18. Capsules with peristome, teeth often perforate or divided above; leaves percurrent to excurrent as a
6 i
.t
t. ¡_
19..Leaves elliptical, obovate spathulate, obtuse to rounded; peristome either with I 6 teeth in 8 pairs or
tl' exostorie extremely reduced
u 20
I I i9. Leaves mostiy ovate-lanceolate to narrowiy lanceolate; margins entire to serate; peristome with teeth,
;
when present not paired nor with exostome extremely reduced 22
5 I
20.Leaves mostly 2 mm long or more: capsules with a modihed, enlarged neck, often inflated or largerl
I

20. Leaves mostly 1.2 mm long or less; capsules lacking a modified neck ¿t
I
I
l
j A
 246 MOSSES IMEM. NEW YORK BOT. GARD. voL

21. Leaves obovate to ellipticai, leaf apices not apiculate; perichaetial leaves similar to stem leaves, not long B I
sheathing; capsules smooth Splachnobryaceae (
21. Leaves obovate to spathulate, leafapices apiculate (most leaves); perichaetial ieaves strongly differenti- 8 I
ated fiom stem leaves, long sheathing; capsules 8-ribbed Rhachitheciaceae p.p. (
22. Capsule neck elongate, i.5-3 times longer than urn, stomata numerous in neck region; peristome absent 9
or single (vertically striate below); costa subpercurrent; plants of disturbed sites, rather uncoÍrmon..Bruchiaceae (
22. Capsule neck distinctly shoiter than um, or absent; peristome double or single (and often divided above); ó
costa subpercurrent to excurrent; plants ofvarioús habitats 23 (
23. Perichaetia at mid-stem or base; leafmargins bistratose and doubly toothed, ifsingly toolhed then distal 0 I
stem tips with cylindrical, papillose geÍrmae; plants epiphytic or terrestrial Rhizogoniaceae p.p. f
23. Perichaetia terminal on stems; lacking the above combinations of characters; plants on soil or rocks ..................24 -10 I
24. Capsules ribbed; annulus compound, exceeding or nearly so the length of the peristome; leaves setace- l1 I
ous from an ovate base; plants rare................ Seligeriaceae p.p. 11 I
24. Plants lacking the above combination of characteristics 25 i
25. Peristome teeth terete, often divided to near base, into 32 filaments Ditrichaceae p.p: t2.I
25. Peristome teeth flat below, entire or divided above Dicranaceae p.p. 12.1

PLANTS PLEUROCARPOUS >

Plants pleurocalpous; stems creeping, spreading, forming mats, or pendent, frondose or dendroid,
often forming fufts, from a creeping primary stem; sporophy'tes lateral on stems, rarely terminal
I
on short branches; peristome double or variously reduced, rarely absent.
t
u

Section 6. Plants pleurocarpous. Leaves 2-ranked, strongly folded throughout or with sheath-like base
2. t
1. Leaves exhibiting an extended dorsal and ventral lamina from a sheath-like leaf base (vagiaant laminae);
2. t
.). L
margins bordered, cells very large, subrectangular, thin-walied; capsules immersed...:-,...................,.SorapiIIaceae
3. L
-1. Leaves strongly folded, conduplicate, lacking a dorsal and ventral lamina; marginal border lacking;
4. L
capsuies shortly to fuliy exserted 2
o
2. Leavestoi.5mmlong; apexmostlymucronateandreflexed, laminalcelisenti¡e,notporose........Catagoniaceae
2..Leaves3mmormorelong; apicestruncateorerectapiculate;laminal cellsporose......................Phyllogoniaceae
4. L
5. L
Section 7. Plants pleurocarpous. Leaves ecostate ir
1. Plants aquatic, attached to rocks in streams or epiph¡ic irrinundated lowlands; capsules immersed .....................2 5. L
1. Plants ter¡estrial, on soil, rocks, and logs, or epiphytic, if aquatic then alar cells inflated; capsules most- 6. A
ly exserted 3 6. A
2. Leaves mostly distant, crowded only at stem or branch tips, oblanceolate to obovate or elliptical; plants 7. L
confined to lowlands I{ydropogonadeae 7. L
2. Leaves mostly crowded, lanceolate; plants confined to montane habitats or páramo Fontinalaceae 8. D
3. Leaves broadly ovate, acute to piliferous, narrowly bordered, border differentiated in.color, often dark
red; laminal cells appearing finely pluripapillose, papillae minute and numerous over and between cells 8. ;
Rhacocarpaeeae 1É 9. L
3. Leaves variously shaped, acuminate, acute, or.obtuse roun{ed; laminal ceiis smooth or if papillose, cr
papillae single or few over cell lumina 4 { 9. L
4. AIar cells inflated, oval to oblong, mostly thick-walled po-rose or no! often dark yellow or golden-red; s1

