BRYOLOGIE-LICHÉNOLOGIE

ANCIENNE REVUE BRYOLOGIQUE ET LICHÉNOLOGIQUE

Fondée par T. HUSNOT en 1874

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Bryologie: J. BERTHIER, J.L. DE SLOOVER, P. GEISSLER, S.R. GRADSTEIN, JP.

HÉBRARD, S. JOVET-AST, D. LAMY, М.С. NOAILLES, C. SUIRE.

Lichénologie: J. ASTA, T. BERNARD, B. BODO, W.L. CULBERSON, М.С, JANEX-

FAVRE, J. LAMBINON, M.A. LETROUIT-GALINOU, Cl. ROUX.

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Source : MNHN. Paris

CRYPTOGAMIE

BRYOLOGIE

LICHENOLOGIE

TOME 10 Fascicule 3 1989

Publié avec le concours du Muséum National d'Histoire Naturelle

Source : MNHN, Paris

Cryptogamie, Bryol. 1989, 10 (3): 189-233 189

ASCOMATAL DEVELOPMENT IN LICHENS:

А REVIEW

М.А. LETROUIT-GALINOU* & A. BELLEMERE**

* Laboratoire de Cryptogamie B t 50 6ème EL,

Université Pierre et Marie Curie, 4 place Jussieu,

F-15252 Paris Cedex 05 France

** Laboratoire de Mycologie-Lichénologie, Ecole Normale Supérieure de

Lyon,

Centre de Saint-Cloud, Grille d'honneur Parc de Saint-Cloud,

F-92211 Saint-Cloud.Cedex, France.

ABSTRACT - From some examples, four of which being studied with details, а compar-

alive study of ascomatal development in Lichens has been attempted. Three fundamental

stages are distinguished: primordium stage, "ébauche" stage, and stage with parathecial for-

mations. The structural components of ascomata are analysed as well as the different

growth modalities during the ontogenetical stages. The opening of the fruiting bodies and

some peculiarities in the development are then studied. ‘The different types of adult asco-

mata are compared and the main ontogenetical types of ascomata are succinctly described.

The systematical value of ascoma development is, at last, briefly discussed

RESUME - A partir d'exemples, dont quatre sont étudiés en détail, on a tenté de faire une

étude comparée du développement de l'ascoma chez les Lichens. Trois stades fondamen-

taux sont distingués (le stade primordium, le stade ébauche et le stade avec formations

parathéciales). Les éléments structuraux des ascomata sont analysés ainsi que les modalites

de croissance au cours des différents stades du développement. Aprés l'étude de l'ouverture

des fructifications et de quelques particularités du développement, les différents types

d'ascocarpes adultes sont compares. Les principaux types ontogeniques d'ascomata sont

ensuite sommairement décrits. La valeur systématique de l'étude du développement des

ascomata est brièvement discutée.

Since a long time, authors who wished to clarify relations between

Ascolichens, Ascomycetes and Algae, have studied morphology and ontogeny of

ascomata : TULASNE (1852), KRABBE (1882, 1891), REINKE (1895),

BAUR (1898, 1901, 1904), WOLFF (1905), NIENBURG (1908), MOREAU &

MOREAU (1928). After years with no further interest, new researches started,

first with DOPPELBAUR (1959 and 1960), then with LETROUIT-GALINOU

(since 1960), HENSSEN (since 1963) and their students, specially JAHNS (since

1970), JANEX-FAVRE (since 1965), KEUCK (1977, 1979). Besides recent

reviews on the subject by LETROUIT-GALINOU (1973, HENSSEN &

JAHNS (1974), HENSSEN (1976, 1981), PARGUEY-LEDUC &

Source : MNHN. Paris

190 М.А. LETROUIT-GALINOU and A.

JANEX-FAVRE (1981), general considerations are also included in KEUCK

(1977) and SIPMAN (1983) who attempt to harmonize concepts and to clarify

terminology. Of peculiar interest are also the synthetic considerations developed

recently by CHADEFAUD (1982 a,b,c), concerning ascomatal structure and

evolution.

The present paper attempts to a comparative analysis of datas relative to

structure and development of the sterile elements in lichen ascomata.

'oncerning the fertile elements (ascogonial apparatus, sporophytic apparatus,

asci and ascospores), a review has been recently published (BELLEMERE &

LETROUIT, 1988), so these will not be specially considered here.

Morphological and structural terminology will be first briefly summarized in

this paper, then four examples of ascoma development will be developed before

variations of ontogenetical events are considered and concluding remarks

exposed.

1- SHORT RECALL OF SOME MORPHOLOGICAL ANI

CHARAC! OF THE ASCOMA.

Ascomata are usually found on the upper surface of the symbiotic thallus;

they can exceptionally form on the purely fungal hypothallus, for example іп

some Rhizocarpon. Generally, they are either scattered all over the thallus or

grouped at its center. Sometimes, they are purely marginal (Cetraria, Peltigera).

In Nephroma, the mature ascomata seem located at the under surface of the

thallus, but, in fa they form at its margin and turn downwards only later,

because the upper side of the ascoma, of thalline aspect, has a more important

growth. In some genera (Cladonia, Stereocaulon, Baeomyces), ascomata are

born on special formations erected on the thallus (- podetions).

PRUCTURAL

There are two majors morphological types of ascomata: apothecia and

perithecia.

Apothecia have a large exposed hymenial surface, usually circular, the

hymenial dise, surrounded by an excipular margin (Fig. 1 A). Their diameter,

which is often about Imm, may be smaller or larger but rarely reaches up lem.

Apothecia are light, bright- or dark-coloured, with sometimes differences

between the disc and the excipulum. They can be immersed in the thallus (e.g.

Aspicilia) or superficial. Frequently they have a short stipe, narrower than the

disc. Some lichens have apothecia born on podetions. These thallus-like expan-

sions, which are feebly elongated in Baeomyces, can reach several centimeters in

Cladoniaceae or Stereocaulaceae.

Lirellae, important variant of apothecia, are lengthened ascomata, frequently

dark, opening by a longitudinal split; they may be branched (Y- or star-shaped).

They are either superficial or immersed in the thallus.

Perithecia are always small, their diameter exceptionnally reaching up Imm.

They appear as black dots on the thallus in which they are generally immersed

(Fig. 1 B). At maturity they are flask-shaped with an internal cavity; they usually

open by a summital pore (ostiole). Only rarely, they are aggregated in stroma-

like structures (Trypetheliaceae, Asterothyriaceae).

Source : MNHN, Paris

ASCOMATAL DEVELOPMENT IN LICHENS 191

Fig. L- Major elements of lichen ascomata. - A- Apothecium: the margin at the right is a

typical amphithecium sensu CORNER (1929-1930) without algae, but at the left, it is

figured as an amphithecium of thalline appearance including algae. - B- Perithecium.

(Abbreviations: see p. 233)

Ascomata whose characters are intermediate between apothecia and

perithecia, are frequently named “perithecioid apothecia” (e.g. Lichina, some

Pertusariaceae and Thelotremataceae).

In ascomata, tħe hymenium is a regular layer of asci, which also generally

comprises vegetative interascal filaments more or less erected. Those, habitually

named paraphyses, can however be diverse in their ontogeny. When the

summital part of the hymenium is well differentiated, it forms an epithecium.

Besides the hymenium, two others main parts have to be distinguished in

ascomata: one, beneath the hymenium, contains the sporophytic elements (or

their remnants) and сап be named the subhymenium s. L5; the other, the

excipulum, devoided of sporophytic elements, envelops the hymenium and the

subhymenium.

The upper part of the subhymenium s. 1. is paraphysogenous; it contains the

terminal branches of the ascogonial hyphae. Generally well delimited and easily

The “subhymenium 817 was originally simply named "subhymenium" by

LETROUIT-GALINOU (1967) who was the first to distinguish it from the excipular

components by the presence of sporophytic elements.

Source - MNHN, Paris

192 М.А. LETROUIT-GALINOU and A. BELLEMÉRE

recognizable, it will be known here as the subhymenium s. s.*. The lower part of

the subhymenium s.l will be defined as the hypothecium*; its structure, at its

base, often gradually passes to that of the underneath excipulum, so the limit

between these two formations is sometimes difficult to distinguish. For some

authors (e. g. SIPMAN, 1983) this transitional zone is known as the

subhypothecium. It must be noticed that in structural descriptions with pure

systematic purposes, authors frequently use the term hypothecium to describe the

whole underneath part of the ascoma, located under the subhymenium s. s. ,

making no distinction between its excipular and non-excipular parts.

According to species, the excipulum differs by its colour (dark or bright),

thickness, aspect (thallus-like or not), texture (fleshy or carbonaceous) and struc-

ture (more or less complex). Lecanorine apothecia have an excipulum with a

thalline appearance (Fig. 1 Cj). Lecideine apothecia have an excipulum which is

structurally clearly distinct from the thallus (Fig. 1 C,) and often reminds the

hymenial disc; when bright-coloured, apothecia are said biatorine. Іп zeorine

apothecia, the margin shows two parts, an outer thalline margin of thalline

structure and an inner proper margin whose structure differs from that of the

thallus (Fig. 1 Cy). Ontogenetically, the excipulum is in fact usually composed of

several components, some being early formed (ie. the floor), other lately

developed (parathecium and amphithecium). It will be shown later that parts of

the excipulum with similar aspect can differ by their origin.

After developmental studies, several terms, previously used to characterize

parts of the excipulum from a purely morphological point of view, have now got

a modified sense, for instance, “amphithecium”, "periphyses", "subhymenium",

“hypothecium” and "podium". So, the term "amphithecium", which was initially

applied by lichenologists to any marginal elements of thalline aspect (— thalline

margin) and the term “parathecium’, which was employed to designate the non-

thalline elements of the excipulum proper margin), are now used in a

restricted ontogenetical sense defined by CORNER (1929 a, b; 1930 a, b, c) (see

p. 195 & 213). Consequently, in this paper, the term "amphithecium” will be

applied to structural components which may be thallus-like or not (cf. Fig. 1 A).

The term "periphyses” has long been used to define short filaments which, in

perithecia, may be present in the ostiolar canal and also under the ceiling of the

ascomatal cavity; this term must be now restricted to the sole ostiolar filaments

(see p. 197 & p. 200).

* For some former authors, the subhymenium s. s. was the “thecogenous layer", the

hypothecium was the “subhymenium”, the elements underneath their "subhymenium" being

the "hypothecium". Some contemporary authors reduce the term “subhymenium” to the

subhymenium s.s.

Source : MNHN, Paris

ASCOMATAL DEVELOPMENT IN LICHENS 193

ІІД)

Mtr

ШЕ

Fig. 2.- Xanthoria parietina: ascoma development schematized after JANEX-FAVRE &

IBRAHIM-GHALEB (1986) (A to F at the same scale). A- Very young ébauche with

ascogonial apparatus, В- Ebauche with circumcentral zone and sporophytic apparatus;

С- Older ébauche with parathecioid net and parathecial crown; D- Young ascoma still

included in thallus with remnants of the roof over the ascomatal cavity and the

developing hymenium; E- Opening of the ascoma and development of the

amphithecium; F- Mature apothecium with typical parathecial apparatus; G-Detail of

the margin of F; H- Habitus: 1, young stages (A to E); 2, adult apothecium.

(Abbreviations, see p. 233).

Source - MNHN. Paris.

194 М.А. LETROUIT-GALINOU and А. BELLEMÉRE

11.- EXAMPLES OF ASCOMA DEVELOPMENT.*

A - XANTHORIA PARIETINA

(Fig. 2) (after JANEX-FAVRE & IBRAHIM-GHALEB, 1986 )

1.- The youngest stage which has been observed, more or less spherical (0 =

5Ошт), is located at the base of the thallus algal layer (Fig. 2 A). Its vegetative

part is already differentiated into two superposed areas: а plexiform

carpocentrum, with rather upright filaments, covered by a roof, with more or less

horizontal hyphae. Ascogonial filaments, with enlarged siderophilic cells, form a

knob in the basal part of the carpocentrum. This stage, where the vegetative

clements are not all similar but appear differentiated into two parts, will be

defined as the ébauche** stage.

2.- At a next stage (Fig. 2 B), the older ébauche (Ø = 100 um) is somewhat

piriform because a plexiform growing zone develops at its margin (circumcentral

zone). Simultaneously, the roof becomes more or less desintegrated by the

enlarging carpocentrum, Numerous sporophytic hyphae arise from the ascogonial

cells.

3. Later (Fig. 2 С), the top of the ébauche reaches the base of the thallus

cortex. Moreover there is a lateral extension of the ascoma, probably due to the

activity of the circumcentral zone. Now the carpocentrum has got divided in two

superposed parts. The upper one, the paraphysoid net, is made of hyphae more

or less upright and parallel: the paraphysoids. The lower part, meniscus-like,

(basal meniscus), has a plexiform structure and contains the sporophytic elements

in which ascogenous hyphae, with croziers, are now distinct. The basal meniscus

begins to produce new arising free-ended filaments, the paraphyses (=

carpocentral paraphyses or primary paraphyses, sensu LETROUIT-GALINOU,

1966). A new formation, the parathecial crown, organizes; issued from the top of

the circumcentral zone, it is constituted by short free-ended diverging hyphae

whose peculiar disposition results from a sympodial mode of growth. `

4.- The older ascoma (Ø = 150um ) (Fig. 2 D), remains covered by the

thallus cortex. Under some thin remnants of its roof, the paraphysoid filaments

have ruptured and an ascomatal cavity develops, being limited at its base by the

paraphysis tops. Arising over the laterally extending basal meniscus, the

paraphyses are now numerous; they get parallely arranged in a definite layer in

which young asci grow up: the hymenium is now edificated; so the basal

meniscus has to be named subhymenium 5. 1. (the upper part of which, with

ascogenous hyphae, being the subhymenium s.s.). At the periphery of the ascoma,

the subhymenium extends laterally producing hyphae with a subradial dispo

sition. These hyphae are named amphithecioid hyphae (sce BELLEMER

1967). In Xanthoria parietina, these hyphae, intermingled with algae, get a

thallus-like structure and form a verruca around the ascoma. At the margin of

* SANTESSON (1984) is used here as reference for species names.

* "ébauche" is a french word used to define the aspect of a work (as a sculpture for ins-

tance) when this is at its beginning, its shape being still unachieved. This term has been

prefered to “primary corpus” (LETROUIT-GALINOU, 1968) which may be easily

confused, erronously, with "primordium".

Source : MNHN, Paris

ASCOMATAL DEVELOPMENT IN LICHENS 195

the ascoma, the parathecial crown develops basipetally but feebly, its base

possibly contributing to the lateral extension of the subhymenium.

5.- The thallus cortex and the remnants of the roof now rupture over the

enlarging ascoma; so the ascomatal cavity opens at its top (Fig. 2 E ). Lateraly

the sympodial growth of the parathecial crown clearly increases: a typical

parathecial apparatus will now build (Fig. 2 F & G ). On its internal face, a few

parathecial paraphyses develop upwards in a basipetal manner contributing to

the lateral extension of the hymenium (and also of the subhymenium). On its

external face, the parathecial apparatus generates amphithecial hyphae which

compose an amphithecium sensu CORNER (cf. p. 192). The bases of the

parathecial paraphyses and those of the amphithecial hyphae diverge from the

parathecium (sensu CORNER, cf. p. 192); this is a thin sheath made of

tangentially oriented hyphae, which lines the subhymenium outside. The

amphithecium, in Xanthoria parietina, gets а thallus-like structure ( with distinct

cortex, algal layer and medulla ); it contributes to the extension of the verruca

which encloses the ascoma and which now projects over the thallus. This verruca

has a composite origin; whilst its reduced upper part is made of amphithecial

hyphae, its flank is made of amphithecioid hyphae and its basal part of thalline

hyphae. Nevertheless, the whole verruca has a thalline-like structure.

6.- Later the parathecial apparatus becomes very important and largely

contributes to the widening of the ascoma. At the end, the hymenium is well

exposed ( Fig. 2 F & G): the ascoma is an apothecium. In this, numerous asci

have formed on the ascogenous hyphae which extend at the upper part of the

subhymenium.

The main characteristics of ascoma development in Xanthoria parietina can

be summarized as follows:

a) The very young ascoma early differentiates into an ébauche.

b) In the young ébauche, the well-developed carpocentrum is not entirely

surrounded by sterile tissues. These are reduced to a transitory roof: the

pericentral envelope* is not complete. Around the carpocentrum, a circumcentral

zone with a plexiform structure is distinct; it has a growing function.

c) In the older stages, the roof hardly modifies ; the carpocentrum

differentiates into two superposed parts: the transitory paraphysoid net, which

early makes room to the ascomatal cavity, and the subhymenial meniscus which

generates paraphyses (carpocentral paraphyses) Some amphithecioid hyphae

form on the external face of the subhymenial meniscus; they take a part in the

formation of the verruca which surrounds the young ascoma.

d) A parathecial crown, early distinct above the circumcentral zone, develops

in a parathecial apparatus when the ascoma begins to open. This parathecial

apparatus plays a major role in the enlargement of the maturing apothecium; its

amphithecium has a thalline-like structure.

* The term "pericentral envelope" is used here preferentially to the anterior term "primary

envelope” (LET ROUIT-GALINOU, 1968) because it gives a better idea of the location and

formation of this envelope and also because it avoids the ambiguity tied to the qualificative

"primary".

Source : MNHN, Paris

196 М.А. LETROUIT-GALINOU and A. BELLEMERE

Fig. 3.- Pyrenula nitida: ascoma development schematized after JANEX-FAVRE (1971) (А

lo F at the same scale). A- Young primordium. B- Young ébauche. C- Carpocentrum

differentiation . D- Paraphyses and periphyses development. E- Young perithecium. F-

Mature perithecium. G- Habitus. (Abbreviations, see p. 233).

B.- PYRENULA NITIDA (Fig. 3)

(after JANEX- FAVRE,1971 and HENSSEN & JAHNS, 1974).

1.- Ascoma development begins, in the thallus, at the base of the algal layer,

by the formation of a knob of hyphae with a distinct plexiform aspect. This

plexus of homogenous structure is a primordium still devoided of ascogonial

elements (Fig. 3 А).

2.- The young ascoma widens and soon reaches the ébauche stage, two parts

being now distinct in the ascoma: the carpocentrum and the roof (Fig. 3 B). In

Ше carpocentrum, the plexiform structure is maintained but a developing

ascogonial apparatus is now present. The roof, which covers the carpocentrum,

is made of tightened hyphae with thick darkening cell walls. It extends laterally

and downwards around the carpocentrum and, at its top, bears a cylindrical

bundle of erected filaments, the epicentral bundle which reaches the thallus surfa-

Source : MNHN. Paris

ASCOMATAL DEVELOPMENT IN LICHENS 197

ce. At this stage, the roof is the only element of the pericentral envelope to be

present.

3.- The ébauche extends downwards into the thallus. At its base, under the

carpocentrum, a clear and thin pale floor made of tangential hyphae develops

constituting a second element of the pericentral envelope (Fig. 3 C). Three

superposed parts are now distinct in the carpocentrum: its plexiform basal part,

or basal meniscus, contains numerous sporophytic elements ; its middle part is a

paraphysoid net; its conical upper part (apical point) penetrates upwards into the

roof and the overtopping epicentral bundle, and reaches the exterior,

4.- The ascoma, still growing downwards (Fig. 3 D), becomes now flask-

shaped; it has a narrow neck and an enlarged bulge with a flat base. Laterally,

between the dark roof and the clear floor, a short and thin muff, made of clear

tangentially disposed hyphae, differentiates; this circumcentral muff, completes the

pericentral envelope. In the carpocentrum, the paraphysoid net ruptures near its

top: the ascomatal cavity appears. The basal meniscus generates upright the first

paraphyses so the hymenium, still devoided of asci, begins to form. The apical

point, inside the darkened epicentral bundle, now shows two regions; the inferior

one, at the level of the neck base, remains more or less plexiform; the superior

one lyses axially; a narrow ostiolar canal is delimitated, open at its top. This ca-

nal is soon lined by short free-tipped filaments turned upwards, the periphyses;

the lower ones are generated on the remnants of the apical point whilst. the

superior ones are directly born on the epicentral bundle.

5.- Later, the ascoma becomes higher and broader, its base remaining pale

and flat (Fig. 3 E). Its growth, which essentially takes place in the bulge part оГ

the ascoma, seems due to the activity of the clear circumcentral muff.

Upgrowing asci born on the ascogenous hyphae infiltrate the numerous

paraphyses: the hymenium is now typical and the underlaying basal meniscus

can be named subhymenial meniscus. At the base of the ascomatal neck, the

remnants of the inferior part of the apical point vanishes; so, the ostiolar canal,

which was open at its top, also communicates now with the ascomatal cavity.

6.- The fully developed ascoma (Fig. 3 F) is a perithecium opening by an

ostiole at the top of an ostiolar canal. Its neck із relatively short and its enlarged

bulge is more or less rounded, The ascomatal cavity is quite entirely filled by the

hymenium, with numerous paraphyses and mature asci. The subhymenium is

reduced to a clear layer containing the ascogenous hyphae. The sterile part of

the ascoma, the excipulum, now entirely dark, has thickened partly by adjonction

of thalline elements; these are especially noticeable at the neck level. Structurally

the excipulum wall is complex. Besides its thalline elements, it is made of

components of diverse origin: the epicentral bundle is a dependance of the roof,

the floor and the hardly distinct circumcentral muff belong to the pericentral

envelope; moreover, a thin clear sheath of tangentially disposed hyphae has

developed inside the lateral part of the excipulum wall at the hymenium level.

This sheath, probably built of peripherical carpocentral remnants, has been

named “secondary envelope" (JANEX- FAVRE,1971; see p. 209); it possibly

produces paraphyses. The periphyses, which are other components of the

excipulum, have also different origin; most of them are born on the carpocentral

apical point, some on the epicentral bundle and others on the thalline envelope.

Source - ММНМ, Paris

198 М.А. LETROUIT-GALINOU and А. BELLEMÉRE

Major traits of the ascoma development in Pyrenula nitida differs from

Xanthoria parietina because:

a) a primordial stage is encountered;

b) the ébauche has a more complicated structure. The carpocentrum (well

developed) shows an apical point. The primary envelope is complete with a well-

developed floor, a reduced muff and an important roof overtopped by an

epicentral bundle;

c) the mature ascoma, remaining included in the thallus, is a perithecium with

an ostiolar canal lined by numerous periphyses;

d) at the mature stage, ébauche structures are persisting and there are neither

amphithecioid nor parathecial elements.

Despite the above differences, there are however some similar characteristics

in the ascomatal development of Pyrenula nitida and Xanthoria parietina. So, in

both species:

a) the young ascoma forms into the thallus under the algal layer;

b) there is an ébauche stage with a plexiform carpocentrum and a pericentral

envelope;

c) the carpocentrum differenciates into а paraphysoid net and a basal

meniscus; this has an important marginal extension;

d) all the paraphyses are generated by the basal meniscus;

€) an ascomatal cavity is formed by the rupture of the paraphysoid net's

upper part.

C.- ENDOCARPON PUSILLUM (Fig. 4)

(after HENSSEN & JAHNS, 1974 and WAGNER, 1987).

L- The first stage which has been observed (Fig. 4 A), is a primordium

located in the algal layer, underneath the thallus upper cortex. It is a

homogenous knob of coalescent filaments whose cells are short and wide. Its

basal part contains an ascogonial apparatus made of large siderophilic cells from

which trichogynes escape, reaching the upper surface of the thallus. Small algal

cells are present іп the primordium.

2.- Later on, when the enlarged young ascoma becomes piriform (Fig. 4 B), it

is made of three parts and so has reached an advanced ébauche stage. Its

summital part, surrounding the degenerating trichogynes, is a conical newly

differentiated area made of upright coalescent filaments and ori

neighbouring thalline hyphae; this area is probably an epicentral bundle because

it is distinctly separated from the carpocentrum by a narrow plate of

subtangential hyphae which probably represents a poorly differentiated roof. The

median part of the ascoma, or carpocentrum, derived from the initial knob, is

now made of thin filaments; it contains the ascogonial cells. The basal part is

interpreted here as the floor because its cells differ from the surrounding thallus

by their structure and disposition. So, the pericentral envelope is not well

characterized, being rather reduced and incomplete (only floor and roof).

Source : MNHN, Paris

ASCOMATAL DEVELOPMENT IN LICHENS 199

"ШҮ s MERI ee

Fig. 4.- Endocarpon pusillum: ascoma development schematized after WAGNER (1987) (А

lo F at the same scale). A- Primordium. B- Young ébauche. C- Older ébauche. D-

Opening of the young ascoma, E- Ascoma with young asci F- Mature ascoma. G-

Habitus: 1, thalline squamulae on the soil; 2, thallus isolated from the soil.

(Abbreviations, see p. 233).

Source : MNHN. Paris

200 М.А. LETROUIT-GALINOU and А. BELLEMÉRE

3.- The broadened ébauche (Fig. 4 C), which has elongated both downwards

and upwards, now reaches the thallus surface. An ascomatal cavity filled with

jelly and containing a few small algal cells, has formed into the carpocentrum

separating it in two superposed parts which remain in contact only by their

margin. The lower part of the carpocentrum, (basal meniscus), more or less

lenticular, has a plexiform structure and contains fertile elements. The bell-

shaped upper part of the carpocentrum (summital bell) generates centrifugally

free-tipped filaments under its inferior face (descending filaments); over its

superior face (where the roof is no longer recognizable), an apical point of

carpocentral origin develops axially in the enlarged epicentral bundle. Around

this one, the surrounding thalline hyphae dispose regularly into a peripherical

supernumerary envelope (thalline envelope) which extends basipetally.

4.- Important modifications of the ascoma structure occur at the next stage

(Fig. 4 D). The enlarged ascomatal cavity gets piriform. The basal meniscus

which bears no paraphyses, has now a conical shape. Under it, the previously

mentioned floor is no longer distinct. Above the cavity, the summital bell and the

overtopping apical point have ruptured axially and so an ostiolar canal forms

which soon opens at its top. On the inferior face of the ruptured summital bell,

descending filaments develop, overhanging the ascomatal cavity. Around the

ostiolar canal, a thin sheath of rearranged hyphae produces periphyses. The

lowest ones are born on the upper part of the ruptured summital bell; those,

above, are formed on the apical point; near the top of the canal, they form on

the epicentral bundle and, at the very top, on the thalline envelope. This one has

grown downwards, but does not reach the ascomatal base.

5.- The ascoma, whose growth has been very important (Fig. 4 E), now

becomes entirely surrounded by the thalline envelope which darkens at its upper

part. Laterally, at the place where the summital bell and the basal meniscus are

fusing, a short muff (circumcentral muff) has differentiated; it does not bear any

filament. The basal meniscus, which has become cupular, does not generate

any sterile interascal filaments. Тһе ascogenous cells, present only in the dense

upper part of the basal meniscus, produce asci into the ascomatal cavity. Those

arrange into a hymenium and so the basal meniscus must now be named

subhymenium and the summital bell, subhymenial bell. Owing to their hyaline as-

pect and their similar structure, the subhymenial bell, the circumcentral muff and

the lower part of the subhymenium constitute a distinct clear peripherical sheath

inside the enlarged thalline envelope.

6.- Тһе adult ascoma is a perithecium (Fig. 4 F). In the bulge, the important

ascomatal cavity still contains a lot of small hymenial algal cells. At the base оГ

the cavity, the non-exposed hymenium remains devoided of paraphyses. At the

top of the cavity, the descending filaments have псагіу all disappeared. In the

neck, the ostiolar canal is coated by numerous periphyses whose superior ones

become dark. The excipulum is formed by the dark thalline envelope, now quite

complete, and which, inside, has incorporated the sheath of carpocentral origin,

observed at the preceding stage.

In Endocarpon pusillum and Pyrenula nitida, ascomata are perithecia with an

entirely dark excipulum. In both, the neck, whose components are of various

origin, is perforated by an ostiolar canal bearing periphyses. Both, also, are

devoided of parathecial elements, oppositingly to the apothecia of Xanthoria

Source : MNHN, Paris

ASCOMATAL DEVELOPMENT IN LICHENS 201

Fig. 5.- Thelotrema lepadinum: азсота development schematized after

LETROUIT-GALINOU (1967), HENSSEN & JAHNS (1974), HENSSEN (1976) (A

16 E at the same scale). A- Young ébauche. В- Ebauche with fully developed ascogonial

apparatus and trichogynes. C- Fully developed ébauche with paraphyses, lateral wall

and descending filaments. Note its early opening. D- Young apothecium. E- Mature

apothecium. F- Habitus. (Abbreviations, see p. 233).

parietina. Nevertheless, Endocarpon differs fundamentally from Pyremula by the

absence of paraphyses and by the presence of a sushymenial Бей producing

descending filaments. Moreover, іп Endocarpon, the primordium is made of

Source : MNHN. Paris

202 М.А. LETROUIT-GALINOU and А. BELLEMERE

short and wide cells while it is made of intermingled thin filaments in Pyrenula.

In Endocarpon, the dark bottle-shaped excipulum results from the important

development of the thalline envelope; in Pyrenula, this misses, the excipulum

differentiating progressively from the pericentral envelope. During the

development of the two species, a clear sheath exists at some stage on the

internal face of the excipulum, but it is different by its origin: in Endocarpon, it is

formed by the carpocentral elements surrounding the cavity and it is transitory,

but, in Pyrenula, it is a lately formed secondary envelope which persists in the

adult ascoma.

D.- THELOTREMA

(after LEFROUI

1976).

1.- The earliest stage which has been observed is a young ébauche (Fig. 5 А),

located under the algal layer of the thallus between the suberified cells of the

cork. This ébauche shows three superposed zones, all of carpocentral origin, no

pericentral envelope being distinct. The lower zone is a basal meniscus with a

plexiform structure containing a few ascogonial elements; the central zone, more

important, is a paraphysoid net; the upper zone, formed by coalescent hyphae, is

a summital bell.

EPADINUM (Fig. 5)

ALINOU,1966; HENSSEN & JAHNS, 1974; HENSSEN,

2. In the increased ébauche (Fig. 5 B), hyphae are now tightened in the

lateral part of the young ascoma, so the flank of an envelope of carpocentral

origin (carpocentral envelope) is differentiating. Ascogonial elements, more or less

erected into the paraphysoid net, are now numerous; they bear many trichogynes

which, at the top of the ébauche, escape through the summital Бей and point

out over the thallus.

