BFNA Title: Campylopus
Author: J.-P. Frahm
Date: August 16, 2002
Edit Level: R Brum+
Version: 2a January 7, 2005

Bryophyte Flora of North America, Provisional Publication
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XX. CAMPYLOPUS  Bridel, Muscol. Recent., suppl. 4: 71.  1818  *  [Greek campylos, curved, and pous, foot, allunding to curved seta]

 

Jan-Peter Frahm

 

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. Spores ca. 13 µm, smooth or papillose. Calyptra cucullate, ciliate or entire at base.

 

Species ca. 180 (17 in the flora): a genus of tropical and warm temperate areas: North America, Central America, South America, Europe, Asia, Africa, Atlantic Islands, Pacific Islands, Australia.

 

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.

 

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

 

1. Leaf tips ending in a hyaline hairpoint.

2 Hairpoints reflexed . . . . 8. Campylopus introflexus

2. Hairpoints straight.

3. Basal laminal cells thick-walled, chlorophyllose, subquadrate to short-rectangular (2:1).

4. Distal laminal cells rectangular. . . . 14. Campylopus sinensis

4. Distal laminal cells vermicular. . . .  . . . . .3.  Campylopus atrovirens

3. Basal laminal cells thin-walled, hyaline, long-rectangular (>4:1).

5.Abaxial side of costa lamellose with lamellae 3--4 cells high  . . . 10. Campylopus pilifer

5. Abaxial side of costa smooth or ridged.

6. Distal laminal cells oval to elongate oval . . . . 13.Campylopus schmidii

6. Distal laminal cells rectangular to obliquely rectangular.

7. Distal laminal cells 4--6:1, plants < 1 cm, leaves 2.5--4 mm, transverse section of costa with adaxial stereids; white sand . . . .  . . . .  . . 4. Campylopus carolinae

7. Distal laminal cells 1.5--2:1, plants to 3 cm, transverse section of costa with adaxial hyalocysts; earth-covered rocks. . . .  . . . . .9. Campylopus oerstedianus

1. Leaf tips concolorous.

8. Leaf tips cucullate . . . . .3. Campylopus atrovirens var. cucullatifolius.

8. Leaf tips plane.

9. Basal laminal cell walls thick-walled, chlorophyllose.

10. Basal laminal cells pitted; transverse section of costa showing adaxial stereids.  . . 2. Campylopus arctocarpus

10. Basal laminal cells with  smooth walls; transverse section of costa showing adaxial hyalocysts.

11. Distal laminal cells 6--10:1; plants distantly foliate with spreading leaves . . . .  1. Campylopus angustiretis

11. Distal laminal cells shorter; plants densely foliate with appressed leaves.

12. Distal laminal cells short, subquadrate to rhombic; plants equally foliate; soil and rocks, British Columbia and the Appalachian Mountains. . . . 5. 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; white sand, se United States . . . . .16. Campylopus surinamensis

9. Basal laminal cells thin-walled, hyaline.

13. Costa occupying 3/5--4/5 of leaf width. . . . .7. Campylopus gracilis

13. Costa 1/2 of leaf width.

14. Distal laminal cells rectangular, ca. 4:1.

15. Plants in small rosettes on white sand in southeastern United States. Leaves long-subulate . . . . .11. Campylopus pyriformis

15. Plants in dense cushions in arctic-alpine habitats. Leaves shortly pointed. . . . .12. Campylopus schimperi

14. Distal laminal cells shorter.

16. Leaves widest below midleaf; basal and distal laminal cells sharply differentiated; specialized asexual reproduction frequently by boomerang-shaped brood leaves . . . . .6. 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. Costa in transverse section lacking abaxial stereids . . . 15. C. subulatus

17. Costa in transverse section showing distinct groups of  abaxial stereids . . . . . 17. Campylopus tallulensis

 

 

1. Campylopus angustiretis (Austin) Lesquereux & James, Man. 80.  1884

 

Dicranum angustirete Austin, Bot. Gaz. 4: 150.  1879; Campylopus delicatulus Williams; C. gracilicaulis subsp. angustiretis (Austin) Kindberg; Campylopus surinamensis var. angustiretis (Austin) J.-P. Frahm

 

Plants 5--20 mm, in loose light green to gray green mats, evenly foliate with distant, spreading leaves, the distal ones sometimes forming a comal tuft, not tomentose. Leaves 6 mm, lanceolate, keeled, long-decurrent, gradually narrowed into a fine channelled,  concolorous, straight tip that is denticulate at the outermost apex; alar cells large, inflated and auriculate, hyaline; basal laminal cells rectangular, moderately thick-walled, narrower and thin-walled in several marginal rows; distal laminal cells elongate, 6--10:1, not sharply delimited from the basal laminal cells; costa relatively narrow, filling 1/4--1/3 of leaf width, excurrent, in transverse section with a adaxial band of hyalocysts that are slightly smaller than the median deuters, abaxially with groups of stereids, smooth at back. Specialized asexual reproduction not known. Sporophyte not known.