laminal cells smooth or papillose, pápillae over cell lumina ........... Sematophyllaceae p. p.§ 10 L
4, Alar cells absent or ifpresent, then not inflated or oval, mostly subquadratg or oblate and quadrate- 10 L
11 A
c(
5- Leaf apex broadly acute or if acuminate then laminal cells rhomboidal to fusiform-rounded and thick-walled ............ 6
6. Leaves short ovate-lanceoiate, to 0.7 mm long; laminal cells on back projecting at distal angies; alar A
cells equally distributed on either side of leaf base, mostly quadrate; plants forming mats....... ... Myriniaceae p.p. r
6. Leaves variously shaped, ovate-lanceolate, ovate to oblong, mostly greitér than i mm long; laminal cells
. smooth, or occasionally on back projecting at dist¿l angles; alar cells lacking or equally or unequally 12. st
).
qa
distributed on either side of leaf base, quadrate to oblong; plants forming mats, dendroid or pendent
7. Stems complanate foliate, lateral leaves slightly asymmetric or not............. ............... 8 eÍ

- "l . Stems terete foliate .......,.... l0 12. Pl

 ñ

r.D.
*
: vol-. 86,20011 KNYS TO SUBCLASSES & FAMILIES a11

Ii' 8. Leaf margins finely biñd toothed; gemmae ciustered beneath sterns oron specialized terminal branches;
ae epiphytic or epiphyllous Pilotrichaceae p.p
i:
8. Leaf margins entire to subentire; gemmae absent or if present, on leaf tips; epiphytic on base of trunks
I
.p. oftrees, logs, or soil 9
t 9. Alar cells well-diflerentiated, unequally distributed with cells on one side more numerou§; upper laminal
ae cells linear; gemmae absent; moist to wet lowlands Stereophyllacese p.p.

23
i 9. Alar cells undifferentiated; upper laminal celis large, elongate-hexagonal; gemmae few on leaf tips and
distal margins; montane lorests
10. Ireaf margias piaae distally; laminal cells obliquely arranged above, kansversely toward base,or not;
Hookeriaceae

.p. median and upper cells rhomboidai to ñrsiiodn-rouaded, porose or not ..í............. ....................,..... 11
24 I - 10. Leaf margins strongiy incurved distally; laminal cells longitudinaLly ananged, linear, mostly porose................. 12
... Leucodotrtsce#F,
t 11. Leaves broadly lanceoiate and plicate, or ovate and smooth, short acuminate; capsules exserted

.p. 11. Leaves broadly ovate to suborbiculate; smooth, broadly, acute, or if acuminate then tip hyaline; capsules
25 l' immersed Erpodiaeeae
p.
I 12. Plants dend¡oid, or subpendent and rather stiff; gemmae commoniy pr€sent in leaf ¿xi1s ....... Pteryobryaceae p.p. "
.Y. f
'i'
11
Plants pendent or forming léose, spreading mats; gemmae absent Meteoriaceae p.p.

