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funaria moss

Habit Plants minute to medium-sized, gregarious to forming open tufts, light- to yellow-green, annual to biennial. Plants small to medium-sized, gregarious or tufted, bright green to yellowish green.
Stems

short, erect, simple or with a few branches, central strand present, basal rhizoids few.

short, erect, simple except for a short basal antheridial branch.

Leaves

usually larger and more crowded distally, often comose, reduced proximally, usually contorted when dry, spreading when wet, broadly elliptic to obovate, usually concave, margins plane to somewhat incurved, entire to serrate, sometimes limbate, apex acute to acuminate, rarely somewhat blunt, costa single, percurrent to excurrent;

distal and median cells usually irregular-rhombic to hexagonal or rectangular, smooth and rather thin-walled, often lax, weakly chlorophyllose, proximal cells usually longer, oblong to rectangular, sometimes weakly inflated at proximal angles, differentiated alar cells absent.

larger and erect distally, reduced proximally, oblong-ovate to broadly obovate distally;

concave;

apex usually acute or acuminate margins erect, entire to serrate beyond middle;

costa single, ending before the tip to excurrent;

distal and medial laminal cells large, rhombic-hexagonal to rectangular, lax and rather thin-walled, proximal cells oblong-rectangular, differentiated alar cells absent.

Seta

terminal, solitary, short to elongate, erect to somewhat curved, smooth or rarely papillose.

elongate, erect to strongly curved or twisted.

Sexual condition

autoicous, sometimes polygamous, rarely synoicous or paroicous.

autoicous;

antheridial branches 1–2, basal, perigonial paraphyses clavate with an enlarged inflated cell;

perichaetia apparently absent paraphyses.

Capsule

stegocarpous or cleistocarpous, immersed to exserted, globose or pyriform to cupulate, sometimes flaring, symmetric and nearly smooth to asymmetric and striate when dry, usually with a neck;

exothecial cells thick- to thin-walled;

stomata restricted to neck, consisting of a slit in a rounded guard cell, superficial or immersed, annulus present or absent, revoluble, revoluble in fragments, or not;

operculum present or absent, flat, conic-rounded, to rostrate;

peristome double, single, rudimentary, or absent, exostome teeth 16, erect to incurved, papillose-striolate or striate, trabeculate on adaxial surface, endostome segments 16 and opposite the exostome teeth, cilia absent, represented only by the exostome when single.

exserted, usually inclined to pendent, asymmetric and usually curved, yellow to brown, pyriform, often sulcate or plicate when dry and empty, annulus large and revoluble or not differentiated, exothecial cells oblong-hexagonal to linear, walls incrassate especially so on inner tangential wall, stomata immersed;

peristome double, inserted somewhat below the mouth, teeth well developed, obliquely directed, lance-acuminate, papillose-striate, often strongly trabeculate, frequently appendiculate at the tips and fusing with a latticed disk, endostome segments opposite the teeth, 1/6 or more the length of the teeth, papillose or weakly papillose-striate with a basal membrane and cilia absent.

Calyptra

deciduous or persistent, mitrate to cucullate, smooth, usually long-rostrate and inflated towards the base.

large, cucullate, usually smooth, and often long-rostrate.

Spores

spherical or subreniform, strongly ornamented to smooth.

spherical, smooth or papillose to baccate-insulate.

Specialized

asexual reproduction absent.

Perigonia

terminal on short basal branches, bud-like, paraphyses yellowish and club-shaped.

Perichaetia

terminal, paraphyses usually absent and filiform when present, perichaetial leaves often somewhat enlarged.

Operculum

usually oblique to the axis of the capsule, convex to weakly conic, cells in obliquely radial rows.

Funariaceae

Funaria

Distribution
Worldwide except Antarctica
map from USDA
North America; Mexico; Central America; South America; West Indies; Europe; Asia (including Indonesia); Africa; Pacific Islands; Australia
Discussion

Genera ca. 13, species ca. 300 (6 genera, 29 species in the flora).

The Funariaceae is characterized by broad leaves, large, pale laminal cells, opposite peristomes, and the distinctive stomata. The majority of species are found in disturbed or open sites on bare soil. Many of the species are annuals or biennials, and some may be perennials. When sterile, only a few species have distinctive vegetative features that allow positive identification to genus or species level. Fortunately most are usually fertile and sporophytes are common, although seasonal.

(Discussion copyrighted by Flora of North America; reprinted with permission.)

Species ca. 200 (9 in the flora).

