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BFNA | Family List | BFNA Vol. 1 | Dicranaceae

Campylopus Bridel, Mant. Musc. 71. 1819.

Greek campylos curved, and pous, foot, for the curved seta

Jan-Peter Frahm

Plants 3--10 cm or more. 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, a adaxial layer of hyalocysts, substereids or stereids and abaxially layers of stereid or non-stereid cells and a 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. Spores ca. 13 µm, smooth or papillose. Calyptra cucullate, ciliate or entire at base.

Species ca. 180 (17 in the flora): largely tropical and warm temperate areas.

The North American species of this genus 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 number of 17 species, and new names have been introduced for C. aureus, C. japonicus and C. schwarzii.

The genus was formerly divided into three subgenera according to different 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 (see distribution maps). Many species are found only very locally.

Excluded Species:

L. E. Anderson et al. (1990) in their most recent checklist of the mosses of North America list C. zygodonticarpus (C. Müll.) Par. This record is based on an unpublished specimen (Anderson 26656, DUKE) collected in Mississippi in 1992, consisting of C. tallulensis.


Anderson, L. E., H. A. Crum and W. R. Buck. 1990. List of the Mosses of North America north of Mexico. Bryologist 93: 448--499. Arts, T. and J.-P. Frahm. 1990. Campylopus pyriformis new to North America. Bryologist 93: 290--294. Frahm, J.-P. 1980. Synopsis of the genus Campylopus in North America north of Mexico. Bryologist 83: 570--588. Frahm. J.-P. 1993. Campylopus japonicus new to North America north of Mexico. Bryologist 96: 142--144. Frahm, J.-P. 1994. A contribution to the differentiation of Campylopus subulatus and C. tallulensis in North America. Evansia 11: 95--99. 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. and P. L. Redfearn. 1981. Atractylocarpus costaricensis (C. Müll.) Bartr. new to North America North of Mexico, and Campylopus carolinae Grout, new to Florida. Bryologist 84: 226--228.

1 Leaf tips ending in a hyaline hairpoint.   (2)
+ Leaf tips concolorous.   (8)
2 (1) Hairpoints reflexed.   Campylopus introflexus
+ Hairpoints straight.   (3)
3 (2) Basal laminal cells thick-walled, chlorophyllose, subquadrate to short-rectangular (2:1).   (4)
+ Basal laminal cells thin-walled, hyaline, long-rectangular (more than 4:1).   (5)
4 (3) Distal laminal cells rectangular.   Campylopus sinensis
+ Distal laminal cells vermicular.   Campylopus atrovirens
5 (3) Abaxial side of costa lamellose with lamellae 3--4 cells high.   Campylopus pilifer
+ Abaxial side of costa smooth or ridged.   (6)
6 (5) Distal laminal cells oval to elongate oval.   Campylopus schmidii
+ Distal laminal cells rectangular to obliquely rectangular.   (7)
7 (6) Distal laminal cells 4--6:1. Plants less than 1cm, leaves 2.5--4 mm, transverse section of costa with adaxial stereids; white sand.   Campylopus carolinae
+ Distal laminal cells 1.5--2:1, plants to 3 cm, transverse section of costa with adaxial hyalocysts; earth-covered rocks.   Campylopus oerstedianus
8 (1) Leaf tips cucullate.   Campylopus atrovirens var. cucullatifolius
+ Leaf tips plane.   (9)
9 (8) Basal laminal cell walls thick-walled, chlorophyllose.   (10)
+ Basal laminal cells thin-walled, hyaline.   (13)
10 (9) Basal laminal cells pitted; transverse section of costa showing adaxial stereids.   Campylopus arctocarpus
+ Basal laminal cells with smooth walls; transverse section of costa showing adaxial hyalocysts.   (11)
11 (10) Distal laminal cells 6--10:1; plants distantly foliate with spreading leaves.   Campylopus angustiretis
+ Distal laminal cells shorter; plants densely foliate with appressed leaves.   (12)
12 (11) Distal laminal cells short, subquadrate to rhombic; plants equally foliate; soil and rocks, British Columbia and the Appalachian Mountains.   Campylopus flexuosus
+ 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; white sand, se United States.   Campylopus surinamensis
13 (9) Costa occupying 3/5--4/5 of leaf width.   Campylopus gracilis
+ Costa 1/2 of leaf width.   (14)
14 (13) Costa 1/2 of leaf width.   (15)
+ Distal laminal cells shorter.   (16)
15 (14) Plants in small rosettes on white sand in southeastern United States. Leaves longly subulate.   Campylopus pyriformis
+ Plants in dense cushions in arctic-alpine habitats. Leaves shortly pointed.   Campylopus schimperi
16 (14) Leaves widest below midleaf;. basal and distal laminal cells sharply differentiated; Specialized asexual reproduction frequently by boomerang shaped brood leaves.   Campylopus fragilis
+ Leaves widest at leaf base; basal and distal laminal cells not sharply delimited;. Specialized asexual reproduction occasionally by deciduous stem tips.   (17)
17 (16) Costa in transverse section lacking abaxial stereids.   Campylopus subulatus
+ Costa in transverse section showing distinct groups of abaxial stereids.   Campylopus tallulensis

  • List of lower taxa


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