127

EPIGAEOUS LICHEN SYNUSIAE

IN THE YUKON TERRITORY *

P.L. NIMIS **

SUMMARY. — Numerical methods of classification and ordination have heen applied to

relevés of epigacous lichen vegetation from the Yukon Territory (Canada). Seven synusiae

have been described, six of them new. Their syntaxonomical status, ће 01у and the

relations with similar types known from Europe have been discussed.

INTRODUCTION

Although lichens are a very important element in the vegetation of the Yukon

Territory, there is apparently no paper dealing specifically with lichen synusiae.

The only published work in which lichens are listed in phytosociological tables

is the paper of HOEFS, COWAN and KRAJINA (1975) on the vegetation of

Sheep Mountain, in the Kluane National Park, where only associations of phane-

rogams are described.

In the catalogue of lichens reported for the Yukon Territory, BIRD (1971)

gives a list of 145 species. A more detailed list for a narrower area is contained

in the paper of DOUGLAS & VITT (1976) in which 197 species are reported

from the region of Kluane. It is clear from these figures that much has still to

be done on the floristic level, with particular regard to crustose lichens, Crus-

tose lichen species being poorly known in the area, I decided to deal mainly

with epigaeous macrolichen synusiae. Eight synusiae are described herein and

their relationships with corresponding european types discussed.

* The field work was completed in summer 1978, in conjunction with a project directed

by Prof. L. Orloci (University of Western Ontario, London, Ontario, Canada). The field

party included : Prof. L. Orloci, Ms. C. Emo, Mr. P, Fewster (Department of Botany, Univer-

sity of Western Ontario, Canada) and Prof. D. Lausi, Mr РР. Merluzzi and the author (Isti-

tuto Botanico, Università di Trieste, Italy). In Canada, the author was recipient of a post-

doctoral stipend from NSERC funds (L. Orloci).

** Istituto Botanico dell'Università, Trieste (Italy).

Cryptogamie, Bryol. Lichénol., 1981, 2, 2 2127-151.

Source : MNHN. Paris

128 P.L. NIMIS

GENERAL DESCRIPTION OF STUDY AREA

The Yukon Territory is situated in the northwestern part of Canada, between

60° and 69° latitude and 108°-104° longitude. The vertical projection of the

Yukon boundary encloses about 482,681 km?. The physiography ranges from

a coastal plain in the extreme North to the massive St. Elias Mountains in the

southwest corner. The elevation, in general, decreases from South to North.

During the field work relevés have been taken along the principal roads of the

Territory, i.e. the Alaska, Klondyke and Dempster Highways. The relevés are

thus scattered along two transects. The first runs in a SE-NW direction between

60° and 62° latitude for a total of 912 km, the second follows a N-S direction

between 135°-139° longitude, for a total of 830 km. The Alaska and Klondyke

5099139013751 135^ Чүзз” ILS 129012771255

Fig. 1. — The principal routes of the Yukon Territory.

Source : MNHN, Paris

EPIGAEOUS LICHEN SYNUSIAE 129

Highways are mainly located in the montane (boreal) biogeoclimatic zone.

Along the Dempster Highway subalpine and alpine vegetation are prevalent.

The climate is continental, with mean annual temperature below freezing.

Precipitation is low, from 300 to 500 mm/year in the studied portion of the area

(BURNE 1974). The driest section is located at the southwestern corner, on the

leeside of the St. Elias Mountains. The northwestern region of the Territory is

the only one which remained unglaciated during the Wisconsin glaciation.

Permafrost is common throughout the whole area, scattered in depressions and

lowlands in the southern portion and becoming more prevalent northwards.

Permafrost thickness varies from a few meters in the south to about 100 m at

the northern boundary.

VEGETATION

Since the phanerogamic vegetation will be described in a future paper (LAUSI

& NIMIS), only essential points which are relevant to the lichen synusiae are

mentioned. Picea glauca and Picea mariana are the principal elements in climax

communities throughout the montane (boreal) biogeoclimatic zone, Picea

glauca stands occur on permafrost free, well drained sites, They are structurally

and floristically similar to the european Vaccinio-piceetalia associations. Picea

mariana muskegs, widespread on permafrost, are characterized by a complex

microgeomorphological pattern of hummocks (Bülten) and hollows (Schlánke).

The former are constituted mainly by mosses (Sphagnum and Aulacomnium

species) or by sedges. On their top lichens are dominant.

Secondary stands in the boreal zone are dominated either by Pinus contorta

or by Populus tremuloides. Whereas the former is prevalent in the southeastern

corner of the Territory, east of Whitehorse, the latter species is most common

in the southwestern and central parts. Pinus contorta stands are generally rich in

lichens, whereas Populus tremuloides stands are poor in epigaeous lichens,

because of the inhibiting influence of Populus litter on lichen growth. Locally,

the influence of dry winds of Féhn type strongly affects vegetation, either

directly, or indirectly by the continuous deposition of loess. In the Kluane

area, Picea glauca stands are often confined to the principal strings of the hydro-

graphic net and most of the ground covered by a stepp-like vegetation belonging

to the Artemisio-agropyrion yukonenis Hoefs & Krajina in the montane zone.

Deposits of acid sand and granitic rock outcrops are frequent throughout the

area : they are colonized respectively by a Carex filifolia-association and by a

Saxifraga tricuspidata-association.

Along the Dempster Highway Picea glauca stands are rare and confined to

permafrost free soils along rivers, whereas extensive areas are covered by Picea

mariana muskegs. Subalpine vegetation consists mainly of Salix pulchra and

Betula glandulosa thickets on acid soils, and of Dryas integrifolia meadows on

alkaline soils. Alpine tundra is prevalent in the region around North Fork Pass

and after the Polar Circle.

Source : MNHN. Paris

130 P.L. NIMIS

METHODS

During the field work, relevés of lichen vegetation were made following the

phytosociological method of BRAUN-BLANQUET (1964). The cover scale is

the Braun-Blanquet scale as modificd by PIGNATTI (1953).

Relevés were taken in seven landscape units in which lichens were dominant :

— Understory, open Pinus contorta stands.

— Understory, mature Pinus contorta and Picea glauca stands,

— On hummocks, Picea mariana muskegs.

— Arid grassy slopes with mineral acid soil.

— Arid grassy slopes with mineral or organic alkaline soils.

— Subalpine vegetation on calcareous parent material.

— Ground, alpine tundra.

In order to achieve a classification of relevés, these were ordered in a matrix

which was submitted to multivariate methods. For the classification of relevés

Average Linkage Cluster Analysis (ALCA) was adopted. This method was chosen

because it results free from the group size (FEOLI 1977). The dendrogram obtai-

ned by ALCA doesn’t reflect the order of the clusters in the Vegetational Space

(GOODAL 1963). For this reason also an ordination method has been applied.

Classification being the main goal, a method of Non Centered Principal Compo-

nent Analysis was chosen, The method is based on the extraction of eigenvalues

and eigenvectors from the similarity matrix between relevés obtained by Simila-

rity Ratio Function (WESTHOFF & VAN DER MAAREL 1978). This function

gives a similarity measure ranging between 0 and 1, and its reliability in the

solution of phytosociological problems has still been recognized (VAN DER

MAAREL 1979). The cluster and ordination results have been compared and

used jointly for interpretations.

The syntaxonomical range of lichen synusiae is still a highly controversial

field in lichen ecology. In many papers tables are presented, in which only lichen

species are listed, without any reference to eventual phanerogams or bryophytes.

Groups of lichen species occurring together in nature are thus considered as

abstract entities, which can be treated separately from all the other members

of the community in which they are growing. At least for epigaeous lichen

synusiae, this often results in a loss of substantial information.

On the other hand, if lichen species are not separated from phanerogams in

the construction of a syntaxonomical system, the possibilities of comparisons

between types with similar lichen vegetation and different phanerogamic vege-

tation are strongly reduced. Lichen ecology being the principal topic of this

paper, phanerogams have been listed in the tables, and lichen species have

been treated separately.

RESULTS

The results of the numerical treatment of the data are showed in Fig. 2 and 3.

Source - MNHN. Paris

45.

sol

73+

87.

EPIGAEOUS LICHEN SYNUSIAE 131

| ШЕТ

A LRA AREN ARRAY IS SEST SST RISZALI

Fig. 2. — Dendrogram of the relevés (Average Linkage Cluster Analysis).

Source : MNHN. Paris

132 P.L. NIMIS

Cluster analysis

Seven principal cluster have been detected. They are characterized by the

following species and correspond to the following landscape units :

1) Cladonia ecmocyna, Cladonia multiformis. Mature Pinus contorta and Picea

glauca stands.

2) Cladonia mitis, Cladonia stellaris, Stereocaulon tomentosum, Cladonia gra-

cilis var. dilatata. Open Pinus contorta stands.

3) Cladonia amaurocraea, Thamnolia subuliformis. Top of hummocks, in Picea

mariana muskegs.

4) Masonhalea richardsonii, Alectoria ochroleuca. Alpine tundra, on ridges.

5) Cetraria tilesii. Subalpine vegetation on calcareous parent material.

6) Parmelia wyomingica. Stepps on mineral acid soil.

7) Caloplaca tominii, Psora decipiens. Stepps on alkaline soils.

Homogeneity within clusters is rather high, with the exception of cluster 3.

The sequence of the clusters follows the results obtained by the ordination

method.

Ordination

Fig. 3 presents the ordination of relevés according to the Il and Ш compo-

nent of РСА. The first six clusters are disposed along a circular line originating

from the center. The seventh cluster is strongly separated from the others. An

ecological interpretation of these results seems particularly difficult. The se-

quence of clusters 1, 2, 3 and 4 probably reveals a gradient from moderately

coniophilous to clearly coniophobous communities. This gradient is associated

with permafrost thickness and soil stability. А moderate snow cover (more than

40cm) is an effective insulator against frost penetration into the soil, and

prevents the formation of permafrost in winter and early spring (DAHL in litt.).

The latter season is the one in which permafrost formation or thickening is more

probable, particularly in open area in which the snow tends to melt earlier

during warm days and the percolating meltwater freezes during cold nights.

Drainage is impeded by the permafrost layer, and a Picea mariana-muskeg or

а tundralike vegetation may develop. Clusters 1 and 2 correspond to commu-

nities growing in more or less closed coniferous stands, where insulation is

reduced in spring and the snow cover lasts longer. Clusters 3 and 4 correspond

to communities occurring respectively in open Picea mariana-muskegs and in

a treeless tundra, both more exposed to the action of winds, and where the

snow melts earlier in spring. Permafrost is always absent under the communi-

ties corresponding to cluster 1 and 2, is frequent under the community corres-

ponding to cluster 3, and is always present under the community corresponding

to cluster 4. The active layer is generally thicker in the community represented

by cluster 3 than in the one corresponding to cluster 4. Good parallels to this

situation have been described by DAHL (1956) for the mountains of South

Norway.

From cluster 1 to cluster 6 the only factor varying according to the disposi-

Source : MNHN. Paris

EPIGAEOUS LICHEN SYNUSIAE 133

1 *

R 7

D =”

Е :

А 5

1 UNS

. ses

uu I

-5 + 1 i 3 4 EM 1

SEE d +

Жез "

E z aa

-5

-sl

Fig. 3. — Ordination of the relevés (Non Centered Principal Component Analysis based on

Similarity Ratio).

tion of the clusters is the cover of the phanerogamic layer. Ombrophytic synu-

siae are disposed on the left (cluster 1, 2), heliophytic ones on the right (cluster

4, 5, 6). Cluster 1 corresponds to a lichen synusia occurring in closed climax

forests. Clusters 2, 3 and 4 correspond to more or less stable communities,

whereas clusters 5, 6, 7 correspond to pioneer communities on primitive soils.

Source : MNHN. Paris

134 P.L. NIMIS

Acid or neutral soils are characteristic for relevés included in the clusters from

1 to 6, cluster 7 includes all the relevés of vegetation growing on strongly alka-

line soils.

Numerical syntaxonomy

According to the syntaxonomical system proposed by KLEMENT (1955),

lichen synusiae should be included into three broad classes : Epipetretea (on

rock), Epigaetea (on soil) and Epiphytetea, the latter divided into three orders :

Epixiletalia (on bark), Epibryetalia (on mosses) and Epiphylletalia (on leaves).

"This system has been adopted, above all in Europe, by a great number of phyto-

sociologists interested in lichen vegetation, although it is far from being satis-

factory. The higher syntaxa are defined on the basis of deductive procedure

founded on rough ecological criteria, so that the resulting classification, being

more ecological than sociological, is not consistent with the inductive floristic

method followed for the delimitation of the unions. The class Epipetretea

has been splitted into 3 classes by MATTICK (1951) and more recently into

6 classes by WIRTH (1972), and there is no doubt that a better articulation

is needed also for the class Epigaetea. Numerical syntaxonomy could contribute

to the clarification of the problem through the quantification of the various

syntaxonomical levels by measures of similarity. As KORTEKAAS, VAN DER

MAAREL & BEETFINK (1976) and WESTHOFF & VAN DER MAAREL

(1978) pointed out, units which are considered as belonging to the same hierar-

chical level do not vary very much in similarity level. On the basis of Jaccard’s

coefficient (or Similarity Ratio for presence-absence values), the association

level corresponds to a similarity level ranging from 40 to 60, the alliance level

to 30-40 and the order level to 10-30. LAUSI & FEOLI (1979) found that the

similarity level for the delimitation of classes ranges between 5 and 15.

In the dendrogram of Fig. 2 it is possible to observe the following principal

fusions :

— Clusters 1, 2, 3, 4, 5, 6, 7, are formed between the similarity levels of 42 and

50, figures corresponding fairly well to the similarity level characteristic for

the syntaxonomical level of association (union).

— Clusters 2 and 3, and 4 and 5 fuse together at a similarity level of 32, a value

falling within the threshold characteristic for the delimitation of the syntaxo-

nomical level of alliance (foederatio). By cutting the dendrogram at this

similarity level we obtain the delimitation of 5 foederationes : A (cluster 1)

on neutral or slightly acid soils in mature forests, В (clusters 2 and 3) on acid

soils in the montane and subalpine zone, C (clusters 4 and 5) on organic

soils in the alpine region, D (cluster 6) on mineral acid soils, and E (cluster 7)

on alkaline mineral soils.

— Clusters 1, 2, 3 and 4, 5, 6, fuse between similarity values of 25 and 27

forming two clusters, characterized respectively by species considered charac-

teristic of the foederationes Cladonion arbusculae and Cetrarion nivalis in

the system of Klement. The similarity values fall within the treshold charac-

teristic for the delimitation of the syntaxonomical level of the order. The first

Source : ММНМ Paris

EPIGAEOUS LICHEN SYNUSIAE 135

group includes relevés of the montane (boreal) biogeoclimatic zone, the

second includes relevés of the alpine and subalpine biogeoclimatic zones.

— The two latter clusters fuse together at a similarity level of 19. Cluster 7 is

finally jointed to the dendrogram at a similarity level of 13, falling within the

threshold characteristic for the delimitation of classes. Cluster 7 is characte-

rized by species considered characteristic of the foederatio Toninion coeru-

leonigricantis in the system of Klement.

The numerical method suggests the inclusion of our relevés into 7 unions,

5 foederationes, 3 orders and 2 classes. According to the system of Klement

they should be included into 7 unions, 3 foederationes, 1 order and 1 class. I

think that these results should be considered as a further corroboration of the

quantitative values of similarity delimitating the various syntaxonomical levels

presented in the recent literature. Considering the literature about epigaeous

lichen synusiae, it is evident how two units like the foederationes Cetrarion

nivalis and Toninion coeruleonigricantis have such a small number of species

in common, that they hardly could be considered as belonging to the same

order. These floristic differences correspond to differences in fundamental

ecological factors; the inclusion of unions occurring on strongly acid soils

in the same class in which unions of strongly alkaline soils are included cannot

be accepted if this fundamental difference is bound to strong differences at the

floristic level. In my opinion, a general revision of the system of Klement is

urgently needed. Such a task goes beyond the limits of the present paper. As

1 am working at a revision of the class Epigaetea on the basis of all the available

relevés with numerical methods, I prefer to avoid the creation of new syntaxa

in the present paper, so that the phytosociological tables will be ordered follo-

wing provisorely the system of Klement in order to facilitate comparisons with

other types described in the literature.

In the following pages a brief description of the seven unions is given.

PELTIGERO-CLADONIETUM ECMOCYNAE nova

The synusia forms a mosaic with mosses on decaying wood and humus-rich

soil in mature Pinus contorta and Picea glauca stands.

Cladonia ecmocyna is a constant species. Frequent and characteristic are

Cladonia multiformis and Cladonia cenotea. The union clearly belongs to the

Cladonion arbusculae. Cladonia arbuscula is common in the early successional

stages and in open stands of Pinus contorta, where it can be considered as a

transgressive from the Cladonietum mitis. Noteworth is the abundance of large

foliose species, better adapted to the shaded habitat of the synusia. Peltigera

aphtosa is constant, sometimes together with P. leucophlebia. Р. canina occurs

mainly in the sun flecks. Nephroma arcticum is rare in closed stands or on North

facing slopes. A subunion, Cladonietosum arbusculae, can be described, characte-

tized by Cladonia arbuscula, Peltigera canina and Cladonia gracilis var. dilatata,

occurring in more open and dry sites. The most common bryophytes are Hylo-

comium splendens, Abietinella abietina and Pleurozium schreberi.

Source : MNHN. Paris

136 P.L. NIMIS

Fallen decaying trees, often buried and covered by a thin humic layer, are the

main habitat for the synusia. Lichens are often absent at the base ot the trees

in open stands of particularly dry regions. In this case, the ground is barren

of vegetation for a radius from 0,3 to 1 m around the tree trunk, surrounded by

a belt covered by an almost pure stand of Drepanocladus uncinatus. The synusia

can be considered ombrophytic, and moderately hygrophytic. Most of the

species reproduce by means of spores, germination being favoured by humidity

and shade.

Pinus contorta stands are secondary successional stages leading to Picea

glauca stands. The latter are climax communities on slopes or permafrost-free

lowlands throughout the montane zone. The Peltigero-cladonietum ecmocynae

can be considered a stable community; only in particularly closed mature

stands mosses have the tendence of overgrowing lichens, forming a continuous

carpet on the ground. If the Picea glauca stand grows on lowlands with under-

laying permafrost, it tends to be replaced by a Picea mariana-muskeg. In this

case the Peltigero-cladonietum ecmocynae is substituted by the Thammolio-

cladonietum amaurocraeae.

The synusia is common along the Alaska Highway in the montane (boreal)

biogeoclimatic zone. Two relevés were made along the Klondyke Highway.

The synusia was also observed near Ogilvie, on the Dempster Highway. Its

distribution probably follows the area of Picea glauca in the Yukon. A хего-

phytic type of Picea glauca stand, described by HOEFS, COWAN and KRAJINA

(1975) as Hypno-Abietinello-Piceetum glaucae, occurs in the region around

Kluane lake. In this association lichens are rare and the synusia is scarcely

developed.

Species diversity in the Peltigero-cladonietum ecmocynae is greater in the

region in which secundary successions are dominated by Pinus contorta.

CLADONIETUM MITIS Krieger 1937

The synusia is exclusively present as an understory in open Pinus contorta

stands. Our relevés readily fit the Cladonietum mitis, described by KRIEGER

in 1937 for Europe. The four characteristic species of this union (Cladonia

mitis, Cladonia gracilis, Cladonia verticillata and Stereocaulon tomentosum)

can be considered characteristic also in the study area. To these Cladonia cariosa

is added, as a local differential species, being confined to this synusia in the

Yukon. Fructicose Cladoniae of the Cladina subgenus are dominant. Large

foliose species such as Peltigera canina and Nephroma arcticum are also well

represented. Lichen cover goes from 70 to 100%. The most frequent mosses

include Tortula ruralis, Polytrichum juniperinum and Ceratodon purpureus.

The soils are sandy in texture and extremely poor în organic material. A

tendence towards podsolization is often evident, but true podzols are rare

because of the scarcity of precipitation. The upper horizon is constituted by

Pinus litter and lichen residua, underlain by a thin organic layer and a weakly

leached sandy mineral horizon. The synusia can be considered as moderately

Source : MNHN. Paris

EPIGAEOUS LICHEN SYNUSIAE

PELTIGERO-CLADONTETUN ECHOCHIAE nove

Type relevés ¥ 2

137

Relevé Y СО 10

Locality (im Alaska Highway) 1192 1236 1050 1313 1618 1458 1 1156

Exposure А. sare вом È

Slope ( degrees) iy siae da oo в в 3

surface mg peu E син. рк 4

Phaneroganes

Picea glauca (Moench) Voss айн Si sa тиэ Ga ae

Limena borealis L. melasma (forbes) Melt. 902 4.2 fd f Ld 4.2 12 12 632 o? ¥

Vaccinium vitis - idasa L. т Wi 14 34494 139 053 Ay,

Geocsulon lividua ( Richards.) Term. + жй Жие; AISA Ao

Mertensia paniculata ( Ait.) Сов. + ES e in

Pyrola secunda L- sana a ni

Pyrola asarifolia Mich. ¢ : п

Carex concinna R.Br. E pru и

Equisetum scirpoides Miche. sig Ж . n

Cormis canadensis L. of Map i

Lycopodium amotimum L. эа ea D

Monasas uniflora (L.) Gray * а; n

Festuca altaica Tein. жэза +2 n

Calypso bulbosa (L.) Rchb. B P

Arnica cordifolia Book, ani

Charact. species of the Union

Cladonia senoeyna (ach. Jy1. Vire sue end ve еа Y

Cladonia maltiforais Narr. Maroni Ma Ha эз эп av

Cladonia phyliophora Hoffm. эз эл эз за +2 ш

Cladonia cenotea (Ach.) Schaer, sea эз +2 sa m

Differential species subunion

Cladonietostm arbusculae

Cladonia arbuscula ( Wallr.) Rabenh.

ap. beeringiana Anti e: e» oe: 62 +2 m

peltigera caning ( L.) WINN 14 1 эз 54 и

Cladonia gracilis (L.)Willd. dilatata Hoffm. + da зоа и

Charact. species of Cladonion arbuscula

Cladonia cormta (L.) Hoffm. л аа 1 ld 2: LA E +2 19

Cladonia chlorophaes (PIBrk+) Spreng. SES а +2 +1 ш

Cladonia fimbriata (L.) Fr. E : а + m

Cladonia acuminata (Ach.) Norri. +2 E u

Cladonia verticillata ( Hoffa.) Schaer. E т

Charset, species of Epigeetal

Peltigers aphtosa (L.) Willd. Da ыз de жй эз ааа а у

Peltigera leucophlebia 091.) Gyeln. D “2 edo: ona in

Vapheoua arcticum (L.) Tors +2 та и

Loberia Linita (Ach.) Rabenh. +2 D п

Fhysconia miacigens (Ach.) Poelt эз" 1

Bryophytes

Nyloconium splendens (Нейм) 8.5.0. за LE 12 62 L3 ei Li ызаа +

Abietinella abietina (ledv.) Fleisch +2 + эз эп эп э m

Pleurorium schreberi (Brid) Mitt. ЖЕ эт Ша эл эз ui

пауы rugosum (Hedu.) Kind. di D E te

Tomenthypmum nitens (Hedv.) Loeske E за са u

Dicranum undulatım Brid. +2 +з em u

Pohlia mutans (Wedv.) Lindo. CET эз u

Lophotia hatcheri (Evans) Steph. E #2 6а n

Ceratodon purpureus (ledu.) Brid- за э жон

Tortula ruralis (Hedv.) CMS. г 1

Cover ( Phanerogames ) Т so 90 so $0 so во во б 40 эз

Covet dos + 20 25 20 15 25 43 ау 40 20 15

ll 4 35 20 10 10 15 15 33 lo 10

Tab. 1.

Source : MNHN. Paris

CIADONIETIN MITIS Krieger 1937

Relevé S* 28 29 30 31 32 33 34 35

Locality (Ка Alaska Highway) 1047 1048 1140 1353 1445 1169 1102 1074

Exposure: оне ве

Slope (degrees) 9 - Jus» are

Surface mq 70 70 100 70 80 80 100 100

Phaneroganes.

Pinus contorta Loud. dA 31 31 за 41 ал каал +

Arctostaphylos uva-ursi L. L2 13 +2 эз 23 13 13 L3 V

iiu мелт. шу: aper cem (onen) male. ua Hi (2 62 Da I еа и V

Festuca altaica Trin. 12 +12 62 #2 +2 12 +2 42 V

Pulsatilla patens (L.) Mill. + * o: * + w

Gentiana propinqua Richards, 11 1л m

Epilobium angustifolium L. 11 = à + n

Vaccinium vitis-idaea L. asp.minus(Lodd.) Hule. #2 аа nf 62 IV

Zypadenus elegans Pursh + + + + w

Rosa acicularis Lindl. e + SEE

Shepherdia canadensis (L.) Nutt. +2 +2 +2 ш

Geocaulon lividum (Richards.) Fern. + Кн Е

Ribes triste Pall. + ж rn tt,

Salix glauca L. +2 +2 п

Calamagrostis canadensis ( Mich.) + т

parta

Charact. species of the Union

Cladonia mitis Sander. 13 13 42 63 L3 13 +3 +31

Cladonia stellaris (Opiz) Pour. et Vezda Aura uii а т

Stereocaulon tomentosum Fr. FA 53-63 13 «9 n3 тз br N

Cladonia gracilis (L.)Willd, v.dilatata Hoffm. #2 +2 +.2 1.2 ані еа,

Cladonia verticillata (Hoffa.) Scheer. + saree it)

Cladonia cariosa (Ach.) Spreng. La 44 0:2 эз m

Cladonia aymphycarpa ( Ach.) Fr. +2 +2 эт nm

Charact. species of Cladonion arbusculae

та з аз 52 Las t3 23 ж

(les) Wigs. +2 23 13 13 +2 #2 42 V

Cladonia deformis Hoffe. Dada ede mes on

Cladonia rangiferina (L.) Web. 12 ia «d € 23 25 w

Cladonia cornuta (L.) Hoffm. +2 +2 +2 +2 sa +2 N

Cladonia coccifera (L.) Willd +2 +2 421062 Sd м

Cladonia phyllophora Hoffe. +2 ГҮ. +2 ап m

Cladonia clorophaea (Floerke) Zopf +2 +2 з ш

Charact. species of Epigeetalia

Peltigera aphtosa (L.) Willd. sant +2 +2 +22 Y

Peltigera canina (L.) Willd. Xa L3 12 «2 RE у

Cornicularia aculeata (Schreb.)Ach. эр жа +2 +2 +2 +2 y

Cetraria laevigata Ras: #2 +2 +2 +2 w

Nephroma arcticum (L.) Torss. +з »3 ла ост Т

Cetraria cucullata (Beil.) Ach. +2 sar

Cetraria nivalis (L.) Ach. +2 +2 п

Stereocaulon paschale (L.) Hoffa. +2 1

Bryophytes

Tortula ruralis (Gaertn.) G.M.S. i3 їй +242 €i L3 63 42 Y

Ceratodon purpureus (Hedw.) Brid. +2 +2 +2 +2 +2 жас

Polytrichum juniperinum Нейм. +2 +2 nose

Hylocomium splendens (Hedv.) 3.5.6. Fr са п

Polytrichum piliferum Нейм. +2 +2 u

Drepanocladus uncinatus (Hedv.) Warnst. 2 т

cover ( Phanerogames ) 1 во ss so so ss 90 90 100

Cover (Lichens ) 1 во 60 во 50 70 90 во 75

Cover ( Bryophytes ) 4 4e үү 5, 5 dn. 10, 5

Tab. 2.

Source : MNHN. Paris

EPIGAEOUS LICHEN SYNUSIAE 139

photophytic, acidophytic, xerophytic and anemophobic. Reproduction occurs

mainly by fragmentation of the thallus.

Pinus contorta stands tend to develop into Picea glauca communities. The

natural development of the lichen vegetation would therefore involve a change

from the Cladonietum mitis to the Peltigero-cladonietum ecmocynae, after

closure of the tree canopy. Fires are extremely common, resulting in large scale

destruction of mature stands. Succession after a strong fire can be summarized

аз follows :

1) Cladonia cariosa stage : From three years after a fire the first lichen to invade

is Cladonia cariosa and a few other species of the Cladonietum mitis which

survived fire, The ground cover is 10-20%. Small Pinus plants start growing.

Phanerogams are represented mainly by annuals such as Gentiana propinqua

and Epilobium angustifolium.

2) Festuca altaica stage : After four to five years following fire the ground cover

is dominated by Festuca altaica. Ericaceous shrubs start appearing. The

cover is 60-80%.

3) Arctostaphylos stage : Arctostaphylos uva-ursi, Vaccinium vitis-idaea and

Empetrum nigrum form a thick continuous carpet six to ten years after

fire. Cladoniae of the Cladina subgenus are growing on the open ground.

The cover is 70-90%.

4) Cladonietum mitis stage. Lichens are dominant in the ground layer. The

cover is 70-100%.

5) Picea glauca stage : the stand closes, mosses become dominant, and the

Peltigero-cladonietum ecmocynae replaces the Cladonietum mitis. Pinus

contorta is slowly replaced by Picea glauca.

On the top of dry hills, the evolution of the Cladonietum mitis is very slow,

since the stand remains open. It is possible that in these sites the synusia behaves

as a «Dauergesellschaft».

The synusia has been observed along the Alaska Highway, from Watson

Lake to Whitehorse, in the montane (boreal) biogeoclimatic zone. It seems

to be lacking outside the distributional limits of Pinus contorta. Stands of

Cladonietum mitis were also observed in British Columbia, always under Pinus

contorta, in the Prince George Area.

THAMNOLIO-CLADONIETUM AMAUROCRAEAE nova

The synusia occurs on the top of Eriophorum vaginatum-tussocks, and on

Sphagnum and Aulacomnium palustre hummocks in Picea mariana muskegs.

The characteristic species for the synusia include Cladonia amaurocraea and

Thamnolia subuliformis. Iemadophyla ericetorum, although occurring frequently

together with the other species of the synusia, is a saprophyte occurring on

Sphagnum, and should be properly considered as belonging to another synusia,

dominated by mosses and more hygrophytic. The most common phanerogams

аге Ledum decumbens, Vaccinium vitis-idaea, Festuca altaica and Rhododen-

dron lapponicum. Among mosses, Aulacomnium and Sphagnum species are

Source : MNHN, Paris

140 P.L. NIMIS

THAMNOLIO = CLADONIETUM AMAUROCREAE nova

Type relevé : N° 15

Relevé N° n 12 o d 15 16 17 18 19

locality ( Km Alaska HW) 1247 1927 1915 1871 1895 1862 1873 1957 1912

Exposure rue вм a

Slope (Degrees) a ae Е pt

Surface ma К shy сз aus ино

Phaneroganes

13 24 31 21 ш

Vaccinium vitii +2 +2 € 42 42 €2 е IV

Empetrum nigrum L. ermaphroditun(Lange) Böcher 1.2 1.2 +2 #2 +2 эз 1

Eriophorum vaginatuh L. NERI +2 +2 эЛ 10

cularis labradorica Wirsing WEEE Bi: m

Ledum palustre 1. 2.2 эз 2,2 4.2 ш

Carex dioica L. gynocraten(Wormsk.) Hult. + +2 +2 п

Chamaedaphne calyculata (L.) Moench +2 E n

Rhododendron lapponicum (L.) Wahlenb. +2 +2

Selaginella selaginoides (/.) Link + + - п

Andromeda polyfolia L. + т

Char. species of the Union

Cladonia amaurocraea (Flörke) Schaer. 1212 22 42 121211292022 Y

‘Thamnolia subuliformis (Ehrh.)W. Culb. +2 +2 +2 42 4242 w

(lemadophila ericetorum (L.) Zahlbr.) 13 14 юз 16 14 ш

Char. species of Cladonion arbusculae

Cladonia arbuscula (Hallr.)Rabenh,

subsp. beeringiana Ahti +2 +2012 эл +2 +2 АА

Cladonia deformis Hoffa. м. + w

Cladonia rangiferina (L.)Web. +2 +2 12 +2 m

Cladonia mitis Sandst. +2 E тї

Cladonia stellaris(Opiz) Pour et Verda 1212 W

Cladonia cornuta (L.) Hoffa. + + n

Cladonia chlorophaes ( Flörke) Spreng. +2 т

Cladonia gracilis (L.)Willd.dilatata Hoffa. + т

Cladonia uncialis (L.) Wigs +1 1

Cladonia coccifera (L.) Will jb eee

Cladonia symphycarpa (Ach.) Fr. SETS

Charact. species of Cetrarion nivalis

Cetraria laevigata Re +2 xp eus aN N

Cetraria nivalis (L.) Ach. 24 дз +2 +2062 + w

Cetraria cucullata ( Beil.) Ach. *4 12 u

Cornicularia divergens Ach. Fan ©

Charact.

Pektigera aphtosa (L.) Willd. 1.2 2.2 L2 L2 L2 13 13 L3 23 +

Peltigera canina (L.) Willd. FIAS 621€ CIV

Bryophytes

Hylocomium splendens ( Hedw.) B.S. i3 12 «223 13 42 +8 э ы y

Rhytidium rugosum ( Hedw.) Kindb. +2 p ape IN

Aulacomiun palustre (Hedv.) Schvaegr. iati eat er m

Tomenthypmum nitens (Hedw. )Loeske +2 52 +62 п

Drepanocladus uncinatus (Hedw.) Warnet. +2 +2 п

Sphagnum эр. us u

Bryum bimum Schreb. + 1

Cover ( Phaneroganes ) 1 io 30 15 40 10 10 40 45 35

Cover ( Lichens ) X 40 45 70 50 65 50 60 40 60

Cover (Bryophytes ) 1 до 4 5 45 25 40 5 5 5

Tab. 3.

Source : MNHN. Paris

EPIGAEOUS LICHEN SYNUSIAE 141

common, but generally suffering from drought. Hylocomium splendens, Rhyti-

dium rugosum and Polytrichum juniperinum are frequent. Species of both

Cetrarion and Cladonion are well represented, so that a syntaxonomical classi-

fication for this synusia is diffieult. Considering the results of the mathematical

treatment of the data, I assigned it provisorily to the Cladonion arbusculae.

In comparison with the other syhusiae described in this paper, the Thamnolio-

cladonietum amaurocraeae shows the highest degree of variation in floristic

composition. The reason lies in the extreme complexity of the microgeomorpho-

logical pattern in Picea mariana muskegs. A more detailed study would be re-

quired to describe several sub-unions.

From the top to the base of the hummock there is a sharp moisture gradient,

to which the lichen synusia responds by occupying the driest upper portion.

The soil consists mainly of undecomposed organic material in the upper horizon,

followed by acid moss or sedge peat. Permafrost is always present at a depth

varying from 10 to 60 cm (in summer). An ice core is sometimes present in the

internal part of the hummock, preserved by the insulating action of the moss

sheet. The synusia can be considered as heliophytic and moderately xerophytic,

acidophytic and anemophobic.

Since the Picea mariana muskegs are stable communities on lowlands and in

depressions, the lichen synusia can also be considered stable. As often happens

with lichen synusiae, a weak cyclical successional pattern is caused by factors

internal to the association in which the synusia grows, so that, while the situa-

tion can change at the local level, the pattern is statistically unaltered on a larger

portion of ground. In particular, the top of the hummock is often colonized by

Picea mariana, which tends to overgrow the lichens. Subsequently the rising

of permafrost kills the tree and the previous situation is reestablished. Fire is

frequent in the area, and the destruction of the insulating moss sheet is the

principal cause of permafrost melting, followed by subsidence and the forma-

tion of ponds or fens. In the infrequent case of a Picea mariana muskeg deve-

loped on a slope, meltwater drains downwards, permitting a Pinus contorta or

Populus tremuloides stand to initiate a succession towards a Picea glauca stand.

