DESCRIPTIONS AND ILLUSTRATIONS OF BARBULA,

PSEUDOCROSSIDIUM AND BRYOERYTHROPHYLLUM (P.P.)

OF MEXICO

R.H. ZANDER *

SUMMARY. — The genera Barbula and Pseudocrossidium are redescribed and their Mexican

species redescribed and illustrated in a full, modern manner. Two Mexican species of Bryo-

erythrophyllum until recently recognised as taxa of Barbula are fully redescribed and illus-

trated. Keys are given to the three genera and to their Mexican species. Barbula indica var.

gregaria (Mitt.) is a comb. et stat. nov.

Key-words : Mosses, Mexico, Barbula, Pseudocrossidium, Bryoerythrophyllum.

The three genera Barbula Hedw., Pseudocrossidium Williams and Bryoery-

throphyllum Chen each have at least some species that together share several

salient characteristics : twisted peristomes (WILLIAMS 1915, ZANDER 1978a,

b, 1979, 1980); leaves with differentiated basal cells and costae with two stereid

bands and differentiated ad- and abaxial epidermises; laminal papillae multiplex

and crowded; and axillary hairs of hyaline cells alone. The genera are distin-

guishable, however, by the key given below, which emphasizes what are appa-

rently three different evolutionary directions.

KEY TO BARBULA, PSEUDOCROSSIDIUM AND

BRYOERYTHROPHYLLUM

1. Dioicous or monoicous; lower leaves reddish in nature, laminae K (10% KOH)

usually medium to deep red or red-brown; laminal papillae hollow, sometimes

solid; basal laminal cells sometimes hyaline, inflated, and bulging; peristome

usually of short, straight or rudimentary teeth, seldom long and twisted .

ИЛТ

1 Dioicous; ewer leaves For "seldom reddish in nature, laminae K (10%

KOH) usually light to deep yellow or yellow-brown to orange-brown; laminal

papillae solid or hollow; basal laminal cells light brown, with straight walls,

not bulging; peristome usually long and twisted. ................... 2

* Clinton Herbarium, Buffalo Museum of Science, Buffalo, NY 14211 U.S.A.

Cryptog., Bryol. Lichénol., 1981, 2, 1 : 1-22.

Source : MNHN, Paris

% R.H. ZANDER

2. Perichaetial leaves in many species strongly differentiated, convolute-

sheathing and largely prosenchymatous; photosynthetic organs formed

in many species either by highly revolute and chlorophyllose leaf margins

or adaxial coastal cells forming a pad of filaments, or both; adaxial stereid

band of costa usually absent, but hydroids usually present; cells more

papillose on lamina medially than at leaf margins (except in coils of

revolute portion). ................,...,...... Pseudocrossidium

2. Perichaetial leaves not differentiated or sheathing the seta in lower half,

seldom convolute-sheathing, seldom largely prosenchymatous; no part of

costa or lamina specialized as photosynthetic organ; adaxial stereid band

usually present, hydroids absent in most species; lamina usually evenly

euo. up MEI UMS iter Mean Barbula

The full descriptions and illustrations provided here for species of these

genera are intended to support recent re-evaluations of generic limits (SAITO

1975, ZANDER 1978a, b, 1979, 1980) of Barbula and related genera in the

Pottiaceae, and are based on examined specimens from many herbaria, but

chiefly BM, BUF, CANM, FH, MICH, NY, PC, SPA, TENN and US. Synony-

my given here is restricted to names with Mexican types; other names in syno-

nymy may be found in the papers cited above. Mexican distribution is based

largely on specimens examined, but also on citations by CRUM (1951). The

chromosome numbers given are taken from the literature.

BARBULA HEDW., Spec. Musc. : 115, 1801, nom. cons.

Plants loosely caespitose to cushion-forming, green to yellow- or black-

brown. Stems mostly 0.2-3.0 cm long, central strand present. Leaves spathulate

to lanceolate or long-triangular, ventrally broadly concave or narrowly chan-

nelled, margins entire to weakly dentate, plane to recurved, leaf apex rounded

to acute, leaf base weakly differentiated to sheathing the stem; costa ending a

few cells below the apex to short-excurrent, ventral and dorsal surficial cells

quadrate to elongate, papillose or smooth; costa in transverse section round to

semicircular, epidermal layers differentiated, adaxial stereid band usually diffe-

rentiated, guide cells usually 24, hydroids (Begleiter cells) sometimes present,

dorsal stereid band usually strong; upper laminal cells subquadrate to short-

rectangular, walls thin to evenly thickened, surficially flat on both surfaces to

weakly convex or bulging adaxially, laminal papillae absent or solid to hollow,

often centered over the lumens, often multiplex; basal laminal cells usually

differentiated, rectangular smooth to weakly papillose. Dioicous (two species

possibly rhizautoicous). Gametoecia terminal, perigonia gemmate; perichaetial

leaves ovate to longlanceolate, often enlarged, occasionally strongly sheathing

the seta, laminal cells usually prosenchymatous in lower half of leaf. Sporophyte

seta elongate, twisted clockwise below, often counterclockwise above; theca

short- to longcylindrical; stomata at base of theca, phaneropore; annulus of

1-2 rows of vesiculose cells usually differentiated, occasionally revoluble or

Source : MNHN. Paris

BARBULA, PSEUDOCROSSIDIUM AND BRYOERYTHROPHYLLUM 3

deciduous in parts; peristome weakly or strongly twisted, of 32, linear usually

densely spiculose teeth, basal membrane low. Operculum short- to long-conic.

Calyptra cucullate, smooth. Laminal color reactions (after clearing in concen-

trated lactid acid) : Cl (conc. HCl) : medium green to bright yellow-green usually

predominating + light to medium yellow-brown to orange-brown, occasionally

light brown; K (10% KOH) : green seldom present, light to deep yellow-brown

to orange-brown; N (conc. HNO3) : light to deep red or red-brown, occasionally

light to dark orange or light brown; SE (Ha SO4-ethanol, 2 1) : green usually

present + light to medium yellow-brown to orange-brown or light brown, sel-

dom light red throughout or with deep red blotches.

KEY TO SPECIES OF BARBULA IN MEXICO

1. Leaves spathulate, laminal cells smooth, adaxially bulging and abaxially near-

ly plane, annulus revoluble . ......... veres ee e 26. B. agraria Hedw.

1. Leaves lanceolate to ovate, laminal cells papillose to nearly smooth, similar on

both surfaces or somewhat more strongly convex adaxially than abaxially,

unnplus persistent yana a ЖІ etre арен, EDS ET th ткен 2

2. Leaves flaccid when wet, upper laminal cells rectangular, usually smooth,

often adaxially more convex than abaxially .........2... Бе PO

2. Leaves firm when wet, upper laminal cells subquadrate, papillose, both

exposed surfaces similar... . . . . « d abf EE gli SGAM DS Le 4

3. Leaves ovate-lanceolate, margins usually plane, laminal cells 11-15 im

ids) ODP As hk tg Ier E HC e A. B. ehrenbergii (Lor.) Fleisch.

3. Leaves narrowly lanceolate to long-triangular, margins narrowly recur-

ved to near apex, laminal cells 8-12 um wide, 1-2:1 . 5. В. arcuata Griff,

4. Upper leaf margins recurved to near apex. 1. B. orizabensis С. Muell.

4. Upper leaf margins plane. . . . . Qi ae cu RES nC eun ld En 5

5. Abaxial costa surface doubly prorulose (i,e., with both ends of

rectangular surficial cells protruding) near apex, often coarsely

mamillose or very rough in appearance; propagula often present

in leaf axils, occasionally on basal rhizoids . .

Le aS LN aco EME Re 2. B. indica (Hook.) Spreng.

5. Abaxial costa surface with hollow or solid, scattered papillae;

propagula (when present) on basal rhizoids only............

зы NITE Ci meray 3. B. convoluta Hedw.

1. BARBULA ORIZABENSIS C. MUELL., Linnaea 38:638, 1874. Type : Me-

xico, Veracruz, Orizaba, Mohr, 1874 (NY, topotype).

— Barbula stenotheca Thér., Smiths. Misc. Coll. 85 (4) : 21, 1931. Type :

Mexico, Distrito Federal, Río Frfo, Amable 1726 (PC, holotype).

Plants turf-forming, occasionally with branches ascending from procumbent

stems, yellow-green above, brown below. Stems often branching, brown, 1.5-

Source : MNHN, Paris

4 RH. ZANDER

2.5 cm long, in transverse section pentagonal, central strand strong, cortex of

smaller, darker cells, hyalodermis absent; axillary hairs of 6-10 hyaline, unise-

riate cells. Leaves when dry appressed-incurved and spiralled around the stem,

when wet spreading to 60°, ligulate to oblong, mostly 1.5-1.9 mm long, adaxial

surface narrowly grooved along costa; leaf margins revolute or narrowly recurved

to near apex, entire but extreme apex often slightly denticulate: leaf apex obtuse

to rounded; leaf base ovate, basal margins short-decurrent; costa short-excurrent

as а stout mucro, adaxial surficial cells quadrate to short-rectangular, papillose,

adaxial surficial cells elongate, papillose, transverse section of costa semicircular,

adaxial surface flat, adaxial and abaxial epidermises differentiated in one layer,

adaxial stereid band weak to absent, guide cells 3-4 іп one layer, hydroids

(Begleiter cells) present, abaxial stereid band strong; upper laminal cells sub-

quadrate, 7-9 um wide, 1 : 1, walls thin to evenly thickened, bulging on both

surfaces, arranged in longitudinal rows, homogeneous in shape and size; laminal

papillae multiplex, massive, covering lumens; basal laminal cells weakly medially

differentiated, 9-12 um wide, 4-6 : 1, walls evenly thickened. Propagula spherical

or occasionally elliptical, of 5-6 cells, 30-35 um in diameter, borne in clusters on

short stalks in leaf axils.

Dicicous. Perichaetia terminal, inner leaves oblonglanceolate, to 3.0 mm

long, lower half sheathing the seta and prosenchymatous; perigonia terminal

or in pseudolateral series, gemmate, hidden by subtending leaves. Sporophyte

seta 1.5-1.9 cm long, one per perichaetium, red-brown, twisted clockwise

below, counterclockwise above: theca 1.9-4.2 mm long, orange-brown, smooth

when dry, longcylindrical and occasionally curved, neck absent to weakly

differentiated, exothecial cells rectangular to rhomboidal, 20-25 um wide,

2-5 : 1, walls thin; stomates at base of theca, phaneropore; annulus of vesi-

culose cells in 3 rows, not deciduous; peristome twisted counterclockwise

once, of 32 longlinear, red teeth with many articulations, 500-1000 ит long,

densely spiculose, basal membrane very low or absent, to 25 um high, papillose;

operculum long-conic, 1.1-1.5 mm long, cells twisted counterclockwise. Calyptra

cucullate, smooth, 2.5-3.5 mm long. Spores light brown, papillose, 9-12 um in

diameter. Laminal color reactions (after conc. lactic acid clearing) : Cl (conc.

HCl) : bright yellow-green or green + yellow-brown; К (10% KOH) : bright

yellow-orange to medium orange; N (conc. HNOs) : light red to red-brown;

SE (Н, SOs-ethanol, 2 : 1) : green + light brown or yellow-brown + red blot-

ches. Illustration : Pl. I, fig. 1-7.

Barbula orizabensis is very similar to В. unguiculata Hedw. in overall appea-

rance. The latter species, which is very common in Anglo America but has not

yet been found in Mexico, has an obtuse to broadly acute leaf apex and leaf

margins recurved in the lower 1/2-2/3, and propagula are absent, while B. ori-

zabensis is easily distinguished by the abruptly rounded to emarginate leaf

apex, leaf margins recurved to near the apex, and propagula are commonly

evident in the leaf axils.

Habitat : soil, rock, walls, often limestone, 910-1850 m elevation.

Mexican distribution : Baja California, Baja California Sur., Chiapas, Coa-

Source : MNHN. Paris

BARBULA, PSEUDOCROSSIDIUM AND BRYOERYTHROPHY LLUM E

PL. I. — Barbula orizabensis and B. indica. — 1-7 :B. orizabensis. 1:Operculum. 2 : Peri-

stome. 3-4: Leaves. 5:Leafapex. 6 : Upper median leaf cells. 7 : Transverse section

of costa. — 8-15 : B. indica. 8 :Peristome, 9-10 : Leaves. 11 :Leafapex. 12 : Upper

median leaf cells. 13 : Transverse section of costa. 14 : Propagula of var. gregaria. 15:

Propagula of var. indica. A —0.5 mm; B =50 im.

Source : MNHN, Paris

6 R.H. ZANDER

huila, Distrito Federal, Guerrero, Hidalgo, Jalisco, Michoacán, Morelos, Nuevo

León, Puebla, Sonora, Tamaulipas, Veracruz.

Range : Mexico, Guatemala, Jamaica, Dominican Republic.

2. BARBULA INDICA (HOOK.) SPRENG. in Steud., Nomencl. Bot, 2 : 72,

1824.

— Tortula indica Hook., Musci Exot. 2 : 135, 1819, basionym, nom. nov. for

Trichostomum indicum Schwaegr., Spec. Musc. Suppl. 1 (1) : 142, 1811,

hom. Шер. non Trichostomum indicum Willd. ex Schrad., 1803. Type :

India, Madras, Tranquebar, Róttler s.n. (NY, isotype).

— Barbula rufipes Schimp. ex Besch., Mém. Soc. Nat. Sci. Nat. Cherb. 16 : 180,

1872. Type : Mexico, Veracruz, Orizaba, Mueller, 1853 (BM, holotype)

— Semibarbula rufipes (Schimp. ex Besch.) Hilp., Beih. Bot. Centralbl.

50 (2) : 622, 1933.

— Barbula hypselostegia Card., Rev. Bryol. 36 : 84, 1909. Type : Mexico,

Puebla, Honey Station, Pringle 10653 (PC, holotype; TENN, isotype)

- Streblotrichum hypselostegium (Card.) Hilp., Beih. Bot. Centralbl. 50

(2) :635, 1933.

— Barbula muenchii Card., Rev. Bryol. 36 : 84, 1909. Type : Mexico, Chiapas,

San Cristóbal, Muench, 1907 (NY, isotype).

— Barbula pringlei Card., Rev. Bryol. 36 : 85, 1909. Type : Mexico, Morelos,

Cuernavaca, Pringle 10637 (PC, lectotype; BM, FH, TENN, isotypes),

15177 (PC, syntype) - Streblotrichum pringlei (Card.) Hilp., Вей. Bot.

Centralbl. 50 (2) 635, 1933.

A full description and extensive discussion of this polymorphic species are

given by ZANDER (1979). Those who wish to recognise certain intergrading

infraspecific variants as taxa may do so with the following scientific names :

Barbula indica (Hook.) Spreng. var. indica

Plants with small, green, obovoid propagula occurring in masses in the upper

leaf axils, composed of 3 to several cells; leaves usually narrowly oval to ellip-

tical, with margins plane or weakly recurved at midleaf. Found throughout the

range of the species and known in Mexico from Durango, Jalisco, San Luis

Potosi, Sonora, and Veracruz.

Barbula indica var. gregaria (Mitt.) Zander, comb. et stat. nov.

Tortula gregaria Mitt., J. Linn. Soc. Bot. London Suppl. 1 : 29, 1859, basio-

пут. Type : Nepal, Tambar R., Hooker 166 (NY, syntype) — Barbula

gregaria (Mitt.) Jaeg., Ber. S. Gall. Naturw. Ges, 1871-72 : 424, 1873

(Ad. 1 : 272).

Plants with massive, brown, elliptical to spherical, often armed propagula

occurring usually 1-3 in upper leaf axils or solitary on basal rhizoids, or both,

composed of up to ca. 50 cells; leaves usually broadly oval, with plane margins.

Found mainly in tropical and subtropical regions, and im Mexico from the

states of Chiapas, Hidalgo, Morelos, Oaxaca, Puebla, and Veracruz.

Source - MNHN, Paris

BARBULA, PSEUDOCROSSIDIUM AND BR YOERYTHROPHYLLUM 7

Barbula indica sensu lato.

This phrase may be used to name collections that lack propagula or which

have propagula of intermediate morphology. Such specimens do not extend

the known Mexican distribution of the species beyond the states mentioned

above.

The names of infraspecific variants recognised here are presented in conso-

nance with the International Rules of Botanical Nomenclature, but, because the

infraspecific taxa intergrade in morphology, no heterotypic synonyms are

assigned to them. Illustration : Pl. 1, fig. 8-15.

3. BARBULA CONVOLUTA HEDW., Spec. Musc. : 120, 1801.

— Tortula convoluta (Hedw.) Gaertn., Meyer & Scherb., Оек. Techn. Fl.

Wetterau 3 (2) : 92, 1802 — Streblotrichum. convolutum (Hedw.) P.

Beauv., Prodr. : 89, 1805.

Plants turf-forming, light to dark green above, light brown below. Stems

often branching, brown, ca. 1.0-1.2 cm long, transverse section pentagonal,

central strand strong, cortex of small-lumened, thicker-walled cells, hyaloder-

mis absent; axillary hairs of 5-8 hyaline, uniseriate cells; stem with red radicles

or tomentum. Leaves crowded, when dry appressed, incurved, contorted, when

wet weakly to widely spreading, ligulate to ovate, 1.0-1.7 mm long, adaxial

surface narrowly and deeply grooved along the costa; leaf margins weakly

recurved in lower 1/3, plane above, entire but minutely crenulate by papillae;

leaf apex entire or apiculate by a smooth, conical cell, rounded or broadly

acute, occasionally sharply acute; leaf base ovate; costa ending (4-)6-8 cells

below apex, weak, not bulging abaxially, adaxial surficial cells smooth to weak-

ly papillose, elongate, abaxial surficial cells elongate, papillose, transverse section

of costa semicircular, adaxial surface convex, epidermal cells differentiated

both ad- and abaxially, abaxial epidermal cells with ovate to semicircular lu-

mens, adaxial stereid band absent or weak, guide cells 3 in one layer, hydroids

absent, abaxial stereid band strong; upper laminal cells subquadrate, 8-10 um

wide, 1: 1, walls evenly thickened, bulging on both surfaces, arranged in longi-

tudinal and transverse rows, homogeneous in shape and size; laminal papillae

multiplex, massive, centered over and covering the lumens, 4-6 salients per

lumen; basal laminal cells differentiated across leaf base but higher along the

costa, rectangular, 9-12(-18) um wide, 3-5 : 1, walls thin, Propagula as multi-

cellular brood bodies on basal rhizoids, spherical to clongate-elliptical, to 1150 um

long, deep red-brown, individual cells 25-50 um in diameter.

Dioicous (but possibly rhizautoicous). Perichaetia terminal, inner leaves

ovate to ovate-acute, 0.8-1.0 mm long, convolute-sheathing, entirely prosen-

chymatous; perigonia clustered terminally on very small androgametophores

situated at base of gynogametophytes, gemmate. Sporophyte seta one per

perichaetium, 1.0-1.8 cm long, yellow to yellow-brown, twisted clockwise

below and counterclockwise above; theca 0.8-1.3 mm long, red-brown, smooth

when dry, long-lliptical, occasionally curved, neck absent to weakly differen-

tiated, microstomous; exothecial cells rectangular-rhomboidal, 20-30 um wide,

Source : MNHN, Paris

8 RH, ZANDER

2-3 : 1, walls thick; stomates at base of theca, phaneropore; annulus deciduous,

revoluble; peristome twisted 1.5-2.0 times counterclockwise, of 32 long-linear,

red teeth, with many articulations, spiculose, 700-1300 um long, basal mem-

brane low, ca. 45-55 um high, papillose; operculum long-conic, 0.8-1.2 mm long,

cells twisted counterclockwise. Calyptra cucullate, smooth, 1.8-2.1 mm long.

Spores light brown, essentially smooth, 10-12 um in diameter. Lamina color

reactions (after clearing in conc. lactic acid) : Cl (conc. НСІ) : bright yellow-

green; К (10% KOH) : deep yellow-orange; N (conc. HNO3) : medium to deep

red; SE (HaSOs-ethanol, 2 : 1) : green + medium yellow- or orange-brown.

n = 11, 13, 14. Illustration : Pl. II, fig. 1-4.

Barbula convoluta is widely distributed in the North Temperate Zone and

disjunct to New Zealand, but 1 have seen only one collection from Mexico

(reported by KOCH & CRUM 1950) : Baja California Sur., Sierra de la Laguna,

La Laguna, E of Todos Santos, canyon W of cabin, granite gravel under edge

of rack, Carter 2372 (MICH).

4. BARBULA EHRENBERGII (LOR.) FLEISCH., Musci Arch Indic. Exs.

ser. 4, n. 161, 1901.

— Trichostomum ehrenbergii Lor, Abhandl. Akad. Wiss. Berlin 1867 : 25,

1868, basionym.

— Barbula ehrenbergii var. mexicana Thér., Smiths. Misc. Coll. 85 (4) : 19,

1931. Type : Mexico, Nuevo León, Monterrey, Abbon 10969 (PC, holo-

type).

Plants loosely caespitose to turf-forming, green above, brown to light tan

below. Stems branching little, red-brown, 1.0-3.5 cm long, in transverse section

pentagonal, central strand present, cortex weakly differentiated, hyalodermis

absent; axillary hairs of 5-9 uniseriate, hyaline cells; stem often incrusted with

carbonate deposits. Leaves when dry erect and weakly incurved, when wet

patent at 45°, long elliptical to short-ligulate, 2.0-2.7 mm long, adaxial surface

flat to broadly channeled across leaf; leaf margins recurved in lower 1/2-3/4,

entire or denticulate at apex; leaf apex narrowly to broadly obtuse; leaf base scar-

cely differentiated in shape to ovate, basal margins decurrent; costa percurrent or

subpercurrent by 2-3 cells, dorsally bulging, adaxial and abaxial surficial cells

short-rectangular to elongate, smooth, transverse section of costa semicircular,

adaxial surface weakly convex, adaxial and abaxial epidermises differentiated,

abaxial epidermis lumens hexagonal, adaxial and abaxial stereid bands strong,

guide cells 3-5 in one layer, hydroids absent; upper laminal cells subquadrate to

short-rectangular, 11-15(-16) um wide, 1-2 : 1, walls thin to evenly thickened,

bulging more adaxially than abaxially, arranged in longitudinal and, often, in

transverse rows, somewhat larger in size medially; laminal papillae usually

absent, occasionally low, small, hollow, simple, 4-6 over each lumen; basal

laminal cells differentiated across leaf and higher on the margins, rectangular,

mostly 16-27 um wide, 34 : 1, cell walls evenly thickened or thin, hyaline,

marginal cells occasionally prorulose. Propagula borne in leaf axils on stalks,

corniculate (branchingstellate) or filamentous, multicellular, to 180 um long.

Source : MNHN, Paris

BARBULA, PSEUDOCROSSIDIUM AND BRYOERYTHROPHYLLUM 9

РЛ. — Barbula convoluta and В. ehrenbergii. — 1-4 :B. convoluta. 1:Leaf, 2:Leaf

apex. 3:Upper median cells. 4 : Rhizoidal propagula. — 5-8 :B. ehrenbergii. 5:

Leaf. 6 :Leaf apex. 7: Upper median cells. 8 : Transverse section of costa. A =

0,5 mm; В =50 Um.

Sporophyte seta deep red, paler red to yellow above, to 2 cm long; theca oval

to oval-cylindrical, 1-2 mm long; peristome to 600 um long, twisted once clock-

wise, of pale red to transparent, fragile teeth, basal membrane low; operculum

long-rostrate, to 2 mm long. Spores ca. 2 um in diameter (sporophyte and spore

characters from CHEN 1941 and STEERE 1937). Laminal color reactions

(after clearing in conc. lactic acid) : Cl (conc. HCl) : green + light yellow-brown

to tan; K (10% KOH) : light yellow-brown to medium orange-brown; N (conc.

HNO) : light orange-brown to red or tan; SE (H2 SOq-ethanol, 2 : 1) : green +

light orange-brown to yellow-orange-brown or brown. n = 13. Illustration :

РЇ. П, fig. 5-8.

Barbula ehrenbergii is a North Temperate Zone species very closely related to

the more tropically distributed B. arcuata, and some specimens cannot be

assigned to either taxon with certainty. It

B į

MUS Jl

+ . Source : MNHN, Paris

10 RH, ZANDER

Habitat : wet, calcareous rocks, often in waterfalls, springs or running water,

often tufa-forming, 650-1700 m elevation.

Mexican distribution : Chihuahua, Coahuila, Nuevo León, San Luis Potosí.

Range : southern U.S.A., Mexico, Belize, Europe, northern Africa, eastern

Asia and the Middle East.

. BARBULA ARCUATA GRIFF., Calcutta J. Nat. Hist. 2 :491, 1842. Type :

India, Griffith 27 (BM, holotype).

— Hydrogonium arcuatum (Griff.) Wijk & Marg., Taxon 7 : 289, 1958.

— Barbula stillicidiorum Card., Rev. Bryol. 37 : 126, 1910. Type : Mexico,

Veracruz, Barnes & Land, 1906 (NY, isotype).

— Barbula rubricaulis Thér., Smiths. Misc. Coll. 85 (4) : 19, 1931. Type : Me-

xico, Nuevo León, Monterrey, Abbon 10968 (FH, isotype).

Plants loosely caespitose or turf-forming, yellow-green above, tan or occa-

sionally brown to red-brown below. Stems little branching, red-brown to dark

red, 1.0-2.5 cm long, in transverse section pentagonal, central strand distinct

to strong, cortex of smaller, more thick-walled cells, hyalodermis not differen-

tiated; axially hairs of 3-6 cells, all hyaline or basal cell brown. Leaves rather

distant, when dry weakly spreading from insertion, apices curled, often catenu-

late, appressed-twisted, incurved, when wet weakly spreading, long triangular

to lanceolate, 1.1-2.0 mm long, adaxial surface broadly channeled across leaf

to narrowly grooved medially with a channel about twice as wide as the costas

leaf margins narrowly recurved from base to near apex, occasionally revolute

below, entire but denticulate at extreme apex; leaf apex often somewhat cucul-

late: leaf base scarcely differentiated in shape, basal margins scarcely decurrent

to decurrent by hyaline, inflated cells; costa percurrent, often sinuose near

apex, adaxial surficial cells elongate, 3-5 : 1, abaxial surficial cells rectangular,

2-4 : 1, transverse section of costa above midleaf round to semicircular, adaxial

surface convex, ad- and abaxial costal epidermises differentiated, lumens of

abaxial epidermis oval, adaxial stereid band absent, guide cells 3-4 in one layer,

hydroids absent, abaxial stereid band weak to strong; upper laminal cells short-

rectangular, 7-10(-12) um wide, 1-2:1, walls thin to evenly thickened and

somewhat porose, bulging more adaxially than abaxially, arranged in longitudi-

nal rows, relatively homogeneous in size and shape; laminal papillae absent;

basal laminal cells weakly differentiated, short-rectangular, to 14 um wide,

34 :1, walls evenly thickened, medial cells occasionally brownish. Propagula

comiculate (branchingstellate), fusiform or armed, borne on branching brown

stalks in leaf axils, median cells to 45 um in diameter, composed of 12-15 cells,

to 135(-185) um long.

Dioicous. Perichaetia terminal, inner leaves weakly differentiated, to 1.5 mm

long, vaginula often large and prominent; perigonia terminal or in pseudolateral

series, gemmate and often prominent. Sporophyte seta one per perichaetium,

0.8-1.7 mm long, red-brown, twisted clockwise; theca 0.8-2.5 mm long, yellow-

to red-brown, smooth to striated when dry, short-cylindrical, neck weakly

Source : MNHN, Paris

BARBULA, PSEUDOCROSSIDIUM AND BRYOERYTHROPHYLLUM

Pl. Ш. — Barbula arcuata and В, agraria. — 1-

5 : Upper marginal cells.

+B. arcuata. 1-3 : Leaves.

6 : Transverse sec:

4 : Leaf apex.

n of costa. 7 :Corniculate propagula. —

8-12 :B. agraria B:Leaf. 9:Leaf apex. 10:Upper median cells. 11 : Transverse

section of leaf. 12 : Revoluble annulus. А —0.5 mm; B — 50 Jim.

Source : MNHN, Paris

12 RH. ZANDER

differentiated, occasionally ventricose when short; exothecial cells rectangular,

23-25 um wide, 3-4 :1, walls thin to evenly thickened; stomates present at

base of theca, phaneropore; annulus of 2-3 rows of vesiculose cells; peristome

twisted 1.5-2.0 times, of 32 longlinear, red-orange, densely spiculose teeth

with many articulations, to 900-1200 um long, basal membrane low, 45-80 um

high, densely spiculose, often perforate; operculum long-conic, 1.0-1.7 mm

long, cells twisted counterclockwise. Calyptra cucullate, smooth, 2.2-3.0 mm

long. Spores light brown, weakly papillose, 12-14(-16) um in diameter. Lami-

nal color reactions (after clearing in conc. lactic acid) : Cl (conc. НСІ) : yellow-

green (occasionally yellow); K (10% KOH) : deep yellow or bright yellow-

orange; М (conc. HNOs) : light red, occasionally light tan; SE (Ha SO4-ethanol,

2:1) : green + yellow-brown, occasionally light brown, occasionally with red

blotches. n = 13, 13 + 1. Illustration : РІ. III, fig. 1-7.

Barbula arcuata is extremely variable in leaf shape, varying from the ellipti-

cal leaves typical of B. ehrenbergii to very narrow, subulate leaves characteris-

tic of many specimens from the West Indies. This species often grows with and

looks much like narrow-leaved, hygric forms of B. indica but differs in its long-

triangular leaves with 1-3 apical teeth, costa abaxially smooth or sharply crenu-

late by projecting cell cross walls, and laminal cells smooth, short-rectangular.

Habitat :wet cliffs, riverbanks, calcareous soil, wet rock, travertine, 150-

1350 m elevation.

Mexican distribution : Nuevo Leôn, Veracruz.