 

Open sandy soil in wet depressions in coastal lowlands; 0--20 m; Fla.; West Indies; South America.

 

Campylopus angustiretis differs from all other species of the genus by its elongate distal laminal cells. Thus it is not certain if it actually belongs to this genus. On account of the lack of sporophytes a decision cannot be made. It has been treated as a variety of the sympatric C. surinamensis and superficially resembles certain expressions of that species (described as C. donnellii). It differs, however, by a narrower costa, the awn not coarsely serrate, the transverse section of the costa showing larger hyalocysts, and distinctly keeled leaves.

 

2. Campylopus arctocarpus (Hornschuch) Mitten, J. Linn. Soc., Bot. 12: 87.  1869

 

Dicranum arctocarpum Hornschuch in C. F. P. von Martius et al., Fl. Bras. 1: 12.  1840

 

Plants 1--3 cm, in dark green tufts, evenly foliate, stems reddish tomentose. Leaves 6--8 mm, erect patent when wet, crisped when dry, lanceolate, ending in a straight concolorous tip; alar cells inflated, hyaline or (in older leaves) reddish brown; basal laminal cells rectangular, thick-walled with pitted cell walls, ca. 3--6:1, narrower and thinner walled at margins, forming a small border; distal laminal cells subquadrate to oblique or rhombic, arranged in distinct rows; costa filling up to half of the leaf width, shortly excurrent in a dentate awn, in transverse section showing abaxial and adaxial stereid bands. Specialized asexual reproduction by microphyllous branches. Sporophytes absent in flora area.

 

Base of trees and decaying logs, cypress swamps; ca. 5 m; Fla.; Mexico; West Indies; Central America; South America; c Africa.

 

Campylopus arctocarpus much resembles C. flexuosus, to which it seems closely related, but is distinguished mainly by the different transverse section of the costa with adaxial stereids. Other examples of the phenomenon of "sister species" differing only by the transverse section of the costa are found in the genus.

 

3. Campylopus atrovirens De Notaris, Syllab. Musc., 221.  1838

 

Plants 1--10 cm, in tall tufts, blackish below, yellowish brown above. Leaves 4--10 mm, straight in wet and dry state, narrowly lanceolate, those of deciduous stem tips often even longer and narrower, ending in a very long subula; alar cells auriculate, hyaline or red-brown; basal laminal cells usually shortly rectangular to subquadrate, thick-walled with pitted walls, 4--8 rows of elongate hyaline cells at basal margins of leaves; distal laminal cells elongate oval to vermicular, walls incrassate; costa filling 1/2--2/3 of leaf width, excurrent in a long, straight, spinose-dentate hyaline hairpoint, in transverse section showing adaxial hyalocysts as wide as the median deuters, and abaxial groups of stereids, weakly ribbed at back. Specialized asexual reproduction by broken stem tips. Seta 4 mm. Capsule 1.5 mm, ovoid, brownish, operculum obliquely rostrate.

 

Varieties 2 (2 in the flora): North America; Europe; Asia.

 

Campylopus atrovirens is similar to C. sinensis, which differs by shorter, not vermicular distal laminal cells and shorter hyaline leaf tips. The latter has been found only once, in B.C., but it could be that collections of C. sinensis from the west coast of North America have been misidentified as C. atrovirens.

 

1. Leaves ending in a hyaline tip. . . . 3a. Campylopus atrovirens var. atrovirens

1. Leaves ending in a cucullate apex . . . . 3b. Campylopus atrovirens var. cucullatifolius

 

3a. Campylopus atrovirens var. atrovirens

 

Leaves gradually contracted into a long, fine point, hyaline at the extreme apex.

 

Wet rocks, damp cliffs, seepage banks, bogs or wet humic soil, always in open habitats at sea level along the coast, or at about 1500 m in the Appalachian Mountains; 0--1500 m; B.C., Nfld.; Alaska, N.C., Wash.; Europe; Asia.

 

The population in the Appalachian Mountains differs by greenish, not blackish  plants, smaller size, less developed alar cells and less incrassate, basal laminal cell walls. Such plants have been described from similar habitats and similar elevations from the Alps of Europe as C. adustus De Notaris. It is not known whether these populations in non-coastal areas are genotypically different or just modifications associated with higher elevations. It may perhaps deserve to be recognized at the varietal rank. Forms with falcate leaves as occurring in Europe or Asia have not yet been found in North America. Sporophytes, produced very rarely, were found in North America only once, in B.C.

 

3b. Campylopus atrovirens var. cucullatifolius J.-P. Frahm, Bryologist 83: 574.  1980

 

Leaves with concolorous, cucullate tips.

