CRYPTOGAMIE

Bryologie-Lichénologie

ANCIENNE REVUE BRYOLOGIQUE ET LICHÉNOLOGIQUE

Fondéc par T. Husnot en 1874.

Directeur de la publication : Dr Héléne Bischler-Causse

Rédaction :

Bryophytes : Dr Hélène Bischler & M. Denis Lamy, Laboratoire de Cryptogamie, 12 rue

Buffon, F-75005 Paris. Tel. 01.40.79.31.84 — Fax : 01.40.79.35-94 Email : lamy

@ mnhn.fr

Lichens: Dr Chantal Van Haluwyn, Laboratoire de Botanique et de Cryptogamie, Faculté

de Pharmacie, B.P. 83, F-59006 Lille Cedex. Tél. 0320964040 —

Fax : 03.20.95.90.09

Editeur : A.D.A.C. — 12 rue Buffon F-75005 Paris

COMITÉ DE LECTURE

Bryologie : J. Berthier (Clermont-Ferrand), B. Bodo (Paris), R.B. Buck (Bronx), J.L. De Sloover

(Namur), P. Geissler (Genève), S.R. Gradstein (Utrecht), J.P. Hébrard (Marseille), 5. Jovet-Ast

(Paris), A. Lecointe (Caen), M.C. Noailles (Paris), R. Ochyra (Kraków), C. Suire (Bordeaux),

B.C. Tan (Cambridge, USA).

Lichénologie : J. Asta (Grenoble), A. Bellemére (Paris), T. Bernard (Rennes), B. Bodo (Paris), Ј.С.

Boissière (Paris), P. Clerc (Genève), W.L. Culberson (Durham), S. Déruelle (Paris), P. Diederich

(Luxembourg), M.C. Janex-Favre (Paris), J. Lambinon (Liége), M.A. Letrouit-Galinou (Paris), X.

Llimona (Barcelone), L. Nimis (Trieste), G. Rambold (Munich), Cl. Roux (Marseille), M.R.D.

Seaward (Bradford), A. Sérusiaux (Liège), Н. Sipman (Berlin), E. Stocker-Wörgötter (Salzburg),

V. Wirth (Stuttgart).

MANUSCRITS

Les manuscrits doivent étre adressés (en 3 exemplaires) à la Rédaction de Cryptogamie,

Bryologie-Lichénologie. L'avis du Comité de Lecture sera requis avant accord pour publication.

Bien que Cryptogamie soit une revue de langue française, les manuscrits rédigés en français, en

anglais, en allemand, en espagnol et en italien, sont pris en considération dans la limite des

possibilités de la Rédaction à trouver des lecteurs compétents dans ces langues. Les disquettes de

micro-ordinateurs (IBM, IBM compatible et MacIntosh) sont vivement souhaitées. Les instruc-

tions aux auteurs sont publiées dans le fascicule 1 de chaque tome. Les auteurs recevront

25 tirés-à-part gratuits ; les exemplaires supplémentaires seront à leur charge.

TARIFS DES ABONNEMENTS Tome 18, 1997

CRYPTOGAMIE comprend trois sections : Algologie, Bryologie-Lichénologie, Mycologie.

Pour une section : France : (350 F ht) 357,35 F ttc Étranger : 380,00 Е

Pour les 3 sections : (950 F ht) 970,00 F ис Étranger: 1050,00 F

VAT Registration number : FR29302655170

Paiement par chéque bancaire ou postal à l'ordre de : A.D.A.C. — CRYPTOGAMIE (CCP La

Source 34 764 05 S) ; adressé à : A.D.A.C. , 12 rue Buffon, F-75005 Paris. :

Cryptogamie, Bryologie-Lichénologie est indexé par Biological Abstracts, Current Contents,

Index of Fungi et Bibliography of Systematic Mycology, Publications bibliographiques du CNRS

(Pascal).

Couverture : Pore de Marchantia foliacea Mitt. (cliché Н. Bischler)

Source : MNHN, Paris

CRYPTOGAMIE

BRYOLOGIE LICHENOLOGIE

TOME 18 FASCICULE 3 1997

CONTENTS

БООН + б Толе SR MER ER he esp cre td, de e e жи TI

Claritza E. GRADSTEIN-SERNA — Atribute from Colombia to Helen Bischler

in Colombian) ps: tac di lr

David С. LONG — Studies on the genus Asterella P. Beauv. II: Asterella cruciata

(Steph.) Horik. in Eastern Asia аена cine retener

S. Rob GRADSTEIN and Patricia GEISSLER — Not

jeunea (Hepaticae)

Suzanne JOVET-AST — A proliferous Riccia from Australia: Riccia pullulans

sp. nov. (Subgenus Ricciella) (in French)

Riclef GROLLE and M. L. SO — Plagiochila bischleriana, a new species from

лт c HM E ce EE EC с + E]

Tamas PÓCS — New or little known epiphyllous Liverworts, УП. Two new

Lejeuneaceae species from the Mascarene Islands.....................

Benito C. TAN and Jian-Cheng ZHAO — New Moss records and range extensions

of some xeric and alpine moss species in China .......................

Cecilia SERGIO and Ana SÉNECA — The first report of Cryptothallus mirabilis

Malmborg (Hepaticae, Aneuraceae) in Southern Europe (Portugal) . . . .

S. Rob GRADSTEIN — Bromeliophila helenae, a new species of Lejeuneaceae

Tromtüe NEOUODIEH POIL рата V aes ANA RT ee

Barbara M.THIERS — Lejeunea bischlerae, a new species of Lejeunea subgenus

Microlejeunea from Australia ia lisses

William R. BUCK and Bruce ALLEN — Ordinal placement of the Fontinalaceae.

Jan-Peter FRAHM — Which function have the hyalocysts of the leaves of Dicra-

пасове (lpGephan) dem ire er este ee

167

169

177

183

191

195

207

213

217

223

227

Bibliothèque Centrale Muséum

3001 00037812 2

Source : MNHN. Paris

IM490T4j

ЭСТОН Я

амура de ЧМР

EDITORIAL

Ce fascicule a été réalisé à l'occasion du départ à la retraite d'Hélène Bischler-Causse. Les

contributions correspondent à une partie de son activité scientifique: systématique des

bryophytes, bryophytes des régions tropicales.

Qu'Héléne considére ce fascicule comme le témoignage de notre profonde et sincére amitié

Denis Lamy

Marie-Catherine Boisselier-Dubayle

This issue has been done in honour of Héléne Bischler-Causse, just retired. The contribu-

tions represent some aspects of her scientific activity: bryophyte systematics, bryophytes

from the tropical area.

This issue should be considered as a mark of our deep and heartfelt friendship.

Denis Lamy

Marie-Catherine Boisselier-Dubayle

Source : MNHN. Paris

= panne à

AREA ате pd. sig ^

Cryptogamie, Bryol. Lichénol. 1997, 18 (3): 167-168 167

UN TRIBUTO DE COLOMBIA A HELENA BISCHLER

Claritza E. GRADSTEIN-SERNA

Que la criptogamia colombiana esté ligada a nombres como Humboldt, Lindig,

Bischler, Cleef, entre otros, es muy cierto. Lo que no se puede asegurar es que muchas de

las especies por ellos colectadas hace unos años, aún existan hoy, especialmente en

aquellas regiones exploradas por Helena Bischler.

A los 24 años llegó a Colombia con un morral, un español impecable y una vida

por vivir. Para entonces ya se había refugiado en la ciencia. Fuerte, de caminar erguido y

de mirada profunda, Helena Bischler se dió tiempo para vivir lo que ella llama “mis años

en Colombia”. Helena como Humboldt y con el mismo propósito de aportar su grano de

arena a la ciencia, recorrió el territorio colombiano desde Antioquia hasta Vaupés

(incluyendo las hoy intendencias del Casanare y Guaviare), en tiempos en que el país

atravesaba por una seria crisis política, social y económica debido a las tensas relaciones

entre los dos partidos tradicionales, el gobierno y la sociedad civil, situación ésta que

reinaba tanto en la vida urbana como rural.

Corría el año 1956 cuando Helena y un grupo de destacados investigadores

colombianos, entre ellos Jesús Idrobo y Polidoro Pinto, emprendieron cordillera arriba su

misión. Unas veces en mula, otras en canoa y muchas otras a pie, atravesando ríos,

montañas y sobreviviendo a la inclemencia del tiempo y los mosquitos, en pleno río

Guayabero en la intendencia del Meta, por nombrar solo un lugar; acamparon, desvelaron

y colectaron en sectores en los que hoy nadie se atreve a penetrar, no por lo espeso de la

selva, como en aquel entonces, sino por que hoy día son regiones ligadas a las palabras que

tan mal nombre le han dado a Colombia: Violencia, guerrilla, coca, subversión y otros

calificativos que nada tienen que ver con investigación y ciencia.

Pues a estas zonas antaño aisladas, despobladas y “pobres” donde predominaba

como actividad de subsistencia la caza, la pesca, el maíz, el plátano, como en el caso del

Casanare (en esa época parte del departamento de Boyacá y considerada ahora como la

región petrolera de Colombia), la “bonanza de la coca” trajo por la generalización de su

cultivo sustanciales cambios en la región y de hecho importantes transformaciones

socio-culturales en la población. La producción, elaboración y comercialización de la hoja

atrajo a muchos colonos, comerciantes, vendedores y toda clase de aventureros que venían

con la esperanza de entrar en una empresa incierta pero que los “sacara de la pobreza”.

Esta nueva población y los mismos colonos ya no ofrecen sus cerdos, las gallinas,

el ganado, las pieles; la gente en éstas tierras ya no sabe de plátanos, yucas, arroz, de perros

de cacería, hasta el transporte en canoa y a caballo fue cambiado por el de avionetas y

helicópteros. Los campesinos que antes recibían al extranjero con una sonrisa amable y

una taza de guarapo (bebida fermentada de la caña de azúcar) son los mismos que hoy con

un arma en la mano, desconfiados, silenciosos y dispuestos a lo que sea, no admiten al

visitante,

Source : MNHN. Paris

168 C. E. GRADSTEIN-SERNA

Pero todo este aparente “florecimiento” no persistió pues actualmente es una de

las regiones mas conflictivas del país, (como muchas otras afectadas por el fenómeno de la

coca) a donde ya nadie se atreve a acercarse ni siquiera en beneficio de la cienci

La inefectiva intervención estatal y de algunos institutos sólo ha dejado una

cantidad considerable de papel, tratos, negociaciones, convenios, acuerdos y comités que

han actuado no como instrumento de ayuda y progreso sino como medio de represión y

discordia en la población.

A Helena le debemos gran parte de la colección de criptógamas existentes en el

Herbario Nacional Colombiano. Helechos, líquenes, briofitas de muchas regiones del

país: Antioquia, Boyacá, Cundinamarca, Huila, Meta, Quindío, Risaralda, Santanderes,

Tolima, Valle y Vaupés. Este material recobra su importancia y sigue siendo objeto de

estudio aunque muchas de estas especies ya no existen pues factores como la tala de los

bosques, el cultivo de la coca y la amapola, los bombardeos accidentales e intencionados

а los oleoductos, las “barridas” que realiza el gobierno y todas las acciones por sanear la

región, se han encargado de que la fauna y la flora hayan ido desapareciendo.

Es decir que en 40 años el país no sólo ha cambiado en cuanto a su división

política; también han cambiado sus gentes, sus recursos, su economía y hasta sus costum-

bres. Faltaria preguntarle a Helena que opina de la Colombia de hoy, será que todo tiempo

pasado fue mejor? Pues sin duda y aunque 40 años no son mucho y la tierra todavía este

allí, no sería fácil reconocer y encontrar las mismas especies.

Quizá lo único que queda hoy es el recuerdo y el regalo de un par de científicos

y las colecciones en los herbarios que afortunadamente dejan prueba que allí existió un

paisaje sano, de gentes buenas que jamás imaginaron que una semillita alteraría tanto sus

vidas.

Estoy plenamente segura que ni con el boom de la conservación y explotación

racional de los recursos naturales y todos los tratados habidos y por haber, podrán

salvarse ya muchas especies.

Source : MNHN. Paris

Cryptogamie, Bryol. Lichénol. 1997, 18 (3): 169-176 169

STUDIES ON THE GENUS ASTERELLA P. BEAUV. III*:

ASTERELLA CRUCIATA (STEPH.) HORIK. IN EASTERN ASIA.

David G. LONG

Royal Botanic Garden, Edinburgh EH3 SLR, Scotland, U.K.

email: d.long@rbge.org.uk — fax: (0044) 131 552 0382

ABSTRACT — The neglected name Fimbriaria cruciata Steph. is lectotypified and its status clarified

as Asterella cruciata (Steph.) Horik. A. odora S.Hatt., A. chichibuensis Shimizu & S.Hatt. and A.

mitsuminensis Shimizu & S. Hatt. are reported as new synonyms. Evidence from gametophyte

morphology and spore ornamentation is used to support this synonymy. The species is a very rare and

highly localised species, newly reported from China (Yunnan and Sichuan) and elsewhere restricted to

Japan (Honshu) and Korea.

Key моко: bryophyte, liverwort, Asterella, SEM, spores, synonymy, typification, Japan, Korea,

China.

INTRODUCTION

Prior to Volumes 1 and 6 of Stephani’s Species Hepaticarum (Stephani 1898-

1900, 1917-1925), no Asterella (syn. Fimbriaria) was reported from Japan. Stephani

reported two species from Japan, F. pilosa Taylor collected by Faurie and the new species

F. cruciata Steph. based on a collection by Uematsu. Localities or other collection details

were not given by Stephani. Until the revision of Japanese Asterella by Shimizu & Hattori

(1952, 1953a), the genus remained neglected in Japan, except for the description of А.

odora by Hattori (1944).

Shimizu & Hattori (1952, 1953a) adopted a rather narrow species concept within

Asterella which resulted in the description of five new Japanese species. One of these, A.

mitsuminensis, was soon synonymised by them in the second part of their revision, under

A. odora S.Hatt. They only considered Japanese material and did not compare taxa from

other parts of E Asia. A. cruciata (Steph.) Horik. was treated only briefly in their revision

as they were unable to locate any material collected by the cited collector, Uematsu, and

the species remained as a doubtful one. They did, however, refer to two specimens in G

determined as F cruciata by Stephani but collected by Sakurai. One of these they

redetermined as Reboulia hemisphaerica (L.) Raddi and the other as Asterella odora

S.Hatt.

* paper П in this series was published in Journal of Bryology (1994) 18: 287-295.

Source : MNHN, Paris

170 D.G. LONG

In their key to Japanese species, A. cruciata was placed next to A. crassa Shimizu

& S.Hatt. on account of its “stalk of female receptacle on a short obovate ventral branch".

This diagnostic character was taken from the unpublished drawing of F. cruciata made by

Stephani in Geneva, now published as a microfiche (Stephani, 1985). However, this

appears to be a misinterpretation of Stephani's drawing which in fact does not show the

branch bearing the archegoniophore as ventral but as a short terminal branch. This

interpretation agrees with Stephani's description ‘Pedunculis....in ramulo exiguo lateralis

terminalis’ which translates as “peduncle terminal on a weak lateral branch', with no

reference to any ventral insertion. Hence the whole question of the identity and rela-

tionships of Е cruciata deserves re-evaluation.

This study of A. cruciata is part of an on-going revision of Eurasian taxa of

Asterella which has taken a somewhat broader view of species limits as well as attempting

to study all taxa in the field and using SEM of spores and comparing Japanese taxa with

related species from Europe and other regions of Asia. The present study of A. cruciata

aims to (a) typify Fimbriaria cruciata and characterise the taxon, and (b) to establish its

synonymy and status in Asia,

MATERIALS AND METHODS

The present study is based largely on herbarium material, incl uding relevant type

specimens, from a range of herbaria (see acknowledgments). Observations on these

specimens were supplemented by a field trip to Honshu, Japan in 1993 on which all the

taxa known from Honshu were observed and collected.

Preparation of spores for SEM studies follows Taylor et al. (1974), using

herbarium material. A few spores were removed from a ripe capsule using a fine moistened

brush and attached to an aluminium SEM stub using double-sided adhesive tape or a

carbon disc. Stubs were coated with gold/palladium in a sputter-coater prior to study

using both a JEOL JSM T200 Scanning Electron Microscope and a Zeiss DSM 962

Digital Scanning Electron Microscope. The scanning voltage was normally 5kV or 10kV.

Images were photographed on Ilford Technical Pan Film.

THE IDENTITY OF FIMBRIARIA CRUCIATA

For all other Fimbriaria species described by Stephani, ori iginal material has

been located in his herbarium in Geneva. Despite careful searching, no material labelled Е.

cruciata collected by Uematsu could be located in G or elsewhere, although the Sakurai

material mentioned above has survived. A likely explanation is that Stephani’s (1917)

citation of Uematsu as collector is erroneous, as in most instances his specimen citations

were very scanty and not infrequently inaccurate.

The Sakurai specimen identified by Shimizu and Hattori as A. odora S.Hatt. was

annotated by Stephani as ‘F. cruciata St. sp. nov.’ It was collected nine years before he

described Fimbriaria cruciata and he probably had it in his possession when describing the

species in 1917. It should therefore be considered as part of the original material, and thus

Source : MNHN, Paris

ASTERELLA CRUCIATA IN EASTERN ASIA m

eligible as a lectotype (Art. 9). The other Sakurai specimen, identified as Reboulia

hemisphaerica (L.) Raddi (Shimizu & Hattori, 1953a), can be excluded as a possible

lectotype as it clearly conflicts with the original description in lacking a pseudoperianth,

although part of the protologue may be based on this specimen (see below).

It is concluded that Stephani cited the collector erroneously in Species Hepati-

carum. The annotation on his illustration of Е cruciata (Stephani, 1985) offers no

additional clues as to the collector as it merely cites ‘Japan’; however in several important

respects it agrees with his description and the Sakurai specimen. This illustration and

description of the thallus and ventral scale strongly resemble those of Asterella Subg.

Brachyblepharis (Grolle 1976, 1989), particularly the broad thin thalli 7-8mm broad and

0.75mm thick with attenuate wings and the single broad appendage of the ventral scales.

In fact, this type of scale appendage is diagnostic for only one member of

Japanese Marchantiales: the plant described under the names Asterella odora by Hattori

(1944) and A. chichibuensis by Shimizu & Hattori (1952). No other genus or species has

identical scale appendages; in Marchantia, Conocephalum and Lunularia the appendages

can be rounded but are auriculate at their insertion; those of Wiesnerella denudata (Mitt.)

Steph. are quite similar but are crenately toothed (Inoue, 1976). In all other Japanese

Marchantiales the appendages are acute and often more than one per scale (Inoue, 1976;

Mizutani & Hattori, 1969; Shimizu & Hattori, 1953b, 1953c, 1954),

Character Lectotype specimen | Stephani Description | Stephani Illustration |

Thallus t.s. Broad, thin Flat, thin, 7-8 x|Broad, thin

0.75 mm

No. ventral scale |1 1 1

appendages b. 5 ا‎

[Shape ventral scale | Broadly ovate, obtuse | Ovate-elliptic or subro- | Sub-rotund, obtuse

appendage wt tund

Position of archego- Terminal on main | Terminal on weak late- | Terminal on short ter-

niophore _ branch | ral branch minal braneh

Stalk length 1.5-3 mm |10 mm 9 mm

Carpocephalum shape | Umbrella-shaped |Subhemispheric or | Bluntly conical

infundibuliform

Spore reticulations Crenately toothed Crenate-dentate Not illustrate

Tabl. 1. — Diagnostic characters of the lectotype of Fimbriaria cruciata compared with Stephani's

description and illustration

The diagnostic characters given in Stephani's description and illustration are

compared with those of the Sakurai specimen (proposed below as lectotype) in Table 1.

The only serious discrepancies are in the archegoniophore length and receptacle shape. It

is possible that Stephani's description of these is partly based on the Reboulia specimen

which he also identified as Fimbriaria cruciata. However, as discussed above, his descrip-

tion of the thallus could only be based on an Asterella not Reboulia.

Apart from Subg. Brachyblepharis, only two other Asterella species are known

fom Japan (excluding Ryukyu Islands), A. crassa Shimizu & S.Hatt. from Honshu and 4.

gracilis (F.Weber) Underw. from Honshu and Hokkaido; these are both ruled out as the

possible identity of F. cruciata by several characters, such as their thicker more xeromor-

Source : MNHN. Paris

172 D.G. LONG

phic thalli, their lanceolate acute ventral scale appendages and non crenate-dentate spore

reticulations. Within Subg. Brachyblepharis, the only species known to occur in Honshu at

the time of Species Hepaticarum was F. cruciata. The other Japanese members of Subg.

Brachyblepharis (described by Shimizu & Hattori as A. pusilla, A. sanoana, and A.

umbelliformis), were not collected on Honshu before 1951 and no Japanese collections of

these taxa exist in Stephani's herbarium in G. The present species, on the other hand, was

collected as early as 1908 by Sakurai, and is the most commonly-collected species of

Asterella in Honshu.

In spite of the discrepancies mentioned above, most of the available evidence

points to the same conclusion, that the Sakurai material represents one of the original

gatherings of the plant Stephani intended to describe as Fimbriaria cruciata. It is accor-

dingly lectotypified below.

SYNONYMY AND TYPIFICATION OF ASTERELLA CRUCIATA

Asterella cruciata (Steph.) Horik., Hikobia 1: 79, 1951.

Basionym: Fimbriaria cruciata Steph., Species Hepaticarum 6: 12, 1917. Type citation:

“Japan, leg. Uematsu.’ Type specimen: ‘Fimbriaria cruciata St. n.sp. Japonia,

Tokyo, 28 ii 1908, Sakurai 9, (G-15200, lectotype selected here).

Synonyms: Asterella odora S.Hatt., Botanical Magazine ( Tokyo) 58: 44, 1944. Type

citation: "Tokyo: Koisikawa-ku, Kamihuzimae-tyo (S. Hattori, no. 2505, 21. Oct.

1938; по. 3124-typus, 23. Dec. 1939, cum spor.).’ Type specimen: Japan, Honshu,

Tokyo, Koishigawa, Kamifujimaye, on soil, 23 xii 1939, S. Hattori 3124 (NICH-

15906, holotype); syn. nov.

A. chichibuensis Shimizu & S.Hatt., Journal of the Hattori Botanical Laboratory 8: 46,

1952. Type citation: [Japan, Honshu] 'Saitama Prefecture: Chichibu, Hashitate,

ca. 350 m. alt., on rocks and on soil in the calcareous region, Oct. 25, 1950, Coll.

D. Shimizu, no. 18650-Typus! in Herb. Hattori Bot. Lab.’. Type specimen: Japan,

Honshu, Saitama Prefecture, Hashitate, ca. 350m alt., on rocks, 25 x 1950, D.

Shimizu (NICH-18650, holotype); syn. nov.

A. mitsuminensis Shimizu & S.Hatt., Journal. of the Hattori Botanical Laboratory 8: 48,

1952. Type citation: “Japan [Honshu]. Saitama Prefecture: Chichibu, Mt. Mitsu-

mine, ca. 600m. alt., on rocks, Nov. 25, 1951, Coll. D. Shimizu, no. 19642-Typus!

in Herb. Hattori Bot. Lab’. Type specimen: Japan, Honshu, Saitama Pref.,

Chichibu Mts., Mt. Mitsumine, ca. 550 m alt., on rock, 25 xi 1951, coll. D.

Shimizu (NICH-19642, holotype); synonymised with A. odora by Shimizu &

Hattori (1953a).

Following Shimizu & Hattori's identification of the Sakurai material as A.

odora, the above lectotypification of F. cruciata means that Asterella odora S.Hatt.

becomes a synonym of А. cruciata. Two further taxa are also considered to be synonyms

of A. cruciata:A. chichibuensis Shimizu & S.Hatt. and A. mitsuminensis Shimizu & S. Hatt.