The synusia being confined to Picea mariana muskegs, it occurs in areas

with permafrost under 66° latitude. Along the Alaska Highway, it is rare and

scattered in depressions from Watson Lake to Kluane, becoming extremely

common west of Kluane, and especially in the section between Koidern and the

Alaskan border, always whithin the montane or subalpine biogeoclimatic zones.

ALECTORIO-MASONHALEETUM RICHARDSONII nova

The synusia occurs on wind-blown rocky outcrops, ridges and peaks in the

alpine zone.

Masonhalea richardsonii, an amphi-beringian endemic, is constant. Other

frequent and characteristic species are Alectoria ochroleuca, Dactylina ramulosa,

Dactylina arctica and Alectoria nigricans, The character species of the alliance

(Cetrarion nivalis) are very well represented : Cetraria cucullata, Cetraria nivalis

Source : MNHN, Paris

142 P.L. NIMIS

ALECTORIO - NASOMMALEETUM RICHARDSONII nova

Type relevé : N° 20

Relevé N° 20 n 22 m 24 15 2% 2

Elevation m 1050 1000 1200 1150 1100 1100 1100 1200

Exposition зз mw - - - - X

Slope (Degrees) LT een Lee

Locality (Km Dempster НУ) ва si 102 90 88 88 вә 101

Surface mq roe c Sele Seah vr rt

Phaneroganes

Hierochloe alpina (Sv.) Roem. et Schult. эл ка ка ка жа ei o2 +2 Ж

Vaccinium vitis-idaea L. minus (Lodd.)Hult. +2 B +2 +2 m

lustre 1. жа э n

arctica Less. +2 т

parviflora (Ehrh.) Desv. +2 т

tes hyperboreus Rydb. salii

Charact. species of the Union

Masonhalea richardsonii (Hook. )Kiirnef. 12 13 43 $2 «2 14 тї 13 Y

Alectoria ochroleuca (Hoffm.) Mass. 12 +2 +2 +2 +2 +2 N

Dactylina ramılosa (Hook.) Tuck. 23 22 12 12 +2 1

Dactylina arctica (Hook.) Nyl. 12 +2 +2 Tuc

Mectoria nigricans (Ach.) Nyl. +2 + пг

Charact species of Cetrarion nivalis

Cetraria nivalis (L.) Ach. 3.2 +2 12 4.2 L2 пака.

Cetraria cucullata (Bell.) Ach. 13 202 12 az А E

Cetraria laevigata Ras 2 12 +12 +2 +2 m

Cornicularia aculeata (Schreb.) Ach. +2 АЖ:

Cetraria islandica (L.) Ach. ZE u

Cladonia amaurocraen(Flörke) Schaer. +2 +2 +3 п

Thaanolia subuliformis (Ehrh.) Culb. E 1

Charace. species of Cladonion arbusculae

Cladonia rangiferina (L.) Web. atrata +2 Wa y

Cladonia chlorophaea (Flörke) Spreng. жа +2 +2 +2 на. Ж

uncialia (L.) Hoffa: +3 +3 + +2 fn

formis Hoffe xi Thats

atellaris (Opis) Pour.et Verda +2 E

gracilis (L.)Willd. dilatata Hoffa. +2 +з п

coccifera (L.) Willd. n

acuminata (Ach.) Norri. + п

talia

Stereocaulon alpinum Laur. 12 12 12 +233 ош

Peltigera aphtosa (L.) Will 13 +2 13 As

Nephroma arcticum (L.) Torri +213 п

Bryophytes

Hylocomium splendens (Hedw.)B.S.G. da engr arti at ыбыз ж

Anastrophyllum minutua(Schreb.)Schuat, +2 +2 +2 +з m

Pleuroriun schreberi (Brid.) Mitt. +2 +2 +2 +2 +2 иг

Abytidiun rugosum (Hedv.) Kind). +з +3 +2 +2 m

Abietinella abietina (Hedv.) Fl. eror п

Polytrichum commune Hedw- r т

Cover (Phanerogames) X Ба оч +

Cover (Lichens) 1 85 so so 60 70 50 do 80

Cover (Bryophytes) T 1545 05 \ u$ ES LS ([®

Tab. 4.

Source : MNHN. Paris

EPIGAEOUS LICHEN SYNUSIAE 143

and Cetraria laevigata are frequent, whereas Comicularia aculeata is rarer and

confined to very exposed sites. The ground cover varies from 70 to 100%, li-

chens being always dominant. Hierochloe alpina, Ledum decumbens and Vacci-

nium vitisidaea are almost the only phanerogams occurring with the lichens.

The bryophytes are represented chiefly by Hylocomium splendens ssp. alaska-

num, Rhytidium rugosum and Anastrophyllum minutum var. grandis.

The synusia is widespread in the siliceous part of the Ogilvie Mountains

(Precambrian igneous and metamorphic rocks, mainly quartzite and schists)

between mile 84 and 200 on the Dempster Highway. The soils are poorly deve-

loped, mostly gravelly or rocky. The synusia seems to prefer south facing slopes

or ridges. Most of the species are sterile, and reproduce by fragmentation of the

thallus. The synusia is heliophytic, acidophytic, anemophytic and xerofrigid.

It can be considered as a «Dauergesellschaft». A complicated dynamic pattern

is sometimes caused by cryoturbation and solifluction.

Outside the Ogilvie Mountains the synusia is probably common in the alpine

zone on siliceous rocks throughout the Yukon Territory. A specimen of Mason-

halea richardsonii was found near Koidern in southern Yukon, probably carried

by wind from the St. Elias Range.

CLADONIETUM ALPESTRIS Frey 1927

In the same region in which the Alectorio-masonhaleetum richardsonii was

observed, a different lichen synusia occupies large areas on solifluction lobes

in relatively wind-protected sites. Of this vegetation type we have only two

relevés, whose floristic composition is very similar to the one of the Cladonietum

alpestris Frey. Both relevés come from North Fork Pass, on solifluction lobes,

25°SW; the cover was 100%,

1 2

Cladonia stellaris 33 43

Cetraria laevigata 1.2 13

Cladonia gracilis ssp. elongata E +

Cladonia amaurocraea 12 12

Cladonia coccifera +2 +2

Cladonia rangiferina +2 12

Cladonia uncialis г

Dactylina arctica Tu

Cladonia acuminata >

Peltigera aphtosa +3

The synusia is extremely common along the Dempster Highway, from Thomb-

stone campsite to the polar circle, and often behaves as understory of a sub-

alpine Salix pulchra thicket. It replaces the Alectorio-masonhaleetum in more

protected sites, where the soil is deeper, the wind less strong, and the snow lasts

longer.

CETRARIETUM TILESII nova

The synusia is bound to phanerogamic associations of the order Dryadetalia

Source : MNHN, Paris

144 P.L. NIMIS

CETRARIETIN TILESII nova

Type relevé + N° 41

ГЕ 36 37 38 39 40 41 42 43

Elevation 1000 1100 1151 1098 900 950 900 980

Exposition. тагии 4 5 À

Slope (Degrees) 20 25 23 30 10 LS 24 13

Surface mq * "E À 4 74 аа

Phancroganes

Dryas inteprifolia M. Vahl 232512014 12 9:3 ES аа Да ви

Carex scirpoidea Michx. +2 +2 +2 ш

Salix reticulata L. 1.2 Dues m

Saxifraga oppositifolia L. smalliana E.et I. 2 aa) f

Carex glacialis Mack. 1212012022 (HL

Carex petricosa Dew. +2 аа DI

Campanula aurita Greene Ala п

Phlox sibirica 1. +2 +2 п

Senecio hyperborealis Creenm. а. п

Parrya nudicaulis (L.) Regel + EI

Charact. species of the Union

Cetraria tilesii Ach. E LAL HE ea a N

Stereocaulon glareosum (Sav.) Magn. Ld 62 623 42 m

Charact. species of Cetrarion nival:

Cornicularia aculeata (schreb.) Ach. жазаа tales Chiaia жир Sy

Cetraria cucullata (Bell.) Ach. +2 #2 43 13 13 +3 00V

Cetraria nivalis (L.) Ach. жа эз эз +2 12 23 mW

‘Thamnolia subuliformis (Ehrh.) Culb. +2 +2 +2 m

Cetraria laevigata Rass. + + п

Charact. species of Epigeetali

Cladonia pocillum(Ach.) 0.-J. Rich. 2 +2 +2 sia замсыз m

Peltigera canina (L.) Willd. #2 13 +2 +2 та m

Dermatocarpon hepaticum (Ach.) Th.Fr. 42 + a; ш

Dactylina arctica (Hook.) Nyl. r Nue osi

Cladonia stellaris (Opiz) Pouz.et Vezda Sonate в

Cladonia chlorophaea (Fl8rke) Spreng. +2 т

Solorina spongiosa (Sm.) Anzi + т

Physconia muscigena (Ach.) Poelt + т

Parnelia chlorochroa Tuck. tro 1

Bryophytes

Tortula ruralis (Wedw.) Gaertn. 12 bd 52 #2 m

Hypnum vaucheri Lesq. 3:8 6521582 OS m

Ceratodon purpureus (Hedw.) Brid. +2 +2 п

Splachnum luteum Hedw. [2] 1

Ptilidium ciliare (L.) Hampe е а ear

Rhacomitriun lanuginosum (Hedw.) Brid. az 1212 ut

Tetraplodon mnioides (Hedw.) B.S.G. ааа п

Hylocomium splendens (Hedw.)B.S.G. eat 1

Drepanocladus uncinatus (Hedw.) Warnst. «it

Cover (Phanerogames) % 20 25 30 20 35 25 30 40

Cover (Lichens) 1 65 60 60 30 15 20 45 60

Cover ( Bryophytes) X RE М BS

Tab. 5.

Source : MNHN. Paris

EPIGAEOUS LICHEN SYNUSIAE 145

integrifoliae Hoefs et Krajina. Because of the high number of associations in which

the synusia was recorded, in the table only the dominant phanerogams are listed.

Constant and characteristic is Cetraria tilesii. Stereocaulon glareosum can be

considered a local differential species for the Kluane area. The synusia clearly

belongs to the Cetrarion nivalis, its characteristic species being very well repre-

sented. The bryophytic component is not constant in the different regions in

which the synusia was recorded : Tortula ruralis and Hypnum vaucheri are

common in the Kluane area, whereas in the relevés of the Ogilvie Mountains the

bryophytes are represented chiefly by Tetraplodon mnioides, Hylocomium

splendens and Rhacomitrium canescens,

Cetraria tilesii is an alpine species preferring neutral to alkaline soils. The

synusia was found on rendzina or protorendzina soils at an elevation of 1300-

1900 m in Kluane and 800-1500 т in Ogilvie. It can be considered heliophytic,

neutro- to basiphytic, and rather anemophobic.

The Cetrarietum tilesii is clearly a pioneer community on poorly developed

alkaline soils. It occurs on loess deposits in the Kluane area and in soil pockets

on calcareous scree slopes in the Ogilvie Mountains. With increasing acidification

it tends to be replaced by moss synusiae behaving as understory of different

shrub communities, sometimes by the Cladonietum alpestris. The continuous

erosion, together with the influence of eolian loess material (Kluane) or the

downward movement of the scree slopes (Ogilvie) create situations in which

the synusia often behaves as a «Dauergesellschaft».

The synusia has been observed in Kluane National Park and in the region

near Ogilvie, on the Dempster Highway. The distribution is expected to follow

the emergence of calcareous rocks in the alpine or subalpine biogeoclimatic

zones throughout the Yukon Territory.

FULGENSIO-CALOPLACETUM TOMINII nova

The synusia is bound to stepp-like phanerogamic associations belonging

to the order Artemisio-Agropyretalia yukonensis Hoefs et Krajina.

Constant and characteristic is Caloplaca tominii, an euroasiatic species new

for the lichen flora of North America (NIMIS in press). Frequent are Psora

rubiformis, Psora decipiens, Buellia elegans and Fulgensia bracteata. Since this

is the only union belonging to a totally different alliance than Cetrarion nivalis

or Cladonion arbusculae, and since other records of similar synusiae described

for North America are unavailable, it is impossible to give a satisfactory syntaxo-

nomical designation. By analogy with similar synusiae described for Europe,

however, it would seem possible to include the Fulgensio-caloplacetum tominii

in the Toninion coeruleonigricantis, four characteristic species of this foederatio

being present in my table : Psora decipiens, Fulgensia bracteata, Dermatocarpon

hepaticum and Toninia coeruleonigricans. Nothing can be said about other

possible characteristic species of the union until further data on similar unions

in North America will be available.

Source - MNHN, Paris

146 P.L. NIMIS

FULGENSIO - CALOPLACETUM TOMINII nova

Type relevé 1*51

Relevé N° 50 sr

Elevation m 810 900

Exposition SE SE

Slope (degrees) 35 20

Surface mq 4 5

Phanerogami

Artemisia frigida Willd. LİR?

Agropyron yukonensis Scribn.et Merr. +.2 +.2

Artemisia alaskana Rydb. +2

Penstemon gormanii Greene +

Linum perenne L. lewisii (Pursh) Hult. Ж: A,

Aster alpinus L. vierhapperi Onno

Artemisia rupestris L. woddii Nelson

Charact.species of the Union

Caloplaca tominii Savicz 1.2 2.2

Charact.species of Toninion coeruleonigricantis

Psora decipiens (Hedw.) Hoffm.

Dermatocarpon hepaticum (Ach.) Th.

Fulgensia bracteata (Hoffm.) Ris

Toninia coeruleonigricans (Lightf.

Psora rubiformis (Wahl.) Hook.

Charact.species of Epigeetalia

Squamarina lentigera (Web.) Poelt

Buellia elegans Poelt

Cladon:

Physconia muscigena (Ach.) Poelt

Cetraria tilesii Ach.

Leptogium saturninum (Ach.) Nyl.

Collema coccophorum Tuck.

Bryophytes

Tortula ruralis (Hedw.) Gaertn.

Ceratodon purpureus (Hedw;) Brid.

Hypnum vaucheri Lesq

Cover ( Phanerogames) 1

Cover ( Lichens) %

Cover ( Bryophytes) 1

1.2

Fro +.2

) Th.Fr.

pocillum (Ach.) 0.-J.Rich.

Tab. 6.

1.2

52 53

850 890

в N

15 15

+.2 +.2

+2 +2

22032

1.2 12

+.2 1.2

+.2 +2

4,2

+.2 +2

2 «2

15, 2

20 15

54 55 56

950 850 850

в:

10 20 14

cio

ha BÉ 4.2

+.2

+.2 r

1.2

Re E

129.2 4.2

+2 т

dita

АА чө

+ +

1.2

+.2

+2 +2 4.2

#2 г

+2 +2

10 20 10

35 20 10

Sos 59

ти

1ш

ту

ти

Source : MNHN. Paris

EPIGAEOUS LICHEN SYNUSIAE 147

The region of Kluane is characterized by a particularly dry local climate,

since it lies in the rainshadow of the St. Elias Mountains, and is exposed to

dty winds, factors favouring the expansion of southern steppic species. For a

detailed discussion of the climatic situation see HOEFS, COWAN and KRAJINA

(1975). The soils are mostly mineral (loess or sand). The pH range varies from

7.2 to 8.4. South facing slopes are preferred. The synusia is (termo)- xerophytic

and basiphytic.

Generally the ecological conditions are so extreme and erosion so strong that

the synusia behaves mostly as a «Dauergesellschaft». Otherwise a continuous

carpet of Arctostaphylos uva-ursi opens the successional process, progressing

towards the Abietinello-piceetum glaucae, a xeric variant of the Picea glauca

forests. I observed that the woods are confined to the principal strings of the

hydrographic net. It seems therefore probable that most of the area occupied

by the synusia will not develop a woody vegetation until the general and local

climatic conditions will change.

The synusia was recorded only in the Kluane National Park. It is best deve-

loped in the Slims River Valley, near the Kaskawulsh Glacier. I suppose that

most of the species reach their northern distributional limit in the area.

PARMELIETUM WYOMINGICAE nova

Constant species are Parmelia wyomingica and Cornicularia aculeata. The

union has been assigned to the Cetrarion nivalis.

The synusia substitutes the Fulgensio-caloplacetum tominii on acid mineral

soils in the montane and subalpine biogeoclimatic zones. It occurs on sandy

grassy slopes dominated by Carex filifolia, and on mineral soils in pockets among

granitic rocks in an association dominated by Saxifraga tricuspidata. The synusia

is xerophytic, heliophytic and acidophytic. It is widespread, but not frequent in

Southern and Central Yukon, being absent in areas with calcareous rocks ог

surficial loess deposits.

DISCUSSION

The homogeneity in lichen flora whithin the holoarctic region is a well-known

fact. The number of species common to Europe and North America increases

from South to North. According to AHTI (1964), 85% of the species belonging

to the flora of northern Ontario are common with the flora of Fennoscandia.

From the catalogue of lichens reported from the Yukon Territory, compiled

by BIRD (1971), 1 have calculated that 88% of the species are common with

the flora of northern Europe. It seems logical to expect a corresponding simila-

rity among vegetation types, since we can suppose that the same species tend

to occupy the same ecological niche in both continents. On the other hand

it also seems logical to expect substantial differences in the lichen vegetation,

due to the presence of local endemics, to the floristic history, and above all to

the climatic difference between the northern portions of the two continents

Source : MNHN. Paris

148 P.L. NIMIS

PARMELIETUM WYOMINGICAE nova

Type releva : N° 48

Relevé N* 44 45 6 47 48 49

Local.(A:Km Alaska HWY;K:Km Klondyke HWY) А1569 K175 K162 K150 K163 А1506

Exposition ci) Argo som bi E

Slope ( degrees) о овлаш

Surface mq Pig A TA

Phanerogames

Carex filifolia Nutt. moe $251 HES

Saxifraga tricuspidata Rottb. + а м

Penstemon gormani Greene ze + act bay

Calamagrostis purpurascens R.Br. а 4.2 +2 N

Artemisia frigida Willd. РРА In

Juniperus horizontalis Moench. eg ta E

Erigeron caespitosus Nutt. + + п

Potentilla hookeriana Lehm. 3 xs п

Melandrium apetalum Fenzl. + 1

Charact. species of the Union

Parmelia wyomingica (Gjeln.) Hale eisen elek ee

Charact. species of Cetrarion nivalis

cornicularia aculeata (Schreb.) Th.Fr. 1.2 TEC

Cetraria nivalis {L.) Ach. + м

Cetraria cucullata (Bell.) Ach. ту

Cetraria laevigata Rass. + r u

Charact. species of Epigeetalia

Cladonia pyxidata (L.) Fr. +2 42 435358: «5 ы

Cladonia cariosa (Ach.) Spreng. PONES + IM

Peltigera rufescens (Weiss.) Humb. i "ox ти

Cladonia symphycarpa Flörke "iaces N it

Bryophytes

Tortula ruralis (Hedw.) G.M.S. Pos Le N

Ceratodon purpureus (Ach.) Spreng. + ent

Polytrichum juniperinum Hedw. * + п

Cover ( Phanerogames ) X 109 GS Mars ESS sb.

Cover ( Lichens ) X 45 50 50 55 45 30

Cover ( Bryophytes ) % E Wa

Tab. 7.

(continental arctic North America versus oceanic arctic Europe). Of the previous-

ly described lichen synusiae, one or two are floristically so similar to european

types that they can be considered as belonging to the same union described

for Europe. Another four unions, although differing somewhat from their

Source : MNHN. Paris

EPIGAEOUS LICHEN SYNUSIAE 149

european counterparts, appear nothing more than geographical vicariants of the

european units in comparable ecological conditions.

In european Picea stands one of the most common lichen synusiae is the

Cladonietum cenoteae Frey. A geographical vicariant, in the Yukon Terri-

tory, is the Peltigero-cladonietum ecmocynae. Although Cladonia cenotea is

present їп my relevés and Cladonia ecmocyna sometimes occurs in the Clado-

nietum cenoteae, the total floristic composition of the two types shows many

important differences. Some are due to the presence of north american species

not present in Europe, as Cladonia multiformis, some to the presence of species

with a different ecology in the two continents. An example is the case of Clado-

nia ecmocyna, which in Europe is a species occurring mainly in snow beds of

the alpine zone, whereas in the Yukon it seems to obtain its ecological optimum

in closed coniferous stands of the montane zone. The Cladonietum ecmocynae

described by FREY (1927) for Europe is thus quite different floristically and

ecologically from our Peltigero-cladonietum ecmocynae. The Cladonietum mitis,

described by KRIEGER in 1937 for north Germany, is widespread from nor-

thern Scandinavia to the Alps (FREY 1952, LIMPRICHT 1931, KALB 1970).

It generally occurs on sandy mineral soils, mostly podzolized, often under a

Pinus spp. open stand. LIMPRICHT describes a Pinetum montanae cladinetosum,

KALB (1970) a Cladonietum mitis bchaving as understory of a Pinus engadinen-

sis open wood. In the Yukon Territory, the synusia is practically the same as

in Europe from the floristic point of view, and has a comparable ecology. Some

differences in the bryophytic component, for example the constance of Tortula

ruralis in the relevés of the Yukon, are due in first line to the continentality

of the climate.

The Cladonietum alpestris is, in Europe, an alpine and subalpine type, known

from northern Scandinavia (KLEMENT 1959) to the Alps (HARKEMA 1976).

In the Yukon Territory it seems equally restricted to alpine and subalpine areas,

in relatively wind-protected sites. Its distribution, with considerable geographic

variation, extends probably throughout arctic and boreal North America. I ob-

served a lichen synusia whose floristic composition was very similar to the one of

the Cladonietum alpestris in the Montmorency Forest (Quebec). The floristic

composition of the Alectorio-masonhaleetum richardsonii is directly comparable

to the one of the Thamnolietum vermicularis Gams, known from the alpine

region of Europe, from Scandinavia to the Alps. Of the 16 species present in

the table published by KALB (1970), 9 are present in our table. The ecology

and the relationships toward the Cladonietum alpestris are strictly similar to

the ones described for Europe. Differences in the floristic composition are due

in the first place to the presence of local endemics.

A lichen synusia that floristically and ecologically is very similar to the

Fulgensio-caloplacetum tominii has been recorded by KLEIVEN (1959) from

South Norway. Six species are common with the relevés from the Yukon Terri-

tory : Caloplaca tominii, Fulgensia bracteata, Toninia coeruleonigricans, Buellia

epigaea sl., Lecanora lentigera and Physconia muscigena. Climatic and ecolo-

gical conditions in South Norway (Gudsbrandalen) are very similar to the ones

Source : MNHN, Paris

150 P.L. NIMIS

in the Kluane region. Gudsbrandalen is the only known station of Caloplaca

tominii in Europe; outside this area similar vegetation types, belonging to the

Toninion coeruleonigricantis, are known for arid regions on alkaline soils of

northern and central Europe (ALBERTSON 1946, DU RIETZ 1925, GAMS

1938). Considering the fact that the Toninion coeruleonigricantis in Europe is

a relict of a stepp-like vegetation on calcium-rich soils, it is probable that in

North America it could play an important role in sagebrush grasslands on alka-

line soils.

For most of the described types there was no difficulty in including them in

the higher syntaxonomical units of lichen vegetation described from Europe.

The distinction between Cetrarion nivalis and Cladonion arbusculae seems to

be meaningful also for the studied area. An exception is perhaps the Thamnolio-

cladonietum amaurocraeae, whose synsystematic position remains unclear. More

relevés from North America are needed for a better articulation of the system,

which is surely possible.

Literature on north american epigaeous lichen synusiae is still not sufficient

for general conclusions to be drawn, but the available information tends to

suggest a high similarity in lichen vegetation between the northernmost portions

of the two continents.

AKNOWLEDGEMENTS, —I am indebted to Prof. T. Ahti (Helsinki), J.J. Barkman (Wij-

ster), F J. Daniels (Utrecht), R.S. Forbes (Belfast), D. Lausi, E. Feoli, S. Pignatti (Trieste),

L. Orloci (London, Ontario), J. Poelt (Graz), O. Willmanns (Freiburg) and W.C. Steere

(New York), for comments on the manuscript or for identification of critical specimens,

LITERATURE CITED

AHTI T., 1964 — Macrolichens and their zonal distribution in boreal and arctic Ontario

(Canada). Ann. Bot. Fenn. 1 : 1-35,

AHLNER S., 1929 — Contributions to the lichen flora of Norway, I. Svensk Bot. Tidskr.

43 :157-162.

ALBERTSON L., 1946 — Österplanan, ett alvaromrade pà Kinnekulle. Acta Phytogeogr.

Suec. 20.

BIRD СШ. 1971 — A catalogue of the lichens reported from the Yukon Territory. Univ.

of Alberta, Edmonton (stencil).

BRAUN-BLANQUET ]. 1964 — Pflanzensoziologie. Grundzüge der Vegetationskunde

Wien.

BURNE B.M., 1974 — The climate of the Mackenzie Valley, vol. II. Environm. Canada,

Athm. Climatol. Studies n° 24.

DAHL Е. 1956 — Rondane mountain vegetation in South Norway and its relation to the

environment. Skr. Norske Vidensk.-Akad. Oslo, Mat.- Naturvidensk. KI. fasc. 3.

DOUGLAS G.W. and VITT. DH., 1976 — Moss-Lichen Flora of St. Elias-Kluane Ranges,

Southwestern Yukon. Bryologist 79 : 437-456.

Source : MNHN. Paris

EPIGAEOUS LICHEN SYNUSIAE 151

DU RIETZ G.E., 1925 — Gotlándische Vegetationsstudien. Svenska Váxtsociol. Sällsk.

Handl. 2.

FEOLI E., 1977 — On the resolving power of Principal Component Analysis in plant com-

munity ordination. Vegetatio 33 :119-125.

FREY E., 1927 — Bemerkungen über die Flechtenvegetation Skandinaviens, verglichen mit

derjenigen der Alpen. Veróff. Geobot. Inst. Rübel Zürich, 1927, 4.

FREY E., 1952 — Die Flechtenflora und Vegetation des Nationalparks in Unterengadin.

Ergebn. Wiss. Untersuch. Schweiz. Nationalparkes N.F., 3.

GAMS, H., 1938 — Über einige flechtenreiche Trockenrasen Mitteldeutschlands. Hercynia

Ок

GOODAL D.W., 1963 — The continuum and the individualistic association. Vegetatio 11 :

297-316.

HARKEMA W., 1976 — Die Flechten der Gletscherinsel Isla Persa (Bernina, Schweiz).

Nova Hedwigia 27 :619-630.

HOEFS M., COWAN I.M., KRAJINA V.J., 1975 — Phytosociological analysis and synthesis

of Sheep Mountain, Southwestern Yukon Territory. Syesis 8 Sup. 1 : 125-228.

HOEFS М., THOMPSON J., 1972 — Lichens from the Kluane Game Sanctuary, SW Yukon

Territory. Canad. Field Nat. 86 : 249-252.

KALB K., 1970 — Flechtengesellschaften der Vorderen Oeztaler Alpen. Diss. Bot. 9. 118 pp.

KLEIVEN N., 1959 — Studies on the xerophile vegetation in northern Gudbrandsdalen,

Norway. Nytt Mag. Bot. 7 :1-60.

KLEMENT O., 1955 — Prodromus der mitteleuropäischen Flechtengesellschaften. Feddes

Керегі. Вей. 135 :5-194.

KRIEGER H., 1937 — Die Flechtenreichen Pflanzengesellschaften der Mark Brandenburg.

Beih. Bot. Centralbl. Abt B. 57.

KORTEKAAS WM., VAN DER MAAREL E, & BEETFINK W.G., 1976 — A numerical

classification of european Spartina communities. Vegetatio 33 : 51-60.

LAUSI D. & FEOLI E., 1979 — Hierarchical classification of european salt marsh vegetation

based on numerical methods. Vegetatio 39 : 171-184.

LIMPRICHT W., 1931 Die Schneegruben des Riesengebirges als Naturschutzgebiet.

Feddes Repert. Beih. 62.

MATTICK, 1951 — Wuchs- und Lebensformen, Bestand und Gesellschaftsbildung der Flech-

ten. Bot. Jahrb. Syst. 75 : 378-424.

NIMIS P., 1980 — Caloplaca tominii Savicz, new to North America. (Manuscript in press).

ORLOCI L., 1978 — Multivariate analysis în vegetation research, ed. 2. The Hague. 451 pp.

PIGNATTI S., 1953 — Introduzione allo studio fitosociologico della pianura veneta orien-

tale. Forli.

TOMIN PM., 1931 — Bestimmungstabellen der Bodenflechten um den Halbwiisten von

Süd-Ost USSR. Keller Festschrift.

WIRTH V., 1972 — Die Silikatflechten-Gemeinschaften in ausseralpinen Zentraleuropa.

Diss. Bot. 17. 305 pp.

WESTHOFF V. VAN DER MAAREL E., 1978 — The Braun-Blanquet Approach, in :

WHITTAKER (ed.) Classification of Plant communities, ed. 2. The Hague. pp. 287-399.

Source : MNHN, Paris

153

THE TAXONOMIC STATUS

OF ENCALYPTA MICROSTOMA BALS. ET DE NOT.

AND E. CILIATA VAR. MICROSTOMA SCHIMP.

D. G. HORTON*

RESUME. — Encalypta microstoma est une espéce distincte qui a été confondue avec

E. ciliata var. microstoma. E. ciliata var. microstoma ne se différencie de E. ciliata que pour

la forme des capsules qui est plus ou moins aberrante. Des capsules intermédiaires entre

celles qui sont aberrantes et celles qui sont caractéristiques pour E. ciliata existent dans une

méme population, Par conséquent, les plantes á capsules aberrantes ne sont pas isolées

génétiquement. Encalypta microstoma se distingue de E. ciliata par ses coiffes á Бес plus

court et plus épais ét à frange brune, l'absence de péristome, ses capsules cylindriques

légèrement contractdes au sommet, et ses feuilles mutiques à marges planes. Il est endé-

mique: dans les montagnes d'Europe Centrale et du Sud, et a été récolté dans les Pyrénées

et le Massif Central en France, les Alpes suisses, autrichiennes et italiennes, les Tatras tchè-

ques, les Айаз bulgares et le Caucase. Une localité additionnelle dans le sud-est de la Nor-

vège est provisoirement rapportée.

ZUSAMMENFASSUNG. — Encalypta microstoma wurde ши Е. ciliata var. microstoma

verwechselt. Die letztere unterscheidet sich von der typischen E. ciliata nur durch die

Form der Kapsel, die mehr oder weniger abweichend ist. Zwischenformen zwischen diesen

abweichenden Kapseln und den charakteristischen für Е. ciliata kommen in derselben

Population vor; die Pflanzen mit abweichenden Kapseln sind demnach genetisch nicht

isoliert. Encalypta microstoma unterscheidet sich von E. ciliata durch ihre Haube mit

kürzerem, dickerem Schnabel, und mit braunen Fransen, das fehlende Peristom, die zylin-

drischen, oben verängerten Kapseln und die stumpf zugespitzten, flachrandigen Blätter. Sie

ist endemisch in den Bergen von Zentral- und Südeuropa und wurde bisher von den Pyre-

náen und dem Zentralmassif in Frankreich, den Alpen der Schweiz, Oesterreichs und Ita-

liens, der Tatra der Tschechoslowakei, der Rila von Bulgarien und vom Kaukasus bekannt.

Ein zusätzlicher Fundort vom Südosten Norwegens wird provisorisch erwähnt,

INTRODUCTION

BALSAMO and DE NOTARIS (1838) described Encalypta microstoma Bals.

et De Not. from a specimen collected by Brambilla in southern Switzerland.

In the original description the type locality is cited only as «Monte Айша».

* Department of Botany, University of Alberta, Edmonton, Alberta T6G 2E9.

Cryptogamie, Bryol. Lichénol., 1981, 2, 2 : 153-169.

Source : MNHN. Paris

154 D.G. HORTON

Later, in his Epilogo della Briologia Italiana, РЕ NOTARIS (1869) cited the 1о-

cality as «In Monte 5. Gottardo, in Helvetia...», Saint Gotthardo is actually loca-

ted west of the Adula Group, but is also in the Lepontine Alps. Although E. mi-

crostoma was described relatively early, up to the present time no other author has

accorded this taxon the specific status it warrants. (While C. MÜLLER (1848-1849)

included E. microstoma in his Synopsis, it is clear from the description that

it was a different taxon to which he referred). Encalypta microstoma has been

confused with what SCHIMPER (1856) described as E. ciliata Hedw. var. micro-

stoma Schimp.

Encalypta microstoma Balsamo et De Notaris, Mem. Reale Accad. Sci. Torino

40 : 342. 1838. TYPE : «In monte Adula, BRAMBILLA.» (Holotype : «Enca-

Тура microstoma DNtrs in monte Adula leg. Brambilla 1836. ex Prof. Balsa-

mo.» RO-De Notaris! Possible Isotype : «fr. Gottardo Encalypta microstoma

DNtrs Bals.» BM-Hampe!).

Figs. 1-5, 11-22.

— Encalypta ciliata Hedwig var. microstoma (Bals. et De Not.) Husnot, Musc.

Gall. : 198. 1888. Hom. illeg. of E. ciliata var. microstoma Schimp.

— Encalypta laciniata Lindberg var. microstoma (Bals. et De Not.) Brotherus,

Act. Soc. Sc. Fenn. 19 (12) :37. 1892.

— Encalypta ciliata Hedw. subsp. microstoma (Bals. et De Not.) Kindberg, Биг.

М. Am. Bryin. 2: 297. 1897.

— Leersia laciniata Hedw. ex Lindb. var. microstoma (Bals. et De Not.) Möller,

Bot. Not. 1907 : 142. 1907.

Plants to 20 mm tall, olive-green to bright-green above, brown to blackish

below: branched fasciculately, only very small plants unbranched. Stem in trans-

verse section with cells parenchymatous, somewhat irregular in size, walls

+ thickened, particularly at corners, and yellow to orange; epidermis 1-2 rows

smaller cells, walls thickened and dark orange; central strand absent. Leaves

when dry keeled by rather prominent costa, + sheathing below, somewhat

spread above, incurved and + twisted, laminae plane to slightly undulate and

incurved to conduplicate; when moist erect-spread to recurved-spread, lamina

inflexed; 2.34.3 mm long, 1.0-1.4 mm wide, oblong to ovate-, obovate- or

elliptic-oblong to elliptic, apex muticous and broadly acute to acute or very

shortly acuminate; margins plane; costa strong, subpercurrent to ending well

below apex, abaxial surface shiny and yellow to olive-yellow or brown, spar-

sely prorulose distally with low papillae, adaxial surface smooth below, papil-

lose laminal-type cells override in upper two-thirds, in transverse section cell

walls thickened, abaxial epidermous 2-3 rows cells with small lumina, guide

cells 1-2 rows with enlarged lumina and on either side 1 row cells with somewhat

smaller lumina, adaxial epidermous 1 row cells papillose as laminal cells but

slightly smaller. Upper laminal cells + isodiametric to subquadrate or short-

oblong, slightly angular with + rounded comers, (7)12-14(18) ит wide, (7)12-

14(20) um long, walls thin with some corners slightly thickened, papillae 3-6 per

Source - MNHN, Paris

ENCALYPTA MICROSTOMA AND E, СПЛАТА VAR. MICROSTOMA 155

cell, bifid to trifid with some indistinctly c-shaped, in transverse section walls even-

ly thickened, protrude more on adaxial surface, cell lumina do not extend into

base of papillae, or only very slightly; upper marginal cells differentiated in one

row, + rounded-subquadrate, 9-14 ит wide, 7-10 ит long, walls somewhat

thickened, otherwise as laminal cells; transitional cells between upper and basal

cells begin 2/3 to 3/4 below apex, arch over basal cells and form broad marginal

border almost to leaf base, + chlorophyllose, quadrate to oblong basipetally, outer-

most row marginal cells short to + elongate and narrowly oblong, grade into

broader cells towards costa, walls evenly thickened and smooth basipetally, pa-

pillae extend farther down abaxial surface; basal laminal cells markedly differen-

tiated by smooth, dark-orange walls thickened transversely, particularly at corners,

outer walls on both surfaces often аге porose or have membrane pleats, oblong

to slightly irregularly oblong, few quadrate, 16-48 ит long, 9-20 ит wide.