 

Wet rocks; elevation not known (mountain); Alaska; Europe; China.

 

In North America this variety is known only from the type locality on Kuju Island, Alaska, and otherwise only from three collections from Great Britain and China. Cucullate instead of hyaline tipped leaves are found in a number of species of this genus.

 

4. Campylopus carolinae  Grout, Moss Fl. N. Amer. 1: 249.  1939

 

Plants usually less than 1 cm, in loose mats, dark green to brownish green or blackish; leaves erect patent; stems sparsely tomentose. Leaves 2.5--4 mm, small, lanceolate, ending in a concolorous straight tip, convolute in the distal part, with entire margins; alar cells not or only slightly differentiated; basal laminal cells rectangular, firm walled, hyaline, 2.5--3.5:1, indistinctly bordered at margins; distal laminal cells oblique to oval, incrassate, ca. 3--5:1; costa filling 1/3 of leaf width, excurrent in a straight, toothed, hyaline point, in transverse section showing abaxial and adaxial stereids, ridged abaxially with prominent cells. Specialized asexual reproduction occasionally by means of deciduous stem tips. Sporophytes not known in area of the flora.

 

Typically buried in white sand in depressions, in open pine and pine-oak forests and open grassland, coastal lowlands; low elevations; Fla., N.C.; South America (Brazil). 

 

The disjunction of western South America--southeastern North America is also found for Campylopus angustiretis, C. surinamensis and C. pyriformis, which grow in similar habitats in white sand. The type material from Brunswick, North Carolina. is mixed with C. surinamensis, which caused confusion and recognition of this species as a variety of C. delicatulus Williams (= C. angustiretis).

 

5. Campylopus flexuosus (Hedwig) Bridel, Muscol. Recent., suppl. 4: 71.  1819

 

Dicranum flexuosum Hedwig, Spec. Musc. Frond. 145.  1801; Campylopus paradoxus Wilson

 

Plants in dense, 1--3 cm, dark green mats, usually reddish tomentose below. Leaves 5--7 mm, erect patent when wet, flexuose when dry, the distal leaves sometimes curved and secund, lanceolate, ending in a straight concolorous tip, which is serrate in the distal part; alar cells hyaline or reddish; basal laminal cells thick-walled, rectangular, ca. 4--5:1, narrower toward the margins; distal laminal cells quadrate to oblique or short rhombic; costa filling 1/2--2/3 of leaf width, in transverse section showing abaxial groups of stereids and adaxial small substereidal hyalocysts which are smaller than the median deuters. Specialized asexual reproduction by microphyllous branches in the axils of the distal leaves.  Sporophytes not known in North America.

 

Rocks, humus covered boulders and outcrops, also humic or peaty soil; coastal lowlands in B.C., 1500 m in N.C.; B.C.; N.C.; Central America; South America; Europe; c Africa; Asia (China).

 

Campylopus flexuosus has been only found in a few localities in British Columbia and a single locality in the Appalachian Mountains. The occurrences in East Asia and B.C. may be interpreted as relictual from the Tertiary, from which area C. flexuosus was---in contrast to Europe---not able to spread after the Pleistocene. The only record from the Appalachian Mountains on Flat Rock, Blue Ridge Parkway, is difficult to explain because many similar habitats exist near that vicinity in which the species has not been found. Before 1980, all specimens from North America, except for three labelled as C. flexuosus, belonged in fact to C. tallulensis or rarely to C. surinamensis. Campylopus flexuosus, however, differs from C. tallulensis by thick-walled, chlorophyllose basal laminal cells and small adaxial hyalocysts and in appearence by dark green color. Campylopus tallulensis has hyaline thin-walled basal laminal cells, large adaxial hyalocysts (even visible in surface view of the costa) and commonly a golden yellowish color. Campylopus surinamensis has longer distal laminal cells and the costa ends in a strongly dentate often subhyaline awn.

 

6. Campylopus fragilis (Bridel) Bruch & Schimper, Bryol. Europ. 1: 164.  1847

 

Dicranum fragile Bridel, J. Bot. (Schrader) 1800: 296.  1801

 

Plants 0.5--2 cm, yellowish green, in tufts, very densely foliate, often ending in a penicillate comal tuft, whitish tomentose,. Leaves 4--5 mm, ovate lanceolate, widest below the middle and contracted at base, narrowed into a more or less long straight, concolorous subula; alar cells hardly developed; basal laminal cells thin-walled and hyaline, very distinctly differentiated from the distal thick-walled and quadrate laminal cells; costa filling 1/2--2/3 of leaf width, shortly excurrent, slightly serrate at tip, in transverse section showing very wide adaxial hyalocysts (easily recognizable in surface view of the costa), filling half of the width of the costa, and abaxial substereids, slightly abaxially ridged. Specialized asexual reproduction by small, boomerang-shaped leaves produced in the axils of the distal leaves. Sporophytes not known in North America.