§ection 8. Plants pleurocarpous. Leaves costate, costae short and forked or rrr
double and elongate (often 112 or mare of lamina length)
Leaves strongly cordate-auriculate; paraphyllia present, several branched ......... Hylocomiaceae p.p.
Leaves not cordate-auriculate, ifauriculate only weakly so; paraphyllia absent or ifpresent then iineal
unb¡anched -1_

a Leaf costa double and e loi-igate, o{lre:rr 112 or rrore ihe lamila iength . ........ Pilotrichaceae p.p.
2. Leaf costa mostly short and forked, mostly less tban ll2 the lamina lergth ........ .............3
3. Leaves compianate 4
ae
3. Leaves teretely foliate, falcate-secund or not ............. 10
4. Leaf margins bordered by fe.*, to severai rows of narrow celis, irnei celis large, rhomboidal to hexag-
z
onal; costa often indistinct or distally forked ñ'om a short single costa; seta papillose and ciliate distally
te
..... Daltoniaceae p.p.
le 4. Leaf margins lacking a border, occasionally ceils of lamina progressively narrowed toward margin.................,.,.. 5
5. Leaves ovate-oblong to oblong-lingulate, often unduiate; apices mostly broadiy acute to obtuse; capsules
immersed to shortly exserted Neckeraceae p.p.
2 5. Leaves mostly ovate to ovate-lanceoiate, smooth; apices mostiy acute to long acurni-nate; capsules exseried ............ 6
6. Alar ceils numerous, extending along leaf margirt to 115-113 lergth .. 7
3 6. Alar cells feri, to several, rareiy extending more tlaa a few cells along margin; capsules erect to horizontal ............. 8
7. Lami¡al cells snióoth; capsules erect ................ Entodontaceae p.p.
¡e 7. Laminal cells papillose at cell angles; capsules inclined Pterigyrandraceae
¡e E. Dislal brasehes §agellare and readiiy deci<iuous; seta distaily roughened to papiilose; urn spinose
§ymphodontaceae
8. Distal br¿nches not flagellate, if so not readily deciduous; seta srnooth; capsules lacking ornament¿tioa ."..........., 9
¡e it 9. Leaves weakly serrulate to subentire distaliy, short to long decurrent; capsules suberect to horizontal,
Plagiotheciaceae
A
9. Leaves ofteo sem:iate to serrate throughout, not or weakly decurrent; capsules ovoid and horizontal to
subpendent or cylindrical and erect Hypnaceae p.p.
p.? 10. Leaves faicate-secund or branches and stems distally curved or cu¡led .......,.... lt
10. Leaves mostly erect to spreading, not falcate-secund, branches and stems not curved .......... t2
5 1 1. Aiar cells quadrate to short rectangular, moderately thick-wal1ed, concolorous or yellow to golden, or

¡e.i * ceils inflated a-nd hyaline, thi¡-walled; exothecial celi walls of urn mostly evenly thickened; operculum
6 conic-apiculate to conic-short rostrate Hyp*aceae p.p.
i 1. Alar cells inflated, thick-walled, often golden-red, supra-alar cells often present, subquadrate; costa
p. .e weakly shart and forked or absent; exothecial ceil walls often collenchyrnatous; opercuium long rostrate
Sematophyllaeeae p.p
12. Stems either dark red with leaves concave and ovate with alar celis subquadrate a¡d thick-walled (often
7 dark red), or sterns wiry with leaves concave and ovate, weakly plicate, costa separated at base; alar cells
I enlarged and thin-walled Hylocomiaceae p.p.
0 i2. Plants lacking the combination ofcharacteristics given above 13
248 MOSSES [MEM. NEWYORK BOT. GARD. vol

13. Stems spreading to subascending, evenly foliate; leaves mostly crowded, ovate-short lanceolate to ovate t2.
(and concave); capsules long exserted; confined to montane sites or high elevations ............ Amblystegiaceae p.p.
13. Stems often pendent or lax and elongate; leaves often distart, often conspicuously so, except at branch
13.
and stem tips; leaves obovate to oblanceolate or elliptical, not concave or weakly so; capsules immersed;-
13.
confined to lowlands Hydropogonaceae p.p.
14.

