BRYOLOGIE
LICHENOLOGIE
TOME 2 Fascicule 4 1981
M6 182
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Copyright © 1981, CRYPTOGAMIE Bryologie Lichénologie
Source : MNHN, Paris.
G 6 1 03
CRYPTOGAMIIE
BRYOLOGIE
LICHÉNOLOGIE
TOME2 Fascicule 4 1981
COMITÉ DE LECTURE
Bryologie : J.L. DE SLOOVER, P. GEISSLER, S.R. GRADSTEIN, J.P. HÉBRARD, S.
JOVET-AST, D. LAMY.
Lichénologie : J. ASTA, B. BODO, W.L. CULBERSON, М.С. JANEX-FAVRE, J. LAM-
BINON, M.A. LETROUIT-GALINOU.
MANUSCRITS
Les auteurs sont priés d'adresser leurs manuscrits á la Rédaction de CRYPTOGAMIE
Bryologie et Lichénologie, Laboratoire de Cryptogamie, 12 rue de Buffon, 75005 Paris
Les tirages à part sont à la charge des auteurs
Ouvrage publié le concours du Centre National de la Recherche Scientifique
Dani мар вува ае M. "59062 - MNHN. Paris
379
DIDYMODON (POTTIACEAE) IN MEXICO AND CALIFORNIA :
TAXONOMY AND NOMENCLATURE
OF DISCONTINUOUS AND NONDISCONTINUOUS ТАХА!
В.Н. ZANDER *
ABSTRACT. — Eight species of Didymodon (Pottiaceae, Bryopsida), including thirteen
varieties, are recognized for Mexico and California. New combinations include Didymodon
sect. Asteriscium (C. Muell.), D. rigidulus var. gracilis (Schleich. ex Hook. & Grev.), D. т. var.
icmadophila (Schimp. ех С. Muell.), D. г. var. subulatus (Thér. & Bartr, ex Bartr.), D. austra-
lasiae var. umbrosus (C. Muell.), D. vinealis var. brachyphyllus (Sull. in Whipple), D. v. var.
luridus (Hornsch. in Spreng), D. v. var. nicholsonii (Culm.) and D. v. var. rubiginosus
(Mitt.). Thirty-five taxa are new synonyms. Didymodon vinealis var. nicholsonii is reported
new to the New World. Unusually large concepts of certain taxa are presented, Whenever
the names of these taxa are used by future bryologists, these and narrower taxonomic
concepts of other authors should be clearly distinguished by indicating the concept «sensu»
or «emendavit» a particular author and publication as part of the citation of the scientific
name for precision in biological applications. An авупопупис method is used to provide
scientific names for nondiscontinuous infraspecific taxa. Certain problems in typification
of bryophytes are largely due to ease of making isotypes and of unmaking holotypes and
lectotypes by past (and future) division of type specimens consisting of more than one
gametophore.
The genus Didymodon Hedw. has long been a problem for bryologists be-
cause it is only poorly distinguished from Barbula Hedw. The species included
in Didymodon are also often difficult to distinguish from each other, both
because of apparent intra- and interspecific variation in morphology and because
laminal papillae obscure the areolation (a character important to the «look» of
a taxon under the microscope) in many species (one competent bryologist
jocularly refers to gametophores of desert species of Pottiaceae as «black dot»,
anon., pers. comm.). I have discussed the differences between Didymodon and
1. Lam grateful to the staff of the British Museum (Natural History), the Linnean Society
of London, and the Conservatoire et Jardin botaniques, Genève, for initial permission and
subsequent aid in study of the original collections of early botanists. Thanks are given го
the curators of the herbaria mentioned in the text for loan of specimens. Р.М. Eckel did
the illustrations. This study was partially supported by a grant from the American Philoso-
phical Society to the author.
* Clinton Herbarium, Buffalo Museum of Science, Buffalo, NY 14211, USA.
Cryptogamie, Bryol. Lichenol., 1981, 2, 4 : 379-422.
Source : MNHN, Paris
380 R.H. ZANDER
Barbula previously (ZANDER 1978a) and here attempt to render some solution
to the former problem by recognizing only relatively discontinuous taxa as
species (i.e. basic taxonomic units) and providing a nomenclatural system
for dealing with nondiscontinuous taxa, and the latter problem by urging the use
of Hoyers Solution (ANDERSON 1954) rather than water for mounting
specimens because it clarifies areolation.
The area of study is all of Mexico and the state of California in the U.S.A.
Major floristic studies of these areas are by CRUM (1951) for the former and
KOCH (1950) for the latter. Major regional taxonomic studies that include
treatments of many of the species discussed here are by GROUT (1928-1940),
STEERE (1938), BARTRAM (1949), LAWTON (1971) and FLOWERS (1973).
This study was done to provide input of a specialist for the proposed moss
floras of Mexico (ed. A.J. SHARP and H. CRUM, in prep.) and of California
(D. NORRIS, in prep. The sections labeled «Distribution» given with the
discussions of taxa below refer only to Mexico and California. The descriptions
are based on examined specimens from several herbaria :notably BM, BUF, FH,
MICH, NY, PC, SPA, TENN and US. Complete lists of specimens examined are
given for the few relatively rare taxa. The chromosome numbers cited are
taken from the literature.
Both SAITO (1975) and ZANDER (1978a) assert that peristomes in Didymo-
don are rather variable in length in some species. Such statements usually refer
only to the peristome of the deoperculate capsule. Although specimens with
short opercula have short peristomes, some specimens with long opercula have
long, but highly fragmented peristome teeth when observed in cleared operculate
capsules — these teeth would appear short when the operculum, together with
the distal peristome fragments, is removed. A morphological feature not pre-
viously seen in Didymodon is the presence of transverse slits in the medial
basal portion of the leaves on both sides of the costa of some specimens of D.
australasiae var. umbrosus. Such slits — resorption channels reaching across
several cells and perforating the leaf — have been previously noted in Kingio-
bryum paramicola Robins. (Dicranaceae) (ROBINSON 1967; ZANDER &
CLEEF, in prep.) and simply perforate basal cells have been described for a few
Pottiaceae species by PROCTOR (1979).
The systematic framework presented here is a model reflecting what are
thought to be natural relationships of taxa inferred from observations on a series
of specimens. There are no arbitrary limits given to the taxa so future workers
may easily test the model and perhaps better it. Some may question the possible
advantage of reducing to varietal status many names that have been in use at
the species level in Europe and North America. First, I affirm that the present
taxonomic scheme accurately reflects the inferred relationships of Didymodon
taxa in Mexico and California — there is no borrowing of European concepts.
Second, the concepts accepted are quite similar, albeit at different taxonomic
levels, to those in modern European identification manuals and may represent
parallel speciative relationships. Third, this exercise in «lumping» has demons-
trated what are surely real, biologically-based relationships between many pre-
Source - MNHN. Paris
DIDYMODON IN MEXICO AND CALIFORNIA 381
viously poorly understood taxa and have brought some order to Barbula sensu
lato of recent manuals.
Varieties of Didymodon species are here presented as intergrading in several
characters. It may be argued that closely related species might also be expected
to show this. However, there are no clear discontinuities between most varieties
in the species treated here and a major inference in this paper is that many
species normally consist of pairs (or trios, etc.) of intergrading variants usually
sympatric in a major portion of the species’ range and disjunctive as colligations.
Such varietal groups may be simply disparate in extreme form (e. g. D. australa-
siae varieties) or may form parallel gradients in plant size and morphology
(e. р. D. vinealis varieties).
There are three nomenclatural inadequacies of the International Code of
Botanical Nomanclature (STAFLEU et al. 1978) that have made this treatment
of Didymodon difficult in execution and needful of unusual solutions. First,
as most taxonomists are well aware, the 1.C,B.N. provides rules that are in-
tended to stabilize usage of names, not to provide different names for differing
concepts of individual taxa. This may become a problem in cases when taxono-
mists’ concepts of taxa differ widely although including the same types. Here,
several species are recognized as vastly larger concepts than are usually held.
This may be a source of some ill ease on the part of other bryologists who now
must determine if an annotation of one such species name on an herbarium
specimen is «sensu Zander (this paper)» or sensu earlier authors. Certainly,
unless this distinction is clear, bryologists who favor narrow species limits in
Didymodon will perceive a chaotic situation in the herbarium. This problem
is not just related to the size of the taxon recognized but also to the characters
by which taxa are recognized. For instance, from descriptions, Barbula icmado-
phila (treated here as Didymodon rigidulus var. icmadophila) sensu NYHOLM
(1956) is quite different from Barbula icmadophila sensu STEERE (1938) or
SAITO (1975). Also, various authors have emphasized rather different characters
in their circumscriptions of Didymodon rigidulus. But, for each of these con-
cepts, the type specimens and, therefore, the scientific name remains the same.
At the generic level, Barbula Hedw. sensu NYHOLM (1956) and Barbula Hedw.
sensu SAITO (1975) are rather different concepts although the generitype is the
same. One might also note that the name Mnium Hedw, is now inexact as far
as its significance to biology is concerned unless one designates whether it is
to be used in the sense of KOPONEN (1968) or of earlier (and some later)
authors. The same is true of Polytrichum Hedw. (sensu SMITH 1971, or of
earlier and some later authors). In fact, stable names for taxa in conceptually
changing taxonomic frameworks result in confusion unless differing concepts
are identified individually. There are no means for distinguishing between
different taxonomic concepts that have the same scientific name unless such
concepts are cited as «sensu» or «emendavit» (Article 47A, 1.С.В.М.) the author
responsible for the concept. ї
Second, implicit in the 1.С.В.М. is the assumption that all taxa are disconti-
nuous. Most taxonomists «follow the rules» and make the best of it, but HES-
Source : MNHN, Paris
382 В.Н. ZANDER
LOP-HARRISON (1960) has stated quite correctly that «clinal variation, but its
very nature, is untreatable by taxonomic methods which demand the recogni-
tion of «types»». Nondiscontinuous taxa, intergrading in morphological charac-
ters, are assigned scientific names on the present paper by an unusual method
that does not, however, contravene provisions of the 1.C.B.N. The usual revisio-
nary nomenclatural treatment is to divide all synonyms of the species s. lat.
among the infraspecific taxa recognized and then determine priority of publica-
tion of legitimate names in each set of synonyms to find the correct name of
each infraspecific taxon. Since such initial segregation of synonyms is impossible
or at least arbitrary for what are apparently nondiscontinuous taxa, the scientific
name for each such infraspecific taxon is here based on the earliest legitimate
name among all synonyms of the species s. lat. with a type specimen clearly
representative of (or «within») the nondiscontinuous infraspecific concept
recognized. Of course, such names are selected first from those at the taxonomic
level recognized, and the type of the typical variety (or other taxonomic catego-
ту) must be included among the correct names of the infraspecific taxa reco-
gnized. All other heterotypic names are then simply left as synonyms of the
species s. lat. Additional discussion, with examples, is given in the treatment of
Didymodon rigidulus below.
The intergrading infraspecific taxa recognized in this paper are somewhat
similar to HERZOG's (1907) «ldealtypen» of the collective species Trichosto-
mum mutabile Bruch (= Т. brachydontium Bruch in F.A, Muell.). He recogni-
zed four informal names for arbitrarily circumscribed intergrading variants along
à morphological cline from large to small stature plants (involving several charac-
ters) that paralleled a north to south cline in distribution of the species in Europe.
The taxonomic scheme used here would treat the variants of the same collective
species in a non-arbitrary fashion, recognizing the large and small ends of the
cline as biological phenomena worthy of names and using a monotypic method
of providing scientific names for them. Informal names as used by HERZOG
(1907) have never proved popular, and the 1.C.B.N. should be used if at all
possible in modern taxonomic studies to provide a bridge between alpha taxono-
mists and concept-oriented experimentalists and biosystematists. Also, for the
biologist, there is no shame and certainly much value in appending «aff.» (=
affinis) before an epithet in an identification on an herbarium label when in fact
the taxon, though often clearly marked, is definitely perceived by a revisionist to
be open-ended, intergrading with another taxon
Third, ISOVIITA's (1966) statement «...the section on typification in the
present Code allows excessive freedom of interpretation in many cases...» is
still true. The Guide for the Determination of Types is extraordinarily yague
and anyone’s attempt at lectotypification may be challenged by a later worker
with different opinions. Because types of bryophyte names may consist of many
plants (or gametophores), any designation of holotype or lectotype may be
obviated by the subsequent division of the plants in a packet, or by the disco-
very somewhere of an additional isotype not seen during previous lectotypifica-
tion(s). If, for example, three different bryologists have proposed three different
lectotypes for the same scientific name of a taxon, whom does one follow ? The
Source : ММНМ Paris
DIDYMODON IN MEXICO AND CALIFORNIA 383
one who is best called a «specialist» ? The one whose lectotypification was
most «scientific» ? The one who saw the most isotypes ? Although designation
of a single gametophore as lectotype might solve some problems, identification
necessarily involving dissection then becomes hazardous or inappropriate. It
may well be that lectotypification of very early scientific names that involve
many, scattered, poorly labeled isosyntypes may ultimately have to involve
conservation of type specimens, now not provided for in the LC.B.N. The
discussion of Didymodon vinealis var. luridus, below, gives examples of problems
in typification in part due to the ease of creating isotypes and of obviating
designations of holotypes and lectotypes by division of a type specimen of
more than one gametophore, or the discovery of such at a later date. A recent
review and discussion of problems in typification of early bryophyte names
is given by KOPONEN (1979), and VITT (1981) has typified the species of
Macrocoma (Orthotrichaceae) exhaustively in a modern fashion.
SYNOPSIS OF TAXA OF DIDYMODON IN MEXICO AND CALIFORNIA
Didymodon Hedw. sect. Didymodon
D. incrassatolimbatus Card.
D. rigidulus Hedw. emend. Zander
var. rigidulus
var. gracilis (Schleich. ex Hook. & Grev.) Zander
var. icmadophila (Schimp. ex C. Muell.) Zander
var. subulatus (Thér. & Bartr. ex Bartr.) Zander
Didymodon sect. Asteriscium (C. Muell.) Zander, comb. nov. *
D. australasiae (Hook. & Grev.) Zander emend. Zander
var. australasiae
var. umbrosus (C. Muell.) Zander
D. revolutus (Card.) Williams
Didymodon sect. Fallaces (De Not.) Zander
D. fallax Hedw.
var. reflexus (Brid.) Zander
D. michiganensis (Steere) Saito
D. tophaceus (Brid.) Lisa
Didymodon sect. Vineales (Steere) Zander
D. vinealis (Brid.) Zander emend. Zander
var. vinealis
var. brachyphyllus (Sull. in Whippl.) Zander
var. luridus (Hornsch. in Spreng.) Zander
var. nicholsonii (Culm.) Zander
var. rubiginosus (Mitt.) Zander
var. flaccidus (B.S.G.) Zander
* Barbula sect. Asteriscium C. Muell., Linnaea 42 : 342, 1879, basionym. — Asteriscium
(C. Muell) Hilp., Beih. Bot. Centralbl. 50 (2) : 618. 1935, hom. illeg. non Asteriscium
Cham. & Schlecht., 1826. — Didymodon sect. Craspedophyllon Card., Rev. Bryol. 36 : 81.
1909, syn. nov.
Source : MNHN. Paris
384 В.Н, ZANDER
Descriptions of sect. Didymodon, sect. Vineales and sect. Fallaces (as sect.
Graciles, synonym fide ZANDER 1979) are provided by ZANDER 19783.
Section Asteriscium is characterized by the leaves spreading to spreading-recur-
ved when moist, broadly channelled; leaf margins not decurrent, plane to broadly
recurved throughout; costa ending 1-6 cells below apex to short-excurrent,
often rather broad at midleaf, adaxial surface convex; upper laminal cells uni.
stratose to evenly or in patches bistratose along the leaf margins, papillae absent
to large, low, simple to bifid 1(-4) per cell lumen each side; adaxial surficial cells
of costa quadrate to elongate, adaxial stereid band absent, hydroid (Begleiter
cell) groups often present between guide cells and abaxial stereid band: stem
occasionally with hyalodermis.
DIDYMODON HEDW., Spec. Musc. : 104, 1801
Generitype : D. rigidulus Hedw.
Husnotiella Card., Rev. Bryol. 36 : 71. 1909. Type : D. revolutus Card.
Trichostomopsis Card., Rev. Bryol, 36 : 73. 1909. Type : T. crispifolia Card.
For additional synonymy sce SAITO (1975),
Plants forming cushions or turf, light to dark green or olive-green. Stems
mostly to 2(-6) cm long, central strand present, occasionally very strong, axillary
hairs usually with 1-2 brown basal cells, Leaves ovate to long-lanceolate or long-
triangular, ventrally usually broadly concave, occasionally somewhat channelled
along the costa above midleaf, margins entire or occasionally weakly dentate
or crenulate, plane to recurved or revolute, leaf apex rounded to narrowly
acute, leaf base weakly differentiated to oblong and half-sheathing the stem;
costa ending several cells below the leaf apex to long-excurrent, ад. and abaxial
surficial cells quadrate to elongate, smooth or occasionally papillose; costa in
transverse section ovate, semicircular or reniform, epidermal layers differentia-
ted, adaxial stereid band present, absent or represented by a layer of large-
lumened substereid cells, guide cells 2-4, hydroids (Begleiter cells) seldom
present, dorsal stereid band usually weak; upper laminal cells subquadrate to
rounded-angular, occasionally short-rectangular or rhomboidal, walls thin to
evenly or unevenly thickened, usually bulging surficially on both surfaces,
laminal papillae often absent, usually solid, simple to bifid, centered over the
lumens to apparently scattered; basal laminal cells not or weakly differentiated
to strongly differentiated, quadrate to rectangular, smooth to weakly papillose.
Propagula occasionally present, usually spherical to elliptical, of 1-10 cells.
Dioicous (some species possibly rhizautoicous). Gametoecia terminal, perigonia
gemmate; perichaetial leaves ovate to long-lanceolate, often enlarged, often
loosely sheathing the seta, laminal cells usually prosenchymatous to above mid-
leaf. Sporophyte seta elongate, twisted clockwise below, occasionally counter-
clockwise above; theca ovate to long-cylindrical; stomata at base of theca,
phaneropore; annulus of 1-3 rows of weakly to strongly vesiculose cells, not
deciduous; peristome rudimentary or short, occasionally long and twisted, of
Source - MNHN, Paris
DIDYMODON IN MEXICO AND CALIFORNIA 385
16 short teeth or 32 triangular to linear rami, usually densely spiculose, occasio-
nally spiral-striate or papillose, basal membrane absent or low; operculum
short- to long-conic or conic-rostrate. Calyptra cucullate, smooth. Laminal
color reactions (after clearing in concentrated lactic acid) : Cl (conc. НО) -
green + light to medium yellow-brown, occasionally light to medium orange-
brown, rarely deep yellow or orange: К (10% KOH) - usually light to medium
yellow-brown or light to deep red-brown or red-orange-brown, occasionally
light to deep red, orange or yellow-orange, or light brown; N (conc. НМОз) -
light yellow-brown or light to medium red-brown or red-orange-brown, осса-
sionally light brown or medium orange; SE (H3SO4-ethanol, 2:1) - green + deep
red or red-orange-brown, occasionally medium orange-brown,
KEY TO SPECIES OF DIDYMODON IN MEXICO AND CALIFORNIA
1. Leaves with elongate upper adaxial costal cells . ......,,.......... их
1. Leaves with quadrate (occasionally short-rectangular) upper adaxial costal
baten? e iere we calde ee MM. eor o Re A cu El 5.
2. Upper laminal cells short-rectangular, upper laminal margins bistratose . . .
EE A чир ечи D, australasiae (Hook. & Grev.) Zander, р.р
2. Upper laminal cells subquadrate to rhomboidal, upper laminal margins
Meet очна + poid tone. фа неко EA d
3. Hygrophyte; leaves ovate to longelliptica, apex obtuse to acute, basal mar-
gins usually broadly decurrent, costa ending before the apex to short-excur-
rent, upper laminal cells often short-rectangular to elongate-rhomboidal |...
a eg E olin, КОЛИ E D. tophaceus (Brid.) Lisa
3. Mesophyte; leaves short- to long-lanceolate, apex acute, basal margins short
and broadly decurrent to little decurrent, costa percurrent to short-excurrent,
upper laminal cells mostly ovate to subquadrate .
4. Leaves catenulate when dry, not keeled or recurved when moist, basal
margins strongly recurved, upper laminal cells in very distinctive longitu-
dinal rows, propagula often present. . . . D. michiganensis (Steere) К. Saito
4. Leaves appressed to weakly spreading when dry, usually recurved and
keeled when moist, basal margins weakly recurved, upper laminal cells
in weakly differentiated longitudinal rows, propagula absent . . ........
FRAC SOME ДРА aN . . . D. fallax (Hedw.) Zander
5. Median basal laminal cells strongly differentiated, thin-walled, hyaline, often
inflated or transversely elongate, costa often with hydroids (Begleiter cells),
and adaxial stereid band absent; leaf apex usually broadly acute and weakly
cucullate; stem often with hyalodermis; plants often black green . ........
Fa be уредена её + «+ D australasiae (Hook. & Grev.) Zander, р.р.
5. Median basal laminal cells (if differentiated from upper laminal cells) weakly
differentiated, with thim to evenly thickened walls, costa lacking hydroids
(except D. revolutus), adaxial stereid Band often present; leaf apex not or
little cucullate, stem lacking hyalodermis; plants olive to light green ... . . 6.
6. Leaves short-ovate, costa spurred, wide, to 8 adaxial epidermal costal
Source : MNHN, Paris
386 R.H. ZANDER
cells across at midleaf, margins strongly recurved to revolute to near apex;
propagula, when present, unicellular; peristome absent to rudimentary ...
EEE aca M lt Кб ORNE D. revolutus (Card.) Williams
6. Leaves ovate to long-lanceolate, costa not spurred, narrower, 2-6 adaxial
epidermal costal cells across at midleaf, margins plane or recurved below
or to above midleaf; propagula mostly multicellular; peristome rudimen-
tary to well-developed ........... PAS Ta Au А
7. Leaves usually yellowish-green at high magnification, usually narrowly chan-
nelled adaxially along costa near apex, costa evenly thick to apex, 5-6 adaxial
epidermal costal cells across at midleaf (in large plants at least), percurrent to
short-excurrent in a conical mucro, adaxial stereid band usually represented
by substereid cells, guide cells often in 2-3 layers, upper laminal cell walls
thin to evenly thickened, lumens subquadrate, papillae often present, bifid or
multiplex and flattened; propagula seldom present. .D. vinealis (Brid.) Zander
7. Leaves usually bluish-green at high magnification, plane to broadly channelled
adaxially near leaf apex; costa narrowing to apex, 2-4 adaxial epidermal
costal cells across at midleaf, percurrent to strongly excurrent as a cylindrical
mucro, adaxial stereid band absent or present, guide cells in one layer, upper
laminal cell walls thin to unevenly thickened, lumens subquadrate to oval
or many-sided, papillae when present usually simple, seldom bifid; propagula
Given РДЫ, НЕ ext УА E гы СЛ e 8.
8. Peristome of long-triangular, red-orange, spirally striate or finely spiculose
to low-papillose teeth; leaves long-triangular to ovate-lanceolate, gradually
narrowed above, laminal margins evenly bistratose from midleaf to apex,
upper laminal cells heterogeneous in size and shape ................
AA Oc ere lage D. incrassatolimbatus Card.
8. Peristome of long-linear (or occasionally ovate, rudimentary), yellow or
orange, densely and strongly spiculose teeth; leaves ovate-lanceolate to
narrowly lanceolate, usually acuminate, laminal margins unistratose or
bistratose near apex or in patches to midleaf, upper laminal cells homo-
geneous in size and shape .. D. rigidulus Hedw.
+ DIDYMODON INCRASSATOLIMBATUS CARD., Rev. Bryol. 36 : 81. 1909.
Type : Mexico, Distrito Federal, Cañada, Pringle 10588 (PC - lectotype: BM,
FH, NY - isotypes); Michoacán, Patzcuaro, Pringle 748 (FH, BM, PC - isosyn-
types).
Plants forming cushions, green, yellow-green or olive-gréen above, brown
to reddish-brown below. Stems with few branches, brown, to 1.0(-2.0) cm long,
rounded pentagonal in transverse section, central strand usually strong, cortical
cells smaller, with walls thin or slightly thickened or cortex little differentiated,
hyalodermis absent; axillary hairs of 3-5 uniseriate cells, basal cell brown. Leaves
when dry appressed to laxly spreading, when wet widely spreading 45-90",
ovatelanceolate to long-triangular, (1.5-)2.0-2.5 mm long, adaxial surface
Source : MNHN, Paris
DIDYMODON IN MEXICO AND CALIFORNIA 387
broadly and weakly convex; leaf margins recurved basally or to midleaf, entire,
2(-4)-stratose along upper margins and in patches in medial portion of lamina;
leaf apex acute; leaf base scarcely differentiated to short-ovate, margins not or
shortly decurrent; costa percurrent to short-excurrent, ad- and abaxial surficial
cells quadrate, in 5-6 rows adaxially at midleaf, transverse section of costa
flattened-semicircular, adaxial surface weakly concave to weakly convex, adaxial
stereid band weak or absent, ad- and abaxial epidermal cells differentiated in
one layer, lumens of abaxial epidermal cells oval, guide cells 2-4 in one layer,
hydroids absent; upper laminal cells hexagonal to subquadrate, occasionally
reniform, 6-8 um wide, 1:1, walls thin to evenly thickened, often wavy, super-
ficially bulging, not organized in a distinct pattern, heterogeneous in size and
shape; laminal papillae usually absent, occasionally weakly differentiated, low
lenses, 1(-2) per lumen per size; basal laminal cells differentiated medially,
rectangular to rhomboidal, to 15 ит wide, 2-4:1, walls thin, hyaline. Propagula
not seen. Apparently dioicous, perigonia not seen. Perichaetia terminal, inner
leaves ovate-lanceolate, 2.0-2.9 mm long, sheathing the seta below, prosenchy-
matous near base. Sporophyte seta 1(-2) per perichaetium, 0.9-1.0 mm long,
orange-brown, twisted clockwise; theca 1.5-2.3 mm long, orange- or yellow-
brown, weakly sulcate when dry, long-elliptical to long-cylindrical, occasionally
curved, neck weakly differentiated, exothecial cells rectangular, 25-30 uim wide,
3-5:1, walls thin; stomates phaneropore, at base of theca; annulus of one layer
of vesiculose cells; peristome of 16 long-triangular, red- to yellow-orange teeth,
270-450 um long, straight or weakly twisted counterclockwise, spirally striate
or low papillose to finely spiculose, with several articulations, basal membrane
low or absent; operculum long-conic to conic-rostrate, 0.5-0.7 mm long, cells
weakly twisted counterclockwise. Calyptra cucullate, smooth, ca, 2 mm long.
Spores light brown, weakly papillose, (7-)10-12(-15) um in diameter. Laminal
color reactions (after clearing in conc. lactic acid) : Cl (conc. HCl) - green +
light yellow-brown; К (10% KOH) - medium orange-brown, occasionally red-
orange-brown; М (conc. НМОз) - light yellow-brown to red-brown; SE (conc.
Но SO, ethanol, 2:1) - green + deep red. Illustration : Pl. I, fig. 1-5.
Didymodon incrassatolimbatus differs from the very similar taxon D. vinealis
var. nicholsoni by the short-excurrent costa, costal transverse section with
adaxial stereid band, leaf apex only weakly channelled, perichaetial leaves ovate-
lanceolate, and peristome red-orange, of long-triangular teeth. Didymodon
australasiae var. australasiae is also similar in appearance, but can be distingui-
shed by the characters emphasized in the Key to Species. The syntype (Mexico,
Pringle 748) of D. incrassatolimbatus is Didymodon rigidulus var. subulatus.
Habitat : rock, soil, wet areas, riverside, occasionally submerged.
Distribution : Mexico : Distrito Federal, México, Michoacán, Morelos.
Range : endemic to Mexico.
Specimens examined : Mexico. Distrito Federal : Tlalpan, Amable 1349 (NY);
Canada, above Contreras, Pringle 10556 (FH), 10588 (BM, FH, NY), Barnes &
Land 442 (NY). México : 3 km E of San Rafael, Cárdenas 47 (MEXU, TENN).
Source : MNHN, Paris
388
К.Н. ZANDER
PLT. 1-5 : Didymodon incrassatolimbatus. 1-2 . Leaves. 3 : Leaf apex. 4 : Upper margi-
nal cells. 5 : Transverse section. 6 : Capsules. 7-10 : D, rigidulus var. rigidulus. 7-8 :
Leaves, 9 ; Leaf apex, note bistratose cells easily seen on edge at margins. 10 : Propa-
gula. Magnifications : leaves x 43 ; leaf apices, upper marginal cells, transverse sections
X 100; propagula x 100; capsules and peristomes x 28.
Michoacán : Morelia, Loma Santa María, Arséne 4866 (FH); Rincón, Arséne
4567 (FH); Morelos, Zempoala, 3.5 km from Tres Cubres, road to Cuernavaca,
Patrick 237 (TENN).
Source : MNHN. LA
DIDYMODON IN MEXICO AND CALIFORNIA 389
2. DIDYMODON RIGIDULUS НЕБУ. EMEND. ZANDER, Spec. Muse. : 104.
1801.
Trichostomum rigidulum (Hedw.) Tarn., Musc. Hib. : 34. 1804. — Barbula
rigidula (Hedw.) Milde, Bryol. Siles. : 118. 1863. — Tortula rigidula (Hedw.)
Lindb., Oefv. К. Vet. Ak. Foerh. 21 (4) :249. 1864.
Tortula acuta Brid., Musc. Recent. Suppl. 1 : 265. 1806. — Barbula acuta
(Brid.) Brid., Mant. Musc. : 96. 1819 — Didymodon acutus (Brid.) Saito,
J. Hattori Bot. Lab. 39 : 519, 1975.
Barbula teretiuscula Schimp. ex C. Muell., Syn. Musc. 1 : 614. 1849, syn. nov.
Type : Mexico, Veracruz, «ad montem Orizaba, Liebmann» (BM - iso-
type). — Tortula teretiuscula (Schimp. ex C. Muell.) Mitt., J. Linn. Soc.
Bot. 12:160. 1869.
Barbula mobilis C. Muell., Linnaea 42 : 482. 1879, syn. nov. Type : Venezue-
la, Fendler, 1855 (NY - isotype).
Barbula flaccidiseta Lor., Moosst. : 161. 1864, syn. nov. Туре: Mexico,
México, Schmitz s. n. (BM, NY - isotypes).
Barbula graciliformis Schimp. ex Besch., Mém. Soc. Nat. Sci. Nat. Cherb. 16 :
179. 1872, syn. nov. Type : Mexico, Distrito Federal, San Nicolas, Bour-
geau 1355 (PC - lectotype, BM - isotype).
Didymodon mexicanus Besch., Mém. Soc. Nat. Sci. Nat. Cherb. 16 : 172.
1872, syn. nov. Type : Mexico, Bourgeau, 27 Sept. 1865 (BM, NY - iso-
types). — Trichostomum mexicanum (Besch) C. Muell., Gen. Musc.
Frond. : 420. 1900.
Barbula leptocarpa Besch., Mém. Soc. Nat. Sci. Nat. Cherb. 16 : 179. 1879,
syn. nov. Type : Mexico, Veracruz, Orizaba, Bourgeau s. n. (BM, PC -
isotypes).
Barbula rigidula Besch., Mém. Soc. Nat. Sci. Nat. Cherb. 16 : 180. 1872,
hom. Шер. non B. rigidula (Hedw.) Mild., 1863; syn. nov. Type : Mexico,
Distrito Federal, Guadalupe, Bourgeau 1321 (PC - lectotype, NY - isotype).
Barbula erythropoda Schimp. ex Besch., Mém. Soc. Nat. Sci. Nat. Cherb. 16 :
180. 1872, syn. nov. Type : Mexico, San Cristobal, Mueller s. n. (NY, PC -
isotypes).
Barbula gracilescens Schimp. ex Besch., Mém. Soc. Nat. Sci. Nat. Cherb. 16 :
178. 1872, syn. nov. Type : Mexico, Veracruz, Mueller, 1853 (BM -
holotype).
Trichostomum ramulosum Schimp. ex Besch, Мет. Soc. Nat. Sci. Nat.
Cherb. 16 : 177. 1872, syn. nov. Type : Mexico, Mueller s. n. (BM - iso-
type). = Didymodon ramulosus (Schimp. ех Besch.) Card., Rev. Bryol. 36 :
82.1909.
Barbula bescherellei Sauerb. in Jaeg., Ber. S. Gall. Naturw. Ges. 1877-78 :
409. 1880 (Ad. 2 : 673), nom. nov. for B. rigidula Besch., 1872, hom.
Шер. — Barbula acuta var. bescherellei (Sauerb. in Jaeg.) Crum, Bryologist
72 : 241. 1969. é
Barbula lagunicola C. Muell., Bull. Herb. Boiss. 5 : 194. 1897, syn. nov.
Type : Guatemala, Laguna del Pino, Tuerkheim 114 (NY).
Source : MNHN. Paris
390 В.Н. ZANDER.
Barbula godmaniana C. Muell., Bull. Herb. Boiss. 5 : 193. 1897, syn. nov.
Type : Guatemala, Volcan de Fuego, Godman & Salvin (NY, SPA - isosyn-
types). — Didymodon godmanianus (C. Muell.) Bartr., Bryologist 49 : 113,
1946.
Barbula strictidens C. Muell, Bull. Herb. Boiss. 5 : 193. 1897, syn. nov.
Type : Guatemala, Laguna del Pinar, Tuerkheim 113 (NY - isotype).
Barbula subteretiuscula Card., Rev. Bryol. 36 : 85. 1909, syn. nov. Type :
Mexico, Hidalgo, Cuzamaloza, Pringle 10621 (РС - lectotype; FH, TENN -
isotypes).
Barbula altiseta Card., Rev. Bryol. 36 : 85. 1909, syn nov. Type : Mexico,
Michoacán, Patzcuaro, Pringle 751 (PC - lectotype; ВМ, FH, TENN - iso-
types).
Didymodon fuscoviridis Card., Rev. Bryol. 36 : 83. 1909, syn. nov. Type :
Mexico, Veracruz, Maltrata, Trelease, 1905 (PC - holotype).
Didymodon viridissimus Card., Rev. Bryol. 36 : 82. 1909, syn. nov. Type :
Mexico, Jalisco, Barranca de Guadalajara, Pringle 15227 (PC - lectotype,
NY - isotype).
Didymodon pusillus Card., Rev. Bryol. 36 : 82. 1909, hom. Шер. non D.
pusillus Hedw., 1801; syn. nov. Type : Mexico, Jalisco, Guadalajara,
Pringle 10574 (PC - holotype).
Barbula bescherellei var. stenocarpa Card., Rev. Bryol. 37 : 126. 1910, syn.
nov. Type : Mexico, México, Ixtaccihuatl, Purpus 3716 (PC - lectotype);
Distrito Federal, Cima, Barnes & Land 390 (NY, PC - isosyntypes); Сайа-
da, San Magdelena, Barnes & Land 161 (NY, PC - isosyntypes).
Didymodon heribaudii Card., Rev. Bryol. 40 : 35. 1913, syn. nov. Type :
Puebla, Rancho Posada, Nicolas 5976 (PC - lectotype, BM, NY - isotypes).
Barbula bescherellei var. crassinervia Thér., Smiths. Misc. Coll. 85 (4) : 17.
1931, syn. nov. Type : Mexico, Distrito Federal, Mixcoac, Arsène 9470
(PC - syntype), 9473 (FH - syntype).
Plants forming cushions or turf, green to dark or blackish green above, light
to dark brown below. Stems irregularly and seldom branching, light brown to
red-brown, mostly 1-2 cm long, in transverse section rounded-pentagonal, central
strand usually very strong, cortex of thick-walled cells with small lumens, seldom
not differentiated, hyalodermis absent or rarely weakly distinguishable in
patches; axillary hairs of 3-7 uniseriate cells, basal 1(-2) cells brown. Leaves
when dry appressed to weakly spreading, when wet weakly to widely spreading,
ovate- to longlanceolate, 0.8-1.9(-3.0) mm long, adaxial surface broadly chan.
nelled across leaf; leaf margins usually narrowly recurved in lower 1/2-3/4,
entire, unistratose or bistratose evenly or in patches above midleaf; leaf apex
obtuse to acuminate, occasionally entirely bistratose, occasionally bearing
thizoids; leaf base scarcely differentiated to ovate or oblong, basal margins
not or shortly and narrowly decurrent; costa percurrent to long-excurrent as a
cylindrical, sharp subula, often flexuose, ad- and abaxial surficial cells quadrate
above midleaf, usually smooth, in 2-5(-6) longitudinal rows at midleaf, transverse
section of costa reniform to elliptical, adaxial surface usually convex, ad- and
Source - MNHN, Paris
DIDYMODON IN MEXICO AND CALIFORNIA 391
abaxial stereid bands usually present but weak, ad- and abaxial epidermal cells
present in one layer, abaxial lumens oval, guide cells 2-4 in one layer, hydroids
absent; upper laminal cells subquadrate to hexagonal, (5-)7-9(-12) um wide, 1:1,
walls evenly thickened to thicker at the cell corners, surficially bulging, often
only abaxially, lumens rounded-quadrate to oval, arranged in distinct longitudi-
nal rows, homogeneous in size and shape; laminal papillae absent to simple or
bifid, often appearing as low lenses, 1-2 per lumen on each side or only abaxial-
ly, usually appearing centered over the lumens, occasionally fused in longitudinal
rows; basal laminal cells not differentiated or weakly differentiated, quadrate to
short-rectangular, 7-12(-36) um wide, 1-4:1, walls evenly thickened, occasionally
thin. Propagula often present, ovate to elliptical, green to brown, of mostly 3-8
cells, borne on rhizoids in leaf axils. Dioicous; perichaetia terminal, inner leaves
scarcely differentiated to longlanceolate, to 1.9-2.7 mm long, often loosely
sheathing the seta, prosenchymatous in lower 1/4-3/4; perigonia terminal, often
in series, on equal-sized or smaller gametophores, gemmate. Sporophyte seta
1(-2) per perichaetium, 0.7-1.7 mm long, red-brown to yellow, twisted clock-
wise, occasionally also counterclockwise above; theca 1.0-2.1 mm long, red-
brown, smooth when dry, long-elliptical to cylindrical, occasionally ovoid or
curved, neck weakly differentiated; exothecial cells rectangular, 18-35 ит wide,
4-5:1, walls evenly thickened or thin; stomates at base of theca, phaneropore;
annulus of 1-3 rows of weakly to strongly vesiculose cells; peristome of 16
rudimentary teeth or 32 filamentous rami, 180-740 ит long, yellow-brown,
occasionally orange, straight to twisted to 1.5 times, usually densely spiculose,
occasionally granulose, basal membrane low, 15-30 ит high, or absent, spiculose
or papillose; operculum long-conic to conic-rostrate, 0.4-0.9 (-1.5) mm long,
cells not or weakly twisted counterclockwise. Calyptra cucullate, smooth,
1.7-2.8 mm long. Spores brown to yellow-brown, smooth, 9-12(-15) um in
diameter. n = 12, 13. Laminal color reactions (after clearing in conc. lactic
acid) : Cl (conc. HCl) - green + light yellow-brown, seldom medium orange; К
(10% KOH) - light to dark orange, occasionally red- to yellow-orange, seldom
light red-brown; М (conc. HNO3) - light yellow- to orange-brown, occasionally
red-orange-brown, yellow-orange or orange; SE (conc. На SOs-ethanol, 2:1) -
green + dark red-orange-brown or dark red. Illustrations : Pl. 1, fig. 7-10; Pl. IL,
fig. 1-14.
Didymodon rigidulus is distinguishable from D. vinealis often only with great
difficulty. All relevant characters emphasized in the above Key to Species must
be carefully evaluated because various character states may be poorly expressed
in examined specimens.
Habitat : soil, gravel, acidic and calcareous rock, walls, occasionally bark,
moist or dry situation, 900-3600 m elevation.
Distribution : Mexico : Baja California, Baja California Sur, Chiapas, Chihua-
hua, Coahuila, Distrito Federal, Hidalgo, Jalisco, México, Michoacan, Morelos,
Nuevo León, Oaxaca, Puebla, San Luis Potosí, Sonora, Tamaulipas, Tlaxcala,
Veracruz, Zacatecas; widespread in California.
Range : nearly worldwide, but in tropical areas mainly found at high eleva-
tions.
Source : MNHN, Paris
392 В.Н. ZANDER
I here provide а key and columbariate names (those to which heterotypic
synonyms cannot be assigned with confidence — ZANDER 1978a) at the
varietal level for those bryologists who may wish to assign their collections to
«pigeonholes» corresponding to (1) concepts of these taxa that may be valid
in some areas of the world but are not discontinuous in Mexico and California,
or (2) to local variants that are of unusual morphology in some specimens.
Key toVarieties of Didymodon rigidulus Hedw. in Mexico and California
1, Peristome teeth rudimentary; entire upper lamina evenly bistratose........
crm те rms ie В Jerez var. subulatus (Bartr.) Zander
1. Peristome teeth short and straight to long and twisted; leaf apex or leaf
margins unistratose or bistratose, but medial cells unistratose.......... 2.
2. Leaves longelliptical to long-triangular, costa usually short-excurrent, leaf
apex and upper margins usually bistratose; propagula commonly present
түе Amat HS V SDN er A em . «var. rigidulus
2. Leaves lanceolate, costa percurrent to long-excurrent, upper laminal mar-
gins occasionally bistratose in patches; propagula commonly absent ... 3.
3. Leaves shortlanceolate to long-lanceolate, leaf base not sharply dilated,
rounded-square to rectangular, costa percurrent to long-excurrent as a rigid
awn, upper laminal cells commonly papillose, lumens oval or rounded-
quadrate, basal cells short-rectangular; propagula occasionally present... . . .
рее, бавария, E Lei var. gracilis (Hook, & Grev.) Zander
3. Leaves long-lanceolate, leaf base sharply dilated, oval, costa long-excurrent
as an often flexuose or fragile awn, upper laminal cells usually smooth,
lumens usually angular, basal cells usually quadrate; propagula absent . . . . . .
WET TENA RAT var. icmadophila (C. Muell.) Zander
— Didymodon rigidulus Hedw. var. rigidulus
Leaves long-elliptical to long-triangular, occasionally broadly ovate-lanceolate,
to 2.5 mm long, leaf base weakly differentiated to rectangular, upper margins
usually recurved, usually bistratose evenly or in long patches, lamina near
apex often completely bistratose; costa usually shortexcurrent as a thick,
blunt mucro, upper laminal cells usually papillose, basal laminal cells rectangular;
propagula commonly present; peristome teeth short and straight to weakly
twisted. Illustration : Pl. 1, fig. 7-10.
The var. rigidulus is relatively stenotypic in eastern North America, having
characteristic features of weakly spreading leaves, broad leaf apex, evenly bistra-
tose leaf margins and apex, short-excurrent to percurrent costa, and leaf cells
with rather thick walls. It intergrades with the other varieties, however, in
western North America. A specimen from eastern United States (New York,
Zander 4635 - BUF) with the above morphology, when cultivated at BUF in
а humid environment, produced branches with new leaves that approached
somewhat the morphology of those of var. gracilis; they were strongly recurved,
the apex was narrowly acute, the leaf margins were bistratose only in occasional
Source : MNHN, Paris
DIDYMODON IN MEXICO AND CALIFORNIA 393
patches, the lamina at the apex was unistratose, the costa was short-excurrent,
and the upper laminal cell walls were thin. It is evident that some collections
of D. rigidulus var. rigidulus, under certain environmental conditions, may grow
to look similar to var. gracilis. The degree to which morphological characteristics
of the varieties and of particular collections of D. rigidulus are stable under
changing environmental conditions is unknown, and broadly based studies
are important for adequate biological evaluation of the elements here treated
as varieties.
Habitat : soil, calcareous rock, moist areas, 900-2500 m elevation.
Distribution : Mexico : Baja California, Michoacán, Puebla and Veracruz.
Range : North and South America, Europe, northern Africa and Asia.
— Didymodon rigidulus var. gracilis (Schleich. ex Hook. & Grev.) Zander,
comb. & stat. nov.
Tortula gracilis Schleich. ex Hook. & Grev., Edinburgh J. Sci. 1 : 300.
1824, basionym, nom. nov. for Barbula gracilis Schwaegr., Spec. Musc.
Suppl. 1(1) :125. 1811, hom. illeg. поп B. gracilis Schum., 1803.
Leaves short- to long-lanceolate, 0.8-1.5(-3.0) mm long, leaf base not strongly
differentiated, rounded-square to rectangular, upper margins plane or recurved,
uni- or bistratose in patches, costa subpercurrent to long-excurrent as a rigid
awn, upper laminal cells commonly smooth, basal cells short-rectangular, propa-
gula occasionally present; peristome teeth short and straight to long and twisted.
Illustration : Pl. II, fig. 1-9.
Barbula gracilis has long been considered a later synonym of the more fami-
liar B. acuta (Brid.) Brid.; however, the appropriate combination of the latter
is preoccupied by Didymodon rigidulus var. acutus Biz. Although nomenclatural
synonymy of the first two names is here avoided for reasons discussed below,
the names are, historically, conceptual synonyms. There are no available varietal
names that unambiguously refer to the taxon described above, and a new status
is given the epithet gracilis.