 

Rocks, humus covered boulders and outcrops, also humic or peaty soil; 0--200 m; Ark.; Central America; South America; w Europe; Asia  (China, Japan); c Africa; Atlantic Islands (Azores, Madeira, Canary Islands).

 

Campylopus fragilis has only been found in a few localities in B.C. and a single locality in Ark. It has been reported from southeastern North America based on the presence of boomerang-shaped brood leaves at the stem tips, however, these records belong to C. surinamensis with similar brood leaves. Campylopus surinamensis differs by rectangular and not quadrate distal laminal cells, a sharply toothed, denticulate, excurrent costa and small adaxial hyalocysts. Brood leaves are produced especially "under stress" in unfavorable conditions, especially on small, depauperate plants.

 

7. Campylopus gracilis (Mitten) Jaeger, Ber. Thätigk. St. Gallischen Naturwiss. Ges. 1870--1871: 427.  1872

 

Dicranum gracile Mitten, J. Linn. Soc., Bot., suppl. 1: 17.  1859; Campylopus schwarzii Schimper

 

Plants 1--8 cm, glossy yellowish green or golden green, not tomentose. Leaves 5--8 mm, erect  when wet, appressed when dry, rarely falcate, narrowly lanceolate, ending in a long and fine, concolorous subula; alar cells conspicuous, hyaline, projecting into the costa; basal laminal cells hyaline, thin-walled, long-rectangular, lamina very short, ending at midleaf, distal laminal cells short-rectangular or oblique, very small; costa very broad, occupying 3/4--4/5 of leaf width, long-excurrent in a fine almost entire subula, in transverse section showing very large, lax adaxial hyalocysts filling half of the costa thickness and no abaxial stereids, faintly abaxially ridged. Specialized asexual reproduction by deciduous leaves or broken leaf tips. Sporophyte not known.

 

Wet soil and soil covered rocks, wet cliffs in boggy slopes in subalpine habitats; low elevations; B.C. (Queen Charlotte Islands and adjacent mainland); w Europe; e Asia .

 

Campylopus gracilis is easily recognized by a very broad costa, occupying 3/4 or more of the leaf width, very small shortly rectangular, distal laminal cells and large auricles projecting distinctly into the costa. In Europe, the species is found in similar situations in the highly oceanic parts as in the west coast of North America but also (as in C. atrovirens) in the Alps.

 

8. Campylopus introflexus (Hedwig) Bridel, Muscol. Recent., suppl. 4: 72.  1818

 

Dicranum introflexum Hedwig, Sp. Musc. Frond., 141. 1801

 

Plants 0.5--5 cm, in dense mats, yellowish to olive green, tomentum present or almost absent. Leaves 4--6 mm, erect patent when wet, appressed when dry, lanceolate, straight, with entire margins; alar cells lacking or formed by thin-walled, hyaline to reddish, inflated cells; basal laminal cells hyaline rectangular, thin-walled, extending higher at margins and forming a V-shaped area; distal laminal cells incrassate, shortly rectangular to oblique, chlorophyllose; costa filling 1/2--3/4 of leaf width, excurrent in a hyaline hair tip, which is conspicuously 90° reflexed, in transverse section showing adaxial hyalocysts and abaxial stereids, shortly lamellose at back with ribs 1--2 cells high. Specialized asexual reproduction occasionally by deciduous stem tips. Seta 7--12 mm, yellowish brown to brownish in age, often several sporophytes from the same plant, curved or sinuose. Capsule brown, 1.5 mm, slightly asymmetric and curved when empty. Calyptra ciliate at base. Spores 12--14 µm.

 

Soil along trails, base of trees, flat roofs of buildings, peat in bogs, sand; 0--200 m; B.C.; Calif, Oreg., Wash.; South America (Chile, Argentina, se Brazil); Europe; s Africa; Australia; Pacific Islands (New Caledonia, subantarctic Islands, New Zealand).

 

Campylopus introflexus occurs in masses in sand dunes along the west coast of North America and throughout the Southern Hemisphere. The species was introduced in Great Britain in 1942, and since the beginning of the 1970's has been aggressively spreading through Europe. It now ranges from Iceland to Spain and from Ireland to Poland. The first record in North America dates from August, 1975, and was made on a gravel roof of a building of Humboldt University, Arcata, California. The species is undoubtedly introduced in North America and is spreading as rapidly as in Europe. The name C. introflexus was used previously for C. pilifer, thus all old references for C. introflexus in North America have to be referred to that species. Also, specimens of C. surinamensis and C. oerstedianus from North America were named as C. introflexus. Campylopus introflexus is easily recognized by the reflexed hair points. Female plants have terminal perichaetial buds. Problems may rarely arise with forms from shaded habitats, in which the hairpoints are lacking or so short that they are not reflexed.