In describing A. chichibuensis and A. mitsuminensis, Shimizu & Hattori (1952)

distinguished them primarily on the number of cell rings in the epidermal pores (3-4 and

Source - MNHN. Paris

ASTERELLA CRUCIATA IN EASTERN ASIA 173

2 rings respectively); the scaliness of the archegoniophore stalk (scaly and non-scaly) and

shape of androecia (large, elongate and small, ovate). All these are characters which vary

even within single populations of A. cruciata. A. mitsuminensis was quickly synonymized

under A. odora by Shimizu & Hattori (1953a). The great similarity of A. chichibuensis to A.

odora was already noted by Inoue (1976). The proposed synonymy is supported by SEM

study of spores of the type specimens of the three taxa (Fig. 1), and by the par-autoicous

sexual condition, with the androecia located close to the base of the archegoniophore on

the main thallus branches.

A. cruciata has been described and illustrated in several works: Shimizu &

Hattori (1952) as A. chichibuensis and A. mitsuminensis; Hattori (1944); Shimizu & Hattori

(1953a); Miyoshi (1966) and by Inoue (1976) as A. odora. Because of its broad thin delicate

thalli, ventral scales with a single appendage, relatively flattened carpocephalum and

yellow reticulate spores with dissimilar sculpturing on distal and proximal surfaces, it can

be confused only with other members of Subg. Brachyblepharis (Grolle 1976, 1989).

Within Subg. Brachyblepharis the par-autoicous sexual condition considerably

narrows the field of taxa related to A. cruciata: in east Asia the only related species is A.

khasyana (Griff.) Pande et al., in Europe A. africana (Mont.) A.Evans, in Africa A.

dissoluta (Steph.) Grolle, in America A. chilensis (Nees & Mont.) A.Evans and A. venosa

(Lehm. & Lindenb.) A.Evans and in Australasia A. tenera (Mitt.) R.M.Schust. Of these

only A. africana and A. dissoluta have a deeply-lobed carpocephalum; all have more

lanceolate acute ventral scale appendages, all have much longer archegoniophores and

none has the highly crenate-dentate spore areolations of A. cruciata.

DISTRIBUTION AND ECOLOGY

Asterella cruciata is a highly localised species. In Japan it is restricted to the

Chichibu Mountains of Saitama Prefecture to the west of Tokyo (Shimizu & Hattori 1952,

1953a). Its distribution in Japan was mapped by Inoue (1976). Within this restricted area,

however, it is locally abundant in suitable habitats.

It has not previously been reported from China, where it is known from only two

collections, one from Yunnanfu [=Kunming], Yunnan collected by Handel-Mazzetti in

1915 and the other from Chungking, Sichuan collected by P.C. Chen in 1940. Hence its

present status in China requires clarification. The Yunnan collection was reported by

Herzog (1930) as ‘Fimbriaria spec. zollingeri Steph. et viridis Steph. affinis’ but correctly

identified in 1977 by Grolle as A. odora. The Sichuan specimen was misidentified as

Sauteria alpina (Nees) Nees.

The status in Korea is equally uncertain. It has been reported from Mt. Gyerong

by Choe (1975) as Asterella chichibuensis and the specimen in JE may be from this locality.

Further information is much needed.

In the Chichibu Mountains of Honshu 4. cruciata is restricted to calcareous

metamorphic substrates, where it grows in the broad-leaved forest zone, on damp shady

rocks and banks, often in stream valleys, sometimes on terrace walls of cultivated slopes.

Recorded altitudes are from 350 to 900m. In Sichuan the species is recorded from clay soil,

and in Yunnan from a clay wall at 1920m. Like other members of Subg. Brachyblepharis,

A. cruciata shows no xeromorphic adaptations and is restricted to moist shaded habi-

tats.

Source : MNHN, Paris

ASTERELLA CRUCIATA IN EASTERN ASIA 175

SPECIMENS STUDIED (EXCLUDING TYPES)

JAPAN, HONSHU: Japonia, No. 1562 as Asterella minuta Lindb. ex herb

Lindberg (BM, S); Prov. Tokyo, 21 xi 1909, H. Sakurai s.n. (YE); Tokyo Prefecture, foot of

Mt. Tenso, along R. Akigawa, alt. 600-700m, deciduous forest, on moist shaded limestone

rock, 10 x 1979, S. R. Gradstein & H. Inoue 3094 (U); Saitama Pref., Chichibu, Kagemori,

25 xi 1950, D. Shimizu NICH 58650 (JE); Saitama County, Ochiai of Otaki, Chichibu

Mts., 480m, on rock crevices and on soil in shade, 31 viii 1952, Shimizu in Hep. Jap. Exsicc.

Ser. 6, 252 (B, BM, G, H, JE, L, LD, NY, S, TENN, UBC, W); Saitama Pref., Chichibu

Mts., 600m, 2 xi 1952, Hikuma NICH-64335 (JE); ibid., 570m, 2 xi 1952, Hikuma NICH

60725 (JE), 60726 (JE); Saitama Pref., Chichibu Mts., 570m, 1952, Hikuma NICH 60725

(JE); Saitama Pref., Chichibu Mts., Mt. Mitsumine, 900m alt., on wet rock in shady places

with Conocephalum conicum, x 1959, Inoue s.n. (S); Saitama Pref., Chichibu Mts., Mt.

Mitsumine ca. 800m, Inoue 8523 (TENN); Saitama Pref., Chichibu-gun, Hinata-ohya

below Mt. Ryokami, 705m, terraced cultivated slope, on base of wall, 5 ix 1993, Long & Т.

Furuki 24716 (E); Saitama Pref., Chichibu-gun, Ohwa, Arakawa River below Mt. Mitsu-

mine, 375m, forested rocky river bank, on damp mossy rocks by road, 6 ix 1993, Long &

Furuki 24789 (E).

CHINA, SICHUAN PROV: Hwa-lung-chiao prope Chungking, in terra

argillosa, skiophila, v 1940, PC. Chen s.n. (JE-H3346); YUNNAN PROV.: in urbe

Yünnanfu, muris, 1920m, 22 xii 1915, Н. Handel- Mazzetti 8603 (E, JE, W, WU).

KOREA: Korea, 13 xii 1978, S. Hattori (JE-H2444).

ACKNOWLEDGMENTS — The author is grateful to the Curators of Bryophytes at B, BM, H, G,

JE, L, LD, NY, S, TENN, U, UBC, W and WU, for loan of specimens. T. Furuki, Chiba, Japan is

thanked for arranging field work in Honshu, Riclef Grolle, Jena and Helene Bischler-Causse, Paris

are thanked for generous help with the Asterella research project. Sally Rae, Edinburgh, is thanked for

technical assistance with SEM work.

REFERENCES

CHOE D.M., 1975 — Floral Studies on the Hepaticae of Korea, Report of the Science and Technology

Ministry of Korea 75-70: 1-25.

GROLLE R., 1976 — Verzeichnis der Lebermoose Europas und benachbarter Gebiete. Feddes

Repertorium specierum novarum regni vegetabilis 87: 171-279.

ОВОШЕ R., 1989 — Über Asterella subg. Brachyblepharis in Lateinamerika. Wissenschaftliche

Zeitschrift der Friedrich-Schiller-Universitát Jenal Thüringen. Mathematischnaturwiss-

enschaftliche Reihe 38: 231-239.

Fig. 1. — Spores of Asterella cruciata and its synonyms. a,b, c, f, g, lateral view; d, distal view; e,

proximal view; h, detail of distal view. a — Fimbriaria cruciata, lectotype (С); b — A. odora, holotype

(NICH); е — A. chichibuensis, holotype (NICH); d-e — A. mitsuminensis, holotype (NICH); f — A.

odora, Japan, Hikuma NICH-60725 (JE); а — A. odora, Yunnan, Handel- Маггені 8603 (WU); h —

A. odora, Japan, Long & Furuki 24716 (E). Scale: a-f, bar = 50 um; g, bar = 20 jim; h, bar = 5 шп.

Source : MNHN, Paris

176 D.G. LONG

HATTORIS. 1944 — Hepaticarum species novae et minus cognitae nipponenses III. The Botanical

Magazine, Tokyo 58: 38-46.

HERZOG T., 1930 — Marchantiales. In: Handel-Mazzetti, H. Symbolae Sinicae 5: 1-6.

INOUE H., 1976 — Illustrations of Japanese Hepaticae. Vol. 2. Tokyo: Tsukiji Shokan. 193 p.

MIYOSHI N., 1966 — Spore morphology of Hepaticae in Japan. Bulletin of the Okayama College of

Science 2: 1-46.

MIZUTANI M. & HATTORI S., 1969 — Check list of Japanese Hepaticae and Anthocerotae.

Miscellanea Bryologica et Lichenologica 5: 33-43.

SHIMIZU D. & HATTORIS., 1952 — Studies on the Japanese species of Asterella (1). Journal of the

Hattori Botanical Laboratory 8: 46-54.

SHIMIZU D. & HATTORI S., 1953a — Studies on the Japanese species of Asterella (2). Journal of

the Hattori Botanical Laboratory 9: 25-31.

SHIMIZU D. & HATTORI S., 1953b — Marchantiales of Japan, I. Journal of the Hattori Botanical

Laboratory 9: 32-44.

SHIMIZU D. & HATTORI S., 19530 —Marchantiales of Japan, (2). Journal of the Hattori Botanical

Laboratory 10: 49-55.

SHIMIZU D. & HATTORI S., 1954— Marchantiales of Japan. Ш. Journal of the Hattori Botanical

Laboratory 12: 53-75.

STEPHANI E, 1898-1900 — Species Hepaticarum. Vol. 1. Genàve, Bále & Lyon: Herbier Boissier.

STEPHANI E, 1917-1925 — Species Hepaticarum. Vol. 6. Genéve, Bále & Lyon: Herbier Boissier,.

STEPHANI Е, 1985 — Icones Hepaticarum. 165 microfiches. Zug, I.D.C.

TAYLOR J., HOLLINGSWORTH PJ. & BIGELOW W.C., 1974 — Scanning electron microscopy

of liverwort spores and elaters. The Bryologist 77: 281-327.

Source : MNHN. Paris

Cryptogamie, Bryol. Lichénol. 1997, 18 (3): 177-182 177

NOTES ON THE GENUS LEUCOLEJEUNEA (HEPATICAE)

S. Rob GRADSTEIN! & Patricia GEISSLER?

! Systematisch-Geobotanisches Institut der Universitát Góttingen,

Untere Karspüle 2, D-37073 Góttingen (Germany)

? Conservatoire et jardin botaniques de la Ville de Genéve,

C. P. 60, CH-1292 Chambésy (Switzerland)

ABSTRACT — The status of 7 poorly known species of Leucolejeunea is discussed. New synony-

mies: L. ecuadorensis — Archilejeunea ludoviciana, L. quitensis — Anoplolejeunea conferta, L. trista-

niana — L. unciloba, Archi-Lejeunea erronea (South Africa) — L. clypeata. The latter synonymy

suggests a label error, since L. clypeata is endemic to North America. Furthermore, Cyrtolejeunea

inermis from SE Brazil is referred to Leucolejeunea (as L. sp.) and Leucolejeunea rotalis from St.

Helena is transferred to Aureolejeunea as A. rotalis. The latter constitutes the first record of

Aureolejeunea from the Old World.

RÉSUMÉ — Le statut de 7 espéces mal connues du genre Leucolejeunea est discuté. Synonymies

nouvelles: L. ecuadorensis = Archilejeunea ludoviciana, L. quitensis = Anoplolejeunea conferta, L.

tristaniana = L. unciloba, Archi-Lejeunea erronea (Afrique du Sud) = L. clypeata. Cette dernière

synonymie suggère une erreur d'étiquette, étant donné que L. clypeata est une plante endémique de

l'Amérique du Nord. En plus, Cyrtolejeunea inermis du Sud-Est du Brésil est considérée comme étant

une espèce du genre Leucolejeunea (comme L. sp.) et Leucolejeunea rotalis de St. Helena est transférée

dans le genre Aureolejeunea (A. rotalis). Tl s'agit ici de la première indication d' Aureolejeunea dans

l'Ancien Monde.

Leucolejeunea Evans is a small genus of the Lejeuneaceae subfamily Lejeuneoi-

deae with a pantropical distribution. The genus has never been monographed although

useful regional treatments have been published for the New World by Schuster (1980), for

South East Asia by Grolle & Piippo (1990) and for Africa by Jones (1974). Twenty five

species have been validly described in Leucolejeunea , including twenty that were described

before 1974 and are therefore listed in Index Hepaticarum (Geissler & Bischler, 1985) as

well as five that have been described more recently. They are the following species

(references are given for names not appearing in Index Hepaticarum and for those no

longer accepted in Leucolejeunea; accepted names are in bold face).

L. antiqua Grolle (fossil in Dominican amber, Grolle, 1983a, 1990).

L. caducifolia Gradst. & Schäf.-Verw. (Brazil, Gradstein & al., 1993)

L. capensis S.W.Arnell = L. xanthocarpa (Jones, 1974)

L. clypeata (Schwein.) A.Evans

L. conchifolia (Evans) A.Evans

Source : MNHN, Paris

178 S.R. GRADSTEIN & P. GEISSLER

L. decurrens (Steph.) Mizut. (New Guinea, Grolle & Piippo, 1990)

L. ecuadorensis Steph.

L. flavescens (S.Hatt.) S.Hatt. = Thysananthus flavescens (S.Hatt.) Gradst. (Gradstein,

1991)

L. gradsteinii Grolle & Piippo (New Guinea, Grolle & Piippo, 1990)

L. japonica (Horik.) Verd.

L. knysnana S.W.Arnell = L. unciloba (Arnell, 1961a)

L. loriana (Steph.) Mizut.

L. nana S. Winkl.

L. paroica N. Kitag.

L. planifolia Horik. = Lopholejeunea kiushiana Horik. (Mizutani, 1961; Thiers $: Grads-

tein, 1989)

. quitensis Steph.

L. rotalis (Hook.f. & Taylor) S. Arnell

L. rotundistipula (Lindenb. ex Lehm.) A.Evans

L. sellowiana (Steph.) Steph. = L. unciloba (Evans, 1907)

L. subalpina S. Най. = L. japonica (Mizutani ,1961)

L. suprema Grolle & Piippo (New Guinea, Grolle & Piippo, 1990)

І.

. tristaniana S.W.Arnell

. turgida (Mitt.) Verd.

. unciloba (Lindenb.) A.Evans,

. xanthocarpa (Lehm. & Lindenb.) A.Evans.

It thus appears that at this moment 19 species remain in Leucolejeunea. A further

species, L. brunneorhiza R.M.Schust. from Venezuela (Schuster, 1980), has never been

validly described, and L. hossei (Steph.) “Verd.” and L. sikkimensis (Steph.) “Verd.”, cited

by Mizutani (1976) as synonyms of the Asiatic L. turgida, are invalid names because

Verdoorn (1934) did not actually make these combinations.

In this paper we would like to present the results of our study of the types of 5

species of Leucolejeunea which were not treated in recent revisions and whose status has

therefore remained dubious: Leucolejeunea ecuadorensis, L. nana, L. quitensis, L. rotalis,

and L. tristaniana, Furthermore, we will deal with the identity of two further species that

belong to Leucolejeunea but were described in the wrong genus: Archi-Lejeunea erronea

and Cyrtolejeunea inermis.

1. Leucolejeunea ecuadorensis Steph., Spec. Hep. 4: 737. 1912,

"Type: Ecuador. Prov. Azuay: Gualaquiza, “in silva montis Sapotilla ad truncorum putri-

dum" 2 Sep 1909, Allioni s.n. hb. Levier 6523 (G-25350, holotype) syn. nov.

= Archilejeunea ludoviciana (De Not. ex Lehm.) Geissler & Gradst. subsp. ludoviciana

Good material, mostly dioicous (male) shoots as usual in this species. In addi-

tion, a single autoicous plant with juvenile gynoecia and with androecia on separate

branches, was observed. The occurrence of autoicous sex distribution had not yet been

reported in this species.

2. Leucolejeunea nana S. Winkl., Rev. Bryol. Lichénol. 35: 326. “1967” 1968.

Type: El Salvador. "SE-hang des Cerro Miramundo bei der Hazienda Montecristo,

Metapan, Dep. Santa Ana, 2240m, А/5/4 “, leg. S. Winkler (ULM slide, holotype). The

Source : MNHN, Paris

NOTES ON LEUCOLEJEUNEA (HEPATICAE) 179

other (paratype) collection cited by Winkler in his original publication could not be

located (A. Lücking, pers. comm.)

The type material of Leucolejeunea nana consists of a sterile shoot-fragment, +

3 mm long, 0.5 mm wide. The leaf lobes are suborbicular and deeply concave, upto0.3mm

long, rounded at apex and with a + plane ventral margin; the lobules are very large,

semicircular, ca. 2/3-3/4 x the length of the lobe, the free margin of the lobule is plane, and

the apical margin of the lobule is oblique, continuing slightly into the ventral margin of the

lobe, and has 1-2 short teeth, the second tooth being located at the apex and being smaller

than the first one, sometimes rudimentary developped only. The leaf cells are + isodia-

metrical and rather small, ca. 16-20 um long in the leaf middle, with subquadrate lumina

and uniformily thickened walls (trigones lacking or minute). The underleaves are rather

small and suborbicular, ca. 3 x stem width.

Leucolejeunea nana seems to be a genuine member of the genus Leucolejeunea.

The species seems to differ from other described species of the genus by its huge 2-toothed

lobule and its very small size. Due to the poorness of the type material the identity of L.

nana remains doubtful, however. The possibility that the plant is a mere branchlet of

another described neotropical species cannot be ruled out.

3. Leucolejeunea quitensis Steph., Spec. Hep. 4: 738. 1912.

Type: Ecuador. Prov. Pichincha: Quito, Jameson s.n. (G-25351, holotype) syn. nov.

= Anoplolejeunea conferta (C.F.W.Meissn.) A.Evans

1 The type specimen consists of only a single shoot fragment, with a few androe-

cia.

4. Leucolejeunea rotalis (Hook.f. & Taylor) S.W.Arnell, Svensk Bot. Tidskr. 55: 210. 1961.

(Jungermannia rotalis Hook.f. & Taylor 1845; for further homotypic synonymy see Wig-

ginton & Grolle, 1996).

Type: St. Helena. Dianas Peak. 2500 *, terricola, Febr. 1840, unknown collector (G-25286,

isotype). Further isotypes were seen in S and W by Grolle (Wigginton & Grolle, 1996)

= Aureolejeunea rotalis (Hook.f. & Taylor) Gradst. & Geissler, comb. nov.

Plants (pale) brown in the herbarium. Leaves imbricate, nearly flat, leaf apex

rounded. Lobules oblong, inflated near the base, keel straight to slightly convex, in lower

leaves continuous with free dorsal lobe margin, the free margin of the lobule flat to slightly

incurved with a blunt apical tooth. Median cells ca. 16-20 um wide, + isodiametric,

trigones bulging and becoming confluent at the leaf margin, intermediate thickenings

scarce. Underleaves about 4 x the stem width, orbicular, decurrent, with a circular

insertion line.

Autoicous. Androecia on short lateral branches, bracteoles limited to the base

(see also Arnell, 1961b). Gynoecia on short lateral branches with one lejeuneoid innova-

tion, perianths almost eplicate.

The isotype in G consists of 9 stem fragments, 5 of which are fertile. The

brownish colour of the plants and the lejeuneoid (instead of pycnolejeuneoid) innovations

indicate that this species belongs in Aureolejeunea rather than in Leucolejeunea. The

lejeuneoid innovation of Aureolejeunea rotalis has already been noted by Thiers (1985, sub

Archilejeunea rotalis (Hook. & Taylor) Steph) and by Grolle (in Wigginton & Grolle,

1996). Aureolejeunea rotalis seems most closely related to the widespread neotropical A.

Source : MNHN, Paris

180 S.R. GRADSTEIN & P. GEISSLER

fulva R.M.Schust., but the leaves of the latter are much more convex and the underleaves

somewhat larger.

The genus Aureolejeunea (5-6 spp.) is widespread at high elevations in Tropical

America and, in addition, occurs in southern Chile. This is the first record of the genus

from the Old World.

5. Leucolejeunea tristaniana S.W.Arnell, Christophersen, Results Могу. Sci. Exped. Tris-

tan da Cunha 1937-38, 42: 15. 1958.

Type: Tristan da Cunha. Inaccessible Island: Plateau at W. end, steep rock face, 475m, leg.

E. Christophersen de Y. Mejland, n.2493, 23/2/1938. (S, isotype) syn. nov.

=L. unciloba (Lindenb.) A.Evans

Leucolejeunea tristaniana seems to be a robust form of L. unciloba. The plants

are about 2 mm wide.

The isotype from S consists of a nice patch of whitish to light brown plants,

resembling Brazilian populations which are generally more vigorous than those of North

America.

6. Archi-Lejeunea erronea Steph., Hedwigia 27: 113. 1888

Type: “South Africa. Cape of Good Hope, leg. Ecklon” [?, probably label confusion]

(G-11008, holotype) syn. nov.

= Leucolejeunea clypeata (Schwein.) Evans

Already Gradstein & Buskes (1985) reported that the type specimen belongs to

the genus Leucolejeunea, probably to Leucolejeunea rotundistipula (Lindenb.) Steph.

Grolle (1995) published the formal synonymy and noted the difficulty to determine the

identity of the scrappy type specimen. Our study of the specimen revealed that the lobules

never bear a discrete tooth, they are ovate-triangular, only slightly inflated; the lobes are

flat, the keel is nearly straight and in an almost continuous line with the dorsal lobe

margin. These features correspond more to Leucolejeunea clypeata than to L. rotundisti-

pula; in fact the plants are inseparable from L. clypeata. We therefore cannot agree with the

synonymy published by Grolle (1995). It is quite improbable that Leucolejeunea clypeata,

endemic to eastern North America, should occur in South Africa besides L. rotundistipula

and L. xanthocarpa. We presume that some specimens were confused by Stephani during

herbarium manipulations and that a label error is at hand.

Archilejeunea (Spruce) Schiffn. has not yet been conserved against Archi-

Lejeunea Steph. 1888, created by a 'descriptio generico-specifica', and therefore the

publication of a conservation proposal becomes urgent (Grolle, 1983b, 1995).

7. Cyrtolejeunea inermis Herzog, Memoranda Soc. Fauna Fl. Fenn. 25: 70. 1949.

Type: Brazil, 540 Paulo, Estaçäo Biologica do Alto da Serra, epiphyll, Е C. Hoehne 561

p.p. (JE, holotype)

— Leucolejeunea sp.

Like Leucolejeunea nana, the type material of Cyrtolejeunea inermis consists of

a sterile shoot-fragment with ca. 12 pairs of leaves, + 4 mm long, 0.5 mm wide. The leaf

lobesare ovate, up to 0.3 mm long, rounded at apex and with a + hollow (but not inflexed)

ventral margin; the lobules are subrectangular , ca. 1/2 x the length of the lobe, the keel is

almost straight and forms a wide angle of ca. 140? with the ventral margin of the lobe, the

Source : MNHN, Paris

NOTES ON LEUCOLEJEUNEA (HEPATICAE) 181

free margin of the lobule is plane, the apical margin of the lobule is truncate, and the apex

is broad and blunt (a discrete tooth is lacking). The leaf cells are slightly elongated, ca.

25-30 um long in the leaf middle, with ellipsoid lumina and distinct trigones. The under-

leaves are rather small and suborbicular, 3-4 x stem width.

By thecharacters of its leaf lobules and cells, Cyrtolejeunea inermis appears to be

a member of Leucolejeunea rather than Cyrtolejeunea. The same conclusion was drawn by

Grolle (in sched.). Although similar in size to L. nana S. Winkler, Cyrtolejeunea inermis is

quite different from L. nana by its lobule and its leaf cells (see descriptions). Like L. nana,

the identity of Cyrtolejeunea inermis remains doubtful due to the poorness of the material;

this problem may be resolved in the framework of a careful study into the variation of the

neotropical species of the genus Leucolejeunea. It could well be that the specimen is a mere

branchlet of L. conchifolia since we have observed that the lobule tooth may be poorly

developed in secondary shoots of the latter species.