Range : Mexico, Guatemala, Cuba, Haiti, Dominican Republic, Jamaica,

Puerto Rico, Guadeloupe, Venezuela, India, Burma, Indonesia, Molucca, New

Guinea, Philippines, China, Japan, western Oceania.

6. BARBULA AGRARIA HEDW., Spec. Musc. : 116, 1801. Type : «Jamaica et

Domingo», Swartz s.n. (С, syntype; BM, syntype ?).

— Tortula agraria (Hedw.) P. Beauv., Prodr. : 91, 1805.

Plants loosely caespitose to turf-forming, green above, brown below. Stems

little branched, to 0.2 cm long, in transverse section irregularly pentagonal,

central strand very strong or stem hollow, cortex not differentiated or of one

layer of thick-walled cells, hyalodermis absent; axillary hairs of ca. 2 uniseriate,

hyaline cells; stem often highly radiculose. Leaves when dry appressed to weakly

spreading, incurved, when wet weakly spreading to 45°, oblong, elliptical or

spathulate, 1.2-2.0 mm long, adaxial surface broadly channelled to concave; leaf

margins plane, entire to weakly serrulate apically, marginal cells thick-walled and

thomboidal near apex; leaf apex broadly acute; leaf base oblong, margins often

winged; costa stoutly short-excurrent as a sharp, often curved mucro, occasio-

nally percurrent, ad- and abaxial surficial cells elongate, smooth, transverse

section of costa oval, adaxial surface convex, adaxial epidermis present, abaxial

epidermis present, with oval lumens, or occasionally absent, adaxial stereid

band weak, guide cells 2 in one layer, hydroids absent, abaxial stereid band

strong; upper laminal cells subquadrate to short-rectangular or rhomboidal,

Source : MNHN, Paris

BARBULA, PSEUDOCROSSIDIUM AND BRYOERYTHROPHYLLUM 13

(811-1315) um wide, 1-2 : 1, occasionally bistratose in patches along margins

or medially, walls thin, occasionally evenly thickened, adaxially bulging but

abaxially flat, arranged in longitudinal and occasionally transverse rows, homo-

geneous in size; laminal papillae absent; basal laminal cells differentiated in

small area or up to 1/3 of leaf, rectangular, 20-25 um wide, 2-4 : 1, walls thin,

occasionally evenly thickened, occasionally brownish.

Dioicous (or possibly rhizautoicous, androgametophytes never found except

in proximity to gynogametophytes). Gametoecia terminal, inner perichaetial

leaves smaller than cauline, oval, sheathing, largely prosenchymatous, occasio-

nally serrulate, ca. 1.5 mm long; perigonia on small plants, gemmate; seta one

per perichaetium, 0.4-1.0 mm long, red-brown, twisted clockwise below, stron-

gly counterclockwise above; theca 0.3-1.7 mm long, red-brown to brown,

sulcate or strongly ribbed when dry, cylindrical to elliptical, neck weakly diffe-

tentiated; exothecial cells rectangular, 16-23 um wide, 3-5 : 1, walls thin; stoma-

tes present at base of theca, phaneropore; annulus revoluble, deciduous, of 1-2

rows of strongly vesiculose cells; peristome twisted counterclockwise 1.5-2.0 ti-

mes, of 32 long.linear, orange, densely spiculose teeth with many articulations,

500-1200 um long, basal membrane low, 45-100 um high, closely and evenly gra

nulate or densely spiculose, trabeculate; operculum long-conic, 0.8-1.5 mm

long, cells twisted 2-3 times counterclockwise. Calyptra cucullate, smooth,

1.4-2.1 mm long. Spores light brown, weakly papillose, 9-11 ит in diameter.

Laminal color reactions (after clearing in conc, lactic acid) : Cl (conc. НСІ):

deep yellow or yellow-green + medium yellow-brown; K (10% KOH) : medium

orange to deep yellow-orange; N (HNO3) : light red; SE (Hz SO4-ethanol, 2 : 1) :

light orange-brown or medium yellow-brown to medium brown. Illustration :

РІ. Ш, fig. 8-12.

Barbula agraria differs widely from other species of Barbula by the spathu-

late leaf shape, the epapillose, adaxially bulging laminal cells (but those of B.

arcuatum and B. ehrenbergii are somewhat similar), and the revoluble annulus,

and has been assigned to a monotypic subgenus, Barbula subg. Hyophiladel-

phus (C. Muell.) Zander.

Habitat : soil, rock, walls, coral, limestone, sandstone, brick, to 350 m ele-

vation.

Mexican distribution : Campeche, Hidalgo, Quintana Roo, San Luis Potosí,

Veracruz, Yucatán.

Range : Mexico, Guatemala, West Indies, northern South America.

EXCLUDED TAXA

Several taxa of Barbula have recently been excluded from the genus. These

are discussed by ZANDER (19782, 1979, 1980); two of these species are fully

described and illustrated later in the present paper as species of Bryoerythro-

phyllum.

Source : MNHN, Paris

14 RH. ZANDER

PSEUDOCROSSIDIUM WILLIAMS,

Bull. Torrey Bot. Club 42 : 396, 1915.

— Barbula sect: Revolutae BSG, Bryol. Eur. 2 : 89, 1842 (fase, 15-15 Mon.

27).— Barbula subsect. Revolutae (B.S.G.) Chen, Hedwigia 80 : 209,

1941.

Plants in cushions or turf, green to brown or red-brown. Stems mostly 0.3-

1.5 em long, central strand usually present. Leaves ovate to lanceolate, ventrally

channelled along the costa, margins entire, recurved and little differentiated

to strongly revolute and differentiated as a photosynthetic organ of thin-walled,

highly chlorophyllose, hollow-papillose cells; leaf apex acute to rounded; leaf

base scarcely differentiated to oblong; costa excurrent, mucronate or short-

awned, adaxial surficial cells quadrate to short-rectangular, papillose, in South

American species differentiated as a photosynthetic organ of a pad of filaments

of uniseriate cells, abaxial surficial cells differentiated as an epidermis of cells

Sith lumens larger than those of the stereid cells, adaxial stereid band usually

absent, guide cells 4(9) in one layer, hydroids (Begleiter cells) often present,

dorsal stercid band usually strong; upper laminal cells subquadrate to hexago-

nal, walls evenly thickened, surficially bulging; laminal papillae crowded, low,

hollow, multiplex; basal laminal cells differentiated medially, rectangular. Dioi-

cous, Gametoecia terminal, perigonia gemmate, perichaetial leaves little diffe-

rent from the cauline to highly differentiated, enlarged, convolute-sheathing,

largely prosenchymatous. Sporophyte seta elongate, twisted clockwise; theca

elliptical to cylindrical; stomata at base of theca, phaneropore; annulus of 3-4

conte of vesiculose cells; peristome weakly to once twisted, of 32 linear, densely

spiculose teeth, basal membrane absent or low; operculum short- to long-conic.

Calyptra cucullate, smooth, Laminal color reactions (after clearing in concen-

trated lactic acid) : Cl (conc. HCl) : green or yellow-green + bright to medium

yellow or yellow-orange, occasionally orange-brown, leaf tips occasionally

orange; К (10% KOH) : deep yellow-orange or yellow, occasionally medium

orange or orange-brown; М (HNOs) : light to deep ted, occasionally light brown;

SE (HSOa-ethanol, 2:1) : green + medium yellow-brown or light orange-

brown, occasionally red-orange at leaf tips or medial blotches.

Pseudocrossidium and Barbula species are rather alike in morphology,

and in acid-base laminal color reactions, except for the technical characters

given in the key above. ZANDER (1979) distinguishes all North American

species of Pseudocrossidium and Barbula in a single key and also provides

a key to South American species of Pseudocrossidium.

KEY TO MEXICAN SPECIES OF PSEUDOCROSSIDIUM

Leaves short-awned, leaf margins recurved, not markedly differentiated... . .. +

BOUT 22.2. P. aureum (Bartr.) Zander

lute, of thin-walled, hollow-papillose,

PAL 0 ЗК 1. Р. replicatum (Tayl.) Zander

Source : MNHN, Paris

BARBULA, PSEUDOCROSSIDIUM AND BRYOERYTHROPHYLLUM 15

1. PSEUDOCROSSIDIUM REPLICATUM (TAYL.) ZANDER, Phytologia 44 :

206, 1979.

— Barbula replicata Tayl., London J. Bot. 5 :49, 1846, basionym. Type : Ecua-

dor, Pichincha, Quito, Jameson 103 (BM, isotype) — Tortula replicata

(Tayl.) Wils., London J. Bot. 5 :454, 1846.

— Barbula spiralis Schimp. ex C. Muell., Syn. Musc. 1 : 622. 1849. Type : Mexi-

co, Veracruz, Yarrea, Mirador, Liebmann, 1842 (BM, isotype) — Tortula

spiralis (Schimp. ex C. Muell.) Mitt., J. Linn. Soc. Bot. 12 : 151, 1869.

— Barbula spiralis var. emarginata Card., Rev. Bryol. 36 : 84, 1909. Type : Me-

xico, Mexico, Amecameca, Pringle 10611 (M, MEXU, Е, FH, isotypes).

Plants in cushions or turf-forming, yellow-green above, red below. Stems often

branching, red-brown, 0.3-1.5 cm long, in transverse section pentagonal, central

strand strong, stem occasionally hollow, cortex of one layer of substereid cells,

hyalodermis not differentiated; axillary hairs of 5-7 uniseriate, hyaline cells;

stem sparsely radiculose. Leaves when dry appressed, spiralled, when wet sprea-

ding-recurved to 45°, ovate-ligulate to ovate-lanceolate, 1,0-1.8(-2.6) mm long,

adaxial surface narrowly and deeply grooved along the costa; leaf margins

spiral-revolute to near apex, entire, verrucose with small papillae or smooth

on exposed surfaces, internally differentiated as a photosynthetic organ of

thin-walled, hollow-papillose, highly chlorophyllose cells; leaf apex broadly

acute to rounded; leaf base ovate to oblong; costa of even width to near apex

or wider above midleaf, short-excurrent as a sharp, smooth mucro of 1-3 cells,

occasionally cuspidate, adaxial surficial cells quadrate above midleaf, papillose,

abaxial surficial cells rectangular, papillose, transverse section of costa circular

to reniform, adaxial surface convex to concave, ad- and abaxial epidermises

differentiated in one layer, lumens of abaxial epidermis semicircular, cells of

abaxial epidermis lateral to costa wide-lumened, similar to adaxial surficial

cells, adaxial stereid band absent or represented by a few substereid cells, guide

cells 4(-6) in 1(-2) layers, hydroids (Begleiter cells) present, abaxial stereid

band strong; upper laminal cells subquadrate to hexagonal, medially 8-9 um

wide, 1:1, occasionally transversely elongate on margins, 1 : 1(-3), walls thin,

occasionally thickened on adaxial surface, surficially bulging on both sides,

not distinctively patterned; laminal papillae, low, hollow, crowded, multiplex,

several salients per lumen; basal laminal cells differentiated medially in leaf

base, occasionally across leaf base, short-rectangular, to 11 um wide, walls

evenly thickened to hyaline, somewhat bulging, inner medial cells often orange.

Dioicous, androgametophytes often in separate turf. Gametoecia terminal;

perichaetial leaves little different from cauline leaves; perigonia gemmate. Sporo-

phyte seta one per perichaetium, 1.0-1.4 mm long, red- to yellow-brown, twisted

clockwise; theca 1.6-2.5(-3.5) mm long, red- to yellow-brown, smooth when dry,

elliptical to cylindrical, neck weakly differentiated; exothecial cells short-rectan-

gular, 16-20 um wide, 2-3 : 1, walls thin; stomates on base of theca, phaneropore;

annulus of 3-4 rows of vesciculose cells, not deciduous; peristome weakly to

once twisted counterclockwise, of 32 filamentous, orange to yellow, densely

spiculose teeth, with many articulations, (700-)900-1000 um long, basal mem-

Source : MNHN, Paris

P1. IV. Pseudocrossidium replicatum, P. aureum,

inaequalifolium. — 1-6 :P. replicatum 1-3 : Leaves. 4:Leafapex, 5 :Оррег median

leaf cells. 6 : Transverse section of costa. — 7-10 :P. aureum, 7 :Leaf, 8 : Leaf apex

9 : Upper median leaf cells. 10 : Transverse section of leaf, — 11-15 : B. caleareume 11-

12 Leaves. 13 :Leafapex. 14: ррег marginal cells, 15 : Transverse section of

costa. — 1619 : B. inaequalifolium. 16-17 : Leaves. 18 : Upper median leaf cells. 19

Unicellular propagula. А = 0.5 mm; В = 50 4m; С = 100 Mm.

Bryoerythrophyllum calcareum and В.

Source : MNHN, Paris

BARBULA, PSEUDOCROSSIDIUM AND BRYOERYTHROPHYLLUM 17

brane absent or low, to 35(-70) um high, weakly spiculose; operculum short- to

longconic, 0.6-1.0(-2.1) mm long, cells twisted counterclockwise. Calyptra

cucullate, smooth, 3.2-3.5 mm long. Spores light brown, weakly papillose,

9-10 um in diameter. Laminal color reactions (after clearing in conc. lactic

acid) : CI (conc. HCl) : green to yellow-green + bright yellow or yellow-orange,

occasionally leaf tips orange; K (10% KOH) : deep yellow or yellow-orange,

occasionally medium orange-brown; М (conc. HNO3) : deep red, occasionally

light to medium red; SE (HSOs-ethanol, 2:1) : green + medium yellow-

brown or light orange-brown, occasionally leaf tips red-orange. Illustration :

PL. IV, fig. 1-6.

Collections of Pseudocrossidium replicatum in Mexico аге on the whole

somewhat smaller in plant size than are those from South America. Pseudo-

crossidium revolutum (Brid. in Schrad.) Zander is known from southern Cali-

fornia (ZANDER 1979) and should be looked for in Mexico — it differs from

P. replicatum by its only once-revolute leaf margins, propagula born on the

adaxial surface of the costa, and highly differentiated perichaetial leaves.

Habitat : soil, rock, concrete, lava, adobe, walls, 750-2590 m elevation.

Mexican distribution : Chiapas, Chihuahua, Coahuila, Colima, Distrito Fede-

ral, Durango, Hidalgo, Jalisco, Mexico, Michoacán, Morelos, Nuevo León, Oaxa-

ca, Puebla, San Luis Potosí, Sonora, Tamaulipas, Tlaxcala, Veracruz.

Range : southwestern U.S.A., Mexico, northern Andes of South America.

2. PSEUDOCROSSIDIUM AUREUM (BARTR.) ZANDER, Phytologia 44 : 207,

1979.

— Tortula aurea Bartr., Bull. Torrey Bot. Club 51 : 339, 1924, basionym. Type :

U.S.A., Arizona, Pima Co., Santa Catalina Mts., Bear Canyon, Bartram

307 (Bartram, Mosses of So. Arizona 98) (FH, holotype; CU, isotype) —

Barbula aurea (Bartr.) Zander in Zander & Steere, The Bryologist 81 : 466,

1978.

Plants in cushions or turf-forming, green above and brown below. Stems

often branching, brown, to 1.0 cm long, in transverse section rounded-penta-

gonal, central strand very strong, cortex weakly differentiated, hyalodermis

absent; axillary hairs of 5-8 hyaline, uniseriate cells; stems weakly radiculose.

Leaves when dry appressed, spiralled, when wet spreading 45-60°, lanceolate

to ovate, 1.5-1.8 mm long, adaxial surface shallowly grooved along costa; leaf

margins broadly and strongly recurved to near apex, entire, occasionally orange;

leaf apex acute to rounded-acute; leaf base scarcely differentiated in shape to

oblong; costa strong to apex, excurrent as a short, or seldom long, awn, smooth,

orange, 185-360(-450) um long, adaxial surficial cells quadrate to short-rectan-

gular, рарШозе, abaxial surficial cells elongate, smooth to papillose, transverse

section of costa reniform, adaxial surface concave, ad- and abaxial epidermises

differentiated, lumens of abaxial epidermis semicircular, abaxial epidermal cells

lateral to costa enlarged, similar to adaxial epidermal cells, often brown, thicke-

Source - MNHN, Paris

18 RH. ZANDER

ned on surficial wall and colored with appearance of ocelli, adaxial stereid band

absent, guide cells 4 in 1(-2) layers, hydroids (Begleiter cells) present, abaxial

stereid band strong; upper laminal cells subquadrate to hexagonal, 12-16(-18) um.

wide, 1:1-2 (occasionally transversely elongate along margins, walls evenly

thickened, bulging on both sides, arranged in longitudinal rows; laminal papillae

hollow, crowded, multiplex, with several salients per lumen; basal laminal cells

usually differentiated medially in leaf base, rectangular, 11-13(-23) um wide,

2-5 : 1, walls thin to evenly thickened.

Apparently dioicous; perichaetia terminal, inner leaves not different from

cauline; perigonia and sporophyte not seen. Laminal color reactions (after

clearing in conc. lactic acid) : Cl (conc. HCl) : green to yellow-green + medium

yellow-brown, occasionally orange-brown; K (10% KOH) : strong yellow-brown,

occasionally orange; N (conc. HNOs) : light red + light brown; SE (Hz S04-

ethanol, 2:1) : green + light to medium yellow-brown or light orange-brown,

occasionally red blotches. Illustration : РІ. IV, fig. 7-10.

Pseudocrossidium aureum lacks most of the salient characteristics of the

genus — the perichaetial leaves are not differentiated (presence of a seta might

make a difference, but the sporophyte is unknown), there are no laminal or

costal specialized photosynthetic organs, and there is no marked differentiated

on medial and marginal laminal papillae. However, the costa transverse section

shows a strong abaxial stereid band and the adaxial band is lacking, which

distinguishes the species from Barbula (although small. plants of some species of

Barbula also lack the adaxial stereid band), while the abaxial costal epidermis is

differentiated, which distinguishes it from most Tortula species. Hydroids are

present in the costa. Pseudocrossidium is similar to P. replicatum in commonly

developing a characteristic line of large-lumened, surficially thick-walled cells,

often of a deep yellow or orange color, abaxially at the juncture of the lamina

and the costa, These cells may have the appearance of a longitudinal row of

ocelli on each side of the costa.

Habitat : soil, rock, canyon walls, ledges, 800-2350 m elevation.

Mexican distribution : Chihuahua, Coahuila, Hidalgo, Puebla, San Luis

Potosi, Sonora, Zacatecas.

Range : southwestern U.S.A., Mexico.

BRYOERYTHROPHYLLUM CHEN (Supplement)

Two Mexican species were recently excluded from Barbula (ZANDER 1980),

and are fully described and illustrated here as a supplement to my synopsis of

Bryoery throphyllum in the New World (ZANDER 19785).

Source : MNHN, Paris

BARBULA, PSEUDOCROSSIDIUM AND BRYOERYTHROPHYLLUM 19

KEY TO BRYOERYTHROPHYLLUM SPECIES IN MEXICO

1. Leaf apex narrowly acute, entire; leaf base ovate, basal cells quadrate, 1:1 ..

есен оа ааа ооа оо. Brrecurvum (Griff.) К. Saito

1. Leaf apex bluntly acute to broadly rounded, entire to dentate; leaf base

elliptical to long-oblong, basal cells rectangular, usually 2 : 1 or longer. . ... 2

2. Leaf apex entire, not apiculate, leaf margins recurved to near apex; propa-

gula when present mostly unicellular, ахкШагу.............. EE

2. Leaf apex entire or dentate, usually apiculate by 1-4 clear cells (i£ not,

then margins plane above midleaf); propagula when present multicellu-

lar, borne on basal rhizoids ............, AEST РАТ"

3. Leaves ovate, marginal cells not differentiated as а border; propagula

common, yellow-brown, borne in axillary masses. . ..............

"та Leser ele SR ud ES B. inaequalifolium (Tayl.) Zander

3. Leaves ligulate, marginal cells often thick-walled in 2-3 rows; propa-

gula uncommon, red-brown, few in leaf axils ............ DU,

«d s rani eI ta RE OON, +... .B: calcareum (Thér.) Zander

4. Leaf margins strongly recurved above midleaf, often to near apex:

usually monoicous, sometimes Шоісовв....................

елер ааа LR I ie a В. recurvirostrum (Hedw.) Chen

4. Leaf margins plane above midleaf or seldom weakly recurved to

near apex; dioicous ......... Ж NUS

5. Leaves oblong to longlanceolate, 1.0-3.5(-5.0) mm long, apex

usually broadly acute to broadly obtuse, upper margins often

dentate, often weakly bordered by thick-walled cells, propagula

absent. ..... pale D. TION QU 2... В. jamesonii (Tayl.) Crum

5. Leaves ovate-triangular to short-lanceolate, 1.0-1.8 mm long, apex

narrowly acute to narrowly obtuse, margins entire, not bordered,

propagula often present on basal rhizoids........ ENTUM 5

22. B. ferruginascens (Stirt.) Giac.

1. BRYOERYTHROPHYLLUM CALCAREUM (THÉR.) ZANDER, The Bryo-

logist 83 :232, 1980.

= Barbula calcarea Thér., Smiths. Misc. Coll. 85 (4) : 20, 1931, basionym.

Type : Mexico, Distrito Federal, Desierto, Amable 1620 (PC, lectotype:

NY, isotype); Michoacán, Morelia, Loma Santa María, Arséne 4891 (PC,

syntype).

Plants cushion- and turf-forming, green above, yellow- to red-brown below.

Stem seldom branching, red-brown, 1.0-1.4 cm long, in transverse section roun-

ded-pentagonal, central strand present, cortex of smaller, darker cells, hyaloder-

mis absent; axiallary hairs of ca. 7 hyaline, uniseriate cells; stems sparsely ra-

diculose. Leaves when dry appressed to weakly spreading, when wet weakly

spreading, ligulate, 0.5-1.5(-1.7) mm long, adaxial surface nearly flat across

Source : MNHN. Paris

20 RH, ZANDER

leaf; leaf margins recurved to apex, entire, cells occasionally bistratose in patches,

thick-walled, yellowish; leaf apex rounded; leaf base scarsely differentiated to

oblong, short-decurrent; costa ending one to several cells below the apex and

decurrent at leaf base, adaxial surficial cells quadrate to short-rectangular, pa-

pillose, abaxial surficial cells elongate, occasionally quadrate or short-rectan-

gular, papillose, transverse section of costa reniform to elliptical, adaxial surface

concave to convex, ad- and abaxial epidermises differentiated in one layer,

abaxial epidermal cells with elliptical lumens, adaxial stereid band absent or

represented by a few substereid cells, guide cells 2 in one layer, hydroids (Be-

gleiter cells) present, abaxial stereid band distinct; upper laminal cells subqua-

drate to hexagonal, 7-9 um wide, 1 : 1, walls thin, occasionally evenly thicke-

ned, bulging on both sides, arranged in much-broken longitudinal rows; lami-

nal papillae crowded, hollow, multiplex, with several salients per lumen; basal

laminal cells differentiated medially or across leaf base, rectangular, 9-12 um

wide, 2-3 : 1, walls thin. Propagula borne іп leaf axils, unicellular, red-brown,

angular-elliptical ог elliptical, 14-20 um wide, 1-2 : 1, not common.

Dioicous. Perichaetia terminal, inner leaves long-oblong, 1.5-2.5 mm long,

convolute-sheathing in lower 1/2; perigonia terminal on budlike plants ca.

1.0 mm high, gemmate. Sporophyte seta 1(-2) per perichaetium, 0.7-1.0 mm

long, light red-brown, twisted clockwise below, counterclockwise above; theca

0.7-2.0(-3.0) mm long, light brown or red-brown, cylindrical, occasionally

curved, neck weakly differentiated; exothecial cells long-rectangular, 16-20 рт

wide, 4-8 :1, walls thin to weakly thickened; stomates on neck, phaneropore;

annulus of ca. 2 rows of vesiculose cells, adherant; peristome twisted once

counterclockwise, of 32 filamentous, densely spiculose, orange teeth, ca. 500 um

long, with many articulations, basal membrane low, 10-15 um high, papillose;

operculum long-conie, 0.5-0.8 mm long, cells twisted counterclockwise. Calyp-

tra cucullate, smooth, 1.7-1.8 mm long. Spores yellow-brown, essentially smooth,

9-12 um in diameter. Laminal color reactions (after clearing in conc. lactic

acid) : Cl (conc. НСІ) : light orange-brown to medium orange; K (10% KOH) :

light orange-red to medium orange; N (conc. HNOs) :light brown; SE (Ha 504-

ethanol, 2 : 1) :light to dark red-brown. Illustration : PL. IV, fig. 11-15.

This species is similar to В. inaequalifolium and it is sometimes difficult to

distinguish the two species. In addition to the key characters, it apparently

differs from the latter by the annulus not revoluble and the laminae K orange

or orange-red, not dark red. These characters may be variable, however, and

need to be checked in additional collections when these become available.

Sterile specimens may be confused with Didymodon species with similar leaf

shapes, but the hollow-multiplex papillae will usually serve to distinguish В.

calcareum from these.

Habitat : soil, calcareous rock, 2600-2800 m elevation.

Mexican distribution : Distrito Federal, Mexico, Michoacan.

Range : Mexico and Guatemala.

Source - MNHN, Paris

BARBULA, PSEUDOCROSSIDIUM AND BRYOERYTHROPHYLLUM 21

2. BRYOERYTHROPHYLLUM INAEQUALIFOLIUM (TAYL.) ZANDER,

The Bryologist 83 : 232, 1980.

— Barbula inaequalifolia Tayl., London J. Bot. 5 : 49, 1846, basionym.

Plants turf-forming, green above, yellow- to red-brown below. Stems often

branching, orange-brown, to 1.5 cm long, in transverse section rounded penta-

gonal, central strand distinct, cortex of smaller, thicker-walled cells, hyalo-

dermis absent; axillary hairs of 6-9, hyaline, uniseriate cells. Leaves when dry

appressed, incurved, when wet erect-spreading, ovate to ovate-lanceolate,

1.0-2.0 mm long, narrowly grooved along costa; leaf margins revolute to apex,

entire; leaf apex rounded to obtusely acute; leaf base ovate to oblong; costa

evenly thick to apex, decurrent at base, ending 1-2 cells below apex, adaxial

surficial cells quadrate, papillose, abaxial surficial cells, elongate, papillose,

transverse section of costa elliptical, adaxial surface convex, ad- and abaxial

epidermises differentiated in one layer, lumens of abaxial epidermis elliptical

to semicircular, ad- and abaxial stereid bands strong, guide cells 4 in one layer,

hydroids absent; upper laminal cells subquadrate to short-rectangular, 7-10 um

wide, 1-2 :1, walls thin to evenly thickened, bulging on both sides, arranged

in longitudinal rows; laminal papillae crowded, hollow, multiplex, several salients

per lumen; basal laminal cells differentiated across leaf base, rectangular, 11-

15m wide, 3-4:1, walls thin to evenly thickened Propagula unicellular,

angular-spherical to elliptical, 11-22-35) um in diameter, borne serially on

branching stalks in axils of upper leaves, forming a massive, brown to black

mass in the axil.

Dioicous. Perichaetia terminal, inner leaves larger, little different in shape

from cauline leaves, to 2.5 mm long, clasping seta, perigonia terminal, sessile

оп soil, gemmate. Sporophyte seta one per perichaetium, (0.7-)2.0-2.5 cm long,

yellow to red-brown, twisted clockwise below, counterclockwise above; theca

1.5-3.0 mm long, red-brown, smooth when dry, cylindrical, neck weakly diffe-

rentiated; exothecial cells rectangular, 22-25 um wide, 3-4 : 1, walls moderately

thickened, stomates at base of theca, phaneropore; annulus of 2-3 rows of revolu-

ble, deciduous, vesiculose cells; peristome twisted counterclockwise 2-4 times,

of 32 filamentous, orange, densely spiculose teeth, 500-950 um long, with

many articulations, basal membrane low, 45-85 um high, spiculose; operculum

long-conic, 0.8-1.3 mm long, cells twisted counterclockwise. Calyptra not seen.

Spores light yellow, essentially smooth, 9-11 um in diameter. Laminal color

reactions (after clearing in conc. lactic acid) : Cl (conc. НСІ) : green + medium

orange- to yellow-brown; К (10% KOH) : dark red; М (conc. HNO3) : light

red-brown to medium orange-brown; SE (H2SO4-ethanol, 2:1) : green +

dark red. Illustration : Pl. IV, fig. 16-19.

The axillary masses of unicellular propagula easily distinguish this species

from other taxa of Bryoerythrophyllum, except B. calcareum, which has ligu-

late leaves with thick-walled marginal cells. Unicellular propagula are apparen-

tly rare in the Pottiaceae; Husnotiella revoluta Card. has similar masses of

unicellular propagula in the leaf axils, but is distinguished by the costa with

Source : MNHN, Paris

22 RH. ZANDER

only one stereid band, the laminal cell walls evenly thickened, and papillae

solid, low, broad, simple to multiplex, usually lens-like.

Habitat : soil, rock, walls, 700-3100 m elevation.

Mexican distribution : Chiapas, Mexico, Michoacán.

Range:U.S.A. (North Carolina) Mexico, Panama, Venezuela, Colombia,

Ecuador, Canary Islands, Java, China, India.

ACKNOWLEDGEMENTS. —I thank the curators of the herbaria mentioned in the

text for loan of specimens. I am grateful to M. R. Crosby, G. S. Mogensen and W. C. Steere

for discussions. The illustrations were prepared by P. M. Eckel.

REFERENCES

CHEN P.-C., 1941 — Studien über die ostasiatischen Arten der Pottiaceae, I-II. Hedwigia

80 : 1-76; 141-322.

SAITO K., 1975 — A monograph of Japanese Pottiaceae (Musci). J. Hattori Bot. Lab. 39:

373-537.

STEERE W.C., 1937 — Mosses of British Honduras and the Department of Peten, Guate-

mala. П. Ann, Bryol. 10 : 115-123.

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

Club 42 : 393-404, pl. 21-25.

ZANDER RH., 1978a— New combinations in Didymodon (Musci) and a key to the

taxa in North America north of Mexico. Phytologia 41 : 11-32.