3.- The older ébauche (Fig. 5 C), remains sunken into the thallus, but opens

at its top, rupturing the thallus cortex. Major changes have occurred in the

ébauche structure. The carpocentral envelope is no longer distinct, making room

to a narrow split which now separates the young ascoma from the thallus. The

subhymenial bell has extended downwards; its new-formed part, made of close

appressed parallel hyphae tangentially disposed, now constitutes the ascomatal

wall. On its internal face, this is lined with short filaments, obliquely oriented

upwards, their length progressively increasing from the base to the top of the

ascoma. Owing to their origin and despite of their orientation, these filaments are

nding filaments. The paraphysoid net is no longer visible; at its place,

upright paraphyses have developed, generated by the basal meniscus in which a

few sporophytic elements are present. These paraphyses, produced centrifugally,

are shorter and shorter towards the margin, forming a conical hymenium, In a

small area at the base of the ascoma, between the ascomatal wall and the

subhymenial meniscus, a peculiar structure appears made, inside, of parallel

hyphae disposed similarly to those of a circumcentral muff and showing, outside,

a plexiform disposition; it constitutes the circumcentral zone.

4.- During the ulterior growth (Fig. 5 D), the ascoma continues to push up

the surrounding thallus; its form and structure are feebly modified. The

hymenium, where young asci develop, widens consecutively to a marginal

growth. So does the subhymenium, whose base, however, remains flat. The

ascomatal wall extends above the short circumcentral muff which bears no ra-

Source : MNHN, Paris

ASCOMATAL DEVELOPMENT IN LICHENS 203

сх

—

qum

toe m

v TY QN

osmm

Fig. 6. Pelügera rufescens: ascoma development schematized after

LETROUIT-GALINOU & LALLEMANT (1971), HENSSEN 4 JAHNS (1974),

HENSSEN (1976) (A to C at the same Scale) A- Sterile thallus margin. B-

Primordium. C- Ebauche. D- Opening of the hemi-angiocarpic apothecium. E-

Habitus. (Abbreviations, see p. 233).

mification. In its basal part, this grows downwards; at its top, it grows upwards

and generates filaments whose length shortens upwards, oppositingly to the

underlying descending filaments whose length shortens downwards. As the

ascomatal wall is of carpocentral origin, its newly formed top may be

considered as an apical point; consequently the filaments that it produces, have to

be interpreted as periphyses. So an ostiolar apparatus is present; this is entirely of

carpocentral origin (PARGUEY-LEDUC, 1966,1967; JANEX- FAVRE,1971).

5.- The mature ascoma, with a widened and largely exposed hymenium (Fig.

5 EL is an apothecium. The numerous paraphyses have now nearly all the same

length, except the marginal ones, so the hymenium has a trapezoidal section. The

surrounding thallus, increased in a prominent wart, is separated from the

ascomatal wall by the narrow split precedingly mentioned; this has got deeper

and more distinct. The apothecium is enlarged by lateral extension of its base,

probably due to the generative activity of the persisting circumcentral muff.

The mature ascomata of Thelotrema lepadinum and Xanthoria parietina, are

apothecia, but have a strongly different structure. First, in Thelotrema lepadinum,

there is no parathecial apparatus. Secondly, the apothecial development exhibits

perithecial characteristics, such as the formation of ап ostiolar apparatus with

periphyses and the development of descending filaments under the sushymenial

Source : MNHN, Paris

204 М.А. LETROUIT-GALINOU and A. BELLEMERE

bell; however, oppositingly to what happens in typical perithecia, those elements

do not overhang the hymenium, but are pushed laterally on its flank. Thirdly,

during the apothecial development of Thelotrema lepadinum, a paraphysoid net,

paraphyses, periphyses and descending filaments are formed; in the three

previous examples, none of these structural components were encountered

altogether.

Finally, only one character is in common in the development of the four

studied examples: the presence of a clearly recognizable ébauche. The importan-

ce of the concept of "ébauche" in studies of ascoma development is thus

confirmed.

HI- VARIATIONS IN ASCOMA DEVELOPMENT.

The study of the four preceding examples shows that diverse developmental

schedules, but also common features exist in lichen ascomatal development.

Enlarging the comparisons 19 other species, considerations will now be developed

in attempt to point out the major developmental trends of the sterile components

of the ascomata.

Successively will be considered:

A. Variations during the ascoma edification, i. e. the variations of the structural

elements, of the modes of growth, in the opening of the ascomata.

B.- Main characteristics of the mature ascomata.

C.- Main ontogenetical types of ascomata.

A.- VARIATIONS DURING THE ASCOMA EDIFICATION.

1.- Variations of the structural components.

The preceding examples have shown that during the development of the

sterile elements of the ascoma, several structurally defined main stages can be

observed. These are:

a) the primordium stage, in which the sterile elements have an uniform structure;

b) the ébauche stage, in which these components begin to differentiate in several

parts;

c) the parathecial stage in which secondary elements with a sympodial mode of

growth develop, forming a parathecial apparatus;

d) the mature stage, in. which no new type of structural element appears (even if

the growth goes on).

a) The primordium stage and its structural variations.

The primordium generally forms in the algal layer of the thallus or close to it,

either over (Parmelia) or under it ( Lecania sulfureofusca, SIPMAN, 1983).

exceptionally, it develops on the thallus ( Lichinodium, HENSSEN, 1981) or in

small outgrowths of thallus lobes (Wavea, HENSSEN & KANTVILAS, 1985).

Source : MNHN, Paris

205

ASCOMATAL DEVELOPMENT IN LICH

- A to D. Cladonia floerkeana: ascoma development schematized after

ROUIT-GALINOU (1967); see also KRABBE (1891), JAHNS (19702), JAHNS &

BELTMAN (1973) (A to C at the same scale). А- Primordium of podetion. B- Young

podetion with young apical ascoma. C-Mature ascoma at the top of a podetion. D-

Habitus. E to 1. Ваеотусез rufus; ascoma development schematized after

LETROUIT-GALINOU (1967); see also JAHNS (1970) (E to Н at the same scale).

Primordium. Е- Ebauche with carpocentrum. G- Young apothecium with

developing hymenium and subhymenium. H- Mature apothecium at the top of a

podium. l- Habitus. (Abbreviations, see p. 233).

The primordium сап be distinguished from the neighbouring thallus by some

histological characters: in most cases, it is a plexus of vegetative hyphae

(=generative tissue, HENSSEN and JAHNS, 1974) usually surrounding several

ascogonial coils ( Endocarpon pusillum, Fig. 4 A). It may also differ from the

thallus by cytochemical characters (such as gelified walls or cytoplasm reactivity).

The primordium generally contains no alga; sometimes, it is the only character

which distinguishes the primordium from the thallus as in Peltigera (Fig. 6 B), in

Rhizocarpon and in Lichinaceae.

In some Lichens, mature ascomata are gathered in specialized areas of the

thallus. In Laurera (JOHNSON, 1940; GALINOU, 1957), Enterographa and

Chiodecton, these areas result from a late coalescence of maturing ascoma

elements and are not true stromas preexisting to the primordium formation.

Oppositingly to non-lichenized Ascomycetes, primordia of Lichens have never

been encountered in, or over, stromas. The defined pulverulent areas on which

the groups of primordia of Phlyctis are born (LETROUIT-GALINOU, 1967)

have not been usually considered as stromatic. The opinion of some authors

Source : MNHN. Paris

206 M.A. LETROUIT-GALINOU and A. BELLEMERE

(VAINIO,1894; CHADEFAUD, 1984) considering the whole organized lichen

thallus as a stroma per se can be also mentioned.

Generally the primordium has a plexiform structure; not rarely its hyphae

have an arbuscular disposition ( Cladonia, Fig. 7А). Only exceptionally, the

primordium shows the plectenchymatous structure common to many non-

lichenized Ascomycetes (Arthopyrenia lapponina and A. submicans, JANEX-

FAVRE,I971; Steinera glaucella, HENSSEN & JAMES, 1982). Sometimes it

has been observed, for instance in the genus Collema (BAUR, 1898; HENSSEN,

1981), that the vegetative filaments of the primordium are distinctly issued from

the foot cells of ascogonial coils.

In the primordium, the sterile filaments generally exist before the ascogonial

elements are distinct. It happens however that these are the first to appear (c.g. in

Physciaceae and Collemataceae). Eventually, the primordium remains devoided

of ascogonial coils, these forming only later, at the ébauche stage, eg. in

Pyrenula (Fig. 3 A), Thelotrema (Fig. 5 B), some Gyalectaceae (Fig. 9 A) and

Physcia tenella (ОТТ, 1987).

In some cases, an ascogonial apparatus develop in a preexisting functional

pycnidium: then, no typical primordium is present and the resulting ascoma is

known as руспосагр (eg. Phyllisciella, HENSSEN, 1963). New primordia can

also appear іп ageing ascomata, eg. іп Lecanora subfuscata

(LETROUIT-GALINOU, ` 1967, Baeomyces roseus (JAHNS &

SMITTENBERG, 1970) or some Caloplaca (MALONE, 1977).

Generally, the primordium soon differentiates into an ébauche. However, in

some cases, the structure of the primordium persists unchanged till the ascoma is

mature (e. g. Arthonia; cf. HENSSEN & JAHNS, 1974).

Тһе biology of primordia is badly known. Exceptionnaly, authors mentioned

that primordia are long-lived ( Megalospora, SIPMAN, 1983). The physiological

conditions of the primordium transformation into an ébauche are unknown; in

some cases (Collema, BAUR, 1901; Cladonia, HONEGGER, 1984) the role of

spermatization seems clear.

b) The ébauche stage and its structural variations.

^ complex differentiation of the sterile elements of the ascoma takes place at

the ébauche stage. It consists, on one hand, in the formation of differentiated

concentric structures: essentially a carpocentrum surrounded by an external

pericentral envelope. Into each of the concentric constitutive elements, a

dorsi-ventralization also happens resulting in a storied disposition with distinction

of a basal, a median and а tectal parts. So, the basal meniscus, the paraphysoid

net, the sushymenial bell individualize in the carpocentrum, while in the primary

envelope, the floor, the circumcentral muff and the roof, with eventually its

epicentral bundle, form. Moreover, a bilateral symetry may also appear

(primordium of lirellac).

« Variations in the carpocentrum.

At the beginning, the carpocentrum, central part of the ébauche in which are

located the ascogonial elements, has the same structure as the primordium.

Its. differentiation in three superposed parts is usually precocious.

Source : MNHN, Paris

ASCOMATAL DEVELOPMENT IN LICHENS 207

* 'The paraphysoid net proceeds from the median part of the carpocentrum

(c. Fig. 2 A & B, 3 C, 5 A), and is devoided of sporophytic elements. It is

generally unique but, in rare cases, several paraphysoid nets may differentiate

side by side into the carpocentrum, each of them being later replaced by a dis-

tinct hymenium (e.g. in Pertusaria pertusa, Fig. 11 K & L). No paraphysoid net

forms in the Verrucariaceae where an ascomatal cavity early appears at its place

by lysis.

Sometimes, the structure of the paraphysoid net persists unchanged until the

ascoma opens ( Opegrapha (Fig. 10 А-В), Phiyctis, Gyalectidium). Frequently,

the paraphysoid net ruptures below the overtopping summital bell, either early

(Lecidella) or late ( Lobaria) and a more or less developed ascomatal cavity

results. Then, the remnants of the paraphysoid net may vanish or persist in the

hymenium as a part of the interascal filaments.

Fig. 8.- A to E. Ramalina ecklonii: ascoma development schematized after KEUCK (1979)

(A to D at the same scale). A- Primordium. B- Ebauche. C- Very young apothecium.

D- Mature apothecium. Habitus. - F to K. Parmelia conspersa: ascoma

development schematized after L ETROUIT-GALINOU (1970); see also P. exasperata

in HENSSEN & JAHNS (1974), HENSSEN (1981) (F to J at the same scale). F-

Primordium. G- Developing husk and disappearing carpocentrum. H- Secondary

carpocentrum. l- Growing apothecium with upper sterile filaments. J- Mature

apothecium. K- Habitus. (Abbreviations, see p. 233).

Source : MNHN. Paris

208 М.А. LETROUIT-GALINOU and А. BE

H Ch vumm

Fig. 9- Gyalecta carneolutea: аѕсота development _schematized after

LETROUIT-GALINOU (1974); see also HENSSEN & JAHNS (1974), HENSSEN

(1976) for others Gyalectaceae (A to G at the same scale). A- Young ébauche with

irichogynes. B- Carpocentrum differentiation: paraphysoid net and first paraphyses. C-

Ascomatal cavity and paraphyses. D- Descending filaments, Е- Young apothecium with

opening roof. F- Maturing apothecium with, at one side, a developing parathecial

crown. G- Mature apothecium. H- Habitus. (Abbreviations, see p. 233).

* The upper part of the carpocentrum, or summital bell, is generally thin. In

ébauche of perithecia, it may produce, upwards, the apical point and,

downwards, descending filaments as in Verrucariaceae (Fig.4 C). When present,

the apical point soon dissociates axially and an ostiolar canal results; at its top,

this opens out and, at its base, it communicates with the ascomatal cavity. This

canal can become lined by periphyses. In ébauche of apothecia, where the apical

point is unknown, descending filaments may exist (Fig. 5 C, 6 D, 9 D); however

the presence of a summital bell has not been mentioned under the roof.

Descending filaments usually vanish, either early, as іп Lasallia

(ANEX-FAVRE, 1973) or late, as in Gyalectaceae (Fig. 9 G) and

Peltigeraceae. They rarely persist; then they generally remain short, e.g. іп

Thelotrema (Fig. 5 E) and Diploschistes (HENSSEN & JAHNS, 1974);

exceptionally, however, they may extend downwards through the ascomatal

cavity and reach the basal meniscus; then they are known as pseudoparaphyses

{Arthopyrenia lapponina (= A. fallax), A. submicans; Fig. 12 А-В).

* The basal part of the carpocentrum or basal meniscus which contains the

sporophytic hyphae generated by the ascogonial cells, becomes the subhymenium

Source : MNHN, Paris

ASCOMATAL DEVELOPMENT IN LICHENS 209

ЇЇ

ПП

БШ li i

pude асарын a E

RANI

Fig. 10. A to C. Opegrapha rufescens: structure of а lirella schematized after

LETROUIT-GALINOU (1967) (A and В at the same scale). A- Extremity of a lirella

(longitudinal section). B- Transversal section of a mature part of a lirella. C- Habitus.

` D to F. Graphis scripta : structure of a lirella schematized after JANEX-FAVRE

(1964) (D and Е at the same scale). D- Extremity of a lirella (longitudinal section). E-

‘Transversal section of mature part of alirella. F- Habitus. (Abbreviations, see p. 233).

when the hymenium forms. In most cases, the basal meniscus produces upwards

free-tipped paraphyses; frequently the first paraphyses grow up into the

paraphysoid net whose ruptured elements may persist as paraphysoids amongst

the layer of paraphyses (Lobaria; LETROUIT-GALINOU, 1971). In several

families or genera (e. е. Verrucariaceae, Opegraphaceae, Pertusariaceae), the

basal meniscus does not generate paraphyses.

* Peripherical hyphae of the carpocentrum sometimes organise in a clear

sheath whose structure is distinct from the internal part of the carpocentrum and

also from the pericentral envelope. If this structure forms early, it is named the

carpocentral envelope (e.g. Verrucaria cazzae, JANEX-FAVRE,1971; BEL-

LEMERE & LETROUIT, 1988); if it forms lately, it is named the secondary

envelope (e.g. Pyrenula nitida, Fig. 3 F).

fi The pericentral envelope and its structural variations.

The pericentral envelope encloses the carpocentrum. Differing structurally

from this and from the surrounding thallus, it is always devoided of sporophytic

elements; rarely it contains a few algae.

‘The pericentral envelope is generally present and comprizes three superposed

parts: the roof, the median part of the envelope and the floor. It exceptionnaly

lacks, the ébauche being only made of the carpocentrum as in Lecanora

subfuscata (LETROUIT-GALINOU, 1967).

* The roof.

Source : MNHN, Paris

100 um J

Fig. A to E. Umbilicaria cylindrica schematized after HENSSEN (1970) &

JANEX-FAVRE (1974): development of secondary and tertiary lirelliform new

apothecia into an initial apothecium (А and D at the same scale). А- Ebauche with a

roof. B- Young apothecium. C- Secondary apothecia in a primary one. D- Secondary

and tertiary apothecia. E- Habitus. - F to H. Graphis elegans schematized after

JANEX-FAVRE (1965): successive development of new lirellae in the remnants of an

initial one (F and G at the same scale). Е- Ebauche of a new lirella into three successive

ones. G. Maturation of the new lirella of F. H. Habitus. - I and J. Superposed

apothecia successively developed in Cladia aggregata: only the last one is fertile

(schematized after JAHNS 19706). 1- Habitus. J- Section; note the successive sterile

hymenia. (hy, hy) and the fertile one (hy). - К to L. Composite apothecium in

Pertusarla pertusa schematized after LETROUIT-GALINOU (1967) K- New

hymenium developed in an old degenerated initial one. L- Habitus. - М to O. Thallus

reaction around an apothecium of Parmeliella coronata (schematized after HENSSEN,

1969). М- Young apothecium. N- Mature apothecium. O- Habitus. (Abbreviations,

see p. 233).

Source : MNHN, Paris

ASCOMATAL DEVELOPMENT IN LICHENS 211

КОШ

зн анди

mum

КҮШТІ

ЗО

BVT

Wel

Fig. 12. Some peculiar ontogenetical types in Lichens. A and В. Arthopyrenia lapponina

(= A. fallax) after JANEX- FAVRE, 1971. А- Ebauche stage. B- Adult stage. - С.

Lichina confinis, schematized after JANEX-FAVRE (1967). - D. Arthothelium sp.,

schematized after HENSSEN & JAHNS, 1974. - Е. Edwardiella mirabilis, drawn after

HENSSEN (1986). - F. Aspicilia calcarea, drawn after JANEX-FAVRE, 1985. - G.

Hertella after HENSSEN 1985. Gl- Hertella chilensis, young stage. G2- Hertella

subantarctica, adult stage. (Abbreviations, see p. 233).

Source : MNHN, Paris

212 М.А. LETROUIT-GALINOU and А. BELLEMERE

- In many Pyrenolichens, the roof, upper part of the pericentral envelope, is

generally well developed. Often, as in Pyrenula nitida (Fig. 3 B) and in

Endocarpon pusillum (Fig. 4 B), it grows upwards forming an epicentral bundle.

This, which incorporates surrounding thalline components, forms the main part

of the ébauche neck, around the ostiolar canal (which results from the disso-

ciation of the apical point) and is lined by periphyses.

- In the ébauche of Discolichens (lirellae inclusively), the roof lacks in some

genera of various families, е. g. Arthonia, Lecanora, Cladonia. When present,

the roof is generally clear and poorly developed. In some cases, it early stops

growing and then vanishes as in Xanthoria (Fig. 2). The roof is rarely well-

developed, being then cither light-coloured, as in Peltigera (Fig. 6) and Gyalecta

(Fig. 9) or dark, as in Graphis and Opegrapha (Fig. 10). The persisting roof rup-

tures early or late; then, it persists some time before to disappear, as in Peltigera

(Fig. 6 С); rarely, it is pushed laterally and remains distinct on the flank of the

mature ascoma (e. р. Graphis, Opegrapha, Vig. 10; Gyalecta, Fig. 9). No

epicentral bundle has been described in Discolichens. However, some formations

are perhaps to assimilate to an epicentral bundle, as, for instance, the "epicentral

filaments” of Baeomyces rufus (Fig. 7 F & С) or also as the parathecial crown

which, in some genera, precocely develops at the top of the young ébauche.

* The floor, basal part of the envelope, may form somewhat later than the

roof, as in Pyrenula (Fig. 3 C). It is sometimes lacking, as in Xanthoria parietina

(Fig. 2 B & C) or in Thelotrema lepadinum (Fig. 5 B). In various Graphidaceae,

Opegraphaceae (Fig. 10) and Pyrenolichens, the floor is reduced or remains

light-coloured whilst the roof is dark and well-developed (dimidiate ébauche). In

some cases, the floor develops strongly in a well distinct podium; this is

clearlight-coloured іп Baeomyces rufus, (Fig. 7 F to D, but it is black in

Opegrapha calcarea (CLAUZADE & ROUX, 1985).

* The median part of the pericentral envelope is casy to distinguish when it is

well-developed and looks like a muff (generally bearing no filaments). This muff

is rather long in Verrucaria cazzae (JANEX- FAVRE,1971) and in the sides of

lirellae (Fig. 10), but it is short in Pyrenula (Fig. 3 D). However, when a muff-

like structure exists in the lateral part of the ébauche, it has not always a

pericentral origin, but may correspond to a carpocentral or a secondary

envelope (see p. 209). Ontogenctical studies are therefore necessary to precisely

define the structural value of such a convergent type of structures.

In some cases, the median part of the pericentral envelope keeps the juvenile

and plexiform structure of the lateral part of the primordium; then it is generally

reduced (е. g. Xanthoria parietina, Fig. 2 С; Thelotrema lepadinum, Fig. 5 C),

but it is more developed in the extremities of young lirellae (Fig. 10 A).

Often the median part of the envelope is not distinct.

c) The parathecial stage and its structural variations.

Parathecial formations are not necessarily present. For instance, they do not

exist in the Arthoniales, Opegraphales and Graphidales whose mature ascomata

remain at the ébauche stage or even at the primordium stage (some Arthoniales).

Where they are present, these formations are characterized by a sympodial type

of ramification.

Source : ММНМ, Paris

ASCOMATAL DEVELOPMENT IN LICHENS 213

* At the beginning, they are always reduced to a parathecial crown of

divaricated hyphae, located at the margin of the ascoma. The parathecial crown

may appear at any ontogenctical stage of development. In some cases (е. g. in

Lecanora or in Megalospora), it forms сапу around the primordium. Usually it

develops at the ébauche stage, either early (e.g. Xanthoria parietina, Fig. 2 С),

or, more frequently, later, when the roof ruptures, as in Peltigera (Fig. 6 D)

where the ébauche diameter has reached more than Imm when the crown forms;

then, it appears at the periphery of the subhymenium (е. в. Gyalecta, Fig. 9;

ELLEMERE & LETROUIT-GALINOU, 1988; Lobaria

Diploicia canescens, BE

laetevirens, LETROUIT-GALINOU, 1971; HENSSEN, 1981).

* Rarely the parathecial formations remain reduced to a parathecial crown

(eg. Gyalectaceae, Fig. 9 G). They generally develop, either early or with some

delay ( Xanthoria parietina, Fig. 2 Û to G), into a parathecial apparatus. This

apparatus is formed by a thin sheath of compact and tangentially oriented

hyphae, the parathecium, with, inside, an increased production of paraphyses

(parathecial paraphyses) and, outside, the formation of an amphithecium sensu

CORNER (1929 a & b; 1930 a, b & c) (cf. p. 192; see also SIPMAN 1983).

When the parathecial apparatus is specially developed, a strong lateral amplifi-

cation of the ascoma results as in many Lecanorales s. 1. (e.g. Xanthoria

parietina, Fig. 2 F; Ramalina, Fig. 8 D; Umbilicaria, Fig. 11 C; Parmeliella, Fig.

11 N). The presence of a parathecial apparatus is generally easy to state because

the amphithecial hyphae have a typical sympodial disposition and differ by their

aspect from the thalline hyphae (Le. the lecideine apothecia). But it happens that

the amphithecium looks like the surrounding thallus and eventually contains

algae, as in lecanorine apothecia (DUGHI 1955) or in immersed ascomata of

Peltigera (Fig. 6 D) or Nephroma (LETROUIT-GALINOU & LALLEMANT,

1970). Meanwhile, in such cases, the parathecial origin of the apothecial margin

may be discerned because, at least in its upper part, the sympodial disposition of

the hyphae is always clearly distinct (Fig. 2 G).

d) Additional structural components.

Growing reactions may take place in the thallus at some distance around the

base of the young ascoma, and may generate additional structures. For instance,

in Thelotrema (Fig. 5 E ) or Parmeliella (Fig. 11 N ), a cupular formation

develops around the ascoma: in Peltigera polydactyla, the finger-like part of the

thallus bearing the ascoma results from a stimulation of thallus growth located

under the apothecium (LETROUIT-GALINOU & LALLEMANT, 1970;

JAHNS & FREY, 1982).

2- Modes of growth in the sterile components of ascomata.

a) Growth during the primordium stage.

In the plexiform primordium, growth proceeds by adjonction of elements

similar to the preexisting ones; those originates cither by intercalary elongation or

by ramification of the preexisting primordial filaments, or also by assimilation of

Source : MNHN, Paris

214 М.А. LETROUIT-GALINOU and А. BELLEMÉRE

the neighbouring thalline hyphae. When there is an equal and diffuse growth,

the shape of the primordium remains unchanged as, for instance, in primordia

of perithecia; when the growth is lateraly more important, the primordium

becomes discoid. If the renforced marginal growth is limited to some points, the

young ascoma turns lirelliform (Fig. 10 C and F, Fig. 11 H) or asteriform. In the

primordia of Cladonia and Baeomyces, horizontal growth is relatively feeble but

vertical growth becomes important: a small podetion forms (Fi

b) Growth during the ébauche stage.

In some cases, the mode of growth of the primordium is maintained during

the ébauche stage. More frequently, new areas with peculiar growth differentiate

in the ébauche. Generally, these areas only concern a part of the preexisting

constitutive structural elements. Each of these areas shows eventually a special

modality of growth. Usually, the growth is engaged, either in a perithecial or in

an apothecial way, as soon as the ébauche stage.

* When the ébauche will become a perithecium, a diametral enlargement

affects essentially the bulge part whilst an usually noticeable elongation takes pla-

ce in the upper part of the ébauche (future neck region); so the young ascoma

becomes flask-shaped. In Pyrenula (Fig. 3 F) the enlargement of the bulge takes

place in the subhymenium and is directed upwards; in Endocarpon (Fig. 4 F) it

takes place in the margin of the sushymenial bell and is directed downwards. A

part of the bulge enlargement may also be due to intercalary growth in the

circumcentral muff (old ébauches of Verrucaria cazzae, JANEX- FAVRE,1971).

* When the ébauche will become an apothecium, the enlargement is essentially

located at the periphery of the ébauche (marginal growth) but a dilfuse

intercalary growth also exists; sometimes, all the structural components of the

ébauche have an equal marginal growth; for instance, in Peltigera Fig. 6), the

paraphysogenous basal meniscus, the paraphysoid net and the roof are similarly

affected. In most cases, the extension of each structural component of the

ébauche differs; so іп Lobaria, the widening of the roof early stops, the

paraphysoid net feebly grows on laterally while the floor and the meniscus

strongly extend becoming concave (Fig. 13 A); in Diploicia canescens, (cf.

BELLEMERE & LETROUIT, 1988), the roof and the paraphysoid net do not

enlarge (and quickly vanish); only the basal meniscus and the floor extend, their

newly formed parts being known as the "proparathecium" (LETROUIT-

GALINOU, 1967).

* The lengthening of the ébauche of a lirella (Fig. 10) is due to the persistance

at its extremities of the type of growth established in the primordial stage.

Oppositingly, on the flank of the lirella, the restricted diametral growth is

probably consecutive to the differentiation of the muff (Fig. 10).

* Ebauches of perithecioid apothecia, show simultaneously apothecial and

perithecial types of growth, the modalities differing with species. For instance, in

Thelotrema (Fig. 5), the enlargement of the ébauche at its base is as important as

in an apothecium, but at its upper part there is a strong vertical elongation as in

a young perithecium. In Lichina (HENSSEN, 1963; JANEX-FAVRE, 1967), the

roof does not develop (as it generally does in a young apothecium) but the lateral

extension of the meniscus and of the floor is reduced (as in a perithecium).

Source : MNHN, Paris

ASCOMATAL DEV

¿LOPMENT IN LICHENS 215

Fig. 13.- Comparison of ascoma and conidioma ébauches in Lobaria laetevirens schematized

after LETROUIT-GALINOU (1971, 1972). А- Ascoma ébauche. B- Conidioma

ébauche. (Abbreviations, see p. 233).

* Sometimes, certain parts of the ébauche have a peculiar growth. For instan-

ce, in Baeomyces (Fig. 7), the meniscus and the floor thicken strongly and a

podium results. In Verrucaria cazzae, a special and important growth of the

upper part of the neck occurs and an involucrum develops (cf. BELLEMERE &

LETROUIT, 1988).

с) Growth during the parathecial stage.

When the sympodial mode of growth characterizing the secondary for-

mations of ascomata is important, with formation of a complete parathecial

apparatus, there is a marginal growth which can lead to a strong diametral en-

largement of the ascoma (с. в. Xanthoria, Parmelia, Usnea). Oppositingly, when

only a parathecial crown develops, the marginal sympodial growth is reduced;

then it is the type of growth established in the ébauche which is the major cause

of the enlargement of the maturating ascoma (Gyalecta, Fig. 9; Pachyphiale

cornea, LETROUIT-GALINOU, 1977).

d) Special localized growths.

New vegetative growth points, with a restricted importance, sometimes appear

externally on thallus-like amphithecium; they may produce different types оГ

marginal hyphae ( as ciliae in Usnea or as excipular irregularities in Lecanora

allophana).

3 - Variations in the modalities of opening in ascomata.

According to species, the opening of the ascoma takes place at different stages

of the development.

Source : MNHN, Paris

216 M.A. LETROUIT-GALINOU and А. BELLEMÉRE

* When no roof develops in the young ascoma (eg. Cladonia (Fig. 7 А),

Lecanora, Arthonia), or if the roof vanishes early, before the beginning of the

hymenium formation (eg. Nephroma, Lecidella, Ramalina, Fig. 8 B-C), the

development of the ascoma is said gymnocarpic. One must keep in mind that

gymnocarpy is independant of an eventual persistance of thalline elements over

the developing ascoma.

* If the roof ruptures after the hymenium is constituted, but before it contains

mature asci, the ascoma is said hemiangiocarpic, e.g. Peltigera (Fig. 6), Gyalecta

(Fig. 9), Stereocaulon (WOLFF, 1905; JAHNS, 1970а), Diploicia

(LÉTROUIT-GALINOU, 1967).

* The ascoma development is said angiocarpie if the roof lately persists, till

many mature asci are present in the hymenium. This is the case of perithecia

(eg. Pyrenula, Fig. 3 and Endocarpon, Vig. 4); these generally open by an

ostiole formed at the top of the neck; when, rarely, perithecia open by a simple

hole, then there is no neck ( Arthopyrenia halodytes, JANEX-FAVRE, 1971).