ACKNOWLEDGEMENTS — We are grateful to Dr. Andrea Lücking (ULM) for the loan of the

type of Leucolejeunea nana, to Dr. H.-J. Zündorf, curator of the Herbarium Haussknecht (JE) for the

loan of Cyrtolejeunea inermis, and to Dr. L. Hedenás and Dr. U. Malmgren (S) for the loan of

Leucolejeunea tristaniana.

REFERENCES

ARNELL S., 1961a — Notes on South African Hepaticae VI. Botaniska Notiser 114 (4): 400-402.

ARNELL S., 1961b — Some new American species of hepatics. Svensk Botanisk Tidskrift 55:

205-210.

EVANS A. W., 1907 — Leucolejeunea, a new genus of Hepaticae. Torreya 7: 225-229.

GEISSLER P. & BISCHLER H. (eds.), 1985 — Index Hepaticarum 10 (Lembidium to Mytilopsis).

Vaduz — Genéve.

GRADSTEIN S. К. & BUSKES G.M.C., 1985 — A revision of neotropical Archilejeunea (Spruce)

Schiffn. Beihefte zur Nova Hedwigia 80: 89-112.

GRADSTEIN S. R., 1991 — Diversity and distribution of Asian Lejeuneaceae subfamily Ptychan-

thoideae. Tropical Bryology 4: 1-16.

GRADSTEINS. R., GROLLE R. & SCHÁFER-VERWIMP A., 1993 — Two interesting species of

Lejeuneaceae from Brazil. Journal of the Hattori Botanical Laboratory 74: 59-70.

GROLLE R., 1983a — Leucolejeunea antiqua n. sp., das erste Lebermoos aus Dominikanischem

Bernstein. Stuttgarter Beitráge zur Naturkunde, Ser. B, 96: 1-9.

GROLLE R., 1983b — Nomina generica Hepaticarum. Acta Botanica Fennica 121: 1-62.

GROLLE R., 1990 - Leucolejeunea antiqua (cin Lebermoos in Dominikanischem Bernstein) erstmals

mit Gynözium, Perianth und Andrózium. Nova Hedwigia 50: 473-479.

GROLLE R. & РПРРО S., 1990 — Bryophyte flora of the Huon Peninsula, Papua New Guinea.

XXXVII. Leucolejeunea (Lejeuneaceae, Hepaticae). Annales Botanici Fennici 27: 119-129.

GROLLE R., 1995 — The Hepaticae and Anthocerotae of the East African Islands. An annotated

Catalogue. Bryophytorum Bibliotheca 48: 1-178.

JONES E. W., 1974— African Hepatics. XXIV. Lejeuneaceae: some new or little-known species and

extensions of range. Journal of Bryology 7: 545-561.

MIZUTANI M., 1961 — A revision of Japanese Lejeuneaceae. Journal of the Hattori Botanical

Laboratory 30: 115-302.

MIZUTANI M., 1976 — Studies of little known Asiatic species of Hepaticae in the Stephani

herbarium. 9. Some little-known species of the family Lejeuneaceae. Journal of the Hattori

Botanical Laboratory 40: 441-446.

Source : MNHN, Paris

182 S.R. GRADSTEIN & P. GEISSLER

SCHUSTER R. M., 1980 — The Hepaticae and Anthocerotae of North America. New York, vol. IV:

1-1334.

THIERS B. M., 1985 — Branching in Lejeuneaceae III: Ptychanthoideae. Beihefte zur Nova Hedwi-

gia 80: 31-61.

THIERS B. M. & GRADSTEIN S.R.,1989 — Lejeuneaceae (Hepaticae) of Australia. I. Subfamily

Ptychanthoideae. Memoirs of the New York Botanical Garden 52: 1-82.

VERDOORN Е, 1934 — Studien über asiatische Jubuleae. Annales Bryologici 4 (suppl.): 1-231.

WIGGINTON M. J. & GROLLE R., 1996 — Catalogue of the Hepaticae and Anthocerotae of

Sub-Saharan Africa. Bryophytorum Bibliotheca 50: 1-267.

Source : MNHN. Paris

Cryptogamie, Bryol. Lichénol. 1997, 18 (3): 183-189 183

UN RICCIA PROLIFÈRE D'AUSTRALIE :

RICCIA PULLULANS SP. NOV. (SOUS-GENRE RICCIELLA)

Suzanne JOVET-AST

Résidence Cháteau d'Arcadie, 34 bis, avenue de la République

F-64200 BIARRITZ — FRANCE

RÉSUMÉ — Description d'une nouvelle espéce récoltée par O. Volk dans le Western Australia.

Principaux caractéres : épiderme dorsal couvert d'une mince pellicule transparente et granuleuse ;

pores nombreux sur la face dorsale ; cellules épidermiques proches des cellules annexes des pores

contenant un corps orangé ; multiplication végétative par la face ventrale d'un thalle mère d’où prend

naissance une suite de thalles fils de plusieurs générations successives.

SUMMARY — Description of a new species collected by O. Volk in Western Australia. Principal

characters : dorsal epidermis covered with a thin, transparent, granulose pellicle ; epidermal pores

numerous ; epidermal cells close to the annexed cells of pores containing an orange body. The species

displays a unique type of vegetative multiplication : from the ventral face of mother thallus arises a

succession of young thalli each one born on the ventral face of the previous one.

Parmi les Riccia australiens, une espéce, inconnue jusqu'alors, se distingue par

son comportement biologique qui se traduit par une multiplication végétative non obser-

Vee chez les autres espèces du genre Riccia (Jovet-Ast, 1991). A ce mode très particulier de

croissance végétative, l'espéce nouvelle doit son nom « pullulans ».

Riccia pullulans sp. nov.

TYPE — Australie, Western Australia, noch feuchter Boden an der Basis einer grossen

Granitglatze, frisch-trocken nitratreich (Mist), offen sehr grosse Fláchen deckend. « The

Humps » bei Heyden. pH: 6.3. Leg. Otto Volk, 23.3.1979, n? 071. (PC, holotype).

Diagnose latine — Thallus monoicus. Lobus albidus vel cyanescens in sicco, 3-5 mm longus,

2-3.5 mm latus, crassus, paulo bulbosus, lucens, translucida pellicula obtectus. Epidermidis

pori numerosi, 55 um diam. Epidermis cellulae proximae ab annexis cellulis pori aurantia-

cum corpus includunt. Squamae hyalinae, aequales margine lobi. Sporae 72-77-(80) um

diam. ; ala irregulariter dentata, 2.5-5 um lata ; distalis superficies cum 8-10 alveolis in

Source : MNHN. Paris

184 S. JOVET-AST

diametro ; proximalis superficies cum triradiata nota distincta, alveolis et tuberculis conspi-

cuis.

Description — Thalle (Pl.1 ; PI. 2, fig. 1-6) monoique, en grandes plaques composées de

lobes grégaires. Sur le sec, lobes blanchátres ou gris bleuté, un peu bulleux, brillants, à face

dorsale couverte d'une trés mince pellicule finement granuleuse, incolore, transparente,

chatoyante, recouvrant méme les pores alors visibles par transparence, se déchirant par les

chocset par l'áge ; à bifurcation basale profonde ; certains courtement bifides au sommet ;

longs de 3-5 mm, larges de 2-3.5 mm, épais, étalés en 2 ailes à la partie supérieure puis

brusquement et longuement rétrécis ; à sillon très étroit au sommet puis s'élargissant et

délimitant une partie médiane un peu bombée. Face dorsale à pores nombreux. Pores de

55 um de diamètre environ, bordés de 6 cellules annexes ; les cellules épidermiques qui

entourent les cellules annexes contiennent un gros corps brun orangé. Faces latérales

obliques, portant des écailles atteignant le bord du lobe mais ne le dépassant pas, à cellules

violettes ou rouge sombre, éparses parmi des cellules incolores ou formant de grandes

taches violettes surtout vers la marge.

Face ventrale couverte de très nombreux rhizoides lisses et tuberculés.

Section transversale 2 fois plus large que haute, parfois à peine plus large que haute,

comprenant : a) sous la fine pellicule granuleuse, la section des cellules annexes du pore et

celle des cellules épidermiques ; dans les cellules épidermiques situées contre les cellules

annexes, un corps brun orangé ; b) le tissu dorsal composé de 2-4 assises de chambres

aériféres limitées par un réseau de cloisons épaisses d'une seule cellule, soit, dans les parties

les plus larges du lobe, de hauts tubes aérifères s'ouvrant directement à l'extérieur ; c) le

tissu ventral dense à cellules petites et à parois minces. Anthéridies trés abondantes sur le

type ; archégones beaucoup plus rares.

Spores (P1. 2, fig. 8-11 ; Ph.1-4) brun rouge, de 72-77-(80) pm diam. ; aile irrégulièrement

dentée, large de 2.5-5 um, à 3 pores trés petits ; face distale à 8-10 alvéoles dans le diamétre,

à murets assez hauts, portant des tubercules dans les angles ; face proximale à marque

triradiée nette, à facettes portant des alvéoles de 4-5 um de diamètre, un peu plus petites

que celles de la face distale, à murets assez élevés munis, aux angles, de tubercules saillants.

Multiplication végétative — Chez le type, elle s'est effectuée selon trois modes :

1) selon le mode déjà observé chez quelques espéces du genre Riccia. Sur la face ventrale

d'un lobe mére, nait un lobe fils qui posséde la méme structure que la mére et qui se libére

rapidement (Pl. 3, fig. 1, 2).

2) par des lobes à structure particuliére (P1. 3, fig. 3-7) détachés de leur mere, mesurant 2

mm de hauteur, larges de 2.5 mm. Ils se composent de différents tissus : un tissu dorsal

dense formé de files paralléles de cellules, entouré de chambres aériféres ; un tissu ventral

de petites cellules compactes ; au-dessous, un pied incolore sur le sec, haut de 1 mm,

cylindrique, composé de longues cellules de 60-100 x 25-38 jim, à parois minces ; à la base

de ce pied, une masse de cellules isodiamétriques, de 50-60 x 50 um, à parois minces, qui

semblent provenir de la face ventrale du lobe mére.

3) par multiplication en chaine (P1. 3, fig. 8), forme curieuse jamais observée chez d'autres

Riccia. Un lobe bien développé porte, sur la face ventrale, un lobe fils un peu plus petit que

lui. Ce dernier ne se libére pas, devient, à son tour, un lobe mére donnant naissance à un

lobe fils sur sa face ventrale. Dans le cas observé, cette multiplication se produit 5 fois de

suite, Il en résulte la formation d'une masse allongée, un peu gélatineuse, dure, liée au lobe

mère d'origine, formée d'une chaîne de petits lobes solidement attachés les uns aux autres.

Cette masse trés résistante ne se brise pas : il faut couper les lobes pour les séparer.

Source : MNHN, Paris

Pl. 1 — Riccia pullulans sp. nov. Australie, Volk 071 (type). 1, 2 : lobes, face dorsale. Les points

représentent les pores vus par transparence. 3, 4 : sommets de lobes, face dorsale. 5 : aspect d'un

fragment de face dorsale, vu en plan. Les cercles représentent les pores ; les points, les fins granules de

la pellicule couvrant la face dorsale d'un lobe. 6-13 : sections transversales de lobes.

Source : MNHN, Paris

PI. 2— Riccia pullulans sp. nov. Australie, Volk 071 (type). 1 : section transversale d'un lobe au niveau

de chambres aériféres. 2 : section transversale d'un lobe dans une partie ayant des canaux aériféres.

3: un pore épidermique entouré des cellules annexes et des cellules épidermiques, vu en plan. Corps

orangés, en pointillé, sous la paroi des cellules épidermiques. 4, 5 : pores, cellules annexes et cellules

épidermiques, vus en plan. 6, 7 : pores, cellules annexes et cellules épidermiques contenant le corps

orangé, en section transversale. 8 : spore, face distale. 9 : alvéoles et tubercules de la face distale d'une

spore. 10, 11 : spores, face proximale à deux mises au point différentes.

Source : MNHN, Paris

30y m

Pl. 3 — Riccia pullulans sp. nov. Australie, Volk 071 (type). — Multiplication végétative. 1, 2: lobes fils

nés sur la face ventrale d'un lobe mère (mode 1). 3 : lobe fils détaché d’un lobe mere (mode 2). 4, 5:

section transversale du lobe de la fig. 3. 6: cellules du pied des lobes 4 et 5. 7: cellules du pied et cellules

de la base du pied des lobes 4 et 5. 8: un chaine de 5 lobes fils (2 à 6) nés sur un lobe mere (1), (mode

3). m : lobe mère ; f : lobe fils ; td ; tissu dorsal ; tv : tissu ventral ; p : pied ; ca : canaux aériféres.

tv: tissu ventral ; саб : chambre aérifére ; el : canal aérifère ; p : pore ; ca : cellule annexe ; cé : cellule

épidermique ; b : corps orangé dans des cellules annexes ; pg : pellicule granuleuse couvrant l'épi-

derme ; r : par transparence, limite d'une chambre aérifère.

Source : MNHN. Paris

188 S. JOVET-AST

Photos 1-4 — Riccia pullulans sp. nov. Australie, Volk 071 (type). 1, 2 : spore, face distale. 3, 4: spore,

face proximale. Diamètre : 75 шт.

Distribution — Australie, Western Australia, The Humps, prés de Hyden (le type).

Comportement biologique — D'aprés l'étiquette de récolte de O. Volk, R. pullulans

occupait au sol une grande surface. Les belles dimensions du spécimen recueilli et la

densité des thalles le prouvent. Ce Riccia était dans un état de reproduction sexuée et de

multiplication végétative intenses. Il vivait donc dans une station écologiquement favora-

ble, suffisamment humide. Sa localisation dans le Western Australia en grande partie

désertique peut surprendre, mais la récolte a été effectuée prés de Hyden situé dans la

pointe sud-occidentale du Western Australia. Les pluies d'été de cette région qui abrite des

foréts déterminent un climat favorable au R. pullulans, espéce hygrophile, peut-étre méme

hydrophile.

Affinités — R. pullulans appartient au sous-genre Ricciella (Braun) Boulay. On peut le

classer dans le sous-groupe Vesiculosa créé par Na-Thalang (1980) et qui rassemble 6

espèces à lobes étroits (larges de 2-4 mm) et spores petites (75-85 um de diamètre),

terrestres, hygrophiles, souvent résistantes à la chaleur.

Source : MNHN, Paris

RICCIA PULLULANS SP. NOV. 189

Trois espéces monoiques de ce sous-groupe sont proches de R. pullulans :

a) К. muscicola Steph., mais cette езресе, localisée dans les New South Wales, a des lobes

trés minces (4-5 fois plus larges que hauts) ; des spores dont le diamétre mesure 75-80 um

et méme 100 um ; des alvéoles au nombre de 12 dans le diametre de la face distale et 14

d'aprés la photographie D, planche 3, de la publication de Na-Thalang (1980).

b) R. deserticola Howe distinct de R. pullulans par ses lobes minces, par ses spores à aile

étroite (2 um) et ses pores épidermiques bordés de 4 cellules annexes.

c) R. crassa Steph. différent de R. pullulans par ses lobes trés caverneux et ses pores

épidermiques bordés de 4 cellules annexes.

Le corps orangé contenu dans certaines cellules épidermiques rappelle les idio-

blastes présents chez trois espéces de Riccia : R. macrocarpa Lev., R. campbelliana Howe,

R. dictyospora Howe. Il n'indique cependant aucune affinité précise avec ces trois espéces

саг R. pullulans appartient au sous-genre Ricciella alors que les autres espéces se classent

dans le sous-genre Riccia.

CONCLUSION

Le type de R. pullulans est remarquable d'une part par sa vitalité peu commune

attestée par l'abondance de ses thalles juxtaposés, par l'intensité de sa reproduction sexuée,

la particularité de sa multiplication végétative, d'autre part par la présence, dans certaines

cellules épidermiques, d'un corps orangé rappelant les idioblastes présents chez quelques

Riccia.

R. pullulans contribue à mettre en lumiére la richesse et la diversité du genre

Riccia en Australie et la valeur exceptionnelle des récoltes de Otto Volk.

BIBLIOGRAPHIE

JOVET-AST S., 1991 — Riccia (Hépatiques, Marchantiales) d'Amérique Latine, Taxons du sous-

genre Riccia. Cryptogamie, Bryologie-Lichénologie 12 (3) : 189-370.

NA-THALANG O., 1980 — A revision of the genus Riccia (Hepaticae) in Australia. Brunonia 3 :

61-140.

Source : MNHN, Paris

Cryptogamie, Bryol. Lichénol. 1997, 18 (3): 191-193 191

PLAGIOCHILA BISCHLERIANA, A NEW SPECIES FROM NEPAL

R. GROLLE ! and M. L. SO?

"Institut für Spezielle Botanik, Friedrich-Schiller-Universität, D-07743 Jena, Germany

?Biology Department, Hong Kong Baptist University, 224 Waterloo Road, Hong Kong

ABSTRACT — Plagiochila bischleriana spec. nov. is described from Nepal. It is recognized by the

following characteristics: 1) plants large, 2) leaves triangularly ovate, 3) leaf margin with long ciliate

teeth, 4) leaf apex subacute, 5) leaf-cells without trigones and 6) perianth long cylindrical with dorsal

keel narrowly winged.

In 1989 D. G. Long (E) gathered a large collection of bryophytes in Nepal. We

are indebted to him for sending us the Plagiochila specimens for study. Although botanists

from N. Wallich (1786-1854) up to some recent Japanese expeditions have collected

bryophytes extensively in Nepal, Long's material contained a handsome, easily distinguis-

hable new species of Plagiochila, that is described and figured here. We are pleased to name

itin honour of our colleague Dr. Hélène Bischler-Causse (Paris).

Plagiochila bischleriana Grolle & M.L.So sp. nov.

Plagiochilae fuscae Sande Lac. similis, sed differt 1) foliis triangulatim ovatis

(non /ate ovatis), 2) cellulis foliorum trigonis nullis vel minutis (non trigonis distinctis

subnodulosis), 3) perianthio dorsaliter humiliter alato (non exalato).

HOLOTYPE: NEPAL, N-facing slope of Dobala Danda above Kabeli Khola, ravine in

wet mossy oak/laurel forest; on mossy boulders, 2350 m, 1989, Long 17412; E (c. per.).

ISOTYPE in JE.

Further specimen studied: Same locality, Long 17380; E (sterile).

Plants large, soft-textured, occurring in loose tufts, 10-12 cm long and 5.9-

6.4 mm wide with leaves, light green, somewhat glossy, no creeping rhizomatous caulid

seen; sparingly branched, branches exclusively lateral-intercalary; stem 280-300 um thick,

light brown, 18-20 cells across, cortical cells in 2 layers, moderately thick-walled, oblong to

rectangular, 16-20 x 20-24 шт, medullary cells 20-24 x 24-36 um, thin-walled, trigones

minute; rhizoids only at the bases of shoots.

Leaves delicate, almost membranous, moderately imbricate to contiguous, obli-

quely to horizontally spreading, never secund nor revolute, oblong-ovate to ovate, dis-

tinctly longer than wide (length/width ratio 1.3-1.71:1), 2.62-2.8 mm long, 1.6-1.9 mm

Source : MNHN, Paris

192 R. GROLLE & M.L. SO

dee

Bra os

аа ze z

Fig 1. — Plagiochila bischleriana sp. nov. — a. Portion of plant, dorsal side. — b. Portion of plant,

ventral side. - c, d. Leaves. — e. Female bract. - f. Marginal tooth of leaf, — g. Perianth. — h. Median

cells of leaf. All drawn from holotype (Long 17412). Scale bars: A = 1 mm (ae, g), B = 0.1 mm (f, h).

Source : MNHN. Paris

PLAGIOCHILA BISCHLERIANA SP. NOV. 193

wide, widest near the base; dorsal margin distinctly arched, with 9-11 spinose teeth, leaf

base moderately decurrent along dorsal midline, leaving stem surface widely exposed; leaf

apex broadly rounded to truncate, with 5-8 ciliate-spinose teeth, ventral margin arched,

moderately ampliate at the base, bearing 12-14 spinose teeth, ventral stem surface com-

pletely hidden, leaf-base hardly decurrent, teeth 24-26 in total, spinose-dentate, 2-6 cells

long in a uniseriate row, and 2-3(4) cells wide at the base, terminal cell elongated,

sometimes deciduous, 12 x 72 um and almost 6 x as long as wide; cells along the leaf apex

30-36 x 30-36(40) um, those in the leaf middle to base 36-40 x 46-60(76) um, vitta absent,

walls evenly thickened throughout, trigones absent or small, cuticle smooth. Underleaves

absent. Paraphyllia absent. Asexual reproduction not seen.

Male plants not seen. Female inflorescence terminal on stem with 2 innovations

arising from the bases of bracts, bracts in 2 pairs, almost similar to leaves in shape and

marginal dentation, but much larger, measuring 3.05 mm long, 2.38 mm wide. Perianth

long cylindrical, 3.77 mm long, 1.30 mm wide, cells of perianth similar to that of leaves,

without trigones, mouth bilabiate, arched, irregularly spinose — dentate, with over 40

teeth, teeth slightly longer than those on the leaves, dorsal keel narrowly winged. Sporo-

phytes not seen.

Superficially, this species resembles an over-sized P. denticulata Steph. because

of the longciliate-spinose teeth along the whole margin of the leaf, and the evenly, slightly

thickened walls of leaf cells, But Р. denticulata is only 1/10 of the size of P. bischleriana, is

dark brown, soft in texture and glossy in appearance. P bischleriana also resembles

members of the Ciliatae in having long ciliate teeth, almost translucent leaves and large

leaf cells. But the Ciliatae have distinct trigones in the leaf cells and the cells are mostly

thin-walled.

In size, P. bischleriana more resembles P. gymnoclada Sande Lac. (Java, Philip-

pines, Sri Lanka, Japan, New Guinea) and Р. fusca Sande Lac. (Java, Celebes, Philippines),

both belonging to section Plagiochila (Inoue, 1984), but these species are very rigid in

texture and the leaves are broadly ovate with a broadly rounded apex. Furthermore, the

total number of teeth is 20-35 per leaf and the leaf cells have + large nodulose trigones.

The three closely related species in the section Plagiochila can be separated by the

following key:

l. Plants soft in texture, leaves triangularly ovate, ventral leaf base ampliate, teeth

spinose-dentate, 2-6 cells long, terminal cell 6 x as long as wide, dorsal keel of perianth

narrowly winged P. bischleriana

1. Plants rigid in texture, leaves broadly ovate, ventral leaf base not ampliate ....... 2

2. Plants very glossy, teeth small, 2-5 cells long, terminal cell 1.1-1.5 x as long as wide,

dorsal keel of perianth with a distinct wing . P gymnoclada

2. Plants dull green to brown, teeth 5-9 cells long, terminal cell 34. 5 xas long as wide,

keels of perianth unyiged ha ER VOOR LR Eh ДИЛ ead P. fusca

REFERENCES

INOUE H., 1965 — Contributions to the knowledge of the Plagiochilaceae of Southeastern Asia.

VII. Synopsis of Plagiochila Dum. in the Himalayan region. Bulletin of the Naional

Science Museum (Tokyo) 8: 375-403.

INOUE H., 1984 — The genus Plagiochila (Dum.) Dum. in Southeast Asia. Tokyo. 142 p., 76 pls.

Source : MNHN, Paris

me e MOST RETAMA E: ОТОК.

