BRYOLOGIE-LICHENOLOGIE

ANCIENNE REVUE BRYOLOGIQUE ET LICHENOLOGIQUE

Fondée par T. HUSNOT en 1874

Directeur : Mme S. JOVET-AST

Rédaction : Mme H. BISCHLER, M. D. LAMY

Éditeur : A.D.A.C.

COMITÉ DE LECTURE

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

HEBRARD, S. JOVET-AST, D. LAMY, M.C. NOAILLES, C. SUIRE.

Lichénologie : J. ASTA, T. BERNARD, B. BODO, W.L. CULBERSON, M.C. JANEX-

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

MANUSCRITS

Les instructions aux auteurs sont publiées dans le premier fascicule de chaque tome.

Les auteurs sont priés d'adresser leurs manuscrits (en double exemplaire) à la Rédaction de

CRYPTOGAMIE, Bryologie-Lichénologie, Laboratoire de Cryptogamie, 12 rue Buffon,

75005 Paris.

Les tirages à part et les planches photographiques sont à la charge des auteurs.

ABONNEMENTS ANNUELS

CRYPTOGAMIE comprend trois Sections : Cryptogamie, Algologie; Cryptogamie,

Bryologie-Lichénologie; Cryptogamie, Mycologie.

Abonnement à l'une ou l'autre Section pour 1988:

France 305 Е HT (317,20 F ТТС) Etranger 335 F HT

Abonnement aux 3 Sections pour 1988 :

France 870 F HT (904,80 F TTC) Etranger 950 F HT

Prière de bien vouloir envoyer le montant par chèque bancaire ou par chèque postal

libellé à l'ordre de : A.D. A.C. — CRYPTOGAMIE, et adressé à :

A.D.A.C. — CRYPTOGAMIE

12, rue Buffon, F-75005 Paris.

CRYPTOGAMIE, Bryologie - Lichénologie est indexé par Biological Abstracts, Chemical

Abstracts, Publications bibliographiques du CDST (Pascal).

Source : MNHN, Pat

CRYPTOGAMIE

BRYOLOGIE

LICHENOLOGIE

ТОМЕ9 Fascicule 2 1988

Publié avec le concours du Muséum National d’Histoire Naturelle

Source : ММНМ. Paris

Mer مت‎ em ant par ۶

Cryptogamie, Bryol. Lichénol. 1988 9 (2) : 95-102 95

PSEUDOCROSSIDIUM REPLICATUM (TAYL.) ZANDER

REPLACES BARBULA ACUTATA C. MULL.

A NOTE ON ITS SYNONYMY, DISTRIBUTION

AND THE XEROPOTTIOID LIFE SYNDROME

W. FREY and H. KURSCHNER*

ABSTRACT. — The xerotropical Barbula acuta is known as endemic from South Africa.

‘There it occurs in semi-arid grasslands. Finds in the Arabian Peninsula (Asir Mountains of

Saudi Arabia) and comparison with Pseudocrossidium replicatum specimens from Mexico,

Peru and Kenya have shown that Barbula acutata is conspecific with Pseudocrossidium re

plicatum and should be considered as synonym (synon. nov.).

Pseudocrossidium replicatum belongs бо the Xerothermic Pangaean element, characte-

rized by the Xeropottioid life syndrome. The assumption of Delgadillo is supported that

the genus together with Aloina and Crossidium derived from a Tortula-Desmatodon like

ancestor. In this case, great systematic relevance shouldn’t be attributed anymore to the

stereid band in the costa of the members of the Pottiaceae s. L in order to define the fami-

lies Trichostomaceae and Pottiaceae.

ZUSAMMENFASSUNG. — Barbula acutata ist als xerotropischer Endemit aus der Bryoflora

Südafrikas bekannt. Die Art tritt dort in semiariden Grasfluren auf. Funde im Asir-Gebirge

der Arabischen Halbinsel und der Vergleich mit Material von Pseudocrossidium replicatum

aus Mexiko, Peru und Kenia ergaben, dass es sich bei Barbula acutata und Pseudocrossidium

replicatum um dieselbe Art handelt. Barbula acutata muss demnach als Synonym von Pseu-

docrossidium replicatum betrachtet werden

Pseudocrossidium replicatum wird dem xerothermen, pangäischen Element zugeordnet,

dessen Vertreter durch das «Xeropottioid life syndrome» charakterisiert sind. Die Annahme

Delgadillos, dass Pseudocrossidium zusammen mit Aloina und Crossidium von einem Tortu-

la-Desmatodon-ähnlichen Vorfahren abstamme, wird unterstützt. Unter dieser Annahme

würde dem Merkmal ein oder zwei Stereidenbänder bei den Pottiaceae s. |. nicht mehr die

systematische Bedeutung wie bisher zukommen.

TAXONOMY

The genus Pseudocrossidium was established by Williams (1915) on the basis

of P. chilense Williams. Following the genus concept of Zander (1979), its

characteristic features are the strongly revolute to spiral-revolute upper leaf

* Institut für Systematische Botanik und Pflanzengeographie der Freien Universitát Berlin,

Altensteinstr. 6, D-1000 Berlin 33.

Source : MNHN. Paris

Fig. 1. — 1-4, Pseudocrossidium replicatum (Saudi Arabia, Frey & Kürschner 84-290). 1.

Leaves. 2. Cross-section of middle part of leaf. 3. Cross-section of basal part of leaf. 4.

Spirally-coiled leaf margin. 5-7. P. replicatum (Peru, Hegewald по, 5035). 5. Cross-section

of upper part of leaf. 6. Cross-section of middle part of leaf. 7. Spirally-coiled leaf mar-

gin. 8.9. P. replicatum (Sudan, redrawn from Townsend 1984). 8. Cross-section of leaf at

about halfway. 9. Leaf. 10-11. P. replicatum (South Africa, syn. Barbula acutata, redrawn

from Magill 1981). 10. Cross-section of leaf. 11. Leaf.

Source - ММНМ, Pans

PSEUDOCROSSIDIUM REPLICATUM 97

margins with cells often more or less strongly chlorophyllose and differentiated

as a photosynthetic organ (cf. also Herzog 1926), the usually strong dorsal

(abaxial) stereid band of the costa, the 4 (-6) guide cells in one layer, the often

present hydroids, and the weak (or often absent) ventral (adaxial) stereid band

(Figs. 1, 2). According to Zander & Steere (1978), Pseudocrossidium is distin-

guished from Barbula Hedw. mainly by the trend to develop parts of the leaf

into photosynthetic organs. From Tortula specimens (e. g. Tortula porphyro-

neura (C. Mill.) Townsend, cf, Townsend 1979, Magill 1981, Frey & Kürschner

1988) Pseudocrossidium is distinguished by the differentiation of an adaxial

costal epidermis, which is either lacking or rare and poorly differentiated in Tor-

tula (Zander 1979) as well as in the number of guide cells.

Originally, the genus Pseudocrossidium shows an American distribution pat-

tern. But the revision by Zander (1979), which showed the synonymy of Bar-

bula sect. Revolutae B.S.G., extended the area to Europe. Townsend (1984)

reported Pseudocrossidium replicatum from E. Africa (Kenya, Somalia, Sudan).

Our collections from the Asir Mountains in Saudi Arabia include three

samples the taxonomical position of which present difficulties at first sight.

Their examination showed a considerable identity with Barbula acutata, des-

ctibed by Miiller (1899) and regarded as endemic in southern Africa (Magill

1981). The description given by Magill (1981) («. . . margins spirally revolute,

entire, marginal cells differentiated into specialized photosynthetic regions; ۰

Costa short excurrent to percurrent, strong towards the apex, 60-70 um wide

at apex; ... guide cells 4-6, large, incrassate, ventral stereid band small, 1-4 cells

over central guide cells, ... dorsal stereid band strong, 5-6 cells thick...») is in

full accordance with the Arabian specimens and differs only slightly from the

description of Pseudocrossidium replicatum given by Zander (1979) («... Leaf

margins spiral-revolute, of thin-walled, hollow-papillose, highly chlorophyllose

cells; ... leaf apex broadly acute to rounded; ... adaxial stereid band absent or

represented by a few substereid cells, guide cells 4 (-6), in 1 (-2) layers, hydroids

(Begleiter cells) present, abaxial stereid band strong»). These nearly identical

descriptions might be the reason, why Magill (pers. comm., and 1981, р. 239)

noted : «The highly specialized marginal cells and the presence of the ventral

stereid band in the costa characterize this species (B. acutata). These features

also place the species within Zander’s (1979) concept of Pseudocrossidium

Williams».

The comparison of the Arabian material with the collections of Pseudocros-

sidium replicatum from Peru (Hegewald no. 5035), Mexico (Zander no. 4950)

and Kenya (Townsend no. 80/176) and Barbula acutata from 5۰ Africa (Magill

no. 6715) shows, that they all are identical (Fig. 1). Characteristics of the Ara-

bian material and the investigated samples from S. Africa, Peru, Mexico and

Kenya are the spirally revolute margins of the leaves with the thin-walled inner

cells, differentiated as a photosynthetic organ (Fig. 1.4, 2.2), and the two stereid

bands (Figs. 1.2, 1.10) in the middle part of the costa one of which is lacking

in the lower part of the leaf (Fig. 1.3).

Source : ММНМ. Paris

Fig. 2. — Pseudocrossidium hornschuchianum (Jordan, Kürschner 86-26). 1. Cross-section

and longitudinal section of leaf margins. 2-4. Pseudocrossidium replicatum (Saudi Ara

bia, Frey & Kiirschner 84-290). 2. Cross-section of leaf margin with photosynthetic

region. 3. Dorsal (abaxial) surface of leaf with strong costa. 4. Papillae of the adaxial

surface of the costa.

Source : MNHN, Paris

PSEUDOCROSSIDIUM REPLICATUM 99

The variability of the stereid layers and the sometimes lacking ventral stereid

band causes the problems in the systematic classification of this taxon. Both in

the taxa from E. Africa (cf. Townsend 1984, Fig. 16, redrawn in Fig. 1.8) as

well as in the one from Peru (Fig. 1.6, confirmed by Zander as D. replicatum)

there is a ventral stereid band present, while with Zander (1981, pl. IV, Fig. 6,

P. replicatum) this ventral stereid band is lacking. In our opinion, and in accor-

dance with Delgadillo (1975), the taxonomic value of the presence of one dorsal

stereid band (subfamily Pottioideae, e. g. Brotherus 1924) and the presence of

one ventral and one dorsal stereid band (subfamily Trichostomoideae) is questio-

nable, especially in Pseudocrossidium (cf. Delgadillo 1975; Zander 1979, 1981).

Therefore, on account of the identically transverse sections of the costae as

well as of other mentioned features, Barbula acutata can be reduced as a syno-

nym to Pseudocrossidium replicatum.

Synonymy (cf. Zander 1979, Townsend 1984) :

Pseudocrossidium replicatum (TAYL.) ZANDER in Phytologia 44 : 206 (1979).

Barbula acutata C. Muell. hal. in Hedwigia 38 : 109 (1899), Broth, in Natürl. Pfanzenfam.

10 : 280 (1924), synon. nov. Type : Transvaal, Spitzkop, near Lydenburg, Wilms s. n.,

1887 (G, holotype).

Barbula apiculata Hampe іп Linnaea 31 :519 (1862) non В. apiculata Hedw., 1801. Tortula

apiculata Mitt. in J. Linn. Soc. Bot. 12 : 153 (1869) nom. nov. pro В. apiculata Hampe

non T. apiculata (Hedw.) Turn., 1804.

Barbula perlinearis С. Muell. hal. in Bull. Herb. Boissier 5 : 195 (1897). Type : Guatemala,

Quezaltenango, Bernouilli & Cario по. 118 (NY, isotype).

Barbula replicata Tayl. in London J, Bot. 5 : 49 (1846). Basionym. Type : Ecuador, Pi-

chincha, Quito, walls, Jameson 1847, no. 103 (BM, isotype), Tortula replicata (Tayl.)

Wils. in London J. Bot. 5 : 454 (1846).

Barbula spiralis Schimp. ex C. Muell. hal. in Syn. Musc. 1 : 622 (1849). Type : Mexico,

Veracruz, Yarrea, Mirador, Liebmann 1842 (BM, isotype). Tortula spiralis (Schimp. ex

C. Muell.) Mitt. in J. Linn, Soc. Bot. 12 : 151 (1869).

Barbula spiralis Schimp. ех С. Muell. var. emarginata Card. іп Rev. Bryol. 36 : 84 (1909).

Type : Mexico, Mexico, Amecameca, Pringle no. 10611 (BM, MEXU, F, FH, isotypes).

Trichostomum fusco-mucronatum C. Muell. hal. in Linnaea 40 : 295 (1876). Type : Soma-

lia, Meid, April 1875, Hildebrandt no. 1497 (B, holotype).

DISTRIBUTION AND ORIGIN

Fig. 3 shows the distribution of Pseudocrossidium replicatum. This distribu-

tion pattern (southwestern USA, Mexico, N. Andes (Peru, Argentina), E. Africa

(Kenya, Somalia, Sudan - Jebel Marra, Jebel Nami), S. Africa, and southwestern

Arabia (Saudi Arabia)) is shared by a lot of taxa (е. р. Aloina spp., Crossidium

spp., Plagiochasma spp., Targionia spp.) and reflects an old distribution pattern

introduced as «Xerothermic Pangaean» by Frey & Kürschner (1988). Characte-

ristic of nearly all the old taxa is the tendency to establish xeromorphic life

strategies evolved by parallel evolution within the Pottiaceae and Marchantiidae

and which were summarized by Frey & Kiirschner (1988) as «Xeropottioid

Source : ММНМ, Paris

100 W. FREY and H. KÜRSCHNER

Fig. 3. — Distribution of Pseudocrossidium replicatum.

and Xerothalloid life syndroms». Within the Pottiaceae the Xeropottioid life

syndrome is concentrated to Aloina, Aloinella, Barbula, Crossidium, Pseuda-

loina, Pseudocrossidium, Pterygoneurum and Tortula species. These genera are

believed to have derived from permo-triassic, highly developed progenitors,

which were able to colonize the arid xerothermic areas of the Pangaean conti-

nent (Frey & Kürschner 1988). This would support the assumption of Delga-

dillo (1975) that the trichostomoid genus Pseudocrossidium is related to the

pottioid genera Aloina and Crossidium and these taxa and those one of Aloi-

nella, Pterygoneurum, Desmatodon and Pottia have derived from a Tortula-

Desmatodon like ancestor and form a unit. The consequence would be to think

about the systematic value of the occurrence of ventral and dorsal stereid bands

within the Pottiaceae 5. 1., which would not anymore be of such a great syste-

matic relevance to define the families Trichostomaceae and Pottiaceae.

Common to all these taxa is the tendency to dislocate the photosynthetic

active organs away from the leaf lamina which is done in two separate ways :

One evolutionary line resulted in the leaf outgrowth and appearance of Ніз-

ments restricted to the costa, and restriction of photosynthesis to the filaments

(е. в. Aloinella, Crossidium, Pseudaloina, Pterygoneurum). Highly specialized

is the hyaline leaf lamina of e. g. Aloina covering the photosynthetic active

filament-layer and protecting it against high light intensities. The ventral out-

growth on the costa (in the form of filaments or lamellae) is a unique character

among mosses (cf. also Polytrichaceae and Grimmiaceae).

The other evolutionary line resulted in the spirally revolute leaf margin (e. g.

Pseudocrossidium, Tortula), in which the photosynthetic active cells are concen-

trated («Assimilationsstreifen» cf. Herzog 1926, p. 24). In addition, the marginal

cells are much more thick-walled on the outside than on the inside (Figs. 1.4,

1.7) which is an additional protection.

Source : MNHN, Paris

PSEUDOCROSSIDIUM REPLICATUM 101

The two evolutionary lines must be considered as an adaptation to arid and

semi-arid habitats (desert shrublands, semi-arid grasslands and heathlands) (cf.

Magill 1981, p. 240 «The presence of specialized marginal cells in Tortula por-

phyroneura suggests that modification of marginal or costal cells into different

photosynthetic tissues is an adaptation to harsh environment. , .», Herzog 1926,

Frey & Kürschner 1988). According to Herzog (1926) the ventral stereid band

(е. в. Pseudocrossidium) is also an expression of arid environments. Тһе Xero-

pottioid life syndrome enabled these old xerothermic Pangaean taxa to survive

and to evolve under semidesert conditions.

SPECIMENS STUDIED

Barbula acutata C. Muell. Southern Africa, Natal, Drakensberg, Royal National

Park, «The Crack», above Gudu Falls, 2200 m. Alpine heath/grassland with

sandstone canyon walls and boulders. Podocarpus and Leucosidea. In soil

overhang. Leg. R.E. Magill, 9 Febr. 1984 (6715). Det. R.E. Magill 1987.

Missouri Botanical Garden Herbarium (MO).

Pseudocrossidium replicatum (Tayl.) Zander. Mexico. Est. Puebla, 8 km М of

Azumbilla, on calcareous rock, thin soil, gully, с. 2240 m. Leg. R.H. Zander,

23 Oct. 1982 (4950). The Clinton Herbarium, Buffalo Soc. of Nat. Sciences,

Buffalo, New York.

Pseudocrossidium replicatum (Tayl.) Zander. Kenya. Naivasha District, on S. -

facing sandy places on slope c. 2 km NW of «Kinja» nursery, South Lake

Navaisha, property of Е. Magius, alt. с. 1840m. Leg. & det. : С.С. Townsend,

19 May 1980 (80/176). Conf. : R.H. Zander.

Pseudocrossidium replicatum (Tayl.) Zander. Peru. Depto. La Libertad, Prov.

Otuzco, Casmiche, 2000 m, leg. P. & E. Hegewald, 8 April 1973 (5035).

Det. : RH. Zander. B, Herb. Hegewald.

Pseudocrossidium replicatum (Tayl.) Zander. Saudi Arabia : Asir Mts., al Qa'raa

S. of Kamish Mushayt, 2100 m, on soil, leg. Kürschner & Kónig, 17 Aug.

1982 (82-2292 b); Jebel Qarn near al-Farhan, 5. of Kamish Mushayt, 2400 m,

in rocks crevices, leg. Frey & Kürschner, 27 March 1984 (84-290); Jebel

Fayfa, 60 km NE. of Gizan, 1500 m, in cultivated fields, leg. Frey & Kürsch-

ner, 30 March 1984 (84425). Herb. W. Frey, Herb. H. Kürschner.

ACKNOWLEDGEMENTS. — For valuable suggestions and for bringing certain specimens to

our attention we thank R.E. Magill (St. Louis) and R.H. Zander (Buffalo). We also thank

Mrs. C. Graber and Mrs. Н. Ritter for preparing the SEM specimens and photos and Mrs I.

Pohl for looking at the text and typing the manuscript.

Source : MNHN, Paris

102 W. FREY and Н. KÚRSCHNER

REFERENCES

BROTHERUS V.F., 1924 — Bryales. In : ENGLER, A. & PRANTL, K., Die natürlichen

Pflanzenfamilien 2. Aufl. 10. Leipzig.

DELGADILLO M.C., 1975 — Taxonomic revision of Aloina, Aloinella and Crossidium

(Musci). Bryologist 78 : 245-303.

FREY W. & KURSCHNER H., 1988 — Bryophytes of the Arabian Peninsula and Socotra.

Floristics, phytogeography and definition of the Xerothermic Pangaean element. Studies

in Arabian bryophytes 12. Nova Hedwigia 46 : 37-120.

HERZOG T., 1926 — Geographie der Moose. Jena.

MAGILL R.E., 1981 — Bryophyta Part 1, Mosses, Fasc. 1 Sphagnaceae - Grimmiaceae. In :

LEISTNER, O.A. (ed.), Flora of Southern Africa, Part I. Pretoria.

MULLER C., 1899 — Contributiones ad Bryologicam austro-afram. Hedwigia 38 : 52-155.

TOWNSEND C.C., 1979 — A new combination in African Tortula. J. Bryol. 10 :576.

TOWNSEND C.C., 1984 — A small collection of mosses from Jebel Marra, Sudan Republic.

Lindbergia 10 : 175-180.

WILLIAMS R.S., 1915 — Mosses from the west coast of South America. Bull. Torrey Bot.

Club. 42 : 393-404.

ZANDER R.H. & STEERE W.C., 1978 — Tortula scotteri sp. nov. from the Northwest

Territories of Canada. Bryologist 81 : 463-467.

ZANDER R.H., 1979 — Notes on Barbula and Pseudocrossidium (Bryopsida) in North

‘America and an annotated key to the taxa. Phytologia 44 :177-214.

ZANDER R.H. 1981 Descriptions and illustrations of Barbula, Pseudocrossidium and

Bryoerythrophyllum (p. р.) of Mexico. Cryptogamie, Bryol. Lichénol. 2 : 1-22.

Source : MNHN. Paris

Cryptogamie, Bryol. Lichénol. 1988 9 (2) : 103-107 103

BRIOFITOS DEL PIRINEO DE NAVARRA (ESPANA)

A. EDERRA INDURAIN*

RESUMEN. — Presentamos un catálogo de 82 taxones de briofitos recolectados en el Piri-

neo navarro. Nueve de ellos son primeras citas para la provincia, y entre ellos destaca Tortel-

la densa (Lor. & Mol.) Crundw. & Nyh., nuevo para el Pirineo español, e Hygrohypnum

cochlearifolium (Vent.) Broth., que no había sido citado desde el siglo pasado.

ABSTRACT. — A catalogue with 82 taxa of bryophytes collected in the Pyrenees of Navarra

is presented. Nine of them are new records for the province. Among them, Tortella densa

(Lor. & Mol.) Crundw. & Nyh., new for the Spanish Pyrenees, and Hygrohypnum cochleari-

folium (Vent.) Broth., not recorded since the last century, are pointed out.

INTRODUCCION

La brioflora del extremo nororiental de Navarra, donde se encuentran las

cotas pirenaicas más altas de la provincia, ha sido poco estudiada. De esta zona

sólo se conocían las aportaciones de Allorge y Casas (1963) que citaron 27

especies de Uztárroz y su entorno, y de Villar (1980), que daba cuenta de 47

especies de Belagua-Larra y alrededores. También P. Allorge, en varios trabajos,

señaló algunas especies de Isaba, que fueron recopiladas por V. Allorge (1955-

56). Puesto que en el reciente trabajo de Casas (1986) no se incluye la zona

pirenaica de Navarra, hemos creído interesante dar a conocer en este trabajo

los resultados de varias recolecciones efectuadas en los últimos años en dicha

zona,

Los muestreos se han llevado a cabo, siempre por encima de 1600 m de altitud,

en los siguientes lugares : Monte Orhi (30TXN66, 2021m de altitud máxima),

monte La Cartxela (30TXN75, 1982m de altitud máxima), Larra (30TXN85,

entre 1500 y 2000m de altitud), monte Lákora (30TXN75, 1847m de altitud

máxima) y monte Arlás (30ТХМ85, 2062m de altitud máxima).

El sustrato más extendido en las áreas de muestreo es de carácter básico,

bien calizas duras del Cretácico, dominantes en el karst de Larra, bien estratos

de flysch margo-calizo, que aparece en forma de bandas entre las calizas en

* Depto. Botánica, Facultad Ciencias, Universidad de Navarra, Pamplona, España,

Source : MNHN, Paris

104 A. EDERRA INDURAIN

Orhi y La Cartxela. Sólo aparecen rocas ácidas en Lákora, donde afloran conglo-

merados del Cretácico Inferior.

En cuanto a la vegetación, casi todas las zonas muestradas están ocupadas por

prados de diente dominados por Nardus stricta L. y Trifolium alpinum L., en

mosaico con prebezales marcados por la abundancia de Calluna vulgaris (L.)

Hull. Zonas arboladas (por encima de 1600 m) perduran en Larra, donde se

encuentran masas de hayedo-abetal, que hacia cotas más altas dan paso a pinares

claros de Pinus uncinata Miller ex Mirbel. Los afloramientos rocosos calizos

están colonizados por una rica flora saxicola y casmófita, con especies tan intere-

santes como Saxifraga longifolia Lapeyr. o Petrocoptis pyrenaica (J. P. Bergeret)

A. Braun,

CATALOGO

A continuación se relacionan por orden alfabético los briofitos identificados,

en primer lugar las hepáticas y a continuación los musgos. Para cada especie se

da el lugar, hábitat y altitud aproximada en que se recolectó. Las especies nuevas

para el catálogo de Navarra se indican con un asterisco.

Hepaticas

*Barbilophozia barbata (Schmid. ex Schreb.) Loeske : Larra, suelo entre rocas

calizas, 1700 m.

Barbilophozia hatcheri (Evans) Loeske : Lákora, rocas ácidas húmedas cerca de

la cima, 1800m; Arlás, suelo muy húmedo acidificado, 1750m. Los ejempla-

res de Lákora fueron citados como novedad provincial por Ederra (1986).

Cololejeunea calcarea (Libert) Schiffn. : Larra, entre otros briofitos en suelo,

1700 m.

Conocephalum conicum (L.) Underw. : Larra, suelo entre calizas, 1700 m.

Diplophyllum albicans (L.) Dum. : Lákora, sobre conglomerados húmedos y en

suelo entre rocas, desde 1600 a 1800 m.

Frullania tamarisci (L.) Dum, : Larra, sobre rocas en el interior del hayedo-

abetal, 1650 m.

*Gymnomitrion concinnatum (Lightf.) Corda in Opiz : Lákora, mezclado con

otros briofitos sobre conglomerados húmedos cerca de la cima, 1800 m.

Jungermannia hyalina Lyell in Hook. : Lákora, sobre rocas ácidas rezumantes,

1800 m.

*Lophozia sudetica (Nees ex Hüb.) Grolle : Lákora, sobre rocas cerca de la

cumbre, 1800 m.

Lunularia cruciata (L.) Lindb. : Lákora, rellanos de rocas alteradas sombrías

y húmedas, 1600 m.

Marsupella emarginata (Ehrh.) Dum. : Lákora, sobre conglomerados húmedos,

1800 m.

Plagiochila porelloides (Torrey ex Nees) Lindenb. : Larra, sobre calizas en el

hayedo-abetal, 1650 т.

Source - MNHN, Paris

BRIOFITOS DE NAVARRA 105

Porella platyphylla (L.) Pfeiff. : La Cartxela, cavidades entre rocas, 1650 m; Ar-

lds, grietas entre rocas, 1750 m.

Preissia quadrata (Scop.) Nees : Larra, en grietas de rocas calizas, 1650 m.

Ptilidium ciliare (L.) Hampe : Lákora, sobre rocas ácidas cerca de la cima,

1800 m.

Riccardia chamedryfolia (With.) Grolle : Larra, еп suelo htimedo, 1650 m.

Scapania aequiloba (Schwaegr.) Dum. : Larra, sobre rocas calizas, 1650 т.

Scapania aspera M. et H. Bern. in H. Bernet : Orhi, sobre calizas sombrias,

1850 m; Larra, en suelo y sobre rocas calizas en el hayedo-abetal, 1650 т.

Musgos

Andreaea rupestris Hedw. var. alpestris (Thed.) Sharp : Lákora, sobre rocas

ácidas cerca de la cima, 1800 т. Se dió a conocer como novedad para Navarra

en Ederra (1986).

Barbula unguiculata Hedw. : Orhi, rocas calizas soleadas, 1700 m.

Bartramia ithyphylla Brid. : Lákora, sobre conglomerados, rezumantes, cerca

de la cima, 1800 m.

Brachythecium rutabulum (Hedw.) B., S. & С. : Larra, sobre rocas en el hayedo-

abetal, 1650 m.

Bryum argenteum Hedw. : Lákora, en suelo somero sobre rocas, 1600 m.

*Bryum elegans Nees ex Brid. : Orhi, paredes de una cueva en calizas, bastante

sombrías, 1850 m.

*Bryum inclinatum (Brid.) Bland, (= B. stenotrichum C. Müll.) : La Cartxela,

suelo de pradera húmeda y con gran inclinación, 1920 m.

Bryum pseudotriquetrum (Hedw.) Gaertn., Meyer & Scherb. : Lákora, suelo

entre rocas rezumantes, 1700 m.

Campylium calcareum Crundw. & Nyh. : Larra, suelo entre calizas, 1700 т.

Campylium chrysophyllum (Brid.) J. Lange : Lákora, rocas básicas soleadas,

1700 m.

Ceratodon purpureus (Hedw.) Brid. : Lákora, rocas soleadas, 1750 m.

Ctenidium molluscum (Hedw.) Mitt. : Larra, rocas dentro del hayedo-abetal,

1650 m; Lákora, calizas soleadas, 1700 m; Orhi, rocas algo sombrías, 1800m.

Dicranella heteromalla (Hedw.) Schimp. : Lákora, sobre rocas ácidas rezu-

mantes, 1800 m.

Dicranum fuscescens Sm. : Larra, sobre rocas y en grietas someras, 1700 m.

Dicranum scoparium Hedw. : Larra, sobre calizas erosionadas en el hayedo-abetal,

1650 m; Lákora, sobre rocas ácidas cerca de la cumbre y en grietas poco

profundas, desde 1600 a 1800 m.

Didymodon fallax (Hedw.) Zander : Orhi, sobre rocas soleadas, 1700 m.

Distichium capillaceum (Hedw.) B., S. & С. : Lákora, sobre calizas, 1750 т;

Arlás, grietas entre rocas calizas, 1750 m.

Ditrichum flexicaule (Schwaegr.) Hampe : Lákora, sobre calizas, 1750 m.

Dryptodon patens (Hedw.) Brid. : Lákora, sobre rocas ácidas rezumantes,

1800 m.

Encalypta streptocarpa Hedw. : Larra, grietas de calizas y suelo, 1700 т,

Source - MNHN, Paris

106 А. EDERRA INDURAIN

Fissidens cristatus Wils. ex Mitt. + Larra, suelo del fondo de una dolina а 1700m.

Funaria muhlenbergii Turn, : Lákora, en suelos someros sobre rocas, а 1600 ۰

Grimmia orbicularis Bruch ех Wils. : Orhi, en rocas calizas soleadas, 1700 m.

Gymnostomum aeruginosum Sm. : Orhi, rocas calizas rezumantes, 1850 m.

Homalothecium lutescens (Hedw.) Robins. : Arlás, formando fieltros en el suelo

de prados alpinizados, 1800 m.

