Campylopus

Plants usually 3–10 cm, occasionally longer. Stems usually simple, not tomentose or with dense reddish or whitish tomentum. Leaves 3–12 mm, erect-patent or appressed foliate, narrowly lanceolate, ending in a smooth or denticulate, straight or reflexed tip; alar cells large, inflated, hyaline or reddish brown, or not differentiated; basal laminal cells thin-walled, hyaline, or thick-walled, chlorophyllose, sometimes with pitted walls, rectangular to subquadrate; distal laminal cell walls incrassate, quadrate to short-rectangular, oblique, or oval to elongate oval; costa strong, filling 1/3–4/5 of leaf width, excurrent in a more or less long, chlorophyllose or hyaline awn, in transverse section showing a median band of deuters, an adaxial layer of hyalocysts, substereids or stereids, and abaxially layers of stereid or non-stereid cells, and an abaxial row of chlorocysts; abaxial side of the costa smooth, ridged or lamellose. Specialized asexual reproduction by brood leaves, microphyllous branches, deciduous leaves or stem tips or rhizoidal tubers. Sexual condition dioicous. Perichaetia terminal, often bud like, rarely pseudolateral; perichaetial leaves with a broader, sheathing base and a long, narrow subula. Seta 5–10 mm, those of young sporophytes curved downward, pushing the immature capsule between the comal leaves and leaving the calyptra behind when the mature capsule curves upward, sinuose, twisted, cygneous when wet and performing uncoiling movements. Capsule erect and symmetric or curved and asymmetric, sometimes strumose, furrowed when empty; annulus present but not dehiscent; operculum rostrate, half as long as the capsule; peristome teeth divided to the middle in two prongs, reddish or orange and horizontally striate proximally, hyaline and papillose distally. Calyptra cucullate, ciliate or entire at base. Spores ca. 13 µm, smooth or papillose.

Distribution

North America, Mexico, West Indies, Central America, South America, Europe, Asia, Africa, Atlantic Islands, Pacific Islands, Australia.

Discussion

Species ca. 180 (17 in the flora).

The North American species of Campylopus were revised by J.-P. Frahm (1980) based on a study of more than 1000 herbarium specimens. At that time, four species of Campylopus were recorded as new to North America and two species were placed into synonymy. The most recent checklist of the mosses of North America (L. E. Anderson et al. 1990) lists 18 species. Of these, the record of C. zygodonticarpus is based on a misidentification and C. paradoxus is a superfluous name for C. flexuosus, which is also included in the list. Since that time, C. japonicus has been newly recorded for North America, resulting in a total of 17 species, and new names have been introduced for C. aureus, C. japonicus, and C. schwarzii.

Campylopus was formerly divided into three subgenera on the basis of morphology of the transverse section of the costa. Although this classification is no longer used, the anatomy of the costa is still an important character for identification. It is, however, not in all cases necessary to prepare cross sections. The presence of adaxial stereids or hyalocysts can also be observed under the microscope by surface view of the adaxial side of the costa. Since the perichaetial leaves vary in both form and by the presence of thin-walled cells walls in species that usually have thick-walled cells, the study of such characters should be avoided. Alar cells are generally also not a valuable character in Campylopus; they are little differentiated in plants growing in damp habitats but are well developed in plants in exposed habitats with water uptake from the underground. The same is true for the presence or absence of a tomentum. Identification is facilitated if the ranges of the species are considered. Many species are found only very locally.

Excluded Species:

Campylopus zygodonticarpus (Müller Hal.) Paris

L. E. Anderson et al. (1990) in their most recent checklist of the mosses of North America listed this species, based on a specimen (Anderson 26656, DUKE) collected in Mississippi in 1992, consisting of C. tallulensis.

Campylopus subporodictyon (Brotherus) B. H. Allen & Ireland

Known in North America only from British Columbia, this species has recently been transferred to Campylopus from Dicranoweisia.