Lirellae (e.g, Opegrapha, Graphis, Vig. 10) are also angiocarpic: they open by a

longitudinal split which is apparently devoided of some special dehiscence

mechanism, oppositingly to some non-lichenized Discomycetes (Lophodermium).

Angiocarpic apothecia are apparently very rare іп Lichens; they are however

encountered in several non-lichenized Discomycetes ( Phacidium).

In some cases, it may be difficult to clearly assign the development to a

gymno-, a hemiangio- or ап angiocarpic type. For instance in Phlyctis and

Roccella (LETROLIT-GALINOL, 1967), as in Dirina (FEHLER, 1983) and in

Lecanactis (MENSSEN & JAHNS, 1974), there is а persisting roof over the

hymenium as in angiocarpic types, but its peculiar structure allows spore release

through it.

4. Repeated development of ascomata.

In some lichens, new apothecia may form on, or in developing ones according

to different processes. For instance, in Graphis elegans (Fig. 11 F & G), the

repeated lirellae develop from new ébauches which successively differentiate

inside the preceding ones, probably in areas where the sporophytic apparatus is

locally reactivated. In Umbilicariaceae (Fig.ll A to D), the new secondarily

developed apothecia also form around reactivited parts of the sporophytic

apparatus, but, in this case, the initial apothecium has developed from a

complete ébauche whilst the secondary apothecia (which become lirelliform) are,

from the beginning, only constituted of parathecial structures. In Pertusaria

pertusa, new ascomata, reduced to a more or less extended hymenium, here and

there differentiate into old degenerated hymenia (Fig.ll K). Clearly, such

ascomata are ontogenetically distinct from those which, in the same genus, result

from the formation of several paraphysoid nets in one single ébauche (sce p.

208). In Cladia, with storied apothecia these are devoided of asci except the last

one (Fig. 11 1 & J).

Source : MNHN, Paris

ASCOMATAL DEVELOPMENT IN LICHENS 217

B. VARIATIONS IN THE ADULT STAGE.

1. The realization of the adult ascomata.

The ascoma is adult when its sterile components have got their definitive

structure; no new structural elements being added later; however the ascomatal

growth can still go on for some time.

The developmental stage at which the ascoma reaches its mature structure

differs according to species. For instance, the mature ascoma keeps the structure

of a primordium in Arthonia (Fig. 12D); it has the structure of an ébauche in

Thelotrema (Fig. 5), Opegrapha (Fig. 10), Graphis (Fig. 10), Pyrenula (Fig. 3); it

has that of a stage with a marginal parathecial crown in the Gyalectaceae (Fig.

9) or in some Lichina (HENSSEN & BUDEL, 1984); it reaches a stage with a

developed parathecial apparatus in Xanthoria (Fig. 11) and many Lecanorales s.

1. (LETROUIT-GALINOU, 1967; HENSSEN & JAHNS, 1974).

Frequently structural components can have disappeared at the adult stage

whatever the stage they individualize in the ascoma development, Examples are:

the roof in Xanthoria parietina, (Fig. 2 А to E); Peltigera, (Fig. 6); Gyalecta,

(Fig. 9 A to Е) or the descending filaments of the lateral part of the envelope іп

Endocarpon (Fig. 4 C to E) or in some Gyalectaceae (Fig. 9 D to F).

2. Structural differences in the adult.

a) The excipulum.

Since a long time, morphological excipular characters (consistance, colour, as-

pect of hyphal walls), which largely differ according to species, have been used

іп systematics. Later, structural characters have been considered at a great

extend. However, one must keep in mind that, structurally, the excipulum may

be composed of several elements which have differentiated at diverse stages of

the development. Those are: the pericentral envelope (and also the carpocentral

envelope or the late secondary envelope), the amphithecioid hyphae (generally

appeared at the ébauche stage), the amphithecium and the parathecium (which

form at the parathecial stage). Morcover, thalline clements may also add to the

excipulum at any stage in the development. This diversity and complexity of the

structural composition of the excipulum, can be illustrated with four examples.

For instance, in Endocarpon (Fig. 4), the excipulum essentially comprizes the

thalline envelope and the remnants of the epicentral bundle. In the Graphidaceae

and the Opegraphaceae (Fig. 10), the excipulum is made of the persisting

pericentral envelope, laterally thickened with black amphithecioid hyphae. In

Diploicia canescens (cf. BELLEMERE & LETROUIT-GALINOU, 1988), the

excipulum is composed by remnants of the pericentral envelope, amphithecioid

hyphae and a complete parathecial apparatus with a black amphithecium. In

Xanthoria parietina (Fig. 2 F), the purely parathecial excipulum, is made of a

thin parathecium and of a thick amphithecium of thalline aspect. Owing to this

Source : MNHN, Paris

218 M.A. LETROUIT-GALINOU and А. BELLEMÉRE

complexity, the excipulum structure has to be used in systematics only after

precise ontogenctical studies.

Thin external hyphae are sometimes produced downwards by the excipulum.

They may plunge into the underneath thallus and eventually reach the

substratum. These hyphae which can be associated in bundles, contribute to

anchor the ascoma and perhaps also to feed it (e.g. Collemataceae, HENSSEN,

1981; Megalosporaceae, SIPMAN,1983). As already mentioned, the excipulum

may also produce ciliae or have an irregular outline.

b) The subhymenium s.l. .

The subhymenium s. L may remain thin, as in Руғелша (Fig. 3) and

Thelotrema (Fig. 5), but is generally relatively thick. Its upper part (the

subhymenium s.s.) contains the ascogenous hyphae and is thin. Its lower part, the

hypothecium, more or less developed, is specially thick in most Lecanorales s.l.

where it progressively incorporates the paraphysis bases (LETROUIT-

GALINOU, 1967). The limit between the hypothecium and the floor of the

primary envelope is often difficult to discern, specially when both are darkened.

So one must be cautious in using hypothecial anatomy as a systematical

character.

c) The hymenium.

Interascal filaments are generally numerous; rarely there are only a few

(Phyllisciella, HENSSEN & BÜDEL, 1984); sometimes, they completely lack as

іп Endocarpon (Fig.4 F) and the Verrucariaceae.

* There are several types of interascal filaments according to their origi

- paraphyses (i.e. erected hyphae) are produced, either by the basal meniscus

of the ébauche (primary paraphyses: LETROUIT-GALINOU, 1961) or by the

inner part of the parathecium (secondary paraphyses*: LETROUIT-GALINOU,

1961);

- paraphysoids result of the rupture, near their top, of the constitutive elements

of the paraphysoid net ( Opegrapha, Fig. 10 A & B; Roccella,

LETROUIT-GALINOU, 1967; Dirina, FEHLER, 1983; Lecanactis, HENSSEN

& JAHNS, 1974);

- residual persisting hyphae of the primordium, only slightly streched by the

growth of the asci, become interascal filaments in some rare cases, е. 5.

Arthonia (HENSSEN & JAHNS, 1974);

- vegetative thallus hyphae, merely modified in their wall aspect and pigmen-

tation rarely become interascal filaments and only in thallinocarpic ascomata (е.

в. Rhizocarpon, HONEGGER, 1978);

- pseudoparaphyses (i.e. elongated descending filaments) exceptionally form in

Lichens JANEX- FAVRE, 1971);

* This term has been also used later to qualify the free-tipped filaments lately formed in

pycnocarps (HENSSEN, 1963)

Source : MNHN, Paris

ASCOMATAL DEVELOPMENT IN LICHENS 219

- conidiophores, functional or not, may become the initial interascal filaments

of ascomata resulting from pycnidial transformation (pycnocarps) Such

conidiophores have been sometimes ambiguously named "primary paraphyses”

(HENSSEN, 1963);

Several types of interascal filaments may coexist in one single ascoma (some

Graphidaceae).

* The aspect of the interascal filaments, which strongly differs іп Lichens,

results from their differentiation in the course of their development and is largely

independant of their origin. Frequently, interascal filaments are very thin and

slender, Sometimes, they аге stiff, with a thick wall of cartilaginous appearance

(c.g. Aspicilia, ROUX, 1977). They may be simple or branched or variously

anastomosed. In Lichina (HENSSEN et al., 1985), they are moliniform at their

top. Conidiogenous рагарһувев have been exceptionally mentioned

(Thelotrema lepadinum, LETROUIT-GALINOU, 1967). In Lichens, interascal

filaments are generally embedded in an abundant hymenial jelly which is often

iodine reactive and makes the hymenium compact (eg. Psora, TIMDAL,

1984b); it may eventually contain oil droplets (c.g. some Trypetheliaceae and

Mogalosporaceae). The upper part of the layer of interascal filaments is

frequently differentiated in an epithecium. This generally results from the

embedment of the modified tips of the filaments by rather dark pigments.

Sometimes, the epithecial layer only consists of a deposit of substances (e.g.

calcium oxalate, lichen phenols) over the top of the interascal filaments. In

Phiyctis, the epithecium is of a different origin; it results from the persistance

above the interascal filaments of cells pertaining to the ascomatal roof. The

reactivity of the epithecium to certain chemical agents is frequently used as a test

for diagnosis. Free tips of the interascal filaments overtopping the epithecium, or

presence of cristal deposits over the hymenium, may give a pruinose aspect to

the hymenial surface.

+ Modalities of growth are poorly known in interascal filaments. Their

elongation contributes to increase the thickness of the subhymenium rather than

that of the hymenium whose height generally remains unchanged during the ma-

turation of the ascoma. In the Caliciales, paraphyses сап have unlimited growth

(TIBELL, 1984).

Generally, asci begin to appear early, before the ascoma is adult ( Xanthoria,

Fig. 2 D). In some cases, however, the fully-developed ascoma remains long

devoided of mature asci, for instance in Lecidella elaeochroma (GALLOE 1927;

LETROUIT-GALINOU, 1967), Lecidea fusco-atra (HERTEL, 1977) and in

Hypocenomyce (TIMDAL, 1984a). Usually, asci are present in the whole

hymenium; eventually they may be absent in its axial part where persisting

tightened sterile filaments, similar to paraphysoids, form a pillar-like structure

(Thyrea rotunda, HENSSEN & BUDEL, 1985).

* Algae are present in the hymenium of a few genera ( Endocarpon, Fig. 4 F;

Staurothele), These hymenial algae are smaller than thallus algae; they however

belong to the same species, but exhibit characters of the non-symbiotic state

(WAGNER, 1984).

Source : MNHN, Paris

220 M.A. LETROUIT-GALINOU and А. BELLEMÉRE

3.- Structural characters of the main types of mature ascomata.

a) Perithecia.

Тһе excipulum of perithecia is generally entirely dark. It may lack or remain

pale at the base (dimidiate perithecia); it is entirely pale in Dermatocarpon. The

structural components of the perithecial excipulum differs according to specie:

eg. primary and secondary envelopes in Pyrenula nitida (Fig. 3), thalline

envelope in Endocarpon (Fig. 4). The neck surrounding the ostiolar canal of the

perithecia is more or less developed; it eventually extends іп an involucrum

which may be important ( Staurothele sapaudica, JANEX-FAVRI 1976;

Verrucaria cazzae, cf. BELLEMERE & LETROUIT, 1988). Periphyses line the

ostiolar canal: the upper ones are directly born on the epicentral bundle, the

lowest ones are produced on the ruptured part of the summital bell and on the

remnants of the apical point. In perithecia, interascal filaments are generally

paraphyses ( Pyrenula, Fig. 3); pseudoparaphyses are very rare ( Arthopyrenia

submicans, JANEX- FAVRE,1971); paraphysoids аге exceptionnaly present (e.g.

Trypetheliaceae, GALINOU, 1957). In pycnocarps, the interascal filaments may

be eventually persisting conidiophores. Interascal filaments are lacking in

Verrucariaces are generally relatively numerous in perithecia; in nearly

all cases, they are of the bitunicate-fissitunicate type even if the interascal fila-

ments are paraphyses.

b) Lirellae.

Тһе excipulum of the lirellae, generally black and well-developed, is made of

intermingled coalescent hyphae with thick and dark wall. The basal excipulum

may develop in a more or less important podium (ер. Opegrapha calcarea,

CLAUZADE & ROUX, 1985); it is missing in dimidiate lirellae (e. g-

Opegrapha saxatilis). Excipulum is quite reduced or entirely lacks in

Arthoniaceae. Interascal filaments, always present in lirellae of lichens, can be

either paraphyses (Graphis) ог paraphysoids (Opegrapha); interascal

pseudoparaphyses and descending filaments are unknown. Ап epithecium is

exceptionnaly developed which may be pruinose or eventually bright coloured

(Arthonia cinnabarina). Enchased lirellae are known in some species (e.g.

Graphis elegans, Fig.11 F to M).

©) Apothecia.

Excipulum of apothecia, gencrally fleshy, has rarely a carbonaceous texture.

In Gyalectaceae (Fig. 9 G), the excipulum is formed by the parathecial crown

and possibly by some remnants of the roof; in most Lecanorales, it is generally

a part of the parathecial apparatus. Thalline aspect of the excipulum may be due

either to a thalloid structure of the amphithecium ( Xanthoria, Fig. 2) or to an

additional thalline envelope ( Parmeliella diplomarginata, Fig. 11 N). Interascal

filaments are always present in apothecia; they are generally paraphyses; more

rarely they are thalline hyphae, as in thallinocarps, or paraphysoids as in

Aspicilia. When there is a distinct stipe, this is either a local swelling of the

thallus (some Lecanora) or the thickened floor of the primary envelope

(podium: eg. Baeomyces) or also а part of the hypothecium (some Lecidella).

Source : MNHN, Paris

ASCOMATAL DEVE

OPMENT IN LICHENS. 221

Although they are thallus-like, the podetions are ontogenetically a part of the

ascoma (except perhaps in Stereocaulaceae); in Cladoniaceae, they result of an

important vertical elongation of the whole primordium (Fig. 7 В); in

Bacomycetaceae, the podetions are thickened podia (Fig. 7 С).

d) Perithecioid apothecia.

Perithecioid apothecia are of two different types. One is encountered in

Thelotremataceae where the disc, large as in apothecia and made of paraphyses,

is surrounded by the sushymenial bell (the so-called ascomatal wall) and by a

persistent surplumbing apical point as in perithecia (Fig. 5). The other type of

perithecioid apothecia exists, for instance, in Lichina pygmaea (HENSSEN, 1963;

JANEX-FAVRE, 1967; HENSSEN et al., 1985) or in Pertusaria pertusa (Fig.11

K). There, the hymenium, which contains paraphysoids, is narrow, as it is in

perithecia, and is more or less enclosed in a well-developed circumcentral muff

(generating externally thalloid amphithecioid hyphae); however the disc is

exposed as in an apothecium.

С - ONTOGENETICAL TYPES OF ASCOMATA

From the combination of the precedingly described possible structural va-

riations and types of growth during ascomatal development, a great number of

ontogenetical types result. Some of these deserve special interest, either because

they are frequent or because they show some remarkable features. They will be

considered now.

1.- Types without any differentiation of a parathecial apparatus.

Most of these types have a perithecial type of mature ascoma, but some have

a perithecioid apothecium or an apothecium.

a) Types with perithecia.

EMERE & LETROUIT, 1988).

This type, illustrated in the present paper by Endocarpon pusillum (Fig. 4),

was first recognized by DOPPELBAUER (1959) and then confirmed by

JANEX-FAVRE (1971) and HENSSEN & JAHNS (1974). Its main characters

are: a plexiform primordium, the absence or quasi-absence of paraphysoids in

the ébauche, the existence of short descending filaments, the absence of

paraphyses (or, if present, lately and poorly developed). The adult perithecium

has a well developed ostiolar apparatus. The stage which precedes the formation

of asci is characteristic (verrucariacean stage) (WAGNER, 1987). Variations are

limited; they concern the constitution of the ascomatal wall, the importance of

the involucrum and also the eventual presence of paraphyses which has been

mentioned by some authors (Verrucaria (Amphoridium) ` calcicedum,

DOPPELBAUER, 1959; Verrucaria controversa, JANEX-FAVRE, 1970;

Thelidium ` immersum, DOPPELBAUER, 1959; Thrombium —aoristum,

а) The Verrucaria type (cf. BELL

Source : MNHN, Paris

222 М.А. LETROUIT-GALINOU and А. BEL!

CLAUZADE & ROUX, 1985). The Verrucaria type is encountered in a large

range of lichen genera, presently included in the Verrucariaceae; it was unknown

in non-lichenized Pyrenomycetes till it has been recently recognized in

Herpotrichiellaceae JANEX-FAVRE, 1988). Owing to the presence of a large

and early formed ascomatal cavity with short descending filaments and without

paraphyses, the Verrucaria-type has been often included in the Ascoloculares

(SANTESSON, 1950; JANEX-FAVRE, 1970). This however has been

questioned because the ascomatal cavity does not form by lysis and also because

the primordium has a plexiform structure, a character considered as

ascohymenial by HENSSEN & JAHNS (1974).

In Arthopyrenia lapponina (= A. fallax) and A. submicans (JANEX-

FAVRE,1971), the ascomatal ontogeny is a variant of the Verrucaria-type, but

rather distant owing to the non-plexiform structure of the primordium and to the

well-developed descending filaments becoming later interascal filaments. The

ébauche stage reminds that of the Microthyriaceae (Pleosporales) (Fig. 12 A-B).

P) The Pyrenula type (Fig. 3).

In this type, paraphysoids are well-developed in the ébauche and numerous

paraphyses are present at the adult stage. An internal and late developed

secondary envelope forms which lately is included in the excipulum. An ostiolar

apparatus may be present ог not. Some Porina, i.e. P. heterospora (= P.nucula

HENSSEN & JAHNS, 1974; HENSSEN, 1976) P. byssophila (JANEX-

FAVRE, 1981), some Acrocordia, i.e. А. conoidea (JANEX-FAVRE,1971) and

some Arthopyrenia, i.e. A. sublittoralis (JANE FAVRE,1971) are probably to

be included in this type. A few minor variations are known in the Pyrenula type

(cf. JANEX-FAVRE, 1971).

y) The Trypethelium type.

This type, also observed in Laurera, is probably to distinguish from the

Pyrenula type, because the interascal filaments are paraphysoids JOHNSON,

1940; GALINOU, 1957); however new precise developmental datas are needed.

b) Types with perithecioid apothecia.

а) The Pertusaria type (Fig. 11 К).

The primordium is plexiform. Paraphysoids are present in the ébauche, and

persist as interascal filaments after the roof has disappeared. The ascomatal wall,

lately formed, consists in a secondary circumcentral muff (cf. р. 212)ithe mature

ascoma, included in a thalloid verruca, is more or less perithecioid (KRABBE,

1882; LETROUIT-GALINOU, 1967; HENSSEN & JAHNS, 1974).

В) The Lichina type (Fig. 12 C).

In this type, similar to the Pertusaria type, some periphysoid filaments form a

reduced ostiolar apparatus, reminding that of typical perithecial types. АШ the

Lichinaceae are not of this type (HENSSEN, 1963; HENSSEN et al., 1985).

y) The Thelotrema type (Fig. 5).

Numerous paraphyses are present. The ascomatal wall, built from the sus-

hymenial bell of the ébauche (Fig. 5) bears descending filaments and an ostiolar

Source - MNHN, Paris

ASCOMATAL DEVELOPMENT IN LICHENS 223

apparatus with periphyses is present at its top. This type, characteristic of the

Thelotremataceae (g. Thelotrema, Diploschistes, Ocellularia), reminds one of

the ascomatal development in the major genera of the Stictidaceae, a mostly

non-lichenized family.

c) Types with apothecia (or lirellae).

x) The thallinocarp type.

In thallinocarps, there is no differentiated primordium, the asci form in cer-

tain parts of the thallus where hyphae get slightly differentiated, becoming the

interascal filaments. This type seems to exist in some Lichinaceae as Edwardiella

(Fig. 12 E) (HENSSEN, 1986) and Lichinodium (HENSSEN, 1963, 1981), but is

in doubt in Lichina (sec above). Rhizocarpon has been mentioned to belong to

this type (HONEGGER, 1978), but the mature ascomata, with an apparent

margin, have probably а more complicated structure than thallinocarps.

В) The Arthonia type (Fig. 12 D).

In this type (Arthonia, Arthothelium ...), the hyphae of the plexiform

primordium get rearranged in the hymenium when asci develop, and become

paraphysoid-like threads. There is no differentiated ascomatal wall. The mature

ascoma may be circular, lirelliform or asteriform. First described as typically

ascolocular (SANTESSON, 1950), this type is more recently considered as

ascohymenial (HENSSEN & JAHNS, 1974).

y) The Opegrapha type (Fig. 10 А to B).

The plexiform primordium extends in an horizontally elongated ébauche. This

differentiates into a meniscus and a paraphysoid net and becomes surrounded by

a distinct envelope. The mature ascoma is a lirella with no ostiolar apparatus,

opening by a split; its interascal filaments are paraphysoids. This type of

development is somewhat alike that of Therrya fuckelü, a non-lichenized

Ascomycete (BELLEMERE, 1967). Іп Roccella montagnei, the development,

difficult to interprete, is perhaps a variant of this type.

6) The Graphis type (Fig. 10 D to F).

The young stages remind those of Opegrapha, but later typical paraphyses

develop; in the mature lirelliform apothecia there are no descending filaments

nor ostiolar apparatus. Some Tholurna (Caliciales) (HENSSEN & JAHNS,

1974; TIBELL, 1984), are perhaps a variant of this type with a developed stipe.

Types reminding that of Graphis are known in some non-lichenized

Ascomycetes, but with some differences: in Rhytisma acerinum, there are short

descending filaments whilst in Pseudopeziza and Bulgaria, the mature apothecia

is not lirelloid (BELLEMERE, 1967); moreover the latter has an important

gelatinous stroma.

£) The Baeomyces type (Fig. 7 E-l).

The important primordium has a plexiform structure; its lower part elongates

upwards in a podium; at its upper part, under a thin veil (= roof) a net of

hyphae containing ascogonial coils differentiates. Later, upright filaments

develop at the top of the net through the veil; only those surrounding the asci,

become paraphyses.

Source : MNHN, Paris

224 М.А. LETROUIT-GALINOU and А. BELLEMÉRE

2. Types with differentiation of a parathecial apparatus.

а) Types with a parathecial apparatus reduced to а parathecial crown.

x) The Aspicilia type (Fig. 12 F).

It is characterized by the precocious differentiation of a marginal parathecial

crown around the primordium. This crown remains reduced in the subsequent

development. The interascal filaments are reputed to be paraphysoids.

$) The Gyalecta type (Fig. 9)

Paraphysoids, which exist in the young ébauche, disappear in the old ones.

Numerous paraphyses develop. Some short descending filaments are also

present. After the subcircular opening of the ébauche, the persisting roof is

pushed lateraly on the flank of the developing ascoma; then it generally

disappears, replaced by a parathecial crown which takes no part in the

subsequent ascoma enlargement, А similar type of development is known in

non-lichenized Ascomycetes (е. g. Calloria, BELLEMERE, 1967)

b) Types with a typical parathecial apparatus.

x) The Peltigera type (Fig. 6).

At first, the primordium has a thalline structure; later it becomes a typical

ébauche with a paraphysoid net. This ébauche enlarges whilst numerous

paraphyses and descending filaments develop. At last, the roof vanishes; a well-

developed parathecial apparatus differentiates at the margin of the hymenium

and takes a large part in the peripherical extension of the apothecium.

B) The Diploicia type (LETROUIT-GALINOU, 1961, 1967; HENSSEN

1968; ef. BELLEMERE & LETROUIT, 1988).

The primordium has a plexiform structure. The young ébauche, with a

paraphysoid net, enlarges only by its basal part which forms a lateral pro-

parathecium. Then the paraphysoid net ruptures near its top and an ascomatal

cavity develops under the roof; paraphyses and asci are generated by the

subhymenium. Later the roof vanishes; a peripherical parathecial apparatus

differentiates, widening the apothecium. A similar hemi-angiocarpic

development, frequent in Lichens, exists, for instance, іп Hertella (HENSSEN,

1985 b), Polychidium (KEUCK, 1977), Lobaria (LETROUIT-GALINOU, 1971;

KEUCK, 1977; HENSSEN, 1981) and іп Lasallia (Н SSEN, 1970;

JANEX-FAVRE, 1973). It is also known in Umbilicaria (HENSSEN, 1970;

JANEX-FAVRE, 1974), where, moreover, secondary hymenia form (Fig. 11 A

to D; cf. p. 216) and in Stereocaulaceae, where apothecia develop on thallus ex-

pansions miming the typical podetions of Cladoniaceae (WOLFF, 1905;

JAHNS, 1970 a).

y) The Xanthoria type (Fig. 2).

In this type, the ébauche is characterized by the early formation of a

parathecial apparatus with a tendancy to a precocious gymnocarpic ascoma.

The parathecial apparatus plays an important role in the growth of the ascoma.

A number of slight variants are encountered, for instance іп Buellia (excl.

Source : MNHN, Paris

ASCOMATAL DEVELOPMENT IN LICHENS. 225

Diploicia), Lecanora, Lecidea, Lecidella, Parmeliella, Lecania, Megalospora,

Ramalina (LETROUIT-GALINOU, 1967; HENSSEN, 1968; KEUCK, 1977,

1979; SIPMAN, 1983). They differ by the reduction of the ébauche, the impor-

tance and details in the structure of the parathecial apparatus.

Тһе Diploicia- and the Xanthoria-types (= parathecial types, BELLEMERE,

1967) are well-represented in the non-lichenized Ascomycetes.

с) Peculiar types with presumably atypical parathecial apparatus.

In these types, difficulties of interpretation arise because dispositions of

hyphae encountered in certain structural components, at definite stages of the

development, are not alike those which are usually observed at such stages in

the same components, but they more or less remind one of dispositions generally

encountered either at other stages or in other structural components.

x) The Cladonia type (Fig. 7 А to D).

The initium of the podetion which will bear the hymenium, is arbuscular. It

can be interpretated either as а non typical primordium or as a precocious

parathecial crown developed on a primordium very reduced (more than in

Xanthoria parietina). With the first hypothesis, the resulting podetion has to be

interpreted as an elongated carpocentrum (LETROUIT-GALINOU, 1967); with

the second hypothesis (CHADEFAUD ct al., 1969), it would be equivalent to

the parathecial discopodium described in various stipitate apothecia of non-

lichenized Discomycetes (cf. BELLEMERE, 1967).

The Parmelia type (LETROUIT-GALINOU, 1970; HENSSEN &

JAHNS, 1974).

Known in the genera Parmelia (Fig. 8, F to K), Usnea, Evernia, Cetraria, this

type shows a plexiform primordium which quickly vanishes whilst a

paraplectenchymatous husk forms at its periphery. Later, paraphyses are

produced on a plexus generated on the internal face of the husk. The origin of

the husk is a subject of controversy. According to IIENSSEN & JAHNS (1974)

and HENSSEN (1981) the husk derives from the external part of the

primordium; consequently it should have to be considered as a part of a

pericentral envelope. According to LETROUIT-GALINOU (1970), the inwards

growth of the husk results from a parathecial structure located at the margin of

the primordium; so this husk would be a new formation, added to the

primordium, and would represent the internal part of a strongly modified

parathecial apparatus. This parathecial hypothesis seems to be more appropriate

as Parmelia asci are similar to those of Lecanora, a genus in which the

ascomatal development is accelerated, with no recognizable ébauche and with

the formation of a precocious parathecial apparatus. The Parmelia type of

development could represent an accelerated (more evoluted ?) Lecanora type.

From these two examples, Cladonia and Parmelia, the question of possible

convergences in ascomatal development arises. At the present time, this question

cannot be correctly answered as detailed ontogenetical studies are still

insufficient; consequently this problem will not be further developed here.

It has been mentioned above that some of the ontogenetical types of lichen

ascomata remind of those known in non-lichenized Ascomycetes. However, the

Source : MNHN, Paris

226 М.А. LETROUIT-GALINOU and А. ВЕ

types encountered in some major taxa of lichens, are frequently original and

unknown in non-lichenized Ascomycetes. Such are, for instance, the Pyrenula

type and its variants in the Pyrenulales, the Peltigera type of the Peltigeraceae,

the Parmelia type of the Parmeliaceae sensu HENSSEN & JAHNS (1974).

Oppositingly, ontogenetical types common in non-lichenized Ascomycetes are

rare or unknown in lichens, as for instance, the Dothidea type (with nutritious

cells), the Nectria type, the discopodial type of the Helotiales. Nevertheless, same

fundamental processes іп ascomatal development exist in the Ascomycetes,

lichenized or not. The originality observed in Lichens, may result either from the

persistence of primitive traits (е. Graphidaceae comparitively to

Rhytismataceae) or from a specific evolution towards types whose setting up is

favoured by lichenized life (e. в. Cladoniaceae, Parmeliaceae, Peltigeraceae).

ТУ.- FINAL CONSIDERATIONS.

Stimulating hypotheses concerning the evolution of Ascolichens can be

elaborated from structural and ontogenetical datas relative to ascomata (see

HENSSEN & JAHNS, 1974; SIPMAN, 1983; HERTEL, 1984). These hypo-

theses however have to be compared with those resulting from the consideration

of others components of the developmental cycle: vegetative apparatus,

conidiomata and fertile elements, specially asci.

‘The processes of hyphal differentiation occuring in ascomatal ontogeny have

to be interpreted by comparison with the development of the non-lichenized

mycelium and with the thallus edification. On these problems, new concepts and

interesting hypotheses have been recently developed (see CHADEFAUD, 1960,

1984; HENSSEN, 1963; LETROUIT-GALINOU, 1969; LALLEMANT, 1983,

1985: WAGNER & LETROUIT, 1988).

Similarities between the development of ascomata and conidiomata have been

recognized in non-lichenized Ascomycetes (CHADEFAUD 1965, 1982 a). In

Lichens, they have been mentioned by LETROUIT-GALINOU (1972, 1984)

who observes, that during the development of Lobaria laetevirens (Fig. 13), а

system of diverging hyphae extends between a reduced roof and a developed

floor as well in the young ascoma as in the young conidioma; in the ascoma,

these hyphae become paraphysoids; in the pycnidium, they become branched

conidiophores. However, the growth is mostly marginal in the ascoma whilst it is

basilar in the conidioma.

The correlations between the development of the sterile elements of the

ascoma and that of the sporophytic apparatus have not been clearly established.