ZANDER RH., 1978b — A synopsis of Bryoerythrophyllum and Morinia (Pottiaceae) in

the New World. Bryologist 81 : 539-560.

ZANDER RH, 1979 — Notes on Barbula and Pseudocrossidium (Bryopsida) in North

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

ZANDER RH., 1980 — Acid-base color reactions : the status of Triquetrella ferruginea,

Barbula inaequalifolia and B. calcarea. Bryologist 83 : 228-233.

Source : MNHN. Paris

ULTRASTRUCTURE OF THE SPOROPHYTE FOOT

IN PHAEOCEROS

R. GAMBARDELLA, R. LIGRONE and R. CASTALDO *

SUMMARY, — The ultrastructure of the sporophyte foot of Phaeoceros laevis (L.) Prosk.

subsp. laevis Prosk. (Anthocerotopsida) is described. In the foot a central bulbous region,

constituted by large vacuolated cells and without intercellular spaces, is surrounded by the

placental region. In the latter two deeply intermingled cell types were recognized, the

branched haustorial cells and the transfer cells, belonging to the sporophyte and to the

gametophyte respectively. This was ascertained on the basis of the plasmodesma distribution

and of other ultrastructural features, mainly of the plastids. The two generations are sepa-

rated in the placental region by an extensive system of intercellular spaces which do not

disappear even where the transfer cells and the haustorial cells are closely approached.

Peculiar crystal bodies were observed in the intercellular spaces and in transfer cell vacuoles.

The unusual organization of the placental region is discussed in relation to the nutrient

translocation.

INTRODUCTION

The contact region between the sporophyte and the gametophyte exhibits

a distinctive differentiation in the bryophytes. In the mosses and liverworts

the two generations are sharply outlined in the placental region (ROSANDER

1906; LORCH 19252, 1925b, 1931; MOTTE 1928; BLAIKLEY 1933). Transfer

cells have been reported on the gametophyte and/or sporophyte side of this

region (HEBANT 1977, WIENCKE & SCHULZ 1978).

In the anthocerotes a high degree of histological variability has been reco-

gnized as for the overall structure of the mature foot, generation outlines, muci-

lage layers, and foot and gametophyte cell size differences (CAMPBELL 1918,

BARTLETT 1928, PROSKAUER 1951). Today, however, ultrastructural

informations on anthocerote foot are not available.

A Phaeoceros representative was chosen for the ultrastructural study reported

in this paper. Since in this genus the foot and gametophyte cells are closely

intermingled in the placental region (RENZAGLIA 1978) the ultrastructural

* Istituto di Botanica, via Foria 223, 80139 Napoli, Italy.

Cryptog., Bryol. Lichénol., 1981, 2, 1 : 23-45.

Source : MNHN, Paris

24 R.GAMBARDELLA, R. LIGRONE and R. CASTALDO

observation may provide more precise criteria for ascertaining the relationships

between the cells of the two generations.

MATERIAL AND METHODS

Phaeoceros laevis (L.) Prosk. subsp. laevis Prosk. grows spontaneously in the

Botanical Gardens of the University of Naples. The sporophytes develop from

February to May.

Feet of sporophytes about 1 ст emerging from the involucre were isolated

with a razor blade together with a small portion of the surrounding gametophyte

parenchyme. After careful washing the samples were fixed either with 3%

glutaraldehyde in 0.05 M phosphate buffer рН 7 or 2% glutaraldehyde + 1%

acrolein in 0.05 M cacodylate buffer pH 7 for 2 hours at room temperature.

In both cases the samples were postfixed for 2 hours at room temperature with

2% OsO4 + 0.7% Ka Cr Oz in the suitable buffers. The samples were then dehy-

drated in ethanol and propylene oxide and embedded in the SPURR's resin

(1969). Ultrathin sections, cut by a diamond knife, were stained with uranyle

acetate and lead citrate and observed by a Siemens Elmiskop IA electron micro-

scope.

Sections 2-3 um thick, obtained by glass knives and stained with 0.1% tolui-

dine bleu, were observed by a Leitz Dialux 20 EB light microscope.

The ultrastructural observations were made at the Centro di Studio di Micro-

scopia Elettronica of the Faculty of Sciences (University of Naples).

RESULTS

Optical observations (Pl. I)

The foot of P. laevis is formed by a central bulbous region surrounded by a

well developed placental region (fig. 1). The former consists of large vacuolated

cells and lacks intercellular spaces. In the latter numerous small cells provided

with wall protuberances (transfer cells) are intermingled with oblonged and

irregularly branched cells rising from the central region of the foot (fig. 2).

These cells are termed haustorial cells, owing to their haustorial-like projections.

The placenta is rich in intercellular spaces, whose content is intensely stained

with toluidine bleu. Close to the placenta the gametophyte parenchymatous

cells show a flattened shape, especially at the bottom of the foot (fig. 1).

Ultrastructural observations (Pl. П-ХІУ)

Central region

"The central region of the foot (PL, II) is constituted by large parenchymatous

cells and, according to the light microscope observations, lacks intercellular

spaces. The cells have thin unilayered walls and usually contain one large vacuole

and abundant lipid reservoirs. Typically their plastids have an irregular shape,

contain many starch granules and a scarcely developed lamellar apparatus and

Source - MNHN. Paris

THE SPOROPHYTE FOOT IN PHAEOCEROS 25

lack the pyrenoid. Furthermore these plastids often show large membrane-

bound cavities filled with a granular content less electrondense than the stroma

(fig. 3). The plastids of the uppermost cells of the central region have a lamellar

apparatus more developed than those of the inner and peripheral cells (fig, 4).

Placental region

Transfer cells. — The transfer cells have a polyhedral shape with variable but

generally small sizes. Their wall protuberances usually are on the cell sides

towards the intercellular spaces (fig. 5, 6, 7, 11). At high magnifications these

protuberances appear to be formed by a central electrondense core surrounded

by a sheath of concentrically arranged fibrils (fig. 8). These fray at the periphery

and give rise to a loose net occupying a large light area delimited by the plasma

membrane (fig. 9). This area extends along all the inner wall surface and is

interrupted only at the level of the plasmodesmata (fig. 10).

Typically the transfer cell plastids have an oblonged and irregular shape,

do not contain starch granules and show a greatly reduced lamellar apparatus.

This is constituted by short thylakoids filled with an electrondense matrix

and arranged in small grana among which several plastoglobules are present.

The stroma occupies most of the plastid lumen. No pyrenoid was ever observed

in such plastids (fig. 12, 16).

The plastids of the more external transfer cells retain their oblonged shape

but show a rather developed lamellar apparatus and sometimes contain starch

granules and a small pyrenoid (fig. 14, 15). So they appear more similar to the

chloroplasts of the surrounding parenchymatous cells of the gametophyte

(fig. 7, 13).

The cytoplasm of the transfer cells contains an abundant reticulum, several

dictyosomes and numerous spheroidal mitochondria. The fairly developed

vacuolar apparatus is constituted by several extremely irregular vesicles where

numerous cytoplasmic digitations penetrate (fig. 6). The nucleus shows an

evenly granular nucleoplasma with a small nucleolus and is sometimes associated

with rough endoplasmic reticulum (fig. 12).

Haustorial cells. — These are oblonged cells always devoid of wall protube-

rances and larger than the transfer cells, among which they creep branching out

irregularly (fig. 5, 7).

The haustorial cells contain one or few large vacuoles that push the cytoplasm

close to the cell walls. Most of their cytoplasm, however, is gathered at the

branch extremities (fig. 5, 11). These are rich in mitochondria, dictyosomes and

rough endoplasmic reticulum and often appear, in the sections, quite isolated

inside the intercellular spaces (fig. 17, 19). The nucleus show a well developed

nucleolus and is often associated with regularly arranged sheets of rough endo-

plasmic reticulum (fig. 18).

The plastids of the haustorial cells are somewhat similar to those of the cen-

tral region cells. In fact they have a spheroidal shape, are quite devoid of grana

and contain several starch granules. These are apparently bound by one mem-

brane and partly or completely surrounded by thylakoids filled with a dark

material (fig. 21, 22). One or more electrondense areas often can be seen at the

Source : MNHN. Paris

26 R.GAMBARDELLA, R. LIGRONE and R. CASTALDO

periphery of the plastids (fig. 19, 21). They seem constituted by an irregular

network of anastomosed tubular vesicles, by plastoglobules and by a dense

granular material that reminds the pyrenoidal matrix (fig. 23), Nevertheless

à typical pyrenoid was never seen in such plastids. The stroma has а granular

appearance and contains numerous oblonged or spheroidal vesicles apparently

rising from the inner membrane of the plastid envelope (fig. 19). Membrane-

bound cavities like those of the plastids of the central region cells sometimes

were observed in the haustorial cell plastids (fig. 18).

Intercellular spaces. — The intercellular spaces constitute an uninterrupted

system of intercommunicating cavities (fig. 5) filled with a loose net of fibrillar

material and. numerous dense bodies that show a crystalline substructure at

high magnifications (fig. 25, 27). The size of these bodies is very variable, ran-

ging from 0.1 um or less to 1 um or more. In longisection they appear to be

constituted Бу а number of parallely arranged electrondense bands about 5 nm

thick and separated from one another by light spaces about 6 nm thick (fig. 28,

30). In transection an hexagonal symmetry can be recognized in the arrangement

of these bands (fig. 29, 31). Crystal bodies have often been seen also inside

transfer cells, where they are within vacuoles (fig. 25, 26).

The intercellular spaces often contain large membrane-bound cavities and

numerous other irregularly scattered membranous structures (fig. 24). The haus-

torial cell walls towards the intercellular spaces are externally covered by a

compact fibrillar layer about 50 nm thick separated by a light space 10-15 nm

thick (fig. 9). In these regions therefore the haustorial cell walls appear bilayered

(fig. 19, 32). The external layer disappears where the haustorial cells are adjoi-

ned each other. The transfer cell walls facing the intercellular spaces usually

lack such external layer but fray peripherically and give rise to a loose net that

mixes with the intercellular material (fig. 9, 32).

Symplastic connections

"The presence and distribution of the plasmodesmata were carefully investi-

gated on a great number of sections. Our observations showed that plasmodes-

mata are present between the haustorial cells and between the transfer cells,

but not between the haustorial and the transfer cells. The walls of these two cell

types are always separated by intercellular spaces that may reduce greatly

but do not disappear even where these cells are tightly approached (fig. 11, 32).

‘An intercellular space also surrounds the projections of the haustorial cells that

sometimes come up to the parenchymatous cells of the gametophyte, outside

the placenta, In these points the gametophyte cells may appear even more flat-

tened (fig. 5).

On the inner side of the placenta plasmodesmata connect the haustorial

cells with the cells of the central region of the foot (fig. 20). On the other hand

the transfer cells show symplastic connections with the parenchymatous gameto-

phyte cells, on the external side of the placenta (fig. 6, 7, 11).

Source : MNHN, Paris

THE SPOROPHYTE FOOT IN PHAEOCEROS 27

DISCUSSION

The constant absence of plasmodesmata and common septa between the

haustorial and the transfer cells shows these two cell types belong to different

generations. The plasmodesma distribution indicates that the haustorial cells

belong to the sporophyte whereas, according to GUNNING & PATE (1969),

the transfer cells belong to the gametophyte. This assumption is strengthened

by the ultrastructural features of the various cell types. The haustorial and the

transfer cells always differ very markedly in their shape, wall, and plastid ultra-

structure. Furthermore there are meaningful similarities, as for the plastid

ultrastructure, between the haustorial and the central region cells, and the

transfer and the surrounding gametophyte cells, respectively. In these two cellu-

lar lines two oppositely directed gradients of more and more developed plastids

сап be recognized, that start from the placental region (fig. 21, 4, 3 and Pl. VII).

This also suggests the two generations may exercise a reciprocal influence on

the respective morphogenetic potentialities during the foot development.

The number of plastids per cell — another potentially useful criterion for

valuating whether a cell of the placental region of Phaeoceros is sporophytic

or gametophytic (SMITH 1955, BURR 1970) — was not extensively investiga-

ted. Nevertheless our sections showed that at the most two plastid profiles are

present in the foot cells, only one in the gametophyte parenchymatous cells

but one, two or more in the transfer cells. In the last case, however, probably

they are profiles of the lobes of a single irregularly shaped plastid. On the

other hand the hypothesis that the transfer cells are sporophytic must presup-

pose, on the basis of our observations, that symplastic connections link the two

generations in Phaeoceros. This appears quite unlikely especially if the develop-

mental stages of the foot formation from the zygote ate considered (CAMP-

BELL 1918, BARTLETT 1928).

Though in Phaeoceros the sporophyte and the gametophyte interpenetrate

each other deeply, the two generations remain always separated by the inter-

cellular spaces, that constitute an extensive acellular system between them.

Such organization suggests the sporophyte draws water and nutrients directly

from the intercellular spaces through the haustorial cells, whose oblonged shape

produces an extensive increase of the absorbing surface of the foot. Likely these

substances (THOMAS et al. 1978) are supplied by the transfer cells that pump

them from the gametophyte parenchyme. In this respect it is interesting to stress

the transfer cell wall protuberances are not constantly localized on a single cell

side, as in the mosses and liverworts (HEBANT 1977), but preferably on the

cell sides towards the larger intercellular spaces. This constitutes, together

with the presence of transfer cells only on the gametophytic side of the placenta,

a quite unusual situation among the bryophytes.

Another peculiarity of the foot of Phaeoceros is the presence of dense bodies

in the intercellular spaces and in the transfer cells, that have never been reported

in the gametophyte-sporophyte interphases of the bryophytes ultrastructurally

Source : MNHN, Paris

28 R. GAMBARDELLA, R. LIGRONE and R. CASTALDO

investigated up to date (EYMÉ & SUIRE 1967, MAIER 1967, KELLEY 1969,

HÉBANT 1975, WIENCKE & SCHULZ 1975, 1978). The crystalline structure

of these bodies suggests they are proteinaceous in nature. The presence of crystal

bodies inside the transfer cells may indicate these cells as the site of their synthe-

sis. The development of crystal bodies and intercellular spaces, however, could

not be recognized in this study. Crystal bodies are also present in the inter-

cellular spaces delimited only by haustorial cells. This leads to believe the

fibrillar matrix of the spaces may have a fluid consistence in vivo, which allows

their diffusion.

The apparent aptitude of the sporophyte of Phaeoceros to take up nutrients

from an acellular environment might be an example of an intermediate evolu-

tionary stage toward the trophic independence of the sporophytic generation.

Our results provide further confirmation of a phyletic gap between the

Anthocerotopsida and the other Bryophyta. Moreover the reported histological

variability of the foot in the different anthocerote genera (CAMPBELL 1918,

BARTLETT 1928, RENZAGLIA 1978) needs further ultrastructural investi-

gations.

ACKNOWLEDGEMENTS. — The authors thank the technical staff of the Centro di Studio

di Microscopia Elettronica of the Faculty of Sciences (University of Naples) for their

competent and kind assistance.

REFERENCES

BARTLETT EM., 1928 — A comparative study of the development of the sporophyte in

the Anthocerotaceae, with especial reference to the genus Anthoceros. Ann. Bot.(Lon-

don) 42 : 409-430.

BLAIKLEY N.M., 1933 — The structure of the foot in certain mosses and in Anthoceros

laevis. Trans. Roy. Soc. Edinburgh 57 : 699-709.

BURR F.A., 1970 — Phylogenetic transitions in the chloroplasts of the Anthocerotales. 1.

The number and ultrastructure of the mature plastids. Amer. J. Bot. 57 : 97-110.

CAMPBELL DH. 1918 — The structure and development of Mosses and Ferns (Arche-

goniatae). 3rd ей. New York.

EYMÉ J. & SUIRE C., 1967 — Au sujet de l'infrastructure de la région placentaire de

Mnium cuspidatum Hedw. C.R. Acad. Sci. Paris, Sér. D, 265 : 1788-1791.

GUNNING B.S.E. & PATE J.S., 1969 — «Transfer cells». Plant cells with wall ingrowths,

specialized in relation to short distance transport. of solutes - Their occurrence, structure

and development. Protoplasma 68 : 107-133.

HÉBANT C., 1975 — Organization of the conducting tissue-system in the sporophytes of

Dawsonia and Dendroligotrichum. J. Hattori Bot. Lab. 39 : 235-254.

HEBANT C., 1977 — The conducting tissues of Bryophytes. Bryoph. Biblioth. 10.

KELLEY C., 1969 — Wall projection in the sporophyte and gametophyte of Sphaerocarpos.

J. Cell. Biol. 41 :910-914.

Source : MNHN, Paris

THE SPOROPHYTE FOOT IN PHAEOCEROS 29

LORCH W., 1925a — Ueber die Saugzellen im Fusse und in der Vaginula bei den Laub-

moosen. Ber. Deutsch. Bot. Ges. 43 : 120-127.

LORCH W., 1925b — Ueber die Haustorialschläuche am Fusse der Laubmoose. Ber. Deutsch.

Bot. Ges. 43 :262-270.

LORCH W., 1931 — Anatomie der Laubmoose. In : LINSBAUER K., Handbuch der Pflan-

zenanatomie VII. Berlin.

MAIER K., 1967 — Wandlabyrinthe im Sporophyten von Polytrichum. Planta 77 : 108-126.

MOTTE J., 1928 - Contributions à la connaissance cytologique des Muscinées. Ann. Sci.

Nat., Bot., sér. 10, 10 : 293-543.

PROSKAUER J., 1951 — Studies оп Anthocerotales. ІП. The genera Anthoceros and

Phaeoceros. Bull. Torrey Bot. Club 78 : 331-349.

RENZAGLIA K.S., 1978 — A comparative morphology and developmental anatomy of the

Anthocerotophyta. J. Hattori Bot. Lab. 44 : 31-90.

ROSANDER H.A., 1906 — Studier ofver Bladmossornas Organisation. Mössa, Vaginula och

Sporogon. Akad. Afh., Uppsala.

SMITH С.М. 1955 — Cryptogamic Botany. П. Bryophytes and Pteridophytes. London.

SPURR A., 1969 — A low-viscosity epoxy resin embedding medium for electron microsco-

ру. J. Ultrastruct. Res. 26 :31-43.

THOMAS R.J., STANTON D.S., LONGENDORFER D.H. & FARR M.E., 1978 — Physiolo-

gical evaluation of the nutritional autonomy of a hornwort sporophyte, Bot, Gaz, 139:

306-311.

WIENCKE C. & SCHULZ D., 1975 — Sporophytentwicklung von Funaria hygrometrica

Sibth. 1. Strukturelle Grundlagen der Wasser- und Nährstoffaufnahme im Haustorium.

Protoplasma 86 : 107-117.

WIENCKE C. & SCHULZ D., 1978 — The development of transfer cells in the haustorium

of the Funaria hygrometrica sporophyte. Bryoph. Biblioth. 13 : 147-167.

PLATE LEGENDS

Abbreviations. — CB : crystal body, CR : central region, CRe : central region cell, Dy :

dictyosome, ЕВА : plastid electrondense area, GP : gametophyte parenchyme, GPc : game-

tophyte parenchymatous cell, Hc : haustorial cell, HcW : haustorial cell wall, Is : intercellular

space, LD : lipid droplet, M : mitochondrion, MR : meristematic region, N : nucleus, Nu :

nucleolus, P : plastid, Pd : plasmodesma, PR : placental region, Py : pyrenoid, S : starch

granule, Tc : transfer cell, TcW : transfer cell wall, V:vacuole, WP : wall protuberance.

Pl. I. — Light microscope. 1 : Overall view of the sporophyte foot of Phaeoceros embed-

ded in the gametophyte parenchyme. It is constituted by a central bulbous region surroun-

ded by the placenta. Note the flattened shape of the gametophyte parenchymatous cells at

the bottom of the foot (x 165). 2 : Higher magnification of the placenta, constituted by

haustorial cells intermingled with transfer cells. The intercellular spaces between them are

intensely stained (x 790).

Pl. П. — Central region of the foot. 3 : Peripheral cells, containing one large vacuole

and plastids with starch granules and a scarcely developed lamellar apparatus. Large mem-

brane-bound cavities (asterisks) are often present in these plastids (x 1750). 4 : Upper-

Source : MNHN. Paris

30 R.GAMBARDELLA, R. LIGRONE and R. CASTALDO

most cells. The plastids show a more developed lamellar apparatus (x 1750).

Pl. Ill, —5 ; Placental region. This is constituted by transfer cells among which the

haustorial cells penetrate. Intercellular spaces separate these cells. The gametophyte pa-

renchymatous cells are markedly flattened in the points of contact with haustorial cells

(x 2250).

Pl, IV. — Peripheral transfer cells, Note the wall protuberances toward the intercellular

spaces. The transfer cell plastids retain a rather developed lamellar apparatus, starch granules

(fig. 6, x 6000) and, sometimes, the pyrenoid (fig. 7, x 2500). Plasmodesmata (arrows)

connect the transfer cells with each other and with the gametophyte parenchymatous cells.

Pl, V.— Details of transfer and haustorial cell walls. 8 : Transversal section of the wall

protuberances in a transfer cell (x 25000). 9 : Comparison between the transfer and the

haustorial cell walls. The external surface of the haustorial cell walls towards the inter-

cellular spaces is covered by a thin electrondense layer (arrow) that is absent on the transfer

cell walls (x 42000). 10 : High magnification of plasmodesmata between two transfer

cells. The light area extending along the inner wall surface is interrupted only at the level

of plasmodesmata (x 35000).

Pl. VI—11 : Relations between transfer and haustorial cells. The cytoplasm of the

haustorial cells is mostly gathered at the branch extremities. Large intercellular spaces

usually separates the two cell types (X 6000). 12 : Plastid and nucleus of a transfer cell.

Several regularly arranged sheets of rough endoplasmic reticulum are associated with the

nucletis (х 14000).

Pl, VIL — Gradient of the plastid ultrastructure from the typical gametophyte cells to

the inner transfer cells. The progressive reduction of the lamellar apparatus, starch content

and pyrenoid, and the increase of the plastoglobules are evident. 13 : Detail of the typical

chloroplast of a gametophyte cell (x 15000). 14 : Chloroplast of a gametophyte cell

near the placental region (x 13000). 15 : Plastid of a peripheral transfer cell (x 39000).

16 : Plastid of an inner transfer cell, containing few thylakoids filled with a dark material

and quite devoid of starch and pyrenoid (x 44000).

Pl. VII. — 17 : Branches of haustorial cells in transection. They are surrounded by

the intercellular spaces (x 5000). 18: Detail of the above micrograph, showing the nucleus

of an haustorial cell provided with a large nucleolus and associated with several regularly

arranged sheets of rough endoplasmic reticulum, Note the membrane-bound cavity (asterisk)

within the plastid (x 17000).

PI. 1X. — 19 : High magnification of an haustorial cell branch extremity in transection.

It is filled with a dense cytoplasm containing numerous mitochondria. The tangentially

cut plastid shows an electrondense area rich in plastoglobules, and several small vesicles

rising from the inner membrane of the envelope (arrows). The bilayered structure of the

wall is evident (x 20000). 20 : Septum between а haustorial cell and a central region

cell. It is crossed by several plasmodesmata (arrows) (x 12000).

PL.X.— 21 : Typical plastid of a haustorial cell. It contains numerous membrane-

bound starch granules surrounded by thylakoids filled with a dark material (x 25000).

22 : Detail of a starch granule (x 41000). 23 : High magnification of an electrondense

area, apparently constituted by irregularly anastomosed vesicles, numerous plastoglobules

and by a dense granular matrix (x 81000).

Pl. XI. —24 : An intercellular space, delimited in the section by three transfer cells.

It contains a loose fibrillar matrix, some small crystal bodies and numerous irregularly

Source : MNHN, Paris

THE SPOROPHYTE FOOT IN PHAEOCEROS 31

scattered membranous structures (х 36000). 25 : Intravacuolar crystal bodies in a transfer

cell (x 4500).

Pl. XI. — 26 : High magnification of two crystal bodies within a transfer cell vacuole.

Note their polyhedral shape (x 62000).

РІ. ХШ. — 27 : Intercellular space, filled with numerous crystal bodies which were cut

both transversally and longitudinally (x 72000). 28 : High magnification of a crystal

body in longitudinal section (x 90000). 29 and 31 : transversal sections of crystal bodies.

Note the hexagonal network (x 96000, x 163000). 30 : Detail of fig. 28 (x 187000).

Pl. XIV.— 32 : À haustorial cell closely approached to a transfer cell. An intercellular

space remains between them though it is greatly reduced in some points. Note the marked

differences in the plastid ultrastructure between the two cell types (x 18000).

Source : MNHN. Paris

PLANCHE І

Source : MNHN. Paris

34 PLANCHE Ш

Source : MNHN. Paris

PLANCHE IV

)

IBL.DU

MUSEUM

PARIS

(байв

Source ММ!

PLANCHE V

Source : MNHN, Paris

38 PLANCHE VII

Source : MNHN, Paris

PLANCHE УШ

Source : MNHN. Paris

PLANCHE IX

Source : MNHN. Paris

PLANCHE X a

aus

Source : MNHN, Paris

PLANCHE XI

Source - MNHN. Paris

PLANCHE ХП 43

Source : MNHN, Paris

44 PLANCHE ХШ

Source : MNHN, Paris

М.

818, DU

(MUSEUM

PARIS,

ж;

Source : MNHN, Paris

LA NOTION D'ESPECE

CHEZ LE GENRE COLOLEJEUNEA

Le complexe Cololejeunea floccosa (Lehm. & Lindenb.) Schiffn.

P, TIXIER *

SUMMARY, — The author divides the species C. floccosa in eight varieties viz. : C. floccosa

var. aurita, C. floccosa var. amoenoides, С. floccosa vat. floccosa, C. floccosa var. trivittata,

С. floccosa var. plicata, C. floccosa var. angustibracteata, C. floccosa var. ocellata, С. flocco-

sa vat. auriculata.

Nous étudierons la notion d'espèce à propos du sous-genre Taeniolejeunea

(Zwick.) Ben. et de son espéce-type : Cololejeunea floccosa (Lehm. & Lindenb.)

Schiffn. Rappelons que le sous-genre Taeniolejeunea est défini par le tissu

foliaire à petites cellules papilleuses, par la marge parfois hyaline, par la présence

d'une vitta constituée d'ocelles à contenu rouge-orangé.

Dans ce sous-genre, en ce qui concerne la biogéographie, on doit distinguer

des espéces et des sections à endémicité plus ou moins élevée, comme nous

l'avons écrit en 1968. Cette endémicité semble d'ailleurs être concentrée en

Asie du Sud-Est. Ailleurs on ne rencontre guére que des taxons à répartition

pantropicale comme Cololejeunea appressa (Evans) Ben. Cololejeunea floccosa

est une espèce à vaste répartition et, en principe, à définition facile. Elle a été

décrite en 1832 par LEHMANN & LINDENBERG sous le nom de Jungermannia

floccosa. Nous reverrons plus loin les tribulations de nomenclature qu'a subi се

taxon. BENEDIX, en 1953, dans son étude consacrée au genre Cololejeunea,

a tenté de le délimiter sur le plan systématique. Il a décrit trois variétés : C.

floccosa var. floccosa, C. floccosa var. aurita, C. floccosa var. conivens. L'aire de

distribution des récoltes étudiées était centrée sur la «Malesia» occidentale :

Java et Malaisie avec, accessoirement, des récoltes des Célébes et de Bornéo.

HORIKAWA (1931-1932) et SCHIFFNER (1929) ont signalé l'espèce à

Formose et au Japon. CHEN et WU l'ont isolée (1964) en Chine méridionale.

* Laboratoire de Cryptogamie, М.М.Н.М., 12 rue Buffon, 75005 Paris.

Cryptog., Bryol. Lichénol., 1981, 2, 1 : 47-76. *

Source : MNHN, Paris

48 P, TIXIER

MIZUTANI (1965) Гіпсіш dans les florules épiphylles du Sabah (Nord Bornéo).

Enfin JONES (1968) l'a trouvée dans du matériel provenant de Sierra Leone,

en Afrique de l'Ouest, Nous l'avons personnellement récoltée, au Bangladesh

(1965), en Thailande (1965), au Laos (1968), au Cambodge (1967-1970), au

Vietnam (1958-1963), aux Philippines (1965), à Java (1969-1970), en Malaisie

(1969-1972) et à Madagascar (1976-1978). Nous avons disposé, en plus, de ré-

coltes diverses : Humbert et Saboureau de Madagascar, Tirvengadum et Koster-

mans de Ceylan, des récoltes néo-calédoniennes de Huerlimann, Schmid et

Mac Kee, du matériel japonais de Hattori et de l’herbier d'Hiroshima (HIRO).

Nous débuterons, comme dans nos articles précédents, par une énumération du

matériel examiné. Le signe (*) indique qu'il ne sera pas analysé dans le texte.

MATÉRIEL EXAMINÉ

Bangladesh : Kaptai, épiphylle en forét, 50 m, 21.2.1965, Tixier 122, 123.

Cambodge : Koh Cong, Kirirom, 700 m, épiphylle en forêt, 18.4.1967, Tixier

2494 — idem, Peach Nil, 120 m, épiphylle en forêt, 20.5.1967, Tixier 2383 (*)

- idem, 5: Ambel, 20 m, épiphylle en forêt dégradée, 19.4.1968, Tixier 3712 —

Kampot, Mt Bokor, piste de Kamchay, épiphylle en forét, 900 m, 18.11.1967,

Tixier 2767 (*) — idem, 29.3.1968, Tixier 3522, 3523, 3524, 3525, 3526,

3527, 3528 — idem, 1969, Tixier 4293, 4544.

Ceylan : Southern Province, Galle district, Hiniduma Kande, 400 m, épiphylle

sur Litsea longifolia, 6.10.1971, Tirvengadum & Kostermans 620 (a, b), 650 —

idem, Kennelyia forest, clairière en basse région, 7.10.1971, Tirvengadum 681.