Homalothecium philippeanum (Spruce) B., 5. & С. : Lákora, sobre rocas y entre

grietas, 1700 m.

Homalothecium sericeum (Hedw.), B., S. & G. : Larra, sobre rocas calizas en

el interior del pinar de P. uncinata, 1700 m.

Hygrohypnum cochlearifolium (Vent.) Broth. : Orhi, suelo de cueva poco

profunda, sobre calizas rezumantes, 1850 m. Según Casas (1986) y Casares

Gil (1915), esta especie no había sido citada en el Pirineo español desde el

siglo pasado.

Hylocomium splendens (Hedw.) B., S. & С. : Lákora, suelo de praderas de Nar-

dus stricta, 1700 m.

Hypnum cupressiforme Hedw. : Larra, sobre rocas en el hayedo-abetal, 1650 m.

Isothecium alopecuroides (Dubois) Isov. : Larra, sobre rocas calizas en hayedo-

abetal, 1650 m.

*Lescuraea incurvata (Hedw.) Lawt. : Lákora, en grietas entre calizas, 1700 m;

La Cartxela, sobre rocas, 1700 m; Orhi, sobre rocas, entre 1700 y 1900 m.

Mnium marginatum (With.) P. Beauv. : Lákora, rocas calizas y grietas, 1700 m;

Larra, suelo de los claros del hayedo-abetal, 1650 m.

Mnium stellare Hedw. : Lákora, grietas y suelo somero entre rocas calizas a

1700 m.

Myurella julacea (Schwaegr.) B. S. & С. : Orhi, rocas entre 1700 y 1900 m;

Lákora, rocas calizas, 1700 т; Arlás, grietas de calizas, 1750 т,

Neckera complanata (Hedw.) Hib. : Larra, sobre rocas en hayedo-abetal, 1650m.

Neckera crispa Hedw. : Larra, sobre rocas calizas en hayedo-abetal, 1650 m;

Larra, sobre rocas sombrías en el pinar de pino negro, 1700 m.

Orthothecium rufescens (Brid.) B., S. & С. : Larra, en suelo y entre rocas calizas,

1750m.

Orthotrichum cupulatum Brid. : Orhi, sobre rocas soleadas, 1700 m.

Philonotis fontana (Hedw.) Brid. : Lákora, grietas de rocas rezumantes, a 1800m.

Pogonatum aloides (Hedw.) P. Beauv. : Orhi, suelos someros sobre rocas alte-

radas, 1700 ۰

Polytrichum alpinum Hedw. : Lákora, en grietas de rocas soleadas, 1650 m.

Polytrichum formosum Hedw. : Lákora, sobre rocas ácidas alteradas y rezuman-

tes, 1800 m.

Polytrichum juniperinum Hedw. : Larra, suelo de claros de hayedo-abetal,

1650 m.

Polytrichum piliferum Hedw. : Lákora, rocas ácidas cerca de la cumbre, 1800m.

Pseudoleskeella catenulata (Schrad.) Kindb. : Lákora, sobre rocas calizas y en

grietas, 1700 m.

Pterigynandrum filiforme Hedw. : Lákora, sobre conglomerados soleados y

húmedos, 1800 m.

Source : MNHN. Paris

BRIOFITOS DE NAVARRA 107

Pterygoneurum ovatum. (Hedw.) Dix. var. incanum (Nees & Hornsch.) Broth

Orhi, rocas soleadas algo alteradas en superficie, 1700 m.

Racomitrium aquaticum (Schrad.) Brid. : Lákora, sobre conglomerados rezu-

mantes y soleados cerca de la cumbre, 1800 тп.

Racomitrium heterostichum (Hedw.) Brid, : Lákora, sobre rocas ácidas cerca

de la cumbre, 1800 m; Orhi, sobre rocas soleadas, 1700 т.

Racomitrium lanuginosum (Hedw.) Brid. : Lákora, sobre rocas ácidas rezuman-

tes, 1800 m.

Rhizomnium punctatum (Hedw.) T. Kop. : Lákora, sobre conglomerados rezu-

mantes próximos a la cima, 1820 m,

Rhynchostegium riparioides (Нейм) Card. : Lákora, en rocas rezumantes, 1800m.

Rhytidiadelphus loreus (Hedw.) Warnst. : Lákora, en rocas próximas a la cum-

bre, 1800 т.

Rhytidiadelphus triquetrus (Hedw.) Warnst,: Lákora, suelo y grietas de rocas

calizas, 1700m.

Thuidium delicatulum (Hedw.) Mitt. : Arlás, entre otros briofitos formando

céspedes sobre el suelo, 1750 m.

Thuidium philibertii Limpr. : Lákora, suelo entre rocas calizas, 1750 m.

Thuidium tamariscinum (Hedw.) B., S. & G : Larra, suelo еп hayedo-abetal,

entre 1600 y 1700 m.

*Tortella densa (Lor. & Mol.) Crundw. & Nyh. : Orhi, rocas calizas soleadas,

entre 1700 y 1900 m. Primera cita para el Pirineo español.

Tortella inclinata (Hedw. f.) Limpr. : La Cartxela, sobre rocas, 1750 m.

Tortella tortuosa (Hedw.) Limpr. : Arlás, suelo de prado alpinizado, 1700 m;

Larra, sobre rocas calizas, 1700 m;Orhi, sobre rocas calizas, 1850 m; Lákora,

sobre rocas calizas, 1750 m.

*Tortula norvegica (Web.) Wahlenb. ex Lindb. : Orhi, rocas sombrías rezuman-

tes, entre 1700 y 1900 m.

Tortula ruralis (Hedw.) Gaertn., Meyer & Scherb. : Lákora, sobre rocas calizas

soleadas, a 1700 m.

BIBLIOGRAFIA

ALLORGE V., 1955-56 — Catalogue préliminaire des Muscinées du Pays basque français

et espagnol. Rev. Bryol. Lichénol. 24 : 96-131;248-333.

ALLORGE V. £ CASAS DE PUIG C., 1963 — Au sujet des Bryophytes récoltés au cours de

Vexcursion de l'Association Internationale де Phytosociologie dans les Pyrénées franco-

espagnoles, (22-29 mai 1960). Rev. Bryol. Lichénol. «1962» 1963, 31:213-238.

CASARES GIL А., 1915 — Enumeración y distribución geográfica de las muscineas de la

Península Ibérica. Trab. Mus. Nac, Ci. Nat., Ser. Bot. 8 : 1-179.

CASAS SICART C., 1986 — Catálogo de los briofitos de la vertiente española del Pirineo

Central y de Andorra. Collect, Bot. (Barcelona) 16 (2) : 255-321.

EDERRA INDURAIN A., 1986 — Aportaciones a la flora briológica de Navarra (España).

Publ. Biol. Univ. Navarra, Ser. Bot. 6 : 39.

VILLAR L., 1980 — Briofitos, en + Catálogo florístico del Pirineo Occidental Español.

Publ. Centr. Pir. Biol. Exp. Jaca 11 : 392-397.

Source - MNHN, Paris

AAA

A Ei

UE Le

Free

Cryptogamie, Bryol. Lichénol. 1988 9 (2) : 109-127 109

VEGETACION BRIOFITICA DEL MACIZO ORIENTAL

DE LOS PICOS DE EUROPA (ANDARA) EN CANTABRIA

(ESPANA).

I. COMUNIDADES SAXICOLAS, ACUATICAS Y SUBACUATICAS

Е. FUERTES LASALA & Е. MARTINEZ-CONDE*

ВЕЗОМЕМ. — En este trabajo se describen 4 asociaciones y 2 subasociaciones de la clase

Tortulo-Homalothecietea sericei Hertel 1974, 2 asociaciones de la Montio-Cardaminetea

Braun-Blanquet & Tüxen 1943, 1 de la Fontinaletea antipyreticae v. Hübschmann 1957

y 1 de Adiantetea Braun-Blanquet 1947, en Cantabria (España). En la descripción de los

sintaxones se hace especial consideración de su estructura y especies características, Se dan

las tablas fitosociológicas y un catálogo alfabético de los taxones citados.

SUMMARY. — In this paper are described 4 associations and 2 subassociations of the class

Tortulo-Homalothecietea sericei Hertel 1974, 2 associations of the Montio-Cardaminetea

Braun-Blanquet & Tüxen 1943, 1 of the Fontinaletea antipyreticae v. Hübschmann 1957

and 1 of the Adiantetea Braun-Blanquet 1943, in Cantabria (Spain). Structure and carac-

teristic species are especially considered in the description of the syntaxa. Phytosociological

tables of the associations and an alphabetic catalogue of the species are given.

INTRODUCCION

El territorio, objeto de nuestro estudio, está situado en la parte nororiental

del macizo de Ándara y comprende los valles de los rios Sobra, Urdón, Corvera y

Deva, este último, en el tramo comprendido entre los kilómetros 423-432 de la

carretera que discurre por el desfiladero de la Hermida (Fig. 1). Los límites,

vienen señalados por las coordenadas 43°16/07 y 4371258” de latitud norte

para los puntos septentrional y meridional, las coordenadas 4°45'00” رز‎

de longitud oeste para los límites occidental y oriental respectivamente. Las

coordenadas U.T.M. (50 x 50) son : UNs.

* Dpto. de Biología Vegetal I. Facultad de Ciencias Biológicas. Universidad Complutense.

28040 Madrid (España).

Source : MNHN, Paris

See

ient uat t аза

TRESYISO

4 44¥ LIMITE Provincia

mancanoiw a

pubes Reo monocnanen

CARRETERA

{+ SENDA

Fig. 1. — Mapa general de la zona estudiada y situación de las localidades donde se han

recolectado los taxones que se citan en el catálogo florístico. Escala 1: 50.000.

SS СА

Fig. 2. — Esquema geológico 1:50.000. 1: Caliza ‘Picos de Urbión'. 2: Caliza ‘de Monta

: Caliza sin especificar. 4: Conglomerados y detritos silíceos. 5: Arcillas y limos. 6:

pésitos glaciares : conglomerados con matriz. 7: Cabalgamientos.

10: Red fluvial. 11: Divisorias Principales, 12: Carretera.

Fallas. 9: Contactos.

Source : MNHN. Paris

VEGETACION BRIOFITICA EN ANDARA I 111

CARACTERISTICAS DEL TERRITORIO

GEOLOGIA

El sustrato de la mayor parte de la zona está constituido por “calizas de

montaña? (calizas negras de grano fino, fétidas y blanqueadas; brechas y calizas

masivas de grano grueso y grises) de edad namuriense A-westfaliense A inferior,

y calizas de formación ‘Picos de Urbión' (calizas blancas de grano fino y calizas

rosadas brechoides) del periodo westfaliense B-cantabriense (Julivert et Font-

bote 1972).

A ambos lados del collado de la Hoja (Sierra de Beges) afloran conglomerados

silíceos basales (Permo-Trias) que han actuado como área fuente de coluviones

en la márgen derecha del Urdón, A estos materiales y a los coluviones que de

ellos se derivan, se asocia la flora y vegetación acidófila de esta zona.

Además, aparecen formaciones superficiales que recubren buena parte del

sustrato, principalmente coluviones de origen periglaciar y depósitos glaciares.

Destacan las morrenas del valle del Urdón, aunque no faltan otras de menor

tamaño en los demás valles. Las morrenas estan formadas por conglomerados

calizos con abundante matriz arcillosa (Fig. 2).

CLIMATOLOGÍA

Los datos termopluviométricos corresponden a las estaciones de La Hermida

y Tresviso. En dichas estaciones sólo constan datos termopluviométricos a partir

de los años 1975 (La Hermida) y 1980 (Tresviso) debido quizá a la escasa

población y al dificil acceso a este territorio desde el desfiladero de La Hermida.

Los valores medios de temperatura (Tm) y precipitación (Pm) anual, así como la

situación geográfica de las estaciones se resúme en la Tabla 1.

Tabla 1. — Situación geográfica de las estaciones termo pluviométricas y valores de tempera-

tura y precipitación media anual.

Estación Altitud Longitud ^ Latitud. años Tm.anual Рт, anual

La Hermida 100m 0950У 43°15N 197583 13,59С 1333,64mm

Tresviso 900m 095990 43°16N 198083 9,896 2103,17mm

Los diagramas ombrotérmicos (Fig. 3) indican que la estación de Tresviso

acusa un corto periodo de sequía y subsequía en el mes de septiembre en tanto

que la estación de La Hermida presenta sequía en el mes de julio y subsequía

desde finales de mayo a agosto (Martinez-Conde 1986).

COROLOGÍA Y VEGETACÍON VASCULAR

Corológicamente, el área estudiada está comprendida en el subsector Pico-

europeano, sector Ubiñense-Picoeuropeano, en la provincia Orocantábrica de

Source : MNHN. Paris

112 Е. FUERTES LASALA у E. MARTINEZ-CONDE

TRESVISO

Рю

re eds 1% | зоо

sr} |р

pat] |Р-2т EA le و‎

LA HERMIDA |

200 200

100 100

204 t 204

bso 50

i 5

ло w

Ам) JASON M AS N

Fig. 3. — Diagramas ombrotérmicos de las estaciones de La Hermida y Tresviso (Cantabria).

la Región Eurosiberiana de la Península Ibérica. En dicha área aparecen repre-

sentados los pisos bioclimáticos colino (90450 m), montano (450-1400 m) y

subalpino (1400-2000 m) y hemos podido distinguir las siguientes series de

vegetación :

Source : MNHN, Paris

VEGETACIÓN BRIOFÍTICA EN ANDARA 1 113

— Serie subalpina orocantábrica basófila del enebro rastrero (Daphno canta-

bricae - Arctostaphyleto uva-ursi sigmetum).

— Serie montana orocantábrica y cántabro-euskalduna baséfila у ombrófila

del haya (Carici sylvaticae - Fageto sigmetum).

— Serie colino-montana orocantábrica, cántabro-euskalduna y galaico-asturia-

na mesofitica del fresno (Polysticho setiferi-Fraxineto excelsioris sigme-

tum)

Serie colino-montana orocantábrica de la encina (Cephalanthero longifoliae

- Querceto rotundifoliae sigmetum).

El piso subalpino, en nuestra zona muy pedegroso, presenta una vegetación

formada por caméfitos y nanofanerófitos constituida por enebrales ricos en

P у р

gayubas, forman la asociacién Daphno cantabricae - Arctostaphyletum uva-ursi.

El piso montano, predominante en nuestro territorio, aparece claramente

delimitado : el submontano, cuyo área corresponde a la Sierra de Beges y el alti-

montano que corresponde a la Sierra de la Corta. La vegetación de la Sierra de

Beges está formada por robledales de Quercus robur y Quercus petraea, este

último muy escaso. Dichos robledades, que se extienden por el collado de la

Hoja, alcanzan su óptimo vital en los niveles bajos de suelos húmedos y profun-

dos de los valles y conviven con fresnos, castaños y tilos. Estos bosques mixtos,

han sido definidos como asociación Polysticho setiferi - Fraxinetum excelsioris,

que en las hoces y gargantas profundas de los ríos, en laderas orientadas al

Norte se enriquece en tilos formando la subasociación tilietosum platyphyllae

(Rivas Martinez et al. 1984).

El piso montano superior (Sierra de la Corta) está ocupado por un gran bos-

que de hayas situado en posiciones de ombroclima húmedo o hiperhúmedo, en

exposición Norte, que por sus características florísticas y climático-edáficas se

incluyeron en la asociación Carici sylvaticae-Fagetum.

La etapa serial del hayedo está constituida por matorrales de genisteas (Ge-

nista hispanica subsp. occidentalis, Genista legionensis) ricos en brezales de Erica

vagans en las zonas más húmedas y sombrías, que forman la asociación Lithodoro

diffusae - Genistetum legionensis, subas. ericetosum vagantis, que caracteriza

a estas laderas de suelos profundos y descarbonatados del monte de la Llama

y Monte de Valdediezma,

Desde el desfiladero de la Hermida, se puede acceder a Beges, por la cuenca

del río Corvera, y a Tresviso por la del río Urdon. En estos tramos, aparecen

como carácter reliquial encinas de carrasca, que Rivas Martinez et al. (1984)

definen como Cephalantero longifolit - Quercetum rotundifoliae. La etapa de

sustitución de estos encinares está formada por matorrales densos de arbustos

esclerófilos (Phillyrea media, P. latifolia) y caducifolios espinosos (Rosa semper-

virens), junto con algunas lianas (Smilax aspera, Tamus comunis). Una mayor

degradación puede conducir a la existencia de matorrales de Lithodoro diffusae -

Genistetum legionensis de carácter más xérico (Fig. 4).

Source - MNHN, Paris

114 E. FUERTES LASALA y E. MARTINEZ-CONDE

RESULTADOS

Las localidades donde se realizaron inventarios y se recolectaron los taxones

que se indican en el catálogo floristico son :

1 - Balneario de la Hermida.

2 - Cuenca del río Deva, Desfiladero de la Hermida, entre la Hermida y Urdén.

3 - Alrededores de la Central Eléctrica.

4 - Margen izquierda del río Urdon, hacia Tresviso.

5 - Margen derecha del río Urdon.

6 -Collado de la Hoja.

7 - Salto de la Cabra.

p- 28, ی‎ de Polysticho setiferi-Fraxinetum excelsioris, subas. tilietosum platy-

phyllae.

9 - Cuetodave.

10 - Monte de Valdediezma.

11 - El Dobrillo.

12 - Monte de la Llama.

3 - Minas de Mazarrasa, alrededores del Mancondiú (1999 m).

14 - Riega del Vau de los Lobos.

15 - Riega del Vau de las Vacas.

16 - Quintana.

17 - La Horcá de Entrelegua (Hayedo).

18 - Varga de los Mollares.

19 - Cuenca del Corvera.

20 - Alrededores de la Aldea.

Ver situación de las localidades en el territorio en la Fig. 1.

SINTAXONOMIA

El conspecto sintaxonómico se ha realizado segiin el criterio establecido por

v. Huebschmann 1986.

I. СІ. Tortulo-Homalothecietea sericei Hertel 1974

O. Schistidietalia apocarpi Ježek & Vondraëek 1962

Al. Schistidion apocarpi Ježek & Vondragek 1962

1. as. Orthotricho-Grimmietum pulvinatae Stodiek 1937

variante ctenidietosum mollusci Marstaller 1983

2. as. Pseudoleskeelletum catenulatae Ježek & Vondradek 1962

О. Ctenidietalia mollusci Šmarda & Hadad 1944

Al. Ctenidion mollusci Stefureac 1941

3. as. Tortello-Ctenidietum mollusci (Gams 1927) Stodiek 1937

subas. scapanietosum asperae (Neumayr 1971) Strasser 1972

O. Neckeretalia complanatae Ježek & Vondràtek 1962

Al. Neckerion complanatae Hadad & Šmarda 1944

4. as. Neckero-Anomodontetum viticulosi (Wisniewski 1929) Philippi

1965

subas. thamnobryetosum alopecuri Marstaller 1985.

Source : MNHN. Paris

VEGETACION BRIOFITICA EN ANDARA I 115

П. Cl. Fontinaletea antipyreticae v. Huebschmann 1957

О. Leptodictyetalia riparii Philippi 1956

Al. Rhynchostegion riparioidis Waldheim 1944

5. as. Rhynchostegietum riparioidis Gams 1927

III. Cl. Montio-Cardaminetea Braun-Blanquet & Tüxen 1943

O. Cardamino-Cratoneuretalia Maas 1959

Al. Cratoneurion commutati W. Koch 1928

6. as. Cratoneuretum commutati (Gams 1927) Walther 1942

7. as. Eucladietum verticillati (Allorge 1922) Giacomini 1951

IV. Cl. Adiantetea capillus-veneris Braun-Blanquet 1947

O. Adiantetalia Braun-Blanquet 1931

Al. Adiantion Braun-Blanquet 1931

8. as. Eucladio-Adiantetum Braun-Blanquet 1931

COMUNIDADES SAXÍCOLAS

Dado el predominio de rocas básicas en la zona estudiada, las comunidades

briofíticas saxícolas que hemos inventariado se incluyen en la clase Tortulo-

Homalothecietea sericei Hertel 1974. Dependiendo de su mayor o menor exposi-

ción a la luz solar y a las condiciones meso- y microclimáticas hemos reconocido

los órdenes Schistidietalia apocarpi que reune a las comunidades más xérofilas,

que se asientan sobre rocas calizas secas y soleadas; Neckeretalia complanatae

que engloba a las comunidades más esciófilas y mesófilas y el orden Ctenidietalia

mollusci Šmarda & Hadad que incluye a comunidades esciófilas y meso-xerófilas.

Orthotricho-Grimmietum pulvinatae Stodiek 1937

variante ctenidietosum mollusci Marstaller 1983

Sinecología y Sinfisionomía. — La comunidad es muy frecuente en rocas

calizas secas y soleadas de la vertiente Sur de Sierra de Beges y margen izquierda

del Urdon (piso colino a submontano) así como en las etapas degradadas de

Sierra de la Corta.

El conjunto de especies: Grimmia pulvinata, Orthotrichum anomalum, Schis-

tidium apocarpum y Tortula muralis forman socies pulvinares, discontinuas,

donde abundan las especies acrocárpicas. El único pleurocárpico, caracterís-

tico del orden es Homalothecium sericeum (Tabla 2).

Sindinámica y Sincorología. — En las rocas de bordes de caminos y sendas,

escasamente protegidas por el bosque, poco denso, hemos diferenciado la va-

riante ctenidietosum mollusci Marstaller 1983, de exigencias más mesofiticas, mar-

cando la transición hacia las comunidades mesoxerófilas de Ctenidietalia mol-

lusci.

El sintaxon se encuentra ampliamente representado en centroeuropa y en

la Región Mediterránea de la Península Ibérica (Fuertes & Alonso 1984, Guerra

Source : MNHN. Paris

116 Е. FUERTES LASALA у Е. MARTINEZ-CONDE

Altitud (1-10 m) 110. 120 72 74 73 70 60 91

Area (dm?) 4 41.22.28). RES ET 16

Orientación S — SW Е NE N NW NE S

Cobertura (%) 50 30 50 10 50 20 10 20

Inclinación (2) AREA do (455 دای‎ 207240

Número de orden 1 EEE a

Características de asociación:

Orthotrichum anomalum

Grimmia pulvinata

Diferencial de variante:

Ctenidium molluscum 2:3 2:3 1:2 2:8

Características de unid.sup.:

Schistidium apocarpum 1:2

12 2 +2 1:2

Orthotrichum cupulatum 1:2 1:

Encalypta vulgaris چڊ ي‎

Schistidium apocarpum confert.1:2 +2 1:2

Tortula muralis +1 +2 هب‎ +2

Tortella tortuosa 1:2 1:2 1:2 1:2

Ditrichum flexicaule +2 1:2 1:2 +2

Thuidium abietinum 1:2

Homalothecium sericeum 1:2 1:2 1:2 1:2

Tortula intermedia *2 ۵

Compañeras:

Grimmia orbicularis US +2

Didymodon fallax 1:2 1:2 +1

Hypnum cupressi forme +2 1:2

Bryum argenteum 1:2 +2

Campylium calcareum 3:2

Además: Tortula ruralis +2 еп 5; Leucodon sciuroides 1:2 en 5; Barbula

unguiculata +1 en 3; Rhytidium rugosum 1:1 en 2; Campylium chrysophyllum

1:1 еп 3; Eurhynchium praelongum +2 en 1; Homalothecium lutescens 1:2 en

Localidades: 1,2,7, Monte de la Llama

3, valle del Urdén

8, valle del Sobra

4,5,6, collado de la Hoja

Tabla 2. — Orthotricho-Grimmietum pulvinatae Stodiek 1937

variante Ctenidietosum mollusci Marstaller 1983

& Varo 1981). Se cita por primera vez en la Región Eurosiberiana de la Penin-

sula Ibérica. (En la Fig. 4, se indica la distribución del sintaxon en el territorio).

Pseudoleskeelletum catenulatae Jezek & Vondrátek 1962

Sincorología y sinecología. — Descrita para centroeuropa por Pilous (1961),

en nuestra zona se desarrolla sobre las calizas del piso altimontano y subalpino

de la vertiente Norte de Sierra de la Corta, cuya vegetación está formada por

matorrales de escaso porte (Lithodoro diffusae-Genistetum legionensis). Tam-

bién la hemos localizado, en medios más mesofíticos, del piso colino, sobre rocas

Source : MNHN. Paris

SeN

| SUBALP]NO.

MONTANO

COLNO |

Fig. 4 — Distribución de las comunidades briofiticas en el área estudiada y catena de la vege-

tación vascular. 1 (U), Oxyrrhynchietum rusciformis; 2 (0), Cratoneuretum com mutati;

3 (9), Eucladietum verticillati; 4 (@), Eucladio-Adiantetum; 5 (+), Orthotricho-Grimmie-

~ tum pulvinati; 6 (0), Pseudoleskeelletum catenulatae; 7 (2), Tortello-Ctenidietum mollus-

ci; 8 (A), Neckero-Anomodontetum viticulosi. 1, Polysticho-Fraxinetum excelsioris; la,

tilietosum platyphyllae; I1, Cephalanthero-Quercetum rotundifoliae; Ш, Lithodoro-Ge-

nistetum legionensis; Ша, ericetosum vagantis; IV, Carici-Fagetum; V, Daphno canta-

bricae-Arctostaphylletum uva-ursi.

Altitud (1-10 m) 90 67 120 140 150

Area (dm?) 4 4 B 9 4

Orientacién NE N N N NE

Cobertura (€) 40 70 40 50 80

Inclinaciém (2) 50 60 во 45 50

Número de inventario 1 2 3 4 5

Caracteríticas de asociación:

Pseudoleskeella catenulata 2:2 3:4 3:3 3:4 4:4

Características de un.sup.:

Schistidium apocarpum 1:2 пе 5

Tortula intermedia 2 2% +2

Encalypta streptocarpa 1:2 i

Lescurea incurvata 1:2

Tortula muralis Lon 2

Grimmia pulvinata 219 асе

Tortella tortuosa euo RUE

Homalothecium sericeum 2:2 1:2

Compañeras :

Bryum argenteum +2 +2 1:2

Bryum capillare +2

Hypnum cupressiforme 332 2%2

Tortella densa nz

Tortula ruralis 1:2 1:2

Además: Distichium capillaceum +2 en 5; Thuidium abietinum 1:2

en 3; Bryoerytrophyllum recurvirostrum +1 en 1; Tortula subula-

en 3; Ditrichum subulatum +2 en 2; Cladonia pyxidata 1:2

en 1.

Localidades: 1,3, el Dobrillo (Sierra de la Corta); 2, Sierra de

Beges (Collado de la Hoja); 4,5, Monte de la Llama (Sierra de la

Corta).

Tabla 3 — Pseudoleskeelletum catenulatae Ježek & Vondratek 1962

Source : MNHN, Paris

118 Е. FUERTES LASALA у Е. MARTINEZ-CONDE

calizas de desfiladeros y gargantas, cerca del cauce de los ríos (Tabla 3).

Sinfisionomía y sindinámica. — En la comunidad Pseudoleskeella catenulata

de color negruzco, contrasta con la blancura de la caliza, forma con Homalothe-

cium sericeum el estrato basal de la asociación. El resto de los taxones está

formado por briocaméfitos erectos (Schistidium apocarpum, Tortula muralis,

etc.) de distribución pulvinular sobre el sustrato basal.

Cuando las condiciones ambientales varian (aumento de temperatura, menor

humedad atmosférica, etc.) hacia un mesoclima más xerico, la comunidad es

sustituida por Orthotricho-Grimmietum pulvinatae (distribución en la Fig. 4).

Neckero-Anomodontetum viticulosi (Wisniewski 1929) Philippi 1965

Sinfisionomia y sinecologia. — Comunidad formada por grandes musgos

pleurocárpicos (Neckera complanata, Anomodon viticulosus) y hepaticocamé-

fitos foliosos (Porella platyphylla, Scapania aspera) de comportamiento basó-

Aret (шей) AU a ae A e dec

lo to yo so so 10 do 70 an eo po 70 soos 70 ви

täractentaticns ač niveiactdhı

A cuin жаға sd на ма ae "m

Laractertatses de subas

varacterfatices dé unid: superis

Weekend ce apu E aes

Korella pleyphylia sese in La m

eres مومس رنه‎ 12 ae m

Semistiarun spacarpum ê aa ve

ی‎ eragsinerviun us ЕН ne

Hlagiocnttla porettotdes за urs in m

ractithecium ilareosum m m

Encalypts ciliata а а

Tantun Cau ой ux en Ai Hueynchium hiana 112 en 9 -

Xen. ta}. dosthecius myomirotis ار همیب‎ am $3 arb

Tabla 4. — Neckero-Anomodontetum viticulosi (Wisniewski 1929) Philippi 1965.

subas. thamnobryetosum alopecurii Marstaller 1985

Source : MNHN, Paris

VEGETACION BRIOFITICA EN АМРАВА 1 119

filo y mesófilo. Se situa en lugares donde la vegetación cormofítica es densa.

Está muy bien representada en la vertiente norte de la Sierra de Beges, donde

el robledal espeso se encuentra mezclado con fresnos. En Sierra de la Corta se

situa dentro del hayedo en las laderas orientadas a septentrión (Tabla 4).

Sincorologia y sindinámica. — Bien representada en centroeuropa en el piso

montano (Wilczynska 1974, Szafran 1955, Marstaller 1985) amplía hasta Canta-

bria su área de distribución, aunque puede extenderse a toda la Región Euro-

siberiana de la Península.

En las rocas donde la exposición es N-NW, más ombrófila, así como en

fondos de dolinas donde se acumulan las nieblas, hemos diferenciado la subaso-

ciación : thamnobryetosum alopecuri Marstaller 1985, que presenta como espe-

cies diferenciales : Metzgeria conjugata, Conocephalum conicum, Lejeunea

cavifolia, Plagiomnium undulatum, Pedinophyllum interruptum. En las áreas

desforestadas o en bosques aclarados esta comunidad es sustituida por sintaxones

de Ctenidietalia mollusci (distribución en la Fig. 4).

Sintaxonomia. — La comunidad fué descrita por Szafran (1955) en las

montañas calizas del Jura (Polonia). Marstaller (1985), considera a esta comuni

dad como sinónima a la dada por Wisniewski (1930) como Anomodonto viti-

culosi-Leucodontetum sciuroides.