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Cryptogamie, Bryol. Lichénol. 1997, 18 (3): 195-205 195

NEW OR LITTLE KNOWN EPIPHYLLOUS LIVERWORTS,

VII. TWO NEW LEJEUNEACEAE SPECIES

FROM THE MASCARENE ISLANDS

Tamás PÓCS

Eszterházy Teachers'College, Dept. of Botany, EGER, Pf.43,H-3301,

HUNGARY, Fax: (36)36-321-016, e-mail: colura@gemini.ektf.hu

RÉSUMÉ — Colura mauritiana et Drepanolejeunea helenae, hépatiques épiphylles et corticoles des

Mascareignes, sont décrites comme espèces nouvelles pour la science. Colura mauritiana, de l'ile

Maurice, appartient à la section Harmophyllum bien qu'elle ait une ressemblance superficielle avec C.

meijeri, membre asiatique de la section Gamolepis. Drepanolejeunea helenae, de Vile de la Réunion, est

affine de D. ankasica de l'Afrique occidentale (Ghana). Sa place dans un sous-genre demeure

indéterminée. Les deux espéces sont restreintes à une aire limitée sur les iles concernées et semblent

endémiques.

SUMMARY — Colura mauritiana and Drepanolejeunea helenae are described as new to science.

Both species occur as epiphyllous and corticolous in the Mascarene islands, the first from Mau-

ritius and the second from Réunion. Colura mauritiana belongs to the section Harmophyllum

although showing superficial resemblance to C. meijeri, an Asian member of section Gamolepis.

Drepanolejeunea helenae is related to the West African (Ghana) D. ankasica. Its subgeneric place is

uncertain. Both species are restricted to a small area within the concerned islands and seem to be

endemic.

Key worps: epiphylls, Lejeuneaceae, Colura, Drepanolejeunea, Mascarenes.

The bryological exploration of the Mascarene Islands is the part of the Bryolo-

gia Africana project initiated by Prof. P.W. Richards (1984). I paid special attention to the

rich hepatic flora of the Indian Ocean islands (recently catalogued by Grolle 1995), which,

according to studies made by different authors in the last two decades, is still quite

underexplored and may yield many novelties (Pócs, 1993, 1995).

Members of our Department made intensive studies on the bryoflora of Mas-

carene Islands, in collaboration with the Muséum d'Histoire Naturelle de Saint Denis,

Réunion, with the Conservatoire Botanique and with the Université de la Réunion,

France and with the Herbarium of Sugar Research Institute, Mauritius. In 1994, 1995 and

1996 three consecutive collecting trips took place in Réunion and two on Mauritius

Island. Our aim, apart from the floristic and taxonomic exploration, was to study in detail

the cryptogamic vegetation of Réunion Island. The present publication deals with 2 new

species resulting from these studies, when numerous relevés were made (cf. also Pócs &

Tóthmérész in press) and hundreds of samples were identified.

Source : MNHN, Paris

196 T. PÓCS

By the description of the habitats of new species, the nomenclature of accom-

panying species is according to Grolle (1995), Wigginton & Grolle (1996) and O'Shea

(1995), where the authors' names also can be found.

Colura mauritiana Pócs sp. nov. (Sectio Harmophyllum Grolle)

Pl. 1 (Figs. 1-9)

Diagnosis — Planta epiphylla vel ramicola pallide virens vel flavovirens caulibus

irregulariter ramosis, ad 5 mm longis, 50-90 um crassis et cum foliis 1.5-2 mm latis. Folia

contigua, 0.6-1.2 mm longa et 0.3-0.5 mm lata sacculis claviformibus apicibus late hemis-

phaericis. Clypeus late orbicularis cellulis 7-11 centralibus et 14-15 marginalibus composita.

Amphigastria acutilobata lobis 4-5 cellulae lata. Gemmae 32 cellulosae. Ceterae desunt.

Holotypus: Mauritius, Pócs 9530/H (EGR; Isotypi: MAU, PC). Paratypi: Pócs

95291BJ (EGR) et 9531/C (EGR).

Description — Epiphyllous or ramicolous, pale or yellowish green with solitary

or slightly branching shoots of 1.5-2 mm width and up to 5 mm length or forming 2-3 mm

large, dense patches. Stems 50-90 um thick, with 7 rows of cortical and 3 rows of

medullary cells (4 rows near the leaf insertion), with evenly thickened walls. Lobes with

entire margin, sac claviform, with a broad hemisphaeric and sometimes crestlike, conically

protuberant group of cells. Lobe and sac cells 20-30 x 30-40 um, with heart-shaped

trigones and 1-3 intermediate thickenings per wall. Valve broad ovate or orbicular, with

14-15 hyaline marginal, 6-10 central cells and with 1 basal median cell. Hinge formed by

3 cells, frame around the valve by 7-9 cells and with a hyaline papilla.

Underleaves with 60-95 um broad, lanceolate lobes that are 4-5 cells wide at their

bases, Gemmae of 32 cells, with 4 growing points. Gametangia and sporophyte not

observed.

Specimens examined — Mauritius: Perrier Nature Reserve in the neighbourhood of Mare

aux Vacoas, submontane Sideroxylon thicket with Trochetia and with Cyathea borbonica, at 540 m

alt., 20°21/50” S, 57°29’30” E. epiphyllous, on Syderoxylon puberulum leaf, associated with Cheilole-

jeunea serpentina, Ch. surrepens, Cololejeunea marginata, C. cristata, Colura digitalis, C. heimii, C.

obvoluta, Diplasiolejeunea cavifolia, D. comorensis, Drepanolejeunea madagascariensis, D. trematodes

and all transitions between the two, Lejeunea tabularis s.l., L. caespitosa, Leptolejeunea maculata and

others (S.& T. Pócs 9529/BJ, Paratypus, EGR). Black River Gorges National Park: Le Petrin Nature

Reserve. Philippia brachyphylla heath with scattered trees, at 650 m alt. 209255, 57*28/20" E, creeping

on Philippia twigs, accompanied by Cheilolejeunea trifaria, Drepanolejeunea vesiculosa, Frullania

onraedtii, F. serrata, Leucoloma dichelymoides and by Leucoloma grimmioides (T. Pécs 9530/H,

Holotype: EGR, Isotypes: MAU, PC). Macchabée Upland Climax Forest Reserve in the canopy with

Labourdonnaisia at 640 m alt. near the road junction, 2024/30” S, 57%27'50” E. Epiphyllous on

naturalized Psidium cattleyanum leaves, accompanied by Cololejeunea zenkeri and by Frullania

purpurea and corticolous, accompanied by Drepanolejeunea cambouena and by Jamesoniella

contracta (T. Pócs 9531/С , Paratypus in the microslide collection of EGR).

All the above collections were made on the 17th March, 1995, when we were with

my wife guided and accompanied by Mr. Joseph Guého and by Miss Danielle Florens,

from the National Herbarium in Mauritius Sugar Industry Research Institute, Réduit.

Colura mauritiana has a superficial resemblance to the Asian Colura meijeri

Jovet-Ast, but the latter belongs to Section Gamolepis, which according to Jovet-Ast

(1953) has a valve fused to the hinge and of many more (40-60) cells. The valve of C.

Source : MNHN, Paris

TWO NEW LEJEUNEACEAE SPECIES 197

PL 1— Colura mauritiana sp. nov., Mauritius. 1: Ventral view of shoot. — 2: Conically protuberant

sac cells, — 3: Leaf, dorsal view. — 4: Valves (in different magnifications). — 5: Cells of lobe margin.

6: of lobe centre. — 7: of the sac. — 8: Underleaves. — 9: Gemma. (Figs 1, 2 and 4 upper two are

drawn from Pócs 9529/BJ and all others from the Holotype: Pócs 9530/H).

Source : MNHN. Paris

198 T.PÓCS

mauritiana is very easily detachable and is composed of fewer (20-35) cells. Their under-

leaves are also different, C. mauritiana having underleaf lobes 4-6 cells broad well above

their mid length while the underleaf lobes of C. meijeri from a 4 cells broad base

immediately taper to 2 cells width. The closest relatives of our new species are probably the

Asian Colura hemisphaerica Jovet-Ast and Colura cymbalifera Herz. from the Section

Harmophyllum, but these species differ again by the much higher number of valve cells

(40-60) and by the different shape of their underleaves. Colura mauritiana has no relatives

in Africa and seems to be an endemic species in Mauritius, with Asian affiliation and

origin, strengthening the Asian affinities of the flora of Indian Ocean islands (Pócs, in

press).

Drepanolejeunea helenae Pócs sp. nov.

PI. 2-4 (Figs. 10-29), Photo

Diagnosis — Planta autoica, corticola vel epiphylla, in statu vivo prasina ocellis

lucentibus, sicca flavovirens. Caules 50-70 um crassi, cum foliis 0.6-0.85 mm lati, irregulari-

ter ramulosi, caespites densi diametro 5-40 mm formans. Folia falcata, apice acuminato,

margine irregulariter obtuse eroso-dentata. Cellulae medianae 15-25 x 28-40 um, ocelli 2-25,

cellulas alias lobi aequantes.

Lobulus oblonge ovatus, margine quatrocellularis saepe incurvo. Dens primus

lobuli unicellularis falcatus, dens secundus obtusus. Papilla hyalina inter dentes duos ad

marginem lobuli crescens. Amphigastria biloba caulibus 3-4-plo latiora lobis acuto-

lanceolatis sinu obtuso.

Androecia in spicis lateralibus saepe sub gynoecio, 2-4 jugis antheridiis singulis.

Perianthia 0.6-0.7 x 0.4 mm, obconica, quinquecarinata carinis spinoso-crenulatis saepe

cornuta. Rostrum elongatum (ad 100 um) apice dilatata. Seta articulata, capsula diametro

250 um, elaterae unispiralae.

Species nova in honorem Doctoris Helenae Bischler-Causse, investigatoris illustris-

simae Drepanolejeunearum Neotropicae dedicata.

Holotypus: Réunion, Pócs 9651/DH (EGR; Isotypi: REU, G, H, JE, PC, MAU,

MO, NY). Paratypi:Vojtkö 9433/AN (microslide collection of EGR); Kis & Vojtkó

9651/CC (EGR).

Description — Autoicous, forming vivid green, 5-40 mm large patches on bark or

more scarcely on living leaves. The glistening dots of the numerous ocelli well visible even

by a handlens. When dry, in herbarium, yellowish green and the ocelli hardly visible.

Shoots 0.6-0.85 mm wide, loosely attached to the substrate. Stems irregularly branching,

50-70 um thick, composed of 7 cortical and 3 medullary cells with evenly thickened walls,

as usual in Drepanolejeunea (Bischler, 1967). Leaves contiguous or imbricate, 3-500 um

long and 2-300 um wide, asymmetric, falcate, with undulately obtuso-dentate margin and

acuminate, often incurved apex ending in 1-2 short cells. Leaf base variable, well exceeding

the stem or just reaching its median line and parallel to it. Central lobe cells 15-25 x

28-40 um, with irregular intermediate thickenings. Ocelli 2-25 (in most cases 10-16) per

leaf, similar to ordinary lobe cells in size but their walls almost evenly thickened, 2 ocelli

usually forming a short basal vitta above the lobule margin, others solitary or geminate

and scattered or arranged in a few irregular lines; ocelli in living plants containing one

large, strongly refractive, slightly granulose oil body of the Leptolejeunea type (in sense of

Kis & Pécs, 1997), while the normal lobe and lobule cells have 1-6 Calypogeia type oil

Source : MNHN, Paris

PI. 2— Drepanolejeunea helenae sp. nov., La Réunion. 10: dorsal view of shoot. On the lower piece the

position of ocelli is shown. — 11: Ventral view. — 12: Underleaves. — 13: Lobule with incurved free

margin. — 14: Part of shoot with male branches. Ventral view. — 15: Male branches with antheridia.

15а: Crest of a male bract. (All drawn from the Holotype).

Source : MNHN. Paris

PI. 3 — Drepanolejeunea helenae sp. nov., La Réunion. 16: Leaves with ocelli. — 17: Oil bodies of

ocelli and of a normal lobe cell. In the third cell the chloroplasts are shown. — 18: Lobules with the

variation of lobule teeth and papilla position. — 19: Underleaf. (All drawn from the Holotype with

the exception of 17:G Kis 9651/CC).

Source : MNHN. Paris

№

PL 4 — Drepanolejeunea helenae sp. nov., La Réunion. 20: Female branch with perianth and

pycnolejeuncoid subfloral innovation (perichaetial leaves removed for better visibility). — 21: Peri-

chaetium. — 22: Perichaetial leaves. — 23: Perianths, — 24: Perianth beak. — 25: Perianth horn. —

26: Keel of a perianth. — 27: Sporophyte. — 28: Elater. — 29: Sporangium wall thickenings. (All

drawn from the type).

Source : MNHN. Paris

202 T. PÓCS

Photo — Type specimen of Drepanolejeunea helenae alive, with well visible ocelli. (Photo Andras

Szabó).

bodies (in sense of Gradstein et al., 1977). Leaf apex and margins with smaller, rectangu-

lar cells of 12-20 um.

Lobules oblong ovate, inflated and with often incurved, 4-celled free margin.

Lobule cells similar to lobe cells but intermediate thickenings more scarce. First (distal)

tooth unicellular, acute and falcate, second (proximal) tooth blunt and separated from the

first tooth by one cell, which usually develops at the inner margin surface and is adnate to

the first tooth. Hyaline papilla in the notch between the first or second teeth or on the tip

of the intermediate cell. Base of the distal lobule margin usually with a cell bulging,

opposing the apex of the falcate first tooth (Fig. 18).

Underleaves bilobed with acute lobes of 2-3 cells width at their base and with a

wide open sinus between them. Adhesive disk rarely developing, from the median cells of

underleaf base (Fig. 12), a real paramphigastrium (Bischler, 1969) not observed.

Androecia on short branches, often more than one near to a gynoecium or on

sterile parts of the stem, with 2-4 pairs of bracts, each with one antheridium of 55-70 um

diameter, upper bracts usually with a crest of protuberant cells; 2-3 bracteoles present

(Fig. 14).

Gynoecia (Pl. 4) on short side branches, each with one pycnolejeuneoid innova-

tion (Fig. 20). Perichatium about half the length of the perianth, with one pair of

unequally bilobed bracts with a few large, irregular teeth. Bracteole acutely bilobed,

Source : MNHN, Paris

TWO NEW LEJEUNEACEAE SPECIES 203

partially connate with the bracts, in cases with an additional tooth and a hyaline papilla.

Perianth obconic or urne-shaped, 0.6-0.7 x 0.4 mm, with scattered ocelli, almost truncate

above with an elongated (up to 100 um), dilatate beak and with 5 spinoso-crenulate keels

ending often in an acute horn.

Seta strongly articulate. Capsule 250 um in diameter, capsule wall with very pale,

irregular thickenings (Fig. 29). Elaters 180 um long, with well developed unispiral band

(Fig. 28). Spores unknown.

The new species is dedicated to the Dr. Héléne Bischler-Causse, well known

monographer of the Neotropical species of genus Drepanolejeunea and many other critical

groups of Hepaticae.

Specimens examined — Réunion: W of Ste Anne, epiphyllous in remnants of tropical

rainforest in the valley around “le Grand Etang”, at 500-550 m altitude, 214/42" S, 5593850" E, 29

Aug.1994. Associated with Ceratolejeunea belangeriana, Cololejeunea af. bebourensis, C. borbonica,

С. duvignaudii, C. marginata, C. obliqua, C. tanzaniae, Colura digitalis, Diplasiolejeunea cavifolia, D.

cornuta, Drepanolejeunea madagascariensis, D. trematodes and all transitions between the two,

Lejeunea caespitosa, Frullania apicalis, Taxilejeunea furciornuta and others. A. Vojtkó 9433/AN (Рата.

type, in the microslide collection of EGR). SSW slope of Piton de la Fournaise volcano, at the W edge

of Basse Vallée. Secundary, naturalized Psidium cattleyanum stand with remnants of former montane

rain forest trees along the trail GR R2 leading towards Piton Lardé, at 750 m alt., 21919'S, 55°41'Е.

Corticolous on the smooth bark of small Psidium cattleyanum and Aphloia theaeformis trees in open

or in shade. Accompanied by Ceratolejeunea, Diplasiolejeunea, Lopholejeunea and Marchesinia spp.

Acrolejeunea aulacophora, Bazzania nitida, Drepanolejeunea vesiculosa, Frullania usambarana, Lep

dolejeunea delesserti, Plagiochila repanda and others, (T. Pécs 9651/DH, 23 July 1996, Holotype:

EGR; Isotypes: REU, G, H, JE, PC, MAU, MO, NY; G Kis & A. Vojtkó 96511CC, 16 July 1996, from

the same site, Paratype, EGR).

The appearence of this plant in the field and under microscope at the first glance

suggested an unknown Harpalejeunea species because of the falcate, pointed leaves with

basal twin ocelli and underleaves with shallow lunate incision. However, Dr. Riclef Grolle

(Jena) was kind to call my attention to the pycnolejeuneoid innovations below the

perianths, which indicate obviously its proper place in the genus Drepanolejeunea. There is

only one species within the genus, which shows closer affinity to our species: Drepanolejeu-

nea ankasica E.W. Jones (in Jones & Harrington, 1983), described from Ghana, West

Africa. The two species have several important features in common, including the falcate

leaves with acute, often incurved apex, the irregularly dentate leaf margin, the geminate

basal ocelli, four-celled free lobule margin, and the arrangement of the lobule teeth. The

autoicous sex distribution (rare feature within Drepanolejeunea, cf. Bischler, 1964) is also

shared by the two species. Other characters, like the ocellate, obconical perianths and the

bulging, distal lobule keel cell opposite the first tooth occur in many other species too (cf.

Bischler, 1964, 1968; Herzog, 1930, 1934, 1937, 1939).

The combination of characters of the two species makes it difficult to interpret

their subgeneric position. Due to their lobule teeth arrangement, subgen. Kolpolejeunea

Grolle (Grolle, 1976) is out of question. The free lobule margin formed by four cells.

suggests subgen. Drepanolejeunea Grolle, but the many ocelli (at least in D. helenae), the

gynoecial arrangement and the leaf shape are typical for subgen. Pristolejeunea Grolle,

especially the tropical Asian Drepanolejeunea thwaitesiana and its relatives (Mizutani,

1990). From these, the two African species are well distinguished by the four-celled free

lobule margin, the totally different underleaf lobes. These are horizontally widely sprea-

ding in D. thwaitesiana group, trapezoid to V shaped in the two African species, like in the

Species group VI of Bischler, 1964) of neotropical Drepanolejeunea. Further studies might

Source : MNHN. Paris

204 T. PÓCS

justify to delimitate a new subgenus or at least a section to accommodate D. ankasica and

D. helenae.

The new species, which seems to be endemic in la Réunion, differs sufficiently

from D. ankasica by the many (up to 25) ocelli in the leaf lobe (only 2-3 in D. ankasica), by

the very different shape of underleaves (obtrapeziform with widely diverging lobes in D.

ankasica, V shaped with narrower lobes in D. helenae), the different perianth keels and

beak and by the ocelli in perianth (not reported in D. ankasica).

ACKNOWLEDGEMENTS — I wish to recognize with thanks the funding of our field research in the

Indian Ocean Islands by the following sponsoring agencies: National Geographic Society (USA),

Grant 5201/94; Hungarian National Science Foundation, Grants No. OTKA 1/3-941, OTKA U

18611, OTKA W 015072 and the Hungarian Ministry of Education and Culture, K & F (Research

and Development) Grant.

We are very grateful to the local scientific institutions and to the persons for organizing and

helping our activities in the field, especially to Dr. Sonja Ribes, Conservateur Principal du Muséum

d'Histoire Naturelle de la Réunion, Philippe Pagezy, Directeur Régional de l'Office National des

Foréts, la Réunion, France and to Mr. Joseph Guého and to Miss Danielle Florens, from the National

Herbarium in the Sugar Industry Research Institute, Réduit, Mauritius and to many others. Finally

Tam specially indebted to Drs. Riclef Grolle and 5. Rob Gradstein for their advise in the treatment of

the new species.

REFERENCES

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Revue Bryologique et Lichénologique 33 (1-2): 15-179.

BISCHLER H., 1967 — Recherches sur l'anatomie de la tige chez les Lejeuneaceae. Ш. Les

sous-familles Lejeuneoideae, tribu Lejeuneeae, et Tuyamaelloideae. Revue Bryologique et

Lichénologique « 1966 > 1967, 34 (3-4): 601-675.

BISCHLER H., 1968 — Le genre Drepanolejeunea Steph. en Amérique Centrale et Meridionale. П.

Revue Bryologique et Lichénologique « 1967 » 1968, 35 (1-4): 95-134.

BISCHLER H., 1969 — Notes sur l'anatomie des amphigastres et sur le développement du param-

phigastre et des rhizoides chez Drepano-, Rhaphido- et Leptolejeunea. Revue Bryologique et

Lichénologique « 1968 » 1969, 36 (1-2): 45-55.

GRADSTEINSR., CLEEF A.M. & FULFORD M.H., 1977 — Studies on Colombian cryptogams.

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Wetenschappen, Ser. C, 80 (5): 377-420.

GROLLE R., 1976 — Drepanolejeunea Subgen. Kolpolejeunea — Fine neue Untergattung aus der

Paläotropis. Journal of the Hattori Botanical Laboratory 40: 191-216.

GROLLE R., 1995 — The Hepaticae and Anthocerotae of the East African Islands. An Annotated

Catalogue. Bryophytorum Bibliotheca 48, 178 p.

HERZOG Th., 1930 — Studien über Drepanolejeunea 1. Annales Bryologici 3: 126-149.

HERZOG Th., 1934 — Studien über Drepanolejeunea 11. Annales Bryologici 7: 57-94.

HERZOG Th., 1937 — Studien über Drepanolejeunea ШТ. Annales Bryologici 9: 115-130.

HERZOG Th., 1939 — Studien über Drepanolejeunea IV. Annales Bryologici 12: 98-122.

JONES E.W. & HARRINGTON A.J., 1983 — The hepatics of Sierra Leone and Ghana. Bulletin of

the British Museum (Natural History), Botany series 11 (3): 215-289.

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Lichénologique 22 (3-4): 206-312; supplément. Revue Bryologique et Lichénologique 23

(1-2): 1-22.

Source : MNHN, Paris

TWO NEW LEJEUNEACEAE SPECIES 205

KIS G. & PÓCS T., 1997 — Ой body studies on African Hepaticae. Journal of the Hattori Botanical

Laboratory 81: 175-242.

MIZUTANI, M., 1990 — Notes on the Lejeuneaceae. 16. Drepanolejeunea thwaitesiana and its

related species from Asia. Journal of the Hattori Botanical Laboratory 68: 367-380.

O'SHEA B., 1995 — Checklist of the mosses of sub-saharan Africa. Tropical Bryology 10: 91-198.

PÓCS T., 1993 — New or little known epiphyllous liverworts, IV. Two new Cololejeuneoideae from the

i Comoro Archipelago. Journal of the Hattori Botanical Laboratory 14: 45-57.

PÓCS T., 1995— East African bryophytes, XIV. Hepaticae from the Indian Ocean islands. Fragmenta

Floristica et Geobotanica 40: 251-277.

PÓCS T., in press — The distribution and origin of the foliicolous bryophyta in the Indian Ocean

islands. Proceedings of the ТАВ & IAL Symposium of Foliicolous Cryptogams, 29 August —

2 September 1995, Eger, Hungary. Abstracta Botanica.

PÓCS T. & TOTHMERESZ B,, in press — Foliicolous bryophyte diversity in tropical rainforests.

Proceedings of the IAB 4 IAL Symposium of Foliicolous Cryptogams, 29 August — 2

September 1995, Eger, Hungary. Abstracta Botanica.

RICHARDS PW, 1984 — The bryologically under-worked regions of the world, with special

reference to West Africa and a proposal for a Bryologia Africana. Journal of the Hattori

Botanical Laboratory 55: 165-172.

WIGGINTON M.J. & GROLLE R. (supplemented by A. Gyarmati), 1996 — Catalogue of the

Hepaticae of Sub-Saharan Africa. Bryophytorum Bibliotheca 50, 267 p.

