BFNA Title: Anomodontaceae
Author: I. Granzow-de la Cerda 
Date: November 8, 2007
Edit Level: R
Version: 1

Bryophyte Flora of North America, Provisional Publication
Missouri Botanical Garden

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Íñigo Granzow-de la Cerda


Plants delicate to robust, forming glaucous-green, dense or loose mats, freely branched and irregularly pinnate from a creeping stem. Stems prostrate, sparsely branching; paraphyllia none.  Stem leaves minute and scale-like, differing in size and shape from branch leaves, apex acute-acuminate to rounded; costa single, long and thick, usually pellucid, ending below the apex or short and double; laminal cells short. Sexual condition dioicous. Capsule exerted, erect. Peristome reduced, exostome whitish yellow to pale brown, often striolate at the base and papillose. Seta flexuouse, dark to lightly reddish or  brown.


Genera 4 (2 in the flora), species ca. 20 (8 in the flora):  mostly panboreal, e North America, South America, e Asia (India, Japan, China), Africa, Australia, Pacific Islands.


1.  Leaves abruptly narrowed near midleaf from a broadly ovate to lanceolate base; laminal cells with multiple, branched papillae or a single, unbranched papilla . . .                          1. Anomodon

1.  Leaves lanceolate, gradually tapering; laminal cells smooth . . .                     2. Herpetineuron




1.  Anomodon Hooker & Taylor, Musc. Britt. 79. 1818  *  [Greek, anomalos, abnormal, and odontos, tooth]


Haplohymenium Dozy & Molkenboer, Musci Frond. Ined. Archip. Indici, 127. 1846


Plants delicate to robust, forming more or less glaucous-green to ferrugineous-brown, dense and thick to loose and delicate mats. Stems with  erect to arcuate branches, sometimes attenuate to flagellate at the end, scarcely to profusely branching secondarily; central strand of stem with differenciated cells in some species; pseudoparaphyllia absent in all but two species. Branch leaves adpressed to moderately secund or crispate when dry, complanate or erect to imbricate, when moist, broadly ovate to lanceolate not plicate proximally, more or less abruptly narrowed toward the middle, distal portion beyond the shoulders with parallel margins to tapering; margins flat, entire throughout; apex rounded to obtuse to acute or narrowly acuminate; costa single, usually strong, pellucid, smooth or papillose abaxially, ending sharply at or near the apex, but sometimes obscured by lamina cells and ending  before the middle of the leaf, sometimes discretely 2-fid at the tip; laminal cells  small, hexagonal, thin-walled and obscure to irregular, oblong, incrassate, smooth and pellucid at the base, with single or multiple, mostly well-developed papillae on both surfaces, but not prorulate. Sexual condition dioicous; perichaetial leaves well differentiated, narrow and longer. Seta long, to 20 mm. Capsule erect, symmetric, ovoid, cylindric to oblong-cylindric, sometimes with stomata; operculum conic to obliquely short-rostrate. Peristome reduced, exostome white to pale brown, teeth narrowly lanceolate, densely papillose, occasionally cross-striolate and sometimes slightly trabeculate; endostome sometimes very reduced or wanting (sect. Haplohymenium), pale with low basal membrane, delicately papillose, segments keeled to linear and reduced or absent, cilia absent or nearly so. Calyptra smooth to papillose or hirsute (sect. Haplohymenium). Spores 9 --20(--23) /um.


Species 16 (7 in the flora): temperate, circumboreal, North America, Central America, South America (Bolivia), s Africa, Australia, Pacific Islands (New Zealand).


Regarding the name Anomodon, it was once believed that the exostome was formed by a double set of teeth: the second one is the endostome segments, certainly an anomalous condition for Neckera, from which Anomodon was segregated. Anomodon viticulosus grows on shaded calcareous outcrops, but the remainder of the species are found on  tree trunks, including the base, logs, sometimes soil or rocks. The species are an important element of the eastern North American deciduous forest. Often several (up to 4) of the species grow together: A. attenuatus, A. rostratus and A. minor---our most abundant species---may grow on the same tree, with A. rugelii, sometimes joining them in sub-montane regions. Although A. tristis may be found on the same trees as either of the above four species, it usually forms much thinner and delicate mats higher on the tree. Anomodon thraustus, also an epiphyte is here recognized as a rare but distinct species which has been consistently misidentied as A. minor in North America, and A. longifolius, only known from a locality in New York growing in rock crevices.All species of Anomodon are dioicous. In North America, at least two, A. rostratus and A. attenuatus, fruit profusely. On the other hand, A. rugelii and A. minor fruit less abundantly and perhaps less frequently, while sporophytes of A. viticulosus are extremely rare in North America (only one fertile specimen of A. viticulosus seen, none of A. tristis).  In Europe, however, A. viticulosus is the most common species and often produces sporophytes.


Selected references  Granzow-de la Cerda, Í. 1989. Notes on five species of Anomodon, some with erroneous identity, including two new combinations. Bryologist 92: 381--386. Granzow-de la Cerda, Í. 1997. Revision and phylogenetic study of Anomodon and Herpetineuron. (Anomodontaceae, Musci). Contr. Univ. Michigan Herb. 21: 205--275. Iwatsuki, Z. 1963. A revision of the east Asiatic species of the genus Anomodon. J. Hattori Bot. Lab. 26: 27--62. Noguchi, A. 1957. A revision of the genus Haplohymenium Doz. & Molk. (Musci). Kumamoto J. Sci. ser. b. sect. 2, 3(1): 20--35.  Schumacker, R., P. De Zuttere and L. Leclercq. 1982. Anomodon rostratus (Hedw.) Schimp. (Thuidiaceae), new for the Belgian bryoflora, in the south of the Ardenne massif (Rochehaut, prov. Luxembourg, Belgium). J. Bryol. 12: 171--177.  