Goniautoicous. Perichaetia terminal, perigonia very short-stalked and late-

ral. Perichaetial leaves 2.0-3.0 mm long, sheathing, broadly ovate-lanceolate

to oblong-lanceolate, apex muticous and acute to shortly acuminate; margins

plane; costa ends in apex; basal cells less distinctly differentiated; otherwise

as vegetative leaves. Perigonial leaves 0.7-0.8 mm long, sheathing, oblong to

very broadly oblong, abruptly narrowed to shortly acuminate apex or to lanceo-

late upper portion and acute apex; margins plane; uppermost cells + papillose,

lower cell walls smooth and orange; otherwise + as vegetative leaves. Paraphyses

uniseriate throughout with upper cell walls firm and smooth.

Setae 2.5-6.0 mm long, erect to slightly flexuose, slightly twisted sinistrorsely

below, + strongly twisted dextrorsely in upper part into base of capsule, shiny

and yellow below and red just at base of capsule to dull orange or brownish

with age; in transverse section + 184 um in diameter, with central strand of small

cluster thin-walled cells surrounded by enlarged cells with thick walls, epidermal

cells 1-3 rows with very small lumina and strongly thickened walls. Capsules

1.2-3.0 mm long, when dry narrowly cylindric to cylindric, + tapered distally

and in some + contracted + below mouth to indistinctly constricted just below

slightly oblique, + narrow mouth, basally slightly constricted then gradually

tapered into seta, abruptly contracted to seta in very few populations, smooth,

young capsules greenish-yellow with rim red or undifferentiated, mature capsules

yellowish to copper-colored with rim red or undifferentiated, older capsules

dull copper-color; when moist as when dry; when old + wrinkled, collapsed

and longitudinally split; exothecial cells 41-93 um long, 12-20 um wide, long

to short-oblong with some slightly irregular, in + regular, longitudinal rows,

shorter near base of capsule and just below rim cells, in transverse section walls

strongly thickened on radial-longitudinal and adjacent tangential walls, < 9 um

thick; cells of rim in 2-5 rows, + regular rows longitudinally, but irregular trans-

versely, 5-23 ит long, 10-20 um wide, + quadrate to subquadrate to short-

oblong, walls + thickened; stomata scattered over surface, superficial, guard

cells 32-53 um long, 23-41 um wide, subsidiary cells undifferentiated, but pro-

minent ring -£ thickening outlines guard cells; peristome absent in most popula-

tions, when present very poorly developed with few, broken, hyaline, + smooth

Source : MNHN. Paris

156 D.G. HORTON

teeth in single row. Operculum when dry convex to slightly concave, rostrate,

1.0-1.2 mm long, lowermost 2-3 rows cells as rim cells of capsule. Annulus

persistent, undifferentiated from uppermost rim cells. Spores in polar view ra-

dially symmetric and circular, 3241 um, in equatorial view polarly asymmetric

and plano-convex, 23-28 ит x 3241 um, exine on proximal surface with nume-

rous radial placae and + distinct trilete mark, in some + irregularly puckered

around trilete mark; on distal surface 5-7 + prominent radial plicae extend from

+ distinct rim bordering + distinct central pit, in some pit, rim and plicae are

overlain by verrucose reticulum; both surfaces are minutely, * distinctly pitted.

Calyptrae 3.2-5.3 mm long, rostrum 1.0-1.8 mm long, cylindric, gradually

tapered above and then rather distinctly, but only slightly contracted to slightly

curved, erect in very few, thick rostrum, at base slightly expanded horizontally

to obliquely above pendent to flaring fringe of narrow to broad trapezoidal

segments; dark-golden with rostrum and fringe dark-brown, pale in few popula-

tions. Rostrum + smooth on outer surface, in transverse section of 5-6 rows

cells with very heavily thickened walls and small lumina, 2 innermost rows with

thinner walls and somewhat enlarged lumina; cylinder smooth on outermost

surface, in transverse section of 3 rows cells with very heavily thickened walls

and 1.2 innermost rows with thinner walls and enlarged lumina; fringe cells

short- to long-oblong, in transverse section of 2-3 rows cells, outermost with

exceptionally thick walls with 2 distinct layers of thickening, the outer yellow,

the inner hyaline, inner 1-2 rows have thin walls and enlarged lumina

SELECTED SPECIMENS EXAMINED. — AUSTRIA. Kärnten : «Hohe

Tauern, Wolligerhütte-Aüernig bei Mallnitz», Thyssen 18.7.1969 (as E. ciliata

var. microstoma М); «Goldegg Бей Spittal a.d. Drau», Froehl. 18.8.1960 (as E.

ciliata S-Froehlich); «Franz Josef Höhe, Gross Glockner», Nicholson & Dixon

21.8.1904 (as E. ciliata var. microstoma BM-Dixon, G-Delessert). Salzburg .

«Südseite des Pihaper 6. Mittersill im Pinzgau», Breidler 25.8.1879 (as E.

ciliata var. microstoma BP-Limpricht); «Alpe 'Storz bei Mur im Lungau»,

Breidler 26.8.1878 (as E. ciliata В microstoma FI). Steiermark : «'Hasenkar' in

den Schieferalpen bei Schladming», Breidler 6.9.1877 (as E. microstoma W + E.

ciliata with «aberrant» capsules); «Arkogel bei Schöder», Breidler 3 8.1875 (as

E. microstoma BP-Limpricht, H-Lindberg; as E. microstoma ciliata W-Juratzka);

«Paalgraben bei Stadl», Breidler 4.7.1878 (as E. ciliata var. microstoma W, as

E. ciliata c. forma gymnostoma H-Lindberg + E. ciliata, as E. ciliata in E. miero-

stoma übergehend BP-Limpricht). Tirol : «Innervillgraten», Gander 14.8.1884

(as E. microstoma O, M-Hiendlmayr - some «aberrant» capsules); «Pitztal/Imst»,

Рай 24.6.1977 (as E. ciliata DUIS).

BULGARIA. «M. Rila Orientalis, Kamenliv Čal», Petrov 25.7.1954 (as E.

ciliata var. microstoma BP, KRAM, 5).

CZECHOSLOVAKIA. «Vysoké Tatry, Predná Bašta», Krajina 2.8.1932 (as

E. rhabdocarpa var. leptodon PRC + E. brevipes).

FRANCE. Haute-Savoie : «col l'Aiguillet du Brevent, Massif du Mont Blanc»,

Guinet 29.7.1904 (as E. ciliata var. microstoma G-Guinet). Puy de Döme :

Source : MNHN, Paris

PL. I. — Figs, 1-5 : Encalypta microstoma. Fig.1. Habit (Düll 24.6.1977 DUIS). Fig. 2.

Variation in calyptrae (left to right : Gander 14.8.1884 М, Brotherus 87 H-Lindberg,

Coll. illegible L-Sande Lacoste, Breidler 4.7.1878 H-Lindberg, Brotherus 87 H-Lindberg).

Fig. 3. Variation in capsules. Capsule in centre is young, the one on extreme right ap-

pears slightly «aberrant» at the mouth (left to right : Petrov 25.7.1954 KRAM, Gander

14.8.1884 М, Brotherus 87 H-Lindberg, J.B. 3.8.1875 H-Lindberg, Gander 14.8.1884

М). Fig. 4. Variation in vegetative leaves (left to right : Gander 14.8.1884 М, Breidler

4.7.1878 H-Lindberg, Breidler 4.7.1878 H-Lindberg, Juratzka 56 M, Coll. illegible NY).

Fig. 5. Variation in perichaetial leaves (left to right : Düll 24.6.1977 DUIS, Breidler

4.7.1878 H-Lindberg). Lines =1 mm.

Source : MNHN. Paris

158 D.G. HORTON

«Mont Dore», Lamy (as E. ciliata M-Progel). Pyrénées Atlantiques . «Pic de la

Sagette», Allorge 8.7.1951 (as E. ciliata PC); «au-dessus du 3me lac d'Ayous»,

Allorge 10.7.1951 (as E. ciliata PC). Pyrénées Orientales : «Vallée d’Eyne»,

sin. coll. 49 (as E. ciliata ВМ + E. ciliata).

ITALY. Lombardy : «Rabbi», Venturi (as E. microstoma W-Juratzka);

«Monte Adamello», Kern 27.7.95 (as E. microstoma PC). Lombardy-Trentino-

Alto Adige : «Tonale», sin. coll. 27.8.1865 (as E. ciliata forma depauperata М),

Trentino-Alto Adige : «Brenner», Arnold (as E. ciliata microstoma S-Möller).

Valle d'Aosta : «Parco Nazionale Gran Paradiso; between Valnontey and Rifu-

gio Vittorio Sella, ca. 5 km S.W. of Cogne», van Driel 29.7.1966 (as E. ciliata

DUIS).

NORWAY. Buskerud : «Lige op for Skjønne i Моге» sin. coll. 27.7.1890 (as

E. mutica O-Kiaer).

SWITZERLAND. Bern : «Faulhornspitze», sin. coll. (as E. microstoma BM-

Hampe). Graubünden : «Muottas Murail», Fürbringer 14.8.1910 (as E. ciliata Н);

«beim Berninahospiz», Herzog 10.1909 (as E. microstoma M-Paul). Ticino :

«in monte Adula», Brambilla 1836 (as E. microstoma RO-De Notaris); «Canton

Tessin Fusio», Trautmann 5.7.1908 (as E. ciliata В microstoma S-Möller). Va-

lais : «Bérisal (Valais)», Philibert 20.7.1889 (as E. ciliata var. gymnostoma PC);

«Simplon Pass», Nash 19.7.1978 (ALTA, priv. herb. Р.С. Horton + E. vulgaris

s. lat.); Val d'Orny, above Orsiéres, Horton 19018 (ALTA, priv. herb. Р.С. Hor-

ton); Région de Crans et Montana, Chetseron, above Sierre, Horton 19054

(ALTA, priv. herb. D.G. Horton).

US.S.R. «Plantae Caucasicae, Мен, Brotherus» (as Leersia laciniata-micro-

stoma H-Lindberg; as E. ciliata var. microstoma H-Brotherus); «Kuban montis

Elbrus», Sommier & Levier 9.9.1890 (as Е. ciliata FI + Е. ciliata); «Svania, in

jugo alpino Latpari», Sommier & Levier 5.8.1890 (as Б. ciliata FI).

DISTINGUISHING FEATURES. — Encalypta microstoma is chatacterized

by smooth, golden-brown calyptrae, each with a relatively broad, slightly curved

or erect rostrum and a dark-brown, glisteny, well-defined fringe, which hangs

more-or-less pendent from a narrow, horizontal or slightly oblique expansion

at the base of the cylinder; gymnostomous capsules (а fragmentary, rudimentary

peristome has been observed in very few populations) that are narrowly cylindric

to cylindric, tapered slightly distally and more-or-less distinctly contracted

distally from just below the small mouth, tapered broadly into the seta, and

yellowish to copper in color; and muticous leaves with plane margins and а

prominent costa which ends just below the apex, except in the perichaetial

leaves (in these it is percurrent). Microscopically, the transverse section of the

capsule wall is distinctive. The radial longitudinal and outer tangential walls

of the exothecial cells are very strongly thickened giving the appearance of a

cut-out paper chain of tulips; in superficial view these thickenings almost obs-

cure all but a narrow, central strip of the cell lumina. The transitional cells

with evenly thickened, yellowish walls arch over the basal laminal cells and form

a broad, marginal border almost to the base of the leaf. The basal laminal cells,

Source : MNHN, Paris

ENCALYPTA MICROSTOMA AND Е. СИЛАТА VAR. MICROSTOMA 159

which are broader, are differentiated by dark-orange walls that are thickened

transversely, particularly at the corners. Generally, a single large and somewhat

irregular pore occurs in the cell wall on both the abaxial and adaxial surfaces

of some of these lower cells. The spores are distinctly polarized with a more-or-

less distinct trilete mark and numerous, radial plicae on the proximal surface;

the distal surface is more variable with a more-or-less distinct central pit borde-

red by a thick, more-or-less prominent rim connected to 5 to 7 more-or-less

prominent plicae. In some cases, the pit, rim and plicae are overlain by anasto-

mosing verrucae, which give a reticulate impression under the light microscope.

Both surfaces are more-or-less distinctly, minutely pitted.

The striking morphological similarity between Е. microstoma and Е. ciliata

leaves no doubt as to the phylogenetic derivation of the former taxon, Indeed,

these two species are but narrowly differentiated from one another; however,

there are a number of character-states in which the two taxa consistently differ.

Encalyptra microstoma can be differentiated from E. ciliata by the darker color

of the calyptrae, including the fringe, which is a rich brown; calyptrae of Е.

ciliata are generally pale-golden throughout, although the fringe may be pale-

brown in some populations. Also, the shape of the calyptra is different; in E.

ciliata the cylinder is broader, which makes the rostrum appear narrower than

in E. microstoma (Fig. 8). The length of the rostrum of E. ciliata is quite consis-

tently 1/3 the overall length of the calyptra, while in many populations of E.

microstoma the rostrum is only 1/4 the total length. Capsules of E. ciliata are

distinctly constricted just below the mouth and abruptly narrowed to the

seta; in contrast, those of E. microstoma are contracted to the mouth (only

rarely constricted and then less distinctly so than in В. ciliata) and more gra-

dually narrowed into the seta (Fig. 7). A well-developed peristome is characte-

ristic of E. ciliata, but absent in E. microstoma (when a fragmentary peristome

does occur, it is always hyaline, never with dark-orange teeth as in E. ciliata).

The setae in populations of E. ciliata are generally longer (up to 15 mm, while

those of Е. microstoma are only up to 6 mm); although they are shorter in some

alpine populations of Е. ciliata. However, in a mixed collection of Е. ciliata

and E. microstoma, setae of the former taxon varied from 5-7 mm in lenght,

while those of the latter were 4 mm. In populations of E. ciliata, the leaves are

mucronate and the margins are always narrowly recurved in the lower 1/2 to

2/3, in contrast to the muticous leaves with plane margins of Е. microstoma.

Leaves of E. ciliata tend to be longer and therefore are more strongly contorted

than those of E. microstoma. Microscopically, E. microstoma and E. ciliata

differ in the transverse section of the capsule wall with the outer tangential

walls of the exothecial cells in Е. ciliata strongly and evenly thickened in con-

trast to the strong, localized thickenings characteristic of E. microstoma.

These consistent morphological differences and the occurrence of mixed popu-

lations indicate genetic isolation of Е. microstoma and Е. ciliata. There is no

doubt that these taxa are distinct at the specific level.

The vast majority of populations of E. ciliata are concentrated in the Nor-

thern Hemisphere north of 30°N. These are relatively uniform in the character-

states discussed above, except that «aberrant» capsules (see below) occur quite

Source : MNHN, Paris

160 D.G.HORTON

pl, I1. — Figs. 6-9 : Comparison of Encalypta microstoma with E. ciliata. Fig. 6. Capsule

of E. ciliata from Peruvian collection that parallels Е. microstoma in the lack of a peri-

stome and the narrowed mouth (Р. & Е. Hegewald 9191). Fig. 7. Capsules of E. micro-

stoma (left) and E. ciliata (right) from mixed collection (Breidler 4.7.1878 H-Lindberg).

Fig. 8. Calyptrae of E. ciliata (left) and E. microstoma (right) from mixed collection

(Breidler 4.7.1878 H-Lindberg). Fig. 9. Calyptra of E. ciliata from Mexican collection

that parallels Е, microstoma in the less distinctly defined fringe and rostrum (Delgadillo

2317a MEXU). Fig. 10. Series of capsules from a single collection of E. ciliata to

illustrate intergradation from those «typical» of E. ciliata (left) to extreme expression of

«aberrant» E. ciliata capsules that are cleistocarpous (right) (Dúll 5.8.1968 DUIS).

Line =1 mm.

Source : MNHN, Paris

ЕМСАГҮРТА MICROSTOMA AND Е. СПЛАТА VAR. MICROSTOMA 161

frequently. However, in populations of E. ciliata from the Southern Hemisphere

and south of 30°N in the Northern Hemisphere, there is considerable variation

in calyptra, capsule, peristome and leaf structure. This variability is exemplified

in collections from Mexico and South America. Some of these populations

mimic E. microstoma in calyptra and capsule shape, and the lack of a peristome

(Figs. 6 + 9). As well, the calyptra fringe is pale brown in many populations

and, in some, is even darker brown than in E. microstoma. Populations with

muticous leaves are of sporadic occurrence. However, these «Southern Hemi-

sphere» populations of E. ciliata are consistently differentiated from E. micro-

stoma by recurved leaf margins.

The identity of Е. microstoma has been obscured because the forms of its

capsule parallel those which occur relatively frequently in populations of E.

ciliata. Generally, E. ciliata is characterized by a well-developed peristome,

and smooth, cylindric and turgid-looking capsules with a distinct constriction

just below the mouth, the diameter of the latter being more-or-less equal to that

of the capsule below the constriction (Fig. 7 - capsule on right). Such capsules

are quite distinct from those of E. microstoma; however, there are populations

of E. ciliata that are characterized completely or in part by capsules with quite

a different morphology, which I refer to as «aberrant». The range of morpholo-

gical variation of «aberrant» capsules of Е. ciliata is considerable and grades

imperceptibly into capsules with morphology typical for Е. ciliata, described

above. The forms that resemble capsules of E. microstoma represent only a

small portion of this variability.

Capsules of E. ciliata that represent the most extreme expression of «aber-

rant» capsules are cylindric and more-or-less smooth and turgid-looking, some

are tapered distally, and there is always a distinct constriction or an abrupt

contraction just below the minute mouth, the latter being considerably less

than the diameter of the central part of the capsule. In some cases, the capsule

is cleistocarpous (with the operculum remaining attached and no line of dehis-

cence apparent); in others, the operculum dehisces, but the peristome teeth

are very irregular or lacking altogether. The mouth is similarly very irregular.

Often «aberrant» capsules occur intermixed in populations that have capsules

of the shape typical for E. ciliata, and in a few specimens a complete morpho-

logical gradient from the extreme «aberrant» capsules to those characteristic

of E. ciliata occurs (Fig. 9). Some of these less strikingly «aberrant», interme-

diate forms are virtually indistinguishable from capsules of Е. microstoma,

except for the presence of a peristome (Fig. 10 - middle).

The occurrence of intergrading forms between typical E. ciliata capsules and

«aberrant» capsules within individual populations indicates that the latter

probably do not represent a habitat modification. In all other macroscopic

and microscopic features, the plants with «aberrant» capsules are indistin-

guishable from those typical of E. ciliata in the narrowest sense, The abnormal

appearance of the extreme «aberrant» capsule forms (Fig. 9) suggests that

such capsules may be the product of some developmental irregularity. While

there may be some genetic basis causing this irregularity, it is clear that such

Source : MNHN, Paris

162 D.G.HORTON

Р.Ш. — Figs. 11-16 : Encalypta microstoma, Fig. 11. Mouth of gymnostomous capsule

with narrow rim differentiated by short exothecial cells (Breidler 4.7.1878 H-SOL),

x 150. Fig. 12. Transverse section through central part of capsules showing the charac

teristic «tulip» thickenings of the radiallongitudinal and outer tangential walls of the

exothecial cells (Dull 24.6.1977 DUIS), x 425. Figs. 13-14. Variation in proximal

surface of spores. Both аге characterized by a + distinct trilete mark, numerous radial

plicae and + distinctly, finely pitted surface. Fig. 13. Note irregularity of exine around

indistinct trilete mark (Juratzka 56 М). x 1300. Fig. 14. Note distinct trilete mark

and distinctly, finely pitted surface (Petrov 25.7.1954 KRAM), x 1400. Figs. 15-16.

Variation in distal surface of spores. Both have + prominent, radial plicae extending

from + distinct central rim and + distinctly, finely pitted surface, Fig. 15. The rim and

radial plicae are distinct with some gross irregularity toward upper part of fig. and the

surface is indistinctly, finely pitted (Breidler 4.7.1878 H-SOL), x 1200. Fig. 16. Plicae

and rim are indistinct and are overlain by verrucose reticulum. Surface is distinctly,

finely pitted (Düll 24.6.1977 DUIS), x 1300.

Source - ММНМ. Paris

ENCALYPTA MICROSTOMA AND Е. СПЛАТА VAR. MICROSTOMA 163

PL IV. — Figs. 17-22 : Encalypta microstoma (Petrov 25.7.1954 KRAM). Fig. 17. Superfi-

cial view of pluripapillose upper laminal cells, adaxial surface, x 800. Fig. 18. Trans-

verse section through upper laminal cells, adaxial surface uppermost. Note that cell walls

protrude more strongly on adaxial than on abaxial surface, x 1200. Fig. 19. Lower la-

minal cells, abaxial surface. Note gradation from upper laminal cells with strongly pluri

papillose walls through transitional cells to markedly differentiated basal cells with

smooth, porose walls. Transitional cells with walls sparsely papillose or smooth arch

over basal cells and form broad marginal border, x 110, Fig. 20. Closeup of abaxial

surface of basal cells. Note large «pores» on both surfaces where the wall appears rather

irregularly torn, x 375. Fig, 21. Short-stalked perigonium attached to base of costa on

abaxial surface, as is characteristic for many species of Encalypta,x 40. Fig. 22. Trans-

verse section of stem. Cells parenchymatous and thin-walled, walls somewhat thickened

in outermost 2-3 layers, Central strand absent, x 225

Source : MNHN, Paris

164 D.G. HORTON

forms are not in any way genetically isolated from «normal» E. ciliata. There-

fore, they should not be given any formal nomenclatural recognition. Such a

conclusion is further supported by the occurrence of similarly «aberrant»

capsules in such other species of Encalypta as E. affinis Hedw. £., E. alpina Smith

and E. brevicolla (B.S.G.) Bruch ex Aongstr., which indicates that the genetic

potential for the formation of such capsules traverses interspecific as well as

infraspecific boundaries. I have also observed what might be interpreted as

«aberrant» capsules (Fig. 3 - extreme right) in a few populations of E. micro-

stoma.

HABITAT. — Populations of E. microstoma have been quite frequently

collected with plants of E. ciliata intermixed, which suggests that these two

closely related species have very similar habitat preferences. Although popula-

tions of E. ciliata occur in alpine and arctic tundra, they are more frequently

associated with more mesic, forested habitats at lower elevations. Therefore,

this species is widespread outside of mountainous regions. It is apparent from

the distribution of E. microstoma that this species is, in contrast, restricted to

montane habitats. The data on collection labels of E. microstoma are meagre, but

elevations of 1000 to 2700 m have been recorded so it can be inferred that popu-

lations occur from below treeline up into alpine tundra. I have collected E. mi-

crostoma at two different localities in the Swiss Alps; one was in alpine tundra

at approximately 2700 m where it occurred on a dry, rocky slope with west

aspect, and the other was at 2100 m in the upper subalpine Larix europaea DC.

zone where it occurred on a dry rock outcrop with west aspect. The apparent

dryness of both habitats was notable by comparison to E. ciliata. The latter

species tends to grow їп slightly mesic habitats where there will be a little inter-

mittent seepage. At the first locality E. microstoma was growing on soil beneath

the edge of a small solifluction terrace where it was well protected by an over-

hanging tuft of Festuca haleri All. and by surrounding granite outcrops. In the

immediate area large mounds of Silene acaulis L. were a prominent feature

indicative of the exposed nature of the vegetation, which also included Agrostis

rupestris АП. Alchemilla alpina (L.) Scopoli, Juncus trifidus L. Phyteuma

hemisphaerica L. and Saxifraga bryoides L. Tortella tortuosa (Hedw.) Limpr.

was relatively abundant on soil in the vicinity. At the second locality, E. micro-

stoma was also well-protected, growing back in the crevices of the siliceous

outcrop with Tortella tortuosa, and with Pterigynandrum filiforme Hedw.

in more exposed positions on the face of the rock. Particularly indicative of the

xeric nature of this habitat was the dominance of Juniperus communis L.

bushes, Also present were Campanula cochlaerifolia Lam., Minuartia laricifolia

(L.) Schinz et Thell., Saxifraga paniculata Miller and Sempervivum arachnoidum

L. The relative dryness of both these habitats and the restriction of E. microsto-

ma to montane regions may indicate a shift in some aspects of the habitat

preferences of this species away from those of E. ciliata.

Another important feature of the habitat of E. ciliata is that this species

frequently grows where the substrate is siliceous, although it also occurs on cal-

careous substrates. In western North America the pH of soil on which popula-

Source : MNHN. Paris

ENCALYPTA MICROSTOMA AND Е. CILIATA VAR. MICROSTOMA 165

tions of E. ciliata were growing ranged from 5.3 to 7.1 (п = 50), but the mean

was slightly acidic at 6.3. Species that are frequently associated with E. micro-

stoma include such indicators of more siliceous conditions as Bartramia ithy-

phylla Brid., while such others as Eurhynchium pulchellum (Hedw.) Jenn. and

Myurella julacea (Schwaegr.) В.5.С. are generally associated with calcareous

conditions. As noted above, the rocks at both the localities where I collected

E. microstoma were siliceous, and pH of the soil was 5.7 at the first locality and

5.8 at the second. Therefore, populations of E. microstoma probably occur

under very similar conditions to those ofE.ciliata with regard to substrate-types.

DISTRIBUTION. — Encalypta microstoma has a relatively restricted distri-

bution across the montane regions of central and southern Europe from the

Pyrénées Atlantiques, the Pyrénées Orientales, and the Massif Central in France,

through the Swiss, Austrian and northern Italian Alps, the High Tatras of Cze-

choslovakia, and some what disjunct in the Rila Mountains of Bulgaria and the

Caucasus Mountains. An additional locality disjunct in southeastern Norway is

tentatively reported (open circle) (Fig. 23).

РІ. V.— Fig. 23. The distribution of E. microstoma as it is presently known.

Source : MNHN. Paris

166 D.G. HORTON

DISCUSSION

As a result of the confusion of capsules of E. microstoma with «aberrant»

capsules of E. ciliata, and the occurrence of morphological intergradation

between these «aberrant» capsules and those typical for E. ciliata, E. microsto-

ma has generally been regarded as a variant of E. ciliata. With the exceptions

of the original authors and HUSNOT (1888), bryologists have generally consi-

dered E. microstoma to be a synonym of E. ciliata var. microstoma Schimp.

The unfortunate, coincidental use of the epithet microstoma by SCHIMPER

(1856) when he described E. ciliata var. microstoma has only increased the

complexity of this problem. SCHIMPER (1856) described E. ciliata var. micro-

stoma in the «Corollarium». No type specimen is cited and I have been unable to

locate any specimens in his herbarium in BM. However, from his description

«capsulae orificium valde angustatum dentibus minoribus irregularibus», I think

that it is very likely that SCHIMPER was describing «aberrant» capsules of

Е. ciliata and not Е. microstoma. The capsules of Е. microstoma lack a peristo-

me, except in a few cases when rudimentary, hyaline bits of teeth are present;

in «aberrant» capsules of E. ciliata, the teeth, when present, are always dark-

orange, although they are characteristically more-or-less irregular in shape and

number. The peristome is also entirely lacking in some «aberrant» capsules.

As I have observed bits of a peristome in only a few populations of E. microsto-

ma, and as their transparency makes them difficult to discern, I think it much

more likely that the small, irregular teeth described by SCHIMPER referred

to the peristome of «aberrant» capsules of E. ciliata, which is frequently present

and readily apparent. Furthermore, since there is no mention of BALSAMO and

DE NOTARIS' (1838) E. microstoma in the text of either Bryologia Europaea

(BRUCH, SCHIMPER & GÜMBEL 1838) or in the Corollarium (1856), it also

seems probable that SCHIMPER was unaware that the epithet microstoma had

been used prior to his own description of E. ciliata var. microstoma. (Encalypta

microstoma is listed in the Index to Bryologia Europaea with the notation

«= E. ciliata var.». The Index was published in 1855 (cf. LIMPRICHT 1895),

which just predates publication of the Corollarium (1856), so it seems quite

feasible that the short Index might actually have been written after the Corol-

larium was completed.) Therefore, I have concluded that E. ciliata var. microsto-

ma was not a new combination of BALSAMO and DE NOTARIS' name by

SCHIMPER as indicated in Index Muscorum (VAN DER WIJK et al. 1962).

Rather, E. microstoma Balsamo et De Notaris and E. ciliata Hedwig var. micro-

stoma Schimp. are considered to represent heterotypic names.

In the Synopsis Muscorum Europaeorum, SCHIMPER (1876) included E.

microstoma in the synonymy of his E. ciliata var. microstoma and changed his

description slightly to include (in translation) «peristome teeth very small and

irregular to clearly lacking». In view of this and the fact that one of the figures

in Bryologia Europaea (BRUCH, SCHIMPER & GÚMBEL 1838), PI. 200,

Fig. 20, appears to be a capsule of E. microstoma, 1 suspect that SCHIMPER

probably did see specimens of E. microstoma, but was confused by the com-

Source : MNHN. Paris

ENCALYPTA MICROSTOMA AND Е. СПЛАТА УАК. MICROSTOMA 167

plexity of the inter-relationships between E. microstoma and «aberrant» cap-

sules of E. ciliata (his E. ciliata var. microstoma). Even before SCHIMPER

described E. ciliata var. microstoma, С. MULLER’s (1849) description of E.

microstoma with mucronate leaves and revolute margins clearly refers not to

this species, but to E. ciliata with «aberrant» capsules.

In the Epilogo della Briologia Italiana, DE NOTARIS (1869) reaffirmed his

belief in the distinctness of E. microstoma. Under E. ciliata, he included the

var. microstoma Schimp., but made it very clear that he considered this to be

something different than his and BALSAMO’s E. microstoma, with the notations

«Schimp. 1. с. excl. E. microstoma Bals. et DNtrs.» and «E. microstoma. Bals et

DNtrs a varietate В. hujusce speciei mihi distincta videtur.». E. microstoma Ваз.

et De Not. is treated later in this same work and among the character-states

that he listed as differentiating this species from E. ciliata is the nerve ending

before the leaf apex. This feature differentiates E. microstoma from all forms

of E. ciliata, except some Southern Hemisphere populations (see Distinguishing

Features). LIMPRICHT (1895) appears to have been confused by DE NOTARIS’

contention that his E. microstoma was different from SCHIMPER's E. ciliata

var. microstoma. Under E. ciliata var. microstoma, which is listed as a new com-

bination by SCHIMPER of BALSAMO and DE NOTARIS’ name, LIMPRICHT

noted (in translation) «In the higher mountain exposures transitions frequently

occur to the variety В; De Notaris knew these intermediate forms for he still

separated in Epil. р. 323 from his E. microstoma an E. ciliata В microstoma

Schimp.». However, LIMPRICHT concluded (in translation) «Also against the

specific nature [of var. microstoma] is the occurrence of a parallel form in E.

rhabdocarpa.».

HUSNOT (1884-1890) clearly considered E. microstoma Bals. et De Not.

and E. ciliata var. microstoma Schimp. to be two different entities. He made a

new combination of Е. ciliata var. microstoma (Bals. et De Not.) Husnot, with

the notation «non Schimpr.» and the inclusion of E. microstoma Bals. et De

Not. in synonymy, making it clear to what he was referring. Then, perhaps in an

effort to end the confusion with names, he described E. ciliata var. gymnosto-

ma Bruch ex Husnot and included in its synonymy E. ciliata var. microstoma

Schimp. (thereby invalidating his name), and pointed out «non De Not». HUS-

NOT cited several specimens of both E. ciliata var. gymnostoma and his E. cilia-

ta var. microstoma. One of the specimens of the var. gymnostoma cited by him

is in his herbarium in PC (!). Contrary to HUSNOT's notation (see above),

which indicated that this specimen should be Е. ciliata with «aberrant» capsules,

it is actually E. microstoma. It is apparent not only from this specimen, but also

from the illustrations in the Muscologia Gallica that HUSNOT did not under-

stand the difference between E. microstoma and E. ciliata var. microstoma

Schimp.

Encalypta microstoma is a rare species, endemic to central and southern

Europe. In each of the herbaria examined, there exist only a few specimens,

and less than 10 in total have been collected since 1950. The concentration

of collections from localities in central and southern Austria-Switzerland and

Source : MNHN. Paris

168 D.G. HORTON

northern Italy indicates that this region can be considered the centre of its

range. It is to be expected from Jugoslavia and possibly Hungary. The disjunct

locality in southeastern Norway is reported with some hesitation, as there is

only a single specimen from 1890 and the collector is not indicated. Possibly

the occurrence of E. microstoma in Scandinavia can be confirmed in the future

on the basis of further collections. By comparison to most other species of

Encalypta, the distribution of E. microstoma 15 markedly restricted. For ехат-

ple, È. ciliata and E. rhabdocarpa Schwaegr. are relatively common and wide-

spread in montane and boreal habitats of the Northern Hemisphere. Even such

species as E. brevipes Schljak., E. brevicolla (B.S.G.) Bruch, E. longicolla Bruch,

E. mutica Hagen and E. spathulata C. Müll., all of which have been suggested

by various authors to have relictual distribution patterns, occur both in North

America and Eurasia. Only two recently described species, E. vittiana Horton

and E. flowersiana Horton (HORTON 1979), appear to have distributions as

restricted as that of E. microstoma. Encalypta vittiana is known from northwes-

tern North America in Alaska, the Yukon Territory and western Northwest

Territories, while E. flowersiana has been reported from Guatemala and Haiti,

and an additional locality in Texas can now be added (U.S.A. Texas : Brewster

Co., Chisos Mtns., Big Bend Ntn'l. Pk., Magill 2036, 2040 MO!). The restriction

of each of these three species to a single continent, in contrast to almost all

other species of Encalypta, suggests that E. microstoma, E. flowersiana and

E. vittiana have evolved relatively recently.

ACKNOWLEDGEMENTS. — am especially grateful to the following curators who

made this study possible by the loan of specimens : R. Ross (BM), M. Rajczy (BP), J.-P.

Frahm (DUIS), J. Engel (E), С. Steinberg (FI), Р. Geissler (С), Р. Isoviita and Т. Koponen

(н), В. Ochyra (KRAM), А. Touw (L), G. Müller (LAU), Н. Hertel (М), Р. Holmgren (NY),

H. Krog (0), В. Baudoin (PC), J. Váňa (PRC), I. Ricci (RO), E. Nyholm (S), U. Passauer

(W), and to Tom Nash for the specimen from Simplon Pass. Also particular thanks to

Carlo Steinberg for assistance їп obtaining the type of E. microstoma, to Walter Moser for

translation of old German script and assistance with some Austrian localities, and to Hélène

Bischler for localities in the Pyrénées and for rendering the French and German summaries

comprehensible. Patricia Geissler was most kind in taking me to collect E. microstoma at

the Caban d’Orny and the identifications of associated vascular plants at both Swiss loca-

lities are hers. Dale Vitt kindly reviewed the manuscript.