Source : MNHN, Paris

Cryptogamie, Bryol. Lichénol. 1997, 18 (3): 207-212 207

NEW MOSS RECORDS AND RANGE EXTENSIONS

OF SOME XERIC AND ALPINE MOSS SPECIES IN CHINA

Benito C. TAN! & Jian-cheng ZHAO?

"Harvard University Herbaria, 22 Divinity Avenue, Cambridge, MA, USA 02138

"Department of Biology, Hebei Normal University, Shijiazhuang, Hebei, China 050016

ABSTRACT — Crossidium crassinerve, Crossidium seriatum and Polytrichastrum sexangulare are

three mosses reported new to China. Range extension and correction of several rare, xeric and alpine

moss taxa in China are reported. Tortula scotteri and Polytrichum crassilamellatum are proposed as

synonyms of Hilpertia velenovskyi and Polytrichastrum sexangulare respectively.

Our study of many collections of bryophytes made recently from arid and alpine

localities in western and northern China, notably Qinghai and Xinjiang Provinces, yielded

significant range extensions for seven rare mosses in China. Of these, Crossidium crassi-

nerve, C. seriatum and Polytrichastrum sexangulare are new to China. Hilpertia velenovskyi

is new to Xinjiang and Qinghai. Grimmia subanodon, G. obtusifolia and Schistidium chenii

are three new moss records for Qinghai Province. In addition, reports of Crossidium

chloronotos from Gansu, Xinjiang and Inner Mongolia are corrected to C. crassinerve.

Lastly, Tortula scotteri (=Hilpertia velenovskyi) and Polytrichum crassilamellatum (=Poly-

trichastrum sexangulare) are two new synonyms proposed.

1. Crossidium seriatum Crum and Steere and C. crassinerve (De Not.) Jur.

Chinese record of Crossidium was first recorded by P.-C. Chen who reported C.

chloronotos (Brid.) Jur, from Gansu Province (P.-C. Chen et al., 1963). Chen and subse-

quent Chinese authors have overlooked the incorrect application of C. chloronotos ЕС

squamiferum (Viv.) Jur.] on European specimens of C. crassinerve in the old literature. This

taxonomic confusion was later clarified by Delgadillo (1975). Unaware of this report,

Zhao (1993) continued to use the incorrect name when he reviewed the morphology and

ecological adaptation of C. chloronotos in China and published a nation-wide distribution

map for this species. Following Delgadillo (1975), Redfearn et al. (1 996) synonymized the

name with C. squamiferum (Viv.) Jur. and, inadvertently, included the provinces of Gansu,

Inner Mongolia, Sichuan and Xinjiang in the range of C. squamiferum for China. Recently

Z.-H. Zhang (1996) reported C. squamiferum from the Huangguoshu karst area in

Guizhou Province.

Source : MNHN, Paris

208 B.C. TAN & JIAN-CHENG ZHAO

According to Delgadillo (1975) and Cano et al. (1993), Crossidium squamiferum

differs from C. crassinerve in having very thick-walled upper laminal cells. Frey and

Kürschner (1984) further observed that the terminal cell of the leaf costal chlorophyllose

filaments of C. crassinerve has 1 to 2 low papillae whereas that of C. squamiferum has 2 to

5 high papillae. To clarify the situation, we re-examined specimens of C. chloronotos from

Gansu, Xinjiang and Inner Mongolia and found them to be C. crassinerve (De Not.) Jur.

Our conclusion is supported further by the illustrations and descriptions of C. chloronotos

published in P. C. Chen er al. (1963), Tung (1963) and Zhao (1993); all of which represent

clearly C. crassinerve. Thus, the total range of C. crassinerve in China encompasses the arid

provinces of Gansu, Inner Mongolia, Xinjiang, and in addition, Qinghai (Tan 95-1698b,

FH, PE). The Sichuan and Guizhou records of C. chloronotos (=C. squamiferum)

published in Redfearn et al. (1994) and Zhang (1996) need confirmation.

Crossidium seriatum, a species distinguished from other congeners by the pre-

sence of 4 to 8 c-shaped papillae per leaf cell, is also found in Qinghai Province (Tan

95-1731, FH, PE). It is a new addition to the moss flora of China. Although the Qinghai

specimens comprise only young sporophytes, the leaf outline with a long hyaline awn and

the morphology of the costal filaments of the leaves clearly identify the specimen as C.

seriatum. Furthermore, the leaf cells bear the characteristically c-shaped papillae (cf.

Delgadillo, 1975). The discovery of C. seriatum on the Qinghai-Tibetan plateau has

extended the Old World range of this species from western Asia eastward to western

China.

2. Hilpertia velenovskyi (Schiffn.) Zand. [Tortula scotteri Zand. & Steere; Hilpertia scotteri

(Zand. & Steere) Zand., syn. nov.] — Figs 1-16.

The genus Hilpertia was proposed by Zander (1989) to accomodate two pottia-

ceous taxa, the Eurasian Tortula velenovskyi Schiffn. and the North American T. scotteri

Zand. & Steere. Both taxa are characterized by (1) a strongly revolute, chlorophyllose leaf

margin and (2) the nearly smooth laminal cells, except those at the leaf margins. According

to Zander and Steere (1978), the two species differ primarily in the presence of a few teeth

near leaf apex in H. velenovskyi (absent in H. scotteri), and also in the rostrate opercular lid

in the former and a conical lid in H. scotteri. Two recent collections from Xinjiang

Province (J.-C. Zhao 5676, 5679, HBNU, FH) and several collections from Qinghai

Province (Tan 95-1703, 95-1723, 95-1725, FH, PE) were determined with difficulty as H.

velenovskyi because the specimens show a suite of combined characters of H. velenovskyi

and Н. scotteri.

The Xinjiang collections are from Qira Xian in Kunlun Mountains and the

Qinghai collections are from Yushu Prefecture bordered with Sichuan to the south. The

plants are common on calcareous walls around villages and on exposed soil growing mixed

with species of Aloina and Crossidium. Individual plants are 1 to 2 cm high. The stem cross

section shows a large central strand with one layer of colored, thick-walled epidermis.

Leaf apices are mostly entire, with a few leaves showing a few teeth at the base of the long

awn. The opercular lids are long-conical, although a few are conical. The same variation

in opercular lid shape can be observed in the holotype of Tortula velenovskyi Schiffn.

preserved at FH. The isotype of Tortula scotteri Zand. & Steere from northern Canada

preserved at BUF has a few individual plants with a long conical lid. Considering the

overlapping morphological variation exhibited by the two species under comparison, we

think that it is meaningless to maintain Н. scotteri as a separate species.

Source : MNHN, Paris

NEW MOSS RECORDS IN CHINA 209

:25mm 13 14

pum 4 15

Figures 1-16 — Hilpertia velenovskyi (Z.-C. Zhao 5676). — 1: Plant habit. 2: Capsule. 3: Peristome

tooth. 4: Spores. 5-12: Leaves. 13. Leaf apex. 14. Cross-section of leaf. 15. Middle leaf cells. 16. Lower

leaf cells.

Source : MNHN, Paris

210 B.C. TAN & JIAN-CHENG ZHAO

Prior to our study, Hilpertia velenovskyi was known from central and eastern

Europe, Siberia (Ignatov and Afonina, 1992) and Inner Mongolia of China (Tung, 1963;

Bai, 1987). We report it here new to the provinces of Xinjiang and Qinghai. Its occurrence

in arid habitats in the neighboring provinces of Gansu, Shaanxi and Xizang can be

expected. As now recognized, the genus is monotypic with a broad northern boreal range.

The report of H. velenovskyi from South America in Bai (1987) is a typographical error for

North America.

3. Polytrichastrum sexangulare (Floerke ех Brid.) G. L. Smith [Polytichum crassilamellatum

W.-X. Xu & R.-L. Xiong, syn. nov.)

A recently collected polytrichaceous specimen (R.-L. Hu 95-80b, FH, SHNU)

from No. 1 Glacier Research Station in the Tianshan Range of Xinjiang Province proved

to be Polytrichastrum sexangulare, a new record for the province, and China as well.

Polytrichastrum sexangulare is distinguished from all other congeners in China

by its somewhat cucullate leaf apex and entire to weakly toothed leaf margins. The

presence of small teeth or appendages on the abaxial side of the leaf costa near the apex is

an additional diagnostic feature. In cross section, the apical cells of the leaf lamellae are

typically pyriform and with smooth, thick, distal cell walls.

The Xinjiang specimen was originally identified as the Tibetan endemic, Poly-

trichum crassilamellatum W.-X. Xu & R.-L. Xiong. However, a critical review of published

descriptions and illustrations of these two taxa (Osada, 1966 as Polytrichum norvegicum

Hedw.; Nyholm, 1969; Lawton, 1971; Xu and Xiong, 1984; Li, 1985; Ignatov and

Smith-Merrill, 1995) and careful examination of herbarium specimens, including the

isotype of Polytrichum crassilamellatum preserved at K UN, did not reveal any meaningful

gametophytic differences between P. crassilamellatum and Polytrichastrum sexangulare.

Although sporophyte is unknown in Р. crassilamellatum, we believe that the two taxa are

conspecific.

Specimens of Polytrichastrum sexangulare can easily be confused with sterile

plants of Polytrichum sphaerothecium (Besch.) C. Muell., which has a similar leaf lamellar

morphology. According to Osada (1965), the leaf sheath of P. sexangulare, seen in the

Xinjiang specimens and in the isotype of Polytrichum crassilamellatum, is more expanded

than the leaf sheath of Polytrichum sphaerothecium. In contrast, Schofield (1966), who

had observed the variation of leaf characters in British Columbian plants of P. sphaero-

thecium, concluded that many specimens would be indistinguishable from P. sexangulare

on gametophytic characters alone. Not withstanding, fruiting individuals of Р sphaero-

thecium differ distinctively from P. sexangulare in having round, non-angular capsules

(Schofield, 1966). Seen in this light, the newly described Polytrichastrum altaicum Ignatov

& Merr. from the Russian Altai (Ignatov and Smith-Merrill, 1995) is more closely related

to P. sphaerothecium than to P. sexangulare.

Polytrichastrum sexangulare is a circum-arctic and alpine species growing on

moist soil in open tundra and in late snow areas. Its presence near a glacier at 3500 m

elevation in Xinjiang, China, is typical of its habitat preference.

Source : MNHN, Paris

NEW MOSS RECORDS IN CHINA 21

4. Schistidium chenii (H.-S. Lin) T. Cao, C. Gao & J.-C. Zhao [syn. Grimmia himalayana

Chen, nom. inval.]

A population of grimmiaceous plants with entirely immersed capsules was

collected from a creek bank in a limey gorge above 4000 m elevation in Nanggen Xian,

Qinghai Province (Tan 95-1612, FH, PE). The specimen has imbricate, concave, and

broadly ovate leaves with obtuse and cucullate leaf apices. A hyaline apical mucro is

observed on several perichaetial leaves. Bistratose areas also appear sporadically across

the upper 1/3 of the lamina and along the leaf margins. The collection was identified as

Schistidium chenii, a Chinese endemic known previously from Xizang (Tibet) and Xinjiang

provinces. Qinghai is the third province for this distinctive, rare Chinese endemic. Cao et

al. (1992) have published a good illustration for this species.

5. Grimmia obtusifolia C. Gao & T. Cao and Grimmia subanodon Ochyra [syn. Schistidium

obtusifolium Irel. & Crum]

A small grimmiaceous specimen with the leaf apex obtuse from Qinghai Pro-

vince (Tan 95-176, FH, PE) was confirmed to be Schistidium obtusifolium (=Grimmia

subanodon Ochyra), formerly an eastern North American endemic known only from

Quebec, Canada. Both the Chinese and Canadian specimens were collected from dry,

calcareous substrates. The leaves of the Qinghai plants, which are unistratose and ovate-

oblong to oblong-lingulate, compare well with the illustration of the leaves published in

the protologue of the type species (Ireland and Crum, 1984). The Qinghai specimen is the

second Chinese collection of this disjunctive eastern North American and western Chi-

nese species. Its first Chinese report was made by Bai and Hao ( 1996) who collected the

species from Inner Mongolia.

Another Qinghai specimen (Tan 95-198, FH, PE), with a similar obtuse leaf apex

but with multi-stratose leaf lamina, proves to be Grimmia obtusifolia, a Chinese endemic

reported from Inner Mongolia, Sichuan, Xinjiang and Xizang (Redfearn er al., 1996). Cao

and Vitt (1985) had discussed and illustrated well the differences between this taxon and

Grimmia subanodon. A new record for the province, the Qinghai collection of G. obtusifo-

lia, as in all other collections, does not have sporophytes. However, the specimen bears

several perigonia, indicating a dioicous sexual condition.

ACKNOWLEDGEMENTS — We are grateful to Dr. D. Boufford for reading and correcting the

English text. Mr. Jia Yu of Beijing Herbarium (PE) helped in the preparation of final illustrations.

Field assistance of Prof. Mamtimim Sulayman of Xinjiang University, Prof; R.-L. Hu of East China

Normal University and Prof. Lu Xie-Feng of Northwest Plateau Institute of Biology are appreciated.

Funding provided by the National Geographic Society for the expeditions to Xinjiang and Qinghai

provinces is equally acknowledged.

REFERENCES

BAI X.-L., 1987 — Preliminary report of the mosses in Inner Mongolia, China. Acta Scientiarum

Naturalium Universitatis Intramongolicae 18: 311-350.

Source : MNHN. Paris

212 B.C. TAN & JIAN-CHENG ZHAO

BAI X.-L. & HAO L.-F, 1996 — Some moss species new to China. Acta Scientiarum Naturalium

Universitatis Nei-Monggol 27: 412-416.

CANO M.J., GUERRA J. & ROS R.M., 1993 — A revision of the moss genus Crossidium (Pottia-

седе) with the description of the new genus Microcrossidium. Plant Systematics and

Evolution 188: 213-235.

CAO T., GAO C. & ZHAO J.-C., 1992 — Studies of Chinese bryophytes (3). Revision of Grimmia

chenii Lin (Bryopsida, Grimmiaceae). Journal of the Hattori Botanical Laboratory 7:

69-73.

CAO T. & VITT D.H., 1985 — A taxonomic assessment of Grimmia obtusifolia Gao et Cao and

Schistidium obtusifolium Irel. et Crum. The Bryologist 88: 362-364.

CHEN P-C. et al., 1963 — Genera Muscorum Sinicorum, Pars Prima. Beijing, Science Publisher.

DELGADILLO C.M., 1975 — Taxonomic revision of Aloina, Aloinella and Crossidium (Musci). The

Bryologist 78: 245-303.

IGNATOV M.S. & AFONINA O.M., 1992 — Checklist of mosses of the former USSR. Arctoa 1:

1-85.

IGNATOV M.S. & SMITH-MERRILL G.L., 1995 — Bryophytes of Altai Mountains. VI. The

family Polytrichaceae (Musci). Arctoa 5: 61-97.

IRELAND R.R. & CRUM H.A., 1984 — Schistidium obtusifolium, a new species of the Grimmia-

ceae from Quebec. The Bryologist 87: 371-373.

FREY W. & KÜRSCHNER H., 1984 — Studies in Arabian bryophytes 3:Crossidium asirense

(Pottiaceae), a new species from Asir Mountains (Saudi Arabia). Journal of Bryology 13:

25-31.

LAWTON E., 1971 — Moss Flora of the Pacific Northwest. Nichinan, Hattori Botanical Laboratory.

LI X.-J. (ed.), 1985 — Bryoflora of Xizang. Beijing, Science Press.

NYHOLM E., 1969 — Illustrated Moss Flora of Fennoscandia. 11. Musci. Fasc. 6. Lund, Natural

Science Research Council.

OSADA T, 1965 — Japanese Polytrichaceae. I. Introduction and the genus Pogonatum. Journal of the

Hattori Botanical Laboratory 28: 171-201.

OSADA T., 1966 — Japanese Polytrichaceae. П. The genera Polytrichum, Oligotrichum, Bartramiop-

sis and Atrichum and phytogeography. Journal of the Hattori Botanical Laboratory 29:

1-52,

REDFEARN PL.JR., ALLEN B. & HE S., 1994 — New distributional records for Chinese mosses.

The Bryologist 97: 275-276.

REDFEARN PLIR., TAN B.C. & HE S., 1996 — A newly updated and annotated checklist of

Chinese mosses. Journal of the Hattori Botanical Laboratory 79: 163-357.

SCHOFIELD W.B., 1966 — The identity of Polytrichum sphaerothecium (Besch.) Broth. Miscellanea

Bryologica et Lichenologica 4; 33-35.

TUNG C.-K., 1963 — Notes on some xerophytic mosses in China. Acta Scientiarum Naturalium

Universitatis Intramongolicae 2: 73-85.

XU W.-X. & XIONG R.-L., 1984 — Taxa nova polytrichacearum sinensium. Acta Botanica Yunna-

nica 6: 173-184.

ZANDER R.H., 1989 — Seven new genera in Pottiaceae (Musci) and a lectotype for Syntrichia.

Phytologia 65: 424-436.

ZANDER В.Н. & STEERE W.C., 1978 — Tortula scotteri, sp. nov. from the Northwest Territories of

Canada. The Bryologist 81: 463-467.

ZHANG Z.-H., 1996 — Contributions to the bryoflora of Guizhou, SW China: new records and

habitat notes on mosses from Huangguoshu karst area. Journal of Bryology 19: 149-152.

ZHAO J.-C., 1993 — A preliminary study on the morphological characters and geographical

distribution of Crossidium chloronotos (Brid.) Limpr. Arid Zone Research 10: 49-52.

Source : MNHN. Paris

Cryptogamie, Bryol. Lichénol. 1997, 18 (3): 213-215 213

THE FIRST REPORT

OF CRYPTOTHALLUS MIRABILIS MALMBORG

(HEPATICAE, ANEURACEAE)

IN SOUTHERN EUROPE (PORTUGAL)

Cecilia SÉRGIO! & Ana SÉNECA?

"Museu, Laboratório e Jardim Botánico, Rua da Escola Politécnica,

58, 1294 Lisboa Codex, Portugal.

? Instituto de Botánica Dr. Gongalo Sampaio, Rua do Campo Alegre 1191, 4100 Porto, Portugal.

ABSTRACT — Cryptothallus mirabilis Malmborg was discovered in Sphagnum communities in the

central eastern part of Portugal. The presence of C. mirabilis in Portugal extends the range of the

species to southern Europe and to the vicinity of the Mediterranean area.

RÉSUMÉ — Cryptothallus mirabilis Malmborg a été trouvé au centre du Portugal. Cette découverte

étend l'aire de répartition de C. mirabilis au Sud de l'Europe.

Key wonps: Cryptothallus mirabilis, Portugal.

During a field work on Sphagnum communities in June 1995, Cryptothallus

mirabilis Malmborg was discovered in the area of a Natural Park, in the central eastern

part of Portugal.

This plant is neither reported from any country of South Europe, nor from any

Iberian locality.

C. mirabilis, first described in 1933 by Malmborg from Scandinavia, is conside-

red to be a North Oceanic hepatic (Duell, 1983). At present, it is known only from the

British Isles, Denmark, Finland, Norway, Sweden. Some new localities have been added

by Dickson & al. (1975) from Russia (Leningrad and Karelia) and recently from Brittany

(Bates & Hodgetts, 1995). However, the majority of records still belong to the British Isles

(Hill & al., 1991). The species has also a very restricted distribution in North America and

Western Greenland (Petersen, 1972).

C. mirabilis is a very interesting parasitic liverwort (Pocock & Duckett, 1984)

inhabiting typically mires and bogs, under Sphagnum or associated to Molinia litter (Hill

& al., 1991).

The first locality in Portugal is located in the Sào Mamede Natural Park. The

area includes a great part of Säo Mamede mountain (altitudes from 1025 m to 350 m,

average 500m) and presents a complex physiography. The highest points occur on ridge

formations made up of quartzites (Devonian) and the bedrock is predominantly formed

Source : MNHN, Paris.

214 C. SÉRGIO & A. SÉNECA

Fig. 1 — Cryptothallus mirabilis Malmborg. Small male plants (A), antheridium (B). Specimen:

Sérgio de Séneca 9727 (LISU).

by granites and schists. The climate shows a wide diversity: mean temperatures range from

5.8 C to 8.5" C in January and 21.5? C to 23.3? С in July, with annual rainfall of 1000 to

1200 mm. The flora is diverse, ranging from Mediterranean to Atlantic oak forests. These

forests consist mainly of Quercus suber L., Q. rotundifolia Lam., Q. pyrenaica Willd. but a

large part of the area is covered by plantations of Pinus pinaster Aiton. Deciduous forests

with Quercus robur L., Q. faginea Lam. and Castanea sativa Miller still cover important

areas of the Park.

Because of the wide diversity of habitats, the bryophyte flora of this area is

particularly rich, not only for Mediterranean elements, but also for Atlantic bryophytes. It

corresponds to the southern limits of the geographical distributions in Portugal of many

bryophytes, such as Lophozia excisa (Dicks.) Dum. and Racomitrium elongatum Frisvoll

(Sérgio & al., 1994).

The vegetation of the new locality of Cryptothallus mirabilis is an conifer

plantation (Pinus pinaster Aiton) and the substrate is on the acid litter at the margin of a

stream near a small pond with Salix atrocinerea Brot., Molinia caerulea (L.) Moench,

Erica ciliaris L., Erica tetralix L., Juncus effusus L., J. tenageia L. fil., Ulex minor Roth,

Calluna vulgaris (L.) Hull., Pteridium aquilinum (L.) Kuhn, Lobelia urens L., Blechnum

spicant (L.) Roth, Wahlenbergia hederacea (L.) Reichenb., Potentilla erecta (L.) Ráuschel,

and Pinguicula lusitanica L. On dried margins of the pond, there Drosophyllum lusitani-

cum (L.) Link and Mediterranean species such as Lavandula stoechas L. subsp. luisieri

(Rozeira) Rozeira and Cistus spp. are present.

The margins of the stream and pond have a rich bryophyte flora with Calypogeia

arguta Nees & Mont., C. suecica (Н. Arn. & J. Perss.) К. Muell., Cephalozia bicuspidata

(L.) Dum., Jungermannia gracillima Sm., Fissidens curnovii Mitt., F. polyphyllus Wils. ex

B., 5. & G., Pseudotaxiphyllum elegans (Brid.) Iwats. and different forms of Sphagnum

auriculatum Schimp.

Source - MNHN. Paris

CRYPTOTHALLUS MIRABILIS IN PORTUGAL 215

The population which was first found consisted of small male plants. Later in the

same locality, two larger colonies without sporophytes were discovered (Fig. 1).

The presence of Cryptothallus mirabilis in Portugal extends the range of the

species to southern Europe and to the vicinity of the Mediterranean area.

The species is easily overlooked because of its subterranean growth (Schuster,

1992). More collections are needed, especially from Atlantic parts of the Iberian Penin-

sula, to elucidate the present distribution of this interesting species.

Specimens: PORTUGAL. Alto Alentejo: Parque Natural da Serra de Sáo

Mamede, entre a Cruz dos Cumes e a Feiteira, 750 m, 298 PD415, 26.06.1995, Sérgio &

Séneca 9727 (LISU); ibidem, 13.07.1995, Sérgio 9738 (LISU).

ACKNOWLEDGMENTS — Our sincerest thanks to M. P. Jones for correcting the English text.

Thanks are also due to the authorities Natural Park and to Eng. Castro Antunes for assistance during

the field work.

REFERENCES

BATES J. W. & HODGETTS N. G., 1995 — New and interesting Bryophyte records from Brittany

including Cryptothallus mirabilis, Ulota calvescens and Weissia perssonii new to France.

Cryptogamie, Bryologie-Lichénologie 16(3): 191-211.

DICKSON J. H., KOPONEN T. & ULVINEN T., 1975 — Cryptothallus mirabilis Мать, löydetty

jälleen Suomesta. Luonnon Tutkija 79: 53-56.

DUELL R., 1983 — Distribution of European and Macaronesian Liverworts (Hepaticophytina).

Bryologische Beitraege 2: 1-115.