Vitt, D. H. and P. Lee. 1984. Anomodon minor (Musci: Leskeaceae) in North America. Bryologist 87: 338--339.  



1.  Leaves long-lanceolate, narrowly acuminate or ending in a long hairpoint.

2.  Plants dull green, slender, sparingly branched; branches prostrate or pendulous, somewhat attenuated; leaf cells with one single, short, unbranched  papilla . . .

8. Anomodon longifolius

2.  Plants glaucous, profusely and iregularly branched; branches julaceous, erect; leaves small, less than 0.8 mm, apex acute, ending in a long hairpoint . . .

6. Anomodon rostratus

1.  Leaves ligulate, constricted near mid-leaf, rounded, obtuse, acute or apiculate, but not ending in a long hairpoint.

3.  Plants slender, delicate, stems less than 1.1 mm thick when dry, leaves less than 1.2 mm, with apices broken off; costa obscured by laminal cells distally.

4. Stems less than 0.5 mm thick when dry, leaves less than 1.2 mm; basal cells few, not reaching the margin; costa weak, obscured by laminal cells almost throughout   (sect. Haplohymenium)  4. Anomodon tristis

4. Stems 0.5--1.1 mm thick when dry, leaves 1.2--1.8(--2.1) mm, with apices; basal cells numerous, reaching the margin; costa somewhat strong but obscured by laminal cells in its distal 1/2 (sect. Thraustus) . . .  5. Anomodon thraustus

3.  Plants rather robust, branches more than 0.8 mm thick when dry, leaves greater than 2 mm, intact; costa ending near the apex, often asymmetrically 2-fid at the end; basal cells almost reaching the margins and extending ± 1/3 of the lamina.

5.  Plants rusty-brown; leaves crispate when dry, broadly  auriculate; stem and costa dark brown to reddish; pseudoparaphyllia present . . . 2. Anomodon rugelii

5.  Plants dull-green to yellow; leaves not crispate when dry, broadly decurrent, not auriculate;  stem light brown, costa yellowish to light green; pseudoparaphyllia absent.

6.  Plants robust, branches ca. 1.3 mm thick or more when dry; leaves flexuose, falcate-secund, spreading when moist, not appresed when dry, greater than 2 mm . . . 1. Anomodon viticulosus

6.  Plants of various sizes, but branches never more than 1 mm thick when dry; leaves complanate when moist, adpressed when dry, less than 2 mm.

7.  Plants profusely branching, irregularly pinnate, apex of secondary branches attenuate; inflorescencences never present on terminal branches but proximally from the most distal branching points; leaves vaguely constricted at middle, tapering to a triangular distal portion, acute, occasionally obtuse, often dentate at the apex, almost always apiculate; abaxial cell of costa not papillose . . . 7. Anomodon attenuatus

7.  Plants poorly branching, not pinnately so, apex of secondary branches not attenuate but often slightly clavate; inflorescences present on terminal branches, distally from the most distal branching points; leaves abruptly constricted at middle into a distinctly lingulate distal portion, rounded, entire at the apex, never apiculate; abaxial cells of costa with large papillae arranged in a row . . . 3. Anomodon minor


1a. ANOMODON Hooker & Taylor subg. ANOMODON


Stems and primary branches scarcely branching, in a simple, parallel pattern; secondary branches not tapering at the apex in most species. Branch leaves with apices rounded to broadly obtuse (narrowly obtuse to acuminate in sect. Haplohymenium), abruptly narrowed proximal to midleaf from an ovate-lanceolate base; laminal cells isodiametric, hexagonal, quadrate or round in all taxa. Peristome usually poorly developed, exostome smooth or faintly striolate at the base and papillose distally in some species, endostome very reduced or wanting, consisting of a low basal membrane, 2--4 cells high, with segments rudimentary in most species, often reduced to 2--3 cells or absent, never keeled and mostly smooth throughout.


1a.1. Anomodon Hooker & Taylor sect. Anomodon


Plants rather robust to somewhat slender,forming rather thick, dense mats. Stems with secondary branches terete, never flagelliform or attenuate. Branch leaves not delicate, oblong-lingulate (1.2--)1.4 mm; costa subpercurrent or ending sharply near the apex, well beyond the middle of the leaf, not obscured by laminal cells. Seta  5--22 mm. Capsule 2--4 mm, urn greater than (1.5--)1.8 mm. Calyptra smooth and glabrous.