La subasociación thamnobryetosum alopecuri definida por Marstaller (1985),

sustituye a la variante del mismo nombre dada por Wilczynska (1974). Otros

autores (Smarda 1947) definieron a esta última subasociación como Neckero

complanatae-Thamnobryetum alopecuri.

En nuestra zona, la comunidad Neckero complanatae-Anomodontetum

viticulosi subas. thamnobryetosum alopecuri queda definida según el criterio

de Marstaller (1985), ya que Anomodon viticulosus preside todas las comuni-

dades como indicadora oceánica y nemoral.

Tortello tortuosae-Ctenidietum mollusci (Gams 1927) Stodiek 1937

Sinecología. — De exigencias esciófilas y mesoxerófilas, se desarrolla sobre

roquedos calizos del interior del bosque caducifolio, en zonas más o menos

soleadas. En nuestro territorio es la comunidad más frecuente del collado de

la Hoja a Cuetodave donde el robledal está más aclarado en esa explanada. En

el hayedo, se situa en los bloques calizos más expuestos a la luz, siempre en o-

rientación Norte, Nordeste o Noroeste. Es una comunidad típica del piso biocli-

mático montano (sub- y altimontano).

La subasociación scapanietosum asperae (Neumayr 1971) Strasser 1972,

se situa en las rocas del bosque con orientación N-NW de claras apetencias

nemorales y oceánicas (Inventario. Tabla 5).

Sinfisionomía y sincorología. — Está definida por briocaméfitos y pleuro-

cárpicos : Ctenidium molluscum, Tortella tortuosa у por Homalothecium

sericeum, Encalypta streptocarpa, Fissidens cristatus y Ditrichum flexicaule en

Source : MNHN, Paris

120 E, FUERTES LASALA y E. MARTINEZ-CONDE

moms i nre £e ға аі аз és an ns аз meon

Ares (ane) hod oe per tea n Oeo de

Camere 15) jo T6 wm 6 M) m X6 yo СЕ

Minero de orden НЧ mn

Vacacterfaticas de asociación:

carnet. de subasóciaeión:

Características de uni р

Ditrichum fiente aoz ne na

Trishostamun criepulun uo am us

Toeslothecium sericeun na an heu

Fiedioshille poretloides а nana nio on

н aro аа 20014

Gymnestomum seruginonim E а

Palena cristatue за ма ata 2

Teichostonum brachydontium "OR а

Schineidium apocerpus E а

Peainophylion interruptus шта

Madana. сверл лата ۳ 3

Canpy sun chrysophyllus uo ona

Mein mar dinatur T ha

Además: Lophozia collarin +i en 8; Scapania aequilube +1 en 61 Pohtia cruda +1 on 93 Campy ion

mtellatun 151 en 10: ‘on 11; Tortula ма

Teen 12) Campylium halleri «2 еп 8; Dicranum scoparium «1 en 1

Monte de Матаса целта: 3, Cuetoda

e del urdén (Cantabria).

Vertiente norte de Sierra de Begesi 4,

Tabla 5. — Tortello-Ctenidietum mollusci Stodiek 1937,

subas. scapanietosum asperae Marstaller 1981

las unidades superiores. Debido al carácter exocomófito particularmente acen-

tuado en nuestra zona, Ctenidium molluscum domina ampliamente la comuni-

dad. Bien representada en rocas calizas de la región mediterránea de la Península

Ibérica, amplía su areal hasta la Región Eurosiberiana.

COMUNIDADES ACUATICAS Y SUBACUATICAS

En nuestro territorio, dichas comunidades se incluyen en las clases Fontina-

letea antipyreticae v. Hübschmann 1957, Montio-Cardaminetea Braun-Blanquet

& Tüxen 1943 y Adiantetea Braun-Blanquet 1947.

Rhynchostegietum riparioidis Gams 1927

Sinecologia, sinfisionomia y sindinamica. — Muy frecuente en riachuelos de

montaña, cascadas naturales o artificiales sobre sustrato calizo o ácido. En ge-

neral, es pobre en nümero de especies y muy sensible a diversos factores limi-

tantes (contaminación, variaciones del pH, caudal de agua, etc.), por lo que

suele presentar un desarrollo fragmentario.

Source : MNHN. Paris

УЕСЕТАСЮМ BRIOFITICA EN ANDARA 1 121

La comunidad es frondosa, formada por briocaméfitos reptantes (Rhynchos-

tegium riparioides, Fontinalis antipyretica, Hygrohypnum luridum, etc.), que se

asientan en las rocas del lecho de los rfos (Urdon, Sobra, Corvera y arroyos de

montafia). Durante los meses de verano, al secarse los cursos de agua, se aprecia

el contacto con comunidades de Brachythecietum rivularis, mal representada en

cuanto a especies, en la zona. (Tabla 6).

Altitud (1-10 m) 35 45 38 38 а аз

Area (dm?) А HEEL 5

Orientacién NW NE NW Е NE Е

Cobertura (%) 80 70 60 80 зо во

Inclinación )۶( 70 45 45 90 70 60

Número de orden AM a Ne COM 4

Características de asociación:

Rhynchostegium riparioides 3:4 4:4 4:4 4:5 5:5 414

Caracteristicas de unid.superiores:

Fontinalis antipyretica 2:3 2:2 2:2

Hygroamblystegium tenax а. 1:2

Pellia epiphylla 2:3 2:3 2:2

Eurhynchium speciosum +2 1:1 1:1 1:2

Compañeras:

Rhizomnium punctatum 1:2 1:2

Brachythecium rivulare 1:2 32 1:2

Hygrohypnum ochraceum 1:2 1:2

Cinclidotus fontinaloides 2:2 2:2

Eurhynchium hians 1:2 2:2

Además: Hygrohypnum luridum 1:1 en 5; Cratoneuron filicinum 1:2 en 5;

Eurhynchium schleicheri +1 en 4

Localidades: 1,3, valle del rio Corvera; 2,4,5,6, valle del rio Urdón;

Tabla 6. — Rhynchostegietum riparioidis Gams 1927

Sincorologia y sintaxonomía. — Descrita para centroeuropa por Gams (1927),

v. Huebschmann (1953), Philippi (1956); presenta un areal muy amplio, desde

Escandinavia hasta los Pirineos, Macaronesia (Islas Canarias) y Canadá (Huebsch-

mann 1967). Se cita por primera vez en la región Eurosiberiana de la Península

Ibérica. Se incluye en la clase Fontinaletea antipyreticae (distribución en

la Fig. 4).

Cratoneuretum commutati (Gams 1927) Walther 1942

Sinecología. — Comunidad helo-higrófila y basófila muy común en el piso

colino y montano del área estudiada, se localiza desde los 100m de altitud

(Desfiladero de la Hermida) hasta los 1200 m en el Alto de la Llama (Sierra de

la Corta) siempre en cascadas, fuentes y rocas húmedas а lo largo del año. El pH

de las aguas oscila entre 7 a 8,5. (Tabla 7).

Source : MNHN. Paris

122 Е. FUERTES LASALA у Е. MARTINEZ-CONDE

Altitud (1-10 m) 67 65 60 111 120 118

Area (dm?) EEE

Orientación NE ON N NE N NE

Cobertura (4) то 90 70 90 90 вв

inclinación (2) 45 50 80 70 80 60

Número de orden 1 AE

Características de asociación:

Cratoneuron commutatum 3

Cratoneuron filicinum 1

Philonotis calcarea 2

550

Características de unid. super.:

Bryum pseudotriquetrum 1:2 uid 1:

Pellia endiviifolia

Aneura pinguis 1:2 2

Riccardia chamaedryfolia 2:2 1

Compañeras:

Gymnostomum aeruginosum 1:2

Eucladium verticillatum

Fissidens grandifrons

Lophozia badensis 138 +2

Conecephalum conicum

Marchantia polymorpha +2

Brachythecium rivulare 1:2 1:2

Además: Cirriphyllum crassinervium 1:1 en 6; Cephaloziella baumgart—

neri +2 en 6; Eurhynchium pumilum +1 en 3; Fissidens adianthoides +1

en 4; Soutbya tophacea 1:1 en 3.

Localidades: 1,2, Quintana; 3, valle del rio Urdón; 4,5, Monte de la

Llama; 5, Desfiladero de la Hermida (Valle del Deva), (Cantabria).

Tabla 7. — Cratoneuretum commutati (Gams 1927) Walther 1942

Sinfisionomía y sindinámica. — Forma céspedes densos de briocaméfitos

reptantes (Cratoneuron commutatum, С. filicinum) y hepatico-teréfitos (РеШа

endiviifolia, Aneura pinguis). Cuando la comunidad recibe un aporte hídrico

mayor, aparecen dominantes las especies reófilas Fissidens grandifrons, F.

rivularis, Cinclidotus fontinaloides, etc. Sí queremos destacar, la ausencia en

estas cascadas, con agua permanente a lo largo del año, de Fontinalis antipyre-

tica, que aparece en cursos de ríos, en aguas tranquilas, cerca de la desemboca-

dura. Por el contrario, si el aporte hídrico es menor, la comunidad es sustituida

por Eucladietum verticillati о Eucladio-Adiantetum.

Eucladietum verticillati (Allorge 1921) Giacomini 1951

Sinecología, sindinámica y sinfisionomía. — La comunidad se asienta sobre

paredes y rocas calizas rezumantes que proporcionan al agua un pH compren-

dido entre 6,5-8. (Tabla 8a). Si la humedad permanece constante a lo largo del

año, la comunidad evoluciona hacia Cratoneuretum commutati.

Source : ММНМ, Paris

VEGETACION BRIOFITICA EN ANDARA I 123

(а) (b)

Altitud (1-10 m) 35 38 68 65 10 21 8 10 12

Area (dm?) ВО Ша ۳3۲ 4 Mal tos 201 9 16

Cobertura (4) 60 70 90 90 во 80 90 90 70

Inclinación (2) 70 60 45 50 45 80 80 90 80

Orientación SE SW E Е NE NE NE SW NE

Número de orden CE 5.4. Re LE ee es

Caract. de asociacién:

Eucladium verticillatum 3:3 3:3 4:4 4:4 3:3

Adiantum capillus-veneris

ев

Caract.de unidades sup.

(Montio-Cardaminetea):

Cratoneuron commutatum 2:2

Cratoneuron filicinum

Pellia endiviifolia

Bryum pseudotriquetrum

Marchantia paleacea 7:3 258 s 1:2

EIS

Características de Adian-

tetea:

Soutbya tophacea 121

Didymodon tophaceus

Samolus valerandi +2 1:2

Hypericum nummularium +1 du 1:1

Pinguicula grandiflora

subsp. coenocantabrica EI 1:1

Compañeras:

Gymnostomum aeruginosum +2

Philonotis calcarea 1:1

Ricardia chamaedryfolia

Lophozia badensis

Fissidens limbatus var.

bambert geri

1:2

Además: Conocephalum conicum 1:1 en 7; Cephaloziella baumgartneri+i еп

5; Cirriphyllum crassinervium +2 en 9; Eurhynchium pumilum +1 en 9;

Schistidium apocarpum var.confertum +1 en 6.

Localidades: 1,2, cuenca del rio Urdón; 3,4, Quintana; 5,7,8,9, Desfi-

ladero de la Hermida; 6, cuenca del rio Corvera (Cantabria).

Tabla 8. — Eucladietum verticillati (Allorge 1922) Giacomini (a)

Eucladio-Adiantetum Braun-Blanquet 1931 (b).

En nuestro territorio, en los taludes que presentan períodos de sequía prolon-

gados (dos meses), la asociación Eucladietum verticillati se configura como

comunidad estable. Cuando las paredes colonizadas por Eucladietum verticillati

soportan menor insolación y mantienen la humedad más tiempo, se favorece

el desarrollo de la comunidad Eucladio-Adiantetum (Adiantetea).

Sintaxonomía. — Se incluye, al igual que el sintaxon precedente, en la alianza

Cratoneurion commutati, clase Montio-Cardaminetea Braun-Blanquet & Tüxen

1943 (distribución en la Fig. 4).

Source : MNHN. Paris

124 Е. FUERTES LASALA y Е. MARTINEZ-CONDE

Eucladio verticillati-Adiantetum capillus veneris Braun-Blanquet

Comunidad meso-higréfila, esciófila y basófila que se desarrolla sobre proto-

suelos de rocas y paredones calizos pudiendo tolerar periodos estivales de sequía

más o menos prolongados.

Sinfisionomia y sincorologia. — La comunidad está formada por briocamé-

fitos (Eucladium verticillatum) y el pteridofito Adiantum capillus veneris. La

comunidad alcanza su óptimo en paredones rezumantes de lugares sombríos,

protegidos por la vegetación arbustiva o arbolada de estos valles.

En nuestro territorio se extiende por el piso colino. Bien representada en el

Desfiladero de la Hermida y valles del río Corvera y Urdon, cerca de su desem-

bocadura en el río Deva. En la vertiente SW aparece cerca de las cascadas, que

en verano, mantienen cursos de agua; en las laderas NE, el suelo acusa sequía

en verano (Tabla 8b).

Sindinamica y sintaxonomia. — Cuando el aporte hídrico es mayor la comu-

nidad es sustituida por el sintaxon Cratoneuretum commutati. Se incluye en la

alianza Adiantion, clase Adiantetea capillus-veneris (distribución en la Fig. 4).

CATALOGO FLORISTICO

Se incluye, en este apartado, todos los briofitos mencionados en los inventa-

rios. Se indica numéricamente entre paréntesis las zonas donde se han recolec-

tado en el Macizo de Andara (Fig. 1). En algunos casos se precisa el sintaxon

que caracteriza la planta de referencia. La nomenclatura se rige por el criterio

de Düll (1983, 1984, 1985).

Aneura pinguis (L.) Dum. Característica de Cratoneurion commutati (1, 2,3);

Anomodon attenuatus (Hedw.) Hüb. (6, 8); A. viticulosus (Hedw.) Hook.& Tayl.

(6,7, 8, 17, 18); Apometzgeria pubescens (Schrank) Kuwah. (6, 12);

Barbilophozia barbata (Schmid. ex Schreb.) Loeske (8, 10, 11, 12); Barbula

unguiculata Hedw. (3, 4, 6); Brachythecium glareosum (Spruce) B., S. & G. var.

glareosum (6, 8); В. rivulare B., S., & G. (3, 5, 18, 19); B. rutabulum (Hedw.)

B.S. & G. var. rutabulum (6, 11, 18); Bryoerythrophyllum recurvirostrum (Нейм)

Chen (6, 11, 12); Bryum argenteum Hedw. (1, 3, 9, 12, 13); B. capillare Hedw.

(2,5, 6, 12); B. pseudotriquetrum (Hedw.) Gaertn, Meyer & Schreb, var. pseudo-

triquetrum (1,2, 3, 5, 14, 15);

Campylium calcareum Crundw, & Nyh. (8, 12); С. chrysophyllum (Brid.)

J. Lange (3, 5); С. halleri (Hedw.) Lindb. (6, 9, 12); С. stellatum (Hedw.) J.

Lange & C. Jens. var. stellatum (6, 14, 16); С. s. var. protensum (Brid.) Bryhn

ex Grout (10, 12); Cephaloziella baumgartneri Schiffn. (2); Cinclidotus fontina-

loides (Hedw.) P. Beauv. (1, 2, 3, 5, 18, 19); Cirriphyllum crassinervium (Тау!)

Loeske & Fleisch. (2, 3, 19); Cololejeunea calcarea (Libert) Schiffn. (12);

Conocephalum conicum (L.) Lindb. (1, 2, 5, 7, 11, 18); Cratoneuron commu-

Source : MNHN, Paris

VEGETACIÓN BRIOFÍTICA EN ANDARA I 125

tatum (Hedw.) б. Roth var. commutatum (1, 2, 3, 5, 12); C. filicinum (Hedw.)

Spruce var. filicinum (1, 2, 3, 18); Ctenidium molluscum (Hedw.) Mitt. var.

molluscum, Caracteristica de Ctenidietum mollusci (2, 5, 6, 12, 13);

Dicranum scoparium Hedw. (2, 3, 5, 12, 16); Didymodon fallax (Hedw.)

Zander (3, 4); D. tophaceus (Brid.) Lisa (1, 2); Distichium capillaceum (Hedw.)

B., $. & С. (13); Ditrichum flexicaule (Schwaegr.) Hampe, Característica de la

clase Tortulo-Homalothecietea sericei (2, 4, 6, 12, 18);D. subulatum Hampe (6);

Encalypta ciliata Hedw. (10, 12); Е. streptocarpa Hedw. (6, 11); Б. vulgaris

Hedw. (3, 4, 9); Eucladium verticillatum (Brid.) В., S. & С. (1, 2, 3, 5, 19);

Eurhynchium hians (Hedw.) Sande Lac. var. hians (5, 8); E. meridionale (B., S.

& G.) De Not. (2, 3, 5); E. praelongum (Hedw.) B., S. & G. var. praelongum (6,

10, 12); E. pumilum (Wils.) Schimp. (2, 5); E. schleicheri (Hedw.) Jur. (5); E.

speciosum (Brid.) Jur. (5, 18, 19);

Fissidens cristatus Wils. ex Mitt. (5, 16, 18, 19); F. grandifrons Brid. (2, 5);

Е. limbatus Sull. var. bambergeri (Schimp. ex Milde) Düll, Cratoneuretalia (1,

2, 3); Fontinalis antipyretica Hedw. subsp. antipyretica (2, 3, 5, 18);

Grimmia orbicularis Bruch ех Wils. (3, 1, 2, 4, 7, 10); С. pulvinata (Hedw.)

Sm., Característica de la clase Tortulo-Homalothecietea sericei (1, 2, 3, 4, 5,

6, 11, 13); Gymnostomum aeruginosum Sm. (3, 5):

Homalia trichomanoides (Hedw.) Brid. var. trichomanoides (2, 3, 5); Homalo-

thecium lutescens (Hedw.) Robins, (2, 3, 6, 12, 18);H. philipeanum (Spruce)

В., S. & С. (12); Н. sericeum (Hedw.) B., S. & G., Característica de la clase Tor-

tulo-Homalothecietea sericei; Hygroamblystegium tenax (Hedw.) Jenn. (3, 5,

19); Hygrohypnum luridum (Hedw.) Jenn. var. luridum (1, 14), H. ochraceum

(Turn. ex Wils.) Loeske (3, 5);

Isothecium myosuroides Brid. var. myosuroides (6, 8);

Lescuraea incurvata (Hedw.) Lawt. (12, 13); Leskea polycarpa Hedw. (1, 2,

5, 12, 13); Leucodon sciuroides (Hedw.) Schwaegr. var. sciuroides (6, 9, 10, 12,

17); Lophozia badensis (Gott. ex Rabenh.) Schiffn., en comunidades de Crato-

neurion (2, 3); L. collaris (Nees in Mart.) Dum., Característica de Ctenidietalia

mollusci (8, 10, 12);

Marchantia paleacea Bertol. (1, 2, 3, 5, 16); М. polymorpha L. emend.

Burgeff (2, 5, 6, 12); Metsgeria conjugata Lindb. (8); M. furcata (L.) Dum.

(8, 9, 10, 12, 17); Mnium marginatum (with.) P. Beauv. (1, 2, 10, 12, 15);

M. stellare Hedw. (6);

Neckera complanata (Hedw.) Hiib., Característica de Neckerion complanatae,

se extiende por Sierra de la Corta y Sierra de Beges; N. crispa Hedw., Caracte-

ristica de Neckeretalia complanatae;

Orthotrichum anomalum Hedw., Característica de Tortulo-Homalothecietea

sericei (1, 2, 3, 4); О. cupulatum Brid. var. cupulatum, Característica de Schisti-

dion apocarpi (1, 18, 19);

Source : MNHN, Paris

126 E. FUERTES LASALA y E. MARTINEZ-CONDE

Pedinophyllum interruptum (Nees) Kaal. (8, 10, 12); Pellia endiviifolia

(Dicks.) Dum., Caracteristica de las comunidades muscinales de Cratoneurion

y Adiantion (1, 2, 3, 5, 12); P. epiphylla (L.) Corda in Opiz (5, 6, 7); Philonotis

calcarea (В. & S.) Schimp. (2, 16); Plagiochila porelloides (Torrey ex Nees)

Lindenb., Característica de Tortulo-Homalothecietea sericei; Plagiomnium

rostratum (Schrad.) T. Kop. (5, 8, 14, 15); P. undulatum (Hedw.) T. Kop. (2,

3, 5, 6, 13); Plagiopus oederi (Brid.) Limpr. var. oederi (6, 12, 17); Platydicta

confervoides (Brid.) Crum (12); Pohlia cruda (Hedw.) Lindb. (12, 13); Porella

platyphylla (L.) Pfeiff.,Característica de Tortulo-Homalothecietea sericei; Preis-

sia quadrata (Scop.) Nees (1, 2, 5, 12); Pseudoleskeella catenulata (Schrad.)

Kindb. subsp. catenulata (2, 12, 13, 18);

Rhacomitrium elongatum (Ehr.) Erisvoll (12); Radula complanata (L.) Dum.

(9, 6, 8); Rhizomnium punctatum (Hedw.) Т. Кор. (1, 2, 3, 7, 10, 12); Riccar-

dia chamaedryfolia (With.) Grolle, Caracteristica de Cratoneurion commutati

(2, 3, 16); Rhynchostegium riparioides (Hedw.) Card., Característica de Fonti-

naletea antipyreticae (3, 4, 7, 19); Rhytidiadelphus triquetrus (Hedw.) Warnst.

(6, 12); Rhytidium rugosum (Hedw.) Kindb. (10, 11, 12, 17);

Scapania aequiloba (Schwaegr.) Dum. (8, 11, 12); S. aspera H. Bernet (6, 8,

11); Schistidium apocarpum (Hedw.) B. & S., Catacteristica de Tortulo-Homalo-

thecietea sericei (4, 6, 9, 11, 13, 18);S. apocarpum (Hedw.) B. & S. var. confer-

tum (Funck) Moell. (2, 18, 19); Scleropodium purum (Hedw.) Limpr. var.

purum (2, 5, 6, 12); Southbya tophacea (Spruce) Spruce, Característica de Cra-

toneurion commutati (2, 3, 19);

Thamnobryum alopecurum (Hedw.) Nieuwl. (8, 10, 12, 14, 15); Thuidium

abietinum (Hedw.) B., S. & G. (6, 9, 10, 12, 14); T. tamariscinum (Hedw.)

B., S. & G. (2, 5, 6, 12, 17): Tortella densa (Lor. & Mol.) Crundw. & Nyl. (12);

T. tortuosa (Hedw.) Limpr., Característica de Tortulo-Homalothecietea sericei;

Tortula intermedia (Brid.) De Not. (6, 11); T. muralis Hedw. (1, 2, 3, 5, 6, 10);

T. ruralis (Hedw.) Gaertn, Meyer & Schreb. (6, 10, 11, 14, 15); T. subulata

Hedw. (1,2, 5); Trichostomum crispulum Bruch (5, 7); T. brachydontium Bruch

(12,16, 17).

BIBLIOGRAFIA

DUELL R., 1985 — Distribution of the European and Macaronesian Liverworts (Hepatico-

phytina). Bryol. Beitr. 2 : 1-115.

DUELL R., 1984-1985 — Distribution of the European and Macaronesian mosses (Bryophy-

tina), Bryol. Beitr. 4-5 : 1-233.

FUERTES E. & ALONSO M., 1984 — Estudio fitosociológico de las comunidades de brio-

fitos, saxícolas y epifitas de la Hoz de Beteta (Cuenca, España). Webbia 38 : 695-703.

Source - MNHN. Paris

VEGETACION BRIOFITICA EN ANDARA I 127

GAMS H., 1927 — Von den Follatéres zur Dent de Morcles. Beitr. Geobot. Landesaufn.

Schweiz 15 : 1-760.

GUERRA J. & VARO J., 1981 — Datos sobre la clase Tortulo-Homalothecietea sericei en

las Sierras Béticas (Andalucía, España). Phytocoenologia 9 (4) : 443-463.

HUEBSCHMANN A. v., 1967 — Uber die Moosgesellschaften und das Vorkommen der

Moose in den übrigen Pflanzengesellschaften des Moseltales. Schriftenreihe Vegetationsk.

2: 63-121.

HUEBSCHMANN A. v., 1986 — Prodromus der Moosgesellschaften Zentraleuropas. Stutt-

gart : J. Cramer; 413 p.

JULIVERT М. & FONTBOTE J.M., 1972 — Mapa tectónico de la Península Ibérica y Balea-

res. Inst. Geol. y Min. de España. Mapa y Memoria.

LEVIER R., 1880 — Mousses récoltées еп 1878 en Espagne et au Portugal. In : Leresche L.

& Levier Е., Deux excursions botaniques dans le Nord de l'Espagne et le Portugal en

1878-1879. Lausanne : Imp. Bridel. Pp. 164-179.

MARSTALLER J., 1981 — Die Moosgesellschaften des Naturschutzgebietes ‘Leutratal’

bei Jena. Veróff. Mus. Stadt. Gera 9 : 41-46.

MARSTALLER R., 1985 — Zur Verbreitung und Soziologie von Pedinophyllum interrup-

tum (Nees) Kaal. Gleditschia 13 (2) : 289-309.

MARTINEZ-CONDE E., 1986 — Contribución al estudio de la flora y vegetación briofitica

del Macizo de Andara : valles del Sobra, Urdón, Corvera y Deva (Picos de Europa, Can-

tabria). Tesis Doctoral. Univ. Complutense. Madrid. 274 p.

PHILIPPI G., 1956 — Einige Moosgesellschaften des Súdschwarzwaldes und der angrenzen-

den Rheinebene. Beitr. Naturk. Forsch. Südwestdeutschl, 15 : 91-124.

PILOUS Z., 1961 — Mechová vegetace Demánovské doliny y Nizych Zazrách. Rozpr. Ces-

koslov. Akad. Véd. 71 (2) : 1-99.

RIVAS MARTINEZ S., DIAZ T.E., PRIETO J.A.F., LOIDI J. & PENAS A., 1984 — La

vegetación de la alta montaña cantábrica. Los Picos de Europa. León (España) : Ed.

Leonesas. 295 p.

ŠMARDA J., 1947 — Mechova a ligejnikova společenstva. CSR . Cas. Zemsk, Mus. Brne 31:

39-88.

SZAFRAN B., 1955 — Mechy Jury Krakowsko-Wielunskiej z uwzgednienem rezerwatow

przyrody. Ochr. Przyr. 23 : 219-254.

WILCZYNSKA W., 1974 — Flora mchéw i zbiorowisk mszakéw Gér Kaczawskich. Monogr.

Bot. 44 : 3-111.

WISZNIEWSKI W., 1930 — Zespoly mszaków epifitycznych Polski ze szczególnym uwzgled-

nieniem Puszcz Bialowieskiej. Bull. Int. Acad. Polon. Sci., Cl. Sci. Math., Sér. B, Sci. Nat.

3 : 293-342.

Source : MNHN. Paris

Cryptogamie, Bryol. Lichénol, 1988 9 (2) : 129-135 129

CULTURE STUDIES ON

PLAGIOCHASMA APPENDICULATUM LEHM. ET LINDENB.

AND REBOULIA HEMISPHAERICA (L.) RADDI POPULATIONS

OF PACHMARHI (CENTRAL INDIA)

IN RELATION TO pH ON A COMPARATIVE BASIS

K.S. VISHVAKARMA and A. KAUL*

ABSTRACT. — Comparative studies on the vegetative growth and spore germination of

Plagiochasma appendiculatum and Reboulia hemisphaerica from central India (Pachmarhi)

in response to different pH have been conducted. In culture both species showed vegetative

growth at pH 3.0 to 7.0, Maximum numbers and dry weight increase of newly formed

branches were obtained at pH 6.0. Spores of P. appendiculatum germinated at pH 4.0 to

7.0 and spores of R. hemisphaerica at pH 5.0 to 7.0. Maximum germination in both species

was obtained at pH 6.0.

INTRODUCTION

Kachroo (1954) stated that members of the Rebouliaceae have their origin

in the Himalayas. If so, then P. appendiculatum and R. hemisphaerica have

migrated from the Himalayas to the different bryogeographical regions of the

Indian sub-continent as suggested by Pande (1958). Their wide occurrence indi-

cates that they have adjusted well under different environmental conditions.

Hydrogen ion concentration (pH) is one of the important physical factors of

the environment that influence the distribution of bryophytes (Steere 1976).

Influence of pH on corticolous bryophytes in culture was studied by Pitkin

(1973) and correlated with the distribution pattern of bryophytes in Britain.

Effect of pH on the growth of some hepatics has been studied by Voth (1943),

Machlis (1962) & Woodfin (1976) and on mosses by Ikenberry (1936), Heitz

(1942), Benson-Evans (1953), Noguchi & Miyata (1957), Sheridan & Rosen

Streter (1973) and others.

* Department of Botany, Madhav Science College, Vikram University, Ujjain 456010, India.

Source : MNHN. Paris

130 K.S. VISHVAKARMA and А. KAUL

In India the effect of physical factors on growth and spore germination of

hepatics have been studied by Chopra & Sood (1973), Kaul (1974), Bhavsar

(1980), Vishvakarma (1981), Patidar (1982). In the present paper an attempt

has been made to determine the influence of pH on growth and spore germina-

tion in population of P. appendiculatum and R. hemisphaerica adapted to the

conditions in central India at Pachmarhi.

MATERIAL AND METHODS

Fresh thalli and mature sporophytes of P. appendiculatum and R. hemisphae-

rica were collected in Pachmarhi in October and February respectively. They

were brought to Ujjain in polythene bags. Thalli were washed in running water.

Three replicates of five thalli (soil free) each were kept in petri-dishes containing

equal amounts of sterilized sand. Natural soil suspension was prepared in distil-

led water in a ratio of 1:10. It was allowed to remain for 24 hours, then main-

tained at different pH ranging from 2.0 to 9.0 with the help of phosphate

buffer and used as a culture medium for growth and spore germination experi-

Petri-dishes with thalli were kept in a closed desiccator at a relative

ity of 80 to 85%. Results in Table 1 are based on 60 days old cultures

and represent the means of the 3 replicates.