HILL M. О., PRESTON C. D. & SMITH A. J. E., 1991 — Atlas of the Bryophytes of Britain and

Ireland. vol. 1 Liverworts ( Hepaticae and Anthocerotae). Colchester, 351p.

PETERSEN P. M., 1972 — Cryptothallus mirabilis Malmb. found on Disko Island, West Greenland

(6091 5" N., 53934" W). Lindbergia 1: 189-190.

POCOCK K. & DUCKETT J. G., 1984 — A comparative ultrastructural analysis of the fungal

endophytes in Cryptothallus mirabilis Malmb. and other British thalloid hepatics. Journal

ы of Bryology 13: 227-233.

SERGIO C., BRUGUES M. & CROS R., 1994 — Duas espécies boreais, encontradas no Parque

Natural da Serra de São Mamede. Lophozia excisa (Dicks.) Dum. nova para Portugal, е

Racomitrium elongatum (Ehrh.) Frisvoll primeira referéncia para o sul do Tejo. In:

SERGIO C., Notulae Bryoflorae Lusitanicae VA. Revista de Biologia (Lisboa) 15: 197-198.

SCHUSTER R. M., 1992 — The Hepaticae and Anthocerotae of North America. Vol. 5. Chicago,

Field Museum of Natural History, 854 p.

Source : MNHN, Paris

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Cryptogamie, Bryol. Lichénol. 1997, 18 (3): 217-221 217

BROMELIOPHILA HELENAE,

A NEW SPECIES OF LEJEUNEACEAE FROM THE NEOTROPICS

S. Rob GRADSTEIN

Systematisch-Geobotanisches Institut, University of Góttingen,

Untere Karspüle 2, D-37073 Góttingen, Germany

SUMMARY — Bromeliophila helenae Gradst. sp. nov. (Lejeuneaceae), described and illustrated,

based on recent collections from the Guayana Highland (Venezuela) and Dominica (Lesser Antilles),

is the second species of this unique neotropical liverwort genus occuring exclusively in the leaf axils of

bromeliads. The new species differs from the type species B. natans (Steph.) Schust. (SE Brazil) by its

rounded leaf apex and rather short leaf cells. The hyaline papilla in the genus appears to be

“proximal”, not “distal” as was suggested previously. The species of Bromeliophila are morphologi-

cally highly variable and some forms are difficult to separate from Lejeunea, The generic status of

Bromeliophila should therefore be considered critical.

Bromeliophila is a unique genus of the Lejeuneaceae that occurs exclusively in

the leaf axils of bromeliads. The genus was described by Schuster in 1994 and contains a

single species, B. natans (Steph.) Schust., endemic to the “Mata Atlantica” of southeas-

tern Brazil. Bromeliophila is related to Lejeunea but is recognized by the leaf lobes that are

usually much longer than wide, having conspicuously elongated cells and a very long

insertion line, and by the long-stalked perianths with 5 double-winged keels. Plants with

shorter, ovate leaves do occur as well, however, and these become almost inseparable from

Lejeunea. Bromeliophila is usually copiously fertile, with gynoecia and numerous slender

male spikes occurring side by side on the stem. The gametoecial arrangement and the long

leaf-insertion are also characteristic of Potamolejeunea and other rheophytic Lejeunea-

ceae, and are probably adaptations to their aquatic habitat.

Among specimens collected on the Macizo del Chimanta in the Guayana

Highland, Venezuela by J. Steyermark and O. Huber in 1983 and sent to me for naming

some ten years ago, is a second species of Bromeliophila. It is very similar to B. natans in

general habit but has rounded leaf apices; in B. natans the leaves are acuminate. Very nice

material of the new species, with sporophytes, was recently collected on the island of

Dominica in the West Indies by A. Scháfer- Verwimp and his wife.

I wish to dedicate the new Bromeliophila to Dr. Héléne Bischler-Causse in

honour of her birthday and as a token of our lasting friendship.

Source : MNHN. Paris

218 S.R. GRADSTEIN

Bromeliophila helenae Gradst., sp. nov. (Fig. 1).

Bromeliophila rotundata Gradst., A Guide to the bryophytes of Tropical America 1.

Liverworts and hornworts (draft version), p. 115, 1995. Brussels, nom. nud.

Type — Venezuela. State of Bolivar, District Piar: Macizo del Chimantá,

"altiplanicie en la base meridional de los farallones superiores del Aparacá-tepui, sector

Norte del Macizo,” 5° 20’ N, 62° 12’ W, ca. 2200 m alt., 30 Jan — 1 Feb 1983, Julian A.

Steyermark, Otto Huber & Victor Carreño E. 128383 (holotype, GOET; isotypes, U,

VEN). Paratype — West Indies. Dominica: “Weg von Laudat zum Valley of Desolation,

in exponierten Bromelientrichtern am Rande des Gratweges oberhalb des Valley of

Desolation, meist submers, 880 m," 24 May 1996, Alfons Scháfer- Verwimp & Inge Verwimp

17887 (СОЕТ, hb. Schäfer-Verwimp).

A B. natans differt foliis apice rotundatis et cellulis foliorum brevioribus.

Description — Plants long and slender, 1.5-2 mm wide, pale yellow-green,

turning dark brownish to black when dry, creeping or floating in water, the older stem

parts often devoid of leaves. Stems flaccid, with hyalodermis; ventral merophyte 2 cells

wide. Leaf lobes wide-spreading, plane or convex, sometimes squarrose, distant to subim-

bricate, highly variable in outline, in some plants narrowly oblong, in others asymmetri-

cally broad-ovate, sometimes both types of leaves on one plant, lobes 0.8-1.3 x 0.35-1.1

mm, 1.2-2 times longer than wide, with a long insertion line (11-16 cells long), apex

rounded, plane or recurved, margins + sinuate to crenulate. Cells in the narrow-oblong

leaves conspicuously elongated throughout the lobe (from base to apex), 50-70 x 25-35 um,

(1.5-)2 times as long as wide, in the broad-ovate leaves shorter, mostly as long as wide,

elongated only towards the base of the lobe, all cells + thin-walled, without trigones,

cuticles smooth; oil bodies unknown; ocelli lacking. Lobules ovate-triangular with a broad

base, small, 1/5-1/3 times lobe length, slightly swollen, insertion line long (ca. 8-10 cells),

keel straight ot slightly curved, forming an almost straight line with the ventral margin of

the lobe, free margin crenulate, with (1-)2(-3) bluntish, 1-celled teeth formed by projecting

cells, the first tooth largest, hyaline papilla inserted at the inner side of the proximal margin

of the first tooth, in a small sinus between the first tooth and the adjacent free margin cell.

Underleaves obovate-orbicular, small, ca. 3 times stem width, 0.25-0.3 mm wide, bifid to

1/3 (-1/2), tips blunt, one-celled, insertion line shallowly curved.

Autoicous, usually copiously fertile, with numerous short-specialized male spi-

kes and a few gynoecia. Male branches tiny, usually shorter than the leaves, with 3-8 pairs

of strongly swollen bracts, each bract with 2 antheridia; bracteoles tiny, limited to the base

of the spike. Gynoecia terminating a short or long branch, with 1-2 lejeuneoid innova-

tions, the innovations usually sterile, female bracts and bracteoles in one series, bracts +

similar to adjacent vegetative leaves but usually with larger lobules, bracteole conspi-

cuously elongated, oblong to ligulate, apex entire or short-bifid with rounded tips, the base

Fig. | — Bromeliophila helenae Gradst. — A. Habit. B. Mid-leaf cells. C. Underleaf. D. Leaf, ventral

view. E. Leaf, dorsal view. F. Leaf apex. G. Lobule, showing first tooth and proximal hyaline papilla.

H. Gynoecium with long-stalked perianth. I. Portion of stem with 2 male branches. All from the type

(drawn by E. M. Reiner-Drehwald).

Source : MNHN Paris

EAS

MNHN. Paris

Source

220 S.R. GRADSTEIN

of the bracteole sometimes connate with a bract on one side. Perianths obpyriform,

usually longly stalked at the base, inflated, 5-keeled, the keels undulate and conspicuously

double-winged, beak short. Sporophytes lejeuneoid, with pale-colored valves that remain

almost erect after dehiscence, valve surfaces with irregular knot-like thickenings in the

middle and upper part, turning slightly brownish in the middle of the valve; elaters 18 per

capsule, 4-5 per valve, attached to upper valve margins, spirals reduced, rudiments of

spiral thickenings becoming more conspicuous and brownish near the free tips of the

elaters. Vegetative reproduction unknown.

Distribution and ecology — Bromeliophila helenae is thus far known only from

Dominica and Venezuela. Its distribution suggests that the species is more widespread in

the Neotropics and has been overlooked. Like the Brazilian B. natans, B. helenae occurs

exclusively in the water-filled leaf axils ("tanks") of Bromeliaceae, in rather open environ-

ments. Often, the species grows submerged. Bromeliophila natans is a lowland species

(0-800 m) and has exclusively been found in terrestrial bromeliads, including Vriesea

glutinosa, Aechmea nudicaulis, and Quesnelia arvensis. Most of the collections are from

open sites in coastal rain forest on sandy soil (“restinga“). Bromeliophila helenae, on the

other hand, is a montane species and has been found associated with terrestrial as well as

epiphytic bromeliads. In the Guayana Highland it occurred submerged in the water-filled

leaf axils of terrestrial Brocchinia hechtioides, in a wet, swampy rock-savanna at ca. 2200

m. On Dominica, the species occurred scattered in terrestrial and epiphytic bromeliads on

a mountain ridge at about 880 m.

Discussion — The most striking feature of Bromeliophila helenae is probably the

variation of leaf shape and cell areolation. In the type specimen the leaf lobes are always

narrowly oblong and the cells are narrowly elongated throughout the leaf. In the material

from Dominica, however, the leaf lobes are mostly broad-ovate and with short cells that

are scarcely longer than wide (elongated only near the base of the lobe). Only some weaker

shoots have elongated leaves and cells as found in the type specimen. A somewhat similar

variation in leaf shape was observed by Schuster (1994) in B. natans. Since elongated leaves

and cells are the main feature separating Bromeliophila from Lejeunea, some doubt ma

be cast on the status of Bromeliophila as a separate genus. This matter requires further

study.

Bromeliophila helenae and B. natans can be distinguished at first glance by their

very diferent leaf apex (rounded versus acuminate). In addition, the leaf cells in B. helenae

seem to be shorter and broader than those in B. natans. In the latter leaf cells are

13.5-16 um wide and up to 4.5 times longer than wide (Schuster, 1.c.). In B. helenae,

however, they are wider, 25-35 шт wide, and at most 2 times longer than wide.

According to Schuster (l.c.), in B. natans the lobule tooth bearing the hyaline

papilla (= first tooth) is situated almost at the keel apex. He therefore interpretes the

position of the hyaline papilla as "distal" (even though the associated tooth is not

reduced). In B. helenae, however, the first tooth is always inserted at some distance from

the keel apex, as usual in the Lejeuneaceae, and the hyaline papilla is clearly “proximal”.

It should be noted that in one of Schuster's figures (Fig. 3-9) a “normal” position of the

first tooth in B. natans, as found in B. helenae, is shown. Apparently, the position of the

first tooth in Bromeliophila is variable. Since the first tooth is never reduced (as is the case

in species of Lejeuneaceae with a "distal" papilla), Bromeliophila should rather be

considered as having a proximal papilla like in Lejeunea.

Source : MNHN. Paris

BROMELIOPHILA HELENAE SP. NOV. 221

ACKNOWLEDGMENT — I am grateful to Dr. E. Reiner-Drehwald for the figures of B. helenae and

for critically reading the manuscript.

REFERENCE

SCHUSTER R. M., 1994 — Studies on Lejeuneaceae, I. Preliminary studies on new genera of

Lejeuneaceae. Journal of the Hattori Botanical Laboratory 75: 211-235.

Source : MNHN. Paris

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Cryptogamie, Bryol. Lichénol. 1997, 18 (3): 223-226 223

LEJEUNEA BISCHLERAE, A NEW SPECIES OF LEJEUNEA

SUBGENUS MICROLEJEUNEA FROM AUSTRALIA

Barbara M. THIERS

New York Botanical Garden, Bronx, NY 10458 U.S.A.

fax: 718-562-6780; email: bthiers@nybg.org

SUMMARY — Lejeunea bischlerae, a new species of Lejeunea subgenus Microlejeunea from Aus-

tralia is described and illustrated. A key to the five species of Lejeunea subgenus Microlejeunea in

Australia is also provided.

Lejeunea sensu lato is a genus of probably at least 200 species, most of which

occur in tropical or subtropical regions of the world. The genus has never been monogra-

phed on a worldwide basis, nor is there widespread agreement on the limits of the genus.

Microlejeunea was accepted as an autonomous genus by authors earlier this century

(Evans, 1900; VandenBerghen, 1948) but the trend in recent times has been to reduce it to

subgeneric status (Schuster, 1980; Mizutani, 1962; Grolle, 1976). The characteristics of the

subgenus were summarized by Schuster (1980) as: a stem cortex of three cells (as viewed in

cross-section); erect to narrowly spreading leaves; the tendency of leaf cells to be small and

have equally thickwalled cells; ocelli (in some species); a leaf lobule that is large relative to.

the leaf lobe, and the tendency for the gynoecial bract keels to be winged.

In the course of work on a treatment of the Lejeuneaceae for the Flora of

Australia project (Thiers, 1990), I have encountered approximately 25 species of Lejeunea,

including an undescribed species of subgenus of Microlejeunea.

Lejeunea bischlerae B. Thiers, sp. nov.

TYPE: AUSTRALIA. Queensland. Yungaburra, Gillies Road, near Mt. Nomico, 8 Sept.

1985, G. A. M. Scott s.n. (holotype: MUCV 6939; isotype: NY).

Latin diagnosis — Ab aliis Lejeunea speciebus apice folii cucullatis, lobis ocellatis,

lobulo magno, cellula magna sinus lobuli necnon androeciis bracteolatis distinguenda.

Description — Monoicous to paroicous. Plants 0.4-0.5 mm wide. Stems pros-

trate, loosely interwoven over substrate. Green to yellow in the dried condition. Stems 50.

um in diam., epidermal cells 7, cortical cells 3. Leaves narrowly spreading, somewhat

uplifted from substrate when dry, distant to imbricate; lobe strongly convex when moist,

orbicular, 0.2-0.3 mm long, 0.2-0.3 mm wide, apex rounded, cucullate, margin entire,

acroscopic base cuneate, basiscopic base straight, incurved (figs. 1, 4, 7); cells isodiametric

Source : MNHN. Paris

Lejeunea bischlerae B. Thiers. 1. Fertile stem sector, ventral view, — 2. Perianth and bracts, dorsal

view. — 3. Underleaf. — 4. Lateral leaf. — 5. Androecium. — 6. Apex of leaf lobule. — 7. Fertile stem

sector, ventral view. — 8. Gynoecial bracteole, — 9. Cells of leaf base showing ocelli. — Figs. 1, 2, 4,

5, 7, 8: standard line = 0.15 mm. Figs. 3, 6, 9: standard line = 30 ит. a = lobule tooth; s= sinus cell

adjacent to tooth; o= ocelli. (All from the type, МОСУ 6939).

Source : MNHN. Paris

LEJEUNEA BISCHLERAE SP. NOV. 225

to oblong, 15-25 um long, 15-30 um wide, walls uniformly thin, cells weakly to strongly

rounded; ocelli present in lateral leaves, forming a basal group, 1-3 per leaf. Lobules

uniform in shape, ovate to dimidiate, 0.2-0.23 mm long, 0.1-0.14 mm wide, as long as leaf

lobe, inflated basally but abruptly flattened above, apex attenuate; free margin sinus

narrowly v-shaped, bordered by 2-3 cells, sinus cell adjacent to tooth 2-3 x larger than

other apical sinus margin cells, free margin tooth 1-celled, oblong to slightly falcate (fig. 6);

lobule keel rounded. Underleaves on vegetative stems distant, oblong, plane, 0.1-0.123 mm

long, 0.075-0.08 mm wide, 2 x stem length, bifid, sinus extending 0.5 underleaf length,

v-shaped, lobes parallel, linear to triangular, lobes 3-4 cells wide at base, apex acute, lateral

margins crenulate, base cuneate, flanking cells not differentiated from other basal cells

(fig. 3).

Androecia terminal, antheridial bracts in 3-6 pairs, bract lobules epistatic,

bracteoles present throughout (fig. 5). Gynoecia restricted to short branches, innovations

present throughout, always single, sterile, leaf sequence lejeuneoid; gynoecial bract lobe

spathulate, 1.5-2 lateral leaf length, apex rounded, gynoecial bract lobe margins entire or

crenate; lobule ligulate, 0.6-0.75 x bract lobe length, apex rounded, gynoecial bract lobule

margins entire to crenulate, keel length 0.75-1 x lobule length, lobule keels winged (wing

1-4 cell wide); gynoecial bracteole oblong, 2-3 x underleaf length, bracteole apex bifid,

sinus extending 0.3 bracteole length, v-shaped, bracteole margins entire to crenulate;

perianths exserted beyond bracts, clavate, circular in cross-section, perianth keels 5, all

keels equally well-defined, extending 0.5 perianth length, keels and perianth surface

smooth.

Paratype — Australia. Queensland. Millaa Millaa, Zillies Falls Rd., 6 Sept.

1985, G. A. M. Scott s.n. (MUCV 7068).

Discussion — Lejeunea bischlerae is a very distinctive species because of its very

large leaf lobules and the cucullate leaf apex. The presence of androecial bracteoles

throughout the androecium is also unusual in subgenus Microlejeunea, although this

feature is characteristic of other subgenera of Lejeunea, namely Apolejeunea and Nanole-

jeunea (Schuster, 1980). In leaf shape, L. bischlerae is reminiscent of Austrolejeunea

bidentata B. Thiers (Thiers, 1985). However, the lateral leaf is very narrowly inserted in A.

bidentata, the lobule bears two teeth on the lobule, and the underleaf lobes are filiform

(i.e., one to two cells wide).

It is a pleasure to name this species for Héléne Bischler, in recognition of her

great contributions to bryology, including work on Australian Microlejeunea (Miller et al.,

1967). It is a pleasure to have her as a colleague and a friend.

Key to Species of Lejeunea subgenus Microlejeunea in Australia

1 — Leaves lanceolate, 2 x longer than wide, apices sharply acute to acuminate ........

vom L. microstipula Steph.

l — Leaves ovate or oblong, 5 x longer than wide, apices broadly acute to rounded.

2 — Cells of lobe midportion and base more or less homogeneous in size and

wall thickness (i.e., no evidence of ocelli).

3 — Lobule tooth acutely pointed, underleaves orbicular, 1.5 x stem

width, sinus extending 0.3 underleaf length, lobes erect, bracts

triangular, gynoecial innovation leaf sequence pycnolejeuneoid. . .

Т Р до MEL ЕБ L. cucullata (Reinw. et al.) Nees

3 — Lobule tooth bluntly pointed, underleaves orbicular to oblong, 2x

Source : ММНМ Paris

226 B.M. THIERS

stem width, sinus extending 0.5 underleaf length, lobes divergent,

narrowly triangular, gynoecial innovation leaf sequence lejeu-

neoid.. L. primordialis (Hook. f. & Tayl.) Gott. et al.

2 — Cells of lobe midportion and base not homogeneous in size and wall

thickness (i.e., ocelli present).

Leaf apex cucullate, lobule as long as leaf, lobule sinus cell

adjacent to tooth 2-3 x larger than other apical sinus

margin cells; androecial bracteoles present throughout

o A L. bischlerae B. Thiers

4 — Leaf apex plane, lobule up to 0.5 lobe length, lobule sinus

cell adjacent to tooth not distinctly larger than other sinus

margin cells; androecial bracteoles at base of androecium

only (1 or 2 bracteoles)......... L. ulicina (Tayl.) Tayl.

Acknowledgments — I gratefully acknowledge the Research Council of the Adelaide Botanic Gar-

dens for a grant that supported my field and herbarium work in Australia in 1984. I thank Roy E.

Halling or his assistance with my fieldwork, and J. Eggers, R. Grolle, G. A. M. Scott, I. Stone and Н

Streimann for making collections of Australian Lejeuneaceae available to me. The curators of FH, G,

JE, L, MELU, MUCY, NICH, and NSW provided loans of specimens critical to this study.

BIBLIOGRAPHY

EVANS A. W, 1900 — The Hawaiian Hepaticae of the tribe Jubuloideae. Transactions of the

Connecticut Academy of Arts and Sciences 10: 387-462.

ОВОШЕ R., 1976 — Verzeichnis der Lebermoose Europas und benachbarter Gebiete. Feddes

Repertorium 86: 71-74.

MILLER H., BONNER C. E. B. & BISCHLER Н. 1967 — Studies in Lejeuneaceae VIII. Microle-

jeunea in Asia and Australia. Nova Hedwigia 14: 61-67.

MIZUTANI M., 1962 — A revision of Japanese Lejeuneaceae, Journal of the Hattori Botanical

Laboratory 24; 115-302.

SCHUSTER R. M., 1980 — Hepaticae and Anthocerotae of North America east of the Hundredth

Meridian. vol. 4. New York, Columbia University Press.

THIERS B. M., 1985 — Austrolejeunea bidentata, a new species of Lejeuneaceae subfamily Tuya-

maelloideae from Australia. The Bryologist 88: 350-352.

THIERS B. M., 1990 — An overview of the Lejeuneaceae in Australia. Tropical Bryology 2: 273-283.

VANDEN BERGHEN C., 1948 — Genera des Lejeuneaceae. Lejeunea Memoirs 6: 1-59.

Source : MNHN. Paris

Cryptogamie, Bryol. Lichénol. 1997, 18 (3): 227-234 227

ORDINAL PLACEMENT OF THE FONTINALACEAE

William R. BUCK! & Bruce ALLEN?

“Institute of Systematic Botany, New York Botanical Garden,

Bronx, NY 10458-5126, U.S.A. E-mail: bbuck@nybg. org

? Missouri Botanical Garden, PO. Box 299,

St. Louis, MO 63166-0299, U.S.A. E-mail: ballen@admin.mobot.org

ABSTRACT — The relationships of the Fontinalaceae have been difficult to discern because of the

extreme morphological modifications to an aquatic habitat undergone by the plants. The family is

shown to possess creeping primary stems with reduced leaves that turn ca. 90? from the substrate to

form the leafy secondary stems. A creeping stem is continued from an adventitous bud near the base

of the secondary stem. This type of branching is indicative of a relationship within the Leucodonta-

les. Stem anatomy of the three genera and paraphyllia in Brachelyma indicate a relationship to the

Climaciaceae.

INTRODUCTION

Fontinalis and its allies, Dichelyma and Brachelyma, have garnered much atten-

tion because of the rarity of truly aquatic mosses. These three genera are held together, in

addition to their habitat preference, by the unusual peristomial feature of a trellised

endostome, otherwise approached only in the Timmiaceae and, to a lesser extent, in the

Prionodontaceae. The exostome teeth are linear with the dorsal (outer) and ventral (inner)

lamellae equally wide. The dorsal lamellae are vertically elongate, but deposition is thicker

on the ventral rather than the dorsal surface with the ventral trabecullae especially heavily

thickened. Both surfaces are papillose rather than horizontally striate. For SEM photo-

graphs see Vitt & Glime (1984). This type of exostome is encountered in a number of

unrelated mosses and also is closely approximated by the “daltoniaceous” exostome type

found in many genera of the Hookeriales. This exostome type curves inward when the

teeth are dry and outward when they are moist (i.e., hygrocastique). The mode of action of

the teeth is therefore opposite that of the “hypnaceous” exostome type which curves

outward when dry and inward when moist (i.e., xerocastique). This functional difference

may be responsible for the parallel derivation in a number of widely separated mosses with

the exostome type found in the Fontinalaceae.