1.  Anomodon viticulosus (Hedwig) Hooker & Taylor, Muscol. Brit., 79. 1818


Neckera viticulosa Hedwig, Spec. Musc., 209. 1801; Hypnum viticulosum (Hedwig) L. ex Withering


Plants robust, forming thick mats, dull green. Stems 6--8 cm, occasionally more, 1--1.8 mm thick when dry, with primary branches erect-ascending to arcuate, somewhat branched; central strand of stem without differentiated cells; pseudoparaphyllia absent; rhizoids few. Branch leaves erect when dry, always secund and somewhat flexuose, spreading to reflexed when moist, oblong-ligulate, 2.2--4 mm; base broadly decurrent; margin plane; apex obtuse to rounded, sometimes acute; costa strong, pellucid, lighter green, ending sharply (only occasionally obscured by laminal cells) 0.05--0.15 mm before the apex, sometimes flexuose, rarely asymmetrically 2-fid at the end; basal laminal cells hyaline, barely extending beyond 1/4 the length of the leaf base, each cell bearing a single papilla, the basal-most with walls sinuose, often perforate; medial and distal laminal cells obscure, hexagonal, with multiple branched papillae, those that are abaxial to the costa long, each with several thick papillae in rows. Inflorescences borne distally from the last branching points; perichaetial leaves similar in shape and size to vegetative leaves, with costa ending closer to the apex, cells papillose. [Seta 10--20 mm.  Capsule long-elliptic; urn (1.7--)2--3.1(--3.3) mm, stomata lacking; annulus well differentiated; operculum obliquely rostrate, 0.7--1 mm; exostome teeth irregular, 0.3--0.5 mm, nearly smooth, with the base yellow, faintly striolate, not trabeculate, with inconspicuous papillae toward the apex; endostome with basal membrane 2--4 cells (0.07--0.09 mm) high, almost smooth, segments moderately developed, 0.07--0.15(--0.2) mm, several cells high, smooth or variably papillose. Spores very variable in diameter between capsules, in some (19--)20.5--23(--25) /um, in others 15--16 /um, densely papillose. Only one specimen seen with sporophytes in North America; description from European and Asian material].


Capsules mature early--mid fall. Almost restricted to rather mesic calcareous environments of montane deciduous forests, limestone rocks or vertical walls, sometimes also as an epiphyte; 200--1700 m; N.B., N.S., Ont., Que.; Ark., Ill., Iowa, Ky., Mass., Mich., Mo., N.H., N.J., N.Y., Pa., Tenn., Vt., Va., Wis.; Mexico (Guerrero, Oaxaca); Europe; Asia.


Anomodon viticulosus is the most robust species of the genus, forming thick mats on rocks, sometimes on tree trunks. Its robust habit and thickness of the mats make it clearly distinguishable from other species with which it could be mistaken. In depauperate populations, A. viticulosus  can always be distiguished from A. rugelii by its falcate-secund leaves, decurrent and tapering from the shoulders toward the apex, instead of the leaves incurved when dry of A. rugelii, with their characteristic auricles at the insertion, and lingulate beyond the shoulders. Unlike A. rugelii or A. minor, the apex of A. viticulosus is never rounded nor are its terminal branches complanate. Anomodon viticulosus is seldom mistaken for A. attenuatus, although the latter has more prostrate stems and its branching is more profuse, in a pattern consisting of successive orders of branhing, with the terminal branches attenuate, complanate, crowded and somewhat fasciculate. In A. attenuatus the abaxial surface of the costa is smooth and perichaetia do not develop on the last year's branches (but only in older portions of branches). This species seldom fruits in North America (only one fruiting specimen seen) or Asia, most likely because of the lack of male gametophytes in these regions (I. Granzow-de la Cerda (1989).


2.  Anomodon rugelii (Müller Hal.) K. Keissler, Ann. Naturh. Hofmus. Wien 15: 214. 1900


Hypnum rugelii Müller Hal., Syn. 2: 472. 1851


Plants rather slender, forming thick, dense mats, dark green to rusty-brown. Stems to 3.5(--5) cm, 0.8--1.5 mm thick when dry, with few primary branches, erect-ascending, somewhat arcuate, poorly branched; central strand of stem with differentiated cells; pseudoparaphyllia foliose, often narrow to 1-seriate, golden-yellow; rhizoids abundant. Branch leaves incurved-contorted when dry, erect-spreading, somewhat secund when moist, broadly oblong-ligulate; (1.2--)1.4--2.3(--2.5) mm; base auriculate with margins of auricles with long-branched spiny papillae; margin plane; apex broadly obtuse to rounded, often apiculate, sometimes minutely denticulate; costa strong, pellucid, golden yellow to rusty brown, ending sharply near the apex, sometimes slightly flexuose and generally asymmetrically 2-fid at the end; basal laminal cells hyaline and well differentiated, occupying more than half the basal portion of the lamina, with walls not papillose, often sinuose, marginal cells of the auricles with 1--2 strong branched spine-like papilla; medial and distal laminal cells hexagonal, 7--12 /um, with multiple high branched papillae, those adaxial to the costa elongate, 40--70 /um, the abaxial ones smooth. Inflorescences at the end of terminal secondary branches; perigonial leaves with distal margins crenulate, cells with few or no papillae; perichaetial leaves, abruptly narrowed toward the apex, becoming almost subulate, cells papillose. Seta (5--)9--22 mm. Capsule elongate, urn (1--)1.8--2.3(--2.5) mm, stomata at the base; annulus absent; operculum obliquely short-rostrate, 0.5--0.8 mm; exostome teeth regular, 0.15--0.3 mm, papillose throughout, often inconspicuously horizontally striolate at the base, trabeculate above; endostome rudimentary, 0.042--0.057 mm, with a basal membrane 2--4(--6) cells high, segments very reduced (1 cell high) or wanting. Spores 9.5--14 /um., slightly papillose.


Capsules mature by mid fall. Extensive, dense and thick mats on tree trunks (generally, although not always, 1--2 m above the base) and on basic as well as acidic rocks; montane deciduous forests; 500--1800 m; N.B., N.S., Ont., Que.; Ga., Ill., Ind., Ky., Maine, Mass., Mich., Mo., N.H., N.Y., N.C., Ohio, Pa., Tenn., Vt., Va., Wis.; c, n Europe, Scandinavia; Asia (Azerbaijan, c, ne China, Japan, n India, Korea, Nepal, e Russia, Vietnam) .