However, in some cases, a regulation seems to exist: so, in Parmelia conspersa,

ascogonial elements are only observed in the primordium, whilst the sporophytic

elements are only found in the basal meniscus, the dicaryotic cells forming later,

when the paraphyses establish. Generally, the development of the asco-

sporophytic apparatus is too poorly known to allow any valuable conclusion.

Ascomatal characteristics have for long been largely used in Lichen

systematics. At first, only morphology and general anatomy have been taken

into account; then detailed structure and histology of the hymenium and

Source : MNHN, Paris

ASCOMATAL DEVELOPMENT IN LICHENS 227

subhymenium were specially considered. Nowadays, ontogeny is recognized to

have the best systematical value amongst ascomatal criteria and so this has led,

either to reconsider families, or to split some of them or also to join some to

others. For instance, the Opegraphaceae have been separated from the

Graphidaceae, the Bacomycetaceae and the Stereocaulaceae have been excluded

from the Cladoniaceae; oppositingly, the Usneaceae have been joined to the

Parmeliaceae (cf. HENSSEN & JAHNS, 1974), and genera precedingly placed

in different orders are now gathered in Lichinales (HENSSEN & BUDEL,

1988).

Important elements to our knowledge of evolutive traits in the Lichens have

been also brought by the study of the fine structure and development of asci

(BELLEMERE & LETROUIT, 1987). Recently, several lichen families have

been redefined and numerous new ones built by the consideration of ascal struc-

ture (HAFFELNER, 1984). It appears that evolutive trends in Lichens would be

more accurately defined if systematical divisions are founded altogether on ascal

and ascomatal datas. Consequently, it would be desirable that in the same

family, genera have similar ascal type and similar ascomatal ontogeny. This is

effectively satisfied in the new delimitation of the Parmeliaceae which joins, for

instance, Рағтейа, Usnea and other genera. Another example is the justified

reestablishment of the Ramalinaceae differing from the Parmeliaceae by asco-

matal ontogeny (Fig. 3) and also by the ascal apex structure. But rather

frequently ascal and ascomatal datas lead to antagonistic systematic conclusions.

Then, other characters must be considered to delimit taxa. For instance

Endocarpon and Dermatocarpon, which һауе similar bitunicate asci and were

precedingly placed in the same family (Verrucariaceae); however they differ іп

their ascomatal development (WAGNER, 1987); differences in their conodioma

types JANEX-FAVRE and WAGNER, 1986), argues to place them in two dis-

tinct. families, respectively the Verrucariaceae and the Dermatocarpaceac

(WAGNER, 1987).

Outlines of ascomatal evolution in Lichens will not be discussed here, the

purpose of the present paper being restricted to expose some reflective

considerations concerning the development of the ascomata. We only hope to

have shown that datas relative to ascomatal ontogeny, though they are long and

difficult to obtain, are fundamental for a better knowledge of evolutionary trends

in Lichens and higher Ascomycetes.

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ASCOMATAL DEVELOPMENT IN LICHENS 233

LIST OF ABBREVIATIONS USED IN THE FIGURES 1 to 13.

a, algae; af, ascogonial filaments; ah, ascogenous hyphae; am, amphithecium; amh,

amphithecioid hyphae; ap, apical point; as, asci; asp, ascospore; at, ascomata; aw, ascomatal

wall; bu, bulge; с, carpocentrum; ce, carpocentral envelope; cm, circumcentral muff, cn,

diverging conidiophores; ct, thalline envelope initium; cv, ascomatal cavity; cx, thallus

cortex; cz, circumcentral zone; dc, disappearing carpocentrum; df, descending filaments; eb,

epicentral bundle; ec, epicentral cone; em, ectal meniscus; ep, epithecium; et, ectal part of

the primordium; ex, excipulum; f, floor; в, growing zone; h, hypothecium; ha, hymenial

algae; hu, husk; hy, hymenium; i, initium of the ascomata; inv, involucrum; Im, lateral

muff, m, thallus medulla; mn, meniscus; n, neck; oc, ostiolar canal; p, paraphyses; pa,

parathecium; pap, parathecial apparatus; pe, parathecial crown; pd, podium; pe, periphyses;

pf, paraphysoid filaments; pm, proper margin; pn, paraphysoid net; po, podetion; pp,

proparathecium; pr, primordium; r, roof; rh, rhizomorph; sa, sporophytic apparatus; sb,

sushymenial bell; sc, secondary carpocentrum; se, secondary envelope; sf, sterile filaments;

sh, subhymenium; sp, split; su, suber; 1, thallus; tm, thalline margin; te, thalline envelope; tr,

trichogyne; sf, sterile filaments.

Source : MNHN. Paris

Cryptogamie, Bryol. Lichénol., 1989, 10 (3): 235-245 235

ADDITIONS AND CORRECTIONS FOR PHILIPPINE

MOSS FLORA

B.C. TAN" and T. KOPONEN**

ж clo ALCON, 629 T. Alonzo st., Sta. Cruz, Manila, The Philippines.

** Department of Botany, University of Helsinki, Unioninkatu 44,

SF-00170 Helsinki, Finland.

ABSTRACT - Bryobrothera crenulata (Broth. et Par.) Thér., Bryum erythropilum Fleisch.,

B. clavatum (Schimp.) C. Muell., Diaphanodon blandus (Нагу. in Hook.) Ren. et Card. var.

recurvedentatus Zant, Entosthodon physcomitrioides (Mont) Mitt, and Philonoris socia

Mitt, are reported new to the Philippine moss flora. In addition, four new synonyms are

proposed for the genus Bryum Hedw., and one new combination, Anomobryum erectum

(Broth.) Tan et T. Kop. is made.

Owing to its geographic location and geologic history, the Philippine archipe-

lago has received its moss taxa from several sources: East Asiatic mainland,

maritime Pacific coast, the Himalayas, tropical Asia, Australasia and Oceania,

not to mention the widespread northern and southern temperate taxa (Tan

1984).

The study of Philippine moss flora started near the turn of this century when

several European and North American explorers and naturalists collected in the

Philippines. The results which were published in widely scattered journals were

ably summarized by Bartram (1939). In spite of this, later collections continue to

yield new records including several species new to science. The recent checklist

by Iwatsuki and Tan (1979) includes 625 species belonging to 218 genera.

Progress of Philippine bryology has been specially rapid in the last ten years.

This is evident in the discoveries of twelve genera new to the country:

Physcomitrium, Hageniella, Orthodontium ` (hwatsuki & Тап 1980),

Orthorrhynchium (Tan 1981), Dixonia, Meteoriella, Mnium (del Rosario & Van

Zanten 1982), Tristichella (Тап & Iwatsuki 1983), Haplohymenium, Solmsiella,

Leptostomum (Тап 1987) and Grimmia (Tan & Deguchi 1987).

No doubt, future exploration in remote mountains on distant islands in the

country will expand substantially the size of this diverse flora. Such exploration

is most necessary because of the on-going destruction of the local forest vege-

tation.

Source : MNHN, Paris

236 B.C. TAN and Т. KOPONEN

Below we present 5 species as noteworthy additions to the Philippine moss

flora. Bryobrothera Thér. is a new generic record. In addition, four new syno-

nyms are clarified and proposed for the genus Bryum Hedw. One new combina-

tion is likewise made in Anomobryum Schimp. Specimens of these new records

are deposited at the herbaria of the University of the Philippines at Los Banos

(САНР) and the University of Helsinki (H).

Bryobrothera crenulata (Broth. et Par.) Thér.

Rev. Bryol. 47: 26. 1920.

The discovery of this species in the mountains of Mindanao Island is а re-

markable extension of another Australasian floral element into the southern Phi-

lippines. For that matter, the species is a new record for the western Malesian

subprovince.

Bryobrothera crenulata has an interesting history having been described at

various times as a member of Mesochaete, Calomnium and Rhizogonium (Norris

& Robinson 1979, Koponen et al, 1986). Its biosystematic position and familial

status has only recently been clarified (Norris & Robinson 1979). As a member

of Hookeriaceae, it was missed out in the recent revision of the family for the

Philippine moss flora (Tan & Robinson 1989).

Locally, this species forms extensive mats covering the branches of tree in

Agathis-Podocarpus montane forest and appears superficially like a hepatic.

Dr. D. Norris kindly confirmed our determination of the Philippine specimen

of Bryobrothera crenulata.

Specimen examined: Mindanao Is. Mt Hilong-Hilong, Cabadbaran, Agusan

Norte Province, Tan & Navarez 84-520 (CAHP, H). New Philippine generic and

species record!

Genus Bryum Hedw.

This large and difficult genus is poorly understood and equally undercollected

in the Philippines. Iwatsuki and Тап (1979) listed 11 species with two endemic

taxa: B. microtheca C. Muell. апа B. chrysobasilare Broth. 1908, non Broth.

1924. Unfortunately, these two Philippine endemics were overlooked by Ochi

(1983) in his summary revision of subfamily Bryoideae for South, Southeast and

East Asia. Nor were the two taxa mentioned by the same author in his earlier

studies (Ochi 1959, 1960, 1968a, 1970).

We examined the types and reference collections of Brpum microtheca, B.

leucophylium Dozy et Molk. and В. chrysobasilare and interpreted the first two

taxa as synonyms of 8. argenteum Hedw., and the last one synonymous to B.

erythrophyllum Fleisch.

We also studied the type оГ Bryum erectum Broth. and found it conspecific

with B. petelotii Ther. et Henry. Earlier, Bartram (1939) had synonymized B.

erectum with B. microtheca, a conclusion with which we disagree.

Source : MNHN, Paris

PHILIPPINE MOSS FLORA 237

Likewise, we studied the Philippine specimens of Bryum australe Hampe cited

in Bartram (1939) and would consider them В. paradoxum Schwaegr. sensu Ochi

(1985).

Lastly, Bryum clavatum (Schimp.) C. Muell. sensu Ochi (1985) was identified

from a recent collection made by the first author from Mindoro Island (Figs.

1-5).

The detailed information is as follows:

Bryum argenteum Hedw.

Sp. musc. frond.: 181. 1801.

Bryum microtheca C. Muell., Syn. muse. frond. 1: 314. 1848. syn. nov. - Type:

Philippines, Manila, Meyer s.n. (not seen).

Bryum leucophyllum Dozy et Molk., Ann. Sei. Nat. Bot. ser. 3, 2: 301. 1844.

syn. nov. - Type: Java, no collector (isotype, H-BR!).

There is a wealth of discussions on the gametophytic variation of Bryum

argenteum (Fleischer 1902-1904, Sainsbury 1955, Nyholm 1958, Gangulee 1974,

Scott & Stone 1976, Crum & Anderson 1981, Frahm & Frey 1983), but curious-

ly, little attention is given to the sporophytes which have been assumed to be uni-

form in this species.

We attempted to fill this gap of sporophytic information by studying two large

population of Bryum argenteum collected from a burnt and cultivated limey out-

crop in Trinidad Valley, Benguet Province, Luzon ls. (Tan 86-465, 86-461,

CAMP, ID. The two populations were selected on the basis of their relatively

uniform gametophytes which clearly identify them as the var. /anatum (P.

Beauv.) Hampe.

In both populations, the number of suberect or slightly inclined capsules in

the semi-dry state were calculated to be about 10% based on a total of randomly

selected 50 capsules in each population. The rest are either horizontal or pendant

with various degrees of angle formed between the theca and the seta.

Subsequently, five suberect to weakly inclined capsules were chosen and dis-

sected for peristomial details. The results were then compared with the peristomi-

al structures observed in five pendant capsules. True enough, there is a trend in

the reduction of peristomial structures from the perfect type seen in typical, pen-

dant capsules to the variously imperfectly developed types seen in the suberect

capsules.

It appears that the tiny transverse bars or appendiculae of the endostomial ci-

lia are the first to disappear in a non-pendant capsule. This is followed by a re-

duction of cilial length. In both suberect and pendant capsules, the endostomial

segments are rather fragile, often left broken in the long dehisced capsules.

"The opercular lid, likewise, was observed to vary from markedly apiculate to

nearly flat with only a vestigial hump. Length of thecae varies between 1-2mm

and the setal length measures from 10-20mm. The variable characters mentioned

in this paragraph, however, have no correlation with the posture of the capsules.

On the other hand, exothecial cells and stomatal apparati are observed to be

rather uniform in all capsules studied.

Source : MNHN, Paris

238 B.C. TAN and Т. KOPONEN

Thus, the separation of Bryum leucophyllum from B. argenteum on the basis

of characters such as appendicular versus nodulose cilia, and apiculate versus

blunt opercular lid (Fleischer 1902-1904, Ochi 1985) is clearly untenable.

Similarly, Bryum microtheca, which was separated from В. argenteum by

characters such as suberect or pendant capsules and reduced cilia (Mueller 1848,

Bartram 1939), is equally an ill-founded species.

We think that it is best to combine these three taxa. Although we failed to lo-

cate the C. Mueller type of Bryum microtheca, the reference specimens so

named at the Brotherus Herbarium, including an isotype of B. leucophyllum,

support our view.

Bryum erythropilum Fleisch.

Musci Buitenzorg 2: 553. 1904. - Type: Indonesia, West Java, Tjibodas,

Fleischer s.n. (syntype, H-BR!).

Bryum chrysobasilare Broth., Philipp. J. Sci. 3 С: 19. 1908, syn. nov. - Type:

Philippines, Luzon Is., Mt. Data, Merrill 4956 (holotype, H-BR!).

Bryum erythropilum was reduced to a synonym of Bryum clavatum (Schimp.)

C. Muell. by Koponen and Norris (1984) who had adopted a broad species con-

cept for the genus because of the great variation seen in their large New Guinean

moss collections. Ochi (1985), however, maintains the two as separate species on

the bases of their differences in sexuality (autoicous іп В. erythropilum and dioi-

cous іп B. clavatum), in the degree of leaf border differentiation (1-2 rows of

elongate border cells in B. erythropilum versus 2-5 rows of linear border cells in

B. clavatum), and also in the size of median to basal leaf cells (longer and larger

in В. clavatum). Seen in this light, the type of Philippine В. chrysobasilare is con-

specific with B. erythropilum.

On the other hand, the remarks made by Bartram (1939) regarding the co-

lored basal leaf cells and deep red capsules of B. chrysobasilare, in our evalu-

ation, has little taxonomic value. The same features are seen in В. clavatum sen-

su stricto and also in Bryum paradoxum (see below).

In addition, numerous deep red, sphaerical rhizoidal gemmae of different

sizes were observed in the other Philippine specimen of B. eryrhropilum (Tan et

al. 84-380).

Specimen examined: Mindanao Island, burnt mossy forest near the peak of

Mt Kitanglad, Butuan Province, mixed with Funaria hygrometrica, Tan, Navarez

& Amoroso 84-380 (CAHP, H). New Philippine species record!

Bryum paradoxum Schwaegr.

Sp. musc. frond. Suppl. 3: 224a. 1827.

Bryum “tenuisculum” Hook. in Iwatsuki et Tan, Kalikasan 8: 186. 1979, error

pro Bryum teretiusculum Hook., Icon. РІ. 1: 20f. 1836, synonymized by Ochi

(1969).

Bryum australe Hampe sensu Bartram, Philipp. J. Sci. 68: 141. 1939.

Source : MNHN, Paris

PHILIPPINE MOSS FLORA 239

Bartram (1939) cited two Philippine collections of Bryum australe (Williams

1782, 3148, H-BR) and corrected the species determination to B. ambiguum

Duby (= B. apiculatum Schwaegr.). Our examination of these two collections

shows that the specimens have strongly excurrent leaf costae. The leaf cell areo-

lation consists of narrowly rhomboidal to elongate-linear upper and median leaf

cells that are (60-)75-135(150)um long, with sharp angular ends and thin-walled,

and becoming abruptly broadly rectangular at base. The leaf margins are weakly

serrulate towards the apex, consisting of 1-3 rows of linear border cells, and are

partly revolute. These characters suggest to us that these two Philippine col

lections are B. paradoxum.

Bryum paradoxum differs from B. apiculatum principally in having a much

stronger excurrent leaf costa, less concave leaves and more revolute leaf margins.

On the other hand, it differs from the true Bryum australe, according to Ochi

(1970, 1985), in having longer excurrent leaf costae and a more or less abrupt

transition of leaf cell shape from the elongate cells in the upper 2/3 of the lamina

to broadly rectangular ones near leaf base. Ochi (1985) considers the two as vi-

carious taxa in temperate Asia and Australasia.

Bryum paradoxum can also be confused with Bryum erythropilum. Based on

our limited experience with the Philippine specimens, the former is best distin-

guished from the latter by its longer leaf cells in the upper half of the lamina, and

by a revolute and more strongly differentiated leaf border consisting of 1-3 rows

of linear cells, instead of the 1-2 rows of rectangular to elongate leaf border cells

seen іп B. erythropilum. Ochi (1985) further distinguishes the two taxa by their

sexuality, autoicous in B. erythropilum and dioicous іп B. paradoxum.

Bryum paradoxum is a variable species. Defined іп a broad sense, it can be

considered a synonym of B. clavatum sensu Koponen and Norris (1984). We

have, however, not seen enough material of B. paradoxum to offer a definitive

taxonomic judgement on this matter, Ochi (1985) distinguishes B. paradoxum

from B. clavatum mainly because of the presence of a more markedly differen-

tiated leaf border іп B. clavatum.

The biosystematic relationship among these four taxa needs more clarifica-

tion. Ochi (1970, 1985) has illustrated well his concepts for these four species.

Specimens examined: 1) Bryum paradoxum - Luzon Is., Benguet Prov., Ba-

guio. on rock, Williams 1782 (П-ВЮ); ibid, Sablan, on rock, Williams 3148

(H-BR). 2) Bryum clavatum - Mindoros Is., Mt Halcon vicinity, Tan 87-042

(CAHP, H). New Philippine species record!

Anobryum erectum (Broth.) Tan et Kop., comb. nov.

Basionym: Bryum erectum Broth., Philipp. J. Sci. 3 C: 19. 1908. - Type: Philip-

pines, Luzon 15., Benguet Province, Kabayan, Merrill 4968 (holotype,

H-BR)).

Bryum petelotii Ther. et Henry in Henry, Rev. Bryol. n. ser. 1: 43. 1928, syn.

nov. - Type: Vietnam, Tonkin, massif du Pia-Quac, dv. Cao-Ouac, Petelot

s.n. (isotype, H-BR!).

Source - MNHN, Paris

PHILIPPINE MOSS FLORA, 241

The types of these two bryaceous taxa described with erect capsules match

perfectly in both their gametophytic and sporophytic details. Chronologically,

Bryum erectum has priority over B. petelotii.

Although the type specimen of Bryum erectum has silvery, terete branches

with leaves that closely resemble that of Bryum argenteum, the two species сап be

separated by several characters. In B. erectum, the median and submedial leaf

cells are oblong-rhomboidal to spindle-linear shaped, measuring 60-130ит long,

with sharp terminal ends and thin-walled. Іп Bryum argenteum, they are shortly

oblong to rectangular, mostly 30-S0um long, with blunt cell ends and moderately

thick-walled. Additionally, the leaf costa of B. erectum is percurrent to only

weakly excurrent from a gradually acuminate leaf apex. Contrastingly, іп 8.

argenteum, especially the var. (апат, the leaf costa is strongly excurrent, filling

the subula of the abruptly contracted cuspidate leaf apex. Indeed, even under a

10x field lens, the piliferous hairpoints are more clearly visible in Bryum

argenteum than in. B. erectum. Most importantly, the capsules of B. erectum are

erect and somewhat oblong, reminiscent of the capsular type of Brachymenium

or Anomobryum.

In discussing the placement of Bryum albo-imbricatum Ochi (syn. B. albidum

Broth. in Herz, 1916, hom. illeg.) from Bolivia, Ochi (1980) disagreed with

Brotherus (Le, see protologue) who allied this taxon to Bryum argenteum on the

basis of their gametophytic similarity. Instead, because of exothecial cell charac-

ters, Ochi (1980) considered it to be a member of his subgenus Anomobryum.

According to him, these cells in Bryum albo-imbricatum are more or less elon-

gate-rectangular with uniformly thick walls except near the oral region of the

capsule. In Bryum argenteum and its allies, the exothecial cells are mostly nar-

rowly rectangular with very thick vertical walls, becoming abruptly hexagonal to

short rectangular with thin cell walls near the oral region.

Interestingly, the same differences exist in the exothecial cell morphology be-

tween Bryum erectum and B. argenteum. Further comparison of capsules of

some species of Anomobryum and Bryum made by us shows that in Bryum, there

are 1-4 rows of horizontally oriented, narrowly rectangular shaped cells border-

ing the capsular mouth. In Anomobryum, the oral border of the capsule appears

to consist of only 1-2 rows of such type of exothecial cells.

Bryum erectum is, extraordinarily, the only species among the Southeast and

East Asiatic members of Ochi's subgenus Bryum (1985) that has erect capsules.

Also, its upper leaf cells tend to be more linear-elongate than usual in Bryum.

Furthermore, the leaf costa of B. erectum is without guide cells. We are, for all

these reasons mentioned above, accepting Bryum erectum as an Anomobryum

sensu Koponen and Norris (1984).

Admittedly, species of Anomobryum with long acuminate leaf apices are rare,

but not unknown. In addition to Anomobryum albo-imbricatum (Ochi) T. Kop. et

Norris cited above, we can mention Anomobryum kasmirense (Broth.) Broth.

from the Himalayas.

Bartram (1939) seem to have overlooked these critical morphological differ-

ences when he synonymized Bryum erectum with B. microtheca (= В. argenteum

in this paper). Of the four collections Bartram (1939) listed under B. microtheca,

only the two specimens from Kabayan town (Merrill 4968, Bacani, For. Bur.

Source ММНМ Paris

242 B.C. TAN and Т. KOPONEN

15988, H-BR) are truly Anomobryum erectum. The other two are typical of

Bryum argenteum var. lanatum.

On several occasions, Ochi (1954, 1959, 1963a, 1985) has suggested that

Bryum petelotii (= Anomobryum erectum) resulted from ап intergeneric cross be-

tween Bryum argenteum and Brachythecium exile (Dozy et Molk.) Lac. He sup-

ported his claim by evidence of aborted spores, among others. However, spores

from the Philippine specimens of Anomobryum erectum which measure 6-9um in

diameter appear highly viable. Furthermore, a few Philippine plants were ob-

served to bear axillary globose bulbils measuring 180-250m long and 1204m

wide.

The range of Anomobryum erectum now includes several localities in warm

temperate Asia and tropical Asia, reaching as far as Costa Rica and Mexico (see

Ochi 1985).

Diaphanodon blandus (Harv. in Hook.) Кеп. et Card. var. recurvedentus Zant.

Blumea 9: 544. 1959. - Type: India, Sikkim, Kurseong, Decoly & Schaul 2564

(isotype, H-BR)).

This variety was described to consist mainly of long, flaccid and flagelliform

stems and branches with few lateral innovations. The leaf margins are coarsely

serrated with patent to recurved teeth (Van Zanten 1959). Although flagelliform

modified branches are seen also in specimens of var. blandus, they are never so

profusely developed as in the present variety.

Despite the remarkable morphological distinction, we think that the var.

recurvedentatus is probably a widespread eco-variety caused by the shade and

highly humid condition at high elevation.

Diaphanodon blandus var. recurvedentatus has been recorded only from Sik-

kim (Van Zanten 1959). We have now seen many more specimens of this varie-

ty, including an isotype, at the Brotherus Herbarium. The total range of it can be

gleaned from the specimens cited below and this includes a new Philippine re-

cord.

Specimens examined: India, Manipur, 1899, Fraser s.n. (H-BR); Upper Bur-

ma, near Ruby mine, Olivier s.n. (H-BR); Sri Lanka, Kirigalpota, Herzog 80a

(H-BR); Indonesia, Java, Pangerango, 1897, Moller 139 (H-BR); Philippines,

Luzon Is., Benguet Prov., Mt Santo Tomas, 30 Хоу. 1986, Tan s.n. (CAHP, Н).

Entosthodon physcomitrioides (Mont.) Mitt.

J. Linn. Soc. Bot. Suppl. 1: 55. 1859.

The key and illustrations to the species of Entosthodon (Ochi 1986b) works

well with the Philippine specimen. However, the leaf apices of Philippine E.

physcomitrioides vary from piliferous to short acuminate, with more leaves on the

piliferous side. Length of capsules also varies from 1.5 to 2mm. Like all other

members of the genus, the local populations of E. physcomitrioides grow on dis-

turbed clayey road cut along a mountain cliff.

Entosthodon dozyanus С. Muell. from Java differs from the present species іп

having a markedly differentiated leaf border and a capsule with only short neck.

Source - MNHN. Paris

PHILIPPINE MOSS FLORA 243

Presently, Entosthodon physcomitrioides has a known distribution from India,

Indochina, Taiwan and New Caledonia (Ochi 1968b). Its presence in the Philip-

pines is not totally unexpected.

Specimen examined: Luzon is., Benguet Ргоу., Buguias, on way to Ballay,

Tan & Hernaez 85-123 (CAHP, H). New Philippine species record!

Philonotis socia Mitt.

J. Linn. Soc. Bot. 8: 151. 1864.

The key and illustrations prepared for the species of Philonotis Brid. in Ochi

(1962, 1963b) and Iwatsuki and Mizutani (1972) should be consulted for the dif-

ferences between P. socia and its related taxa.

Among the Philippine congeners, Philonotis socia is characterized by a lan-

ceolate-triangular leaf outline with predominantly short rectangular basal leaf

cells and a short excurrent leaf costa. The leaf margins are strongly revolute

nearly throughout.

The local habitat is a wet, partly shaded rocky ledge at the mouth of a spring

in a newly opened forest clearing.

The range of Philonotis socia now covers Japan, Taiwan, China mainland,

India and the Philippines.

Prof. Z. Iwatsuki of Hiroshima University kindly confirmed our species deter-

mination.

Specimen examined: Luzon Is., Laguna Prov., Louisiana, Tan 84-37 (CAHP,

H). New Philippine species record!

ACKNOWLEDGEMENT

The first author acknowledges the financial support given by the Finnish Ministry of

Education during his visit-stay at the Botany Department of the University of Helsinki in

summer of 1988. He further expresses his gratitude to Drs P. Isoviita and S. Piippo for the

use of library and equipments at the Brotherus Herbarium of the University of Helsinki.

Lastly, both authors thank Dr. D.H. Norris for his comments on the manuscript and his

help in editing the English text.

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cent regions, J. Fac. Educ., Tottori Univ., Nat. Sci. 19: 24-40.

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regions. Jap. J. Bot. 20: 1-34.

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land and the adjacent islands. J. Fac. Educ., Tottori Univ., Nat. Sci. 21: 7-67.

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PHILIPPINE MOSS FLORA 245

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Bot. Lab. 55: 13-22.

TAN B.C., 1987 - New records of Philippine mosses. Mem. New York Bot. Gard. 45:

446-454.

TAN B.C. & DEGUCHI H., 1987 - The genus Grimmia (Musci) in the Philippines.

Yushania 4: 11-12.

TAN В.С. & ROBINSON H., 1989 - A revision of Philippine Hookeriaceae. US Smithso-

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Source : MNHN. Paris

Cryptogamie, Bryol. Lichénol. 10 (3): 247-252 247

LIQUENES CON CIANOFICEAS DE CAAVEIRO,

LA CORUNA (N-O DE ESPANA)

М.Е. LOPEZ DE SILVANES y R. CARBALLAL

Departamento de Biología Vegetal, Facultad de Biología,

Universidad de Santiago, España.

RESUMEN - Continuamos dando a conocer los resultados del estudio sobre la flora

liquénica de la “fraga” de Caaveiro (La Coruña), reuniendo en este trabajo los táxones per-

tenecientes a las familias: Collemataceae Nannf., Lichinaceae Chev., Lobariaceae Chev.

Pannariaceae Tuck. y Peltigeraceae W. Watson. Consideramos especies interesante:

Polychidium dendriscum (Nyl.) Hensen, nueva cita para Europa continental; Pannaria

tavaresii P.M. Jørg., nueva Cita para España peninsular; Leprogium microphylloides Nyl. y

Parmeliella jamesii Ahlner & P.M. Jørg. muy poco citadas en España. Otras 8 especies son

novedades para la flora de Galicia.

ABSTRACT - New results of the study on the lichen flora of the “fraga” of Caaveiro (La

Corufia).The taxa of the families: Collemataceae Nannf., Lichinaceae Chev., Lobariaceae

Chev., Pannariaceae Tuck. and Peltigeraceae W. Watson are treated. Interesting species are:

Polychidium dendriscum (N y1.) Hensen new for continental Europe, Pannaria (avaresii Р.М.

Jorg. new for the peninsular Spain, and Leptogium microphylloides Nyl. and Parmelia

jamesii Ahlner & P.M. Jørg., seldom mentioned from Spain. Additionnal $ species are new

for the flora of Galicia.

INTRODUCCION

Hemos reunido en este trabajo los liquenes con cianoficeas recogidos en la

fraga de Caaveiro, La Coruña (N-O de España). La zona ha sido caracterizada

en un trabajo anterior (Lopez de Silanes & Carballal 1987); añadiremos aquí

que los materiales geológicos corresponden fundamentalmente a esquistos y gra-

nitos y las altitudes oscilan entre los 20 y 500 metros. La mayor parte del territo-

rio estudiado está ocupado por un bosque perteneciente a la asociación Blechno

spicantis-Quercetum roboris Tx. 6: Oberd. (1958).

Para cada especie indicamos su ecología y distribución para Europa, tomado

esta última de Jorgensen (1978) y Clauzade & Roux (1985). Es de destacar el

alto número de especies de distribución restringida, en su mayoría atlánticas o de

tendencia oceánica y otras de distribución puntual, lo que da a la zona un gran

interés Погізісо. En aquellas especies que son nuevas citas o están poco citadas

Source : MNHN, Paris

248 М.Е. LOPEZ DE SILANES y R. CARBALLAL

en España, así como en las que presentan сага

su morfología se efectua una breve descripción.

teristicas peculiares en cuanto a

Las muestras se hallan en el herbario del Departamento de Biología Vegetal

de la universidad de Santiago (SAN T-Lich.) cuyo número de pliego se especifica.

La recogida de material ha sido efectuada por el equipo de liquenólogos de di-

cho departamento.

CATALOGO

Collema flacidum (Ach.) Ach. - Encontrado en las grietas de un muro orienta-

do al norte y en rocas a ras de suelo. Amplia distribución en Europa. 9m, es

quistos, SANT-Lich. 227

Collema furfuraceum (am) Du Rietz - Corticicola y muscicola-saxícola.