Funaria comprises mainly small to medium seasonal mosses growing on moist mineral or peaty soils in strong light. For the most part, they are relatively short-lived pioneer species adapted to complete the life cycle by producing many spores quickly, in a cool, moist, bright (but not sunny for long periods), exposed, disturbed habitat. In North America, the best time to look for members of the family is spring before the soil dries out. The most common species can be recognized by the production of large numbers of sporophytes bearing a double peristome with inner and outer teeth opposite rather than alternate as is typical for most mosses. The teeth tend to be torqued in one direction with the tips of the exostome adhering weakly to a few-celled disk. Because the sporophyte shows more morphologic diversity than the gametophyte, it is often essential for identification. H. A. Crum and L. E. Anderson (1981) discussed the indistinct generic limits between Funaria and Entosthodon and the application of generic names.

Etymology: Latin funis, rope, alluding to cord-like twisted seta

(Discussion copyrighted by Flora of North America; reprinted with permission.)

Parent taxa Funariaceae
Subordinate taxa
Aphanorrhegma, Entosthodon, Funaria, Physcomitrella, Physcomitrium, Pyramidula
F. americana, F. apiculatopilosa, F. arctica, F. flavicans, F. hygrometrica, F. microstoma, F. muhlenbergii, F. polaris, F. serrata
Key
1. Calyptra large and four-angled, completely enclosing the mature capsule.
Pyramidula
1. Calyptra smaller than mature capsule, not angled or persistent
→ 2
2. Capsules immersed
→ 3
2. Capsules emergent or long-exserted
→ 5
3. Capsules rupturing irregularly; exothecial cells delicate, thin-walled.
Physcomitrella
3. Capsules operculate; exothecial cells firm, thick-walled or collenchymatous
→ 4
4. Exothecial cells collenchymatous; spores spiculose-papillose.
Aphanorrhegma
4. Exothecial cells not collenchymatous; spores papillose.
Physcomitrium
5. Capsules inclined and asymmetric; peristome double, endostome well developed to somewhat rudimentary.
Funaria
5. Capsules erect and more or less symmetric; peristome either single, rudimentary, or absent
→ 6
6. Capsules sub-cylindric to narrowly pyriform; most exothecial cells oblong to oblong-linear, rarely isodiametric; calyptra cucullate.
Entosthodon
6. Capsules urn-shaped, broadly pyriform, to cupulate; operculum rostrate; most exothecial cells irregularly hexagonal, ± isodiametric; calyptra mitrate to irregularly mitrate, sometimes appearing cucullate.
Physcomitrium
1. Annulus absent
→ 2
1. Annulus large, revoluble
→ 5
2. Distal leaves with a long, flexuose, excurrent costa with a hyaline tip.
F. apiculatopilosa
2. Distal leaves acute to narrowly acuminate, costa percurrent or, if excurrent, straight and concolorous
→ 3
3. Leaves with acute tips, distal marginal cells serrate from a series of elongate blade cells.
F. serrata
3. Leaves with acuminate tips, marginal cells not different from the blade cells
→ 4
4. Costa ending in a narrowly acuminate tip or excurrent, endostome segments broadly triangular proximally and nearly as long as the exostome teeth, east of Rockies.
F. americana
4. Costa ending before the slender, filiform acumination, endostome segments narrowly lanceolate and about 2/3 the length of the exostome teeth, western states.
F. muhlenbergii
5. Endostome segments lanceolate, slender pointed, at least 2/3 the length of the exostome teeth
→ 6
5. Endostome segments narrow, less than 1/2 the length of the exostome teeth, or rudimentary
→ 7
6. Seta strongly twisted and hygroscopic, 20-80 mm, capsule with an oblique mouth sometimes almost parallel to the axis of the capsule, leaves acute to acuminate, often with a short excurrent costa.
F. hygrometrica
6. Seta curved to cygneous, 8-12 mm, capsule mouth scarcely oblique to the axis of the capsule, leaves broadly acute to obtuse, costa ending before or in the tip.
F. arctica
7. Capsule symmetric or nearly so, seta strongly curved, axis of the capsule aligned with the seta.
F. polaris
7. Capsule curved to nearly straight, horizontal to pendent from the seta
→ 8
8. Capsule nearly straight and cylindrical, mouth 2/3 or more the diameter of the capsule, leaves tending to be broadly oblong-ovate.
F. flavicans
8. Capsule curved and pyriform, mouth 1/2 or less the diameter of the capsule, leaves tending to be oblong-lanceolate to oblanceolate.
F. microstoma
Name authority Schwägrichen Hedwig: Sp. Musc. Frond., 172. 1801 ,
Source FNA vol. 27, p. 180. Treatment author: Terry T. McIntosh. FNA vol. 27, p. 188. Treatment authors: Donna H. Miller, Harvey A. Miller.
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