During the experiment on spore germination, spores of five capsules were

removed in a cavity block. This spore mixture was used in 3 replicates. Experi-

ment on spore germination was conducted in sterilized petri-dishes 2” in dia-

meter, containing the culture medium. Data in Table 2 represent average germi-

nation percentage of 3 replicates based on 15 days old cultures.

Culture experiments were conducted in laboratory : the temperature was

20°C + 3°C, light intensity, 1500 + 200 lux for growth experiments and 800

+ 60 lux for spore germination. Analysis of variance were performed on the

data.

Number of Fresh weight uf Dry weight of

newly formed newly formed newly formed

pl newly formed branches branches branches

branches (m9) (mg)

в» Re pe Re pr Re pe RY

3.0 60 40 7 5 5.0 5.4 T RIED

4.4 80 60 7 6 11.0 7.3 ZA t

5.0 100 80 10 E 15.9 8.2 ف‎ ау

6.0 100 100 20 16 64.0 56.5 92% 72

7.0 во 100 ви 9.5 18.0 Tio 59

8.0 = 4 - E Е E E "

Table 1. — Effect of pH on growth. P* - P. appendiculatum, R* - R. hemisphaerica.

Source : MNHN, Paris

PLAGIOCHASMA APPENDICULATUM AND REBOULIA HEMISPHAERICA 131

OBSERVATIONS

It is seen from Table 1 that thalli of P. appendiculatum and В. hemisphaerica

showed development of newly formed branches in pH ranging 3.0 to 7.0. Be-

yond that thalli failed to develop new branches. In P. appendiculatum 100 %

FIG. IB

Fig. 1 A — Development of newly formed branches in P. appendiculatum at different pH.

Fig. 1 B — Development of newly formed branches in R. hemisphaerica at different pH.

A : at pH 3.0. B :at pH 4.0. C : at pH 5.0. D : at pH 6.0. Е: at pH 7.0. Diagrams are

drawn with camera lucida from one of the replicates.

Source : MNHN. Paris

132 K.S. VISHVAKARMA and A. KAUL

thalli showed development of new branches at pH 5.0 and 6.0, whereas іп R,

hemisphaerica it was at pH 6.0 and 7.0. In both species maximum number

of newly formed branches occurred at pH 6.0; fresh and dry weight were maxi-

mum at pH 6.0.

100 7

GERMINATION (7)

9 — Р.А PPENDICULATUM

O-- R.HEMISPHAERICA

FIG.2

Fig.2. — Spore germination of P. appendiculatum and R. hemisphaerica at different pH.

Germination (%) at pH

3.0 4.0 5.0 6.0 7.0 8.0

P - 19.0 56.6 100.0 25.6 =

34.6 92.3 28.3 -

Table 2. — Effect of pH on spore germination. P* - P. appendiculatum, R* - К. hemisphae-

rica.

Source : MNHN, Paris

PLAGIOCHASMA APPENDICULATUM AND REBOULIA HEMISPHAERICA 133

It is seen from Table 2 that spores of P. appendiculatum germinated at pH

4.0 to 7.0, whereas spores of R. hemisphaerica, at pH 5.0 to 7.0. In both species

maximum germination occurred at pH 6.0.

Source of Degree of Freedom Sum of squares Мей sun of equator [TET

Rati Peur E С 2

ШЫТ № رو‎ mo и мм

% 45.05 50.02 5.99

within py аш pur mono»

Totar * 1 CEN

Table 3. — Analysis of variance. Effect of pH on growth. P* - P. appendiculatum, R* -

R. hemisphaerica.

ТЕТІ

eow

Source of бегаа of freedom Sum of Squares Mean sum of Squares

Саа 2 0 ” 3 ” r

Due to ph 7 2 F0 614.09 6096.77 IITA 227,09 552,0 3.99 10.92

Within он D B мала Ann 18.06 3.66

Ten " D 13474 .07 6108.89

Table 4. — Analysis of variance. Effect of pH on spore germination. P* - P. appendiculatum,

R* - R. hemisphaerica.

ganea grer RE — JET son eem GE зв cos ке)

Table 5. — Multiple range tests. Note : Any two means not underscored by the same line

are significantly different. Any two means underscored by the same line are not signifi-

cantly different. A : 3.0 pH. B : 4.0 pH. C : 5.0 pH. D : 6.0 pH. E : 7.0 pH. DW : Dry

weight (mg). G : Germination (%).

Analysis of variance revealed that the effect of pH on growth and spore

germination of P. appendiculatum and R. hemisphaerica is significant.

DISCUSSION

Bryophytes are small plants and microclimatic factors play an important

tole in their growth and distribution. Phenotypic plasticity and acclimatization

Source : MNHN. Paris

134 K.S. VISHVAKARMA and А. KAUL

have played a major role in the adaptation of certain mosses and hepatic species

to climatic condition (Longton 1979). P. appendiculatum and R. hemisphaerica

are widely distributed under different ecological conditions in the Indian sub-

continent. In central India at Pachmarhi P. appendiculatum is much more

common than R. hemisphaerica. Both species show much similarity in their

life cycles. Their populations propagate by vegetative means as a result of the

development of new branches.

Physical factors of the environment play an important role in the growth

of bryophytes. Allsopp (1967) has reviewed regeneration and development of

newly formed branches in many thalloid forms of bryophytes in culture. In the

present study it is seen that P. appendiculatum and R. hemisphaerica popula-

tions of Pachmarhi are well adjusted to an acidic range of pH (3.0 to 7.0) for

vegetative propagation; maximum number of branches and dry weight in both

species were obtained at pH 6.0. Chopra & Sood (1973) observed optimum

growth in Riccia crystallina at pH 6.5. Voth (1943) also observed better growth

of Marchantia polymorpha at acidic pH. Alkaline pH inhibited the growth of

Sphaerocarpos donnelli (Machlis 1962). Patidar & Kaul (1984) found that in

Riccia discolor, the number of newly formed branches and their dry weight

production were maximum at pH 4.5.

Spores are formed in abundance in both species (reproductive capacity is

very high, Vishvakarma 1981). Spores of В. hemisphaerica are adjusted to a nar-

rower range of pH for germination than those of P. appendiculatum. pH appears

to be one of the important limiting factors for the growth and spore germi-

nation of both species. Inoue (1960) observed that spores of R. hemisphaerica

germinate within pH 5.0 to 8.0, But higher germination percentage was obtained

in acidic pH. Kaul (1974) studied the influence of pH on the spore germination

of Plagiochasma intermedium collected from wall and rocks. The spores collec-

ted from rocks revealed germination within pH 3.0 to 6.0. Studies conducted

by Shukla (1977), Bhavsar (1980), Patidar (1982) on the vegetative propagation

and spore germination of certain Marchantiales also revealed that liverworts

of Pachmarhi are better adjusted to acidic pH rather than to basic pH for growth

and spore germination.

Authors are grateful to the Principal, Dr. S.L. Chhajlani, Madhav Science College, for

providing necessary facilities.

REFERENCES

ALLSOPP A., 1967 — Regeneration and polarity in the Marchantiales. Phytomorphology

17 : 364-370.

BENSON-EVANS K., 1953 — Some notes on spore germination in Mnium hornum Hedw.

Trans. Brit. Bryol. Soc. 2 :291.

Source : MNHN, Paris

PLAGIOCHASMA APPENDICULATUM AND REBOULIA HEMISPHAERICA 135

BHAVSAR S., 1980 — Studies on bryophytes of Ujjain and some aspects of experimental

studies on Plagiochasma intermedium L. et G. Ph. D. Thesis, Vikram University, Ujjain.

CHOPRA В.М. & SOOD S., 1973 — In vitro studies in Marchantiales 1. Effects of some

carbohydrates, agar, pH, light and growth regulators on the growth and sexuality in

Riccia crystallina. Phytomorphology 23 : 230-244,

HEITZ E., 1942 — Die Keimenden Funaria Sporen als physiologisches Versuchsobjekt.

Ber. Deutsch. Bot. Ges. 60 : 17-27.

IKENBERRY G.J., 1936 — The relation of Hydrogen ion concentration to the growth and

distribution of mosses. Amer. J. Bot. 3 : 271-279.

INOUE Н., 1960 — Studies in spore germination and the earlier stages of gametophyte

development in the Marchantiales. J. Hattori Bot. Lab. 23 : 148-191.

KACHROO Р., 1954 — Distribution of the Rebouliaceae in India. The Bryologist 57 :

159-166.

KAUL А., 1974 - Some aspects of spore germination of liverworts. J. Hattori Bot. Lab.

38 : 283-297.

LONGTON R.E., 1979 — Climatic adaptation of Bryophytes in relation to systematics. In +

G.CS. Clarke & Ј.С. Duckett, Bryophyte Systematics, London & New York : Academic

Press. Pp. 511-531.

MACHLIS L., 1962 — The effects of minerals salts, glucose and light on the growth of the

liverwort, Sphaerocarpos donnelli, Physiol. Pl, 15 : 354-362.

NOGUCHI А. & MIYATA І., 1957 — Sporelings and regenerants in some mosses. Kuma-

moto J. Sci., Ser. B., Sect. 2, 3 :1-19.

PANDE S.K., 1958 — Some aspects of Indian Hepaticology. J. Indian Bot. Soc. 37 : 1-27.

PATIDAR K.C., 1982 — Ecological studies on Ricciaceae. Ph. D. Thesis, Vikram University,

Ujjain.

PATIDAR К.С. & KAUL A., 1984 — Effect of pH on the growth of Riccia discolor L. et

L. Hikobia 9 : 61-63.

PITKIN P.H., 1973 — Aspects of the Ecology and Distribution of some Widespread Corti-

colous Bryophytes. M. Phil. Thesis, Univ. Oxford.

SHERIDAN R.P. & ROSENSTRETER R., 1973 — The effect of hydrogen ion concentra-

tion in simulated rain on the moss Tortula ruralis (Hedw.) Sm. Bryologist 76 :168-173.

SHUKLA R.B., 1977 — Ecological studies of two liverworts. Ph, D. Thesis, Vikram Uni-

versity, Ujjain.

STEERE W.C., 1976 — Ecology, Phytogeography and Floristics of Arctic Alaskan bryo-

phytes. J. Hattori Bot. Lab. 41 : 42-72.

VISHVAKARMA K.S., 1981 — Ecological studies of Plagiochasma appendiculatum Lehm.

et Lindenb. and Reboulia hemisphaerica (L.) Raddi on comparative basis. Ph. D. Thesis,

Vikram University, Ujjain.

VOTH P.D., 1943 — Effect of nutrient solution concentration on the growth of Marchantia

polymorpha. Bot. Gaz. (Crawfordsville) 104 : 591-601.

WOODFIN C.M., 1976 — Physiological studies on selected species of the liverwort family

Ricciaceae. J. Hattori Bot. Lab. 41 : 179-183.

Source : MNHN. Paris

Rem نی و‎ Фи eer À

+ ac en ond

Cryptogamie, Bryol. Lichénol. 1988 9 (2) : 137-139 137

CLADONIA GALINDEZII,

A NEW ANTARCTIC LICHEN SPECIES

D.O. OVSTEDAL*

ABSTRACT. — Cladonia galindezii Qvst., sp. nov., from the western antarctic Galindez

Island is characterized by the densely tufted primary thallus of upright, narrow lobes, the

cortical hyphae which are anticlinal to the surface, and the content of porphyrilic acid and

atranorin. No podetia were found.

When examining lichen collections from some western antarctic islands, I

came across some specimens of a Cladonia which could not be identified with

any known species. As there has been collected many specimens from two is-

lands, the material has been regarded as sufficient for the description of a new

species.

Cladonia galindezii Qvst. sp. nov.

Thallus primarius confertim caespitosus, lobis ligulatis, longitudine 8-10mm,

crassitudine 1-1.2 mm, subramosis, subinvolutis, ochraceis; hyphae corticales

perpendiculares; podetia et pycnidia ignota; acidum porphyrilicum et atra-

norinam continentia.

Primary thallus in dense tufts (Fig. 1), up to 10cm diam., and 6cm high,

pale yellow-brown (ochraceous) with a greenish tinge. Individual squamules

ascending, ca. 1 mm wide, usually partially involute, often finger-like or irregu-

larly divided in upper part. Upper side weakly tuberculate, cortex 60-704m

thick, composed of anticlinal hyphae with rather large lumina (Fig. 2). Lower

surface ecorticate, white to grey.

Type : Antarctica, West Graham Land, Argentine Islands, Galindez Island,

65°15’ S, 64°16’ W, from dry depression among rocks on north-facing inland

cliff slope, 1935, Brit. Graham Land («Penola») Exped. 1934-37, no. 1108

(BM-holotype). Other collections : 1) as type, no. 1108 a (BM); 2) West Graham

Land, Wiencke Island, Port Lockroy, Goudier Islet, halfway up island, alt. ca.

6 m, in crevice іп S-facing granodiorite face, 1944, LM. Lamb, Op. Tabarin

no. 1996 (BM).

* ARBOHA, University of Bergen, Allégt. 41, N-5007 Bergen, Norway.

Source : MNHN. Paris

138 D.0. QVSTEDAL

Fig. 12. — Cladonia galindezii Фуз. 1 : habitus (Holotype). Scale : 2 em. 2 : transverse

section of thalline lobe. Scale : 25 Um.

DISCUSSION

The anatomy of C. galindezii generally conforms with that of the primary

thallus of a typical Cladonia, except that the hyphae of the upper cortex are

strongly gelatinized and distinctly oriented towards the surface. For comparison

sections of the cortex of the primary thallus of some occasional Cladonia species

were examined. The cortex varies from typically paraplectenchymatous tissue

with + isodiametric cells, as in C. subcervicornis (Vain.) Kernst. to tissues with

hyphae vaguely oriented towards the surface, as in С. macrophylla (Schaer.)

Stenh. and С. turgida Hoffm., but in no species a cortex with hyphae with

such a distinct orientation towards the surface as in C. galindezii was found.

C. galindezii contains the B-orcinol p-depside atranorin and the dibenzofuran

porphyrilic acid, which have different precursors in their biochemical pathways

(Huovinen & Ahti 1982). In the genus Cladonia, dibenzofurans appear to occur

only in the group Cocciferae (Huovinen & Ahti 1982) and C. strepsilis (Ach.)

Vain., and the combination of atranorin and porphyrilic acid appears to be

previously unknown in the genus. A survey through recent papers on Cladonia

from southern Chile and New Zealand (Ahti & Kashiwadani 1984, Archer &

Bartlett 1986, Galloway 1985) did not reveal any taxon which could be identi-

fied with C. galindezii.

ACKNOWLEDGEMENTS. — I am indebted to Mr. T. Ténsberg, Bergen, for determi-

nation of the chemical content and for comments on the manuscript, and to Professor R.

Santesson, Uppsala and Professor T. Ahti, Helsinki, for comments on some of the speci-

mens.

Source - MNHN, Paris

CLADONIA GALINDEZII, SP, NOV. 139

REFERENCES

АНТІ Т. & KASHIWADANI H., 1984 — The Lichen Genera Cladia, Cladina and Cladonia

in Southern Chile. In : Inoue H., Studies on Cryptogams in Southern Chile. Tokyo.

Pp. 125-149.

ARCHER A.W. & BARTLETT J.K., 1986 — New species and distributions of the lichen

genus Cladonia in New Zealand together with a revised key. New Zealand J. Bot. 24 :

581-587.

GALLOWAY D.J., 1985 — Flora of New Zealand Lichens. Wellington : Government Pu-

blishing. 662 p.

HUOVINEN K. & AHTI T., 1982 — Biosequential patterns for the formation of depsides,

depsidones and dibenzofurans in the genus Cladonia (lichen-forming ascomycetes).

Ann. Bot. Fenn. 19 : 225-234,

Source : MNHN. Paris

Cryptogamie, Bryol. Lichénol. 1988 9 (2) : 141-147 141

LICHENS OF MADAGASCAR :

THE PYXINACEAE (syn. PHYSCIACEAE)

A. APTROOT*

ABSTRACT. — Thirty-six species of the lichen family Pyxinaceae are reported from Mada-

gascar, including Pyxine glaucovirescens (Nyl.) Aptroot comb. nov. Affinities with the flora

of adjacent regions are briefly discussed.

Since a long time, Madagascar is known to botanists as a remarkable environ-

ment for plantlife with a very high degree of endemism. Unfortunately, reports

on the lichen flora and vegetation are very scarce, due to the fact that most

lichen collecting has not been carried out by lichenologists but by non experts.

During april and may, 1984 I have been in the opportunity to make a field

trip to Madagascar and to collect lichens on many sites all over the island. This

is the first paper dealing with material from this trip, treating the Pyxinaceae

(with the exception of ВиеШа 5. 1.). Some material from other collectors has

been included, as well as all literature references found.

In order to facilitate spotting of the localities a brief account on the main

collecting-sites from the field trip is given below. Unless otherwise stated, all

collections are made by A. Aptroot and R.V. Hensen in april/may 1984 and

preserved in the private herbarium of the author. To some species, taxonomical

notes or other remarks are added. Keys and descriptions of the species cited can

be found in Swinscow and Krog (1975a, 1975b, 1976, 1978), Moberg (1983,

1986) and Aptroot (1987). The material of Rinodina has been examined by Dr.

Н. Mayrhofer. АП material cited has been examined by me, unless otherwise

stated. In addition type specimens of most of the species have been investigated

(Aptroot 1987).

A comparison with the Pyxinaceae floras of other areas shows a striking

similarity with the flora of East Africa; nearly all species reported here from

Madagascar are already known from there. The similarity with the flora of

Australasia is less significant. Thus, the Pyxinaceae flora is not believed to repre-

sent a Gondwana-element but has probably been established more recently. No

* Institute of Systematic Botany, Heidelberglaan 2, Utrecht. The Netherlands.

Source : MNHN, Paris

142 A. APTROOT

species seems to be endemic, contrary to several other thoroughly studied groups

of cryptogams, е. в. Musci (Crosby et al. 1983), Cololejeuneoideae (Tixier

1985), Cladonia (Des Abbayes 1948) and Parmeliaceae (Des Abbayes 1956).

This remarkable fact may be explained by the preference of most Pyxinaceae for

open or even artificial habitats. About 80 % of the species cited here are known

to have a wide tropical or even pantropical distribution (Awasthi 1975, Kuro-

kawa 1962 & 1973, Aptroot 1987).

Within Madagascar some different distribution patterns may be recognized.

The majority of the species occurs in the relatively humid central part of the

country, e. g. all Heterodermia spp. Some species are more or less restricted to

the Northern part of the country, usually the coastal regions, е. g. Dirinaria

aegialita and D. picta. Other species are restricted to the extremely dry Southern

area, e. g. Dirinaria complicata and D. confluens.

COLLECTING LOCALITIES

1. Nosy Be, an islet NW of Madagascar, Om. ۲ 139228. Mangrove forest along

the coast. А А

2. Nosy Komba, small islet between Nosy Be and Madagascar, Om. 48 2ГЕ 13 27%.

Mangrove forest along the coast. % v

3. Col de Radama, 60km S of Antalaha, 500m. 5001۴ 15 165, Primary tropical

lowland forest, the largest still existing on Madagascar. |,

4. Maroantsetra, E coast of Madagascar, Om. 49 45'E 15726%. Wooden cabin near the

beach, wind-exposed. Ж

5. Foulpointe (= Mahavelona), E Coast of Madagascar, Om. 49°29'Е 17 4178. On Philip-

pia (Ericaceae) in sandy dune area. 5 5

6. Tamatave, E coast of Madagascar, Om. 49%25'E 18°10'S. Palm stems in the coastal

area of the town. 6 5

7. Tle Ste Marie, an islet E of Madagascar, Om. 49°52'E 16 58'S. Palm stems in the coastal

area of Ambodifototra. 5 f.

8. Ambohimanga, central highlands, 1550m. 47"38'E 187455. On clay along forest

track in forest relict, ۳ ۳

9. Ambohidratrimo, central highlands, 1300 m. 47^26'E 187495. On granite rock out-

crop in forest relict. р к

10. Antananarivo (= Tananarive), the capital, 1350 m. 47^31'E 18 5615, On various trees

in parks in the E suburbs. 7 Я

11. Angavokely mountain near Carion, E of Antananarivo, 1550 т. 47°43'E 18 55'S.

On granite boulders in open area. ۳ Pa

12. Périnet (= Andasibe), on the E facing slope of Madagascar, 950 m. 48 16'E 18 56%.

On various trees in primary tropical mountain forest, ^

13. Col de Tapia, 45km М of Ambositra, 1500 m. 47 07'E 20^ 16S. On Uapaca trees in

dry forest type called transition forest. — ۳

14. Ambositra, central highlands, 1400 m. 47°14Е 209317, On various trees in cultivated

area.

15. Ambalamanakana, 30 km S of Ambositra, 1800m. 47°07°E 20° 51'S. On various trees

in undisturbed cloud forest. 1 n

16. Fianarantsoa, central highlands, 1250 m. 47°03'E 21°26'S. On various trees in cultiva-

ted area.

17. lfaty,25 km N of Tulear, Om. 43"38'E 23 1 LS. Qn various trees in semidesert woods.

18. Ankilibe, 10km SE of Tulear, Om. 43 46'E 23 25S. On various trees in semidesert

woods.

19. Betioky, Southern province, 300 m. 44”21'E 2374495. On deciduous trees in savannah.

Source : MNHN, Paris

PYXINACEAE OF MADAGASCAR 143

Dirinaria aegialita (Afz. in Ach.) Moore — loc. 2, Petit s. n. (PC); loc. 4,

13141; loc. 6, 13087; loc. 7, 13098; loc. 10, Grandidier s. n. (PC); Antanimoro,

near Fort Dauphin, Decary s. n. (PC).

This species occurs predominantly in coastal areas in the Northern part of the

country. The Grandidier specimen is cited by Awasthi (1975) as D. consimilis

(Stirt.) Awashi and erroneously said to be preserved in REN.

D. applanata (Fée) Awasthi — loc. 10, Lemaitre s. n. (H); Benoist 1074 (LD,

not seen, cited by Awasthi 1975).

This species has not been found recently even though the original locality

has been visited.

D. complicata Awasthi — loc. 17, 12604; loc. 18, 12566; loc. 19, 12586,

12588, 12591; Ambanja, Pervillé 561 (H-NYL 31792).

This species is apparently restricted to the dry Southern area, where it is not

uncommon on deciduous trees and shrubs, It also occurs in the spines of Didiera

spp. together with Roccella montagnei Mont. It is known from the African

continent only.

D. confluens (Fr.) Awasthi — loc. 10, 12314; loc. 18, 12565; Antanimoro,

near Fort Dauphin, Decary s. п. (PC); Ampanihy, Humbert s. п. (PC).

This species occurs predominantly on deciduous trees in the South.

D. picta (Sw.) Clem. et Shear — loc. 1, 13699; loc. 2, 12768; loc. 4, 13142;

loc. 16, 12549.

This species is common in coastal areas in the North, whereas it is lacking

in the South.

Heterodermia albicans (Pers.) Swinsc. et Krog — loc. 10, 12329; loc. 14,

12652.

Only found in the central highlands, whereas it occurs preferably coastal in

other areas (E. Africa, С. & S. America).

H. comosa (Eschw.) Follm. et Redôn — loc. 12, 13551; loc. 14, 12647; loc.

16, 12540; Lac Mantasoa, Tixier s. n. (PC).

Not uncommon on branches in the central highlands.

Н. diademata (Tayl.) Awasthi — loc. 12, 13683; loc. 15, 12672, 12901.

Only found in the larger forest areas visited.

H. flabellata (Fée) Awasthi — loc. 15, 12669.

Found only once in one of the larger forest relicts.

H. galactopbylla (Tuck.) Trev. — loc. 8, 12503; loc. 12, 13290, 13293; loc.

15, 12908; Betsileo, Bois de Uranie, Catala s. n. (PC).

This species is not cited for East Africa by Swinscow & Krog (1976) and is

reported here for the first time from Africa. Part of the material cited above is

Somewhat atypical in having narrow corticated margins on the lower surface of

the lobes. It occurs in forest relicts, mainly on branches.

Source : MNHN, Paris

144 A. APTROOT

Н. hypoleuca (Ach.) Trev. — Type : Amer. bor., Muehlenberg 33-2 (H,

lectotype nov.) Dr. O. Vitikainen (in litt.) kindly drew my attention to the exis-

tence of a collection no. 20-2 from the same species and author, also preserved

in H. Therefore, a lectotype had to be chosen. loc. 12, 13282, 13289, 13681.

All material cited here has small spores without sporoblastidia. The species

was reported previously by Vainio (1898). The resembling species H. magellanica

(Zahlbr.) Swinsc. et Krog is reported for Madagascar by Kurokawa (1973 : 604)

by a dot on a map without any reference in the text.

H. isidiopbora (Vain.) Awasthi — loc. 9, 13004; loc. 12, 13287; loc. 15,

12878, 12910.

This species is found only in the larger forest relicts.

H. japonica (Sato) Swinsc. et Krog — loc. 2, Petit s. n. (PC); loc. 3, 13163;

loc. 5, 13040; loc. 8, 12503a; loc. 12, 13279, 13280, 13283, 13288, 13294,

13296, 13555, 13556, 13559, 13560; loc. 15, 12675, 12741b, 12909, 12538.

This is the most common species of the Pyxinaceae in Madagascar, occurring

on various substrates throughout the country.

H. leucomela (L.) Poelt subsp. boryi (Fée) Swinsc. et Krog — loc. 12, 13286,

13292, 13682; loc. 14, Humbert s. n. (PC); loc. 15, 12671, 12900, 12904,

12905; loc. 16, 12523, 12539; Betsileo, Bois de Uranie, Catala s. n. (PC).

This common and conspicuous lichen has been reported several times from

Madagascar, viz. by Crombie (1877), Mueller Arg. (1884), Stizenberger (1890)

and Vainio (1898).

H. lutescens (Kurok.) Follm. — loc. 11, 12416; loc. 15, 12670a.

À conspicuous but uncommon species from both rock and bark.

Н. obscurata (Nyl.) Trev. — loc. 4, 13146; loc. 11, 12417; loc. 12, 13285;

loc. 13, 12937; loc. 14, 12632; loc. 15, 12911, 12912.

This common species is previously reported for Madagascar by Mueller Arg.

(1884) as Physcia speciosa var. sorediifera.

H. podocarpa (Bél) Awasthi — loc. 15, 12673; Imerina, Andrangsloaka,

Hildebrandt 2169 (FH, M, not seen).

This species, described from the Malagasy region is reported from Madagascar

by Mueller Arg. (1884), Stizenberger (1890) and Vainio (1898).

Н. speciosa (Wulf.) Trev. — loc. 8, 12501; loc. 9, 13003; loc, 11, 12415; loc.

14, 12649; loc. 15, 12907.

This rather common species of inland habitats is previously reported for

Madagascar by Stizenberger (1890, var. dactyliza Nyl.) and Vainio (1898).

Н. verrucifera (Kurok.) W. A. Weber — loc. 9, 13013; loc. 12, 13558; loc. 15,

12670b.

This is the first report of this species from Africa of this tropical American

species. It has been frequently confused with H. leucomela.

Source : MNHN. Paris

PYXINACEAE OF MADAGASCAR 145

Н. vulgaris (Vain.) Follm. et Redon — loc. 12, 13284, 13291; loc. 15, 12906.

Unlike the related H. leucomela this species is found only in sheltered habi-

tats,

Hyperphyscia adglutinata (Floerke) Mayrh. et Poelt — loc. 8, 12436; loc. 10,

12305, 12336; loc. 14, 12656.

This species is found only inland in Madagascar, whereas it is commonly

found in coastal tropical areas. The morphology agrees well with the European

material.

Phaeophyscia bispidula (Ach.) Moberg — loc. 8, 12502; loc. 10, 12318.

The only representative of the genus Phaeophyscia in Madagascar. Stizen-

berger (1893) reported Physcia setosa Ach., which probably refers to the same

species.

Physcia atrostriata Moberg — loc. 2, 13759; loc. 5, 13053; loc. 9, 13012;

Mandraka, unknown collector (L ).

A northern and mainly coastal species of exposed habitats.

P. erumpens Moberg — loc. 10, 12310.

Only one record from this recently described species.

Р. fragilescens A. Zahlbr. — loc. 10, 12315.

This distinctive and much overlooked species is also only known from one

collection in Madagascar.

P. krogiae Moberg — loc. 8, 12466.

Also only one collection.

Р. stellaris (L.) Nyl. — loc. 10, 12328, 12367; Waterlot 43 р. р. (PC); loc. 14,

12650.

Remarkably the most common species of Physcia in Madagascar, previously

reported by Mueller Arg. (1894) and Stizenberger (1890), who also reported

P. integrata (Nyl.) from Madagascar.

Pyxine berteriana (Fée) Imshaug — loc. 10, Waterlot 43 p. p. (PC).

Probably extinct now. Visiting the same locality as Waterlot 1 was not able

to retrace this species.

P. cocoes (Sw.) Nyl. — loc. 1, Hildebrandt 2189 (PC); loc. 6, 13087a; loc. 7,

13096, 13097; loc. 10, 12310a, 12334.

This species is common in the Northern part of the country only. It is repor-

ted for Madagascar already by Stizenberger (1890).

Р. consocians Vain. — loc. 2, 13769.

Only found once on the North coast.

P. copelandii Vain. — loc. 1, 13698; loc. 5, 13056.

Also only on the North coast.

Source : MNHN, Paris

146 А. APTROOT

Pyxine glaucovirescens (Nyl.) Aptroot comb, nov.

Bas. : Physcia glaucovirescens Nyl., Synopsis Lich. Vol. 1, 1860 : 419. Туре:

Nouvelle Hollande (= Australia), Morton Bay, Verreaux 1846 (PC-lectotype).

Syn. : Р, endochrysina Nyl., Lich. Jap., 1890 : 34. Type : Japan, Hiroshima,

Almquist s. n. (H-NYL 31782).

loc.15,12688, 12903; Ankazobe, Bosser 12676 (TAN).

An uncommon inland species in Madagascar.

P. kibweziensis Swinsc. et Krog — loc. 8, 12504; loc. 11, 12406.

This species was only known from East Africa, It may belong to the African

mountain flora, like several Cladonia spp. (Ahti 1977). It occurs in Madagascar

on rock outcrops but also on acidic soil.

P. meissnerina Nyl. — loc. 1, 13695.

Only one record on the North coast.