BIBLIOGRAPHY

BALSAMO J. & DE NOTARIS J., 1838 — Pugillus muscorum Italiae novorum vel minus

cognitorum. Mem. Reale Accad, Sci. Torino 40 : 327-351.

BRUCH Ph., SCHIMPER W. Ph & GÜMBEL Th., 1838 — Bryologia Europaea seu Genera

Muscorum Europaeorum Monographice Illustrata III (4) : 1-16, pl. 1-8.

DE NOTARIS J., 1869 — Epilogo della Briologia Italiana. Genova.

Source : MNHN, Paris

ENCALYPTA MICROSTOMA AND Е. CILIATA VAR. MICROSTOMA 169

HORTON D.G., 1979 — Encalypta vittiana sp. nov. and E. flowersiana sp. nov. from North

America. Bryologist 82 : 368-381.

HUSNOT T., 1884-1890 — Muscologia Gallica I. Paris.

LIMPRICHT K.G., 1895 — Die Laubmoose Deutschlands, Oesterreichs und der Schweiz.

In RABENHORST, Kryptogamen-Flora von Deutschland, Oesterreich und der Schweiz

4 11-853.

MÜLLER C., 1848-1849 — Synopsis Muscorum Frondosorum Omnium Hucusque Cogni-

torum. I. Berlin.

SCHIMPER W. Ph., 1856 — Corollarium Bryologiae Europaeae, Conspectum Diagnosticum

Familiarum, Generum et Specierum, Adnotationes Novas Atque Emendationes Complec-

tens. Stuttgart, 1855 «1856».

SCHIMPER W. Ph., 1876 — Synopsis Muscorum Europaeorum Praemissa Introductione de

Elementis Bryologicis Tractante. Stuttgart.

Source : MNHN. Paris

171

GROWTH RATES OF XANTHORIA PARIETINA

AND THEIR RELATIONSHIP TO SUBSTRATE TEXTURE

ТН. МОХНАМ*

SUMMARY. — Growth rates of Xanthoria parietina are reported and comparisons are made

with previously recorded growth rates for this species. The effect of substrate texture on

growth rate is also discussed.

INTRODUCTION

Lichens, for their final size at maturity, are a relatively slow growing group of

plants, and studies on their growth and growth rates are inevitably long-term

projects. However, it becomes apparent from even short term observations of

the growth of a number of individuals of the same species, that variations in

the rate of growth occur throughout the seasons (FISHER & PROCTOR 1978).

This is, of course, due to a multiplicity of factors (HARRIS 1972), of which

rainfall has been shown to be the most important (е. р. FISHER and PROCTOR

1978, KARENLAMPI 1971). HALE (1973), ARMSTRONG (1975) and TO-

PHAM (1977), in reviewing our current knowledge of lichen growth, discuss the

different factors affecting growth, the many methods of growth measurement,

and various methods of interpreting results. Bearing in mind the different

aspects they discuss, the published results of growth measurements on Xanthoria

parietina have been recalculated, and it is suggested from the data presented

below that the growth rate of X. parietina, especially in juvenile thalli, is influen-

ced by the texture of the substrate.

METHODS

Five lichens, initially between 2-22 mm diameter, growing on brushed, buff

concrete pavers were photographed at approximately monthly intervals during

the period November 1976-April 1978 using a Nikon F camera with a micro-

* Department of Plant Sciences, School of Biological Sciences, University of Bath, Claverton

Down, Bath, Avon, BA2 7AY.

Cryptogamie, Bryol. Lichénol., 1981, 2, 2 : 171-180.

Source : ММНМ. Paris

172 T.H. MOXHAM

Nikkor lens on Kodak plus X pan film (ASA 125). The camera was mounted

via a bosshead and screw to a retort stand with an «A» base. This guaranteed

а 90° angle between camera and thallus. The camera was positioned approxi-

mately 25 cm above the substrate giving a final reduction on the negative of

about 1 : 0.35. Photographs were taken only on dry days, and during wet

weather a full dry day was allowed to elapse before photographs were taken.

А clear plastic rule, graduated in 1mm units was included in each photograph.

The negatives were enlarged 1x-4x original size onto single weight glossy paper

and the graduations of the rule were measured with pointed vernier calipers

to produce an individual magnification factor for each print. The area of the

thallus was measured with a planimeter and the thallus, for the sake of equa-

tion b) was considered to be circular.

lanimeter readin;

a) toe E as cia de

(magnification)?

and a mean diameter from :

b) diameter nf x

Several (5) diameters were measured from each photograph to calculate a

mean diameter for comparison with the diameter derived from the area (Tab. 1).

Meteorological data (which included mean vapour pressure, wind run, wet and

dry bulb temperatures, sunshine hours and precipitation, from which a modified

Penman equation was calculated) was recorded during the period of investigation

from the Wessex Water Authority’s Weather Station at Bath University (grid

ref. ST76/776643, alt. с. 180 та) approximately 500 m from the lichens investi-

gated.

RESULTS

Fig. 1 shows the growth of one of the thalli investigated (A/1), and Tab. 1

shows the area and growth of the thalli at monthly intervals. Even though

care was taken to allow at least one dry day to elapse before taking photographs,

there were times, especially during the dry summer months, when the thallus

had contracted more than the previous measurement. Consequently only posi-

tive readings have been used when presenting the data in the figures, Rainfall

had the greatest influence on growth, showing a positive, though weak (r = 0.61,

0.02 > p > 0.01) correlation with growth rate. The other weather data showed

no significant correlation with growth, although it was noted that there was

more thallus contraction with increased sunshine hours and low relative humi-

dity. The Relative Growth Rate of X. parietina was found to be greater in smal-

ler thalli than in larger ones (Fig. 2).

Source - MNHN. Paris

173

GROWTH RATES ОЕ XANTHORIA PARIETINA

Month Measurement

date

NOV 1976 26/11

DEC 23/12

JAN 1977 8/2

FEB 2/3

MAR 23/3

APR 22/4

MAY 25/5

JUN 22/6

JUL 26/7

AUG

SEP 28/9

ocr 10/11

Nov 2/12

DEC 23/12

JAN 1978 24/1

FEB 3/3

MAR 23/3

JUN 16/6

TOTAL GROWTH

325

355

363

387

405

418

455

593

665

705

747

860

892

943

Growth

(mm?)

37*

138

113

51%

702

A/2

Thallus

Growth

(am?)

4/3

Thallus

area

(шш?)

моо ч

Growth

(am?)

(5)

A/4

Thallus

area

(шш?)

366

403

411

459

484

456

454

486

478

495

586

561

566

595

633

623

Growth

(m?)

]

17*

-10

257

А/5

Thallus

area

93.0

102.5

112.0

134.2

165.5

174.5

201.0

172.0

187.0

204.5

218.0

9.5

0.0

22.2

31.3

9.0

26.5

29.0*

15.0

ins

13.5

125

Tab. 1. — Growth measurements of Xanthoria parietina obtained by measuring the area with a planimeter. * represents damage

to the thallus. Only positive growth increments have been used for plotting points in Fig. 2.

Source : MNHN. Paris:

174 T.H. MOXHAM

Source : MNHN, Paris

GROWTH RATES OF XANTHORIA PARIETINA 175

Fig. 1b (cont.)

Fig. 1. Sequential photographs of Xanthoria parietina showing monthly increments of

growth

Source - MNHN. Paris

176 T.H. MOXHAM

Rydzak 1

Rydzak 5

x Ayazak 6 aa

Mohan aa

14 `

da Beschel

Lotsy

In A.

Fig. 2. — The Relative Growth Rates of Xanthoria parietina plotted against the initial thallus

arca, From RYDZAK (1966) 1 —г —0.81, p => 0.001, 5 ~r =0.43, 0.02 >p >0.01,

6—r— 0.74, p=>0.001, BESCHEL (1958) r=0.74, 0.2>p>0.1 and LOTSY

(1890) r—0.72, 0.02>p>0.01. The Moxham results are obtained from Tab. 1,

r=0.74,p=>0.001.

DISCUSSION

‘There are numerous and often confusing ways of expressing growth rates of

lichens because of the different methods of measurement and the different

approaches to analysing the results; the relative merits of each are discussed

by TOPHAM (1977). In order, therefore, to compare the techniques of others,

the photographs of this study were measured in different ways.

1. The measurement of single lobes growing towards a fixed point. (This method

was discarded as it does not take into account the whole thallus size, and

cannot be used, therefore, to calculate the Relative Growth Rate — see

below).

2. The mean of several diameters.

3. The area.

The results in Tab. 1 show the growth of the thalli measured. Tab. 2 shows

the diameter calculated from the area, and the mean diameter calculated from

Source : MNHN, Paris

GROWTH RATES OF XANTHORIA PARIETINA 177

several diameters and how they are comparable over a period of a year. The

greatest monthly discrepancies arose from thalli that were not strictly circular,

i.e. those that had indentations in the thallus (e.g. A/1 — see Fig. 1). The diffe-

rences which occur in the monthly measurements became, over a greater period

of time, less and less so that by the end of the investigation (or a period of a

year) the two methods produced similar results (Tab, 2). This suggests that it

is possible to take the results of others, where measurements have been made

by a variety of methods, to calculate the areas of the lichens, from data presen-

ted in diameters or radii.

Date af 4/2 1/3 4/4 Date 45

Toy ‘76 241 (243) 17 (23) 6 (1) 366 (374) Apr."

Hov."11 665 (107) 36 (40) т(ю) 561 (566) Ap. T] 93 (94)

Jun.'78 — 943*(1005) 43 (41) 6*(6) 623 (629) Har.'78 204 (195)

Growth 702 (762) 26 (24) om 257 (255) 111 (101)

Tab. 2. — Comparison of the two methods of measuring the diameters of Xanthoria parie-

tina. The figure in the column is the area measurement by planimetry, and the figure in

brackets is the area calculated from the measurement of several (5) diameters of the

thallus.

In order to analyse the results, the method of WOOLHOUSE (1968) was used

as it expresses growth rate as a function of initial thallus area, and demonstrates

the productivity of the thallus throughout its growth phases.

log Ar log Ai ^02

mm ? mm ? yr.

c) Relative Growth Rate (R.G.R.) —

ieu tu

Although there has been some criticism of this formula on the grounds that

the whole thallus does not contribute to lichen growth, it has been used here

because it enables a good comparison to be made between similarly sized thalli.

The recent models of APLIN & HILL (1979) and CHILDRESS & KELLER

(1980) were not used, firstly because of the effects of «averaging out» environ-

mental factors when one is seeking to hightlight particular factors which affect

growth, and secondly, because the models, based on the movement of carbo-

hydrate evenly towards the outer edge of the thallus, does not accomodate the

presence of lobes in foliose lichens, e.g. Xanthoria parietina.

Fig.2 shows the results of recalculating the data of LOTSY (1890), BES-

СНЕГ (1958), and RYDZAK (1966) and comparing it with the data from

Tab. 1. A simple linear regression line has been drawn through the points, and

demonstrates the differences in growth rates between the samples. The relati-

vely small samples available (see Tab.3) did not allow thorough statistical

analysis of the data and so it is impossible to draw precise conclusions from

Source : MNHN. Paris

178 T.H. MOXHAM

these results although some reasonable assumptions can be made based on the

data.

It is also impossible, because of the limited data available, to be able to

state quantitatively any of the ecological or environmental parameters (except

rainfall) that affected the lichens measured, and outside the scope of this study

to examine the chemistry, pH, nutrient potential or water retention capacities

of the substrates concerned. However, rainfall has been shown (FISCHER

& PROCTOR 1978, KARENLAMPI 1971) to influence lichen growth more

than any other single climatic factor, and the rainfalls during the growth periods

of the lichens in Fig. 2 differed greatly from one another. Average monthly

rainfalls during the periods of growth = Moxham 90.2 mm, Beschel 81.6 mm,

and Rydzak 57,8 mm. If rainfall alone accounted for differences in growth rates

in Fig, 2, the Rydzak 1 slope would be expected to lie with the Rydzak 5 & 6

slopes and all growth would be directly related to rainfall, which is not the case.

But if the substrate їз taken into consideration, it is noted that of the three

highest growth rate slopes, two are from lichens on saxicolous substrates and

one is from lichens on smooth barked Betula, whereas the three lower growth

rate slopes are derived from lichens growing on coarser-barked aspen and pine-

wood. This trend is further reflected in Tab. 3 where saxicolous lichens and

smooth-barked corticolous lichens (Meyer, Beschel, Degelius, Rydzak 1, 2 & 3

and Moxham) tend to grow faster than about 2.5 mm ‘diam yr., whereas

rough-barked lichens (Lotsy, Hakulinen and Rydzak 5 & 6) tend to grow slower

than about 2.5 mm ‘diam ‘yr. The single exception to this trend is the result

of HOOKER & BROWN, where the period of growth (8 months) on which the

yearly average was calculated, excludes the 4 months autumn and early winter

when growth in England is usually at a maximum (cf. Tab. 1).

X. parietina is а ubiquitous lichen found growing on a wide range of substra-

tes and previous investigations have highlighted differences in lichens growing

on different substrates. HILL and WOOLHOUSE (1966), for instance, in their

autecological studies on X. parietina growing on rocks, roofs and trees, found

significant differences in thallus thickness, chlorophyll content and algal num-

bers between the lichens on different substrates. The differences were mainly

attributed to light intensity and nutrient supply, although these relationships

are far more complicated, as demonstrated by HARRIS (1972) in his model

simulating net assimilation and lichen productivity of corticolous lichens. Others

(e.g. SHOWMAN 1976) have found no correlation between substrate and lichen

growth rates.

The effect of the texture of a substrate in a lichen thallus is due to many

physical and chemical properties of the individual substrate, such as its capacity

to retain water and provide nutrient as well as the pH and chemical composition

of that substrate. Much attention has been paid to the relationships between

substrate and lichens (BRIGHTMAN & SEAWARD 1977) but many more con-

trolled investigations involving far larger samples and many more known ecologi-

cal and environmental parameters are needed in this area. However, the texture

of the substrate is undoubtedly one of the major factors influencing lichen

growth.

Source : ММНМ Paris

GROWTH RATES OF XANTHORIA PARIETINA 179

‘AUTHOR DATE SUBSTRATE COUNTRY NO. OF

SAMPLES

Meyer 1825 Pine, poplar, aspen Germany 2.65 26 9

lotsy 1890 Mainly pinewood Netherlands 0.93 16 9

Beschel 1958 Grave stones Austria 5.00 1 в

Degelius 1964 Ash Scandinavia 5.00 3 т

Hakulinen 1966 Aspen Scandinavia 2.15 2 1

Rydzak 1 1966 Silver birch Poland 3.93 6 8

ha 1966 Paga) " 2.77 6 3

eS 1966 Y m S 2.55 в 2

"= 1966 Aspen s 2.45 6 19

e 1966 = E 1.31 6 u

Hooker & Brom 1977 Slate England 0.52 В mths. 6 lobes

aliss

Noxham 1980 Concrete pavers E 5.54 1 5

Tab. 3. — Average growth rates of Xanthoria parietina (mnT1diamrlyr.) obtained from

published results of the authors in column 1. The calculation of Meyer's data is based

on the assumption that the «Paris Linien» (’”), when translated from the German, is the

equivalent of the «Paris ligne», a unit of measurement used prior to the French Revolu-

tion. The metric equivalent is 1™ = 2.256 mm.

Erratum : Average growth (mn diam! yr) Beschel (1958) =4.71, not 5.00.

CONCLUSIONS

Tt is possible, using data derived from different methods of measurement,

to demonstrate different trends in foliose lichen growth. From the data avai-

lable, the growth rate of Xanthoria parietina is shown to be influenced by the

texture of the substrate; saxicolous and smooth-barked corticolous lichens

tended to have a higher growth rate than rough-barked corticolous lichens.

ACKNOWLEDGEMENTS. — Grateful acknowledgement is made to the World Wildlife

Fund for some financial assistance made available through the British Lichen Society,

to Т. Fletcher of the Division of Mechanical and Optical Metrology, National Physics

Laboratory, for his information on the «Paris Linien», and to Dr. Р.Н. Brown for his help-

ful comments and criticisms.

Source : MNHN. Paris

180 T.H. MOXHAM

REFERENCES

АРЫМ PS. and HILL D.

Biol. 78 :347-363.

ARMSTRONG R.A., 1975 — Studies on the Growth Rates of Lichens, pp. 309-322. In «Li-

chenology : Progress and Problems». Eds. D.H. BROWN, D.L. HAWKSWORTH and RH.

BAILEY, Academic Press, London and New York.

BESCHEL ВЕ. 1958 — Flechtenvereine der Städte, Stadtflechten und ihr Wachstum. Ber.

Naturwiss.-Med. Vereins Innsbruck 52 : 1-158.

BRIGHTMAN ЕН. and SEAWARD M.R.D., 1977 — Lichens of Man-made Substrates,

рр. 253-293. In «Lichen Ecology». Ed. M.R.D. SEAWARD, Academic Press, London,

New York and San Francisco.

CHILDRESS $. and KELLER J.B., 1980 — Lichen Growth. J. Theor. Biol, 82 : 157-165.

DEGELIUS G., 1964 — Biological Studies on the Epiphytic Vegetation of Fraxinus excel-

sior. Acta Horti Gothob. 27 :11-55.

FISHER P.J. and PROCTOR M.C.F., 1978 — Observations on a season's growth in Parmelia

caperata and Р. sulcata in South Devon. Lichenologist 10 (1) : 81-89.

HAKULINEN R., 1966 — Über die Wachstumsgeschwindigkeit einiger Laubflechten. Ann.

Bot. Fenn. 3 : 167-179.

HALE ME. 1973 — Growth in «The Lichens», pp. 473-492. Eds. V. AHMADJIAN and

M.E. HALE, Academic Press, New York.

HARRIS СР. 1972 — The Ecology of corticolous lichens. Ш. A simulation model of pro-

ductivity as a function of light intensity and water availability. J. Ecol. 60 : 19-40.

HILL DJ. and WOOLHOUSE НМ. 1966 — Aspects of the Autecology of Xanthoria

parietina agg. Lichenologist 3 (2) : 207-214.

HOOKER TN, and BROWN РН. 1977 — A photographic method for accurately measu-

ring the growth of crustose and foliose saxicolous lichens. Lichenologist 9 (1) : 65-75.

KARENLAMPI L., 1971 — Studies on the relative growth rate of some fruticose lichens.

Rep. Kevo Subarctic Res. Sta. 7 : 33-39.

LOTSY J.P., 1890 — Beiträge zur Biologie der Flechtenflora des Hainbergs bei Göttingen.

Dissertation Georg-Augusts-Universität zu Göttingen.

MEYER СЕМ. 1825 Nebenstunden meiner Beschäftigungen im Gebiete der Pflanzen-

kunde : 29-41. Göttingen.

RYDZAK J., 1966 — Investigations on the Growth Rate of Lichens. II. Ann. Univ. Mariae

Curie-Sklodowska, Sect. C, Biol. 21 (13) :167-182. (In Polish),

SHOWMAN ВЕ. 1976 — Seasonal Growth of Parmelia caperata. Bryologist 79 (3) : 360-

363.

TOPHAM P.B., 1977 — Colonization, Growth, Succession and Competition, рр. 31-68.

In «Lichen Ecology». Ed. M.R.D. SEAWARD, Academic Press, London, New York and

San Francisco.

WOOLHOUSE H.W., 1968 — The measurement of growth rates in lichens. Lichenologist 4 :

32-33.

1979 — Growth Analysis of Circular Lichen Thalli. J. Theor.

Source : MNHN, Paris

181

VEGETAZIONE MUSCINALE DELLA FAGGETA

DEL BOSCO DI MALABOTTA (PELORITANI)

M. PRIVITERA е В. LO GIUDICE*

SUMMARY. — The authors present a study on the moss vegetation of the Malabotta beech-

woods on the Peloritani. After a brief survey on the environment, a study of moss communi-

ties is carried out. А moss community dominated by Thamnobryum alopecurum has been

recognized on moist and shady rocks, another one, which is referred to Oxyrrhynchietum

rusciformis Gams 1927, on periodically flooded rocks, Grimmia pulvinata and Tortula

muralis у. Hibschmann 1950 moss community appears on dry rocks. On soil, only frag-

mentary examples of vegetation have been recognized because of on important litter of

dead leaves. Finally, on trunks, two types of vegetation have been pointed out : one which

prefers the median and high parts of the barks (Orthotrichetum Iyellii Lecointe 1975)

and another which extends up from the tree basis (Isothecium myurum and Homalothecium

sericeum moss community).

In conclusion, some informations on the biogeography, soil affinities and altitudinal

types of species are given.

Nel quadro delle ricerche già intraprese sulle Briofite della Sicilia abbiamo

voluto fermare la nostra attenzione sulla flora e vegetazione muscinale della

faggeta di Malabotta, sita sui Peloritani occidentali.

In questo territorio limitato, ma interessante, dell’Isola, già oggetto di uno

studio fitosociologico sulla vegetazione superiore (RONSISVALLE & SIGNO-

RELLO 1977), lo strato muscinale vegeta lussureggiante insediandosi su rocce,

terreno scoperto, tronchi e rami delle essenze forestali.

La scarsa letteratura sulle Briofite della Sicilia, che in nessun modo riesce a

dare un quadro completo nè sulla flora nè tantomeno sulla vegetazione musci-

nale, unitamente all’interesse suscitato dal territorio stesso per l'abbondanza

e rigogliosità della brioflora, ci ha indotto ad intraprendere tale studio allo scopo

di apportare un modesto contributo a questo settore di ricerca che ancor oggi

è scarsamente studiato.

* Istituto di Botanica dell’Università di Catania, via A. Longo 19 (Italia).

Crvptogamie, Bryol. Lichénol., 1981, 2, 2 : 181-200.

Source : MNHN. Paris

182 M. PRIVITERA е В. LO GIUDICE

CENNI SULL’AMBIENTE

Il territorio oggetto della nostra ricerca (fig. 1) è localizzato sui Peloritani

occidentali nella fascia altitudinale compresa tra 900 e 1300 m ca. In partico-

lare la faggeta, costituita da aree discontinue, è compresa tra Pizzo Palo, M. Cer-

reto e Punta Tre Carrini. La zona è interessata da modesti corsi d’acqua a carat-

tere torrentizio come ad esempio il torrente Pistone e il torrente Licopeti.

Dal punto di vista geologico nella Sicilia Nord-orientale sono stati indivi-

duati (OGNIBEN 1960) vari «complessi» di terreni, costituiti da varie succes-

Fig. 1. — Schizzo topografico del territorio studiato (scala 1:25 000). L'area punteggiata

indica l'estensione della faggeta pura, le aree tratteggiate la faggeta diradata (da RONSIS-

VALLE С.А, e SIGNORELLO P., 1977 modif.).

Source : MNHN, Paris

MUSCINEE DELLA FAGGETA DI MALABOTTA 183

sioni sedimentarie suddivise in una o piü falde di trasporto o da successsioni

molto affini appartenenti alla stessa falda о a più falde contigue.

Secondo l’autore sopra citato il territorio da noi esaminato appartiene al

«Complesso Calabride», costituito da varie falde di ricoprimento a basamento

metamorfico con interposti livelli mesozoici prevalentemente calcarei. Il «Flysch

di Capo d’ Orlando», che affiora in tutta l’area da noi esaminata, rappresenta

un sedimento detritico di età oligocene superiore-miocene inferiore, costituito

alla base da potenti livelli conglomeratici e verso l’alto da alternanze ar;

arenacee. I sedimenti prevalenti sono di tipo subarcosico cioé contenenti dal

75% all’85% di quarzo, buone percentuali di feldspati alcalini sodico-potassici

e scarsi frammenti litici e/o miche. Si tratta di un sedimento marino tipicamente

sin-orogeno.

Per quanto riguarda la nostra indagine, i rilievi floristici sono stati effettuati

tutti in zone interessate dai terreni di cui sopra, che da quanto esposto risultano

ricchi in SiO).

Per ció che concerne il clima nel territorio studiato non esistono stazioni

meteorologiche; si fa riferimento pertanto ai dati climatici delle vicine stazioni

TUO) FLORESTA (1250 m) 9,6° 1390 Р (тт)

(10)

220

Go 4 маме са шо

Fig. 2. — Diagramma ombrotermico di Floresta (1250 m). Le osservazioni si riferiscono al

periodo 1963-1973.

Source : MNHN. Paris

184 М. PRIVITERA е В. LO GIUDICE

ТСС) | ROCCELLA VD. (655m) Р (тт)

] L140

104

[Es M

БОЕМ, А RES 4 Ао

Fig. 3. — Diagramma pluviometrico di Roccella Valdemone (655 m). Le osservazioni si

riferiscono al periodo 1959-1970.

di «Floresta» e «Roccella Valdemone» (fig. 2 e 3). Di quest'ultima stazione

disponiamo solo dei dati relativi alle precipitazioni (1959-1970), che sono mag-

giormente concentrate nei mesi invernali e nei primi mesi primaverili.

Per la stazione di Floresta (1250 m di altitudine), sulla base di un decennio

di osservazioni (1963-1973), si è rilevato che la temperatura ha un valore medio

annuo di 9,6°C: in particolare le temperature medie massime mensili si riscon-

trano in luglio con valori di 19,1°C, mentre le temperature medie minime si

hanno in gennaio con —1,0°С raggiungendo valori estremi di -15,3°C.

Le precipitazioni medie annue (1300-1400 mm) sono più abbondanti in

autunno e їп inverno, diminuiscono nei mesi primaverili ed estivi.

Dai valori di temperatura e piovosità, riportati nella fig. 2, si rileva che il

clima nella stazione di Floresta, secondo la tipologia proposta da WALTER

& LIETH (1960), rientra nel tipo mediterraneo montano.

Vegetazione fanerogamica

Il territorio in esame è caratterizzato da una vegetazione forestale in cui

domina il faggio! che assume per le favorevoli condizioni climatiche, in parti-

colare l'umidità, dimensioni considerevoli e aspetto rigoglioso. Oltre al faggio,

che è la specie arborea più comune, compaiono sporadicamente Quercus pu-

bescens, О. cerris, Acer campestre, A. obtusatum.

1. Le stazioni di faggio sui Peloritani sono stati per la prima volta segnalati da GRAMUGLIO

nel 1968.

Source : MNHN. Paris

MUSCINEE DELLA FAGGETA DI MALABOTTA 185

Il sottobosco è discretamente popolato da essenze arbustive ed erbacee.

Fra le prime particolare significato assume Ilex aquifolium, che spesso ai margi-

ni della faggeta, si addensa in fitti popolamenti. Fra le essenze a portamento

arbustivo, oltre all'Ilex aquifolium ricordiamo Crataegus monogyna, Clematis

vitalba, Rubus glandulosus, Cytisus villosus.

Tra le essenze erbacee più frequenti ed abbondanti oltre che significative

citiamo Lathyrus venetus, Euphorbia amygdaloides, Daphne laureola, Мейса

uniflora, Allium pendulinum, Geranium versicolor, oltre che Lamium flexuosum,

Cyclamen hederifolium, Ranunculus lanuginosus, Doronicum orientale, ecc.

La vegetazione di cui sopra si può riferire, come già precisato da RONSIS-

VALLE e SIGNORELLO (l. c.) all’Aquifolio-Fagetum, associazione già descritta

da GENTILE (1969) per l'Appennino meridionale e per la Sicilia, e appartenente

al Geranio-Fagion Gentile 1969, alleanza a carattere spiccatamente meridionale.

LA VEGETAZIONE MUSCINALE

Lo studio della vegetazione è stato effettuato secondo il metodo fitosociolo-

gico della scuola sigmatista di Zurigo-Montpellier; secondo questa metodologia

i due coefficienti attribuiti a ciascuna specie esprimono rispettivamente la

«dominanza» e la «sociabilità » della specie stessa.

La tipologia effettuata, anche se non sempre ha consentito un inquadramento

fitosociologico, ha dato tuttavia la possibilità di tracciare un quadro espressivo

degli aggruppamenti di Briofite che colonizzano i diversi biotopi.

Viene qui di seguito presentato lo studio degli aggruppamenti distinti secondo

l'habitat in aggruppamenti del suolo, delle rocce, dei tronchi.

La vegetazione delle rocce

Uno degli aspetti più interessanti del bosco è rappresentato dalla vegetazione

muscinale delle rocce che, indisturbata dalla presenza delle piante superiori,

tende alla esclusività formando consorzi più o meno omogenei.

Per lo studio degli aggruppamenti rupicoli sono stati considerati tre ambienti

più significativi : rocce umide ed ombrose, rocce periodicamente inondate,

rocce secche e soleggiate.

La vegetazione muscinale delle rocce umide ed ombrose

Sulle rocce ombrose prospera rigogliosa la florula muscinale che si addensa

in fitti popolamenti ricoprendo talora l’intera superficie rocciosa.

La vegetazione muscinale rinvenuta in questo biotopo è caratterizzata dalla

presenza di Thamnobryum alopecurum che spesso prevale su tutte le altre specie,

caratterizzando fisionomicamente la vegetazione. 1 ciuffi lassi, di un verde-cupo,

di questo grande pleurocarpo, in armonia con la debole luminosità del luogo,

presentano un fusto stolonifero da cui si dipartono dei fusti secondari che hanno

Source : MNHN, Paris

186 M. PRIVITERA е В. LO GIUDICE

il portamento di piccoli arbuscoli di 10-11 cm di altezza.

Alla specie sopra citata si associano altri grandi pleurocarpi fra cui particolare

importanza assumono Plasteurhynchium striatulum, Cirriphyllum crassinervium

e ancora Leucodon sciuroides, Isothecium myurum, Homalothecium sericeum,

Brachythecium rutabulum. Le altre entità rinvenute nel nostro aggruppamento

come Fissidens cristatus, Brachythecium velutinum, Plagiothecium nemorale,

ecc. hanno fisionomicamente un ruolo accessorio.

Gli esempi più significativi della vegetazione in esame sono stati riuniti nella

tab. 1 che si compone di 6 rilevamenti effettuati su superfici esposte per lo

più a N-NW.

Dell'esame della tabella sembra difficile accostare la vegetazione rilevata ad

unità fitosociologiche già note sia per la carenza di letteratura riguardante la

briovegetazione del territorio italiano, sia per l'insufficienza di dati a disposi.

zione. Tuttavia essa può essere differenziata da Thamnobryum alopecurum, qui

particolarmente abbondante. Un certo significato nell'aggruppamento studiato

assume anche la presenza di Plasteurhynchium striatulum, Cirriphyllum crassi-

nervium, Isothecium myurum, Homalothecium philippeanum che sembrano

trovare Poptimum climatico ed altitudinale nel dominio dei Querco-Fagetea.

La vegetazione muscinale delle rocce periodicamente inondate

Lungo il fondo valle, sulle rocce e sui grossi blocchi che ingombrano il letto

del torrente Licopeti, si stabilisce una vegetazione di Briofite a carattere igro-

filo, Le specie rinvenute sono poche trattandosi di un ambiente alquanto spe-

cializzato. La vegetazione che vi si insedia è dominata fisionomicamente da

Platyhypnidium riparioides, muschio acquatico, che riesce talora a formare

aggruppamenti monofitici. Ad esso si accompagnano Leptodictyum riparium,

anch'esso legato all'ambiente acquatico; Thamnobryum alopecurum, Oxyrrhyn-

chium praelongum var. praelongum, Plagiochila asplenioides, ecc. maggiormente

diffusi ai bordi del torrente.

In tali stazioni, che riteniamo di un certo significato, abbiamo eseguito, in

periodi particolarmente favorevoli, i rilevamenti riportati nella tab. 2. Da essa

si rileva che la vegetazione indagata è da riferire all’Oxyrrhynchietum rusci-

formis Gams 1927, associazione appartenente ai Fontinaletalia antipyreticae

у. Hübschmann 1957.

Oltre a Platyhypnidium riparioides, caratteristica dell'associazione che è sem-

pre ed abbondantemente presente sia immerso, sia lambito o inondato dalle acque,

ricordiamo Amblystegium varium e Leptodictyum riparium, caratteristiche dei

Fontinaletalia antipyreticae. All'interno dell’aggruppamento è stata individuata

la subassociazione a Thamnobryum alopecurum già descritta da v. HÜBSCH-

MANN 1967, che nel nostro territorio sembra differenziare un aspetto di vegeta-

zione meno legato all'ambiente acquatico preferendo piuttosto i substrati umidi.

L'associazione di cui sopra interessa l'Europa dalla Scandinavia sino ai Pirenei

e frammentariamente il Mediterraneo e le Canarie. Questo nostro rinvenimento

Source - ММНМ. Paris

ор

“штипэәй,

124;

шпбидоишоці, v orusweddnssße : эвозашо po әргшп 99901 apap ouo;

Numero dei rilievi

Superficie (n*e om?)

Esposizione

Inclinazione (°)

Altitudine (m)

Ricoprinento muscinale (4)

Nunero delle specie

Thamnobryum alopecurun

Altre specie +

~ Specie sassicole soiafile

Plasteurbynohiun etriatulun

Cirriphyllum craseinervium

Fissidens cristatus

Orimmia ap.

7 Specie cortico-sensicole

Leucodon aciuroides

Isothectus nyurun

Honalotheoiun sericeum

Isotheciun myurun fo.robustum

Honalothectun philippeanun

~ Specie terricole sassicole

Brachythecium rutabulun

Plagicohila asplenioide

Braohytheoiun velutinum

Plagiotheoiun nemorale

- Specie che vivono su terrene,recce, trenohi

Hyjnum cupressiforne

Sobistidium apocarpun

Hygus cupressiforme var. unoinulatua

1 2 3 4 5 6

1. — $000 1. 2500 2500 3500

x x жом x a

50 30 40 20 во та

950 1150 1000 1200 1050 1150

100 80 100 б во 100

10 8 8 8 6 4

N E їз ки Жы

1.4 : 3 4 м +

14 24 14 я 4

+ + > 5 un mem

À 14 DEA 14 : P

+ : 1.3 5 1.4 $

1 + 4 1-4 :

2.3 + 1.4 a 1.3

+ s 1.4 : : i

Я + 2 л 1.4 $

о.

ur.

Baran.

po altitudinale

Plan.

от.

Ind.

Ina.

Ind.

таа.

Ind.

or.

Ind.

Ina.

Ind.

Ina.

Ind.

Ina.

Ind

м.

Ina,

таа.

VLLOIVIWVW Id VLH99VA ултна FINIOSNW

181

Source : MNHN. Paris

liunero dei rilievi

Superficie (сп?)

Esposizione

Inclinazione (°)

Atitudine (m)

Ricoprimento muscinale (7)

Numero delle specie

Caratt. di associazione

Platyhypnidium riparoides

Diff. della subassociazione

Thamobryum alopecurum

Caratt. di unità superiore

Anblystegium variun

Leptodiotyun ripari

Altre spe

Oxyrrhynchium praelongum var. praelongun

Plagiochila asplenioides

Brachytheoium rutabulun

Rhynchostegiun confertum

Tab. 2. — Vegetazione delle rocce periodicamente inondate

1 3 4

2000 1500 1500

N NE N

60 50 40

950 910 900 2

85 15 10 3

6 3 3 8

3.5 4.4 4.4 01.

2.4 . Bur.

+ 1.2 + 01.

. * * Cosm.