Distribución: especie frecuente en lugares húmedos de Europa. Castanea sativa,

SANT-Lich. 2261; 40m, Fraxinus excelsior, SANT-Lich. 2279; rocas graniticas

con musgo, 90m, SANT-Lich. 2260.

Collema aff. nigrescens (Нийѕ.) DC. - Nuestros ejemplares difieren de la

descripción dada por Degelius (1954) en el grosor del talo, siendo de

120-130ym. El material ha sido revisado por el Dr. P.M. Jorgensen comunican-

donos que pudieran ser ejemplares de esta especie especialmente gruesos. Sobre

musgos en troncos de árboles. Distribución: en Europa de tendencia oceanica,

en Espana de amplia distribución. Castanea sativa, SANT-Lich. 2283.

Ephebe (апаға (L.) Vain. - Especie abundante y frecuente en las partes de la

roca que están próximas al río o arroyos y en ciertas épocas pueden estar inun

dadas. Amplia distribución en Europa. 30m, esquistos, SANT-Lich. 2276.

Leptogium brebissonü Mont. (Syn: L. chloromelum Midd) - Corticicola y

muscicola. Distribución: Europa atlántica y mediterránea. De España conoce-

mos citas en Asturias (Vazquez S Crespo 1978), Mallorca (Font & Fiol 1985),

Navarra (Etayo 1986), Guipúzcoa (B. Aguirre, com. pers.) y Cataluña (X. Lli-

mona, com. pers.). Primera cita para Galicia. Quercus robur, SANT-Lich. 2249;

Castanea sativa, SANT-Lich. 2285.

Leptogium aff. cochleatum (Dickson) P.M. Jorg. (Syn.: L. tremelloides (L.

fil.) S.F. Gray, L. azureum auct. р.р.) - Nuestros ejemplares presentan un grosor

del talo menor que el descrito por Jorgensen & James (1983). El Dr. Р.М.

Jorgensen al que le hemos enviado material lo considera conferible a dicha espe-

cie. Muscicola y corticicola. Distribución: regiones atlánticas de Europa. Еп

Espana conocemos citas de Canarias (Follmann 1976) como Leptogium azureum

(Swans.) Mont, de Navarra (Lacoizqueta 1885, Colmeiro 1889) y Lugo (Bellot

1952) como Leptogium tremelloides Fr., no sabemos con certeza si se trata de la

misma especie al no poder estudiar los ejemplares de dicha citas. Castanea

sativa, SANT-Lich 2289; Fraxinus excelsior, SANT-Lich, 2290.

Leptogium cyanescens (Rabenh.) Körber (Syn.: L. caesium (Ach.) Vain.) - Еп

zonas humedas, entremezclado con musgos, grietas de muros y directamente so-

bre la roca en fuentes donde la influencia del agua es directa. Especie oceanica

Source : MNHN, Paris

LIQUENES CON CIANOFICEAS 249

de amplia distribución en Europa. Esquistos, SANT-Lich. 2286; granito, SANT-

Lich. 2288.

Leptogium hibernicum Mitchell ex Jorg. - Muscicola en troncos de árboles.

Distribución: regiones atlánticas europeas. En España conocemos citas de

Huesca (Llimona 1976), Salamanca (Marcos 1985) y de Guipúzcoa (B. Aguirre,

com. pers.). Primera cita para Galicia. 90m, Castanea sativa, SANT-Lich. 2291;

Laurus nobilis, SANT-Lich. 2292.

Leptogium liquenoides (L.) Zahlbr. (Syn: L. lacerum (Retz.) S-F. Gray) - En

rocas cubiertas de musgos y piedras de muros y suelos. Amplia distribución en

Europa. Primera cita para Galicia. 90m, esquistos, SANT-Lich. 2293.

Leptogium microphylloides Nyl. - Talo marrón verdoso. Lóbulos alargados y

estrechos no alcanzando los 0,5mm de ancho, casi siempre cubiertos de isidios

granulosos o papilosos normalmente más estrechos en la base, de simples a ra-

mificados, empiezan saliendo en el borde del lóbulo hasta llegar a ocultarlo,

Siempre esteril. Nuestros ejemplares son pequeños, aproximadamente de 5mm

de diámetro, sus talos aparecen aislados casi siempre en grietas de la corteza, La

distribución que conocemos es la dada por Clauzade & Roux (1985) en el oeste,

centro y sur de Francia. En España la cita Carballal & Garcia-Molares (1988 b)

en las cercanias de la ciudad de Pontevedra. Fraxinus excelsior, SANT-Lich.

2296.

Leptogium sinuatum (luds.) Massal. (Syn: L. scotimum (Ach) Frey) -

Muscicola en la parte vertical de un muro y en rocas a ras del suelo, Especie de

amplia distribución en Europa. No conocemos citas de Сі Esqui

nitos, SANT-Lich. 2294.

Leptogium tenuissimum (Dicks. Körb. (Sync L. sutile (Schrad.) Torss.) -

Saxicola-muscicola. Distribución: amplia en Europa. En España conocemos ci-

tas de Baleares (Maheu & Gillet 1921), Málaga (Seaward 1983) y de Cataluña

(X. Llimona, com. pers.). Primera cita para Galicia. 90m, rocas graníticas,

SANT-Lich, 2295.

Lobaria scrobiculata (Scop.) DC. - Corticicola, poco frecuente en la zona de

estudio, Ampliamente distribuida por Europa. 230m, Quercus robur,

SANT-Lich. 2297.

Nephroma laevigatum Ach. - Corticicola. Distribución: en Europa atlántico-

mediterránea, muy extendida en España. 130m, Quercus robur y Castanea

sativa, SANT-Lich. 2298.

Pannaria conoplea (Ach.) Bory - Muscicola-corticicola. Distribución amplia

en Europa con tendencia atlántica; en España se limita su distribución al tercio

norte. Castanea sativa, SANT-Lich. 2341.

Pannaria mediterranea С. Тау. - Corticicola y muscicola, a veces invade el

talo de otros liquenes. Distribución: atlántico-mediterránea. Castanea sativa,

SANT-Lich. 2329.

Pannaria rubiginosa (Ach.) Bory - Corticicola. Especie frecuente en la zona.

Amplia distribución en Europa. Quercus robur, SANT-Lich. 2348; Corylus avel-

lana, Laurus nobilis, Castanea sativa, SANT-Lich. 2346, 2347, 2349; Fraxinus

excelsior, SANT-Lich. 2327,

Source : MNHN, Paris

250 М.Е. LOPEZ DE SILANES y К. CARBALLAL

Pannaria sampaiana C. Tav. - Corticicola, Un solo ejemplar de pequeñas di-

mensiones. Distribución: en Europa atlántica y mediterránea. Primera cita para

Galicia. Castanea sativa, SANT-Lich. 2321.

Pannaria tavaresii P.M. Jorg. - Talo foliäceo en roseta, marrón claro o ama-

rillento, Р + naranja. Lóbulos ligeramente cóncavos con margenes que tienden

a ser ascendentes y más pálidos. Isidios marginales simples o coraloides con fre-

cuencia algo más ensanchados en el ápice. Especie corticicola encontrada en zo-

nas sombrias del interior del bosque. Distribución: altantica y mediterránea en

Europa. En España esta citada de Canarias (J'ogensen 1978, Sanchez-Pinto & al.

1983, Hernandez-Padron 1985, Hernandez-Padron & al. 1987). Nuestra cita

amplia su distribución al territorio de Espana peninsular. Castanea sativa,

SANT-Lich. 2322.

Parmeliella jamesii Ahlner & Р.М. Jorg. - Talo formado por pequeñas

escuamulas ligeramente alargadas, 0,5-2mm de largo, más o menos continuas,

de color gris azulado claro. Soralios granulosos difusos y marginales, pudiendo

invadir la superficie del talo. Especie frecuente que puede confundirse con

Pannaria mediterranea C. Tay. que es más abundante, cuando esta última pre-

senta una coloración clara. Muscicola sobre troncos de árboles. Distribución:

Europa atläntica. En España conocemos las citas de Canarias (Hernandez-Pa-

dron & al. 1987), Pontevedra (Carballal & Garcia-Molares 1988 a) y Guipúzcoa

(B. Aguirre, com. pers.). Corylus avellana, Пех aquifolium, SANT-Lich. 2323,

2333; Castanea sativa y Eucalyptus globulus, SANT-Lich. 2331, 2332.

Parmeliella plumbea (Lightf.) Vain. - Corticicola, especie frecuente y la más

abundante del género Parmeliella. Distribución: toda Europa con tendencia

atlántica, menos frecuente еп el centro. Corylus avellana, SANT-Lich. 2335;

‘Alnus glutinosa, Quercus robur, SANT-Lich. 2338, 2336; Castanea sativa,

SANT-Lich. 2334; Fraxinus excelsior, SANT-Lich. 2337.

Parmeliella testacea P. Jorg. - Corticicola y muscicola, frecuente sobre

hepáticas foliosas. Distribución: Europa atlántica con algunos enclaves en el

norte de Italia. Primera cita para Gali Castanea sativa, SANT-Lich. 2324;

Quercus robur, SANT-Lich. 2342; Fraxinus excelsior, SANT-Lich, 2331.

Parmeliella triptophylla (Ach.) Müll. Arg. - Corticicola. Distribución: toda

Europa excepto en la zona ärtica. Castanea sativa, SANT-Lich. 2340.

Peltigera horizontalis (Huds.) Baumg. - Muscicola-saxicola. Distribución:

ia en Europa. No conocemos citas de Galicia. 40-400m, taludes,

ich. 2301.

Peltigera polydactyla (Necker) Hoffm. - Sobre restos vegetales, musgos y ro-

cas. Distribución: amplia en Europa. 400m, granitos, SANT-Lich. 2300.

Peltigera praetexta (Flörk. ex Sommerf.) Zopf. - Es la especie de Peltigera

más abundante y frecuente. Saxicola y muscicola. Distribución: casi toda Euro-

pa. 90m, rocas graniticas con musgo, SANT-Lich. 2303; 30-40m, talud rocoso,

SANT-Lich. 2302.

Polychidium dendriscum (Nyl.) Hensen - Se diferencia de P. muscicola (Sw.)

Gray en su talo de ramificación dicótoma que no presenta tomento y el fico-

bionte del género Scptonema. Corticicola. Distribución: Gran Bretaña, Irlanda y

Source : MNHN, Paris

LIQUENES CON CIANOFICEAS 251

Azores; No conocemos citas de Europa continental. Castanea sativa, SANT-Lich.

2304.

Sticta limbata (Sm.) Ach. - Corticícola y muscicola. Distribución: en Europa

atlántico-mediterránea. Corylus avellana, Castanea sativa, Quercus robur,

SANT-Lich. 2306, 2309, 2310; Fraxinus excelsior, Eucalyptus globulus, Laurus

nobilis, SANT-Lich. 2305, 2307, 2308.

Sticta sylvatica (Huds.) Ach. - Muscicola y corticicola. Distribució: en Europa

media y mediterranea con tendencia oceánica. De España conocemos citas de

La Coruña (Colmeiro 1889, Crespo & al. 1981), Cataluña (X. Llimona, com.

pers.), Navarra y Guipúzcoa (B. Aguirre, com. pers.). Corylus avellana, Quercus

robur, Castanea sativa, SANT-Lich. 2311, 2318, 2316; Alnus glutinosa, Laurus

nobilis, SANT-Lich. 2314, 2315; Fraxinus excelsior y Eucalyptus globulus,

SANT-Lich. 2312, 2313.

AGRADECIMIENTOS Al Dr. Р.М. Jorgensen quien ha revisado las especies de los

géneros Collema, Leptogium, Pannaria y Parmeliella y al Dr. B.J. Coppins por la

confirmación de Polychidium dendriscum (Nyl.) Hensen.

BIBLIOGRAFIA

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(Santiago) 6: 17-29.

CARBALLAL R. & GARCIA-MOLARES A., 1988 a - Fragmenta Chorologica Occiden-

talia (Liquenes), 1852-1866. Anales Jard. Bot. Madrid 45 (2): 521-523.

CARBALLAL В. & GARCIA-MOLARES A., 1988 b - Acercamiento a las comunidades

liquénicas epifiticas del entorno urbano de la ciudad de Pontevedra. Bol. Soc, Brot. 61

(En prensa).

CLAUZADE G. & ROUX С., 1985 - Likenoj de Okcidenta Europo. Royan: Société Bota-

nique du Centre-Ouest. 893p.

COLMEIRO M., 1889 - Enumeración y revisión de las plantas de la peninsula hispano-lusi-

tana e islas Baleares (tomo V, Liquenes 758-875). Madrid.

CRESPO A., BARRENO E., SANCHO C.G. & BUENO A.G., 1981 - Establecimiento de

una red de valoración de la pureza atmosférica en la provincia de la Согийа (Espana)

mediante bioindicadores liquénicos. Lazaroa 2: 287-311.

DEGELIUS G., 1954 - The lichen genus Collema in Europe: morphology, taxonomy and

ecology. Symb. Bot. Upsal. 13 (2): 1-499.

ETAYO J., 1986 - Liquenes epifitos navarros nuevos o interesantes para la peninsula.

Publ. Biol. Univ. Navarra. Ser. Bot. 6: 29-39

FOLLMANN G., 1976 - Lichen Flora and Lichen Vegetation of the Canary Islands. In G.

KUNKEL, Biogeography and Ecology in the Canary Island. The Hague. pp. 267-286.

FONT М.А. & FIOL LL.A., 1985 - Liquens epifits de Quercus ilex de Mallorca (П). Boll.

Soc. Hist. Nat. Baleares 28: 41-58.

HERNANDEZ-PADRON C., 1985 - Novedades corológicas para la flora liquénica epifita

de Canarias. I: Sabinares de El Hierro. Vieraea 14 (1-2): 113-130.

Source : MNHN, Paris

252 М.Е. LOPEZ DE SILANES y R. CARBALLAL

HERNANDEZ-PADRON C., SANCHEZ-PINTO L. & FOLLMANN G., 1987 - Zur

Kenntnis der Flechtenflora und Flechtenvegetation der Kanarischen Inseln VIL. Areal-

typen und Verbreitungsmuster einiger Neufunde. Cour. Forsch. Inst. Senckenberg 95:

189-199.

JORGENSEN P.M., 1978 - The lichen family Pannariaceae in Europe. Opera Bot. 45:

1-113.

JORGENSEN P.M. & JAMES P.W., 1983 - Studies on some Leprogium species of western

Europe. Lichenologist 15 (2): 109-125.

LACOIZQUETA J.M., 1885 - Catálogo de las plantas que espontáneamente crecen en el

Valle de Vertizarana. Anales Soc. Esp. Hist. Nat. Mem., ser. 1, 14: 185-238.

LLIMONA X., 1976 - Prospecciones liquenológicas en el alto Aragón occidental. Collect.

Bot. (Barcelona) 10: 281-328.

LOPEZ DE SILANES М.Е. & CARBALLAL R., 1987 - Líquenes epifitos de Caaveiro,

La Coruña (España), I. Cryptogamie, Bryol. Lichénol. 8 (4): 359-367.

MAHEU J. & GILLET A. 1921 - Contribution à l'étude des lichens des iles Baléares.

Bull. Soc. Bot. France 68: 426-436, 516-525.

MARCOS В., 1985 - Flora y vegetación liquénicas de las sierras meridionales salmantinas.

Tesis Doctoral. Univ. Salamanca.

SANCHEZ-PINTO L., HERNANDEZ-PADRON С., PEREZ DE PAZ 11. & FOLL-

MANN G., 1983 - Notas corológicas sobre la flora liquénica de las islas Canarias П.

Vieraea 12 (1-2): 233-248.

SEAWARD M.R.D., 1983 - Lichens of Málaga province, S. Spain. Nova Hedwigia 37:

325-345.

VAZQUEZ V. & CRESPO A., 1978 - Catálogo de liquenes de Asturias. I. Epifitos. Acta

Bot. Malacitana 4: 11-26.

Source : MNHN. Paris

Cryptogamie, Bryol. Lichénol., 1989, 10 (3): 253-266 253

ETUDE COMPARÉE DE LA VEGETATION

BRYOPHYTIQUE

DES TRONCS DE CHÉNE VERT ЕТ DE CHEN

PUBESCENT (PEUPLEMENTS ÁGÉS)

DANS LA FORET DOMANIALE DE LA GARDIOLE

DE RIANS (VAR, FRANCE)

J.P. HEBRARD

Institut méditerranéen d'écologie et de paléoécologie,

Laboratoire de botanique et d'écologie méditerranéenne,

Faculté des sciences et techniques de Saint-Jéróme,

avenue de l'escadrille Normandie-Niemen, F-13397 Marseille.

RESUME - Étude de la végétation bryophytique des trones de chêne vert et de chêne

pubescent (аде: environ 61-66 ans en janvier 1984), sur deux niveaux de hauteur, dans la

forêt domaniale de la Gardiole de Rians (Var, France). La surreprésentation des mousses

(au moins 75% du total des taxons) par rapport aux iques et la trs grande pauvreté

des communautés traduisent la xéricité du milieu ambiant, peu favorable aux épiphytes.

Les résultats des inventaires montrent qu'un seul ensemble de bryophytes, constitué

d'un fond constant de trois taxons à large amplitude écologique, d'une hépatique xérophile

et d'une mousse mésophile, occupe le niveau inférieur des troncs des deux espèces de ché-

mes. Il en est de même pour les parties hautes où la communauté, constituée de plusieurs

Orthotrichum, est trés spécialisée (сһепе vert: 62% de corticoles, chéne pubescent: 72,73%),

mais demeure extrémement fragmentaire sur Quercus ilex.

Enfin, quatre groupements (à Frullania dilatata et Radula complanata sur Quercus ilex,

à Homalothecium sericeum et Frullania dilatata sur Quercus pubescens, à Hypnum

cupressiforme var. filiforme ou à Hypnum cupressiforme var. cupressiforme sur les deux

chénes) ont été décrits antérieurement pour la classe d'áge 6-31 ans. On peut déduire de la

comparaison des données, que le vieillissement des phorophytes s'accompagne d'une dimi-

nution de la richesse des communautés, dont le recouvrement reste important

ABSTRACT - A study of the bryophytic vegetation of holm-oak and pubescent oak trunks

(age of stands: ca. 61-66 years in january 1984), on two height levels in the forest of La

Gardiole de Rians (Var, France). The overrepresentation of mosses (at least 75% of the

total number of taxa), compared to liverworts, and the very great poverty of the communi-

ties, express the dryness of the environment, Which is unfavourable to epiphytes

The results of the inventories show that a single unit of bryophytes, made of a constant

set of three taxa with a wide ecological amplitude, a xerophilous hepatic and a mesophilous

moss, occupies the lower level of trunks on both oak species. The same is true for the up-

Source : MNHN, Paris

254 J.P. HÉBRARD

per level whose community, constituted of several Orthotrichum, is very specialized (holm-

oak: 62% of corticoles, pubescent oak: 72,2%), but remains fragmentary on Quercus ilex.

Finally, four groupings (Frullania dilatata - Radula complanata community on Quercus

ilex, Homalothecium sericeum - Frullania dilatata community on Quercus pubescens, Hyp-

mum cupressiforme var. filiforme or Hypnum cupressiforme var. cupressiforme ones on

both tree species) have been distinguished for the lower level of trunks, according to the

frequency and measured dominance of their components. The first three have been previ-

ously described for forests belonging to the 6-31 years age group. It can be deduced from

the comparison of the data, that phorophyte ageing is accompanied by a decrease in the

richness of the communities, whose cover remains important.

INTRODUCTION

Dans le cadre des activités du GRECO 130043 du C.N.RS. "Ecologie des

forêts méditerranéennes”, des recherches pluridisciplinaires ont été entreprises sur

plusicurs sites de basse Provence.

Elles comportent, entre autres, une analyse typologique des formations

végétales qui correspondent aux différentes étapes de maturation forestiére des

écosystémes, étude dans laquelle les bryophytes ont été prises en considération,

compte tenu de leur importance en tant qu'indicateurs biologiques.

La forêt domaniale de la Gardiole, située entre Pourrières et Rians (Var)

s'étend, sur environ 714һа, à l'intérieur d'une aire limitée du nord au sud et

d'ouest en est par des points dont les coordonnées (en grades) sont les suivantes:

3,720 E х 48,415 №, 3,746,E x 48,395 №, 3,776 E x 48,429/N et 3785 E x

48,401 Х.

L'altitude varie de 380 à 630m et les affleurements géologiques, constitués de

calcaires compacts, appartiennent au Jurassique supérieur.

Du point de vue climatique, par référence à la localité la plus proche

(Pourrières, période 1951-1970), la moyenne annuelle atteint 746,7mm pour la

pluviométrie (régime AHPE) et + 13,1%C pour la temperature.

Nos investigations n'ont concerné que des parcelles forestiéres correspondant

à des taillis de Quercus ilex L. (Quercetalia ilicis Braun-Blanquet (1931) 1936)

ou à des chênaies pubescentes (Quercetalia pubescentis Braun-Blanquet 1932)

ägés d'environ 61-66 ans en janvier 1984 (d'après Miglioretti 1983).

La localisation géographique des stations étudiées ici, dont la numérotation

(tableaux 1 et 3) est identique à celle que nous avons utilisée dans deux contribu-

tions antérieures, n'a pas été rappelée.

Nous renvoyons donc le lecteur à ces travaux: Hébrard et Rolando (1985: 93,

fig. 1, stations 16 à 20) pour les taillis de chêne vert et Hébrard (1987а: 115, fig.

1, stations 11 à 15) pour les chénaies pubescentes.

(1) Ces limites d'âge sont évidemment approximatives et ne concernent que l'ensemble du

peuplement arborescent de chaque station. Précisons que, pour des raisons matérielles,

nous n'avons pas sondê chacun des arbres étudiés afin d'en connaitre l'age.

Source : MNHN, Paris

BRYOPHYTES DE LA FORET VAROISE 255

Dans chaque station, afin de disposer de données exhaustives sur la compos

tion floristique des groupements bryophytiques de l'écorce, un inventaire détaillé

a été effectué à deux niveaux de hauteur par rapport au sol (0-(46) 60cm et

42-(300) 700cm) sur les troncs (pente 90°) de 4 à 5 individus de chêne vert ou de

chêne pubescent, dont la hauteur évaluée ( Quercus ilex: 3 à 4,5m, Quercus

pubescens: 5 à Tm) et la circonférence à la base correspondaient à la moyenne

[нле quercus ler wu

pp | ° | à [ense] эз | me Lk [ien | er] 6

Жаша مت‎ en cat” aote | жем | ous | мәз | нен |] 20-05 | же | 50-60 | 10:50 | 50.60

(sait sonptansce КІР EP e ТТР ы

GE si) Pese ENE x еШ emend ise

[sur caprezeiforme var.

Ee e eee Je Sere ІЗ el a Na

|rruttania dilatato 4 ЭЖ e | = O | «ғ

Los Soungarenart К i] | 4 fem |. cm:

(pur cupraesifome var.

(Ste ‚|+ Ite cos. | т

бойо эттм . SSC ы

een nésopnties

[ipeum сирена farm var

дим S 5 p

asia sonptanara А RS Sg

ec affine var.

GE 5 . E

Ke Autoren ai pes

zl 42-300] mell 16-400] 51-800] 61-100) 51=600| 61-500

[axons zët

гата dilatata s fx] sm temi ЕЕЕ ЕЕ

[ota tamiptia var

Ge ^ ala > | &.

eener i 1 ыш

be diaphanum " 5 piro

(bre perpuett tue Е E»

EE + |e

(Киото tenet ae + =

tasona эркез

bert affine var,

Ge а : E Мы

(нта ө 3

ауаға theatre ee] oe

Tableau 1 - Inventaire des bryophytes rencontrées sur les trones (pente: 90°, 4 à 5 individus

par station) de chéne vert et de chéne pubescent, depuis la base jusqu'à 2-7m au-dessus

du sol, dans la forêt domaniale de la Gardiole de Rians (peuplements ages d'environ 61-

66 ans en janvier 1984). A: affinités selon le type de substrat, Co. = corticole, Co.Sax

= cortico-saxicole, In. = indifférent; B: distribution mondiale des taxons, Circb.

circumboréale, Cosm. cosmopolite, Em. = euryméditerranéenne, М.А.

méditerranéenne - atlantique, Or.A. = oréoatlantique, Scosm. = subcosmopolite, Sm.

— subméditerranéenne.

Source : MNHN. Paris

256 J.P. HÉBRARD

du peuplement. N'ont été envisagées, dans les conditions d'exposition globale de

la station (chéne vert, N + NW + ENE: 80% des observations, W: 20%; chéne

pubescent, N + NW: 100%), que les parties occupées par les bryophytes à

l'exclusion de celles ой dominent les lichens.

Enfin, la typologie des groupements muscinaux du niveau inférieur des troncs

(jusqu'à 40-50ст au-dessus du sol), faisant intervenir la dominance des taxons, a

été étudiée.

А cet effet, après avoir noté l'exposition de la surface choisie (entre 451 et

950cm?, moyenne: 685,70 + 151,27cm?), оп a mesuré le recouvrement de chaque

bryophyte, en ramenant les aires à des figures géométriques simples (principale-

ment rectangle et carré, plus rarement triangles).

STRUCTURE DE LA VEGETATION BRYOPHYTIQUE DES ÉCORCES

DE СНЕХЕ VERT ET DE CHENE PUBESCEN

DANS LA FORÉT DOMANIALE DE LA GARDIOLE DE RIANS

(tableaux 1 et 2, figures 1, 2 et 3)

Nous avons volontairement limité cette étude aux chénaies ágées d'environ

61-66 ans à l'époque de nos travaux de terrain. En effet, le dépouillement des

inventaires réalisés pour la classe d'áge 41-46 ans apporte peu de choses.

Toutefois, Tortula virescens, Pottiaceae non encore signalée dans le secteur, а

été découvert sur l'écorce de jeunes Quercus pubescens (hauteur: 5-6m) dans la

station 2-2284 (altitude: 460m, exposition: indéfinie), en compagnie des taxons

suivants.

- Depuis le niveau du sol jusqu'à 40cm: Bryum flaccidum, Frullania dilatata,

Homalothecium lutescens, H. sericeum, Hypnum cupressiforme var. cupres-

siforme et var. strictifolium, Leucodon sciuroides, Orthotrichum acuminatum, O.

affine var. fastigiatum, O. diaphanum, O. lyellü, O. striatum, O. tenellum, Tortu-

la laevipila var. laevipila, T. princeps, T. virescens.

- entre 40 et 100cm: Frullania dilatata.

Précisons enfin que les observations ont été effectuées dans des limites altitudi-

nales peu différentes (maximum: 620m pour le chéne vert et 520m pour le chéne

pubescent!" minimum: 510 et 490m, moyenne: 548 et 500m).

Végétation bryophytique du niveau inférieur des troncs

de chéne vert (0-46cm) et de chéne pubescent (0-60cm)

Dans les stations inventories, le peuplement muscinal propre au niveau

inférieur des troncs a été observé depuis la surface du sol jusqu'à une hauteur

maximale de 46cm sur chéne vert et 60cm sur chéne pubescent.

(1) Par la suite, nous désignerons ces deux chénes par C.V. et C.P.

Source : MNHN, Paris

Taxons

Xérophiles

Mésophiles

A trés large amplitude, vis-à vis de l'humidité

Total

Corticoles

Cortico-saxicoles

Indifférents

Aire de distribution mondiale des taxons :

éléments

Cosmopolite et subcosmopolite

Circumboréal

Eurynéditerranéen

Subméditerranéen

Méditerranéen-atlantique

Oréo-atlantique

Autres

Mousses

Acrocarpes

Pleurocarpes

Hépatiques

Niveau inférieur

10,00

60,00

10,00

80,00

25,00*

15,00%

20,00

30,00

40,00

30,00

100,00

20,00

50,00

30,00

10,00

70,00

10,00

80,00

37,508

62,50%

20,00

33,33

41,67

25,00

100,00

16,67

58,33

25,00

8,33

66,67

8,33

83,33

30,00%

то;00*

16,67

Niveau supérieur

50,00

100,00

62,50

37,50

12,50

37,50

12,50

75,00

83,33*

16,67"

25,00

54,55

45,45

100,00

12,73

27,27

9,09

45,45

9,09

18,18

9,09

90,91

90,00%

10,00%

9,09

53,85

46,15

100,00

69,23

30,77

15,38

38,16

1,69

15,38

7,69

HSIOYVA 32103 УЛ ЗА 541АНАОАЯЯ

84,62

81,82%

18,18*

15,38

Tableau 2 - Nombre et pourcentage de taxons de bryophytes rencontrés sur les trones (niveaux inférieur et supérieur) dans les

taillis de chêne vert (C.V.) et les peuplements de chêne pubescent (C.P.) agés d'environ 61-66 ans en janvier 1984 (Forêt do-

le de la Gardiole de Rians). Répartition selon les préférences écologiques (humidité, type de substrat) et l'appartenance

mania

aux aires de distribution mondiale; mousses acrocarpes, pleurocarpes, et hépatiques. *: en % du nombre total de mousses.

LST

Source : MNHN, Paris

258 J.P. HÉBRARD

Néanmoins, dans toutes les yeuseraies considérées et dans deux chénaies pu-

bescentes (stations 11 et 12), il ne s'étend. souvent que jusqu'à 20 (30)cm sur

beaucoup d'arbres.

Le tableau 2 fait apparaitre de grandes analogies dans la structure du peuple-

ment bryophytique épiphyte des deux Quercus.

On peut noter en particulier les faibles pourcentages de corticoles (10 à 20%

du total de taxons), la forte proportion de muscinées à aire de distribution mon-

diale vaste (70 à 80%), notamment les circumboréales (C.V.: 60%, С.Р: 70%)

et la rareté des méditerranéennes au sens large (C.V.: 20%, C.P.: 10%).

En outre, la prépondérance des mousses (80%) sur les hépatiques traduit bien

la xéricité du milieu ambiant (bioclimat méditerranéen). Réciproquement, les

differences sont discrétes. Tout au plus peut-on admettre que la représentation

des xérophiles (C.V.: 40% du total de taxons, C.P.: 30%) et des cortico-saxicoles

(С.У.: 70%, C.P.: 50%) est meilleure sur Quercus ilex qui colonise des biotopes

plus secs et plus dégradés (affleurements rocheux importants).

D'autre part, le peuplement muscinal est trés pauvre (nombre maximum de

taxons par relevé, C.V.: 6, С.Р.: 7, nombre minimum: 4 et 5, nombre moyen:

5,4 + 0,80 et 5,8 + 0,98).

