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BFNA Title: Campylopus |
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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. |