Section 9. Plants pleurocarpous. Leaves costate; costa single. Leaves strongly dimorphiq, upper 14.
or lower leaves on stem smaller and differing in shape from larger láteral leaves 15.
l. Leaves on upper side of stem smaller; leaf margins lacking a border; costa of lateral leaves stoutly short

1. Leaves on lower side ofstem smaller; leafmargins bordered or at least differentiated; costa oflateral
leaves ending well below apex or weakly short excurrent 2
2. Plants procumbent; laminal cells unipapillose; apex ofupper larger leaves obtuse-rounded to broadly
1
acute; margins partially recurved; sporoph¡es immersed; semi-dry sites Ilelicophyllaceae 1
2. Plants stipate-frondose; laminal cells smooth; apex ofupper larger leaves rather sharply acute to short
2
acuminate; margins plane; sporophytes exserted; moist montane sites Hypopterygiaceae

2.
[Sections 10-l l. Plants pleurocarpous. Leaves monomorphic, usually differing only in that branch leaves
are somewhat smaller, differing in shape or not; or lateral leaves asymmetric and median leaves
symmetric, or primary and secondary stem leaves differentiated; costa single.]
-)-

Section 10. Plants pleurocarpous. Leaves costate; costa single; iaminal cells mammillose or papillose
3.
1. Laminal cells pluripapillose (cells with a single papillae mixed with some celli occasionally 2-3-papillose keyed
out here also) 2
4.
1. Laminal cells uniformly unipapillose or mammillose 7
2. Papillae rather numerous! over cell lumen and side walls J
4.

3. Leaves smooth, not plicate, erect-appressed or wide-spreading; apices either acute to short acuminate or
5.
narrowly long acuminate Meteoriaceae p.p
6.
3. Leaves plicate, loosely e¡ect to spreading; apices broadly long acuminate Trachypodaceae p.p.
6.
4. Paraphyllia present on stems; leaves ofstem and branch differentiated, the former ovate or triangular
and acuminate, the latter ovate and acute or obtuse-rounded; stems regularly l-3 pinnately branched; -
leaf apices ending in one or more papillae Thuidiaceae p.p
4. Paraphyllia absent; leaves of secondary stems and branches diflering mostly in size; stems irregularly 6.

branched; leaf apiceg not papillose ---.................... 5

5. Upper laminal ceils isodiamet¡ic, mostly obscurely hexagonal Anomodontaceae p.p. i..
5. Upper laminal celis elongate; linear to oblong-linear 6
7
(
6. Secondary stems rather rigidiy erect.- solitary^ or few branched; leaf margins skongly recr¡rved .... Pterabryaceae p.p.
6. Secondary stem soft or rigid, not erect, several to numerousty branched; leafmargins not recurved, or if
(
so only weakl¡ at base Meteoriaceae p.p. 8
(
7. Stems differentiated between creeping primary stems and erect, subpendent or frondose/dendroid second- 8

(
7 - Stems not differentiated between prinury and secondary stems, plants mostly forming mats or dease tuftq .......... 13 9
B. Creeping stems densely tom€ntose, leaves mostly obscurely hidden among tomentum; elongate branches (

(appearing as secondary stems) erect and often nunerous; seta elongate, on terminal branches; calyptra
mitrate-campanulate, often deeply iobed and plicate Macromitriaceae 9. ,l
8. Creeping stems tadiculose beneath, or naked; secondary stems erect to suberect, pendent o¡ frondose to ¿