Les proportions de bryophytes à fréquence trés faible (F = 20% des relevés,

C.V.: 30% du nombre total de taxons, C.P.: 40%) ou au contraire forte et trés

forte (Е = 80-100% des relevés, C.V.: 40% du nombre total de taxons, C.P:

50%) sont voisines pour les deux phorophytes (figure 1). Les secondes, un peu

plus nombreuses, sont représentées essentiellement par trois ubiquistes (Radula

complanata, Homalothecim sericeum, Hypnum cupressiforme var. cupressiforme)

et une hépatique xérophile ( Frullania dilatata), auxquels s'adjoint Hypnum

cupressiforme var. filiforme (mésophile), constant sur chêne pubescent (F =

60% des relevés en С.У. et 100% en C.P.).

Enfin, le nombre de familles de bryophytes est très réduit (C.

leur richesse en taxons différe peu d'un chêne à l'autre (figure 3A).

6, C.P.: 5) et

Végétation bryophytique du niveau supérieur des troncs

de chêne vert (42-300cm) et de chêne pubescent (46-700cm)

Sur les deux Quercus (tableau 2), les pourcentages de xérophiles (50% et

54,55% du nombre total de taxons) et de mésophiles (50% et 45,45%) sont

proches, alors que les muscinées à large amplitude écologique et les indifférentes

au type de substrat font défaut.

En ce qui concerne les xérophiles et les mésophiles, ces résultats peuvent

s'expliquer du fait des caractéristiques du substrat et du milieu ambiant (compen-

sation): écorce retenant moins bien l'eau, mais éclairement plus réduit (feuillage

persistant en hiver) dans le cas de Quercus ilex et au contraire écorce plus riche

en eau mais éclairement intense des parties moyennes et hautes des troncs en

hiver et au printemps, dans le cas de Quercus pubescens.

Source : MNHN, Paris

BRYOPHYTES DE LA FORET VAROISE 259

Nombre de taxons (% di tal: 10) Nombre de taxons(% du otl 10)

A" CN. С.Р.

» poy

n Els

0 | 4 |

| | Es

E GEET EE

Fréquence (% du nombre de stations . 5)

Fig.1. NIVEAU INFERIEUR DES TRONCS

Figure 1 - Histogrammes des fréquences des bryophytes rencontrées sur le niveau inférieur

des troncs de chéne vert (C.V.) et de chéne pubescent (С.Р.) dans la forét domaniale de

la Gardiole de Rians (inventaire portant sur 4 à 5 individus par station).

De méme, la représentation des cortico-saxicoles (37,50% et 27,27%), des

éléments à vaste aire de distribution mondiale (50% et 54,54%, circumboréales:

37,50% et 45,45%), des méditerranéennes au sens large (euryméditerranéennes

+ submediterraneennes + méditerranéennes - atlantiques, C.V.: 37,50%, C.P.:

27,27%) et des mousses (C.V.: 75%, C.P.: 90,91%) par rapport aux hépatiques

sont trés comparables.

Toutefois, les écorces lisses du chéne vert semblent un peu moins propices que

celles du chéne pubescent, épaisses et crevassées, à l'installation des corticoles

strictes (C.V.: 62,50% du total de taxons, С.Р.: 72,73%). D'ailleurs, la richesse

en bryophytes est plus grande avec Quercus pubescens, notamment si l'on com-

pare les nombres moyens par relevé (C.V.: 2,8 + 2,23, C.P.: 5,4 + 1,85).

Le peuplement est aussi mieux structuré sur cet arbre (figure 2) puisque, par

rapport à ce que l'on note pour le chéne vert, les taxons à fréquence trés faible et

faible (20-40% des relevés) ont une importance plus réduite (C.V.: 87,50% du

nombre total de taxons, C.P.: 54,4%), contrairement à ceux dont la fréquence

est forte et trés forte (80-100% des relevés, C.V.: 12,5% du nombre total des tax-

ons, C.P.: 27,27%).

Mis à part l'hépatique cortico-saxicole et xérophile Frullania dilatata, qui est

absolument constante sur les deux chénes, et méme parfois seule présente sur le

niveau supérieur des troncs de certaines yeuseraies (stations 16 et 18, tableau 1);

le groupement est constitué de corticoles strictes comme Orthotrichum lyellii (F,

C.V.: 20%, des relevés, C.P.: 100%), O. affine var. fastigiatum (F, C.V.: 40%,

C.P.: 80%) et plus accessoirement Orthotrichum striatum (F, CAN: 4 SP.

60%) ou Tortula laevipila var. laevipila (F, C.V.: 20%, C.P.: 60%). П n'es!

Source : MNHN. Paris

260 J.P. HEBRARD

m} Nombre de taxons (% du total.: 8) Nombre de taxons (% du total :11)

cv ce

шош a mw шош D “ mo w

Fréquence (% du nombre de stations : 5)

Fig. 2.NIVEAU SUPERIEUR DES TRONCS

Figure 2 - Histogrammes des fréquences des bryophytes rencontrées sur le niveau supérieur

des troncs de chêne vert (C. V.) ou de chêne pubescent (C.P.) dans la forêt domaniale de

la Gardiole de Rians (inventaire portant sur 4 à 5 individus par station).

done parfaitement individualisé que sur Quercus pubescens. Parmi les espéces ac-

cidentelles, il convient de signaler Ulora crispa, rencontré une seule fois sur un

gros chéne pubescent dans un ravin encaissé (station 11). Cette mousse, dont la

présence indique une humidité atmosphérique élevée, du moins en dehors de la

saison estivale, n'avait encore jamais été signalée en basse Provence.

Notons enfin (figure 3B) que le nombre de familles de bryophytes est faible

(familles représentées par un seul taxon, C.V.: 4, C.P.: 3), celle des Orthotricha-

ceae regroupant le plus grand nombre de taxons, presque tous corticoles (C.V.: 4

taxons, C.P.: 8).

PRINCIPAUX GROUPEMENTS BRYOPHYTIQUES

DU NIVEAU INFÉRIEUR DES TRONCS DE CHÉNE VERT

_ ET DE CHÊNE PUBESCENT, ETABLIS D'APRES

LA FRÉQUENCE ET LA DOMINANCE DE LEURS COMPOSANTES

(tableau 3)

Hormis le groupement à Hypnum cupressiforme var. cupressiforme, les unités

décrites dans un travail antérieur (Hébrard 1987b) pour des peuplements

Source : MNHN. Paris

BRYOPHYTES DE LA FORET VAROISE 261

d'arbres jeunes (6-31ans) de la forét domaniale de la Gardiole de Rians, se re-

trouvent sous une forme appauvrie dans les chénaies plus ágées (61-66 ans).

Toutes s‘intégrent aux Leucodontetalia (Von Hübschmann 1952) Lecointe

1975 et au Frullania dilatatae Lecointe 1975.

Groupement à Frullania dilatata et Radula complanata

H n’a été rencontré que sur Quercus ilex (sta

ах

Le recouvrement muscinal représente 45 et 60% de la surface de mesure. Се

groupement, avec seulement 3 taxons constants, est constitué par les xérophiles

Vrullania dilatata et Hypnum cupressiforme var. strictifolium, dominants par

rapport à l'hépatique à vaste amplitude écologique Radula complanata.

ions 19 et 20), en exposition N

Groupement à Hypnum cupressiforme var. filiforme

Il est fréquent sur les parties des troncs exposées au nord ou à l'est et son re-

couvrement atteint des valeurs plutót élevées (38,95% à 88,55% de la surface de

mesure, moyenne: 65,19 + 17,79%). Т ‘outefois, il est d'une grande pauvreté, avec

À Nombre де!атйев

A Me, в

на

122

Nombre de taxons

Figure 3 - Histogrammes de distribution du nombre de familles de bryophytes selon leur si

‘chesse en taxons présents sur les troncs de chêne vert (C.V.) et de chêne pubescent

(CP) dans la forêt domaniale de la Gardiole de Rians (inventaire dans des peuplements

forestiers ages d'environ 61-66 ans en janvier 1984) A: niveau inférieur des dones

(EV: 0-46em, C.P 0-60em), B: niveau supérieur (С.У 42-300cm, C.P.: 46-700ст);

Brachytheciaceae, Fa. - Fabroniaceae, Fr. = Frullaniaceae, Ну. = Hypnaceae,

LE = Laucodontaceae, Me. = Metzgeriaceae, Ne. = Neckeraceae, Or. = Orthotricha-

сеге, Po. = Pottiaceae, Ra. = Radulaceae.

Source : MNHN. Paris

797

Numéros des stations

Code des stations

Exposition (surface de mesure)

Intervalle de hauteur à partir du

sol (сп)

Surface totale étudiée (cm2)

Recouvrement muscinal (X de la surface

totale étudiée)

Différentielles des groupements

Frullania dilatata

Radula complanata

Hypnum cupressiforme var. filiforme

Hypnum cupressiforme var. cupressiforme

Homalothecium sericeun

Autres taxons

Hypnum cupressiforme var. strietifolium

Neckera complanata

Orthotrichum striatum

Orthotrichum affine var. fastigiatum

19 20 16 18 14 12

3-153 | 4-263 | 2-143 | 2-153 | 2-263 | 4-63

N NE NE E NW N

0-41 0-41 0-45 0-41 0-50 0-45

451 533 | 585 615 950 585

59,87 | 45,40 | 88,55 | 62,93 | 38,95 | 70,34

52,96 | 78,51

бт 3,31

46,67 | 18,18 T .

17 16 11 13

1-153 | 3-263 | 3-63 | 1-263

NE SE H SE

0-46 0-50 0-19 0-40. 3

736 800 882 720 ч

30,03 | 58,03 | 28,40 | 44,03) m.

31,14 | 3,47 S

17,96 4

98,87 | 50,30 2

л Т 96,53

21,15 ;

и 1,13 ?

. D 0,60

Tableau 3 - Groupements museinaux du niveau inférieur des trones de chêne vert et de chêne pubescent (peuplements âgés de

61-66 ans en janvier 1984) dans la forét domaniale de la Gardiole de

en % de la surface totale occupée par les bryophytes.

Rians. Pour chaque taxon, la dominance est exprimée

Source : MNHN. Paris

BRYOPHYTES DE LA FORET VAROISE 263

seulement les constantes Hypnum cupressiforme var. filiforme (mésophile), Frul-

lania dilatata et Radula complanata.

Sur chéne vert (stations 16 et 18), la seconde de ces muscinées, nettement

xérophile, domine (D: 47,68% et 80,62% de la surface totale occupée par les

bryophytes, contre 41,89% et 18,60% pour Hypnum cupressiforme var.

filiforme), alors que l'inverse est constate sur chéne pubescent (stations 14 et ner

D: 90.27% et 75,82% pour Hypnum cupressiforme var. filiforme contre 9,46%

et 21,87% pour Frullania dilatata).

Groupements à Hypnum cupressiforme var. cupressiforme

et à Homalothecium sericeum et Frullania dilatata

Ils remplacent les groupements précédents dans des biotopes plus xériques et

notamment sur les parties basses des troncs de Quercus pubescens exposées au

sud et à l'ouest. Tous deux sont floristiquement très pauvres (2 à 4 taxons par

relevé).

Le premier (recouvrement muscinal: 28,40% à 58,03% de la surface de me-

sure) est caractérisé par la dominance de l'ubiquiste Hypnum cupressiforme var.

cupressiforme dont la valeur, plutót faible sur Quercus ilex (D: 3,62%, station

17), est au contraire considérable sur Quercus pubescens (D: 50, 30% et 98,87%,

stations 11 et 13).

Enfin, le second groupement, rencontré sur Quercus pubescens, est presque

entiérement constitue de colonies denses 4” Homalothecium sericeum (D: 96,53%

et recouvrement muscinal: 44,03% de la surface de mesure).

CONCLUSION

Des prospections effectuées dans les taillis de Quercus ilex et des foréts de

Quercus pubescens âgées d'environ 61-66 ans (janvier 1984) sur le site de la Gar-

diole de Rians (Var) ont permis d'étudier, sur deux niveaux de hauteur par rap-

port au sol, la structure de la végétation bryophytique peuplant les troncs de ces

Penes. L'inventaire approfondi des muscinées épiphytes a été réalisé en 10 sta-

tions (4 à 5 arbres par station).

Les résultats obtenus montrent que, pour le niveau inférieur (46 à 60cm au-

dessus du sol) le peuplement bryophytique est pratiquement identique sur les

deux phorophytes.

D'une grande pauvreté (total, С.У. ou C.P.: 10 taxons, C.V, + C.P.: 12) il

est caractérisé fidelement par Radula complanata, Homalothecium sericeum,

Hypnum cupressiforme var. cupressiforme, à large amplitude écologique,

accompagnés par Frullania dilatata (xérophile) et Hypnum cupressiforme var.

filiforme (mésophile), ce dernier plus constant avec le chêne pubescent.

En ce qui concerne le niveau supérieur (entre 42 et 300-700cm au-dessus du

sol), le peuplement muscinal n'est bien individualisé que sur Quercus pubescens

Source : MNHN, Paris

264. J.P. HÉBRARD

(total, C.V.: 8 taxons, C.P.: 11, С.У. + C.P.: 13), dont l'écorce rugueuse est plus

favorable à l'installation des bryophytes.

Les muscinées les plus constantes sur ce chéne sont en particulier:

Orthotrichum Iyellü, O. affine var. fastigiatum (corticoles, mésophiles et photo-

philes) et Frullania dilatata (cortico-saxicole et xérophile), auxquels s'adjoignent

Tortula laevipila var. leavipila (corticole et xérophile) et Orthotrichum striatum

(corticole, mésophile et photophile).

D'autre part, dans une ambiance plutót xérique (bioclimat méditerranéen,

dégradation récente, Z.P.: 83,33% de mousses pour le niveau inférieur

des trones et 84,62% pour le niveau supérieur, xérophiles: 33,33% et 53,85%),

des différences apparaissent lorsqu'on examine la structure de la végétation bryo-

phytique des deux niveaux de hauteur, qui correspondent chacun à des condi-

tions écologiques particuliéres (humidité et apport d'éléments minéraux prove-

nant du sol plus importants, mais irement beaucoup plus faible pour les

parties basses des troncs).

Ainsi, le peuplement du niveau supérieur est trés spécialis

ticoles: 69,23% du nombre total de taxons, indifferentes au type de substrat: 0,

contre 16,67% et 25% pour le niveau inférieur), puisque les Orthotrichaceae

représentent 61,54% des taxons dénombrés pour l'ensemble С.У. + С.Р. (25%

pour le niveau inférieur).

En conséquence, les mousses acrocarpes (C.V. + C.P.: 81,82% du total des

mousses) y sont bien représentées, alors qu'au contraire les Pleurocarpes,

favorisées par la proximité du sol et indiquant une humification plus avancée du

substrat, l'emportent largement sur le niveau inférieur des troncs (C.V. + C.P.,

Brachytheciaceae + Hypnaceae + Neckeraceae: 70% du total des mousses).

Enfin, l'analyse de relevés comportant la mesure précise de la dominance de

chaque muscinée a permis de distinguer quatre groupements sintegrant au

Frullanion dilatatae Lecointe 1975 | Leucodontetalia (Von Hübschmann 1952)

Lecointe 1975].

Mis à part le groupement à Hypnum cupressiforme var. cupressiforme (Quer-

cus ilex et Q. pubescens), les communautés à Frullania dilatata et Radula com-

planata (Quercus ilex), à Hypnum cupressiforme var. filiforme (sur les deux

chénes) et à Homalothecium sericeum et Frullania dilatata (Quercus pubescens),

trés appauvries, mais dont le recouvrement est important, ont déjà été décrites

pour la classe d'áge 6-31 ans (Hebrard 1987b).

Liste des bryophytes citées

Nomenclature conforme dans l'ensemble à Corley et al. (1982) et à Smith

(1978) ou à l'Index Muscorum: taxons infraspécifiques, pour les mousses, ainsi

qu'à Grolle (1983) pour les hépatiques.

Mousses. - Bryum flaccidum Brid., Habrodon perpusillus (De Not.) Lindb.,

Homalothecium lutescens (Hedw.) Robins., Homalothecium sericeum (Hedw.) B.,

S. et G., Hypnum cupressiforme Hedw. var. cupressiforme, Hypnum cu-

pressiforme Небу. var. filiforme Brid., Hypnum cupressiforme Hedw. var.

Source : MNHN, Paris

BRYOPHYTES DE LA FORET VAROISE 265

strictifolium Warnst, Leptodon smithii (Hedw.) Web. et Mohr, Leucodon

sciuroides (Hedw.) Schwaegr., Neckera complanata (Hedw.) Hüb., Orthotrichum

acuminatum Philib., Orthotrichum affine Brid. var. fastigiatum (Brid.) Hüb.,

Orthotrichum lyellii Hook, et Tayl., Orthotrichum stramineum Horsch, ex Brid.,

Orthotrichum striatum Medw., Orthotrichum tenellum Bruch ex Brid., Tortula

laevipila (Brid.) Schwaegr. var. laevipila, Tortula princeps De Not, Tortula

virescens (De Not.) De Not., Ulota crispa (Hedw.) Brid., Zygodon baumgartneri

Malta.

Mépatiques. - Frullania dilatata (L.) Dum., Metzgeria furcata (L.) Dum.,

Radula complanata (L.) Dum.

BIBLIOGRAPHIE

BARKMAN J.J., 1958 - Phytosociology and ecology of cryptogamic epiphytes. Assen:

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CORLEY M.F.V., CRUNDWELL A.C., DULL R., HILL М.О. & SMITH A.J.E., 1982 -

Mosses of Europe and the Azores; an annotated list of species, with synonyms from the

recent literature. J. Bryol. "1981" 11 (4): 609-689.

GOUNOT M., 1969 - Méthodes d'étude quantitative de la végétation. Paris: Masson.

314p.

GROLLE R., 1983 - Hepatics of Europe including the Azores: an annotated list of species,

with synonyms from the recent literature. J. Bryol. 12 (3): 403-459.

HEBRARD J.P., 1973 - Étude des bryoassociations du Sud-Est de la France et de leur con-

texte écologique. Thèse Doct. Etat, Marseille, tome I: 422p., tome Il: 75 tabl., 17 pl.

HEBRARD J.P. et ROLANDO C., 1985 - Btude comparée du peuplement bryophytique

de taillis de chêne vert d'âge different en forêt domaniale de la Gardiole de Rians (Var,

France). Ecol. Medit. 11 (2-3); 87-110, 14 tabl., 6 fig.

HEBRARD J.P., 1987a - Étude comparée du peuplement bryophytique des chénaies pu-

bescentes de "bas fond" et de "plateau" еп forêt domaniale de la Gardiole de Rians (Var,

France). Cryptogamie, Bryol. Lichénol. 8 (2): 109-146, 13 tabl., 7 fig.

HEBRARD J.P., 1987b - Etude comparée de la végétation bryophytique des parties basses

et moyennes des troncs de chéne vert et de chéne pubescent (peuplements jeunes) dans

la forét domaniale de la Gardiole de Rians (Var, France). Bull. Soc. Bot. Centre-Ouest

n.s. 18: 125-144, 6 tabl., 7 fig.

LECOINTE A., 1975 - Étude phytosociologique des groupements de bryophytes épiphytes

de la Brenne Doc. Phytosociol. (Lille) 9-14: 165-195, 1 tabl., 1 fig. + 10 tabl. hors texte.

LECOINTE A., 1979 - Intérêts phytogéographiques de la bryoflore normande: 1 ~ Les

cortéges cosmopolite et méditerranéen s. 1. Bull. Soc, Linn. Normandie 107: 61-70, 1 fig.

LECOINTE A., 1981a - Intérêts phytogéographiques de la bryoflore normande: 2 - Le

cortège atlantique s. 1. Bull. Soc. Linn. Normandie 108: 51-60, | tabl.

LECOINTE A., 1981b - Intérêts phytogéographiques de la bryoflore normande: 3 - Le

cortège circumboréal s. |. Bull. Soc. Linn. Normandie 109: 55-66, 1 tabl

MIGLIORETTI F., 1983 - Phytoécologie des peuplements à Quercus ilex et Quercus

pubescens Willd. en Gardiole de Rians (Var). Approche méthodologique pour évaluer la

phytomasse des taillis de chéne vert. Thése Doct. Зе cycle, Marseille, tome 1: 90р.,

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Source : MNHN. Paris

266 J.P. HÉBRARD

SLACK N.G., 1977 - Species diversity and community structure in Bryophytes: New

York state studies. New York State Museum Bull. 428: 76р., 9 tabl., 30 fig., 40 phot.

SLACK N.G., 1984 - A new look at bryophyte community analysis: field and statistical

methods. J. Hattori Bot. Lab. 55: 113-132, 11 fig.

SMITH A.LE, 1978 - The moss flora of Britain and Ireland. Cambridge: University

Press. 706p., 333 Пр,

VAN DER WIJK R., MARGADANT W.D. et FLORSCHÜTZ P.A., 1959-1969 - Index

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Source : MNHN. Paris

Cryptogamie, Bryol. Lichénol. 1989, 10 (3): 267-275 267

BIBLIOGRAPHIE BRYOLOGIQUE

D. LAMY

Laboratoire de Cryptogamie, 12 rue Buffon, F-75005 Paris

Systématique, Nomenclature

89-249 ABOLIN A.A. - Polytrichum strictum (Polytrichaceae) - an original species or а

modificant P. juniperinum ? Bot. Zurn. (Moscow & Leningrad) 1985, 70(11):

1503-1511, 3 fig., 2 tabl., en russe.

89-250 GUERRA J. y PUCHE F. - Contribucién al estudio de Orthotrichum baldaccii y О.

sardagnanum (Мәсі). Acta Bot. Malacitana 1985, 10: 11-16, 2 pl. (Dept. Bot., Fac.

Ci., Univ. Malaga, Malaga, España).

MEB de la surface foliaire, des stomales de la capsules, des dents du péristome et des

spores d Orthotrichum cupulatum var. sardagnanum et O. cupulatum var. baldaccii, Propo-

sition de O. cupulatum subsp. baldaccii (Bott. et Vent.) comb. nov.

89-251 OCHYRA R. - What is Dichelyma antarcticum C. Muell. ? Polar Research 1987,

8(4); 403-410, 11 fig. (Dept. Bryol. & Lichenol, Inst. Bot, Pol. Acad. Sci, Lubicz 46,

31-512 Krakow, Poland)

Dichelyma antarcticum est syn. de Blindia magellanica C. Muell.

89-252 OCHYRA R. and LIGHTOWLERS P.J. - The South Georgian moss flora: Pittia.

Brit. Aniarct. Surv. Bull. 1988, 80: 121-127, 3 fig. (Ibidem).

Descr., ill, hab., distr. de Virria pachyloma (Mont.) Ochyra précédemment connu en

Géorgie du Sud sous le nom Sciaromium conspissatum (Hook. f. & Wils.) Vitt.

89-253 SCHUSTER К.М. - The aims and achievements of bryopyte taxonomists. Bor. J.

Linn. Soc 1988, 98(3) 185-202 (Dept. Bot, Univ. Massachusetts, Amherst,

Massachusetts 01002, USA).

Dans l'évaluation des phénomènes d'évolution, les données tectoniques peuvent être

pertinentes; la compréhension des mécanismes de dispersion, la sexualité et la biologie de la

reproduction sont essentielles.

Voir aussi: 89-256, 89-259, 89-278, 89-314, 89-317.

Morphologie, Anatomie

Voir: 898-250, 89-252, 89-277, 89-295.

Source : MNHN. Paris

268 BIBLIOGRAPHIE BRYOLOGIQUE

Cytologie, Ultrastructure

89-254 APOSTOLAKOS P. and GALATIS B. - Studies on the development of air pores

and air chambers of Marchantia paleacea. Ш. Microtubule organiz: preprophase-

prophase initial aperture cells - Formation of incomplete preprophase microtubule bands.

Protoplasma 1985, 128(2/3): 120-135, 26 fig. (Inst. Gen. Bot, Univ. Athens, Athens,

GR-15701).

89-255 APOSTOLAKOS P. and GALATIS B. - Studies on the development of the air pores

and air chambers of Marchantia paleacea. IV. Cell plate arrangement in the initial

aperture cells. Protoplasma 1985, 128(2/3): 136-146, 19 fig. (Ibidem)

89-256 BROWN R.C. and LEMMON B.E. - Sporogenesis іп bryophytes. Advances Bryol.

1988, 3: 159-223, 70 fig. (Dept. Biol., Univ. Southwestern Louisiana, Lafayette, LA

70504-2451, USA).

Les spores libres de bryophytes résultent de la sporogenése, caractérisée par des divi-

sions nucléaires et cytoplasmiques précises. La cytomorphogenése génétiquement contrôlée

est programmée dans la prophase 1 du sporocyte. L'ontogénie et la configuration du fuseau

pendant la métaphase I chez les mousses sont inhabituelles et sont liées à la polarité établie

dans la prophase 1. Formation de l'endo- et de l'exospore. Les études ultrastructurales de la

sporogenése revélent quelques caractéristiques importantes pour l'étude de la phylogénie

des bryophytes: 3 types de parois caractérisent les groupes de mousses (Andreaeopsida,

Sphagnopsida et Bryopsida); l'ontogénie de la paroi sporale des Marchantiidae diffère de

celle des Jungermanniidae; les Anthocerotae présentent des affinités avec les mousses et les

hépatiques.

89-257 CAROTHERS Z.B. and RUSHING

bryophyte blepharoplast. Advances Bryol.

Univ. Illinois, Urbana, Illinois 61801, USA).

Les auteurs font état des travaux récents en morphologie du blépharoplaste des

bryophytes ainsi que de leurs recherches personnelles. Comparaison entre les 14 hépatiques,

2 Anthocérotes et les 6 genres de mousses étudiés. Premiere observation du blepharoplaste

d' Haplomitrium gibbsiae. Les caractères essentiels utilisés pour la comparaison sont: forme,

nombre de microtubules et ouverture du spline, strip lamellaire, corps basal avec zones de

transition. Alors que les blépharoplaste des Jungermanniidae presente une grande diversité,

celui des Marchantiidae est d'une relative uniformite.

89-258 DOONAN J.H. and DUCKETT J.G. - The bryophyte cystoskeleton: experimental

and immunofluorescence studies of morphogenesis. Advances Bryol. 1988, 3: 1-31, 32

fig. (Pharmacol. Dept, Robert Wood Johnson Medical School, Busch Campus,

Piscataway, New Jersey, USA).

Investigations immunologiques et microscopie électronique pour l'étude du róle du

cytosquelette dans la morphogenése des bryophytes, notamment chez le protonéma des

Funariales. Polarité pendant la germination de la spore et la régénération du protoplaste,

présence et rôle des microtubules et des microfilaments dans les dómes apicaux du

protonéma, mitose asymétrique. Perspectives de recherches, amélioration des techniques.

89-259 DUCKETT J.G. and RENZAGLIA K.S. - Cell and molecular biology of bryophytes:

ultimate limits to the resolution of phylogenetic problems. Bot. J. Linn. Soc. 1988, 98(3):

225-246 (School Biol. Sci, Queen Mary College, Mile End Road, London El 4NS,

UK).

Il est évident que l'ultrastructure, la biochimie, la morphologie et la morphogenèse

contribuent à une meilleure connaissance de l'origine phylogénétique des bryophytes

L'ultrastructure devrait être plus largement utilisée dans la systématique des bryophytes.

89-260 DUCKETT J.G. and RENZAGLIA K.S. - Ultrastructure and development of

plastids in the bryophytes. Advances Bryol. 1988, 3: 33-93, 63 fig. (Ibidem)

Analyse des micrographies électroniques des plastes de bryophytes publices depuis 25

ans, et données personnelles sur les régions méristématiques de divers bryophytes pour

donner un nouvel éclairage sur les processus de développement des plastes. il n'y a pas de

différences évidentes dans l'ultrastructure des plastes des mousses et des hépatiques ni entre

- Comparative morphology of the

88, 3: 95-134, 1 tabl. (Dept. Pl. Biol,

Source - MNHN, Paris

BIBLIOGRAPHIE BRYOLOGIQUE 269

les gamétophytes et les sporophytes. Sous divers régimes lumineux, la morphologie des

plastes est moins susceptible de changements chez les mousses que chez les plantes

supérieures.

89-261 LIGRONE R. and GAMBARDELLA R. - The sporophyte-gametophyte junction in

bryophytes. Advances Bryol. 1988, 3: 225-274, 42 fig. (Dipto. Biol. veg., Univ. Napoli,

Via Foria 223, 1-80139 Napoli).

Ultrastructure de la région histologique comprenant le pied du sporophyte et la

vaginule entourant le gamétophyte; la région de contact est appelée placenta. Activités

enzymatiques et róle de cette région.

89-262 MUELLER D.M.J. and NEUMANN A.J. - Peristome structure and the regulation

of spore release in Arthrodontous mosses. Advances Bryol. 1988, 3: 135-158, 12 fig.

(Dept. Biol., Texas A & M Univ., College Station, Texas 71843-3258, USA).

Dans la plupart des mousses le peristome est partie intégrante du mécanisme

d'éjection des spores et sert généralement à réguler cette ejection, Les péristomes

arthrodontes sont composés de restes de parois cellulaires des couches du tissu spécifique de

ja capsule qui sont exposés par dehiscence de l'opercule. L'endostome est généralement

considéré comme jouant un rôle passif dans l'éjection des spores, tandis que l'exostome

joue fréquemment un rôle actif, soit en gênant soit en facilitant cette éjection. Les mouve-

ments hygroscopiques qui tendent à ouvrir ou fermer l'ouverture de la capsule sous

différentes conditions spécifiques d'humidité résultent de la construction et de la compo-

sition spécifiques des dents

89-263 NEWTON М.Е. - Heterochromatin diversity in two species of Реа (Hepaticae) as

revealed by C-, О-, N- and Hoechst 33258-banding. Protoplasma 1985, 92(5): 378-386,

1 tabl, 7 fig. (Dept. Cell & Structur. Biol, Williamson Build, Univ. Manchester,

Manchester, M13 9PL, UK).

Mise en évidence de 4 types d'hétérochtomatine chez Pellia neesiana et 2 autres chez

P. epiphylla. Evolution cytologique entre ces deux espèces.

89-264 NEWTON М.Е. - Chromosomes as indicators of bryophytes reproductive performan-

ce. Bot. J. Linn. Soc. 1988, 98(3): 269-275 (Ibidem).