Distinctive characters for Anomodon rugelii include the rusty brownish color of the plant, felt-like primary branches due to the abundance of rhizoids, more slender branches with strongly incurved leaves when dry, redness of the costa, but above all, conspicuous auricles at the leaf base. Apiculate leaves are not a reliable character, as there are many plants the leaves of which are obtuse to rounded and lack an apiculus. Anomodon rugelii usually has foliose pseudoparaphyllia, although they can sometimes be completely 1-seriate. Fruiting mats are infrequent, but when found, sporophytes are produced in abundance.


3. Anomodon minor (Hedwig) Fürnrohr, Fora 12 (Erg. 2): 49. 1829


Neckera viticulosa var. minor Hedwig, Spec. Musc., 210. 1801; Anomodon platyphyllus Kindberg; Neckera minor (Hedwig) Palisot de Beauvois


Plants of medium size, forming loose mats, dull green when dry, bright green when moist. Stems up to 2.5 cm, exceptionally to 6 cm, (0.6--)1--1.7 mm thick when dry, with primary branches erect-ascending to arcuate, julaceous when dry, not or seldom branched; central strand of stem with poorly or not differentiated cells; pseudoparaphyllia absent, rhizoids few. Branch leaves imbricate, rarely slightly crisped when dry, erect to spreading, complanate, not secund when moist, broadly oblong-ligulate, sometimes slightly spatulate; (1--)1.5--2.3(--3) mm; base broadly decurrent, sometimes dentate to spinulose at the insertion because of very high papillae and sometimes infolded; margins undulate; apex rounded; costa moderately robust and prominent abaxially, pellucid, ending (0.04--)0.08--0.15(--0.2) mm before the apex, often asymmetrically 2-fid at the end, not obscured by laminal cells near the apex; basal hyaline cells extending just 1/3 of the basal leaf portion or less, mostly papillose; medial and distal laminal cells hexagonal, 7--12 /um wide, densely papillose, with high branched papillae; abaxial cells of the costa elongate (40--68 /um), with globulose papillae in rows. Inflorescences on terminal branches, beyond the distalmost branching points; perichaetial leaves with papillose cells toward the apex. Seta 4--16 mm. Capsule ovoid; urn 1.5--2.2 mm, without stomata; annulus well developed; operculum obliquely short-rostrate, 0.35--0.6 mm; exostome reduced, teeth irregular, up to 0.35 mm, not striolate proximally, papillose distally, often trabeculate; endostome very reduced, consisting of a papillose basal membrane, no more than 2 cells high (ca. 0.035 mm) and segments ca. 0.045 mm, irregular and fragmentary. Spores (11)13--18(21) /um, practically smooth to faintly papillose.


Capsules mature early fall. Deciduous forests on calcareous rocks as well as bark of trees, forming loose thin mats, somewhat discontinuous; 200--1600 m; Alta., Man., N.B., Ont., Que.; Ala., Ariz., Ark., Conn., D.C., Del., Fla., Ga., Ill., Ind., Iowa, Kans., Ky., La., Maine, Md., Mass., Mich., Minn., Miss., Mo., Nebr., N.H., N.J., N.Mex., N.Y., N.C., N.Dak., Ohio, Okla., Pa., S.C., S.Dak., Tenn., Tex., Vt., Va., W.Va., Wis.; Mexico, Central America (Costa Rica, Guatemala); Asia (Bhutan, n Burma,  n India, s, e, ne China, Japan, Korea, , n Pakistan, sc Russia ).


Large specimens of Anomodon minor with occasional non-complanate leaves can be told from A. viticulosus by their  more slender habit and smaller leaves, imbricate when dry and not secund when moist, while A. viticulosus has more congested, secund branch leaves. Often specimens of A. minor have strongly spinulose basal marginal cells, like those of the auricles of A. rugelii. The lack of auricles and the thick papillae on the back of the costa remain good characters to separate both species, although it is also true that the presence of obvious papillae abaxial on the costa in A. minor is not absolutely consistent. Characters that traditionally have been taken as of great taxonomic value for both A. minor and A. rugelii may be found mixed, and therefore are questionable. The only characters that remain congruent with each other are the auricles (sometimes extremely reduced) and foliose pseudoparaphylla of A. rugelii (both structures absent in all specimens of A. minor). Specimens of A. thraustus, a rare species in North America, have been misidentified as A. minor, but the distal part of the costa of the latter remains pellucid for most of its length and often becomes asymmetrically 2-fid before ending, while in A. thraustus the costa is obscured by abaxial laminal cells for the distal (1/2--)1/3 of the lamina, and the distal portion of its delicate, narrowly lingulate leaves is broken off. In North America A. minor requires mesic habitats and fruits infrequently.


1a. 2. ANOMODON sect. HAPLOHYMENIUM (Dozy & Molkenboer) Granzow-de la Cerda


Haplohymenium Dozy & Molkenboer, Musci Frond. Ined. Archip. Ind., 127. 1846


Plants slender, very delicate, forming loose, thin mats.  Stems with secondary branches often flagelliform. Branch leaves very delicate, less than 1 mm; costa ending near or before the middle of the leaf. Seta 2--5 mm.  Capsule 1--2 mm, urn 1 mm or greater. Calyptra papillose and hirsute.