Japon : Riou Kiou, Okinawa, Hanjéi, sur fougères, 25.6.1961, Kora s.n. —

Kiou Siou, Kumamoto Pref, Kuma-gum, Ichibu, 11.60, Mayebara 2691 —

Hondo, Hyogo Pref., Kato-gum, Yaschiro Sho, base de tronc, 150 m, 1.3.1964,

Kodama 23681 — Mt Ishushima, ex hb. Schiffner — Hondo, Aki prov., Mt

Misen, ile Miyajima, 23.4.1931, Horikawa, HIRO 3378 — idem, 20.9.1931,

Iwamasa, HIRO 5496 — idem, Iwasama 1025, HIRO 5489 — Shikoku, prov.

Tosa, Honokawa-yama, Yoshigana 6, HIRO 5519 — Kiou Siou, prov. Higo,

Mt Ichibara, 22.3.1933, K. Mayebara, HIRO 10869 — Riou Kiou, Okinawa,

Haneji, sur fougères, 21.5.1931, Kora s.n. — idem, Okinawa, Mt Gengadaké,

30.3.1931, Maki, HIRO 3601 — idem, Okinawa, Mt Benoki, 28.3.1934, Horika-

wa, HIRO 13178 — idem, Okinawa, Mt Yonaha, 27.7.1933, Tamura, HIRO

12218 — Riou Kiou, île Iriomoté, Mt Sonaidaké, 6.4.1934, Horikawa, HIRO

13433 — idem, Mt Fedov, 3.4.1934, Horikawa, HIRO 13362 — Riou Kiou,

ile Ishigaki, Mt Bannadaké, 31.3.1934, Horikawa, HIRO 13281 — Formose,

Prov. de Taihoku, Urai, 7.1928, Noguchi, HIRO 6202 — Botel Tobago, 17.6.

1934, Fukuyama & Suzuki, HIRO 14498.

Java : Tjibodas, 1400 m, épiphylle en forét, 15.4.1969, Tixier 4164, 4349.

Laos : Paksé, Paksong, 1100 m, épiphylle en forêt inondable, 30.12.1968,

Tixier 4156, 4159, 4161.

Source : MNHN, Paris

COLOLEJEUNEA FLOCCOSA 49

Madagascar : Diégo-Suarez, massif du Marojejy, Ouest d'Andapa, Haute Am-

batorahana, bassin inférieur de la Antasakaberoka, 1000 m, 9.11.1959, Humbert

& Saboureau s.n. — Tananarive, Ambatoloana, reste de la forêt primaire, 1250 m,

18.5.1976, Tixier 8347 — Tamatave, Ivaloina, 100 m, 17.2.1977, Tixier s.n.

Malaisie : Kedah, Kedah Peak, 800 m, épiphylle en forét, 13.4.1972, Tixier

5965, 5967, 5971 — Pérak, Maxwell's Hill, 900 m, épiphylle en forêt, 17.4.

1972, Tixier 6128, 6136 — idem, 1100 m, épiphylle en forêt sous falaise, 18.4.

1972, Tixier 6154, 6153, 6156 - Pulau Pinang, Penang Hill, 700 m, bas fond à

Cyathea , 13.4.1971, Tixier 5544, 5545, 5546, 5547, 5549, 5551, 5552, 5586,

5590 — Sélangor, Fraser's Hill, 1300 m, épiphylle en forêt, 26.12.1969, Tixier

4688 — Johore, Mt Ophir, 300 m, épiphylle en forêt secondaire au bord du lac,

20.4.1972, Tixier 6246.

Nouvelle-Calédonie : Dumbéa, vallée de la Sunshine, vers le Mt To, 700 m,

27.7.1951, Huerlimann 2748 — Наше Yaté, Rivière Bleue, épiphylle en forêt,

16.6.1974, Schmid s.n. —Riviére Thi, épiphylle en forêt, 11.1974, Schmid s.n.

- Poindimié, Povila, 400 m, forêt humide sur schistes, épiphylle sur Freycinetia,

26.9.1977, Мас Kee 33858 pp — Base du col de Negacia, versant de Camboui,

150 m, s.d., Veillon sn. — Touho, Ponandou, forêt galerie, sur schistes, 30 m,

épiphylle sur Psychotria sp., 22.3.1975, Mac Kee 29906.

Nouvelle-Guinée : Bismarck Gebirge, 30 m, 18.10.1899, Lauterbach 3115a

(ex hb Stephani 19123).

Philippines : Luçon, Laguna, Mt Maquiling, épiphylle en forêt, 1000 m,

20.5.1965, Tixier 1194 — idem, Sorzogon, ex hb Lindenberg 6180 (ex hb Ste-

phani 19121) (isotype).

Thailande : Nakhon Ratchasima, Khao Yai, épiphylle en forêt galerie, 700 m,

20.3.1965, Tixier 1194 (*) — Surat Thani, Thanom Phrao, 40 m, épiphylle en

forét dense, 18.4.1965, Tixier 677, 678.

Vietnam : Khanh Hoa, Suoi Lau, 100 m, 30.10.1959, Tixier s.n. — Lam

Dông, Bao Lóc, 700 m, 20.3.1958, Tixier s.n. — idem, 18.11.1959 (a, b, c),

Tixier s.n. — idem, 3.12.1959, Tixier 2278 — idem, épiphylle en forét, 14.1.

1960, Tixier s.n. — idem, forét du pk 184, 2.1.1960, Tixier s.n. — idem, 10.1.

1960, Tixier s.n. — idem, derriére l'Ecole d'Agriculture, 11.1.1960, Tixier s.n.

idem, 21.1.1960, Tixier s.n. — idem, piste forestière de la Dalao, 11.11.1960

et 22.3.1961, Tixier s.n. — idem, Da Brinh, 700 m, 30.11.1960, Tixier s.n. -

idem, Bao Lóc, 700 m, en forét dense, 1.6.1961, Tixier s.n. — idem, Santa

Maria, 150 m, 17.1.1961, Tixier s.n. - idem, Col des Bananiers, 120 m, 3.3.

1961, Tixier s.n. — idem, Klong Riol, 600 m, épiphylle en forêt galerie, 10.3.

1961, Tixier sn. — Tuyên Duc, Dalat, station forestière de Manline, épiphylle

en forêt, 20.1.1958, Tixier s.n. — idem, Dalat, Mt Lang Bian, 1900 m, épiphylle

en forêt, 1958, Tixier s.n. — Long Khanh, Dinh Quán, pk 128 de la route Sai-

gon-Dalat, 100 m, 3.3.1962, Tixier s.n. — Ninh Binh, parc de Сис Phuong,

épiphylle en forêt, 20.11.1965, Pocs, Tran Ninh et Bich 3049/3.

En faisant abstraction du nombre exact de récoltes, nous avons examiné

Source : MNHN, Paris

50 P.TIXIER

du matériel provenant d'une soixantaine de localités et revu la plupart du ma-

tériel des grands herbiers. Signalons cependant que l'herbier Stephani contient

sept échantillons sous le binôme Cololejeunea floccosa. Seuls les numéros

19121 et 19123 font partie du complexe, 19122 et 19123 appartiennent au

complexe Cololejeunea mutabilis Ben., 19124 et 19126 contiennent Colo-

lejeunea appressa (Evans) Ben., et 14342 est un taxon proche de Cololejeunea

bontocensis P. Tx.

DESCRIPTION BIBLIOGRAPHIQUE DE L'ESPECE

Dans la bibliographie l'espèce est définie de façon taxonomique et iconogra-

phique comme suit :

Cololejeunea floccosa (Lehm. & Lindenb.) Schiffn., Conspect. Hep. Archip.

Ind. : 243, 1898.

Synonymes : Jungermannia floccosa Lehm. & Lindenb. in Lehm, Pugillus

5 :26, 1832.

Lejeunea floccosa (Lehm. et Lindenb.) Lehm. et Lindenb. in Gott., Lindenb.

et Nees, Syn. Hep. : 324, 1845.

Symbiezidium floccosum (Lehm. et Lindenb.) Trev., Mem. Ist. Lombardo,

ser. 3,4 :403, 1877.

Leptocolea floccosa (Lehm. et Lindenb.) Steph., Sp. Hep. 5 : 850, 1916.

Taeniolejeunea floccosa (Lehm. et Lindenb.) Zwick., Ann. Bryol. 6 : 107,

1935.

Icones : BENEDIX, Feddes Rep. 134, tab. 4, a-d, 1953 — CHEN & WU,

Acta Phytotax. Sin. 9 (3) : 247, fig. 15, 1-8, 1963 — MIZUTANI, J. Hattori

Bot, Lab. 24 : 275, fig. 37, 20-29, 1961 — JONES, Trans. Brit. Bryol. Soc.

3 (3) : 37, fig. 5, ab, 1968 — IWATSUKI & MIZUTANI, Bryophytes of Japan :

351, fig. 181, 1972. On peut remarquer, au passage, que cette espéce promenée

de genre en genre, décrite en 1832, n'a été dessinée qu'en 1953!!!

MORPHOLOGIE ET CARACTERES DE L'ESPECE

Nous donnons ici une description succincte et générale du taxon : plante

petite, à l'état sec blanchátre (d'oà le nom), épiphylle, appliquée au support.

Cellules marginales petites, carrées, papilleuses. Cellules basales plus allongées.

Vitta d'au moins quatre cellules. Lobe arrondi, dissymétrique ne recouvrant

pas la tige, avec ou sans sinus entre la caréne et la marge inférieure du lobe.

Lobule en sac, dent apicale généralement nulle, dent médiane composée d'une

cellule ronde et d'une cellule allongée plus ou moins en crochet. Nous ne pen-

sons pas que le type soit fructifié. Nous reviendrons sur la morphologie du

périanthe. Plante monoique ou synoïque. Propagules presque toujours à 16

cellules. L'espèce appartient à la section Floccosae Ben. Elle est définie par la

forme de la feville, la forme de la dent médiane et la longueur de la vitta.

La vitta demeure une caractéristique du sous-genre Taeniolejeunea (Zwick.)

Source : MNHN, Paris

COLOLEJEUNEA FLOCCOSA 51

Ben. mais on la retrouve chez certaines sections du sous-genre Lasiolejeunea

Ben. Nous avons décrit une vitta à 1-3 cellules chez Cololejeunea schmidtii

Steph. (TIXIER 1977). On la rencontre aussi chez Cololejeunea preciosa Ben.,

ces deux espéces appartenant respectivement aux sect. Globigerae et Venustae

du sous-genre Lasiolejeunea Ben. La vitta de C. floccosa reste proche des vitta

de C. amoena, de C. appressa et de C. gynophthalma. Elle se différencie de

celles de C. mutabilis Ben. et C. falcata (Horik.) Ben. par sa moindre largeur.

Le sinus de la base du lobe, entre la caréne et la marge inférieure du lobe,

est nul ou faible ce qui sépare C. floccosa de C. mutabilis, C. appressa, C. gy-

nophthalma et C. falcata. Nous avons décrit, de Malaisie, C. mutabilis fo. Йоссо-

soides qui diffère de C. floccosa par la largeur de sa vitta.

L'organisation de la dent médiane éloigne C. floccosa de C. appressa, de С.

gynophthalma et de C. falcata où Гарех du lobule est organisé de façon diffé-

rente. Cette méme morphologie la rapproche de C. amoena (espèce à sinus

entre marge et caréne marqué) et de C. tenella (sect. Globigerae du sous-genre

Lasiolejeunea) qui ont la méme organisation de l'apex du lobule mais un tissu

cellulaire différent, à grosses papilles, avec ou sans vitta. Enfin C. mutabilis et

ses différentes formes possèdent une dent médiane proche de celle de C. floccosa

mais cette dent est généralement obtuse.

Nous voyons que les sections Floccosae et Falcatae du sous-genre Taeniole-

jeunea et Globigerae du sous-genre Lasiolejeunea (au moins au niveau de cer-

taines espéces) forment un tout continu.

Nous allons examiner les caractéres suivants :

Morphologie de la feuille : 1 — forme, 2 — taille, 3 — distance entre les

feuilles, 4 — rapport longueur du lobule/longueur de lobe.

Tissu foliaire : 5 — cellules marginales, 6 — longueur de la vitta, 7 — largeur

de la vitta, 8 — ornementation des ocelles, 9 — organisation de Гарех du lobule,

10 — propagules.

Périanthe : 11 — rapport entre la longueur du périanthe et la longueur des

bractées, 12 — morphologie de Гарех du lobule des bractées périanthaires,

13 — présence ou absence d’oreillettes, 14 — plis ventraux, 15 — présence

d'ocelles sur le périanthe, 16 — forme du périanthe, 17 — épi mâle.

Nous allons reprendre les différents caractéres selon l'ordre indiqué. La

végétation du genre Cololejeunea est assez connue. Les tiges immatures demeu-

rent généralement peu ramifiées; les ramifications proviennent de l'apparition

des périanthes sous lesquels naissent les innovations sympodiales. La protandrie

est souvent de mise surtout dans le sous-genre Pedinolejeunea (Ben.) Mizutani.

Elle est parfois trés précoce, puisque les bractées máles fertiles apparaissent

dès le troisième étage de feuilles sur le rameau issu du propagule. Chez certaines

espèces, il existe un polymorphisme lié à la sexualité, les rameaux juvéniles

étant entièrement constitués de feuilles bractéales mâles (cf. Cololejeunea

abnormis Mizutani). Par ailleurs, dans le sous-genre Lasiolejeunea, la fructifica-

tion mâle et la fructification femelle apparaissent la plupart du temps à l'extré-

mité de rameaux secondaires.

Source : MNHN, Paris

52 P. TIXIER

1 — Forme des feuilles. En principe les feuilles de Cololejeunea Йоссова sont

elliptiques avec une légère dissymétrie et sans sinus entre la carène du lobule

et la marge inférieure du lobe. BENEDIX (1953), dans sa clé, sépare les Flocco-

sae des Falcatae par l'existence d’un sinus chez les espèces de la seconde section.

Dans un certain nombre de taxons, ceux 4 longue vitta spécialement, on trou-

verait une légère inversion entre la courbe de la caréne et la courbe de la marge

inférieure de la feuille. Les caractères concernant les sinus ne sont pas absolu-

ment constants, pas plus d’ailleurs que la forme de la feuille qui passe parfois à

une forme subtriangulaire.

2 — Taille des feuilles. Nous avons déjà expliqué que la taille des Lejeunéacées

épiphylles est d’un médiocre intérêt taxonomique. On peut, en effet, rencontrer

des «formes miniatures» des taxons typiques sans que l’on puisse en tirer des

conclusions d'ordre systématique. La longueur du lobe varie entre 0,5 mm et

0,2 mm. Notons cependant que les échantillons japonais et néo-calédoniens,

qui correspondent aux limites de la distribution, ont des lobes généralement

plus petits (0,3 mm).

3 — Distance entre les feuilles. Elle est généralement de 0,2 mm (mesurée de

caréne à carène ou de style à style). Ce caractère, d'une façon générale, est très

constant chez les «Paradoxae » et présente un intérét taxonomique certain.

4 — Cellules marginales. Elles sont petites, généralement isodiamétriques et

plus où moins papilleuses. Les papilles apparaissent, en effet, plus ou moins loin

de la marge (jusqu'à trois rangs de la marge). Quelques échantillons ont aussi,

réparties plus ou moins groupées le long de la marge, des cellules rectangulaires

plus longues qui représentent une amorce vers l'organisation d'une marge semi-

hyaline. Nous n'insisterons pas sur la distinction se rapportant à la grosseur des

papilles, ni sur le fait que les papilles se rencontrent plus ou moins bas le long de

la vitta.

5 — Longueur de la vitta. La vitta se compose d’ocelles plus ou moins remplis,

non pas comme chez la plupart des Lejeunéacées d'un ou de plusieurs globules

d'essence, mais par une sorte de dépót de couleur plus ou moins orangée à la

loupe binoculaire ou au microscope, les ocelles n'ayant peut-étre pas la méme

signification chimique chez tous les genres (nos observations ont toujours

porté sur du matériel frais ou quasiment frais). La vitta est rectiligne quand

elle est courte, plus ou moins sigmoide quand elle s'allonge. Sa longueur s'étage

de 4 cellules et de 130-160 um à 7-8 cellules et jusqu'à 420 um. Nous avons,

arbitrairement délimité trois classes : a — 4 cellules, longue jusqu'à 190 um;

b — 5-7 cellules, longue jusqu'à 220 à 300 um; c — 7-8 cellules, longue de plus

de 300 um. Сене division quelque peu arbitraire se complète par l'étude de la

largeur de la vitta (tab. 1).

6 — Largeur de la vitta. BENEDIX (1953) distingue une variété à vitta longue

et à une série unique d’ocelles : Cololejeunea floccosa var. floccosa; une variété à

vitta courte et à un rang d'ocelles : Cololejeunea floccosa var. aurita; et enfin

une variété à vitta composée de 2-3 rangs d'ocelles : Cololejeunea floccosa var.

conivens, sans indication précise de la longueur de la vitta. N'ayant pu voir le

Source : MNHN, Paris

COLOLEJEUNEA FLOCCOSA 53

Nombre

Longueur

de iavitta

en um

210-230

230-250

250-270

270-290

290-310

310-330

330-350

Tab. 1. — Longueur de la vitta.

type de la var. conivens, la définition de cette variété nous laisse perplexe. Le

type de l'espèce, provenant de Luçon (Philippines), possède une vitta courte,

à 4 cellules et à deux rangs d'ocelles (ou un rang d'ocelles et un rang de cellules

compagnes). Par conséquent, C. floccosa var. conivens semble être la variété se

rapprochant le plus du type. Malgré l'abondance de notre matériel, nous n'avons

jamais observé un taxon à vitta longue et sigmoïde à un seul rang d'ocelles.

Source : MNHN, Paris

54 Р.ТІХЕК

Généralement, la classe «a» ne possède qu'un rang d'ocelles (C. floccosa var.

aurita) mais parfois (Luçon, Riou Kiou), elle а deux rangs d’ocelles. Les classes

«b» et «c» possèdent 2-3 rangs d’ocelles ou plutôt un rang d'ocelles et 1 ou 2

rangs de cellules compagnes qui sont généralement plus étroites que les ocelles

s. str.

7 — Parois des cellules de la vitta. Elles peuvent étre minces ou munies de

trigones et d'épaissements intermédiaires. La classe «a» possède des parois

minces, les classes «b» et «c» des parois à trigones. Les caractères «longueur

de la vitta», «doublement du nombre des rangs» et «épaisseur des parois»

semblent liés.

8- Apex du lobule. La dent apicale est presque toujours pratiquement

nulle. La papille hyaline, plus ou moins piriforme, est externe et située au-

dessus de la dent médiane. Celle-ci est généralement composée d'une cellule

basale globuleuse et d'une cellule apicale. Cette dernière est d'habitude allongée,

parfois arquée, assez raremént courte. BENEDIX а doté, dans son dessin, С.

floccosa d'une dent apicale de trois cellules. Nous reviendrons plus loin sur

ce fait, assez rarement observé.

9 — Propagules. Nous savons que le nombre de cellules des propagules est

assez constant, surtout quand il est peu élevé, et nous avons montré que dans la

section Leonidentes il existait une corrélation positive entre le nombre de

cellules du propagule et la longueur de la vitta. Le nombre de cellules du propa-

gule demeure un caractère complémentaire et ne peut, à lui seul, différencier

une espèce. Le propagule de C. floccosa apporte peu de renseignements intéres-

sants, On le rencontre assez rarement : 22 fois sur 69 échantillons examinés.

Il est composé de 16 cellules dont trois cellules «crampons».

10 — Rapport entre la longueur des bractées périanthaires et la hauteur du

périanthe. Cette caractéristique morphologique est à retenir sur le plan systéma-

tique. Nous distinguerons ici deux types d'organisation :

— le périanthe dépasse largement les bractées,

— le haut du périanthe est tangent au sommet des bractées, ou méme sous

les bractées.

11— Morphologie du lobule de la bractée périanthaire. Les lobules sont

généralement larges et arrondis. Dans une seule de nos variétés, ils se présentent

en bandelettes. Il est à regretter que la plupart des auteurs s'intéressant aux

Lejeunéacées se préoccupent assez peu de la morphologie des bractées périan-

thaires et de l'organisation de Гарех du lobule. Nous avons séparé C. vidaliana de

C. punctata et des taxons proches par la morphologie du style de la bractée

périanthaire. Dans le genre Diplasiolejeunea, au niveau du groupe des Cornutae

(TIXIER 1977), le haut du lobule de la bractée fournit des données intéres-

santes. Dans notre étude de la section Leonidentes (TIXIER 1968), nous avons

observé des différences dans la morphologie de Гарех de la bractée périanthaire.

Nous avons, avec la restriction que les périanthes sont, hélas, peu nombreux

dans les populations de C. floccosa, deux types de lobule : haut du lobule denti-

culé, — haut du lobule non denticulé.

Source : MNHN, Paris

COLOLEJEUNEA FLOCCOSA 55

12 — Présence d'oreillettes. Les oreillettes du haut du périanthe sont fré-

quentes dans certaines sections du sous-genre Lasiolejeunea, mais généralement

moins marquées dans le sous-genre Taeniolejeunea. Ici seuls les taxons à longue

vitta peuvent étre pourvus d'oreillettes.

13 — Existence d'une aile ventrale. On trouve cette aile ventrale formée par

la confusion des plis ventraux sur les variétés plicata et ocellata.

14 — Existence de vitta sur le périanthe. BENEDIX a décrit des ocelles, ou

plutôt des vitta, sur le périanthe de Cololejeunea gynophthalma, Nous avons

rencontré cette caractéristique anatomique sur quatre des taxons à longue

vitta (classe c).

15 — Forme du périanthe. On note des formes en toupie et des formes en

cœur, selon la configuration de l'apex du périanthe.

16 —Inflorescence mále. Bien que ce groupe de taxons soit monoique ou

méme synoique, l'inflorescence mâle se rencontre rarement (6 fois sur 69

échantillons). Elle forme généralement un épi latéral, avec peu d'étages de

bractées fertiles (2 ou 3).

CLÉ DES VARIÉTÉS ET DESCRIPTION

Vitta courte, de 4 cellules, mesurant jusqu'à 190 um (a)

Papilles petites (1-2 um) 1 Cololejeunea floccosa var. aurita

Papilles grosses (3-5 um) 2 Cololejeunea floccosa var. amoenoides

Vitta moyenne, de 5-7 cellules, mesurant jusqu'à 220-300 шт (b)

Périanthe sans aile ventrale

Vitta à deux rangs d'ocelles 3 Cololejeunea floccosa var. floccosa

Vitta à trois rangs d'ocelles 4 Cololejeunea floccosa var. trivittata

Périanthe à aile ventrale 5 Cololejeunea floccosa var. plicata

Vitta longue, de 7-8 cellules, mesurant plus de 300 um (c)

Lobule des bractées périanthaires étroit

6 Cololejeunea floccosa var. angustibracteata

Lobule des bractées périanthaires large

Périanthe à aile ventrale 7 Cololejeunea floccosa var. ocellata

Périanthe à oreillettes 8 Cololejeunea floccosa var. auriculata

1 — COLOLEJEUNEA FLOCCOSA (Lehm. et Lindenb.) Schiffn. var. AURITA

Ben. Feddes Rep. 134 : 24, 1953. (Fig. 1-3).

Description. - Plante petite, épiphylle, appliquée au support. Tiges plus ou

moins ramifiées; feuilles insérées sous un angle de 70° et distantes entre elles de

0,1 à 0,2 mm. Lobe foliaire ovale à réniforme, de 0,25 à 0,4 mm de long;

lobule petit, discoide, le rapport de la longueur du lobule à celle du lobe variant

de 1/3 à 1/4. Lobule parfois réduit (échantillon laotien); léger sinus entre la

caréne et la marge inférieure du lobe. Cellules de la marge isodiamétriques,

de 7 um de diamètre. Le tissu papilleux descend pratiquement jusqu'à la moitié

de la vitta. Extrémité de la carène parfois crénelée. Vitta à 4 ocelles, longue de

Source : MNHN, Paris

56 P.TIXIER

- Cololejeunea floccosa (Lehm. et Lindenb.) Schiffn. var. aurita Ben. (Ceylan 630).

ige. 2:apex du lobe, 3 :vitta, 4-5 :lobule. 6-7 :périanthe. 8 :apex de lobule

périanthaire.

Fig

140 à 200 um. Dent apicale nulle; dent médiane à cellule terminale longue,

atteignant parfois le bord du lobe. Bractées périanthaires plus courtes que le

périanthe, à lobule peu ou pas denticulé au sommet. Périanthe еп toupie; bec

peu apparent et à sommet plan (ce dernier point n'est pas conforme à la des-

cription de BENEDIX).

Échantillons examinés. - CAMBODGE : Koh Kong, Sré Ambel, épiphylle en

forét dégradée, 20 m, 19.4.1968, Tixier 3712 (lobule de bractée périanthaire à

apex denticulé) — Kampot, Mont Bokor, piste de Kamchay, 900 m, 29.3.1968,

Source : MNHN. Paris

COLOLEJEUNEA FLOCCOSA 57

Fig. 2. — Cololejeunea floccosa (Lehm. et Lindenb.) Schiffn. var. aurita Ben. (Sré Ambel

3712). 1 feuille. 2:vitta. 3:apex du lobe, 4-5 ; lobules. 6 :épi mâle. 7 :арех du

lobule de bractée périanthaire. 8 : bractées périanthaires.

Tixier 3522.

CEYLAN : Galle District, Hiniduma Kande, Tirvengadum et Kostermans

630 (a).

JAPON : Mt Ishukushima ex hb Schiffner — Hondo, Hyogo Pref., Kato-gum,

Yashiro Sho, corticole, 150 m, 1.3.1964, Kodama 23681 — Riou Kiou, Okina-

wa, Mt Yonata, 27.7.1933, Tamura, HIRO 12218 — Kiou Siou, Kumamoto

Pref., Kuma-gum, Ichibu, 11.1960, Mayebara 2691 — Riou Kiou, Okinawa, Mt

Gengadaké, 30.3.1931, Maki 3601 — Riou Kiou, Okinawa, Mt Yonaha, 22.3.

1933, Mayebara, HIRO 10869 — Kiou Siou, ile Iriomoté, Mt Fedov, 3.4.1934,

Horikawa, HIRO 13362.

LAOS : Paksé, Paksong, épiphylle en forêt inondable, 30.12.1968, Tixier

4122.

MALAISIE : Kedah, Kedah Peak, 800 m, 13.4.1972, Tixier 5965, 5967,

5971.

THAILANDE : Surat Thani, Thanom Phrao, 40 m, épiphylle en forét, 18.5.

1965: Tixier 677.

Source : MNHN, Paris

58 P. TIXIER

VIETNAM : Lam Dông, Вао Lôc, forêt du km 184, 700 т, 2.1.1960 (а),

Tixier s.n. — idem, 19.2.1960, Tixier s.n.

Une dent médiane à extrémité composée de deux cellules s'observe assez

fréquemment chez des échantillons japonais. Cependant, ce caractère n'est

en liaison avec aucun autre caractére, en particulier de la vitta. On ne peut donc

Fig. 3. — Cololejeunea floccosa (Lehm. et Lindenb.) Schiffn. var. aurita Ben. (Hondo). 1:

tige. 2 :vitta, 3 :apex du lobe. 4-5 : lobules. (Kiou Siou), 1':tige. 2' :vitta. 3° :арех

du lobule. 4-6 :lobules.

Source : MNHN, Paris

COLOLEJEUNEA FLOCCOSA 59

guére en tenir compte au point de vue taxonomique, bien que BENEDIX y ait

attaché une grande importance en ce qui concerne le complexe mutabilis (fig. 3).

2- COLOLEJEUNEA FLOCCOSA (Lehm. & Lindenb.) Schiffn. var. AMOE-

NOIDES P. Tx. var. nov. (fig. 4).

Nous avons des doutes sur la valeur réelle de l'espéce C. amoena Ben. Notre

matériel vietnamien ne se différencie pas essentiellement des taxons qui lui sont.

sympatrides sauf pour l'agencement de la vitta (caractére mal précisé par BENE-

DIX pour C. amoena), à cellules un peu plus grandes dans le haut, et pour les

papilles, plus grosses. Les caractéres représentés dans les dessins de BENEDIX :

Fig. 4. — Cololejeunea floccosa (Lehm. et Lindenb.) Schiffn. var. amoenoides P. Tx. (Bao

Léc, 3.10.1961). l:tige. 2:feuille. 3 :vitta, 4:apex du lobe. 5-6 :lobules. 7 : pé-

rianthe. 8 : apex de bractée périanthaire.

Source : MNHN, Paris

60 P, TIXIER

sinus entre carène du lobule et marge inférieure de la feuille, densité des papilles

sur la carène, forme de l'inflorescence femelle, ne se retrouvent pas sur le maté-

riel vietnamien. Nous avons signalé, pour les Taeniolejeunea malaisiens,

combien dans certains cas la séparation de C. floccosa et de C. tenella était

difficile.

Differt a varietate anteriore cellulis magnis papillosis, lobulo elongatiore,

vitta cum pluribus ordinibus ocellarum, perianthoque cum una ruda ventrali.

Description. — Plante petite, épiphylle, appliquée au support, de la méme

taille que les autres taxons. Feuilles insérées sous un angle de 70-90° et distantes

entre elles de 0,2 mm. Lobe ovale à réniforme, avec un sinus entre la carène et

la marge inférieure du lobe, long de 0,4 mm. Lobule en sac, parfois réduit (Bao

Lóc, 5.12.1959), allongé, long jusqu'au 1/3 du lobe. Carène arrondie, plus ou

moins crénelée (pas de papilles sur la caréne, Bao Lóc, 3.11.1961). Dent apicale

nulle: рарШе hyaline piriforme au-dessus de la dent médiane, cette dernière à

cellule terminale longue. Cellules apicales du lobe de la même taille et de la

même forme que chez les variétés précédentes (7 um de diamètre). Cellules

basales un peu plus grandes. Vitta longue de 140 à 260 um, assez indistincte,

à plusieurs rangs de cellules en «escalier», certaines cellules de la vitta papil-

leuses. Nous n'avons pas observé de propagules sur ce taxon. Bractées périan-

thaires plus courtes que le périanthe. Périanthe en toupie avec une aile ventrale,

à apex aplati et à bec peu visible. Épi mâle latéral, court, à deux étages de

bractées fertiles.