DIXON (1924) and SMITH (1978, as B. acuta) noted the occasional presence
of propagula in var. gracilis, while other major authors did not and considered
the presence of propagula a diagnostic characteristic of var. rigidulus (as D. rigi-
dulus s. str.). CRUNDWELL and NYHOLM (1965) «reluctantly» synonymized
the European species Barbula valida (Limpr.) Moell. with var. gracilis (as B. acu-
га), noting correlated gradients in plant morphology and stature; the characteris-
tic leaf shapes ascribed to these taxa occurred at the extremes in plant size
variation but intermediate leaf shapes were found in plants of intermediate size.
MOENKEMEYER (1927) treated В. valida as a variety of D. rigidulus and
asserted that there were many transitional forms between it and var. rigidulus.
Specimens from western North America identified as «Barbula bescherellei»
have much the same morphological facies as European specimens I have seen
that were identified as «Barbula valida».
Source : MNHN, Paris
394 RH.ZANDER
PI. II. — 1-9 : Didymodon rigidulus var. gracilis. 1-4 : Leaves. 5-6 : Leaf apices. 7 : Upper
marginal cells. 8 : Transverse section. 9 : Peristome. 10 : D. rigidulus var. subulatus,
leaf. 11-13 : D. rigidulus var. icmadophila. 11 : Leaf. 12 : Leaf apex. 13 : Upper cells.
14 : Transverse section. Magnifications : leaves x 43; leaf apices, upper marginal cells,
transverse sections x 100; propagula x 100; capsules and peristomes x 28.
Habitat : soil, gravel, sandstone, limestone, walls, banks, 1300-3360 m
elevation.
Distribution : Mexico : Baja California, Chihuahua, Colima, Distrito Federal,
Hidalgo, Jalisco, México, Michoacán, Nuevo León, Oaxaca, Puebla, San Luis
Potosí, Sonora, Tamaulipas, Tlaxcala, Veracruz.
Range : North and Central America, Europe, the Middle East, and Asia.
— Didymodon rigidulus var. істадорЬйа (Schimp. ex C. Muell.) Zander,
comb. nov.
Barbula icmadophila Schimp. ex C. Muell., Syn. Musc. 1 :614. 1849, basio-
Source : MNHN, Pans
DIDYMODON IN MEXICO AND CALIFORNIA 395
пут, — Tortula icmadophila (Schimp. ex С. Muell.) Lindb., Oefv. К. Vet.
Ak. Foerh. 21 : 249. 1864. — Barbula gracilis ssp. icmadophila (Schimp.
ex C. Muell.) Amann, Fl. Mouss. Suisse 2 : 105. 1919 — Barbula gracilis
var. icmadophila (Schimp. ex С, Muell) Moenk., Laubm. Eur, : 289.
1927. — Barbula acuta ssp. icmadophila (Schimp. ex C. Muell.) Podp.,
Consp. Musc. Eur. : 209. 1954. — Didymodon «icmadophyllus» (Schimp
ex C. Muell.) Saito, J. Hattori Bot. Lab. 39 : 519. 1975, typographic
error.
Leaves longlanceolate, mostly 1.5-3.5 mm long, leaf base sharply dilated,
short-ovate, upper margins usually plane, occasionally recurved, usually unistra-
tose, costa excurrent as a long, often thick or flexuose subula, upper laminal
cells commonly smooth, basal cells usually quadrate, little differentiated; propa-
gula rarely present; peristome teeth short and straight to long and twisted.
Illustration : Pl. II, fig. 11-14.
The var. icmadophila may rarely have fragile or swollen awns of the excurrent
costae, but these are not both caducous early and swollen as in Didymodon
johansenii (Williams) Crum of New and Old World arctic and northern alpine
areas.
Habitat : soil, gravel, rhyolite, cement, brick, limestone, bark, generally
at higher elevations than var. gracilis, 1750-3600 m elevation.
Distribution : North and Central America, Europe, Asia; recognized in Mexico
from the states of Chiapas, Chihuahua, Distrito Federal, Hidalgo, Jalisco, Méxi-
co, Morelos, Oaxaca, Puebla, San Luis Potosí, Tlaxcala, Veracruz, Zacatecas;
California.
Range : North and Central America, Europe, Asia.
— Didymodon rigidulus var. subulatus (Thér. & Bartr. ех Bartr.) Zander,
comb. nov.
Didymodon mexicanus var. subulatus Thér. & Bartr. ex Bartr., Bryologist
29 :1. 1926. Type : U.S.A., Arizona, Pima Co., Santa Catalina Mts.,
Window Trail, Bartram 174 (CU - isotype).
Leaves long-lanceolate, 1.5-2.5 mm long, leaf base long-ovate, upper margins
usually plane, upper lamina entirely bistratose, costa excurrent as a subula,
upper laminal cells smooth, basal cells short-ovate; propagula apparently absent;
peristome very short, rudimentary, of 16 oval teeth with 1-2(-4) articulations.
Illustration : Pl. II, fig. 10.
In leaf shape and other characters, var. subulatus is similar to var. icmado-
phila. Specimens with upper lamina unistratose in large patches represent inter-
grades. The peristome characters were taken from very few specimens, but, in
all, var. subulatus appears to be a distinctive geographic variant.
Habitat : soil, rock, 900-2500 m elevation.
Source : MNHN, Paris
396 R.H, ZANDER
Distribution : Mexico : Distrito Federal, Michoacän.
Range : Mexico and U.S.A. : Arizona.
Specimens examined : Mexico : S. loc. , Hahn, 1868 (NY - as Barbula ery-
thropoda); Distrito Federal, Bourgeau s. n. (BM, NY - as Didymodon mexica-
nus); Vallée de Mexico, San Juanico, Amable 1332 (FH); Michoacán, Patzcuaro,
Pringle 748 (FH, NY - isosyntypes of Didymodon incrassatolimbatus). U.S.A. :
Arizona, Pima Co., Bartram 174 (CU).
The characters of Didymodon rigidulus appear to vary independently of each
other except for modalities of correlation at extremes of plant morphological
variation that are here recognized as varieties. The bryologist should resist the
temptation to distinguish between these varieties by only one character. In
eastern U.S.A., var, rigidulus usually has bistratose upper leaf margins, has
propagula and seems comparatively stenomorphic, but in western North Ameri-
ca, these characters are less well correlated and the bryologist is faced with such
problems as having to decide whether to identify one specimen as either an
unistratose var. rigidulus or a propaguliferous var. gracilis, or another specimen
as either a non-propaguliferous var. rigidulus or a bistratose-margined var. gracilis.
Previous authors have had differing concepts (under names in other combina-
tions) of var. gracilis and var. icmadophila depending on whether they emphasi-
zed relative length of the excurrent costa (DIXON 1924, STEERE 1938, SAITO
1975 among others) or the relative appearance of the median upper laminal
cells (NYHOLM 1956, SMITH 1978). Character states of most characters inter-
grade within the character and correlate only poorly between characters. Al-
though varietal concepts are recognized here because extreme variants are often
more common than intergrades and their appearance is rather distinctive, and
because the varieties may have some evolutionary and ecological importance,
a majority of the type specimens of heterotypic names that clearly belong to
D. rigidulus s. lat. cannot be assigned — with confidence in repeatibility —
to varietal synonymies on the basis of any combination of characters. For
purposes of determining priority, the name of each of the varieties here regarded
as correct is a combination with the earliest legitimate name or epithet (selected
from the synonymy of the species s. lat., and chosen first from those available
at the varietal level) whose type clearly represents the varietal concept reco-
gnized, and all other heterotypic names are assigned to the synonymy of the
species sensu lato. Thus, in a field of continuous variation, concepts of extreme
or distinctive morphology can be efficiently identified by the same number
of type specimens as there are names recognized as correct, of course necessarily
inclusive of the type specimen of the typical variety (or other infraspecific level
recognized). It is this author's opinion that stability of names is — in this particu-
lar case, at least — enhanced by this legal method of circumventing a major
deficiency of the 1.C.B.N., namely that Principle IV and Articles 25 and 57,
taken together, imply that all taxa are discontinuous. This is not the case with
the varieties of D. rigidulus, yet they are here recognized as (at least potential)
biological realities worthy of names. Since duplicability of results is a major
criterion of scientific worth, and because designation of synonymy is here
Source : MNHN, Paris
DIDYMODON IN MEXICO AND CALIFORNIA 397
considered a scientific study that should be repeatible by future workers, no
heterotypic synonymy is provided here for nondiscontinuous taxa.
3. DIDYMODON AUSTRALASIAE (-«П») (HOOK & GREV.) ZANDER
EMEND. ZANDER, Phytologia 41 : 21. 1978.
Tortula australasiae Hook. & Grev., Edinburgh J. Sci. 1 : 301. 1824. — Bar-
bula australasiae (Hook. & Grev.) Brid., Bryol. Univ. 1 : 828. 1827. — Tri-
chostomopsis australasiae (Hook. & Grev.) Robinson, Phytologia 20 :
187. 1970.
Barbula graminicolor C. Muell., Syn. Musc. 1 : 611. 1849, syn. nov. Type :
Chile, Racangua, Bertero, 1828 (BM, PC - isotypes). — Tortula graminico-
lor (C. Muell.) Mont. in Gay, Hist. Fis. Polit. Chile Bot. 7 : 156. 1850
Didymodon torquescens Card., Rev. Bryol. 36 : 83. 1909, syn. nov. Type :
Mexico, Michoacán, Bosque de San Pedro, Solorzano, 1908 (PC - holo-
type). — Husnotiella torquescens (Card.) Bartr.. Bryologist 29 :45. 1926 —
Asteriscium torquescens (Card.) Hilp., Вей. Bot. Centralbl, 50 (2) : 620.
1933.
Didymodon craspedophyllus Card., Rev. Bryol. 36 : 81. 1909, syn. nov.
Type : Mexico, Jalisco, Barranca de Guadalajara, Pringle 15227 (PC -
holotype).
Trichostomopsis crispifolia Card., Rev. Bryol. 36 : 74. 1909. Type : Mexico,
Hidalgo, Tula, Pringle 15273 (NY - syntype).
Didymodon diaphanobasis Card., Rev. Bryol. 37 : 125, 1910. Type : Mexico,
México, Ixtaccihuatl, Purpus 3721 (PC - holotype). — Trichostomopsis
diaphanobasis (Card.) Grout, Moss Fl. N. Amer. 1 : 228. 1939.
Didymodon patentifolius Thér., Smithsonian Misc, Coll. 85(4) : 15. 1931,
gyn. nov. Type : Mexico, D.F., Xoquiapán, Amable 1676 (PC - syntype).
Trichostomopsis brevifolia Bartr., Bryologist 34 : 61. 1932.
Trichostomopsis fayae Grout, Moss Fl. М. Amer. 1 : 228. 1939. Туре:
U.S.A., California, Los Angeles Co., MacFadden 8172 (DUKE - holotype).
Plants forming turf and cushions, green to black-green above, brown below.
Stems often branching, brown, to 0.6(-1.5) cm long, rounded-pentagonal in
transverse section, central strand strong, cortical cells not differentiated or with
thicker walls, hyalodermis not differentiated or weakly developed or developed
in patches to distinct, strongly differentiated; axillary hairs of 3-5 cells, basal
cell brown. Leaves crowded, when dry spreading incurved and twisted to incur-
ved-appressed, when wet spreading to spreading-recurved, ovate-triangular to
longelliptical or long-lanceolate, usually 1.0-2.0(-4.0) mm long, adaxial surface
broadly channelled across leaf; leaf margins plane to recurved at midleaf to
throughout upper half of leaf, entire, bistratose in one to several rows; leaf
apex acute, often somewhat cucullate; leaf base scarcely differentiated in shape
to ovate and sharply differentiated, basal leaf margins often with one or more
Source : MNHN, Paris
398 R.H. ZANDER
tows of narrow cells bordering median basal cells; costa subpercurrent to short-
excurrent, adaxial surficial cells quadrate to short-rectangular or elongate,
smooth to weakly papillose, abaxial surficial cells elongate or seldom quadrate
near apex, smooth to papillose, transverse section of costa elliptical to sub-hemi-
spherical, adaxial surface convex, adaxial epidermis present, abaxial epidermis
usually present, lumens oval to semicircular, adaxial stereid band absent, guide
cells 3-6 in 1-2 layers, hydroids occasionally present, abaxial stereid band strong
to weak or substereid or absent; upper laminal cells subquadrate to short-rectan-
gular, 7-12 um wide, 1-2:1, walls evenly thickened to thickened at the corners,
surficially weakly bulging, patterned in longitudinal rows; laminal papillae
absent to large, entered over the lumens, solid, simple to bifid, occ. trifid, flatte-
ned-rounded, often massive, 1-4 per lumen each side; basal laminal cells often
sharply differentiated, medially or across leaf base, quadrate to rectangular,
often transversely elongate, to 18(-23) um wide, 2-5:1, walls thin, hyaline,
somewhat to distinctly bulging, occasionally resorbed to form transverse slits in
the medial portion of the leaf base. Propagula occasionally present, red-brown,
spherical to elliptical or irregularly shaped, of ca. 2-7 cells, to 85(-100) um
long, borne on rhizoids between lower leaves. Dioicous; perichaetia terminal,
inner leaves long-elliptical, to 2.5(-4.0) mm long, little to half sheathing, pro-
senchymatous below ot to 1/2 of leaf length; perigonia terminal on smaller
plants, often in series, gemmate, inner leaves entire to serrate. Sporophyte
seta 1 per perichaetium, 0.7-1.0 mm long, red-brown to yellow, twisted clock-
wise, occasionally counterclockwise above; theca 1.0-1.9 mm long, red-brown
to dark yellow-brown, ovoid to elliptical-cylindrical, neck weakly differentiated;
exothecial cells rectangular to rhomboidal, 20-25 um wide, 2-4:1, walls thin:
stomates phaneropore, at base of theca; annulus strongly vesiculose, 1-2 rows
of cells; peristome of 32 teeth, occasionally rudimentary, linear, ca. 600 um
long, with many articulations, yellow, nearly straight to twisted counterclock-
wise 0.5 times, closely spiculose, basal membrane absent or low, to 60 ит high,
branching-spiculose. Operculum long-conic, often curved, 0.7-1.2 mm long, cells
nearly straight to twisted counterclockwise to one time. Calyptra cucullate,
smooth, 2.0-2.5 mm long. Spores light brown, weakly papillose, 11-15 ит in
diameter, possibly occasionally anisosporous in some specimens, 6-9 and 12-
15 um wide. Laminal color reactions (after clearing in conc. lactic acid.) : Cl
(conc. HCl) - green or blue-green + light to medium brown or yellow-brown,
or medium orange-brown; К (10% KOH) - green + light to medium yellow.
brown, light to dark red-brown, or dark orange-brown; N (сопс HNO3) - light
brown or light to medium red-brown or light orange-brown; SE (H2SOq-ethanol,
2:1) - green + red-orange-brown to medium brown. Illustration : Pl. Ш, fig. 1-8.
In the genus Didymodon, hydroids (Begleiter cells) have been seen in the
costae of only this species and D. revolutus. Hydroids are common in the Pot-
tioideae, and the lack of an adaxial stereid band may also indicate a relation to
that subfamily. However, the narrow leaves, presence of an abaxial epidermis
in the costa, and general appearance, together with the presence of hydroids
in both Barbula and Bryoerythrophyllum spp., support retention of this species
in the Barbuloideae. Both varieties recognized below bear tubers. The varieties
Source - ММНМ. Paris
DIDYMODON IN MEXICO AND CALIFORNIA 399
Р.Ш. — 1-4 : Didymodon australasiae var. australasiae. 1-2 : Leaves. 3 : Leaf apex. 4:
Transverse section. 5-8 : D. australasiae var. umbrosus. 5 : Leaf, 6 : Leaf apex. 7:
Upper marginal cells. 8 : Transverse section. 9-13 : D. revolutus. 9-10 : Leaves. 11 :
Leaf apex. 12 : Transverse section. 13 : Propagula. Magnifications : leaves x 43; leaf
apices, upper marginal cells, transverse sections x 100; propagula x 100; capsules and pe-
ristomes x 28.
are distinctive in extreme form, but intergrade completely in morphology.
Their descriptions given below are nondiscontinuous and heterotypic synonyms
are not recognized at the varietal level but are given instead for the species
sensu lato, above.
Source : MNHN, Paris
400 К.Н, ZANDER
Habitat : on rock, soil, 137-3750 m elevation.
Distribution : Mexico : Baja California, Chihuahua, Distrito Federal, Jalisco,
México, Michoacán, Puebla, San Luis Potosí, Sonora; California.
Range : North, Central and South America, Europe, Canary Islands, South
Africa, Australasia.
Key to Varieties of Didymodon australasiae (Hook. & Grev.) Zander
(Note : Only salient characters of the idealized taxa are given here.
See descriptions for variation in these and other characters.)
Leaves blackish-green, short-ovate to long-elliptical, apex broadly acute, weakly
cucullate, leaf base not sharply differentiated in shape, adaxial costal cells
quadrate, upper laminal cells rounded-quadrate. ......... var. australasiae
Leaves green to very bright green, long-lanceolate, apex narrowly acute, not
cucullate, leaf base sharply differentiated, ovate, adaxial costal cells rectan-
gular, upper laminal cells rectangular. ....... Маг. umbrosus (C. Muell.) Zander
— Didymodon australasiae (Hook. & Grev.) Zander var. australasiae
Stem hyalodermis absent, occasionally present in patches: plants often blackish-
green; costa bulging usually adaxially: leaves short-ovate to long-elliptical, occa-
sionally lanceolate, apex broadly or occasionally narrowly acute, often some.
what cucullate; leaf margins usually recurved, leaf base not or weakly differentia-
ted in shape, not perforated medially; adaxial cells of costa quadrate, occasio-
nally short-rectangular; upper laminal cells rounded-quadrate; upper laminal
papillae usually distinct, occasionally very coarse, often absent; basal marginal
laminal cells quadrate to rectangular, seldom in distinct rows, medial basal
cells quadrate, occasionally longitudinally or transversely short-rectangular,
little bulging, Illustration : Pl. IIT, fig. 1-4.
Habitat : rock, soil, 137-3750 m elevation.
Distribution : Mexico : Baja California, Chihuahua, Distrito Federal, Jalisco,
México, Michoacán, San Luis Potosi, Sonora; California.
Range : North, Central and South America, Europe, southern Africa, Austra-
lasia.
— Didymodon australasiae var. umbrosus (C. Muell.) Zander, comb. & stat.
nov.
Barbula umbrosa C, Muell., Linnaea 42 : 340. 1879, basionym. — Tricho-
stomopsis umbrosa (C. Muell.) Robinson, Phytologia 20 : 185, 1970.
Stem hyalodermis usually present; plants green to very bright green; costa
adaxially flattened; leaves long-lanceolate, apex narrowly acute, not cucullate;
leaf margins plane, leaf base usually sharply differentiated in shape, ovate,
occasionally medially perforated with transverse slits; adaxial cells of costa
usually rectangular; upper laminal cells quadrate to rectangular; upper laminal
papillae absent, occasionally weakly developed; basal marginal cells narrowly
Source - MNHN. Paris
DIDYMODON IN MEXICO AND CALIFORNIA 401
rectangular in several rows bordering the inflated inner basal cells, occasionally
this not clearly evident, medial basal cells short-rectangular, usually somewhat
bulging. Illustration : Pl. III, fig. 5-8.
Habitat : soil, river margin, 500-2160 m elevation.
Distribution : Mexico : Distrito Federal, Hidalgo, Jalisco, México, Michoacán,
Puebla, San Luis Potosí; California.
Range : North and South America, Europe.
4. DIDYMODON REVOLUTUS (CARD.) WILLIAMS, Bryologist 16 : 25. 1922.
Husnotiella revoluta Card., Rev. Bryol. 36 : 71. 1909, basionym.
Husnotiella palmeri Card., Rev. Bryol. 37 : 121. 1910 — Husnotiella revoluta
var. palmeri (Card.) Thér., Smiths. Misc. Coll. 85 (4) : 7. 1931.
Plants bulbiform, gregarious to forming turf, green to black-green above,
light brown below. Stems seldom branching, brown, to 0.6 cm long, rounded-
pentagonal in transverse section, central strand present, often strong or dark
brown or both, cortex scarcely differentiated or cortical cells with darker walls,
hyalodermis absent or weakly differentiated as a layer of cells larger than the
cortical cells; axillary hairs of 3-9 cells, basal one brown. Leaves when dry appres-
sed, imbricated, incurved, when wet widely spreading, ovate, 0.4-1.2(-2.0) mm
long, adaxial surface broadly channelled across leaf; leaf margins recurved to
revolute to near apex, occasionally recurved to just above midleaf, entire, bistra-
tose in small patches or unistratose; leaf apex rounded-obtuse, often strongly
cucullate; leaf base scarcely differentiated in shape, margins weakly decurrent;
costa ending 1-6 cells below the apex, broad and spurred above midleaf, adaxial
surficial cells quadrate, papillose, abaxial surficial cells quadrate to elongate,
smooth or papillose; transverse section of costa elliptical, adaxial surface bul-
ging-convex, adaxial stereid band absent, guide cells 2-4 in one layer, hydroids
(Begleiter cells) occasionally present, adaxial stereid band present, occasionally
substereid; upper laminal cells rounded-quadrate to rhomboidal, (6-)8-10 um
wide, 1:1, walls thick, surficially bulging, lumens rounded-quadrate, arranged
in longitudinal rows; papillae often best developed in the upper medial portion
of lamina, low, simple or occasionally multiplex or plate-like, usually one per
lumen each side, occasionally absent; basal laminal cells differentiated medially
or across the leaf base, short-rectangular, elongated longitudinally or transverse-
ly, 8-11 ит wide, 1:1-2 ог 1-2:1, walls thin to weakly thickened. Propagula
seldom present, unicellular, spherical to elliptical, red-brown, 16-20 um
wide, 1:1(-2), borne in large clusters in axils of upper leaves. Dioicous; perichae-
tia terminal, inner leaves occasionally smaller, shorter than cauline leaves,
weakly sheathing seta, prosenchymatous in lower third; perigonia on smaller
gametophores, gemmate. Sporophyte seta 1 per perichaetium, 0.35-0.9 mm long,
red- to yellow-brown, twisted clockwise, counterclockwise above; theca 0.7-
1.5 mm long, red-brown to dark brown, smooth when dry, elliptical to short-
Source : MNHN. Paris
402 В.Н. ZANDER
cylindrical, neck weakly differentiated; exothecial cells rectangular, 16-22 ит
wide, 2-3:1, walls weakly thickened; stomates present at base of theca, phanero-
pore: annulus of 1-3 rows of highly vesiculose cells, deciduous in pieces; peri-
stome absent to rudimentary, teeth irregularly ligulate, to 65 um long, to 4 arti-
culations, straight, papillose, yellow to red-brown; operculum conic, rather
oblique, 0.3-0.4 mm long, cells not twisted. Calyptra cucullate, smooth, 1.7-
2.0 mm long. Spores yellow to brown, smooth to lightly papillose, 9-11 um in
diameter. Laminal color reactions (after clearing in conc. lactic acid) : Cl (conc.
НС) - green + medium to dark yellow-brown; К (10% KOH) - light brown to
medium red-orange-brown; М (conc. HNO3) - medium brown to red; SE (conc.
H¿SO4-ethanol, 2:1) - dark green + medium orange-brown or medium red-
brown. Illustration : Pl. IIT, fig. 9-13.
1 concur with WILLIAMS’ (1913) opinion that Husnotiella revoluta belongs
with Didymodon. Husnotiella torquescens is a synonym of D. australasiae.
ROBINSON (1970) noted that some specimens of D. revolutus (as Husnotiella
revoluta) resemble D. australasiae (as Trichostomopsis australasiae). There is
indeed a close relationship between the two taxa as at least some specimens
of each share the following character states : leaves occasionally with plane
margins, apex often cucullate, marginal laminal cells often bistratose, adaxial
costal surface bulging and hydroids evident in the costal transverse section.
Didymodon revolutus differs from D. australasiae, however, in these respects :
rudimentary or absent peristome, rounded-obtuse leaf apex, often revolute
laminal margins, subpercurrent costa with spurs, and one layer of guide cells
instead of two. Bryoerythrophyllum calcareum (Ther.) Zander is similar to
D. revolutus in gametophytic characters but may be distinguished by the for-
mers crowded, hollow, multiplex рарШас with several salients per lumen each
side.
Habitat : soil, rock, walls, to 2280 m elevation.
Distribution : Mexico : Baja California, Distrito Federal, Durango, Hidalgo,
Jalisco, México, Michoacán, Morelos, Puebla, San Luis Potosí, Tlaxcala, Zacate-
cas; California.
Range : southwestern U.S.A. : California, Arizona, Texas, Oklahoma; Mexico;
Guatemala; Andes of South America.
5. DIDYMODON FALLAX (HEDW.) ZANDER, Phytologia 41 : 28. 1978.
Barbula fallax Hedw., Spec. Musc. : 120. 1801.
Barbula ferruginea Schimp. ex Besch., Mém. Soc. Nat. Sci. Nat. Cherb. 16 +
181. 1872. Type : Mexico, San Cristobal, Mueller s. n. (PC - holotype:
BM, NY - isotypes). — Triquetrella ferruginea (Schimp. ex Besch.) Thér.,
Smiths. Misc. Coll. 85 (4) :9. 1931.
Plants in turf or loose cushions, green to reddish-brown above, brown to
reddish-brown below. Stems irregularly branching, yellow- to red-brown or
Source : MNHN. Paris
DIDYMODON IN MEXICO AND CALIFORNIA 403
dark brown, to 2.0 cm long, rounded pentagonal in transverse section, central
strand present, cortex differentiated as 2-3 layers of stereid cells, hyalodermis
absent; axillary hairs of 4-5 cells, basal one brown; stem often flattened. Leaves
when dry appressed to weakly spreading, when wet spreading to strongly refle-
xed-recurved, ovate-triangular to lanceolate, 1.2-2.0 mm long, keeled; leaf mar-
gins nearly plane to recurved to midleaf, entire; leaf apex acute; leaf base scarce-
ly differentiated to ovate, basal margins narrowly to broadly decurrent, occasio-
nally somewhat auriculate; costa short-excurrent in an often papillose mucro,
adaxial surficial cells short-rectangular to elongate, smooth, abaxial surficial
cells quadrate to near base, often papillose, transverse section of costa semi-
circular, adaxial surface mostly concave, adaxial epidermis absent, abaxial epi-
dermis present, lumens of cells oval to hexagonal, adaxial stereid band absent
or of few cells, guide cells 2-4 in one layer, abaxial stereid band seldom strong,
often reduced or absent; upper laminal cells subcircular to rhomboidal or pen-
tagonal, (5-)8-11 um wide, 1:1 or 1.5.1, occasionally transversely elongate,
walls thickened at the corners, surficially bulging, lumens oval to triangular or
diamond-shaped, arranged on longitudinal rows; laminal papillae absent or sim-
ple, solid, 1-3 centered over lumens on each side, occasionally bi- to trifid;
basal laminal cells not or weakly differentiated, quadrate or short-rectangular.
Dioicous; perichaetia terminal, inner leaves enlarged, to 3.0 mm long, leaf
base rectangular, sheathing seta, prosenchymatous in lower 2/3; perigonia termi-
nal, occasionally in series, gemmate, inner leaves ovate. Sporophyte seta 1 per
perichaetium, 0.6-1.2 ст long, yellow-brown, twisted clockwise; theca 1.2-
1.5 mm long, brown, smooth when dry, cylindrical to elliptical, neck weakly
differentiated; exothecial cells rectangular, 14-27 um wide, 2-3:1, walls thin;
stomates present at base of theca, phaneropore; annulus weakly differentiated,
of 2-3 rows of vesiculose cells; peristome of 16 teeth cleft to near base, linear,
275-1100 ит long, straight to twisted counterclockwise once, yellow to yellow-
orange, densely papillose to weakly spiculose, basal membrane low to rather
high, 25-70 um high, papillose; operculum long-conic, 1.2-1.3 mm long, cells
in straight rows or twisted to one time. Calyptra not seen. Spores light yellow-
brown, weakly papillose, 7-9 ит in diameter. п = 9, 10, 11, 13, 13 + 1. Laminal
color reactions (after clearing in conc. lactic acid) : Cl (conc. HCl) - green +
medium yellow-brown to light orange; K (10% KOH) - light to dark red-brown,
occasionally medium brown or deep red-orange-brown; N (conc. HNO3) -
light brown to red-brown; SE (Hz SO4-ethanol, 2:1) - blue-green + light to dark
red-brown, occasionally orange-brown.
Distinctions between true Triquetrella, known only from California, and
D. fallax are discussed by ZANDER (1980). All collections of D. fallax seen
were aff. var. reflexus but probably material aff. var. fallax will eventually turn
up in Mexico, I agree with FLOWERS’ (1973) observations that character states
typical of the varieties as recognized here are not constant. The varieties are
characteristic only in extreme form — for this reason the synonym Triquetrella
ferruginea is placed with D. fallax sensu lato, although the type of the former
is aff. var. reflexus.
Source : MNHN, Paris
404 R.H. ZANDER
— Didymodon fallax var. reflexus (Brid.) Zander, Bryologist 83 : 230. 1980.
Tortula reflexa Brid., Musc. Recent. Suppl. 1 : 255. 1806. — Barbula
reflexa (Brid.) Brid., Mant. Musc. : 93. 1819. — Barbula fallax var. reflexa
(Brid.) Brid., Bryol. Univ. 1 : 558. 1826.
Leaves reddish-brown, strongly recurved, apex broadly acute, laminal cell
walls strongly thickened, laminal papillae usually present, often high. Plate IV,
fig. 1-5.
Habitat : rock, limestone, bark, 2200-2800 m elevation.
Distribution : Mexico : Distrito Federal, Nuevo León, Oaxaca, Puebla, Vera-
cruz.
Range : eastern and central North America, Europe, North Africa, Asia.
6. DIDYMODON MICHIGANENSIS (STEERE) SAITO, J. Hattori Bot. Lab
39 :516. 1975.
Barbula michiganensis Steere in Grout, Moss Fl. М. Amer. 1 180. 1938.
Type : U.S.A., Michigan, Alger Co., Pictured Rocks, Nichols & Steere, 1935
(DUKE - isotype).
Plants in loose turf, green or glossy green above, brown below. Stems seldom
branching, brown, to 2(-6) cm long, rounded-pentagonal in transverse section,
central strand present, cortex of 2-3 layers of stereid cells, hyalodermis absent;
axillary hairs of 4 cells, basal one brown; stem somewhat flattened. Leaves
when dry catenulate-incurved, when wet spreading ca. 45°, lanceolate to
ovatedanceolate, 0.9-1.1, keeled in upper third; leaf margins narrowly to
broadly recurved in lower 1/2-2/3, entire; leaf apex acute to acuminate; leaf
base broadly ovate, basal margins short and broadly decurrent, often weakly
but distinctly auricled; costa percurrent to very shortly excurrent, adaxial sur-
ficial cells rectangular, smooth, abaxial surficial cells short-rectangular or qua-
drate, smooth, transverse section of costa semicircular, adaxial surface concave,
adaxial epidermis lacking, abaxial epidermis present, cells with oval lumens,
adaxial stereid band lacking (apparently represented by 1-2 layers of substereid
or parenchymatous cells), guide cells 2 in one layer, abaxial stereid band absent
or weak; upper laminal cells rhomboidal to rounded pentagonal, 7-9(-12) ит
wide, 1-2:1, walls thickened at the corners, often very thick, lumens angular to
oval, in very distinctive longitudinal rows; laminal papillae absent to low, simple;
basal laminal cells not to weakly differentiated, quadrate to rectangular, 9-13 um
wide, 1-2:1, walls thick. Propagula usually present, spherical to elliptical, red to
orange, 13-25 ит wide, 1-2:1, of 1-4 cells, borne in masses in the leaf axils.
Sporophyte not seen. Laminal color reactions (after clearing in conc. lactic
acid) : Cl (conc. HCl) - green + light to deep yellow ог yellow-orange; К (10%
KOH) - light yellow-orange-brown to deep orange; N (conc. HNO3) - light
yellow-brown to medium orange; SE (conc. HaSOs-ethanol, 2:1) - green +
medium red-brown. Illustration : Pl. IV, fig. 6-10
Source : MNHN, Paris
DIDYMODON IN MEXICO AND CALIFORNIA 405
Didymodon michiganensis is similar to D. fallax in morphology and may
be mistaken for the latter when propagula are absent; however, the following
characters are diagnostic : leaves not keeled or recurved when moist, strongly
catenulate when dry, lower margins strongly recurved, and upper laminal cells
in distinct longitudinal rows and very thick-walled.
Habitat : cliffs, stone walls, boulder, to 2440 m elevation.
Distribution : Mexico : Chihuahua.
Range : Canada : Northwest Territories; U.S.A. : Michigan; Mexico; India :
Assam; Japan.
PLIV.— 1-5 :Didymodon fallax var. reflexus. 1 : Leaf. 2 ; Leaf apex, lateral view. 3 :
Upper marginal cells. 4 : Elongate adaxial costal cells. 5 : Transverse section. 6-10 :
D. michiganensis. 6-7 : Leaves. 8 : Leaf apen? 9 : Upper marginal cells. 10 : Propagula,
1144 : D. tophaceus. 11-12 : Leaves. 13 : Leaf apex. 14 : Upper marginal cells. Magni-
fications : leaves x 43; leaf apices, upper marginal cells, transverse sections x 100; propa-
gula x 100; capsules and peristomes x 28.
Source : ММНМ. Paris
406 К.Н. ZANDER
7. DIDYMODON TOPHACEUS (BRID.) LISA, Elenco Muschi Torino : 31.1837.
Trichostomum tophaceum Brid., Mant. Musc. : 84. 1819. — Barbula tophacea
(Brid.) Mitt., J. Linn. Soc. Bot. Suppl. 1 : 34. 1859.
Dactylhymenium pringlei Card., Rev. Bryol. 36 : 72. 1909. Type : Mexico,
Chihuahua, Chihuahua, Pringle 4 (PC - lectotype; ВМ - isotype). — Barbula
pringlei (Card.) Hilp., Вей. Bot. Centralbl. 50 (2) :645. 1933, hom. illeg.
non Card. 1909. — Husnotiella pringlei (Card.) Grout, Moss Fl. N. Amer.
1:219. 1939.
Barbula abbonii Thér., Smiths. Misc. Coll. 85 (4) : 20. 1931. Type : Mexico,
Nuevo León, Monterrey, Abbon 10970 (PC - lectotype; BM, FH - iso-
types).
Plants in turf or cushions, green to olive-green above, tan below, costa often
dark brown. Stems irregularly branching, brown, to 1.5 ст long, transverse
Section rounded pentagonal, central strand present, cortex of 2-3 layers of
darker, smaller cells, hyalodermis absent; axillary hairs of 3-5 cells, basal one
brown. Leaves when dry spreading to 45°, occasionally appressed or nearly so,
when wet spreading, ligulate to ovatelanceolate, 1.0-1.9 mm long, adaxial
surface broadly channelled to keeled; leaf margins weakly recurved in lower
3/4, entire or occasionally crenulate apically, occasionally bistratose in patches;
leaf apex broadly acute to obtuse or rounded; leaf base ovate, basal margins
usually broadly decurrrent, rarely low, small lamellae present medially abaxially;
costa percurrent or ending up to 6 cells below apex, occasionally short-excur-
rent, adaxial surficial cells rectangular, smooth, abaxial surficial cells quadrate
ог rectangular, usually smooth, transverse section of costa semicircular, adaxial
surface concave to convex, adaxial epidermis absent, abaxial epidermis present,
cell lumens oval, adaxial stereid band absent or represented by substereid cells,
guide cells 24 in one layer, abaxial stereid band weak or absent; upper laminal
cells oval to rhomboidal or triangular, (7-)9-12(-14) um wide, 2-1:1 or 1:1-2,
often longitudinally and occasionally transversely elongate, surficially bulging,
lumens angular to dumbbell-shaped, arranged in longitudinal rows, heteroge-
neous in size and shape; laminal papillae low, broad, simple, 1-2 salients per
lumen each side, sometimes apparently absent; basal laminal cells little diffe-
rentiated, 10-12 um wide, 2-1:1, walls irregularly thickened. Dioicous; perichae-
tia terminal, inner leaves ovate-lanceolate, to 1.5 mm long, somewhat sheathing
the seta, prosenchymatous to near apex; perigonia terminal, gemmate, inner
leaves ovate. Sporophyte seta 1 per perichaetium, 0.7-0.9 mm long, red-brown,
twisted clockwise; theca 0.6-1.3 mm long, red-brown, smooth when dry, ovate
to short-cylindrical, neck weakly differentiated, mouth occasionally oblique:
exothecial cells rhomboidal to rectangular, 15-25 um wide, 1-3:1, walls evenly
thickened; stomates present on neck, phaneropore; annulus of 2-3 rows of
weakly vesiculose cells; peristome absent, rudimentary or short, of 16 teeth cleft
to near base, occasionally perforated below, ligulate, to 220 um long, straight,
1-5 articulations, light yellow, spiculose to papillose, basal membrane absent:
operculum long-conic, 1.0-1.2 mm long, cells in straight rows. Calyptra cucul-
Source : MNHN. Paris
DIDYMODON IN MEXICO AND CALIFORNIA 407
late, smooth, са, 2.5 mm long. Spores brown, weakly papillose, 12-14(-20) um
in diameter. п = 12, 13. Laminal color reactions (after clearing in сопс. lactic
acid) : Cl (conc. HCl) - green + medium yellow, yellow-brown or yellow-orange-
brown; К (10% КОН) - light to dark red-orange-brown, occasionally light to
deep red-orange or orange, seldom light red-brown or deep yellow-orange;
N (conc. НМОз) - tan to light orange-brown; SE (conc. На $04 - ethanol, 2:1) -
blue-green + deep red or yellow-orange-brown ог red-orange-brown. Illustra-
tion : Pl. IV, fig. 11-14.
The broad decurrencies of the basal margins of the leaves of D. tophaceus
are usually best developed in large plants — smallsized specimens may lack
this characteristic of the species that some authors consider diagnostic (discus-
sion by ZANDER 1978a). The type of the synonym Barbula abbonii has small
lamellae on the medial portion of the leaf bases of many leaves, abaxially; this
has been seen in no other specimen examined. «Didymodon luridus» has mor-
phology similar to D. tophaceus, but is here treated as a variety of D. vinealis,
to which it keys in the key to species (above). Didymodon fallax intergrades
somewhat with D. tophaceus gametophytically in eastern U.S.A., but interme-
diate specimens have not been seen in Mexico or California.
Habitat : soil, rock, calcareous rock, wet areas, riversides, springs, 1100-
2200 m elevation.
Distribution : Mexico : Baja California, Baja California Sur, Chiapas, Chihua-
hua, Coahuila, Durango, Hidalgo, Jalisco, México, Morelos, Nuevo León, Puebla,
San Luis Potosí, Sonora; California.
Range : North, Central and South America, Europe, Africa, and Asia.
8. DIDYMODON VINEALIS (BRID.) ZANDER EMEND. ZANDER, Phytologia
41 :25. 1978.
Barbula vinealis Brid., Bryol. Univ. 1 : 830. 1827. — Barbula fallax var. vinea-
lis (Brid.) Hueb., Musc. Germ. : 327. 1833 — Tortula fallax var. vinealis
(Brid.) De Not., Mem. К. Acc. Sc. Torino 40 : 319. 1838. — Tortula vinealis
(Brid.) Spruce, London J. Bot. 4 : 194. 1845. — Barbula cylindrica var.
vinealis (Brid.) Lindb., Musci Scand. : 22. 1879, nom. Шер. prior. ut spec. —
Barbula cylindrica ssp. vinealis (Brid.) Dix., Stud. Handb Brit. Moss. : 197.
1896.
Barbula rectifolia Tayl., London J. Bot. 5 : 49. 1849, syn. nov. Type : Ecua-
dor, Pichincha, Jameson, 1827 (NY - isotype). — Tortula rectifolia (Tayl.)
Мис. J. Linn. Soc. Bot. 12 : 158. 1869.
Tortula olivacea Besch. ex Mitt., J. Linn. Soc. Bot. 12 : 631. 1869, syn. nov.
Type : Mexico, D.F., prope Santa Fe, Bourgeau 1331 (BM, NY, PC - iso-
types). — Barbula olivacea (Mitt.) Besch., Mém. Soc. Sc. Nat. Cherbourg
16 :181. 1872. t
Barbula bourgaeana Besch., Mém. Soc. Sc. Nat. Cherbourg 16 : 179. 1872,
syn. nov. Type : Mexico, D.F., in monte Zacoalco prope Guadalupe,
Source : MNHN, Paris
408 В.Н. ZANDER
Bourgeau 1321 (BM, NY, PC - isotypes).
Barbula lozanoi Card., Rev. Bryol. 36 : 85. 1909, syn. nov. Type : Mexico :
Puebla, Honey Station, Pringle 10653 (PC - lectotype; TENN - isotype);
Nuevo León, Monterey, Pringle 10459 (BM, FH, РС isosyntypes); Hidal-
go, Tula, Pringle 15216 (FH, PC - isosyntypes).
Barbula salazarensis Thér., Rev. Bryol. Lichénol. 5 (1932) : 96. 1933, syn.
nov. Type : Mexico, D.F., Salazar, St. Pierre 1919 (PC - holotype).
Plants forming turf or cushions, green to dark green, occasionally reddish-
brown above, light brown below. Stems occasionally branching, red-brown,
to 2.0 ст long, rounded pentagonal, central strand present, often strong or
brown or hollow, cortex little differentiated or of 1-2 layers of smaller, thick-
walled cells, hyalodermis absent, axillary hairs of 4-5 cells, basal one brown;
stem occasionally flattened. Leaves when dry appressed-twisted to incurved-
twisted or spreading-flexuose, when wet spreading to spreading-recurved, short-
to long-lanceolate or long-oval, 2.5-3.5(-4.0) mm long, adaxial surface broadly
grooved along the costa, narrowly so near apex; leaf margins recurved at leaf
base to recurved to near apex, entire, unistratose to bistratose in patches or
evenly; leaf apex obtuse to narrowly acute, often apiculate by a smooth, conical
cell; leaf base scarcely differentiated to oblong, basal margins short and broadly
decurrent; costa percurrent to short-excurrent, evenly thick to apex, abaxially
somewhat bulging, ad- and abaxial surficial cells quadrate to short-rectangular,
smooth to papillose, transverse section of costa semicircular to elliptical, adaxial
surface weakly convex to flat, adaxial stereid band absent, usually represented
by thin-walled to substereid cells, guide cells 4(-6) in 1(-2) layers, abaxial stereid
band distinct, strong, ad- and abaxial epidermal layers differentiated, lumens
of abaxial cells quadrate to semicircular; upper laminal cells quadrate, 7-9(-14) um
wide, 1:1, walls thin to evenly thickened, occasionally porose, weakly surficially
bulging, arranged in longitudinal rows, often in distinct patch-work, relatively
homogeneous in size and shape; laminal papillae seldom absent, weakly deve-
loped to strong, simple to bifid, occasionally appearing multiplex, solid or
hollow, centered over the lumens, 1-5 per lumen cach side; basal laminal cells
scarcely differentiated to weakly differentiated medially, quadrate to short-
rectangular, to 10-11 um wide, 1-2:1, walls thin to evenly thickened. Propagula
rare, spherical to elliptical, brown, 21-30(-55) ит long, of 1-2(-8) cells, born in
clusters on crowded branching stalks in leaf axils. Dioicous; perichaetia terminal,
inner leaves oblong-lanceolate, little differentiated, loosely sheathing the seta,
prosenchymatous in lower half; perigonia terminal, often in series, gemmate,
inner leaves ovate. Sporophyte seta 1 per perichaetium, 0.8-1.0 mm long, red-
brown to yellow, weakly twisted clockwise; theca 1.5-2.5 mm long, light red-
brown to brown, nearly smooth when dry, cylindrical, neck absent; exothecial
cells rectangular, 16-20-25) um wide, 4-6:1, walls thin to evenly thickened;
stomates at base of urn, phaneropore; annulus of 1-2 rows of vesiculose cells;
peristome of 32 linear, yellow to orange-red, densely spiculose teeth, 185-
1300 ит long, straight to twisted 0.25-0.50 times, basal membrane low, distinct,
50-70 ит high, spiculose; operculum long-conic, 1.1-1.4 mm long, cells weakly
Source : MNHN. Paris
DIDYMODON IN MEXICO AND CALIFORNIA 409
twisted counterclockwise. Calyptra cucullate, smooth, 2.8 mm long. Spores
yellow-brown, smooth, 9-12 um wide. n — 10,12, 13, 13 + 1m, 14. Laminal
color reactions (after clearing in conc. lactic acid) : Cl (conc. HCl) - green or
yellow-green + light to medium yellow-brown, occasionally light-brown, orange,
or yellow-orange; K (10% KOH) - light to dark red to red-brown, often medium
to deep red-orange-brown, occasionally medium orange-brown; N (conc. HNO3)
- light brown to red-brown, light yellow to orange-brown; SE (conc. Ha SO4-
ethanol, 2:1) - dark green + medium red-brown to dark red, Illustration PL V,
fig. 1-12.