The genus Haplohymenium was created to accommodate plants that resembled Anomodon but are more slender and have a papillose calyptra with long, hyaline scattered hairs. Segregating Haplohymenium as a separate genus would make the rest of Anomodon paraphyletic, as Haplohymenium is a sister group of A. rugelii, A. viticulosus and A. minor, all of which are part of subg. Anomodon (I. Granzow-de la Cerda 1997).  All these taxa are more distantly related to subg. Pseudoanomodon, to which A. rostratus and A. attenuatus belong.


4. Anomodon tristis (Cesati) Sullivant & Lesquereux, Musci Bor. Amer. 52. 1856


Leskea tristis Cesati in G. De Notaris, Syll. Musc., 67. 1838; Haplohymenium triste (Cesati) Kindberg; Hypnum triste (Cesati) Müller Hal.


Plants  filiform to wiry, dull dark brownish green. Stems 0.5--1.5 cm, 0.3--0.5 mm thick when dry, wiry, with main stem creeping, sparsely branching into prostrate primary branches, which rarely branch themselves; central strand of stem without differentiated cells; pseudoparaphyllia absent; rhizoids few. Branch leaves julaceous when dry, spreading, not complanate when moist, very delicate, narrowly ligulate to tapering, often with apical 1/3 of the leaf broken off; 0.5--0.9 mm; base narrowly decurrent; margin flat, crenulate toward the apex; apex narrowly obtuse to acuminate, with very prominent marginal papillae; costa short and delicate, pellucid proximally, ending before the middle of the leaf, obscured by laminal cells; basal laminal cells elongate, not papillose at the insertion; medial laminal cells round ca. 4 /um, with multiple prominent papillae, thin-walled cells of margin becoming mammillose toward the apex. Inflorescences rare, on terminal branches; perichaetial leaves oblong, acuminate.


Sporophytes not found in North America. Bark of trees in deciduous forests; 300--1800 m;  N.B., N.S., Ont., Que.; Ala., Ariz., Ark., Conn., Del., Fla., Ga., Ill., Ind., Iowa, Ky., La., Mass., Maine, Md., Mich., Minn., Miss., Mo., N.H., N.J., N.Y., N.C., Ohio, Okla., Pa., S.C., Tenn., Tex., Vt., Va., W.Va., Wis.; Mexico (Jalisco, Nuevo León, Sonora, Veracruz); Central America (Costa Rica); South America (Bolivia); Europe (restricted to Alps of Italy, Switzerland, Austria); Asia (Turkey, c, e Russia, e China, Korea, Japan, Taiwan); Pacific Islands (Hawaii).


Although the degree to which the apex breaks off is variable within plants of Anomodon tristis, the character allows for easy identification of this species and others in section Haplohymenium. Other taxa outside the section, however, also present this feature. 


1a.3. ANOMODON sect. THRAUSTUS Granzow de la Cerda, Contr. Univ. Michigan Herb 21: 239. 1997


Plants slender, forming loose mats. Stems with secondary branches terete. Branch leaves delicate, narrowly lingulate, usually broken off at the middle; more than 1.2 mm; costa ending well before the apex, 2/3--3/4 the length of the leaf, fading, obscured by laminal cells abaxially, (1/2--)1/3 before the apex . Seta 5--13 mm.  Capsule 1.5--2.5 mm, urn 1.3--1.8 mm. Calyptra smooth and glabrous.


5. Anomodon thraustus Müller Hal., Nuovo Giorn. Bot. Ital., n. ser. 5: 207. 1898


Plants dark when dry, bright green when moist. Stems creeping, 1.5--3 cm, 0.5--1 mm thick, with primary branches erect to ascending, julaceous when dry, somewhat densely branched and sometimes irregularly pinnate, with branches up to 2 cm; central strand of stem with differentiated cells; pseudoparaphyllia absent, rhizoids few. Branch leaves slightly crisped, incurved when dry, secund, reflexed when moist, not complanate;  1.2--1.8(--2.1) mm; base ovate; margin flat, not papillose-crenulate at the insertion; apex obtuse to rounded; basal pellucid cells extending up to half the basal portion of the lamina, or beyond; laminal cells with multiple high-branched papillae; abaxial cells of the costa (where not obscured by laminal cells) with large papillae in rows, larger than those of the laminal cells. Inflorescences on terminal branches, beyond the most distal branching points, near the apices. [Seta 5--13 mm. Capsule ovoid; urn 1.3--1.8 mm, without stomata; annulus differentiated; operculum short-rostrate, ca. 0.7 mm; exostome teeth well developed, ca. 0.3 mm, papillose to the base; endostome papillose, basal membrane very low, 2--3 cells high (ca. 0.045 mm), with very reduced segments (not more than 0.035 mm). Spores 16--19.5 /um, green, finely papillose. Description based on sporophytes from China and Japan.]


Sporophytes not seen in North America. Bark of trees in deciduous forests; moderate elevations; N.C., N.J., N.Y.; Mexico (Hidalgo, Jalisco, San Luis Potosí); Asia (China, Japan, India, Nepal, e Russia).