Echantillons examinés. - CAMBODGE : Kampot, Mt Bokor, piste de Kam-

chay, 900 m, 29.3.1968, Tixier 3524 A.

JAPON : Riou Kiou, ile Iriomoté, Mt Sonaidaké, 6.4.1934, Horikawa, HIRO

13433.

MALAISIE : Pulau Pinang, Penang Hill, 700 m, épiphylle en bas fond à Cya-

thea, 13.4.1971, Tixier 5545, 5546, 5549, 5586 — Johore, Mt Ophir, 300 m,

épiphylle en forét secondaire en bordure du lac, 20.4.1972, Tixier 6246.

VIETNAM : Lam Dông, Bao Lôc, 750 m, épiphylle en forêt dense, 18.11.

1959, Tixier s.n. — idem, 5.12.1959, Tixier s.n. — idem, forét du pk 184, 700 m,

2.1.1960 (b), Tixier s.n. — idem, forét derriére l’École d'Agriculture, 3.2.1960,

Tixier s.n. — idem, 1.6.1961., Tixier s.n. — idem, 3.10.1961, Tixier s.n. (holo-

type, PC) — idem, Col des Bananiers, 120 m, épiphylle en forêt dense, 3.3.1961,

Tixier s.n.

3 — COLOLEJEUNEA FLOCCOSA (Lehm. et Lindenb.) Schiffn. var. FLOCCO-

SA (fig. 5 et 6) Feddes Rep. 124 : 21, 1953, pro parte.

C'est dans cette variété que se classe l'isotype de l’herbier de Genève, il faut

donc lui réserver le nom de la variété typique.

Description. — Plante petite, épiphylle, appliquée au support, de la même

taille que les taxons précédents. Feuilles insérées sous un angle de 70-90° ес

distantes entre elles de 0,2 mm. Lobe de forme ovale, long de 0,5 mm, sans

sinus entre la carène du lobule et la marge inférieure du lobe. Lobule en sac:

Source : MNHN, Paris

COLOLEJEUNEA FLOCCOSA 61

Fig. 5. - Cololejeunea floccosa (Lehm. et Lindenb.) Schiffn. var. floccosa (Sorzogon). 1:

tige. 2-3 :vitta. 4 apex du lobe. 5-6 :lobules. 7 : lobule réduit.

discoide, parfois réduit (Kamchay, Maxwell’s Hill), à carène arrondie, long de

1/3 à 2/5 du lobe. Dent apicale nulle. Papille hyaline piriforme, au-dessus de la

dent médiane; dent médiane à cellule terminale courte. Cellules de l'apex du

lobe isodiamétriques, de 7 um de diamètre; cellules basales de la méme taille,

mais un peu plus grandes sur l'échantillon malais. Vitta longue de 3-4 cellules,

avec un rang d'ocelles et un rang de cellules compagnes allongées, plus étroites

et vides. Parois de la vitta à trigones et épaississements intermédiaires. Propagules

à 16 cellules. Plante monoique. Inflorescence femelle à bractées périanthaires

plus courtes que le périanthe. Périanthe cordiforme du fait du faible développe-

ment des oreillettes, plis ventraux peu marqués, bec court. Ері axial mâle à trois

étages de bractées fertiles (Kamchay).

Échantillons examinés. - CAMBODGE : Kampot, Mt Bokor, Kamchay,

900 m, épiphylle en forêt, 29.3.1968, Tixier 3424 - idem, 1969, Tixier 4293,

4544.

FORMOSE : Prov. de Taihoku, Urai, 7.1928, Noguchi, HIRO 6203 — Botel

Tobago, 17.2.1934, Fukuyama et Suzuki, HIRO 14498.

JAPON : Hondo, prov. d'Aki, ile Miyajima, Mt Misen, 24.4.1931, Horikawa,

HIRO 3378 — idem, 20.9.1931, Iwamasa, HIRO 5496 — idem, Iwamasa 1025,

HIRO 5489.

MALAISIE : Pérak, Maxwell's Hill, 800 m, épiphylle en forêt, 17.4.1972,

Source - MNHN. Paris

62 P. TIXIER

iuo

OOE

ive

54)

ПА)

2225

oes

UOS

= ж

qui

ол

0400

Fig. 6. — Cololejeunea floccosa (Lehm. et Lindenb.) Schiffn. var. floccosa (Maxwell Hill

ge. 2 feuille. 3 sapex du lobe. 4 :vitta. 5-6 :apex du lobule. 7 : propa-

apex de bractée périanthaire.

Tixier 6128.

PHILIPPINES : Luçon, Sorzogon, ех hb Lindenberg 6180, ex hb Stephani

19121 (isotype).

4 — COLOLEJEUNEA FLOCCOSA (Lehm. et Lindenb.) Schiffn. var. TRIVIT-

TATA P. Tx. var. nov. (fig. 7 & 8).

Differt a C. floccosa vitta cum tribus ordinibus ocellorum.

Description. - Plante petite, épiphylle, appliquée au support. Feuilles insérées

sous un angle de 70-90% et distantes entre elles de 0,2 mm. Lobe petit pour le

groupe, de 0,3 mm de longueur, de forme variée, ovale, avec ou sans sinus entre

la caréne et la marge inférieure de la feuille. Lobule en sac, généralement discoi-

de, à carène arrondie, rarement crénelée (Fraser's Hill), parfois réduit (Bao Lôc).

Dent apicale nulle; papille hyaline piriforme, au-dessus de la dent médiane.

Source : MNHN. Paris

COLOLEJEUNEA FLOCCOSA 63

Dent médiane à cellule terminale courte et unique (sauf Ceylan 650, à deux

cellules terminales). Cellules de l'apex du lobe isodiamétriques, de 7 um de dia-

métre, sur plusieurs rangs sans papilles (Ceylan 650, Ceylan 681, Marojejy).

Cellules basales papilleuses jusqu'à la naissance de la vitta dans la plupart des cas

(sauf l'échantillon de Touho où les papilles ont déjà disparu dés le haut de la

vitta). Vitta sigmoide, atteignant 220 шп, composée de 7 ocelles accompagnés

d'un premier rang de cellules allongées et plus étroites, puis d'un rang de cellules

plus courtes, de longueur variable. Propagules fréquents, à 16 cellules. Taxon

rarement fructifié. Périanthe en toupie, à apex plat, situé au niveau des bractées

périanthaires. Lobule de la bractée denticulé à l'apex. Périanthe haut de 0,4 mm,

Fig. 7. — Cololejeunea floccosa (Lehm. et Lindenb.) Schiffn. var. trivittata P. Tx. (Fraser's

Hill 4634). 1 :tige. 2 :vitta, 3 : feuille. 4-5 :lobules. 6 : périanthe. 7 :épi mâle.

Source : MNHN, Paris

64 P. TIXIER

doar

Fig. 8. Cololejeunea floccosa (Lehm. et Lindenb.) Schiffn. var. trivittata Р. Tx. (Bao

Léc 2278). 1 :tige. 2 feuille. 3 сарех du lobe. 4 :vitta. 5-7 :lobules réduits. 8:

périanthe et bractées périanthaires. 9 : apex du lobule de la bractée. 10 : archégone.

à apex plan et bec court. Epi mâle latéral, court, à 3 étages de bractées fertiles.

Échantillons examinés. — CEYLAN : Southern Province, Galle District, Hini-

duma Kanda, épiphylle sur Litsea longifolia, 400 m, 6.10.1974, Tirvengadum

et Kostermans 630 (b), 650 — idem, Kennelyia Reserve Forest, clairiére еп

basse région, 7.10.1974, Tirvengadum 681.

MADAGASCAR : Diégo-Suarez, massif du Marojejy, épiphylle en forêt,

1000m, 9.11.1958, Humbert & Saboureau s.n. — Tananarive, Ambatoloana,

1100 m, 16.5.1976, Tixier 8347.

Source : MNHN. Paris

COLOLEJEUNEA FLOCCOSA 65

MALAISIE : Pérak, Maxwell's Hill, épiphylle en forêt sous falaise, 1100 m,

18.4.1972, Tixier 6154, 6153, 6156 — Sélangor, Fraser's Hill, 1300 m, épiphylle

en forét, 29.12.1969, Tixier 4634 (holotype, PC), 4688.

NOUVELLE-CALEDONIE : Touho, Ponandou, 30 m, épiphylle sur Psycho-

tria en forêt galerie, 22. 3.1975, Mac Kee 29906 — idem, 23.3.1975, Mac Kee

29917 — Poindimi Povila, forêt humide sur schistes, épiphylle sur Freycinetia,

26.9.1977, Мас Кес 33858 — Base du Col de Negacia, versant ouest de Camboui,

150, s.d., Veillon s.n. — Riviére Bleue, s.d., s.n. — Vallée de la Sunshine, 22.8.

1951, Huerlimann 2748.

VIETNAM : Lam Dóng, Bao Lóc, épiphylle en forét dense, 700 m, 5.12.1959,

Tixier 2278 — Tuyên Duc, Dalat, Mt Lang Bian, 1900 m, 1959, Tixier s.n.

5 — COLOLEJEUNEA FLOCCOSA (Lehm. et Lindenb.) Schiffn. var. PLICATA

P. Tx. var. nov. (fig. 9 & 10).

Differt a varietate typica anteriore periantho cum una ruda alata ventrali.

Description. — Plante petite, épiphylle, appliquée au support. Feuilles insérées

sous un angle de 70-907 et distantes entre elles de 0,2 mm. Lobe long de 0,3-

0,4 mm, ovale, sans sinus entre la caréne et la marge inférieure de la feuille.

Lobule en sac, discoide, à carène lisse ou crénelée (Kaptaï), parfois réduit

(Kaptai), atteignant le 1/3 de la longueur du lobe; dent apicale nulle, papille

hyaline piriforme, au-dessus de la dent médiane. Dent médiane à cellule termi-

nale longue. Cellules de Гарех du lobe de 7-10 um de diamètre, les rangs margi-

naux sans papilles. Cellules basales plus grandes, papilleuses jusqu'au milieu de

la feuille. Vitta droite, longue de 170- 250 um, à 4-8 cellules accompagnées

de 1-2 rangs de cellules allongées plus étroites, Parois des ocelles à trigones et

épaississements intermédiaires plus ou moins marqués. Propagules à 16 cellules.

Plante monoïque. Périanthe en toupie, de 0,4-0,5 mm de haut, les plis ventraux

formant une aile. Bractées périanthaires plus courtes que le périanthe; apex

du lobule périanthaire généralement denticulé. Épi mâle latéral, à 2 étages de

bractées fertiles.

Échantillons examinés.— BANGLADESH : Kaptai, épiphylle en forêt,

27.2.1965, Tixier 122.

CAMBODGE : Koh Kong, Kirirom, Phnom Prom, 700 m, épiphylle en forét,

18.4.1967, Tixier 2494.

JAVA : Tjibodas, 1400 m, épiphylle en forêt, 15.4.1969, Tixier 4349 (holo-

type, PC).

Cette variété semble étre celle qui se rapproche le plus de la variété conivens

Ben. Bien que la diagnose n'en fasse pas état, le périanthe de cette variété est

représenté, semble-t-il, avec une aile ventrale.

6. — COLOLEJEUNEA FLOCCOSA (Lehm. et Lindenb.) Schiffn. var. ANGUS-

TIBRACTEATA P. Tx. var. nov. (fig. 11 & 12)

Differt a varietate typica vitta longiore, dente apicali parva, lobuloque brac-

tearum crenulato angustoque.

Source : MNHN, Paris

66 P. TIXIER

Fig. 9. — Cololejeunea floccosa (Lehm. et Lindenb.) Schiffn. var. plicata P. Tx. (Tjibodas

4349). 1:tige. 2 feuille, 3 :арех du lobe. 4 :vitta. 5-6 :lobules. 7 : périanthe, 8:

apex de bractée périanthaire.

Description. — Plante petite, épiphylle, appliquée au support. Feuilles insérées

sous un angle de 70-90° et distantes entre elles de 0,2 mm. Lobe ovale, sans

sinus entre la marge inférieure du lobe et la caréne du lobule, relativement grand,

long de 0,5 mm. Lobule en sac, parfois réduit (Mt Maquiling), discoide, à

carène arrondie, atteignant du 1/3 au 1/5 de la longueur du lobe. Dent apicale

nulle; papille hyaline au-dessus de la dent médiane; dent médiane à cellule ter-

minale plus ou moins développée. Cellules de Гарех du lobe pappilleuses, isodia-

Source : MNHN, Paris

COLOLEJEUNEA FLOCCOSA 67

Fig. 10. — Cololejeunea floccosa (Lehm. et Lindenb.) Schiffn, var. plicata P. Tx. (Kaptai).

1:tge. 2 :feuille. 3:apex du lobe. 4:vitta. 5-6 :lobules. 7-8 :lobules réduits. 9:

périanthe. 10 :apex du lobule de bractée périanthaire.

métriques, de 7 um, avec une tendance à différencier sur la marge des cellules

rectangulaires allongées dont l'axe est dirigé dans le sens radial (comme chez

Cololejeunea peraffinis Schiffn.). Cellules basales papilleuses jusqu'à la base de

la vitta. Vitta sigmoide, constituée de 7 cellules et longue jusqu'à 300-350 um.

Ocelles accompagnés par deux séries de cellules allongées et étroites. Parois des

ocelles sans ornementation bien marquée. Propagules non observés, Plante

parfois synoique (Maxwell's Hill). Périanthe en forme de toupie, à apex aplati

Source : MNHN. Paris

68 P.TIXIER

et à bec peu marqué, haut de 0,4-0,5 mm. Bractées périanthaires atteignant plus

ou moins la hauteur du périanthe. Lobule périanthaire étroit et denticulé au

sommet. Épi mále latéral, à 2-3 étages de bractées fertiles.

Échantillons examinés.— MALAISIE : Pérak, Maxwell's Hill, 800 m, épi-

phylle en forét, 18.4.1972, Tixier 6136.

Fig. 11. — Cololejeunea floccosa (Lehm. et Lindenb.) Schiffn. var. angustibracteata P. Tx.

(Mt Maquiling). 1 tige. 2 :vitta. 3 :apex du lobe. 4 :lobule. 5 :lobule réduit. 6 : pé-

rianthe. 7 :apex du lobule de bractée périanthaire. 8 : épi mâle.

Source - MNHN. Paris

COLOLEJEUNEA FLOCCOSA 69

Fig. 12. — Cololejeunea floccosa (Lehm. et Lindenb.) Schiffn. var. angustibracteata P. Tx.

(Maxwell's Hill 6136). 1 :tige. 2 : feuille. 3 :vitta. 4 :арех du lobe, 5-7 :lobules. 8:

inflorescence synoïque.

PHILIPPINES : Luçon, Mt Maquiling, 1000 m, 10.7.1965, Tixier 1393

(holotype). Ce taxon semble le plus proche de celui représenté par BENEDIX

comme étant Cololejeunea floccosa typique.

7 — COLOLEJEUNEA FLOCCOSA (Lehm. et Lindenb.) Schiffn. var. OCELLA-

TA P. Tx. var. nov. (fig. 13 & 14).

Differt a specie typica, vitta longiore, ocellis perianthii,

Source : MNHN, Paris

70 P. TIXIER

Fig. 13. — Cololejeunea floccosa (Lehm. et Lindenb.) Schiffn. var. ocellata P. Tx. (Klong

Riol). 1 :tige et épi mâle. 2:vitta. 3:lobule. 4 :vitta du périanthe. 5 :apex du lo-

bule. 6-7 : lobules réduits. 8 : périanthe. 9 : apex du lobule périanthaire.

Description. — Plante petite, épiphylle, appliquée au support. Feuilles insérées

sous un angle de 70° et distantes entre elles de 0,2-0,3 mm. Lobe ovale, avec un

léger sinus entre la marge inférieure du lobe et la carène, long de 0,45-0,5 mm.

Lobule en sac, discoïde, à carène arrondie et lisse, parfois réduit, atteignant 1/3

Source : MNHN, Paris

COLOLEJEUNEA FLOCCOSA 71

Fig. 14. — Cololejeunea floccosa (Lehm. et Lindenb.) Schiffn. var. ocellata P. Tx. (Bao Lóc,

18.11.1959). l:tige. 2:feuille, 3 :vitta, 4:apex du lobe, 5-6 :lobules. 7 :lobules

réduits. 8 :périanthe. 9 :vitta périanthaire. 10 :épi mâle.

Source : MNHN, Paris

72 P, TIXIER

à 2/5 de la longueur du lobe. Dent apicale visible, formant épaule; papille hya-

line piriforme, au-dessus de la dent médiane. Dent médiane à cellule terminale

longue, parfois dédoublée (Bao Lóc 18.11.1959). Cellules de l'apex du lobe

isodiamétriques, de 7 um de diamètre, les papilles n'apparaissent qu'à la seconde

ou à la troisième rangée de cellules. Cellules basales papilleuses jusqu'à la base

de la vitta. Vitta grande, sigmoïde, longue de 300 à 370 um, à 7-8 grands

ocelles accompagnés de deux séries de cellules longues et étroites. Trigones

et épaississements intermédiaires marqués sur les parois des ocelles. Propagules

à 16 cellules. Périanthe cordiforme, haut de 0,4-0,5 mm, à ailes peu marquées et

plis ventraux formant aile, orné, dès la mi-hauteur, d'ocelles plus ou moins

organisés en vitta. Bractées périanthaires à hauteur du périanthe; lobule des

bractées large et denticulé au sommet. Épi mâle axial, à 3 étages de bractées

fertiles.

Echantillons examinés. - CAMBODGE : Kampot, Mt Bokor, 900 m, piste

forestière de Kamchay, 29.3.1968, Tixier 3524 B.

VIETNAM : Lam Dông, Bao Lóc, épiphylle en forêt, 20.3.1958, Tixier s.n.

idem, 18.11.1959, Tixier s.n. — idem, 11.1.1960, Tixier s.n. — idem, 19.1.

1960, Tixier s.n. — idem, 21.1.1960, Tixier s.n. — idem, piste du Da Brinh,

épiphylle en forêt, 10.11.1960, Tixier s.n. — idem, piste de la Dalao, 900 m,

11.1.1960 et 21.2.1961, Tixier s.n. — idem, Santa Maria, forét secondaire,

150m, 17.1.1961, Tixier s.n. — idem, Klong Riol, 600 m, 10.3.1961, Tixier

s.n. (holotype, PC).

8—COLOLEJEUNEA FLOCCOSA (Lehm. et Lindenb.) Schiffn. var. AURI-

CULATA P. Tx. var. nov. (fig. 15).

Differt a varietate anteriore aliis perianthii, dente apicali minore, ocellis

lobi paucioribus majoribusque.

Description. — Plante petite, épiphylle, appliquée au support. Feuilles insé-

rées sous un angle de 70° et distantes entre elles de 0,2 mm. Lobe ovale, sans

sinus entre la carène du lobule et la marge inférieure du lobe, long de 0,4 mm.

Lobule en sac, entier ou réduit, discoïde, à caréne arrondie, Dent apicale non

visible; рарШе hyaline piriforme; dent médiane à cellule terminale relativement

courte. Cellules de l'apex du lobe isodiamétriques, de 7 um, à papilles seulement

à partir du second rang de cellules. Cellules basales plus ou moins grandes selon

l'échantillon et papilleuses jusqu'à la base de la vitta, au moins du côté de la

marge supérieure du lobe. Vitta peu sigmoide, longue de 270 à 300 um. Ocelles

moins nombreux et plus grands que chez la variété précédente, accompagnés

d'un seul rang de cellules allongées. L'ornement des parois des ocelles est nul.

Propagules non observés. Périanthe à oreillettes marquées. Bec apparent et long.

Ocelles en vitta à mi-hauteur du périanthe, lui-même haut de 0,5-0,6 mm.

Bractées périanthaires à lobule peu denticulé au sommet. Épi mâle à trois étages

de bractées fertiles.

Échantillon examiné. - NOUVELLE GUINÉE : Bismarck Gebirge, 18.10.

1890, Lauterbach s.n. ex hb Steph. 19123 (holotype).

Source : MNHN, Paris

COLOLEJEUNEA FLOCCOSA

(Nile-

2:vitta. 3-4 :lobules. 5 :apex du lobe. 6 : périanthe,

face ventrale et dorsale. 7 : apex du lobule périanthaire. 8 : vitta périanthaire.

- Cololejeunea floccosa (Lehm. et Lindenb.) Schiffn. var. auriculata P. Tx.

Guinée). 1 : tige et épi mâle.

Fig. 15.

Source : MNHN, Paris

74 P. TIXIER

CONCLUSIONS

Ce travail est la somme d’une vingtaine d'années de récoltes et de réflexions.

Quelques-unes de nos observations sont en contradiction avec celles de BENE-

DIX : nous n’avons pas pu voir le matériel étudié par cet auteur. Cependant,

notre matériel a des origines plus variées et des aires de distribution plus vastes

que celui de notre prédécesseur,

Nous n'insisterons pas ici sur la spéciation et la biogéographie dans le sous-

genre Taeniolejeunea dont nous avons déjà donné une idée à propos de la

section Leonidentes. Rappelons seulement les passages entre les différents

taxons des sections Floccosae, Falcatae et Globigerae de BENEDIX. Nous

donnons (tab. 2) la répartition des différentes variétés au point de vue géogra-

phique, la variété dont l'aire est la plus étendue restant la var. aurita. Notons

aussi que c'est dans les régions oà nous avons le plus récolté que le matériel le

plus abondant a été recueilli, c'est-à-dire au Vietnam. Étant affecté à l'École

d'Agriculture de Bao Lóc, nous avons eu, durant plusieurs années, la forêt

tropicale à notre porte.

Nous terminerons par deux remarques écologiques. Comme nous l'avons

indiqué dans nos listes des mousses de Thailande (1971), il existe, sous les

Tropiques en Asie du Sud-Est, trois zones climatiques : une zone équatoriale

sans saison séche, une zone tropicale à saison séche chaude, une région tropicale

à saison sèche fraiche. Dans la seconde de ces régions se rencontre la plus forte

richesse spécifique chez les Cololejeunoidées. Cette région prend l'Asie du Sud-

Est en écharpe, des Philippines à la Malaisie. La seconde remarque concerne

l'influence de l'altitude. Dans la plupart des groupes de plantes épiphytes que

nous avons étudiées au Vietnam, la richesse spécifique passait par un maximum

dans l'étage de 500-1200 m. C. floccosa n'échappe pas à cette règle. La réparti-

tion des taxons suivant l'altitude (pour autant que nous connaissions celle-ci)

montre que les diverses variétés ont un maximum de richesse spécifique entre

600 et 1200 m. Cependant, une certaine richesse en basse région est à noter.

Les florules hépaticologiques épiphylles diminuent avec l'altitude, soit qu'elles

disparaissent, soit qu'elles s'appauvrissent. Nous basons cette opinion sur nos

récoltes au-dessus de 2000 m (2000 m au Mt Lang Bian au Vietnam, 1900 m

au Mt Polis, à Luçon, 2100 m aux Palni Hills dans le Deccan, et enfin 2100 m

dans l'Ankaratra, à Madagascar). Sur un point plus précis nous avons récolté

15 espèces de Cololejeunea au Mont Maquiling à Luçon, à 1000 m, et seule-

ment 4 au Mont Polis.

Nous remercions bien vivement S. Hattori et H. Ando qui ont eu l'amabilité de nous

prêter le précieux matériel du Japon. Notre reconnaissance va aussi à notre ami D,D. Tirven-

gadum qui nous a spécialement récolté du matériel épiphylle à Ceylan, ce qui a permis

d'accroître sérieusement la connaissance des Lejeunéacées cinghalaises (TIXIER 1977).

Nous ne pouvons pas oublier ici les contributions remarquables qu'ont constitué les récoltes

néo-calédoniennes. Enfin, nous remercions P. Geissler, conservateur de l'herbier de Genève,

qui nous a adressé le matériel de l'herbier Stephani.

Source : MNHN, Paris

COLOLEJEUNEA FLOCCOSA 75

5 H 5

БЕЛ ы 20:15

Sole са 2

Variétés үле ІНЕ ЕЕ

= | S| 9] = sS

EE ЕЛІ 2|2|8

cu ers I =] als

ORIGINES -

BANGLADESH 1

CAMBODGE . . 3

CEYLAN . 2

JAPON . б 1

JAVA б 1

Laos . 1

MADAGASCAR . 1

MALAISIE .1. . 3

Nile CALE DONIE . 1

Nile GUINEE “|1

PHILIPPINES . . 2

THALI ANDE . 1

VIET NAM ele ele lene,

БРО Е Eee cuin:

ALTITUDE

0-600m 4 3 1 1 8

400-1200 ОП ІРІП»

SE tad 2

Tab. 2. - Répartition des taxons (géographie et altitude).

BIBLIOGRAPHIE

BENEDIX EH., 1953 — Indomalayische Cololejeuneen. Feddes Керегі. 134 : 1-96.

CHEN P.C. & WU P.C., 1964 — Studies on epiphyllous Liverworts. Act. Phytotax. Sin. 9 :

137-153.

Source : MNHN, Paris

76 P. TIXIER

HORIKAWA Y., 1931 — Studies on the Hepaticae of Japan. IV. J. Sci. Hiroshima Univ.

Ser. B, Div. 2, 1 (2) :13-35.

HORIKAWA Y., 1931a — Idem. V. Ibidem 2, 1 (4) : 55-76.

HORIKAWA Y., 1932 — Idem. VI. Ibidem 2, 1 (5) : 77-94.

IWATSUKI Z. & MIZUTANI M., 1972 — Coloured illustrations of Bryophytes of Japan,

Osaka, Hoikusha : 1-405.

JONES E.W., 1968 — African Hepatics. XX. Some little known species and some extension

of range. Trans. Brit. Bryol. Soc. 5 : 563-572.

LEHMANN J.G.C., 1832 — Novarum et minus cognitarum stirpium pugillus 5 : 26.

MIZUTANI M., 1961 — A revision of Japanese Lejeuneaceae. J. Hattori Bot. Lab. 24 :113-

302.

MIZUTANI M., 1965 — Epiphyllous species of Lejeuneaceae from Sabah (North Borneo).

J. Hattori Bot. Lab. 29 : 153-172.

SCHIFFNER V., 1929 — Ueber ерірһуШе Lebermoose aus Japan nebst einigen Beobach-

tungen ueber Rhizoiden, Elateren und Brutkoerper. Ann. Bryol, 2 : 87-106.

STEPHANI F., 1912-1917 — Species Hepaticarum. 5 : 1-1044. Georg, Genève.

TIXIER P., 1968 — Cololejeunea de l'Asie du Sud-Est. I. Leonidentes et espèces affines.

Rev. Bryol. Lichénol. 36 : 343-391.

TIXIER, P., 1971 — Bryophytae indosinicae. Mousses de Thailande. Ann. Fac. Sci. Phnom

Penh 4 :91-166.

TIXIER P., 1972 - Mount Maquiling Bryoflora (Luzon). Gard. Bull. Singapore 25 :137-153.

TIXIER P., 1975 - Contribution à l'étude de la phyllosphére. Les Lejeuneacées et l'altitude

en Asie méridionale et en Insulinde. Compt. Rend. Sommaire Séances Soc. Biogéogr.

440-442 :91-95.

TIXIER P., 1977 — La notion d'espèce chez le genre Cololejeunea. C. schmidtii St. et les

taxa voisins. Rev. Bryol. Lichénol. 43 : 35-52.

TIXIER P., 1977 — Lejeuneaceae from Sri Lanka collected by D.D. Tirvengadum. Ceylon J.

Sci. Bio. Sci. 12, 2 :14-17.

Source : MNHN. Paris

CYTOLOGICAL OBSERVATIONS ON SOME

WEST HIMALAYAN MOSSES. VI

S.S. KUMAR & S.K. VERMA

SUMMARY. — The meiotic chromosomes of 2 species each of Mielichhoferia, Pohlia, and

Anomobryum, and of 5 species of Brachymenium are studied Five of the investigated

species, namely, Mielichhoferia sasaokae : п = 11, Mielichhoferia sp. :n = 10, Brachyme-

nium acuminatum :n = 10, 11, B. leptophyllum : п = 20 and B. sikkimense : n =10, are

studied cytologically for the first time. Intraspecific polyploidy and aneuploidy is recorded

in Pohlia longicollis (п —11, 20, 22, 22 + m), Brachymenium exile (n =10, 11, 20), and B.

nepalense (п = 22, 22 +m). Heteromorphic bivalents are found in Brachymenium acumina-

tum, B. exile, B. sikkimense, Anomobryum auratum and Anomobryum sp. Triads are

observed in Brachymenium exile and B. sikkimense.

The present note which deals with the cytological results in 11 species of 4

genera of the family Bryaceae is a continuation of our earlier communication

(1980, in press) on the cytology of 11 species of Bryum from the Western

Himalayas.

MATERIALS AND METHODS

The materials were collected from the Western Himalayas during the mon-

soon months (July 1975 to September 1977). The meiotic study was made by

squashing the archesporial tissue in 2% acetocarmine. The slides were made

permanent in Euparol. The vouchers are in the Herbarium of the Department of

Botany, Panjab University, Chandigarh, India.

OBSERVATIONS AND RESULTS

Mielichhoferia sasaokae Broth. : п = 11 (Figs. 1-2) — This is the first cytologi-

cal study of the genus. The investigated species (PU 3156), a new record for the

Western Himalayas, was collected from Nainital (Kilbery forest, on wet shady

rocks, 2300 т). The frequency of sporophyte formation was rather very low.

Department of Botany, Panjab University, Chandigarh 160014, India

Cryptog., Bryol. Lichénol., 1981, 2, 1 : 77-89.

Source : MNHN. Paris

78 S.S. KUMAR & S.K. VERMA

The reproduction appeared to be mainly through the perennation of old gameto-

phores. The 11-chromosome complement included three relatively large biva-

lents, one of which was noticeably larger than the others (Figs. 1-2). In some

sporocytes, a few laggards were observed at anaphase-I because of late disjunc-

tion in the first telophase nuclei. As judged from the morphology of the biva-

lents (Fig. 1), at least four chromosomes appear to be acrocentric.