Selected References

Frahm, J.-P. and D. H. Vitt. 1978. A taxonomic study of Campylopus schimperi and C. subulatus in North America. Brittonia 30: 365–372.

Frahm, J.-P. 1980. Synopsis of the genus Campylopus in North America north of Mexico. Bryologist 83: 570–588.

Frahm, J.-P. 1994. A contribution to the differentiation of Campylopus subulatus and C. tallulensis in North America. Evansia 11: 95–99.

Lower Taxa

Campylopus angustiretis, Campylopus arctocarpus, Campylopus atrovirens, Campylopus carolinae, Campylopus flexuosus, Campylopus fragilis, Campylopus gracilis, Campylopus introflexus, Campylopus oerstedianus, Campylopus pilifer, Campylopus pyriformis, Campylopus schimperi, Campylopus schmidii, Campylopus sinensis, Campylopus subulatus, Campylopus surinamensis, Campylopus tallulensis

Key

1	Leaf tips ending in a hyaline hairpoint	> 2
1	Leaf tips concolorous	> 8
2	Hairpoints reflexed.	Campylopus introflexus
2	Hairpoints straight	> 3
3	Basal laminal cells thick-walled, chlorophyllose, subquadrate to short-rectangular (2:1)	> 4
3	Basal laminal cells thin-walled, hyaline, long-rectangular (more than 4:1)	> 5
4	Distal laminal cells vermicular.	Campylopus atrovirens
4	Distal laminal cells rectangular.	Campylopus sinensis
5	Abaxial side of costa lamellose with lamellae 3-4 cells high.	Campylopus pilifer
5	Abaxial side of costa smooth or ridged	> 6
6	Distal laminal cells oval to elongate oval.	Campylopus schmidii
6	Distal laminal cells rectangular to obliquely rectangular	> 7
7	Distal laminal cells 4-6:1, plants less than 1 cm, leaves 2.5-4 mm, transverse section of costa with adaxial stereids.	Campylopus carolinae
7	Distal laminal cells 1.5-2:1, plants to 3 cm, transverse section of costa with adaxial hyalocysts.	Campylopus oerstedianus
8	Leaf tips cucullate.	Campylopus atrovirens
8	Leaf tips plane	> 9
9	Basal laminal cell walls thick-walled, chlorophyllose	> 10
9	Basal laminal cells thin-walled, hyaline	> 13
10	Basal laminal cells pitted; transverse section of costa showing adaxial stereids	Campylopus arctocarpus
10	Basal laminal cells with smooth walls; transverse section of costa showing adaxial hyalocysts	> 11
11	Distal laminal cells 6-10:1; plants distantly foliate with spreading leaves.	Campylopus angustiretis
11	Distal laminal cells shorter; plants densely foliate with appressed leaves	> 12
12	Distal laminal cells short, subquadrate to rhombic; plants equally foliate	Campylopus flexuosus
12	Distal laminal cells short- to long-rectangular or oblique, 2-4(-6):1; plants in small rosettes or with appressed foliate leaves in a terminal comal tuft.	Campylopus surinamensis
13	Costa occupying 3/5-4/5 of leaf width.	Campylopus gracilis
13	Costa 1/2 of leaf width	> 14
14	Distal laminal cells rectangular, ca. 4:1	> 15
14	Distal laminal cells shorter	> 16
15	Plants in small rosettes; leaves long-subulate.	Campylopus pyriformis
15	Plants in dense cushions; leaves shortly pointed.	Campylopus schimperi
16	Leaves widest below mid leaf; basal and distal laminal cells sharply differentiated; specialized asexual reproduction frequently by boomerang-shaped brood leaves.	Campylopus fragilis
16	Leaves widest at leaf base; basal and distal laminal cells not sharply delimited; specialized asexual reproduction occasionally by deciduous stem tips	> 17
17	Costa in transverse section without abaxial stereids.	Campylopus subulatus
17	Costa in transverse section showing distinct groups of abaxial stereids.	Campylopus tallulensis