A majority of specimens of Didymodon vinealis have a bright to dark red
color with KOH. Although this is not a constant feature and is occasionally
duplicated in.other species of Didymodon, color in KOH may be used as an
additional character when identifying small specimens of the genus, which
can appear rather characterless. Propagula are rare in D. vinealis. They are
present in var. vinealis (Mexico, Michoacän, Huerta, 1969 - TENN, as D. god-
manianus) and var. luridus (U.S.A. Arizona, Mariposa Co., Zander 4366 -
BUF). Like D. rigidulus, D. vinealis is treated here as a complex of morpholo-
gically intergrading variants that are distinctive in extreme appearance and
some of which have characteristic geographic ranges. Likewise, heterotypic
synonyms are not assigned to the varieties of D. vinealis but are placed with
the species sensu lato. Good summations of some expressions of variation
are given by STEERE (1938) and FLOWERS (1973), both of whom seem
to recognize an intergradation of short- and long-leaved plants.
Habitat : soil, rock, acidic to calcareous situations, moist to dry, sealevel
to 2130 m elevation.
Distribution : Mexico : Baja California, Coahuila, Distrito Federal, Hidalgo,
México, Nuevo León, Puebla; California.
Range : western North America, Central America, South America, Europe,
Asia and Africa.
Key to Varieties of Didymodon vinealis in Mexico and California
1. Leaves short-ovate to ovate-triangular, usually short, to 1.5 mm long... ... pn
1. Leaves long-ovate to long-lanceolate, often long, to 4.0 mm long ....... 3.
2. Upper laminal cells thin-walled, papillose, apex blunt and often apiculate
by а conical cell . „D. vinealis var. brachyphyllus (Sull. in Whipple) Zander
2. Upper laminal cells with evenly thickened walls, smooth, apex acute to
zoudded. s ss lees PECORIS ..... D. vinealis var. luridus Hornsch.
3. Upper laminal cells large, 10-13 um wide ............... AE |
"ые e РОТ .,.,.. D. vinealis var. rubiginosus (C. Muell.) Zander
3. Upper laminal cells small, 7-10 um wide. ........... baron e gn E 4.
4. Leaves long-ovate to broadly lanceolate, apex blunt to broadly acute,
upper marginal cells evenly bistratosé or occasionally bistratose in patches;
laminal color reaction K deep yellow- or orange-brown, seldom red-brown .
TR a Praia лее D. vinealis var. nicholsonii (Culm.) Zander
Source : MNHN, Paris
410 R.H. ZANDER
4. Leaves narrowly lanceolate, apex narrowly acute, upper marginal cells
unistratose to bistratose in patches; laminal color reaction К deep red to
red-brown ..,. ‚ы да ES o
5. Leaves long, often to 4.0 mm, twisted ог flexuose above, margins recurved
alongleaf bas D, vinealis var. flaccidus (B.S.G.) Zander
5. Leaves shorter, to 3.0 mm, not or little twisted, margins usually recurved to
above midleaf. SPO ЙИ coo. D. vinealis (Brid.) Zander var. vinealis
PL V,—1-3 : Didymodon vinealis var, vinealis. 1 : Leaf 2 : Leaf apex. 3 : Upper marginal
cells. 4 : D. vinealis var. brachyphyllus, leaf. 5 : D. vinealis var. luridus, leaf. 6-7 :D.
vinealis var. rubiginosus. 6 : Leaf, 7 : Transverse section. 8 : D. vinealis var. flaccidus,
leaf. 9-12 : D. vinealis var. nicholsonii. 9-10 : Leaves. 11 : Transverse section near
apex. 12 : Transverse section near midleaf. Magnifications : leaves x 43; leaf apices,
upper marginal cells, transverse sections x 100; propagula x 100; capsules and peri-
stomes x 28.
Source : MNHN. Paris
DIDYMODON IN MEXICO AND CALIFORNIA 411
— Didymodon vinealis (Brid.) Zander var. vinealis
Leaves lanceolate, to 3.0 mm long; leaf apex acute, narrowly grooved to
terete; leaf margins usually recurved to above midleaf; upper laminal cells
7-10 ит wide, nearly smooth to highly papillose, often thick-walled, upper
marginal cells unistratose or bistratose in patches. n = 14. Illustration : Pl. V,
fig. 1-3.
This taxon represents central or intermediate morphology (from which the
other varieties may be conceptually derived by elaboration or reduction with
minimum morphological change). No such morphologically central taxon is
recognized here for the other major complex, D. rigidulus s. lat.; however, if
sure characters can be found to distinguish a D. rigidulus-related variant from
D. vinealis var. luridus as conceived here, then D. rigidulus var. gracilis would
represent that central morphology within D. rigidulus s. lat.
Habitat : soil, rock, sea level to 2130 m elevation,
Distribution : Mexico : Baja California, Coahuila, Distrito Federal, Hidalgo,
México, Nuevo León, Puebla; California.
Range : western North America, Central and South America, Europe, Asia,
Africa.
— Didymodon vinealis var. brachyphyllus (бий. in Whippl.) Zander, comb. nov.
Barbula brachyphylla бай. in Whippl., Rep. Pacif. Вайт. Surv. Bot. 4 : 186.
1856, basionym. — Didymodon brachyphyllus (бий. in Whippl.) Zander,
Phytologia 41 :24. 1978.
Leaves ovate to ovate-triangular, to 1.5 mm long; leaf apex broadly acute to
obtuse, usually apiculate, often rather cucullate; leaf margins usually recurved
to near apex; upper laminal cells 7-10 um wide, weakly papillose, thin-walled,
upper marginal laminal cells unistratose to occasionally bistratose in patches.
n = 10, 12. Illustration : Pl. V, fig. 4. -
This variety is closely related to, or may be the same as D. reedii Robins. of
eastern U.S.A. and D. tectorum (C. Muell.) Saito of Asia.
Habitat : soil, rock, 135-1600 m elevation.
Distribution : Mexico : Distrito Federal; California.
Range : Mexico; western U.S.A. : Washington, Oregon, California, Montana,
Utah, Colorado, Arizona (cf. LAWTON 1971); Canada : British Columbia,
Specimens examined : Mexico : D.F. near Santa Fé, Bourgeau 1331 (BM,
Y). U.S.A. : California : s. loc., Bigelow s. n., Sull. & Lesq., Musc. Bor. Amer.
100c (CU); San Diego Co., Anza Borrego Desert State Park, Stark 472-D (PAC):
Riverside Co., Joshua Tree National Monument, Norris 50513 (HSC); Tehama
Co., near Manton, bluff over Battle Creek, Norris 21332 (HSC). Utah : Millard
Co. Boardman U.S. Navy Bombing and Gunnery Range, just S of Boardman,
Christy 1398-2 (herb. J.A. Christy; BUF). Canada : British Columbia : S side of
Thompson R., between Cache Creek and Savonah, Wallachin, Roadside, McIn-
tosh & Kruckeberg 5250 (UBC).
Source : MNHN, Paris
412 К.Н. ZANDER
= Didymodon vinealis var. luridus (Hornsch. in Spreng.) Zander, comb. nov.
Didymodon luridus Hornsch. in Spreng., Syst. Veg. 4 (1) : 173. 1827,
basionym. — Barbula lurida (Hornsch. in Spreng.) Lindb., Musci Scand. :
22. 1879. — Didymodon trifarius var luridus (Hornsch. in Spreng.) Mont.,
Arch. Bot. 1 : 139. 1833. — Trichostomum luridum (Hornsch. in Spreng.)
Spruce, Ann. Mag. Nat. Hist. Ser. 2, 3 : 379. 1849.
Leaves short-lanceolate to ovate-triangular; leaf apex broadly acute to roun-
ded-obtuse; leaf margins usually recurved to above midleaf, often strongly
recurved above; upper laminal cells 7-10 um wide, smooth to weakly papillose,
walls usually irregularly thickened, lumens often rounded or triangular to penta-
gonal; upper marginal cells unistratose. Illustration : Pl. V, fig. 5.
Plants with similar leaf morphology but elongate adaxial surficial costal
cells are D. tophaceus. The leaf apex of var. luridus usually has an adaxial costal
groove, the costa is about equally wide to the apex, and at least European
specimens are mostly bright red in color with KOH — all characteristics fairly
constant in the D. vinealis group. It may well be that variants of D. rigidulus
— which is occasionally red in KOH — with short, blunt leaves may be here
included in var. luridus, but, to date, no sure way of distinguishing these has
been found,
CRUM et al. (1973) and ZANDER (1978a) did not recognize D. luridus
in North America north of Mexico, referring most collections identified as this
to D. tophaceus (with acute leaf apices). After seeing more collections and using
laminal color reactions to some extent, I present here a concept of the taxon
somewhat different and at a different taxonomic level than that of FLOWERS
(1973), STEERE (1938) and other American authors who have treated it under
the name D. trifarius (Hedw.) Roehl., they indicating that it has a close relation-
ship with D. tophaceus, which it has not. Material from U.S.A. : Washington,
California, Arizona; Mexico and Guatemala matches that from Europe in BUF
and keys to D. vinealis in the above key to species of Didymodon.
Didymodon luridus is often placed in the synonymy of D. trifarius. Didymo-
don trifarius (Hedw.) Roehl. is based on Cynodontium trifarium Hedw. HEDWIG
(1801) characterized the genus Cynodontium as «Flos terminalis hermaphrodi-
tus (i.e. monoicous)», including what are now Didymodon trifarius, Distichium
capillaceum (Hedw.) B.S.G., D. inclinatum (Hedw.) B.S.G. and Bryum uligino-
sum (Brid.) В.5.С. (fide VAN DER WIJK et al. 1959-1969). The last two taxa
are autoicous and D. capillaceum is paroicous, but species of Didymodon are
always dioicous. HEDWIG included references to the following elements : speci-
mens «Upsaliae legit Ehrharty and «in Anglia, Scotia Dicksonus,» and a refe-
rence «Hedw. St. Cr. р. 76 t. 28. Swartzia trifaria. Brid. Muscol. П p. 120» to
his (HEDWIG 1789) description and illustration of «Swartzia trifariay. The
BRIDEL reference (BRIDEL 1798) cites other authors but these lead back
to HEDWIG (KOPONEN 1979),
KOPONEN (1979) quite rightly noted that my assumption (ZANDER
1978c) was incorrect that the specimen labeled «Swartzia trifariay in Hed-
Source : MNHN. Paris
DIDYMODON IN MEXICO AND CALIFORNIA 413
wig's herbarium at С was the holotype of the species, because other elements
were mentioned in the protologue. He indicated that it may be possible to
choose a lectotype for Didymodon trifarius in such a way as to preserve current
usage. The simple specimen labeled «Swartzia trifaria in G-Hedwig proved to be
taxonomically Saelania glaucescens (Hedw.) Broth, (ZANDER 1978c). In an
effort to track down additional elements, I was permitted to examine the ori-
ginal herbarium of J. Dickson at BM. The only specimens possibly a syntype
was labeled (but not in Dickson's handwriting — see Pl. УП, fig. 4) «Weisia
trifaria». This specimen was taxonomically D. tophaceus. 1 was also allowed to
examine specimens (Pl. VI, fig. 1-3) at the J.E. Smith herbarium (LINN), which
Pl. VI. — 1-3 : «Swartzia trifaria» collections — sheet 1671.1 — in herbarium of J.E. Smith
at LINN 1 : The specimens, of which numbets 1 and 3 may be isosyntypes of Cynodon-
tium trifarium Hedw. 2 : Close up of the Ehrhart 174 plants. 3 : Label for sheet 1671.1.
4 : Possible isosyntype of Cynodontium trifarium Hedw. in herbarium of J. Dickson at
BM.
Source : MNHN. Paris
414 В.Н. ZANDER
included а specimen labeled «Mr. Dickson. 1802. «Bryum trifarium»» and ano-
ther labeled both «Ehrh. Crypt. 174. Herb. Davall. 1802» and «174. Swartzia
trifaria Ehrh. Upsaliae ». These may or may not be syntypes as «1802» may refer
to date of collection or date of communication (but probably the latter). Regu-
lations at LINN do not allow dissection or examination of specimens with
a high-power microscope; however, the Dickson specimen was similar to that
in his herbarium, and the Ehrhart specimen matched that in С, With the dissec-
ting microscope available at LINN, the latter looked quite like specimens at BUF
of Saelania glaucescens with rather short capsules. The illustrations of S. glau-
cescens and «Swartzia trifaria» of HEDWIG (1789, 1792) are very similar
(contrary to the view of KOPONEN 1979), except for capsule shape (anyway
variable in S. glaucescens) and flexion of peristome teeth (straight when dry
but variably incurved at first when wet). The exsiccatum of Ehrhart 174 in
Ehrhart's original herbarium at GOET is not available by loan for study. My
discussion (ZANDER 1978c) of past use of the name D. trifarius by early
authors after HEDWIG (1801) shows that many regarded D. trifarius and D.
luridus as different. Some thought the former name was a nomen ambiguum.
Others ignored it. LINDBERG (1864) apparently examined the Ehrhart syntype
and placed it in the synonymy of Trichostomum viridulum Bruch in F. Muell.
(as T. crispulum var. angustifolium B.S.G.) (perhaps a mixture is involved ? ).
Inasmuch as major isosyntype specimens (1) are unavailable for study, ог
labeled in an ambiguous fashion, or apparently mixed or scattered, or their
identification is hampered by overly restrictive curatorial regulations, (2) the
protologue of the first valid publication (1801) surely requires a lectotype with
a monoicous condition (preferably synoicous — Saelania glaucescens is autoi-
cous, but HEDWIG 1789 may have made a mistake in this although this is most
unlikely), and (3) Didymodon trifarius (in the modern sense) is not known for
Finland (KOPONEN 1979) but S. glaucescens is (KOPONEN et al. 1977), it
seems certain that no bryologist in the near future will sucessfully designate a
lectotype for D. trifarius that will preserve current usage. Following the lead
of DIXON (1924) and LIMPRICHT (1890), I here treat the name D. trifarius
as a nomen ambiguum. The unambiguous name D. vinealis var. luridus is appro-
priate for that taxon of Didymodon commonly identified as «D. trifarius» in
European herbaria but seldom so in American herbaria.
The J.E. Smith herbarium (LINN) is extraordinary, by the way, for its
richness in welllabeled (Pl. УП, fig. 2) early specimens, many of which are
possible isotypes or are otherwise representative of taxonomic concepts of
early authors of bryophyte names. Some persons who apparently corresponded
with Smith and sent him specimens include : Davall, Davies, Dickson, Hedwig,
Hooker, Humboldt, Linneus, Mackay, Menzies, Muehlenberg, Schrader, Swartz,
Torrey and Turner. The Smith bryophyte herbarium is presently under the
same regulations as those for the Linnean collections at LINN. These allow
low-power microscopic examination only (without manipulation or dissection
of the specimen) and are not appropriate for taxonomic study of bryophytes.
One might hope that the Smith herbarium might someday be transferred to
BM where facilities are more suited to the needs of specialists on the bryophytes.
Source - MNHN. Paris
DIDYMODON IN MEXICO AND CALIFORNIA 415
Pl, ҮП, — 1 : Sample specimens in J. Dickson herbarium at BM showing typical lack of
collection data. 2 : Sample label of a sheet in J.E. Smith herbarium at LINN showing
relative excellence of collection data.
The name Barbula lurida Hornsch. in Mart. may be confused with Didymo-
don luridus Hornsch. in Spreng. The former is based on a different type (from
Source : MNHN, Paris
416 А.Н. ZANDER
Brazil) and the specimen reported (CRUM 1951) under this name from Mexico
(Nuevo León, Harvey 1061 - MICH) is Barbula arcuata Griff.
Habitat : soil, rock, riverside, dry to moist areas, to 1570 m elevation.
Distribution : Mexico : México; California.
Range : western North America, Central America, Europe, Middle East, Asia,
Africa.
North American specimens examined : U.S.A. : Washington, Klickitat-Fran-
klin counties border, ca. 48 km E of Plymouth, soil and steppe, McIntosh &
Kruckeburg 4902 (UBC). Arizona, Cococino Co., Oak Creek Gorge, West Fork,
Zander 4150 (BUF). California, Lassen Co., 24 km S of Adin, limestone out-
crop, Norris 21462 (HSC); Tehama Co., Woodson Bridge State Park, Norris
50430 (HSC). Mexico : México, Tlalmanalco, 3 km E of San Rafael, Cárdenas 56
(MEXU, TENN). Guatemala : Quetzaltenango, below Zunil, along Rio Samalá,
Steyermark 34985 (FH).
— Didymodon vinealis var. nicbolsonii (Culm.) Zander, comb. nov.
Didymodon nicholsonii Culm., Rev. Bryol. 34 : 100. 1907, basionym.
Type : England, Sussex, Wild Brooks, Amberley, Nicholson, 1905 (BM). —
Barbula nicholsonii Culm., Rev. Bryol. 34 : 100. 1907. — Didymodon
rigidulus var. nicholsonii (Culm.) Roth, Hedwigia 50 : 303. 1911. — Barbu-
la rigidula ssp. nicholsonit (Culm.) Dix., Stud. Handb. Brit. Moss. ed. 3 :
413. 1924, — Didymodon luridus ssp. nicholsonii (Culm.) Loesk., Ber.
Naturh. Ver. Preuss. Rheinl. Westf. 1932-33 : 16. 1934. — Didymodon
luridus var. nicholsonii (Culm.) Loesk. in WIRTGEN, Herb. Fl. Rhen.
n. 826. 1934. — Didymodon trifarius ssp. nicholsonii (Culm.) Wijk A
Marg., Taxon 7 : 289. 1958.
Leaves rigid, long-ovate, occasionally ovatelanceolate, to 3.5 mm long;
leaf apex broadly acute to blunt, somewhat cucullate; leaf margins recurved
to near apex; upper laminal cells 7-9 um wide, weakly papillose, walls evenly
thickened, upper marginal cells evenly bistratose or nearly so. Illustration :
Pl. V, fig. 9-12.
Except fot the type specimen, which is K red-brown, the laminal color
reaction of this variety is K orange-brown to red-orange-brown. The var. nichol-
sonii is fairly sharply differentiated from var. luridus by its evenly bistratose
upper marginal cells, but it intergrades with var. vinealis. Didymodon rigidulus
var. incrassatolimbatus is very similar in appearance to var. nicholsonii; however,
the latter differs var. incrassatolimbatus by its filamentous, yellow peristome
(vs. the long-triangular, red-orange peristome of var. incrassatolimbatus); peri-
chaetial leaves ovate-lanceolate, weakly differentiated (vs. narrowly lanceolate,
unlike the cauline leaves); cauline leaves keeled (vs. plane); leaf apex somewhat
cucullate, occasionally apiculate by one conical cell (vs. apex plane, costa
short-excurrent as a cone or cylinder); costa bulging dorsally, brown (vs. little
bulging dorsally, green); and adaxial stereid band represented only by thin-
walled or substereid cells (vs. stereid cells usually present adaxially).
Source - MNHN. Paris
DIDYMODON IN MEXICO AND CALIFORNIA 417
Contrary to indication in the Index Muscorum (VAN DER WIJK et al.
1959-69), Didymodon nicholsonii Culm. was validly published together with
Barbula nicholsonii Culm., being one of two «alternative names» (I.C.B.N.,
Art. 34).
Habitat : soil, rock, dry to moist situations, 90-1370 m elevation.
Distribution : California.
Range : western North America, western Europe. Reported here as new to
the New World.
Specimens examined : U.S.A. California : Humboldt Co., Mad R., near Maple
Creek, Schofield 28662 (UBC); Mendocina Co., Navaro R. near entrance to
Hendy Woods State Park, Norris 53101 (ОВС); Ventura Co., Howard Creek
Trail, Norris 55495 (HSC); Chorro Grande Canyon, Norris 55543 (HSC). Ore-
gon : Lake Co., Lakeview, canyon at end of Center St., streamside, Zander &
Eckel 4422 (BUF), Canada. British Columbia : S of Cristina Lake, Kettle R.
Gorge, Tan & Teng 78-119 (UBC). England. Gloucestershire : Symonds Yat,
Townsend, 1957 (US); Northumberland : Linshiels, Coquet R., Long 4903 (E).
Germany (BRD). Neuwied, Düll 7.4.1979 (BUF).
— Didymodon vinealis var. rubiginosus (Mitt.) Zander, comb. & stat. nov.
Barbula rubiginosa Mitt., J. Linn. Soc. Bot. 8 : 27. 1864, basionym. —
Didymodon occidentalis Zander, Phytologia 41 : 26. 1978, non Didymo-
don rubiginosus (C. Muell.) Broth., Nat. Pfl. 1 (3) : 405.1902.
Leaves rather rigid, long-lanceolate, to 3.0 mm long; leaf apex narrowly
acute, grooved to terete, margins recurved to above midleaf, upper laminal
cells relatively large, 10-13 ит wide, weakly papillose, pellucid, walls evenly
thickened to somewhat porose, upper marginal cells unistratose or more com-
monly bistratose in patches or entirely. Illustration : Pl. V, fig. 6-7.
This taxon is confined to the West Coast of North America. It is characte-
rized by the acuminate leaf apex that is often bistratose marginally or entirely,
occasionally somewhat notched and appearing fragile, and the rather large upper
laminal cells. The peristome often appears to be absent, but actually is removed
with the operculum. This and the character of quadrate basal laminal cells are
variable and present in other varieties occasionally. LAWTON (1971) notes that
a rudimentary peristome is occasionally present.
Habitat : bark, rock, soil, wet to dry situations, to 1100 m elevation.
Distribution : California.
Range : I have seen specimens from Canada : British Columbia; U.S.A, : Ore-
gon and California; LAWTON (1971) reports it also from Washington and Idaho.
— Didymodon vinealis var. flaccidus (B.S.G.) Zander, Phytologia 41 :25. 1978.
Barbula vinealis var. flaccidus B.S.G!, Bryol. Eur. 2 : 86. 1842 (fasc. 13-
15 Mon. 24).
Leaves long-lanceolate, to 4.0 mm long, often flexuose or twisted when wet;
Source - MNHN. Paris
418 В.Н. ZANDER
leaf apex narrowly acute to blunt, grooved adaxially; laminal margins plane above
midleaf; upper laminal cells 7-10 um wide, usually distinctly papillose, walls
usually thin, upper marginal cells usually unistratose. n = 13 or 13 + 1m. Illus-
tration : Pl. V, fig. 8.
The var. flaccidus is red-brown with KOH. I agree with STEERE's (1938)
evaluation that «the longer and much more flexuous leaves, which are less
strongly revolute, and the more papillose cells...» are characteristic of this
taxon; STEERE, who treated this under the name Barbula cylindrica (Tayl.)
Schimp., also recognized a degree of intergradation with what is here var. vinea-
lis, such being more apparent in the south of its range in North America.
Habitat : soil, rock, to 3000 m elevation.
Distribution : California.
Range : western U.S.A., western Canada, Europe, northern Africa, Asia.
EXCLUDED NAMES
Barbula and Didymodon species reported for Mexico and California that
have not been previously treated (ZANDER 1978a, b, 1979) are discussed
below, together with a few other names of taxa from Central America or the
Antilles.
Barbula lonchostega C. Muell., Bull. Herb. Boiss. 5 : 195. 1897. Type: Guatemala,
between Cubulco and Soyabaji, Bernoulli & Cario 104 (NY - isotype). = Bar-
bula indica (Hook.) Spreng. emend. Zander (1979) (syn. nov.).
Barbula graminicolor C. Muell., a synonym of Didymodon australasiae (Hook. &
Grev.) Zander, was reported from Mexico (BESCHERELLE 1872) on the
basis of a specimen (Veracruz, Orizaba, Liebmann s. n. - BM!) that is actually
Barbula orizabensis C. Muell.
Barbula lurida Hornsch. in Mart. This is a Brazilian species reported (CRUM
1951) from Mexico based on a specimen (Nuevo León, Hacienda Hermosa,
Harvey 1061 - MICH!) that proved to be Barbula arcuata Griff. The former
name may, however, be a synonym and predates the latter.
Barbula saint-pierrei Ther., Rev. Bryol. et Lichénol. 5 (1932) :97. 1933. Туре:
Mexico, D.F., Cuautzin, Saint-Pierre 1879 (NY, PC - isotypes). — Bryoery-
throphyllum ferruginascens (Stirt.) Giac. (syn. nov.).
Didymodon filicaulis Card., Rev. Bryol. 37 : 126. 1910. Type : Mexico, México,
Ixtaccihuatl, Purpus 3721 (PC - holotype; NY - isotype). = Bryoerythrophyl-
lum recurvirostre (Hedw.) Chen var. recurvirostre (syn. nov.), malformed
leaves.
Didymodon juniperinus (C. Muell.) Broth., a Peruvian species, was reported
(BOWERS et al. 1973) from Mexico on the basis of a specimen (Puebla,
Sharp 948-a - ТЕММ!) that proved to be Oxystegus tenuirostris (Hook. &
Tayl.) A.J.E. Sm. var. tenuirostris.
Source : MNHN. Paris
DIDYMODON IN MEXICO AND CALIFORNIA 419
Didymodon planifolius P. Varde & Thér., Rev. Bryol. Lichénol. 14 : 11. 1944.
Type : Haiti, Morne de la Selle, Eckman 3178 (MICH). — Trichostomum
brachydontium Bruch in F.A. Muell. (syn. nov.).
Husnotiella obtusifolia (Hampe) Zander. This combination (ZANDER 1977)
has proved to be inappropriate and the correct name for the taxon is Gyro-
weisia obtusifolia Broth. (nom. nov. for Trichostomum obtusifolium Hampe,
1870, hom. Шер. non T. obtusifolium P. Beauv., 1805). Relationship to the
genus Husnotiella (treated here as a synonym of Didymodon and for which
the generitype is D. revolutus) was misevaluated because of similarities in leaf
shape and papillae; mixture of G. obtusifolia and D. revolutus in the type of
G. papillosa Thér., a synonym, and in other specimens; both species have
leaf margins bistratose in small patches, convex adaxial costal surfaces and
quadrate .adaxial costa cells; and because of the fact that specimens of D.
revolutus may rarely have plane leaf margins. Gyroweísia obtusifolia differs
from D. revolutus by the following combination of characters : capsule
microstomous, annulus revoluble or adherent; peristome red-brown, some-
what twisted, teeth long, filamentous and acute; leaves ligulate, plane, basal
cells enlarged, swollen, hyaline and thin-walled; adaxial costal cells short-
rectangular to elongate; costa with two stereid bands (the adaxial one occasio-
nally substereid), Although my comments (ZANDER 1977) that Gyroweisia
is a «wastebasket» genus still hold, С. obtusifolia can be refered to the genus
Gyroweisia by the following combination of characters : annulus often
revoluble; peristome teeth short; leaves occasionally cucullate, margins plane,
basal cells much enlarged; costa subpercurrent, adaxial costal cells occasio-
nally elongate: upper laminal cell papillae several over each lumen each side.
SUMMARY OF NEW SYNONYMS
(Basionyms arranged by epithet; emendations refer to this paper unless otherwise noted)
Barbula altiseta Card. =Didymodon rigidulus Hedw. emend. Zander
B. bescherellei var. crassinervia Thér. =D. rigidulus Hedw. emend. Zander
B. bescherellei var. stenocarpa Card. =D. rigidulus Hedw. emend. Zander
В. bourgaeana Besch. — D. vinealis (Brid.) Zander emend. Zander
D. craspedophyllus Card. = D. australasiae (Hook. & Grev.) Zander emend. Zander
B. erythropoda Schimp. ex Besch. =D. rigidulus Hedw. emend, Zander
D. filicaulis Card, = Bryoerythrophyllum recurvirostre (Hedw.) Chen var. recurvirostre
B. flaccidiseta Lor. =D. rigidulus Hedw. emend. Zander
D. fusco-viridis Card. = D. rigidulus Hedw. emend. Zander
B. godmaniana C. Muell. =D. rigidulus Hedw. emend. Zander
B. gracilescens Schimp. ex Besch. — D. rigidulus Hedw. emend. Zander
B. graciliformis Schimp. ex Besch. =D. rigidulus Hedw. emend. Zander
B. graminicolor C. Muell. = D. australasiae (Hook. & Grev.) Zander emend. Zander
D. heribaudii Card. = D. rigidulus Hedw. emend. Zander
B. lagunicola C. Muell. =D. rigidulus Hedw. emend. Zander
B. leptocarpa Besch. = D. rigidulus Hedw. emend. Zander
B. lonchostega C. Muell. = В. indica (Hook.) Spreng. emend. Zander (1979)
B. lozanoi Card. — D. vinealis (Brid.) Zander emend. Zander
D. mexicanus Besch. = D. rigidulus Hedw. emend. Zander
Source : MNHN, Paris
420 В.Н. ZANDER
B. mobilis C. Muell. =D. rigidulus Hedw. emend. Zander
Tortula olivacea Besch. ex Миг. =D. vinealis (Brid.) Zander emend. Zander
D. patentifolius Thér. = Р. australasiae (Hook. & Grev.) Zander emend. Zander
D. planifolius P. Varde & Thér. = Trichostomum brachydontium Bruch in F.A. Muell.
D. pusillus Card., hom. illeg. =D. rigidulus Hedw. emend. Zander
Trichostomum ramulosum Schimp. ex Besch. = D. rigidulus Hedw. emend. Zander
rectifolia Tayl. = D. vinealis (Brid.) Zander emend. Zander
rigidula Besch., hom. Шер. =D. rigidulus Hedw. emend. Zander
saint-pierrei Thér. — Bryoerythrophyllum ferruginascens (Stirt.) Giac.
salazarensis Thér. = D. vinealis (Brid.) Zander emend. Zander
strictidens C. Muell. — D. rigidulus Hedw. emend. Zander
subteretiuscula Card. =D. rigidulus Hedw. emend. Zander
teretiuscula Schimp. ex C. Muell, =D. rigidulus Hedw. emend. Zander
torquescens Card. =D, australasiae (Hook. & Grev.) Zander emend. Zander
viridissimus Card. — D. rigidulus Hedw. emend. Zander
sect. Craspedophyllon Card. =D. sect. Asteriscium (C. Muell.) Zander
BLESSE EEE
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DIDYMODON IN MEXICO AND CALIFORNIA 421
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423
ESTUDIO BRIOSOCIOLOGICO
DE LAS COMUNIDADES REOFILAS DE SIERRA NEVADA
(ESPANA)
J.A. GIL & J. VARO*
RÉSUMÉ. — On fait une étude bryosociologique des groupements rhéophiles de la Sierra
Nevada, avec la description des associations suivantes : Scapanietum undulatae Schwickerath
1944, Dichodontietum pellucidi v. Hübschmann 1967, Fontinaleto-Pachyfissidentetum
grandifrondis W. Koch 1936, à rattacher à la classe Platyhypnidio-Fontinaletea Philippi
1956.
De méme on a étudié les groupements rheophiles alpins de la classe Hygrohypnetea v.
Hübschmann 1957, avec les associations Schistidio-Hygrohypnetum dilatati Geissler 1976
et Solenostomo-Hygrohypnetum Geissler 1976.
GENERALIDADES
1. Geología y climatología. — El desarollo de Sierra Nevada como conjunto
montañoso netamente individualizado, tiene lugar principalmente durante el
Plioceno y, es en el Cuaternario cuando los períodos activos periglaciares deter-
minan el decisivo modelado de su relieve. El resultado es una zona 6 región
central constituida por micasquistos, que poseen un considerable contenido
en granates y grafitos, además de cuarcitas, gneis, anfibolitas, serpentinitas y
mármoles, materiales de edad anterior a la era secundaria; este nücleo central
queda envuelto por una zona periférica u orla calizo-dolomítica del Triásico.
Las diferencias altitudinales determinan las lógicas variaciones climáticas;
asi, las zonas comprendidas en el piso supramediterraneo (RIVAS MARTINEZ,
1981) registran una precipitación media anual de 650 mm (a 1084 m), mientras
las zonas situadas en el piso oromediterráneo (2000 m) suelen sobrepasar los
D
* Departamento de Botánica, Facultad de Ciencias, Universidad de Granada, España,
Cryptogamie, Bryol. Lichénol., 1981, 2, 4 : 423-440.
Source : MNHN. Paris
^ J.A. GIL & J. VARO
900 mm (по se dispone de datos pluviométricos fiables para el crioromediterrá-
neo, es decir, entre 2800 y 3000 m). La temperatura media anual del piso
mesomediterráneo es de 15,8°C (a 900 m), descendiendo a 4,7°C (a 2700 m)
en el límite entre los pisos oro- y crioromediterráneo, Para las zonas situadas
entre 2000 y 2500 m la duración de la nieve comprende de cuatro a cinco
meses al año, mientras que en altitudes superiores oscila entre siete y ocho
meses y permaneciendo de forma permanente tan sólo en algunos ventisqueros.
2. Hidrografía e hidrología. — La mayoría de los cursos nevadenses tienen
sus nacimientos a partir de lagunas que proceden del deshielo estando situadas,
entre 2700 y 3000 m, al pié de accidentadas crestas que denotan las huellas
dejadas en la montaña por los glaciares cuaternarios del Pleistoceno; los ríos
procedentes de ellas muestran caracteres erosivos incompletos, de lo que se
deduce la fase de inmadurez que aún poseen, susceptibles por tanto de trans-
formarse y desgastarse todavia más.
En las cuencas nevadenses el deshielo actúa como regulador del caudal, ya
que las cantidades sustraidas en los meses de máxima precipitación son luego
aportadas en los meses de mayor sequía. Los ríos nevadenses poseen pues, un
régimen de tipo pluvial pero de intensa influencia nival en verano, haciéndose
torrenciales y de caudal constante en sus tramos más elevados (entre 2000 y
3000 m); sin embargo en los tramos medio y basal, el caudal experimenta
notables variaciones desde principios a final del estío, También se aprecia un
fuerte contraste respecto a las temperaturas y calidades quimico-biolögicas
de las aguas, frías (rara vez superan 4°C), it (РН = 4,5-5) y oligótrofas
al principio, haciendose después más cälidas (10°C), mesötrofas y de pH próximo
a la neutralidad.
ESTUDIO BRIOSOCIOLOGICO
Los trabajos briosociológicos realizados por KRAJINA (1933), HÜBSCH-
MANN (1953, 1957, 1967, 1973), PHILIPPI (1956, 1965), HERTEL (1974) y
GEISSLER (1976), constituyen la base de nuestro estudio sobre las comuni-
dades reöfilas nevadenses, agrupándolas para ello en las clases Platyhypnidio-
Fontinaletea Philippi 1956 e Hygrohypnetea v. Hübschmann 1957.
Ambas clases se caracterizan por ser hidrofilo-reófilas y pioneras en la colo-
nización de cursos de agua, Son pobres en especies y particularmente sensibles
a diversos factores limitantes (contaminación, fuertes oscilaciones del caudal,
variaciones del pH, etc), por lo que habitualmente presentan un desarrollo
fragmentario.
GEISSLER (1976) ha establecido en los Alpes suizos una disposición catenal
de ambas clases, localizändose la clase Hygrohypnetea entre los pisos alpino y
subalpino centroeuropeos (no desciende por debajo de 1300 m), mientras que la
clase Platyhypnidio-Fontinaletea se halla relegada a los pisos montano y colino.
Source : MNHN. Paris
COMUNIDADES REOFILAS DE SIERRA NEVADA 425
Esta situaciôn no se reproduce tan exactamente en Sierra Nevada, pues а partir
de los inventarios levantados por encima de 2400 m, se puede comprobar la
presencia de una u otra clase con independencia de su altitud. Probablemente la
latitud del macizo nevadense determina que ambas clases se hallen interpuestas
en el seno de los pisos oromediterráneo y crioromeditarráneo, actuando como
factor limitante en el establecimiento de las mismas la mayor o menor torren-
cialidad del agua.
1. CLASE PLATYHYPNIDIO-FONTINALETEA Philippi 1956
Las comunidades reofilo-acidéfilas del piso montano centroeuropeo se han
incluido tradicionalmente en la clase Brachythecietea plumosi v. Hübschmann
1957, así como las basöfilas de llanuras en la clase Fontinaletea antipyreticae
v. Hübschmann 1957; no obstante, a partir de los estudios realizados por PHI-
LIPPI (1956) y HERTEL (1974), ambas clases podrían quedar reunidas en una
única clase Platyhypnidio-Fontinaletea Philippi 1956. Dicha clase está repre-
sentada en Sierra Nevada por los ordenes Brachythecietalia plumosi Philippi
1956 y Leptodyctietalia прати Philippi 1956.
1.1. ORDEN BRACHYTHECIETALIA PLUMOSI Philippi 1956
Se halla representado por la alianza de comunidades sumergidas y acidófilas
Scapanion undulatae Philippi 1956. La alianza de comunidades anfibias y acidó-
filas Racomitrion acicularis Krusenstjerna 1945 no alcanza su óptimo en Sierra
Nevada, pues el deshielo mantiene constante el nivel de los ríos en los tramos
superiores, así como el carácter básico del agua en los tramos basales impide
el establecimiento de la comunidad. La alianza Scapanion undulatae está repre-
sentada a su vez por la asociación Scapanietum undulatae Schwickerath 1944.
A) ASOCIACION SCAPANIETUM UNDULATAE Schwickerath 1944
Estructura y composición florística ; tab. 1. Origen de los inventarios :
tab. 1. № 1, 11 : 2840, 2800 m - Rio Dílar : 4.9.74. № 2, 3, 14 : 2380, 3000,
2850 m - Rio Seco : 6.8.75, N° 4, 5, 6 : 2450, 2600, 2400 m - Chorreras del
Monte Cuna : 26.7.75. N° 7, 8, 9 : 1800, 1860, 2900 m - Arroyo de la Ragua :
21.6.76. N° 10, 12 : 2700, 2900 m - Siete Lagunas : 13.8.76. № 13 : 2700 m -
Rio Trevélez : 13.8.76.
Sinecología. — Comunidad acidéfila y fotéfila, de aguas frías que rara vez
sobrepasan los 15°C de temperatura media, y que no soporta fuertes variaciones
hidrostáticas; es por esto que suele situarse sobre esquistos sumergidos o semi-
sumergidos, que constituyen los tramos superiores de los ríos de Sierra Nevada
(1800-3000 m), preferentemente en estaciones donde la escorrentia no es
excesivamente torrencial. 4
Sinfisionomía. — Comunidad abierta (cobertura media = 619/o), para una
Source : MNHN. Paris
426 Т.А. GIL & J. VARO
Minero de invgntario ue He pe E E ИРИ Ce
Superficie (dm?) 12 10 16 10 12 16 15 12 18 11 12 25 18 16
Cobertura(x) 60 40 40 30 80 80 70 то 70 60 60 60 60 80
Exposición WR OQ SE з E Mi apo SO EK OM e
Tnelinaciön(®) 90 10 20 0 45 45 45 90 90 90 10 60 90 60
Sustrato Xj eM) Or Им Жы ы Me Go Музеи (М
Número de especies hart ЧЕ de d а:
Característica de asociación y alianza(Scopontetum unduLotae, Scapanion
UnduLotae!
Scapanta undulata TOUR TE RE IS E
Características de unidades superiores (Brochythecietolio pLumosL, PLotuhupnidio-
FontinoLeteo] :
Brachythectun rtvutore Е JE, Щщ. EN
SchLstidlum rivulare WT ў > R Ps o
Fontinalis ontipyretica sut S 4 Е ae > ptt
GTA ME RE Zë № ле н. ër a Ae, т 1
Dichodontiun pellucidum ee Ma Of tero er К x
Compañeras
Bryum schleichert Май» т... PR A
PhiLonotie sertota mM
SoLenostoms cordifoliun И TIROL MINI PESCE T 1
Crotoneuron decipiens eub И ei rn que IT LS E 1
Chyloseyphus palyantnus
war. eivularts dard scies C eer e M би 1
Leyenda: M-Micasquistos
Tab. 1 — Scapanietum undulatae Schwickerath 1944.
superficie de inventarios media de 14 dm?, con un nümero específico medio
de 2,9 (variaciones de 2 a 5). La asociación viene esencialmente configurada
en su fisionomía por los Bryochamaephyta reptantia, Scapania undulata y
Brachythecium rivulare, aspecto que se halla reforzado por las restantes especies
de la asociación como, Platyhypnidium riparioides y Fontinalis antipyretica.
La ünica especie de biótipo cespitoso, Schistidium rivulare, no alcanza índices
suficientemente grandes que puedan influir en forma decisiva en la fisionomia.
Lo más destacable en la fisionomía de la asociación es su carácter fragmentario,
pues son pocas las veces que pueden inventariarse la totalidad de las especies
diferenciales.
Sindinämica. — Las especies pioneras Scapania undulata (V) y Brachythecium
rivulare (III) son las que tienden a ocupar el mayor área posible en las rocas
sumergidas. El desarrollo de las restantes especies de la comunidad es fragmen-
tario y variable, pues mientras Fontinalis antipyretica (П) suele establecerse
en lugares algo protegidos del impacto directo de la corriente, Schistidium
rivulare (П) y Platyhypnidium riparioides (1) prefieren sin embargo estaciones
donde es superior la torrencialidad del agua. Es precisamente en dichas estacio-
nes donde se produce el tránsito hacia las comunidades de Hygrohypnetea,
Source : ММНМ. Paris
COMUNIDADES REOFILAS DE SIERRA NEVADA 427
la mayor parte de las veces hacia Schistidio-Hygrohypnetum dilatati Geissler
1976; sélo en aquellas estaciones donde la escorrentia no es excesivamente
grande, la comunidad evoluciona hacia Solenostomo-Hygrohypnetum Geissler
1976.
En rocas próximas a los bordes de los ríos, la asociación es rápidamente
invadida por las comunidades de la alianza Montion (Br. Bl. 1925) Maas 1959,
produciéndose la sucesión desde comunidades saxicolas y pioneras hacia comuni-
dades helofiticas, terri-saxicolas y terricolas secundarias.
Sintaxonomía. — La asociación descrita por SCHWICKERATH (1944), fué
posteriormente confirmada por KRUSENSTJERNA (1945) quien la incluyó en
la alianza Racomitrion acicularis. Sin embargo, el carácter anfibio de que
goza esta alianza determinó que PHILIPPI (1956) situara a la asociación en su
nueva alianza Scapanion undulatae, más acorde con la tendencia de la misma
a permanecer sumergida.
Sincorología. — La asociación se halla ampliamente distribuida en Europa,
con preferencias en las zonas montafiosas de Centroeuropa y regiones escandi-
navas. También ha sido descrita por HUBSCHMANN (1971) en la isla de Madera.
En la Península Ibérica disponemos de algunas referencias en estudios realizados
por V. et P. ALLORGE (1947) y CASAS (1958); a partir de dichos trabajos
se deduce una distribución adscrita a las regiones de abundante precipitación
y sustrato ácido, como Galicia, noroeste de Portugal y afloramientos graníticos
de las montañas astures, vascas y catalanas, sierras de Guadarrama y Albarracín.
En el sur de la Península sólo tenemos las referencias de У. et Р, ALLORGE
(1945) para las sierras de Algeciras, lo que unido a nuestra cita correspondería
a su localisación más meridional en Europa.
1.2. ORDEN LEPTODYCTIETALIA RIPARII Philippi 1956
El orden de comunidades basófilas Leptodyctietalia riparii, está representado
en Sierra Nevada por la alianza de comunidades anfibias submontanas Brachy-
thecion rivularis Hertel 1974, y probablemente la alianza de comunidades
sumergidas Fontinalion antipyreticae Koch 1936.
La alianza Brachythecion rivularis comprende asociaciones anfibias, capaces
de resistir un período de desecación más o menos prolongado. En las zonas
basales de la sierra, los caudales acusan extremadas fluctuaciones durante el
verano, llegando en ocasiones a una sequía casi total, con lo que se anula el
posible desarrollo de la alianza; en las partes altas sin embargo el aforo es conti-
nuo, por lo que apenas si varía el caudal. En consecuencia, las estaciones que
potencialmente debiera ocupar la alianza son normalmente colonizadas por
Scapanion undulatae о Montion, exceptuando algunos nichos más protegidos
de la corriente en que se establece Brachythecion rivularis, representada por la
asociación Dichodontietum pellucidi v. Hübschmann 1967.
La presencia de la alianza Fontinalion antipyreticae se manifiesta probable-
mente mediante la asociación Fontinaleto-Pachyfissidentetum grandifrondis
Source : MNHN, Paris
428 J.A.GIL & J. VARO
Koch 1936, comunidad que permanece sumergida gran parte del año y de
netas preferencias basófilas. Sin embargo, de acuerdo con lo expresado por
BOROS (1968) y HERTEL (1974), hay que tener presente que la especie
que da nombre a la alianza, Fontinalis antipyretica, es en realidad indiferente
respecto del pH (entre 5,4 y 7,7). Todo ello hace de la alianza en cuestión
una unidad sintaxonómica incierta y heterogénea; no obstante, seguimos adop-
tando de momento la tradicional inclusión de Fontinaleto-Pachyfissidentetum
grandifrondis dentro de la alianza Fontinalis antipyreticae.
Dicha alianza está ausente de la zona esquistosa de la sierra, relegándose a
las formaciones calizas basales, donde alcanza un precario desarrollo en conse-
cuencia а la escasa calidad biológica de las aguas así como por las fuertes
variaciones de los caudales en el período estival.
B) ASOCIACION DICHODONTIETUM PELLUCIDI v. Hübschmann 1967
Estructura y composición florística : tab. 2, Origen de los inventarios : tab. 2.