P. petricola Nyl. in Cromb. — loc. 10, 12300, 12308, 12945; loc, 18, 12563.

А common species of Pyxine in Madagascar, with an inland distribution.

Rinodina dissa (Stirt.) Mayrh, — loc. 4, 13147; loc. 15, 12705.

The specimen from Ambalamanakana has very finely ornamented spores.

Since this is believed to be a valuable character in the Rinodina Callispora -

group, it makes the determination somewhat doubtful.

ACKNOWLEDGEMENTS. — I like to thank Dr. Н. J. M. Sipman for reading the manuscript

and the curators of H, L, PC and TAN for providing valuable material.

REFERENCES

AHTI T., 1977 — The Cladonia gorgonina group and C. gigantea in East Africa, Lichenolo-

gist 9:145.

APTROOT A., 1987 — Flora of the Guianas, The Pyxinaceae. Utrecht. 50 p.

AWASTHI D.D., 1975 — A Monograph of the Lichen Genus Dirinaria. Biblioth. Lichenol.

2:1-108.

CROMBIE J.M., 1877 - Lichens collected by W. Pools, Esq. in Madagascar. J. Linn. Soc.

Bot. 15 : 409-410.

CROSBY М.В. et al., 1983 — Katalog der Laubmoose von Madagascar und den umliegenden

Inseln. Willdenowia 13 : 187-255.

DES ABBAYES H., 1948 — Caractères et affinités de la flore des Cladonia (lichens) de la

région Malgache. Мет. Inst. Sci. Madagascar, Sér. В, Biol. Vég. 1 (2) :57-63.

DES ABBAYES H., 1956 — Lichens de la région Malgache I. Ibid. 7 :1-26.

Source : MNHN. Paris

PYXINACEAE OF MADAGASCAR. 147

DES ABBAYES Н., 1961 — Lichens récoltés à Madagascar et à la Réunion I et II. Ibid.

10 :81-121.

KUROKAWA S., 1962 — A Monograph of the Lichen Genus Anaptychia. Вей. Nova Hed-

wigia 6 : 1-115.

KUROKAWA S., 1973 — Supplementary notes on the genus Anaptychia. J. Hattori Bot.

Lab. 37 : 563-607.

MOBERG R., 1983 — The genus Phaeophyscia in East Africa. Nord. J. Bot. 3 : 509-516.

MOBERG R., 1986 — The genus Physcia in East Africa. Nord. J. Bot. 6 :843-864.

MUELLER ARGOVIENSIS J., 1884 — Lichenologische Beiträge XX. Flora 67 : 613-621.

ROGERS R.W., 1986 — The Genus Pyxine (Physciaceae, Lichenized Ascomycetes) in

Australia. Austral. J. Bot. 34 : 131-154.

STIZENBERGER E., 1890 — Lichenaea Africana. Ber. Thätigk. St. Gallisch. Naturwiss.

Ges. 1888/1889. St. Gallen. 120 p.

STIZENBERGER E., 1893 — Idem, Supplementa I. Ibid. 1891/1892. 40 p.

SWINSCOW T.D.V. & KROG H., 1975a — The genus Pyxine in East Africa. Norweg. J.

Bot. 22 :43-68.

SWINSCOW T.D.V. & KROG H., 1975b — Further observations on Pyxine in Africa.

Norweg. J. Bot. 22 : 125-128.

SWINSCOW T.D.V. & KROG H., 1976 — The genera Anaptychia and Heterodermia in

East Africa. Lichenologist 8 : 103-138.

SWINSCOW T.D.V. & KROG H., 1978 — The genus Dirinaria in East Africa. Norweg. J.

Bot. 25 : 157-168.

TIXIER P., 1985 — Contribution à la Connaissance des Cololejeuneoideae. Bryophyt.

Biblioth. 27. 439 р.

VAINIO E.A., 1898 — Lichenes quos in Madagascaria centrali Dr. C. Forsyth Major a 1896

collegit. Beiblatt zur Hedwigia 37 : 33-37.

Source : MNHN. Paris

Cryptogamie, Bryol, Lichénol, 1988 9 (2) : 149-154 149

CULTURE STUDIES ON RICCIA GANGETICA AHMAD

V - INFLUENCE OF MINERAL NUTRIENTS ON GROWTH

К.С. PATIDAR!, C.M. SOLANKI! and A. KAUL?

ABSTRACT — The effects of five mineral nutrients on vegetative growth of Riccia gangetica

Ahmad. was studied under controlled condition. Data are based on 60-day-old cultures.

When the concentration of all five nutrients were increased, there was no significant increase

in dry weight of thalli and results in abnormal growth. Growth was noted between 0.02% to

0.1% in Potassium nitrate, 0.02% to 1.0% in Potassium dihydrogen orthophosphate, 0.02%

to 0.5% in Calcium nitrate, 0.02% to 0.5% in Calcium chloride and 0.02% to 1.0% in Ma-

gnesium sulphate. The increase in nutrients concentration of 0.02% adversely influenced

growth, In the present experiments, 0.02% concentration was generally optimum to sustain

healthy growth as measured by revival capacity, number, average length and breath and

fresh and dry weight production.

INTRODUCTION

The mineralion concentration within the plants varies considerably, not so

much in relation to species but to environment, and to some extent to growth

form which is strongly correlated with environment. Different species associated

with particular habitat tend to have similar concentrations of specific mineral

elements, whereas the same species growing in a variety of sandy habitat tends

to have a range of mineral-ion concentrations (Smith 1978). The role of mineral

nutrition on the growth of bryophytes has been studied by Garjeanne (1932),

Mägdefrau (1933), Voth & Hamner (1940), Voth (1943), Fulford et al. (1947),

Machlis (1962), Hoffmann (1966), Sood (1974), Woodfin (1976), Shukla

(1977), Selkirk (1979), Patidar (1982). This communication deals with the

effect of five mineral nutrients on growth performance of Riccia gangetica

Ahmad, a common species of Ricciaceae (Patidar & Kaul 1982) at Pachmarhi

(22° 28° N and 78° 36° E). During the experiment, growth performance such

as revival capacity, length and breadth of thalli and fresh and dry weight pro-

duction of newly formed branches were noted.

1, Department of Botany, Gujarati Science College, Indore 452001, India.

2. Department of Botany, Madhav Science College, Ujjain 456010, India,

Source : MNHN, Paris

150 K.C. PATIDAR, C.M. SOLANKI and A. KAUL

MATERIALS AND METHODS

Thalli of Riccia gangetica Ahmad were collected from Pachmarhi (22°28 N

and 78° 26° E, 1067m) and brought to the laboratory in polythene bags. The

thalli were washed thoroughly to remove adhering soil particles. The experiment

was conducted on sand which was treated with acid and thoroughly washed

by water, then dried at 100°C for 24 hrs., transferred into sterilized petri dishes

and moistened with sterilized solutions of different concentrations of mineral

nutrients, as shown in tab. 1 through 5. Soil free thalli were cut transversely

into 4-5 mm long parts with a razor blade and grown at each concentration

of mineral nutrient. Potassium nitrate (KNO3), Potassium dihydrogen ortho-

phosphate (KHz PO), Calcium nitrate (Ca(NO3)2), Calcium chloride (CaCl2)

and Magnesium sulphate (MgSO4), were used as mineral nutrients. Three re-

plicates of five thalli were used for each experiment. The experiments were

conducted and discountinued for KNO3 from 11.8.80 to 9.10.80, for KH2PO4

from 22.8.80 to 22.10.80, for Ca(NO3)2 from 28.8.80 to 26.10.80, for CaCh

from 24.8.80 to 22.10.80, and for MgSO, from 20.8.80 to 18.10.80. The

control contained equal amount of water. Results in tab. 1 through tab. 5

are based on 6-day-old cultures. During the 60-day-growth period the cultures

were exposed to 12 hrs light (5000 Lux, maintained by incandescent and cool

white fluorescent light) and 12 hrs dark, to temperature of 22 + 5°C and humi-

dity of 80 to 85%. Dry weight production data were analysed by analysis of

variance, and by Duncan’s multiple Range Test (Duncan 1955).

OBSERVATION

Potassium nitrate. — The first sign of regeneration was seen about one month

after treatment.Growth occurred between 0.02% to 0.1% concentration (tab. 1).

The best growth performance on the basis of dry weight production of newly

formed branches, was at 0.02% concentration. No growth was noted above

0.1% because all thalli turned pale to dark black after 15 days.

Concentration Percentage Number of Average length Average breadth Fresh weight Dry weight "

revival of the newly of the newly оғ the newly ОҒ the newly о? the newly

ПИН ^ forme! formed branches formed branches formed branches formed branches

* branches (mm) (т) (na) (n

Control 50 4 3.0 2.00 9.25 2.152

0.02 80 4 4.6 2.80 15,39 3.51 b

0.05 100 5 4.0 1,90 10.6? La

ot 40 3 2.0 1.90 4.14 130€

5.5 ۰ - = x 5 E

Lo > В E à -

15 E = ж 5

2.0 - = B 3 :

“jeans within colums followed by sane letters are not significantly different at the 5° level as de-

termined by Duncan's Mult iole Rance Test

Tab. 1 — Effect of Potassium nitrate on vegetative growth of R. gangetica.

Source - MNHN, Paris

GROWTH OF RICCIA GANGETICA

151

Potassium dihydrogen orthophosphate. — Growth occurred between 0.02%

to 1.0% concentration (tab. 2). Above 1.0% concentration, the thalli we:

re tinged

with reddish-brown along the margins after 10 days. The dry weight production

was significantly higher at 0.02% that at any other concentration tested.

Concentration Percentage Number of Average length Average breadth Fresh weight Dry weight *

revival of the newly of the newly of the newly of the newly о? the newly

thalli formed formed branches formed branches formed branches formed branches

1 branches (т) (om) (ns) (т)

Control 3 9.5 30 20 9.25 2.15 a

0.02 80 В 4.6 23 16 2.18

0.05 80 5 22 12 5 1.69»

0.1 70 3 2.6 La ам LM b

0.5 so 4 12 1.1 4.20 1,09 с

0 E 2 10 14 0.84 алта

Means within columns followed by same letters are mot sionificantly different at the 5 level

termined by Duncan's Multiple Range Test

Tab. 2 — Effect of Potassium dihydrogen orthophosphate on vegetative growth

of R. gangetica

Concentration Percentage Number of Average length Average breadth Fresh weight Dry weight ^

revival of the newly of the newly of the newly of the newly of the newly

thalli formed formed branches formed branches formed branches formed branches

1 branches. (т) (т) (mg) (na)

Control % 5 3.0 2.0 9.25 2.15 а

o2 100 в 4.9 23 13.46 2.52 b

.05 70 6 3.2 1.7 1.15 3.30»

50 4 2.0 1.6 nu 2362

т 3 LB 1.0 5.36 1.91 €

Means within the colums followed by sane letters are not sionificantly different at the 5” level de

termined by Duncan's Multiple Pange Test.

Tab. 3 — Effect of Calcium nitrate on vegetative growth of R. gangetica

Calcium nitrate. — The first signs of regeneration were seen after two weeks.

Regeneration took place from the dorsal side and the cutted face of the

thallus.

Growth occurred between 0.02% to 0.5% concentration (tab, 3). In higher

concentrations the thalli were pale to brown and finally died after 7 days. The

best growth performance, on the basis of dry weight production, was noted at

0.02% concentration. At a higher concentration, dry weight production

sed significantly,

decrea-

Source : MNHN. Paris

152 K.C. PATIDAR, C.M. SOLANKI and A. KAUL

Calcium chloride. - The first sign of regeneration was seen after 15 days.

Regeneration took place from the cutted face of the thallus. Growth occurred

between 0.02% to 0.5% concentration (tab. 4). All parameters were highest

at 0.02% concentration.

Concentration Percentage Number of Average length Average breadth Fresh weight Dry weight "

revival of the newly of the newly of the newly ОҒ the newly О? the newly

thalli ^ formed formed branches formed branches formed branches formed branches

s branches (т) (т) (ne) (me)

% 5 3.0 2.0 9.25 таға

90 E 47 2 13.39 ns

20 6 4.2 11 зло 2.51 a

5 17; 1.3 4.61 1.83 €

4 14 1.0 2.2 өле

2.0 E 2 2. r E а

* Deans within the colums followed by sane letters are not significantly different at the 5 ۱ Y

determined by Duncan's Multiple Rane Test

Tab. 4 — Effect of Calcium chloride on vegetative growth of R. gangetica

Concentration Percentage Number of Average length Average breadth Fresh weight Dry weight *

revival of the newly of the newly of the newly of the newly ОҒ the newly

thalli formed formed branches formed branches formed branches formed branches

1 branches. (т) (тп) (ng) (та)

Control 90 5 3.0 2 10.28 2452

02 100 7 3.9 2: 15.36 3.78 b

05 90 5 4.3 24 12.40 злоһ

1 то 4 n9 11 2.69 220%

5 в 3 1.8 1.0 1.65 с

0 т 5 16 no ie

5 5 1 È Е =

o : 2 7

Means within the colums followed by sane letters are not significantly different at the 5 level as

determined by Duncan's Multiple Ranae Test.

Tab. 5 — Effect of Magnesium sulphate on vegetative growth of R. gangetica

Magnesium sulphate. — Regeneration was observed in about one week,

from both the ventral and dorsal surfaces of thallus. Growth was noted bet-

ween 0.02% to 1.0% concentration (tab. 5). The increase in concentration up

to 1.0% brought about a decrease in revival capacity, number, length, breadth

and fresh and dry weight production. Above 1% concentration, growth was

completely inhibited; only a few stunted regenerants developed, which turned

pale within a month and developed reddish-brown coloration on the ventral

surface.

Source : MNHN. Paris

GROWTH OF RICCIA GANGETICA 153

DISCUSSION

Most of the Marchantiales are ecologically specialized, not only in respect to

climate and microclimatic factors, but also to mineral nutrition. The experiment

reveals that the different nutrients at different concentrations affect the growth

performance of newly formed branches. Growth may be regarded as revival

capacity, number, average length and breadth, and fresh and dry weight produc-

tion of newly formed branches (tab. 1 to 5). Riccia gangetica survives from

0.02% to 0.1% of Potassium nitrate (tab. 1), from 0.02% to 1.0% of Potassium

dihydrogen orthophosphate (tab. 2), from 0.02% to 0.5% of Calcium nitrate

(tab. 3), from 0.02% to 0.5% of Calcium chloride (tab. 4) and from 0.02% to

1.0% of Magnesium sulphate (tab. 5). On the basis of statistical analysis of dry

weight production data, the growth performance was best at 0.02% concentra-

tion. The increase in concentration level brought about decrease of all parame-

ters.

The present experimental results suggest that effect of mineral nutrients is

particularly important for controlling growth іп R. gangetica, Sood (1974) sta-

ted that almost all nutrients are toxic when present at high concentration, and

are ineffective when present in extremely small amounts. Our findings on R.

gangetica are in agreement with those of Sood (1974) on Riccia crystallina.

Voth (1943) and Machlis (1962) reported that dry weight production decrea-

sed at higher concentration of minerals in Marchantia polymorpha and Sphaero-

carpos donnellii, respectively. Woodfin (1976) demonstrated that several species

of Riccia showed better vegetative growth when media contained less nitrogen.

Shukla (1977) found that fresh and dry weight production of Marchantia poly-

morpha and Plagiochasma appendiculatum were inhibited above 0.05% nutrients

concentration. In the present studies, R. gangetica showed good vegetative

growth at 0.02% concentration of Potassium dihydrogen orthophosphate (tab.

2), Calcium nitrate (tab. 3), Calcium chloride (tab. 4), Magnesium sulphate

(tab. 5) and Potassium nitrate (tab. 1). Selkirk (1979) found that growth rates

and number of branches of Riccia duplex and R. fluitans considerably increased

when treated with diluted nutrient solutions. Dacknowski (1907) reported

that in Marchantia transferred from field to water or weak nutrient solution

(0.1%), the formation of gemmae and sex organs was depressed. Patidar (1982)

found that Riccia discolor Lehm. et Lindenb. and R. fluitans L. showed dry

weight production and growth significantly higher at 0.02% nutrients concen-

tration,

The present results provide clear evidence that R. gangetica requires all essen-

tial elements for growth at reduced concentrations. Further investigations are

required to specify optimum concentration and range of tolerance between

zero to 0.02%,

ACKNOWLEDGEMENT. — Authors are grateful to the former principal Dr. H.N. Sharma,

Madhav Science College, Ujjain for providing facilities and encouragements.

Source : MNHN, Paris

154 К.С. PATIDAR, С.М. SOLANKI and А. KAUL

REFERENCES

DACKNOWSKI A., 1907 — Zur Kenntnis der Entwicklungsphysiologie von Marchantia

polymorpha L, Jahrb. Wiss. Bot. 44 :244-286.

DUNCAN D.B., 1955 — Multiple Range and Multiple F Test. Biometrics 11 : 1-42.

FULFORD M., CARROLL С. & COBBE T, 1947 — The response of Leucolejeunea clypea-

ta to variations in the nutrient solution. Bryologist 50 : 113-146.

GARJEANNE AJ.M., 1932 — Physiology. In : Е. VERDOORN, Manual of bryology. The

Hague. Pp. 207-232.

HOFFMAN G.R., 1966 - Observation on the mineral nutrition of Funaria hygrometrica.

Hedw. Bryologist 69 : 182-192.

MACHLIS L., 1962 — The effects of mineral salts, glucose and light on the growth of the

liverwort, Sphaerocarpos donnellii. Physiol. Pl. (Copenhagen) 15 : 354-362.

MAGDEFRAU K., 1933 — Versuche über den Einfluss der Ernährung auf den Thallus-

bau von Marchantia polymorpha. Ber. Deutsch. Bot, Ges, 51 : 106-111.

PATIDAR K.C., 1982 — Ecological studies on Ricciaceae. Ph. D. Thesis, Vikram University,

India.

PATIDAR К.С. & KAUL A., 1982 — Culture studies on Riccia gangetica Ahmad. I. Effect

of pH on growth. Cryptogamie, Bryol. Lichénol, 2 : 157-160.

SELKIRK P.M., 1979 — Effect of nutritional conditions on sexual reproduction in Riccia.

Bryologist 82 : 37-46.

SHUKLA R.M., 1977 — Ecological studies on two liverworts . Ph. D. Thesis, Vikram Uni-

versity, India.

SMITH R.LL., 1978 — Summer and Winter concentration of Sodium, Potassium and Cal-

cium in some Maritime Antarctic cryptogams. J. Ecol. 66 : 891-909.

SOOD S., 1974 — In vitro studies in Marchantiales - II. Effect of mineral nutrients, chelates

and organic nitrogenous sources on the growth and sexuality in Riccia crystallina. Phyto-

morphology 24 : 186-197.

VOTH P.D, & HAMNER K.C., 1940 — Responses of Marchantia polymorpha to nutrient

supply and photoperiod. Bot. Gaz. (Crawfordsville) 102 : 169-204.

VOTH P.D., 1943 — Effect of nutrient solution concentration on the growth of Marchantia

polymorpha, Bot. Gaz. (Crawfordsville) 104 : 591-601.

WOODFIN C.M., 1976 — Physiological studies on selected species of the liverwort family

Ricciaceae, J. Hattori Bot. Lab, 41 : 179-183.

Source : MNHN. Paris

Cryptogamie, Bryol. Lichénol. 1988 9 (2) : 155-172 155

ULTRACYTOCHEMICAL LOCALIZATION

OF ACID AND ALKALINE PHOSPHATASES

IN THE PLACENTAL REGION

OF THE MOSS PHYSCOMITRIUM CYATHICARPUM

Е. CHAUHAN

SUMMARY. — The distribution of acid (Bglycerophosphatase) and alkaline phosphatase

activity has been determined at the EM level in the transfer cells comprising the placental

region of the moss Physcomitrium cyathicarpum Mitt. Reaction products of these phospho-

hydrolases have been found to be associated with the amplified plasma membranes of the

foot and the vaginula cells at the sporophyte-gametophyte junction, Activity of Bglycero-

phosphatase is also seen in golgi, vesicles and some microbodies. In addition, the wall-mem-

brane apparatus of the vaginula cells shows higher alkaline phosphatase activity than in cells

of the haustorial foot. These findings have been presented in relation to the role of transfer

cells in active transport. Also discussed are some of the important problems in the cyto-

chemistry of phosphatases.

INTRODUCTION

The placental zone in bryophytes constitutes an ideal site for studying the

short distance transport of materials from cells of one generation to the other.

The sporophyte-gametophyte junction is characterized by distinct wall in-

growths either only in cells of the foot, or exclusively in the vaginula cells (as

in hornworts Phaeoceros (Gambardella et al. 1981) and Anthoceros (Chauhan

& Schraudolf 1986), or in both. Several ultrastructural and physiological studies

have been made to elucidate the structural and functional specialization of

transfer cells of the foot-vaginula complex (for a review of literature see Hébant

1977, Chauhan & Lal 1987b). A survey of literature reveals that very few

histoenzymological studies have been made so far in bryophytes in general, and

on the placental zone in particular. At the light microscopy level, there are just

three reports on the localization of phosphatases and respiratory enzymes in

the haustorial foot and its surrounding gametophytic tissue in mosses (Hébant

& Suire 1974, Chauhan & Lal 1980, 1982). Just one such report exists on a

hornwort Phaeoceros laevis (Thomas et al. 1978). At the EM level only two

* Plant Cytochemistry and Ultrastructure Laboratory, Department of Botany, University

of Delhi, Delhi- 110007, India.

Source : MNHN. Paris

156 Е. CHAUHAN

studies have been made, viz., localization of B-glycerophosphatase and Mg” -

activated adenosine tri-phosphatase in Polytrichum piliferum (Maier & Maier

1972) and adenylate cyclase in Physcomitrium cyathicarpum (Chauhan 1987).

In some of the earlier papers of the ultrastructure of the sporophyte-gameto-

phyte junction in Physcomitrium cyathicarpum (Chauhan & Lal 1980, 1984,

1987b, Lal & Chauhan 1981), it has been shown that extensive cell wall invagi-

nations exist in the peripheral cells of the haustorial foot in the abutting cells of

the vaginula. The present investigations, involving ultratsructural enzyme cyto-

chemistry, reveal high activities of acid (B-glycerophosphatase) and alkaline

phosphatases associated with the amplified plasma membrane of the transfer

cells at the foot vaginula interface. In addition, the reaction products of these

enzymes were found to occur in some cell organelles and the cytoplasmic islands

trapped in the cell wall labyrinth. Also discussed are the problems encountered

in the cytochemistry of phosphatases.

MATERIALS AND METHODS

Fresh plants of Physcomitrium cyathicarpum Mitt. (Funariaceae) were collec-

ted from the Delhi University Botanical Gardens. The haustorial foot, along

with the surrounding gametophytic tissue was excised and sliced with the help

of a sharp razor blade. These slices were subjected to the following two pro-

cedures :

a. The material was immediately fixed in 1.5% glutaraldehyde in 0.1 M so-

dium cacodylate buffer (pH 7.2), and kept in crushed ice (Roland 1978).

After 2 hour fixation, the samples were washed in 0.1 M sodium cacodylate

buffer (pH 7.2) at 4°C, rinsed briefly in 0.1 M tris-maleate buffer at pH 6.0 and

subjected to incubation for acid and alkaline phosphatase localization (Barka &

Anderson 1963).

b. Alternatively, the material was first directly incubated to localize acid

phosphatases (EC 3.1 3.2) by the modified lead phosphate method (Shaykh &

Roberts 1974) using sodium f-glycerophosphate as substrate, and alkaline phos-

phatases (EC 3.1 3.1) were localized by the procedure of Lojda et al. (1979)

using naphthol AS-MS phosphate as substrate. Incubations in the complete

reaction media in both cases were carried out at 35-37°C.

The enzyme reaction was tested light microscopically and incubation was

found to be adequate after 60 min and 45 min for acid and alkaline phospha-

tases respectively (see Chauhan & Lal 1982), Controls included : (i) samples

incubated in the reaction medium minus substrate, (ii) material pretreated with

0.01 M sodium fluoride for 5 min followed by incubation in complete reaction

medium plus NaF (Bitensky 1963), and (iii) without the incubation step. After

incubation, the material was washed with 0.1 M sodium cacodylate buffer

(pH 7.2) and fixed according to the method described in a. at room temperature.

Post-fixation, dehydration, embedding and thin sectioning was done accor-

ding to the procedure already outlined (Chauhan 1987). The sections were

Source : ММНМ, Paris

PHOSPHATASES IN THE PLACENTAL REGION OF PHYSCOMITRIUM 157

post stained with only aqueous uranyl acetate or with uranyl acetate followed

by alkaline lead citrate (Reynolds 1963). Sections were observed and photo-

graphed in Carl Zeiss EM 109 or Philips EM 300 transmission electron micro-

scopes at 80 kV.

Understandibly so, incubation in the reaction medium and short fixation

resulted in some sacrifice in the quality of fine structure of the tissue,

OBSERVATIONS

As reported earlier, the peripheral cells of the foot and those of the vaginula

abutting the foot in Physcomitrium cyathicarpum possess wall labyrinth (Chau-

han 1981, Chauhan & Lal 1980, Lal & Chauhan 1981). Also, the haustorial tip,

which is represented by a quadrate of cells, shows characteristic wall invagina-

tions (Chauhan & Lal 1984). Present ultracytochemical studies show that GPase

(GPase: Bglycerophosphatase) activity is associated with the cell walls and

invaginations of the peripheral foot cells and those of the abutting vaginula

(Figs. 1, 2). Lead precipitate is localized in the plasmalemma following the

contours of the labyrinth in cells of both the generations at the sporophyte

gametophyte junction. However, with the NaF control, almost all activity is

eliminated from these sites (Fig. 8). GPase activity is also associated with the

golgi (Figs. 1, 2, 3, 4), vesicles (Fig. 4) and some microbodies (Figs. 3, 4) in

transfer cells of the foot. However, no lead phosphate deposits are detected in

the ER cisternae. Some amount of lead precipitate is also visualized in the

multivesicular bodies of the vaginula cell (Fig. 2).

Deposits of lead precipitate are scattered throughout the nucleus of the peri-

pheral foot cells (Fig. 5). Fine particles of lead are, however, concentrated in the

nucleolus. Surprisingly, the nuclear membrane in foot as well as the vaginula

transfer cells along the entire length of the haustorium shows no enzyme activity

(Figs. 5, 10). A similar staining pattern for GPase activity is also observed in the

nucleus of the axial cell (Fig. 6). No such lead deposits are seen in the nucleus

after use of NaF in controls (Fig. 9) indicating non-specificity of deposits in the

earlier photographs. Golgi and vesicles in the abutting vaginula cells also show

indications of GPase activity (Fig. 7). Enzyme activity is also associated with

walls of the axial cells (Fig. 6).

Relatively more AlPase (AlPase : alkaline phosphatase) activity is seen at the

wall-membrane apparatus of the vaginula cells (Fig. 11) as compared to that in

the peripheral foot cells, There are distinct electron-dense deposits of the reac-

tion product on membranes of vacuoles and those following the contours of the

wall invaginations (Fig. 11, 12). The substrate- free control, however, eliminated

all deposits from the vacuolar membranes (Fig. 14). In addition, the outer face

of the cell wall of the haustorial tip shows heavy deposits of reaction product for

AlPase activity (Fig. 13).

Source - MNHN. Paris

158 E. CHAUHAN

DISCUSSION

The present tltracytochemical studies in Physcomitrium cyathicarpum ex-

hibit strong GPase and AlPase activities associated with the wall invaginations of

the transfer cells at the foot-vaginula interface. These phosphohydrolases were

earlier localized at the light microscopy level in the peripheral foot cells and the

abutting vaginula cells in the same species (Chauhan & Lal 1982). Several studies

have shown phosphatase activities associated with the cell walls (De Jong et al.

1967, Hall & Butt 1968, Hall 1969, Tripodi 1970, Hall & Davie 1971, Sexton

et al. 1971, Matile 1975, Joshi et al. 1985). Among bryophytes, similar pattern

of acid phosphatase localization has been reported in the placental region of

Polytrichum piliferum (Maier & Maier 1972), and in Dawsonia papuana, Dicra-

num scoparium, Polytrichum commune and Scapania undulata (Hébant & Suire

1974). The role of these enzymes in the cell walls has been speculated to involve

intercellular transport mechanism, phosphate transfer reactions and cell wall

metabolism (Halperin 1969, Sexton et al. 1971, Onofeghara 1972). This view

is also supported by histoenzymological studies on the conducting tissues of

some bryophytes (Hébant 1973, 1977), In other situations, acid phosphatases

may also be involved in the dephosphorylation processes associated with the

degradation of starch, and thus having a possible role during the stomatal ope-

ning, as suggested by Couot-Gastelier & Louguet (1985) in Tradescantia virgi-

niana.

In P. cyathicarpum, the reaction product for GPase is also observed in the

golgi cisternae, golgi vesicles, microbodies and some multivesicular bodies. The

small cytoplasmic vesicles showing GPase activity may have a lysosomal function

in these cells (see Matile 1975). Activity of this enzyme in components of the

endomembrane system points towards a possible secretory activity. In this con-

text, it is pertinent to note that a similar pattern of distribution of acid phospha-

tase activity has been reported in the digestive glands of some insectivorous

plants like Dionaea muscipula (see Henry & Steer 1985, and references cited

therein).

Acid phosphatases in plant cells can be demonstrated by the lead salt method

(Gahan 1965, Gahan et al. 1978), the azo dye method and the bromoindoxyl

method (see Gahan 1984). The commonly used substrates are B-glycerophos-

phate, p-nitrophenyl phosphate and naphthol AS-BI phosphate. Each method

demonstrates a multiplicity of acid phosphatases and, in addition, does not

necessarily demonstrate the same enzymes (Gahan et al. 1978). Therefore, large

variations exist in the methodology used to localize acid phosphatases and right-

ly so, since their localization at the EM level involves significant limitations

and variations in the visualization of the final precipitate, viz., effect of fixatives,

simultaneous preservation of both morphology and enzyme activity, and the

nature of the electron-dense reaction product (for a discussion on the problems

of acid phosphatase cytochemistry see Pearse 1968, Sexton & Hall 1978, Gahan

1984 and Borgers & Verheyen 1985). According to Gahan (1984) there is no

totally reliable method for the localization of acid phosphatases that has been

Source - MNHN. Paris

PHOSPHATASES IN THE PLACENTAL REGION OF PHYSCOMITRIUM 159

shown to be suitable for a broad range of tissues.