Mielichboferia sp. : n = 10 (Figs. 3-4) — This unnamed species (PU 3157),

also gathered from Nainital (Kilbery forest, on rocks, 2300 m), showed abun-

dant fruiting. The chromosome complement (Fig. 3) as in the previous species,

showed three relatively large bivalents. In about 5% of the studied sporocytes,

1-2 laggards were observed at first and second telophases. In a few sporocytes

abnormal meiotic behaviour, probably caused by the weak spindle, resulted in

three apparently normal nuclei and three micronuclei (Fig. 4). It would be

desirable to undertake chromosome sampling of several populations of this

species to find out if the expected aneuploid numbers exist.

Poblia elongata Hedw. : n = 11 (Fig. 5) — The material studied (PU 3158)

was collected from Nainital (Kilbery forest, on wet soil, 2300 m). One of the

two large bivalents in the complement showed a tendency to disjoin precocious-

ly. As judged from the morphology of the bivalents, at least four chromosomes

appear to be acrocentric. Earlier, the same chromosome number was reported

for some other populations of this species (YANO 1957, PANDE & CHOPRA

1958, and SMITH & NEWTON 1967). In the European materials, however,

SMITH & NEWTON (1967) observed irregular disjunction of the bivalents.

Poblia longicollis (Hedw.) Lindb. : n = 11, 20, 22, 23 (22 + m) (Figs. 6-12) —

Three collections of this nearly cosmopolitan species gave different chromo-

some counts. The material (PU 3159) gathered from Ranikhet (on soil, 1850 m),

showed 11 bivalents at metaphase-I (Fig. 6). The smallest member of the set,

recognizable as the h-bivalent, was lightly stained and anticipated disjunction

of the set. Of the other two materials collected from Didihat (Ghurpatta,1 350 т),

one (PU 3160, on soil) showed 20 bivalents (Fig. 7) which tended to lie in pairs,

while in the other gathering (PU 3161, on dead logs of wood) 22 bivalents

were counted at metaphase-I (Figs. 9-10). As judged from the morphology

of the bivalents, four of the six large bivalents appeared to be metacentric. In

few sporocytes from some of the capsules of this population, in addition to

the normal 22 chromosomes, a faintly stained, dot-like, precociously disjoined

m-bivalent was also observed (Fig. 11). This tiny chromosome appears to be

floating in this population and is probably on its way to elimination. In 15-20%

of the studied sporocytes from eudiploid and hypodiploid gatherings, 1-3

laggards were observed: at anaphase-I (Fig. 8) and telophase-l: Sometimes an

anaphase bridge was seen at the end of the first meiotic division in the eudi-

ploid materials (Fig. 12).

KHANNA (1967) counted 11 bivalents in a population from Colorado and

YANO (1957) reported n = 22 for the Japanese population of this species.

Source : MNHN. Paris

CYTOLOGY OF SOME WEST HIMALAYAN MOSSES 79

P1. I. — 1-2 : Mielichhoferia sasaokae —n = 11. ‘1 : Metaphase-I; note the noticeably large

bivalent. 2 :Metaphase-I; the arrow marked bivalent has disjoined 3-4 : Mielichhoferia

sp.—n =10. 3 :Metaphasel. 4: Telophase-Il, proceeding towards the constitution of

3 apparently normal and 3 micronuclei. 5 : Pohlia elongata — n = 11. Metaphase-l; note

the precocious disjunction of one bivalent. 6 :P. longicollis —n = 11. Metaphase-l;

note 2 half-bivalents (arrow marked) of the h-bivalent (x 2720).

Source : MNHN, Paris

80 S.S. KUMAR & S.K, VERMA

% T S adi

20. 7 :Metaphase-l; note the tendency of a few biva-

lents to lie in pairs. 8 :Anaphase-l, showing laggards. 9-12 :P. longicollis — n = 22, 23

(22 +m). 9-10 :Metaphase-l. 11 :Metaphase-I; note two (arrow marked) half-biva-

lents of the m-bivalent. 12 : First anaphase bridge (х 2720).

PL.IL. — 7-8 : Pohlia longicollis — n =

Source : MNHN, Paris

CYTOLOGY OF SOME WEST HIMALAY AN MOSSES 81

The quantitative differences in morphological characters (plant length, seta

length and capsule size) between the haploid (n = 11, Fig. 32), diploid (n = 22,

Fig. 33) and the hypodiploid (n — 20) materials appear to be environmentally

controlled.

Of the reported chromosome numbers in 16 species of Pohlia, n = 10, 11,

10 + 4 m, 20, 21, 22, 22 + m, 30, 44 (FRITSCH 1972, DANYLKIV & VI-

SOTSKAYA 1975, VISOTSKAYA 1975), n — 11 (or its multiple) is found

in 13 taxa and may be regarded as the basic number for this genus. Further

evolution and speciation in Pohlia seems to have been accomplished through

polyploidy and aneuploidy.

Brachymenium acuminatum Hedw. : п = 10, 11 (Figs. 13-16) — Of the two

gatherings of this so far cytologically uninvestigated species, the one (PU 3162)

collected from Bhimtal (on soil, 1390 m) showed 10 bivalents with the largest,

dimorphic, H-bivalent located at the periphery and anticipating disjunction of

the set (Figs. 13-14). The smallest bivalent showed faint stainability and a

tendency towards precocious disjunction (Fig. 14). The plants of the second

collection (PU 3163) from Dharamsala (Tea Estate, 1500 m) showed 11 biva-

lents at first metaphase (Fig. 16). The largest member of the set, unlike that of

the previous population, was not heteromorphic. As judged from the morpho-

logy of the bivalents, at least five chromosomes appear to be metacentric.

Brachymenium exile (Doz. & Molk.) Bosch. et Lac. : n = 10, 11, 20 (Figs.

17-24) — Of the six populations studied, three (PU 3164 - Almora, 1605 m;

PU 3165 - Dharamsala, Tea Estate, 1500 m; PU 3166 - Narkanda, Hatoo Peak,

2700 m) showed n — 10. The complement included 3-4 large bivalents, of which

the largest one anticipated disjunction of the set (Fig. 18). The resulting half-

bivalents appeared slightly dissimilar. One large chromosome (arrow marked)

appears to be involved in the organization of the nucleolus (Fig. 17). In two

other populations (PU 3167 - Almora, 1605 m; PU 3168 - Didihat, 1450 m)

11 bivalents were counted at first metaphase (Figs. 19-20). In the Didihat popu-

lation the largest bivalent was rod-shaped (Fig. 19) and, as in materials with

n = 10, it disjuncted precociously into two dissimilar half-bivalents. Occasio-

nally the three smallest members of the set also anticipated disjunction. One or

two chromosomes were observed as laggards at telophase-I (5-8% of the sporo-

cytes) and telophase-Il (12-15% of the sporocytes) in the Almora population.

In the Didihat population, triads (Fig. 21) with one of the three nuclei (possibly

of diploid constitution) of approximately double the size of the remaining two

were observed in 5-8% of the studied sporocytes. The sixth population (PU

3169), gathered from Didihat and separated from plants (PU 3168) with n = 11

by a distance of nearly 200 m, proved diploid with n = 20. The bivalents could

be grouped into 10 pairs with 3 pairs of relatively larger size (Figs. 22-23). Some

of the bivalents tended to lie in pairs. Occasionally 2-3 bivalents failed to show

normal orientation at the equatorial plate (Fig. 22). In about 37% of the sporo-

cytes, 1-2 laggards were observed at telophase-II (Fig. 24), and in about 4% of

the sporocytes triads were formed with one large (apparently diploid) and two

small (apparently haploid) nuclei.

Source : MNHN, Paris

РІ. Ш. — 13-15 : Brachymenium acuminatum —n = 10. 13 :Metaphase-l; note the large

heteromorphic H-bivalent, and also the precocious disjunction of the small h-bivalent.

14 :Metaphase-I; note the large heteromorphic H-bivalent. 15 : Anaphase-I; note the

difference in the size of X and Ұ components (arrow marked) of the heteromorphic

bivalent. 16 :В. acuminatum — п = 11. Metaphase-l. 17-18 : B. exile — n =10. 17

Prometaphase; note the largest bivalent in association with the nucleolus. 18 : Meta-

phase-I; note the large, slightly heteromorphic H-bivalent (x 2720).

Source - MNHN, Paris

АБ

РІ. ТУ, — 19-21 :B. exile —n —11. 19-20 : Metaphase-l; note the heteromorphic bivalent

in 19. 21:А triad with one large (apparently diploid) and two small (apparently

haploid) nuclei. 22-24 +B, exile —n = 20. 22-23 : Metaphase-l, showing 20 bivalents

with 3 bivalents not oriented on the metaphase plate in 22. 24 : Telophase-II, showing

laggards. 25 :В. leptophyllum —n = 20. Metaphase-l; note 2 h-bivalents (2720).

Source : MNHN. Paris

84 5.5. KUMAR & S.K, VERMA

A morphological comparison of the haploid (PU 3164, PU 3165, PU 3166

Fig. 34) and the diploid (PU 3169 : Fig. 35) populations of B. exile revealec

that the diploid plants are taller (5.5-6.5 mm versus 1.7-2.0 mm) and posses:

larger basal areolations (34-51 x 17 um versus 17-36 х 10-17 um). Likewise

plants of the Didihat population (PU 3168 : Fig. 36) with n = 11 are alsc

taller (3.5-5.0 mm versus 1.7-2.0 mm) and have longer setae (1.6-2.5 cm versu:

1,0-1.8 cm) than those of the plants (Fig. 34) with n = 10. However, plant:

of the Almora population (PU 3167 : Fig. 37) which also show n = 11 are noi

differentiable from plants (PU 3164, PU 3165, PU 3166) with n = 10. These

quantitative differences in size in different populations seem to be linked wit

available humidity and shade conditions.

The close similarity between the plants of haploid and diploid constitutior

in essential morphological characteristics coupled with the cytological trait:

(20 bivalents falling into 10 pairs, tendency of the bivalents to lie in pairs

displayed by plants with n = 20 hints at the autoploid nature of the diploic

materials. The formation of triads in the polyploid population is indicative о

the receptivity of the genome to further polyploidy.

YANO (1957) counted 11 chromosomes in the somatic complement of the

Japanese materials. GANGULEE & CHATTERJEE (1962) observed 11 bivalent:

in a East Himalayan population of this species.

Brachymenium leptophyllum (C. Muell.) Jaeg. : n = 20 (Fig. 25) — Unlike

the preceding species of Brachymenium, which are either terrestrial or litho

phytes in our area, this species is corticolous growing on the trunks of broad

leaved trees. The material studied (PU 3170) was collected from Bhimtal (1390m

in a relatively open forest. This is the first cytological study of the species. The

twenty bivalents observed at metaphase-I (Fig. 25) are grouped into ten pair:

with three pairs of relatively large size. As judged from the morphology of the

bivalents, one of the larger pairs appears to be acrocentric and another meta

centric. Two members of the set, recognized as h-bivalents, showed faint staina

bility and displayed a tendency to anticipate disjunction. A careful survey of the

area of colonisation of this seemingly autopolyploid taxon may result in the

discovery of plants of haploid constitution.

Brachymenium nepalense Hook. :n = 22, 23 (22 + m) (Figs. 26-28) — Of the

three studied gatherings of this common epiphytic moss, two (PU 3171 - Dha

ramsala, Bhagsunath, 1900 m; PU 3172 - Dharamsala, on way to Triund, 1900 т

showed 22 bivalents which could be grouped into eleven pairs with one of the

four large-sized pairs faintly stained and recognizable as the H-pair (Fig. 26)

The smallest pair of the set, also faintly stained, corresponded to the h-bivalent

The duplicated genome is indicative of the autoploid nature of this species. А:

judged from the morphology of the bivalents, at least 6 chromosomes appeai

to be acrocentric. In the third population (PU 3173 - Bhimtal, 1390 m) a dot

like, faintly stained m-bivalent showing precocious disjunction (Figs. 27-28)

augmented the chromosome count, As judged from the morphology of the

bivalents, at least 8 chromosomes (3 large and 5 small) appear to be metacentri

(Fig. 28).

Source - MNHN, Paris

CYTOLOGY OF SOME WEST HIMALAYAN MOSSES 85

i % “.. % ,

едь

le а “ 27

i "

=? * а

£N T є +

У * + »

P dd

+ 26 qe

в.

22%. І

.* <-

С +,

Á --ө..

29 v 31

Pl. V. — 26 : B. nepalense — п = 22. Metaphase-I; note 2 large H-bivalents and 2 h-bivalents.

27-28 : B. nepalense — 23 (22 +m). Metaphase-I; note the precocious disjunction of

the m-bivalent. 29 :B. sikkimense — n = 10. Metaphase-l; note the large heteromorphic

bivalent. 30 :Anomobryum auratum —n = 10 + асс. Metaphase-l; note the small

dot-like peripherally located chromosome. Also note the precocious disjunction of the

large heteromorphic bivalent (arrow marked). 31 : Anomobryum sp. — n = 10. Meta-

phase-I; note the large heteromorphic H-bivalent and one small h-bivalent (x 2720).

Source : MNHN, Paris

86 S.S. KUMAR & S.K. VERMA

GANGULEE & CHATTERJEE (1962) reported n = 11, 12 (11 + m) in two

East Himalayan populations of this species.

Brachymenium sikkimense Ren. & Card. : n = 10 (Fig. 29) — Cytological

examination of two populations (PU 3174 - Ranikhet, on soil, 1850 m; PU 3175

- Hemkunt, on way from Gobind Ghat to Gobind Dham, on soil, 2850 m) of this

previously uninvestigated species gave a consistant count of n — 10 (Fig. 29).

One of the three relatively large bivalents of the complement was faintly stained

and anticipated disjunction of the set. The resulting half-bivalents were dissimi-

lar. In about 40% of the studied sporocytes, 1-2 laggards were observed at ana-

phase-I and telophase-II. Occasionally erratic meiotic behaviour resulted in the

formation of triads.

The chromosome numbers, n = 10, 11, 12, 20, 22, 23, 30 (FRITSCH 1972,

RAMSAY 1977), known in 8 out of the 170 species included in Brachymenium

suggest that polyploidy may be widely spread in this genus of largely tropical

distribution. The basic number of the genus appears to be x = 10.

Anomobryum auratum (Mitt.) Jaeg. : п = 10 + acc. (Fig. 30) - The material

studied (PU 3176) was gathered from Didihat (Pam Tori, on soil, 1350 m). The

largest bivalent of the set was slightly heteromorphic, faintly stained, and anti-

cipated disjunction. A dot-like faintly stained accessory chromosome, in addi-

tion to the normal complement, was seen in the peripheral position in the sporo-

cytes (Fig. 30). Occasionally 1-2 laggards were also observed at anaphase-I and

anaphase-II. Earlier, GANGULEE & CHATTERJEE (1962) counted 10 biva-

lents in a Darjeeling population of this species, They, however, did not observe

the distinctly large, heteromorphic bivalent in their materials. KUMAR (1973)

reported a long chromosome in the somatic complement of a West Himalayan

population which appears to correspond with the large bivalent seen in the

presently examined material.

Anomobryum sp. : п = 10 (Fig. 31) — The investigated material (PU 3177)

was collected from Nainital (Laria Kanta, on wet rocks, 2600 m). As in the

previous species, the chromosome complement included a distinctly large,

faintly stained, heteromorphic H-bivalent which disjoined precociously. One of

the smaller bivalents, recognisable as the h-bivalent, was weakly stained at me-

taphase-l.

Three other investigated species of Anomobryum (A. filiforme var. concinna-

tum, À. gemmigerum, A. nitidum) also show n — 10 (FRITSCH 1972) which

may be regarded as the basic number for this genus.

The chromosome numbers reported for 90 species included in 11 genera of

Bryaceae are :n - 10, 11, 12, 14, 15, 20, 21, 22, 23, 24, 26, 27, 30, 33, 40 and

50 (FRITSCH 1972, BRYAN 1973, DANYLKIV & VISOTSKAYA 1975,

INOUE 1976, KUMAR 1973, KUMAR & GARG 1974, KUMAR & VERMA

1980, NEWTON 1972, RAMSAY 1974, 1977, SNIDER 1970, TSUTSUMI et

al. 1973, VISOTSKAYA 1972, 1975, WIGH 1972, WIGH & STRANDHEDE

Source : MNHN, Paris

CYTOLOGY OF SOME WEST HIMALAY AN MOSSES 87

Е os

82 ug ‘ 35

jx 5

ІШ

РІ. VI. — 32-33 : Pohlia longicollis. 32 :Plants with n = 11. 33 :Plants with n = 22.

34-37 : Brachymenium exile. 34: Plants with n = 10. 35 :Plants with n —20. 36:

Plants with n —11. 37 :Plants with n —11 (x2).

Source : MNHN, Paris

88 S.S. KUMAR & S.K. VERMA

504

40-

NUMBER ОҒ ТАХА

5 10 15 22 25 30 35 40 45 50

СНВОМО5ОМЕ NUMBERS 38

Fig. 38. — Histogram showing the distribution of chromosome numbers іп Bryaceae.

1971). Of these numbers, n — 10 is found in 44 taxa and n — 11 in 35 taxa

(Fig. 38). The original basic number of the family appears to be x — 5 which

has either been lost or is still awaiting discovery. Thirty-seven species exist

as euploids and 23 species are found as aneuploids. The only other family of

acrocarpous mosses which shows a striking parallelism with Bryaceae in posses-

sing the afore mentioned cytological traits is Pottiaceae (FRITSCH 1972,

VERMA & KUMAR 1980). Incidently the latter family also appears to be hight-

ly evolved in the haplolepidous pool. The existing morphological plasticity

obscuring generic, subgeneric and specific boundaries, coupled with cytological

diversity, indicates that Bryaceae are in an active stage of evolution and spe-

ciation.

ACKNOWLEDGEMENT. — The authors are indebted to Prof. Н. Ochi, Department of

Botany, Tottori University, Japan, for his kind help in identifying the specimens.

LITERATURE CITED

BRYAN VS. 1973 — Chromosome studies on mosses from Austria, Czechoslovakia and

other parts of Central Europe. Österr. Bot. 2. 121 : 187-226.

DENYLKIV LS. & VISOTSKAYA E.I., 1975 — Chromosome numbers of leafy mosses of

the Lithunian SSR. Ukrajins'k Bot. Zurn. 32, 2 : 246-250.

Source : MNHN. Paris

CYTOLOGY OF SOME WEST HIMALAYAN MOSSES 89

FRITSCH R., 1972 — Chromosomen der Bryophyten. Wiss. Z. Friedrich-Schiller-Univ.

Jena, Math.-Naturwiss. Reihe 21, 5/6 : 839-944.

GANGULEE H.C. & CHATTERJEE N.K. 1962 — Cytological studies in the mosses of

Eastern India. III. Caryologia 15 : 367-400.

INOUE S., 1976 — Chromosome studies on five species of Antarctic mosses. Kumamoto

J. Sci., Biol, 13,1 : 1-5.

KHANNA K.R., 1967 — А cytological investigation of the mosses of the Rocky Mountains.

Univ. Colorado Stud., Ser. Biol. 26 :1-39.

KUMAR S.S., 1973 — Cytological studies on some West Himalayan mosses. Bryologist 76 :

201-206.

KUMAR S.S. & GARG R.G., 1974 — Cytological studies on some West Himalayan mosses.

Misc. Bryol. Lichenol. 6,9 :153-156.

KUMAR 5.5. & VERMA S.K., 1980 — Cytological observations on some West Himalayan

mosses. Misc. Bryol. Lichenol. (in press).

NEWTON M.E., 1972 — Chromosome studies in some South Georgian bryophytes. Brit.

Antarct, Surv, Bull. 30 : 41-49.

PANDE S.K. & CHOPRA N., 1958 — Cytological studies in Indian mosses. III, Proc. Natl.

Inst. Sci. India Pt. B, Biol. Sci. 24 : 94-99.

RAMSAY H.P., 1974 — Cytological studies of Australian mosses. Austral. J. Bot. 22 :293-

348.

RAMSAY H.P., 1977 — Chromosome numbers of some mosses from Western Australia.

J. Bryol. 9 : 343-347.

SMITH A.J.E. & NEWTON M.E., 1967 — Chromosome studies on some British and Irish

mosses. II. Trans. Brit. Bryol. Soc. 5, 3 :463-522.

SNIDER J.A., 1970 — Chromosome studies of some mosses of the Douglas Lake Region. I.

Michigan Bot. 9 :67-71.

TSUTSUMI S., TAGUCHI M. & INOUE S., 1973 — Karyological studies on Swedish mosses.

Misc. Bryol, Lichenol. 6, 5 : 82-84.

VERMA S.K. & KUMAR S.S., 1980 — Cytological observations in West Himalayan mosses.

J. Hattori Bot. Lab. 47 : 245-268.

VISOTSKAYA EJ., 1972 — New data on chromosome numbers for mosses from the Ukrai-

ne. Ukrajins’k Bot. Zurn. 29,1 : 84-87.

VISOTSKAYA EJ., 1975 — New data on chromosome numbers of Bryopsida in the Ukrai-

ne. Ukrajins'k Bot. Zurn. 32, 4 : 498-503.

WIGH K. & STRANDHEDE S.O., 1971 — Chromosome numbers in some Swedish and

Danish mosses. Bot. Notiser 124 : 213-227.

WIGH K., 1972 — Cytotaxonomical and modification studies in some Scandinavian mosses,

Lindbergia 1 : 130-152.

YANO K., 1957 — Cytological studies on Japanese mosses. 1. Fissidentales, Dicranales,

Grimmiales, Eubryales. Мет. Fac. Educ. Niigata Univ. 6 : 1-31.

Source : MNHN. Paris

Nena дылығы «іе әженің

DORE арым

REPRODUCTIVE PHENOLOGY ОЕ 5ОМЕ TROPICAL

AFRICAN MOSSES

E.A. ODU*

SUMMARY. — Four mosses, Racopilum africanum, Fissidens glauculus, Thuidium gratum

and Stereophyllum sp. were selected for field and laboratory studies on gametangia and

sporophyte development over а period of 18-21 months in south-western Nigeria. Maturity

indices of reproductive organs show that gametangia develop at the onset of the rainy

season (March/April) while sporophytes develop later and mature towards and at the end

of the same rainy season (October/November and December). Spores from the mature

capsules of all species are dispersed during and after the dry season (November-April).

The reproductive process from gametangia initiation to spore dispersal is completed in every

12-month cycle in both the perennial shoots of Racopilum, Thuidium and Stereophyllum

and annual shoots of Fissidens.

INTRODUCTION

Reproductive phenology for a given species of moss is the sequence of deve-

lopmental stages from the start of gametangia production until the fall of

dehisced capsules. The ultimate goal of phenological studies is to explain factors

which permit the development of a given stage in a species or in another taxon

(FORMAN 1965). Thus formation of sex organs in bryophytes may be influen-

ced by photoperiod (BENSON-EVANS 1961, 1964, ZEHR 1979) or tempera-

ture (BENSON-EVANS & HUGHES 1955, MONROE 1965, ANTHONY 1962)

or even an interaction between climatic factors and endogenous rhythms in

some species (NEWTON 1972, LONGTON 1979).

There is a great deal of information on patterns of gametangia and sporo-

phyte development in temperate and polar bryophytes e.g BENSON-EVANS

and HUGHES (1955) on Lunularia cruciata (L.) Dum., GREENE (1960) on six

British mosses, HUGHES (1962) on Polytrichum aloides Hedw. and P. piliferum

Hedw., LONGTON and GREENE (1968) on Pleurozium schreberi (Brid.)

Mitt., JOHNSEN (1969) on Orthotrichum anomalum Hedw. and HANCOCK

and BRASSARD (1974) on Buxbaumia aphylla Hedw.

* Department of Botany, University of Ife, Ile-Ife, Nigeria.

Cryptog., Bryol. Lichénol., 1981, 2, 1 : 91-99.

Source : MNHN, Paris

92 Е.А. ODU

There is little information available on the reproductive phenology of tro-

pical African bryophytes. Equatorial Africa provide nearly non-seasonal condi-

tions, Nigeria having only wet and dry seasons. Different parts of the country

experience different durations of each season. In Western Nigeria vegetative

growth of epiphyllous liverworts during the dry season was found to be much

less than that in the wet (OLARINMOYE 1974) indicating the regulatory

influence of moisture potential on growth.The present phenological study of

selected mosses was undertaken to discern the timing of sexual developmental

stages in relation to environmental factors in tropical African mosses.

MATERIALS AND METHODS

Moss shoots were sampled from natural populations of Racopilum africanum

Mitt., Thuidium gratum (P. Beauv.) Jaeg., Fissidens glauculus C. Müll. and Stereo-

phyllum sp. at different locations in Ile-Ife (0723074, 04^40'E). Collections

were made continuously at one month intervals for a period of 18-21 months

(April 1978-December 1979) and the shoots examined fresh in the laboratory

for gametangia and sporophyte development. Additional information was

recorded for field collections of R. africanum and T. gratum between January

and March 1979 at Ibadan (079257М, 03°53'Е) and Owena (07^12'N, 05°E),

T. gratum in February at Olokemeji (0722574, 03^ 32'E).

The reproductive stages recognized in the samples were those of GREENE

(1960) slightly modified by LONGTON and GREENE (1969) and CLARKE

and GREENE (1970). Index ratings were assigned to the developmental stages

(Tab. 1).

Maturation stage Maturity index

GAMETANGIA

Juvenile (J) 1

Immature (1) 2

Mature (M) 3

Dehisced (D) 4

SPOROPHYTE

Swollen Venter (SV)

Calyptra in perichaetium, early (ECP)

Calyptra in perichaetium, late (LCP)

Calyptra intact, early (ECI)

Calyptra intact, late (LCI)

Operculum intact, early (EOI)

Operculum intact, late (LOI)

Operculum fallen (OF)

Empty and fresh (EF)

Tab. 1. Stages in maturation of gametangia and sporophytes in mosses (Source : CLARKE

and GREENE 1970).

Source : MNHN, Paris

REPRODUCTIVE PHENOLOGY OF MOSSES 93

Gametangia and sporophyte stages for each species were scored monthly

from 5 current cycle perichaetia/perigonia from each of 10 shoots and 50

sporophytes respectively. Mean index values or maturity indices (LONGTON

and GREENE 1967) were then calculated from the formula :

E (Index rating x no. of gametangia/sporophytes)

Total no. of gametangia/sporophytes

Climatic data showing monthly total rainfall, number of rain days, relative

humidity and temperature of the area around Ile-Ife during the period of

study were obtained from the weather station of the Faculty of Agriculture,

University of Ife.

RESULTS

Analysis of between 900 and 1050 perichaetia/perigonia in each of the spe-

cies showed that there is a clearly defined seasonal cycle of gametangial develop-

ment just as has been reported for several temperate mosses. Juvenile gametangia

were usually first sited at the start of the rainy season between March/April and

May of each year; majority of the gametangia dehisce in May (Fig. 2a) or

June/July (Figs. 1a, 3a, 4a). It is apparent from Figs. 1a and 3a that antheridia

develop earlier than archegonia in Racopilum and Thuidium. This phenomenon

may also be possible in Fissidens and Stereophyllum where both sex organs were

reported to develop in the same month (Figs. 2a and 4a) but was not detected

probably due to the spacing (monthly) of field sampling. The differential deve-

lopment of male and female sex organs in these bryophytes may possibly be

a device for preventing infraspecific self fertilization.

The time of fertilization is expected to be during the mature stage of the

gametangia which is difficult to observe from field collections in large numbers.

The low frequency of mature gametangia may be due to the short duration

of the mature stage as has been suggested for some other mosses (GREENE

1960, LONGTON and GREENE 1969, ZEHR 1979). Thus the shortest period

of fertilization occurs in May in Fissidens (Fig. 2a.). Fertilization occurs in May

in Racopilum (Fig. 1а) but is earlier (March/April) in Thuidium and Stereophyl-

lum(Figs. За and 4a). The apparently long duration between juvenile and dehis-

ced gametangia in Thuidium is probably due to its shaded, highly humid habitat

(Fig. 6b). This may favour increased development of sex organs and fertilization.

Sporophytes in the swollen venter (SV) stage were first recorded for Stereo-

phyllum immediately after fertilization in March-May 1979 (Fig. 4b). Develop-

ment progresses rapidly in the same rainy season, the capsules maturing (LOI-

OF) towards the end of the rainy season in October (viz. Fig. 5).

After fertilization in April/May in Fissidens and Thuidium, sporophytes

developed rapidly to ECI stage within the same month (Figs. 2b and 3b).

However, sporophytes in all the mosses matured in October/November.

Spore dispersal (EF) occurred largely in Fissidens, Thuidium and Stereophyl-

lum during the dry season, November-February/March (viz. Fig. 5). In Raco-

Source : ММНМ, Paris

94 E.A. ODU

044 id

O ANTHERIDIA

71 5 е ARCHEGONIA

1 €—€———

Ü (ы)

EF 9

OF 8:

LOI 7

EOI 64 Т

0615]

ECI 4

LCP3 4

ЕСЕ2ІЗ

5У1 1

La --------

AM JJ TAS ON DTP MA MI TASO NO

1978 1979

Fig. 1. — Maturity indices for Racopilum africanum; development of gametangia (above)

and sporophyte (below). Vertical bars indicate the range of stages present.

(a)

i О ANTHERIDIA

ES анта DET

(в)

r mm

Fig. 2. — Maturity indices for Fissidens glauculus; development of gametangia (above) and

sporophyte (below). Vertical bars indicate the range of stages present.

Source : MNHN, Paris

REPRODUCTIVE PHENOLOGY OF MOSSES 95

(а)

M3 О ANTHERIDIA

€ ARCHEGONIA

24 —— TĒ т ES LE

(b)

EES |а

OF8

LOI 7

EOI 6

LCI 5 f

ECI 4 Е

LCP 3:

ECP2

5У1

——— Е , E

АМІТА5ОМОТЕМАМТ7А5О0М0

1978 1979

Fig. 3. — Maturity indices for Thuidium gratum; development of gametangia (above) and

sporophyte (below). Vertical bars indicate the range of stages present.