Philibert (1888a, 1888b, see Taylor, 1962) was the first to precisely describe the

morphology of the Fontinalaceae endostome. As in other endostomes it is a two-layered

structure consisting of thickenings and wall remnants deposited by the primary peristo-

mial layer (PPL) on the outer (dorsal side) and by the inner peristomial layer (IPL) on the

inner (ventral side) surfaces of the endostome. Unlike most other endostomes, secondary

Source : MNHN. Paris

228 WR. BUCK & B. ALLEN

thickening is relatively heavy and as a result the endostome is firm rather than membra-

nous. But, secondary deposition is extremely localized and occurs primarily on the outer

surface at and very near the wall pair sutures on the PPL. Deposition on the inner surface

occurs only where the cell walls of the IPL cells are contiguous to the thickenings

deposited by the PPL. Thus, as stated by Philibert, it is PPL that has determined the

direction and location of thickening on the Fontinalaceae endostome. A significant

difference between the Fontinalaceae endostome and that of most other endostomes is its

homogeneous structure, i.e., there is no division of the endostome into the basal mem-

brane and the segment/cilia complex (see Vitt & Glime, 1984, figs. 10 & 11). Asa result of

its localized thickening and its homogeneous structure, the Fontinalaceae endostome

resembles a net consisting of 16 vertical filaments (each corresponding to the median line

on a normal endostomial segment) that are connected by nearly equidistant horizontal

filaments (each corresponding to an anticlinal wall-pair) (see Vitt & Glime, 1984, figs.

10-12). The entire net-like structure is termed a trellis,

When all the wall pair sutures on the PPL of a Fontinalaceae endostome are

secondarily thickened, an unbroken net called a perfect trellis is formed (see Vitt & Glime

1984, figs. 10 & 11). However, when some wall pair sutures lack secondary thickening, an

imperfect trellis results. Whether the trellis is perfect or imperfect, all trellises have 16

secondarily thickened vertical wall pair sutures. Thus, imperfect trellises are formed when

some (as a rule basal) horizontal wall pair sutures are unthickened. Regardless of the

condition of the trellis, the cellular pattern on the PPL can always be determined because

the 16 vertical wall pair sutures are present. This cannot be said for the cellular pattern on

the IPL since its pattern is irregular and reconstructing it depends upon observing wall

fragments deposited on the inside of the horizontal and vertical wall pair sutures. But, if

the endostomes of species with perfect trellises, e.g., Dichelyma falcatum (Hedw.) Myrin

and Fontinalis antipyretica Hedw., are examined it can be seen that the number of cells in

the IPL varies from 32 to 48 or 64, i.e., with 2, 3, or 4 cells between each pair of vertical

filaments. Vastly more important than the number of cells in the IPL is the observation

that the vertical wall pairs on the PPL align with vertical wall pairs on the IPL. Thus, the

vertical filaments of the endostome have median lines on both the front and back surfaces

(see Fig. 1, especially the far left filament). In contrast, most moss endostomes have

vertical wall pairs (a median line) on the PPL that straddle the middle of a single column

of IPL cells. This offsetting of the IPL cellular pattern so that some IPL vertical wall pairs

line up with the median line on the PPL, although uncommon, is also encountered in

Funaria, Encalypta, Splachnum, Orthotrichum, Mittenia (see Edwards, 1984, fig. 7) and

Timmia (see Shaw & Rohrer, 1984, figs. 15 & 17). The combination, however, of an

offsetting IPL cellular pattern, and a firm endostome having a homogeneous structure

consisting of vertical filaments connected by equidistant horizontal filaments makes the

Fontinalaceae endostome unique within the mosses.

Taxonomic work on the group has focused primarily at the species level, and

little attention has been given to relationships of the family. It was first described by

Schimper (1855) to include Fontinalis and Dichelyma. Shortly thereafter he (Schimper,

1860) segregated Dichelyma (and the North American Brachelyma) into a separate family,

but retained all three genera in the same “Tribus” as the Fontinalaceae. No relationships

were discussed, but the tribe was the first one listed within the pleurocarps and immedia-

tely preceded the Cryphaeaceae (included in a tribe with the Leptodontaceae, Neckera-

сеае, and Leucodontaceae).

In 1879 Lindberg placed Fontinalis and Dichelyma with Climacium in the subfa-

mily Meteorieae within the Neckeraceae. No explanation was provided, but this is the

Source : MNHN, Paris

ORDINAL PLACEMENT OF THE FONTINALACEAE 229

Fig. 1. — Portion of the endostome of Dichelyma falcatum (Allen, Fontinalaceae Exsiccatae 3, MO)

viewed with a light microscope and showing cellular pattern of the IPL (dotted lines) superimposed

on the cellular pattern of the PPL (solid lines). Scale bar = 0.05 mm.

origin of the association of the Fontinalaceae with the Climaciaceae. Brotherus (1905) in

the first edition of Die natürlichen Pflanzenfamilien, followed Lindberg in claiming a

relationship between the Fontinalaceae and Climacium. The reasons he provided are

3-sided branches as viewed in transverse section, decurrent leaves with lax, hyaline

thin-walled alar cells and thin-walled, elongate laminal cells, brown-red exostome teeth

with papillose dorsal plates, low basal membrane, segments perforate ladder-like, and the

long, somewhat twisted calyptra covering the whole capsule.

Subsequently Fleischer (1908) followed Brotherus' (1905) lead, but offered no

new information. However, since neither the Fontinalaceae nor the Climaciaceae occur in

Java it is not surprising that Fleischer treated the two in a superficial manner. In the second

edition of Die natürlichen Pflanzenfamilien Brotherus (1925) went a step further and

Source : MNHN. Paris

230 WR. BUCK & B. ALLEN

placed the Fontinalaceae and the Climaciaceae in their own suborder, nestled between the

Racopilaceae and the Hedwigiaceae. However, such a placement engenders a certain

amount of distrust because today both the Racopilaceae and Hedwigiaceae are conside-

red to be acrocarpous (Buck & Vitt, 1986; De Luna García, 1992).

The acceptance of a relationship between the Fontinalaceae and Climaceae has

not been universal. Andrews (1954: 261) was very blunt with his dissention. He argued for

a possible relationship for the Climaciaceae with the Hylocomiaceae because “Any

possible indication of a closer relationship of the Climaciaceae to anything should be

welcome, as the current idea that it stands near the Fontinalaceae has always seemed to the

present writer the height of the impossible." Following Andrews, Crum and Anderson

(1981) placed Climacium near the Hylocomiaceae (Hypnales) but kept the Fontinalaceae

within the Leucodontales between the Leucodontaceae and the Pterobryaceae.

The composition of the Climaciaceae has been called into question and is

beyond the scope of this paper. However, we agree with Ireland (1968) that Pleuroziopsis

is not very closely related to Climacium. However, it should be noted that Pleuroziopsis

also has a leucodontalean branching pattern, and despite the horizontal capsule and

cross-striolate exostome teeth, it is probably not properly placed in the Hypnales. A similar

situation can be cited in the Neckeraceae where Porothamnium has cross-striolate exos-

tome teeth.

Most recently Buck & Vitt (1986) placed the Fontinalaceae in the Hypnales in its

own superfamily. Their justification was linear leaf cells, well developed alar cells, and an

interpretation of the endostomial nodal projections as amplified lateral walls of the IPL,

as well as a north temperate distribution.

Curiously, monographers of the Fontinalaceae have not addressed the problems

of the family's relationships. Cardot (1892), followed by Welch (1960), only treated the

taxa within the family. Cardot, in addition to the three genera discussed here, Fontinalis,

Dichelyma, and Brachelyma, also included Hydropogon, Hydropogonella (as Cryptan-

gium), and Wardia in the family. Welch (1943), excluded the latter three genera into the

Hydropogonaceae and the monogeneric Wardiaceae.

What kinds of evidence might we look for in determining the relationships of the

Fontinalaceae to other families?

ORDINAL PLACEMENT

Recently Buck (1997) found that the character that best binds the families of the

Leucodontales into a coherent group is branching pattern. Basically, there is a creeping

primary stem with reduced, often hyaline leaves. The primary stem eventually turns about

90* from the substrate and differentiates into an erect, leafy, often branched, upright stem

(the secondary stem). Subsequently an adventitious bud forms near the bend of the stem,

elongates and continues the creeping stem. This branching pattern would best be defined

as sympodial sensu La Farge-England (1996). Every family of Leucodontales examined

has this branching pattern although not every genus of every family does. In contrast, the

Hypnales show a creeping stem with or without differentiated leaves, but never with

reduced, hyaline leaves. Most often the leaves of the creeping stem are larger than those of

subsequent levels of branching. Branches are formed laterally from branch primordia or

from buds in leaf axils. The creeping stem is more or less indeterminate. This pattern is

referred to as monopodial (La Farge-England, 1996).

Source : MNHN, Paris

ORDINAL PLACEMENT OF THE FONTINALACEAE 231

| YA

Fig. 2-8. — Diagramatic sketches of stems of Fontinalaceae. — Fig. 2-3. — Fontinalis duriaei

(Buck 30061). Fig. 2. Branching pattern. Fig. 3. Portion of transverse section of creeping stem. — Fig.

-— Fontinalis novae-angliae var. latifolia (Buck 30113), branching pattern. — Fig. 5-6. — Fontinalis

welchiana. Fig. 5. Branching pattern (Buck 30055). Fig. 6. Branching pattern (Buck 30059). — Fig. 7.

— Brachelyma subulatum (Buck 30083), branching pattern. — Fig. 8. — Dichelyma capillaceum

(Buck 30088), branching pattern. All specimens at NY.

Source : MNHN, Paris

232 WR. BUCK & B. ALLEN

Fontinalis, and to a lesser extent Dichelyma and Brachelyma, live in environments

which require that plants be securely anchored to their substrates in order to prevent being

swept away by water currents. As a consequence, herbarium material of Fontinalaceae

rarely includes creeping stems because substrate is rarely collected, especially that which

seems to support what appears to be juvenile plants. Therefore, we made a special field trip

into southeastern Missouri, U.S.A. in late March-early April, 1996 specifically to collect

Fontinalaceae still attached to their substrates. Dichelyma capillaceum (With.) Hartm.,

Brachelyma subulatum (P. Beauv.) Card., and four species of Fontinalis, Е. duriaei Schimp.,

F. novae-angliae Sull. var. latifolia Card., F. sullivantii Lindb., and F. welchiana B. Allen,

were collected, all of which but F. sullivantii provided suitable material for study. In the

laboratory, material was rehydrated and individual stems were removed from the substra-

tes (usually rock but sometimes wood) under the dissecting microscope.

In every specimen examined (Figs. 2, 4-8), typical leucodontalean branching was

found. In the cases where typical hynalean branching was exclusively found, the creeping

stems did not have reduced leaves and were interpreted as secondary stems that were not

erect. Sometimes (e.g., Figs. 5-7) hynalean branches arose from leucodontalean creeping

stems. On creeping primary stems of the leucodontalean type, leaves were widely spaced,

hyaline, and greatly reduced, usually less than the length of typical leaves.

Transverse sections of the primary stems were particularly interesting. In the

Fontinalaceae there is no central strand. In the upright shoots the relatively large, thin- to

firm-walled interior cells are surrounded by 2-4 rows of small, thick-walled cells, the

thickest being the outermost. However, in the creeping stem the outermost row of cells are

somewhat thinner-walled than the rows immediately subtending them. Each of these

epidermal cells seems to be capable of becoming a rhizoidal initial (Fig. 3). As a conse-

quence, rhizoids can be found all around the stems for their entire lengths. Only after the

stems bend and become secondary stems does the anatomy change and the rhizoids

disappear. The ability to form rhizoids along the length of the stems is also a character of

the Leucodontales. In the Hypnales rhizoids typically are formed in discrete patches.

Axillary hairs of all material were also examined. In every case the hairs had 1-2

quadrate to short-rectangular, brown basal cells and (3-)4-12 hyaline apical cells. The

distinguishing feature of the axillary hairs, though, was the length of the hairs. They

ranged from 350 um to 860 um long. These elongate axillary hairs are yet another

leucodontalean feature. In the Leucodontales axillary hairs are often very elongate,

whereas in the Hypnales (and Hookeriales) they are typically quite short.

Therefore, branching pattern, rhizoid-associated stem anatomy, and axillary

hairs all point toward a position within the Leucodontales for the Fontinalaceae.

FAMILIAL ALLIANCES

The Fontinalaceae are highly modified in response to their specialized habitats.

Therefore, many of their unique features (synapomorphies), such as the trellised endos-

tome, are of little use in assessing possible relationships to other mosses. Rather, we need

to look for other kinds of characters.

Recently Allen (1995) noted the presence of paraphyllia in Brachelyma.

Although his illustration (fig. 1, no. 4) and even his discussion might lead one to think the

structures to which he referred are pseudoparaphyllia, this is not the case. They are not

clustered around branch primorida but rather are scattered along the stems. Curiously

Source : MNHN, Paris

ORDINAL PLACEMENT OF THE FONTINALACEAE 233

they also occur on decurrent leaf bases. Within the Leucodontales paraphyllia are rare and

might be suggestive of a relationship.

The stem anatomy where every epidermal cell can become a rhizoidal initial is

also unusual. If another group could be found with such a feature it could be significant.

Within the Leucodontales branching of the upright, secondary stems usually

results in a frondose-stipitate habit. In other words, branching only occurs in a single plane

and not all around the stem. True dendroid branching is relatively rare. However, in the

Fontinalaceae branches emerge from all sides of the stem. Not uncommonly the branches

bend shortly thereafter in order for the plant to be more or less flat, presumably in order to

be more streamlined in the water, but the branches truly do emerge from all sides of the

stem.

The only genus found in the Leucodontales which combines paraphyllia, the

distinctive stem anatomy, and a dendroid habit is Climacium. Interestingly, this conclusion

is based on evidence entirely independent from those features that convinced Brotherus

(1905) of a relationship. Among those characteristics listed by Brotherus, the long, twisted

calyptra covering the entire capsule seems most significant.

In Climacium the creeping stem is typically leucodontalean, with the primary

stem becoming an erect, secondary stem, and then a new primary stem originating from an

adventitious primordium at the base of the secondary stem. The secondary stem is

dendroidly branched.

In transverse section the primary stem is similar to that of the Fontinalaceae. In

common with the Fontinalaceae every cell can become a rhizoidal initial and rhizoids

emerge from all sides, all along the stem. The difference is that in the Fontinalaceae the

epidermal cell itself elongates so that each rhizoid is more or less embedded in the stem

(Fig. 3). In Climacium the rhizoids arise from atop the epidermis with the epidermal cells

themselves not elongating. This difference may reflect the greater need of the Fontinala-

ceae for secure attachment to the substrate.

Although the peristome of Climacium has some similarity to that found in the

Fontinalaceae, overall it is significantly different. As in the Fontinalaceae, the exostome

teeth are dark red and papillose rather than striate. However, the outer lamellae are

narrower, relatively wider and more heavily thickened than the inner lamellae, the inner

trabeculae are weakly thickened, and the teeth are relatively broad at base. The endostome,

as in the Fontinalaceae, is dark red to orange, firm, and often united at the tips of the

segments. But, unlike the Fontinalaceae endostome, that of Climacium is differentiated

into 16 segments (cilia are absent) and a low basal membrane. The segments are broadly

perforate and there are 32 cells in the IPL. Most significantly, the IPL cells are not offset so

that, as in most other mosses, the median line on the segments straddles a single column of

IPL cells.

In conclusion, despite peristomial dissimilarities, the original speculation of

Lindberg (1879) that the Fontinalaceae are related to Climacium still seems to be the most

reasonable phylogenetic hypothesis. At least, a relationship between the two is the most

logical based on morphological evidence. This problem seems an ideal one for molecular

investigation. Although molecular data can also be deceiving, sometimes it can resolve

problems where morphological modification has been extreme.

Source : MNHN. Paris

234 WR. ВОСК & B. ALLEN

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SCHIMPER W. P., 1855 [1856] — Corollarium Bryologiae Europaeae, conspectum diagnosticum

familiarum, generum et specierum, adnotationes novas atque emendationes complectens.

Stuttgartiae, E. Schweizerbart, 140 p.

SCHIMPER W. P., 1860 — Synopsis Muscorum europaeorum praemissa introductione de elementis.

bryologicis tractante. Stuttgartiae, E. Schweizerbart, 735 p.

SHAW J. & ROHRER J. R., 1984 — Endostomial architecture in diplolepideous mosses. Journal of

the Hattori Botanical Laboratory 57: 41-61.

TAYLOR E. C. SR., 1962 — The Philibert peristome articles. An abridged translation. The

Bryologist 62: 175-212.

VITT D. H. & GLIME J. M., 1984 — The structural adaptations of aquatic Musci. Lindbergia 10:

95-110.

WELCH W. H., 1943 — The systematic position of the genera Wardia, Hydropogon, and Hydropo-

gonella. The Bryologist 46: 25-46.

WELCH WH., 1960 — А monograph of the Fontinalaceae. The Hague, Martinus Nijhoff, 357 p.

ае and Hylocomiaceae.

Source : MNHN, Paris

Cryptogamie, Bryol. Lichénol. 1997, 18 (3): 235-242 235

WELCHE FUNKTION HABEN DIE HYALOCYTEN

IN DEN BLATTERN DER DICRANACEAE ?

Jan-Peter FRAHM

Botanisches Institut der Rheinischen Friedrich-Wilhelms Universität, Meckenheimer Allee

170, D-53115 Bonn, Germany, e-mail frahm@uni-bonn.de

ABSTRACT — The function of the hyalocysts of Leucobryum and other Dicranaceae (Paraleuco-

bryum, Campylopus) is still dubious. As possible explanations, (1) water storage, (2) protection of the

enclosed chlorocysts against desiccation, (3) a rôle for gas exchange, (4) a rôle for nutrient uptake and

(5) an increased evaporation for the concentration of nutrients are discussed.

The fact that predominantly species of wet habitats possess hyalocysts, which do not need to

store water, seems to indicate that water storing is not the main function of this anatomical structure.

The observation that species with hyalocysts of mesic habitats compete with species without hyalo-

cysts without having an advantage seems to indicate that such structures were evolved under different

ecological conditions than today.

Protection of the enclosed chlorocysts against desiccation works only for a short period

when the water stored in the hyalocysts is already evaporated but the chlorocysts are not yet dried out.

It is therefore also not a sufficient explanation for its function. Air bubbles observed in the hyalocysts

are not a result of photosynthesis as suggested in the literature but a result of dehydration.

Cation exchange was so far regarded as a method for nutrient uptake typical for Sphagnum.

Cation exchange experiments with Sphagnum palustre, 5. fimbriatum, Leucobryum glaucum, Dicra-

num scoparium and Mnium hornum revealed that all these species have about the same cation exchange

capacity regardless their different structures of their leaves. Thus the special anatomy of Sphagmum

leaves does not increase the cation exchange.

Evaporation of a Leucobryum tuft is not higher than that of a water table. However, the

amount of water stored is 2.3 times higher than that of Mnium hornum and the desiccation takes 3

times as long. During desiccation, the leaves undergo three phases with the hyalocysts filled with

water, with air bubbles and with air, which provide different conditions for gas exchange. It is not yet

known, whether the prolongation of the period of desiccation balances or succeeds the net photo-

synthesis reduced by the diffusion of gas through the water filled hyalocysts as compared with other

mosses.

The function of hyalocysts may perhaps be explained as a combination of the different

explanations mentioned above. The enormous water uptake made possible by the hyalocysts princi-

pally extends the time of desiccation by factor three. Photosynthesis is reduced as long as the

hyalocysts are filled with water but is suboptimal when the hyalocysts are filled with air bubbles and

optimal when the empty hyalocysts protect the chlorocysts from desiccation. This could finally result

in a higher net-photosynthesis as compared with other mosses growing in the same habitat.

ZUSAMMENFASSUNG — Die Funktion der Hyalocyten in den Blatter von Leucobryum und

anderen Dicranaceen (Paraleucobryum, Campylopus) ist weitgehend unklar. Als mégliche Erklárun-

gen werden (1) Wasserspeicherung, (2) Austrocknungsschutz, (3) eine Rolle fiir den Gasaustausch, (4)

Source : MNHN, Paris

236 J.-P. FRAHM

eine Rolle für die Náhrstoffaufnahme und (5) eine gesteigerte Verdunstung zur Konzentration der

Nährstoffe diskutiert.

EINLEITUNG

Hyalocyten sind vergrößerte tote Zellen, Sie finden sich in der Stengelrinde

(Hyalodermis) von einigen aber nicht allen Arten der Gattungen Sphagnum, Drepanocla-

dus, Hypnum und anderen Laubmoosgattungen. Sie finden sich zudem in den Bláttern der

Sphagnaceae, den Blattbasen von Encalyptaceae, Calymperaceae und einigen Pottiaceae

und in der Rippe von Dicranaceae (Campylopus z.Tl., Paraleucobryum und speziell den

Leucobryoideae).

Hyalocyten kónnen dabei in einigen systematischen Gruppen perforiert sein. So

sind die Hyalocyten von einem Teil der Sphagnen, von Calymperaceae, Encalyptaceae

oder Tortula in den áuBeren Zellwánden, die der Leucobryaceae in den inneren Zellwán-

den perforiert.

Hyalocyten sind phylogenetisch offenbar schon eine sehr alte Struktur, die

bereits in Moosen des Paläozoikums vorhanden war. Smoot & Taylor (1986) publizierten

Photographien von Gesteinsdünnschliffen aus dem Perm der Antarktis, die Querschnitte

von breiten Rippen mit mehrschichtigen Hyalocyten zeigen.

Obgleich Hyalocyten ein auffálliges Element in der Blattanatomie sind, ist ihre

Funktion weitgehend unklar. Manche Autoren von Lehrbüchern (van der Wijk, 1932;

Watson, 1978; Schofield, 1985) geben nur morphologische Beschreibungen aber keine

Erklárungen der Funktion. Und in dem jüngsten und wohl umfangreichsten Kompen-

dium zur Bryologie, dem *New Manual of Bryology" (Schuster, 1983), ist die Physiologie

oder funktionelle Morphologie der Bryophyten (welche der wesentlichste Aspekt zum

Verstándnis dieser Pflanzengruppe ist) nicht einmal eingeschlossen. Daher sollen hier

einige Hypothesen zur Funktion dieser Hyalocyten diskutiert werden.

HYPOTHESEN ZUR FUNKTION DR HYALOCYTEN

1. Wasserspeicherung

Bereits Goebel (1915) bemerkte, daB die Calymperaceae und Leucobryaceae

besonders typisch für die tropischen Tieflánder sein und daB daher Einrichtungen zur

Wasserspeicherung an einem permanent feuchtem Standort ohne VerdunstungsstreB

keinen Sinn machen. Goebel (1915) führte ferner an, daB die Hyalocyten die Chlorocyten

vor der Austrocknung schützen. Das wiederum macht aber physiologisch keinen Sinn, da

die von Hyalocyten eingeschlossenen Chlorocyten den Gasaustausch herabsetzen.

Wie bereits Goebel (1915) bemerkte, kommen Hyalocyten sowohl in Moosen

von feuchten als auch von trockenen Standorten vor. Das scheint anzudeuten, daB die

Funktion bei den Pflanzen beider Standorte nicht dieselbe ist. Torfmoose sind Beispiele

für Moose mit Hyalocyten von nassen Standorten. Bei ihnen muB bezweifelt werden, ob

Source : MNHN. Paris

FUNKTION DER HYALOCYTEN 237

Hyalocyten wirklich Mechanismen für die Wasserspeicherung sind, da diese Arten

teilweise im Wasser wachsen, keine Wasserdefizite haben und auch (vielleicht mit Ausna-

hme von auf Waldboden wachsenden Arten) auch keiner Austrocknung unterliegen

Daher erscheint eine Wasserspeichereinrichtung bei Arten nasser Standorte paradox zu

sein.

Auch bei Campylopus sind die Arten mit den größten ventralen Hyalocyten (z.B.