10. I
9. Upper laminal cells with papillae projecting at distal angles 10 t
9. Upper laminal cells with papillae over cell lumen 10. I
10. Plants not stipate, mostly irregularly branched; gemmae absent; seta very short, capsules immersed 11. I
Cryphaeaceae p.p. (
10. Plants mostly stipate, often frondose; gemmae often present, seta elongate, capsules exserted ..... lleryobryaceae p.p. 11. I
I 1. Leaf margins forming a border of somewhat elongate, smooth cells ............... .. Trachypodaceae p.p. (
11. Leaf margins lacking a border, cells similar to intralaminai cells ............. ................. i2 t2. I
12. Plants small, frondose; leaves ovate, smooth; upper laminal cells rhombic, walls entire...,.. Thamnobryaceae p.p. 12. f
 I

AR'. i vol.. 86,20011 KEYS TO SUBCL,{SSES & FhMILIES 245

I

i 12. Plants mediur¡l sized to large, irregularly branched to subdendroid; leaves ovate-lanceolate. plicate;
) p.p I upper laminal cells rhomboidal to linear-rhomboidal, stellate .....,.... Prio¡rodontaceae
I
13. Stem leaf margins strongly ciliate; laminal cell papillae strongly curved Anomodontaceae p.p.
I 13. Stem leaf margins serrulate to serrate, not ciliate; laminal cell papillae erect or projecting .....,...... 14
p.p 14. Plants forming tufts; leaves falcate-secund; some cells with papil14e strongly projecting at distal angies
I

t
per I 14. Plants forming mats; leaves not falcate-secund; papillae ovrer cel1 lumen or weakly projecting 15
15. Leaves loosely complanate, lateral leaves somewhat as¡rmmetric, oblong acute-rounded; alar cells un-
I equally disfibuted, rnore Dumerous on one side of costa .......... Stereophyllaceae p.p.
.ceae 15. Leaves not complanate, ovate acuminata; alzr cells equally distributed on either side of costa ..... Leskeaceae p. p,
I
t
i
,....2 I Section I 1. Plants pleurocarpous. Leaves costate; costa single; laminal celis smooth
I 1 Laminal cells elongate, mostly 8 or more times longer than wide 2*ñ
i
ceSe 1 Laminal cells isodiamekic to short oval or rhomboidal, mostly 5 or less times longer than rvide ....,..........,........... 1
I

I
2 Secondary stems erect, arising &om densely tomentose short creeping primary stems; leaves oblong- .

ceae lanceolate, to 5 mm long, ending in a long capillary tip, finely plicate; laminal cells lineai and porose;
I
i
epiph¡ic in high montane to zacatonal, páramo, and puna .............. I-epyrodo*taceae
2 Stems and branches creeping or subascending, not densely tomentose below; leaves variously shaped, .*
rves
I mostly ovate- to oblong-lanceolate, apex acute to long acuminate, not endiag in a capillary tip, if piicate
I
I
I
then ratler coarsely so; laminal cells weakly porose or not; various substrates and elevations ......... 3
3 Plants pendent, derdroid, or frondose, from a creeping primary stem (occasionally forming loose mats
on banks or logs); stem and branch leaves often dimorphic or polymorphic in repeated series ............................... 4
lose
3 Plants spreading or creeping, stems not differentiated b€tween primary and secondary; stem and branch
yed
leaves differiag only by degree of size and often by width
....2
.... "l
4. Flants mostlyerecf and ffondose or dendroid, if pendent then leaves in 5-spiral rorvs; nume¡ous ñlamen-
tous pseudoparaphyllia present Pteryobryaceae p.p.
..-- I
.... 4
4. Plants pendent or occasionally forming ioose spreading mats; leaves rot an-anged ia 5-spiral rows; fila-
mentous pseudoparaphyllia absent ... Meteoriaceae p.p,
5. Leaf alar cells asl,rnmetrically distributed, more numerous on one side of costa Stereophyllaceae p.p.
p-p.
5. Lezf a\ar cells, if present, equallydistributedoneithe¡side of costa ..:........-.....:..-............................ 6.
6. Plants rnostiy of wet habitats; leaves smooth; alar cells usually present, either thin-walled, enlarged and
rounded or if quadrate to short rectangular then cells small and few; capsules horizontal.s*ongly asym-
p.p. mekic and curved Arnblystegiaceas p.p.
6. Piants of moist or semidry. habitats, if aquatic then leaves broadly ovate, leaf margins semrlate through-
5
out, upper laminal cells twice as long as wide; leaves plicate or smooth; alar cells quadrate to short rect-
p-p. angulal often rather numerol§; capsules incli¡ed to erect. weakly asymmetric Brachltheciaceae p.p.
7 Stems differentiated between creeping primary stems and erect to suberect secondary stems
---..\
.,.6 8