La capacité de reproduction chez les bryophytes est un équilibre entre la monoécie et

la dioécie, entre l'autofécondation et la fécondation croisée, entre la dispersion et le maintien

de la variation génétique adaptative.

89-265 VYSOTSKAYA - Karyotypie structure of the genus Brachythecium B.S

species. Ukrajinsk. Bot. Zurn, 1985, 42(4). 44-47, en russe, гё. angl. (L'viv. Bid-nja,

Int. Bot, im M.G. Holodnogo, AN URSR, Kiev, USSR).

Nombre chromosomique de 16 Brachythecium, diversité intraspécifique pour 10 d'en-

tre eux. П est supposé que x=6 et x = 10 sont des nombres chromosomiques ancestraux.

Physiologie, Chimie

89-266 COVE D.J. and ASHTON N.W. - Growth regulation and development in

Physcomitrella patens : an insight into growth regulation and development of bryophytes.

Bot. J. Linn. Soc. 1988, 98(3): 247-252 (Dept. Genet, Univ. Leeds, Leeds LS2 9/7,

UK)

89-267 ELLIS J.G. and THOMAS R.J. - Phototropism of Pellia : evidence for mediation by

auxin-stimulated acid efflux. J. PL Physiol. 1985, 121(3): 259-264, 3 fig. (Dept. Biol.,

Bater College, Lewiston, Maine 04240, USA).

89-268 LONGTON R.E. - Adaptations and strategies of polar bryophytes. Bot, J. Linn.

Soc. 1988, 98(3): 253-268, 8 fig., 2 tabl. (Dept. Bot., School РІ. Sci., The University,

Reading, RG6 2AS, UK).

Caractères physiol. des bryophytes polaires en relation avec les modes de distr. géogr.,

le climat polaire, et l'histoire de l'environ. des regions polaires. Plasticité phénotypique,

Source : MNHN, Paris

270 BIBLIOGRAPHIE BRYOLOGIQUE

réponses opportunistes dans l'échange du CO, et la circulation de l'eau procurent aux

bryophytes une tolérance considérable à la dessiccation et au froid.

89-269 NEHIRA K. - Germination of gemmae in some mosses collected from North

America. In: Papers of Plant ecology and taxonomy to the memory of Dr Satoshi

Nakanishi. The Kobe Geobotanical Society 1987: 355-360, 2 fig. (Dept. Biol, Fac.

Integrat. Arts & Sci., Hiroshima Univ., Higashi-senda, Naka-ku, Hiroshima 730 Japan).

89-270 NEHIRA К. - Germination and protonemata. In: СІЛМЕ J.M., Methods in

bryology. (Proc. Bryol Meth. Workshop Mainz). Nichinan: Hattori Botanical

Laboratory, 1988, pp. 113-117, 2 fig. (Ibidem).

89-271 RYDIN H. - Effect of water level on desiccation of Sphagnum in relation to

surrounding Sphagna. Oikos 1985, 45(3): 374-379, 1 tabl., 4 fig. (Inst. Ecol. Bot., Univ.

Uppsala, Box 559, S-751 22 Uppsala)

Mesure de la quantité d'eau contenue dans le capitulum des sphaignes pour suivre la

dessiccation pendant les périodes de climat sec et de basses eaux. Variation d'une esp. à

l'autre. Influence de la présence de certaines esp. pour freiner la dessiccation des esp. voisi-

nes.

Répartition, Ecologie, Sociologie

89-272 BOCH M KUZMINA Е.О. - On Sphagnum mosses from the North-West of the

RSFSR. Bot. Zurn. (Moscow & Leningrad) 1985, 70(10); 1337-1346, en russe (Bot.

Inst. V.L. Komarov ANSSSR, Leningrad, USSR).

89-273 CANALÍS V. & CASAS С. - Novetas per а la brioflora dels Pirineus centrals.

Collect. Bot. (Barcelona) 1985, 16(1): 59-61 (Dept. Bot, Fac. Biol, Univ. Barcelona,

E-08028 Barcelona).

Liste de 10 esp. de mousses du Val d'Aran et d'Alta Ribagorça dont cert. sont nouv.

pour la Catalogne.

89-274 DIA M.G., MICELI G. e NOT R. - Check-list delle Epatiche note in Sicilia. Webbia

1985, 39(1): 163-177 (Inst, & Orto Bot. dell'Univ., via Archirafi 38, 1-90123 Palermo).

Liste de 107 hépatiques récoltes en Sicile (d'aprés la littérature et les récoltes récentes).

89-275 DOLL R. - Verbreitung und soziologisches Verhalten von Orthotrichum lineare

Schwaegr. im Norden der DDR. Gleditschia 1985, 13(1): 141-145, 2 tabl., 1 carte (1020

Berlin, Rochstr. 9, DDR).

89-276 FRAHM J.P. (en coll. avec LAMY D, SCHUMACKER R., PHILIPPI G.,

RASTETTER У.) - La bryoflore des Vosges et des zones limitrophes. — Duisbur:

Universitát-Gesamthochschule. 1989. Non paginé, 680 cartes, (Univ. Duisburg,

Fachber. 6, Bot., Postfach 101503, D-4100 Duisburg).

Remarques générales, écol, des bryophytes de la région étudiée: Vosges cristallines et

région du gres bigarre, region des collines, plaines du Rhin, Sundgau. Liste des bryophytes

avec loc. d'après la littérature et des récoltes récentes de Frahm, Philippi, Rastetter ... Cha-

que taxon est cartographié pour les Vosges.

89-277 GEISSLER P. und BISANG I. - Frullania inflata Gott., ein neues thermophiles

Element der Schweizer Moosflora. Saussurea 1985, 16: 95-100, 3 fig. (Conserv. & Jard.

Bot., С.Р. 60, CH-1292 Chambesy/GE).

Descr. de la loc, (Tessin méridional) où a été trouvé Frullania inflata pour la 4° fois en

Europe

89-278 GUERRA J. y РОСНЕ F. - Bryum dunense Smith & Whitehouse en la Peninsula

iberica y Baleares. Observaciones taxonomicas, corologicas y fitosociologicas. Acta Hor.

Malacitana 1984, 9: 85-92, 2 tabl., 1 pl., 1 fig., 1 carte.

89-279 HEBRARD J.P. et ROLANDO С. - Etude comparée du peuplement bryophytique de

taillis de chêne vert d'âge différent en forêt domaniale de la Gardiole de Rians (Var,

Source : MNHN, Paris

BIBLIOGRAPHIE BRYOLOGIQUE 271

France). Ecol, Меди. 1985, 11(2/3): 87-110, 14 tabl. (Lab. Bot. & Ecol. Medit., Fac.

Sci. & Techn. St Jérôme, av. de l'Escadrille Normandie-Niemen, F-13397 Marseille Ce-

dex 13).

Forte proportion de mousses photoxerophiles, de saxicoles, d'esp. de l'étage

mésomédit., de cosmopolites et de circumboréales dans les taillis de chêne vert d'âge

différent en forét domaniale de la Gardiole de Rians. Influence de l'exposition des stations

sur la composition spécifique. Effet litière et xéricité des microclimats limitent les possiblités

d'implantation des bryophytes. Seules 2 mousses à trés large amplitude sont à la fois

constantes et dominantes: Homalothecium sericeum et Hypnum cupressiforme var.

cupressiforme.

89-280 KHMELJEV К.Е., POPOVA N.N. - Rare species of bryophytes іп the Voronezh

Region. Bot. Zurn. (Moscow de Leningrad) 1985, 70(9); 1208-1214, 1 fig., en russe.

89-281 MARSTALLER R. - Die Moosgesellschaften der Ordnung Orthotrichetalia Hadac

in Klika et Hadac 1944. 19. Beitrag zur Moosvegetation Thüringens. Gleditschia 1985,

13(2): 311-355, 10 fig, 13 tabl. (Sekt. Biol, Friedrich-Schiller-Univ., WB Okol., 6900

Jena, DDR).

Structure, composition floristique, des 11 associations de l' Orthotrichetalia en

Thuringe: influence de la pollution; discussion synsystématique, Nouvelle révision de la

classe Frullanio dilatatae-Leucodontetea sciuroidis Mohan 1978 en Europe.

89-282 MIHAI Gh., KAPTANIS E. - Some bryophytes of Lesbos (Greece). Analele Stünt.

Univ. AL I. Cuza lasi, Sect. П а. Biol. (ser. noua), 1984, 30: 55-57.

89-283 NESCHATEYEV V. Yu. - Some associations of paludified pine forests in Leningrad

region. Bot. Zurn. (Moscow 4 Leningrad) 1985, 70(10): 1362-1373, 1 fig, 1 tabl., en

russe.

Bryophytes et lichens associés

89-284 OCHYRA R. and BARYLA J. - Wyginiecie Skorpionowea oblego Scorpidium

turgescens (Musei) w Polsce. (Extinction of the turgid-feather-moss Scorpidium

turgescens (Musci) іп Poland. Chronmy Przyrade Ojczysta 1988, 44(3): 68-69, 3 fig., еп

polon. (Dept. Bryol. LichenoL, Inst. Bot, Pol. Acad. Sci., Lubicz 46, 31-512 Krakow,

Poland)

89-285 OVSTEDAL D.O. - The vegetation of Lindas and Austrheim, western Norway.

Phytocoenologia 1985, 13(3): 323-449, 4 fig., 42 tabl. (Bot, Inst, Р.О. Box 12, N-5014

Bergen).

Descr. des communautés végétales dans les cantons de Lindas et d'Austrheim. Descr.

de nouv. ass. et sous-ass. Bryophytes et lichens associés.

89-286 OTNYUKOVA T.N. - Ecology of some moss species of above-soil cover іп the

forests of Muisk Кеше (Bam Zone). Bot. Zurn. (Moscow & Leningrad) 1985, 70(10)

1373-1380, 3 fig. 2 tabl., en russe (Bot. Inst. V.L. Komarova ANSSSR, Prof. Popof

Str. 2, Leningrad P-22, USSR).

89-287 OTNYUKOVA T.N. - The ecology and phytocoenology of some moss synusia in the

forests of the Muiskaya hollow (Baikal-Amur railway region). Bot. Zurn. (Moscow &

Leningrad) 1985, 70(11): 1465-1476, 6 tabl., en russe, rés. angl. (Ibidem).

Structure et dynamique des synusies à Rhytidium rugosum, Ptilidium ciliare, Pleurozium

schreberi, Aulacomnium turgidum, A. palustre et Tomenthypnum nitens dans 6 types de fo-

réts de la région du train de Baikal-Amur.

89-288 PEÑUELAS J, CANALÍS V. € CASAS C. - Aportació al coneixement de la

brioflora aquatica de Valta muntanya pirinenca. Collect. Bot. (Barcelona) 1985, 16(1):

51-57 (Dept. Ecol., Fac. Biol, Univ. Barcelona, Е-08028 Barcelona).

Bryophytes aquatiques des Pyrénées centrales et orientales, au-dessus de 1600m, avec

hab. Adaptation à l'environnement.

89-289 ROSSELLÓ J.A. - Notes sobre la Brioflora Balear Ш. Collect. Bor. (Barcelona)

1985, 16(1): 63-66, 2 fig. (Dept. Bot., Univ. Autonoma, Bellaterra, Barcelona, Espafia).

Source : MNHN, Paris

272 BIBLIOGRAPHIE BRYOLOGIQUE

Notes chorologiques de 16 bryophytes aux Baléares. Athalamia hyalina, A. spathysit

et Bryum alpinum sont nouv. pour les Baléares.

89-290 SCHOFIELD W.B. - Bryophyte disjunctions in the Northern Hemisphere: Europe

and North America. Bot. J. Linn. Soc. 1988, 98(3): 211-224, 8 fig. (Dept. Bot., Univ.

Brit, Columbia, University Boulevard, Vancouver, B.C., V6T 2B1, Canada).

Disjonctions majeures des bryophytes: amphiatlanuque et Europe W - Amérique NW.

Relations avec les régimes climatiques. La connaissance du passé continental et de la biolo-

gie des bryophytes sont nécessaires pour une meilleure interprétation de ces disjonctions.

89-291 SCOTT G.A.M. - Australasian bryogeography: fact, fallacy and fantasy. Bot. J.

Linn. Soc. 1988, 98(3): 203-210 (Queen's College, Univ. Melbourne, Parkville, Victoria

3052, Australia),

Les mécanismes possibles conduisant aux modéles actuels de distribution des

bryophytes en Australasie ne peuvent étre rigoureusement déduits des modéles eux-mémes.

Les données de distr. sont toutes imparfaites et changeantes. Causes possibles de

disjonction. Imperfections de la théorie de l'endémisme.

89-292 THEURILLAT J.P. et BEGUIN C. - Les groupements végétaux du canton de

Neuchâtel (Jura, Suisse). Saussurea 1985, 16; 67-93 (Cons. et Jard. Bot, C.P. 60,

CH-1292 Chambésy GE).

Présentation systématique des groupements végétaux du canton de Neuchatel, Vali-

dation de certaines unités. Bryophytes associé

89-293 WATTEZ J.R. et VAN HALUWYN Ch. - Contribution à l'étude de la végétation

épiphytique (Lichens et Bryophytes) de la région guérandaise et des abords de l'estuaire

de la Loire. Bull Soc. Sci. Nat. Ouest France n.s., 1985, 7(2): 70-93, 13 cartes, tabl.

(Lab. Bot., Fac. Pharmacie, pl. Dewailly, F-80000 Amiens).

Les lichens et les bryophytes les plus intéressants (une dizaine) de la région

guérandaise et des abords de l'estuaire de la Loire ont été cartographiés et font l'objet de

commentaires sur leur rareté, leur répartitions géogr. et stationnelles. Noter la présence de

Lecanactis subabietina. Descr. des principaux groupements de lichens épiphytes; influence

de la pollution atmosphérique.

89-294 WERNER J. - Vorkommen und Verbreitung der Grimmiales (Musci) im

Grossherzogtum Luxemburg, im Westlichen Saarland und in einigen angränzenden

Gegenden. "Faun.-Florist. Notizen Saarland 1985, 17(3) 355-376, 5 fig., 2 tabl. (32 rue

Michel-Rodange, L-7248 Bereldange).

Liste de 24 Grimmiales avec loc. dans le Grand-Duché-de-Luxembourg et en Sarre oc-

cidentale. Phytogéogr. et écol. des Grimmiales importance des facteurs géol. et pétrogr.

dans cette distr. Liste des Grimmiales menacees

89-295 WERNER J. - Lophozia turbinata (Raddi) Steph. (Hepaticae) dans le sud-est du

Srand-Duché de Luxembourg. Dumortiera 1988, 42; 11-16 (Ibidem).

Descr. et ill. de Lophozia turbinara nouveau pour le Grand-Duché de Luxembourg.

Ecologie et distr. en Europe.

Voir aussi: 89-252, 89-314, 89-317.

Pollution

89-296 KISS Т. - Regressive succession induced by acid rain in cryptogamic communities

inhabiting Juglans bark. Symp. Biol. Hung. 1987, 35: 865-882, 8 tabl., 4 fig. (Dept.

Nat. Hist., Savaria Mus., Szombathely, Kisfaludy 9, 9701 Hungary).

Les correlations significatives entre le pH de l'écorce et le SO; , entre le pH de la pluie

et le SO, „entre le pH de l'écorce et le pH de la pluie reflètent des effets directs et indirects

des polluants sur les plantes épiphytes et leurs substrats. Les thalles des lichens sont plus di-

rectement exposés et les plus touchés. Chez les mousses, sauf Hypnum cupressiforme , seuls

les colonisateurs sont capables de survivre au stress.

Source : MNHN, Paris

BIBLIOGRAPHIE BRYOLOGIQUE 273

89-297 KWAPULINSKI J. and SAROSIEK J. - Radioecotoxicological influence of a power

station on mosses. Symp. Biol. Hung. 1987, 35: 815-826, 7 tabl., 1 fig. (Silesian Univ.

Med., Dept. Toxicology, ul. Jagiellonska 4, 41200 Sosnowiec, Poland).

Etude de la variation du contenu en ra, Ra et Cs dans Polytrichum commune

dans la région de Beskidy. L'influence de la minéralisation est directement et inversement

proportionnelle à la précipitation annuelle sur la station. Relation avec la distance de la

mousse par rapport à l'usine.

89-298 MAKINEN A. - Sphagnum moss-bags im air pollution monitoring іп the city of

Helsinki. Symp. Biol Hung. 1987, 35: 755-776, 12 fip. 2 tabl. (Dept. Bot., Univ.

Helsinki, Unioninkatu 44, SF-00170 Helsinki).

Sur la base de l'analyse atomique de l'absorption de Sphagnum girgensohnii, calcul et

cartographie des valeurs mensuelles d'accumulation de Cd, Cr, Cu, Fe, Pb, Ni, V et Zn. 3

sources: traffic routier, usine thermique, incinération des ordures. La 1* est responsable du

Fe, cendres, Pb, Cr, la 2° du NI et du V, la 3° du Zn et du Cd.

89-299 MAKINEN A. - Use of Hylocomium splendens for regional and local heavy metal

monitoring around a coal-fired power plant in Southern Finland. Symp. Biol. Hung.

1987, 35: 777-793, 5 fig., 3 tabl. (Ibidem).

Par spectroscopie atomique d'absorption, concentrations en Cd, Cr, Cu, Mn, Ni, Pb,

V et Zn dans Hylocomium splendens récolté à proximité d'une usine électrique. Distr. des

métaux dans la plante, relation avec la distance à l'usine.

39-300 RON E., MAZIMPAKA V., VICENTE J. and GRANZOW DE LA CERDA I. -

Urban bryophytes in spanish towns. Symp. Biol. Hung., 1987, 35: 727-753, 10 tabl., 4

fig. (Dept. Bot., Fac. Biol., Univ. Complutense, Madrid 3, Spain).

Présence absence dans différents habitats, résistance aux conditions urbaines des

bryophytes dans 4 villes du centre de l'Espagne.

89-301 SAROSIEK J., WIEWIÓRKA Z. and MRÓZ L. - Bioindication of heavy metals

toxicity of water by the liverwort Ricciocarpus natans (L.) Corda. Symp. Biol. Hung.

1987, 35: 827-833, 3 tabl. (Dept. Ecol. & Nat. Prod, Inst. Bot, Wroclaw Univ., ul.

Kanonina 618, 50-328 Wroclaw, Poland).

Etude des réactions symptomatiques de Ricciocarpus natans aux métaux lourds (Zn,

Cu, Pb) pour l'utiliser comme indicateur biologique de pollution des eaux.

89-302 SAROSIEK J. and SAMECKA-CYMERMAN A. - The bioindication of ethylene

glycol in water by the mosses Fontinalis antipyretica L. and Platyhypnidium rusciforme

(Neck.) Fleisch. Symp. Biol. Hung. 1987, 35: 835-841, 2 tabl. (Ibidem).

Etude en laboratoire. Fonrinalis antipyretica et Platyhypnidium rusciforme different

dans leur vulnérabilité à l'action toxique du glycol éthyléne. Les populations de ces mous-

ses ont une susceptibilité écologique au glycol éthyléne et sont capables de le décomposer.

89-303 SAROSIEK J. and WIEWIORKA Z. - The electric properties of the aquatic moss

Fontinalis antipyretica L. in the bioindication of environmental contamination by ethylene

glycol. Symp. Biol, Hung. 1987, 35: 843-848, 2 fig (Ibidem).

89-304 SAROSIEK J., WOZAKOWSKA NATKANIEC H. and WIEWIORKA Z. - The

effect of heavy metals on the dynamics of Ricciocarpus natans (L.) Corda population.

Symp. Biol. Hung. 1987, 35: 857-863, 3 tabl. (Ibidem).

Relations entre nombre de pieds de Ricciocarpus natans d'une population, la biomasse

et les niveaux des contenus en métaux (V, Ni, Cr, Co) dans un environnement donné, Utili-

sation de Riccioc. natans comme biodincateur de la pollution des eaux par ces métaux.

89-305 SERGIO C. - Epiphytic bryophytes and air quality іп the Tejo Estuary. Symp. Biol.

Hung. 1987, 35: 795-314, 7 fig., 2 tabl. (Inst. Bot., Fac. Ci., 1294 Lisboa, Portugal).

Bryoflore epiphyte (32 taxons), principalement sur Olea europea, de 200 sites de

l'estuaire du Tejo; présence, fertilité en fonction du SO. Noter la raréfaction et l'extinction

de certaines espèces. Toxitolérance depaupérisation chez Tortula laevipila.

89-306 WIEWIÓRKA 7. and SAROSIEK J. - The effects of non-ioning radiation on the

aquatic liverwort Ricciocarpus natans (L.) Corda. Symp. Biol. Hung. 1987, 35:

Source : MNHN, Paris

274 BIBLIOGRAPHIE BRYOLOGIQUE

349-856, 2 tabl. (Dept. Ecol. & Natur. Prod., Inst. Bot., Wroclaw Univ., ul. Kanonina

618, 50-328, Wroclaw, Poland).

Voir aussi: 89-281

Paléobryologie

89-307 BOSE М.Х. and BANERJI J. - The fossil foras of Kachchh. I- Mesozoic

megafossils. Palaeobotanist 1984, 33: 1-189, 71 fig., 55 pl.

Descr. de 44 genres dont 3 sont nouv. Les plantes du mésozoique du Kachchh peu-

vent appartenir au Jurassique moyen et supérieur. Descr. et ill. des taxons. Noter Thallites

sp. et Hepaticites sukhpurensis sp. nov.

Bryophilie

89-308 DAVIDSON A.J. and LONGTON R.E. - Acceptability of mosses as food for a

herbivore, the slug, Arion hortensis. Symp. Biol, Hung. 1987, 35: 707-719, 5 tabl., 1 fig.

(Dept. Bot., Univ. Reading, Whitekmghts, Reading RG6 2AS, UK).

D'aprés le comportement de la limace, les auteurs suggérent la présence de barriéres

chimiques et physiques empêchant la consommation des pieds de Polytrichum commune et

de Mnium hornum.

89-309 DÓBBELER P. - Moosbewohnende Ascomycten VIL. Neufunde einiger Arten der

Gattung Epibryon. Mitt. Bot. Staatssamml, Munchen 1985, 21(1): 757-773, 6 fig. (Inst.

Syst. Bot., Univ. München, Menzingerstr. 67, D-8000 München 19).

Nouv. récoltes de 5 esp. communes 4” Epibryon (Dothideales), ascomycétes bryophiles.

Descr., ill. de certains caractères, Le genre Thuidium est un nouvel hôte pour Epibryon

diaphanum, et Pogonatum capillare pour E. pogonati-urnigeri.

89-310 DOBBELER P. und TRIEBEL D. - Hepaticole Vertreter der Gattungen Muellerella

und Dactylospora (Ascomycetes). Bor. Jahrb. Syst. 1985, 107(1-4): 503-519, 4 fig., 2

tabl. (Ibidem).

Descr., ill. de quelques espéces hépaticoles des genres Muellerella et Dactylospora es-

sentiellement lichénicoles: М. frullaniae sp. nov. et M. rubescens sp. nov. sur Frullania,

Dacıylospora heimelii (Zuka) c.n. ( Paryphydria у sur Jungermanniales. Lichénisation non

demontrée chez cette derniére esp.

89-311 DOBBELER P., POELT J. und VEZDA А. - Lopadium hepaticola spec. nov. ein

moosparasitisches echtes Lopadium von der Südhalbkugel. Herzogia 1985, 71-2); 81-91,

2 fig. (Ibidem).

Diagn., descr., ill. de Lopadium hepaticola, parasite d'hépatiques en Tasmanie. Clés aux

genres séparés de Lopadium s. lat.

89-312 GERSON U. - Mites which feed on mosses. Symp. Biol Hung. 1987, 35: 721-724

(Dept. Entomol., Fac. Agric., Hebrew Univ., Rehovot 76100, Israel).

89-313 ORTEGA A. y BUENDIA A.Ga. оп al estudio de la tribu Aleurieae

Seaver emend. Korf. en la Peninsula iberica. Cryprogamie, Mycol. 1987, 8(2): 125-140,

26 fig. (Depto. Biol. Veg. (Bot), Fac. Ci., Univ. Granada, Granada, España).

Etudes morphol. de 20 esp. d’ Aleurieae Seaver emend. Korf (Pezizales) dont certaines

sont bryicoles. Problémes taxonomiques pour 7 autres.

Voir aussi: 89-325, 89-236.

Source : MNHN, Paris

BIBLIOGRAPHIE BRYOLOGIQUE 275

Ouvrages généraux

89-314 ENGEL J.J. and HATTORI S. - Bryological contributions presented in Celebration

of the distinguished Scholarship of Rudolph M. Schuster. Beth. Nova Hedwigia 1988, 90,

402 p. ill (Dept, Bot, Field Mus. Nat, Mist, Roosvelt Road at Lake Shore Drive,

Chicago, Ш 60605, USA).

6 contributions biogr., témoignage de ses éléves et amis, honorent R.M. Schuster (né

еп 1921) en tant qu'explorateur, enseignant et taxonomiste tout entier voué à la botanique

et plus spécialement aux hépatiques. Les 23 contributions scientifiques reflètent le role

éminent que tient R.M. Schuster en taxonomie et en systématique des hépatiques et aussi

des mousses, Noter une bibliographie complöte avec les dates effectives de publication. А

l'occasion de ce jubilé, les espèces suivantes lui sont dédiées: Fossombronia rudis G.A.M.

Scott et D.C. Pike d'Australie, Frullania (Trachycolea) schusreri Hattori de Queensland,

Plagiochila rudischusteri Robinson du Venezuela, Plagiochila rudolfii Pocs de Tanzanie et

Fissidens schusteri wats. et Wu de Chine.

89-315 MILLER N.G. - Bryophyte ultrastructure. Advances Bryol. 1988, 3: i-vii, 1-281, ill.

(Biol Surv., New York State Mus, Albany, NY 12230, USA) 6 contributions:

cystosquelette, plastes, blépharoplaste, péristome, sporogenese, jonction sporophyte-

gamétophyte chez les bryophytes, Etat des connaissances, perpectives de recherches.

89-316 NEWTON М.Е.) WANSTALL P.J. and JURY S.L. - Bryology: modern research

and the ways forward. Bot. J. Linn. Soc. 1988, 98(3): 183-275, ill, (Dept. Cell & Struct.

Biol, Williamson Building, Univ. Manchester, Manchester M13 9PL, UK).

A l'occasion du bicentenaire de la Linnean Society of London, un meeting commun,

Linnean Society et British Bryological Society a été organisé. 7 contributions furent

présentées, exposant les approches contemporaines ct les perspectives de recherches en

taxonomie, cytologie, physiologie et distribution des bryophytes.

89-317 SCHUSTER К.М. - The Hepaticae of South Greenland. Вей. Nova Hedwigia

71988" 1989, 92: 1-255, 13 tabl., 27 fig. (Cryptog. Lab., Hadley, Massachusetts, USA).

Historique, paramètres biologiques, géologiques et climatiques du Groenland S. Re-

production, écologie, phytogéographie, dissemination des spores en environnement arctique

Probléme taxonomique posé par les hépatiques arctiques. Traitement systématique des

taxons avec loc., notes écol. et morphol.; clé aux esp. de certains genres. Bibliographie (5р.),

index (3p.).

Documentation, Histoire des Sciences

Voir: 89-358.

Nouveau périodique. - ACTA BRYOLICHENOLOGICA ASIATICA

А new journal to report on short, original research findings on Asiatic bryophytes and

lichens is scheduled to be published in January of 1990. The journal will appear twice a

year, and be published in Chinese and English languages. All English articles shall have а

Chinese abstract prepared by the author(s) or editor, and vice versa. Submission of

manuscripts for the first issue shall be no later than August 30, this year. Manuscripts

should be sent to the editor or the publisher at the following addresses: Dr. Benito C. Tan,

c'o Cryptogam Herbarium, New York Botanical Garden, Bronx, NY 10458, USA; or Dr.

Ming-lou Lai, Р.О. Box 190004, Taipei, Taiwan 24199. Subscription rate is about USS

10.00 per year.

Source : MNHN Paris

Cryptogamie, Bryol. Lichénol. 1989, 10 (3): 276-281

BIBLIOGRAPHIE LICHENOLOGIQUE

D. LAMY

Laboratoire de Cryptogamie, 12 rue Buffon, F-75005 Paris

Systématique, Nomenclature

89-318 ABASSI MAAF L., ROUX Cl. - Hypocenomyce stoechadiana nova lekenspecio

(Hypocenomyce stoechadiana espèce nouvelle de lichen). Bull. Soc. Linn. Provence

71984” 1985, 36: 189-194, 2 fig., en espéranto, гёз. franc. (Greco 43, Lab. Bot. & Ecol.

Médit, Fac. Sci. & Techn. St Jérôme, Av. Escadrille Normandie-Niemen, F-13397

Marseille Cedex 13).

Diagn. descr, ill, chimie de Hypocenomyce stoechadiana esp. nouv. de Port-Cros

(France).

89-319 AHTI T. - Correction to Proposal to conserve Roccella against Thamnium

(lichenized fungi). Taxon 1985, 34(4): 709 (Dept. Bot., Univ. Helsinki, Unioninkatu 44,

SF-00170 Helsinki).

Le type de Roccella est Rocella fuciformis (Linnaeus) A.P. de Candolle ( Lichen

Juciformis Linnaeus). (cf. Taxon 1984, 33: 330).

89-320 HAFELLNER J. - Studien über lichenicole Pilze und Flechten ІП. Die Gattung

Roselliniella Vainio emend. Haf. (Ascomycotina, Dothideales). Herzogia 1985, 7(1-2):

145-162, 12 fig., 1 tabl. (Inst. Bot., Karl-Franzens-Univ. Graz, Holteigasse 6, A-8010

Graz).

Reinstallation de Roselliniella Vainio, distinct des genres Adelococcus Theiss. et Syd. et

Muellerella Hepp ex Müll. Arg. - 4 esp. lichönicoles: К. haplospora (Th. Fr. et Айтам.) (=

Endococcus ), R. frustulosae (Vouaux) (= Миейегейа ), В. kalbil sp. nov. et R. lopadit

(Vouaux) (= Миейегейа ). Clés aux esp. de Roselliniella et genres voisins. Descr., ill,

laxonom., distr. de chaque taxon.

89-321 HAFELLNER J. - Studien über lichenicole Pilze und Flechten IV. Die auf

Brigantiaea -Arten beobachteten Ascomyceten. Herzogia 1985, 7(1-2) 163-180, 4 fig.

(Ibidem).