Campylopus pittieri Williams, C. albidovirens Herz., C. cavifolius Mitt., C. shawii Wils. in

Hunt.) charakteristisch für nasse oder moorige Standorte. Im Gegenteil dazu besitzen

Arten von trockenen Standorten, die eine Wasserspeicherung nótig hátten, keine Hyalo-

cyten, wie z .B. die wenigen epiphytischen Arten in dieser Gattung. Bei Paaren nah

verwandter Arten, von denen die eine Art epigáisch lebt, die andere epiphytisch (z.B. C.

cuspidatus (Hornsch.) Mitt. — C. valerioi Allen , C. andersonii (C. Müll.) Jaeg. — C.

atlanticus Allen , C. flexuosus (Hedw.) Brid. — C. flagelliferus (C. Müll.) Jaeg.), hat die

epiphytische Art ventrale Stereiden, die epigäische aber ventrale Hyalocyten.

Vielfach kommen auch Arten mit und ohne Hyalocyten in der Natur an dem-

selben Standort vor (z.B. in Mitteleuropa Paraleucobryum longifolium (Hedw.) Loeske and

Dicranum fulvum Hook. auf Silikatblócken, Leucobryum glaucum (Hedw.) Aongstr. und

Campylopus flexuosus (Hedw.) Brid. auf stark sauren Waldbóden). In diesen Fällen

konkurrieren die Arten mit und ohne Hyalocyten im Rippenquerschnitt an demselben

Standort, ohne daß eine Art von beiden offensichtlich einen Vorteil von seiner Anatomie

hätte. Das könnte darauf schließen, daß solche anatomischen Strukturen wie Hyalocyten

sich unter anderen ökologischen Bedingungen gebildet haben und daß sie unter den

heutigen ökologischen Bedingungen keinen Vorteil mehr bilden.

2. Schutz der Chlorocyten

Goebel (1915) argumentierte, daß viele der Leucobryaceen in den Tropen Epi-

phyten sind. Ihre weißliche Farbe, die in manchen Gattungsnahmen (Leucobryum, Leu-

cophanes) und Artnamen (albidum) ihren Ausdruck findet, soll nach Goebel belegen, daß

die Hyalocyten in der Regel nicht wassergefüllt sondern luftgefüllt sind. Dadurch sollen

die Hyalocyten als Schutz für die Chlorocyten gegen Austrocknung und als Isolation

gegen Insolation und die dadurch hervorgerufene Aufhitzung dienen. Diese Hypothese

scheint dadurch gestützt zu werden, daß die Hyalocyten der Leucobryaceen im Gegensatz

zu den Sphagnen keine Poren in den äußeren Zellwánden besitzen.

3. Gasstoffwechsel

Robinson (1985, 1990) interpretierte die Rippenstruktur der Leucobryaceae

unter einem neuen stoffwechselphysiologischen Aspekt. Er führte aus, дай die Hyalocyten

der Leucobryaceae nicht nur Wasser sondern auch Gasblasen enthielten und schrieb

dazu:: “the characteristic leucocyst-chlorocyst cellular pattern in the moss family Leuco-

bryaceae depends functionally on the presence of air inside the leaf. Such leaves prove

basically different in structure and function from those of other mosses, including the

leucocyst-containing Sphagnum”. Diese Hypothese schien so interessant und neu zu sein,

daB sie den Anlaß zu der vorliegenden Studie gab. Nach Robinson kann man davon

ausgehen, daß die Hyalocyten eine spezielle Funktion für den Gasaustausch haben.

Source : MNHN Paris

238 J.-P. FRAHM

Dadurch wird suggeriert, daB die Hyalocyten über Tag mit O, als Produkt des Gasstof-

fwechsels gefüllt werden und nachts CO, aufnehmen und speichern und somit einen

Gaspuffer bilden.

Um diese Hypothese zu testen wurden Polstern von Leucobryum glaucum mit

Wasser gesättigt und in einer Plastikdose direktem Sonnenlicht ausgesetzt. Zur Kontrolle

der Gasblasen wurden Blátter entnommen und direkt unter dem Mikroskop studiert ohne

zusützliches Wasser zuzugeben. Es zeigte sich, daB die Blatter trotz starker Nettophoto-

synthese keine Gasblasen in den Hyalocyten enthielten. Auch wenn das PlastikgefáB mit

einer Aluminiumfolie umwickelt wurde und die Pflanzen 16 Stunden lang im Dunkeln

gehalten wurden, zeigten sich keine Gasblasen in den Hyalocyten. Gasblasen wurden erst

gebildet, wenn die Pflanzen abtrockneten. Damit ist erwiesen, daß Gasblasen sich nicht

durch den Gasstoffwechsel bilden sondern erst wenn das verdunstete Wasser in den

Hyalocyten durch Luft ersetzt wird. In diesem Stadium kónnen sie allerdings den Gass-

toffwechsel erleichtern.

4. Nührstoffaufnahme

Auffälligerweise sind alle Dicranaceae mit Hyalocyten in den Rippen (wie auch

Torfmoose) starke Acidophyten, die an nährstoffarmen Standorten wachsen. Hyalocyten

würden — ähnlich wie bei Torfmoosen — die für die Nahrstoffaufnahme zur Verfügung

stehende Oberfläche vergrößern und in eine effektivere Nährstoffaufnahme ermöglichen.

Basierend auf der Beobachtung, daß Sphagnum-Arten das 20-30fache ihres Trocken-

gewichtes an Wasser aufnehmen können, postulierte erstmalig Sachs (zitiert nach Goebel,

1915), daß Torfmoose in der Lage sein müßten, größere Nährstoffmengen aus dem

nährstoffarmen Milieu aufzunehmen. Das betrifft insbesondere Sphagnum-Arten von

Hochmooren, die fast ausschließlich von Regenwasser als Nährstoffquelle abhängig sind.

Im Prinzip betrifft die Abhängigkeit von der Nährstoffversorgung aus dem Regen jedoch

auch die Epiphyten, von denen speziell Leucobryaceae und Calymperaceae über Hyalo-

cyten verfügen. Jedoch besitzen die meisten epiphytischen Moose keine Hyalocyten und

kommen damit gut aus. Ferner besitzen die Hyalocyten von Torfmoosen Poren in den

äußeren Zellwánden, die der Leucobryaceen jedoch in den inneren Zellwänden, was an

eine unterschiedliche Funktion denken läßt. Daß die Poren der Sphagnen zur Wasse-

raufnahme dienen, wird daraus ersichtlich, daB die Poren bei aquatischen Modifikationen

reduziert werden oder ganz fehlen.

Wenn die Hyalocyten von Leucobryum zur Nährstoffaufnahme áhnlich wie bei

Sphagnum dienen, müßte der Kationenaustausch von beiden auch ähnlich sein. Um diese

Hypothese zu testen, wurden Ionenaustauschversuche mit Leucobryum glaucum (Hedw.)

Aongstr. durchgeführt. Zum Vergleich wurde Sphagnum fimbriatum Wils. und 5. palustre

L. und als Vertreter von Moosen ohne Hyalocyten Dicranum scoparium Hedw. und

Mnium hornum Hedw. herangezogen.

Für den Ionenaustauschversuch wurde Material der genannten Arten in einem

Wald nahe Eupen (Belgien) gesammelt. Für den Versuch wurden nur die lebenden oberen

Pflanzenteile benutzt. Das Pflanzenmaterial wurde bei 50° C 24 Stunden getrocknet,

gewogen und in ein Glasrohr gefüllt. Durch das Glasrohr wurde 0.005mol CaCl,.-Lésung

gegossen. Dabei wurde darauf geachtet, daB der DurchfluB immer etwa dieselbe Zeit

(150-160 sec.) dauerte. In der durchgeflossenen Lósung wurden pH und Leitfáhigkeit

gemessen und titrimetrisch Kalzium bestimmt. Die Ergebnisse wurden auf das Trocken-

gewicht bezogen.

Source : MNHN. Paris

FUNKTION DER HYALOCYTEN 239

Überraschenderweise war der Kationenaustausch bei allen untersuchten Arten etwa

derselbe ungeachtet der unterschiedlichen anatomischen Strukturen ihrer Blátter (Tab. 1).

Aqua CaCl, Sphagnum Sphagnum Leuco- Dicranum Mnium

dest. palustre fim- bryum scopa- hornum

briatum — glaucum. rium

Gewicht(g) 3,36 422 10,26 5,36 547

pH 7,25 6,24 3,56 3,64 3,42 3,35 3,58

Leitfähigkeit 1,54 1058 933 962 903 984 939

(uS)

absorbiertes Ca 8,62 9,81 545 10,10 9,00

(mg)

absorbiertes Ca 256 232 53 187 164

(mg/100g)

Tab. 1 — Ergebnisse von Ionenaustauschversuchen von Moosen mit und ohne Hyalocyten. / Results

of cation exchange experiments with mosses without and with hyalocysts. ]

Der pH wurde von urspriinglich 7.25 bei Sphagnum palustre auf 3,56, bei S.

fimbriatum auf 3,64, bei Leucobryum glaucum auf 3,42, bei Dicranum scoparium auf 3,35

und bei Mnium hornum auf 3,58 gesenkt. Die Leitfähigkeit sank von ursprünglich 1058 uS

auf 933 uS bei Sphagnum palustre, 962 uS bei S. fimbriatum, 903 uS bei Leucobryum

glaucum, 984 uS bei Dicranum scoparium und 939 uS bei Mnium hornum. Die Kalzium

aufnahme betrug 8.62 mg bei Sphagnum palustre, 9.81 mg bei S. fimbriatum, 5.45 mg bei

Leucobryum glaucum, 10.1 mg bei Dicranum scoparium und 9 mg bei Mnium hornum.

Auffällig ist, daß die Kalziumaufnahme bei Leucobryum trotz des Vorhandenseins von

Hyalocyten am geringsten ist und nicht nur geringer als bei Sphagnen sondern auch

geringer als bei Mnium hornum und Dicranum scoparium.

Die Ergebnisse stimmen mit denen von Clymo (1963) überein, nachdem der

höchste Ionenaustausch abgesehen von Sphagnum bei Mnium hornum zu verzeichnen war,

der geringste bei dem (partiell endohydrischen) Polytrichum commune Hedw. Es ist zu

berücksichtigen, daß die für den Ionenaustauschversuch verwendeten Sphagnen aus

einem Wald stammen, da nach Clymo (1963) die Ionenaustauschkapazitát bei Hoch-

moorsphagnen am größten ist. Dennoch ist festzuhalten, daB der Ionenaustausch bei

Sphagnum im Verháltnis zu der durch die Hyalocyten viel gróBeren Oberfláche nicht sehr

wesentlich größer ist als bei Leucobryum mit Hyalocyten, aber ohne äußere Poren, und bei

Dicranum und Mnium ohne Hyalocyten. Eine Verbesserung der Nährstoffaufnahme

durch Hyalocyten ist daher keine ausreichende Erklárung für ihre Funktion.

5. Verdunstung

Die Speicherung von gróBeren Wassermengen kónnte auch zu einer gróBeren

Verdunstung führen. Wenn nährstoffarmes Wasser (wie z.B. Regenwasser) aufgenommen

wird, so ist die Náhrstoffkonzentration zunáchst sehr niedrig. Die Konzentration würde

durch Verdunstung steigen.

Source : MNHN. Paris

240 J.-P. FRAHM

—*—— Mnium

70%

= %

3 60 4 z a

g 50 7 à

E 40 A 2 B

à 30 + >

| = н п o = п m =

20 4 9,

B а

10 + D

гяазвеззезвзвв

110

120

150

140

150

hours

Fig. | — Austrocknung von Polstern von Mnium hornum und Leucobryum glaucum. | Desiccation of

tufts of Mnium hornum and Leucobryum glaucum. J

Um die unterschiedliche Wasserabgabe von Moosarten mit und ohne Hyalocy-

ten zu testen, wurde die Verdunstung von Polstern von Leucobryum glaucum im Vergleich

zu Mnium hornum bei Raumklima (21? C + 1% and 45 % rH) gemessen (Fig. 1). Mnium

hornum trocknete innerhalb 54 Stunden vóllig aus und reduzierte dabei sein Gewicht auf

26%. Leucobryum brauchte hingegen 93 Stunden um 26% seines Nafgewichtes zu

erreichen und war erst nach 142 Stunden vóllig ausgetrocknet. Zu diesem Zeitpunkt

betrug das Trockengewicht nur noch 12 % des NaBgewichtes. Die Wasserspeicherkapazi-

tüt von Mnium hornum betrug das Dreifache des Trockengewichtes, was der Mittelwert für

die meisten Moose ohne spezielle Wasserspeichereinrichtungen darstellt, da derselbe

Faktor auch bei Untersuchungen zur Wasserspeicherkapazitát von tropischen

Regenwaldmoosen gemessen wurde (Frahm, 1990). Leucobryum glaucum speichert hin-

gegen das Siebenfache des Trockengewichtes und verdunstet die Feuchtigkeit in der

dreimal längeren Zeit. Daraus geht hervor, daß eine hohe Wasserspeicherkapazität, wie sie

bei Moosen mit Hyalocyten gegeben ist, nicht auch gleichzeitig eine hohe Verdunstung

einschlieBt, da diese über eine lángere Zeit geht.

Erst durch Gasstoffwechselversuche kónnte getestet werden, ob die Verlánge-

rung der turgeszenten Phase durch eine langsame Austrocknung nach der Befeuchtung

von Leucobryum (also z.B. nach einem Regenfall) in einer hóhere Nettophotosynthese

resultiert, welche die Reduktion der Photosynthese durch die Diffusion von CO, durch die

Source : MNHN, Paris

FUNKTION DER HYALOCYTEN 241

wassergefüllten Hyalocyten wieder ausgleichen würde. Bei der Befeuchtung und an-

schlieBenden Austrocknung von Leucobryum Blátter konnten drei unterschiedliche Pha-

sen beobachtet werden:

1. Die Blátter sind mit Wasser vollgesogen und die Hyalocyten sind mit Wasser gefüllt. Die

Photosynthese ist durch die Lage der Chlorocyten in der Mitte der Rippe, eingeschlossen

durch die wassergefüllten Hyalocyten, eingeschránkt.

2. Steigende Austrocknung führt zu einer Verdunstung in den Hyalocyten. Das Wasser in

den Hyalocyten wird durch Luftblasen ersetzt, die sich allmählich vergrößern. Es kann

angenommen werden, daB die Photosynthese durch bessere Verfügbarkeit von CO, steigt.

3. SchlieBlich ist das Wasser in den Hyalocyten vóllig verdunstet, die Chlorocyten bleiben

jedoch noch eine Zeit lang turgeszent. Das diirfte die optimale Phase fiir die Photosyn-

these sein.

Die Frage bleibt, ob Arten mit Hyalocyten im Endeffekt eine hóhere Nettopho-

tosynthese haben als Arten ohne Hyalocyten. Die Antwort hángt vermutlich weitgehend

von den ókologischen Bedingungen ab. Wenn Hyalocyten durch feuchtes Klima und

dauernde Befeuchtung immer wassergefüllt bleiben, so ist das sicher ein Nachteil. Es

kónnte ein Vorteil sein, wenn das Klima oft zwischen trockenen und feuchten Phasen

wechselt. Unter diesem Aspekt ist es aber unverstándlich, warum die meisten Leucobrya-

сееп in den Feuchttropen vorkommen. Auch wenn man postuliert, daB die Leucobryaceen

in temperaten Klimaten entstanden sind und erst spáter in die Feuch ttropen vorgedrungen

sind, so ist es kaum verständlich, daß sie unter diesen Bedingungen diese Strukturen nicht

reduziert hátten, wenn sie einen Nachteil bedeutet hátten. Im Gegenteil haben sich in den

Tropen neue Gattungen gebildet.

Berechungen aus dem Verdunstungsversuch zeigen, daf ein Quadratmeter von

Leucobryum etwa 17 Liter Wasser speichert. Die Verdunstungsrate eines Leucobryum

Polsters von 100 cm? betrágt zwischen 0,6 g/h und 2,6 g/h, im Durchschnitt 1,6 gh

wáhrend der ersten 54 Stunden und weniger gegen das Ende der Austrocknung. Die

durchschnittliche Verdunstung wáhrend der gesamten Austrocknung betrágt 1,3 g/h/

100 cm”. Auf diese Weise verdunstet Leucobryum zwischen 100 und 200 ml Wasser per

Stunde und m? unter Testbedingungen (21? C, 45% rH). Die Verdunstungsrate von

Mnium hornum ist ungefáhr dieselbe wie bei Leucobryum glaucum. Ein Polster von 100 cm?

verdunstet im Schnitt 1,2 g/hr, hat aber eine Verdunstungsperiode, die nur ein Drittel so

lang ist wie bei Leucobryum. Unter denselben Bedingungen verdunstet eine Wasserfláche

von 100 cm? zwischen 1,1 and 1.8 g/h. Daher ist die Verdunstung eines Moospolsters nicht

signifikant höher als die einer gleich großen Wasserfläche. Sphagnen sollen nach Oltmanns

(zitiert nach Goebel, 1915) fünf mal so viel Wasser verdunsten wie eine entsprechende

Wasserfläche.

DISKUSSION

Zusammenfassend kann man sagen, daB die Funktion der Hyalocyten nicht

befriedigend erklárt ist. Wie ausgeführt wurde, kann man Vorteile durch Wasserspeicher-

ung, einen besseren Kationenaustausch oder besseren Gasaustausch ausschließen. Ein

Vorteil für die Photosynthese müßte belegt werden. Es könnte allenfalls sein, daß eine

Kombination der genannten Faktoren einen Vorteil für Moose mit Hyalocyten haben.

Vielleicht spielen auch ganz andere Gründe eine Rolle. Zum Beispiel haben

Source : MNHN. Paris

242 J-P. FRAHM

Arten mit groBen Hyalocyten (Leucobryum, Paraleucobryum, bei Campylopus nur die

Arten mit großen Hyalocyten wie C. pittieri) auffálligerweise keine Flavonoide. Das

kónnte andeuten, daf diese Arten einen anderen Stoffwechsel haben, in denen die Hyalo-

cyten eine Rolle spielen.

Eine sehr spekulative Erklärung könnte auch sein, daß Hyalocyten ursprünglich

für endophytische Algen, Cyanobakterien oder Bakterien angelegt wurden. Endophyt-

ische Algen wurden in den Zellen von mehreren Moosarten gefunden und wurden auch in

den Hyalocyten von Campylopus aureonitens (C. Müll.) Jaeg. aus Malawi gefunden.

Eine letzte Erklärung könnte sein, daß diese anatomische Struktur der Hyalo-

cyten unter anderen ökologischen Bedingungen als heute gebildet wurde und heute keine

spezielle Funktion mehr hat. Diese Erklärung drängt sich speziell auf bei an demselben

Standort koexistierenden Arten mit und ohne Hyalocyten, wie sie für Leucobryum glau-

cum und Campylopus flexuosus sowie Paraleucobryum longifolium und Dicranum fulvum

geschildert wurden.

Ich danke Herrn H. Ibsch, Anorganische Chemie der Universität Duisburg, für die Durchführung

der lonenaustauschversuche, und B.J. O'Shea für die Durchsicht des Abstract.

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SCHOFIELD W.B., 1985 — Introduction to Bryology. New York.

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BIBL.DU.

MUSEUM

PARIS

Commission paritaire 15-9-1981 - N° 58611 - Dépôt légal 3° trimestre 1997 - Imprimerie F. Paillart

Sortie des presses le 31 juillet 1997 - Imprimé en France.

Éditeur : A.D.A.C. (Association des Amis des Cryptogames)

Lamy ; Secrétaire : B. Dennétière

- Directeur de la publication : H. Causse

Source : MNHN, Paris

Société Francaise de Systématique <3

La Société Française de Systématique réunit les systématiciens ou les personnes

intéressées par la Systématique et les informe en publiant un Bulletin. Elle convie ses

membres à des colloques annuels transdisciplinaires, au cours desquels les systématiciens

et d'autres scientifiques peuvent s'exprimer et débattre.

Cotisation annuelle: 100F

Demande d'adhésion à adresser au:

Secrétariat de la Société Française de Systématique, 45 rue Buffon, F-75005 Paris.

CCP 7-367-80 D PARIS

La Société édite aussi la série Biosystema.

Prix TTC du Biosystema (France, Etranger): 150 FF, membre SFS : 100 FF.

Biosystema 1 - Introduction à la systématique zoologique - (Concepts. Principes, Méthodes) par

L. Matile. P. Tassy & D. Goujet. 1987.

Biosystema 2 - Systématique Cladistique - Quelques textes fondamentaux. Glossaire. Traduction

et adaptation de D. Goujet, L. Matile, P. Janvier & J.P. Hugot. 1988

Biosystema 3 - La systématique et l'évolution de Lamarck aux théoriciens modernes. par

S. Lovirup. 1988.

Biosystema 4 - L'analyse cladistique: probléme et solutions heuristiques informatisées, par

M. d'Udekem-Gevers. 1990.

Biosystema 5 - Les introuvables de J.B. Lamarck- Discours d'ouverture du cours de zoologie et

articles du Dictionnaire d' Histoire naturelle. Edition préparée par D. Goujet. 1990.

Biosystema 6 - Systématique et Ecologie, par R. Barbault, Cl. Combes, F. Renaud, N. Le Brun

& A. Dubois. Edition coordonnée par J.P. Hugot. 1991.

Biosystema 7 - Systématique et Biogéographie Historique. Textes historiques et

méthodologiques. Traduction et adaptation de P. Janvier, L. Matile & Th. Bourgoin.

1991.

Biosystema 8 - Systématique et Société. Edition coordonnée par G. Pasteur. 1993.

Biosystema 9 - Les Monocotylédones, par J. Mathez. 1993.

Biosystema 10 - Systématique botanique : problémes actuels. Edition coordonnée par O. Poncy.

1993,

Biosystema 11 - Systématique et Phylogénie: modéles d'évolution biologique. Edition

coordonnée par P. Tassy et H. Leliévre. 1994.

Biosystema 12 - Phylsyst: logiciel de reconstruction phylogénétique, par I. Bichindaritz,

S. Potter & B. Sigwalt *. 1994.

Biosystema 13 - Systématique et Biodiversité. Edition coordonnée par Th. Bourgoin. 1995.

Biosystema 14 - Systématique et Informatique. Edition coordonnée par J. Lebbe. en préparation.

Le Conseil de la SFS. XII 1995

Source : MNHN, Paris

SOMMAIRE

Editorial

Claritza E. GRADSTEIN-SERNA — Un tributo de Colombia a Helena Bischler .

David С. LONG — Studies on the genus Asterella P. Beauv. III Asterella cruciata

(Steph.) Horik. in Eastern Asia.

S. Rob GRADSTEIN and Patricia GEISSLER — Notes on the genus Leuco-

lejeunea (Hepaticae)

Suzanne JOVET-AST — Un Riccia prolifére d'Australie: Riccia pullulans sp. nov.

AE AS оди за ee АЛДЕ

Riclef GROLLE and M. L. SO — Plagiochila bischleriana, a new species from

NN ET AR TER N ree atari ee ср лгы

Tamas PÓCS — New or little known epiphyllous Liverworts, VIL. Two new

Lejeuneaceae species from the Mascarene Islands .

Benito C, TAN and Jian-Cheng ZHAO — New Moss records and range exten-

sions of some xeric and alpine moss species in China..................

Cecilia SERGIO and Ana SÉNECA — The first report of Cryptothallus mirabilis

Malmborg (Hepaticae, Aneuraceae) in Southern Europe (Portugal) ....

5. Rob GRADSTEIN — Bromeliophila helenae, a new species of Lejeuneaceae

from the Neotropics.

Barbara M.THIERS — Lejeunea bischlerae, a new species of Lejeunea Ed

Microlejeunea from Australia .

William R. BUCK and Bruce ALLEN — Ordinal placement of the Fontinalaceae.

Jan-Peter FRAHM — Welche Funktion Haben die Hyalocyten in den Bláttern

Cryptogamie, Bryologie-Lichénologie 1997, 18(3): 165-242.

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