1.4 . . 01.

1.4 . 01.

. E . Cosn.

+ . Buras.

: Oxyrrhynchietum rusciformis Gams 1927.

Tipo altitudinale

Ind.

Plan.

Ind.

Plan.

Ind.

Plan.

edafiche

Aff:

Ind.

м.

Ind.

881

ялало OT 'Y ? VILLIATIA ‘N

Source : MNHN. Paris

MUSCINEE DELLA FAGGETA DIMALABOTTA 189

ci sembra di un certo significato confermando la presenza dell’associazione fino

alle nostre latitudini.

La vegetazione muscinale delle rocce secche

L'ambiente umido e fresco della faggeta condiziona lo sviluppo di una florula

crittogamica xerofila che riesce ad insediarsi solo su rocce secche e soleggiate,

poco diffuse nel terrotorio in esame. Queste rocce, site ai margini o nelle schia-

rite del bosco, accolgono una vegetazione muscinale costituita da specie preva-

lentemente acrocarpe, eliofile, formanti dei pulvini sparsi su substrato che

spesso non è la roccia viva ma che trattiene una piccola quantità di terriccio.

Allo scopo di caratterizzare questo tipo di vegetazione abbiamo eseguito nel

corso dei nostri sopralluoghi quattro rilevamenti fitosociologici riportati nella

tab. 3.

La vagetazione rupicola in esame sarebbe da riferire all’associazione a Grimmia

pulvinata e Tortula muralis v. Hibschmann 1950, dei Grimmietalia.

mere del rilievi 1 Sure: 4

Superficie (on?) 2500 2500 2500 2500

Jeposisione Som m de à 3

Inclinazione (°) 50 35 20 40 Е

stendine (n) 950 1050 1150 1050 3

Rooprinente muscinale (f) ا‎ д E

Fumero delle specie 1 1 6 4 E

Sani delltassooiasime

orienta pulvinata d ed e rE me | me

Tortula moralis 2 aie : NET

Carati, di unità superiore

башыны» pia а а" mà | me

беш trionopayila È Sdi ; Bee | зы

ice mete ,

Puebla um

Rhynchestegiella tenella fo.neridionalis + . . . Med. Plan. ба.

Tortella tertuoca fo, fregilifelia E S Е о Da | жо

- Busse terzicola sessioni

беззен Gozo AE uM dine

frishorimus crispulum і : ЖР a rim: | 9

аа mine ын»

Grthetrioh эр. ; : p чё А " 5

Heselotheous seriou +: x Ў poe КГ. Ж

= эша ole vivene si terres, recco. Ызы

Tortula талаца nes ч com | m | me

Wun oupressiforne yar-unoimalatun гм i $ m | ы | ы

Bs овраге E u : р ы |ы.

Nus Oupmenniferne fe. teoterun i Sun кау ў tom |ы | m

Tab. 3. — Vegetazione delle rocce secche : ass. а Grimmia pulvinata-Tortula muralis v.

Hübschmann 1950.

Source : MNHN, Paris

190 M. PRIVITERA e В. LO GIUDICE

Fra le specie caratteristiche dell’associazione si notano Grimmia pulvinata e

Tortula muralis e fra le specie dell'ordine (Grimmietalia) Schistidium apocarpum

e Grimmia trichophylla. Ad esse si accompagnano Ceratodon purpureus, Tortula

ruralis e le pleurocarpe Homalothecium sericeum, Rhynchostegiella tenella fo.

meridionalis, Hypnum cupressiforme var. uncinulatum, ecc.

La vegetazione muscinale del suolo

Sul terreno le Briofite sono scarsamente rappresentate per la spessa lettiera

delle foglie di faggio che occupa la gran parte del suolo (copertura massima 90%,

copertura minima 60%) e più localmente per lo sviluppo dello strato erbaceo.

La povertà del ricoprimento muscinale ci ha indotti a scegliere per i rilevamenti

briosociologici superfici molto limitate ricercate nelle piccole schiarite, nei

pendii più inclinati, ai piedi degli alberi, alla base dei massi. 1 dati relativi alla

vegetazione del suolo constano di 7 rilevamenti riuniti nella tab. 4. Dall’ esame

di essa si nota che trattasi più che altro di esempi frammentari di vegetazione

rappresentati per lo più da entità di vasta ampiezza ecologica, ne risulta pertanto

жөө» des rilievi ft Me MU AN Pt 3

Баана (om?) axo тую 200 100 soa 100 м®

Ташын» (+) nm - жш m m È

лання (a) Wo» riso шю sco too 950 мю à 3

Risepeinente модные (f) a € D D » » 4% i 3

Banane dallo speste ТАЕП и Tre в 3 4

pecie del ceriegzie tion

октав sonetos (orga ЫЗЫ. жору no |- | м

Оооо угы төш vas. praetonem | = 2 i ө [ru | me

орали presiengos Var. state exo ES moe. | ыы | mo

Brest eta mutus oe АӨ? а Oe

Moles estratos E n ARCAM À

Magie sepienteite балл

am ou dE NE о? |

soie ohe sirme mu terrano ros, trat

тшен мамыны,

Aud mez Lr a ETE

Bern capitare a Ie

Arpa сами Wee NL ا‎ Е E

Magnesia street JEN RUE ARE ү Je po

ee pen x E OSS

Tab. 4. — Vegetazione del suolo.

Source : MNHN, Paris

MUSCINEE DELLA FAGGETA DI MALABOTTA 191

difficile la tipizzazione. E' quanto già rilevato da altri autori a proposito della

vegetazione muscinale dell’habitat preso qui in considerazione.

Dal punto di vista floristico, fra le specie del «corteggio silvatico», segnaliamo

per la discreta presenza le pleurocarpe Oxyrrhynchium schleicheri, Brachythe-

cium rutabulum, Oxyrrhynchium praelongum var. praelongum, muschi forestali

che prediligono il terreno umido, ricco di humus.

Fra le specie rinvenute particolare significato assumono nell'ambiente studia-

to Brachythecium rutabulum, Plagiochila asplenioides, Brachythecium veluti-

num, le quali, benchè non siano legate all'ambiente delle foreste, sembrano

qui trovare l'optimum climatico; per tale motivo vengono considerate caratte-

ristiche preferenziali dei Querco-Fagetea (GAMISANS & HEBRARD 1979), ecc.

La vegetazione muscinale epifita

Ben rappresentati sono i muschi epifiti nel territorio in oggetto che ora

ricoprono sparsamente o a chiazze le cortecce, ora formano lussureggianti

tappeti che rivestono ampie superfici sino ad altezze considerevoli, se l’esposi-

zione è favorevole. Per quanto riguarda la copertura, come è noto da tempo,

il lato Nord della corteccia degli alberi è più densamente popolato, potendo

qui le specie proteggersi da un'insolazione eccessiva. Alcuni muschi corticicoli,

per resistere all’essiccamento, mettono in atto degli adattamenti morfologici che

sono generalmente delle «xeromorfosi»; ne è esempio il Leptodon smithii.

Fra le specie rinvenute alcune sono arboricole (come Orthotrichum schimperi,

O. affine e O. Iyellii), altre sono entità che vivono indifferentemente su rocce

don sciuroides, Leptodon smithii, ecc.) altre ancora sono poco sensibili ai

diversi substrati.

Allo scopo di meglio conoscere la composizione della vegetazione muscinale

di questo ambiente abbiamo effettuato alcuni rilevamenti fitosociologici che

sono stati riuniti nella tab. 5 e nella tab. 6.

Nella tab. 5 vengono presentati i rilievi eseguiti essenzialmente nelle parti

mediane e superiori dei tronchi di faggio, tra 1000 e 1300 m di altitudine e nelle

condizioni di esposizione ottimale.

Dall’esame della tabella si rileva che l'aggruppamento individuato è da riferire

all'Orthotrichetum lyellii Lecointe 1975, associazione già descritta dallo stesso

autore per la «Brenne», territorio situato a Sud della Loira, In questo territorio

POrthotrichetum lyellii è stato rinvenuto su tronchi di Quercus, Carpinus, Fagus,

ecc. negli aggruppamenti del Quercion occidentale Braun-Blanquet 1967 e del

Carpinion betulii Oberdorfer 1943.

Oltre alle caratteristiche dell’associazione, Orthotrichum lyellit e Hypnum

cupressiforme fo. filiforme, ben rappresentate sono le specie dell'ordine (Leuco-

dontetalia) fra le quali citiamo le briocamefite cespitose Orthotrichum affine,

О. schimperi, Leucodon sciuroides. Meno rappresentate sono le briocamefite

Source : MNHN. Paris

+6461 әзщоэәт #1124 шизацизоцзао : ed2 әчоргезәВәд — `S ‘QUI

Nunero dei rilievi 1 2 3 4 t ae

Superficie (n°) 1000 1300 1200 1300 1000 1000

Esposizione н NRO и им 0

Inclinazione (*) 100 80 100 100 100 100

Atitudine (m) 1050 1100 1000 1200 1100 1150

Ricoprimento muscinale (£) Te 8609 п ю юу ею

Numero delle specie 8 6 8 5 6 6

Caratt. ө diff. di assooiagione

Orthotrichun 1961244 au аи o ПО

Hymum cupressiforne fo. filiforne 4H paeem an 5 d

Caratt. e diff. di unità superiore

— Specie arboricole

Orthotrichun affine al E zi fua dg x

Orthotriohum sohimperi , 3 hy Se St me

= Specie cortice-sassicole

Honalotheoiun serioeum = PC Бый. сый + 5

Leptoden smithii б ота À 3

Leucodon soturoides 13 + 5 + ۴

Pterigman&rum filiforme M. 5 x к 5

Badula conplanata SÙ К 5 ; +

Zygodon viridissimus 5 a mm Й Ё

Pterogontun gracile 5 К 3 ^ +

- Specie che vivono au terreno,recce,irenchi

Tortula ruralis + б + 4 s

Tortula subulata + + z 5 > E

Barbula vinealis дор. cylindrica j К + 5 1 ў

Lobaria pulmonaria 3 à

Aum.

bati,

01.

où

Зарвеа.

ЕН:

Sabri

Conn.

situa ins

Ina.

Ind.

Ina.

Ina. ?

Ind.

Ina.

Ind.

Plan.

ma.

Ind.

Ina.?

dar ihe

Ina.

таа.

ma.

таа.

Ina.

таа.

Ina.

Source

TANID ол "м ә учамлгча "W

MNHN, Paris

“wnaouas

: wide әшогтзә8әд — ‘9 qe

штоэщіојршон ә wninku штәәцозү e озпэшедатаа8е

Nunero dei rilievi

Superficie (on?)

Esposizione

Inclinazione (°)

Atitudine (а)

Ricoprimento muscinale (Я)

Hunere delle specie

Isotheciun murum

Homalotheoium sericeun

Caratt. e diff. dei Leucedentetalia

- Specie certico-sassicole

Leuceden sciureides

Leptodon smithii

Pterogenium graoile

Radula conplanata

Pierigmandrun filiforme

Specie arboricole

Hymum cupressiforme fo.filiferne

Orthetrichum lyellii

Altre specie 1

Eymun oupressiferne

Brachythecium rutabulum

Brachytheciun starkei

Cirriphyllun orassinerviun

Leskeella nervo!

1 2 3 4 5 6

2500 2000 2000 1500 1500 2000

NW NE ч ош x x

70 80 100 100 90 100

1100 1050 1200 1150 1100 1000

90. "i95 ду X90: Сш =

1 6 6 6 1 5

3X4 ca E

4:3 E 3-4 23 E

RITO ah Mo +

19302: E È + 4

* - al AR: 2%

+ + z Xie Т. >

* + + + J >

> E = + + ‘

: EE Я ne

È E * E E 1.2

doo

Elenente biogeo;

Сева.

Submed.

subatl-

altitudinale

Ina.

Ind.

Plan.

Ind.

Ina.

Affinità edafiche

Ind.

Ina.

Ind.

Ina.

Ind.

м.

Ina.

VLLOIVIVA Id V.I399V4 VITIO TINIOSNW

£61

Source : MNHN. Paris

194 M. PRIVITERA е R. LO GIUDICE

rampanti ad eccezione di Homalothecium sericeum, specie alquanto diffusa

nell’ambiente corticicolo.

Esempi simili di vegetazione crittogamica sono stati rinvenuti da GAMISANS

& HÉBRARD (1979) nella Grecia Nord-occidentale, su tronchi di Quercus. Gli

stessi autori rilevano inoltre che tale vegetazione è rappresentata in numerosi

ecosistemi forestali della parte settentrionale ed orientale della regione medi-

terranea, dove il corteggio epifita raggiunge il piano montano.

Un altro aspetto individuato su vecchi tronchi di faggio (tab. 6) è rappre-

sentato dall'aggruppamento a «lsothecium myurum ed Homalothecium seri-

ceum», che dalla base dei tronchi si spinge talora sino in alto.

I rilevamenti effettuati su tronchi, non di rado inclinati, rivelano una coper-

tura muscosa assai densa per il prevalere dei pleurocarpi Isothecium myurum ed

Homalothecium sericeum aventi un grado di abbondaza-dominanza particolar-

mente elevato. Numerose sono le specie appartenenti ai Leucodontetalia, alcune

delle quali abbastanza frequenti come Leucodon sciuroides, Leptodon smithii,

Pterogonium gracile, Pterigynandrum filiforme, quest'ultimo ritenuto partico-

larmente abbondante nel dominio del Fagion nell’ Europa meridionale.

La presenza (50% dei rilievi) di Pterogonium gracile e Leptodon smithii

suggerisce un avvicinamento possibile alla suballeanza del Fabronion pusillae

Barkman 1958 dei Leucodontetalia largamente distribuita nell'Europa meridio-

nale. L’aggruppamento in esame nel suo insieme potrebbe avvicinarsi anche

al’Homalothecio-Porelletum platyphyllae Lecointe 1975, riferito anch'esso

ai Leucodontetalia, ma dal quale si discosta per certi caratteri floristici.

CENNI CONCLUSIVI

Lo studio condotto ha consentito di cogliere alcuni tra i più significativi e

caratteristici aspetti della vegetazione crittogamica della faggeta del bosco di

Malabotta, come si può rilevare dalla tab. 7. In particolare nel suolo si riscontra-

no esempi frammentari di vegetazione che non riescono a costituire aggruppa-

menti specializzati a causa dello spesso strato di lettiera che occupa gran parte

del sottobosco, oltre che per la notevole presenza di entità a grande ampiezza

ecologica.

A differenza del suolo, le rocce umide disseminate fra gli alberi accolgono

una lussureggiante vegetazione muscosa, caratterizzata da Thamnobryum alope-

curum che si impone per la sua rigogliosità e vitalità, lasciando poco spazio

ad altri grandi pleurocarpi che, pur tuttavia, riescono ad insediarsi colonizzando

il substrato roccioso.

Sulle rocce periodicamente inondate si stabilisce una vegetazione muscinale

a carattere prevalentemente igrofilo, caratterizzata da Platyhypnidium riparioi-

des, specie idrofila acquatica.

Le rocce secche e scoperte, rare nel territorio in esame, perdono ogni carat-

tere mesofilo nella flora ospitata, rivestendosi di una vegetazione prevalentemen-

Source : MNHN. Paris

AMBIENTE LEUCODONTETALIA FONTINALETALIA ANTIP, GRIMMIETALIA

чу,

terreno esempi frammentari

umide aggr.a Thamnobryum alopecurum

rocce inondate

secche

Oxyrrhynchietum rusciformis

ass.a Grimmia pulvinata

e Tortula muralis

tronchi

Orthotrichetum lyellii '

aggr.a Isothecium myurum

е Homalothecium sericeum

V.LLOSVTVAW Id V.L3899 V3 VITA HANIOSNN

$61

Source : MNHN. Paris

196 M. PRIVITERA е К. LO GIUDICE

te xerofila da riferire all'associazione a Grimmia pulvinata e Tortula muralis

у. Hübschmann 1950.

Sugli alberi, infine, sono stati individuati due aspetti di vegetazione; uno

caratterizzato da specie prevalentemente acrocarpe che prediligono le parti

mediane e più alte della corteccia (Orthotrichetum lyellii Lecointe 1975) l’altro,

rappresentato in massima parte da pleurocarpi che dalla base degli alberi risal-

gono a tappeto sino ad altezze considerevoli (aggruppamento a Isothecium

myurum ed Homalothecium sericeum).

Dal punto di vista biogeografico (fig. 4a) lo strato muscinale dei differenti

biotopi studiati è maggiormente rappresentato dall’elemento oloartico (38%),

segue l’elemento cosmopolita (24%), che assieme all'elemento oloartico raggrup-

pa crittogame a vasta ripartizione geografica, capaci di adattarsi ai climi più vari.

Discretamente rappresentati sono gli elementi europeo (16%) е submediterra-

neo-subatlantico (10%), trascurabili gli altri elementi che complessivamente

rappresentano il 12% delle entità rinvenute.

Per quanto riguarda le affinità altitudinali (fig. 4b) poco abbondanti sono le

specie planiziarie (27%) e le orofite (11%) rispetto alle entità indifferenti all’alti

tudine, che costituiscono il 62% del ricoprimento muscinale.

Anche per ciò che riguarda le affinità edafiche (fig. 4c) si nota una elevata

proporzione di specie indifferenti alla natura del substrato (68%); molto meno

fappresentate sono le specie acidofile (20%), trascurabili infine le calcifile e le

silicicole che insieme con le acidofile e calcicole costituiscono il 12% della

copertura muscosa.

Lo studio condotto sulla faggeta del bosco di Malabotta ha consentito di

conoscere alcuni aspetti della vegetazione muscinale rappresentati nel nostro

territorio anche se limitati alla zona studiata. I risultati a cui siamo pervenuti

riteniamo possano rappresentare un modesto contributo e allo stesso tempo

una premessa che possa costituire da base per ulteriori indagini sul vasto ed

inesauribile mondo crittogamico.

APPENDICE FLORISTICA

POLYTRICHACEAE :Pogonatum aloides (Hedw.) Р. Beauv.

FISSIDENTACEAE : Fissidens minutulus Sull. ssp. pusillus (Wils.) Wijk et

Marg. — F. taxifolius Hedw. ssp. taxifolius — F. cristatus Mitt.

DITRICHACEAE :Pleuridium subulatum (Hedw.) Rabenh. — Ceratodon

purpureus (Hedw.) Brid. ssp. purpureus.

ARCHIDIACEAE : Archidium alternifolium (Hedw.) Мих.

POTTIACEAE : Tortula subulata Hedw. ssp. subulata — T. muralis Hedw. —

T. ruralis (Hedw.) Gaertn, Meyer et Scherb. ssp. ruralis — Barbula vinealis Brid-

ssp. cylindrica (Tayl) Podp. — Trichostomum crispulum Bruch — Tortella

tortuosa (Hedw.) Limpr. fo. fragilifolia (Jur.) Moenk.

Source : MNHN. Paris

MUSCINEE DELLA ЕАССЕТА DI MALABOTTA 197

Europee

Euroamericane

Oloartiche

38%

Med. Atl. e Med

Eurasiatiche

Cosmopolite

Planiziarie

Orofite

Indifferenti

Acidofile

Sili

Acid. е calcic

Calcifile

Indifferenti 68%

Fig. 4. — a) Spettro biogeografico della vegetazione muscinale della faggeta di Malabotta.

b) Tipi altitudinali. c) Affinità edafiche.

Source : MNHN, Paris

198 M. PRIVITERA е R. LO GIUDICE

GRIMMIACEAE : Schistidium apocarpum (Hedw.) B.S.G. — Grimmia pulvi-

nata (Hedw.) Sm. — G. trichophylla Grev. ssp. trichophylla — С. sp.

ORTHOTRICHACEAE : Zygodon viridissimus (Dicks.) Brid. ssp. viridissimus

— Orthotrichum lyellii Hook. et Tayl. — О. affine Brid. — O. schimperi Hammar.

BRYACEAE : Bryum capillare Hedw. ssp. capillare.

MNIACEAE : Mnium rostratum Schrad.

BARTRAMIACEAE : Bartramia stricta Brid. — В. pomiformis Hedw.

LEUCODONTACEAE :Pterogonium gracile (Hedw.) Sm. — Leucodon sciu-

roides (Hedw.) Schwaegr.

LEMBOPHYLLACEAE :Isothecium myurum Brid. I. myurum Brid. fo.

robustum (B.S.G.) C. Jens. — Plasteurhynchium striatulum (Spruce) Fleisch. —

P. striatulum (Spruce) Fleisch. fo. cavernarum (Mol.) Podp.

NECKERACEAE : Leptodon smithii (Hedw.) Web. et Mohr — Thamnobryum

alopecurum (Hedw.) Nieuwl.

LESKEACEAE : Leskeella nervosa (Brid.) Loeske.

AMBLYSTEGIACEAE : Amblystegium varium (Hedw.) Lindb. — Leptodic-

tyum riparium (Hedw.) Warnst. — Platyhypnidium riparioides (Hedw.) Dix.

BRACHYTHECIACEAE : Homalothecium sericeum (Hedw.) В.5.6. — Н.

philippeanum (Spruce) B.S.G. — Brachythecium velutinum (Hedw.) B.S.G. —

В. velutinum (Hedw.) B.S.G. var. salicinum (B.S.G.) Moenk. — B. salebrosum

(Web. et Mohr) B.S.G. — В. starkei (Brid.) B.S.G.—B. rutabulum (Hedw.)

B.S.G. — Rhynchostegium confertum (Dicks.) B.S.G. — Rhynchostegiella tenella

(Dicks.) Limpr. fo. meridionalis (Boul.) Podp. —Cirriphyllum crassinervium

(Tayl) Loeske et Fleisch. — Oxyrrhynchium praelongum (Hedw.) Warnst. var.

praelongum — O. praelongum (Hedw.) Warnst. var. stokesii (Turn.) Podp. — О.

schleicheri (Hedw.) Roell — Eurhynchium striatum (Hedw.) Schimp. var. stria-

tum.

ENTODONTACEAE : Pterigynandrum filiforme Hedw.

PLAGIOTHECIACEAE : Plagiothecium nemorale (Mitt.) Jaeg.

НУРМАСЕАЕ :Нурпит cupressiforme Hedw. ssp. cupressiforme — Н.

cupressiforme Hedw. ssp. cupressiforme fo. tectorum (Brid.) Moenk. — H. cu-

pressiforme Hedw. ssp. cupressiforme var. uncinulatum B.S.G. — H. cupressifor-

те Hedw. ssp. cupressiforme fo. filiforme (Brid.) Krahmer.

RADULACEAE : Radula complanata (L.) Dum.

PLAGIOCHILACEAE : Plagiochila asplenioides (L.) Dum.

LICHENE : Lobaria pulmonaria (L.) Hoffm.

Source : MNHN. Paris

MUSCINEE DELLA ЕАССЕТА DI MALABOTTA 199

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201

EL GENERO SPHAGNUM EN LOS ANDES DE COLOMBIA

Y VENEZUELA

Clave para las especies frecuentes u ocasionales

con notas ecologicas y taxonomicas

Dana GRIFFIN, Ш*

SUMMARY. — An illustrated key is offered to 15 frequent or occasional species of Spha-

gnum known from the Andes of Colombia and Venezuela, Some ecological and taxonomic

notes are appended.

INTRODUCCION

Aunque el género Sphagnum se encuentra hasta en climas cälidos, como la

cuenca del Amazonas у orillas de sus afluentes, alcanza su mayor desarrollo

y diversidad en zonas más frescas. En el norte de Sudamérica esto corresponde

a los bosques andinos y a los páramos. En los páramos más húmedos, Sphagnum

representa uno de los componentes más importantes y más conspicuos de la

flora, formando grandes cojines y a veces llenando los charcos y lagunas con su

proliferación,

A pesar de la contribución a la flora norandina del género Sphagnum, es

muy escasa la literatura que trata las especies más frecuentes de esta región.

Considerando esto, el autor del presente trabajo estimó conveniente organizar

la información registrada en nuestro y otros herbarios y notas del campo, Esta

información se adjunta a la lista de especies que se exponen a continuación

de la clave y comprende notas taxonómicas y ecológicas.

* Department of Botany and Florida State Museum, University of Florida, Gainesville,

Florida, E.U.A. 32611.

Cryptogamie, Bryol. Lichénol., 1981, 2, 2 : 201-211.

Source : ММНМ. Paris

202 D. GRIFFIN

No se pretende abarcar todas las especies conocidas ni citadas de esta region,

La idea principal ha sido incluir las especies frecuentes u ocasionales. La litera-

tura a que se hace referencia incluye las listas de PURSELL (1973) y FLOR-

SCHÙTZ-DE WAARD y FLORSCHUTZ (1979) y el estudio preliminar de

ROBINSON (1967); ademäs, los trabajos más detallados de WARNSTORF

(1911), ANDREWS (1941a, b, 1947 y 1949), BLOMQUIST (1938) y CRUM

(1976). Los trabajos de WARNSTORF, CRUM y BLOMQUIST son de interés

para quien desea empezar el estudio de los Sphagnum porque presentan trata-

mientos generales sobre la morfología y taxonomía del género. El trabajo

de BLOMQUIST (1938) comprende también algo de la metodología básica

para el trabajo con las plantas en el laboratorio,

Conviene señalar que las características de las fibrillas y los poros deben obser-

varse a través de las hialocistes sin seccionar las hojas o, cuando se trata de los

tallos, en sección longitudinal. Cuando esas características no son muy destaca-

das facilita su observación teñir el material primero con unas gotas de violeta

de genciana (solución acuosa). En lo que se refiere a la forma y posición que

ocupan las clorocistes entre las hialocistes, la observación se hace en cortes

transversales de las hojas. El ‘poro anillado” a que se hace referencia en la clave

es el que se presenta circundado por un espesamiento que tiene la forma de

un halo,

CLAVE

1. Células corticales de los tallos y ramas con fibrillas, hojas rameales con ápice

redondeado y + cuculado, márgenes denticulados . ,.............. 2

1. Células corticales de los tallos y ramas sin fibrillas, hojas rameales que se

estrechan apicalmente, con ápice + truncado-dentado, márgenes usualmente

enteros o algo aserrados cerca del ápice 4

2. Clorocistes usualmente alcanzando ambos lados (de la hoja rameal) pero

más expuestas por el lado adaxial, de manera típica isösceles-triangulares,

hojas caulinares liguladas o subespatuladas, + del mismo tamaño que las

hojas rameales ...,..................... EE . 5. palustre

2. Clorocistes completamente incluidas о de + igual exposicién por ambos

и асый A S E O 3

3, Clorocistes elípticas e incluidas, hojas caulinares + uniformes en cuanto a

format liguladaz o suBespatuladasy ne do sles. =e 5. magellanicum

3. Clorocistes angustirectangulares o lenticulares, de + igual exposición, hojas

caulinares que varían en la misma planta, las inferiores anchiovadas, las

superiores ligulado-espatuladas ................... S. erythrocalyx

4. Células corticales de las ramas homogéneas, cada una con un poro en el

polo anterior, clorocistes incluidas, hojas caulinares menos de 1mm de largo,

triángulo-liguladas o trapezoidales, épices chato ......... 5. compactum

4. Células corticales de las ramas heterogéneas, unas con poros y otras sin poros,

Source - MNHN. Paris

CLAVE DEL GENERO SPHAGNUM 203

clorocistes y hojas caulinares varias .. u., poiesis ees 5

. Plantas simples o poco ramificadas, ramas simples o con 2-3 ramas por

fascículo, las hojas caulinares y rameales parecidas en forma ........ 6

. Plantas copiosamente ramificadas, los fascículos con 4-5 ramas cada uno,

hojas caulinares + distintas en forma de las rameales .......... т

. Hojas caulinares mas de 5mm de largo, hialocistes portando 2 о mäs poros

НВ abaxial Ir oS seg ANNE. абме н S. cyclophyllum

. Hojas caulinares hasta 2mm de largo, hialocistes portando de 0-2 poros

opel lado abaxjak Е о an 5. pylaesii

. Clorocistes más expuestas por el lado abaxial o de + igual exposición ... 8

. Clorocistes más expuestas por el lado adaxial (raramente con + igual expo-

E P 8 P

ON) SOON SPEA DRE PRIM. MS AME

. Clorocistes truncado-elípticas o trapezoidales, de igual exposición o algo

más expuestas por el lado abaxial, hialocistes usualmente con numerosos

poros por el lado abaxial, la corteza caulinar de 1 (3) estrato de células

hinchadas, hojas caulinares usualmente menos de 1mm de largo, triangula-

res о elípticas (pero con una forma que puede variar segtin las condiciones

Е III Meer RADAR А S. subsecundum

. Clorocistes triangulares y más expuestas por el lado abaxial, plantas sin

10.

11.

12,

12.

las demás combinaciones de caracteres ..............,‚....... e

. Hojas rameales no marcadamente plano-onduladas cuando secas, clorocistes

trapezoidales o isösceles-triangulares, hojas caulinares triangulares o ob-

longo-triangulares, hasta 1,8mm de largo . . ... S. cuspidatum

Hojas plano-onduladas cuando secas, clorocistes equilátero-triangulares,

hojas caulinares varias Dee EIEL T AAE TEN 10

Corteza caulinar apenas distinta, células corticales pequeñas, de paredes

gruesas, hojas caulinares triangulares o ligulado-triangulares, 1,3-1,5 mm

de largo, hialocistes con 4-7 poros en los ángulos y a lo largo de los lados

(lado adaxial), usualmente difíciles de distinguir sin teñir... . . S. recurvum

Corteza caulinar distinta, las células hinchadas, en 2 estratos, hojas caulina-

res oblongo-ovadas, 1,2-1,5mm de largo, hialocistes con 4-11 poros a lo

largo de los lados (lado adaxial), usualmente fáciles de distinguir .......

RE SS III $. sancto-josephense

Poros de las hojas rameales (lado abaxial) hasta 104m diam., redondeados

y fuertemente anillados, clorocistes a veces con + igual exposición ......

je RE S. sparsum

Poros de las hojas rameales (lado abaxial) usualmente más de 104m diam.,

redondeados o elf pticos, no o muy debilmente anillados ......... 12

Hojas rameales patentes, falcadas, dispuestas claramente en 5 hileras, hojas

caulinares triangulares, hasta 12mm de largo ,........ 5. oxyphyllum

Hojas rameales no dispuestas claramente en 5 hileras, hojas caulinares usual-

mente más de 1,2mm de largo ............... E ORA 13

Source : MNHN, Paris

204 D.GRIFFIN

13. Hialocistes portando 5-12 poros grandes у elipticos a lo largo de los lados

(lado abaxial), clorocistes trapezoidales o triangulares, hojas rameales

gradualmente involuto-acuminadas, hojas caulinares oblongo-ovadas о

triangulo- liguladas, hasta 1,4mm de largo, raramente más S. capillifolium

13. Hialocistes con 0-5 poros por el lado abaxial, clorocistes triangulares, hojas

rameales abruptamente involuto-acuminadas, hojas caulinares triangulares

о ligulado-triangulares, 1,5-1,6 mm de largo ....... lare А, 14

+. S. limbatum

...5. meridense

14. Corteza caulinar porosa . .

14. Corteza caulinar no porosa

NOTAS ECOLOGICAS Y TAXONOMICAS

El género Sphagnum compatte ciertas caracteristicas con otras plantas acuä-

ticas o de sitios muy húmedos, entre ellas tenemos las variaciones en el nivel o perma-

nencia del agua que pueda dar lugar a cambios morfológicos en las plantas.

Como es de esperar eso ha provocado serios trastornos en la taxonomía del

género. Como consecuencia hay muchos sinónimos y hasta hoy en día se discuten

los límites morfológicos de muchas especies.

En las notas que siguen se indica los patrones geográficos y algunos sinónimos

que son aceptados por varios autores modernos, como también, indico la gra-

diente altitudinal de las colecciones nuestras, representativas de las especies

en los Andes de Colombia y Venezuela. Por último, se incluye un cuadro des-

criptivo, muy breve, del nicho ecológico de las especies y láminas con dibujos

de las características más diagnósticas para la determinación de cada una. Las

descripciones ecológicas se basan en los datos que aparecen en las colecciones

realizadas en Venezuela por el autor de este trabajo o por sus colegas Drs.Manuel

López Figueiras y Luis Ruiz-Terán de Mérida y de Colombia por Antoine M.

Cleef (Utrecht), Tomás van der Hammen (Amsterdam) y otros . El Sr. Cleef

está por concluir un trabajo bastante detallado sobre las comunidades de plantas

que habitan los páramos de Colombia. Cuando ese trabajo esté publicado debe

arrojar mucha luz sobre la cuestión del papel de Sphagnum en las partes altas

de los Andes colombianos.

Sphagnum capillifolium (Weiss) Schrank. (Sinon. S. capillaceum (Weiss)

Schrank.). Cosmopolita. 3000-3600 т, Terrestre, en suelos húmedos.

Sphagnum compactum DC. Cosmopolita. 3000-4000m, Terrestre, en panta-

nos o suelos muy húmedos.

Sphagnum cuspidatum Ehrh. ex Hoffm. (Sinon. S. lehmannii Warnst., S.

serratum (Schlieph.) Warnst. hom. illeg., S. trinitense C. Müll.). Cosmopolita.

500-4400m. Es una especie de gran amplitud ecológica, pero, por regla general,

se encuentra en charcos, lagunas o en suelos muy mojados. Cuando crece sumer-

gido puede tener una forma muy delgada con hojas estrechas y aserradas.

Source - MNHN. Paris

CLAVE DEL СЕМЕВО SPHAGNUM 205

Sphagnum cyclopbyllum Sull. et Геза. Holárticos. 3300-3800 m. Conocido

de pantanos y lagunas, frecuentemente sumergido.

Sphagnum erytbrocalyx Hampe. (Sinon. S. perichaetiale Hampe in С. Müll.,

S. peruvianum Mitt.). Neotropical. 700-3500 m, En suelos muy húmedos.

Sphagnum limbatum Mitt. Neotropical. 2300-4000 m. En suelos muy hüme-

dos o pantanosos (tratada por algunos autores como una forma de S. meridense).

Sphagnum magellanicum Brid. (Sinon. $. andinum Hampe, S. medium Limpr.,

5. sanguinale Warnst.). Cosmopolita, 1500-3900 m. En suelos húmedos o hasta

mojados,

Sphagnum meridense (Hampe) C. Müll. (Sinon. S. acutifolium Schrad. var.

meridense Hampe, 5. coryphaeum Warnst., 5. gracilescens Hampe ex С. Miill.,

S. flavicaule Warnst., S. limbatum Mitt.). Neotropical. 1200-3300 m. En suelos

hümedos hasta mojados.

Sphagnum oxyphyllum Warnst. Neotropical. 1200-3300 m. En suelos muy

húmedos.

Sphagnum palustre L. Cosmopolita. 50-2000. En suelos húmedos. Esta

es una especie polimörfica. Dentro del complejo ‘palustre’ hay muchos sinóni-

mos o especies aliadas pero no basadas en material de Colombia o Venezuela.

El trabajo de ANDREWS (1941b) ofrece detalles sobre este polimorfismo.

Sphagnum pylaesii Brid. Holärtico. 3500-4400 т. Normalmente sumergido

en charcos y lagunas.

Sphagnum recurvum P, Beauv, (Sinon. 5. pulchricoma С. Müll.). Cosmopolita,

Alt, 1900-4100 m. En suelos muy húmedos.

Sphagnum sanctojosephense Crum et Crosby. Neotropical. 1900-4100m.