№ 1, 2, 10 : 2880 m. N° 3 : 2870 m - Laguna de las Yeguas : 2.8.73. N° 4:
2770 m. № 5, 8, 9: 2700 m - Rio Guarnón : 10.7.76. № 6, 7 : 2450, 2470 m -
Rio Maitena : 13.7.77.
Sinecología . — Asociación reófila que muestra notables diferencias ecológi-
cas respecto a otras comunidades de carácter semejante. Estas diferencias son,
unà patente esciofília que provoca su localización en oquedades protegidas y
wmbrías, un marcado carácter anfibio que le hace mostrarse sumergida en los
primeros momentos del estío y expuesta a desecación en los meses finales;
por ültimo, es asimismo notable su indiferencia respecto del pH del agua. Dicho-
dontietum pellucidi responde pues a un comportamiento higro-hidrófilo, esció-
filo e indiferente respecto al sustrato, a diferencia de las otras asociaciones
reófilas de carácter exclusivamente hidrófilo, fotófilo y acidófilo estricto.
Sinfisionomía, — A la diferencia de la comunidad anterior, Dichodontietum
pellucidi se muestra como asociación un tanto más cerrada (cobertura media
= 519/0), para una superficie de inventarios media de 12,3 dm? , con un número
específico medio de 4 (variaciones de 2 a 6). Asimismo, las diferencias fisiond-
micas se ven potenciadas por el predominio que establece Dichodontium pelluci-
dum (V) (briocaméfito cespitoso-pulvinular) sobre los briocaméfitos reptantes,
como Cratoneuron decipiens (П), Brachythecium rivulare (II), Cratoneuron
commutatum var. irrigatum (П), Solenostoma pumilum (П) у Solenostoma
cordifolium (11).
Sindinámica. — La asociacion descrita por HUBSCHMANN (1967), posee
entre otras las siguientes especies : Dichodontium pellucidum, Platyhypnidium
riparioides, Schistidium rivulare, Scapania undulata у Brachythecium rivulare;
de todas ellas, la única que aparece regularmente en la asociación nevadense
es Dichodontium pellucidum, pues Brachythecium rivulare está presente tan
solo en cuatro de los inventarios, Es evidente por otra parte la presencia de un
elevado nümero de especies, compañeras de las anteriores, que en su mayoría
Source : ММНМ. Paris
COMUNIDADES REOFILAS DE SIERRA NEVADA
429
tienen carácter terricola o terrisaxícola, así como un comportamiento meso-
higrófilo. La mayor parte de ellas pertenecen a diferentes asociaciones de
Montio-Cardaminetea Br. Bl. et Tx. 1943, que son las que generalmente
la clase
ocupan
las estaciones potencialmente colonizables por la asociación Dichodontietum
pellucidi. En definitiva, podemos concluir que en Sierra Nevada sólo disponemos
de una representación fragmentaria de la asociación estudiada,
Nümero de inventario qq qa "eg gt our Legg d
Superficie(dm ) 12 10 10 10 12 20 10 15 12 12
Cobertura(%) 70 60 40 40 80 80 60 30 30 20
Exposición NE ON N N N NE ON N N м
Inelinación(?) 40 70 80 70 80 40 5 90 90 60
Sustrato MOM M M M M M M OM
Nümero de especies quer fl Een EE С ЕЕ
Caracteristica de asociación:
Dichodontium pellucidum
fo. fogimontonum [UE a т Па
Características de unidades superiores (Brochythecion rivularis,
Leptodyctietalia riporii,Plotyhyenidio-Fontinaleteo)
Brachytnectum rlvüLene “а „1. E e e
Scoponio undulata e e er SET E
Compañeras :
Solenostomo cordifoltum + + + + + + + 2 1 .
Solenostoma pumiLum UR з, ж, CR eg uie EE
Bryum schleicheri O cT АБ eS
Crotoneuron decipiens pues LE рч,
Crotoneuron commutotum
var. Lrrigatum qM woe oe to oem
Philonotis seriota Gomes cx A aen ie
AmphidLum mougeotit i cim oem pole СРОКА NES
Leyenda: M-Micasquistos
Tab. 2. — Dichodontietum pellucidi v. Hübschmann 1967.
Sintaxonomía. — Dado su carácter montano la asociación fué incl
її
II
III
11
п
11
luida en
la alianza Scapanion undulatae por НОВЗСНМАММ (1967). Dicho autor (1973),
reestructura la sintaxonomia de las comunidades briofíticas reófilas y e
n conso-
Source : MNHN. Paris
430 J.A.GIL & J. VARO
папсіа con su carácter anfibio sitúa a la asociación en la alianza Racomitrion
acicularis. Sin embargo, este sintaxon sólo reúne asociaciones estrictamente
acidófilas, lo que no es consecuente con la indiferencia que manifiesta Dicho-
dontietum pellucidi respecto al sustrato: este aspecto ya ha sido anteriormente
reseñado por HÜBSCHMANN (1967, 1973).
La presencia simultánea de Dichodontium pellucidum y Brachythecium
rivulare que evidencia nuestra asociación, constituye para nosotros un aspecto
bastante significativo, pues también se puede constatar en las comunidades
reófilas estudiadas por HERZOG (1944), HÜBSCHMANN (1973), NORR
(1969) y HERTEL (1974). Todo ello nos parece lógico, si consideramos que
Brachythecium rivulare también se comporta como anfibia e indiferente al
sustrato. La cocxistencia de ambas, parece relacionada con una tendencia por
parte de las especies de Montio-Cardaminetea para invadir las comunidades ya
establecidas de Dichodontietum pellucidi. En este sentido, en opinión de
HÜBSCHMANN (1973), Brachythecium rivulare sería más bien un vínculo
entre las comunidades briofiticas reófilas y las helofíticas de las alianzas Montion
(Br. Bl. 1925) Maas 1959, Cardaminion Maas 1959 y Cratoneurion commutati
W. Koch 1928. En consecuencia, la alianza Brachythecion rivularis incluiría
asociaciones que, como Dichodontietum pellucidi, representan la transición
entre las comunidades estrictamente reófilas, y las de la clase Montio-Cardami-
netea que están en intimo contacto con éllas.
Sincorología. — Según HÜBSCHMANN (1967), la asociación se extiende
por toda Europa en ríos de alta y media montaña, Cáucaso, Japón y América
del Norte. No conocemos con precisión su areal peninsular, pero la distribución
y características ecológicas de la especie directriz, permiten suponer su presencia
en algunas de las regiones montañosas del norte de la Península Ibérica.
€) ASOCIACION FONTINALETO-PACHYFISSIDENTETUM GRANDIFRONDIS
W. Koch 1936
Estructura y composición florística : tab. 3. Orígen de los inventarios : tab. 3.
N° 1 : 1000 m - Rio Genil : 13.2.76. № 2 al 8 : entre 1200 y 1370 m - Rio
Aguas Blancas : 20.4.77.
Sinecología. — Comunidad reófila que permanece siempre sumergida, coloni-
zadora de arroyos y ríos de aguas apenas contaminadas y ricas en cal (pH = 7-
8,5). Por tanto, actúan como factores limitantes la eutrofía del agua y la oscila-
ción del caudal.
Aunque de forma precaria, la asociación aparece en la orla calizo-dolomítica
basal de la sierra; el limitado desarrollo que alcanza se debe en parte a la fuerte
sequía estival, así como a la notable filtración y absorción de aguas que caracte-
riza a la topografía calcárea; existe además una extensa red de acequias y una
creciente contaminación.
Sinfisionomía. — La comunidad participa de un doble aspecto fisionómico,
Source - MNHN. Paris
COMUNIDADES REOFILAS DE SIERRA NEVADA
431
conferido por la presencia simultánea de Fissidens grandifrons (Bryochamae-
phyta caespitosa) (V) y Platyhypnidium riparioides (Bryochamaephyta reptantia)
(У). En cualquier caso, la asociación es abierta (cobertura media = 520/0), para
una superficie de inventarios media de 16,1 dm?, con un número específico
medio de 3,5 (variaciones de 2 a 5).
Nümero de inventario
superficie (dm)
Cobertura(%)
Exposición
Inclinacién
Sustrato
Namero de especies
Caracteristica de asociacié:
Fissidens grandifrons
Caracteristicas de unidades
Leptodyctietolio riparti,PlatyhypnLdio-Fontinaletea):
PLotghupnidium Fiparioides
Compañeras:
Cratoneuron commutatum
var. commutatum
Philonotis calcarea
Aptum nodiflorum
Nostoc Sp.
Cardamine sylvatica
Plagtomnium rostratum
Bryum pseudotriquetrum
Calizas
Leyend
10
80
90
1
15
20
NO
D
Ке
16
20
10
10
50
6
18
Tab. 3. — Fontinaleto -Pachyfissidentetum grandifrondis W.
7
80
чоо =
8
25
70
аа
Koch 1936.
3,5
superiores(Fontinalion antipyreticae,
тїї
п
її
II
Sindinámica. — La asociación descrita por KOCH (1936) tiene como carac-
terísticas a Fissidens grandifrons y Fontinalis antipyretica. La comunidad
estudiada en Sierra Nevada sólo lleva la primera de estas especies, siendo el cauce
del río Aguas Blancas la única localidad conocida para dicha especie. La altitud
relativamente baja del nacimiento de este tío, así como el efecto del deshielo,
permite а sus aguas mantener un alto régimen de caudal además de una esti-
Source : MNHN. Paris
432 J.A.GIL & J, VARO
mable pureza y en consecuencia, condiciones idéneas para fijarse una especie
tan exigente como Fissidens grandifrons; es por tanto la especie pionera que,
junto a Platyhypnidium riparioides, inicia la colonización de las rocas sumergidas
en el cauce del río Aguas Blancas.
Las rocas próximas a los márgenes del río, muestran la sucesión de Fontina-
leto-Pachyfissidentetum grandifrondis hacia la alianza helofitico-basöfila Crato-
neurion commutati, con la que se halla en contacto e incluso solapada la mayo-
ría de las veces.
Sintaxonomía. — La asociación ha sido tradicionalmente incluida en la
alianza Fontinalion antipyreticae Koch 1936, de la clase Fontinaletea antipy-
reticae v. Hübschmann 1957. Sin embargo, en la reestructuración realizada
por HERTEL (1974), la alianza Fontinalion no se contempla de forma defini-
tiva, pues la especie directriz de la alianza, Fontinalis antipyretica, es realmente
indiferente al sustrato; en consecuencia, no es del todo satisfactoria la situación
de Fontinalion dentro del orden basófilo Leptodyctietalia riparii. Concluimos
de todo ello que es necesaria una posible reconsideración de la estructuración
de la clase Platyhypnidio-Fontinaletea.
Sincorología. — Se halla generalizada su presencia en los cauces de aguas
poco contaminadas de las llanuras calizas europeas. Los datos aportados por
P. ALLORGE (1947) nos hacen suponer su existencia en el occidente y centro
de la region cántabra, en las montañas calizas del País Vasco y en algunos
macizos de Levante y Andalucia, En la provincia Bética, la asociación ha sido
localizada en el lecho calizo del río Plines, próximo a Loja (Granada), plena-
mente desarrollada, aunque con Fontinalis duriaei sustituyendo a Fontinalis
antipyretica, creemos que como asociación vicariante meridional de Fontinaleto-
Pachyfissidentetum grandifrondis.
2. CLASE HYGROHYPNETEA v. Hübschmann 1957
Reúne aquellas comunidades réofilas de carácter alpino. Los estudios de
ALLORGE (1925), KRAJINA (1933), HÜBSCHMANN (1957), así como la
revisión realizada por GEISSLER (1976), nos han permitido el reconocimiento
y estudio de dicha clase en Sierra Nevada, estando representada por las asocia-
ciones Schistidio-Hygrohypnetum dilatati Geissler 1976 y Solenostomo-Hygro-
hypnetum Geissler 1976. Ambas asociaciones presentan el comportamiento
ecológico que caracteriza a la alianza Hygrohypnion Krajina 1933, es decir, una
gran adaptación frente a la torrencialidad del agua y acusada psicrofilia (los
valores medios de la temperatura del agua no llegan a superar los 8 С en verano);
las restantes características ecológicas coinciden en su mayor parte con las ya
descritas para la clase Platyhypnidio-Fontinaletea, puesto que ambas clases se
hallan directamente en contacto en Sierra Nevada.
Source : MNHN, Paris
COMUNIDADES REOFILAS DE SIERRA NEVADA 433
D) ASOCIACION SCHISTIDIO-HYGROHYPNETUM DILATATI Geissler 1976
Sinónimo : Hygrohypnetum dilatati tatrense Krajina 1933 p.p.
Estructura y composición floristica : tab. 4. Origen de los inventarios : tab. 4.
№ 1, 2, 3, 16, 17, 18 : 2700, 2760, 2600, 2840, 2680, 2650 m - Rio Dilar :
2.9.74. NO 4, 5 : 2370, 2380 m - Campos de Otero : 15.9.74. N° 6 : 3050 m -
Laguna de Aguas Verdes : 6.8.75. N° 7, 8, 9, 10 : 2950, 2890, 2870, 2840 т -
Rio Seco : 6.8.75. № 11, 19 : 2720, 2700 m - Вю Veleta : 26.8.75. № 12 :
2880 m - Laguna de las Yeguas : 2.8.73. N? 13, 14 : 2880, 2870 m - Rio Guar-
nón : 10.7.76. № 15 : 2720 m - Rio Trevélez : 17.8.76. № 20, 21 : 2950,
2900 m - Rio Lanjarón : 27.7.77. № 22, 23 : 2450, 2890 m - Rio Valdecasillas :
11.8.77.
Sinecología. — La característica fundamental de la asociación es su marcada
reofilia, situándose en forma preferente en los rápidos, allí donde la corriente
golpea más intensamente, Su carácter psicréfilo también es acentuado, siendo
normal su presencia en aguas cuya temperatura no supera un valor medio de
8°C. La asociación es asimismo acidófila, estando siempre sumergida o semi-
sumergida en los tramos silíceos comprendidos entre 2400 y 3000 m.
Sinfisionomía. — Comunidad abierta (cobertura media = 65,60/0), para
una superficie de inventarios media de 21 dm”, con un número específico
medio de 2,9 (variaciones de 2 a 5). La asociación generalmente presenta un
aspecto mixto, pues tiene lugar el contraste entre Hygrohypnum dilatatum
(Bryochamaephyta reptantia) y Schistidium rivulare (Bryochamaephyta caespi-
tosa), pues si bien las restantes especies de la comunidad (pertenecientes à la
clase Platyhypnidio-Fontinaletea) poseen un bidtipo rastrero, éstas no alcanzan
indices muy altos. Este equilibrio fisionómico se altera en la subasociación
Platyhypnidietosum; es decir, la entrada de Platyhypnidium riparioides (Bryo-
chamaephyta reptantia) se produce en tal grado, que llega a ocultar la super-
ficie previamente colonizada por las especies pioneras antes mencionadas.
Es asimismo significativa la forma fragmentaria en que suele presentarse la
asociación, en consecuencia con la considerable competencia que se establece
entre sus especies integrantes.
Sindinámica. — Hygrohypnum dilatatum (V) comunica un neto carácter
reófilo a la comunidad, estando generalmente asociada a élla Schistidium rivulare
(Ш) asimismo adaptada а la torrencialidad del río. Ambas especies зе comportan
pues, como pioneras en la colonización de rocas con una franca exposición a
la corriente. A las anteriores suelen agregarse después, aunque en forma espora-
dica, las especies que caracterizan a la clase Platyhypnidio-Fontinaletea, tales
como Brachythecium rivulare (П), Scapania undulata (П), Cladophora glomerata
(1), asi como Fontinalis antipyretica (1) en estaciones más protegidas: en todos
los casos va a ser la fuerza del agua, ya que no la temperatura, quien delimite
las estaciones a colonizar por las especies de una u otra clase.
Source : MNHN. Paris
Námero de inventario Голи вст ее
Superficie(dm ) 12 10 20 15 40 50 12 20 20
Cobertura(X) 30 70 70 60 90 70 40 50 80
Exposición NE NO NO N NS S SS
Inclinación(*) 45 40 10 60 70 90 50 45 70
Sustrato MMMM MM MOM OM
Námero de especies Ps s $4 s 32 4
Características de asociación y unidades superiores(Schistidio-Hygrohypnetum diLatati,Hygrohyenton
HygrohypnetaLta,Hygrohypnetea) :
Hygrohypnum diLototum аз: E а
Schistidium rivulare E e BAT упа
Diferencial de subasociacién(PLatyhypnidietosum) :
Platyhypnidium riparloides + + + + e + eoe
Especies de PLotyhypnidio-FontinoLeteo :
Brochythecium rivulare T nee SCH e
Scoponto undulata 5 5 пара на
Fontinalis antipyretica о а XM
Cladophora glomerata SEE CEE mI:
Compañeras:
Crotoneuron decipLens TIN EE э ЖЕШ.
Hygrohypnum ochroceum WE OS
Hygrohypnum Luridum JR 413.5
Porello cordaeana ge аи
Leyenda: M-Micasquistos
10
20
90
s
30
и
3
1
12
90
50
12
30
80
45
13
15
40
30
14
20
90
70
15
12
60
SE
90
=
16
10
90
NE
70
=
17
15
40
45
Tab. 4 — Schistidio-Hygrohypnetum dilatati Geissler 1976.
18
15
ко
60
19
12
90
so
20
21 22 25
30 20 40
40 80 80
S0 SE NO
50 70 60
мом м
Мм
diletati,
ара
^c SET
3 ET
тїт
її
п
и
РЕР
Омул ‘Le TD v'f
Source : ММНМ. Paris
COMUNIDADES REOFILAS DE SIERRA NEVADA 435
La transición hacia las comunidades de Montio-Cardaminetea se produce
en las rocas próximas a las márgenes del río, siendo generalmente Brachythecium
rivulare quien sirve de nexo entre las comunidades теб аз y las helófitas.
Finalmente, a medida que el río se va haciendo más caudaloso y como conse-
cuencia de un aumento progressivo en la temperatura del agua, en los pisos
inferiores de la sierra se produce la definitiva desaparición de Hygrohypnum
dilatatum, quedando tan solo Platyhypnidium riparioides (1) en poblaciones
puras que a veces participan también de la presencia de Brachythecium rivulare.
Por todo ello proponemos la subasociación Platyhypnidietosum riparioidis
(Holotypus : inventario n° 22, tab. 4) que caracterizaría los tramos a menor
altitud, es decir, en el límite del areal ocupado en nuestro territorio por la
asociación Schistidio-Hygrohypnetum dilatati.
Sintaxonomia. — La asociación es incluida por GEISSLER (1976) en la
alianza Hygrohypnion dilatati Krajina 1933, orden Hygrohypnetalia Krajina
1933, clase Hygrohypnetea v. Hübschmann 1957.
Sincorología. — La asociación ha sido localizada al este de los Alpes (AL-
LORGE, 1925), Trata (KRAJINA, 1933), Alpes Réticos (BRAUN BLANQUET,
1948) y recientemente al oeste de los Alpes (GEISSLER, 1976).
En relación a su distribución peninsular, ALLORGE (1947) hace referencia
а una comunidad de Hygrohypnum dilatatum e Hygrohypnum molle, propia
de los pisos montano y alpino de las montañas silicicolas de la Peninsula, si-
tuándola concretamente por encima de 1600 m en los Pirineos y Cordillera
Central, y por encima de 2500 m en Sierra Nevada. CASAS (1975) ha estable-
cido la corología peninsular de Hygrohypnum dilatatum, permitiéndonos supo-
ner la existencia de la comunidad en distintos macizos montañosos de la pro-
vincias de León, Soria, Lérida, Gerona y la estudiada por nosotros en Sierra
Nevada.
E) ASOCIACION SOLENOSTOMO-HYGROHYPNETUM Geissler 1976
Estructura y composición florística : tab. 5. Orígen de los inventarios : tab. 5.
№ 1 : 2880 m- Laguna de las Yeguas : 2.9.74. N? 2, 3,4 : 2710, 2700, 2500 m
- Rio Dílar : 4.9.74. № 5, 6 : 3000, 2730 m - Rio Veleta : 6.8.75. N9 7 :
2500 m - Rio Maitena : 13.7.77.
Sinecologia. — Comunidad psicrófila y acidófila que sustituye a Schistidio-
Hygrohypnetum dilatati en aquellos puntos donde el curso se hace menos
caudaloso y torrencial. GEISSLER (1976) diferencia ambas comunidades
a partir de la velocidad media de los arroyos, siendo ésta muy superior en Schisti-
dio-Hygrohypnetum dilatati; asimismo establece diferencias altitudinales, de
forma que Solenostomo-Hygrohypnetum corresponde al piso subalpino mientras
que Schistidio-Hygrohypnetum dilatati lo es del alpino.
En Sierra Nevada también se puede observar una dependencia entre la fuerza
del agua y la aparición de una и otra asociación: sin embargo no sucede lo mismo
Source : MNHN, Paris
436
Némero de inventario
Superficie(dm-)
Cobertura (%)
Exposición
Inclinación(*)
Sustrato
Número de especies
J-A.GIL & J. VARO
1
10
80
70
2
20
70
NO
90
3.
15
90
NO
40
4
5
30
60
SE
45
6
10
50
10
7
40
70
NE
20
3 3,8
Caracteristicas de asociacién y de unidades superiores
(Solenostomo-Hygrohypnet um,Hygrohypnton dilatatl,Hygro
hypnetalta,Hygrohypnetea):
Solenostoma cordifolium
Hygrohypnum dilatatum
Características de Plotyhypnidio-Fontinaletea:
Brachythecium rivulare
Fontinalis antipyretica
Scapanta unduLata
Cladophora glomerata
Compañeras:
Crotoneuron decipiens
Hygrohypnum ochraceum
Hygrohypnum Luridum
Porella cordaeana
Leyenda: M-Micasquistos
Tab. 5. — Solenostomo-Hygrohypnetum Geissler 1976.
»
Yir
тї
en cuanto a su distribución altitudinal, ya que las aguas del piso oromediterráneo
nevadense no retinen condiciones idéneas para el asentamiento de Solenostomo-
Hygrohypnetum (con temperaturas muy superiores a la media requerida para
la comunidad), de ahí que ambas comunidades coexistan dentro de una franja
comprendida entre 2400 y 3000 m.
Sinfisionomía. — Se trata de la comunidad menos abierta de todas las estu-
diadas (cobertura media = 710/0), pues suele formar tapices continuos sobre
lechos pedregosos (en menor proporción sobre lechos rocosos); no obstante, al
igual que en las asociaciones anteriores la comunidad se caracteriza por ser
Source : MNHN. Paris
COMUNIDADES REOFILAS DE SIERRA NEVADA 437
fragmentaria, así como por el neto dominio que establecen las especies de bió-
tipo rastrero (Bryochamaephyta reptantia), en total ausencia de las de biótipo
cespitoso. La superficie de inventarios media es de 20 dm” , con número especi-
fico medio de 3,8 (variaciones de 2 a 5).
Sindinámica, — El óptimo de la asociación corresponde а los nacimientos
de los ríos, es decir, donde el agua conserva la temperatura de deshielo y un
régimen no excesivamente torrencial ni caudaloso. En semejantes condiciones,
Hygrohypnum dilatatum (V) y Solenostoma cordifolium (V) actúan como
pioneras en la colonización de lechos pedregosos que al adquirir un régimen
torrencial más acentuado, evolucionan hacia la asociación Schistidio-Hygro-
hypnetum dilatati. La asociación Solenostomo-Hygrohypnetum establece
asimismo unas condiciones que favorecen la entrada de especies de Platyhyp-
nidio-Fontinaletea, tales como Brachythecium rivulare (ШШ), Fontinalis anti-
pyretica (11), Scapania undulata (1) y Cladophora glomerata (1).
También es frecuente en contacto entre la asociación y las comunidades de
la clase Montio-Cardaminetea, puesto que los factores limitantes de éstas no
actüan tan intensamente como en asociaciones anteriores.
Sintaxonomía. — Por su distribución y características ecológicas es afín
a Schistidio-Hygrohypnetum dilatati, por lo que GEISSLER (1976) la incluye
asimismo en la alianza Hygrohypnion dilatati.
Sincorología. — Solenostoma cordifolium es especie nórdica, cuyo areal
se extiende a lo largo de Escandinavia, norte de Gran Bretaña e Islandia; también
alcanza las partes más elevadas de diferentes cadenas montañosas de Centro-
europa, norte de Italia, Caúcaso, Trata y Cárpatos. La presencia conjunta de
dicha especie con Hygrohypnum dilatatum, tan sólo ha sido descrita por GEISS-
LER (1976) para el oeste de los Alpes suizos, aunque posiblemente también
se encuentre la comunidad en algunas de las estaciones antes mencionadas.
De forma semejante a Hygrohypnum dilatatum, Solenostoma cordifolium
posee una distribución peninsular disyunta y limitada a la Sierra de Guadarrama,
Pirineos y Sierra Nevada; según MÜLLER (1954-1957), esta última localidad
constituye una estación relíctica y es en ella donde describimos la asociación
por primera vez para la Península Ibérica,
ESQUEMA SINTAXONOMICO DE LAS COMUNIDADES ESTUDIADAS
Clase Platyhypnidio-Fontinaletea Philippi 1956
Ord. Brachythecietalia plumosi Philippi 1956
Al. Scapanion undulatae Philippi 1956
As. Scapanietum undulatae Schwickerath 1944
Ord. Leptodyctietalia riparii Philippi 1956
Al. Brachythecion rivularis Hertel 1974
As. Dichodontietum pellucidi v. Hübschmann 1967
Source : MNHN, Paris
438 J.A.GIL & J. VARO
Al. Fontinalion antipyreticae W. Koch 1936
As. Fontinaleto-Pachyfissidentetum grandifrondis W. Koch 1936
Clase Hygrohypnetea v. Hübschmann 1957
Ord. Hygrohypnetalia Krajina 1933
Al. Hygrohypnion dilatati Krajina 1933
As. Schistidio-Hygrohypnetum dilatati Geissler 1976
As. Solenostomo-Hygrohypnetum Geissler 1976
La lista que a continuación detallamos, corresponde a los musgos y hepáticas
que integran las comunidades estudiadas :
Hepáticas
Chiloscyphus polyanthus (L.) Corda var. rivularis (Schrad.) Nees
Porella cordaeana (Hüb.) Evans
Scapania undulata (L.) Dum.
Solenostoma cordifolium (Hook.) Steph.
Solenostoma pumilum (With.) K. Mueller
Musgos
Amphidium mougeotii (B.S.G.) Schimp.
Brachythecium rivulare B.S.G.
Bryum pseudotriquetrum (Hedw.) Schwaegr.
Bryum schleicheri Schwaegr.
Cratoneuron commutatum (Hedw.) Roth, var, commutatum
Cratoneuron commutatum (Hedw.) Roth var. irrigatum (Zett.) Monk.
Cratoneuron decipiens (De Not.) Loesk.
Dichodontium pellucidum (Hedw.) Schimp. fo. fagimontanum C. Jens.
Fissidens grandifrons Brid.
Fontinalis antipyretica L.
Fontinalis hypnoides Hartm, var. duriaei (Schimp.) Husnot
Hygrohypnum dilatatum (Wils.) Loesk.
Hygrohypnum luridum (Hedw.) Jenn.
Hygrohypnum ochraceum (Wils.) Loesk.
Philonotis calcarea (B.S.G.) Schimp.
Philonotis seriata Mitt.
Plagiomnium rostratum (Schrad.) Kop.
Platyhypnidium riparioides (Hedw.) Dix.
Schistidium rivulare (Brid.) Podp.
AGRADECIMIENTOS
Nuestro particular agradecimiento a la Dra. С, Casas Sicart, quien amablemente revisó
la determinación de algunas especies, así como por su asesoramiento bibliográfico en nues-
tro estudio briosociológico.
Source : MNHN. Paris
COMUNIDADES REOFILAS DE SIERRA NEVADA 439
BIBLIOGRAFIA
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Briançonnais. Verdff. Geobot. Inst. Rübel Zürich 3 : 108-126.
ALLORGE P., 1928 — Remarques sur la flore muscinale des hauts sommets de la Péninsule
Ibérique. Mem. Soc. Biogéogr., le série, 2 : 252-259.
ALLORGE P., 1947 — Essai de Bryogéographie de la Péninsule Ibérique. Paris, P. Lecheva-
lier, 105 p.
ALLORGE V. et P. 1945 — La végétation et les groupements muscinaux des montagnes
d'Algésiras. Mém. Mus. Natl. Hist, Nat. попу, sér., 21 (3) : 85-116,
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ALLORGE V. & CASAS C., 1963 — Au sujet des Bryophytes récoltées au cours de l'excur-
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BOROS A., 1968 — Bryogeographie und Bryoflora Ungarns, Budapest, Akadém. Kiadó,
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Península Ibérica. Trab. Mus. Мас. С. Nat., Ser. Bot. 8 : 1-179.
CASAS DE PUIG C., 1953 — Una excursión briolôgica al Valle de Nuria. Collect. Bot.
(Barcelona) 3 (2) : 199-206.
CASAS DE PUIG C., 1956 — Aportaciones a la flora briolögica de los Pirineos Centrales.
Collect. Bot. (Barcelona) 5 : 419-424.
CASAS DE PUIG C., 1958 — Aportaciones a la flora briológica de Cátaluña. Anales Inst.
Bot. Cavánilles 16 : 121-226.
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y Hepáticas de las provincias de Soria, Logroño, Burgos y Segovia. Anales Inst. Bot.
Cavdnilles 32 (2) : 731-762.
DEGUCHI H., 1979 — Les véritables caractères de Schistidium alpicola (Sw. ex Hedw.)
Limpr. et son nouveau synonyme, Schistidium agassizii Sull. et Lesq. Rev. Bryol. Liché-
nol. 45 (4) : 425-435.
DUNK К. von der, 1972 — Moosgesellschaften im Bereich des Sandsteinkeupers in Mittel-
und Oberfranken, Ber. Naturwiss. Ges. Bayreuth 13 : 7-100.
GEISSLER P., 1976 — Zur Vegetation alpiner Fliessgewasser. Mater. Flore Cryptog. Suisse
14 (2) : 1-52, 25 tabl.
GEHEEB A., 1862 — Beitrag zur Moosflora von Spanien. Flora 5 : 516-521.
GIL J.A. & VARO J., 1973 — Contribución al estudio briológico de Sierra Nevada. II.
El Barranco de San Juan. Trab. Dep. Bot. Univ. Granada 2 (2) : 63-79.
HERTEL E., 1974 — Epilithische Moose und Moosgesellschaften im nord-óstlichen Bayern.
Beih. Ber. Naturwiss. Сез. Bayreuth 1 : 1-489, 52 tabl.
HERZOG T., 1944 — Die Mooswelt des Kódnitztales in den Hohen Tauern. Wiener Bot.
Z. 93 (1) : 1-65.
Source : ММНМ. Paris
440 J.A.GIL & J. VARO
HONEL F., 1895 — Beitrag zur Kenntnis der Laubmoosflora des Hochgebirgstheiles der
Sierra Nevada in Spanien, Sitzungsber. Kaiserl Akad. Wiss, Math. - Naturwiss. CL,
Abt. 1,104 (1) : 297-336.
HÜBSCHMANN A. von, 1953.— Einige hygro-und hydrophyle Moosgesellschaften Nord.
deutschlands. Mitt. Florist. Soziol. Arbeitsgem., N.F. 4 : 15-25.
HÜBSCHMANN A. von, 1957 — Zur Systematik der Wassermoosgesellschaften, Mitt.
Florist. Soziol. Arbeitsgem., N.F. 6/7 : 147-151,
HÜBSCHMANN А. von, 1967 — Uber die Moosgesellschaften und das Vorkommen der
Moose in den übrigen Pflanzengesellschaften des Moseltales. Schriftenreihe Vegeta-
tionsk, 2 1 63-121.
HÜBSCHMANN A. von, 1973 — Moosgesellschaften des Nordwestdeutschen Tieflandes
zwischen Ems und Weser. Herzogia 3 : 111-130.
KOCH W., 1936 — Über einige Wassermoosgesellschaften der Linth. Ber. Schweiz. Bot.
Ges. 46 : 355-364,
KRAJINA V., 1933 — Die Pflanzengesellschaften des Mlynica-Tales in den Vysoké-Tatry.
Вей. zum Bot. Centralbl. 50 : 774-957.
KRUSENSTJERNA E. von, 1945 — Bladmossvegetation och Bladmossflora i Uppsala-
Trakten. Acta Phytogeogr. Suec. 19 : 1-250.
NORR M., 1969 — Die Moosvegetation des Naturschutzgebietes Bodetal. Hercynia 6 (4) +
345-435.
PASCUAL A., 1975 — Estudio hidrogeológico de las cuencas medias de los rios Monachil
y Dilar (Granada). Tesis de Licenciatura, Universidad de Granada. Mscr., 134 p.
PHILIPPI G., 1956 — Einige Moosgesellschaften des Südschwarzwaldes und der angren-
zenden Rheinebene, Beitr. Naturk. Forsch. Südwestdeutschl. 15 (2) + 91-124.
PHILIPPI G., 1965 — Die Moosgesellschaften der Wutachschlucht. Mitt. Bad. Landesvereins
Naturk. Naturschutz, N.F. 8 (4) : 625-668.
POELT J. 1954 — Moosgesellschaften im Alpenvorland. 1 und IT, Osterr. Akad. Wiss.
Math. - Naturwiss. Kl., Sitzungsber., Abt. 1, Biol. 163 : 141-174 y 495-539.
RIVAS-MARTINEZ S., 1981 — Les étages bioclimatiques de la végétation de l'Espagne.
(Conferencia inaugural. III? Congreso OPTIMA, Madrid). Anales Jard. Bot. Madrid
1980 (1981), 37 (2) : 251-268.
SCHWICKERATH M., 1944 — Das Hohe Venn und seine Randgebiete. Vegefation, Boden
und Landschaft. Pflanzensociol. 6 : 1-278.
VELILLA N., 1976 — Estudio hidrogeológico de la cuenca del río Aguas Blancas (Granada).
Tesis de Licenciatura, Universidad de Granada. Mscr., 208 p.
WALTHER K., 1942 — Die Моозйога der Cratoneurum commutatum-Gesellschaft in den
Karawanken. Hedwigia 81 : 127-130.
Source : MNHN, Paris”
441
ERGÂNZUNGEN ZUR CAMPYLOPUS-FLORA
VON BRASILIEN
JP. FRAHM*
SUMMARY. — Campylopus luetzelburgii Herz. ex J.P. Frahm and C. tortilipilus J.
Frahm are described as new. C. minarum Par. has proved to be identical with C. surinamen-
sis C. Müll., C. praealtus (C. Müll.) Par. with C. subcuspidatus (Hamp.) Jaeg., C. rectisetus
(Hamp.) Jaeg. and C. discriminatus (Hamp.) Jaeg. with C. arctocarpus (Hornsch.) Mitt.,
D. rabenii Lor. as well as C. rubricaulis Lindb. ex C. Müll, with C. filifolius (Hornsch.) Mitt.
var. humilis (Mont.) J-P, Frahm, C. brachyphyllulus (C. Müll.) Broth. with C. savannarum
(C. Müll.) Mitt. and C. brasiliensis Broth, nom. nud. with Atractylocarpus brasiliensis
(C. Müll.) Williams,
Herbarstudien anlässlich Aufenthalten am Botanischen Museum in Berlin
(B) 1979 und am Muséum National d'Histoire Naturelle in Paris (PC) 1980
brachten unter anderem eine Reihe interessanter Belege aus Brasilien zum
Vorschein, die bei der Bearbeitung der Campylopus-Arten Brasiliens (FRAHM
1979) nicht berücksichtigt worden waren.
Dr. Jovet-Ast, Dr. Bischler und Dr. Schultze-Motel sei an dieser Stelle für die
gebotenen Arbeitsmöglichkeiten gedankt.
Ausserdem ist die Beschreibung einer neuen Art nach Material, welches D.M.
Vital (Sao Paulo) in Brasilien gesammelt hatte, mit aufgenommen.
1. CAMPYLOPUS LUETZELBURGII Herz. ex J.-P. Frahm
Unter den Campylopus-Proben im Herbar Herzog (JE) sowie im Herbar
Thériot (PC) befand sich auch ein mit «Campylopus luetzelburgii Herz. n. sp.
ad interim» beschrifteter Beleg aus der Serra dos Orgaos in Brasilien. Offenbar
hatte Herzog gezögert, diese Art neu zu beschreiben. Ein Vergleich mit den
übrigen aus Brasilien beschriebenen Arten (FRAHM 1979) zeigt, dass es sich
dabei um eine gewissen anderen sehr ähnliche aber durchaus distinkte eigene
Art handelt. Nachdem schliesslich im Herbar des Botanischen Museums Berlin
(B) noch ein weiterer Beleg dieser Art, von Dusén in der Serra do Itatiaia gesam-
melt und als «Campylopus concolor» bestimmt, aufgefunden wurde, ist es
`
* Universität Duisburg, Fachbereich 6, Botanik, Postfach 101629, D-4100 Duisburg.
Cryptogamie, Bryol. Lichénol., 1981, 2, 4 : 441-448.
Source : MNHN. Paris
442 J-P. FRAHM
angebracht, diese Art zu veröffentlichen. Dabei soll auf die von Herzog gewählte
Bezeichnung Campylopus luetzelburgii zurückgegriffen werden. Auf Grund des
besseren Herbarmaterials wird jedoch die Probe von Dusén als Holotypus benutzt.
Campylopus luetzelburgii Herz. ex J.-P. Frahm
Plantae steriles, 2-2,5 cm altae, flavescentes, dense foliosae. Folia concava,
siccitate appressa, 5 mm longa, ovata, breviter lanceolata, in subulam lanceola-
tam excurrentes, nervo excurrente, summo apice dentato, basi 3/4 folii occu-
pante, in sectione transversali cellulis ventralibus inanibus, stereidibus nullis.
Cellulae laminae basi hyalinae, teneres, rectangulares, 70-90 x 10-14 um, supe-
riores breviter rectangulares (1:2-3), incrassatae, 13-18(-25) x 9-10 um (Fig. 1).
Material ; Brasilien, Serra do Itatiaia, Agulhas Negras, leg. Dusén 2.7 1902
snr. (Holotypus В); Serra dos Orgaos, Morro Assu, auf Granit 2400 m, leg.
von Lützelburg Juli 1916 nr. 6793 (Paratypus JE, PC hb. Thériot).
Campylopus luetzelburgii kommt an beiden etwa 200 km Luftlinie ausein-
anderliegenden Lokalitäten oberhalb der Waldgrenze vor.
Die Art ist an der durch breite, etwas konkav eingebogene Blátter dicht
wurmfórmige Beblätterung und der hellen, an Frischmaterial wohl gelbgrünen
bis weisslichen Beblätterung habituell kenntlich. Anatomisch weist sie Beziehun-
gen zu Campylopus fragilis (Brid.) B.S.G. auf, mit der sie die unterhalb der
Blattmitte verbreiterte Rippe und die zur Basis hin kontrahierten Blätter gemein-
sam hat. C. luetzelburgii unterscheidet sich aber von C. fragilis durch sehr viel
kürzere und breitere Blatter, eine breite Lamina mit 8-12 Zellreihen (C. fragili
3-5) und verlängerte (1:2-3) rechteckige obere Laminazellen (bei C. fragilis
* quadratisch), sowie schliesslich dem Rippenquerschnitt ohne Stereiden.
Auffilligerweise existiert im Gebiet noch eine weitere, wie durch den Namen
schon angedeutet ebenfalls in die Verwandtschaft von C. fragilis gehörende Art,
Campylopus fragiliformis J.-P. Frahm. Diese Art hat deutlich von C. fragilis und
C. luetzelburgii unterschiedene verlängert ovale obere Laminazellen (Fig. 2).
Beide Arten sind aus dem Stidostbrasilianischen Bergland bekannt : C. fragi-
liformis aus der Serra do Itatiaia (Typus) und von D.M. Vital neuerdings aus
der Serra do Caraca nachgewiesen, C. luetzelburgii aus der Serra dos Orgaos und
der Serra do Itatiaia, Beiden fehlen die für C. fragilis typischen Brutblatter und
die durch apikale Kurztriebe pinselfórmigen Spitzen der Pflanzen.
2. CAMPYLOPUS MINARUM Par. (Dicranum laxobasis C. Müll.)
Die von C. MÜLLER 1900 in Hedwigia 39 : 252 unter dem Homonym
Dicranum laxobasis beschriebene Art wurde von PARIS 1904 in Campylopus
minarum umbenannt. Von dem nicht auffindbaren Holotypus fand sich in PC
(Hb. Thériot) eine Doublette, aus der hervorgeht, dass es sich bei dieser Art
um Campylopus surinamensis C. Müll. handelt. Diese Art war ursprünglich aus
den Guianas, dem Amazonastiefland (als C. marmellensis und C. gracilicaulis),
der Karibik und dem Südosten der USA (als C. gracilicaulis) bekannt, wo sie
auf Sandbóden im Tiefland vorkommt, tritt aber auch in Südostbrasilien in
Source : MNHN, Paris
CAMPYLOPUS VON BRASILIEN 443
ВО
BOS ROSS
аө roses =
6 Fig-1. — Campylopus luet-
zelburgii Herz. ex J.-P.
Frahm, Holotypus. 1 :
Blatt aus der Sten-
gelspitze (nat. Grósse
5 mm) 2 : Blatt-
spitze, 3 : Basale La-
minazellen, 4 : Obere
Laminazellen, 5 : Ha-
bitus (nat. Grösse 2
cm) in trockenem Zu-
stand, 6 : Rippen-
querschnitt aus der
Blattmitte. Massstrich
= 50 Um.
Fig. 2. — Obere Lamina-
zellen von 1. Campy-
lopus fragiliformis (a :
Vital 7706, b : Iso-
typus im Hb. des Ver-
fassers), 2. Campylo-
| L| pus fragilis (a : Earl
Bishop 14.2.1975, b :
Vitt 12374, Doubl.
E aus FLAS bzw. ALTA
im Hb. des Verf.). 3.
ux Campylopus luetzel-
burgii (a : Holotypus,
b : Paratypus).
3
Fig. 1
Fig. 2
QQ os #0300 009990
ШО
фу зе ce C (0р «ШИП
ai 200 52 ш УП
2а 2b 3a 3b
Source : MNHN. Paris
444 J.-P. FRAHM
grôsseren Нёһеп bis 1500 т auf, von wo sie schon früher als Campylopus ca-
tumbensis Broth. beschrieben worden war. Auch der Typus von Campylopus
minarum stammt hierher (Minas Geraes, Serra Caraca, in rupibus, leg. Ule
3/1892 nr. 1353).
— Campylopus surinamensis C. Müll. Linnaea 21 : 186, 1848.
— Campylopus minarum Par. Ind. Bryol. ed. 2, 1 : 318, 1904 syn. nov (Dicra-
num laxobasis C. Müll. Hedwigia 39 : 252, 1900).
3. CAMPYLOPUS BRACHYPHYLLULUS (C. Müll.) Broth.
Im Herbar Thériot (PC) fand sich ein Isotypus dieser nur von der Typusloka-
lität bekannt gewordenen Art (ein Holotypus in B existiert nicht mehr), der sich
als Campylopus savannarum erwies :
— Campylopus savannarum (C. Müll.) Mitt. J. Linn. Soc., Bot. 12 : 85, 1809
(Dicranum savannarum C. Müll. Syn. Musc. 2 : 596, 1851).
— Campylopus brachyphyllulus (C. Müll.) Broth. Nat. Pfl. 1(3) : 332, 1901
(Dicranum brachyphyllulum С. Müll. Hedwigia 39 : 260, 1900) syn. nov.
Typus : Brasilien, Minas Geraes, Caraca, ad rupes, Ule 1360 (PC, ISO).
4. CAMPYLOPUS BRASILIENSIS Broth. nom. nud.
Das von BROTHERUS in der Hedwigia 38 Beibl. 1 : 58, 1899 erwähnte
Campylopus brasiliensis basiert auf der Nr. 109 der Bryotheca brasiliensis
von Ше. Ein Beleg dieser Nummer im Herbar Fleischer (FH) : Brasilien, Rio
de Janeiro, Serra do Itatiaia, in paludibus 2300 m, leg. Ule 2/1894, beinhal-
tet jedoch kein Campylopus, sondern ein Atractylocarpus. Die gleiche Probe
wurde bereits 1898 von C. MÜLLER im Bull. Herb. Boissier 6 : 39 als Dicranum
brasiliense publiziert und von WILLIAMS 1928 im Bryologist 31 : 110 zu
Atractylocarpus gestellt. Das Material von Atractylocarpus brasiliensis gleicht
in jeder Hinsicht Herbarbelegen von A. longisetus (Hook.) Bartr., sodass A. brasi-
liensis mutmasslich mit A. longisetus identisch ist und dieses eine weitere,
sowohl in den Anden als auch im südostbrasilianischen Bergland vorkommende
Art wäre.
— Atractylocarpus brasiliensis (C. Müll.) Williams Bryologist 31 : 110, 1928
(Dicranum brasiliense С. Müll. Bull. Herb. Boissier 6 : 39, 1898).
— Campylopus brasiliensis Broth. Hedwigia 38 Beibl. 1 : 58, 1899 пот, nud.
5. DICRANUM RABENII Lor. und CAMPYLOPUS RUBRICAULIS Lindb.
in C. Müll.