Interesting in this context are some of the recent innovations made for im-

proved localization of intracellular sites of phosphatases. Over the years, the pro-

cedures for the demonstration of acid phosphatases (first developed in 1952 for

light microscopy by Gomori) have undergone several modifications for electron

microscopy (see Borgers & Verheyen 1985) involving a variety of newer sub-

strates, buffers and also the trapping agents. For example, Veenhuis et al. (1980

— cited in Borgers & Verheyen 1985) used cerium chloride to produce a final

precipitation of cerium phosphate, which can be visualized as very fine electron-

dense deposits and allows a more accurate localization of the sites of enzyme

activity than the conventional lead-based procedures. Aluminium nitrate, when

used as a trapping agent, also shows such fine deposits (Berry et al. 1982).

Taking into account the advantages of cerium as a capture metal over lead,

Robinson (1985) has shown that permeation with low levels of detergents in

conjunction with cerium cytochemistry of acid and alkaline phosphatase gave

more consistent intracellular localization and a better preservation of ultrastruc-

tural morphology. Also permeabilization and certain techniques together re-

vealed intracellular sites of enzyme activity not otherwise detectable.

Likewise, although the metal salt technique for the localization for alkaline

phosphatases was reported as early as in 1939, independently by Gomori and

Takamatsu (1939), the problems in the demonstration of this enzyme in plant

tissues are still far from resolved. Even in animal tissues, it has not been possible

to describe precisely a function for this enzyme because it is found in such

diverse cell types and because it consists of several isoenzymes. Details of metho-

dology and interpretation of alkaline phosphatase localization at the EM level

in animal tissues has been reviewed by Essner (1973) and Borgers & Verheyen

(1985). Like in acid phosphatases, here too, the phosphate released combines

with the metal ions like lead or calcium present in the incubation medium.

One of the crucial factors here is pH since at pH above 8.0, lead phosphate is

increasingly mobile in an aqueous medium. It was, therefore, recommended

that calcium should be used at high pH values which is to be subsequently re-

placed by cobalt prior to the formation of metal sulfide (Gomori 1952). Earlier,

В- naphthyl phosphate was used as a substrate (Menten et al. 1944) and subse-

quent studies revealed that the naphthyl compounds were more satisfactory

since the precipitates formed were much more insoluble than lead or calcium

phosphates. Various diazo compounds have also been studied in relation to alka-

line phosphatase localization by Pearse (1968) with regard to their colour,

decomposition, coupling speed and enzyme inhibition. Recently, Doty (1980)

has discussed the problems encountered in the cytochemistry of alkaline phos-

phatases. He has also stressed the need to find out specific inhibitors in order to

differentiate these enzyme activities at lower pH. Literature on localization of

this enzyme in plants is discussed in the later paragraphs.

In the present investigation, a nuclear reaction for GPase is also recorded.

The reaction product is seen in both the conditions, viz., slices of the haustorial

foot with prefixation, and material incubated and subsequently fixed. Similar

Source : MNHN, Paris

160 Е. CHAUHAN

nuclear staining has been reported in a wide variety of plants (see Sexton & Hall

1978, Gahan 1984). The presence of lead precipitates in the nucleus can be

attributed to a random, non-enzymatic deposition (Gomori 1952) or to diffu-

sion of either the enzyme (Bitensky 1962), or the reaction product itself (Bi-

tensky 1963) to the nucleus with prolonged incubation periods.

In the case of plant tissues, there is the distinct possibility of the presence of

high concentrations of endogenous phosphate that can also lead to the forma-

tion of non-enzymic lead phosphate (Gahan 1984). The nuclear reaction is how-

ever, very inconsistent and depends on several experimental variables (Barka &

Anderson 1962) and may pose some additional problems in phosphatase locali-

zation at the electron microscopy level. Some of such artefacts are easily identi-

fiable. Coupled with adequate controls (like the ones employed in the present

study), these methods have been explained in order to recommend the metal

salt techniques with confidence (see Ericsson & Trump 1964, Cornelisse & van

Duijn 1973a, b, Essner 1973, Sexton & Hall 1978).

In a recent report, GPase activity localized in the nucleolus of the developing

fibre initial of cotton has been thought to serve a developmental function, and

its transitory occurrence in the nucleus is suggestive of dephosphorylation of

the nuclear protein (Joshi et al. 1985). According to these workers, GPase may

function as a phosphoprotein phosphatase which activates the enzymes involved

in synthetic pathways by its phosphorolytic activity. Joshi et al. (1985) have

made this speculation in the light of the growing concept that the phosphory-

lation-dephosphorylation of critical enzymes is one of the controlling mecha-

nisms in biological systems (see Krebs & Beavo 1979, Ingebritsen & Cohen

1983).

Present studies reveal the reaction product of GPase below the walls of the

axial cells (Fig. 6), whereas the hadrom of the foot in Polytrichum piliferum

did not show any lead deposits for beta-glycero-phosphatase (Maier & Maier

1972), though there was an occasional unspecific accumulation of the reaction

product in the walls of the dead hydroids. This can be explained on account

of the fact that the haustorium in P. cyathicarpum reveals a simplified histolo-

gical pattern characteristic of the highly reduced forms and shows no demar-

cation of the axial tissues into ground cortex, leptoids and/or hydroids, as

described in polytrichaceous mosses. It is therefore, suggestive that the entire

haustorium in this species represents a single, composite functional unit (for a

discussion on axial cells see Chauhan & Lal 1986).

With reference to alkaline phosphatase, distinct electron-dense deposits of

AlPase have been observed оп the membranes of the wall invaginations in P.

cyathicarpum. It is seen that the reaction for AlPase is stronger in the vaginula

transfer cells as compared to those of the foot. These studies regarding the

presence of AlPase confirm the earlier investigations in the same species made

at the light microscopy level (Chauhan & Lal 1982). So far, there are no other

reports of this enzyme in bryophytes, and the literature on alkaline phosphatases

in plants is, as such, scanty. AlPase activity has been reported in the rhizoids

of the young gametophytes of Polypodium vulgare (Smith 1972a, b), stem

Source : MNHN, Paris

PHOSPHATASES IN THE PLACENTAL REGION OF PHYSCOMITRIUM 161

tissues of cucurbits (Onofeghara & Koroma 1974), onion roots and the myc

rhiza of Glomus mosseae (Gianinazzi-Pearson & Gianinazzi 1976, 1977, Gia

nazzi et al. 1979). On the other hand, some negative reports also exist (see

Chauhan 1984).

The presence of alkaline phosphatase in the sporophyte-gametophyte junc-

tion points towards its possible role in intercellular and apoplastic transport.

In addition, accumulation of the reaction product on the outer face of the wall

of the haustorial tip strengthens the possibility that some hydrolytic agents

secreted by the tip are involved in the process of dissolution of the surrounding

tissue in order to obtain early nutritional requirements for the young embryo

(see also Browning & Gunning 1979a, Chauhan & Lal 1982). Since this enzyme

is found primarily in cell membranes where active transport processes take

place, the present studies on the foot-vaginula interface in P. cyathicarpum

provide further evidence to the notion that the transfer of materials at this zone

are mediated by active, apoplastic transport mechanisms.

In some mosses like Funaria hygrometrica and Polytrichum formosum, the

conduction of organic compounds has been shown to be an active, energy

dependent process (Browning & Gunning 1979b, c, Caussin et al., 1979, 1983).

This is in contrast to the water uptake by the haustorial foot (Bopp & Weniger

1971), The present ultracytochemical studies involving the activities of phospho-

hydrolases, therefore, are corroborative to our earlier investigations on Physco-

mitrium wherein activities of Са?" — and Mg?* — activated ATPases (Chauhan &

Lal 1980), succinate dehydrogenase (Chauhan & Lal 1982) and adenylate

cyclase (Chauhan 1987) have been localized in the transfer cells making the foot-

vaginula interface a seat of active transport between the apoplast and symplast.

In addition, the transfer cells of the foot show cytochemical attributes which are

commensurate with the nutritional needs of the growing capsule (Chauhan &

Lal 1987a).

ACKNOWLEDGEMENTS. — The author thanks Professor M. Lal (University of Delhi)

and Professor H. Schraudolf (Universitat ULM, Federal Republic of Germany) for providing

laboratory facilities. Award of a DAAD Fellowship (1984-85) is also gratefully acknow-

ledged.

REFERENCES

BARKA Т. & ANDERSON P.J., 1962 — Histochemical methods for acid phosphatase using

hexazonium pararosanalin as coupler. J. Histochem. Cytochem. 10 :741-753.

BARKA Т. & ANDERSON Р.)., 1963 — Histochemistry : Theory, Practice and Bibliogra-

phy. New York : Harper & Row. 660 p.

BERRY J.P., HOURDRY J.. STENBERG М. & GALLE P., 1982 — Aluminium phosphatase

visualization of acid phosphatase activity. J. Histochem. Cytochem. 30 :86-90,

BITENSKY L., 1962 — The reversible activation of lysosomes in normal cells and the effects

Source : MNHN, Paris

162 E. CHAUHAN

of pathological conditions. In : DE REUCK A.V.S. & CAMERON М.Р. (eds.), Ciba

Found, Symp. Lysosomes. Boston : Little Brown, Pp. 362-375.

BITENSKY L., 1963 — Modifications to the Gomori acid phosphatase technique for con-

trolled-temperature frozen sections. Quart. J. Microsc, Sci, 104 ; 193-196.

BOPP M. & WENIGER H.P., 1971 — Wassertransport vom Gametophyten zum Sporophyten

bei Laubmoosen. 2. Pflanzenphysiol. 64 : 190-198.

BORGERS M. & VERHEYEN A., 1985 — Enzyme cytochemistry. Int. Rev. Cytol. 95 :

163-227.

BROWNING АЈ. & GUNNING B.E.S., 1979a — Structure and function of transfer cells in

the sporophyte haustorium of Funaria hygrometrica Hedw. 1. The development and

ultrastructure of haustorium. J. Exp. Bot. 30 : 1233-1246.

BROWNING A.J. & GUNNING B.E.S., 1979b — Structure and function of transfer cells in

the sporophyte haustorium of Funaria hygrometrica Hedw. И. Kinetics of uptake of

labelled sugars and localization of absorbed products by freeze-substitution and auto-

radiography. J. Exp. Bot. 30 : 1247-1264.

BROWNING AJ. & GUNNING B:ES., 1979c — Structure and function of transfer cells in

the sporophyte haustorium of Funaria hygrometrica Hedw. Ш, Translocation of assimi-

late into the attached sporophyte and along the seta of attached and excised sporo-

phytes. J. Exp. Bot. 30 : 1265-1273.

CAUSSIN C., DESPEGHEL J.P., FANCHER M., КЕСЕК A. & BONNEMAIN J.L., 1979 —

Étude du mécanisme des échanges entre le gamétophyte et le sporophyte chez les Bryo-

phytes. Compt. Rend. Hebd. Séanc. Acad. Sci., Série D 289 : 1329-1334.

CAUSSIN C., FLEURAT-LESSARD P., BONNEMAIN J.L., 1983 — Absorption of some

amino acids by sporophytes isolated from Polytrichum formosum and ultrastructural

characteristics of the haustorium transfer cells, Ann. Bot. (London) 51 : 167-173.

CHAUHAN E. & LAL M., 1980 — Histochemical and ultrastructural investigations on the

foot of the moss Physcomitrium cyathicarpum Mitt. Evidence for the presence of a

Ca?* — activated ATPase. J. Cytol. Genet. 15 :190-194.

CHAUHAN E., 1981 — Ultrastructural, Cytochemical and Histochemical Studies on Some

‘Aspects of Sporophyte Development in the Moss Physcomitrium cyathicarpum Mitt.

Ph. D. Thesis, Univ. of Delhi, India. 183 p.

CHAUHAN Е. & LAL M., 1982 — Localization of some hydrolases and succinate dehydro-

genase in the sporophyte-gametophyte junction in Physcomitrium cyathicarpum Mitt.

Ann, Bot. (London) 50 : 763-769.

CHAUHAN Е., 1984 — Ultrastructural investigations on the haustorial foot in the moss

Physcomitrium cyathicarpum. Cytobios 41 : 85-93.

CHAUHAN E. & LAL M., 1986 — Ultrastructure of the haustorial axis in the moss Physco-

mitrium cyathicarpum Mitt. In : KUMAR 5.5. (ed.), Proceedings of the All India Confe-

rence on Bryology (Feb. 25-27, 1986), Chandigarh, India. Pp. 13-14 (abstr.).

CHAUHAN E. & SCHRAUDOLF H., 1986 — Ultrastructural studies on the placental

region in Anthoceros punctatus L. Beitr. Biol. РЯ. 61 : 357-372.

CHAUHAN E., 1987 — Ultracytochemical localization of adenylate cyclase in the placental

region of Physcomitrium cyathicarpum. Cryptogamie, Bryol. Lichénol. 8 : 189-198.

CHAUHAN E. & LAL M., 1987a — Cytochemical studies on the developing sporophyte in

the moss Physcomitrium cyathicarpum. Phytomorphology 37 :17-27.

CHAUHAN E. & LAL M., 1987b — Development of transfer cells in the haustorium-vagin-

nula complex of Physcomitrium cyathicarpum Mitt. — An ultrastructural study. J.

Hattori Bot. Lab. 63 : 373-394.

Source : MNHN, Paris

PHOSPHATASES IN THE PLACENTAL REGION OF PHYSCOMITRIUM 163

CORNELISSE C.J. & VAN DUIJN P., 1973a — A new method for the investigation of the

Kinetics of the capture reaction in phosphatase cytochemistry. I. Theoretical aspects

of the local formation of crystalline precipitates. J. Histochem. Cytochem. 21 : 607-613.

CORNELISSE C.J. & VAN DUIJN P., 1973b — A new method for the investigation of the

kinetics of the capture reaction in phosphatase cytochemistry. II. Theoretical and ex-

perimental study of phosphate diffusion from thin polyacrylamide films. J. Histochem.

Cytochem. 21 1614-622.

COUOT-GASTELIER J. & LOUGUET P., 1985 — Localisation infrastructurale de la phos-

phatase acide dans les cellules stomatiques de Tradescantia virginiana L. Cytobios 43 :

8795.

DE JONG D.W., JANSEN Е.Ғ. & OLSON A.C., 1967 — Oxidoreductive and hydrolytic

enzyme patterns in plant suspension culture cells. Exp. Cell Res, 47 : 139-156,

DOTY S.B., 1980 — Problems inherent in obtaining the alkaline phosphatase reaction. J,

Histochem. Cytochem. 28 : 66-68.

ERICSSON J.L.E. & TRUMP B.F., 1964 — Observations on the application of electron

microscopy of the lead phosphate technique for the demonstration of acid phosphatase.

Histochemie 4 : 470-487.

ESSNER E., 1973 - Phosphatases. In : HAYAT М.А. (ed.), Electron Microscopy of Enzy-

mes, Vol. 1. New York : Van Nostrand Reinhold Company. Pp. 44-76.

GAHAN P.B., 1965 — Histochemical evidence for the presence of lysosome-like particles

in root meristem cells of Vicia faba. J. Exp. Bot. 16 : 350-355.

GAHAN P.B., DAWSON A.L. & FIELDING J., 1978 — Paranitrophenyl phosphate as a

substrate for some acid phosphatases in roots of Vicia faba. Ann, Bot. (London) 42 :

1413-1420.

GAHAN P.B., 1984 — Plant Histochemistry and Cytochemistry, London : Academic Press,

301 p. (Exp. Bot. Monographs, Series 18).

GAMBARDELLA R., LIGRONE R. & CASTALDO R., 1981 — Ultrastructure of the sporo-

phyte foot in Phaeoceros. Cryptogamie, Bryol. Lichénol, 2 :23-45.

GIANINAZZI-PEARSON V. & GIANINAZZI S., 1976 — Enzymatic studies on the meta-

bolism of vesicular-arbuscular mycorrhiza, I. Effect of mycorrhiza formation and phos-

phorus nutrition on soluble phosphatase activities in onion roots. Physiol. Vég. 14 :

833-841.

GIANINAZZI-PEARSON V. & GIANINAZZI S., 1977 — Enzymatic studies on the meta-

bolism of vesicular-arbuscular mycorrhiza. II. Soluble alkaline phosphatase specific to

mycorrhizal infection in onion roots. Physiol. Pl. Pathol. 12 : 45-53.

GIANINAZZI S$., GIANINAZZI-PEARSON V. & DEXHEIMER J., 1979 — Enzymatic

studies on the metabolism of vesicular-arbuscular mycorrhiza. Ш. Ultrastructural loca-

lization of acid and alkaline phosphatase in onion roots infected by Glomus mosseae

(Nicol et Gerd.). New Phytol. 82 :127-132.

GOMORI G., 1939 — Microtechnical demonstration of phosphatase in tissue section. Proc,

Soc. Exp. Biol. Med. 42 : 23-26.

GOMORI G., 1952 — Microscopic Histochemistry : Principles and Practice. Chicago : Univ.

of Chicago Press. 273 p.

HALL J.L. & BUTT V.S., 1968 — Localization and kinetic properties of B-glycerophospha-

tase in barley roots. J. Exp. Bot. 19 :276-287.

HALL J.L., 1969 — Histochemical localization of B-glycerophosphatase activity in young

root tips. Ann. Bot. (London) 33 : 399-406.

HALL J.L. & DAVIE С.А.М., 1971 — Localization of acid hydrolase activity in Zea mays

Source : ММНМ. Paris

164 E. CHAUHAN

root tips. Ann. Bot. (London) 35 : 849-855.

HALPERIN W., 1969 —Ultrastructural localization of acid phosphatase in cultured cells of

Daucus carota. Planta 88 : 91-102.

HEBANT C., 1973 — Acid phosphomonoesterase activities (B-glycero phosphatase and

naphthol AS-MX phosphatase) in conducting tissues of bryophytes. Protoplasma 77 :

231-241.

HEBANT С. & SUIRE C., 1974 — Mise en évidence d'activités enzymatiques au niveau de

la zone de transfert gamétophyte-sporophyte chez quelques Bryophytes. Rev. Bryol.

Lichénol. 40 : 171-181.

HÉBANT C., 1977 — The conducting Tissues of Bryophytes. Bryophyt. Biblioth. 10 : I-

ХТ, 1-157.

HENRY Y. & STEER M.W., 1985 — Acid phosphatase localization in the digestive glands

of Dionaea muscipula Ellis flytraps. J. Histochem. Cytochem. 33 : 339-344.

INGEBRITSEN T.S. & COHEN Ph., 1983 — Protein phosphatases : properties and role in

cellular regulation. Science 221 : 331-338.

JOSHI P.A., STEWART J. Мер. & GRAHAM E.T., 1985 — Localization of B-glycerophos-

phatase activity in cotton fiber during differentiation. Protoplasma 125 :75-85.

KREBS E.G. & BEAVO J.A., 1979 — Phosphorylation-dephosphorylation of enzymes.

Annual Rev. Biochem, 48 :923959.

LAL M. & CHAUHAN E., 1981 — Transfer cells in the sporophyte-gametophyte junction

of Physcomitrium cyathicarpum Mitt. Protoplasma 107 : 79-83.

LOJDA Z., GOSSRAU R. & SCHIEBLER T.H., 1979 — Enzyme Histochemistry — A Labo-

ratory Manual. Berlin, Heidelberg, New York : Springer-Verlag. 339 p.

MAIER К. & MAIER U., 1972 — Localization of beta-glycerophosphatase and Mg" — acti-

vated adenosine triphosphatase in a moss haustorium, and the relation of these enzymes

to the cell wall labyrinth. Protoplasma 75 :91-112.

MATILE P., 1975 — The Lytic Compartment of Plant Cells. Vienna, New York : Springer-

Verlag. 183 p.

MENTEN M.L., JUNGE J. & GREEN M.H., 1944 — A coupling histochemical azo dye test

for alkaline phosphatase in kidney. J. Biol. Chem. 153 : 471477.

ONOFEGHARA F.A., 1972 — Histochemical localization of enzymes in Tapinanthus beng-

wensis : Acid phosphatase. Amer. J. Bot. 59 : 549-556.

ONOFEGHARA F.A. & KOROMA S.A., 1974 — Histochemical localization of alkaline

phosphatases and f-glucosidases in Cucurbitaceae. Ann. Вог. (London) 38 : 705-710.

PEARSE A.G.E., 1968 — Histochemistry : Theoretical and Applied. Ed. 3. London : J. & A.

Churchill. Vol. I. 759 p.

REYNOLDS E.S., 1963 — The use of lead citrate at high pH as an electron-opaque stain

in electron microscopy: J. Cell Biol. 17 : 208-212.

ROBINSON J.M., 1985 — Improved localization of intracellular sites of phosphatases using

cerium and cell permeabilization. J. Histochem. Cytochem. 33 :749-754.

ROLAND J.C., 1978 — General preparation and staining of thin sections. In : HALL J-L.

(ed.), Electron Microscopy and Cytochemistry of Plant Cells. Amsterdam, Oxford, New

York : Elsevier/North Holland Biomedical Press. Pp. 1-62.

SEXTON R., CRONSHAW J. & HALL J.L., 1971 — A study of the biochemistry and cyto-

chemical localization of Bglycerophosphatase in root tips of maize and pea. Protoplasma

73 :417-442.

SEXTON R. & HALL J.L., 1978 — Enzyme histochemistry. In : HALL J.L. (ed.), Electron

Source : MNHN, Paris

PHOSPHATASES IN THE PLACENTAL REGION OF PHYSCOMITRIUM 165

Microscopy and Cytochemistry of Plant Cells. Amsterdam, Oxford, New York : Elsevier/

North Holland Biomedical Press. Pp. 63-147.

SHAYKH М.М. & ROBERTS L.W., 1974 — A histochemical study of phosphatases in root

apical meristems. Ann. Bot. (London) 38 : 165-174.

SMITH D.L., 1972a — Localization of phosphatases in young gametophytes of Polypodium

vulgare L. Protoplasma 74 : 133-148.

SMITH D.L., 1972b — Staining and osmotic properties of young gametophytes of Polypo-

dium vulgare L. and their bearing on rhizoid function. Protoplasma 74 : 465-469.

TAKAMATSU Н., 1939 — Histologische und biochemische Studien über die Phosphatase.

Histochemische Untersuchungsmethodik der Phosphatase und deren Verteilung in ver-

schiedenen Organen und Geweben. Trans. Soc. Pathol. Japan 29 : 492-498.

THOMAS R.J., STANTON D.S., LONGENDORFER D.H. & FARR M.E., 1978 — Physiolo-

gical evaluation of the nutritional autonomy of a hornwort sporophyte. Bot. Gaz.

(Crawfordsville) 139 : 306-311.

TRIPODI G., 1970 — Localization of tryptophan rich proteins and Bglycerophosphatase

activity in Cuscuta haustorial cells. Protoplasma 71 : 191-196.

Source : MNHN. Paris

166 Е. CHAUHAN

Fig. 1 : Portion of haustorial foot in longisection shows the granular product of enzyme

reaction associated with the wall invaginations of the peripheral foot cells and those of the

abutting vaginula. Lead precipitate is seen associated with the plasmalemma (arrows) lining

the wall labyrinth and the wall separating the two peripheral foot cells. The golgi cisternae

also show deposits of the reaction product. In contrast, wall-membrane apparatus of the

vaginula cell has less lead precipitate. B-glycerophosphatase (GPase), I; F; P (x 12280).

Fig. 2 : Profile of another sporophyte-gametophyte junction from the middle zone of the

haustorium exhibits differential enzyme activity associated with the wall invaginations of

the foot and the vaginula. GPase, I; F; P (x 7720).

Fig. 34 : Cytoplasm of peripheral foot cell shows enzyme activity associated with golgi,

vesicles and some microbodies (arrows), ER cisternae show no lead phosphate deposits.

GPase, F; I; P (3 : x 18200; 4 : x 29000).

Fig. 5 : Nucleus of the peripheral foot cell reveals fine particles of lead phosphate

scattered throughout, but more concentrated in the nucleolus. Interestingly, deposits of

the reaction product are negligible at the nuclear membrane. GPase, E; I; P (x 7415). Fig. 6:

Nucleus of an axial cell of the haustorium shows a similar pattern of lead precipitate as in

Fig. 5. Accumulation of the reaction product is visible below the walls of the cells of the

axis. GPase, I; F; P (x 5505).

Fig. 7 : Cytoplasm of vaginula cell exhibits enzyme activity associated with golgi and

vesicles. GPase, 1: F; Р (x 20370). Fig. 8 : Control for GPase shows absence of lead deposits

on the membrane lining the wall labyrinth, These sites, however, have fine particles of lead

acetate staining. NaF; 1; F; P (x 14815). Fig. 9 : No indication of GPase activity is visible in

the nucleus after the use of inhibitor. NaF; I; F; P (x 24074).

Fig. 10 : Low magnification picture of Fig. 1 reveals the reaction product deposits

associated with the wall invaginations. Some deposits are also visible in regions external to

the plasmalemma (arrows). Note the fine particles of lead sulphide in the nucleus of the va-

ginula cell and the peripheral foot cell. GPase, I; Е; P (x 6000). Fig. 11 : Region similar to

the one shown in Fig. 10 with more alkaline phosphatase (AlPase) activity associated with

the wall-membrane apparatus of the vaginula cell as compared to that in the peripheral foot

cell. AlPase, 1: F; P (x 14000).

Fig. 12 : Distinct electron-dense deposits along the vacuoles and the membrane which

follows the contours of the wall invagination in the vaginula cell. AlPase, 1: F; P (x 12946).

Fig. 13 : Heavy deposits of the reaction product are seen associated with the cell wall of

the haustorial tip. AlPase. I; F; P (x 12500). Fig. 14 : Substrate-free control for alkaline

phosphatase shows no non-specific deposits of the reaction product on the vacuoles corres-

ponding to Fig. 12. AlPase, F; I-$; P (x 23215).

ABBREVIATIONS

Enzyme localization — Е; ЦР : Prefixation; incubation with substrate; processing.

Е; LS; P : Prefixation; incubation; minus substrate; processing. 1; F; Р : Incubation with

substrate; fixation: processing. NaF; 1; F; P : Treatment with sodium fluoride; incubation

with substrate plus sodium fluoride; fixation; processing.

Figures — er : endoplasmic reticulum, g : golgi, m : mitochondria, mb : microbody, mvb :

multivesicular body, n : nucleus, nu : nucleolus, p : plastid, pfc : peripheral foot cell, sgj :

sporophyte gametophyte junction, y : vacuole, vag : vaginula, ves : vesicle, wl : wall laby-

rinth.

Source - MNHN Paris

fi AA

BIBL.DU

MUSEUM

\ PARIS

hipa

Source : MNHN, Paris

Source : ММНМ, Paris

Source : MNHN. Paris

Source : MNHN, Paris

Cryptogamie, Bryol. Lichénol. 1988, 9 (2) : 173-181. 173

BIBLIOGRAPHIE BR YOLOGIQUE

D.LAMY!

Systématique, Nomenclature

88-132 DEGUCHI Н. - Neomeesia Deguchi, a new genus of the family Meesiaceae

from Southern South America. Bull. Natl. Sci. Mus., Ser. B (Bot) 1983, 9(4): 143-148, 2 fig.

(Dept. Biol., Fac. Sci., Kochi Univ., Kochi 780, Japan).

Diagn., descr., ill. de Neomeesia gen. nov., esp. type: Dichodontium paludella Besch.,

endémique d'Amérique du Sud.

88-133 DEGUCHI Н. - Diphyscium unipapillosum, sp. nov. (Diphysciaceae, Musci)

from Japan. J. Jap. Bot. 1984, 59(4): 97-103, 3 fig. (Ibidem).

Diagn., descr., Ш. de Diphyscium unipapillosum sp. nov. de la Préf. de Kochi, aff. de D.

fulvifolium Mitt.

88-134 INOUE Н. - Taxonomic miscellany on hepatics (2).J. Jap. Bot, 1983, 58(3): 65-71,

11 fig. (Dept. Bot, Natl. Sci. Mus., Shinjuku, Tokyo, Japan).

Cephalozia subgen. Haplocephalozia Schust. est syn. de Cephalozia (Dum.) Dum. et С.

uniloba М. Kitag. de С. leucantha Spruce - Gymnomitrium integerrimum М. Kitag. est syn. de

Cryptocoleopsis imbricata Amak. Descr., distr. des taxons étudiés.

88-135 LI X. - A new species of Sphagnum from Xizang (Tibet). Acta Bot. Yunnanica

1984, 6(1): 77-78, 1 fig. (Kumming Inst. Bot., Acad. Sinica, Kumming, Yunnan, People's Rep.

China).

Diagn., descr., ill. de Sphagnum multifibrosum Li et Zang sp. nov. du Tibet, aff. de 5.

palustre L.

88-136 SCHUSTER В.М. - Diagnoses of some new taxa of Hepaticae. Phytologia 1984,

56(2): 65-74 (Cryptog. Lab., Hadley, Massachusetts, USA).

Diagn. de taxons supra- et infragénériques parmi les Metzgeriales, Jungermanniales et

Marchantiales. Noter Radula yanoella sp. nov. et Pteroriccia gen. nov. (esp. type: Riccia villosa

Steph).

88-137 VYSOTSKAYA EJ. - [Comparative characteristics of Brachythecium starkei

(Brid.) B.S.G. and В. custum (Lindb.) Limpr. from the Western part of the USSR].

Ulrajins'k. Bot. Zurn. 1984, 41(5): 29-33, 4 fig., rés. angl. (Inst. Bot. im M. G. Kolodnogo AN

URSR, 290000 Lvov, URSS).

Morphol., distr., statut taxonom. de Brachythecium starkei (n=10) et B. curtum (n-20).

Voir aussi: 88-140, 88-156, 88-159, 88-164, 88-168, 88-169, 88-171, 88-172, 88-173, 88-

174, 88-176, 88-184, 88-185, 88-188.

Morphologie, Anatomie

88-138 IRELAND R.R. - The genus Plagiothecium in North America. Evansia 1985, 2(1):

4-9, 29 fig. (Natl. Mus. Nat. Hist., Natl. Mus. Canada, Ottawa, Canada K1 A 0M8).