D4 TRIS al

1 ==

M3 О ANTHERIDIA

12 ө ARCHEGONIA

ie ee

(b)

EF94

OF 84 >

101 74

EOI 6

LCI 5

ECI 4

LCP3

ECP2]

svi]

желілі аі еі а, aped eg e

AMTTASONSOIFMAMJIJASO ND

1978 1979

Fig. 4. — Maturity indices for Stereophyllum sp.; development of gametangia (above) and

sporophyte (below). Vertical bars indicate the range of stages present.

Source : MNHN, Paris

E.A. ODU

30] =.

20] os.

в-а. aaa a A ss à ge

pu ee XUI |

recent)

604 Minimum

numigity (pe.

TT.

Relative

os 8

és

e oe

(b)

mers.

Total rainfall (mm)

а»

Number of rain

1978

1979

T ——

AMJIJIJASONDJFMAMJIIASONO

Fig. 5. — Climatic data recorded at Ile-Ife (0793074, 04940). (a) Maximum and minimum

temperatures;

of rain days.

(b) 1000 h and minimum relative humidity; (c) € total rainfall, А no.

pilum capsules matured fully between December and February with the spores

being dispersed as from March.

Source : MNHN, Paris

REPRODUCTIVE PHENOLOGY OF MOSSES 97

Fig. 6. — Microclimatic data recorded at ground level in one of the sampling sites at Ile-Ife

in 1979. (а) 31 Juanuary-7 February (b) 18-25 June (с) 3-10 September (4) 17-23

December. Broken lines, relative humidity; continuous line, temperature.

DISCUSSION

The phenological data recorded for these selected mosses indicate that there

seems to be clearly defined seasonal cycle of gametangia and sporophyte deve-

lopment in tropical African mosses. Similar patterns of cyclic behaviour have

been reported for many temperate and polar bryophytes. In these latter bryo-

phytes, however, cycles differ markedly between species (LONGTON 1979)

and in the duration of sexual cycle. For instance while it takes 18 months for

capsules of Pylaisia polyantha to mature (GREENE 1960) capsules of Ptilium

crista-castrensis require over 3 years (Van der WIJK 1960 cited by LONGTON

and GREENE 1969); spores from mature capsules of Pleurozium schreberi are

Source : MNHN, Paris

98 E.A. ODU

dispersed 15-20 months after gametangial initiation (LONGTON and GREENE

1969) but a similar cycle in the bipolar moss Polytrichum alpestre takes 24

months (LONGTON 1979). In the present studies the period between game-

tangial initiation and spore dispersal is a much shorter time, 7-10 months.

The regular alternate development of gametangia and sporophytes within

a 12-month cycle in these mosses seems to be controlled mainly by climatic

factors. This conclusion is based on this study and similar observations in seve-

ral other mosses, both field and herbarium specimens, from a wide area of

southern Nigeria over an 11-year period, 1968-1979; the observations show

a consistent occurrence of mature sporophytes only in shoots collected between

October/November and February. In the area of study, gametangia development,

fertilization and sporophyte development occur only during the extended cool

and wet months (March-October) of the biseasonal annual cycle (Fig. 5).

It is difficult to ascertain the role of photoperiod in the induction of sexual

organs without laboratory or field experimental data. The area covered in the

field study experiences equinoxes in March and September, slightly long days

during May-July and slightly shorter days in December-February. Indications are

that gametangial development during April-May is probably partially controlled

by the slightly long days but this needs to be verified experimentally. Additional

stimulus for gametophore production could also be provided by the diurnal

fluctuations in temperature (Figs. 5a and 6a-b) as has been suggested for male

Mnium undulatum (NEWTON 1972).

Sporophytes of all the species that matured at the end of the rainy season

disperse their spores during and at the end of the dry season. The intervening

dry period is necessary for shedding of the opercula (JOHNSEN 1969), effec-

tive maturation and dispersal of the spores ready for germination at the onset

of the next rainy season. This is of survival value for Fissidens glauculus, an

annual that has been suggested to become established in the field only by

spore germination (ODU 1979).

The salient point emerging from this study indicates that in a 12-month

cycle, maturity of sex organs, fertilization and sporophyte development are

all coordinated to take place during the rainy season while capsule dehiscence

and spore dispersal occur in the dry season, Gametangia and sporophyte deve-

lopment take approximately 1-4 and 6-8 months respectively with no resting

phase of any kind.

An endogenous rhythm in coordination with the season seems to influence

the reproductive process in Racopilum where capsules matured during the dry

season (December-February) even though fertilization occurred early in the

rainy season (May/June). Further studies are therefore necessary on the repro-

ductive phenology via-a-viz shoot growth of mosses and the tropical climatic

factors.

ACKNOWLEDGEMENTS. —I wish to thank the Faculty of Agriculture, University of Ife,

for supplying the weather data from the University of Ife, Ile-Ife. I would also like to thank

Source : MNHN, Paris

REPRODUCTIVE PHENOLOGY OF MOSSES 99

Drs. А.О. Lawanson and O. Olorode for their useful comments. A research grant was provi-

ded by the University of Ife, Nigeria.

REFERENCES

ANTHONY R.E., 1962 — Greenhouse culture of Marchantia polymorpha and the induction

of sexual reproduction structures. Turtox News 40 : 2-5.

BENSON-EVANS K., 1961 — Environmental factors and bryophytes. Nature, Lond. 191 :

255-260.

BENSON-EVANS K., 1964 — Physiology of the reproduction of bryophytes. Bryologist

67 :431-445.

BENSON-EVANS K. and HUGHES J.G., 1955 — The physiology of sexual reproduction in

Lunularia cruciata (L.) Dum. Trans. Brit. Bryol. Soc. 2 : 513-522.

CLARKE G.C.G. and GREENE S.W., 1970 — Reproductive performance of two species

of Pohlia at widely separated stations. Trans. Brit. Bryol. Soc. 6 : 114-128.

FORMAN R.T.T., 1965 — A system for studying moss phenology. Bryologist 68 : 289-300.

GREENE S.W., 1960 — The maturation cycle, or the stages of development of gametangia

and capsules in mosses. Trans. Brit. Bryol. Soc. 3 : 736-745.

HANCOCK J.A. and BRASSARD G.R., 1974 — Phenology, sporophyte production and life

history of Buxbaumia aphylla in Newfoundland, Canada. Bryologist 77 : 501-513;

HUGHES J.G., 1962 — The effects of day-length on the development of the sporophytes

of Polytrichum aloides Hedw., and P. piliferum Hedw. New Phytol. 61 : 266-273.

JOHNSEN AB., 1969 — Phenological and environmental observations on stands of Ortho-

trichum. Bryologist 72 : 397-403.

LONGTON R.E., 1979 — Studies on growth, reproduction and population ecology in rela-

tion to microclimate in the bipolar moss Polytrichum alpestre. Bryologist 82 . 325-367.

LONGTON К.Е. and GREENE S.W., 1967 — The growth and reproduction of Polytrichum

alpestre Hoppe on South Georgia. Phil. Trans, Roy, Soc. B252 : 295-322.

LONGTON R.E. and GREENE S.W., 1969 — The growth and reproductive cycle of Pleuro-

zium schreberi (Brid.) Mitt, Ann. Bot, 33 : 107-126.

MONROE J.H., 1965 - Some factors evoking formation of sex organs in Funaria, Bryolo-

gist 68 :337-339.

NEWTON М.Е., 1972 — An investigation of photoperiod and temperature in relation to

the life cycles of Mnium hornum Hedw. and М. undulatum Sw. (Musci) with reference

to their histology Bot. Jour. Linn. Soc. 67 : 189-209.

ODU E.A., 1979 — Spore germination in two tropical mosses : Fissidens sp. and Racopilum

sp. Ann. Bot. 14 : 147-152.

OLARINMOYE S.O., 1974 — Ecology of epiphyllous liverworts : growth in three natural

habitats in western Nigeria. J. Bryol. 8 : 275-289,

WIJK R. Van der, 1960 — De periodiciteit inde ontwikkeling der bladmossen. Buxbaumia

14 : 25-39,

ZEHR D.R., 1979 — Phenology of selected bryophytes in southern Illinois. Bryologist 82 :

29-36.

(раст

|MUSEUM

Source : MNHN, Paris

101

SPORE ORNAMENTATION IN LEIOMELA

(Musci : Bartramiaceae)

D. GRIFFIN, III *

SUMMARY. - The spores of 4 species of Leiomela were examined by light microscopy, and

1 species, L. bartramioides, was examined also by scanning electron microscopy. АП 4

species produce spores with a reticulate pattern of surface ornamentation. This type of

ornamentation has not been reported previously in the Bartramiaceae.

INTRODUCTION

The utility of spore ornamentation in moss systematics differs from one

taxonomic group to another (for general review see CLARKE 1979), and for

many families and genera it is not yet possible to state what that utility might

be. In Encalypta (JARAI-KOMLODI and ORBAN 1975, VITT and HAMIL-

TON 1974), Bruchia (McCLYMONT 1955) and some other genera there appear

to be significant differences in spore ornamentation at the species or species

complex level. On the other hand, SAITO and HIROHAMA (1974) discovered

that in several genera of Pottiaceae the patterns of spore ornamentation do not

always correlate with taxonomic concepts. They argued that on the basis of

similarities in spore ornamentation the genera Astomum and Hymenostomum

could not be maintained apart from Weissia. In the Bartramiaceae it is known

that several major discontinuities in spore ornamentation occur (BOROS and

JARALKOMLODI 1975, ERDTMAN 1965, MATTERI 1968, 1973, MIYOSHI

1973, HIROHAMA 1977), especially at the generic level; however, many gaps

remain in our knowledge of spore morphology in this family. Relatively little is

known of spore morphology in Leiomela, for instance, and with this in mind the

present study was undertaken.

METHODS AND MATERIALS

Spores of the following 4 species were used in this investigation : Leiomela

africana Thér. et Nav., L. aristifolia (Jaeg.) Wijk et Marg. L. bartramioides

(Hook.) Par. and L. javanica (Ren. et Card.) Broth.

* Department of Botany and Florida State Museum, University of Florida, Gainesville, Flo-

rida, U.S.A. 32611.

Cryptog., Bryol. Lichénol., 1981,2, 1 : 101-106.

Source : MNHN, Paris

102 D. GRIFFIN

Specimens examined. — L. africana, TANZANIA, Ukaguru Mts N of Kilosa

town, on Cyathea stems, T. Pócs, B.F. Harris and Mrs. Mwansabe 6592/K, 25

May 1972. L. aristifolia, ECUADOR, summit of Pichincha, in rock crevices,

Jameson sn, Sept. 1847 (type). L. bartramioides, VENEZUELA, Mucuy

above the town of Tabay, on trunk of tree fern, Griffin, López F. and L. Ruiz-

Terán 1838, 7 Aug. 1972. L. javanica, JAVA, am Gedeh bei Kandang-Badak

an Baumrinde, Fleischer 17286, 18 Juli 1898.

Light microscopy. - Spores from mature capsules were heated gently in

water and then mounted on glass slides in Hoyer's solution. Ten spores of each

species were measured and the measurements averaged. A few spores of each

species were heated and measured in water mounts to check on possible size

changes in water versus Hoyer's-mounted material. No perceptible changes were

observed, and the Hoyer’s-mounted spores were used for measuring and for

photomicrography.

Scanning electron microscopy. - Spores of L. bartramioides were studied by

SEM. Spores for this study were mounted dry on squares of 2-sided adhesive

tape affixed to aluminium stubs. The spores were gold coated using an Eiko

sputter-coater and later examined at 20 KV with a Hitachi S 450 scanning

electron microscope.

RESULTS

Spores of the 4 species of Leiomela examined in this study were of similar

shape and surface ornamentation but differed somewhat in size. The mature

spores are subspherical to spherical, apolar, with little or no discernable diffe-

rence in surface ornamentation between distal and proximal faces. The averaged

sizes are as follows (length/width — 10 spores measured for each species) : L.

africana, 31 x 29 y; L. aristifolia, 31 и x 30 s L. bartramioides, 34 u x 320:

L, javanica, 38 u x 36 u.

The basic type of ornamentation is found in all 4 species examined (Figs. 1,

I and Ш). The sculpturing is a reticulum with the muri showing perfect or

imperfect fusions. The lacunae are short-oblong to rectangular and vary from

0.5 р to 4 u in long dimension,

DISCUSSION

Because of the small sample size no significance is attributed to the differen- N,

ces in spore sizes among the 4 species. The reticulate ornamentation is of interest |

in that this type has not been reported previously for ће Bartramiaceae. Several

other genera (eg, Bartramia, Conostomum, Breutelia and Plagiopus) show

some form of verrucate ornamentation while in Philonotis a number of types

are known, including baculate, clavate and pilulate.

Leiomela is a small and morphologically somewhat isolated genus of the

Bartramiaceae. It is now seen to have an unusual, if not unique, type of spore

Source : MNHN. Paris

Fig. I. — Spores of Leiomela bartramioides (Hook.) Par., Venezuela, Griffin et al., 7.VIII.

1972. — 1 : Whole spore, x 2450. 2 : Detail of surface ornamentation, x 5000.

Source : MNHN, Paris

Spores of Leiomela. — 3 : L, africana Thér. et Nav., Tanzania, Pécs et al., 25.V.

1972, x 2000. 4: L. aristifolia (Jaeg.) Wijk et Marg., Ecuador, Jameson, 1847, x 2000.

ource - MNHN. Paris

Fig. Ш. — Spores of Leiomela. — 5 : L. bartramioides (Hook.) Par., Venezuela, Griffin et

al., 7.У11,1972, x 2000. 6 : L. javanica (Ren. et Card.) Broth., Fleischer, 1898, x 2000.

Source : MNHN, Paris

106 D. GRIFFIN

ornamentation. This supports the generic concept for Leiomela; however, it

remains to be seen (GRIFFIN 1980) whether the patterns of spore ornamen-

tation can be used to help refine and improve the concepts on which the other

genera of the family are based.

LITERATURE CITED.

BOROS A. and JARAI-KOMLODI M., 1975 — An atlas of recent European moss spores.

Akadémiai Kiadó. Budapest.

CLARKE G.C.S., 1979 — Spore morphology and bryophyte systematics, In «Bryophyte

Systematics», ed. G.C.S. CLARKE and J.G. DUCKETT, pp. 231-250. Academic Press.

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ERDTMAN G., 1965 — Pollen and spore morphology / plant taxonomy, part II, Almqvist

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GRIFFIN D. III, 1980 — Spore morphology and generic concepts in the Bartramiaceae,

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HIROHAMA T., 1977 — Spore morphology of bryophytes observed by scanning electron

microscopy, II. Bartramiaceae. Bull. Nat. Sci. Mus. Tokyo, Ser. B, Bot. 3 : 37-44.

JARALKOMLODI M. and ORBAN S. 1975 — Spore morphological studies on recent

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MATTERI C., 1968 — Las especies de Philonotis (Bartramiaceae) del sur de Argentina.

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MATTERI C., 1973 — El género Breutelia (Bartramiaceae, Musci) en la regíon Andino-

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MIYOSHI N., 1973 — Scanning electron microscopic observation of spores in the Musci.

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Source : MNHN, Paris

107

BIBLIOGRAPHIE BRYOLOGIQUE

D.LAMY*

SYSTÉMATIQUE, NOMENCLATURE

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Cryptog., Bryol. Lichénol., 1981,2, 1 : 107-125.

Source : MNHN, Paris

108 BIBLIOGRAPHIE BRYOLOGIQUE

Grolle) et sect. Metaclasmatocolea (C. navistipula et la var. parceramosa Engel);

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Source : MNHN, Paris

BIBLIOGRAPHIE BRYOLOGIQUE 109

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Source : MNHN, Paris

110 BIBLIOGRAPHIE BRYOLOGIQUE

MORPHOLOGIE, ANATOMIE

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Source : MNHN, Paris

BIBLIOGRAPHIE BRYOLOGIQUE 11

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Source : MNHN, Paris

112 BIBLIOGRAPHIE BRYOLOGIQUE

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Volemitol, sucrose et amidon sont les principaux produits de la photosyn-

thése chez Plagiochila asplenioides var. major; mais les carbohydrates les plus

abondants quantitativement (fructanes) n'incorporent pas rapidement "^C et

CO. Le glucose ехорёпе, le fructose et les sucroses sont absorbés et méta-

bolisés dans le noir comparablement au CO; à la lumière, sauf là où il y a une

incorporation importante en fructanes. Le mannitol exogène est absorbé mais

peu métabolisé.

VANDEKERKHOVE 0, — Stabile Wachstumsanomalien bei Riccia fluitans

nach lingerer Einwirkung von Phenylborsäure auf die Vegetationspunkte.

Naturwissenschaften 1980, 67, 2 : 96-97, 1 fig. (Inst. Allg. Bot., Univ.

D-6500 Mainz).

VIELL B. und SCHAAR I. — Änderungen im Phosphat-Stoffwechsel regene-

rierender Zellen des Lebermooses Riella helicophylla (Bory et Mont.) Mont.

Biochem. Physiol. Pfl. 1979, 174 : 780-793, 3 tabl., 6 fig. (Inst. Entwick-

lungsphysiol. Univ. Köln).

Le phosphate inorganique augmente dans les cellules régénérées, diminue

Source : MNHN, Paris

BIBLIOGRAPHIE BRYOLOGIQUE 113

dans celles non régénérées. Ces dernières apparaissent comme des donateurs de

phosphate, le transfert du 22Р se faisant très rapidement entre les deux types

de cellules.

CYTOLOGIE

INOUE S. — A preliminary report on moss chromosome numbers. Misc. Bryol.

Lichenol. 1979, 8, 6 : 109-113 (Dept. Biol, Fac. Sci., Kumamoto Univ.,

Kurokami 2-chome, Kumamoto-Shi, 860 Japan).

Nombre chromosomique pour 96 esp.; réalisation d'un premier comptage

pour 48 d'entre elles.

MEHRA Р.М. and KUMAR D. — Chromosomes in Aneura indica St. Misc. Bryol.

Lichenol. 1979, 8, 5 : 85-86, 4 fig. (Dept. Bot., Panjab Univ., Chandigarh,

India).

Présence de races haploides et diploides chez Aneura indica.

RÉPARTITION, ÉCOLOGIE, SOCIOLOGIE

ANDO, H.- Ecology of terrestrial plants in the Antarctic with particular

reference to Bryophytes. Mem. Natl. Inst. Polar Research 1979, spec. iss. 11 :

81-103 (Bot. Inst., Fac. Sci., Hiroshima Univ., Hiroshima 730, Japan).

BOIKO M.F. — Moss cover in phytocenoses of Desert Steppe in the blanck Sea

Reservation of the Ukrainian SSR Academy of Sciences. Ukrajins’k Bot.

Zum. 1980, 37, 2 : 59-61, 1 fig., 1 tabl. (Inst. Bot. M.G. Kholodnogo, AN

URSR, Lab. Bryol., Nadijishla).

Selon le gradient d'humidité du sol et de salinité, il y a soit Syntrichia ruralis

(Hedw.) Brid., soit Brachythecium albicans (Hedw.) B.S.G.

DIA M.G., RAIMONDO Е.М. e FERRARELLA A, — Contributi alla brioflora

sicula П. Naturalista Sicil. ser. 4, 1979, 3 : 111-118 (Ist. Orto Bot., Univ.

Palermo, 90123 Palermo, Italia).

Liste de 8 esp. d'hépatiques nouv. pour la Sicile, avec les loc. et données

chorologiques et écologiques.

DUDA J., VANA J. — Rozsireni jatrovek v Ceskoslovensku - XXVI (Die Verbrei-

tung der Lebermoose in der Tschechoslowakei - XXVI). Cas. Slez. Muz.,

Vedy Prirod. Ser. A 1979, 28 : 111-128, 5 cartes, en tchèque, rés. all. et

russe (Slezke Muz., C-746 46 Opava).

Loc. en Tchécoslovaquie, bibliogr. : 125 - Marsupella sprucei (Limpr.) Bern.

- 128 - M. brevissima (Dum.) Grolle - 141 - Gymnomitrion coralloides Nees -

143 - G. obtusum (Lindb.) Pears. - 241 - Porella cordaeana (Hüb.) Moore.

Source : MNHN, Paris

114 BIBLIOGRAPHIE BRYOLOGIQUE

EGUNYOMI A. — Bryophytes and the Nigerian vegetation types. Misc. Bryol.

Lichenol. 1979, 8, 4 : 67-68 (Dept. Bot., Univ. Ibadan, Ibadan, Nigeria).

HIRANO H. and SUZUKI T. — Lejeunea magohuki Mizut. occurs in Shizuoka-

ken, Central Japan. Proc. Bryol. Soc. Japan 1978, 2, 5 : 68, en japonais.

IRELAND R.R. and BELLOLIO-TRUCCO G. — Mosses new to Ontario and

Quebec. Canad. Field-Naturalist 1979, 93, 4 : 431-433.

Liste avec loc. de 5 mousses nouv. pour l'Ontario et 5 pour le Québec.

IWATSUKI Z. — New locality of Acroporium suzukii Sak. Proc. Bryol. Soc.

Japan 1978, 2, 5 : 69, en japonais.

IWATSUKI Z. —Pylaisiella laetoviridis (Okam.) Iwats. et Nog. in Miyazaki.

Proc. Bryol. Soc. Japan 1978, 2, 5 : 69, en japonais.

IWATSUKI Z. and TAKAKI N. — Eccremidium minutum (Mitt.) Stone et Scott

newly found in Japan. Misc. Bryol. Lichenol. 1979, 8, 5 : 98-100, 2 fig.,

en japonais,

IWATSUKI Z. and KIGUCHI Н. — Campylopodium euphorocladum (С. Müll.)

Besch., newly found іп Izu-Schichito Islands, Japan. Misc. Bryol. Lichenol.

1979, 8, 6 : 121-124, 4 fig., en japonais, rés. angl.

JAVORCIKOVA D. — Beitrag zur Bryoflora der Weissen Karpaten. Acta Fac.

Rerum Univ. Comenianae, Bot. 1979, 27 : 163-169 (Lehrst. Syst. Bot.,

Naturwiss. Fak., Kominsky Univ, 80100 Bratislava, Tchécoslovaquie).

20 hépatiques et 78 mousses rencontrées au cours de l'excursion.

KIGUCHI H. — Fissidens in Agano district, Saitama Prefecture, Central Japan.

Proc. Bryol. Soc. Japan 1978, 2, 5 : 62-64, en japonais (11 esp.).

KIGUCHI Н. and OKABE T. — Fissidens hyalinus Hook. et Wils. found in Shin-

juku-ku, Tokyo, Central Japan. Proc. Bryol. Soc. Japan 1978, 2, 5 : 69, en

japonais.

KITAGAWA N. — Lunularia cruciata and Monoselenium tenerum in Nara City.

Proc. Bryol. Soc. Japan 1978, 2, 5 : 69, en japonais.

KUMAR S.S. — Fissidens geppii Fleisch. — A new record of India. Misc. Bryol.

Lichenol. 1979, 8, 4 : 68-69, 1 fig. (Dept. Bot., Panjab Univ., Chandigarh-14,

India).

LECOINTE A. — Southbya nigrella (De Not.) Spruce, Cephaloziella baumgart-

neri Schiffn. et Tortella inflexa (Bruch) Broth., bryophytes nouvelles pour la

Basse-Normandie, aux Carrières d'Orival, près de Creully (Calvados). Bull.

Soc. Linn. Normandie 1979, 107 : 47-60, 5 cartes (Lab. Phytogéogr., UER

Sci. Terre et Aménagem. Région., Univ. Caen, F-14302 Caen Cedex).

Biotopes et groupements bryologiques de cette localité menacée de destruc-

Source : MNHN, Paris

BIBLIOGRAPHIE BRYOLOGIQUE 115

tion. 28% des esp. appartiennent au cortège méditerranéen 5.1. 3 esp. sont nouv.

pour la Basse-Normandie.

LECOINTE A. — Intérêts phytogéographiques de la bryoflore normande : 1 - Les

cortèges cosmopolite et méditerranéen s. 1. Bull. Soc. Linn. Normandie

1979, 107 :61-70, 1 carte (Idem).

A l'aide de l'étude des cortèges cosmopolite et méditerranéen s. 1. (respective-

ment 10 et 22% de l'ensemble), ГА, précise certaines de ses conceptions person-

nelles sur la définition et la désignation des aires biogéographiques pour les

bryophytes.

LEE T.D. and LA ROI G.H. — Bryophyte and understory vascular plant beta

diversity in relation to moisture and elevation gradients. Vegetatio 1979, 40,

1 : 29-38, 2 fig., 3 tabl. (Dept. Bot., Univ. Alberta, Edmonton, Alberta T6G

2E9, Canada).

La diversité beta (changement dans la composition en esp. selon un gradient

d'habitat) des bryophytes et des plantes vasculaires est semblable pour l'humi-

dité dans les 3 zones de végétation étudiées. Elle est plus importante chez les

plantes vasculaires selon le gradient de l'élévation dans les 4 régimes d'humidité.

Les bryophytes apparaissent plus tolérants aux facteurs liés à l'altitude, incluant

la température.

NAKAMURA T., SUGITA H. and INOUE H. — Flora and ecology of Bryo-

phytes in the National Park for Nature Study. Misc. Rep. Natl. Park Nat.

Stud. 1979, 9 : 61-73, 9 fig., 2 tabl. (Lab. Forest Bot., Fac. Agricult., Univ.

Tokyo).

Liste des 52 esp. récoltées. Écologie des bryophytes terricoles et épiphytes.

Évolution de la bryoflore depuis 1954.

PECIAR У. — Studia bryofloristica Slovaciae XI. Acta Fac. Rerum Nat. Univ.

Comenianae, Bot. 1979, 27 : 151-161 (Lehrst. Syst. Bot. Naturwiss. Fak.

Komensky Univ., 80100 Bratislava, Tchécoslovaquie).

Liste des esp. d'hépatiques et de mousses avec loc.

PROVOST M. et LECOINTE A. — Compte-rendu de l'excursion botanique faite

par la société linnéenne de Normandie en forêt de Grimbosq (Calvados) le

11 juin 1978. Bull. Soc. Linn. Normandie 1979, 107 :83-90 (Lab. Phyto-

géogr., UER Sci. Terre et Aménagem. Région., Univ. Caen, F-14032 Caen

Cedex).

Stations visitées avec esp. rencontrées (phanérogames, bryophytes, lichens).

REYES D.M. — Nuevos reportes para la flora hepaticologica de America latina, I.

Acta Bot. Acad. Sci. Hung. 1979, 25 (2-3) : 359-360 (Inst. Bot., Acad. Sci.

Cuba, La Havane, Cuba).

Liste de 26 esp. avec loc. en Amérique latine. 11 sont nouv. pour la région de

la Gran Piedra, Cuba. Aphanolejeunea ephemeroides Schust., Colura greig-

Source : MNHN, Paris

116 BIBLIOGRAPHIE BRYOLOGIQUE

smithii Jovet-Ast, C. tenuicomis (Evans) Steph., Drepanolejeunea lichenicola

(Spr.) Steph., Prionolejeunea helleri Evans et Radula pallens (Sw.) Dum. sont

nouv. pour Cuba.

SHIMOSE S. - Cololejeunea aoshimensis (Horik.) Mizut. found in Takashima

Island, Shimane Prefecture, Japan. Proc. Bryol. Soc. Japan 1978, 2, 5 :

61-62, 9 fig., en japonais.

SZMAJDA P. — Bryoflora Gor Stolowych i jej charakterystyka Geobotaniczna.

Prace Komis. Biol. 1979, 52 : 1-80, 14 fig., 4 tabl. (Dept. Geobot., Adam

Mickiewicz Univ., 61-713 Poznan, Pologne).

Bryoflore des Monts Stolowe et ses caractéristiques géobotaniques. Descr.

de la région (géol., climat, végétation). Liste systématique de 272 esp., 20 sous-

esp. et 9 fo. de mousses et sphaignes avec loc. Distrib. verticale. Analyse floris-

tique.

WALTHER J.M.N. — Groupements muscinaux dans les Vosges moyennes. Bull.

Soc. Hist. Nat. Colmar 1975-1977 (1979), 56 : 91-102, 2 fig., 2 phot., 15

tabl. (Inst. Bot., Lab. Ecol. Végét., Univ. Louis Pasteur, F-67000 Strasbourg).

152 relevés sur 3 types de substrat permettent de définir 9 groupements

saxicoles, 3 épixyles et saproliginicoles, 3 sur rochers et troncs d'arbres, 2 épi-

phytes corticoles. Structure et dynamique de ces groupements.

YAMADA К. and LAI М.Ј. – Five Bazzania species new to Taiwan. Misc.

Bryol. Lichenol. 1979, 8, 5 : 87 (Ujiyamada High School, Uraguchi, Ise-shi,

Mie-ken, Japan).

B. conophylla (Sde Lac.) Schiffn., B. imbricata (Mitt.) Hatt., B. ovistipula

(Steph.) Abeyw., B. shikkimensis (Steph.) Herz., et B. vittata (Gott.) Trev. nouv.

pour Taiwan.

POLLUTION

FERGUSON P. and LEE J.A. — Some effects of Bisulphite and Sulphate on

the Growth of Sphagnum species in the Field. Environm. Pollut. Ser. A 1980,

21, 1 : 59-71, 7 tabl., 3 fig. (Dept. Bot., Univ. Manchester, M 139 PL Great

Britain).

Effet principal : réduction de croissance.

ROBERTS B.A., THOMPSON L.K. and SIDHU S.S. — Terrestrial bryophytes

as indicators of fluoride emission from a phosphorus plant, Long Harbour,

Newfoundland, Canada. Canad. J. Bot. 1979, 57, 15 : 1583-1590, 5 fig., 2

tabl. (Newfoundland Forest Res. Centre, Canad. For. Serv., P. O. Box 6028,

Pleasantville, StJohn’s Newfld., Canada АІС 5X8).