Eine Überprüfung der Typen von Dicranum rabenii Lor. (Brasilia, com
Raben, B) und Campylopus rubricaulis Lindb. in C. Müll. (Brasilien, Caldas,
leg. Lindberg 18.9.1894, B) ergab, dass es sich bei beiden Arten um Campylopus
filifolius var. humilis handelt, einer in Südostbrasilien und im gesamten Areal
Source : MNHN. Paris
CAMPYLOPUS VON BRASILIEN 445
von С. filifolius durch das nördliche Südamerika bis nach Mittelamerika häufigen
Varietät im Bereich der Bergregenwälder.
— Campylopus filifolius (Hornsch.) Mitt. J. Linn. Soc. Bot. 12 : 76, 1869
(Dicranum filifolium Hornsch. Fl. Bras. 1 : 12, 1840) var. humilis (Mont.)
J.P. Frahm Nova Hedwigia 29 : 249, 1978 (Campylopus humilis Mont.
Ann. Sci. Nat. Bot. ser. 3, 4 : 110, 1845).
— Dicranum rabenii Lor. Moosstudien : 158, 1864, syn. nov.
— Campylopus rubricaulis Lindb. in C. Müll. Gen. Musc. Fr. : 169, 1900 nom.
nud., syn. nov.
6. CAMPYLOPUS SUBCUSPIDATUS (Hamp.) Jaeg.
Bei dem Typus dieser nur von der Typuslokalität (Brasilien, Pic de la Tijuca,
leg. Glaziou 19.4.1873 nr. 7096, PC) bekannt gewordenen und von HAMPE
(1870) beschriebenen Art handelt es sich um das, was 30 Jahre später als Campy-
lopus praealtus (C. Müll.) Par. beschrieben wurde. HAMPE gibt zur Unterschei-
dung dieser Art an : «vero D.ficrano) cuspidato affinis, sed caule interrupte
tomentoso, laxe foliato, lutescente; splendente, foliis patentibus, nervo laxiore,
triplo latiore omnini diversa.» In der Tat ähneln sich C. cuspidatus und sub-
cuspidatus (= praealtus) habituell bezüglich der Grösse und dem goldgelben
Glanz der Pflanzen sehr. Wie von HAMPE angegeben, ist bei C. subcuspidatus
im Gegensatz zu C. cuspidatus die Rippe wesentlich breiter (3/4 statt 1/3 der
Blattbreite am Blattgrund) und die Blatter stehen bei C. subcuspidatus weiter ab.
Ausserdem besitzt C. cuspidatus auffällige kollenchymatisch verdickte Blatt-
flügelzellen. Bei C. subcuspidatus ist ferner der basale Blattrand von einem Saum
aus dünnwandigen schmalen Zellen gesäumt, welcher bei C. cuspidatus fehlt.
Auch das Zellnetz ist in der oberen Hälfte der Lamina unterschiedlich, und
zwar verlángert, verdickt und deutlich getüpfelt bei С. cuspidatus und verlängert
oval, dickwandig, aber nicht deutlich getüpfelt bei С. subcuspidatus. Schliesslich
besitzt C. cuspidatus vielfach eine hyaline Haarspitze.
Aus Prioritätsgründen muss daher die wohlbekannte Bezeichnung C. praealtus
dem älteren Namen С, subcuspidatus weichen.
— Campylopus subcuspidatus (Hamp.) Jaeg. Ber. Thätigk. St. Gallischen Natur-
wiss. Ges. 1870-71 : 441, 1872 (Dicranum subcuspidatum Hamp. Vidensk.
Meddel. Dansk Naturhist. Foren. Kjébenhavn 1870 : 273).
— Campylopus praealtus (C. Müll.) Par. Ind. Bryol. : 96, 1900, syn. nov. (Dicra-
num praealtum C. Müll. Hedwigia 37 : 227, 1898).
Campylopus subcuspidatus ist unter der jüngeren Bezeichnung C. praealtus
von Puerto Rico, Venezuela, Surinam, Honduras und Brasilien bekannt und nach
einem bislang unpubliziertem Fund auch in Costa Rica nachgewiesen.
7. CAMPYLOPUS DISCRIMINATUS (Hamp.) Jaeg.
Eine Überprüfung der Syntypen von Campylopus discriminatus (Glaziou
Source : MNHN, Paris
446 J-P. FRAHM
7434, 7096, PC) und des Lectotyps (Glaziou 7147, H-BR) ergab, dass keine
artspezifischen Unterschiede zu Campylopus arctocarpus (Hornsch.) Mitt. vor-
liegen. Interessanterweise stellte sich bei Durchsicht der Glaziou-Kollektionen
in Paris heraus, dass Hampe als Bearbeiter dieser Aufsammlungen C. filifolius
(Hornsch.) Mitt. für C. arctocarpus hielt. Dementsprechend hat er die Aufsamm-
lungen des eigentlichen C. arctocarpus als C. discriminatus neu beschrieben.
Ausser von Hampe wurde die Art nur noch von BROTHERUS (1895, 1926)
aus Brasilien angegeben.
— Campylopus arctocarpus (Hornsch.) Mitt. J. Linn. Soc., Bot. 12 : 87, 1869
(Dicranum arctocarpum Hornsch. Fl. Bras. 1(2) : 12, 1840).
— Campylopus discriminatus (Hamp.) Jaeg. Ber. Thätigk. St. Gallischen Natur-
wiss. Ges. 1877-78 : 497, 1880 syn. nov. (Dicranum discriminatum Hamp.
Vidensk. Meddel. Dansk Naturhist. Foren. Kjóbenhawn 1878 : 254).
8. CAMPYLOPUS TORTILIPILUS J.-P, Frahm spec. nov.
Unter Aufsammlungen, die ich von D.M. Vital (Sao Paulo) bekam, befand
sich u.a. eine Probe eines haartragenden Campylopus mit so eindeutig neuen
Merkmalsausprágungen, dass die Art hier, obwohl nur von einer Stelle bekannt,
als neue Art beschrieben werden soll. Kennzeichnend für diese Art sind die
auffällig gelbgrünen niedrigen Rasen, Blätter, die in ein relativ langes (1/3 der
Blattlänge) gezähntes und wellig verbogenes hyalines Glashaar enden und ein
Rippenquerschnitt mit ventralen Substereiden. Abgesehen von den verbogenen
Glashaaren und der hellen Färbung ist C. tortilipilus einem С. pilifer Brid.
ähnlich, weicht aber durch verlängert ovale und in den Ecken verdickte Lamina-
zellen, einen gesägten oberen Blattrand und dickwandige basale Laminazellen
sehr deutlich von dieser Art ab,
Campylopus tortilipilus J.-P. Frahm spec. nov.
Plantae steriles, 5-10 mm altae, luteo-flavescentes, laxe caespitosae. Folia
3 mm longa, e basi contracto lanceolata, in pilum hyalinum longum serratum et
tortuosum apiculata. Costa: dimidiam folii occupante, in sectione transversali
substereidibus ventralibus et stereidibus dorsalibus. Cellulae laminae in parte
superiore ovatae-elongatae, incrassatae, 18-35 x 5-9 ит (1:4-5), т parte inferiore
rectangulares, incrassatae, 20-40 x 15 um, ad marginem subquadratae.
Material : In partial shade, on soil, in a cerrado vegetation, 10°33°5, 45^ 11 W,
Municipio de Corrente, Piaui State, leg. D.M. Vital 28.5.1978 nr. 8232 (Holoty-
pus SP, Isotypus im Herbar des Verfassers).
Der Nordosten Brasiliens ist bryologisch nur wenig bekannt und durchforscht,
sodass anzunehmen ist, dass C. tortilipilus dort noch an weiteren Stellen vor-
kommt. Ebenfalls nur aus diesem Gebiet Brasiliens ist Campylopus gardneri
(C. Müll.) Mitt. bekannt, was eine gewisse Eigenständigkeit dieses Florengebietes
ausdrückt.
Source : MNHN, Paris
CAMPYLOPUS VON BRASILIEN 447
ШИ
ОС
QUE
SOC
С
sa
OO
сш)
Kx o
NE
e
С,
D
©
Q
O
Io
S
[a]
| nnnc s
Fig. 3. — Campylopus tortilipilus J.P. Frahm spec. nov. 1 :Pflanze, Originalgrósse 5 mm,
2 : Blatt, Originalgrósse 3 mm incl. Glashaar, 3 :Blattspitze, 4 : Zellnetz der oberen
Lamina, 5 : Zellnetz des Blattgrundes, 6 : Rippenquerschnitt. Massstrich = 50 Um.
9. CAMPYLOPUS RECTISETUS (Hamp.) Jaeg.
Bei dem Typus dieser Art (Brasilien, Rio de Janeiro, Glaziou 6364, PC)
handelt es sich um C. arctocarpus (Hornsch.) Mitt., einer von Hampe verkannten
Art (vgl. Nummer 7 dieses Beitrages).
— Campylopus arctocarpus (Hornsch.) Mitt. J. Linn. Soc., Bot. 12 : 87, 1869
(Dicranum arctocarpum Hornsch. Fl. Bras. 1(2) : 12, 1840).
Source : MNHN, Paris
448 J-P. FRAHM
— Campylopus rectisetus (Hamp.) Jaeg. Ber. Thätigk. St. Gallischen Naturwiss.
Ges. 1877-78 : 385, 1880 (Dicranum rectisetum Hamp. Vidensk. Meddel.
Dansk Naturhist. Foren. Kjébenhavn 1875 : 137).
Campylopus rectisetus ist später nur noch von BROTHERUS (1895) aus
Minas Geraes angegeben worden; auch bei diesen Belegen (H-BR) handelt es sich
um C. arctocarpus.
LITERATUR
BROTHERUS V.F., 1895 — Nouvelles contributions à la flore bryologique du Brésil. Acta
Soc. Sci. Fenn. 19 :5.
BROTHERUS V.F., 1926 — Musci in : SCHIFFNER & WETTSTEIN, Ergebnisse der Bot.
Expedition der Akad. Wissen. nach Südbrasilien 1901. IL Band. Denkschr. Kaiserl. Akad.
Wiss., Math.-Naturwiss. Kl. 83 : 251-358.
FRAHM Ј.Р., 1979 — Die Campylopus-Arten Brasiliens. Rev. Bryol. Lichenol. 45 :127-178.
HAMPE E., 1870-78 —Symbolae ad floram Brasiliae centralis cognoscendam. Vidensk.
Meddel. Dansk Naturhist. Foren. Kjóbenhavn 1870 : 267, 1872 : 36, 1874 : 129, 1877-
78:251.
Source : MNHN. Paris
449
SYNOPSIS ОЕ RHIZOGONIACEAE BROTH. IN MALAYA'
М.С. MANUEL *
ABSTRACT. — Pyrrhobryum Mitt. emend. Manuel has three species (Р. spiniforme (Нейм)
Mitt., P. longiflorum Mitt., & P. latifolium (Bosch. & Lac.) Mitt.) in Malaya while Rhizogo-
nium Brid. has only a single species, В. novae-hollandiae (Brid.) Brid. A key to the two
genera and four species is provided along with critical notes on each species, and illustrations,
INTRODUCTION
Of the seven genera currently recognized in Rhizogoniaceae (BROTHERUS
1924, CROSBY 1979, NORRIS & ROBINSON 1979), two are present in Malaya
(Peninsular Malaysia and the Republic of Singapore). Pyrrhobryum Mitt. has
three species whereas Rhizogonium Brid. has only a single species. The recent
discovery of P. longiflorum Mitt. in Malaya (MANUEL 1980a) prompted a
critical reexamination of previous collections of Rhizogoniaceae in Malaya.
The results of that reexamination are reported herein.
METHODS AND MATERIALS
This study is based on about 120 specimens in the herbaria of KLU, SING,
and TNS. All observations, measurements, and drawings (except habit of game-
tophores) were made from specimens mounted in Hoyer's Solution (ANDER-
SON 1954).
SYSTEMATIC TREATMENT
Key to the Genera and Species of Rhizogoniaceae in Malaya
1. Leaves distichous, margins unistratose with single teeth, costa smooth dor-
sally; capsule almost without a neck... ............ 1. RHIZOGONIUM
1. Leaves not distichous, margins multistratose with double teeth, costa toothed
dorsally; capsule with distinct neck. ...... 2. PYRRHOBRYUM...... 2.
1. I thank Dr. B.C. Stone (KLU), for reading the manuscript, Dr. N.G. Miller (СН) and Dr.
A. Touw (L) for help with the literature, and the curators of SING and TNS, Financial
assistance from Universiti Malaya (Vote F 128/79 & 110/80) is acknowledged.
* Jabatan Botani, Universiti Malaya, Kuala Lumpur, Malaysia.
Cryptogamie, Bryol. Lichénol., 1981, 2, 4 : 449-455.
Source : MNHN, Paris
450 М.С. MANUEL
2. Medial stem leaves narrowed to insertion, 3.0-4.0(-5.0) mm long; inner
perichaetial bracts over 5.0-8.0 mm long, filiform acuminate. .... а
2. Medial stem leaves not narrowed to insertion (lower and upper le
occasionally slightly narrowed to insertion), 4.0-6.5 mm long; inner
perichaetial bracts 2.0-3.5(-4.1) mm long, acuminate to filiform acumi-
A У TT Par AO rok oh PA 2a. P. spiniforme
3. Medial stem leaves narrowly elliptic-lanceolate, apices long acuminate.
i km Qe s mi E os NO EE 2b. P. longiflorum
3. Medial stem leaves widely elliptic-lanceolate, apices acuminate. ......
vss ed WARNEN FE U КНК RE 2с. P. latifolium
1. RHIZOGONIUM Brid., Bryol. Univ. 2 : 644. 1827.
Note. — The genus Rhizogonium Brid. s.s. (MANUEL 1980b) is pantropic in
distribution and is represented by eight species within Malesia. One species,
К. novae-hollandiae (Brid.) Brid. occurs in Malaya.
Та. Rhizogonium novae-bollandiae (Brid.) Brid., Bryol. Univ. 2 :664. 2f. 1-3.
1827. — Fissidens novae-hollandiae Brid., Bot. Zeitung (Regensburg) 1 :
212, 234. 1802.
Distribution. — Malaya (Pahang), Borneo (Sabah), Australia, Tasmania, New
Zealand, Juan Fernandez, and Patagonia.
Note. — Rhizogonium novae-hollandiae (Brid.) Brid. is an antipodal species
with disjunct populations in Malesia at high elevations. In Malaya the species
has been found only in the montane forests between 1440 and 2100 m while
in Borneo it has been reported from Mt Kinabalu (DIXON 1935).
Specimens examined. — MALAYSIA. PAHANG : Cameron Highlands, Brin-
chang, ca 5000 ft., Clear 1329 (KLU). Cameron Highlands, Tanah Rata, 4800
ft., Henderson 17829 (SING). Gunung Tahan, 5500-7000 ft., Nur 7976 (SING).
Gunong Tahan, + 5500 ft., Holttum 20926a (SING). SABAH (Borneo) : Kam-
borangah ridge forest, 7200 ft., Holttum 25655 (SING).
2. PYRRHOBRYUM Mitt. emend. Manuel, J. Linn. Soc., Bot. 10 : 174. 1868.
Note. — For a discussion of the classification of Rhizogonium and Pyrrho-
bryum see MANUEL (1980b).
2a. Pyrrbobryum spiniforme (Hedw.) Mitt., J. Linn. Soc., Bot. 10 : 174.
1869. — Hypnum spiniforme Hedw., Spec. Musc. : 236. 1801. — Rhizo-
gonium spiniforme (Hedw.) Bruch in KRAUSS, Flora 29 : 134. 1846.
Distribution. — Malaya (Johor, Kedah, Kelantan, Pahang, Penang & Selangor;
Singapore); pantropic.
Notes. — 1. NOGUCHI (1973) reported Rhizogonium badakense Fleisch.
(Pyrrhobryum spiniforme var. badakense (Fleisch.) Manuel) from the state of
Source : MNHN, Paris
RHIZOGONIACEAE IN MALAYA 451
реа SE UC
b 10mm
С Nr OE
e 0.2 mm
Fig. 1. — Rhizogonium novae-hollandiae (Brid.) Brid. a. Gametophore, b. Leaves, c.Me-
dial leaf cells, d. Marginal tooth, e. Leaf apex.
Pahang. The specimen upon which he based his report (Inoue 10797, 10904,
16223 & 16371) are P. spiniforme var. spiniforme.
2. Within the local populations of Р, spiniforme there are some morphologi-
cal «entities» that are more or less distinct and occur in more than one locality.
Whether these «entities» are the result of a similar phenotypic response to
similar microenvironments or the result of genotypic differences, | do not
know and hence have not given them formal recognition under the ICBN.
Further investigations are in progress. i
— ENTITY A. Gametophore small; medial stem leaves short, narrow, and plane.
(Manuel 2419, Nur 9191, Henderson 19640, & JSG 248; vide specim. exam.).
Source : MNHN, Paris
о
\
El Oca
JA 0 mm
c&e 0.2mm
i&h 0.1mm
b,d,f,9 8
c&
Fig. 2.—Pyrrhobryum. P. longiflorum Mitt. a. Gametophore, b. Medial stem leaves,
c. Leaf base, d. Inner perichaetial bracts. Р. spiniforme (Hedw.) Mitt. e. Leaf base,
f. Medial stem leaves, g. Inner perichaetial bract. Р, latifolium (Bosch, & Lac.) Mitt.
i. Medial leaf cells, h. Marginal tooth, j. Medial stem leaves.
Source : MNHN. Paris
RHIZOGONIACEAE IN MALAYA 453
— ENTITY В. Gametophore robust; medial stem leaves long, broad, and slightly
keeled (Holttum 10697 & 18068, Ridley 106 & 526, & Curtis 5; vide specim.
exam.).
The perichaetial bracts of the above two «entities» are well within the normal
range of variation for P. spiniforme var.spiniforme (INOUE & IWATSUKI 1976).
Specimens examined. — SINGAPORE. Chan Chu Kang, J.G.G. Gamble 248
(SING.). Chan Chu Kang, Ridley 113 (SING). Bukit Timah, Nur 9191 (SING).
Kranji Ridley 106 (SING). Brittian Peak, Luerek 675 (SING). Bukit Tuish,
Ridley 250 (SING). MALAYSIA, JOHOR : Gunong Belumut, 3000 ft., Holttum
10698 (SING) & 10697 (SING). Gunong Panti, 1600 ft., Holttum 18068 (SING).
KEDAH : Gunong Raya, Harift & Nur 7144 (SING). Langkawi, Curtis, Sept.
1890 (SING). KELANTAN : Gunong Sitong, 2600 ft., Nur 12236 (SING).
Sungei Keteh, Henderson 19640 (SING). PAHANG : Fraser Hill : 4000-4370 ft.,
Burkill & Holttum 8468 (SING) & 8784 (SING); 4000 ft., Ridley 287 (SING);
Kalong 22430 (SING); Burkill 1166 (SING), 2061 (SING), & 2114 (SING);
Manuel 2778 (KLU); Poore s.m. (KLU); Johnson B1037 (KLU); Poore 48
(KLU). Cameron Highlands : Clear 1195 (KLU) & 1339 (KLU); Tanah Rata,
5200 ft., Inoue 10059 (TNS), 10060 (TNS) & 10061 (TNS); Brinchang, 600 ft.,
Inoue 10797 (TNS) & 10904 (TNS), Poore 116 (KLU); Near Robinson’s Falls,
5000 ft., Inoue 10753 (TNS), 10787 (TNS), & 10788 (TNS); around Robinson’s
Water Fall, Inoue 16223 (TNS) & 16371 (TNS), Henderson 11722 (SING).
Gunong Tahan, 5500 ft., Holttum 20926 (SING). Tahan River, Ridley, 1841
(SING). Gunong Ber (icalica?), Nur 121 (SING). Telom, Nur 1900 (SING),
94 (SING), 142 (SING). Gua Kajang, 3383 ft., Henderson 18271 (SING). Pulau
Tioman, Gunong Kajang, 2000 ft. D.W. Lee, 24 May 1974 (KLU). Genting
Sempah, 3000 ft., Stone 9572a (KLU). PERAK : Taiping Hills, 4500, Anderson
312 (SING) & 314b (SING). Maxwell Hill, Ridley, 1893 (SING). Jor Tapah,
Haniff 16173 (SING). Temengoh, Ridley 182 (SING). Lower Camp, Gunong
Batu Puteh, Wray 1047 (SING). Bird’s Hill, 3800 ft., Burkill 12603 (SING).
PENANG : Penang Hill : 2000 ft., Johnson, May 1966 (KLU); Ridley 526
(SING). Tiger Hill, 2100 ft, W. (Suikiee?) 3315 (SING). Richmond’s Pool,
W. (Suikiee?) 2607 (SING); Haniff 15003 (SING) & 15020 (SING). Govern-
ment Hill, Curtis 5 (SING). SELANGOR : Genting Bidai, Ridley 407 (SING).
Near 21 mile mark on old Genting Highland Rd., ca 1000 m, Manuel 2419
(KLU). Bukit Hitam s. leg., 1896 (SING).
2b. Pyrrbobryum longiflorum Mitt., J. Linn. Soc., Bot. 10 : 174. 1868. —
Rhizogonium longiflorum (МНЕ) Jaeg., Ber. Thätigk. St. Gallischen
Naturwiss. Ges. 1873-74 : 223. 1875 (Ad. 1:685).
Distribution. — Malaya (Johore, Negri Sembilan, & Pahang) and Borneo
(Sabah & Sarawak).
Notes. — 1. IWATSUKI (1969 & 1972) stated that Pyrrhobryum longiflorum
Mitt. (as Rhizogonium longiflorum (Mitt.) Таев.) is a lowland species (below
330 m) in northern Borneo and is clearly separated from P. spiniforme (Hedw.)
Mitt. (600-1500 m elev.) in elevational distribution. From the data on herbarium
Source : MNHN, Paris
454 М.С. MANUEL
labels, the same elevational distribution seems to occur in Malaya except that
Р. spiniforme reaches 1650 m on Gunong Tahan.
2. The species is characterized by a) medial stem leaves narrowed at inser-
tion, b) long, filiform acuminate perichaetial bracts, and c) dioicous sexuality
(IWATSUKI 1969).
Specimens examined. — MALAYSIA. JOHORE : Gunong Kambak, ca. 700
ft., on fallen tree, Holttum 9459 (SING). Mersing, Clear 1362 (KLU) & 1363
(KLU). NEGRI SEMBILAN : Selaru, F.R., on fallen tree, Holttum 9745 (SING).
PAHANG : Taman Negara : Lata Berkoh area on Sungai Tahan, 200-300 ft.,
Manuel 2566 (KLU), 2568a (KLU); Bukit Indah, ca. 250 ft., Manuel 2663
(KLU); & Bukit Tersek Trail ca. 200-1000 ft., Manuel 2535 (KLU). Tahan River,
Ridley, 1893 (SING).
2c. Pyrrhobryum latifolium (Bosch & Lac.) Mitt., J. Linn. Soc., Bot. 10 :
175. 1868. — Rhizogonium latifolium Bosch & Lac., Bryol. Jav. 2 : 2.
133. 1861.
Distribution. — Malaya (Johor, Perak, & Singapore), Banka, Borneo (Sabah
& Kalimantan Barat), Philippines, and New Guinea.
Note. — Pyrrhobryum latifolium (Bosch & Lac.) Mitt. and P. longiflorum are
separated by differences in the apices and the shape of the medial stem leaves.
1 have not seen any overlap in the shape of the medial stem leaves of the two
species, but the apical and basal stem leaves of P. longiflorum sometimes resem-
ble the medial stem leaves of P. latifolium.
Specimens examined. — SINGAPORE : Jungle Fall Valley, on rock, L.H.
Merton, 19 May 1960 (KLU). Bukit Timah, Ridley 652A (SING). Rock Path,
Bukit Timah Forest Reserve, Sinclair 7355 (SING). MALAYSIA. JOHOR :
Sungei Semagot Kanan, 30 mile mark Kota Tinggi-Mersing Rd., Sinclair 10757
(SING). PERAK : Bidor Road (Tapah ?), s.n. 154 (SING).
LITERATURE CITED
ANDERSON L.E., 1954 — Hoyer's solution as a rapid permanent mounting medium for
bryophytes. Bryologist 57 : 242-244.
BROTHERUS V.F., 1924 — Musci. In ENGLER & PRANTL, Natürl. Pflanzenfam., Ed. 2.
10 : 1-478.
CROSBY M.R., 1979 — An alphabetical catalogue of the families of mosses with included
genera listed alphabetically under each family name. St. Louis, 12 pp.
DIXON H.N., 1935 — A contribution to the moss flora of Borneo. J. Linn. Soc., Bot. 50 :
57-140.
INOUE S. & IWATSUKI Z., 1976 — A cytotaxonomic study of the genus Rhizogonium
Brid. (Musci). J. Hattori Bot. Lab. 41 : 389-403.
Source : MNHN, Paris
RHIZOGONIACEAE IN MALAYA 455
IWATSUKI Z., 1969 — Bryological miscellanies XIX, Some interesting mosses from North
Borneo. J. Hattori Bot. Lab. 35 : 269-287.
IWATSUKI Z., 1972 — Geographical isolation and speciation of Bryophytes in some islands
of Eastern Asia. J. Hattori Bot. Lab. 35 : 126-141,
MANUEL M.G., 1980a — A contribution to the moss flora of Taman Negara (Malaysia).
Lindbergia 6 : 37-40.
MANUEL M.G., 1980b — Miscellanea Bryologica 11. Classification of Rhizogonium Brid.,
Penzigiella hookeri Gangulee, and some nomina nuda. Cryptogamie, Bryol. Lichénol.
1(1) :67-72.
NOGUCHI A., 1973 — Mosses of Malaya collected by Dr. Hiroshi Inoue. Bull. Natl. Sci.
Mus. 16(2) : 293-304.
NORRIS DH. & ROBINSON H., 1979 — The Systematic Position of Bryobrothera crenu-
lata. Bryologist 82 : 305-309.
Source : MNHN. Paris
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457
REGENERATION AND GEMMA DEVELOPMENT
IN HYOPHILA CRENULATA C. MUELL. EX DUS.
5.0, OLARINMOYE *
ABSTRACT. — Further culture studies and field observations have proved that gemmae of
Hyophila crenulata C. Muell. ex Dus. are protonematal and not rhizoidal. Leaves of the
moss readily regenerate and the protonemal strands which produced copious gemmae have
supplied the required information on the origin of gemmae in Hyophila. Some of the
gemmae, while still attached to gemmifers, germinated to produce more gemmae in culture.
Gemmae production is related to humidity; higher humidity resulting on more abundant
gemmae production. Flooding of shoots of H. crenulata resulted in abundant production of
basal protonematal gemmae. Gemmae production in the moss, must have contributed
significantly to its extensive cover in its ecological niches, usually gutters, drainage areas
etc. and to its ecological success in such areas.
INTRODUCTION
Abundant production of gemmae has been reported in Hyophila crenulata
(OLARINMOYE 1981). Occurrence of gemmae has also been reported in H.
tortula (ANDREWS & REDFEARN 1965) and in H. propagulifera (NEHIRA
1969). However, the exact origin, protonemal or rhizoidal, of these gemmae,
has not been established.
In nature, gemmae of H. crenulata are produced both on long tufts of gemmi-
fers arizing from the apices of shoots and in grapefashion bunches on exposed
bases of the shoots.
In the present work, the origin of the gemmifers and their gemmae, as well
as the effects of moisture on the production of gemmae has been elucidated.
The possible role of gemmae in the ecological success of the moss, Hyophila,
was discussed.
MATERIALS AND METHODS
Hyophila crenulata was collected from the almost pure stands in the wet
gutter in front of the author's laboratory at Ibadan University. The material
* Department of Botany, University of Ibadan, Ibadan, Nigeria.
Cryptogamie, Bryol. Lichénol., 1981, 2, 4 : 457-460.
Source : MNHN, Paris
458 S.0. OLARINMOYE
was divided into suitable portions and each portion was put in a large evapora-
ting dish and kept moist as required during the period of use.
The shoots used in regeneration experiments were thoroughly washed in run-
ning tap water and later surface-sterilized in chlorex for 3-4 minutes before
being finally washed in several changes of sterile distilled water.
15 detached leaves were plated on pads of two No. 3 Whatman filter papers,
moistened with distilled water, in 10 cm plastic Petri dishes. The culture was
maintained in a 16h-8h light cycle at a light intensity of 1000 lux and a tempe-
rature of 23°С £ 2?C, at the culture level. Each plate was duplicated. The
cultures were observed daily for regeneration, which was recorded as percentage
regeneration per day. Gemmae production, and their positions were also ob-
served.
In another set-up, detached leaves were cut into two approximately equal
halves and plated side by side under a similar set-up as whole leaves. 25 pieces
of each half were plated. Other detached leaves were fragmented into tiny bits
and plated. Not less than 45 fragments were plated in each Petri dish.
In a final set-up, 20 whole shoots of H. crenulata were introduced into a
Petri dish and then flooded with distilled water. The covered Petri dish was
subjected to the same treatment as earlier described. All experiments were
duplicated.
RESULT
In all the set-ups, regeneration commenced on the 3rd day and reached the
peak on the 8th day (Tab. 1). Regeneration occurred in all treatments, though
lower in the detached whole leaf treatment in the early days. A maximum of
less than 100% was recorded in the mutilated leaf segments because a number
of fragments died before regeneration started (Tab. 1). Regeneration occurred
only on the cut ends in cut and fragmented pieces, while in whole leaves the
bulk of regenerants arose at the base, and occasionally on the midrib.
Tab. 1. — Regeneration in leaves of Hyophila crenulata
Treatment % regeneration
Day 3 Day 5 Day 8
` Whole leaf 35 2.0 8052.5 100
Upper half of leaf 90+1.0 95415
Lower half of leaf 881.5 97+2.0
Mutilated leaf 75+2.0 8550.5
In all cultures where regeneration was recorded, gemmae, similar to those
usually found in nature, were produced (Tab. 2). They occurred at different
stages of development. These gemmae were borne on gemmifers produced on
protonemal strands (Pl. 1, 1). In some cases the gemmae were produced in
Source : MNHN, Paris
HYOPHILA CRENULATA 459
Tab. 2. — Gemma production on regenerants from leaves of Hyophila crenulata.
% gemma production was assessed from the number of regenerating leaves or leaf fragments
producing gemmae in a culture dish.
Treatment | Wholeleaf Upper half — Lower half Mutilated leaf
% gemma production | 100 100 100 100
bunches on the protonemal strands. The quantity of gemmae produced was
а function of the ramification of protonemal strands in the cultures. Gemmi-
fers and gemmae however, were not produced on all protonemal strands.
When whole shoots were flooded with water, gemmifers and gemmae were
produced both on the young leaves at the apices and at the base of the shoots.
Gemmae were borne on long tufts of gemmifers when they occur on leaves
at the apices while they were either in grape-like bunches or on a few strands
of protonemata when at the base (Pl. 1, 2).
Pl. 1. — 1 : Bunches of protonemal gemmae from the leaf. 2 : Basal protonemal gemmae.
Some gemmae produced on regenerating leaves germinated and produced
other gemmae, while still attached to the original gemmifer (Fig.).
Source : MNHN, Paris
460 S.O. OLARINMOYE
DISCUSSION
Leaves of H. crenulata regenerated readily in
culture. The protonemal strands produced the
gemmifers on which the gemmae were borne. Pro-
duction of gemmae on long gemmifers on leaves
and in large bunches at the base of whole shoots
has been confirmed in later collections from
different habitats. It has been found that H.
crenulata collected on cement slabs and other
places not regularly inundated, produced apical
gemmae mainly. Those in gutters with regular
inundation produced both apical and basal gem-
mae. This apparently occurs because the base of
shoots are normally more exposed in those gro-
wing in gutters than in those on concrete slabs.
Fig.— Secondary gemma pro-
duction from a primary The observed production of secondary gemmae
one. PG :primary gemma. on gemmifers produced from primary gemmae is
SG : secondary gemma. similar to that observed on Schistostega pennata
Ah (Hedw.) Web. & Mohr (EDWARDS 1978).
The ready production of regenerants and the
abundant production of gemmifers and gemmae
on them must be of tremendous ecological importance to the moss, H. crenulata.
While it would surely serve as an effective means of spread, it also confers to the
moss competitive advantage and thus enchances its survival.
REFERENCES
ANDREWS S. and REDFEARN P.L. Jr., 1965 — Observations on the germination of the
gemmae of Hyophila tortula (Schwaegr.) Hampe. Bryologist 68 : 345-347.
EDWARDS S.R., 1978 — Protonemal gemmae in Schistostega pennata (Hedw.) Wb. & Mohr.
J. Bryol. 10 169-72.
NEHIRA K., 1969 — The germination of gemmae in three mosses. Hikobia 5 : 189-195.
OLARINMOYE S.O., 1981 — Gemmae in Hyophila crenulata C. Muell. ex Dus. Nova
Hedwigia (in press).
Source : MNHN. Paris
461
CONTRIBUTION A L'ÉTUDE DES LICHENS
DU KIVU (ZAÏRE), DU RWANDA ET DU BURUNDI
VI. Les genres COCCOCARPIA Pers. et LOBARIA (Schreb.) Hoffm.
Е. SÉRUSIAUX*
RÉSUMÉ. — Le genre Coccocarpia est représenté dans l'est de l’Afrique centrale par trois
espèces : С. erythroxyli (Spreng.) Swinscow & Krog, répandu dans les milieux de forêts et
savanes boisées jusqu'à 2100 m d'altitude, C. palmicola (Spreng. Arvidss. & D. Gall.,
trouvant son optimum dans la forêt de montagne jusqu'à 2450 m, et C. pellita (Ach.) Müll.
Arg. connu seulement dans la chaîne des Birunga vers 2400 m. Le genre Lobaria compte
trois espèces : L. pulmonaria (L.) Hoffm., épiphyte rare localisé aux stations humides de la
forêt de montagne entre 2000 et 2700 m, L. retigera (Bory) Trev., fréquent à l'état épiphy-
tique et terricole dans des milieux analogues entre 1900 et 2500 m, et Lobaria sublaevis
(NyL) Yoshim. dont c'est la première mention en Afrique continentale.
ABSTRACT. — Revision of the genera Coccocarpia and Lobaria in Kivu (Zaire), Rwanda
and Burundi, The first one is represented by three species : С, erythroxyli (Spreng.) Swins-
cow & Krog widespread in forests and wooded-savannas up to 2100 m elevation, C. palmi-
cola (Spreng. Arvidss. & D. Gall. mostly developped in montane forests up to 2450 m
elevation and С. pellita (Ach.) Miill. Arg.restricted to the Birunga volcanoes around 2400 m.
Three Lobaria species are present : Г. pulmonaria (L.) Hoffm. rare epiphyte confined to
humid stations inside montane forests from 2000 to 2700 m, L. retigera (Bory) Trev.
epiphyte or terricolous in the same biotopes between 1900 and 2500 m but much more
abundant and Lobaria sublaevis (Nyl.) Yoshim. which is mentioned for the first time in
continental Africa.
Nous poursuivons ici la série de travaux consacrés aux lichens du Kivu (Zaïre),
du Rwanda et du Burundi, par l'étude de deux genres (Coccocarpia et Lobaria)
modestement représentés dans cette partie de l'Afrique. La comparaison est
cependant instructive avec d'autres régions intertropicales (Amérique centrale,
Indonésie et Asie du S-E). De plus, les espèces de ces deux genres présentes dans
la dition sont bien connues taxonomiquement. On se reportera donc au remar-
quable travail de YOSHIMURA (1971) pour de plus amples détails sur les
Lobaria et à ceux de SWINSCOW & KROG (1976) et de ARVIDSSON & GAL-
+ Aspirant du FN.RS. - Département de Botanique, Université de Liège, Sart Tilman,
B-4000 Liège, Belgique.
Cryptogamie, Bryol. Lichénol., 1981, 2, 4 : 461-469.
Source : MNHN, Paris
462 Е. SÉRUSIAUX
LOWAY (1979) pour les Coccocarpia. Sauf indication contraire, le matériel est
conservé à LG, mais des doubles doivent être distribués dans plusieurs herbaria.
COCCOCARPIA Persoon
In C. GAUDICHAUD, Voyage autour du monde entrepris par ordre du Roi
(u) exécuté sur les corvettes de S.M. L'Uranie et la Physicienne pendant les
années 1817, 1818, 1819 et 1820 par M. Louis de Freycinet. Paris, Pillet Ainé,
1826. Bot. : 2061.
Espéce-type. — Coccocarpia molybdea Pers., loc. cit. : 206 (lectotyp. : voir
В. SANTESSON, Sym. Bot. Upsal. 12 (1) : 414, 1952, sub «C. molybdina
Pers.»).
COCCOCARPIA ERYTHROXYLI (Spreng.) Swinscow & Krog
Norw. J. Bot. 23 : 256, 1976 (comme «erythroxili
Basionyme. — Lecidea erythroxyli Spreng. Kungl. Vetensk. - Academ. Nya
Handl. 1 :47, 1820.
Type. — Guadeloupe, in cortice Erythroxyli squamati, Martius 1818 (lecto-
type, TO).
= Coccocarpia parmelivides (Hook.) Curtis (pour plus de détails sur la syno-
nymie, voir SWINSCOW & KROG 1976 : 256).
ZAIRE : District forestier central : Irangi, station de recherches de l'I.R.S.A.C.,
près de la rivière Luhoho, env. 850 m, sur racines-échasses moussues d'Uapaca en
lisière de la forêt à la limite de la clairière de la station, Lambinon 72/08 (fer-
tile).
RWANDA : District afro-montagnard : Forêt de Rugege, lieu-dit Ruwanku-
ba, 1950 m, forêt de vallée à Syzygium rowlandii, sur les lianes et les rameaux
ligneux couverts de bryophytes dans le sous-bois, vers 1-2 m de haut, Lambinon
71/1076 (fertile). — Ibid., vers le km 107 de la route Butare-Cyangugu (non loin
de Garamba), env. 1900 m, sur branche de Ficalhoa laurifolia, forêt de mon-
tagne humide en bordure de la route, Lambinon 74/844 (fertile). — Ibid., entre
1. Comme C. GAUDICHAUD l'indique dans sa préface, c'est Persoon qui s'est chargé
de la partie descriptive relative aux champignons et lichens récoltés à l'occasion de ces expé-
ditions. Le matériel étudié était assurément de mauvaise qualité, puisque (voir préface et
note n95 en р. 191), le bateau ayant fait naufrage et échoué aux Îles Malouines, les plantes
ont «macéré» dans l'eau de mer puis ont été reséchées. Par ailleurs, de nombreux taxons
de lichens (Pyrenula, Porina, Arthonia, Verrucaria, Opegrapha, ...) sont cités dans le para-
graphe concernant les Fungi, alors que l'auteur les considére bien comme des lichens. Il
doit s'agir là d'un des tout premiers essais d'intégration de la systématique des lichens au
sein de celle des champignons, méme si ce n'est qu'une démarche implicite. А noter aussi
que Coccocarpia viridescens Pers., cité dans ce texte, est un nomen nudum : il n'y a ni
description, ni diagnose.
Source : MNHN. Paris
LICHENS DU KIVU, DU RWANDA ET DU BURUNDI 463
Garamba et la plantation de Gisakura, env. 1900 m, fragment de forêt de mon-
tagne humide à Mimulopsis arborescens en bordure de la piste, sur branchettes
mortes, Lambinon 74/1059 (fertile). — Vallée de la Bikeneko à l'W du poste
minier de Gikungu (env. 30 km N de Rutsiro), env. 2100 m, grosses branches
d'un Neoboutonia macrocalyx, forêt secondarisée au bord de la rivière, Lambi-
non 74/700 (fertile).
District du Rwanda et du Burundi : Ntyazo (région du Mayaga), env. 1400 m,
fourrés xérophiles, sur branche de Carissa edulis, Lambinon 80/297 (stérile, trés
petit spécimen).
BURUNDI : District du Rwanda et du Burundi : Kinazi, crête entre la Ruvu-
bu et la Chizanye (pointe Е du Burundi vers le territoire tanzanien), env. 1400 m,
sur petite branche de Parinari curatellifolia, savane herbeuse boisée à Entada,
Combretum, Parinari, .., Lambinon 78/425 (fertile). — Cankuzo, route vers
Mugera-Murore près de la bifurcation, env. 1650 m, petite galerie forestière à
Syzygium cordatum, Macaranga schweinfurthii, Carapa, … branches d'arbres
abattus (vers 6-7 m de haut?), Lambinon 80/621 (fertile).
District du Mosso-Malagarasi : Plaine de la Musindozi, entre la plaine d'avia-
tion du Gihofi et Mulamba, env. 1200 m, savane boisée à Oxytenanthera abyssi-
nica, sur chaumes d'Oxytenanthera mort, Lambinon 74/1384 (fertile).
Matériel africain complémentaire :
ZAÏRE : Plateau de la Luweo, Yangambi, env. 470 m, forêt primitive ombro-
phile, sur branche maîtresse et dans la cime d'un Klainedoxa, Louis 6664 (BR).
Commentaires. — 1. Coccocarpia erythroxyli est une espèce pantropicale qui
s'étend jusqu'aux régions tempérées océaniques. Dans la dition, sans être fré-
quent, c'est un épiphyte assez largement distribué dans pratiquement tous les
milieux naturels ou peu altérés, à atmosphére suffisamment humide (foréts,
savanes boisées, parfois bosquets xérophiles) elle ne dépasse toutefois pas
2100 m d’altitude.
2. Certains spécimens, en particulier le n° 72/08, ont une marge lobulée
et sont munis de lobules plus ou moins laminaux. Ils correspondent à С. pellita
var. semiincisa Müll. Arg. (Flora 65 : 321, 1882); nous les considérons comme de
simples variations sans valeur taxonomique.
3. Notons que dans une étude du genre en Afrique orientale, SWINSCOW &
KROG (1976 : 253) signalent C. azurella Nyl. du Ruwenzori. Cette espéce se
distingue de C. erythroxyli par ses lobes beaucoup plus étroits, tronqués à
l'apex, et par ses spores globuleuses.
COCCOCARPIA PALMICOLA (Spreng.) Arvidss. & D. Gall.
Bot. Not. 132 :242, 1979.
Basionyme. — Lecidea palmicola Spreng, Kungl. Vetensk-Academ. Nya
Handl. 1 :46, 1820.
Туре. — Guadeloupe, in cortice Cocos nucifera, 1817 (lectotype, TO).
= Coccocarpia cronia (Tuck.) Vain. (pour plus de détails sur la synonymie,
voir SWINSCOW & KROG 1976 : 253).
Source : MNHN, Paris
464 E. SERUSIAUX
ZAÏRE : District forestier central : Irangi, réserve de I'I.R.S.A.C., sur la rive
droite de la rivière Luhoho, env. 850 т, forêt équatoriale, sur petites branches
de Brazzea longipedicellata vers 5 т de haut, dans le bas d'un petit versant,
Lambinon 72/21 (stérile).
District du Graben occidental : Ile Idjwi, Rwagahe (côte occidentale de la
partie nord de l'île), 1460 m, face latérale + ombragée d'un gros bloc granitique
éclairé en lisière de la forêt secondarisée, Lambinon 78/381 (stérile).
RWANDA : District afro-montagnard : Forêt de Rugege, colline entre le mont
Muzimu et le Bigugu, au-dessus de la rivière Bizumu, 2450 m, tronc de Syzygium
parvifolium en lisière de la forêt, Lambinon 72/1009 (stérile). — Ibid., vers le km
91 de la route Butare-Cyangugu (entre Pindura et Uwinka), environ 2400 m,
talus (schiste métamorphisé «pourri») frais, en lisière de la forêt de montagne
+ secondarisée, Lambinon 74/783 (stérile) et 74/784 (fertile). — Ibid., entre
Garamba et la plantation de Gisakura, env. 1900 m, fragment de forêt de mon-
tagne humide à Mimulopsis arborescens au bord de la piste, sur branchettes
mortes, Lambinon 74/1060 (fertile). — Ibid., km 81 de la route Butare-Cyan-
gugu (un peu à I'W du marais Gasare), env. 2320 m, talus (schiste métamorphisé
«pourri») en bordure de la forêt de montagne + secondarisée, Lambinon 74/
1077 (stérile). — Ibid., lieu-dit Ruwankuba, 1950 m, forêt de vallée à Syzygium
rowlandii, sur les lianes et les rameaux ligneux couverts de bryophytes dans le
sous-bois, à 1-2 m de haut, Lambinon 71/1077 (stérile). — Ibid., sur feuilles
mortes de Symphonia globulifera, Lambinon 71/1083 (stérile).
BURUNDI : District du Rwanda et du Burundi ; Kinazi (entre Cankuzo et Ми-
hinga), env. 1400 m, grosses branches de Parinari dans la savane-parc de versant,
Lambinon 74/1457 (stérile). — Gitwenge, colline Niabitangu (au N de la mission,
pente vers la dépression du Mosso), env. 1780 m, petite galerie forestiére traver-
sant la savane dégradée, tronc ombragé de Syzygium, Lambinon 80/555 (stérile).
Matériel africain complémentaire :
ZAÏRE : Plateau de la Luweo, Yangambi, env. 470 m, forêt primitive ombro-
phile, sur branche maîtresse d'un Tessmannia, Louis 6565 (BR).