Anomodon thraustus greatly resembles A. minor but it can be distinguished by its more delicate habit, with thinner branches, incurved or crisped leaves when dry, and which do not become complanate when moist but rather widely spreading, while branches of A. minor are always complanate distally when moist. Leaves of A. thraustus are narrower and tend to have broken apices, similar to that occuring in A. tristis, and costa is obscured by laminal cells in its distal 1/2--1/3. The delicate, narrowly lingulate distal leaf portion of A. thraustus is still rather distinct, while that of A. minor is wider and proportionally shorter.  Anomodon thraustus has been considered an east Asian species (Japan, e China), but isolated records from central Mexico exist (L. J. Gier 1980). There were no previous reports from elsewhere in the New World, as all North American (and most Mexican) specimens of this taxon have been misidentified as A. minor (and potentially could also be mistaken for A. tristis). Although certainly a rare species, the presence of A. thraustus in the New World has been overlooked, and it may be more common than here indicated. The disjoint distribution of A. thraustus parallels that of A. minor, although in the New World A. thraustus is somewhat more southerly in range.


1b.  Anomodon subg. PSEUDOANOMODON Limpricht, Laubm. Deutschl. 2: 774. 1895


Stems and primary branches branching in a stairway fashion;.secondary branches profusely fasciculate, in some species tapering at the apex. Branch leaves with apices acute to acuminate, only slightly narrowed beyond midleaf from an ovate-lanceolate to lanceolate base; laminal cells rhombic in most taxa. Peristome rather well-developed, exostome striolate at the base and papillose distally, endostome well developed (except for A. longifolius), consisting of a high basal membrane, 4--7 cells high, and strong, keeled segments, often papillose and as high as the exostome.


6. Anomodon rostratus (Hedwig) Schimper, Syn. Musc. Eur., 448. 1860


Leskea rostrata Hedwig, Spec. Musc., 226. 1801; Hypnum rostratum (Hedwig) Palisot de Beauvois


Plants forming thick dense, tight mats, glaucous, light to dull yellowish green. Stems prostrate, profusely branching, with primary branches 0.5--2 cm, ca. 1 mm thick when dry, clustered, with a pseudo-verticillate branching pattern, secondary branches erect; central strand of stem with differentiated smaller cells; pseudoparaphyllia absent. Branch leaves julaceous, imbricate when dry, erect when moist, 0.8--1.4 mm; ovate-lanceolate at base, broadly decurrent, acuminate, ending in a hyaline hair point several cells long (to 0.2 mm or more) and one cell thick, slightly constricted in the distal 1/3, to 1.2 mm, margin revolute; costa pellucid, ending well before the apex, ca. 10 /um, often somewhat flexuose distally; medial laminal cells rhomboidal, scarcely isodiametric in the basal 1/2--2/3 of the lamina, multipapillose, with papillae slightly branched; costa cells with one row of high and thick papillae abaxially, sometimes slightly branched, proximal costa cells in a group of a few with sinuose walls, almost scabrid. Inflorescences distal to the last branching nodes; perichaetial leaves elongate, long-acuminate with cells smooth. Seta 7--13 mm, sometimes shorter, dark red. Capsule short, elliptic; urn ovoid to oval-cylindric, 1--1.5 mm, stomata at the base; annulus of 2 rows of cells; operculum obliquely rostrate, 0.8--1.1 mm, asymmetrically rostrate; peristome well developed, exostome teeth 0.2--0.3 mm high, teeth trabeculate, with a yellowish base, striolate up to the middle or beyond, and apex conspicuously papillose; endostome well developed, with tall basal membrane, 7 cells high or more, as high as the exostome, segments keeled, long and slender, 0.09--0.11 mm, papillose, cilia absent. Calyptra smooth, glabrous. Spores 11.5--15 /um, densely papillose.


Capsules mature early--mid fall. Base or bark of deciduous trees, soil, less commonly on fallen logs, usually in deciduous forests, sometimes in rather open spaces as well as secondary forests, vertical calcareous rocks, occasionally acidic; 100--1600 m;  Alta., B.C., N.B., Nfld., N.S., Ont., Que.; Ala., Ariz., Ark., Colo., Conn., D.C., Del., Fla., Ga., Ill., Ind., Iowa, Kans., Ky., La., Maine, Md., Mass., Mich., Minn., Miss., Mo., Nebr., N.H., N.J., N.Mex., N.Y., N.C., Ohio, Okla., Pa., R.I., S.C., S.Dak., Tenn., Tex., Vt., Va., W.Va., Wis.; e and s Mexico (Chiapas, Hidalgo, Nuevo León, Puebla, San Luis Potosí); West Indies (Hispaniola, Jamaica); Central America (Guatemala, Honduras); Europe; Atlantic Islands (Bermuda).


Anomodon rostratus is absent from Japan and, in general, from eastern Asia, given that most records are misidentifications. It forms dense and thick mats, often very extensive. This species is well distinguishable from congeners in the short branch leaves, ending in a hair point. The leaf areolation is also more lax; the basal portion of rhomboid cells extends beyond 1/2 the leaf length. The species fruits frequently and abundantly in North America. Despite this, male gametophytes in North America are extremely scarce, even in populations fruiting profusely. The paucity of male plants is probably the reason why sporophytes are virtually unknown elsewhere.