En suelos muy húmedos,

Spbagnum sparsum Hampe (Sinon. 5. acutifolium Schrad.?, S. apollinairei

Par, et Warnst.), Neotropical, 3000-3300m. En suelos húmedos.

Sphagnum subsecundum Nees, Cosmopolita. 2800-3700m. Sumergido о

creciendo en sitios muy hümedos. Especie muy polimörfica con una sinonimia

extensiva pero no sobre material de Colombia о Venezuela.

AGRADECIMIENTOS

El autor manifiesta su agradecimiento a los colegas Isidro Bermüdez у Manuel Маһи

por su critica y ayuda con este trabajo.

Source : ММНМ. Paris

206 D. GRIFFIN

BIBLIOGRAFIA

ANDREWS A.L., 1941 a — Notes on the Warnstorf Sphagnum herbarium II. The section

Malacosphagnum, Bryologist 44 : 97-102.

ANDREWS A.L., 1941b — Notes on the Warnstorf Sphagnum herbarium III, The subgenus

Inophloea in South America. Bryologist 44: 155-159.

ANDREWS A.L., 1947 — Studies in the Warnstorf Sphagnum herbarium IV. The group

Acutifolia in South America. Bryologist 50 : 181-186.

ANDREWS A.L., 1949 — Studies in the Warnstorf Sphagnum herbarium V. The group

Cuspidata in South America. Bryologist 52: 124-130.

BLOMQUIST H.L., 1938 — Peatmosses of the Southeastern states. J, Elisha Mitchell

Sci. Soc. 54 (1): 1-21, 5 pl.

CRUM H., 1976 — Mosses of the Great Lakes Forest. University Herbarium, University

of Michigan, Ann Arbor. (edición revisada), i-iii + 404 р.

FLORSCHÜTZ-DE WAARD J. & FLORSCHÜTZ P.A., 1979 — Estudios sobre criptó-

gamas colombianas III. Lista comentada de los musgos de Colombia. Bryologist 82

(2) : 215-259,

PURSELL R.A., 1973 — Un censo de los musgos de Venezuela. Bryologist 76 (4): 473-500.

ROBINSON H., 1967 — Preliminary studies on the bryophytes of Colombia. Bryologist

70 (1): 1-61.

WARNSTORF C., 1911 — Sphagnales-Sphagnaceae. In A. ENGLER, Das Pflanzenreich.

Sphagnologia Universalis. Leipzig.

Source : MNHN. Paris

CLAVE DEL GENERO SPHAGNUM 207

РІ. 1. — 1-4: Sphagnum palustre. 1, células corticales, x 280; 2, corte transversal de una

hoja rameal, x 400; 3, hoja rameal, x 13; 4, hoja caulinar, x 13. 5-8: Sphagnum magella-

nicum. 5, células corticales, x 170; 6, corte transversal de una hoja rameal, x 530; 7, hoja

rameal, x 17; 8, hoja caulinar, x 17. 9-12: Sphagnum erythrocalyx. 9, células corticales,

x 220; 10, corte transversal de una hoja rameal, x 500; 11, hoja rameal, x 17; 12, hoja

caulinar, x 17, 13-17: Sphagnum compactum. 13, células corticales de una rama, x 100;

14, corte transversal de una hoja rameal, x 500; 15-16, hojas rameales, x 17; 17, hoja

caulinar, x 27.

PI. IL. — 18-20: Sphagnum cyclophyllum. 18, hialocistes de una hoja rameal (lado aba-

xial), x 330; 19, corte transversal de una hoja rameal, x 700; 20, hoja caulinar, x 12. 21:

Sphagnum pylaesii. Hialocistes de una hoja rameal (lado abaxial), x 420, 22-25: Sphagnum

recurvum. 22, células corticales, x 200; 23, corte transversal de una hoja rameal, x 700;

24, hoja rameal, x 20; 25, hoja caulinar, x 20, 26-27: Sphagnum sancto-josephense. 26,

hialocistes de una hoja rameal (lado adaxial), x 700; 27, hoja caulinar, x 20. 28-32: Spha-

gnum cuspidatum. 28-29, hojas rameales, x 8; 30-31, cortes transversales de una hoja

Tameal, x 700; 32, hialocistes de una hoja rameal (lado abaxial), x 700.

PI. IH. — 33-37: Sphagnum subsecundum. 33, hialocistes de una hoja rameal (lado

abaxial), x 330; 34, corte transversal de una hoja rameal, x 700; 35-36, hojas rameales,

x 23; 37, hoja caulinar, x 23. 38-42: Sphagnum sparsum. 38, hialocistes de una hoja

rameal (lado adaxial), x 500; 39, corte transversal de una hoja rameal, x 400; 40-41, hojas

rameales, x 23; 42, hoja caulinar, x 23. 43-46: Sphagnum oxyphyllum. 43, hialocistes de

una hoja rameal (lado adaxial), x 500; 44, corte transversal de una hoja rameal, x 700;

45, hoja rameal, x 23; 46, hoja caulinar, x 23. 47-51: Sphagnum capillifolium. 47, hialo-

cistes de una hoja rameal (lado abaxial), x 600; 48, corte transversal de una hoja rameal,

x 700; 49-50, hojas rameales, x 23; 51, hoja caulinar, x 23.

PL IV. — 52-56: Sphagnum limbatum. 52, vista de la corteza caulinar, x 260; 53, corte

transversal de una hoja rameal, x 700; 54, hialocistes de una hoja rameal (lado abaxial),

x 260; 55-56, hojas rameales, x 23. 57-62: Sphagnum meridense. 57, vista de la corteza

caulinar, x 170; 58, corte transversal de una hoja rameal, x 500; 59, hialocistes de una

hoja rameal (lado abaxial), x 200; 60-62, hojas rameales, x 18.

Source : MNHN, Paris

CLAVE DEL GENERO SPHAGNUM 209

Source MNHN, Paris

D. GRIFFIN

source : ММ

211

CLAVE DEL GENERO SPHAGNUM

Source : MNHN. Paris

213

L-USNATE AND PERMEABILITY

С. VICENTE and В. CIFUENTES *

SUMMARY. — Sodium L-usnate included in the culture media produces both structural

and metabolic modifications in Proteus mirabilis cells. By adding the drug to a cell culture,

oxidation rate of fumarate is strongly stimulated. This increase cannot be explained by

changes їп oxidation enzyme levels but by changes in the permeability of the cells to fuma-

rate. Oxidative phosphorylation in membrane vesicles is not altered by the drug at the

assayed concentrations. The cell membrane is disrupted at a polar position and the cell-

wall is enlarged by the action of sodium L-usnate.

INTRODUCTION

Lusnie acid, a natural product of lichen species, has been shown to exert

antibacterial activity especially toward gram positive organisms (MARSHAK et

al. 1947, BUSTINZA 1951). The antibacterial activity could be explained by the

action of L-usnie acid which is not only an uncoupler of the oxidative phos-

phorylation (JOHNSON et al. 1950) but also an inactivator of some enzymes,

eg. glutamate dehydrogenase (VICENTE et al. 1973) and urease (VICENTE

et al. 1978, VICENTE and CIFUENTES 1979). In addition, an effect of the

drug on membrane permeability has been shown in some lichens which syn-

thesize usnic acids (FOLLMAN and VILLAGRAN 1965). Similarly, the growth

of Proteus mirabilis, a gram negative bacterium, was accelerated by adding

sodium L-usnate to the culture media (MANSO and VICENTE 1971). This

effect of the drug could be explained by a change in membrane permeabi-

lity provoking a better nutrient uptake.

In the present paper the modifications of both structure and permeability

of Proteus mirabilis membrane are investigated.

* Cátedra de Fisiología Vegetal, Facultad de Biología, Universidad Complutense, Madrid 3,

Spain.

Cryptogamie, Bryol. Lichénol., 1981, 2, 2 :213-222.

Source : MNHN, Paris

214 C. VICENTE and В. CIFUENTES

MATERIAL AND METHODS

Proteus mirabilis, NCIB 5887, was used throughout this work. The micro-

organism was maintained on slants о the solid medium recommended by MA-

САМА and RUIZ (1967). The basal medium used for liquid culture was 0.7 g

K,HPO,, 0.3 g KH PO4, 0.5 g (МНа)2503, 0.01 g MgSOs 7H0 and 100 ml

distilled water. In some experiments, medium was supplemented with 50 mM

sodium salt of acetic, citric or fumaric acids and sodium L-usnate at а series of

concentrations.

15 mg in dry weight of cells were used to inoculate 100 ml of liquid medium.

To study carbohydrate degradation and related processes a large inoculum is

usual (AMARAL and KORNBERG 1975). Incubations were continued with a

vigorous shaking at 37°C. Growth curves were determined by turbidity measu-

rements at 610 nm.

Intact cells were collected by centrifugying the media at 16,000 x g for 15

minutes at 4°C. The pellet was washed in sterile conditions with 75 mM phos-

phate buffer, pH 6.9, and suspended in the same buffer. To obtain membrane

vesicles the cells were suspended in EDTAdysozyme according to the method

of TSUCHIYA (1976).

To prepare cell-free extracts, intact cells were washed two times with 0.1 М

NaCl, collected by centrifugation and suspended in 0.25 mM Tris-HCl, pH 7.0,

containing 0.14 М 2-mercapto-ethanol. The cells were disrupted by sonication

at 8,000 microns for 1 minute with ice protection and then centrifugated at

29,000 x g for 20 minutes at 4°C. Supernatant was used as cell-free extract.

Oxygen uptake was measured by the conventional WARBURG technique

at 37°C with air as the gas phase and 0.2 ml 40 per cent (w/v) potassium hydro-

xide in the centre cup. The main compartment of the vessel contained 1.0 mg

of intact cells, or membrane vesicles and enough 50 mM phosphate buffer,

pH 6.9, to give a final volume of 2.5 ml. The sodium salt of fumaric acid (25

umol in 0.5 ml) was added from the side arm after equilibration. After 30 mi-

nutes, the cells were removed from the mixtures by filtering through a millipore

filter (0.22 um of pore). Then, remaining fumarate was estimated according to

ALBERTY et al. (1954).

Membrane vesicles were also assayed for the oxidative phosphorylation

according to TSUCHIYA (1976) using fumarate (20 mM sodium salt) instead

of Ddactate. Protein was estimated by the method of LOWRY et al. (1951)

using bovine serum albumin as a standard.

Isocitrate, succinate and malate dehydrogenases were assayed according to

COOK and SAWAL (1969), VEEGER et al. (1969) and MURPHEY and KITT

(1969) respectively, using the enzyme units proposed by the authors.

Osmotic resistance of the cells was determined by suspending the cells in

distilled water, shaking them at 37°C for time periods varying from 0-6 hours

and measuring the optical density at 610 nm

Source - ММНМ. Paris

L-USNATE AND PERMEABILITY 215

For electron microscope, cells were fixed for thin sectioning by adding

glutaraldehyde. After 2 hours of glutaraldehyde fixation at 25°C, the cells were

post-fixed in osmium tetroxide and stained with uranyl acetate-lead citrate

(HIGGINGS 1972).

RESULTS

Cells are cultured on three media supplemented with 50 mM acetate, citrate

or fumarate. A lag phase of growth is observed in all cases. The lag period is

about 8 hours for the cells cultured on acetate or fumarate whereas the lag

period for cells cultured on citrate is more than 10 hours (fig. 1). The number

of viable cells decreases at the start of the lag phase but increased later. Addition

of sodium L-usnate in the culture media changes these results. Active growth

is initiated in the first two hours of incubation using 1.4 or 14.0 ир sodium L-

usnate per ml medium (fig. 2 and 3). The best growth is obtained when acetate is

used as carbon source.

Capacity for oxidation of fumarate by intact cells grown on different carbon

Е от E

È E

5 5

E $

5 d

E E

viable cells/ml (108)

viable cells/ml (+108 )

hours

Fig. 1. — Growth curves of Proteus mirabilis as A) increase in dry weight ог В) number of

viable cells, growing on media supplemented with (А) acetate; (0) fumarate or (№) citrate.

Fig. 2. — Growth curves of Proteus mirabilis growing on media containing 1.4 g/ml sodium

L-usnate. For symbols, see fig. 1.

Source : MNHN. Paris

216 C. VICENTE and B. CIFUENTES

sources is shown in tab. I. When there is no sodium L-usnate in the media,

fumarate is only oxidized by cells grown on fumarate. However, fumarate is

efficiently oxidized by cells grown on the media containing the lowest concen-

tration of sodium L-usnate (1.4 ug/ml), using fumarate or acetate as carbon

source whereas the bacteria grown on citrate consume no oxygen. Cells grown

on the highest concentration of the drug (14.0 ug/ml) oxidize fumarate more

efficiently. In this case, the metabolite oxidation rate is comparable in the cells

grown on fumarate, acetate or citrate. Remaining fumarate in the mixtures,

estimated on the basis of its extinction coefficient after having removed the cells

by filtration, is in parallel with the oxidation rates.

Table I. Fumarate oxidation by intact cells of Proteus mirabilis grown on

several carbon sources. Each value is the average of 4 replicates.

Fumarate oxidation

Carbon source L-usnate in

in the media the media Oxygen uptake Remaining fumarate

(g/m) (l/min) in the mixtures

(umo1)

Fumarate --- 0.49 + 0.03 24.2 + 1.52

1.4 2.73 + 0.31 21.1 + 1.40

14.0 13.96 + 1.10 6.2 + 0.61

Acetate — 0.00 24.7 + 1.58

1.4 2.00 + 0.19 22.0 + 1.49

14.0 9.74 + 0.94 11.6 + 1.11

citrate --- 0.00 24.8 + 1.76

1.4 0.50 + 0.02 24.3 + 1.82

14.0 8.56 + 0.92 13.9 + 1.23

Dehydrogenase activities in cell-free extracts are shown in tab. IL. Although

isocitrate dehydrogenase activity slightly increased as a function of incubation

time, no change in activity is observed after addition of sodium L-usnate. The

malate dehydrogenase level changed somewhat after prolonged incubation time

but succinate dehydrogenase falls in all the cases. This decrease has a greater

extent in bacteria grown in control media with acetate or fumarate than in

organisms grown in the presence of L-usnate.

Both oxygen uptake and oxidative phosphorylation are not changed using

membrane vesicles isolated from the cells grown in presence ог absence of the

drug (tab. Ш).

These results might have a chemical basis which may be reflected in structu-

ral changes. Analysis of the osmotic resistance of the cells as well as a study of

cell membrane structure in the electron microscope were carried out to investi-

gate this hypothesis.

Source : MNHN. Paris

Table II. Levels of several tricarboxylic acid cycle enzymes in cell-free ехїгасїз of Proteus

mirabilis. Each value is the average of 4 replicates.

y Enzyme activities ( units

Carbon source i-usnate in m ( y

in the media the media Tsocitrate Malate Succinate a

(ug/m1) dehydrogenase dehydrogenase denydrogenase а

at zero at 10h at zero at 10h at zero at 10h >

time time time a

Fumarate — 0.46+0.06 0.53+0.05 0.002 0.004 0.33+0.02 0.10+0.01 =

1.4 0.53+0.05 0.002 0.28+0.04 E

14.0 0.48+0.05 0.004 0.30+0.03 =

Acetate == 0.30+0.03 0.49+0.06 0.001 0.001 0.30+0.03 0.04+0.01 E

1.4 0.50+0.05 0.001 0.34+0.02 i

14.0 0.44+0.05 0.001 0.22+0.03

Citrate — 0.34+0.03 0.39+0.03 0.001 0.002 0.51+0.04 0.20+0.01

1.4 0.38+0.03 0.002 0.13+0.01

14.0 o 0.03 0.002 0.12+0.01

LIT

Source : MNHN, Paris

Tab. Ш. Oxygen uptake and ATP formation by membrane vesicles isolated from Proteus

mirabilis. Each value is the average of three replicates.

L-usnate included Oxygen uptake ATP formed

in the culture me- (umol/mg protein (ито1/тд pro-

dia (ug/ml) and minute) tein and min)

== 1.55 + 0.07 4.75 + 0.30

1.4 1.62 + 0.10 5.01 + 0.50

14.0 1.52 + 0.09 4.52 + 0.40

1 mg membrane vesicles loaded with ADP are suspended in 50 mM phosphate buffer,

0.28; sucrose and 5 mM МЕСІ). Reaction is initiated by addition of 20 mM fumarate (final

volume 3.0 ml) and followed for 5 minutes.

dry weight (mg/ml)

optical density at 610 nm

viable cells/ml («109)

ERFORSCHEN |

54:243 PES MB

hours hours

Fig. 3. — Growth curves of Proteus mirabilis growing on media containing 14.0 pg/ml sodium

L-usnate. For symbols, see fig. 1.

Fig. 4. — Decrease in optical density at 610 nm (circles) and number of viable cells (trian-

gles) of cell suspension in distilled water. (0,4) cells grown on 14.0 pg/ml sodium L-

usnate; (О, 4) cells grown on media without the drug.

Source : MNHN. Paris

L-USNATE AND PERMEABILITY 219

Two parallel cultures on fumarate were grown for 10 hours in the presence

of the drug in the highest concentration (14.0 ug/ml) to verify changes in the

osmotic resistance. Bacteria, collected and washed according to the conditions

specified in methods, are dispersed in sterile water and kept vigorously shaking

for 6 hours. A rapid decrease in the optical density at 610 nm is observed in

parallel with a decrease in the number of viable cells (fig. 4). Cells grown in the

presence of L-usnate are less susceptible for osmotic shock.

An electron microscopic study of the cells grown at the same conditions as

above was performed. The cells which are grown on L-usnate show a partially

destroyed membrane (fig. 5b). The disruption of this three-layered structure is

located in a polar position for L-forms of Proteus. Enlargement of the cell-wall

occurred at the same position.

Fig. 5. — Thin sections of Proteus mirabilis. In a) the cells are grown for 10 hours on media

supplemented with 50 mM fumarate. In b), 14.0 ig/ml sodium L-usnate is included in

the media. tlm = three layered membrane; bm = broken membrane; ecw = enlarged

cell wall.

DISCUSSION

Oxidation rate of several tricarboxylic acids cycle metabolites has been cho-

sen as permeability characteristic in Pseudomonas spp. (KOGUT and PODOSKI

1953, BARRET and KALLIO 1953) and Pseudomonas aeruginosa (CLARKE

and MEADOW 1959) considering that oxidation enzyme of TCA cycle are

constitutive enzymes. Therefore, variable oxidation rates of a sole metabolite

Source : MNHN, Paris

220 C. VICENTE and В. CIFUENTES

by cells cultured on disctinct carbon sources are a function of induced permease.

However, this claim is unsubstantiated in Proteus mirabilis without an adequate

experimental basis. The activity of several dehydrogenases can be changed by ca-

tabolite repression (TAKAHASHI 1975) or inactivation (BENNET and HOLMS

1975) in Escherichia coli, or by carbon source uptake (LEWIS and MILLER

1975) in Desulfovibrio. Alteration of dehydrogenase activities has also been

observed in mitochondria of growing and resting yeast (UTTER et al. 1968).

However, the observed changes in succinate, malate and isocitrate dehydroge-

nase activity does not explain the increase in the fumarate oxidation rate when

the organism has been cultured on L-usnate (see tab. 11). Cells grown on fuma-

rate in the presence of 14.0 ug/ml sodium L-usnate oxidize fumarate at rate

about 30 times higher than those cells which are grown in the absence of the

drug (tab. 1). When bacteria are not induced for permease, fumarate oxidation

is only carried out when they are cultured on media containing L-usnate, their

oxidation rates being similar to those obtained by cells containing permease.

Thereby, it is necessary to reject the hypothesis about an action of the drug

on permease levels.

There is no change on both ATP formation and oxygen uptake in membrane

vesicles incubated with fumarate in the presence or absence of sodium L-usnate.

So that increase in oxygen uptake by intact cells treated with the drug cannot

be explained by the action of phenol on the oxidative phosphorylation, as it

has been described for dinitrophenol (LOOMIS and LIPMANN 1948) and con-

firmed for L-usnic acid (JOHNSON et al. 1950).

Alterations of membrane continuity induced by L-usnate could be the basis

of the permeability changes. It is highly improbable that a cell containing gross

ruptures of the cytoplasmic membrane, as claimed from the electron micro-

graphs, would be viable, This disruption may be interpreted as the final stage

in the degradation process, the intermediate states being responsible of the

accelerated uptake of fumarate. But since it is always localised in the thickening

of cell-walls, the disruption cannot be adscribed to a tangential section of the

cells but to a true enlargement of the structure.

ACKNOWLEDGEMENT, — We thank Prof. Е. Bustinza for a generous supply of sodium

L-usnate and Prof. M. Rubio (Instituto Jaime Ferrán, CSIC) for electron micrographs. This

work has been supported, in part, by a grant from INAPE (Ministerio de Universidades,

Espana) to Miss Blanca Cifuentes.

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Source : MNHN, Paris

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Source : MNHN, Paris

223

RESISTANCE А ГА SECHERESSE

ЕТ MULTIPLICATION VEGETATIVE

CHEZ FOSSOMBRONIA CAESPITIFORMIS DE NOT.

С. LEFORESTIER *

RESUME, — Entre mai et aoút, les Fossombronia caespitiformis De Not. de la région nigoise

resistent á la chaleur et А la sécheresse grace & des modifications morphologiques : flétrisse-

ment et chute des feuilles, dessèchement progressif des axes où il ne subsiste que quelques

zones chlorophylliennes. Il semble qu'il y ait un lien entre ces formes de résistance et la

multiplication végétative : les zones chlorophylliennes donnent de jeunes pousses dès les

premiéres pluies de fin d'été; les feuilles tombées au sol peuvent former tout au long de

l’année de jeunes plantes.

SUMMARY. — Between May and August, Fossombronia caespitiformis De Not. around

Nice resist to hot weather and to dryness by morphological modifications : fading and fall

of leaves, progressive withering of axis where only a few chlorophyllian points remain.

It seems that there exists a link between forms of resistance and vegetative multiplication :

chlorophyllian points give young sprouts after the first rains of the end of summer; the

leaves fallen on the ground can make, during all the year, young plants.

Certains mécanismes de résistance à la chaleur et à la sécheresse sont sans

doute liés à ceux de la multiplication végétative. Celle-ci complete efficacement

la reproduction sexuée chez Fossombronia caespitiformis dans la région médi-

terranéenne.

RÉSISTANCE А LA CHALEUR ET А LA SÉCHERESSE

MODIFICATIONS DES CORMUS. — La période de l’année comprise entre

mai et août est caractérisée par une trés faible pluviosité dans la région méditer-

ranéenne. Les gamétophytes de Fossombronia caespitiformis subissent alors des

modifications qui leur permettront de subsister pendant l’été sous diverses

formes d'adaptation. Ils seront ainsi réduits à un état de vie ralentie, puis, dès

* Laboratoire Botanique, Faculté des Sciences et des Techniques, 06034 Nice Cedex. Des-

sins de D. CHIAVERINI.

Cryptogamie, Bryol. Lichenol., 1981, 2, 2 : 223-232.

Source : MNHN. Paris

224 C. LEFORESTIER

les premières pluies de fin d'été, ils redeviendront verts et retrouveront une

vitalité optimale.

Les modifications morphologiques succèderont à la formation des capsules

pendant l'hiver, à celle des jeunes feuilles et, éventuellement, des archégones

et des anthéridies au printemps.

Différents stades de flétrissement ont été observés :

L Les cormus deviennent plus sombres, ternes, moins vigoureux, les feuilles

sont aplaties les unes contre les autres, affaissées au lieu d’être dressées, en

état de fanaison.

— En méme temps la marge des feuilles les plus âgées brunit. Ces feuilles,

puis les moins ägees et enfin celles de Гарех sécheront, et certaines se détache-

ront du cormus dont la base se trouve alors dépourvue de protection. A ce stade

le cormus est encore chlorophyllien (fig. I, 1).

- Ensuite se produit le desséchement des cellules externes du cormus qui

devient gris et se flétrit superficiellement. Le centre de Гахе reste chlorophyllien.

— Un desséchement encore plus poussé est possible. ЇЇ ne reste du cormus

qu'un axe dépourvu de feuilles, gris, aplati, dont le centre est quasiment sec.

Seuls quelques groupes de cellules saillants (bourgeons de résistance) restent

verts et subsistent è la surface de l’axe (fig. I, 2, 3, 4).

Des modifications dans le port du végétal ont été également observées. On

trouve parfois, à la fin de l'été, des apex repliés sur eux-mêmes et ainsi protégés

par le reste du cormus (fig, 1, 5). Cette courbure semble s'amorcer dès la crois-

sance de printemps, avant que les chaleurs ne stoppent le processus.

Les observations suivantes montrent Pimportance de l’eau pour la vie des

cormus :

— Des Fossombronia caespitiformis bien arrosés au printemps restent chloro-

phylliens, feuillés, et conservent leur aspect habituel au moins dans leur extré-

mité antérieure.

— Ces cormus épargnés par la dessication portent en juin-juillet des arché-

gones et des anthéridies (fig. 1, 6) et même des capsules. En général cette géné-

ration de printemps ne s'effectue pas ou est interrompue par les conditions

climatiques estivales dans notre région.

REGENERATION DES СОВМО$. — Après quelques heures de pluie, les

cormus ternes, dépourvus de feuilles, changent d'aspect; le développement

apical reprend, de nouvelles feuilles et de jeunes rhizoides se forment, les cormus

Sallongent, le contour des taches de Fossombronia caespitiformis devient

circulaire. Le centre des taches est constitué par la base des plantes plus ou

moins dégénérées et la circonférence par la multiplicité des apex chlorophylliens.

Anthéridies et archégones apparaîtront ensuite, assurant la formation des cap-

sules pendant l’hiver.

Les bourgeons de résistance vont, eux aussi, se développer. Les premières

feuilles naissent très proches les unes des autres. La poussée de Рахе leur donnera

leur emplacement définitif.

Source : MNHN. Paris

FORMES DE RESISTANCE CHEZ FOSSOMBRONIA 225

Fig. 1. — 1 :Debut de flétrissement d'un cormus. Les feuilles inférieures se dessèchent les

premières. 2, 3, 4 : Bourgeons de résistance sur des cormus flétris. r = rhizoide. 5

Apex chlorophylliens recourbés. Cormus prélevés 2 jours aprés les orages de septembre.

Apex recourbés semblant être ainsi protégés de la dessication. 6 : Cormus porteur de

jeunes antheridies (juillet).

Les observations suivantes mettent en évidence la reprise de croissance des

cormus par leur apex ош par des bourgeons de résistance :

— Sur un ancien cormus, une masse chlorophyllienne а évolué en donnant

deux feuilles et un rhizoïde (fig. II, 1, a, b, c).

Source : ММНМ. Paris

226 C. LEFORESTIER

_ Un cormus conservé au laboratoire, partiellement déshydraté et remis

dans un environnement humide, a formé une plantule feuillée (fig. II, 2).

_ Un cormus observé le jour méme de sa récolte (octobre) portait une jeune

pousse feuillée et garnie de rhizoïdes (fig. 11, 3).

Fig. II, — 1 a, b, c : Évolution d'un apex chlorophyllien sur un ancien cormus (octobre). f =

feuilles, г = rhizoide. 2 : Plantule feuillée en formation (juin). 3 : Pousse feuillée et

rhizoides sur un ancien cormus (octobre). 4, 5 : Bourgeonnement d'anciens cormus (oc-

tobre). 6 :Jeunes feuilles poussant sur un ancien cormus (septembre). 7 : Ancien

cormus avec 4 zones de formation de nouvelles feuilles (septembre).

Source : ММНМ. Paris

FORMES DE RESISTANCE CHEZ FOSSOMBRONIA 227

— Un seul cormus initial peut donner naissance à plusieurs nouvelles pousses;

ainsi la figure II, 4 représente un cormus ancien portant au mois d’octobre deux

nouvelles proliférations, l'une de quatre feuilles et l'autre, plus jeune encore, à

l'état de bourgeon. Entre elles se trouvent quelques rhizoides hyalins.

— Un cormus brunître, recueilli en octobre également, a formé trois jeunes

cormus chlorophylliens; malgré leurs dimensions encore très réduites, des feuilles

commencent á se différencier (fig. II, 5).

— Deux autres cormus forment de nouvelles feuilles en un point (fig. II, 6),

ou en quatre points (fig. II, 7).

Dans la nature, ce renouveau se produit dés l'arrivée de la pluie depuis la fin

du mois d’aoüt et jusqu’en septembre. Le retour à une taille normale, la nais-

sance d’archégones et d’anthéridies, la formation des capsules s’effectuent sans

incident après les pluies d'été, puisque aucune période de sécheresse et de cha-

leur n'intervient dans notre région pendant plusieurs mois. Les cormus conservés

au laboratoire puis remis en atmosphere humide sont également le siège d’une

nouvelle activité après leur dessèchement,

La sécheresse apparaît donc comme le facteur principal du blocage des

phénoménes de la vie active. Il n'existe pas de période de repos obligatoire,

particulière aux F. caespitiformis.

MULTIPLICATION VEGETATIVE

C’est apparemment un mode de résistance à la chaleur et à la sécheresse.

Elle se présente de deux façons :

1) Les bourgeons de résistance formés sur les cormus desséchés, donnent de

nouveaux axes permettant la continuité de la vie et favorisant l’extension des

touffes.

2) De nouvelles plantes se forment sur des feuilles tombées au sol; en un

point de leur surface apparaît un boursouflement susceptible d’évoluer en un

futur cormus. Sur ce bourgeon très chlorophyllien, muni d’un ou de plusieurs

thizoides, apparaissent les jeunes feuilles. Cette multiplication végétative ne

se produit pas seulement à la fin de l'été. Trés importante aussi en mai avant

la grande période de chaleur et de sécheresse, elle s'effectue selon le même

processus, quelle que soit la saison.

Les faits suivants ont été remarqués en février :

— Une feuille isolée, tombée au sol entre des cormus adultes, reste chloro-

phyllienne sur le tiers environ de sa surface, le reste devenant violet. Sur cette

dernière zone se développe une masse cellulaire très chlorophyllienne constituée

de cellules de taille inférieure А celles de la feuille et portant quelques rhizoides

(fig. Ш, 1).

— Une feuille а demi-desséchée (encore attachée au cormus récolté le matin

méme) porte un «bourgeon» chlorophyllien. Trois jeunes rhizoides sont déjà

formés (fig. III, 2).

— Une feuille porte à sa base un amas cellulaire et quatre rhizoides (fig. III,

3).

Source : MNHN, Paris

228 C. LEFORESTIER

Fig. Ш. 1 :Groupe de cellules chlorophylliennes et rhizoides sur une feuille tombée

au sol et à demi-desséchée (mai). 2, 3, 4 : Même formation qu’en 1 (février). 5 : Jeune

pousse provenant d’une feuille tombée au sol (février).

Fig. IV. — Jeune plantule feuillée et munie de rhizoides, encore fixée sur le feuille initiale,

longue de 0,7 mm (mai). A : ensemble feuille-plantule. В : plantule. C : jeunes feuilles.

Source : MNHN. Paris

230 C. LEFORESTIER

_ Une autre feuille porte deux jeunes feuilles en formation et cing rhizoides

(fig. III, 4).

` Dans un cas, la nouvelle formation chlorophyllienne se présente comme

une pousse longue et fine. Peut-étre la feuille initiale était-elle enfouie sous une

pellicule de terre, car seule était visible dans la masse de sol recueilli l'extrémité

de la jeune pousse. Trois rhizoides sont formés (fig. LI, 5).

Les photographies représentent le méme phénoméne remarqué en mai. Le

processus se déroule sans doute à différentes époques de l'année, mais la sé-

cheresse de l'été qui provoque la chute de feuilles de la plante le favorise.

DISCUSSION

La multiplication végétative est très répandue chez les Hépatiques. Citons

quelques exemples :

Calobryales. — La multiplication de cellules du centre des feuilles donnant

naissance à de futures plantes est signalée par FULFORD & DILLER (1957)

chez Haplomitrium hookeri (Lyell) Nees en culture.

Metzgeriales. — Dans le genre Fossombronia Raddi la multiplication vege-

tative a déjà été décrite chez plusieurs espéces. CHALAUD (1929) parle de

«rameaux adventifs des stations naturelles», de «rameaux adventifs de tiges

mutilées» («pousses de régénération»), et de «rameaux foliairesy chez Fossom-

bronia pusilla Dum. SRIVASTAVA & UDAR (1975) citent la présence de «cons-

picuous apical tubers in the plant» (Fossombronia himalayensis Kash.) en pé-

riode de sécheresse: ces tubercules pénètrent même dans le sol. PATON (1974),

dans la note sur Fossombronia fimbriata Paton, pense que cette езрёсе possède

une multiplication végétative semblable à celle qui existe chez d’autres Fossom-

Lronia. SCHUSTER (1966) signale chez Fossombronia lamellata St. la présence

de séries de petits tubercules pédicellés, susceptibles de devenir le siège d’une

germination donnant de nouvelles plantes. D'après ОМО (1979), toutes les

cellules du thalle chez les Fossombronia et les Metzgeria montrent un haut

degré de régénération. Chez Petalophyllum indicum Kash., la présence de «ma-

ture tubers» signalés des le début d'avril, permettent à la plante de subsister

pendant l'été (MEHRA & VASHIST 1950).

Jungermanniales. — Dans le genre Ceratolejeunea, et les espèces suivantes,

Frullania asagrayana St., Isopaches bicrenatus (Schmid.) Buch, Mastigolejeunea

auriculata (Wils. et Hook.) Schiffn., des plantes nouvelles proviennent de cellules

foliaires. Chez Nowellia curvifolia (Dicks.) Mitt., la régénération, après traite-

ment approprié, se produit sur la tige, la feuille et le périanthe (FULFORD

1955).

Marchantiales. - JOVET-AST (1971) expose le cas du thalle de Riccia peren-

nis St. pourvu d’un filament termine par un tubercule résistant lors de la dessi-

cation du gamétophyte et susceptible de se développer en un thalle nouveau.

JOVET-AST & BISCHLER (1971) ont constaté que, pendant l'été, les Riccia

Source - MNHN. Paris

FORMES DE RESISTANCE CHEZ FOSSOMBRONIA 231

perennis St. ne subsistaient que par leurs tubercules enfouis dans le sol. JOVET-

AST (1979) cite le cas de Riccia fluitans L., qui, mis en culture sur des eaux

de pluie, a fabriqué des bourgeonnements donnant de nouveaux thalles. Ce fait

est particuliérement remarquable car, contrairement aux espèces énumérées,

R. fluitans vit en milieu aquatique.

BISCHLER (1977) signale que les Plagiochasma Lehm. et Lindenb. forment

des «rameaux ventraux abondants» dans des conditions de vie difficile ou en

cas de «section volontaire ou accidentelle de l’apex du thalle».

La multiplication végétative existe donc très fréquemment dans tous les

groupes d’Hépatiques, plus souvent chez les Jungermanniales où elle se manifeste

sous des formes diverses : propagules foliaires, disques multicellulaires, fragmen-

tation de souches, fragmentation et chute de feuilles, périanthes caducs. Cepen-

dant chez les Metzgériales, des propagules existent dans certains cas (Blasia)

et des disques propagulifères chez Metzgeria uncigera Evans. La multiplication

végétative par l'intermédiaire des feuilles est très active chez Fossombronia

caespitiformis. La formation de tubercules existe chez les trois genres voisins,

Fossombronia, Petalophyllum et Sewardiella. Il semble que, par des moyens

divers, et a différents endroits des gamétophytes à morphologie dissemblable,

la multiplication végétative soit très importante et quasiment constante des que

les conditions d’environnement l’exigent.