Clé, taxonom., descr, ill, distr. de 7 champignons lichénicoles sur Brigantiaea

(Brigantiaeaceae, Lecanorales) Esp. nouv: Dacrylospora frigida, D. porphyrea, Opegrapha

brigantina, Buelliella pusilla,

89-322 HERTEL H. und RAMBOLD G. - Lecidea sect. Armeniaceae : lecideoide Arten

der Flechtengattungen Lecanora und Tephromela (Lecanorales). Bot. Jahrb. Syst. 1985,

107(1-4) 469-501, 3 tabl, 9 fig. (Bot. Staatssamml., Menzinger Str. 67, D-8000

München 19).

Lecidea sect. Armeniaceae (Th. Fr.) Ras. est divisé en deux entités selon les caractères

des asques, des paraphyses, des conidiophores et des conidies: groupe marginata (membre

des Lecanoraceae s.str.) el groupe armeniaca (membre des Tephroelataceae). Puisque des

caractéres adéquats n'ont pas été trouvés pour les séparer en genres distincts, les especes du

Source : MNHN, Paris

BIBLIOGRAPHIE LICHENOLOGIQUE 277

groupe marginata sont classés sous Lecidea et celles du groupe armeniaca sous Tephromela.

Nouv. comb. Lecanora albicans (Nyl.) ( Lecidea ), L. marginata (Schaer.) ( Lecidea ),

Tephromela aglaea (Sommer) ( Lecidea ), T. armeniaca (DC) ( Rhizocarpon ), T.

testaceoatra (Vainio) ( Lecidea ) et T. septentrionalis sp. nov. du Groenland. Situation ac-

tuelle du genre Lecidea. Evaluation des caractéres utilisés pour séparer les genres des lichens

Jécidéoides.

89-323 KASHIWADANI H. - Genus Hyperphyscia (Lichen) in Japan. Bull. Natl. Sci. Mus.

ser. B ( Bor.) 1985, 11(3): 91-94, 1 fig. (Dept. Bot., Natl. Sci. Mus., Tokyo, Japan).

Hyperphyscia crocata sp. nov. et H. adglutinata nowy. pour le Japon.

89-324 KUROKA WA S. - Studies on Australian and Tasmanian species of Parmelia (2).

Bull. Natl. Sci. Mus. ser. B (Bot.) 1985, 11(3): 77-90, 11 fig. (Tsukuba Bot. Gard.,

Natl. Sci, Mus., Ibaraki, Japan).

Diagn. descr, ill, affinités de 10 Parmelia nouveaux: P. adusta, P. capnoides, P.

cerrussata, P. fumigata, P. interposita, P. lithophila, P. lithophiloides, P. murina, P. nana, P.

pantherina. - Nouvelles loc. en Australie pour P. neoquintaria. P. pseudohypoleia nouv.

pour la Tasmanie.

89-325 MAYRHOFER H. und POELT J. - Die Flechtengattung Microglaena sensu

ahlbruckner in Europa. Herzogia 1985, 7(1-2): 13-79, 25 fig. (Inst. Bot, Karl-

Franzens-Univ. Graz, Holteigasse 6, A-8010 Graz).

La révision du materiel européen des Microglaena sensu Zahlbruckner permet de dis-

tinguer 3 groupes d'espèces reconnus comme genres par les auteurs, Chromatochlamys

Trev. (esp. type: Chr. muscorum (Fr.) c.n. (= Verrucaria ), esp. se développant en parasite

de mousses pleurocarpes, et 2 autres taxons: Chr. larbaleshieri (A.L. Smith) c.n. (=

Microglaena ), Chr. vezdae sp. nov. Protothelenella Räsänen emend. Mayrh. et Poelt

caractérisé, notamment par le phycobionte Elliptochloris (esp. type P. reducta (= Microgl.

sphinctrinoides subsp. reducta , = P. sphinctrinoidella (Nyl.) c.n. (= Verrucaria ) et 5 esp:

P. corrosa (Koerb.) cn. (= Limboria ), P. leucothelia (Nyl.) сл. (= Verrucaria ), P.

polytrichi Dobb. et Mayrh. in Dobb. (sans phycobionte, parasite sur Polytrichum

sexangulare ), P. sphinctrinoides (Nyl) c.n. (= Verrucaria ) et P. xylina sp. nov.). Thelenella

Nyl. (syn. Microglaena Koerb.) (4 esp: T. justil (Servit in Zschacke) сп. (= Microglaena),

Т. modesta (Nyl) Nyl., T. pertusariella (Nyl.) Vainio et T. sampaiana (B. de Lesd.) c.n. (=

Microgalena )) Caractères morphol. et biol., distr. des taxons. I] est difficile d'établir les

affinités de ces genres avec les familles d'Ascomycétes. Protorhelenella appartient à la nouv.

fam. des Protothelenellaceae.

89-326 POELT J. - Rhizocarpon bryonthae spec. nov. aus Grünland, eine parasitische

Flechte auf der epibryen Pertusaria bryontha. Herzogia 1985, 7(1-2): 93-98, fig. (Inst

Bot., Karl-Franzens-Univ. Graz, Holteigasse 6, A-8010 Graz).

Diagn., descr. ill de Rhizocarpon bryonthae sp. nov. du Groenland, parasite de

Pertusaria bryontha bryicole. Cle aux Rhizocarpon croissant sur Pertusaria.

39-327 RUOSS E. - Die Renntierflechte Cladonia stygia in den Alpen. Bot. Helvet. 1985,

9S(2) 239-245, 2 fig, 1 tabl. (Syst. Geobot. Inst. der Universität, Altenbergrain 21,

CH-3013 Bern).

Taxonomie, écol., distr. de Cladonia stygia (Fr.) c.n. (= C. rangiferina f.) nouveau

dans les Préalpes suisses. Comparaison avec C. rangiferina (L.) Web. L'auteur n'accepte pas

la separation des genres Cladonia et Cladina.

89-328 VITIKAINEN O. - Three new species of Peltigera (Lichenized Ascomyctes). Ann.

Bot. Fenn. 1985, 22(4): 291-298, 6 fig. (Bot. Mus., Univ. Helsinki, Unioninkatu 44,

SF-00170 Helsinki ).

Diagn., descr., ill, affinités, distr. de Peltigera kristinssonii d'Islande, P. pacifica du Ca-

nada, et P. retifoveata de Finlande.

Voir aussi: 89-310, 89-311, 89-329, 89-341, 89-351, 89-352.

Source : MNHN, Paris

278 BIBLIOGRAPHIE LICHENOLOGIQUE

Morphologie, Anatomie

89-329 AHTI T. and HYVONEN S. - Cladina stygia, a common overlooked species of

inder lichen. Ann, Bot. Fenn. 1985, 22(3): 223-229, 2 fig. (Dept. Bot., Univ. Helsinki,

Unioninkatu 44, SF-00170 Helsinki).

Taxonomie, descr., chimie, distr. de Cladina stygia caractérisé par la production de la-

mes rouges dans les conidiomata et une forte mélanisation dans les tissus nécrotiques. Prés.

d'atranorine et d'ac, fumarprotocétrarique dans cette esp. habitant les tourbiéres.

89-330 GARTNER С. - Die Gattung Trebouxia Puymaly (Chlorellales, Chlorophyceae).

Arch. Hydrobiol. Suppl. 1985, 71(4); 495-548, 30 fig.

Cytomorphol, taxonom., systémat. des 25 esp. du genre Trebouxia (précédemment

Cystococcus incluant Pseudotrebouxia ) souvent phycobiontes de lichen.

89-331 GODEFROY A. - Rhizocarpons à thalle jaune dans la forét de Fontainebleau. Bull.

Assoc. Franc. Lichénol. 1985, 10(1): 15-20 (Lab. Biol Végét, Route de la Tour

Dénécourt, F-77300 Fontainebleau).

Descr. du thalle, des apothécies, des spores et ill de Rhizocarpon riparium subsp.

lindsayanum, R. viridiairum, R. lecanorinum subsp. lecanorinum et subsp. drepanodes.

89-332 POELT J. - Caloplaca epithallina. Porträt einer parasitischen Flechte. Bor. Jahrb.

Syst. 1985, 107(1-4): 457-468, 3 fig. (Inst. Bot., Holteigasse 6, A-8010 Graz).

Descr., variabilité, écol., distr. de Caloplaca epithallina, parasite strict sur un nombre

d'hôtes taxonomiquement non liés mais bien définis parmi les Lecanorales.

89-333 TSCHERMAK-WOESS E. - Elliptochloris bilobata kein ganz seltener Phycobiont.

Herzogia 1985, 7(1-2): 105-116, 2 fig. (Inst. Bot. & Bot, Gart., Univ. Wien, Rennweg

14, A-1030 Wien).

Descr. d’ Elliptochloris bilobata phycobionte de Catolechia wahlenbergü, Protothelenella

corrosa, Pr. sphinctrinoides, Baeomyces rufus. Relations avec le mycobionte, affinités

systématiques.

Voir aussi: 89-318, 89-320, 89-321, 89-322, 89-323, 89-324, 89-325, 89-326, 89-328, 89-350,

89-351, 89-352, 89-353.

Cytologie, Ultrastructure

89-334 HOLOPAINEN T. and KÂRENLAMPI L. - Characteristic ultrastructural

symptoms caused in lichens by experimental exposure to nitrogen compounds and

fluorides. Ann. Bot. Fenn. 1985, 22(4) 333-342, 3 fig, 3 tabl. (Dept. Environm.

Hygiene, Univ. Kuopio, Р.О. Box 6, SF-70211 Kuopio).

Chez Bryoria capillaris et Hypogymnia physodes, les modifications ultrastructurales

dues aux composés nitrogénes et fluorides sont différents de ceux di à la fumigation de

50,.

335 SCOTT M.G. and LARSON D.W. - The effect of winter field conditions on the dis-

tribution of two species of Umbilicaria. 11. Fine structure and storage body distribution.

New Phytol. 1986, 102(2): 313-326, 3 fig., 4 tabl. (Dept. Bot, Univ. Guelph, Guelph,

Ontario , Canada NG6 2W1),

Effets de la transplantation sur ultrastructure et le mode de distr, des réserves de 2

lichens saxicoles: Umbilicaria vellea et U. deusta. Ce dernier peut accumuler des produits de

photosynthése pendant l'automne.

Physiologie, Chimie

, CULBERSON W.L. and JOHNSON A. - Two new lichen

89-336 CULBERSON C.

inic acid and methylbarbatate, from the genus Haematomma

products, elati

Source - MNHN, Paris

BIBLIOGRAPHIE LICHENOLOGIQUE 279

(Ascomycotina, Haematommataceae). Mycologia 1986, 78(6): 888-891, 1 tabl, 2 fig.

(Dept. Bot., Duke Univ., Durham, North Carolina 27706, USA).

89-337 FIEDLER P, GAMBARO V, GARBARINO J.A. and QUILHOT W, -

Ерірһогейіс acids 1 and 2, two diaryl ethers from the lichen Cornicularia epiphorella.

Phytochemistry 1986, 25(2): 461-465, tabl, (Dept. Quimica, Fac. Ci, Univ. Frederico

Santa Maria, Casilla 110-V, Valparaiso, Chile).

89-338 HUNECKS S., TØNSBERG Т. and BOLHMANN Е. - (--Allo-pertusaric acid and

(-dihydropertusaric acid from the lichen Pertusaria albescens. Phytochemistry 1986,

25(2): 453-459, 2 fig., 2 tabl. (Inst. Pl. Biochem., Res. Dept. BioSci. & Med., Acad. Sci.

GDR, Weinberg, 4010 Halle,Saale, East Germany).

Spectroscopie et chimie pour déterminer la nature de 2 acides carboxyliques lactone

de Pertusaria albescens : acides (-)-allopertusarique et (-)-dihydropertusarique. Mise en

évidence de taraxerone et d'une mixture de longues chaines d'alcools aliphatiques et d'acides

gras chez P. ophthalmiza.

89-339 INGOLFSDOTTIR K., HYLANDS P.J. and SOLBERG Y. - Structure of

vesuvianic acid from Stereocaulon:species Phytochemistry 1986, 25(2): 550-553, 1 tabl.

(Pharmacognosy Res. Lab., Dept. Pharmacy, Chelsea College, Univ. London, Manresa

Road, London $W3 6LX, UK).

89-340 LEUCKERT Ch. and MAYRHOFER H. - Chemische Flechtenanalysen IV.

Herzogia 1985, 7(1-2) 99-104, 3 fig. (Inst. Syst. Bot. & Pflanzengeogr., FU Berlin,

Altensteinstrasse 6, D-1000 Berlin 3).

Présence de depside sulphurelline chez Dimelaena australiensis, d'atranorine, de

chloroatranorine et de sulphurelline chez 6 specimens de Lecanora sulphurella (calycine en

petites quantités dans l'un d'eux).

89-341 MANRIQUE E., BALAGUER L., VALLANDARES F. - Sustancias liquenicas еп

táxones de la provincia de Madrid Il. Hypogymnia gr. intestiniformis. Anales Jard. Bot.

Madrid 1985, 42(1): 81-85, 1 tabl (Dept. Bot, Fac. Farm., Ciudad Univ., E-28040

Madrid).

La présence ou l'absence d'acide furmarprotocétrarique permet de distinguer

Hypogymnia atrofusca d' Н. intestiniformis s.str., qui ont respectivement 2 et 5 races chimi-

ques.

89-342 MAYRHOFER Н. und LEUCKERT Ch. - Beitráge zur Chemie der Flechtengattung

Rinodina (Ach.) Gray Ш. Herzogia 1985, 7(1-2) 117-129, 1 tabl. (Inst. Bot, Karl-

Franzens-Univ. Graz, Holteigasse 6, A-8010 Graz).

Chromatographie en couche mince de 52 spécimens de 26 espéces de Rinodina. Les

acides confluentinique et 2-O-möthylperlatolique sont nouv. chez Rinodina. Distinction de

2 races chimiques chez А. subglaucescens.

89-343 PÉREZ URRIA E. and VICENTE C. - Regulation of urease by urea and its

precursors іп the lichen Evernia prunastri. Physiol. Pl. (Copenhagen) 1985, 65(4):

433-438, 2 tabl., 6 fig. (Dept. РІ. Physiol., The Lichen Team, Fac. Biol., Complutense

Univ., E-28040 Madrid).

89-344 SCOTT М.С. and LARSON D.W. - The effect of winter field conditions on the dis-

tribution of two species of Umbilicaria. 111. CO; exchange in thalli exposed to laboratory

stimulations of winter. New Phytol. 1986, 102(2): 327-343, 10 fig., 3 tabl. (Dept. Bot.,

Univ. Guelph, Guelph, Ontario, Canada N 16 2W1).

Umbilicaria vellea est exclu des habitats à neige parce qu'il est incapable de stocker les

produits de photosynthése sous la neige, contrairement à U. deusta

89-345 SIGAL L.L. and JOHNSTON

J.W. Jr. - Effects of acidic rain and ozone on

nitrogen fixation and photosynthesi in the lichen Lobaria pulmonaria (L.) Hoffm.

Environm. Exper. Bot. 1986, 26(1): 59-64, 2 fig., 1 tabl. (Environm. Sci. Div., Oak

Ridge Natl. Lab., Oak Ridge, TN 37831, USA).

A pH 2,6, la pluie acide entraine une réduction de la fixation de l'azote et de la

Photosynthese; pas de modification à рН 5,6-4,2. L'ozone n'a pas d'action significative,

Source : MNHN, Paris

280 BIBLIOGRAPHIE LICHENOLOGIQUE

mais il y a une tendance à une diminution de la fixation d'azote quant les concentrations

d'O; augmentent. Pas d'interaction Oy -pluje acide.

89-346 TAKALA K. and OLKKONEN H. - Titanium content of ens in Finland. Ann.

Bot. Fenn. 1985, 22(4): 299-305, 5 fig., 4 tabl. (Provincial Gouv. Kuopio, Hallituskatu

12-14, SF-70100 Kuopio).

Le contenu en Ti de lichens épiphytes et terricoles de 31 sites de Finlande dépend

étroitement de l'espéce et varie selon différents facteurs (contenu en sulphure, pourcentage

de dépôt de sulphate, poussières minéroorganiques de l'environnement).

Voir aussi: 89-318, 89-329, 89-335.

Répartition, Ecologie, Sociologie

89-347 ABASSI MAAF L., ROUX СІ. - Champignons lichénisés ou lichénicoles de la Fran-

méridionale: espèces nouvelles ou intéressantes (ІП). Bull. Soc. Linn. Provence

198471985, 36: 195-200 (Greco 43, Bot. & Ecol. Médit., Fac. Sci. & Techn. St Jérôme,

Ау. Escadrille Normandie-Niemen, F-13397 Marseille Cedex 13)

Liste de 32 lichens de la France méridionale avec notes dont Agonimia octospora,

Coccocarpia erythroxyli, Dimerella tavaresiana, Strangospora deplanata et Lecanora

livido-cinerea sont nouv. pour la France.

89-348 ANDREEVA E.1. - Lichenes epiphytici plantarum vascularium principalium regionis

desertorum Kazachstaniae. Bot. Mater. Gerbarija, Inst. Bot. Akad. Nauk Kazakstoj

SSR 1985, 14: 112-123, 2 tabl., en russe.

89-349 APTROOT А. und LUMBSCH Н.Л. - Ergänzungen zur Verbreitung von Cladonia

Sragilissima, Herzogia 1985, 7(1-2): 243-245 (Inst. Syst. Plantk., Heidelberglaan 2,

NL-3508 TC Utrecht),

Cladonia fragilissima nouv. pour les Pays-Bas, et nouv. loc. еп Rép. Féd. d'Allemagne:

89-350 BARRENO E. e RENOBALES G. - Aportaciones a la flora liquenica del País

asco (España): rocas calcáreas, I. Anales Jard. Bot. Madrid 1985, 42(1); 61-80, 1 car-

te, 4 pl. (Dept. Bot., Fac, Farmacia, Univ. Complutense, E-28040 Madrid).

Liste de 21 taxons saxicoles des rochers calcaires du Pays basque espagnol.

Arthopyrenia saxicola, Encephalographa elisae, Gyalecta leucaspis, Lecanora agardhiana

subsp. sapaudica, Opegrapha grumulosa, Petraciis hypoleuca, Polyblastia deminuta, P.

discrepans, P. amota, Staurothele catelepta, S. nantiana, S. rupifraga, Thelidium

absconditum et T. minutulum sont nouv. pour l'Espagne.

89-351 HERTEL Н. - New or little-known New Zealand lecideoid-lichens. Mitt. Вог

Staatssamml. Munchen 1985, 21(1): 301-337, 10 fig.

Carbonea phaeostoma, C. vorticosa, Fuscidea asbolodes, Lecidea endochlora, Poeltiaria

corralensis, Porpidia arthrocarpa, P. macrocarpa, Rhizocarpon disporum, Rimularia insularis,

Sporastatia testudinea sont nouy. pour la Nouvelle-Zélande et les iles subantarctiques.

Nouv. loc. pour 14 autres lichens. Notes taxonom., morphol., répartition géographique;

noter nouv. synonymes.

89-352 KASHIWADANI H. - Lichens of Dokgo Islands, the Oki Islands. In: Natural

History of the Hokuriku and San-in Districts (1). Мет. Май. Sci. Mus. 1985, 18:

95-106, 2 fig.

Liste des lichens des iles Dokgo et Oki, avec loc. Diagn., descr., ill. de P. orientalis sp.

nov.

89-353 LARSON D.W., MATTHES-SFARS U. and NASH T.H. HI - The ecology of

Ramalina menziesii 1. Geographical variation in form. Canad. J. Bot. 1985, 63(11):

2062-2068, 2 tabl., 6 fig. (Dept. Bot, Univ. Guelph, Guelph, Ontario, Canada NIG

2W!)

Source : ММНМ, Paris

BIBLIOGRAPHIE LICHENOLOGIQUE 281

La morphologie de Ramalina menziesii est en relation avec la proximité de la côte et la

latitude, la concentration en NaCl et la variation annuelle de la température, mais pas avec

les plantes hótes.

89-354 MAKRYI Т.У. - The epiphytic lichens of the Baikalsky mountain range. Bot. Zurn.

(Moscow & Leningrad) 1985, 70(11): 1441-1451, 2 tabl., en russe, гёз. angl. (Central.

Sibirsk. Bot. Sad SO, AN SSSR, Novosibirsk).

Liste de 92 esp. épiphytes, relations écologiques avec l'hóte.

89-355 OSORIO ILS. - Contribution to the lichen flora of Uruguay. XXI. Additions to the

Rio de la Plata lichen flora. Mycoraxon 1985, 24: 463-466 (Dept. Bot., Mus. Nac. Hist.

Nat., Casilla de Correo 399, Montevideo, Uruguay).

Liste de 21 lichens du Rio de la Plata avec hab. Noter l'habitat maritime pour Lecidea

icteria, Xanthoparmelia congensis et X. hypopsila.

89-356 POELT J. und MAYRHOFER Die Flechtenflora der Modlinger Klause einst

und jetzt (Niederösterreich). Ber. Deutsch. Bot. Ges. 1985, 98(3-4): 385-392, 1 carte

(Inst. Bot., Karl-Franzens-Univ. Graz, Holteigasse 6, A-8010 Graz).

La flore lichénique des rochers dolomitiques sur les 2 versants de Modlinger Klause a

été étudiée pour la 1° fois en 1856-1857 et pour la 2° fois en 1984. Liste annotée des 82.

esp. saxicoles, terricoles et bryicoles. Noter СайШала scotina et Dermatocarpon

leptophyllum nouveaux pour les Alpes.

89-

57 VIVANT J. - Les lichens des Pyrénées occidentales frangaises et espagnoles. Doc.

Ecol. Pyrén. 1988, 5:3-119, 1 carte (16 rue Guanille, F-64300 Orthez).

304 lichens avec loc. dans les Pyrénées occidentales francaises et espagnoles dont nom-

breuses nouveautes pour les Pyrénées et pour la France.

Voir aussi: 89-283, 89-285, 89-293, 89-318, 89-322, 89-323, 89-324, 89-325, 89-326, 89-327,

89-328, 89-332.

Pollution

Voir: 89-226, 89-334, 89-345, 89-346.

Documentation, Histoire des Sciences

89-358 JOHNSON D.E. - Literature on the history of botany and botanic gardens

1730-1840: A bibliography. Hunneg 1985, 6(1): 3-121 (Rancho Bernardo, 12283 Rouche

House Road, San Diego, CA 92128, USA).

FORMATIONS

Ouvrages récemment regus

ENGEL J.J. and HATTORI 5. - Bryological contributions presented in celebration of the

guished scholarship of Rudolf M. Schuster. BEIHEFTE ZUR NOVA HEDWIGIA

1988, 90, 402p., Ш. (ISBN 3-443-51012-4, Prix DM 280.-, Cramer, Berlin/Stuttgart).

FRAHM J.P. (en collaboration avec LAMY D., SCHUMACKER R., PHILIPPI G.,

RASTETTER V. ) - La bryoflore des Vosges et des zones limitrophes. Duisburg:

Universität-Gesamthochschule, 1989, non paginé, ill. (Chez l'auteur: Univ. Duisburg,

Fachber. 6, Bot., Postfach 101503, D-4100 Duisburg).

Source : MNHN, Paris

282 BIBLIOGRAPHIE LICHENOLOGIQUE

MILLER N.G. - Bryophyté ultrastructure. ADVANCES IN BRYOLOGY 1988, 3, 281р.,

ill. (ISBN 3-443-52001-4, Prix DM 120.-, Cramer, Berlin/Stuttgart).

NEWTON M.E., WANSTALL P.J. and JURY S.L. - Bryology: modern research and the

ways forward. BOTANICAL JOURNAL OF THE LINNEAN SOCIETY 1988, 98(3):

183-275, ill. (S.L. Jury, Dept. Bot., Univ. Reading, Р.О. Box 221, Reading, RG6 2AS,

UK).

SCHUSTER R.M. - The Hepaticae of South Greenland. BEIHEFTE ZUR NOVA

HEDWIGIA "1988" 1989, 92, 255p., ill. (ISBN 3-443-51014-0, Prix DM 170.-, Cramer,

Berlin Stuttgart).

Congrés

3th International colloquium on Lichen Biology - Madrid- Spring 1990, organisé par l'Inter-

national Association of Lichenology. Renseignements; C. Vicente, Lab. Plant

Physiology, Fac. Biology, Complutense Univ., E-28040 Madrid).

Union Internationale des Instituts de Recherches forestières (IUFRO) - 15° Congrés mon-

dial, Montréal, 5-11 août 1990. Renseignements: D.K. Lemkay, IUFRO Montréal 1990

Inc., C.P. 1990, Place d'Armes, Montréal, Canada H2Y 3L9.

Congress of East Asiatic Bryology - Helsinki, 12-19 août 1990 - Organisé par le Dept. Bot.

(Univ. Helsinki) et la Finnish Bryological Society, à l'occasion du 350° anniversaire de

l'Université d'Helsinki. Renseignements: T. Koponen, Dept. Bot, Univ. Helsinki,

Unioninkatu 44, SF-00170 Helsinki.

International Symposium on Endangered Bryophytes іп Furope: Causes and Conservation -

Uppsala, 24-28 septembre 1990 - Organisé par le Swedish Committee for the conser-

vation of endangered bryophytes. Renseignements: N. Cronberg, Dept. Systematic

Botany, Univ. Lund, Ostra Vallg 18-20, S-22361 Lund.

Annonces de décés

Nous avons appris la mort de A. NOGUCHI survenue le 24 septembre 1988. Né en 1907,

A. Noguchi était l'auteur de nombreux travaux bryologiques concernant les mousses

asiatiques, et notamment de /llustrared moss flora of Japan dont les deux premiers fasc.

sont parus et qui sera continue раг Z. lwaisuki. Une notice avec photo est parue dans

le Proc. Bryol. Soc. Japan 1989, 5(1), par S. Inoue, en japonais.

Nous avons appris la mort subite de Stanley Wilson GREENE, survenue le 14 juin 1989.

Né le 19 juillet 1928, S.W. GREENE fut en 1969, parmi les membres fondateurs de

l'International Association of Bryologists dont il était l'actuel Président. En 1980, il créa

“The Bryological Times’ qui, reflet de sa personnalité, plus qu'un simple journal de liai-

son, devint le lien trés vivant (grace à ses éditions européenne, américaine et asiatique)

des bryologues du monde entier. S.W. GREENE s'est consacré aux mousses

antarctiques, mais sa principale préoccupation fut de donner à ses collégues, débutants

et confirmés, des outils documentaires spécifiques; ainsi, le Conspectus of Bryological

Taxonomic Literature dont la premiére partie est parue en 1988 et la deuxiéme vient

d'étre publiée. Nous souhaitons, tous, que les données qu'il a patiemment réunies puis-

sent, malgré sa disparition, être éditées. Nous sommes persuadés que ceci serait son

voeu le plus cher.

Commission paritaire 15-9-1981 - No 58611 - Dépôt légal no 14668 - Imprimerie de Montligeon - Sorti des

1 presses le 25 juillet - Imprimé en France

Éditeur : A.D.A.C. (Association des Amis des Cryptogames)

Président : A. Couté; Secrétaire : D. Lamy

Trésorier : К. Baudouin; Directeur de la publication : Н. Causse

Source : MNHN, Paris

INSTRUCTIONS AUX AUTEURS

Les manuscrits proposés à CRYPTOGAMIE, Bryologie-Lichénologie, doi-

vent être fournis en double exemplaire, dactylographiés à double interligne, sans

rature ni surcharge, et comporter des marges droites et gauches de 25mm, et

hautes et basses de 50mm. Les auteurs sont priés de fournir des textes d'excel-

lente qualité d’encrage. Chaque manuscrit devra comporter:

- le titre de l'article, dans la langue du manuscrit, et sa traduction en anglais;

- e titre courant (haut-de-page) de 30 signes au maximum;

- le nom et les prénoms des auteurs et leurs adresses;

- deux résumés, l'un dans la langue du manuscrit, l'autre en francais ou en an-

glais, d'environ 180 mots ou 15 lignes, faisant ressortir les résultats. essentiels

exposés dans l'article;

- des légendes explicites des figures, planches et tableaux, sur feuilles séparées;

1 une liste bibliographique par ordre alphabétique des auteurs et chronologique

par auteurs sans tenir compte des auteurs secondaires. Les titres des périodiques

devront étre abrégés suivant le B-P-H (Botanico-Periodicum-Huntianum,

Pitsburg: Hunt Botanical Library, 1968), les ouvrages cités selon Ғ.А. Stafleu &

R.S. Cowan, 1976- ... Taxonomic literature. Ed. 2 Utrecht/Antwerpen: Bohn,

Scheltema & Holkema ( Regnum vegetabile 94, 98, 105, 110...).

MONTAGNE C., 1838 - Centurie des plantes cellulaires exotiques nouvelles, Ann. Sci.

Nat., Bot., 2, 9: 38-57.

NEES VON ESENBECK C.G. 1836 - Hepaticae. In: Lindley J., A natural system of

Botany... Ed. 2. London. Pp. 412-414.

WATSON E.V., 1971 - The structure and life history of bryophytes. Ed. 3. London:

Hutschinson University Library. 211 p., 26 fig.

TEXTE. - La présentation du texte devra faire apparaitre clairement ses sub-

divisions et leur hiérarchie ainsi que le début des paragraphes. Les noms des au-

teurs qui suivent les binómes latins devront étre abrégés selon G. Sayre et al.,

1964 (The Bryologist 67 (2): 113-135). Les renvois 4 la liste bibliographique se

feront par le nom de l'auteur et l'année de publication (ex. (Dubois 1980) ou

Dubois (1980) et non par les renvois numériques. La place des illustrations devra

être indiquée dans la marge. Les notes infrapaginales sont à éviter.

ILLUSTRATIONS. - Toutes les illustrations, y compris les tableaux, doivent

étre des originaux de qualité suffisante pour la reproduction directe en offset. El-

les devront comporter les échelles et symboles nécessaires à leur compréhension.

En particulier, les tableaux devront être dactylographiés par une machine

électrique ou composés en lettres de transfert. Les positifs des documents pho-

tographiques devront être montés par planches. Toutes les illustrations doivent

être numérotées dans l'ordre d'appel dans le texte. Les auteurs devront tenir

compte du format de la revue (11 x 18cm) et de la réduction que subissent

éventuellement les originaux en choisissant l'épaisseur des traits et la taille des

lettres et des chiffres.

Les tirages à part et les planches photographiques sont à la charge des au-

teurs.

Source : MNHN, Paris