7.  Anomodon attenuatus (Hedwig)  Hübener, Musc. Germ., 562. 1833


Leskea attenuata Hedwig, Spec. Musc., 230. 1801; Hypnum attenuatum (Hedwig) Smith


Plants forming dense mats, dull yellowish green. Stems 1.5--3 cm, 0.8--1 mm thick when dry, prostrate to arcuate, irregularly pinnate, secund when dry, complanate when wet, mostly attenuate, branching pattern often consisting of several orders of successive branching, in a stepwise fashion; central strand of stem without differentiated cells; pseudoparaphyllia absent. Branch leaves lanceolate, slightly narrowed at the middle, 1.2--2(--2.2) mm, base ovate-lanceolate, broadly decurrent, apex acute, sometimes obtuse or slightly apiculate; margin flat, entire from the base to the apex (although cells moderately mammillose at the insertion), sometimes denticulate near the apex; costa pellucid, sharply ending a few cells before the apex, sometimes fading in youngest leaves; laminal cell walls somewhat sinuose, perforate near the costa; basal cells poorly differentiated, usually hyaline but sometimes chlorophyllose and extending more than 1/2 the length of leaf base; distal laminal cells quadrate, 6--8 /um, multipapillose, papillae branched; abaxial cells of costa mostly short (less than 23 /um, often ca. 35 /um). Inflorescences never beyond the last branching points; perichaetial leaves abruptly narrowed, costa reaching well into the distal half of lamina, ending near the apex in the most interior leaves. Seta 10--15 mm. Capsule oblong; urn 1.6--2.8 mm, stomata at the base; annulus not differentiated; operculum obliquely short-rostrate, ca. 1.3 mm; exostome teeth irregular, 0.35 mm, yellow, becoming white with age, striolate proximally, trabeculate and more or less finely papillose towards the apex; endostome papillose with high basal membrane (3--4 cells high), segments keeled and well developed, up to 0.3 mm, cilia absent.  Spores 10--13 /um, densely papillose.


Capsules mature in early--mid fall. Tree bark, mostly at the base, soil, rocks; low to moderate elevations. Man., N.B., Nfld., N.S., Ont., P.E.I., Que.; Ala., Ariz., Ark., Colo., Conn., D.C., Del., Fla., Ga., Ill., Ind., Iowa, Kans., Ky., La., Maine, Md., Mass., Mich., Minn., Miss., Mo., Nebr., N.H., N.J., N.Mex., N.Y., N.C., Ohio, Okla., Pa., R.I., S.C., S.Dak., Tenn., Tex., Vt., Va., Wash., W.Va., Wis.; Mexico; West Indies (Cuba, Jamaica, Hispaniola); Central America (Guatemala, Belize); Europe; Asia (Turkey, e Russia, India).


Anomodon attenuatus is a rather polymorphic species producing abundant sporophytes in North America. The attenuate branches, sometimes arcuate and seldom ascending, are distinctive. This character, however, is sometimes lacking; instead, in some specimens the branch apices become slightly capitate and incurved. Robust plants of A. attenuatus might be mistaken for A. viticulosus as both have acute leaf apices. They can be told apart, however, by their branch morphology:  ascending and robust in A. viticulosus, prostrate and complanate in A. attenuatus. Also, the leaf shape is different in both species: lanceolate and with no constriction in A. attenuatus, broadly ligulate, abruptly narrowed from a broadly ovate base in A. viticulosus. The costa in A. attenuatus also lacks the thick, aligned papillae on the dorsal surface. A consistent character in fertile plants is the absence of gametangia beyond the most recent branching points (i.e., they are never formed in the youngest branches).


8. Anomodon longifolius (Bridel) Hartman, Handb. Skand. Fl. ed. 3, 300, 1838


Plants variable in size, from somewhat slender to robust, forming thin to rather thick mats, yellowish green. Stems slender, 2--3(--6) cm, 0.5--1 mm thick when dry, pendant, with primary branches, fasciculate or irregularly pinnate into secondary branches that areusually attenuate, often flagelliform, frequently club-shaped and/or recurved at their apices; central strand of stem with well differentiatedand smaller cells; pseudoparaphyllia foliose; rhizoids abundant distally. Branch leaves rather densely arranged, secund, often falcate, adpressed to almost recurved when dry and somewhat plicate at the base; in slender forms julaceous to imbricate when dry; 2.1--2.8 mm and sometimes more in robust forms, 0.3--0.5(--0.6) mm in slender forms; decurrent, wide at the base, narrowing more or less abruptly before the middle of the leaf (0.2--0.35 mm at the widest beyond the shoulders), tapering into a subula-like apex, often 2--3 cells wide at the apex; margins revolute near the shoulder, entire to serrulate and often with shorter and wider cells near the apex; costa yellowish, somewhat decurrent, protruding and making the leaf carinate, often scabrose on the back in the distant half, percurrent; basal cells near the costa generally not hyaline, smooth, longer than those of the rest of the lamina, the basal group extending less than 1/4 the length of the leaf base; medial laminal cells ca. 15 /um, with single, central, rather high papillae on both surfaces, these never exceeding the cell's diameter; abaxial cells of the costa noticeably shorter and incrassate, as long as those of the lamina (7--11 /um).  Inflorescences on the most distal portions of the branches (with just minute or flagelliform branches produced distally from the inflorescences); perigonial leaves smooth, areolation lax; perichaetial leaves with isodiametric cells, smooth near the apex. [Seta 4--8 mm, yellow. Capsule elongate, cylindric, urn 1.2--1.7(--2) mm, stomata lacking; annulus not differenciated; operculum conic obtuse to short-rostrate, somewhat asymmetric, ca. 0.5 mm; exostome teeth irregular, 0.25--0.30 mm, striolate proximally; endostome very rudimentary, 0.08--0.13 mm, generally reduced to a short pellucid basal membrane (1--2 cells high), sometimes bearing a few scattered very slender segments of the same height. Spores 12--17 /um.]