 

9. Campylopus oerstedianus (J. K. A. Müller) Mitten, J. Linn. Soc., Bot. 12: 81.  1869

 

Dicranum oerstedianum J. K. A. Müller, Syn. Musc. Frond. 2: 596.  1851

 

Plants 1--3 cm, in olive green tufts, lighter above and brownish below, evenly foliate, tomentose. Leaves 4--5 mm, lanceolate, gradually narrowed into a subtubulose, straight, concolorous subula; alar cells slightly differentiated, reddish or hyaline; basal laminal cells hyaline, rectangular; distal laminal cells thick-walled, subquadrate to short-rectangular or oblique; costa filling half of the leaf width, excurrent in a short, hyaline tip, which is longer in perichaetial leaves, in transverse section showing adaxial hyalocysts and abaxial stereids in groups of 2 cells, abaxially ridged. Specialized asexual reproduction not seen. Sporophytes unknown.

 

Soil covered rocks; ca. 50 m; N.C.; Central America (Costa Rica); West Indies (Jamaica); Europe (France, Germany, Greece, Italy).

 

Campylopus oerstedianus has been found only once in the flora area, in the piedmont of North Carolina. The overall distribution is very scattered and suggests a circum-Tethyan range (margins of the Caribbean and Mediterranean seas). It has been described from Europe as C. mildei Schimp. Plants of C. oerstedianus resemble C. pilifer in habit, with shorter hairpoints. In shady habitats the hairpoints are sometimes lacking. The plants are microscopically distinguished by the slightly different shape of the distal laminal cells and the transverse section of the costa, by the lack of abaxial lamellae on the costa, smaller adaxial hyalocysts of about the diameter of the median deuter cells, and groups of abaxial stereids with only 2 instead of 4 stereid cells.

 

10. Campylopus pilifer Bridel, Muscol. Recent., suppl. 4: 72. 1819

 

Campylopus leucotrichus Sullivant & Lesquereux

 

Plants 0,5--3 cm long, in tufts, dirty green, olive green or yellowish green, darker below, equally foliate, the fertile ones comose,. Leaves 4--7 mm, erect spreading or loosely appressed, lanceolate, ending in a straight, more or less long serrate hairpoint; alar cells not differentiated or strongly developed, inflated, thin-walled, hyaline or reddish; basal laminal cells hyaline, thin-walled, rectangular, forming a V-shaped area; distal laminal cells oval to rhomboidal, ca. 2:1; costa filling 1/2--3/4 of leaf width, excurrent in a hairpoint, in transverse section showing adaxial hyalocysts and abaxial groups of stereids, abaxially with lamellae 3--4 cells high. Specialized asexual reproduction occasionally by deciduous stem tips. Setae often aggregated, about 5 mm, sinuose. Capsule 1.5 mm, slightly asymmetric, furrowed when dry, brownish; operculum rostrate. Spores ca. 13 µm. Calyptra fringed at base.

 

Acidic sandy soil and acidic rocks (sandstone, granite), rock crevices, exposed, dry habitats; 50--1500 m; Ala., Ariz., Ark., Ga., La., N.C., S.C., Tenn., Tex.; Central America; South America; sw Europe; c Africa; Asia (Sri Lanka).

 

The distribution of Campylopus pilifer in tropical America, tropical Africa, and Sri Lanka (but not other parts of Asia) suggests a Gondwanaland origin, from where the species has extended its range into warmer parts of North America and southwestern Europe. Until 30 years ago this species was not distinguished from C. introflexus, and accordingly all old references from North America must be referred to C. pilifer. The true C. introflexus has been a neophyte in North America since 1975.

 

11. Campylopus pyriformis (Schultz) Bridel, Bryol. Univ.  1: 469.  1826

 

Dicranum pyriforme Schultz, Prodr. Fl. Starg., suppl. 73.  1819

 

Plants 3 mm, gregarious or in loose, low tufts, forming low rosettes, appearing stemless, light to olive green. Leaves 3 mm, erect-patent, flexuose when dry, from lanceolate base gradually contracted into a long, fine, straight, concolorous, distinctly canaliculate subula; margins serrate in the distal part of the leaves; alar cells scarcely differentiated; basal laminal cells hyaline, thin-walled, rectangular; distal laminal cells thick-walled, rectangular, ca. 4:1; costa filling 1/2--2/3 of leaf width, excurrent, in transverse section with large, empty, adaxial hyalocysts and abaxial groups of stereids, abaxially smooth. Specialized asexual reproduction by colorless, multicellular, long cylindric rhizoidal tubers, 300--700 µm long, deciduous leaves and small brood leaves produced at stem tips. Sporophytes not present in North America.