Chez Fossombronia caespitiformis, entre mai et aoüt, un temps de repos est

observé. Il correspond à une période de sécheresse et de chaleur intenses. Un

certain nombre de modifications morphologiques permettent la survie des cor-

mus. Le développement des bourgeons de résistance en jeunes cormus débute

des la reprise des pluies de fin d'été. Cette multiplication végétative est alors

intense. Par contre, le développement de nouveaux cormus sur des feuilles tom-

bées au sol, moins spectaculaire, s’observe à différentes époques de l’année.

Le nombre de plantes filles est bien supérieur à celui des plantes mères en

raison de la présence possible de plusieurs bourgeons de résistance sur un même

cormus — chacun donnant un apex — et du nombre de feuilles détachées, for-

mant par bourgeonnement, chacune un nouveau cormus.

La multiplication végétative peut être interprétée dans le cas des bourgeons

de résistance comme une adaptation à la période sèche.

BIBLIOGRAPHIE

BISCHLER H., 1977 — Plagiochasma Lehm. et Lindenb. Rev. Bryol. Lichénol. 43 : 67-109.

CHALAUD G., 1929 — Le cycle évolutif de Fossombronia pusilla Dum. Étude morpholo-

gique et cytologique suivie d'une révision des espéces européennes des genres Simodon et

Fossombronia. (Thèse Fac. Sciences, Toulouse). Rev, Gén. Bot, 41 : 1-343.

FULFORD M., 1942 — Sporelings and vegetative reproductive structures in Mastigolejeunea

auriculata. Amer. J. Bot. 29 : 848-850, fig. 1-19.

Source : ММНМ. Paris

232 C. LEFORESTIER

FULFORD M., 1944 — The genus Ceratolejeunea. Bull. Torrey Bot. Club 71 : 638-654,

fig. A-J and 1-72.

FULFORD M., 1955 — Sporelings, gemmalings and regeneration in Isopaches bicrenatus

(Schmid.) Buch. Bryologist 58 : 317-322.

FULFORD M., 1955 — The patterns of development of sporelings, gemmalings, and regene-

rants in Nowelia curvifolia (Dicks.) Mitt. Rev. Bryol. Lichénol, 24 : 41-48, fig. 1-28.

FULFORD M., 1956 — The pattern of regeneration in Frullania asagrayana. Bryologist 59 :

265-270, fig. 1-23.

FULFORD M. and DILLER V.M., 1957 — Studies on the Haplomitrium I. Rev. Bryol.

Lichénol. 1956 (1957), 25 : 241-246.

FULFORD M., 1964 — Contemporary thought in plant morphology : Hepaticae and Antho-

cerotae. Phytomorphology 14 : 103-119.

JOVET-AST S., BISCHLER Н. 1971 — Distribution, écologie, sociologie du Riccia perennis

St. Rev. Bryol. Lichénol. 1970 (1971), 37 : 247-264.

JOVET-AST S., 1971 — Riccia perennis Stephani. Nomenclature, morphologie, caryotype,

affinités. Rev. Bryol. Lichénol. 1970 (1971), 37 : 237-245.

JOVET-AST S., 1979 — Récolte en France d’un Riccia fluitans L. fertile. Rev. Bryol. Liché-

nol. 45 :179-183.

MEHRA PN., VASHIST B.R., 1950 — Embryology of Petalophyllum indicum Kash., and

a new suggestion of the evolution of thalloid habit from foliose forms. Bryologist 53 :

89-114, fig. 1-63.

NOAILLES M.C., 1978 — Etude ultrastructurale de la récupération hydrique après une

période de secheresse chez une Hypnobryale : Pleurozium schreberi (Willd.) Mitt. Ann.

Sci. Nat. Bot., sér. 12, 19 : 249-265.

ONOS., 1979 — Studies on vegetative regeneration in the Metzgeriales, Hepaticae. J. Hattori

Bot. Lab. 46 : 169-191.

PATON J.A., 1974 — Fossombronia fimbriata sp. nov. J. Bryol. 8 : 1-4.

SCHUSTER R.M., 1966 — The Hepaticae and Anthocerotae of North America East of the

Hundredth Meridian, Columbia University Press. New York and London.

SRIVASTAVA S.C., UDAR R., 1975 — The genus Fossombronia Raddi in India, with a

Note on the Indian taxa of the family Fossombroniaceae. Nova Hedwigia 26 1 799-845.

Source : MNHN, Paris

233

МОТЕ

FLORULE BRYOLOGIQUE DES ROCHERS BATHONIENS

R.DHIEN*

L'appétence chimique des muscinées se trouve nettement délimitée si l'on

considére des espéces calcicoles ou silicicoles. Mais on peut se demander quelle

préférence manifestent les végétaux pour les différents étages géologiques.

Ainsi, la carte géologique de la France (Beaune по 125) mentionne : Rauracien,

Oxfordien et Callovien, Bajocien, Bathonien. Ce dernier comprend le Bathonien

supérieur, peu étendu, et le Bathonien moyen se subdivisant en 1) calcaire blanc

compact, 2) calcaire blanc oolithique. J’ai recensé, pour ce dernier, les musci-

nées croissant sur les roches situées dans les localités suivantes : Aubaine, Bligny-

sur-Ouche, Lusigny-sur-Ouche, Montceau-Echarnant. On у trouve d’imposantes

formations les unes soumises à une insolation maximum, d’autres abritées par

des foréts qui en modifient la composition floristique.

Rochers secs et ensoleilles

Barbula fallax Hedw. circumboréale

Camptothecium lutescens (Hedw.) B.S.G. circumboréale

Campylium chrysophyllum (Brid.) J. Lange circumboréale

Ditrichum flexicaule (Schwaegr.) Hampe circumboréale

Entodon concinnus (De Not.) Par. (= E. orthocarpus

(Brid.) Lindb.) circumboréale

Fissidens cristatus Wils. ex Mitt. circumboréale

Grimmia pulvinata (Hedw.) Sm. cosmopolite

G. p. var. africana (Hedw.) Hook. f. et Wils.

(= С. orbicularis Bruch in Wils.) méditerranéenne

Homalothecium sericeum (Hedw.) B.S.G. circumboréale

Neckera complanata (Hedw.) Hüb. circumboréale

N. crispa Hedw. subatlantique

Orthotrichum anomalum Hedw. cosmopolite

Plagiopus oederi (Brid.) Limpr. circumboréale

Pleurozium schreberi (Brid.) Mitt. cosmopolite

Pseudoleskeella catenula (Schrad.) Kindb.

(=Pseudoleskea catenulata (Schrad.) B.S.G.) circumboréale

* Lusigny-sur-Ouche, 21360 Bligny-sur-Ouche, France.

Cryptogamie, Bryol. Lichénol., 1981, 2, 2 : 233-234.

Source : MNHN. Paris

234 R. DHIEN

Racomitrium canescens (Hedw.) Brid. circumboréale

Schistidium apocarpum (Hedw.) B.S.G.

(= Grimmia apocarpa Hedw.) cosmopolite

Tortella tortuosa (L.) Limpr. (souvent fructifiée) circumboréale

Tortula muralis Hedw. cosmopolite

T. ruralis (Hedw.) Gaertn. cosmopolite

Rochers plus ou moins humides ou abrités en forét

Aloina rigida var. ambigua (B.S.G.) Craig.

(=A. ericaefolia (Lindb.) Kindb.) méditerranéenne

Amblystegiella confervoides (Brid.) Loeske

(= Amblystegium с. (Brid.) B.S.G.) circumboréale

Anomodon attenuatus (Hedw.) Hüb. circumboréale

A. longifolius (Brid.) Hartm. circumboréale

Bryoerythrophyllum recurvirostre (Hedw.) Chen

(= Didymodon rubellus B.S.G.) circumboréale

Campylium hispidulum var. sommerfeltii (Myr.) Lindb.

(= С. sommerfeltii (Муг.) J. Lange) circumboréale

Ctenidium molluscum (Hedw.) Mitt. circumboréale

Dicranum scoparium Hedw. circumboréale

Encalypta vulgaris Hedw. cosmopolite

Hylocomium splendens (Hedw.) B-S.G. eurasiatique

Hypnum cupressiforme L. ex Hedw. cosmopolite

Lophozia collaris (Nees) Dum. (= L. muelleri (Nees) Dum.) circumboréale

Orthothecium intricatum (Hartm.) B.S.G. circumboréale

Oxyrrhynchium pumilum (Wils.) Loeske méditerranéenne

Pseudoscleropodium purum (Hedw.) Fleisch. circumboréale

Rhynchostegiella tenella (Dicks.) Limpr.

(= В. algiriana (Р. Beauv.) Warnst.) subméditerranéenne

Seligeria pusilla (Hedw.) B.S.G. (rare) circumboréale

Trichostomum brachydontium ssp. mutabile (Bruch) Giac.

(= Т. mutabile Bruch) cosmopolite

Frullania tamarisci (L.) Dum. circumboréale

Metzgeria pubescens (Schrank) Raddi circumboréale

Pedinophyllum interruptum (Nees) Kaal. cosmopolite

Plagiochila asplenioides (L.) Dum. circumboréale

Scapania aspera Bernet cosmopolite

Rochers tuffeux

Eucladium verticillatum (Brid.) B.S.G. circumboréale

Gymnostomum calcareum Nees et Hornsch. subméditerranéenne

Il conviendrait d'examiner si toutes ces espèces sont inféodées au Bathonien,

ou si elles colonisent aussi d'autres formations géologiques.

Source : MNHN, Paris

BIBLIOGRAPHIE BRYOLOGIQUE

D.LAMY*

SYSTEMATIQUE, NOMENCLATURE

ABRAMOVA A.L., ABRAMOV 1.1. — Quid est Ptychomitrium altaicum Broth. ?

Nov. Sist. Nizsh. Rast., Inst. Bot. Komarova, Akad. Nauk SSSR 1980, 17 :

174-178, en russe.

Comparaison entre Р. altaicum Broth. et P. sinense (Mitt.) Jaeg.

CROSBY M.R. — The diversity and relationships of mosses. In TAYLOR RJ.

and LEVITON A.E., ей. — The Mosses of North America. San Francisco,

1980. Voir : Ouvrages généraux : TAYLOR R.J. and LEVITON А.Е.

ENGEL J.J. — Austral Hepaticae X. A revision of Hepatostolonophora Engel et

Schust., nom. nov. (Hepaticae). J. Hattori Bot. Lab. 1979, 46 : 91-108,

8 fig. (Field Mus. Nat. Hist., Lake Shore Drive, Chicago, Illinois 60605

USA)

Hepatostolonophora Engel et Schust. nom. nov. pour Stolonophora Engel et

Schust. 1975, non Nizamuddin 1969 (Phaeophyta). Descr. du genre, sa place

parmi les Lophocoleaceae. Clé aux 3 esp. : Н. abnormis (Besch. et Mass.) c.n.

(Leioscyphus (?) a.), Н. rotata (Hook. f. et Tayl.) c.n. (= Jungermannia г.) avec

les var. rotata et perssonii (Schust.) c.n. (= Calyptrocolea p.) et Н. paucistipula

(Rodw.) c.n. (= Lophocolea p.). Pour chaque taxon : taxonom., synon., descr.,

ill., écol., distr.

FRAHM J.P. — Campylopodes Brasiliae exsiccatae. Herzogia 1979, 5 : 103-110.

(Gesamthochschule Duisburg, Fachbereich VI, Biologie, D-41 Duisburg).

Commentaires pour quelques spécimens de la série d’exsiccata. Noter Сатру-

lopus cuspidatus (Hornsch.) Mitt. var. dicnemioides (С. Müll.) c.n. (= С. dicne-

mioides (С. Müll.) Par.).

GODFREY J.D. and GODFREY G.A. — Jungermannia schusterana, a new hepa-

tic from the Pacific Coast of North America. J. Hattori Bot. Lab. 1979, 46 :

109-117, 3 fig. (Dept. Biol., Univ. Victoria, Р.О. Box 1700, Victoria, British

Columbia, Canada V8W 2Y2).

* Lab. Cryptogamie, 12 rue Buffon, 75005 Paris.

Cryptogamie, Bryol. Lichénol., 1981, 2, 2 : 235-252.

Source : MNHN, Paris

236 BIBLIOGRAPHIE BRYOLOGIQUE

Diagn., descr., Ш. de Jungermannia (Plectocolea) schusterana sp. nov. du

Canada, Washington et Alaska. Plante robuste avec des oléocorps botryoides.

АЙ. de cette nouv. esp. avec J. hattoriana (Amak.) Amak., J. torticalyx Steph.

et J. flagellata (Hatt.) Amak., 3 esp. du Japon.

GROLLE В. — Lebermoose aus Neuguinea. 17. Harpalejeunea. J. Hattori Bot.

Lab. 1979, 46 : 43-47, 1 fig, (Friedrich-Schiller-Univ., Sekt. Biol., 69 Jena,

DDR).

Loc., distr. de Harpalejeunea constricta Grolle, Н. filicuspis (Steph.) Mizut.

(deser.) et de Н. pinaudensis sp. nov. (diagn., descr., ill.) de la Nouvelle-Guinée.

Drepanolejeunea riddleana Steph. et Microlej. acuminata Steph. sont syn. avec

Н. filicuspis.

GROLLE В. — Miscellanea hepaticologica 191-200. J. Hattori Bot. Lab. 1979,

46 : 337-355, 3 fig. (Idem).

191, — Lectotype, descr. Ш. de Trachylejeunea aneogyna (Spruce) Grolle

em. (= Lejeunea a.). = 192. — Cheilolejeunea Steph. nov. 1890 (type : Cheilolej.

kurzii Steph.) doit remplacer Cheilolej. sensu Evans 1906. — 193. — Lectotypifi-

cation de Lejeunea intertexta Lindb. 1845; clé à Cheilolej. intertexta (Lindenb.)

Steph. s. str. et Cheilolej. serpentina (Mitt.) Mizut. Synonymie et distr. de ces

deux esp. Noter Cheiloj. usambarana (Steph.) c.n. (= Strepsilej. и.). — 194. —

Pycnolej. arietina Tixier est synonyme avec Cheilolej. ceylanica (Gott.) Schust.

et Kachr. — 195. — Euosmolej. sayeri Steph. et Euosmolej. trifaria (Reinw. et

al.) Steph. (respectivement esp. types de Euosmolej. Steph. 1889 et Euosmolej.

(Spruce) Schiffn. 1893) sont conspécifiques, Suggestion pour conserver Euosmo-

lej. (Spruce) Schiffn. 1893 et pour annuler le remplacement de Cheilolej. par

Euosmolej. — 196. Placolej. subarrhyncha Herz. est syn. avec Cheilolej. trapezia

(Nees) Kachr. et Schust. — 197. — Descr., Ш. de Drepanolej. pulla (Mitt.) cn.

(= Lej. p.), et diagn. de Drepanolej. subgen. Pristolej. sect, Pullae sect. nov. —

198. — Strepsilej. ocellata Herz., зуп. avec Drepanolej. erecta (Steph.) Mizut.,

est Резр. type de Drepanolej. subgen. Ophthalmolej. (Schust.) c.n. (= Harpalej.

subgen. O.). — 199. — Hygrolej. hebridensis Steph., Cheilolej. samoana Steph.

et Rectolej. samoana Steph. sont зуп. avec Lejeunea cuculliflora (Steph.) Mizut.

— 200. —Lej. exilis (Reinw. et al.) сл. (= Jungermannia e.), avec qui Byssolej.

abnormis Herz. est syn., est l'esp. type de Lejeunea subgen. Byssolej. (Herz.) с.

et stat. п. (= Byssolej. Herz.).

SCHLJAKOV R.N. – De hepaticis in regionibus borealibus URSS inventis

novis et curiosis notula. Nov. Sist. Nizsh. Rast., Inst. Bot. Komarova, Akad.

Nauk SSSR 1980, 17 : 235-240, 3 fig., en russe.

Diagn., descr., ill. de Lophozia (sect. Heteromorphae Schust.) rufescens sp.

nov. de la partie occidentale de Komi RSSA, aff. de L. groenlandica (Nees)

Macoun. Notes taxon. et morph. pour Massula hyperarctica (Schust.) Schljak.

et Nardia japonica St.

TAN B.C. — A new classification for the Genus Physcomitrium B.S.G, J. Hattori

Source - MNHN, Paris

BIBLIOGRAPHIE BRYOLOGIQUE 237

Bot. Lab. 1979, 46 : 327-336, 1 carte, 1 tabl., 12 fig. (Dept. Bot., Univ.

Philippines at College, Los Bafios, Laguna Province, The Philippines).

Observ. morphol. (feuille, calyptre, capsule et stomate, spore) des Physco-

mitrella. Biologie. Hybridation et écologie. Taxonomie de ce genre considéré

comme monospécifique. P. patens (Hedw.) B.S.G. comprend 4 sous-esp. :

subsp. patens, subsp. readeri (C. Müll.) c.n. (= Ephemerella r.), subsp. californica

(Crum et Anderson) c.n. (= Physcom. с.) et subsp. magdalenae (De Sloover) сл.

(= Physcom. m.). Clé et spécimens examinés.

TIXIER P. — La famille des Cololejeuncoideae (Grolle) dans l'Océan Indien

occidental. — Essai monographique. Bull. Trim. Acad. Malgache 1977 (1979),

55, 1-2 :173-247, 45 fig. (Lab. Cryptog., 12 rue Buffon, F-75005 Paris).

Monographie des Cololejeunea (Spruce) Schiffn. et Campylolejeunea Hatt.

dans l'Océan Indien occidental : Madagascar, Seychelles, Comores et Mascarei-

gnes. La plupart des taxons y sont africains, quelques-uns sont asiatiques, peu

sont endémiques. Une cinquantaine d’esp. et var. sont étudiées avec taxonom.,

icon., descr., échantillons examinés, distr. Diagn. des sp. nov. et var. nov. :

Cololejeunea andapania (Madagasc.), C. subinflata (id.), C. saltuum (id.), С.

borbonica (La Réunion), С. lemuriana (Madagasc.), С. tamatavense (id.), С,

adnata (id.), С. succinea (id.), С. onraedtii (Seychelles), С. praeruptorum (Ma-

dagasc.), С. androphylla var. madecassa (id.), С. bandamiae (Comores), С.

duvignaudii var. papillata (Madagasc.), C. camusi (id.), C. mariana (id.), С.

bebourensis (La Réunion), С. crenatiflora var. orientalis (pas d’holotype),

C. bosseriana (La Réunion), C. capuronii (Madagasc.), C. moramangae (id.).

Physocolea decemplicata Steph. et Leptocolea verdoomii Herz. sont transférés

sous le genre Cololejeunea. Clés aux 4 sous-genres de Cololejeunea et aux

esp. des sous-genres.

TOUFFET J. — Clefs pour l'identification préliminaire des Sphaignes armori-

caines. Bot. Rhedonica, Ser. A, 1978-1979, 16 : 17-32, 4 pl. (Lab. Écologie

végét., Univ. Rennes-Beaulieu, 35042 Rennes Cedex, France).

Clés illustrées, basées soit sur des caractéres macroscopiques permettant de

distinguer les sections et d'orienter les récoltes sur le terrain, soit sur des observ.

microscopiques simples permettant l'identification de la plupart des esp. armo-

ricaines.

ZEHR Р.К. — An assessment of variation in Scapania nemorosa and selected

related species (Hepatophyta). Bryophyt. Biblioth. 1980, 15 : 1-140, 125

fig., 14 tabl. (Dept. Bot., Southern Illinois Univ., Carbondale, Illinois 62901

USA).

Morphologie du développement , terpénes, composés methanol, isozymes

(malate déhydrogénase, phosphoglucoisomérase, estérase, anhydrase carboni-

que). La variation des caractéres morphol. est utilisée pour mieux comprendre

les variations de Scapania nemorosa et d'esp. proches telles : S, crassiretis Bryhn,

5. umbrosa (Schrad.) Dum., 5. undulata (L.) Dum., S. subalpina (Nees in Lin-

denb.) Dum., S. obscura (H. Arnell et Jens.) Schiffn., S. serrulata Schust., S.

Source : MNHN, Paris

238 BIBLIOGRAPHIE BRYOLOGIQUE

paludosa (К. Müll.) К. Müll, et 5. uliginosa (Sw. ex Lindenb.) Dum. Il apparait

que les sections Scapania et Nemorosa sont А conserver (chromatographie

des terpènes et morphol. des anthéridies differentes). 5. obscura et 5. serrulata

sont syn. avec 5. alpina et 5. paludosa avec 5. uliginosa. Noter enfin que les

differences dans les terpènes de 5. nemorosa sont dues au substrat et que les

isozymes sont utiles pour differencier les esp. de Scapania.

MORPHOLOGIE, ANATOMIE

ANDERSON L.E. — Cytology and reproductive biology of mosses. In TAYLOR

R.J. and LEVITON А.Е. ed. — The Mosses of North America, San Francisco,

1980. — Voir : Ouvrages généraux : TAYLOR R.J. and LEVITON А.В.

MICHELOT-GERNEZ М.В. — La spermatogenèse chez deux espèces de Bryo-

phytes : le Marchantia polymorpha et le Mnium undulatum; étude ultrastruc-

turale et cytochimique. These Зе cycle, Univ. Pierre et Marie Curie Paris VI,

Paris 1981 ; 1-135, 9 schém., 34 pl.

Historique des recherches sur la spermatogenése - Matériel et techniques -

Gamétogenéses d chez March. polym. et Mnium und.; étude cytochimique de

la différenciation nucléaire au cours de la spermatogenése chez ces 2 esp.i

schéma général de la différenciation des spermatozoides; étude des sécrétions

mucilagineuses des spermatides. — La gamétogenèse dest comparable chez

les 2 esp.; le spermatozoide est typique des Bryophytes. L’évolution nucléaire

s'effectue en 2 temps : une phase de décondensation accompagnée d'une dispa-

rition des RNP et une élimination de protéines vraisemblablement liées antérieu-

rement à l'ADN; une phase de recondensation au cours de laquelle apparaissent

les fibrilles nucléoprotéiques. Par Pisolement de la chromatine et Pobservation

des fibres qui la constituent, il serait possible de vérifier l'existence d'une organi-

sation chromatinienne de type nucléosomale dans les spermatozoides.

ONO $. Studies on vegetative regeneration in the Metzgeriales, Hepaticae.

J. Hattori Bot. Lab. 1979, 46 : 169-191, 12 fig., 6 tabl. (Lab. Biol., Yonezawa

Higashi High School, Yonezawa, Yamagata Pref., 992 Japan).

Étude de la régénération végétative chez 11 esp. de Metzgériales. Cette activité

est liée è la differentiation de la nervure et des ailes du thalle. La régénération

est soit épidermique, soit corticale. Le processus de régénération А partir des

tissus du thalle est dans de nombreux cas très différent des processus de dévelop-

pement des spores ou des gemmules, les protonémas étant uniformément de type

globuleux, sauf pour de rares cas chez Metzgeria où il peuvent étre thalloides.

Comparaison avec les Marchantiales.

PHYSIOLOGIE, CHIMIE

ASAKAWA У., HUNECK S., TOYOTA M., TAKEMOTO Т. and SUIRE C. —

Mono- and Sesquiterpenes from Porella arboris-vitae. J. Hattori Bot. Lab.

Source : MNHN, Paris

BIBLIOGRAPHIE BRYOLOGIQUE 239

1979, 46 : 163-167, 1 fig. (Inst. Pharmacognosy, Tokushima-Bunri Univ.,

Yamashiro-cho, Tokushima, 770 Japan).

Isolement et structure du composé «piquant» : un nouveau sesquiterpéne

hémi-acétal. Mise en évidence de 8 sesquiterpenes déjà connus et detection

d’hydrocarbones mono- et sesquiterpéniques.

BASILIER К. — Nitrogen fixation associated with Sphagnum and some other

musci. Acta Univ, Upsal. 1979, 538 : 1-23, 3 fig., 1 tabl.

Réle de Nostoc et d’autres algues bleues hétérocystes dans la fixation d’azote

par les Sphaignes et autres mousses. Importante fixation chez S. angustifolium,

$. annulatum, S. squarrosum, Drepanocladus aduncus et Calliergon stramineum;

peu de fixation chez S. papillosum, S. fallax, 5. compactum et S. girgensohnii.

Röle des bactéries. Importance écologique.

DELOIRE A. HÉBANT C. and HÉNON J.M. — Vizualization of «external»

conduction in Bryophytes by means of a fluorescent tracer. J. Hattori Bot.

Lab. 1979, 46 : 61-65, 18 fig. (Lab. Paléobot. & Evol. végét., Univ. Sci.

Techn. Languedoc, F-34060 Montpellier Cedex).

Hépatiques et mousses sont irradiées par la lumière bleue. La progression des

solutions de fluorescéine sur les spécimens est observée et photographiée &

travers un filtre jaune-orange.

FLORSCHÜTZ-DE WAARD J. and WORRELL-SCHETS М. — Studies on

colombian Cryptogams. VII. Culture Studies on the taxonomic relevance of

costal anatomy їп the Campylopus leucognodes-subconcolor complex and in

Campylopus pittieri. Proc. Kon. Ned. Akad. Wetensch, Ser. C, Biol. Med.

Sci., 1980, 83, 1 : 37-45, 1 tabl., 4 fig. (Inst. Syst. Bot., Univ. Utrecht, 3584

CS Utrecht, Nederlands).

Relation entre le degré d'humidité de l'habitat et l'épaisseur de la paroi

cellulaire. La présence ou l'absence de pseudostéréides est contrôlée par l'envi-

ronnement. Il s'ensuit que С. leucognodes (С. Mill.) Par. et C. argyrocaulon

(C. Müll.) Mitt. sont syn. avec C. subconcolor (Hamp.) Mitt. En culture, С.

pittieri Williams présente un faible développement des bords apicaux sur la

surface dorsale des nervures. Valeur taxonomique des 2 var. de C. pittieri.

KRZAKOWA M. and SZWEYKOWSKI J. — Isozyme polymorphism in natural

populations of a liverwort, Plagiochila asplenioides. Genetics 1979, 93 :711-

719, 4 tabl., 1 fig. (Dept. Genetics, Adam Mickiewicz Univ., Poznan, Poland).

Les phénotypes apparaissent être gouvernés par 3 locus, chaque locus ayant

2 allèles dont Рип est nul ou muet. Variabilité dans les phénotypes observée

dans et entre les populations. L'implication d'évolution de cette variabilité

serait fonction de Phaplophase persistante et de la diplophase éphémère de cet

organisme.

LONGTON В.Е. — Physiological ecology of mosses. In TAYLOR В.Е. and

LEVITON А.Е. ed. — The mosses of North America. San Francisco, 1980.

— Voir : Ouvrages généraux : TAYLOR R.J. and LEVITON A.E.

Source : MNHN. Paris

240 BIBLIOGRAPHIE BRYOLOGIQUE

TAYLOR J. — Enzymatic isolation of protoplasts from suspension tissue cul-

tures of liverworts. J. Hattori Bot. Lab. 1979, 46 : 55-59, 1 fig. (Dept. Biol.,

Univ. Michigan-Flint, Flint, Michigan 48503 U.S.A.).

Méthode enzymatique pour préparer de bons champs de protoplastes stables,

isolés de cultures de tissus en suspension, d'hépatiques à thalle ou à feuilles.

RÉPARTITION, ÉCOLOGIE, SOCIOLOGIE

ABRAMOV 11., ABRAMOVA A.L., BLAGODATSKICH L.S. — Ad bryofloram

partis boreali-orientalis URSS. Nov. Sist. Nizsh. Rast., Inst. Bot. Komarova,

Akad. Nauk SSSR 1980, 17 : 201-204, en russe.

21 mousses avec loc.

ABRAMOVA A.L., ABRAMOV 11. — Materies ad examinationem bryoflorae

paeninsulae Czukotka. Nov. Sist. Nizsh. Rast., Inst. Bot. Komarova, Akad.

Nauk SSSR 1980, 17 : 179-200, 6 fig., en russe.

71 esp. avec loc. Descr, ill. des esp. décrites par C. Müller.

ABRAMOVA A.L., AFONINA O.M., DUDA J. — Ad floram hepaticarum pae-

ninsulae Czukotka. III. Nov. Sist. Nizsh. Rast., Inst. Bot. Komarova, Akad

Nauk SSSR 1980, 17 : 204-211, en russe.

71 esp. et 25 var. d'hépatiques avec loc. Nouveautés pour la péninsule.

AFONINA О.М. — Addimenta ad bryofloram paeninsulae Czukotka. 1. Nov.

Sist. Nizsh. Rast., Inst. Bot. Komarova, Akad. Nauk SSSR 1980, 17 : 241-

244, en russe.

29 taxa avec loc. dont cert. sont nouv. pour la péninsule.

AFONINA O.M., DUDA J.— Ad йогат hepaticarum in flexu medio fl. Indi-

girka vigentium. Nov. Sist. Nizsh. Rast., Inst. Bot. Komarova, Akad. Nauk

SSSR 1980, 17 : 211-215, en russe.

43 hépatiques avec loc.

ANDO Н. — A phytogeographical account of Hypnum cupressiforme Hedw. in

Japan. Bull. Yokohama Phytosociol. Soc. Jap. 1979, 16 : 339-348, 13 fig.

(Bot. Inst., Hiroshima Univ., Japan).

Distr, écol. de Hyphum cupressiforme au Japon. Comparaison avec distr.

des autres esp. du genre. Fertilité, variabilité (var. cupressiforme et var. fili-

forme).

DE ZUTTERE Ph. Quelques bryophytes nouvelles ou rares pour la Breta-

gne. Deux espéces nouvelles pour la France. Bot. Rhedonica, Ser. A, 1978-

1979, 16 : 3341, 2 fig, 1 саме (16 rue du Bois, B-1430 Wauthier-Braine).

Hépatiques et mousses avec hab. et loc, Noter Gymnomitrion crenulatum

Source - MNHN. Paris

BIBLIOGRAPHIE BRYOLOGIQUE 241

Gott. ex Carringt. et Bryum tenuisetum Limpr. nouv. pour la France.

FENTON J.H.C.— The rate of peat accumulation in Antarctic moss banks.

J. Ecol. 1980, 68, 1 : 211-228, 11 tabl,, 8 fig. (Brit. Antarct. Surv., Madingley

Road, Cambridge, СВЗ ОЕТ, Great Britain).

Etude de la décomposition dans trois types de bancs de mousses. 2 sont

dominés par Polytrichum alpestre Hoppe et un par Chorisodontium aciphyllum

(Hook. f. et Wils.) Broth.

РВАНМ J.P. — Ergänzungen zur Campylopus-Flora von Mexico. Herzogia 1979,

5 : 111-117, 1 fig. (Gesamthochschule Duisburg, Fachbereich VI, Biologie,

D-41 Duisburg).

7 esp. de Campylopus (avec loc. et notes) nouv. pour le Mexique.

FRAHM J.P. — Verbreitungskarten von Moosen in Deutschland I. Amblyodon,

Catoscopium, Meesia. Herzogia 1979, 5 : 119-161, 2 tabl., 8 fig., 6 cartes

(Idem).

Cartographie en Allemagne (sur la base de spécimens d’herbier et de la litté-

rature) d'Amblyodon dealbatus (Dicks.) Р. Beauv., de Catoscopium nigritum

(Hedw.) Brid., de Meesia hexasticha (Funck) Bruch, de М. longiseta Hedw., de

М. triquetra (Hook. et Tayl.) Angstr., et de М. uliginosa Hedw. Mise en évidence

d'un nowy. hybride : Meesia triquetra X uliginosa.

FRAHM J.P. — Zum Vorkommen von Rhodobryum ontariense (Kindb.) Kindb.

in Mitteleuropa. Herzogia 1979, 5 : 163-169, 5 fig. (Idem).

Caractères permettant de distinguer R. roseum de В. ontariense. Distr. et

hab. des 2 esp. R. ontariense est présent en Allemagne, Autriche, Suisse, Tché-

coslovaquie, Italie, Yougoslavie, Pologne, France et Union soviétique,

GUERRA J. — Nota briologica I. Trab. Monogr., Dept. Bot Malaga 1980, 1 :

29-36, 3 fig. (Dept. Bot., Fac. Ci., Univ. Malaga, Езрайа).

Distr. et notes écol. pour Isothecium algarvicum Dix. et Nicholson, Ortho-

thecium durieui (Mont.) Besch. et Fissidens rufulus (Br. et Schimp.) B.S.G.

KARCZMARZ K. — Mszaki Pasma Bukowicy w Beskidzie Wiskim - Bryophytes

of the Bukowice Range in Low Beskids (Polish Western Carpathians). Fragm.

Florist. Geobot. 1979, 25, 1 : 191-206 (Dept. Pl. Taxonom. and Geogr.,

Inst. Biol., Maria-Curie-Sklodowska Univ., Lublin, Poland).

Données écologiques. Liste de 95 mousses et 20 hépatiques récoltées par

ГА. en 1975-76, dans cette région jamais explorée par les bryologues.

MIHAI Gh. — Rhynchostegiella jacquinii (Garov.) Limpr. în brioflora Moldavei

si raspindirca ei pe teritoriul Romäniei. Lucr. Stat. «Stejarul», Ser. Ecol.

Terrestr. Genet. 1976-1977 : 15-18, 1 fig., 1 tabl.

Clé aux Rhynchostegiella tenella (Dicks.) Limpr., В. jacquinii (Garov.)

Limpr., R. teesdalei (Sm.) Limpr. R. jacquinii est попу. pour la Moldavie (distr.

de Jassy).

Source : MNHN, Paris

242 BIBLIOGRAPHIE BRYOLOGIQUE

OCHYRA R., TOMASZEWICZ Н. — Nowe Stanowiska Ricciocarpus natans

(L.) Corda (Ricciaceae, Hepaticopsida) i przeglad jego rozmieszczenia w

Polsce. — New localities of Ricciocarpus natans (L.) Corda (Ricciaceae, Hepa-

ticopsida) and a review of its distribution in Poland. Fragm. Florist. Geobot.

1979, 25, 3 : 429-438, З tabl., 1 fig. (Lab. Bryol. Lichenol., Inst. Bot., Polish

Acad. Sci., 31-512 Krakow, Poland).

Écol. et position syntaxonom. de R. natans. 8 nouv. loc. en Pologne. Distr.

en Pologne.

PHILIPPI С. — Moosflora und Moosvegetation des Bruchswaldes bei Grenzach-

Wyhlen. Natur- u. Landschaftsschutzgebiete Bad.-Wiirttemberg. 1979, 9 :113-

116, 11 tabl., 7 fig. (Landessammlungen f, Naturkunde, D-7500 Karlsruhe).

Vegetation bryophyt. des rochers (8 groupements), des endroits humides,

associations terricoles (9), et épiphytes (11). Liste des esp. d’hépatiques et de

mousses avec écol.

PIERROT В.В. — Bryophytes des dunes du littoral charentais. Bull. Soc. Bot.

Centre-Ouest n.s., 1980, по sp. 4 : 101-111 (Les Andryales, St André, F-

17550 Dolus).

Réle des bryophytes dans la fixation de la dune. Caractère de xérophilie et

de thermophilie. La bryoflore dunaire apparaît pauvre en espèces.

RAIMONDO F.M. e DIA M.G. — Note briogeografiche П. Cryphaea heteromalla

(Hedw.) Mohr in Sicilia, Atti Ассай. Sci. Lett. Arti Palermo, Ser. IV, 1978"

1979 (1979), 38, 1 : 3-18, 1 tabl., 4 fig. (Ist. ed Orti Bot., Univ., 90123

Palermo, Italia).