L'étude porte principalement sur Polytrichum commune. Observation d'une

Source : MNHN, Paris

BIBLIOGRAPHIE BRYOLOGIQUE 117

chlorose bénigne des phyllides allant jusqu'au brunnissement à 95%. Concentra-

tion des fluorures dans les différents tissus; comparaison avec celle des fluorures

dans les aiguilles du sapin baumier, des fluorures présents dans l'humus et dans

Pair. Les Bryophytes apparaissent comme bons indicateurs de pollution.

STEFAN М.В. and RUDOLPH В.р. — Terrestrial Bryophytes as Indicators of

Air Quality in Southeastern Ohio and Adjacent West Virginia. Ohio J. Sci.

1979, 79, 5 : 204-212, 4 fig., 3 tabl. (Dept. Bot., Ohio State Univ., Columbus,

Ohio 43210 U.S.A.).

Cartographie utilisant Dicranum scoparium, Leucobryum albidum et Polytri-

chum ohioense comme indicateurs de pollution par SO; et chrome.

VARIA

DOBBELER P. Untersuchungen an moosparasitischen Pezizales aus der Ver-

wandschaft von Octospora. Nova Hedwigia 1979 (1980), 31, 4 : 817-864, 12

fig. (Inst. Syst. Bot., Menzingerstr. 67, D-8000 München 19).

Octospora et les genres alliés parasitent les rhizoides entraînant la formation

d'innovations et quelquefois les cauloides et les feuilles. Révision systématique

du genre Octospora. Noter que les Humariaceae, apparemment pyrénocarpes,

parasites des hépatiques feuillées, font partie de Octosporella gen. nov. (d’où

les comb. nov. : O. hemicrypta, O. jungermanniarum, О. perforata, O. suboper-

culata) et non du genre Pseudonectria. Relations parasite-plante hôte. Bryo-

philie.

ISOVIITA P.— S.O. Lindberg’s bryological reports in Finnish newspapers.

Critically annotated facsimile collection. Helsingin yliopiston kasvimuseon

monisteita (Pamphlet Bot. Mus., Univ. Helsinki) 1980, 8 : 1-88 (Bot. Mus.,

Univ. Helsinki, SF-00170 Helsinki 17).

Les minutes des réunions de la «Societas pro Fauna et Flora Fennica» ont

paru en suédois dans les journaux : Helsingfors Dagblad, Morgonbladet et Fin-

land; elles comportent la publication de nombreux nouv. noms de plantes.

50. Lindberg y a publié validement autour de 90 nouv. noms ou nouv. combi-

naisons, L'A. nous propose, en facsimilé, les notes bryologiques (1865-1887) de

Lindberg qui ont paru dans ces quotidiens, avec une liste des nouveautés nomen-

claturales, Aucune altération dans la nomenclature établie n'est proposée, sauf

pour Gymnomitrion obtusum Lindb., qu'il faut citer (Lindb.) Pears.; discussion

de cert. autres status. Cet important document est accompagné de notes histo-

riques, et met en valeur l'importance des journaux non-scientifiques dans la

nomenclature, mais aucune proposition formelle n'est présentée pour modifier

les régles actuelles. Le travail de recherche reste à faire pour d'autres auteurs en

d'autres régions.

LECOINTE A. — A propos de la cartographie en réseau : deux exemples, remar-

Source : MNHN, Paris

118 BIBLIOGRAPHIE BRYOLOGIQUE

ques et appel. Bull. Soc. Linn. Normandie 1979, 107 : 46 (Lab. Phytogéogr.

UER Sci. Terre et Aménagem. Régional, Univ. Caen, F-14032 Caen Cedex).

SEAWARD M.R.D. — Two letters of bryological interest from Richard Spruce to

David Moore. Naturalist (N. England) 1980, 105 : 29-33.

D. Moore (1807-1879), était superintendant puis conservateur du National

Bot. Garden., Glasnevin, Dublin.

SYNNOTT D.M. — A catalogue of collectors in the bryophyte herbarium, Natio-

mal Botanic Garden, Glasnevin (DBN). Glasra 1980, 4 : 17-30 (Natl. Bot.

Gard., Glasnevin, Dublin 9, Ireland).

Source : MNHN, Paris

119

BIBLIOGRAPHIE LICHÉNOLOGIQUE

D. LAMY

SYSTÉMATIQUE, NOMENCLATURE

CRESPO А. y SANCHO L.G. - Umbilicariaceae (Lichenes) de la Sierra de Gua-

darrama (España). Anales Inst. Bot. Cavanilles 1978 (1980), 35 : 79-101,

5 fig, 2 tabl. (Dept. Bot., Fac. Farmacia, Ciudad Univ., Madrid-3, España).

Morphol., systématique des Umbilicariaceae. Liste avec descr., hab., écol. de

21 taxons de la Sierra de Guadarrama. Diagn., descr., de Umbilicaria crustulosa

subsp. punctata subsp. nov. Descr. de la forme des spores et du comportement

écologique de U. depressa. Noter les stat. nov. : U. hirsuta subsp. papygiria

(Ach.) (= Gyrophora hirsuta var. pap.), Lasallia brigantium subsp. hispanica

(Frey) (= U. brig. var. hisp.). Clé aux taxons récoltés.

DIBBEN М.Ј. — Тһе chemosystematics of the lichen genus Pertusaria in North

America North of Mexico. Publ. Biol. Geol. (Milwaukee Public Mus.) 1980,

5 : 1-162, 136 fig., 29 tabl. (Dept. Bot., Milwaukee Public Mus., Milwaukee

Wisconsin 53233 USA).

Morphol., chimie, géogr., écol., phylogénie, histor. du genre Pertusaria. Corré-

lations entre les caractères morphol. et chimiques des 6800 spécimens examinés.

Mise en évidence de 40 substances lichéniques dont 6 sont nouv. La forme des

apothécies et la structure de la paroi sporale permettent de diviser le genre en

2 sous-genres : Pertusaria (esp. type :Pertusaria pertusa (L.) Tuck.) et Pionospo-

ra (Th. Fr.) c. n. (esp. type : P. bryontha (Ach.) Nyl.). Dans chaque sous-genre

les similitudes chimiques mettent en évidence des groupes d'esp. Clés aux 2 sous-

genres et aux 66 esp. étudiées. Pour chaque esp. : taxonomie, descr., ill., affini-

tés, données écol. et distr. 10 esp. et ssp. sont exclues de cette étude. Esp.

nouv. : P. floridana (Floride), P. hypothamnolica (Illinois), P. multipunctoides

(Caroline N), P. santamonicae (Californie), P. subambigens (Brit. Columbia),

P. suboculata Brodo et Dibb. (Brit. Columb.), P. sulcata (Maine), P. arizonica

(Texas), P. californica (Californie), P. consocians (Minnesota), P. ostiolata (Caro-

line №), P. paratuberculifera (New York), P. shenandoensis Hale et Dibb. ( Virgi-

nie), P. sinusmexicani (Louisiane), P. valliculata (Arkansas), Comb. nouv. : P.

тасоипй (Lamb) (= Melanaria m. ).

ELIX J.A. and STEVENS G.N. — New species of Parmelia (Lichens) from Aus-

tralia, Austral. J. Bot. 1979 (1980), 27, 6 : 873-883, 6 fig. (Chemistry Dept.,

Austral. Natl. Univ., P. O. Box 4, Canberra A.C.T., 2600 Australia).

Source : MNHN, Paris

120 BIBLIOGRAPHIE LICHÉNOLOGIQUE

Diagn., descr., de Parmelia queenslandica, P. subtabacina, P. butleri, P. numin-

bahensis, P. springtonensis, P. norsticticata, esp. nouv. d'Australie. Discussion

autour de l'intégration des genres Pseudoparmelia Hale et Parmelina Hale. Pseu-

doparmelia conlabrosa Hale, Relicina subconnivens Hale et Pseudoparm. neo-

quintaria Hale sont transférés à Parmelia.

FERRARO L.I. — Una nueva especie de Parmeliaceae para el NE Argentino.

Hickenia 1979, 1 : 191-193, phot. (CONICET, Inst. Bot. del Nordeste, Casil-

la de Correo 209, 3400 Corrientes, Argentina).

Diagn., descr., ill. de Parmotrema masonii sp. nov. de Corrientes, affine de P.

delicatulum (Vain.) Hale. Clé pour les deux esp.

HAFELLNER J. und POELT J. — Die Arten der Gattung Caloplaca mit pluri-

locularen Sporen (Meroplacis, Triophthalmidium, Xanthocarpia). J. Hattori

Bot. Lab. 1979, 46 : 1-41, 44 fig, (Inst. Bot. Univ., A-8010 Graz).

Révision des Caloplaca ayant des spores pluriloculaires (le nombre de locules

n’est pas un caractére taxonom. suffisant pour les diviser en plus. genres). Ces

spores pluriloculaires semblent avoir évolué en 3 groupes. Clé aux 17 esp. et 1

sous-esp. reconnues. Pour chaque taxon : taxonom., descr. du thalle, des apo-

thécies et des spores, distr. La plupart des esp. sont connues de quelques loc.

seulement. Diagn., descr. de С. araguana sp. nov. du Venezuela, C. crocea ssp.

mexicana ssp. nov. du Mexique, С. oahuensis sp. nov. des Iles Hawaï. Nom.

nov. : C. vainioi (= Placodium brebissonii var, microspora Wainio). Lecidea

brebissonii Fée, L. crocea Krempelh., Blastenia kauaiensis H. Magn., Callopisma

tetramera Müll. Arg. sont transférés à Caloplaca. Lecanora brebissonii var.

subdecadens Nyl. est une esp. de Caloplaca.

JORDAN W.P. — Edrudia, a new genus from California (Lichenes : Teloschista-

ceae). Bryologist 1980, 83, 1 : 64-67, 3 fig. (Dept. Biol, Univ. San Fran-

cisco, San Francisco, Ca 94117 USA).

Diagn., descr. de Edrudia Jordan gen. nov., se distinguant de Polycauliona

Hue par des ascospores simples, des pycnidiospores filiformes et des rameaux

du thalle dorsiventralement compressés. E. constipans (Nyl.) c.n. (= Placo-

dium c.).

NOBLE W.J. — Catillaria columbiana comb. nov. from Western North America.

Bryologist 1980, 83 : 71-73 (Dept. Bot., Univ. British Columbia, Vancouver,

BC, Canada V6T 1W5).

Descr., distrib. de Catillaria columbiana (Merr.) c. n. (= Biatora c.), corticole,

de la Colombie britannique à la Californie.

POELT J. — Physcia opuntiella spec. nov. und die Lebensform der sprossenden

Flechten. Flora 1980, 169, 1 : 23-31, 3 fig., (Inst. Bot., Univ. Graz).

Diagn. descr. ill de Physcia opuntiella sp. nov., des localités sèches des

Alpes. Formation de propagules segmentées, semblables à des levures et nom-

mées blastides. Différents types de diaspores végétatives produites par les lichens.

Source - MNHN. Paris

BIBLIOGRAPHIE LICHÉNOLOGIQUE 121

PHYSIOLOGIE, CHIMIE

ASCASO C.A. — Rapid Method for the Quantitative Isolation of Green Algae

from Lichens. Ann. Bot. (London) 1980, 45, 4 : 483 (Inst. Edafologia Biol.

Veget., Serrano 115 bis, Madrid (6), Spain).

CZECZUGA B. - Investigations on Carotenoids іп Lichens IV : Representatives

of the Parmeliaceae Family. Nova Hedwigia 1980, 32, 1 : 105-111, 1 fig,

1 tabl. (Dept. Gener. Biol., Medic. Acad., 15-230 Bialystok, Poland).

Chromatographie en colonne et en couche mince de 11 esp. de Cetraria, Hy-

pogymnia, Parmelia et Pseudevernia. Sont isolés : neurosporéne, lycopéne, @-, B.,

Y- et Ó-caroténe, époxide B-carotène, 0- et B-cryptoxanthène, canthaxanthéne,

lutéine, lutéine époxide, zéa-, isozéa-, rubi- et astaxanthène, ester d'astaxan-

théne, violaxanthine, auro- et mutatochrome, mutatoxanthéne, 3,4-didéhydrol-

colpène, flavochrome et capsanthine. Le plus riche est Parmelia acetabulum.

GREEN Т.С.А., HORSTMANN J., BONNETT H., WILKINS A. and SILVES-

TER W.B. - Nitrogen fixation by members of the Stictaceae (Lichens) of

New Zealand. New Phytol. 1980, 84, 2 : 339-348, 3 fig., 3 tabl. (Biol. Sci.,

Univ. Waikato, Hamilton, New Zealand).

44 esp. des Stictacées sont testées. 10 esp. dont le phycobionte est une algue

verte n'ont pas d'activité nitrogénase.

LARSON D.W. — Seasonal change in the pattern of net CO; exchange in Umbili-

caria Lichens. New Phytol. 1980, 84, 2 : 349-369, 2 tabl., 5 fig. (Dept. Bot.

& Genetics, Univ. Guelph, Guelph, Ontario N1G 2W1).

Chaque esp. est adaptée uniquement aux conditions environnantes dans les-

quelles elle pousse. Chacune peut maintenir une balance énergétique positive

pourvu que sa distribution ne s'étende pas aux sites présentant des températures

en excès par rapport à certaines valeurs.

MATHEY A. — Contribution à l'étude de la famille des Trypéthéliacées (Lichens

Pyrénomycétes). Nova Hedwigia 1979 (1980), 31, 4 : 917-935, 23 fig. (Bot.

Gart. & Bot. Mus., Berlin-Dahlem, D-1000 Berlin 33).

Mise en évidence, chez les Trypéthéliacées, de la pariétine et de la lichéxan-

thone, responsables de la couleur orangée ou jaune sale des verrues thallines ou

de tout le thalle. Première isolation de l'acide sécalonique chez Trypethelium

eluteriae. Analyse d'une coupe de Laurera benguelensis. Culture des mycosym-

biotes de T. kunzei et de T. catervarium. Les substances produites par le myco-

symbiote de T. eluteriae sont antifongiques, inhibant le développement d'Asper-

gillus glaucus et de Penicillium sp.

SHIMADA S., SAITOH T., NAMIKI Y., SANKAWA U. and SHIBATA S. — New

siphulin derivatives from the lichen Siphula ceratites. Phytochemistry 1980,

19, 3 :467-469, 1 tabl. (Fac. Pharmac. Sci., Univ. Tokyo, Hongo, Bunkyo-ku,

Tokyo 113, Japan).

Structure de 2 nouveaux chroménones métabolites : proto- et oxysiphuline.

Source : MNHN, Paris

122 BIBLIOGRAPHIE LICHÉNOLOGIQUE

RÉPARTITION, ÉCOLOGIE, SOCIOLOGIE

ASTA-GIACOMETTI J. — Flore et végétation lichéniques des Alpes nord-occi-

dentales : écologie, biogéographie, écophysiologie, biodétection de la pollu-

tion fluorée. Thèse Univ. Sci. Méd. Grenoble, juin 1980 : 1-249, 71 tabl., 49

fig.

Description (géogr., climat., végétat.) de la région des Alpes nord-occidentales.

Étude de la végétation lichénique (corticole, terricole et humicole, saxicole) et

de ses relations avec l'altitude, les groupements forestiers, la nature du substrat.

Comparaison de la flore lich. des Alpes nord-occidentales avec celle de la France,

des Pays scandinaves. Rappelons que la région compte plus de 1200 espéces, soit

plus de la moitié de la flore franc. Influence de quelques facteurs écologiques

sur cert. associations et esp. lichéniques : СаСОз et pH pour les saxicoles, nutri-

tion minérale (Ca*t, K*, Mg'*) des esp. terricoles et corticoles. Les lichens indi-

cateurs de pollution :le fluor dans ces vallées alpines. Les résultats exposés selon

ce plan sont le fruit de recherches effectuées seule ou en collaboration. Ils per-

mettent de mettre en évidence l'importance de la végétation lichénique dans une

étude écologique complète, et les relations entre le substrat, la végétation phané-

rogamique, la pollution et les lichens. Bibliogr., liste des groupements liché-

niques, liste des esp. (avec leur département de récolte, et, pour les macrolich.,

l'étage auquel ils appartiennent), glossaire viennent compléter cet important

document. Un volume contenant les travaux déjà publiés sur ce sujet par J. Asta,

est joint à la thèse proprement dite.

AWASTHI D.D. and JOSHI M. — Macrolichens of Mussoorie Hills, Uttar Pra-

desh. Geophytology 1977, 7, 1 : 91-97 (Dept. Bot., Lucknow Univ., Luck-

now, India).

Liste avec loc. et quelques notes de 35 macrolichens, récoltées dans les col-

lines de Mussoorie. Clés pour les esp. de Collema, de Leptogium, de Parmelia,

d'Usnea, de Pyxine, de Physcia et d’Heterodermia.

AWASTHI D. D. and SINGH K. P. — Additions to the lichen flora of India IV.

Geophytology 1977, 7, 2 : 276-277, 5 fig. (Dept. Bot., Univ. Lucknow,

India).

Descr., ill. de Phylliscum testudineum A. Henssen nouv. pour l'Inde.

AWASTHI D.D. and DANGE K. — Additions to the lichen flora of India. V.

Natl. Acad. Sci. Letters 1978, 1, 1 : 11-13 (Dept. Bot., Lucknow Univ.,

India).

Descr. d'Acarospora fusca B. de Lesd., A. fuscata (Nyl.) Arn., A. oxytona

(Ach.) Mass. et A. subfuscescens (Nyl) H. Magn. récoltés en Uttar Pradesh,

nouv. pour l'Inde. Clé aux 13 esp. d'Acarospora connues en Inde.

AWASTHI D.D. and SHARMA L.R. — Additions to the lichen flora of Nepal.

1. Indian J. Bot. 1978, 1, 1-2 : 139-141 (Dept. Bot. Lucknow Univ., India).

Descr. et loc. de Anthrocothecium himalayense (Ras.) Awas., Bombyliospora

Source : MNHN, Paris

BIBLIOGRAPHIE LICHÉNOLOGIQUE 123

domingensis (Pers.) Zahlbr., Cetrelia isidiata (Asah.) Culb. et Culb. Coccocarpia

cronia (Tuck.) Vain., Haematomma puniceum (Sw.) Mass., Microglaena thelosto-

moides (Nyl.), Zahlbr., Parmelia subaurulenta Nyl., P. sublaevigata (Nyl.) Nyl.,

P. subsumpta Nyl., Phlyctella indica Awas., Physma byrsinum (Ach.) Müll., et

Tylophoron moderatum Nyl. nouv. pour le Népal.

BREDKINA L. I. — Lichenes montium Tian-Schan Centralis (Districtum Kocz-

kor et Naryn). Nov. Síst. Nizsh. Rast., Bot. Inst. Komarova, Akad. Nauk

SSSR 1979, 16 : 115-127, fig., en russe.

Données écol., végétation selon l'altitude. Liste de 57 lichens avec loc.

BJAZROV L.G. — Usnea sulphurea (Koenig) Th. Fr. ex Respublica Populari

Mongolia. Nov. Sist. Nizsh. Rast., Bot. Inst. Komarova, Akad. Nauk SSSR

1979, 16 : 127-129, en russe.

GOLUBKOVA N.S., SCHAPIRO Т.А. — Species generis Umbilicaria Hoffm. em.

Frey in URSS crescentes et examinatio earum chemotaxonomica. Nov. Sist.

Nizsh. Rast., Inst. Bot. Komarova, Akad. Nauk SSSR 1979, 16 : 133-153, 9

tabl., en russe.

Distr. et chimiotaxonomie de 34 esp. d’Umbilicaria.

KIRBY N.— New additions to the Lichen Flora of the Burren. Irish Nat. J.

1980, 20, 1 :45 (Dept. Bot., Univ. College, Galway).

7 lichens avec loc.

NOVRUSOV V.S. — Lichenes pro flora caucasi novi. Nov. Sist. Nizsh. Rast.,

Inst. Bot. Komarova, Akad. Nauk SSSR 1979, 16 : 155-158, en russe.

Liste de 35 esp. avec loc.

SINGH S.R.— А note on some macrolichen taxa from Pachmarhi, Madhya

Pradesh, India. Geophytology 1977, 7, 1 : 123-125 (Dept. Bot., Lucknow

Univ., India).

Liste de 16 taxons avec courte descr. et loc. au Pachmarhi.

SINGH S.R. and AWASTHI D.D. — A note on some micro-ichens from Pach-

marhi, Madhya Pradesh, India. Geophytology 1978, 8, 1 : 127-128 (Idem).

25 taxons avec loc. Caloplaca festivella (Nyl.) Kieff, C. pellodella (Nyl.)

Hasse, C. subnigricans H. Magn., Graphis subintegra Red., С. vittata Müll. Arg.,

Lecanora glabrata (Ach.) Malme, L. subcarnea (Sw.) Ach. et Ochrolechia africa-

na Zahlbr. sont nouv. pour l'Inde.

PENTECOST А. – Тһе Lichens and Bryophytes of rhyolite and pumice-tuff

rock outcrops in Snowdonia and some factors affecting their distribution.

J. Ecol. 1980, 68, 1 : 251-267, 8 fig., 7 tabl. (Dept. Applied Biol., Chelsea

College, London SW10 0QX).

La végétation de la rhyolite est dominée par Lecidea tenebrica tandis que

Source : MNHN, Paris

124 BIBLIOGRAPHIE LICHÉNOLOGIQUE

celle des pierres ponce volcaniques n'a pas d'esp. prépondérante. Ces dernières

roches apparaissent plus poreuses, mais leur perte en eau est plus rapide que chez

les rhyolites.

VARIA

BOTANICAL SOCIETY OF AMERICA — Abstracts of papers to be presented

at the University of British Columbia, Vancouver, 12-16 July 1980. Miscella-

neous Series Publications 1980, 158.

AHMADJIAN V. — Algal-fungal relationships in lichens; recognition, synthe-

sis, development (Dept. Biol, Clark Univ., Worcester, Massachusetts 01610

USA). — AHTI T. — Morphological intergradation in local populations of Clado-

nia (lichenized Ascomycetes) (Dept. Bot., Univ. Helsinki, SF-00170 Helsinki

17). - EGAN RSS. — Studies on the Lichen Family Parmeliaceae in Texas :

«Progress and Problems» (Dept. Biol, Univ. Nebraska at Omaha, Omaha NE

68182 USA). — EVERSMAN S. — Transmission electron microscope studies of

lichens fumigated with air pollutants (Biol. Dept., Montana State Univ., Boze-

man, Montana 59717 USA). - FAHSELT D. — Electrofocussed protein patterns

in populations of fruticose lichens (Dept. Pl. Sci., Univ. Western Ontario,

London, Ontario, Canada N6AN 5В7).- GARDNER C.R. and MUELLER

D.M.J. — The effects of several lichenic acids on membrane permeability in

Funaria hygrometrica Hedw. (Dept. Biol., Texas A & M Univ., College Station,

Texas 77843 USA). — HALE M. E. — Importance of cortical structure in lichens

(Dept. Bot., Smithsonian Inst., Washington D.C. 20560 USA). — LARSON D. —

Low temperature effects on temperate species of Umbilicaria (Dept. Bot. &

Genet., Univ. Guelph, Guelph, Ontario, Canada N1G 2W1). - LARSON D. —

Morphological control of wetting in some lichens and mosses (Idem). - MALA-

CHOWSKI J.A. — Status of Usnea cavernosa and Usnea trichodea in Eastern

North America (Dept. Bot. & РІ. Pathol., Michigan State Univ., East Lansing

МІС 48824 USA). — MOSER T.J., NASH ІП T.H. and LONK S.O. — Photosyn-

thetic patterns of dominant arctic lichens (Dept. Bot. & Microbiol., Arizona

State Univ., Tempe, Arizona 85281 USA). — OTTO С.Е. — Distribution of the

lichen Tholurna dissimilis (Norm.) Norm. in North America (Dept. Bot , Univ.

British Columbia, Vancouver, B.C., Canada V6T 2В1). - PEARSON L.C. —

Air pollution increases leakage of electrolytes from lichen cells (Dept. Biol.

Sci., Ricks College, Rexburg, Idaho 83440 USA). - RUNDEL P.W. — The rela-

tionships of morphological form and physiological response in coastal fog zone

lichens (Dept. Ecol. & Evolut. Biol., Univ. California, Irvine, Ca 92717 USA). —

SAYCE К. — SEM observations of the algal-fungal interactions : methods for in-

creased clarity (Bot. & Microbiol. Dept., Arizona State Univ., Tempe, Arizona

85281 USA). — SIGAL L.L. — The effects of peroxyacetyl nitrate fumigations

on lichens from southern California forests (Oak Ridge National Lab., Environm.

Sci. Div., P. O. Box X, Oak Ridge, TN 37830 USA). — SIMONSEN P. — Micro-

organisms associated with non-symbiotic nitrogen fixation in boreal forests

Source : MNHN, Paris

BIBLIOGRAPHIE LICHÉNOLOGIQUE 125

and eastern hardwood forests (Dept. Forests Sci. & Soil Sci., Univ. Alberta,

Edmonton, Alberta, Canada T6G 2E3). — SLOCUM R.D. — Light and electron

microscopic investigations in the Dictyonemataceae (Basidiolichens) (Cell

Research Inst. & Dept. Bot., Univ. Texas, Austin, TX 78712 USA). — SMITH

C.W. — The lichen and bryophyte flora of fumaroles in Hawaii Volcanoes

National Park (Dept. Bot., Univ. Hawaii at Manoa, Honolulu, Hi 96822 USA).

— WOOD J. — Relationship between water content and morphology of Ramalina

menziesii Tayl. along an altitudinal gradient (Dept. Biol. Sci., Univ. California,

Santa Barbara, California 93106 USA).

Source : MNHN. Paris

INFORMATIONS

Vient de paraître :

TAYLOR R.J. and LEVITON А.Е. ed. — The Mosses of North America. San

Francisco. Pacific Div., Amer. Assoc. Advanc. Sci., 1980, 170 p., ill. (communi-

cations bryol. au 59e meeting annuel. — Prix : $ 11.95 U.S., auprès de Secretary,

Pacific Division, AAAS, C/o California Academy of Sciences, San Francisco,

Ca 94118, USA).

CATCHESIDE D.G. — Mosses of South Australia. South Australia, Woolman

1980, 364 p. Ш. (prix : $ 13.20, auprès de Publications officer, 282 West

Beach Rd, Netley 5037 South Australia).

LANDWEHR J. - Atlas van de Nederlandse Levermossen. Zutphen, Thieme

et Cie, 1980, 288 р., ill. (prix : DFL. 45, auprès de Konink. Ned. Natuurhist.

Vereniging, B Hoogenboomlaan 24, 1718 BJ Hoogwoud, Holland).

ZEHR D.R. — An assessment of variation in Scapania nemorosa and selected

related species (Hepatophyta). Bryoph. Biblioth. 1980, 15 : 1-140, ill. (prix :

DM. 40, auprès de Cramer J., D-3300 Braunschweig, in den Springäckern 2).

CRUM H.A. and ANDERSON L.I. — Mosses of Eastern North America.

New York, Columbia Univ. Press, 1981, 2 vol., 1328 p., ill. (prix : $ 78.00 US).

Ces ouvrages seront analysés dans Cryptog., Bryol. Lichénol. 1981, 2, 2.

Commission paritaire 26-1-1976 N° 57365

Dépôt légal n° 10829 - Imprimerie de Montligeon

Sorti des presses le 31 mars 1981

Source : MNHN. Paris

COLLOQUE INTERNATIONAL

du CNRS N° 258

ÉCHANGES IONIQUES TRANSMEMBRANAIRES

CHEZ LES VÉGÉTAUX

TRANSMEMBRANE IONIC EXCHANGES IN PLANTS

org. : б. Ducet, R. Heller, M. Thellier

Universités de Rouen et Paris VII - 5-11 juillet 1976

е études électrophysiologiques @ cas particulier des transferts d'anions et de molécules

organiques @ localisation d'ions et aspects structuraux et moléculaires @ intervention

d'échanges ioniques dans les régulations intercellulaires

kinetic and thermodynamic considerations, model systems

metabolic and other couplings, ATPases

particular features of anionic transfers

electrophysiology of the ionic transfer

absorption of organic molécules

localization, molecular and structural aspect of the transfers

interference of the transmembrane transfers in other processes than absorption

€ ion exchanges іп cell organites

(69 communications dont 64 en anglais et 5 en frangais) i

21x29, 7 - 608 pages - broché 180 F

286 fig. - 89 tabl. - 30 phot.

ISBN 2-222-02021-2

(co-édition CNRS-Université de Rouen)

ditions du CNRS

quai Anatole France: 75700 Paris

CCP Paris 9061-11 - Tél. 555.92.25

DR ЕА ЕЕЕ RIED A SSeS

chez son libraire o

5 à défaut aux Editions du CNRS (chèque joint) ГІ

profession et demande votre documentation

0 Sciences humaines — .

adresse C) Sciences exactes et naturelles

O Trésor de la langue Française

ach О Revue de l'Art

BS ILLE TCI LLL й

Source : MNHN, Paris

SOMMAIRE

R.H. ZANDER. — Descriptions and illustrations of Barbula, Pseudocros-

sidium and Brycerythrophyllum (p. р.) of Mexico. .............

R. GAMBARDELLA, R. LIGRONE AND R. CASTALDO. — Ultrastruc-

ture of the sporophyte foot in Риаеосетов.............. %

P. TIXIER. — La notion d'espéce chez le genre Cololejeunea. Le com-

plexe Cololejeunea floccosa (Lehm. & Lindenb.) Schiffn. . ........

5.5. KUMAR & S.K. VERMA. — Cytological observations on some West

НАУАН адм о УШІН АН Сы Med ore VT esie

E.A. ODU. - Reproductive phenology of some tropical African mosses. .

D. GRIFFIN, Ш. — Spore ornamentation in Leiomela (Musci : Bartra-

[SU eSATA «rate gia ACIE RE

BIBLIOGRAPHIE BR YOLOGIQUE: 52,202 2

BIBLIOGRAPHIE LICHÉNOLOGIQUE

INFORMETIONS TR ie eee rate ces ане