Clé, descr., ill., hab., distr. des 7 Plagiothecium présents en Amérique du Nord.

1 Laboratoire de Cryptogamie, 12 rue Buffon, F-75005 Paris.

Source : MNHN, Paris

174 BIBLIOGRAPHIE BRYOLOGIQUE

Voir aussi: 88-132, 88-133, 88-134, 88-135, 88-136, 88-137, 88-140, 88-156, 88-162, 88-

164, 88-170, 88-172, 88-175, 88-184, 88-185, 88-188, 88-189.

Cytologie, Ultrastructure

88-139 SINGH J., BLACKWELL B.A., MILLER R.W. and BEWLEY J.D. - Membrane

organization of the desiccation-tolerant moss Tortula ruralis in deshydrated states,

PL Physiol. (Lancaster) 1984, 754(4): 1075-1079, 4 fig. (Chemistry & Biol. Res. Inst, Agric.

Canada, Ottawa, Ontario K1A 0C6 Canada),

Résonance magnétique nucléaire au ЗІР et ultrastructure des membranes deTortula ruralis

lors de la déshydratation. Observation de vésicules pouvant servir de réservoirs lors des échanges

membranaires, en dessiccation ou réhydratation.

Voir assi: 88-137, 88-140, 88-184.

Génétique

88-140 DEWEY R.M. - Taxonomic and populational studies of the thallose liverworts

Riccia subgenus Riccia . Ph. D. Thesis, Texas A & M Univ., 1986, 141 p., 14 tabl., 48 fig.

Electrophorèse sur gel d'amidon et caryologie pour établir plus précisément les délimitations

des esp. dans le sous-genre Riccia et observer les variations dans une population naturelle de R.

dictyopsora Howe. Remarquer une petite différence caryotypique entre les 12 esp. examinées: n=8

(141ш) chez toutes, sauf А. macallisteri (n=16) et К. nigrella (n=8?). Les phénotypes

électrophorétiques permettent une identification assez sure d'un matériel peu aisément assignable à

une esp. par la seule morphologie. R. hirta et А. eldeeniae seraient synonymes. Les observ.

morphol. et électrophor. permettent la mise en évidence de 2 esp. nouv., non encore identifiées. Un

survol des 8 loci d'enzymes chez R. dictyopsora met en évidence 3 génotypes distincts parmi les 38

localités échantillonnées aux USA. Ce taxon serait composé de plusieurs esp. affines.

88-141 ZIELINSKI В. - Genetic variation in the liverwort genus Pellia with special

reference to Central European Territory. Symp. Biol. Hung. 1987, 35: 175-189, 1 tabl., 6 fig.

(Dept. Genetics, A. Mickiewicz Univ., Dabrowskiego 165, 60-594 Poznan, Poland).

Electrophorèse sur gel d'amidon de 10 enzymes chez Pellia. P. neesiana et P. borealis ont

une structure génétique uniforme. Peu de variabilité dans les populations polonaises de Pellia. P.

borealis, diploïde, est assez différent de Р. epiphylla .

Physiologie, Chimie

88-142 DEISING Н. - In vivo studies on the regulation of nitrate reductase in

Sphagnum species. Symp. Biol. Hung. 1987, 35: 59-69, 5 fig., 1 tabl. (Bot. Inst, Christian-

Albrechts Univ., Biologiezentrum, Olshausenstr. 40-60, D-2300 Kiel 1).

La nitrate réductase chez Sphanum magellanicum est régulée par les substances disponibles

dans le substrat. Le rôle du co-substrat est discuté. Le МН+4 n'a pas d'effet significatif, dans les

conditions naturelles, ni sur l'induction ni sur l'activité de la nitrate réductase de la sphaigne.

88-143 HUNECK S., CONNOLLY J.D., HARRISON L.J., JOSEPH R.S.L and POCS T. - 1-(3,4-

Dihydroxy-5-methoxyphenyl)-3-methylbut-2e,e from the liverwort Plagiochila

rutilans. Phytochemistry 1984, 23(10): 2396-2397 (Inst. РІ. Biochem., Res. Centre Molec. Biol. &

Med., Acad. Sci. GDR, GDR-401 Halle/Saale, Weinberg).

88-144 RUDOLPH H., DEISING H. and VOIGT J.U. - The tolerance of raised bog

Sphagnum species in respect to inorganic nitrogen.Symp. Biol. Hung. 1987, 35: 71-80, 1

tabl., 4 fig. (Bot. Inst., Christian-Albrechts Univ., Olshausenstr. 40-60, D-2300 Kiel 1).

Source - MNHN, Paris

BIBLIOGRAPHIE BR YOLOGIQUE 175

Mesures du pourcentage de croissance, du contenu en chlorophylle, de la photosynthése

nette, de l'activité de la nitrate réductase pour suivre l'influence du NH*4-N et du NO^3-N chez

Sphagnum magellanicum cultivé en conditions contrôlées.

88-145 TUBA Z. - Light, temperature and desiccation response of COp-exchange in

the desiccation-tolerant moss, Torfula ruralis . Symp. Biol. Hung. 1987, 35: 137-149, 4

fig., 2 tabl. (Inst. Ecol. & Bot., Hung. Acad. Sci., Н-2163 Vacrato!).

Tortula ruralis croissant sous de fortes intensités lumineuses présente une capacité

photosynthétique extrémement basse, conséquence de la réponse spécifique à la température,

causée par un effort d'économie d'eau.

88-146 WU P.C, LI D.K, GAO С.Н. - Light and epiphyllous liverworts in the

subtropical evergreen forest of South East China. Symp. Biol. Hung. 1987, 35: 27-32, 3

fig. (Inst. Вог, Acad. Sinica, 141 Hsi Chih Men Wai Ta Chie, Beijing, China).

Voir aussi:88-139, Ecophysiologie, 88-184, 88-190, 88-193,

Ecophysiologie

88-147 BALO К. - Some photosynthesis-ecological characteristics of forest

bryophytes. Symp. Biol. Hung. 1987, 35: 125-135, 2 tabl., 3 fig. (Inst. Ecol. de Bot, Hung.

Асай, Sci., Н-2163 Vacratot).

Contenu total en pigment photosynthétique et courbes de photosynthése nette de 3

mousses (provenant de 2 sites forestiers à régime lumineux différents) par chromatographie en

couche mince ct analyse gazeuse en infrarouge.

88-148 BIRO G. and DEBRECZY Z. - Ecological conditions in a photophilous-

xerothermotolerant moss community (Mannio-(Grimaldio-) Tortelletum inclinatae

(Assoc. поу.)) and microrespirometric test of its species. Symp. Biol. Hung. 1987, 35:

81-101, 4 tabl., 11 fig. (Dept. Bot, Hung. Nat. Hist. Mus., Budapest, Hungary).

La période active des mousses est réduite ici à la période froide de l'année et, avec

photopériode courte, aux heures matinales de la saison chaude. Les mémes facteurs génent

l'adaptation des différentes esp. à la haute-température du site. Lichens associés.

88-149 ELIAS P. and MASAROVICOVA E. - Some ecophgysiological features in

woodlands mosses in SW Slovakia 1. Water relations. Symp. Biol, Hung. 1987, 35: 103-

111, 1 fig., 1 tabl. (Inst. Exp. Biol. & Ecol., Slovak Acad. Sci., Dept. Prod. Ecosystems, Dubravska

cesta 14, Cs-81434 Bratislava).

Mise en évidence parmi les esp. de mousses des Monts Male Karpaty, de 2 groupes: esp. à

bas pourcentage de perte d'eau et esp. à fort pourcentage de perte d'eau.

88150 MASAROVICOVA E. and ELIAS P. - Some ecophysiological features in

woodlands mosses in SW Slovakia 2. Chlorophyll content and photosynthesis, Symp

Biol. Hung. 1987, 35: 113-123, 1 fig., 2 tabl. (Inst. Exp. Biol. & Ecol., Slovak Acad. Sci., Dept.

Prod. Ecosystems, Dubravska cesta 14, Cs-81434 Bratislava).

Contenu en chlorophylle, rendement de la photosynthèse et respiration varient selon les esp.

88-151 RUSSELL $. - Water relations and nutrient status of bryophyte communities at

Marion Island (Subantarctic). Symp. Biol. Hung. 1987, 35: 39-57, 3 fig. (PL Sci. Dept., Fort

Hare Univ., Alice, Ciskei, South Africa).

L'humidité du sol est le facteur principal contrôlant la distribution des bryophytyes. Rôle

additionnel du statut nutritif en relation avec la composition spécifique des communautés, notamment

dans les sites bien drainés ou biologiquement influencés.

Voir aussi: 88-155, 88-160, 88-190,

Source : ММНМ. Paris

176 BIBLIOGRAPHIE BRYOLOGIQUE

Répartition, Ecologie, Sociologie

88-152 BOUDIER Р. - Quelques observations bryologiques: Rhodobryum ontariense

(Kindb.) Kindb. en Auvergne, Fissidens curnowii Mitt. dans les Cévennes. Bull. Soc.

Bot. Centre-Ouest n.s., 1984, 15: 125-126 (Muséum de Chartres, 12 rue St Michel, F-28000

Chartres).

88-153 DUVIVIER J.P., SOTIAUX А. et O., POHL Н. et BRUYNSEELS G. - Intérêt

bryologique du Gros Têne du Bi et de ses environs (commune de айу, province de

Namur, Belgique). Natura Mosana 1984, 37(2): 46-49 (75 ry del Praile, B-6423 Somzée).

88-154 FRAHM ІР. - Struktur und Zusammensetzung der epiphytischen

Moosvegetation in Regenwälderen NO-Perus. Вай. Nova Hedwigia 1987, 88: 115-141, 26

fig. (Univ. Duisburg, FB 6, Bot., Postfach 101629, D-4100 Duisburg).

Etude de la végétation des bryophytes épiphytes autour des troncs dans les forêts humides

du Peru МЕ, entre 200 et 3200 m. Fréquence, abondance, relations mousses/hépatiques, exposition,

formes de vie, zonation verticale des épiphytes le long du transect.

88-155 FRAHM ТР. - Ökologische Studien über die epiphytische Moosvegetation in

Regenwäldern NO-Perus. Beih. Nova Hedwigia 1987, 88: 143-158, 7 fig. (Ibidem).

Mesures de la phytomasse des bryophytes épiphytes, de l'intensité lumineuse dans l'habitat,

de la température et de l'humidité relative, le long d'un transect de 200 à 3200 m dans les forêts

humides du Pérou NE. La phytomasse augmente des basses terres à la limitee de forêt et montre une

correlation avec l'intensité lumineuse et l'humidité. Les mesures de photosynthése en laboratoire avec

différentes combinaisons de température et d'intensité lumineuse révèlent que les hautes températures

associées à de basses intensités lumineuses, comme c'est le cas dans les forêts humides de basse

altiutude, ne permettent aucun gain de phytomasse.

88-156 FURUKI Т. and INOUE H. - The Hepaticae and Anthocerotae of the Mts.

Yatsugatake, Central Japan. Bull. Май. Sci. Mus., Ser. B (Bot) 1984, 10(1): 1-25, 1 сапе

(Bot. Inst., Fac. Sci., Hiroshima Univ., Hiroshima, Japan).

Liste annotée de 194 esp. d'hépatiques et d'anthocérotes des Mts Yatsugatake (Japon

central). Notes morphol. et taxonom. pour Lophozia silvicola М. Kitag,, Nardia breidleri (Limpr.)

Lindb., Cephaloziella byssacea (Roth) Wamst. var. asperifolia (Tayl.) Масу, C. massalongoi (Spr)

K Mall.

88-157 GOBAT J.M. - Importance des bordures de tourbières pour la protection des

Hauts-Marais: exemple de deux tourbières du Jura suisse. Bull. Soc. Neuchâtel. Sci. Nat. ,

4° sér., 1984, 107: 25-38, 2 tabl., 8 fig. (Lab. Ecol. Végét., Inst. Bot., 22 chemin de Chantemerle,

CH-2000 Neuchâtel 7).

88-158 GRADSTEIN S.R. und FRAHM J.P. - Die floristische Hóhengliederung der Moose

entlang des BRYOTROP-Transektes in NO-Peru. Beih. Nova Hedwigia 1987, 88: 105-113,

2 fig. (Inst. Syst. Bot., Heidelberglaan 2, NL-3508 TC Utrecht).

A partir des récoltes de BR YOTROP au Pérou NE, la zonation altitudinale des bryophytes a

été déterminée au moyen des discontinuités floristiques. Description de ces zones en utilisant les

données de la physionomie de la végétation et des données floristiques. Comparaison avec la

Colombie.

88-159 GUERRA J. y PUCHE F. - Bryum dunense Smith et Whitehouse en la Peninsula

iberica y Baleares. Observaciones taxonomicas, corologicas y fitosociologicas. Acta

Bot, Malacitana 1984, 9: 85-92, 1 carte, 1 fig., 1 tabl. (Dept. Bot., Fac. Ci., Univ. Malaga, Malaga,

Espana).

Chorologie, écol., phytosociol. de Bryum dunense dans la Péninsule ibérique et aux

Source : MNHN, Paris

BIBLIOGRAPHIE BRYOLOGIQUE 177

Baléares (nouv. loc. en Espagne et au Portugal). Descr. du Tortello flavovirentis-Bryetum dunensis

commun. arénicole nouy. appartenant au Barbuletea unguiculatae.

88-160 HEBRARD J.P. - Contribution à l'étude du peuplement bryophytique des

pinèdes de pin d'Alep sur substrat calcaire dans les Bouches-du-Rhône et le Var

sud-occidental. Ecol. Medit. 1984, 10(1-2): 183-203, 3 tabl., 4 fig. (Lab. Bot. Ecol. Médit., Fac.

Sci. Techn. St Jérôme, rue Henri-Poncaré, F-13397 Marseille Cedex 13).

Etude écol. (caractér. physico-chimiques de l'horizon édaphique supérieur notamment) et

bryosociol. des pinèdes du Querco-Piceetum halepensis Loisel 1971 sous ass. typicum sur roche

mère calcaire dans les Bouches-du-Rhône et le Var de 0 à 600 т. Rôle de la strate muscinale dans

l'accumulation de matériel organique. Relations bryophytes et pH du sol. Liste des taxons.

88-161 HEBRARD J.P. - Contribution à l'étude des Muscinées du Parc National du

Mercantour. Observations floristiques et écologiques dans le bassin supérieur de la

Tinéee. IT. Inventaire bryoécologique des affleurements sédimentaires de part et

d'autre de la ligne de crête limitant au Nord le bassin supérieur de la Tinéee. Bull.

Soc. Linn. Provence "1983" 1984, 35: 29-51 (Ibidem).

Liste des loc. des affleurements sédimentaires dans le bassin supérieur de la Tinée (Alpes-

Maritimes). Inventaire des 110 mousses et 15 hépatiques avec hab., loc., distr., notes écol. 11

moysses et 3 hépatiques sont ouv. pour td région.

88-162 IWATSUKI Z. and DEGUCHI A. - Three additional localities of Aptychella

robusta (Broth.) FI. in Japan. Proc. Bryol. Soc. Japan 1984, 3(10): 163-165, 1 fig., en japonais,

rés. angl. (Bot. Inst., Fac. Sci., Hirshoma Univ., Higashisenda-machi, Naka-ku, Hiroshima 730,

Japan).

Descr., distr. horizont. et vertic. d'Aptychella robusta au Japon. Comparaison avec

Aerobryum specium.

88-163 IWATSUKI 2. and HATTORI S. - Microclimate at Inotani Valley near Nichinan,

S. Japan. Symp. Biol. Hung. 1987, 35: 11-25, 8 fig., 1 tabl. (Ibidem).

П est raisonnable de considérer les mousses pendantes et les hépatiques épiphylles comme

étant de bons indicateurs d'un certain microclimat.

88-164 JOHANNSSON В. - Notes on some Icelandic bryophytes species. Acta Bot.

Islandica 1984, 7: 37-50 (Icelandic Mus, Nat. Hist., Р.О. Box 5320, Reykjavik, Iceland).

Descr. et révision des esp. appartenant aux groupes Bryum bicolor, В. capillare, B.

erythrocarpum, Racomitrium canescens. 12 taxons sont nouv. pour l'Islande, Notes tax. et morph.

pour certains taxons.

88-165 LECOINTE А., PIERROT К.В. ( et BOUDIER P. et ROGEON М.А) - Bryophytes

observées pendant la dixième session extraordinaire de la S.B.C.O.: Vosges-Alsace.

Bull. Soc. Bot. Centre-Ouest n.s., 1984, 15: 269-300, 13 tabl., 31 fig. (Lab. Phytogéogr., Univ.

Caen, F-14032 Caen Cedex).

Liste des stations visitées, liste des 360 taxons récoltés, notes phytosociol.

88-166 LONG D.G. - The moss Fissidens rivularis (Spruce) B.S.G. in Kerry, new to

Ireland. /rish Naturalists’ J. 1984, 21(8): 347-348 (Royal Bot. Gard., Edinburgh, Scotland).

88-167 MASZA K., KOVACS-LANG E. and SNAKIN V.V. - Changes in soil pH along the

zonation of eryptogamous synusia at Bugac (Hungary). Symp. Biol. Hung. 1987, 35: 33-

37, 1 tabl. (Dept. PI, Tax. & Ecol., L. Eötvös Univ., Kun В. ter 2, Н-1083 Budapest).

88-168 OCHI H. - Revision of the Australasian Bryoideae (Musci). Supplement ۰

Studies on the Bryum type specimens of К. Brown ter. New Zealand J. Bot. 1984, 22(2):

179-182 (Biol. Inst., Fac. Educ., Tottori Univ., Koyama-cho, Tottori 680, Japan).

Révision des spécimens types de Bryum décrits раг К. Brown ter.: Brachymenium exile est

Source : ММНМ Paris

178 BIBLIOGRAPHIE BRYOLOGIQUE

nouv. pour la Nouv. Zélande; Bryum cylindrothecium R. Brown ter. est une bonne esp. Correction

de nombreuses déterminations faites par H.N. Dixon.

88-169 OCHYRA R. - Mchy Skalic Nowotarskich i Spiskich (Pieninski Pas Skalkowy)

- Moss flora of the Nowotarskie and Spiskie Klipper (Pieniny Klippen Belt, Western

Carpathians Mts). Fragm. Florist. Geobot. "1982" 1984, 28(3): 419-489, 16 fig. (Lab. Bryol.

Lichenol., Inst. Bot, Polish Acad. Sci., Lubicz 46, PL-31-512 Krakow).

Distr. horizontale et verticale, éléments écol. des mousses de Nowotarskie et Spiskie Klipper

(Carpathes W). Catalogue de 249 esp. avec loc. Seligeria austriaca Schauer et Rhodobryum

ontariense (Kindb.) Kindb. sont nouv. pour la Pologne. 2 comb. nouv. : Didymodon fallax var.

brevifolius (Brid.) (=Bryum brev.) et Campyliadelphus stellatus var. protensus (Brid.) (= Hypnum

prot.). Clé aux Mniaceae et aux Plagiotheciaceae de Pologne.

88-170 PIERROT R.B., SCHUMACKER В. et WATTEZ J.R. - Lophozia capitata (Hook.)

Macoun (Hepaticae), nouveau pour la bryoflore française, dans le Pas-de-Calais et

en Charente-Maritime. Bull. Soc. Bot. Centre-Ouest n.s., 1984, 15: 103-115, 2 tabl., 2 cartes, 2

fig. (Les Andryales, F-17550 Dolus d'Oléron).

Descr, ill., synécol., répart. en Europe de Lophozia capitata récolté dans le Pas-de-Calais et

en Charente-Maritime. Lichens associés.

88-171 PIERROT R.B. (avec la collab. de ROGEON M.A., SAPALY J., VILKS A.) - L'année

bryologique 1983. Bull. Soc. Bot. Centre-Ouest n.s., 1984, 15: 117-123 (Ibidem).

Espèces intéressantes du Centre-Ouest de la France et du Massif central. Remarques sur

Campylopus oerstedianus (C. МВП.) Mitt, Bryum stirtonit Schimp., В. elegans Nees ex Brid.

Classif, et clés pour les Майит 51. en France.

88-172 ROGEON M.A. et SCHUMACKER R. - Leptodontium flexifolium (With) Hampe

et L. gemmascens (Mitt. ex Hunt) Braithw. sur les toits de chaume de la Haute-

Adour (Hautes-Pyrénées, France). Bull. Soc. Bot. Centre-Ouest n.s., 1984, 15: 81-102, 3 tabl.,

4 cartes, 3 fig. (14 rue Henri-Dunant, F-86400 Civray).

Distr., écol., morphol., répart. en Europe de Leptodontium flexifolium et L. gemmascens

sur toits de chaume entre 650 et 1250 m dans la vallée de l'Adour.

88-173 SCHULTZE-MOTEL W. und MENZEL M. - Die Laubmoosflora im BRYOTROP-

Transekt von Peru. Веі, Nova Hedwigia 1987, 88: 9-59, 2 fig. (Bot. Сап. & Bot. Mus., Berlin-

Dahlem, Kónigin-Luise-str. 6-8, D-1000 Berlin 33)

Liste des loc. Liste des 217 mousses récoltées sur un transect de 190 km entre Yurimaguas

(200 m) et Leimebamba-Balsas (3400 m), Pérou NE. 56 taxons sont nouv. pour le Pérou. Clés aux

genres et aus esp. Diagn., descr., ill. de Tortella bryotropica Zander sp. nov. - Nombreux nowy. syn. -

7 comb. nouv.: Kindbergion oedogonium (C.Müll:) Ochyra (=Cratoneuron ), Meteorium illecebrum

var. latifolium (Lindb.) Menzel (=Neckera lat. ), Rhynchostegium peruviense (Williams) Ochyra

(<Hygrohypnum ), Rh. validum (Herz.) Ochyra (-Hygrohypnum ), Stereobryon elamellosum (Herz.)

Menzel (=Catharinea ), Zelometeorium patens var. allionii (Manuel) Menzel (-Zelometeorium all.),

Z. patens var. recurvifolium (Hornsch.) Menzel (=Pilotrichum rec. ).

88-174 SCHULTZE-MOTEL №. und MENZEL M. - Die Lebermoosflora im BRYOTROP-

Transekt von Peru. Beih. Nova Hedwigia 1987, 88: 61-104, 1 fig. (Ibidem).

Liste des loc., liste de 261 hépatiques sur un transect de 190 km entre Yurimaguas (200 m)

et Leimebamba-Balsas (3400 m). 119 taxons sont nouv. pour le Pérou. Clés aux genres et aux esp. -

Diagn., descr., ill. de Radula peruviana Yamada sp. nov. - Nombreux nouv. syn. - 5 comb. nouv.:

Riccardia hymenophytoides (Spruce) Meensk (=Aneura ), К. plumaeformis (Spruce) Hassel ex

Meenks (=Aneura ), R. tenuicula (Spruce) Schiffn. ex Meenks (= Aneura ), R. trichomanoides

(Spruce) Hässel ех Meenks (=Anewra ), Trichocolea flaccida var. floccosa (Herz, et Hatcher) Menzel

(-Trichoc. floce. ).

88-175 SOTIAUX A. et DUVIVIER J.P. - Lejeunea lamacerina (Steph.) Schiffn.,

Source - MNHN, Paris

BIBLIOGRAPHIE BRYOLOGIQUE 179

hépatique nouvelle pour la Belgique et le département des Ardennes en France.

Dumortiera 1983, 27: 23-26, 2 fig. (Chaussée de Bruxelles 676, В-1410 Waterloo).

Descr., ill., écol,, distr. de Lejeunea lamacerina nouveau pour la Belgique et les Ardennes.

88-176 VANA J. and INOUE Н. - Studies in Taiwan Hepaticae V. Jungermanniaceae.

Bull. Natl. Sci. Mus. , Ser. В (Bot.) 1983, 9(4): 125-142, 5 fig. (Dept. Cryptog. Bot., Charles Univ.,

Praha, Czechoslowakia).

Clés, notes taxon. et loc. pour 2 Mylia, 1 Notoscyphus , \Scaphophyllum, 13 Jungermannia,

2 Nardia. Taxons exclus. Noter syn. nouv. pour J. truncata Nees,

88-177 WERNER 1. et REICHLING L. (en collab. avec SCHUMACKER К.) - Lophozia

heterocolpos (Thed.) Howe, Scapania aequiloba (Schwaegr.) Dum. et Scapania

mucronata Buch (Hepaticae) dans la vallée de la Pétrusse (Grand-Duché de

Luxembourg). Bull. Soc. Roy. Bot. Belgique 1984, 117: 109-121, 4 fig., 1 tabl. (14 rue Henri-

Pensis, L-2322 Luxembourg).

Distr. en Europe de 3 hépatiques récoltées en 1982, dans la vallée de la Pétrusse au centre de

la ville de Luxembourg: Lophozia heterocolpos nouv. pour le Luxembourg, Scapania aequiloba non

revu depuis un siècle, ni dans le Grand-Duché de Luxembourg, ni en Belgique, Scapania mucronata

nouv. pour le Grand-Duché et pour la Belgique. Ecol. et phytosociol. du site de Luxembourg.

Voir aussi: 88-132, 88-133, 88-134, 88-135, 88-136, 88-137, 88-138, 88-184, 88-185, 88-

186 88-187, 88-188, 88-189, 88-191, Ecophysiologie, 88-204, 88-206, 88-210, 88-217.

Bryophilie

88-178 SIMON E. - Lyophyllum palustre , a parasite on Sphagnum. Symp. Biol. Hung.

1987, 35: 165-174, 1 fig., 1 tabl. (Bot. Inst, Christian-Albrechts Univ., Olshausenstr, 40-60, D-2300

Kiel 1).

Etudes microscopiques et enzymatiques du mécanisme de l'infection du Sphagnum par

Lyophyllum palustre .

Techniques

88-179 RAEYMAEKERS б. and LONGWITH LE. - The use of dimethyl sulfoxide

(DMSO) as a solvent to extract chlorophyll from mosses. Symp. Biol. Hung. 1987, 35:

151-164, 5 fig. (Dept. Biol. Sci., Michigan Technol. Univ., Houghton, MI 49931, USA).

Documentation, Histoire des sciences

88-180 ANDO Н. - Moss gardening in Japan. Symp. Biol. Hung. 1987, 35: 3-10 (Bot. Inst,

Hiroshima Univ., Higashisenda-Machi, Naka-ku, Hiroshima 730, Japan).

88-181 FRAHM J.P. - Das Untersuchungsgebiet. Вей. Nova Hedwigia 1987, 88: 1-5, 1

carte, 2 fig. (Univ. Duisburg, FB 6, Bot., Postfach 101629, D-4100 Duisburg).

Carte de la région étudiée et altitude du transect au Pérou NE, lors de l'expédition

BRYOTROP 1982.

88-182 FRAHM J.P. Zur bryologischen Erforschung des Untersuchungsgebietes. Beih.

Nova Hedwigia 1987, 88: 7-8 (Ibidem).

Historique des récoltes au Pérou . Bibliographie,

88-183 IRELAND R.R. and LEY L.M. - Additions to the type specimens of bryophytes in

the National Museum of Natural Sciences, National Museum of Canada- 1 (1986).

(Additions to the list in Syllogeus 47, 1984). Sine dat, 5. loc., 8 p.

Source : MNHN, Paris

180 BIBLIOGRAPHIE BRYOLOGIQUE

Voir aussi: 88-187.

Ouvrages généraux

38-184 BUCK W.R. - Bryostephane steereana: a collection of bryological papers

presented to William Campbell Steere on the occasion of his 80th birthday. Мет.

New York Bot. Gard. 1987, 45: 1-778, ill. (New York Bot. Gard., Bronx, New York 10458, USA)

83 bryologues rendent hommage à W.C. Steere, en lui offrant ce volume composé de 67

articles couvrant les domaines abordés par W.C. Steere durant sa longue carrière: morphologie,

anatomie, cytologie, physiologie, écologie, floristique, paléobryologiue, bryogéographie, mais aussi

monographies, histoire de la bryologie. Cet ensemble est précédé d'un témoignage de Dorothy

Osbome Steere, son épouse, relatant la vie d'un couple de botanistes. L'éditeur a eu le courage d'y

ajouter un index des auteurs et des taxons (30 p.), qui rend cet hommage trés pratique. La lecture de

ces articles offre au bryologue un apercu très actualisé des recherches bryologiques en Amérique,

mais aussi en Australie, en Océanie et dans l'Antarctique. A cette occasion plusieurs éponymes ont

été créés: Anombryum steerei Shaw, Frullania steereana Hattori, Leucomium steerei Allen &

Veling, Orthotrichum steerei Lewinsky, Pylaisiella steerei Ando & Higuchi, Racomitrium steerei

Griffin, Radula steerei Grolle, Schistidium steerei Ochyra, Steerecleus Н. Robinson gen. nov.,

Steereochila ecuadorica Н. Inoue gen. et sp. nov., Steereomitrium minutum Е.О. Campbell gen. et

sp. nov.

88-185 FLATBERG КІ. - Taxonomy, morphovariation, distribution and ecology of the

Sphagnum imbricatum complex with main reference to Norway.Gunneria 1986, 54: 1-

118, 32 fig., 7 tabl. (Dept. Bot., Univ. Trondheim, N-7055 Dragvoll).

Distinction de 2 sous -esp. chez Sphagnum imbricatum: subsp. austinii (Sull.) Flatb. et

subsp. affine (Ren. et Card.) Flatb. Descr., relations, distr. en Norvège, habitat, écol. et

environnement de ces taxons. Etude du dimorphisme sexuel des feuille de la sous-esp. affine .

Histoire paléoécologique. Etude des variations à partir des échantillons d'herbiers, récoltes

récentes et cultures.

88-186 FREY W. - Moosflora und -vegetation іп Regenwäldern NO-Perus.

Ergebnisse der BRYOTROP-Expedition nach Peru 1982. Вей. Nova Hedwigia 1987,

88: 1-159, 41 fig., 2 tabl., 1 carte.

Voir 88-154, 88-155, 88-158, 88-173, 88-174, 88-181, 88-182.

88-187 KANDA H. - Catalog of moss specimens from Antarctica and adjacent

regions housed mainly in the herbarium of the National Institute of Polar

Research. Tokyo: National Institute of Polar Research, 1987. 186 p., 4 tabl., 2 fig., 1 pl. coul.