Distr. en Italie de С. heteromalla (Hedw.) Mohr, mousse épiphyte, atlantico-

méditerranéenne, récolté en Sicile pres d’Erice. Ròle de cette esp. dans les

unités syntaxonomiques connues.

RODRIGUEZ CABRERA М. у RON ALVAREZ М.Е. — Contribucion al conoci-

miento briologico del Barranco del Agua (Guimar) Teneriffe. Vieraea 1979

(1980), 9, 1-2 :49-56 (Dept. Bot., Fac. Biol., Univ. La Laguna, Tenerife, Islas

Canarias).

Liste de 47 taxons avec loc.

ROGEON М. — Inventaire des récoltes bryologiques effectuées par Н. Bouby en

Haute-Vienne dans le but d’esquisser une étude des groupements muscinaux

de la région de Rochechouart. In BOUBY H., Matériaux pour une étude

floristique et phytosociologique du Limousin occidental - Forét de Roche-

chouart et secteurs limitrophes (Haute-Vienne). Bull. Soc. Bot. Centre-Ouest

n.s., 1978, по sp. 2 : 105-118, phot. 15.

Liste de 152 taxons avec loc.; noter nouveautés pour la Vienne.

SCHOFIELD W.B. and GODFREY G.A. — Radula brunnea Steph. in Western

North America. J. Hattori Bot. Lab. 1979, 46 : 285-288, 1 fig. (Dept. Bot,

Source - MNHN. Paris

BIBLIOGRAPHIE BRYOLOGIQUE 243

Univ. British Columbia, Vancouver, B.C., Canada V6T 1W5).

Découverte dans P'Orégon NW de Radula brunnea Steph., trouvé jusque la

en Asie E seulement.

SCHOFIELD W.B. — Phytogeography of the mosses of North America. In TAY-

ТОК RJ. and LEVITON A.E. ed. — The Mosses of North America. San

Francisco, 1980. - Voir : Ouvrages généraux : TAYLOR R.J. and LEVITON

AE.

VJUNOVA G.V. — Materies ad examinationem bryoflorae regionis Leningraden-

sis. Nov. Sist. Nizsh. Rast., Inst. Bot. Komarova, Akad. Nauk SSSR 1980,

17 : 216-230, en russe.

Liste avec ref. bibliogr. de 343 esp.

ZHUKOVA A.L., PREIS Ju.I. — Ad floram hepaticarum in systemate fl. Bureja

(regio Amurensis) notula. Nov. Sist. Nizsh. Rast., Inst. Bot. Komarova, Akad.

Nauk SSSR 1980, 17 :231-234, fig., en russe.

PALÉOBRYOLOGIE

MILLER М.С. — Fossil mosses of North America and their significance. In

TAYLOR RJ. and LEVITON А.Е. ed. — The mosses of North America,

San Francisco, 1980. - Voir : Ouvrages généraux : TAYLOR R.J. and LE-

VITON А.Е.

MILLER М.С. — Mosses as paleoecological indicators of lateglacial terrestrial

environments : some North America studies. Bull. Torrey Bot. Club 1980,

107, 3 : 373-391, 5 fig. 6 tabl. (Harvard Univ. Herb., 22 Divinity Ave.,

Cambridge, Massachusetts 02138 USA).

La presence de mousses fossiles implique que les sites calcaires et plantes

calcicoles fussent dominants dans les prairies lors des dernières glaciations.

OUVRAGES GENERAUX

CATCHESIDE D.G. — Mosses of South Australia. Woolman, Netley (South

Australia) 1980, 364 p., 209 fig., 16 pl. coul.

Généralités - clé générale aux genres et esp. - classification - divisions floris-

tiques de l'Australie du $ - traitement systématique de 200 taxons environ;

pour chacun : nomenclature, descr., ill., distr., affinités. Nombreuses clés,

dessins clairs, pl. coul, glossaire détaillé, bibliogr., index font de cette flore

un outil agréable. Esp. nouv. : Fissidens hebetatus (North-Territory), Е. subhu-

milis (idem), Pottia latzi (idem), Р. globosa (South Australia), Crossidium

davidai (idem), Physcomitrium phymatodeum (idem). Comb. nouv. : Didymo-

don torquatus (Tayl.) (= Barbula t.), D. subtorquatus (С. Müll. et Hampe)

Source : MNHN, Paris

244 BIBLIOGRAPHIE BRYOLOGIQUE

(= B. =), D. luehmanni (Broth. et Geh.) (= B. l), Weissia willisiana (Sainsb.)

= Pottia w.), Bartramia hampei (Mitt.) (= Bartramidula h.). Noter la descr.

des échantillons douteux, mais sans épithéte nouvelle.

СВОМ H.A. and ANDERSON L.E. — Mosses of Eastern North America. New

York, Columbia Univ. Press, 1981, 2 vol., 1328 p., 637 fig.

Flore des mousses de l'est de l'Amérique du Nord, soit l'aire occupée par les

arbres à feuilles caduques de l'Est; sens écologique et phytogéographique de

cette forêt. Dans les grandes lignes, la définition des genres et l’arrangement

des familles suivent BROTHERUS in ENGLER und PRANTL. Cette flore, du

point de vue taxonomique, ne cherche pas á étre une solution universelle, mais

une proposition basée sur la réflexion et l'expérience des auteurs. Conspectus

avec descr. des classes, sous-classes, ordres, familles. Traitement taxonomique,

Chaque taxon supra-générique est décrit. Clés aux différentes subdivisions

supra-spécifiques et aux esp. Pour chaque taxon : synon., descr., hab., distr.,

ill. Ce travail est complété par un glossaire (15 p.), une bibliogr. (14 р.) et un

index (14 p.). Les nombreux dessins et la présentation clairs font de cet ouvrage

un outil de référence agréable & consulter.

IRELAND R.R., BIRD C.D., BRASSARD G.R., SCHOFIELD W.B. and VITT

D.H. — Checklist of the Mosses of Canada. Natl. Mus. Canada, Ottawa 1980,

Publ. Bot. 8 :ixii, 1-75, ill.

Liste de tous les noms de mousses publiés du Canada. 966 esp. et 401 taxons

infraspécifiques sont reconnus. Liste des syn. et des taxons exclus.

LANDWEHR J.- Atlas Nederlandse Levermossen (met Medewerking van

S.R. GRADSTEIN en Н. van MELICK). Zutphen, Thieme et Cie, 1980,

287 p., 119 pl. h.t. en coul. et en noir.

Structure des hépatiques sous forme de glossaire détaillé et illustré. Habitat

des hepatiques avec hors-texte en noir. Classification des esp. néerlandaises

selon Schuster 1966 et Grolle 1976. Étude systématique: pour chaque taxon :

nom latin, nom vernaculaire, courte descr., ill. (légendes des pl. en hollandais,

anglais, allemand et francais), caractéristiques écol. Index des noms latins, des

noms vernaculaires. Liste du matériel utilisé.

SJÓDIN А. - Index to distribution maps of bryophytes 1887-1975. II. Hepati-

cae. Växtekol. Stud. 1980, 12 : 1-143 (Växtbiologiska Inst., Box 559, S-751

22 Uppsala).

Liste alphabétique des genres et esp. (ssp., var. et f.) avec les références des

cartes s'y rapportant. Les références sont données en ordre géographique, ce

qui pour certains taxons ne facilite pas la compréhension ou une recherche

rapide. Cet index est complété par la bibliographie correspondante. 3300 cartes

de distribution ont été inventoriées.

TAYLOR R.J. and LEVITON А.Е. ed. — The mosses of North America. An

inquiry into the biology of mosses based upon a symposium sponsored by

Source : MNHN. Paris

BIBLIOGRAPHIE BRYOLOGIQUE 245

the Pacific Section, Botanical Society of America. San Francisco, Pacific

Division, Amer. Assoc. Advanc. Sci. 1980, 170 p., ill.

TAYLOR R.J. — Préface. - WAGNER D.H, — Introduction (Recherches

bryol. en Amérique du Nord). — MILLER М.С. — Fossil mosses of North Ame-

rica and their significance (: 9-36, 8 fig., 9 tabl.). Les fossiles proviennent peu du

mésozoïque, surtout du Tertiaire; noter toutefois la présence de spores sembla-

bles à celles de Sphagnum et de certaines Bryideae dans le Crétacé. 162 esp.

(14% de la flore actuelle) ont été déterminées du Pleistocène et de l’Holocène.

Importance du site des Grands Lacs. Affinités entre un assemblage de mousses

fossiles du Vermont et de la flore de la Péninsule de Gespé. Utilisation de ces

recherches au niveau phytogéographique. Comb. nouv. proposées : Muscites

coloradensis (Brown) (= Нурпит c.), M. montanensis (Brown) (= Mnium m.),

Plagiopodopsis eocenica (Кис) (= Muscites e.). Paleohypnum arnoldianum

Steere, Р. beckeri Steere, P. brittoniae Steere, P. jovetastii Кис, Р. steerei Kuc,

Hypnum brownii Kirchner, Rhynchostegium knowltonii E, Britton et Archeom-

nium patens Britt. in Knowlt. sont transférés sous le genre Hypnites Ettingshau-

sen. — ANDERSON L.E. — Cytology and reproductive biology of mosses (: 37-

76, 64 fig., 3 tabl.). Nombre chromosomique en relation avec les groupements

taxonomiques. Polyploidies. Hétérochromatine. m-chromosomes. Bivalents

hétéromorphiques. Rôle des chromosomes dans la méiose et la mitose. Repro-

duction sexuée et états sexuels principaux en relation avec la persistance, la

survie et l'évolution. Hybridation interspécifique. Méthodes de reproduction

asexuée et dispersion des spores en relation avec les structures des populations.

— LONGTON В.Е. — Physiological ecology of mosses (: 77-113). Relations

hydriques. Réponses aux stress (désiccation, froid, chaleur). Nutrition minérale

(röle des mousses comme indicateurs des conditions du sol, de la toxicité des

minéraux et de la pollution). Microclimat. Croissance et phénologie de la repro-

duction. Écologie (photosynthèse). Importante bibliogr. — CROSBY М.В. — The

diversity and relationships of mosses (: 115-129, 2 tabl.). Les mousses et les

plantes vasculaires ont probablement un ancétre commun. L’ancétre des mousses

serait proche des Andreaeobryum. Les Bryideae sont rediscutés et reclassifiés en

10 ordres selon les caractéristiques du péristome. — SCHOFIELD W.B. — Phy-

togeography of the mosses of North America (North of Mexico) (: 131-170, 45

cartes). Les mousses de l'Amérique du Nord peuvent être classées selon au moins

20 modes de distribution géographique. Les Pseudoditrichaceae Steere et Iwats.

sont la seule fam. restreinte a cette région; 13 genres sont endémiques dont

10 principalement à l'ouest des Monts Cordilléres. 23% des mousses ne sont

connues que de l'Amérique du Nord et sont réparties selon 10 modes de distr.

Les esp. confinées à l'hémisphére W suivent 3 modes de distr., les disjointes,

10 modes. Interpretation.

VARIA

ABRAMOV LL, ABRAMOVA A.L. — Ad historiam examinationis bryoflorae

URSS поша. Nov. Sist. Nizsh. Rast., Inst. Bot. Komarova, Akad. Nauk

Source : MNHN, Paris

246 BIBLIOGRAPHIE BRYOLOGIQUE

SSSR 1980, 17 : 167-174, en russe.

BAUDOIN R., BISCHLER H., JOVET-AST S. — Exemple d’exploitation infor-

matique des données sur la végétation hépaticologique méditerranéenne.

Actes 70 Colloque Informatique et Biosphére 1979 . 127-142, 4 fig. 5 tabl.

(Lab. Cryptog., 12 rue Buffon, F-75005 Paris).

BRISSE H. et GRANDJOUAN G. — Utilisation de l'ordinateur pour la consti-

tution d’un code floristique (exemple des Bryophytes de la France). Actes

70 Colloque Informatique et Biosphère 1979 : 71-85, 5 fig. (Lab. Morphol.

Exper., Inst. Bot., 28 rue Goethe, F-67083 Strasbourg Cedex).

DULL R. — Moosforschung in Hessen. Hessische Florist. Briefe 1979, 28, 4 :

54-65 (Gesamthochschule Duisburg).

POCS T. — Bryotaxonomical and chorological research in Hungary. Abstr. Bot

(Budapest) 1978 (1979), 5, suppl. 3 (Confer. Bryol. 1978 in Eger) : 43-45

(Res. Inst. Bot., Hungar. Acad. Sci., Vacratot, Hungary).

Activités, orientation de la recherche bryol. en Hongrie.

SZMAJDA P. — Investigations on mosses at Adam Mickiewicz University in

Poznan, Poland. Abstr: Bot. (Budapest) 1978 (1979), 5, suppl. 3 (Confer.

Bryol. 1978 in Eger) : 15-31, 11 fig. (Dept. Geobotany, Adam Mickiewicz

Univ., 61-713 Poznan, Poland).

Historique depuis le Prof. 2. CZUBINSKI (1912-1967), Quelques exemples

de distrib. de mousses en Pologne.

VANA J. — Bryologie in der CSSR. Abstr. Bot. 1978 (1979), 5, suppl. 3 (Con-

fer. Bryol. 1978 in Eger) : 7-13 (Bot. Inst., Karls-Univ., Praha, Czechoslova-

kia).

Source : MNHN. Paris

247

BIBLIOGRAPHIE LICHENOLOGIQUE

D.LAMY

SYSTEMATIQUE, NOMENCLATURE

AWASTHI D.D. and AKHTAR P. — The lichen genus Leptogium (Sects, Lepto-

gium, Leptogiopsis and Homodium) in India. Geophytology 1979, 8 (2) :

189-204, 2 pl., 7 fig. (Dept. Bot., Lucknow Univ., Lucknow, India).

Clé aux 24 esp. de Leptogium représentées en Inde. Taxonom., spécimens

examinés, descr. de 19 d’entre elles. Diagn., descr., ill. de Leptogium indicum

sp. nov. affine de L, azureum (Sw.) Mont. et L. tremelloides (L, f.) S. Gray.

FOLLMANN С. und SANCHEZ-PINTO L, — Zur Kenntnis der Flechtenflora

und Flechtenvegetation der Kanarischen Inseln. III. Ein neue Strauchflechte

aus dem Verwandtschaftskreis von Ramalina fraxinea (L.) Ach. Philippia

1980, 4, 3 : 193-200, 5 fig. (Naturkundesmus. Ottoneum, D-3500 Kassel 1).

Diagn., descr., ill. de Ramalina superfraxinea, esp. nouv. de Tenerife, aff. de

R. fraxinea. Esp. à larges ascocarpes et contenant beaucoup d’ac. divaricatique

accumulé dans la médulle. Elle appartient au Ramalinetum subgeniculatae

Klem.

HAWKSWORTH D.L. — Notes on British lichenicolous fungi : Ш. Notes Roy.

Bot. Gard. Edinburgh 1980, 38, 1 : 165-183 (СМІ, Ferry Lane, Kew, Surrey,

TW9 3AF, Great Britain).

Diagn., descr., ill. de Clypeococcum hypocenomyceae sp. nov. sur Нуросе-

потусе scalaris, et de Ranularia peltigericola sp. nov. sur Peltigera polydactyla.

Comb. nouv. : Spilomela ascaridiella (Nyl.) (= Lecidea a.), Thamnogalla crom-

biei (Mudd) (= Endocarpon с.) et Trichothyrina cetrariicola (Nyl.) (= Sphaeria

с.). 2 esp. sont пошу. pour les Iles Britanniques. Arthonia punctella Nyl. est

distinct de A. clemens (Tul.) Th. Fr. et Microthelia dissepta (Nyl.) Al. Sm.

est probablement syn. avec Pleospora peripherica (Tayl.) D. Hawksworth.

SINGH К.Р. — Awasthiella, а new lichen genus from Manipur. Norweg. J. Bot.

1980, 27, 1 :33-35, 8 fig. (Bot. Surv. India, Howrah-711103, India),

Diagn., descr., ill. de Awasthiella indica gen. et sp. nov. des Verrucariaceae.

Périthèces noirs contenant des cellules algales hyméniales plus petites que celles

du thalle; spores simples.

Source : MNHN, Paris

248 BIBLIOGRAPHIE LICHENOLOGIQUE

PHYSIOLOGIE, CHIMIE

AHMADJIAN V., RUSSELL L.A. and HILDRETH K.C. — Artificial reestablish-

ment of Lichens. I. Morphological interactions between the phycobionts of

different lichens and the mycobionts Cladonia cristatella and Lecanora

chrysoleuca. Mycologia 1980, 72 : 73-89, 16 fig. (Dept. Biol., Clark Univ.,

Worcester, Massachusetts 01610 USA).

Obtention de nouvelles synthéses entre un mycobionte de Cladonia cristatella

et 13 isolats du phycobionte Trebouxia, échec avec 10 isolats de Pseudotre-

bouxia et avec Pleurastrum terrestre, algue libre. Le mycobionte de Lecanora

chrysoleuca forme des sorédies avec 5 isolats de Pseudotreb. et avec 1 isolat de

Treb. Les squamules synthétisées ont une morphol. identique à celle des squa-

mules naturelles de Clad. crist. (produisant des pycnidies müres, de courts po-

détions avec des apothécies immatures). Observation de corps concentriques

dans les hyphes de Clad. crist. synthétisé avec les phycobiontes de Clad. Богу?

et de Lecidea tumida.

BECKER V.E. — Nitrogen fixing lichens in forests of the Southern Appalachian

Mountains of North Carolina. Bryologist 1980, 83, 1 : 29-39, 2 fig., 6 tabl.

(Dept. Biol., Wake Forest Univ., Winston-Salem, МС 27109 USA).

Aesculus octandra Marshal apparaît comme l'habitat optimum particuliére-

ment pour Lobaria quercizans Mich. et Lobaria pulmonaria (L.) Hoffm. La

biomasse des lichens est augmentée dans les foréts de hétres si Aesculus octandra

est present. Lichens moins abondants dans les chénaies et les foréts abritées,

pratiquement inexistants chez les conifères.

CULBERSON С.Е. and AHMADJIAN У. — Artificial reestablishment of lichens.

П. Secondary products of resynthetized Cladonia cristatella and Lecanora

chrysoleuca. Mycologia 1980, 72 : 90-109, 12 fig. 1 tabl. (Dept. Bot., Duke

Univ., Durham, North Carolina 27706 USA).

Les plants obtenus par la synthèse entre le mycobionte de Clad. crist. et les

phycobiontes Trebouxia produisent de l'acide barbatique comme produit

secondaire majeur, et aussi pour la plupart de l'acide dibenzofurane didymique,

mais pas d'acide usnique. Les sorédies formées avec le Lec. chrys. et le Pseudo-

trebouxia montrent des traces d’ac. usnique. La formation de nombreux pro-

duits caractéristiques doit impliquer Pinhibition algale d'enzymes fongiques.

HUNECK S., SCHREIBER К. and SUNDHOLM С. — Ovosáure, eine neues

Tridepsid aus der Flechte Parmelia substygia. Phytochemistry 1980, 19, ph

885-887, 1 tabl. (Inst. Biochem. Pflanz. Forschungszentr. Molekul. Med.,

Akad. Wissensch. DDR, DDR-401 Halle/Saale, Weinberg).

Structure de l'acide ovoique, nouveau tridepside isolé chez Parmelia substy-

gia Каз. : acide 2’-0-méthylgyrophorique.

LANGE O.L. — Moisture content and СО; exchange of Lichens. I. Influence of

temperature on moisture-dependent net photosynthesis and dark respiration

Source : MNHN. Paris

BIBLIOGRAPHIE LICHENOLOGIQUE 249

in Ramalina maciformis. Oecologia 1980, 45, 1 : 82-87, 7 fig., 1 tabl. (Lehrst.

f. Bot. П, Univ. Würzburg, D-8700 Würzburg, FRG).

SCHAPIRO Г.А. — Conspectus chemotaxonomiae lichenum in statu praesenti.

Nov. Sist. Nizsh. Rast., Inst. Bot. Komarova, Akad. Nauk SSSR 1980, 17 :

152-166, 3 fig., en russe.

SIGFRIDSSON В. — Some effects of humidity on the light reaction of photo-

synthesis in the lichens Cladonia impexa and Collema flaccidum. Physiol. PI.

(Copenhagen) 1980, 49, 3 : 320-326, 9 fig., 1 tabl. (Univ. Lund, Dept. Pl.

Physiol., Вох 7007, S-220 07 Lund 7).

Mesures de la luminescence et de la fluorescence А différentes humidites

relatives. Répercussion sur la photosynthese.

TSCHERMAK-WOESS E.— Ueber Plastidenstapel bei Botrydiopsis alpina

sowie Anlage und Vermehrung der Stigmen bei dieser und Heterococcus

(Xanthophyceae). Pl. Syst. Evol. 1979, 131 : 179-192, 6 fig. (Inst. Bot.,

Univ. Wien, Rennweg 14, A-1030 Wien).

Sous certaines conditions il y a formation d'amas de chloroplastes chez

Botrydiopsis alpina Vischer (Botrydiopsis est le phycobionte de Lobaria linita

(Ach.) Rabenh.). Formation et multiplication de stigmates chez Botrydiopsis

et Heterococcus (phycobionte de Verrucaria).

RÉPARTITION, ÉCOLOGIE, SOCIOLOGIE

AFONINA O.M., BREDKINA LI. MAKAROVA LI. — Musci lichenesque

zonae silvo-stepposae in fluxu medio Fl. Indigirka. Nov. Sist. Nizsh. Rast.,

Inst. Bot. Komarova, Akad. Nauk SSSR 1979, 16 : 175-186, en russe.

52 mousses et 96 lichens avec loc.

ALSTRUP V. — Notes on selected Greenlandia Lichens. Bot. Tidsskr. 1979, 74,

2-3 : 155-163 (Inst. Pl. Ecol., Univ. Copenhagen, DK-1353 Copenhagen К).

Liste avec loc. et notes de 57 lichens du sud et de l'ouest du Groenland. 18

taxa sont пошу. pour le Groenland. Lecanora glaucocarpa var. endocarpoides

Vainio est syn. avec Lecidea rhizobola Nyl. Noter aussi que les Physcia grisea

précédemment décrits du Groenland sont en fait des Physconia detersa.

AWASTHI D.D. and SINGH K.P. — Observations on some Graphidaceous lichen

taxa. Phyta s.d., 1 : 34-40, 2 fig., 1 pl. (Idem).

Taxonom., descr., ill. et distr. de 8 esp. de Graphis d'Inde et du Sri Lanka.

AWASTHI D.D. and SINGH S.R, — The lichen genus Rhizocarpon (Ram.) Th.

Fr. in India. Kavaka 1977, 5 : 49-58, 28 fig. (Dept. Bot., Lucknow Univ.,

Lucknow, India).

Descr., ill., distr. de 11 esp. de Rhizocarpon connues en Inde. К. alpicola

Source : MNHN, Paris

250 BIBLIOGRAPHIE LICHENOLOGIQUE

(Hepp) Rabh., R. lindsayanum Räs., R. macrosporum Räs., R. sphaerosporum

Ris, et R. viridiatrum (Wulf.) Korb. sont nouv. pour l'Inde.

BOISSIÈRE J.C. — Quelqueslichens. In BOUBY H., Matériaux pour une étude

floristique et phytosociologique du Limousin occidental - Foret de Roche-

chouart et secteurs limitrophes (Haute-Vienne). Bull. Soc. Bot. Centre-

Ouest, n.s., 1978, по sp. 2 : 119-122.

Lichens récoltés dans les 18 localités étudiées.

BOTINEAU М. et HOUMEAU J.M. — Contribution à l'étude des lichens des

dunes côtières de Charente-Maritime. Bull. Soc. Bot. Centre-Ouest, nus.

1980, по sp. 4 : 83-99, 9 phot. (La Clef d'Or, 16410 Dignac).

Relevé de 70 esp. de lichens dont cert. sont rares pour la région. Présence

desp. méditerranéo-atlantiques comme Cladonia mediterranea et Ramalina

canariensis.

BREDKINA 11. — Toninia groenlandica Lynge em. Bredk. in locis silvi-step-

posis SSSA Jacutiae. Nov. Sist. Nizsh. Rast., Inst. Bot. Komarova, Akad.

Nauk SSSR 1980, 17 : 109-111, 1 tabl., en russe.

Descr., comparaison avec T. tristis (Th. Fr.) Th. Fr. et T. tabacina (Massal.)

Flag., et loc. de T. groenlandica.

BREDKINA LJ. — Lichenes montium Tjan-Schan Centralis pro URSS novi et

curiosi. Nov. Sist. Nizsh. Rast., Inst. Bot. Komarova, Akad. Nauk SSR 1980,

17 :111-116, en russe.

4 taxa avec descr. et loc.

CHRISTIANSEN M.S., RAMKAE K., ROSE Е. and SOCHTING U. — Additions

to the Danish Lichen Flora. Bot. Tidsskr. 1979, 74, 2-3 : 89-115, 18 fig,

5 tabl. (Bot. Mus. & Centralbibl., DK-1123 Kobenhavn К).

Descr., hab., distr. de 30 esp. de lichens nouv. pour le Danemark. Clé aux

Alectoria et Bryoria danois et clé aux Micarea danois.

CRESPO A., BARRENO Е. y VASQUEZ V.M. — Buellia subcanescens (Physcia-

ceae) en el norte de España. Lazaroa 1979, 1 : 139-141, 1 fig. (Dept. Bot

Fac. Farmacia, Univ. Complutense, Madrid 3, España).

B. subcanescens R.G. Werner et Pertusaria gallica В. de Lesd. nouv. pour le

nord de l'Espagne : Cabo de Peñas (Asturies).

CRESPO A., BARRENO Е. RICO V.J. y BUENO A.G. — Catalogo liquenico del

desierto de Calanda (Terruel, España). 1. Anales Jard. Bof. Madrid 1979

(1980), 36 : 43-55, 1 fig., 1 tabl. (Idem).

Avant Vinstallation d'une centrale thermique dans le désert de Calanda, les

AA. ont fait l’inventaire des 70 taxa épiphytes et terricoles qui permettra de

suivre Pinfluence de la pollution sur les lichens. 12 taxa sont nouv. pour PEspa-

gne. Comb. nouv. : Bacidia microbola (Clemente) Crespo et Barreno (= Lecidea

т.).

Source : MNHN, Paris

BIBLIOGRAPHIE LICHENOLOGIQUE 251

GALLE L. — Wirkung der Luftverunreinigung auf die Verarmung der Flechten-

vegetation der Stadt Szeged und ihre Umgebung. Acta Biol., Acta Univ.

Szeged, n.s., 1979, 25, 1-2 : 3-15, 2 tabl., 1 fig. (Н-6722 Szeged, Petofi

Sandor sgt. 6.1.7).

Bioécologie dans les ass. : le Caloplacetum тиготит, le Lecanoretum albo-

marginatae et le Caloplacetum citrinae.

GOLUBKOVA N.S. — Species Acarosporae Massal. Xanthochroae in Mongolia

Inventae. Nov. Sist. Nizsh. Rast., Inst. Bot. Komarova, Akad. Nauk SSSR

1980, 17 : 120-134, en russe.

Le sousgenre Xanthothallia Massal. est représenté par 9 esp. en Mongolie.

Taxonomie et descr, Répartition dans le monde.

INASCHVILI Tz. N. — Species familiae Collemataceae pro URSS novae et

rarae. Nov. Sist. Nizsh. Rast., Inst, Bot. Komarova, Akad. Nauk SSSR 1980,

17 : 134-136, en russe.

Distr., notes morphol, pour Collema japonicum (Mill. Arg.) Hue, С. subcon-

veniens Nyl., Leptogium burnetiae Dodge et L. corticola (Tayl.) Tuck.

KOROLEV Ju. B., TOLPYSCHEVA Т. Ju. — De Lichenoflora stationarii «Con-

tact» (Planities elata verchnekolymskoje dicta) notula. Nov. Sist. Nizsh. Rast.,

Inst. Bot. Komarova, Akad. Nauk SSSR 1980, 17 : 137-150, 2 tabl., en russe.

108 taxa avec loc., notes biogéogr.

LAFLAMME-LEVESQUE M., PERRON J.M. et JOBIN L. — Etude des lichens

appartenant aux genres Bryoria, Alectoria et Ramalina dans les foyers d’in-

festation de Lambdina fiscellaria fiscellaria (Guén.) A Vile d'Anticosti. Natu-

ral. Canad, 1979, 106, 5-6 : 505-510, 5 fig., 1 tabl. (Dept. Biol., Fac. Sci. &

Génie, Univ. Laval, Québec G1K 7Р4).

Distrib. de 9 Bryoria, 1 Alectoria et 4 Ramalina et 3 Usnea dans les peuple-

ments forestiers de l'ile d’Anticosti (Golfe du Saint-Laurent). La densité liché-

nique est mise en relation avec l'áge et la densité des peuplements forestiers.

Les foréts susceptibles d'étre atteintes par les pullulements de Lambdina fiscella-

ria fiscellaria (insecte défoliateur) seraient celles qui auraient atteint leur maturi-

té et qui auraient des lichens corticoles en abondance. Noter Bryoria salazinica

Brodo et D. Hawksw. nouveau pour le Québec.

LEROND М. Les lichens épiphytes en Normandie orientale. Distribution,

sociologie et application à la cartographie de la pollution atmosphérique.

Actes Mus. Rouen 1981, 1 et 2 : 1-298, 114 fig., 11 tabl. (Mus. Rouen, 198

rue Beauvoisine, F-76000 Rouen).

Thése Doct. Univ. Rouen-Haute-Normandie. Données générales biophysiques

(situation géogr. et humaine, topographie, climatol., végétation arborescente).

Lichénographie : prise en compte des informations sur fiches perforées et car-

togr. de la distr. des esp. Répertoire alphabét. de 133 taxons avec indications

générales, carte de distr., chorologie et écol. (14 esp. apparaissent comme попу.

Source - MNHN. Paris

252 BIBLIOGRAPHIE LICHENOLOGIQUE

pour la Normandie). Lichenosociologie : méthode stigmatiste permettant de

reconnaître 16 groupements dont 11 ass. bien caractérisées. Remarques syncho-

rologiques et synécologiques pour chacun. Cartographie de la pollution atmo-

sphérique : utilisation de la methode de Hawksworth et Rose et proposition

dune nouvelle échelle, établissant la corrélation entre les groupements liché-

niques et la pollution. Cette étude donne un coup d’envoi à la connaissance des

groupements de Normandie et dresse le bilan de la qualité de Pair. Les lichens

pourraient être utilisés comme indicateurs de potentialités biologiques.

MAKAROVA 11. — Species lichenum pro URSS et paeninsula Czukotka novae.

Nov. Sist. Nizsh. Rast., Inst. Bot. Komarova, Akad. Nauk SSSR 1980, 17

150-152, en russe.

2 Cetraria et 2 Pertusaria sont nouv. pour la Péninsule Czukotka. Getrelia

alaskana (Culb. et Culb.) Culb. et Culb. пошу. pour l'URSS.

OLECH M. — Materials to the Lichen Flora of the Khangai Mountains (Mong

lia). Fragm. Florist. Geobot. 1979, 25, 3 : 439-443, 1 fig. (Inst. Вог, Univ.

Jagiellonskiego, Krakow, Poland).

Liste des lichens avec loc., récoltés sur les pentes sud du Khangai.

PUEYO С. — Sur quelques localités lichéniques dans le sud-ouest de la France

I1. Littoral atlantique (répercussion de l'intervention de l'homme). Bull.

Centr. Etudes Rech. Sci. 1979, 12, 4 : 643-650 (Lab. Muséum et Centre

Etudes Rech. Sci., Biarritz).

Agrandissement, stabilisation ou récession des surfaces couvertes par les

lichens ont été suivis avec Parmelia, Cladonia et Usnea dans les landes du

littoral, et avec Lichina, Verrucaria et Xanthoria sur les rochers du pays basque.

SOLYMOSI Р. — A Duna hullámteri erdöinek epiphyton zuzmóvegetációja. Bot.

Kózlem. 1978, 65 (1) : 7-13, 3 fig, en hongr., rés. allem. (Növenyvedelmi

Kutatointeset, H-1525 Budapest).

Lichens épiphytes des régions inondées par le Danube entre Rajka et Mohäes.

Pollution atmosphérique en fonction des installations industrielles. Physcia et

Parmelia apparaissent comme les lichens les plus sensibles, Peltigera et Xanthoria

comme les moins sensibles.

SOLYMOSI Р. — A Gerecse-Hegység Zuzmoi. Bot. Közlem. 1979, 66 (3) :185-

193, 4 fig, en hongrois, гёз. allem. (Idem).

Liste de 101 lichens avec loc. de la région de Gerecse. Données écol.

Commission paritaire 15-8-1980 N° 58611

Dépôt légal n° 10949 - Imprimerie de Montligcon

Sorti des presses le 20 juin 1981.

Source - MNHN, Paris

COLLOQUE INTERNATIONAL

du CNRS № 258

ECHANGES IONIQUES TRANSMEMBRANAIRES

CHEZ LES VEGETAUX

TRANSMEMBRANE IONIC EXCHANGES IN PLANTS

org. : G. Ducet, В. Heller, М. Thellier

Universités de Rouen et Paris VII - 5-11 juillet 1976

@ analyse des modèles théoriques e recherche des couplages métaboliques ou autres

® études électrophysiologiques @ cas particulier des transferts d’anions et de molécules

organiques @ localisation d’ions et aspects structuraux et moléculaires @ intervention

d’échanges ioniques dans les régulations intercellulaires

@ kinetic and thermodynamic considerations, model systems

® metabolic and other couplings, ATPases

® particular features of anionic transfers

absorption of organic molécules

localization, molecular and structural aspect of the transfers

interference of the transmembrane transfers in other processes than absorption

ion exchanges in cell organites

(69 communications dont 64 en anglais et 5 en français)

21х29, 7 - 608 pages - broché 180 F

286 fig. - 89 tabl. - 30 phot.

ISBN 2-222-02021-2

(co-édition CNRS-Université de Rouen)

CCP Paris 9061-11 - Tél. 555.92.25

chez son librair

á défaut aux Editions du CNRS (chéque joint) 8

mande votre documentation

iences humaines

ces exactes et naturelles

г de la langue Francaise

В Revue de ГА!

III ш

Source NEIN Paris

»rofession

SOMMAIRE

рл. NIMIS. — Epigaeous lichen synusiae in the Yukon Territory. . .

D.G. HORTON. — The taxonomic status of Encalypta microstoma Bals.

et De Not. and Е. ciliata var. microstoma Schimp...............

TH. MOXHAM. — Growth rates of Xanthoria parietina and their rela-

tionship to SIE AE E rai Зе

M. PRIVITERA e R. LO GIUDICE. — Vegetazione muscinale della

faggeta del bosco di Malabotta (Peloritani). .............. sph

D. GRIFFIN, III. — El genero Sphagnum en Los Andes de Colombia y

Venezuela, Clave para las especies frecuentes u ocasionales con notas

ecologicas y taxonomicas ......... О Е unten

C. VICENTE and B. CIFUENTES. — L-usnate and permeability... . . + »

C. LEFORESTIER. — Résistance è la sécheresse et multiplication végéta-

tive chez Fossombronia caespitiformis De Not

NOTE

В. DHIEN. — Florule bryologique des rochers bathoniens. .

BIBLIOGRAPHIE BRYOLOGIQUE.......................

BIBLIOGRAPHIE LICHÉNOLOGIQUE

urca

127

181 +

201

213

223

UNE