p.p. 7 Stems creepiag or spreading, occasion*lly subascending, ¡rot diff,erenti¿ted bef,veen priuary and sccor¡d-
arv stems 15
p.p. 8. Secondary stems distinctly stipate, subdendroid to &ondose; atlenuate flageliate branches often present ..... .... . . . . . 9
8. Secondary stems not stipitate or weakly so, rigidly erect; tips short aftenuate or not, flagellate branches
... 8 absent, microphyllous branches present or not .....t......... 11
13 9. Secondary stem and branch leaves loosely to strongly complanate; secondary stem leaves ovate to more
commonly oblong or oblong-ligulate, apex mostly acute to acute-¡ounded, often coarsely toothed at apex
Thamnobryaceae p.p
ceae 9. Secondary stem and branch leaves mostly leretely foiiate; leaves ovate to ovate-short or ovate-long
acuminate...... 10
.... 9 i0. Seconda¡v stem leaves abruptly long acuminate from a broad ovate base; margins serrulate to near
.. l0 base .-........ Rigodiaceae
l1 10. Secondary stem leaves gradually short acuminate; margins entire to sem¡late at apex............ Leptodontaeeae p.p.
11. Leaves bordered with narrowly linear cells and conspicuously differentiated from inner iaminal cells;
p.p. calyptra mirate or campanulate and fringed with ciliate hairs.... ...".. Daltoniscese p.p.
p.p. 11. Leaves lacking a border, margioal cells not differing greatly from inner laminal cells; calyptra mitrate or
p.p. cucullate, base lacking hairs 12
.12 12. Secondary stem leaves loosely to strongly complanate 13
p.p- i2. Secondary stem leaves terete foliate 14.

j
 250 MOSSES IMEM. NEW YORK BOT. GARD. VO

13. Leaves broadly ligulate, apex truncate; costa ca. 3l4leaf length; attenuate branches uncoÍrmon; gemmae

13. Leaves broadly e1liptical, apex acute; costa percurrent; stem and branch tips short attenuate with gemmae
in axil ofreduced leaves Adelotheciaceae
14. Microphyllous branches present; median laminal cells fusiform to fusiform-rhomboidal; leaf margins
plane; seta elongate, capsules exserted Leptodontaceae p.p,
14. Microphyllous branches absent; median laminal ceils oval to oblong-oval; leaf margins recurved or plane;
seta very short, capsules immersed....... Cryphaeaceae p.p
15. Leaf costa ca. ll2 leaf length or less; laminal cells thin-walled; plants rather delicate, stems readily
breaking when removing leaves l6
15. Leaf costa mostly 213 leaf length or longer; laminal cells rather thick-walled; stems resilient 17
16. Leaf margins denticulate, ciliate or entire; peristome present or absent; plants common ............ Fabroniaceae p.p.
16. Leaf margins semrlate above aiar regiotr; peristome absent; plants rare My,riniaceae p.p.
17. Upper laminal cells subquadrate; costa strongly flexuose distally Anomodontaceae p.p.
17. Upper laminal cells rhombic to short rhomboidal or oval; costa straight or weakly flexuose 18 N
.!
18. Branches ascending; leaf margins entire; costa often spurred below; exostome teeth ca. 712 the length of
the endostome Regmatodontaceae
18. Branches spreading or creeping; leaf margins dentate or semrlate; costa entire, not spurred; exostome
subequal to or larger than endostome 19
19. Leafapex relativeiy broad, acute; seta rqughened or scabrous; capsule ovoid, to i.1 mm long ..... Myriniaceae p.p.
19. Leaf apex relatively abruptly short to somewhat long acuminate; seta smooth; capsules cylindrical, mostly

ADELOTHECiACEAE
A monotypic family, a member of the Hookeriales.