 

Bare soil, also base of trees and old pine stumps in wet acid meadows and swamp forests; 0--50 m; Fla., La., Miss.; s South America (Argentina, se Brazil, Chile);  w Europe; Asia (China); c and s Africa, Atlantic Islands (Azores, Madiera, s Iceland); Australia; Pacific Islands (New Zealand, New Caledonia).

 

The description above refers to North American plants of Campylopus pyriformis---specimens from other parts of its range have a somewhat different appearance. This species was first recorded for North America (T. Arts and J.-P. Frahm 1990) based on collections made by W. D. Reese. The occurrence in North America at only three localities in Louisiana and Mississippi, and an additional unpublished record from Florida, can perhaps be explained by introduction facilitated by the presence of rhizoidal tubers. It may therefore be doubted whether this species is native in North America. However, the small form found in the United States resembles a form occurring in Brazil in similar habitats, from which area it may have been introduced by birds. Similar disjunctions between Brazil and se North America are also found in C. surinamensis, C. carolinae and C. angustiretis, which all conspicuously grow together on bare, acid, white sand. Campylopus pyriformis was also found mixed with C. surinamensis, but can be distinguished by the more elongate, narrowly lanceolate leaves with a channelled apex, a long-excurrent nerve and a lamina ending at midleaf and colorless rhizoidal tubers instead of the reddish or reddish brown ones as in C. surinamensis.

 

12. Campylopus schimperi Milde, Bot. Zeitung (Berlin) 22: 13.  1864

 

Campylopus subulatus var. schimperi (Milde) Husnot

 

Plants 1--3 cm, in compact tufts, light green above, brownish and tomentose below. Leaves 2,5--5 mm, appressed, from a narrow base gradually contracted to an acute, straight, concolorous subula; alar cells little differentiated, only slightly wider than the basal laminal cells; basal laminal cells thin-walled, hyaline, rectangular, at margins narrower, forming a small band; distal laminal cells chlorophyllose, rectangular, ca. 4:1; costa filling 1/2--2/3 of leaf width, shortly excurrent,in transverse section showing large adaxial hyalocysts, abaxial stereids absent, almost smooth at back. Specialized asexual reproduction frequently by deciduous stem tips. Sporophytes not known in North America.

 

Soil in tundra habitats; in alpine elevations, 2750--3400 m in Colo.; Greenland; B.C., Nfld., Yukon; Alaska, Colo.; Europe; Asia  (Bhutan, China, Japan, Nepal).

 

Records of Campylopus schimperi from Mexico and the Andes are doubtful. The taxonomic value of this species has often been doubted. It was frequently regarded as a variety of C. subulatus, which is similar in several respects. Campylopus schimperi, however, has more elongate, distal laminal cells (1:4 instead of 1:1.5--2) and abaxially nearly smooth costa (ridged in C. subulatus). Furthermore, C. schimperi is a species of arctic-alpine habitats and C. subulatus is found at low elevations. These anatomical as well as ecological differences indicate that C. schimperi should be regarded as a separate species.

 

13. Campylopus schmidii  (J.K.A. Müller) A. Jaeger, Thätigk. St. Gallischen. Ber. Naturwiss. Ges. 1870--71: 439.  1872

 

Dicranum schmidii J.K.A. Müller, Bot. Zeitung (Berlin) 11: 37.  1853; Campylopus aureus Bosch & Sande La Coste

 

Plants 2--5 cm, yellowish green, stiff, evenly foliate, the perichaetia in comal tufts. Leaves 5--6 mm, erect patent when wet, appressed when dry, from oblong base gradually contracted to a long subula, ending in a straight, hyaline, serrate tip; alar cells hardly differentiated; basal laminal cells thin-walled, rectangular, hyaline; distal laminal cells chlorophyllose, oval to narrow or elongate oval, incrassate; costa filling 1/2--2/3 of leaf width, in transverse section showing adaxial hyalocysts as large as the median deuter cells, and abaxial groups of stereids, ribbed abaxially. Specialized asexual reproduction by deciduous buds produced in the distalmost part of the stem. Sporophytes not known from North America.

 

Soil in open pine, cedar and cypress forests; 80--200 m; Calif., Oreg.; Mexico; Asia (China, s India, Sri Lanka, Java, Sulawesi, Borneo, Taiwan); Africa (Madagascar, c Africa); Pacific Islands (Hawaii); n Australia.

 

Campylopus schmidii does not fruit in North America, where apparently only female plants exist. The range of this species is mainly southeastern Asia. From there it extends south to Queensland, west to Madagascar and Central Africa, east to Hawaii, California, Oregon, and Mexico. In California it is known from two localities, in Oregon from one, and in Mexico from one, which suggests that the occurrence of C. schmidii may result from occasional long distance dispersal events across the Pacific Ocean; it may not be native in North America. The first collection was made in California in 1933. Plants of C. schmidii resemble C. pilifer but are distinguished by elongate-oval rather than oval distal laminal cells and costa smooth at the abaxial surface and not with lamellae 3--4 cells high as in C. pilifer. Campylopus introflexus has lamellae 2 cells high and a similar areolation as C. pilifer but is distinguished in the field by reflexed hairpoints. All three species are more or less vicariant sister species, C. introflexus in the subantarctic to subtropical parts of the southern hemisphere, C. pilifer in tropical India, Africa and South America and from there extending to southeastern North America and southwestern Europe, and C. schmidi mainly in southeastern Asia.