Crevices between boulders; 620 m; N.Y.; Asia (Azerbaijan, Georgia and w Siberia, Japan); Europe.


A single collection of Anomodon longifolius, from the Adirondack Mountains of New York, is the only record for the New World. This species is something of a morphological outlier within the genus, as it has a single wide and short, rounded, unbranched central laminal cell papilla. Distinctive characters are the incrassate, isodiametric, irregular or shortly rectangular cells on the back of the costa. It is also the only species in the genus, other than A. rugelii, that has pseudoparaphyllia. Although the specimen from New York has no sporophytes, the exostome of A. longifolius is also unique in being irregularly striolate, alternating with sometimes finely punctate areas, otherwise smooth distally, in some instances divided into two filaments which are generally verruculose-papillose. Because of superficial similarities in areolation and leaf shape, A. longifolius is often mistaken for Leskeella nervosa in Europe and Asia despite the more crowded, shorter, julaceous (instead of secund), more acuminate than subulate, non-papillose leaves of L. nervosa. The sometimes similar Heterocladium heteropterum is easily distinguishable as it has no costa. This species is may be looked for on tree bark and ledges, mostly in calcareous regions, shady places, moist deciduous montane or boreal forests



2. HERPETINEURON (Müller Hal.) Cardot, Beih. Bot. Centralbl. 19(2): 128. 1905  *  [Greek herpes, snake, and neuron, nerve, alluding to the terminally strongly sinuose costa]



Anomodon sect. Herpetineuron Müller Hal., Flora 73: 476. 1890


Plants rather robust, forming yellowish or brownish thick to somewhat loose mats. Stems with rather arcuate branches, circinate or often attenuate to flagellate d, moderately branching secondarily; central strand of stem with well differenciated, smaller cells; pseudoparaphyllia absent.  Branch leaves incurved, secund when dry, erect, not complanate when moist, ovate-lanceolate, broadly decurrent, somewhat plicate proximally, tapering from the base, without an obvious constriction; margins flat, serrate in the distal 1/3--1/4;  apex sharply acuminate; costa single, strong, pellucid, cells smooth abaxially, not obscured by laminal cells, ending sharply at or near the apex, not 2-fid at the end; laminal cells rectangular to rhombic, slightly longer than wide,  walls somewhat thickened, smooth and pellucid throughout the lamina, some prorulate in the distal part. Sexual condition dioicous. Inflorescences with perichaetial leaves somewhat differentiated. [Seta 14--16 mm. Capsule erect, almost symmetrical, stomata few; operculum conic to obliquely short-rostrate. Peristome fairly well developed, exostome teeth yellowish, papillose  and faintly striolate at the base and beyond, endostome with basal membrane to 3--4 cell high, segments papillose throughout. Calyptra smooth and glabrous. Spores 16--22 /um.]


Species 2 (1 in the flora): mostly temperate or montane regions, worldwide.


SELECTED REFERENCES   Granzow-de la Cerda, Í. 1997. Revision and phylogenetic study of Anomodon and Herpetineuron. (Anomodontaceae, Musci). Contr. Univ. Michigan Herb. 21: 205--275.  Norris, D. H. and A. J. Sharp. 1961. The known distribution of Herpetineuron toccoae (Sull. & Lesq.) Card. J. Hattori Bot. Lab. 24: 110--114.


1. Herpetineuron toccoae  (Sullivant & Lesquereux) Cardot, Beih. Bot. Centralbl. 19(2): 128. 1905


Anomodon toccoae Sullivant & Lesquereux, Musci Bor. Amer., 52. 1856



Plants dull green, dark greenish to yellowish brown. Stems (1--)2--4 cm, (0.2--)0.9--1.3 mm thick, terete when dry, branching pattern often consisting of several orders of successive branching, in a stepwise fashion.  Branch leaves involute in a tubular fashion when dry; 1.8--2.8 x  0.5--1.2 mm; margin serrations consisting of 1 to several cells; costa almost always sinuose in the distal 1/3.  Inflorescences on youngest branches; perichaetial leaves 1.5--2.2 mm, subulate, subula flexuose, often somewhat crispate, lamina cells smooth throughout. [Seta 14--16 mm. Capsule erect, urn (1.7--)2--2.8 mm, stomata few at the base; annulus well differentiated; operculum conic; peristoma well developed, exostome teeth ca. 0.5 mm, papillose and faintly striolate at the base and distally; endostome with a basal membrane 0.05--0.15 mm, segments 0.2--0.3 mm, papillose. Calyptra smooth. Spores 16--22 /um, densely papillose.  No sporophytes found in the Flora area; description based on sporophytes from China and Japan.]


Trees, sometimes at their base or, occasionally, rocks, temperate deciduous forests; moderate elevations; Ala., Ariz., Ark., Fla., Ga., Ill., Ky., La., Miss., N.C., Okla., S.C., Tenn., Tex.; Mexico; West Indies (Dominican Republic); Central America (El Salvador, Guatemala, Honduras, Nicaragua); South America (e Brazil); e, se Asia (Pakistan, India, Sri Lanka, Indonesia, China, Taiwan, Japan); w, s Africa (Tanzania, South Africa); Pacific Islands (New Caledonia, Papua New Guinea).




Gier, L. J. 1980. A preliminary study of the Thuidiaceae (Muci) of Latin America. J. Bryology 11: 253--309.

Iwatsuki, Z. 1963. A revision of the East Asiatic species of the genus Anomodon. J. Hattori Bot. Lab. 26: 27--62.