LA RÉUNION : Cirque de Cilaos, Grand Matarum, 1450 m, forêt des bois de
couleur, sur tronc, J.L. De Sloover 17715 (NAM, LG).
Commentaires. — 1. Nous suivons ARVIDSSON & GALLOWAY (1979 :
243-244) dans leur interprétation du type de Lecidea palmicola Spreng. et dès
lors dans la mise еп synonymie de Coccocarpia стопа. С. palmicola s'avère ainsi
une espèce répandue dans toutes les régions intertropicales et débordant sou-
vent largement dans les zones tempérées des deux hémisphères. Dans notre
dition, c'est une espèce trouvant manifestement son optimum dans la forêt
de montagne, où elle ne dépasse toutefois pas 2450 m d'altitude. Elle colonise
tant les branches et lianes du sous-bois que les talus schisteux frais: une fois
elle a été notée comme une espèce pseudofoliicole accidentelle.
2. Trois spécimens (Lambinon 71/1077, 71/1083 et 72/21) ont, par rapport
aux autres collections, un thalle de petite taille (1-2 cm), fragile, d'une couleur
plus bleutée et muni d’isidies très fines et très fragiles, cylindriques et non rami-
fées; ces variations semblent dues à un habitat plus ombragé et ne justifient pas
une distinction taxonomique.
Source - MNHN. Paris
LICHENS DU KIVU, DU RWANDA ET DU BURUNDI 465
COCCOCARPIA PELLITA (Ach.) Müll. Arg.
Flora 65 : 320, 1882.
Basionyme. — Parmelia pellita Ach., Lich. Univ. : 468, 1810.
Type. — Localité inconnue, «Habitat ad truncos arborum Indiae Occidenta-
lis» (lectotype, H-Ach.).
RWANDA : District afro-montagnard : Chaîne des Birunga, pied sud du
Gahinga, 2400 m, forét de montagne secondarisée à dominance de Neoboutonia,
tronc incliné de Neoboutonia macrocalyx, Lambinon 72/657 (stérile). — Ibid.,
Kinigi (NW de Ruhengeri), env. 2300 m, plantation Rops, tronc d'Eucalyptus
planté entre un chemin et des friches, Lambinon 72/711 (fertile). — Ibid., entre
Kinigi et le pied du Gahinga, en contrebas de la selle entre le Sabyinyo et le
Gahinga, 2400 m, forét de montagne broussailleuse, tronc incliné de Maesa
lanceolata var. mildbraedii, Lambinon 72/918 (stérile).
Commentaires. — 1. Dans la dition, cette espéce, pantropicale et occasionnel-
lement tempérée-chaude, est limitée aux horizons supérieurs de la forét de mon-
tagne de la chaine des Birunga. Elle se comporte de la méme façon dans les ré-
gions d'Afrique orientale étudiées par SWINSCOW & KROG (1976); par contre,
en Nouvelle-Zélande, elle est observée entre 0 et 400 m d'altitude. Dans la di-
tion, elle est toujours épiphyte, alors que, en Amérique centrale, elle est parfois
épiphylle (SANTESSON 1952 : 422).
2. SWINSCOW & KROG (1976 : 258) ont soulevé le probléme de la distinc-
tion entre C. pellita et C. erythroxyli, se demandant si C. pellita n'est pas simple-
ment une forme stationnelle de C. erythroxyli. Dans notre dition, méme si ces
deux espéces ne sont pas sympatriques, leur morphologie est suffisamment
différente pour qu'elles soient maintenues comme distinctes. C. pellita possède
des isidies squamuleuses, souvent trés fines et incisées, tandis que C. erythroxyli
a des isidies cylindriques, épaisses à enflées.
LOBARIA (Schreb.) Hoffm.
Deutschl. Fl. 2 : 138, 1796.
Lichen (sect. ?) Lobaria Schreb., Linn. Gen. Pl., éd. 8 : 768, 1791.
Espéce-type. — Lobaria pulmonaria (L.) Hoffm
LOBARIA PULMONARIA (L.) Hoffm.
Deutschl. Fl. 2 : 146, 1796.
Basionyme. — Lichen pulmonarius L., Sp. Plant. : 1145, 1753
Type. — Localité inconnue, herb. C. Linnaeus (LINN 1273. 103b, lectotype :
voir YOSHIMURA & HAWKSWORTH 1970).
— Lobaria africana Dodge (fide YOSHIMURA 1971 : 286).
ZAIRE : District afro-montagnard : Wimbi (26 km S de Lubero), 2200 m,
Source : MNHN. Paris
466 E.SÉRUSIAUX
horizon supérieur de la forêt de montagne, sur un gros Conopharyngia au-dessus
d'un ruisseau, Louis 4694 (BR, LG). — Luemba, 2100 m, galerie forestiére, sur
bambous, Kinet 1537 (BR, LG) (cité par DES ABBAYES 1958 : 5, sub Г.р. var.
meridionalis (Vain.) Zahlbr.).
RWANDA : District afro-montagnard : Forét de Rugege, versant sud du mont
Muzimu (partie N de la forêt), 2720 m, forêt basse humide, branches et petits
troncs moussus, Lambinon 72/946. — Gikungu (env. 30km N de Rutsiro),
versant gauche de la vallée de la Bikeneko, env. 2150 m, forêt de montagne +
secondarisée, sur la liane Schefflera myriantha, vers 10 m de haut, Lambinon
74/477. — Vallée de la Sebeya, à env. 4 km au sud du poste minier de Gikungu,
env. 2050 m, forêt humide de fond de vallée, à épiphytisme bryophytique dense,
sur liane (Schefflera) sur la lisière, Lambinon 74/687. — Vallée de la Bikeneko,
à PW du poste minier de Gikungu, env. 2100 m, grosse branche d'un arbre mort,
dans une clairière au bord de la rivière, Lambinon 74/694. — Gisovu, Centre
forestier suisse, 2200 m, forét de montagne, sur Prunus africana, Troupin 14451.
Matériel africain complémentaire :
TANZANIE : Kilimandjaro-Sud, 2800 m, Gürtelwald, Flechte an Bäumen,
gr. Gruppen, Schlieben 4949 (BR, LG).
LA RÉUNION : Cirque de Cilaos, Sentier du Piton des Neiges, 1450 m, forêt
hygrophile des bois de couleur, épiphyte, J.L. De Sloover 17550 (NAM, LG). —
Ibid., env. 2400 m, formation basse d'éricacées et de composées, entre les blocs
rocheux, sur arbrisseau, J.L. De Sloover 17940 (NAM, LG).
Commentaires. — 1. Espèce répandue dans toutes les zones tempérées de lhé-
misphére nord, Lobaria pulmonaria n'est, dans les régions intertropicales, connu
que des montagnes d'Afrique orientale et méridionale et d'Amérique centrale
(cartes dans YOSHIMURA 1971 : 240 et 250). Dans notre dition, ce lichen
est un épiphyte croissant dans les stations les plus humides de la forét de mon-
tagne entre 2000 et 2700 m; il y est d'ailleurs rare et n'est qu'exceptionnelle-
ment pourvu d'apothécies (seule la récolte 74/477 est fertile, d'ailleurs très peu).
2. Depuis DEGELIUS (1941 : 17), on a souvent identifié à la var. meridio-
nalis (Vain.) Zahlbr. les specimens non sorédiés mais presque uniquement isidiés
de cette espèce, provenant surtout des régions tempérées chaudes de l'Europe
et de Macaronésie. YOSHIMURA (1969 : 68-73) a cependant montré que L.
meridionalis Vain. était une espéce tout à fait différente, du Sud-Est asiatique.
Par ailleurs, il existe, en tout cas dans le matériel africain que nous avons exami-
né, un continuum complet entre les échantillons à sorédies + ales et les
spécimens uniquement isidiés. Certains fragments de thalles montrent d'ailleurs
à la fois des sorédies à реше isidiféres et des isidies s. str П nous paraît donc
inopportun d'accorder une valeur taxonomique à cette variation.
LOBARIA RETIGERA (Bory) Trev.
Lichenotheca Veneta : 75, 1869.
Basionyme. — Lichen retiger Bory de St Vincent, Voyage dans les Quatre
Principales Iles des Mers d'Afrique 1 : 392 et 3 : 101, 1804.
Source : ММНМ Paris
LICHENS DU KIVU, DU RWANDA ET DU BURUNDI 467
Type. — La Réunion, Richard s.n. (lectotype, М; voir YOSHIMURA 1971 :
299-300).
= Lobaria natalensis Räs. (fide YOSHIMURA 1971 : 299).
ZAÏRE : District afro-montagnard : Massif du Kahuzi, env. du km 41 de la
route Bukavu-Walikale, 2300 m, forêt de montagne mêlée de bambous, sur ver-
sant escarpé, branches maîtresses couvertes de mousses, fougères et orchidées
d'un arbre tombé, Lambinon 71/1180 (stérile). — Ibid., lieu-dit Mukaba (km 49
de la route Bukavu-Walikale), 2250 m, tronc ombragé d’Eucalyptus en bordure
d’une plantation, Lambinon 72/66 (stérile). — Mont Biega, piste du versant
sud, 2520 m, branche horizontale dans la forêt mélangée de bambous, Lambinon
72/92 (fertile).
RWANDA : District afro-montagnard ; Chaîne des Birunga, versant sud du
Sabyinyo, 2250m, forét de montagne mélée de bambous, grosse branche
morte, Lambinon 72/450 (stérile). — Forét de Rugege, km 67 de la route Butare-
Cyangugu, 2350 m, talus terreux frais très raide à proximité de la route, Lambi-
non 72/1059 (stérile). — Ibid., vers le km 91 de la route Butare-Cyangugu (entre
Pindura et Uwinka), env. 2400 m, talus (schiste métamorphisé «pourri» ) frais,
en lisière de la forêt de montagne + secondarisée, Lambinon 74/777 (stérile). —
Ibid. vers le km 105 de la route Butare-Cyangugu, env. 1950 m, talus frais
moussu (schiste métamorphisé «pourri») en lisière de la forêt de montagne,
Lambinon 74/826 (stérile). — Ibid., entre Garamba et la plantation de Gisa-
kura, env. 1900 m, talus embroussaillé, piste en bordure de fragments de forêt
de montagne, Lambinon 74/1049 (stérile). — Ibid., collines entre le mont
Muzimu et le Bigugu, 2380 m, branches de la cime d'Apodytes dimidiata dans la
forêt de montagne, Lambinon 72/979 (stérile). — Ibid., lieu-dit Ruwankuba,
1950 m, forêt de vallée à Syzygium rowlandii, sur les lianes et les rameaux
ligneux couverts de bryophytes dans le sous-bois, à env. 1-2 m de haut, Lambi-
non 71/1066 (fertile), — Forét de Nyungwe, km 5 de la piste Pindura-Bweyeye,
env. 2050 m, branches de Cassipourea, vers 11 m de haut, dans la forêt de mon-
tagne de versant, Lambinon 74/864 (stérile). — Ibid., km 22 de la piste Pindura-
Bweyeye, env. 1750 m, branche d’un arbre mort (Bersama ?) dans la forêt de
montagne prés de la rivière Shabwa, Lambinon 74/996 (stérile). — Env. de la
commune de Rugera (SE de la préf. de Gikongoro), colline Uwintaschya, env.
2400 m, dans le bas d'un tronc de Macaranga dans la forêt de bambous, Lambi-
non 74/1122 (stérile) et 74/1123 (fertile). — Ibid., vallon Uwagahunga, env.
2350 m, sur tronc de Rapanea, fourté au bord du ruisseau, Lambinon 74/1165
(stérile). — Gikungu (env. 30 km N de Rutsiro), env. 2200 m, talus argileux
raide, en bord de piste, dans la forêt de montagne secondarisée, Lambinon
74/432 (stérile). — Ibid., versant gauche de la vallée de la Bikeneko, env. 2150 m,
forêt de montagne + secondarisée, sur branches de Conopliaryngia, vers 6 m de
haut, Lambinon 74/464 (stérile). — Vallée de la Sebeya, à env. 3 km au sud
du poste minier de Gikungu, env. 2050 m, grosse branche de Nuxia floribunda
vers 10 m de haut, Lambinon 74/677 (stérile). — Ibid., à env. 4 km au S du
poste minier de Gikungu, env. 2050 m, forêt humide de fond de vallée à épiphy-
tisme bryophytique dense, sur liane (Schefflera) dans le sous-bois sombre,
Source : MNHN, Paris
468 Е. SERUSIAUX
Lambinon 74/688 (stérile). — Ibid., à PW du poste minier de Gikungu, env.
2100 m, forêt de bas de versant à sous-bois dense d'Acanthaceae buissonnantes,
sur le sol riche en débris végétaux, Lambinon 74/708bis (stérile). — Forêt de
Gishwati, au km 39 de la route Gisenyi-Kibuye, 2150 m, talus gréso-terreux
frais et raide de la route, Lambinon 72/515 (stérile).
BURUNDI : District afro-montagnard : Forêt de Bururi, env. 2100 m, forêt
de montagne à Entandrophragma sur sol subhorizontal, branches tombées,
Lambinon 74/1299 (stérile). — Massif du Mont Teza, crête au-dessus de la plan-
tation de thé, env. 2500 m, forêt de montagne + secondarisée, dans le bas d'un
tronc de Macaranga, Lambinon 74/1407 (stérile). — Ibid. Bugarama, env,
2100 m, forêt de montagne, sur tronc de Macaranga, Petit 2407 (stérile) (BR,
LG).
Matériel africain complémentaire :
ZIMBABWE (ex Rhodésie) : Mts Vumba, Leopard Rock, 1600 m, rochers
éclairés dans la forét, Bamps, Symoens et Vanden Berghen s.n. (stérile) (LG). —
Inyanga, Mimunzi, vers 1900 m, épiphytes dans la forét, Bamps, Symoens et
Vanden Berghen s.n. (stérile) (deux collections, LG). — Inyanga, Inyangani,
vers 2070 m, sur Leucosidea, Bamps, Symoens et Vanden Berghen 460d (stérile)
(LG).
LA RÉUNION :Cirque de Cilaos, Grand Matarum, 1500 m, sur tronc, J.L.
De Sloover 17452 (fertile) (NAM, LG). — Ibid., Sentier du Piton des Neiges,
env. 1950 m, formation à Philippia et composées, sur blocs rocheux, J.L De
Sloover 17659 (stérile) (NAM, LG). — Près du sommet de la Grande Montée,
env. 1550 m, forét des bois de couleur, sur gros troncs, J.L. De Sloover 17334
(stérile) (NAM, LG).
Commentaires. — Espèce caractéristique de la forêt de montagne entre 1900
et 2500 m d'altitude, où, à l'état épiphyte ou terricole, elle occupe des stations
humides et ombragées. Elle est nettement plus abondante que la précédente
et est occasionnellement fertile. Mises à part quelques localités signalées sur les
côtes du nord-ouest de l'Amérique du Nord, c'est une espèce paléotropicale,
limitée en Afrique aux montagnes orientales et méridionales (carte dans YOSHI-
MURA 1971 : 254).
LOBARIA SUBLAEVIS (Nyl.) Yoshim.
J. Hattori Bot. Lab. 34 : 315, 1971.
Basionyme. — Ricasolia sublaevis Nyl., in KREMPELH., Flora 51 : 231, 1868.
Type. - Madère, С. Mandon 30 (lectotype, H-Nyl. 33386).
ZAÏRE : District du Graben Occidental : Angi (7 km W de Rutshuru), plaine
de lave, sur arbuste, Bequaert 5734 (BR, LG).
Commentaires. — Le spécimen examiné correspond parfaitement à la descrip-
tion de YOSHIMURA (1971 : 316-317) si ce n'est que le tomentum de la face
inférieure est souvent bien développé. Signalée d'Extréme-Orient, oà elle est
abondante, de La Réunion et de Madére, cette espéce semble nouvelle pour
Source - MNHN. Paris
LICHENS DU KIVU, DU RWANDA ET DU BURUNDI 469
l'Afrique continentale. Elle n'est connue que d'une seule localité dans la plaine
de la Rutshuru.
REMERCIEMENTS. — Nous tenons à remercier les Professeurs J.L. De Sloover et С.
Vanden Berghen qui ont mis leurs collections lichénologiques de La Réunion et du Zim-
babwe à notre disposition, de même que le curateur de lherbier du Jardin Botanique
National de Belgique (BR) qui nous laisse libre accés aux riches collections préservées dans
cette institution. Le Dr I. Yoshimura a examiné bon nombre des Lobaria cités ici et le Dr. L.
Arvidsson a revu plusieurs Coccocarpia : qu'ils en soient tous deux vivement remerciés.
Enfin, toute notre gratitude va au Prof. J. Lambinon qui nous a confié l'étude de l'abondant
matériel qu'il a collecté dans la dition et a relu et amélioré notre manuscrit.
BIBLIOGRAPHIE
ARVIDSSON L. & GALLOWAY D.J., 1979 — The lichen genus Coccocarpia in New Zea-
land. Bot. Not. 132 : 239-246.
DEGELIUS G., 1941 — Lichens from the Azores, mainly collected by Dr. Н. Persson.
Kongl. Gôtheborska Wetensk. Samhällets Handl., Wetensk. Afd. ser. B., 1 : 46 pp.
DES ABBAYES H., 1958 — Lichenes. Exploration hydrobiologique du lac Tankanika
(1946-1947). Résultats scientifiques 4(4), 14 pp. Inst. R. Sci. Nat. Belg., Bruxelles.
SANTESSON В.., 1952 — Foliicolous lichens I. Symb. Bot. Upsal. 12 (1) : 1-590.
SWINSCOW T.D.V. & KROG H., 1976 — The genus Coccocarpia in East Africa. Norweg. J.
Bot. 23 :251-259.
YOSHIMURA I., 1969 — Lichenological Notes 2-6. J. Hattori Bot. Lab. 32 :67-78.
YOSHIMURA I., 1971 — The Genus Lobaria of Eastern Asia. J. Hattori Bot. Lab. 34 :231-
364.
YOSHIMURA I. & HAWKSWORTH D.L., 1970 — The typification and chemical substances
of Lobaria pulmonaria (L.) Hoffm. J. Jap. Bot. 45 :33-41.
Source : MNHN. Paris
ZIMBABWE te À
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Man den 30 race
471
NOTE
OCTOBLEPHARUM ERECTIFOLIUM MITT. ex WILLIAMS
ЕТ STEEREOBRYON SUBULIROSTR UM (SCHIMP. ex BESCH.)
G.L. SMITH EN GUADELOUPE
J.L. DE SLOOVER*
1. Octoblepharum erectifolium Mitt. ex Williams
a) Guadeloupe, Basse-Terre, chemin bordant le Grand Étang, 400 m, forét
dense avec Gommiers et Bambous, épiphyte sur gros tronc, 15 avril 1981, J.L.
De Sloover 33884 (NAM, PC).
b) Suivant DE FOUCAULT (1977 : 23), deux espèces d'Octoblepharum
sont connues en Guadeloupe, à savoir O. albidum Hedw. et O. pulvinatum
(Dozy & Molk.) Mitt., toutes deux assez communes dans leur étage respectif,
Suivant FLORSCHUTZ (1964 : 107), Octoblepharum erectifolium est connu
de la Jamaique, de Trinidad, de la Guyane anglaise et du Surinam. Sa présence
en Guadeloupe n'a donc rien d'étonnant.
Octoblepharum erectifolium, suivant le même auteur, est caractérisé par ses
feuilles très longues (jusque 2 cm), très fragiles, avec une section transversale
à mi-hauteur aussi épaisse que large (semi-circulaire ou triangulaire équilatérale
à angles arrondis). La récolte de la Guadeloupe correspond fort bien à la des-
cription et à l'illustration que donne FLORSCHÜTZ.
2. Steereobryon subulirostrum (Schimp. ex Besch.) G.L. Smith
a) Guadeloupe, Basse-Terre, La Soufrière, chemin des Dames (piste jaune),
sur le sol, 10 avril 1981, J.L. De Sloover 33833 et 33838 (NAM, PC).
* Laboratoire de Botanique, Facultés Universitaires de Namur, Belgique.
Cryptogamie, Bryol. Lichénol., 1981, 2, 4 : 471-472.
Source : MNHN. Paris
472 J.L. DE SLOOVER
b) Le genre Steereobryon est nouveau pour la Guadeloupe, où on ne connais-
sait, de la famille des Polytrichaceae, que Pogonatum tortile (Sw.) Brid., espéce
trés commune et citée sous de nombreux synonymes (DE FOUCAULT 1977 :
18).
Atrichum subulirostrum Schimp. ex Besch. et A. portoricense Crum & Steere
ont été mis en synonymie par SMITH, qui а créé pour cette espèce le genre
Steereobryon (1971 : 56). NYHOLM (1971) gardait encore les deux espèces
séparées. Les deux récoltes citées ici, dont la seconde est fertile, confirment
la mise en synonymie proposée par SMITH. On trouvera des descriptions et des
illustrations de cette espéce dans CRUM & STEERE (1957), FRYE (1948),
NYHOLM (1971) et SMITH (1971).
Steereobryon subulirostrum était connu du Mexique, de la Jamaïque, de
Haïti, Porto Rico, le Venezuela et la Colombie (SMITH 1971 : 56). La Guade-
loupe s'inscrit naturellement dans cette répartition.
TRAVAUX CITÉS
CRUM H.A. & STEERE W.C., 1957 — The mosses of Porto Rico and the Virgin islands.
Sci. Surv. Porto Rico Virgin isl., УП, 4 : 393-599.
DE FOUCAULT B., 1957 — Flore des bryophytes de Guadeloupe, Off. Natl. Forêts Basse-
Terre, 111 pp.
FLORSCHÜTZ P.A., 1964 — Flora of Suriname, VI, 1, Musci, 271 РР.
FRYE T.C., 1948 — Atrichum subulirostrum. Bryologist 51 : 186-188.
NYLHOLM E., 1971 — Studies in the genus Atrichum P. Beauv. A short survey of the
genus and the species. Lindbergia 1 : 1-33.
SMITH G.L., 1971 — A conspectus of the genera of Polytrichaceae. Mem. New York
Bot. Gard. 21 (3), 83 pp.
Source : MNHN. Paris
473
INFORMATIONS
VIENT DE PARAÎTRE
CROSBY М.В. and MAGILL В.Е. — A dictionary of mosses. Third printing.
St Louis, Missouri Botanical Garden. 1981, 43 р. ($ 4.00, auprès du Department
Eleven, Missouri Botanical Garden, P.O. Box 299, St Louis, Mo 63166, USA).
GANGULEE Н.С. — Mosses of Eastern India and adjacent regions — a mono-
graph. Calcutta, Н.С. Gangulee. 3 vol. 1969-1980, xvii + (liii) + 2145 p., 1061
fig., 597 cartes (Bot., Univ. Calcutta, Calcutta 700019 India).
Cet ouvrage comprend 8 fascicules dont voici les dates de parution :
fasc. 1, 28 juin 1969, i-xiii, (iiv), 1-170
fasc. 2, 28 juin 1971, xiv, (v-xiii), 171-566
fasc. 3, 6 oct. 1972, xv, (xiv-xix), 567-830
fasc. 4, 1 nov. 1974, xvi, (xx-xxvi), 831-1134
fasc. 5, 1 nov. 1976, (xxvii-xxxv), 1135-1462
fasc. 6, 30 sept. 1977, (xxxvi-xxxviii), 1463-1546
fasc. 7, 31 déc. 1978, xvii, (xxxix-xliv), 1547-1752
fasc. 8, 16 oct. 1980, (xlv-liii), 1753-2145
MUSCILLANEA — publication du «Werkgroep Bryologie» devant paraître
2 fois par an (en hollandais, avec rés. angl.). Ce groupe de travail existe depuis
1978; il a pour objet l'étude de la bryoflore, spécialement de la partie nord de la
Belgique, et la diffusion de la connaissance des mousses et hépatiques belges.
(ВЕ 100 par an, auprès de : Willy Van Rompu, Bosstraat 86, B-9180 Belsele).
POELT J. und VEZDA А. — Bestimmungschlüssel europäischer Flechten.
Ergänzungsheft IL. Bibliotheca lichenologica 1981, 16 : 1-390 (CRAMER ]., in
den Springackern 2, D-3300 Braunschweig; Prix de souscription : 64 DM, prix
ordinaire : 80 DM).
ANNONCE DE DÉCÈS
Hans HORMANN (7 mai 1902-20 mai 1981) — courte note in Phytologia
1981, 49, 1 : 51-52, 1 photo, par Degener О. ynd I., und Smith D.R.
Monte G. MANUEL est subitement décédé le 8 juin 1981, à Nairobi, Kenya.
Source : MNHN, Paris
474
BIBLIOGRAPHIE BRYOLOGIQUE
D. LAMY*
SYSTÉMATIQUE, NOMENCLATURE
BIZOT M. and POCS T. — East African Bryophytes, Ш. Acta Bot. Acad. Sci.
Hung. 1979, 25, 3-4 : 223-261.
Liste de 330 esp. récoltées par Pocs T. en Tanzanie et par d’autres coll. en
Afrique du S В. Nombreuses esp. sont nouv. pour la région, 6 pour l'Afrique.
Proposition de 2 nom. nouv. : Frullania vandenberghenii Pocs pour Е. epiphylla
Vanden Berghen, et Lejeunea tuberculiflora E.W. Jones ex Pocs pour Eulejeunea
camerunensis Steph. Comb. nouv. : Lophocolea muhavurensis (S. Arn.) S. Arn.
ex Pocs (— Chiloscyphus m.), Campylopus metzlerelloides (P. Varde et Thér.)
Biz. (= Bryohumbertia m.), Fissidens diaphanodonta (P. Varde) Biz. (= Moenke-
meyera d.), F. enervis ssp. hedbergii (P. Varde) Biz. (= F.h.), Rhodobryum pers-
pinidens (Broth.) Pocs (= Bryum p.) et Rh. spathulatum (Hornsch.) Pocs (=
Mnium s.). Noter les remarques taxonom. et les nouv. synon.
DÉGUCHI H. — Keys to the species of Grimmia, Schistidium and Coscinodon
in Japan. Proc. Bryol. Soc. Japan 1979, 2, 8 : 107-110, en japonais (Dept.
Biol., Fac. Sci., Univ. Kochi, 2 chome, Akebono-cho, 780 Japan).
DEGUCHI Н. — Note sur quatre espèces himalayennes de la famille des Grim-
miacées (Musci). Hikobia 1980, 8, 3-4 : 259-268, 4 fig. (Idem).
Notes morphol. et taxon. concernant Grimmia khasiana Mitt. (syn. nouv. : G.
dimorphula C. Müll.), Racomitrium strictifolium (Mitt.) Jaeg. (syn. avec С. atra-
ta Mielich.), Racomitrium fuscescens Wils. et В. himalayanum (Mitt.) Jaeg.
HASEGAWA ]. — Taxonomic studies on asian Anthocerotae. Il, Some asian
species of Dendroceros. J. Hattori Bot. Lab. 1980, 47 : 287-309, 11 fig.
(Lab. Applied Bot., Fac. Agric., Kyoto Univ., Kyoto, 606 Japan).
Clé, descr., ill. des esp. de Dendroceros représentées au Japon : D. japonicus
Steph. et D. tubercularis Hatt. Notes à propos des esp. asiatiques à nervure
pleine : descr., ill. de D. subplanus Steph. (syn. nouv. : D. angustus Steph. et
D. integerrimus Steph.), D. foliicola sp. nov. de Bornéo aff. de la précédente,
D. acutilobus Steph. (syn. пошу. : D. karstenii Schiffn. ex Steph.), D. validus
* Lab, Cryptogamie, 12 rue Buffon, F-75005 Paris.
Cryptogamie, Bryol. Lichénol., 1981, 2, 4 :474-489.
Source : MNHN, Paris
BIBLIOGRAPHIE BRYOLOGIQUE 475
Steph. et D. javanicus (Nees) Nees (nouv. descr., syn. nouv, : D. elegans Steph.).
Notes à propos des esp. asiatiques à nervure non pleine : descr., ill. de D. diffici-
lis Steph. (syn. nouv. : D. cucullatus Steph.), D. pedunculatus Steph., D. caver-
nosus sp. nov. de Bornéo.
HATTORI S. — Dr. Н. Inoue’s Frullania collection made in Formosa. Bull. Natl.
Sci. Mus. Ser. В (Bot.), 1980, 6, 1 : 33-40, 3 fig. (Hatt. Bot. Lab., Nichinan-
shi, Miyazaki-ken, 889-25 Japan).
Liste de 12 esp. avec les loc. Diagn., descr., Ш. de 3 esp. nouv. : Е. (Trachy-
colea) caduca, Е. (Trach.) hiroshii, E. (Trach.) inouei.
HATTORI S. — Notes on Frullania species of Iriomote and Ishigaki Islands, the
Ryukyu Archipelago. J. Jap. Bot. 1980, 55, 5 : 132-135, 1 fig. (Idem).
Liste de 5 taxa avec loc. dont Fr. (Diastaloba) iriomotensis sp. nov. (diagn.,
descr., ill).
HATTORI S. — Notes on the Asiatic species of the genus Frullania, Hepaticae.
XII. J. Hattori Bot. Lab. 1980, 47 : 85-125, fig. 219-240 (Idem).
106. — Diagn., descr., ill. de Е. (Frull.) vittata fo. denticulata, Е. (Trachyc.)
piptophylla et Е. (Trachyc.) piptophylloides, taxons nouv. de Nouvelle-Guinée.
— 107. — Diagn., descr., ill. de F. (Frull.) antaresensis sp. nov. de Nouv.-Guinée.
— 108. — Diagn., descr., Ш. de F. (Trachyc.) subpedicellata, de F. (Trachyc.)
rhytidantha, Е. (Trachyc.) sphaerantha et Е. (Trach.) pseudoschensiana, esp.
nouv. de l'Inde. — 109. — Е. acutiloba Mitt. est considéré comme esp. distincte
de Е. hampeana Nees. Cette dernière doit céder la priorité à Е. monocera (Tayl.)
Tayl. — 110. — Е. (Fusiorillegerae) gaudichaudii var. ceylanica (Nees) c.n. (=
Е. ceyl.). — 111. — F. bonincola Hatt. doit être exclu de la flore de Formose et
remplacé par Е. monocera. — 112. — Lectotypification de Е. (Frull.) apiculata
(Reinw. et al.) Dum. — 113. — Е. armitana var. longe-attenuata (Hatt.) c.n.
(= F. longe-attenuata). — 114. — Descr, de F. polyptera Tayl. récolté à Ceylan.
— 115. — Descr, ill., distr. de F. (Trachyc.) subclavata Steph. — 116. — Diagn.,
descr., ill. де F. (Trachyc.) carrii var. subcarrii Hatt. var. nov. de Nouv.-Guinée.
— 117. — Descr., ill. de Е. (Frull.) serrata subsp. van-zantenii (Kamim. et Hatt.)
cn. (= Е. van-zant.). — 118. — Descr., ill. de F. (Frull.) madens Steph. de Nouv.-
Guinée. — 119. — Descr., ill. de Е, (Trachyc.) sinensis Steph. nouv. pour l'Inde.
— 120. — Е. fusco-virens Steph. doit être considéré comme une var. de Е. valida
Steph. — 121. — F. hamatiloba Steph. pourrait étre conspécifique avec F.
(Trach.) koreana Steph. (descr.). — 122. — Descr. de Е. (Diastal.) hypoleuca
Nees. — 123. — Propagules de F. gemmulosa Hatt. et Thaith. — 124. — Descr.,
Ш. de Е. (Trachyc.) brittoniae subsp. truncatifolia (Steph.) Schust. et Hatt. nouv.
pour la Chine.
KITAGAWA М. — Studies on Asian species of Bazzania, Hepaticae, IL. Bull.
Nara Univ. Educ. 1979, 28, 2 : 71-83, 6 fig. (Biol. Lab., Nara Univ. Educ.,
Nara, 630 Japan).
Descr, ill de Bazzania palmatifida (Steph.) Grolle, B. parvitexta Steph.,
Source - MNHN. Paris
476 BIBLIOGRAPHIE BRYOLOGIQUE
В. calcarata (Sde Lac.) Schiffn., B. stresemannii (Herz.) c.n. (= Mastigobryum
s.), B. asymmetrica (Steph.) c.n. (= Mast. a.), B. subaequitexta (Steph.) c.n. (=
Mast. s.) de Nouvelle-Guinée.
KUMAR S.S. — Taxonomic studies in West Himalayan mosses. 1. Hikobia 1980,
8, 3-4 : 245-258, 6 fig. (Dept. Bot., Panjab Univ., Chandigarh, 160014 India).
Clé, descr., ill. de 6 Encalypta : E. streptocarpa Hedw., E. rhabdocarpa
Schwaegr., E. tibetana Mitt., E. ciliata Hedw., E. vulgaris Hedw. et E. alpina Sm.
KUWAHARA У. — Metzgeria maegdefraui, spec. nov. from the Neotropics.
Hikobia 1980, 8, 34 : 269-273, 1 fig. (10-2139, Mii-machi, Kurume, Fu-
kuoka, 830 Japan).
Diagn., descr., ill. de M. maegdefraui esp. nouv. de Colombie. Comparaison
avec M. mexicana Steph. et M. herminieri Schiffn.
KUWAHARA Y. — Four new species of the Metzgeriaceae (Hepaticae) from the
subantarctic region with enumeration of previously reported taxa of the
family. Hikobia 1980, 8, 3-4 : 274-296, 45 fig. (Idem).
Diagn., descr., ill. de M. engelii esp. nouv. des Iles Falkland, M. sparrei esp.
nouv. du Chili, M. vittii esp. nouv. de Nouvelle-Zélande et M. monoica esp.
nouv. du Chili. Comparaison avec esp. affines. Énumération avec loc. de 49 esp.
et 4 var. de la famille des Metzgeriaceae représentées dans la région subantarc-
tique incluant la Nouvelle-Zélande, la Tasmanie et le Sud de l'Amérique du Sud.
NOGUCHI A. — Notulae bryologicae, X. J. Hattori Bot. Lab. 1980, 47 : 311-
317, 4 fig. (Hatt. Bot. Lab., Nichinan-shi, Miyazaki-ken 889-25 Japan).
1. Forsstroemia tripinnata (Dix.) c.n. (= Porotrichum t.). — 2. — Garovaglia
conchophylla Ren. et Card., Pterobryopsis handelii Broth. et P. morrisonicola
Nog. sont syn. avec Pier. acuminata (Hook.) Fleisch. — 3. — Calyptothecium
auriculatum (Dix.) c.n. (= Рег. aur.). — 4. — Calypt. integrifolium (Bartr.) c.n.
(= Symphysodon i.). — 5. — Descr. du sporophyte de Jaegerina luzonensis
Broth.
TAN B.C. and SCHOFIELD W.B.— On Dichodontium pellucidum and D.
olympicum. Canad. J. Bot. 1980, 58, 19 :2067-2072, 1 carte, 16 fig., 1 tabl.
(Dept. Bot., Univ. British Columbia, Vancouver, B.C., Canada V6T 1W5).
Comparaison morphol. et anatom. entre D. pellucidum (Hedw.) Dix. et D.
olympicum Ren. et Card. Caract. bien spécif. : détails cellulaires de la surf.
ventrale de la nervure de la feuille, cellules de l'exothéque, la sexualité. Les
Oreoweisia serrulata (Funck) De Not. du Canada sont en fait des D. pellucidum.
D. olympicum est endémique de l’ouest de l'Amérique du Nord.
VANDER GRONDE K. — Studies on Colombian Cryptogams VIII, The genus
Jensenia Lindb. (Hepaticae). Proc. Kon. Ned. Akad. Wetensch., Ser. C Biol.
Med. Sci. 1980, 83, 3 : 271-278, 1 fig. (Inst. Syst. Plantkunde, Heidelberglaan
2, Utrecht, Netherlands).
Source : MNHN, Paris
BIBLIOGRAPHIE BRYOLOGIQUE 477
Descr., (diagn.), distr., écol. de Jensenia florschuetzii sp. nov., J. erythropus
(Gott.) Grolle, J. e. var. erythropus et J.e. var. nobandae var. nov.
MORPHOLOGIE, ANATOMIE
DEGUCHI H. — Foot of Hedwigia ciliata. Proc. Bryol. Soc. Japan 1980, 2, 9 :
126, en japonais.
KAWAI I. — Anatomical characteristics of stems in some species of Dicranaceae.
Proc. Bryol. Soc. Japan 1980, 2, 9 : 126, en japonais.
SATO S. and YAMADA М. — Scanning electron microscopy on the antheridium
of Conocephalum conicum. J. Hattori Bot. Lab. 1980, 47 : 333-344, 15 fig.
(Dept. Biol. & Lab. Electr. Microsc., Japan Women's Univ., Mejirodao,
Bunkyo-ku, Tokyo, Japan).
Mise en évidence à la surface des réceptacles d, de pores anthéridiaux et de
pores de ventilation à structure différente. Observation de la spermatogenése au
microsc. à balayage. Présence de cristaux contenant du calcium dans les cellules
antheridiales.
PHYSIOLOGIE, CHIMIE
HUAULT Cl. — Contrôle par le phytochrome de la germination des propagules
de Lunularia cruciata L. Compt.-Rend. Hebd. Séances Acad. Sci., Ser. D
1980, 291, 2 : 307-310, 6 fig., tabl. (Centres Rech. Biol. & Physiol. Cell.,
Lab. Photobiol., Г.А. CNRS по 203, Fac. Sci., F-76130 Mont-Saint-Aignan).
Le besoin de lumière est lié à l'intervention du phytochrome et au rôle des
substances endogènes inhibitrices dans la germination des propagules de Lun.
cruciata.
KARUNEN P., MIKOLA H. and EKMAN R, — Separation and analysis of steryl
and wax esters from Dicranum elongatum. Physiol. Pl. (Copenhagen) 1980,
49, 4 1351-353, 2 tabl., 1 fig. (Dept. Bot., Univ. Turku, SF-20500 Turku 50).
Séparation complète des esters cireux et des esters de stéryle, chez le Dicra-
num elongatum subarctique, faite par chromatographie en plaque mince (MgO).
La composition en acide gras des esters cireux varie en fonction de l'áge du pied.
Par contre la distr. d'acides gras des esters de stéryle ne montre pas de différence
notable entre les différents pieds.
MATSUO A., NAKAYAMA M., HAYASHI S. and МАСА! К. — Fatty acid ethyl
esters in the liverwort Conocephalum conicum. Phytochemistry 1980, 19, 8 :
1848-1849, 1 tabl. (Dept. Chemistry, Fac. Sci. Hiroshima Univ., Hiroshima
730, Japan).
PIHAKASKI S. and PIHAKASKI K. — Effects of glyphosate on ultrastructure
Source : MNHN, Paris
478 BIBLIOGRAPHIE BRYOLOGIQUE
and photosynthesis of Pellia epiphylla. Ann. Bot. (London) 1980, 46, 2 :
133-141, 6 pl., 2 fig. (Dept. Bot., Univ. Turku, SF-20500 Turku 50).
Rapide chute de l'activité photosynthétique aprés aspersion de glyphosate;
tendance à une reprise aprés une semaine. Observation de structures vésiculaires
et tubulaires à la surface du chloroplaste, et de corps granuleux semblant étre
détachés du stroma du chloroplaste. Il y a détérioration des oléocorps, du réti-
culum endoplasmique et des ribosomes, augmentation de sphérules lipidiques
et vacuolisation du cytoplasme. Aprés 2 semaines, détérioration du grana.
SIMOLA L.K. and KOSKIMIES-SOININEN K. — The effect of fluoride on the
growth and fatty acid composition of Sphagnum fimbriatum at two tempera-
tures. Physiol. Pl. (Copenhagen) 1980, 50, 1 : 74-77, 2 tabl. (Dept. Bot.,
Univ. Helsinki, Helsinki, Finland).
THEODOR R., ZINSMEISTER H.D., MUES В. and MARKHAM К.В. — Fla-
vone C-glycosides of Apometzgeria pubescens. Phytochemistry 1980, 19, 8 :
1695-1700, 3 tabl. (Fachber. 16, Bot., Univ. Saarlandes, 6600 Saarbrücken,
West Germany).
Identification de 11 flavones di-C-glycosides dont 9 sont nouv. pour Apo-
metzgeria pubescens. Le 6,8-di-C-glucoside tricétine et le 6-C-arabinoside-8.C-
pentoside tricine sont en proportion majoritaire. Noter la présence du «feru-
lylisoschaftoside». Importance taxonomique.
CYTOLOGIE
KUMAR 5.5. and NARULA M. — Cytological studies on some West Himalayan
mosses. Hikobia 1980, 8, 34 : 355-361, 16 fig. (Dept. Bot., Panjab Univ.,
Chandigarh 160014, India).
9 taxons sont étudiés. Premier comptage pour Fissidens rambii Gang. (n =
10), Е. curvatoxiphioides Dix. et P. Vard. (п = 10), Е. curvatoinvolutus Dix.
(n = 12), F. biformis var. subbryoides Mitt. (n = 10).
KUMAR 8.5. and VERMA S.K. — Chromosome numbers in some West Hima-
layan mosses. Hikobia 1980, 8, 3-4 : 362-364, 1 tabl. (Idem).
Étude de 21 esp. de Bryaceae. Premier comptage pour 9 esp.
RÉPARTITION, ÉCOLOGIE, SOCIOLOGIE.
DEGUCHI Н. — Noteworthy mosses from the Shikoku district of Japan. Mem.
Fac. Sci. Kochi Univ. Ser. D (Biol.) 1980, 1 : 35-43, 5 fig. (Dept. Biol., Fac.
Sci., Kochi Univ., 2 chome, Akebono-cho, 780 Japan).
11 esp. récoltées dans le district de Shikoku, dont 9 sont nouv. pour cette
région. Descr., ill. de Ditrichum flexicaule (Schwaegr.) Hampe; rectification de
la descr. de Heterocladium capillaceum Broth. ex Ihs.
Source : MNHN. Paris
BIBLIOGRAPHIE BRYOLOGIQUE 479
HARTMANN Н. — Beitrag zur Kenntnis der Pflanzengesellschaften Spitzbergens.
Phytocoenologia 1980, 8, 1 : 65-147, 8 phot., 14 fig., 6 tabl. (Sonnenrain
62, CH-8700 Küsnacht).
Descr. des associations. Écosociologie de cert. esp.; mousses, lichens, phané-
rogames cités. Observ. de toundra de mousses : comparaison entre la communau-
té «sèche» à Sphagnum squarrosum et Hypnum revolutum et la communauté
«humide» à Drepanocladus revolvens et Deschampsia alpina. Noter la trouvaille
de larges tapis de Racomitrium lanuginosum.
IWATSUKI Z. — Some noteworthy mosses from South Eastern Shikoku, Japan.
Proc. Bryol. Soc. Japan 1979, 2, 8 : 118, en japonais.
KAMIMURA M., ANDO H., DEGUCHI H., HASEGAWA J., HIGUCHI M.,
IWATSUKI Z., KIGUCHI H., KITAGAWA N., NISHIMURA N., SATO Y.,
UNE K. and YOSHIMURA I. — List of bryophytes collected in the Tengu-
kogen Highland, Shikoku, Southern Japan, during the 7th Foray (1978)
organized by the Bryological Society of Japan. Bull. Kochi Gakuen Junior
Coll. 1979, 10 :25-33, en jap., rés. angl.
154 mousses et 56 hépatiques avec loc.
KANDA Н. — Distributional additions to the Japanese Amblystegiaceae and
allied family. Hikobia 1980, 8, 3-4 : 322-330, 2 fig. (Natl. Inst. Polar Res.,
Kaga, Itabashi, Tokyo, Japan).
Nouv. loc. au Japon pour Heterocladium tenellum Deg. et Suz., Amblyste-
gium serpens (Hedw.) B.S.G., Leptodictyum radicale (P. Beauv.) Kanda, Campy-
lium squarrosulum (Besch. et Card.) Kanda, Drepanocladus schulzei (Limpr.)
Roth et Pseudohygrohypnum purpurascens (Broth.) Kanda.
MARSTALLER R. — Zur Verbreitung und Soziologie einiger Moose der Trocken-
und Halbtrockenrasen im östlichen Thüringen - 3. Beitrag zur Moosvege-
tation Thüringens. Wiss. Z. Friedrich-Schiller-Univ. Jena, Math.-Naturwiss.
Reihe 1980, 29, 1 : 79-88, 6 fig, 1 tabl. (Friedrich-Schiller-Univ., Sekt.
Biol., DDR 69 Jena).
Distr., écol. et sociol. d'Abietinella abietina, Rhytidium rugosum, Camptothe-
cium lutescens, Entodon orthocarpus et Tortella inclinata dans la partie est de
la Thuringe. Descr. de l'ass. Rhytidio-Entodontetum Stodiek en Thuringe.
MARSTALLER В. — Die Bryophytengesellschaften der Jenaer Umgebung —
eine Uebersicht. 4. Beitrag zur Moosvegetation Thüringens. Wiss. Z. Friedrich-
Schiller-Univ. Jena, Math.-Naturwiss. Reihe 1980, 29, 1 : 89-108 (Idem).
Descr. (structure, composition floristique, distr.) de 74 assoc. de bryophytes
naturelles et anthropogéniques des alentours de Jena; 3 ass. nouv. :le Pottietum
davallianae , le Dicranello-Campylopetum flexuosae et l'Hookerietum lucentis.
NISHIMURA N. and HIGUCHI M. — A new locality of Diphyscium perminutum
Tak. Hikobia 1980, 8, 3-4 : 273, en japonais.