 

14. Campylopus sinensis (J.K.A. Müller) J.-P. Frahm, Ann. Bot. Fenn. 34: 202.  1997

 

Dicranum sinense J.K.A. Müller, Nuovo Giorn. Bot. Ital., n. s. 4: 249.  1897; Campylopus japonicus Brotherus; Dicranodontium sinense (J.K.A. Müller.) Paris

 

Plants to 3 cm, in dense tufts, blackish below, golden green above. Leaves 5--10 mm, the distal ones longest, erect patent when wet, appressed when dry, narrowly lanceolate, long-subulate, ending in a straight, fine, almost entire apex, piliferous at least in the distalmost leaves and plants from exposed habitats, rarely subhyaline; alar cells reddish brown, inflated; basal laminal cells thick-walled, rectangular, narrower at margins, thin-walled in perichaetial leaves; distal laminal cells shortly rectangular or oblique, 3--5:1; costa filling 1/2--3/4 of leaf width, excurrent, in transverse section showing abaxial groups of stereids and adaxial firm walled hyalocysts, slightly abaxially ridged. Specialized asexual reproduction by deciduous stem tips or deciduous leaves. Sporophytes not known from the flora area.

 

Usually on soil and rocks; ca. 60 m; B.C.; Mexico; Asia (China, Japan, Korea, Vietnam); Pacific Islands (Tahiti); Australia (Queensland).

 

In North America north of Mexico Campylopus sinensis has been found only once, in a depauperate condition in a blanket bog in the Queen Charlotte Islands, B.C. The species shows a distinct gradient from large to small plants in the tropical to the subtropical or temperate-oceanic parts of its range in East Asia, which seems to be matched also for the North American populations with regard to specimens from Mexico and from British Columbia.  It is not evident whether the record from Queen Charlotte Islands is the result of a long distance dispersal or a relict from the Tertiary, as supposed from some other bryophyte species with amphi-Pacific range or disjunct occurrence in East Asia and Mexico. It is also possible that C. sinensis was hitherto overlooked in North America and (as frequently in China) confused with the similar C. atrovirens (for differences see discussion under the latter species).

 

15. Campylopus subulatus  Schimper in G. L. Rabenhorst and G. Winter, Bryotheca Eur. 9: no. 451.  1861

 

Plants 0.5--3 cm, in loose, slender tufts, yellowish green to green, not tomentose. Leaves 3--4 mm, erect patent when wet, appressed when dry, lanceolate, narrowed into a short, straight subula; margins entire below, faintly serrate at apex; apex of leaf serrate at back; alar cells hardly differentiated, only slightly larger than the basal laminal cells; basal laminal cells thin-walled, hyaline, rectangular; distal laminal cells short, subquadrate; costa filling 1/2--2/3 of leaf width, excurrent in a short concolorous apex, in transverse section showing adaxial hyalocysts that are 1/3 as wide as the costa, without abaxial stereids, ribbed at back.  Specialized asexual reproduction by deciduous stem tips. Sporophytes not known in North America [rare elsewhere].

 

Open soil in oak and Douglas fir forests, also open sand in dunes with Pinus contorta; 80--200 m; Calif., Oreg.; Europe; Asia.

 

Campylopus subulatus is known only from two localities in California and one in Oregon. Although all records of C. subulatus from North America were referred to C. schimperi by J.-P. Frahm and D. H. Vitt (978), collections made later in California and Oregon proved to be this species. Campylopus schimperi grows in compact tufts in alpine habitats and differs from C. subulatus by an abaxially smooth costa, and rectangular, not subquadrate distal laminal cells. Campylopus subulatus resembles much C. tallulensis. The latter differs by distinct groups of  abaxial stereids and adaxial hyalocysts, which are 1/2 as wide as the thickness of the costa.