Adelothecium (Fig. 81) A monotypic genus, with
-
A. bogotense (Hampe) Mitt. rather widespread in the mon-
tane regions of the Neotropics (Mexico, Central America,
Greater Antilles, tropical Andes, Guayana Highlands and
southeastern Brazil) and East Africa.
HABITAT. Epiphytic, commonly on branches or tninks
of shrubs and treelets; primary or slightly disturbed sec-
ondary montane forests, at elevations ñom 114G-3300 m.
DESCRIPTIoN. Plants somewhat large, forming tufts,
olive green to goiden-brown. Primáry stems short, creep-
ing, radiculose beneath. Secondary stems andbranches erect-
spreading, often perpendicular to sub§trate. Stem leaves
complanate, 8-ranked, lateral leaves somewhat asyrnmet-
ric and undulate, obovate, 3-4 mm long, apex rounded and
apiculate, base asymmetric, slightly decurrent; margins
plane, crenulate-dentate; costa single, strong, subpercurrent
to percurrent; apical cells linear-fusiform, porose; median
cells stellate, smooth to bulging; basal cells fusiform to
irregularly rectangular and porose, golden brown; marginal
cells narrow and smaller; dorsal and ventral leaves simi-
lar, symmetrical, oval to obovate. Branch leaves smalier,
gradually attenuated distally, leaves reduced in size. Gem-
mae present at distal branch tips in axil ofhighly reduced
leaves, gemmae short-cylindrical to clavate.on axillary
stalks. Dioicous. Sporophytes lateral; perichaetial leaves
elongate, ovate-lanceolate or -subulate. Seta orect, short,
ca. 2 mm long, smooth to weakly papillose. Capsule erect,
urn ovoid. Operculum rostrate. Peristome double, exostome
teeth 16, cross-striate, appearing furrowed with a hyaline
bolder; endostome basal membrane low, segments 16,
keeled and perforate, cilia absent. Calyptra mitrate, base
weakly lobed, densely covered by long capillary hairs, pli-
cate. Spores spherical, papillose-granulose.
DISCUSSIoN. The compianate leaves that are obovate, single
costate with stellate laminal cells, combined with the gernmiferous
branch and stem tips, should separate this t¿xon from other mon-
tane epiph¡es. Sporophytes are exceedingly rard, and the prin-
ciple rnode ofrep¡oduction is asexual via gemmae. The ¡elation-
ship ofthis taxon within the Hookeriales is currently debated by
several workers. The genus was previously placed in the
Hookeriaceae; see comrnents under the Pilotrichaceae.
LITERATURE. Ochyra, R., H. Bednarek-Ochyra, T. Pócs & M.
I
R. Crosby. 1992. The moss Adelothecium bogotense in continen-
F.U
tal Africa, with a review of its wo¡ld range. The Bryologist 95:
mar[
28'7-295.
mart

AMBLYSTEGIACEAE oblc
lonE
Plants small to somewhat large, forming loose to dense mats or ascending tufts. Stems creeping, decum- Lint
bent, or ascending, irregular to subpinnately branched; central strand usually present; paraphyllia only present tusi.
irt Cratoneuron; pseudoparaphyllia filamentous or foliose; rhizoids sparse. Leaves imbricate to somewhat fTate
,distant, often subjulaceous or faicate-secund, appressed to spreading, short to long ovate-lanceolate, ovate or dioi,