(9-10, Кара 1-chome, Itabashi-ku, Tokyo, Japan).

Inventaire de 6000 spécimens: 5500 mousses et 500 hépatiques. Historique des récoltes,

régions étudiées.

88-188 MAGILL ВЕ. - Bryophyta. Part 1. Mosses. Fascicule 2. Gigaspermaceae-

Bartramiaceae. In: О.А. LEISTNER, Flora of Southern Africa. Pretoria: Botanical Research

Institute, 1987, pp. i-ix, 293-443, fig. 84-125, cartes 111-176 (Missouri Bot, Gard., Р.О. Box 299,

St Louis, Missouri 63166-0299, USA).

Clés, descr., ill, distr. en Afrique du Sud, des genres et esp. des Gigaspermaceae,

Ephemeraceae, Funariaceae, Splachnaceae, Bryaceae, Mniaceae, Eustichiaceae, Aulacomniaceae,

Bartramiaceae présents en Afrique du Sud, Transkei, Lesotho, Swaziland, Bophuthatswana, SW

Africa/Namibie, Bostwana et Venda. Diagn. des esp. nouv.: Breutelia elliptica du Cap, Ephemerum

namaquense du Cap, Qhathlamba debilicostata gen. et sp. nov. du Lesotho. Comb. nouv.: Anacolia

breutelii (C.Müll.) (= Bartramia ), А. b. var. squarrifolia (Sim) (= Ванг. squar. ), Bartramia

Source : MNHN, Paris

BIBLIOGRAPHIE BRYOLOGIQUE 181

compacta var. macowaniana (СМЕП. (= Ванг. macow. ), Breutelia substricta (C.Milll.) (=Bartr. ),

Funaria clavata (Mitt.)(= Entosthodon ), Е. succuleata (Wager & Wright) (=Physcomitrium ),

Physcomitrium spathulatum var. sessile (Shaw) (-Physc. ses. ).

88-189 MURRAY В.М. - Andreaeobryaceae - Tetraphidaceae. In: G.S. MOGENSEN,

Illustrated Mossflora of Arctic North America and Greenland. 3. Meddelelser om

Grønland, Bioscience 1987, 23: 1-36, 16 fig., 13 cartes (Univ. Alaska Mus., Fairbanks, Alaska

99775-1200, USA).

Descr., ill. , distr. d'1 Andreaeobryum ,7 Andreaea , 1 Buxbaumia , 1 Diphyscium , 2

Tetraphis , 1 Tetradontium présents dans l'Arctique. Illustrations de V.G. Friis et G.S. Mogensen.

88-190 POCS T., SIMON T., TUBA Z. and PODANI J. - Proceedings of the IAB Conference

of Bryology. Budapest, Vacratot, Hungary, 5-10 August, 1985. Symp. Biol. Hung.

1987, 35: ХХ, 1-902, ill. (en 2 volumes).

72 contributions réparties en 6 sessions: écologie physiologique, reproduction et dispersion,

écologie des communautés, écologie des populations, bryophytes comme bioindicateurs. Préface de

Т. Pocs et Т. Simon. Le Symposium a été ouvert et cloturé par P.W. Richards. Index matières et liste

des participants.

88-191 WERNER J. - Liste rouge des bryophytes du Grand-Duché de Luxembourg.

Trav. Sci. Mus. Hist. Nat. Luxembourg 1987, 11: 1-42, 10 fig., 3 tabl.

Présentation en 3 parties: epèces probablement disparues (4 hépatiques et 5 mousses), esp.

menacées (25 hépatiques et 34 mousses), espèces potentiellement menacées et dignes de protection

(33 hépatiques et 65 mousses). Recension des principaux biotopes menacés. Causes de ces

disparitions: méthodologie des agriculteurs et viticulteurs, pollution atmopshérique, tourisme.

Source : MNHN. Paris

Cryptogamie, Bryol. Lichénol. 1988, 9 (2) : 182-186.

BIBLIOGRAPHIE LICHENOLOGIQUE

D.LAMY!

Systématique Nomenclature

88-192 YOSHIMURA I. - Taxonomic studies on Lobaria crenulata and its allies. J.

Hattori Bot. Lab. 1984, 57: 97-126, 6 pl., 5 tabl., 5 fig. (Kochi Gakuen College, 292 Asahi-

tenjuncho, Kochi 780, Japan).

Observ. morph. et chimiques sur les Lobaria crenulata sens. lat. Clé, tax., descr., chimie,

spéc. examinés, variation des 5 taxons reconnus: Lobaria crenulata (Hook. in Kunth) Trev., L

exornata (Zahlbr.) c.n. (=Lobaria crenulata var. e. ), L. e. var. corallophora var. nov. de Costa

Rica, L. subexornata sp. nov. de Costa Rica, L. pallida (Hook. in Kunth) Trev. Position

taxonomique de Lobaria crenulata et de ses alliés A pseudocyphelles sur le cortex supérieur du

thalle. Ils forment un groupe naturel. Distr., surtout en Amerique du Sud.

Voir aussi: 88-195, 88-215, 88-224, 88-225, 88-228, 88-230.

Morphologie, Anatomie

Voir: 88-192, 88-224, 88-225, 88-226, 88-227, 88-228, 88-230.

Physiologie, Chimie

88-193 ATTARPOUR N., TURK К. und HOFMANN W. - Radionuklide im Wald-

Okosystem. Forschungsinitiative gegen das Waldsterben Bericht 1987 . Ergebnisse aus der

Immissionforschung 1987: 39-48, 9 fig. (АЫ. Biophys. & Abt. Okophysiol. Pfl., Univ. Salzburg, A-

5020 Salzburg).

Activité 137Cs et 49K dans les lichens, mousses, champignons et cert. plantes vasculaires,

88-194 CHAMPION-ARNAUD P. and LALLEMANT В. - Localisation of ammonia

assimilating enzyme activities in the course of the cycle of Nostoc filaments

isolated from the lichens Peltigera canina (L.) Willd. and Peltigera praetextata

(Flórke ex Sommerf.) Zopf. Lichen Phsysol. Biochem. 1986, 1: 104-116, 6 fig. (Lab. Biol. &

Cytophysiol. Vég., Univ. Nantes, 2 chemin de la Houssinière, F-44072 Nantes Cedex).

Activité de la déhydrogenase glutamique acide dans les cellules végétatives des trichomes,

soit dans toutes, quant la conentretion en ammonium est forte, ou seulement dans les cellules proches

des hétérocystes, lors de la syntèse d'ammonium, Elle n'est jamais présente dans les hormogonies ou

les hétérocystes. La synthétase glutamine est présente dans toutes les cellules végétatives des

trichomes.

88-195 FEIGE G.B., VIETHEN B., GEYER M. und FOLLMANN С. - Untersuchungen zur

Phytochemie der Flechtenfamilie Roccellaceae Chey. 1. Uber die Sekundirstoffe und

Chemotypen von Roccella hypomecha (Ach) Bory. J. Hattori Bot. Lab. 1986, 60: 143-

148, 3 tabl., 2 fig. (Bot. Inst., Univ. Essen, Universitátsstr. 5, D-4300 Essen 1).

Grande variation des produits secondaires dans les échantillons africains de Roccella

hypomecha. Distinction de 5 chemotypes. Acide dibenzofurane schizopeltique décellé pour la 1°

1 Laboratoire de Cryptogamie, 12 rue Buffon, F-75005 Paris.

Source - MNHN, Paris

BIBLIOGRAPHIE LICHENOLOGIQUE 183

chez Roccella, et ac. lécanorique et érythrine pour la 1° fois chez К. hypomecha. Les chémotypes

ayant Гас. schizopeltique sont considérés comme des taxons hautement dérivés.

88-196 RAVINSKAJA A.P. - De valore biologico acidorum lichenim notula. Noy. Sist.

Niz. Rast., Akad. Nauk SSSR Bot. Inst. Komarova 1984, 21: 160-178, 3 fig., en russe (bibliogr. 4

р).

88-197 SOLBERG Y. - Chemical constituents of the lichen species Cetraria islandica.

J. Hattori Bot. Lab. 1986, 60: 391-406, 6 tabl., 4 fig. (Chem. Res. Lab., Agric. Univ. Norway, P.O.

Box 31, N-1432 ÀS-NLH).

La plupart des composés détectés (acides gras, keto-acides, ac. dicarboxyliques, terpénoides,

composés aromatiques) chez Cetraria islandica le sont pour la première fois chez cette espèce.

Descr. de nouveaux composés.

Voir aussi: 88-148, 88-192, 88-225, 88-226, 88-227, 88-228.

Répartition, Ecologie, Sociologie

88-198 ABBASSI MAAF L. et ROUX Cl. - Lichens corticoles du Pare National de l'île de

Port-Cros (Var). Espèces nouvelles et corrections. Sci. Rep. Port-Cros Nail. Park "1985"

1986, 11: 13-21 (Lab. Bot. & Ecol. Medit., Fac. Sci. Techn. St Jérôme, rue Henri-Poincaré, F-13397

Marseille Cedex 13).

Notes, hab., distr. de 27 lichens corticoles et 1 champignon lichénicole nouv. pour Port-

Cros.

88-199 ANDREEV М.Р. - Analysis complexus specierum activarum lichenoflorae

planitiei elatae anjujensis. Nov. Sist. Niz. Rast., Akad. Nauk SSSR, Bot. Insi., Komarova

1984, 21: 123-127, 2 tabl., en russe.

88-200 ANDREEV М.Р. - Lichenes Paeninsulae Jamal. Nov. Sist. №. Rast., Akad, Nauk

SSSR, Bot. Inst. Komarova 1984, 21: 127-136, 1 fig., en nisse.

88-201 ANDREEV М.Р. - De compositione taxorum lichenoflorae planitiei elatae

anjujensis notula. Nov. Sist . Niz. Rast., Akad. Nauk SSSR, Bot. Inst, Komarova 1984, 21: 136-

140, 1 fig., en russe.

88-202 ANDREEV М.Р. - Lichenes epilithici rupestrium ad limitem regionis

leningradensis et Kareliae. Nov, Sist. Niz. Rast., Akad. Nauk SSSR, Bot. Inst. Komarova

1984, 21: 141-142, en russe.

88-203 ANDREEV М.Р. - Lichenes epilithici insulae ionae (mare Ochotense). Nov. Sist.

Niz. Rast , Akad. Nauk SSSR, Bot. Inst. Komarova 1984, 21: 143, en russe.

88-204 ASTA J. - Colonisation par les lichens et les mousses. In: CNRS-PIREN-MAB,

L'aménagement de la haute montagne et ses conséquences sur l'environnement. Le cantan d'Aime

(Savoie). 1985, pp. 266-276, fig. (USTM Grenoble, Lab. Bot. & Biol. Vég., ВР 68, F-38042 Si-

Martin-d'Heres Cedex).

Pendant trois ans, suivi d'une colonisation par Baeomyces rufus puis par Pogonatum

urnigerum d'un talus de bord de piste de ski en Savoie.

88-205 ASTA J. - Flore et végétation lichéniques de la pinède à crochets du Bois du

Sapey (Pellafol, Istre). Bull. Féd. Mycol. Dauphiné-Savoie 1986, 101: 9-12, fig. (Ibidem).

88-206 BLAKE G., ASTA J., GENSAC P. - Le lac noir, tourbière de moyenne altitude. In:

CNRS-PIREN-MAB, L'aménagement de la haute montagne et ses conséquences sur l'envrionnement.

Source : MNHN, Paris

184 BIBLIOGRAPHIE LICHENOLOGIQUE

l'envrionnement. Le canton d'Aime (Savoie). 1985, рр. 285-292, 1 tabl., 5 fig.

Descr. de la végétation, étude limnologique du site du Lac Noir (Savoie). Impact néfaste des

aménagements touristiques sur la stabilité de cette toubière. Lichens associés,

88-207 BEGUINOT J. - Glanures lichénologiques en Haute-Corrèze (secteur

Eymoutiers-Treignac-Bugeat-Gentioux). Bull. Soc. Bot. Centre-Ouest n.s., 1987, 18: 215-

216 (Le Bois Joli, 77 rue Docteur-Rebillard, F-71200 Le Creusot).

88-208 BREDKINA L.I. - Analysis arealogica lichenoflorae montium Tian-Schan

centralis, Nov. Sist. №. Rast., Akad. Nauka SSSR, Bot. Inst. Котағоуа 1984, 21: 144-149, en

russe.

88-209 HENDERSON A. - Lichens. Га: Meanwood valley survey Part УП. Leeds Naturalist’ Club

& Sci. Assoc. Newsletter 1985, 2: 27-42, 4 fig. (Dept. РІ. Sci., Univ. Leeds).

Données écol., liste des lichens avec hab. et loc. de Meanwood valley. Utilisation des

lichens comme bioindicateurs de pollution atmosphérique.

38-210 HYVÓNEN S. and HYVONEN J. - Contributions to the lichen and bryophyte

flora of Aletschwald Nature Reserve and its surroundings (Valais, Switzerland). Bull.

Murithienne 1985, 103: 127-168, 1 fig. (Bot. Mus., Univ. Helsinki, Unioninkatu 44, SF:00170

Helsinki).

Site, végétation, flores lichénique et bryophytique, de la réserve naturelle de l'Aletschwald

(Valais suisse).

88-211 KRIEGER Н. and TURK К. - Floristische und immission-ükologische

Untersuchungen an Rindenflechten im unteren Mühlvierterl, Oberósterreich. Linzer

Biol. Beitr. 1986, 18(2): 241-337, 174 fig., 3 tabl. (Am Gründberg 8, A-4283 Bad Zell).

Végétation et floristique des lichens corticoles épiphytes de la région de Mühlvierterl;

influence de la pollution. Distrib. selon le support. Cartes de répartition des 155 taxons étudiés.

88-212 LADD D. and WILHELM G. - New and noteworthy macrolichens from Missouri.

Trans. Missouri Acad. Sci. 1986, 20: 15-22 (The Morton Arboretum, Lisle, Illinois 60532, USA).

Liste de 20 macrolichens avec habitat dans le Missouri.

88-213 MAKAROVA LL, PERFILJEVA V.I. - Ad lichenofloram partis boreali-

occidentalis Jakutiae. Nov. Sist, №. Rast., Akad. Nauk SSSR, Bot. Inst. Komarova 1984, 21:

150-160, 1 fig., en russe.

88-214 STEFFENS A. - The occurence of epiphytes on Picea excelsa (CV) in the area

of Minster (Westfalen). Biblioth. Lichenol. 1987, 25: 77-80, 9 fig. (Bot. Inst. Univ.,

Schlossgarten 3, D-4400 Münster).

88-215 STENROOS S. - Studies on the genus Cladonia Sect. Cocciferae in Papua New

Guinea and the adjacent regions. Biblioth. Lichenol. 1987, 25: 421-422 (Dept. Bot., Univ.

Helsinki, Unioninkatu 44, SF-00170 Helsinki).

88-216 TITOV AN, - Lichenes caliciales in parte boreali-occidentali planitiei putoran.

Nov. Sist. Niz. Rast., Akad. Nauk SSSR, Bot. Inst. Komarova 1984, 21: 179-183, 3 fig., en russe.

88-217 VASANDER H. - Effect of forest amelioration on diversity in an ombrotrophic

bog. Ann. Bot. Fenn. 1984, 21(1): 7-15, 2 fig., 3 tabl. (Lammi Biol. Station, Univ. Helsinki, SF-

16900 Lammi).

La diversité (index Shannon, nombre d'Hill) totale décroit des parties vierges aux parties

drainées et fertilisées, reflétant une tendance générale dans les écosystèmes eutrophiques ou

desherbées. Bryophytes et lichens associés .

Source : MNHN, Paris

BIBLIOGRAPHIE LICHENOLOGIQUE 185

88-218 VITIKAINEN O. - Distribution patterns of European Peltigera. Biblioth. Lichenol.

1987, 25: 423-425, 4 fig. (Bot. Mus., Univ. Helsinki, Unioninkatu 44, SF-00170 Helsinki).

88-219 WITTMANN Н. und TURK К. - Beiträge zur Flechtenflora von Salzburg VI: Das

Rauriser Tal. Florist. Mitt. Salzburg 1986, 10: 47-68, 1 fig. (Inst. Bot., Univ. Salzburg, Lasserstr.

39, A-5020 Salzburg).

Liste de 440 lichens, champignons lichénisants et lichénicoles avec loc. du Rauriser Tal.

38220 WITTMANN Н. und TURK R. - Zur Flechtenflora Oberösterreichs. Neue und

Bemerkenswerte Flechten und Flechtenparasiten. Linzer Biol. Beitr. 1987, 19(2): 389-399

(Ibidem).

Liste de 31 lichens et champignons lichénicoles avec loc. en Haute-Autriche. 18 sont nouv.

pour la région. Pachyphiale ophiospora et Rinodina efflorescens sont nouv. pour l'Autriche.

Voir aussi: 88-148, 88-167, 88-170, 88-192, 88-224, 88-225, 88-226, 88-227, 88-228, 88-

230.

Pollution

88221 BELANDRIA С. et ASTA J. - Les lichens bioindicateurs: la pollution acide dans

la région lyonnaise.Pollut, Atmosphér. (Paris) 1986, 109: 10-23, 4 tabl., 4 fig. (USTM

Grenoble, Lab. Bot. & Biol. veg., ВР 68, F-38042 St-Martin-d'Heres Cedex).

‘Application de la méthode Hawksworth & Rose. Mise au point d'une échelle de sensibilité

des lichens de la région lyonnaise et cartographie de la pollution estimées (6 zones). Relations

pollution estimée et teneurs en 507 mesurées. Causes possibles des difficultés de recolonisation par

les lichens malgré la baisse sensible de la pollution atmosphérique.

Voir aussi: 88-209, 88-211, 88-228.

Docuemntation, Histoire des Sciences

88222 PEVELING Е. - Lichenology and lichenologists іп Westphalia. Biblioth. Lichenol.

1987, 25: 1-14 (Bot. Inst., Univ., D-4400 Münster).

Grandes lignes de la lichénologie, biographie (portrait et bibliogr.) des lichénologistes en

Westphalie: Johann бой. LAHM (1811-1888), Conrad BECKHAUS (1821-1890), Friedrich ZOPF

(1846-1909), Friedrich TOBLER (1879-1957).

88-223 TURK R. und WITTMANN Н. - Lichenologische Forschungen im Bundesland

Salzburg - ein historischer überblick. Jahrb. Haus der Natur Salzburg 1987, 10: 86-92, 6 fig.

(Univ. Salazburg, Inst. Pflanzenphysiol., Hellbrunner Strasse 34, A-5020 Salzburg).

Ouvrages généraux

88224 CLAUZADE G. kaj ROUX С. - Likenoj de Okcidenta Europo. Supplemento 2а.

Bull. Soc. Bot. Centre-Ouest n.s., 1987, 18: 177-214, en espéranto (rue des Pinsons, F-84300

Cavaillon)

Nouveautés lichénologiques depuis la parution de "Likenoj de Okcidenta Europo” en 1985.

Corrections des erreurs, lacunes et omissions.

88-225 HANSEN E.S., POELT J. and SOCHTING U. - Die Flechtengattung Caloplaca in

Grönland. Meddelelser от Grønland, Bioscience 1987, 25: 1-52, 19 fig. (Bot. Mus., Københavns

Univ., Gothersgade 130, DK-1123 København К).

Clé, morphol., chimie, distr., hab. et écol. des 43 esp (+ 2 incertaines) de Caloplaca

Source - MNHN, Paris

186 BIBLIOGRAPHIE LICHENOLOGIQUE

présentes au Groenland (récoltes de Poelt & Sgchting en 1982 et 1983 et herbiers de C et O).

Délimitation de certaines esp. Essai de classification du genre. Nouveautés pour la région. Diagn.

des taxons nouv. : Caloplaca psoricida, С. citrina var. soropelta, С. jemtlandica var. cerinosora el

C. lithophila var. elaeophora.

88226 MIDDELBORG J. and MATTSSON J. - Crustaceous lichenized species of the

Caliciales in Norway. Sommerfeltia 1987, 5: 1-71, 49 fig., 6 tabl.(Bot. Gard. & Mus., Univ.

Oslo, Trondheimsveien 23B, N-0562 Oslo 5).

Données morphol., anatom., chimiques, écol. et distr. de l'ordre des Caliciales dovisé en 3

fam.: Caliciaceae, Coniocybaceae, Microcaliciaceae. Délimitation des gneres Chaenotheca (Th. Fr.)

Th. Fr. et Sclerophora Chevall. Clé aux genres (Calicium, Chaenotheca, Cyphelium, Microcalicum,

Sclerophora, Thelomma ) et aux 39 esp. présentes en Norvège. Descr., chimie, hab, distr. et notes

pour chaque taxon. Noter Calicium adaequatum, C. adspersum, C. corynellum, C. parvum,

Chaenotheca carthusiae, Microcalicium ahleri nouv. pour la Norvège.

88227 MOBERG R. and HANSEN Е.5. - The lichen genus Physcia and allied genera in

Greenland. Meddelelser om Grønland, Bioscience 1986, 22: 1-32, 17 cartes, 19 fig. (The

Herbarium, Univ. Uppsala, P.O. Box 541, 5-751 21 Uppsala).

Clé, morphol., , chimie, distr., hab. et écol. de 7 Physcia, 6 Phaeophyscia, 4 Physconia

présents au Groenland. Noter Phaeophyscia kairamoi, P. nigricans, Physcia adscendens et

Physconia enteroxantha nowy. pour le Groenland.

88-228 — PEVELING E. - Progress and problems in lichenology іп the Eighties.

Proceedings of an International symposium held at the University of Münster on 16-

21 March 1986. Biblioth. Lichenol, 1987, 25: I-XV, 1-497, ill. (Ibidem).

50 contributions, liste des participants. Index matières et taxonomique

88229 VAILLE L. - Généralités sur les lichens et leur détermination (traduction de la

premiére partie (pp. 9-69) de la flore rédigée en Espéranto par G. Clauzade et C.

Roux: "Likenoj de Okcidenta Europo, ilustrita determinlibro"). Bull. Soc. Bot. Centre-

Quest n.s., 1987, 18: 147-176, 5 tabl, (Mesnil St Père, F-10140 VendeuvreS/ Barse).

88-230 WIRTH У. - Die Flechten Baden-Württembergs: Verbreitungsatlas. Stuttgart:

Verlag Eugen Ulmer, 1987. 528 p., 408 photos coul., 860 cartes (Staatliches Mus. Naturk. Stuttgart,

Rosenstein 1, D-7000 Stuttgart 1).

Les données écol. et répartition des lichens de la région du Bade-Wurttemberg, accumulées

depuis 20 ans, permettent de présenter un atlas des lichens de cette région. Pour chaque genre:

bréve descr., liste des esp., cartes de distr. de la plupart d'entre elles, photographies en coul. des

plus représentatives. Ces photos magnifiques sont de K. et H. Rasbach et de V. Wirth. Un glossaire

(3 p.), une partie taxonomique (incl. les comb. nouv. (est-ce la place dans une flore ?)), une

bibliographie (3p.) et un index des taxons (10 p.) complètent ce merveilleux atlas. - Esp. nouv.

Placopsis lambii Hertel et Wirth; comb. nouv.: Bacidia globulosa (Flórke) Hafellner & Wirth

(=Lecidea ), Carbonea assimilis (Koerb.) Haf. & Hertel (=Lecidella ), Fellhanera vezdae (Coppins

& James) (=Bacidia ), Lecidea lapicida var. lactea (Flórke ex Schaerer) (=Lecidea lact. ),

Loxospora cismonica (Beltram.) Haf. (=Haematomma ), Megaspora (Clauz. & Roux) Haf. & Wirth

(=Aspicilia subgen.), M. verrucosa (Ach.) (-Lecanora ), Mycobilimbia berengeriana (Massal.)

Haf. et Wirth (=Biatora ), M. fusca (Massal.) Haf. et Wirth (=Bilimbia), М. hypnorum (Libert)

Kalb & Haf. (=Lecidea ), M. lobulata (Sommerf.) Haf. (=Lecid. ), Protoparmelia atriseda (Fr.) К.

Sant. & Wirth (=Parmelia bada var.), Trapelia granulosa (Hoffm.) (=Verrucaria ), T. percrenata

(Nyl) (ELecidea ), T. wallrothii (Flórke) (=Lecidea).

Source : MNHN, Paris

187

INFORMATIONS

Ouvrages récemment reçus

BUCK W.R. - Bryostephane Steereana: a collection of bryological papers presented to

William Campbell Steere on the occasion of his 80th birthday. Memoirs of the New

York Botanical Garden 1987, 45.778 p., ill. (New York Botanical Garden, Bronx, New York 10458,

USA, US orders $ 136.05, non US 5 138.40, ISBN 0-89327-322-8).

DEWEY R.M. - Taxonomic and populational studies of the thallose liverworts Riccia

subgenus Riccia. Ph. D. Thesis, Texas A & M University, 1986, 141 p ill.

FLATBERG K.I. - Taxonomy, morphovariation, distribution and ecology of the

Sphagnum imbricatum complex with main reference to Norway. Gunneria 1986, 54: 1-

118, ill.

FREY W. - Moosflora und -vegetation in Regenwäldern NO-Perus. Ergebnisse der

BRYOTROP-Expedition nach Peru 1982. Beihefte zur Nova Hedwigia 1987, 88: 1-159, ill.

(J. Cramer, Berlin/Stuttgart, E. Schweizerbart'sche Verlagsbuchhandl., Johannesstr. 3A, D-7000

Stuttgart 1, DM 130.-, ISBN 3-443-51010-8).

KANDA H. - Catalog of moss specimens from Antarctica and adjacent regions

housed mainly in the herbarium of the National Institute of Polar Research. Tokyo:

National Institute of Polar Research, 1987. 186 p., ill. (9-10, Kaga 1-Chome, Habashi-ku, Tokyo,

MAGILL R.E. - Bryophyta. Part 1. Mosses. Fascicule 2. Gigaspermaceae-

Bartramiaceae. In: О.А. LEISTNER, Flora of Southern Africa. Pretoria: Botanical Research

Insitute, 1987. ix, 293-443 p., ill. (Dept. Agric. & Water Affairs, Private Bag X101, Pretoria, South

Africa, R 31.85 , ISBN O 621 10325 X).

MEDDELELSER ОМ GRONLAND, BIOSCIENCE -

22 - MOBERG К. and HANSEN Е.5. - The lichen genus Physcia and allied genera in

Greenland, 1987, 32 p., ill. (DKK 56,-

23 - MOGENSEN G.S. - Illustrated Moss Flora of Arctic North America and

Greenland. - Andreacobryaceae - Tetraphidaceae (by B.M. MURRAY). 1987. 36 p., ill.

(DKK 50,-).

25 - HANSEN E.S., POELT J. and SÓCHTING U. - Die Flechtengattung Caloplaca in

Grönland. 1987. 52 p., ill. (DKK 85,-).

(ISSN 0106-1054, Arnold Busck-Intern. Booksellers, Købmagergade 49, DK-1150 Copenhagen).

MIDDELBORG J. and MATTSSON J. - Crustaceous lichenized species of the Caliciales

in Norway. Sommerfeltia 1987, 5: 1-71, ill. (Bot. Gard. & Mus., Trondheimsveien 23 В, N-0562

Oslo 5, NOK 55.00, ISBN 82-7420-001-2).

POCS T., SIMON T., TIBA Z. , PODANI J. - Proceedings of the IAB conference of

Bryology. Budapest, Vacratot, Hungary, 5-10 August, 1985. Symposia Biologica

Hungarica 1987, 35, XIX, 902 p. en 2 volumes (Akademiai Kiado, Budapest, Hungary, ISBN 963

054633 7).

WERNER J. - Liste rouge des bryophytes du Grand-Duché de Luxembourg.Travaux

scientifiques du Musée d'Histoire Naturelle du Luxembourg 1987, 11. 42 p., ill (Marché aux

Source : MNHN, Paris

188

Poissons, L-2345 Luxembourg).

WIRTH V. - Die Flechten Baden-Württembergs: Verbreitngsatlas. Stuttgart: Verlag Eugen

Elmer, 1987. 528 p., ill. (Postfach 70 0561, D-7000 Stuttgart 70, DM 78,-; ISBN 3-8001-3305-9).

Commission paritaire 15-9-1981 - No $8611

Dépôt légal пе 13809 - Imprimerie de Montligeon

Sorti des presses le 25 avril 1988

Imprimé en France

Éditeur : A.D.A.C. (Association des Amis des Cryptogames)

Président : A. Couté; Secrétaire : D. Lamy

Trésorier : R. Baudouin; Directeur de la publication : H. Causse

Source : MNHN, Paris

SOMMAIRE

W. FREY and KURSCHNER - Pseudocrossidium replicatum (Tayl.) Zan-

der replaces Barbula acutata C. Müll. A note on its synonymy, distri-

bution and the xeropottioid life syndrome ...................

А. EDERRA INDURAIN — Briofitos del Pirineo de Navarra (España)

Е. FUERTES LASALA y E. MARTINEZ-CONDE - Vegetacion briofitica

del macizo oriental de los Picos de Europa (Andara) en Cantabria

(España). 1. Comunidades saxfcolas, acuáticas y subacuáticas .,.....

К.5. VISHVAKARMA and A, Kaul — Culture studies on Plagiochasma

appendiculatum Lehm. et Lindenb. and Reboulia hemisphaerica (L.)

Raddi populations of Pachmarhi Eor India) in relation to pH on

a comparative basis, .......... ТЕА ОЕ Я

Р.О. OVSTEDAL — Cladonia galindezii, a new antarctic lichen species .

А. APTROOT - Lichens of Madagascar : the Pyxinaceae а.

crac e ۳ ы a КӨНЕК, АРЕН РЕҢІ os

К.С. PATIDAR, C.M. SOLANKI and А. KAUL - Culture studies on Ric-

cia gangetica Ahmad V - Influence of mineral nutrients on growth . . . .

E. CHAUHAN — Ultracytochemical localization of acid and alkaline

phosphatases in the placental region of the moss Physcomitrium cyathi-

ОЕ : х

Bibliographie bryologique .. .

Bibliographie lichénologique . .

Informations ............. [ede o O OUT

Cryptogamie, Bryol. Lichénol. 1988, 9 (2) : 95-188.

Source

103

109

129

137

141

149

155

173

182

187