Source : MNHN, Paris
480 BIBLIOGRAPHIE BRYOLOGIQUE
PETERSON W.L., SMITH D.K. and SHARP A.J. — The mosses of the Kodiak
Archipelago. J. Hattori Bot. Lab. 1980, 47 : 269-285, 1 fig., 1 tabl. (Dept.
Bot., Univ. Tennessee, Knoxville, Tenn. 37916 USA).
Historique de la découverte et de l'exploration de l'archipel. Liste avec loc.
de 240 taxons dont 32 sont nouv. pour l'ile Kodiak (parmi eux 26 sont nouv.
pour l’Archipel).
SEKI T. and MIYAGI C. — An ecological observation on Gymnostomiella longi-
nervis Broth. in Okinawa Island. Hikobia 1980, 8, 34 : 416-423, 3 fig.,
1 tabl., en jap., rés. angl. (The Miyajima Nat. Bot. Gard., Hiroshima Univ.,
Hiroshima, Japan).
SKOLOWSKI A.W. — Zbiorowiska Lesna Polnocno — Wschodniej Polski (Forest
communities of North-Eastern Poland). Monogr. Bot. 1980, 60 : 1-205,
18 fig., 59 tabl.
Étude écologique du territoire de Bialystok. Descr. de 25 syntaxons de rang
d'association représentant 4 classes. Noter 6 nouv. ass. dont le Sphagno girgen-
sohnii-Piceetum. 8 ass. ont une affinité boréale. Mousses et lichens cités.
TAODA Н. — Studies on the Fabroniaceae of Japan. II. Hikobia 1980, 8, 3-4 :
298-321, 12 fig. (Asakawa Herb., Forestry and Forest Prod. Res. Inst., Ha-
chioji, Tokyo 193, Japan).
Clés, descr., loc., Ш. de 3 esp. et 1 var. d’Anacamptodon Brid., 2 esp. de
Fabronia Raddi, 1 esp. d’Iwatsukiella Buck et Cram, 2 esp. de Schwetschkeopsis
Broth.
UETA Н. and DEGUCHI Н. — Mosses for gardening in the Нага district, Kochi
Prefecture. Proc. Bryol. Soc. Japan 1980, 2, 9 : 133-135, fig., en jap., rés. angl.
WILLEMS J.H. — Observations on north-west European limestone grassland
communities. V, a and b. Proc. Kon. Ned. Akad. Wetensch., Ser. C Biol. Med.
Sci. 1980, 83, 3 : 279-306, 7 tabl., 5 fig. (Dept. Pl. Ecol. & Veg. Sci., State
Univ., Utrecht, The Netherlands).
Approche expérimentale de l'étude de la diversité des esp. et de la biomasse
des prairies calcicoles. Róle des bryophytes.
YUZAWA У. — Some species of Lejeuneaceae from Fukushima Prefecture,
Japan. Proc. Bryol. Soc. Japan 1979, 2, 8 : 105-107, en jap. (13 esp. avec loc.).
OUVRAGES GÉNÉRAUX
GANGULEE Н.С. — Mosses of Eastern India and adjacent regions - a mono-
graph. Calcutta, H.C. Gangulee, 3 vol., 1969-1980, xviii, (liii), 2145 p., 1061
fig., 597 cartes (Bot., Univ. Calcutta, Calcutta 700019, India).
L'aire étudiée représente la zone politique orientale de l'Inde, c’est-à-dire :
Source : MNHN, Paris
BIBLIOGRAPHIE BRYOLOGIQUE 481
Arunachal (ex NEFA), Assam, Nagaland, Manipur, Tripura, Bangal W, Bihar et
Orissa avec les régions naturelles adjacentes : Sikkim, Bhutan, Népal et Bangla-
desh; s’y ajoutent les Iles Andaman et Nicobar et une petite partie du Madhya
Pradesh. Cette monographie, qui veut servir de base à des recherches futures,
recouvre 990 esp. réparties en 274 genres et 52 familles. Descr. des familles et
des genres. Clés aux genres et esp. Taxonomie, descr., ill., distr. régionale et
mondiale de chaque esp. Notes cytologiques (nombres chromosomiques et
investigateurs) par genres. Les données géographiques par genre, et un index
taxonomique complétent cette importante monographie, qui est parue en 8
fascicules de 1969 à 1980. Ce travail, unique pour l'Inde orientale, bien illus-
tré, se présente comme un bon outil de référence. Taxons пошу. : Atrichum
longifolium Card. et Dix. ex Gang. - 1969, Calyptothecium dixonii Gang. - 1976,
Daltonia decolyi Broth. ex Gang. - 1977, Distichophyllum decolyi Gang. -
1977, Fissidens allanii Gang. - 1971, F. rambii Gang. - 1971, F. rigidiusculus
Broth. in Bruehl ex Gang. - 1971, Isopterygium andamanicum Gang. - 1980,
Leucophanes nicobaricum С. Müll. ex Gang. - 1971, Neckeropsis darjeelin-
gensis Gang. - 1976, Orontobryum recurvulum Gang, - 1977, Penzigiella hookeri
Gang. - 1976, Pohlia ampullacea Gang. - 1974, Schoenobryum concavifolium
(Griff) Gang. - 1976 (— Orthotrichum), Splachnobryum bengalense Gang. -
1974, Thamnobryum fruticosum (Mitt.) Gang. - 1976 (= Neckera), T. macro-
carpum (Brid.) Gang. - 1976 (— Neckera), Thyridium andamense Besch. ex
Gang. - 1972, T. nicobaricum Broth. ex Gang. - 1972, Trichosteleum puncti-
papillosum Par. ex Gang. - 1980, T. stereodontoides Broth, ex Gang. - 1980. (Les
dates de parution des différents fascicules sont indiquées dans la partie «Infor-
mations» du présent fascicule).
GRIFFIN III D. — Guia preliminar para as Briófitas freqüentes em Manaus e
adjacéncias. Acta Amazonica 1979, 9, 3, suppl. : 1-67, 11 pl. (Univ. Florida,
Gainesville, Florida, USA).
Clés aux esp. de mousses et hépatiques. Descr. des familles et clés aux esp.
de certains genres avec notes descr.
VARIA
ANDO Н. — Bibliography of the use of Bryophytes. Hikobia 1980, 8, 34 :
424-448 (Bot. Inst., Hiroshima Univ., Hiroshima, Japan).
BRUNEAU D. — Regards sur l'étude des Muscinées et des Lichens en Anjou.
Mém. Soc. Études Sci. Anjou «1980» 1981, 4 : 165-169 (44 rue du Pin,
F-49000 Angers).
Historique des récoltes et herb. depuis 1800. Courtes notices des principaux
bryologues et lichénologues qui travaillérent dans la région.
GADEA E.-— La distribució de la nematofauna muscícola i liquenicola als
illots del País Valencia. Treb. Inst. Catalana Hist. Nat. 1981, 9 :69-73, 1 fig.
(Dept. Zool., Fac. Biol. Univ. Barcelona, España).
Source : MNHN, Paris
482 BIBLIOGRAPHIE BRYOLOGIQUE
GÓTZ н. — Wilhelm Philipp SCHIMPER und Karl Friedrich SCHIMPER - zwei
Naturforscher am Oberrhein. Zum 100. Todestag Wilhelm Philipp Schimpers.
Beitr. Naturk. Forsch. Südwestdeutschl. 1980, 39 : 19-35, 2 fig. (Berliner
Strasse 50, D-6830 Schwetzinger).
Liens de parenté et relations scientifiques entre М.Р. Schimper de Strasbourg
et son cousin de Bade, le naturaliste K.F. Schimper. Généalogies, extr. de lettres,
portr.
GRELON 7. A propos de Marchantia polymorpha L., hépatique adventice
nouvelle, en extension dans les pépinières angevines. Mém. Soc. Études Sci.
Anjou «1980» 1981, 4 : 161-163 (ENITH, rue Le Nótre, F-49045 Angers
Cedex).
Essai de cultures sur milieu gélosé ou différents substrats horticoles. Test de la
sensibilité de cette esp. vis-à-vis des produits phytosanitaires.
IWATSUKI 2. — Notes on various methods of cultivation of bryophytes. 2.
Proc. Bryol. Soc. Japan 1979, 2, 8 : 110-113, fig. 4-5, en jap., rés. angl.
KATOH K., ISHIKAWA M., MIYAKE K., OHTA Y., HIROSE Y. and IWAMU-
RA T. — Nutrient utilization and requirement under photoheterotrophic
growth of Marchantia polymorpha : improvement of the culture medium.
Physiol. Pl. (Copenhague) 1980, 49, 2 : 241-247, 6 fig., 6 tabl. (Inst. Food
Chemistry, Shimamoto-cho, Mishima-gun, Osaka 618, Japan).
Descr. d'un nouv. milieu de culture pour les cellules chlorophylliennes de
M.p.
NISHIMURA N., HIGUCHI M. and UNE К. — Bryophytes as materials of the
nest of a songbird Pallass Dipper (Cinclus pallasii hondoensis). Hikobia
1980, 8, 3-4 :350-354, 2 fig., 3 tabl. (Bot. Inst., Hiroshima Univ., Hiroshima,
Japan).
STOTLER В.Е. — A history of Illinois Bryology. Trans. Illinois State Acad. Sci.
197, 72, 2 :16-27.
La bryologie en Illinois : historique, notes biographiques des principaux
botanistes.
WATANABE R.— A biography of Philipp Franz Baltasar von Siebold. Proc.
Bryol. Soc. Japan 1979, 2, 8 : 113-116, 2 fig., en japonais.
Source - MNHN. Paris
483
BIBLIOGRAPHIE LICHÉNOLOGIQUE
D. LAMY
SYSTÉMATIQUE, NOMENCLATURE
АНТИ Т. — Taxonomic revision of Cladonia gracilis and its allies. Ann. Bot.
Fenn. 1980, 17 : 195-243, 35 fig. (Dept. Bot., Univ. Helsinki, Unioninkatu
44, SF-00170 Helsinki 17).
7 esp. sont reconnues appartenir au groupe Cladonia gracilis : С. macroceras
(Delise) Ahti, C. maxima (Asah.) Ahti et C. squamosissima (Müll. Arg.) Ahti c.n.
(= C. gracilis var. s.) sont confinés à l'hémisphére nord, et С. subchordalis A.W.
Evans à lhémisphére sud, tandis que C. cornuta (L.) Hoffm., С. ecmocyna
Leighton et C. gracilis (L.) Willd. ont une distribution bipolaire. C. gracilis est
divisé en 6 sous-esp. : ssp. elongata (Jacq.) Vainio, ssp. gracilis, ssp. nigripes
(Nyl.) Ahti c.n. (= C. ecmocyna f. nigr.), ssp. tenerrima ssp. nov, d'Australie,
ssp. furbinata (Ach.) cn. (= Lichen t.), et ssp. vulnerata ssp. nov. d'Alaska.
Cladonia comuta comprend 2 sous-esp. : ssp comuta et ssp. groenlandica
(E. Dahl) c.n. (= C. comuta var. g.). C. ecmocyna est divisé en ssp. ecmocyna
et ssp. intermedia (Robbins) c.n. (= C. elongata f. i.). C. propagulifera (Vainio)
Dodge est syn. avec C. gracilis ssp. gracilis. C. gracilis var. dilatata (Hoffm.)
Vainio est remplacé par C. gracilis ssp. turbinata. Noter parmi les taxa exclus :
C. campbelliana (Vainio) Gyelnik, C. isabellina Vainio et C. comuta f. subdila-
iata Asah. Présence de types morphol. intermédiaires entre les sous-esp. de C.
gracilis, comparables aux hybrides locaux chez les plantes supérieures. Clé aux
taxons. Pour chaque taxon : taxonom., descr., chimie, ill., spécim. examinés.
La plupart des taxons contiennent les acides fumarprotocétrarique et protocétra-
rique, quelques-uns de l’atranorine. Nouv. substances isolées ; ac. squamatique
chez C. subchordalis et ac. grayanique chez С. isabellina. Noter encore la récolte
de C. alinii Trass. en Alaska, nouv. pour l'Amérique du Nord.
POELT J. und VEZDA А. — Bestimmungsschlüssel europäischer Flechten.
Ergänzungsheft IL. Biblioth. Lichenol. 1981, 16 : 1-390.
Cet ouvrage, fait en collaboration avec d’autres lichénologues, fait suite à
l'Ergánzungsheft 1 paru en 1977, Il comprend : le système des Lichens (genres
européens) : ordres, familles et genres; clés aux lichens à basides, aux lichens à
algues bleues (voir POELT 1979), aux lichens arbusculés et fruticuleux, aux
lichens foliacés et squamuleux, aux lichens à algues vertes stériles, ou avec des
sorédies ou des isidies, aux lichens coniocarpes, aux lichens pyrénocarpes, aux
Source - MNHN, Paris
484 BIBLIOGRAPHIE LICHÉNOLOGIQUE
lichens lirellocarpes et aux lichens discocarpes crustacés. Révision de 64 genres
et de Parmelia subgen. Melanoparmelia. Descr. du genre, clé aux esp. (avec
descr., chimie et syn.) et bibliographie. Corrections et additions au 1° supplé-
ment. Corrections et nouveautés taxonomiques du 2? supplément; on notera :
28 comb. nouv., diagn. de Melanolecia Hertel gen. nov. (esp. type : M. transitoria
(Arnold) c.n. (= Lecidea) et de Lecanora mugosphagneti Poelt et Vézda, de
Bavière, ex aff. Lecanora pallidae. Index pour les 2 suppléments.
SINGH A. — Lichenology in Indian subcontinent 1966-1977. Lucknow, Econo-
mic Bot. Inf. Serv., Natl. Bot. Res. Inst. 1980, 112 p.
Centres de recherches et herbiers de lichens en Inde; études botaniques et
chimiques effectuées. Index des esp. avec loc.; index des esp. avec contenus
chimiques et bibliographie. Comb. nouv. : Heterodermia dactyliza f. serpens
(Vain.) (= Anaptychia obscurata var. s.), Н. dentritica var. propagulifera (Vain.)
(= Anaptychia), Hypogymnia pseudohypotrypa (Asah.) (= Parmelia), Parmelia
coorgiana (Patw. et Prabhu) (= Hypotrachyna), P. dahlii (Hale) (= Pseudoparme-
lia), P. indica (Hale) (= Parmelina), P. hamatii (Patw. et Prabhu) (= Parmotre-
ma), P. mason-halei (Patw. et Prabhu) (= Hypotrachyna), P. rhytidodes (Hale)
(= Parmelina), Phaeophyscia endococcina var. khumbuensis (Poelt) (= Physcia
е. var. k.).
MORPHOLOGIE, ANATOMIE
GIERSBERG M. — Wachstumsmessungen an Cladonia chlorophaea (Flk.) Zopf.
Arch. Freunde Naturgesch. Mecklenburg 1978, 18 : 65-67, 1 fig, 1 tabl.
(Wilhelm-Pieck-Univ. Rostock, Sekt. Biol., WB Terrestrische Oekol., DDR-25
Rostock).
LAMBRIGHT D.D. and TUCKER S.C. — Observations on the Ultrastructure of
Trypethelium eluteriae Spreng. Bryologist 1980, 83, 2 : 170-178, 12 fig.
(Dept. Bot., Louisiana State A & M Univ., Baton Rouge, LA 70603 USA).
Cortex supérieur amorphe, parois fines des hyphes de la médulle et invagina-
tions du plasmalemme suggèrent une relation saprophytique entre le myco-
bionte et les cellules du péridesme. Observ. et descr. du phycobionte (Trente-
pohlia) et du mycobionte ascomycéte.
POELT J.— Eine diózische Flechte. Pl. Syst. Evol. 1980, 135, 1-2 : 81-87,
3 fig. (Inst. Bot., Univ. Graz, Holteigasse 6, A-8010 Graz).
Descr. de Lecidea verruca poussant en parasite sur d'autres lichens crustacés,
qui, contrairement à la plupart des autres ascomycétes lichénisés, est dioique.
Les thalles d portant des spermogonies sont plus petits que les thalles 9.
RÉPARTITION, ÉCOLOGIE, SOCIOLOGIE
ANDREEV M.P. — Lichens of the genera Cladina and Cladonia from Anyui-
Source : MNHN, Paris
BIBLIOGRAPHIE LICHÉNOLOGIQUE 485
plateau. Bot. Zum. (Moscow et Leningrad) 1981, 66, 1 : 31-41, 3 fig., 3 tabl.,
en russe, rés. angl. (Bot. Inst. V.L. Komarova, AN SSSR, Leningrad).
47 esp. de Cladonia et Cladina des Mts Апуш. 12 sont попу. pour la région
de Chukotka. Comparaison avec les flores de l'holarctique М. Comparaison des
esp. d'un habitat à l'autre. Utilisation de l'index Goodal.
BACKEUS I. — Vegetation changes after fertilization on drained peatlands in
Central Sweden. Acta Phytogeogr. Suec. 1980, 68 : 17-30, 4 fig, 5 tabl.
(Inst. Ecol. Bot., Uppsala Univ., Box 559, S-751 22 Uppsala).
Étude des changements à court et long termes des tourbiéres ombrotro-
phiques : diminution de la couverture totale de Sphaignes, baisse de leur vitalité,
influence des conditions de nutrition et de la croissance des arbres. La végétation
devient plus ou moins semblable à celle des forêts humides sur sol minéral.
Bryophytes et lichens cités.
CORTINI-PEDROTI С. — La flora briologica dell'Isola di Montecristo (Arcipelago
Toscano). Webbia 1980, 34, 2 : 707-760, 1 tabl. (Ist. Bot., Univ., 50121
Firenze, Italia).
Descr. de l’île de Montecristo et de sa végétation. Liste de 33 hépatiques et
129 mousses avec données écol. Nouveautés pour l’île. Noter liste de 52 lichens.
DOUGLAS G.W. and PETERSON W.L. — Contributions to the floras of British
Columbia and the Yukon Territory. И. Mosses and Lichens. Canad. J. Bot.
1980, 58, 20 : 2145-2147 (Douglas Ecol. Consultants Ltd., 2049 Crescent
Road, Victoria, BC Canada V8S 2G9).
Mousses et lichens avec loc. 1 mousse et 5 lichens nouv. pour la British
Columbia et 8 mousses pour le Yukon.
HERNANDEZ PADRÓN С. y PÉREZ DE PAZ P.L. — Estudio preliminar de
los líquenes epifitos del Sabinar de la Dehesa en El Hierro (Islas Canarias).
Vieraea «1979» 1980, 9, 1-2 : 15-32, 4 fig., 3 tabl, (Mus. Ins. Ci. Nat., Apdo.
853, Santa Cruz de Tenerife, Tenerife, Islas Canarias).
Identification de 43 esp. épiphytes du Juniperus phoenicea L. dans l'ile
de Hierro. 8 sont nouv. pour Hierro, 1 pour les Iles Canaries (Parmelia stuppea
Tayl.). Analyse des facteurs déterminant leur distribution.
MAKRYI T.V. — New species of lichen flora of the USSR. Bot. Zurn. (Moscow
et Leningrad) 1981, 66, 1 : 129-132, en russe (Zentral'niji Sibirskii Bot. Sad,
CO AN SSSR, Novosibirsk).
Parmelia olivaceoides Krog., P. predisjuncta Essl., P. ruderata Vain., Solari-
nella asteriscus Anzi пошу. pour l'URSS.
MONTACCHINI F. PIERVITTORI В. — Studi sulla vegetazione del Parco
Nazionale del Gran Paradiso. I. Prime osservazioni sulla flora e vegetazione
lichenica nell orizzonte alpino e subalpino del versante Piemontese del
P.N.G.P. Alloinia «1978-1979» 1980, 23 : 161-182, 14 tabl. (Ist. Bot., Univ.
Torino, Italia).
Source : MNHN, Paris
486 BIBLIOGRAPHIE LICHÉNOLOGIQUE
Caractéristiques de la végétation phanérogamique et lichénique. De nombreux
lichens sont nouv. pour la région du Parc national de Gran Paradiso. Associations
lichéniques.
OSORIO H.S. — Contribution to the lichen flora of Uruguay. XIII. Lichens from
Sierra Mahoma, San Jose department. Phytologia 1980, 45, 3 : 217-220
(Dept. Bot., Mus. Nac. Hist. Nat., Montevideo, Uruguay).
51 taxa avec loc. de Uruguay SW. Xanthoria plittii (Gyeln) Hale nouv. pour
PUruguay.
OSORIO H.S. — Contribution to the lichen flora of Uruguay XV. Additional
records to the Rio Uruguay lichen flora. Phytologia 1980, 46, 3 : 137-142
(Idem).
70 taxa avec loc. Candelariella vitellina (Ehrh.) Müll. Arg., Leptogium margi-
nellum (Sw.) S. Gray, Opegrapha bonplandia var. imitans Redgr., О. lichenoides
var. octomera Redgr., Physcia aipolia (Ehrh.) Hpe., Porina subpungens Malme,
Pyrenula brunnea Fée et Rinodina megapotamica Malme sont nouv. pour l'Uru-
guay.
OSORIO H.S. and ARBELO MARTINS C.S. — Contribution to the lichen
flora of Brazil V. Three interesting records from Rio Grande do Sul. Phytolo-
gia 1980, 46, 4 : 229-230 (Idem).
Stereocaulon ramulosum (Sw.) Rausch, est nowy. pour le Rio Grande do Sul.
La présence de Cladonia confusa В. Sant. et celle de Cl. chondrotypa Vain, sont
confirmées.
OSORIO H.S. — Contribution to the lichen flora of Uruguay XVI. Lichens
collected by Mariano B. Berro. Phytologia 1981, 47, 5 : 393-396 (Idem).
Liste de 26 taxons avec loc.
PUEYO G. — Conditions climatiques d'une station lichénique de la Cóte Basque
(suite). Bull. Cent. Etud. Rech. Sci. Biarritz 1980, 13, 1 : 131-138, 3 tabl.
(Lab. Mus., C.E.R.S., Biarritz, France).
SEE М.С. and BLISS L.C. — Alpine lichen-dominated communities in Alberta
and the Yukon. Canad. J. Bot. 1980, 58, 20 : 2148-2170, 10 tabl., 12 fig.
(Dept. Bot., Univ. Alberta, Edmonton, Alta., Canada T6G 2E9).
Descr. de 3 communautés alpines de macrolichens; Cetraria tilesii et Thamno-
lia subuliformis caractérisent les sites les plus secs dans les 2 régions; Cetraria
cucullata, les habitats mésiques; des esp. de Cladonia dominent les sites acides
au Yukon, Stereocaulon alpinum et Peltigera aphthosa les habitats équivalents
dans l'Alberta. Descr. des communautés de pl. vasculaires. Relations avec le pH,
l'humidité du substrat, l'histoire de la glaciation. Comparaison avec d'autres
études nordiques.
SEPPELT R.D. — Lichen collections from Cape Denison, Commonwealth Bay,
King George V Land, Antarctica. Austral. Gov., Antarct. Div., Techn. Mem.
Source : MNHN, Paris
BIBLIOGRAPHIE LICHÉNOLOGIQUE 487
1980, 95 : 14 (Antarctic Div., Dept. Sci. & Environment, Melbourne, Victo-
ria, Australia).
Liste des esp. de Cap Denison.
SEPPELT R.D. — Bryophytes and lichens collected by the visit of Australian
Museum personnel to Macquarie Island, Summer 1977-1978. Austral. Gov.,
Antarct. Div., Techn. Mem. 1980, 94 : 1-11 (Idem).
Liste des loc. visitées. Liste des bryophytes et lichens avec loc.
SKYE E. — Continued investigations of epiphytic Lichen Flora around Kvarn-
torp in Naárke. Acta Phytogeogr. Suec. 1980, 68 : 141-152, 13 fig. (Inst.
Ecol. Bot., Uppsala Univ., Box 559, S-751 22 Uppsala).
Comparaison avec les études faites depuis 1952/53. Division des lichens selon
leur distribution dans l'aire étudiée. Changement en abondance selon le pH et
la pollution.
VALCUVIA PASSADORE M.G. — Contributo alla flora lichenica dell'Isola di
Montecristo (Arcipelago Toscano). Atti Ist. Bot. «Giovanni Briosi» set. 6,
«1978-79» 1980, 13 : 159-169 (Ist. Bot., Univ. Pavia, Italia).
Liste de 57 taxons avec loc., récoltés par S. Filipello et Е. Sartori (1973-
1976) sur l'ile de Montecristo. 30 sont nouv. pour l'Archipel toscan.
POLLUTION
BELTMAN LH., DE KOLK L.J., KUIPER Р.].С. and VAN HASSELT Р.В. —
Fatty acid composition and chlorophyll content of epiphytic lichens and a
possible relation to their sensitivity to air pollution. Oikos 1980, 35, 3 :
321-326, 1 fig, 3 tabl. (Dept. Pl. Physiol., Biol. Centre, Univ. Groningen,
Р.О. Box 14, 9750 Aa'Haren (Gr.) The Netherlands).
Suggestion : la sensibilité des lichens à la pollution atmosphérique serait liée
au degré de dépendance du mycobionte vis-a-vis du phycobionte, aussi long-
temps que l'énergie métabolique est concernée.
CLERC P. et ROH P.D. — Les lichens, indicateurs biologiques de la pollution
atmosphérique, autour de la fabrique d'aluminium de Martigny (Valais,
Suisse). Saussurea 1980, 11 : 107-139, 9 fig., 3 tabl. (Syst. Geobot. Inst.,
Univ. Bern, CH-3013 Bern).
3 méthodes sont utilisées et comparées pour doser les composés fluorés :
dosage des thalles de Xanthoria parietina in situ et nécrosés, étude des trans-
plants de Xanthoria parietina et Parmelia caperata pendant 6 mois, étude phyto-
sociol. des épiphytes de Peupliers dans 35 sites (IAP).
'
KAUPPI M. — Fluorescence microscopy and microfluorimetry for the ехатїпа-
tion of pollution damage in Lichens. Ann. Bot. Fenn. 1980, 17, 2 : 163-173,
Source : MNHN, Paris
488 BIBLIOGRAPHIE LICHÉNOLOGIQUE
16 fig., 1 tabl. (Dept. Bot., Univ. Oulu, Box 191, SF-90101 Oulu 10).
Descr. de la méthode d'investigation avec Hypogymnia physodes (L.) Nyl.
comme matériel. Cette méthode permettrait d'étudier les changements apportés
aux lichens par la pollution atmosphérique, et l'estimation des conditions de vie
de la couche cellulaire algale.
MOSER Т.]., NASH Ш T.H. and CLARK W.D. — Effects of a long-term field
sulphur dioxide fumigation on Arctic Caribou forage lichens. Canad. J. Bot.
1980, 58, 21 : 2235-2240, 3 tabl., 1 fig. (Dept. Bot. & Microbiol., Arizona
State Univ., Tempe, Az. 85281 USA).
Études in situ de l'influence de la fumigation continue de SO; sur la capacité
photosynthétique de Cladonia stellaris (Opiz.) Pouz. et Vezda, C. rangiferina
(L.) Wigg. et Cetraria cucullata (Bell.) Ach. Noter aprés un an, la teneur en chlo-
rophylle plus faible que chez les témoins.
PERKINS D.F., MILLAR R.O. and NEEP Р.Е. — Accumulation of Airbone
Fluoride by Lichens in the Vicinity of an Aluminium Reduction Plant. Envi-
ronm. Pollut. Ser. A, 1980, 21, 1 : 155-168, 4 tabl., 5 fig. (Inst. Terr. Ecol.,
Penrhos Road, Bangor, Gwynedd L57 2LQ, Wales, GB).
Les lichens corticoles semblent accumuler plus rapidement le fluor que les
saxicoles, d'où des dégâts accélérés et une réduction en nombre plus importante.
Róle du vent et de la distribution par rapport à l'usine. Évolution de la popula-
tion lichénique entre 1970 et 1978.
VARIA
BRUNEAU D. — Regards sur l'étude des Muscinées et des Lichens en Anjou.
Mém. Soc. Étud. Sci. Anjou «1980» 1981, 4 : 165-169 (44 rue du Pin,
49000 Angers).
Historique des récoltes et herbiers depuis 1800. Courtes notices des princi-
paux bryologues et lichénologues qui travaillérent sur la région.
HAWKSWORTH D.L. - Notes on some fungi occurring on Peltigera, with a key
to accepted species. Trans. Brit. Mycol, Soc. 1980, 74, 2 : 363-386, 11 fig.
(CMI, Kew Surrey TW9 3AF, Great Britain).
Clé aux 40 champignons exclusivement lichénicoles et notes pour 16 d'entre
eux. Le fait que tant d'esp. soient exclusives à ce genre d'hóte et que 6 genres
soient monotypiques, suggère que Peltigera représente un groupe particuliére-
ment ancien parmi les Lecanorales. Noter les gen., sp. et comb. nov. parmi les
champignons lichénicoles.
HERTEL H. — Index collectorum Lichenum Herbarii Monacensis. Ein Sammler-
Verzeichnis des Flechtenherbars der Botanischen Staatssammlung München.
Mitt. Bot. Staatssamml. München 1980, 16 : 333-462 (Bot. Staatssamml.,
Menziger Str. 67, D-800 München 19).
Source : MNHN, Paris
BIBLIOGRAPHIE LICHÉNOLOGIQUE 489
Liste de 1674 collecteurs de lichens, de l'herb. de Munich (M) : nom, prénom,
date de naiss. et de mort, littérature, biograph., aires de coll., années de coll.,
nb de spécimens récoltés.
MOXHAM Т.Н. — Lichens in the Perfume Industry. Dragoco Rep. 1981, 2 :
31-39, 3 fig., phot. coul. (Dept. Pl. Sci, Univ. Bath, Claverton Down, Bath
ВАР 7АУ, Great Britain).
Evernia prunastri (Oakmoss, mousse de chêne) et Pseudevernia furfuracea
(mousses d'arbre) sont les plus menacés : 7800 à 9200 tonnes par an sont
récoltées pour l'industrie du parfum, au Maroc, en Yougoslavie (principalement),
en France. Destruction des sites par élimination d’autres esp. telles Ramalina
et Usnea,
RENNER B. und GERSTNER E. — Mediumabhängige Anthrachinon-Bildung in
Mycobiontenkulturen von Caloplaca ferruginea. Naturwissenschaften 1980,
67, 7 : 352-353, 2 fig. (Fachber. Biol. & Chem., Univ., D-3550 Marburg).
Source : MNHN. Paris
рез e = Эст A
491
INDEX DU VOLUME 11
compilé par D. LAMY
Il ne figure que la première page de l'article dans lequel est cité la taxon.
Les nouveautés taxonomiques sont indiquées en italiques. Les taxons cités en sy-
nonymie ou comme basionymes sont indiqués par "syn." ou "bas.". Lorsque le numé-
ro de page est suivi d'un nom de région, le taxon est considéré comme nouveau
pour celle-ci (ex. Acaulon triquetrum, 289 Navarre).
BRYOPHYTES
Abietinella abietina, 127
Acer campestre, 181; obtusatum, 181;
pseudoplatanus, 277
Afrique tropicale, 91 Reproductive
phenology of some tropical african
mosses
Agrostis canina, 277; rupestris, 153
Alchemilla alpina, 153
Alicularia compressa, 277
Allium pendulinum, 181
Aloina ericaefolia, 233 syn.; rigida
var. ambigua, 233
Anblystegiella confervoides, 233
Amblystegium confervoides, 233 syn. ;
varium, 181; mougeotii, 423
Anastrophyllum minutum var. grandis,
127
Anemone nemorosa, 277
Anomobryum auratum, 77 cytologie; fi-
liforme var. concinnatum, 77 cytol.;
gemmigerum, 77 cytol.; nitidum, 77
Cytol.; sp.,77 cytol.
Anomodon attenuatus, 233; longifolius,
233
Aqui folio - Fagetum, 181
Archidium alternifolium, 181
Ardenne, 277 Hyocomium armoricum en -
Asteriscium, 379 syn.; torquescens,
379 syn.
Astomum, 101
Atractylocarpus, 441; brasiliensis,
441; longisetus, 441
Atrichum portoricense, 471 syn.; subu-
Tirostrum, 471 syn.; undulatum, 277
Aulacomnium palustre, 127
AWASTHI U.S., voir UDAR В. and AWASTHI
U.S., 345
Bangladesh, 57 Cololejeunea floccosa
Barbula, 1 du Mexique, 379; subgen.
Hyophiladelphus, 1; sect. Asterisci-
um, 379 bas.; sect. Revolutae, 1
syn.; subsect. Revolutae, 1 syn.;
abbonii, 379 syn.; acuta, 379 syn.,
ssp. icmadophila, 379 syn., var.
bescherellei, 379 syn.; agraria, 1;
altiseta, 379 syn. nov.; arcuata, l,
379; aurea, 1 syn.; australasiae,
379 syn.; bescherellei, 379 syn. ‚var.
crassinervia, 379 syn. nov., Var.
stenocarpa, 379 syn. nov.; bourgaea~
па, 379 syn. nov.; brachyphylla, 379
bas.; calcarea, 1 bas.;convoluta, 1;
cylindrica, 379, ssp. vinealis, 379
syn., var. vinealis, 379 syn.; ehren-
bergii, 1, var. mexicana, 1 syn.; e-
rythropoda, 379 syn. nov.; fallax,
233, 379 bas., var. reflexa, 379
syn.,. var. vinealis, 379 syn.; fer-
ruginea, 379 syn.; flaccidiseta, 379
syn. nov.; godmaniana, 379 syn. nov.;
gracilescens, 379 syn. nov.; gracili-
formis, 379 syn. nov.; gracilis ssp.
icmadophila, 379 syn., var. icmado-
phila, 379 syn.; graminicolor, 379
syn. nov.; hypselostegia, 1 syn.;
icmadophila, 379 bas.; inaequalifo-
lía, 1 bas.; indica, 1, 379, var.
gregaria comb. et atat. nov., 1, var.
indica, 1; lagunicola, 379 syn. nov.;
leptocarpa, 379 syn. nov.; lonchos-
tega, 379 syn. nov.; lozanoi, 379
syn. nov.; lurida, 379 syn.; michi-
ganensis, 379 syn.; mobilis, 379 syn.
поу.; muenchii, 1 syn.; nicholsonii,
379 syn.; olivacea, 379 syn.; oriza-
bensis, 1, 379; pringlei, 1 syn.,
379 syn.; rectifolia, 379 syn. nov.;
reflexa, 379 syn.; replicata, 1 bas.;
rigidula, 379 syn., ssp. nicholsonii,
379 syn.; rubiginosa, 379 bas.; ru-
Source : MNHN, Paris
492
ficaulis, 1 syn.; rufipes, 1 syn.;
saint-pierrei, 379 syn. nov.; sal
zarensis, 379 syn. nov.; spiralis,
1 syn., var. emarginata, 1 syn.;
stenotheca, 1 syn.; stillicidiorum,
1 syn.; strictidens, 379 syn. nov.;
subteretiuscula, 379 syn. nov.; te~
retiuscula, 379 syn. nov.; tophacea,
379 syn.; umbrosa, 379 bas.; ungui-
culata, 1; valida, 379; vinealis,
379 syn., ssp. cylindrica, 181,
var. flaccidus, 379 syn.
Bartramia, 101; ithyphylla, 153; po-
miformis, 181; stricta, 181
Bartramiaceae, 101 spore ornamenta-
tion in Leionela
Belgique, 277 Hyocomium armoricum en -
Betula pubescens, 277
Blasia, 223
Blechnum spicant, 277
Brachymenium acuminatum, 77 cytol.:
exile, 77 cytol.; leptophyllum, 77
cytol.; nepalense, 77 cytol.; sik-
Kimense, 77 cytol.
Brachypodium silvaticum, 277
Brachythecietalia plumosi, 277, 423
Brachythecietum plumosi, 277
Brachythecio plumosi - Hyocomietum
flagellaris, 277
Brachythecion rivularis, 423
Brachythecium plumosum - Hyocomium
flagellare, 277 assoc. à -
Brachythecium plumosum, 277; rivulare
277, 423; rutabulum, 181, 277; sa-
lebrosum, 181; starkei, 181; velu-
tinum, 181, var. salicinum, 181
Brésil, 441 Ergänzungen zur Campylo-
pus-Flora von Brasilien
Breutelia, 101
Bruchia, 101
Bryochamaephyta caespitosa, 277; rep-
tantia, 277
Bryoerythrophyllum, 1 du Mexique,
379; calcareum, 1, 379; ferruginas-
cens, 1, 379; jamesonii, 1; inae-
qualifolium, 1; recurvirostre, 233,
var. recurvirostre, 379; recurvum,
1
Вгуип, 77; bimum, 127; capillare,
181, 277, ssp. capillare, 181; pseu-
dotriquetrum, 423; uliginosum, 379
Buxbaumia aphylla, 91
California, 379 Didymodon in Mexico
and - : taxonomy and nomenclature
Callitriche sp., 277
Caltha palustris, 277
Calvados, 277 Hyocomium armoricum au -
Calypogeia arguta, 277; fissa, 277;
muellerana, 277
Cambodge, 47 Cololejeunea floccosa
Campanula cochlaerifolia, 153
Camptothecium lutescens, 233
INDEX
Campylium chrysophyllum, 233; hispidu-
lum yar. sommerfeltii, 233; sommer-
feltii, 233 syn.
Campylopus arctocarpus, 441; brachy-
phyllulus, 441 syn. noy.; brasilien-
sis, 441 syn.; catumbensis, 441; cus-
pidatus, 441; discriminatus, 441 syn.
поу.; filifolius, 441, var. humilis,
441; fragiliformis, 441; fragilis,
441; gardneri, 441; gracilicaulis,
441; introflexus, 349; Luetzelburgii
sp. nov. , 441 Brésil; marmellensis,
441; minarum, 441 syn. nov.; pilifer,
441; praealtus, 441 syn. nov.; recti-
setus, 441 syn.; rubricaulis, 441
syn. nov.; Savannarum, 441; subcus-
pidatus, 441; surinamensis, 441; tor-
tilipilus ap. nov., 441 Brésil
Cardamine flexuosa, 277; hirsuta, 277;
sp. ‚277
Cardaminion, 423
Carex pendula, 277; remota, 277
Carpinion betulii, 181
Carpinus, 181
CASTALDO R., voir GAMBARDELLA R., LI-
GRONE R. and CASTALDO R., 2
Cephalozia bicuspidata, 277
Ceratodon purpureus, 127, 181, ssp.
purpureus, 181
Ceratolejeunea, 223
Ceylan, 47 Cololejeunea floccosa
Chiloscyphus pallescens, 277
Chorologie, 277 Hyocomium armoricum en
Belgique et le nord-ouest de la
France
Chrysosplenium oppositifolium, 277
Cirriphyllum crassinervium, 181
Clematis vitalba, 1
Cololejeunea, 47 Notion d'espéce dans
le genre. -
Cololejeunea subgen. Lasiolejeunea, 47,
sect. Globigerae, 47, sect. Venustae,
47; subgen. Pedinolejeunea, 47; sub-
gen. Taeniolejeunea, 47 notion d'es-
рёсе, sect. Falcatae, 47, sect. Floc-
созае, 47; sect. Leonidentes, 47; a-
bnormis, 47; amoena, 47; арргезза,
47; bontocensis, 47; falcata, 47;
floccosa, 47 étude du complexe, var.
amoenoides var. nov.,47, var. angus-
tibracteata var. nov., 47, var. au-
viculata var. nov., 47,var. aurita,
47, var. floccosa, 47, var. ocellata
var. nov.,47, var. plicata var. nov.,
47, var. trivittata var. не
gynophthalma, 47; mutabilis, 47,
floccosoides, 47; peraffinis, 47;
preciosa, 47; punctata, 47; schmidtii
47; tenella, 47; vidaliana, 47
Colombie, 201 ЕТ genero Sphagnum en
los Andes de Colombia y Venezuela
Comunidades reofilas, 423 Estudio brio-
с 7479
Source : ММНМ Paris
INDEX
sociologico de las - Sierra Nevada
Conocephalum conicum, 277
Conostomum, 101
Coryllus avellana, 277
Cótes-du-Nord, 277 Hyocomium armoricum
dans les -
Crataegus monogyna, 181
Cratoneurion commutati, 423
Cratoneuron commutatum, 423, var. ir-
rigatum, 423; decipiens, 423
Ctenidium, 277; molluscum, 233,277,
f. condensatum, 277
Cyathea, 101 phorophyte
Cyclamen hederifolium, 181
Cynodontium, 379; trifarium, 379 bas.
Cytisus villosus, 181
Cytological observations on some West
Himalayan mosses VI, 77
Dactylhymenium pringlei, 379 syn.
Daphne laureola, 181
Deschampsia caespitosa, 277; flexuosa,
277
Descriptions and illustrations of Bar-
bula, Pseudocrossidium and Bryoery-
throphyllum (p.p.) of Mexico, 1
DE SLOOVER J.L. - Octoblepharum Mitt.
ex Williams et Steereobryon subuli-
rostrum (Schimp. ex Besch.) G.L.
Smith en Guadeloupe, 471
DE ZUTTERE Ph., voir SCHUMACKER R. ,
LECOINTE A., TOUFFET J., DE ZUTTERE
Ph., LECLERCQ L. et FABRI R., 277
DHIEN R. - Florule bryologique des
rochers bathoniens, 233
Dichodontietum pellucidi, 423
Dichodontium pellucidum, 423
Dicranella heteromalla, 277
Dicranum arctocarpum, 441 syn.; brachy-
phyllulum, 441 syn.; brasiliense,
441 syn.; discriminatum, 441 syn.;
filifolium, 441 syn.; humilis, 441
syn.; laxobasis, 441 syn.; praeal-
tum, 441 syn.; rabenii, 441 syn.
nov.; rectisetum, 441 syn.; savan-
narum, 441 syn.; scoparium,233; sub-
cuspidatum, 441 зуп.; undulatum, 127
Didymodon, 379 in Mexico and Califor-
mia : taxonomy and nomenclature;
sect. Asteriscium comb. nov., 379,
sect. Craspedophyllon, 379 syn. nov.,
sect. Didymodon, 379, sect. Fallax,
379, sect. Vineales, 379; acutus,
379 syn.; australasiae, 379, var.
australasiae, 379, var. umbrosus
comb. et stat. nov., 379; brachyphyl-
lus, 379 syn.; craspedophyllus, 379
syn. поу, ; diaphanobasis, 379 syn
fallax, 379, var. reflexus, 379
filicaulis, 3/9 syn. nov.; fuscovi-
ridis, 379 syn. nov.; godmanianus,
379 syn. nov.; heribaudii, 379 syn.
nov.; icamdophyllus, 379 syn.; in-
crassatolimbatus, 379; johansenii,
493
379; juniperinus, 379; luridus, 379
bas., ssp. nicholsonii, 379 syn., var.
nicholsonii, 379 syn.; mexicanus, 379
syn. поу., Var. subulatus, 379 bas.;
michiganensis, 379; nicholsonii, 379
bas.; occidentalis, 379 syn.; paten-
tifolius, 379 syn. nov.; planifolius,
379 syn. nov.; pusillus, 379 syn. nov.;
ramulosus, 379 зуп.; revolutus, 379;
rigidulus, 379, var. gracilis comb.
et stat. nov., 379, var. icnadophila
comb. nov., 379, var. incrassatolim-
batus, 379, var. nicholsonii, 379
syn., var. rigidulus, 379, var. subu-
latus comb. nov., 379; rubellus, 233
syn.; rubiginosus, 379; tectorum, 379;
tophaceus, 379; torquescens, 379 syn.
поу.; trifarius ssp. nicholsonii, 379
syn., var. luridus, 379 syn.; vinea-
Tis, 379, var. brachyphyllus comb.
nov.,379, var. flaccidus, 379, var.
luridus comb. пор. ‚379, var. nichol-
sonii comb. nov., 379, var. rubigino-
sus comb. et atat. nov., var. vinea-
lis, 379; viridissimus, 379 syn. nov.
Digitalis purpurea, 277
Diplasiolejeunea, 47; groupe des Cornu-
tae, 47
Diplophyllum albicans, 277
Distichium inclinatum, 379; palleaceum,
379
Ditrichum flexicaule, 233
Doronicum orientale, 181
Drepanocladus uncinatus, 127
Dryopteria carthusiana, 277
Ecologie, 91 reproduction de mousses
d'Afrique tropicale, 181 végétation
muscinale de Malabotta, 201 Sphagnum
de Colombie et du Venezuela, 277 Hyo-
comium armoricum en Belgique et dans
le nord-ouest de la France, 423, com-
munautés réophiles de la Sierra Ne-
vada
Encalypta, 101; affinis, 153; alpina,
153; brevicolla, 153; brevipes, 153;
ciliata, 153, f. depauperata, 153,
cf. f. gymnostoma, 153, var. gymnos-
toma, 153, var. microstoma, 153;
flowersiana, 153; laciniata ssp. mi-
crostoma, 153 syn.; longicolla, 153;
microstoma, 153 The taxonomic status
of -; mutica, 143; rhabdocarpa, 153,
var. leptodon, 153; spathulata, 153,
vittiana, 153; vulgaris, 233
Entodon concinnus, 233; orthocarpus,
233 syn.
Epilobium sp., 277
Epipetria amphiphytia, 277
Ergänzungen zur Campylopus-Flora von
Brasilien, 441
Espèce, 47 Notion d' - chez le genre
Cololejeunea. Le complexe Cololejeu-
Source : MNHN, Paris
494
nea floccosa (Lehm. et Lindenb.)