 

16. Campylopus surinamensis J. K.A. Müller, Linnaea 21: 186.  1848

 

Campylopus donnellii (Austin) Lesquereux & James; C. gracilicaulis Mitten; C. gracilicaulis var. donnellii (Austin) Grout; C. tallulensis var. subleucogaster (J. K. A. Müller) Grout; Dicranum donnellii Austin

 

Plants 0.5--3 cm, in loose tufts, light green, young plants forming low rosettes, older plants taller, with a stem arising from the rosette with densely appressed leaves, ending in a comal tuft that consists of perichaetia or produces brood leaves. Leaves 4--7 mm, narrow lanceolate, narrowed to a short (in stem leaves) or long (in rosette and comal leaves) serrate subula; alar cells sometimes well developed, forming reddish or hyaline auricles, sometimes not much differentiated; basal laminal cells of appressed stem leaves more or less thick-walled, of comal leaves thin-walled, narrower at margins, forming an indistinct small border; distal laminal cells short to long-rectangular or oblique, 2--5:1; costa filling half of the leaf width, excurrent in a serrate awn that is subhyaline in the comal leaves, in transverse section with adaxial hyalocysts and abaxial groups of stereids, abaxially ribbed. Specialized asexual reproduction by small hooked or boomerang-shaped leaves (similar to those of C. fragilis) in axils of comal leaves. Sporophytes not seen in North America.

 

Open, acidic, sandy soil in sandhills or open forests; 0--50 m; Ala., Fla., Ga., La., N.C.; West Indies (Cuba, Trinidad); Central America (Honduras); South America.

 

This species was named Campylopus gracilicaulis in North America before the identity with the South American C. surinamensis was determined. As expressed in the description, the rosette leaves, appressed stem leaves and comal leaves have a different shape, have different, short or long, concolorous or subhyaline leaf tips and different basal laminal cells. This and the variability of the plant morphology have caused confusion. The almost hairpointed apices of the comal leaves have led to identification as C. introflexus, the production of brood leaves to identification as C. fragilis. The confusion increased when Bartram transferred C. gracilicaulis and C. donnelli to C. flexuosus, as he did with C. subleucogaster. The latter was regarded as a distinct species by Williams but placed as a variety of C. tallulensis by Grout. The type specimen of C. donnellii does not differ from those of C. surinamensis or C. gracilicaulis. However, this name has frequently been used for forms of C. surinamensis without comal tufts or consisting only of small rosettes

 

17. Campylopus tallulensis Sullivant & Lesquereux, Musci Bor.-Amer. ed. 2, 17.  1865

 

Plants to 5 cm, in tufts, yellowish green, rarely green. Stems slender, not or densely reddish tomentose, evenly foliate. Leaves about 5 mm, erect spreading, lanceolate, narrowed to a straight, serrate tip; alar cells hardly differentiated, forming hyaline or reddish auricles; basal laminal cells hyaline, thin-walled, rectangular, often forming a V-shaped area; distal laminal cells short-rectangular, incrassate; costa filling half of the leaf width, shortly excurrent in a concolorous tip, in transverse section showing large adaxial hyalocysts occupying 1/2 of the thickness of the leaf, and abaxial groups of stereids, abaxially ridged. Specialized asexual reproduction by deciduous leaves or stem tips. Sporophytes not known.

 

Acidic rocks (granite, sandstone), exposed boulders, rarely on soil in open woods; 100--600 m; Ala., Ariz., Ark., Del., Ga., Ill., La., Miss., N.C., Ohio, S.C., Tenn., Va., Wyo; Mexico; Central America (Nicaragua); South America (Bolivia, Colombia, Peru, Venezuela)

 

The disjunction of Campylopus tallulensis from southeastern North America to Mexico, which is also met in other bryophytes and flowering plants, is considered to be a result of a former continuous range in the Tertiary. Campylopus tallulensis was included in C. flexuosus by American authors. There is a superficial similarity regarding the habit and the shape of the distal laminal cells. Campylopus flexuosus is, however, easily distinguished by thick-walled basal laminal cells and the presence of microphyllous brood branches. Plants of C. tallulensis from Mexico and eastern North America are robust and yellowish to golden green. In contrast, the specimens collected in Illinois, Mississippi (in part) and Arkansas are more slender and dark green, resembling C. subulatus in appearance. It is not known whether these differences in color depend on a different geological substrate or are the expression of different populations. Both species are anatomically very similar with thin-walled hyaline basal laminal cells, almost quadrate distal laminal cells, a costa excurrent in a sometimes subhyaline point and being roughened at the abaxial side like a rat's tail file and a channeled leaf apex. The only way to distinguish both species seems to be the transverse section of the costa, which shows very distinct groups of abaxial stereids in C. tallulensis but no abaxial stereids in C. subulatus. Furthermore, the adaxial hyalocysts of C. tallulensis are twice as wide as those of C. subulatus (J.-P. Frahm 1994). On the basis of this character, the only records of C. subulatus in North America from California belong to this species and are not extensions of the range of C. tallulensis from Mexico.

 

Excluded Species:

L. E. Anderson et al. (1990) in their most recent checklist of the mosses of North America list Campylopus zygodonticarpus (J. K. A. Müller) Paris.  This record is based on a specimen (Anderson 26656, DUKE) collected in Mississippi in 1992, consisting of C. tallulensis.

 

OTHER REFERENCES

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.