Schiffn.
Estudio briosociologico de Tas comuni-
dades reofilas de Sierra Nevada (Es-
paña), 423
Eucladium verticillatum, 233
Euphorbia amygdaloides, 181
Eurhynchium pulchellum, 153; stokesii,
277; striatum var. striatum, 181
FABRI R. , voir SCHUMACKER R., LECOIN-
TE A., TOUFFET J., DE ZUTTERE Ph.,
LECLERCQ L. et FABRI R., 277
Fabronion pusillae, 181
Fagion, 181
Fagus, 181
Fegatelletum conicae, 277
Festuca haleri, 153
Finistère, 277 Hyocomium armoricum
Fissidens cristatus, 181, 233; curno-
vii, 277; glauculus, 91 reproduction;
minutulus ssp. pusillus, 181; novae-
hollandiae, 449 syn.; polyphyllus,
277, var. lusitanicus, 277; taxifo-
lius, 181, 277, ssp. taxifolius, 181
Fissidens polyphyllus et Hyocomium
flagellare, 277 assoc. à -
Fissidenti polyphylli - Hyocomietum
flagellaris, 277
Floristique, 1 Barbula, Pseudocrossi-
dium, Bryoerythrophyllum du Mexique;
181 Malabotta (Sicile); 201 Sphagnum
des Andes de Colombie et du Venezue-
Та; 233 Côte-d'Or; 379 Didymocon du
Mexique et de Californie;423 Sierra
Nevada; 441 Campylopus du Brésil;
449 Rhizogonium et Pyrrhobryum de
Malaya; 471 Guadeloupe
Florule bryologique des rochers ba-
thoniens, 233
Fontinaletalia antipyreticae, 181, 277
Fontinaletea antipyreticae, 423
Fontinaleto - Pachyfissidentetum gran-
difrondis, 423
Fontinalion antipyreticae, 423
Fontinalis antipyretica, 423, var.
gracilis, 277; squamosa, 277
Fossombronia, 223; caespitiformis, 223
résistance à la sécheresse, multipli-
cation végétative; fimbriata, 223;
himalayensis, 223; lamellata, 223;
pusilla, 223
FRAHM J.P. - Ergänzungen zur Campylo-
pus-Flora von Brasilien, 441
France, Nord-Ouest, 277 Hyocomium ar-
moricum
Freycinetia, 47 phorophyte
Frullania asagrayana, 223; tamarisci,
233
GAMBARDELLA R., LIGRONE R. and CASTAL-
DO R. - Ultrastructure of the sporo-
phyte foot in Phaeoceros, 23
Gemma, 457 Regeneration and - deve-
INDEX
lopment in Hyophila crenulata C.
Muell. ex Dus.
ЕТ genero Sphagnum en los Andes de Co-
lombia y Venezuela. Clave para las
especies frecuentes u ocasionales
con notas ecologicas y taxonomicas,
Geranio - Fagion, 181
Geranium robertianum, 277; versicolor,
181
GIL J.A. y VARO J. - Estudio briosocio-
logico de las comunidades reofilas
de Sierra Nevada (España), 423
Glyceria fluitans, 277
GRIFFIN III D. - Spore ornamentation
in Leiomela (Musci : Bartramiaceae) ,
101 ; El genero Sphagnum en los Andes
de Colombia y Venezuela. Clave para
las especies frecuentes u ocasionales
con notas ecologicas y taxonomicas,
201
Grimmia apocarpa, 233 syn.; orbicula-
ris, 233 syn.; pulvinata, 181, 233,
var. africana, 233; trichophylla,
181, ssp. trichophylla, 181; sp.;
181
Grimmia pulvinata - Tortula muralis,
181 assoc. à -
Grimmietalia, 181
Guadeloupe, 471 Otcoblepharum erecti-
folium Mitt. ex Williams et Steereo-
bryon subulirostrum (Schimp. ex
Besch.) G.L. Smith
Gymnostomum calcareum, 233
Gyroweisia, 379; obtusifolia, 379; pa-
pillosa, 379
Haplomitrium hookeri, 223
Hedera helix, 277
Hedwigidium imberbe, 277
Helvella sp. 277
Heterocladium heteropterum, 277, ssp.
wulfsbergii, 277
Hétraie, 181 végétation muscinale de
Та hétraie de Malabotta (Peloritani)
Himalaya, 77 Cytological observations
on some West Himalayan mosses
Holcus mollis, 277
Homalothecio - Porelletum platyphyllae,
181
Homalothecium philippeanum, 181; seri-
ceum, 181, 233
Hookeria lucens, 277
HORTON D.G. - The taxonomic status of
Encalypta microstoma Bals. et De Not.
and E. ciliata var. microstoma Schimp,
153
Husnotiella, 379 syn.; obtusifolia, 379
syn.; palmeri, 379 syn.; pringlei,
379 зуп, ; revoluta, 1, 379 bas., var.
palmeri, 379 syn.; torquescens, 379
syn.
Hydrocotyle vulgaris, 277
Source : MNHN, Paris
INDEX
Hydrogonium arcuatum, 1 syn.
Hygrohypnetalia, 423
Hygrohypnetea, 423
Hygrohypnetum dilatati tatrense, 423
syn.
Hygrohypnion dilatati, 423
Hygrohypnum lusitanicum, 277; ochra-
Ceum, 277, 423
Hylocomium splendens, 127, 233, ssp.
alaskanum, 127
Hymenostomum, 101
Hyocomium armoricum (Brid.) Wijk et
Marg. en Belgique et dans le nord-
ouest de la France (Ardenne, Breta-
gne, Normandie). Etude chorologique,
écologique et phytosociologique,
277
Hyocomium flagellare, 277 syn.
Hyophila, 457; crenulata, 457 Regene-
ration and gemma development in -;
propagulifera, 457
Hypnum cupressiforme, 181, 233, 277,
ssp. cupressiforme, 181, f. tecto-
rum, 181, f. filiforme, 181, var.
uncinulatum, 181; vaucheri, 127
Ilex aquifolium, 181
Ille-et-Vilaine, 277 Hyocomium armo-
ricum
India, 345 A new species of Lejeunea
from -
Isopaches bicrenatus, 223
Isopterygium elegans, 277
Isothecium myosuroides, 277, var. ri-
vulare, 277; myurum, 181, f. robus-
tum, 181
Isothecium myurum et Homalothecium
sericeum, 181 groupement à -
Jamesoniella autumnalis, 277
Japon, 47 Cololejeunea floccosa
Java, 47 Cololejeunea floccosa
Juncus bulbosus, 277; effusus, 277;
trifidus, 153
Jungermannia floccosa, 47 syn.; sphae-
rocarpa, 277
Juniperus communis, 153
Kingiobryum paramicola, 379
KUMAR 5.5. and VERMA S.K. - Cytologi-
cal observations on some West Hima-
Тауап mosses VI, 77
Lamium flexuosum, 181; galeobdolon, 277
LAMY D. - Bibliographie bryologique,
107, 235, 360, 474
Larix europaea, 127
Lathyrus venetus, 181
LECLERCQ L., voir SCHUMACKER R., LE-
COINTE A., TOUFFET J., DE ZUTTERE
Ph., LECLERCQ L. et FABRI R., 277
LECOINTE A., voir SCHUMACKER R., LE-
COINTE A., TOUFFET J., DE ZUTTERE
Ph., LECLERCQ L. et FABRI R., 277
Leersia laciniata var. microstoma,
153 syn.
495
LEFORESTIER C. - Résistance à la sé-
cheresse et multiplication végétati+
ve chez Fossonbronia caespitiformis
De Not., 223
Leiomela, 101 Spore ornamentation in -;
africana, 101; aristifolia, 101; bar-
tramioides ; javanica, 101
Lejeunea floccosa, 47 syn.; indica sp.
nov, 345 Inde; lamacerina, 277; mo-
niliata, 345; wallichiana, 345
Leptocolea floccosa, 47 syn.
Leptodictyetalia riparii, 423
Leptodictyum riparium, 181
Leptodon smithii, 181
Leskeella nervosa, 181
Leucodon sciuroides, 181
Leucodontetalia, 181
Liège, 277 Hyocomium armoricum
LIGRONE R., voir GAMBARDELLA R., LI-
GRONE R. and CASTALDO R., 23
Listea longifolia, 47 phorophyte
LO GIUDICE R., voir PRIVITERA M. et LO
GIUDICE R., 181
Lophocolea cf. cuspidata, 277; hetero-
phylla, 277
Lophozia collaris, 233; hatcheri, 127;
muelleri, 233 syn.
Lunularia cruciata, 91
Luxembourg, 277 Hyocomium armoricum
Luzula silvatica, 277
Lysimachia nemorum, 277
Madagascar, 47 Cololejeunea floccosa
Malabotta, 181 Vegetazione muscinale
della faggeta del bosco di Malabotta
(Peloritani)
Malaisie, 47 Cololejeunea floccosa
Malaya, 449 Synopsis of Rhizogoniaceae
in-
Manche, 277 Hyocomium armoricum
MANUEL M.G. - Synopsis of Rhizogonia-
ceae Broth. in Malaya, 449
Marsupella aquatica, 277; emarginata,
277
Mastigolejeunea auriculata, 223
Melica uniflora, 181
Metzgeria, 223; furcata, 277; pubescens,
233; uncigera, 223
Mexico, 1 Descriptions and illustra-
tions of Barbula, Pseudocrossidium
and Bryoerythrophyllum (p.p.) of -;
379 Didymodon in - and California:
taxonomy and nomenclature
Mielichhoferia sasaokae, 77 cytologie;
sp., 47 cytol.
Minuaria laricifolia, 153
Unio homi - Hyocomietun ammorioi
ass. поо. 277; brachythectetosun
plumosi subass. nov., 277; fissiden-
‚tetosum curnovit subass. nov., 277;
fissidentetosum polyphylli subase.
hov., 277; heteroeladietosum hetero-
Source : MNHN. Paris
496 INDEX
ptert subass. nov., 277; hookerieto-
sun lucentis subase. nov. ,277; hyo-
comietosun armoriet subass. nov.,
277; marsupelletosum emarginatae
subass. nov., 277;nardietosun com
pressae subaso, пор. ,277;тћасоті-
ec? acteularia subase. nov. ,
277
Mnium, 379; hornum, 277; punctatum,
277; rostratum, 181; undulatum, 277
Moehringia trinervia, 277
Molinia coerulea, 277
Montio - Cardaminetea, 423
Montion, 423
Morbihan, 277 Hyocomium armori cum
Morphologie, 1 Barbula, Pseudocrossi-
dium, Bryoerythrophyllum; 101 spores
de Leiomela; 345 Lejeunea indica sp.
поу.; 379 Didymodon ; 449 Pyrrho-
bryum et Rhizogonium; 457 gemmules
de Hyophila crenulata
Multiplication végétative, 223 Résis-
tance à la sécheresse et - chez
Fossombronia caespitiformis De Not.;
457 gemmules de Hyophila crenulata
Myurella julacea, 153
Namur, 277 Hyocomium armoricum
Nardia compressa, 277
Neckera complanata, 233; crispa, 233
(A) New species of Lejeunea from In-
dia, 345
Nomenclature, 153, statut d'Encalypta
microstoma; 379 Didymodon in Mexico
and California : taxonomy and -
Normandie, 277 Hyocomium armoricum
(La) Notion d'espèce chez le genre Co-
lolejeunea. Le complexe Cololejeunea
floccosa (Lehm. et Lindenb.) Schiffn.
47
Nouvelle-Calédonie, 47 Cololejeunea
floccosa
Nouvelle-Guinée, 47 Cololejeunea floc-
cosa
Nowellia curvifolia, 223
Octoblepharum albidum, 471; erectifo-
Tium, 471 Guadeloupe; pulvinatum, 471
ODU Е.А. - Reproductive phenology of
some tropical african mosses, 91
Oenanthe crocata, 277
OLARINMOYE 5.0. - Regeneration and gem-
ma development in Hyophila crenula-
ta C. Muell. ex Dus., 457
Orne, 277 Hyocomium armoricum
Orthothecium intricatum, 233
Orthotrichetum lyellii, 181
Orthotrichum affine, 181; anomalum, 91,
233; lyellii, 181; schimperi, 181;
sp., 181
Osmunda regalis, 277
Oxalis acetosa, 277; acetosella, 277
Oxyrrhynchietum rusciformis, 181, 277
Oxyrrhynchium praelongum var. praelon-
gum, 181, var. stokesii, 181; pumi-
lum, 233; schleicheri, 181
Oxystegus tenuirostris var. tenuiros-
tris, 379
Pedinophyllum interruptum, 233
Pellia epiphylla, 277
Pellietum epiphyllae, 277
Petalophyllum indicum, 223
Phaeoceros, 23 Ultrastructure of the
sporophyte foot in - ; laevis ssp.
laevis, 23
Phenology, 91 Reproductive -
tropical african mosses
Philippines, 47 Cololejeunea floccosa
Philonotis, 101; seriata, 423
Phyteuma hemisphaerica, 153
Phytosociologie, 277 Hyocomium armori-
cum en Belgique et dans le nord-ouest
de la France
Picea abies, 277
Plagiochasma, 223
Plagiochila asplenioides, 181, 233,277
Plagionnium rostratum, 423
Plagiopus, 101; oederi, 233
Plagiothecium nemorale, 181; silvati-
cum, 277; undulatum, 277
Plasteurhynchium striatulum, 181, f.
cavernarum, 181
Platyhypnidietosum riparioidis, 423
Platyhypnidietum riparioidis, 277 syn.
Platyhypnidio - Fontinaletea, 423
Platyhypnidium riparioides, 181, 277,
423; rusciforme var. lusitanicum,
277
Pleuridium subulatum, 181
Pleurozium schreberi, 91,127, 233
Pogonatum aloides, 181; tortile, 471
Pohlia, 77 cytologie; elongata, 77 су-
tol. longicollis, 77 cytol.; nutans,
127, 277
Polygonatum verticillatum, 277
Polytrichum , 379; aloides, 91; commu-
ne, 127, 277; formosum, 277; junipe-
rinum, 127, 349; piliferum, 91, 127,
349
of some
Pottiaceae, 379 Didymodon in Mexico
and California
PRIVITERA M. e LO GIUDICE В. - Vegeta-
zione muscinale della faggeta del
bosco di Malabotta (Peloritani), 181
Prunus sp., 277
Pseudocrossidium, 1 du Mexique; aureum,
1; replicatum, 1; revolutum, 1
Pseudoleskea catenulata, 233 syn.
Pseudoleskeella catenulata, 233
Pseudoscleropodium ригип, 233
Psychotria sp., 47 phorophyte
Pterigynandrum filiforme, 153, 181
Pterogonium gracile, 181
Ptilidium ciliare, 127
Source - ММНМ Paris
INDEX
Ptilium crista-castrensis, 91
Pylaisia polyantha, 91
Pyrrhobryum, 449 de Malaya; latifo-
lium, 449; longiflorum, 449; spini-
forme, 449, var. badakense, 449,
var. spiniforme, 449
Querco - Fagetea, 181
Quercus, 181; cerris, 181; pubescens,
181; sp., 277
Racomitrion acicularis, 423
Racomitrium aciculare, 277; canescens,
233; lanuginosum, 127
Racopilum africanum, 91 reproduction
Radula complanata, 181
Ranunculus ficaria, 277; lanuginosus,
181; repens, 277
Regeneration and gemma development in
Hyophila crenulata C. Muell. ex Dus.,
457
Reproductive phenology of some tropi-
cal african mosses, 91
Résistance à la sécheresse et multi-
plication végétative chez Fossombro-
nia caespitiformis De Not., 223
Rhizogoniaceae, 449 Synopsis of ~
Malaya
Rhizogonium, 449 de Malaya; badakense,
449 syn.; latifolium, 449 syn.; lon-
giflorum, 449 syn.; novae-hollandiae
449; spiniforme, 449 syn.
Rhynchostegiella algiriana, 233 syn.;
tenella, 233, f. meridionalis, 181
Rhynchostegium confertum, 181
Rhytidiadelphus loreus, 277
Rhytidium rugosum, 127
Riccardia chamaedryfolia, 277; multi-
fida, 277
Riccia fluitans, 223; perennis, 223
Rochers bathoniens, 233 Florule bry-
ologique des -
Rubus glandulosus, 181; sp., 277
Saccogyna viticulosa, 277
Saelania glaucescens, 379
Sambucus nigra, 277
Saxifraga bryoides, 153; paniculata,
in
Scapania aspera, 233; curta, 277; un-
dulata, 277, 423, f. dentata, 277
Scapanietum undulatae, 277, 423
Scapanion undulatae, 423
Schistidio - Hygrohypnetum dilatati,
423
Schistidium apocarpum, 181, 233
Schistostega pennata, 457
SCHUMACKER R., LECOINTE А. , TOUFFET
J., DE ZUTTERE Ph., LECLERCQ L. et
FABRI R. - Hyocomium armoricum
(Brid.) Wijk et Marg. en Belgique
et dans le nord-ouest de la France
(Ardenne, Bretagne, Normandie). Etu-
de chorologique, écologique et phy-
tosociologique, 277
497
Scirpus fluitans, 277
Scleropodium caespitosum, 277
Sécheresse, 223 Résistance à la -
et multiplication végétative chez
Fossombronia caespitiformis De Not.
Seligeria pusilla, 233
Semibarbula rufipes, 1 syn.
Sempervivum arachnoidum, 153
Senecio fuschii, 277
Sewardiella, 223
Sibthorpia europaea, 277
Sicile, 181 végétation muscinale de
Melabotta
Sierra Nevada, 423 Estudio briosocio-
logico de las comunidades reofilas
de - (España)
Silene acaulis, 153
Sociologie, 181 végétation muscinale
де Malabotta; 277 Hyocomium armori-
cum en Belgique et dans le nord-ouest
de la France; 423 communautés réo-
philes de la Sierra Nevada
Solenostomo - Hygrohypnetum, 423
Sphagnum, 201 El genero Sphagnum en
los Andes de Colombia y Venezuela.
Clave para las especies frecuentes
и ocasionales con notas ecologicas
y taxonomicas;acutifolium, 201 syn.
var. meridense, 201 syn.; andinum
201 syn.; apiculatum, 277; apolli-
nairei, 201 syn. sauriculatum,277;
capillaceum, 201 syn.; capillifolium,
201; compactum, 201; coryphaeum, 201
syn.; cuspidatum, 20l;cyclophyllum,
201; erythrocalyx, 201; gracilescens,
201 syn.; flavicaule, 201 syn.; gir-
gensohnii, 277; lehmannii, 201 syn.;
Timbatum, 201 et 201 syn.; magella-
nicum, 201; medium, 201 syn.; meri-
dense, 201; oxyphyllum, 201; palus-
tre, 201, 277; perichaetiale, 201
syn.; peruvianum, 201 syn.; pulchri-
coma, 201 syn.; pylaesii, 201; re-
curvum, 201; sancto-josephense, 201;
sanguinale, 201 syn.; serratum, 201
syn.; sparsum, 201; subnitens, 277;
subsecundum, 201; trinitense, 201
зуп.; sp.» 127, 277
Splachnum luteum, 127
Spore ornamentation in Leiomela (Musci:
Bartramiaceae), 101
Sporophyte foot, 23 Ultrastructure of
the - in Phaeoceros
Steereobryon, 471 Guadeloupe; subuli-
rostrum, 471 Guadeloupe
Stereophyllum sp., 91 reproduction
Streblotrichum convolutum, l syn.;
hypselostegium, 1 syn.; pringlei, 1
syn.
Symbiezidium floccosum, 47 syn.
Synopsis of Rhizoginiaceae Broth. in
Malaya, 419
Source : MNHN, Paris
498 INDEX
Taeniolejeunea floccosa, 47 syn.
(The) Taxonomic status of Encalypta
microstoma Bals. et De Not. and Е.
ciliata var. microstoma Schimp., 153
Taxonomie, 47 le complexe Cololejeunea
floccosa; 153 Encalypta microstoma;
201 Sphagnum de Colombie et du Vene-
zuela; 379 Didymodon in Mexico and
California : - and nomenclature
Taxons nouveaux, l, 47, 277, 345, 379,
1
Tetraplodon mnioides, 127
Teucrium scorodonia, 277
Thaïlande, 47 Cololejeunea floccosa
Thamnium alopecurum var. protensum,
277
Thamnobryum alopecurum, 181 groupement
Thuidium gratum, 91 reproduction; ta-
mariscinum, 277
TIXIER P. - La notion d'espéce chez
le genre Cololejeunea. Le complexe
Cololejeunea floccosa (Lehm. et
Lindenb.) Schiffn., 47
Tomenthypnum mitens, 127
Tortella tortuosa; 153, 233, f. fra-
gilifolia, 181
Tortula, 1; acuta, 379 syn.; agraria,
1 syn.; aurea, ] bas.; australasiae,
379 bas.; convoluta, 1 syn.; fallax
var. vinealis, 379 syn.; gracilis,
379 bas. ;graminicolor, 379 syn.;
gregaria, 1 bas.; icmadophila, 379
syn.; indica, 1 bas.; muralis, 181,
233; olivacea, 379 syn. nov.; rec-
tifolia, 379 syn.; reflexa, 379
syn. ;replicata, 1 syn.; rigidula,
379 syn.; ruralis, 127, 181, 233,
ssp. ruralis, 181; spiralis, 1 syn.;
subulata, 181, ssp. subulata, 181;
teretiuscula, 379 syn.; vinealis,
379 syn.
TOUFFET J., voir SCHUMACKER R., LE-
COINTE A., TOUFFET J., DE ZUTTERE
Ph., LECLERCQ L. et FABRI R., 277
Trichocolea tomentella, 277
Trichostomopsis, 379 syn.; australasi-
ae, 379 syn.; brevifolia, 379 syn.;
crispifolia, 379 syn., diaphanobasis,
379 syn.; fayae, 379 syn.; umbrosa,
379 syn.
Trichostomum brachydontium, 379, ssp.
mutabile, 233; crispulum, 181, var.
angustifolium, 379; ehrenbergii, 1
syn.; indicum, 1 syn.; luridum, 379
syn.: mexicanum, 379 syn.; mutabile,
233 syn., 379 syn.; obtusifolium,
379 syn.; ramulosum, 379 syn. nov.;
rigidulum, 379; tophaceum, 379 syn.;
viridulum, 379
Triquetrella, 379; ferruginea, 379 syn.
UDAR В. and AWASTHI U.S. - A new spe-
cies of Lejeunea from India, 345
Ultrastructure of the sporophyte foot
in Phaeoceros, 23
VARO J., voir GIL J.A. y VARO J., 423
Vegetazione muscinale della faggeta
del bosco di Malabotta (Peloritani),
181
Venezuela, 201 El genero Sphagnum en
los Andes de Colombia y - . Clave
para las especies frecuentes u oca-
Sionales con notas ecologicas y ta-
xonomicas
VERMA S.K., voir KUMAR S.S. and VERMA
STE
Vietnam, 47 Cololejeunea floccosa
Viola sp., 277
Weissia, 101
ZANDER В.Н. - Descriptions and illus-
trations of Barbula, Pseudocrossidi-
um and Bryoerythrophylium (p.p.) of
Mexico, 1; Didymodon (Pottiaceae) in
Mexico and California : taxonomy and
nomenclature of discontinuous and
nondiscontinuous taxa, 379
Zygodon viridissimus, 181, ssp. viri-
dissimus, 181
LICHENS
Abietinello - Piceetum glaucae, 127
Agropyron yukonensis, 127
Alectoria nigricans, 127; ochroleuca,
127
Alectorio - Masonhaleetum richardsonii
nova,
Amaioud guianensis, 323
Anatomie, 253 périthèces de Porina bys-
sophila; 349 Cladonia berghsonii
Andromeda polyfolia, 127
Anthracobia nitida, 253
Apodytes dimidiata, 461 phorophyte
Araucaria angustifolia, 323
Arctostaphylos uva-ursi, 127
Arnica cordifolia, 127
Artemisia alaskana, 127; arctica, 127;
frigida, 127; rupestris, 127
Artemisio - Agropyrion yukonensis, 127
Arthonia, 461; accolens 323 Rio Grande
do Sul; mira, 323 Rio Grande do Sul
Ascomycétes, 253
ASPERGES M. - A new lichen species,
Cladonia berghsonii Asperges sp. no-
va (sect. Cocciferae), 349
Aster alpinus, 127
Aulaxina quadrangula, 323
Bacidia apiahica, 323; brasiliensis,
323; pallidocarnea, 323 Rio Grande
Source : MNHN, Paris
INDEX 499
do Sul
Belgique, 348 Cladonia berghsonii
Bersama, 461 phorophyte
Betula, 171 phorophyte; glandulosa,
127
Brazzea longipedicellata, 461 phoro-
phyte
Brésil, 323 écologie et phytogéogra»
phie des lichens foliicoles de Rio
Grande do Sul
Buellia elegans, 127; epigaea, 127
Burundi, 461 Contribution à l'étude
.des lichens du Kivu (Zaïre), du
Rwanda et du Burundi VI. Les genres
Coccarpia Pers. et Lobaria (Schreb.)
Hof fm.
Byssoloma leucoblepharum, 323; subdis-
cordans, 323
Calamagrostis canadensis, 127; purpu-
rascens, 127
Calluna, 349
Caloplaca tominii, 127
Calypso bulbosa, 127
Campanula aurita, 127
Carapa, 461
Carex concinna, 127; dioica, 127; fi-
lifolia, 127; glacialis, 127; petri-
cosa, 127; scirpoides, 127
Carex filifolia, 127 assoc. à
Carissa edulis, 461 phorophyte
Cassipourea, 461 phorophyte
Catillaria bouteillei, 323, semecarpi,
323
Cetraria cucullata, 127; islandica,
127 laevigata, 127; nivalis, 127;
tilesii, 127
Cetrarietum tilesii, 127
Cetrarion, 127; nivalis, 127
Chamaedaphne calyculata, 127
Chroodiscus coccineus, 323 Rio Grande
do Sul
CIFUENTES B., voir VICENTE C. and CIFU-
ENTES B., 213
Cladonia subgen. Cladina, 127; sect.
Cocciferae, 349; acuminata, 127; a-
maurocraea, 127; arbuscula, 127, ssp.
beeringiana, 127; bacillaria, 349;
herghsonit ep. nov., 349; cariosa,
127; cenotea, 127; chlorophaea, 127;
coccifera, 127; cornuta, 127; defor-
mis, 127; ecmocyna, 127; fimbriata,
127; floerkeana, 349, f. intermedia,
3493 gracilis ssp. elongata, 127,
var. dilatata, 127; incrassata, 349;
mitis, 127; multiformis, 127; phyl-
lophora, 127; pocillum, 127; pyxida-
ta, 127; rangiferina, 127; stellaris,
127; sulphurina, 349; symphycarpa,
127; uncialis, 127; verticillata,
127
Cladonietosum arbusculae, 127
Cladonietum alpestris, 127; cenoteae,
127; еспосупае, 127; mitis, 127
Cladonion, 127; arbusculae, 127
Coccocarpia, 461 Contribution à l'étu-
de des lichens du Kivu (Zalre), du
Rwanda et du Burundi VI. Les genres
- Pers. et Lobaria (Schreb.) Hoffm.
azurella, 461; cronia, 461 syn.; e-
rythoxyli ("erythroxiTi"), 461; mo-
lybdea ("molybdina"), 461; palmicola,
461; parmelioides, 461 syn.; pellita,
461, var. semiincisa, 461; virides-
cens, 461
Cocos nucifera, 461 phorophyte
Collema coccophorum, 127
Combretum, 461
Conopharyngia, 461 phorophyte
Contribution à l'étude des lichens du
Kivu (Zaire), du Rwanda et du Burun-
di VI. Les genres Coccocarpia Pers.
et Lobaria (Schreb.) Hoffm. ‚461
Cornicularia aculeata, 127; divergens,
127
Cornus canadensis, 127
Cyathicula coronata, 253
Dactylina arctica, 127; ramulosa, 127
Dermatocarpon hepaticum, 127
Dimerella epiphylla, 323
Disconycètes, 253
Dryadetalia integrifoliae, 127
Dryas integrifolia, 1
Ecologie, 127 synusies épigées dans le
Yukon Territory; 323 lichens foliico-
les du Rio Grande do Su1; 461 Coccocar-
pia et Lobaria du Kivu,Rwanda ‚Burundi
Enpetrum nigrum, 127
Entada, 461
Entandrophragma, 461
Epibryetalia, 127
Epigaeous lichen synusiae in the Yukon
Territory, 127
tpigaetea, 127
Epilobium angustifolium, 127
Epipetretea, 127
Epiphytetea, 127
Epixiletalia, 127
Equisetum scirpoides, 127
Erigeron caespitosus, 127
Eriophorum vaginatum, 127
Erythroxylon squamatus, 461 phorophyte
Etudes ontogéniques chez le Porina bys-
sophila (Pyrénolichens) II - Les pé-
rithéces, 253
Eucalyptus, 461 phorophyte
Eugenia sp., 323
Festuca altaica, 127
Ficalhoa laurifolia, 461 phorophyte
Floristique, 127 Lichens épigés du
Yukon Territory; 323 lichens folii-
coles du Rio Grande do Sul; 461 Coc-
carpia et Lobaria au Kivu, Rwanda et
Burundi
Fulgensia bracteata, 127
Source : MNHN. Paris
500
Fulgensto - caloplacetun tominii nova,
17
Gentiana propinqua, 127
Geocaulon lividum, 127
Growth rates of Xanthoria parietina
and their relationship to substra-
te texture, 171
Gyalectidium filicinum, 323; rotuli-
forme, 323
Hierochloe alpina, 127
Hollande, 349 Cladonia berghsonii
Hypno - Abietinello - Piceetum glaucae,
127
Icmadophila ericetorum, 127
JANEX-FAVRE M.C. - Etudes ontogéniques
chez le Porina byssophila (Pyrénoli-
chens). II - Les périthéces, 253
Juniperus horizontalis, 127
Kivu (Zaïre), 461 Contribution à 1'étu-
de des lichens du Kivu (Zaïre), du
Rwanda et du Burundi VI. Les genres
Coccocarpia Pers. et Lobaria
(Schreb.) Hoffm.
Klainedoxa, 461 phorophyte
LAWY D. - Bibliographie lichénologi-
que, 119, 247, 372, 483
Lecanora lentigera, 127
Lecidea erythroxyli, 461 bas.; palmi-
cola, 461 bas.
Ledum decumbens, 127; palustre, 127
Leptogium saturninum, 127
Leucosidea, 461 phorophyte
Lichen (sect. ?) Lobaria, 461 syn.;
pulmonarius, 461 bas.; retiger, 461
bas.
Limnaea borealis ssp. americana, 127
Linum perenne, 127
Lobaria, 461 Contribution à l'étude
des lichens du Kivu (Zaïre), du
Rwanda et du Burundi VI. Les genres
Coccocarpia Pers. et ~ (Schreb.)
Hoffm.; africana, 461 syn.; linita,
127; meridionalis, 461; natalensis;
461 syn.; pulmonaria, 181, 461, var.
meridionalis, 461; retigera, 461;
sublaevis, 461 Afrique continentale
Lopadium foliicola, 323; fuscum, 323;
puiggarii, 323
Luzula parviflora, 127
Lycopodium annotinum, 127
Macaranga, 461 phorophyte; schwein-
furthii, 461
Meesa lanceolata var. mildbraedii, 461 _
phorophyte
Martensia paniculata, 127
Masonhalea richardsonii, 127 ,
Mazosia melanophthalma, 323; phyllose-
ma, 323 Rio Grande do Sul; rotula,
323
Melandrium apetalum, 127
Mimulopsis arborescens, 461
Molinia, 349
INDEX
Moneses uniflora, 127
Morphologie, 349 Cladonia berghsonii
MOXHAM T.H. - Growth rates of Xantho-
ria parietina and their relationship
to substrate texture, 171
Neoboutonia, 461; macrocalyx, 461 pho-
rophyte
Nephroma arcticum, 127
(A) New lichen species, Cladonia bergh-
Sonii Asperges sp. nova (sect. Cocci-
ferae), 349
NIMIS P.L. - Epigaeous lichen synusiae
in the Yukon Territory, 127
Nuxia floribunda, 461 phorophyte
Opegrapha, 461
(Zur) Ökologie und Pflanzengeographie
blattbewohnender Flechten von Rio
Grande do Sul, 323
Onotgenie, 253 Etudes ontogéniques chez
le Porina byssophila (Pyrénolichens)
IT. Les périthèces
Oxytenanthera, 461 phorophyte; abyssi-
nica, 461
Parinari, 461 phorophyte; curatellifo-
lia, 461 phorophyte
Parmelia pellita, 461 bas.; wyomingica,
127
Pamelietum wyomingteas nova, 127
Parrya nudicaulis, 127
Pedicularis labradorica, 127
Peltigera aphthosa, 127; canina, 127;
leucophlebia, 127; rufescens, 127
Peltigero - Cladontetun еспогупае nova,
127
Penstemon gormanii, 127
Périthèces, 253 Etudes ontogéniques
chez le Porina byssophila (Pyréno-
Tichens) II. -
Permeability, 213 L-usnate and -
Petasites hyperboreus, 127
Philippia, 461
Physiologie, 213 L-usnate et perméabi-
lité
Phytogéographie, 323 lichens foliicoles
du Rio Grande do Sul; 461 Соссосаг-
pia et Lobaria du Kivu, du Rwanda et
du Burundi
Phlox sibirica, 127
Phyllobathelium epiphyllum, 323
Physconia muscigena, 127
Picea, 349; glauca, 127
Pinetum montanae cladinetosum, 127
;Pinus, 349; contorta, 127; engadinen-
sis, 127; mariana, 127; sp.,127
Populus tremuloides, 127
Porina, 253, 461; atrocoerulea, 323;
byssophila, 253 Etudes ontogéniques
chez le Porina byssophila (Pyréno-
lichens) II. Les périthéces; epiphyl-
Та, 323; fulvella, 323; leptosperma,
323; limbulata, 323; nitidula, 323;
octomera, 323; phyllogena, 323; ru-
Source : MNHN, Paris
INDEX 501
fula, 323; tetramera, 323; sp., 253
Potentilla hookeriana, 127
Proteus mirabilis, 213 modifications
de structures par L-usnate
Protium heptaphyllum, 323
Prunus africana, 461 phorophyte
Pseudomonas aeruginosa, 213; sp., 213
Psora decipiens, 127; rubiformis, 127
Pulsatilla patens, 127
Pyrénolichens, 253
Pyrenula, 253, 461; nitida, 253
Pyrola asarifolia, 127; secunda, 127
Raciborskiella janeirensis, 323 Rio
Grande do Sul
Rapanea, 461 phorophyte
Rhododendron lapponicum, 127
Ribes triste, 127
Ricasolia sublaevis, 461 bas.
Rio Grande do Sul, 323 Zur Okologie
und Pflanzengeographte blattbe-
wohnender Flechten von -
Rosa acicularis, 127
Rwanda, 461 Contribution à l'étude
des lichens du Kivu (Zaïre); du
- et du Burundi VI. Les genres Coc-
cocarpia Pers. et Lobaria (Schreb.)
Hoffm.
Salix glauca, 127; pulchra, 127; reti-
culata, 127
Saxifraga oppositifolia, 127; tricus-
pidata, 127; tricuspidata - ass., 127
Schefflera, 461 phorophyte; myriantha,
461 phorophyte S
SCHELL S. und WINKLER S. - Zur Okologie
und Pflanzengeographie blattbewohnen-
der Flechten von Rio Grande do Sul
(Südbrasilien), 323
Sclerotina sclerotiorum, 253
Selaginella selaginoides, 127
Senecio hyperborealis, 127
SERUSIAUX E. - Contribution à l'étude
des lichens du Kivu (Zatre), du
Rwanda et du Burundi VI. Les genres
Coccocarpia Pers. et Lobaria
(Schreb.) Hoffm., 461
Sociologie, 127 synusies épigées dans
Те Yukon Territory; 323 Tichens foli-
coles du Rio Grande do Sul
Solarina spongiosa, 127
Sporopodium xantholeucum, 323 Rio Gran-
de do Sul
Squamarina lentigera, 127
Stereocaulon alpinum, 127; glareosum.
127; paschale, 127: tomentosum, 127
Strigula complanata, 323; elegans var.
antillarum, 323 Rio Grande do Sul,
var. elegans, 323; nemathora var.
nemathora, 323, var. hypothelia, 323;
nitidula, 323; subtilissima, 323
Substrate texture, 171 Growth rates of
Xanthoria parietina and their rela-
tionship to -
Symphonia globulifera, 461 phorophyte
Syzygium, 461 phorophyte; cordatum,
461; parvifolium, 461 phorophyte;
rowlandii, 461
Tapellaria epiphylla, 323; molleri,323
Rio Grande do Sul; nana, 323
Tapiria guianensis, 323
Taxons nouveaux, 127, 349
Tessmannia, 461 phorophyte
Thamnolia subuliformis, 127
Thamnolietum vermicularis, 127
Thamnolio - cladonietum amaurocraeae ,
127
Tillandsia usnoides, 323
Toninia coeruleonigricans, 127
Toninion coeruleonigricantis, 127
Trebouxia, 349
Tricharia albostrigosa, 323 Rio Grande
do Sul; carnea, 323; vulgaris,323
Trichothelium annulatum, 323; epiphyl-
lum, 323
Цараса, 461 phorophyte
L-usnate and permeability, 213
Vaccinio - piceetalia, 127
Vaccinium vitis-idaea, 127, ssp. minus,
27
Verrucaria, 461
VICENTE C. and CIFUENTES B. - L-usnate
and permeability, 213
Vriesea, 323
WINKLER S. , voir SCHELL S. and WINKLER
5$., 323.
Xanthoria parietina, 171 Growth rates
of - and their relationship to
substrate texture
Yukon Territory, 127 Epigaeous lichen
synusiae in -
Zygadenus elegans, 127
Source : MNHN, Paris
* солена
St
dig
boa a: e
ipei EE re
AL
TABLE DU TOME DEUX
ASPERGES M. — А new lichen species, Cladonia berghsonii Asperges sp. nova (Sect.
E 00 e Res Жу; MM d NU MEAS ANS. 349
FRAHM JP. — Ergänzungen zur Campylopus-Flora von Brasilien ....... DNE
GAMBARDELLA R., LIGRONE В. and CASTALDO В. — Ultrastructure of the sporo-
phyte foot in Phaeoceros. .
GIL ГА. y VARO J. — Estudio briosociologies de las comunidades reofilas de Sierra
Nevada (España) я . 423
GRIFFIN Ш D. — Spore ornamentation in Leiomela (Musc . 101
GRIFFIN Ш D. — El genero Sphagnum en los Andes de Colombia y Venezuela. Clave
para las especies frecuentes u ocasionales con notas ecologicas y taxonomicas
artramiaceae). .
HORTON Р.С. — The taxonomic status of Encalypta microstoma Bals. et De Not. and
E. ciliata var. microstoma Schimp. . ....... пы КАНА d At PONE
JANEX-FAVRE М.С. — Études ontogéniques chez le Porina byssophila (Pyrénolichens)
Il. - Les périthèces ........ Tm. ETC ЫЕ ACA ГӨ 253
KUMAR S.S. and VERMA S.K. — BE) observations on some west Himalayan
Mosses VI M US 2 б Оо 77
LEFORESTIER С. - Résistance à la sécheresse et Жын végétative chez
Fossombronia caespitiformis De Not. ........... A SE PERS
MANUEL М.С. — Synopsis of Rhizogoniaceae Broth. in Malaya... . . » Е: 449
МОХНАМ ТН. — Growth rates of Xanthoria ын and their relationship to
substrate texture . Se E кас em VT eot
NIMIS P.L. — Epigaeous lichen synusiae in the Yukon Territory . . ..... ocean
ODU E.A. — Reproductive phenology of some tropical african mosses .......... 91
OLARINMOYE 5.0. ex os and gemma development in Hyophila crenulata
C. Muell. ex Dus. . DIE EE AN qe E 457
PRIVITERA M. et LO GIUDICE R.— ber e muscinale della pos del bosco
di Malabotta (Peloritani) . m E B ИЕ
SCHELL S. und WINKLER S.— Zur Ökologie und hien ehe blattbewoh-
nender Flechten von Rio Grande do Sul (Südbrasilien) . . .. . . - К
SCHUMACKER R., LECOINTE A., TOUFFET J., DE ZUTTERE Ph., LECLERCQ L.
et FABRI В. — Hyocomium armoricum (Brid.) Wijk et Marg. en Belgique et dans le
nord-ouest de la France (Ardenne, ed Normandie) Etude chorologique,
écologique et phytosociologique . WA PME 277
SÉRUSIAUX Е. — Contribution à l'étude des lichens du Kivu (Zaïre), du Rwanda et
du Burundi VI. Les genres Coccocarpia Pers. et Lobaria (Schreb.) Нот... . . . . 461
Source : MNHN, Paris
TIXIER Р. — La notion d'espèce chez le genre Cololejeunea. Le complexe Cololejeunea
floccosa (Lehm. et Lindenb.) Schiffn. ................ sce NET
UDAR В. апа AWASTHI US. — A new species of Lejeunea from India.......... 345
VICENTE C. and CIFUENTES B. — L-usnate and permeability ...... se] en nn. 213
ZANDER В.Н. — Description and illustrations of Barbula, Pseudocrossidium and
Bryoerythrophyllum (p.p.) of Mexico ...... AW m EN AU “зек еб и
ZANDER Е.Н. — Didymodon (Pottiaceae) in Mexico and California : taxonomy and
nomenclature of discontinuous and nondiscontinuoustaxa . .............. 379
NOTES
DE SLOOVER J.L. — Octoblepharum erectifolium Mitt. ex Williams et Steereobryon
subulirostrum (Schimp. ex Besch.) G.L. Smith en Guadeloupe . . ++... Mone e PARA
DHIEN В. — Florule bryologique des rochers bathoniens . . .......... a 233
Lnformationsr ДИЛ E E E 126, 359, 473
Bibliographie bryologique . ++...
Bibliographie lichénologique. . . .
LOC SERRE i
ses... 107, 235, 360, 474
+ 119, 247, 372, 483
„Commission paritaire 15-12-1981 № 58611.
Dépôt légal n° 11124 - Imprimerie de Montligeon
Sorti des presses le 20 décembre 1981,
Source : MNHN. Paris
COLLOQUE INTERNATIONAL
du CNRS № 258
ÉCHANGES IONIQUES TRANSMEMBRANAIRES
CHEZ LES VÉGÉTAUX
TRANSMEMBRANE IONIC EXCHANGES IN PLANTS
org. : G. Ducet, В. Heller, M. Thellier
Universités de Rouen et Paris VII - 5-11 juillet 1976
@ analyse des modèles théoriques @ recherche des couplages métaboliques ou autres
® études électrophysiologiques @ cas particulier des transferts d'anions et de molécules
organiques @ localisation d'ions et aspects structuraux et moléculaires @ intervention
d'échanges ioniques dans les régulations intercellulaires
kinetic and thermodynamic considerations, model systems
metabolic and other couplings, ATPases
particular features of anionic transfers
electrophysiology of the ionic transfer
absorption of organic molécules
localization, molecular and structural aspect of the transfers
interference of the transmembrane transfers in other processes than absorption
ion exchanges in cell organites
(69 communications dont 64 en anglais et 5 en français)
21х29, 7 - 608 pages - broché 180 Е
286 fig. - 89 tabl. - 30 phot.
ISBN 2-222-02021-2
(co-édition CNRS-Université de Rouen)
ditions du CNRS
5 quai Anatole France. 75700 Paris
CCP Paris 9061-11 - Tél. 555.92.25
chez son libraire
à défaut aux Éditions du CNRS (chèque joint) 8
profession et demande votre documentation
О Sciences humaines
adresse. О Sciences exactes et naturelles
O Trésor de la langue Francaise
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Source : MNHN, Paris
18 JAN. 196€
2 1 JAN. 1982
SOMMAIRE
R.H. ZANDER. — Didymodon (Pottiaceae) in Mexico and California :
Taxonomy and nomenclature of discontinuous and nondiscontinuous
ou Der M TIN RE TP 379
J.A. GIL & J. VARO. — Estudio Briosociologico de las comunidades
reofilas de Sierra Nevada (España) ........... Е a: 423
J.P. FRAHM. — Ergänzungen zur Campylopus-Flora von Brasilien ..... 441
М.С. MANUEL. — Synopsis of Rhizogoniaceae Broth. in Malaya...... 449
S.O. OLARINMOYE. — Regeneration and gemma development in
Hyophila crenulata C. Muell. ех Dus........................ 457
E. SÉRUSIAUX. - Contribution à l'étude des lichens du Kivu (Zaire),
du Rwanda et du Burundi VI. Les genres Coccocarpia Pers. et Lobaria
icit iei LANE Та AR кзн 461
NOTE
J.L. DE SLOOVER. — Octoblepharum erectifolium Mitt. ex Williams
et Stereobryon subulirostrum (Schimp. ex Besch.) G.L. Smith en
Guadeloupe . 47
INFORMATIONS . . . 473
BIBLIOGRAPHIE BRYOLOGIQUE 474
BIBLIOGRAPHIE LICHÉNOLOGIQUE . 483
MEX о ees E (uses DERE US 491
Source MNHN. Pari: