FLORA OF SOUTHERN

BRYOPHYTA

Fascicle 2

Gigaspermaceae— Bartramiaceae

by Robert E. Magill

Botanical Research Institute

Department of Agriculture and Water Supply

Republic of South Af lica

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REPUBLIC OF SOUTH AFRICA

REPUBLIEK VAN SUID-AFRIKA

DEPARTMENT OF AGRICULTURE AND WATER SUPPLY

DEPARTEMENT VAN LANDBOU EN WATER VOORSIENING

FLORA OF SOUTHERN AFRICA

BRYOPHYTA

PARTI, FASCICLE 2

ISBN 0 621 10325 X

G.P.-S.

Digitized by the Internet Archive

in 2016

https://archive.org/details/floraofsoutherna12unse

Dedicated to the memory of

EDMUND ANDRE CHARLES LOUIS ELOI SCHELPE

Bryologist

1924-1985

FLORA OF SOUTHERN AFRICA

which deals with the territories of

SOUTH AFRICA, TRANS KEI, LESOTHO, SWAZILAND, BOPHUTHATSWANA,

SOUTH WEST AFRICA/NAMIBIA, BOTSWANA AND VENDA

BRYOPHYTA

PART 1 MOSSES

Fascicle 2 Gigaspermaceae-Bartramiaceae

by

Robert E. Magill

Edited by

0. A. Leistner

Editorial Committee: B. de Winter, D. J. B. Killick, G. E. Gibbs Russell and 0. A. Leistner

Botanical Research Institute,

Department of Agriculture and Water Supply

1987

CONTENTS

Page

New taxa and new combinations published in part 1 , fascicle 2 vi

Geographical regions referred to in this fascicle vii

Introduction to fascicle 2 ix

Conspectus of classification 293

Literature cited 294

Provisional key to the families of fascicle 2 296

GIGASPERMACEAE 299

EPHEMERACEAE 305

FUNARIACEAE 311

SPLACHNACEAE 331

BRYACEAE (by J. van Rooy & R. E. Magill) 335

MNIACEAE 395

EUSTICHIACEAE 399

RHIZOGONIACEAE 401

AULACOMNIACEAE 405

BARTRAMIACEAE 407

Index to fascicle 2

439

NEW TAXA AND NEW COMBINATIONS PUBLISHED IN PART 1

FASCICLE 2

Anacolia breutelii (C. Mull.) Magill, comb. nov. , p. 41 1

Anacolia breutelii (C. Mull.) Magill var. squarrifolia (Sim) Magill , stat. nov. , p. 412

Bartramia compacta Hornsch. var. macowaniana (C. Mull.) Magill, stat. nov., p. 413

Breuteiia elliptica Magill, sp. nov., p. 435

Breutelia substricta (C. Mull.) Magill, comb, nov., p. 437.

Ephemerum namaquense Magill, sp. nov., p. 307

Funaria clavata (Mitt.) Magill, comb, nov., p. 326.

Funaria succuleata (Wager & Wright) Magill, comb. nov. , p. 321

Physcomitrium spathulatum (Hornsch.) C. Mull. var. sessile (Shaw) Magill, stat. nov., p. 319

Quathlamba Magill, gen. nov., p. 421

Quathlamba debilicostata Magill, sp. nov., p. 421

Date of publication: July, 1987.

GEOGRAPHICAL REGIONS REFERRED TO IN THIS FASCICLE

B — Botswana

CC — central Cape

CE — eastern Cape

CN — northern Cape

CS — southern Cape

CNW — northwestern Cape

CSW — southwestern Cape

L — Lesotho

N — Natal

O — Orange Free State

S — Swaziland

SWA — South West Africa/Namibia

T — Transkei

TC — central Transvaal

TE — eastern Transvaal

TN — northern Transvaal

TS — southern Transvaal

TW — western Transvaal

Z — Zululand

vii

INTRODUCTION TO FASCICLE 2

The following text constitutes Fascicle 2 of Part 1 of Volume Bryophyta in the Flora of

southern Africa Cryptogam series. This fascicle includes the families Gigaspermaceae to Bartra-

miaceae or roughly tne Acrocarpi-diplolepideae (see Conspectus of classification, p. 293).

The notes on Study and identification and Characters used in keys and description of mosses

from Fascicle 1 also apply here. One important difference is that most of the species in Fascicle 2

have double peristomes, i.e. exostome and endostome (see Glossary and Fig. 2: 3 of Fascicle 1).

Occasionally only one of these structures is present e.g. Conostomum (exostome only) or the

capsules of some species are gymnostomous. The capsules in only a few species in this fascicle are

cleistocarpic.

Variation in the structure and development of the endostome is quite common in members

discussed in this fascicle. This variation is expressed in (1) height of the basal membrane, (2)

structure and height of the segments especially in relation to the exostome teeth, and (3) devel-

opment, number and height of the cilia.

Although the endostome can be examined using the method for observing capsule mouth parts

outlined in Fascicle 1 , observation can be enhanced by carefully teasing the endostome away from

the exostome teeth on one or two of the mouth sections. Some of the mouth sections should be left

intact since differences in height of endostome and exostome, and position of segments in

relationship to teeth (opposite or alternate) are occasionally important.

The illustrations were prepared by Mrs Rita Weber and Ms Gillian Condy; for technique and

procedure see Introduction in Fascicle 1 (p.xv).

Research on this fascicle was partly supported by grants to the Missouri Botanical Garden

from the National Science Foundation (BSR- 83 15245) and the National Geographic Society

(2719-83).

Erratum in Part 1 Fascicle 1 :

Page 225: in the caption of Map 86 Weisiopsis pulchriretis and Weisiopsis involuta must be

switched.

IX

293

CONSPECTUS OF CLASSIFICATION

DIVISION BRYOPHYTA

Fascicle 1:

CLASS SPHAGNOPSIDA

ORDER SPHAGNALES

Family Sphagnaceae

CLASS ANDREAEOPSIDA

ORDER ANDREAEALES

Family Andreaeaceae

CLASS BRYOPSIDA

ORDER DICRANALES

Family Fissidentaceae

Nanobryaceae

Archidiaceae

Ditrichaceae

Seligeriaceae

Dicranaceae

ORDER POTTIALES

Family Calymperaceae

Encalyptaceae

Pottiaceae

Bryobartramiaceae

Gnmmiaceae

Fascicle 2:

ORDER FUN ARI ALES

Family Gigaspermaceae

Epnemeraceae

Funariaceae

Splachnaceae

ORDER BRYALES

Family Bryaceae

Mniaceae

Eustichiaceae

Rhizogoniaceae

Aulacomniaceae

Bartramiaceae

Fascicle 3:

ORDER ORTHOTRICHALES

Family Erpodiaceae

Rhachitheciaceae

Ptychomitriaceae

Orthotrichaceae

Rhabdoweisiaceae

Racopilaceae

ORDER ISOBRYALES

Family Fontinalaceae

Wardiaceae

Hedwigiaceae

Cryphaeaceae

Leucodontaceae

Prionodontaceae

Trachypodaceae

Pterobiy'aceae

Meteonaceae

Phyllogoniaceae

Neckeraceae

Lembophyllaceae

ORDER HOOKERIALES

Family Hookeriaceae

Fascicle 4:

ORDER THUIDIALES

Family Fabroniaceae

Leskeaceae

Thuidiaceae

ORDER HYPNOBRYALES

Family Amblvstegiaceae

Brachytheciaceae

Entodontaceae

Plagiotheciaceae

Hypnaceae

Hylocomiaceae

CLASS POLYTRICHOPSIDA

ORDER POLYTRICHALES

Family Polytrichaceae

294

LITERATURE CITED IN FASCICLE 2

Brotherus, V. F., 1924-1925. Musci. In

Engler & Prantl (eds), Die natiirlichen Pfian-

zenfamilien. Edn 2, Vol. 10-11. Leipzig.

Bruch, P., Schimper, W. P. & Gumbel, T.,

1836-1855. Bryologia Europaea seu genera

muscorum europaeorum monographice illu-

strata, 65 fasc. Stuttgart.

Codd, L. E. & Gunn, M., 1982. The collect-

ing activities of Anton Rehmann (1840-

1917) in South Africa. Bothalia 14: 1-14.

Crundwell, A. C. & Nyholm, E., 1964. The

European species of the Bryum erythrocar-

pum complex. Trans. Br. bryol. Soc. 4:

597-637.

Dixon, H. N., 1920. New and interesting

South African Mosses. Trans. R. Soc. S. Afr.

8: 179-224.

Dixon, H. N. & Gepp, A., 1923. Rehmann’s

South African Mosses. Kew Bull. 6:

193-238.

Fife, A. J., 1982. Taxonomic and nomenclatu-

re! observations on the Funariaceae. 2. Lec-

totypification of Physcomitrium subgen.

Cryptopyxis (C. Mull.) Broth. Lindbergia 8:

75-76.

Fife, A. J., 1985. A generic revision of the

Funariaceae (Bryophyta: Musci), part 1. J.

Hattori bot. Lab. 58: 149-196.

Gangulee, H. C., 1974. Mosses of eastern

India and adjacent regions fasc. 4, Calcutta.

Iwatsuki, Z. & Koponen, T., 1972. On the

taxonomy and distribution of Rhodobryum

roseum and its related species (Bryophyta).

Acta Bot. Fenn. 96: 1-22.

Koponen, T., 1972. The east Asiatic species of

Plagiomnium sect. Rostrata (Bryophyta).

Acta Bot. Fenn. 97: 1-29.

Koponen, T., Li, X.-J. & Zang, M., 1982. A

synopsis of Rhodobryum (Musci, Bryaceae)

in China. Ann. Bot. Fenn. 19: 75-80.

Magill, R. E. & Schelpe, E. A., 1979. The

bryophytes of southern Africa: An annotated

checklist. Mem. bot. Surv. S. Afr. 43: 1-38.

Manuel, M., 1980. Miscellanea Bryologica.

II. Classification of Rhizogonium Brid., Pen-

zigiella hookeri Gang, and some nomina

nuda. Cryptogam., bryol. lichen. 1: 67-72.

Mohamed, M. A., 1979. A taxonomic study of

Bryum billardieri Schwaegr. and related

species. J. Bryol. 10: 401-465.

Muller, C., 1899. Contributiones ad Bryolo-

giam austroafram. Hedwigia 38: 52-155.

Ochi, H., 1970. A revision of the Subfamily

Bryoideae in Australia, Tasmania, New Zea-

land and the adjacent islands. J. Fac. Educ.

Tottori Univ. 21: 7-67.

Ochi, H., 1972. A revision of African Bryoi-

deae, Musci (First Part). J. Fac. Educ. Tot-

tori Univ. 23: 1-126.

Sainsbury, G. O. K., 1955. A handbook of

the New Zealand Mosses. Bull. R. Soc. N.Z. 5:

1^190.

Sayre, G., 1978. Musci. In D. H. Pfister

(ed.), Cryptogams of the United States North

Pacific Exploring Expedition 1853-1856.

XV + 196 pp. Farlow Reference Library and

Herbarium of Cryptogamic Botany, Cam-

bridge, Massachusetts.

Scott, G. A. M. & Stone, I. G., 1976. The

Mosses of southern Australia. London: Aca-

demic Press.

Shaw, J., 1878. Catalogue of the mosses of the

Cape Colony. Cape Monthly Magazine 17:

311-320.

Shaw, J., 1985. Nomenclatural Changes in the

Bryaceae Subfamily Mielichhoferioideae.

The Bryologist 88: 28-30.

Shaw, J. & Crum, H., 1984. Peristome homo-

logy in Mielichhoferia and a taxonomic ac-

count of North American species. J. Hattori

bot. Lab. 57: 363-381 .

Sim, T. R., 1926. The Bryophyta of South Af-

rica. Trans. R. Soc. S. Afr. 15: 1^175.

295

Smith, A. J. E. & Whitehouse, H. L. K.,

1978. An account of the British species of the

Bryum bicolor complex including B. dunense

sp. nov. J. Bryol. 10: 29-41.

Sullivant, W. S. & Lesquereux, L., 1859.

Characters of some new Musci collected by

Charles Wright in the North Pacific Explor-

ing Expedition. Proc. Am. Acad. Arts Sci. 4:

275-282.

SYED, H., 1973. A taxonomic study of Bryum

capillare Hedw. and related species. J.

Bryol. 7: 265-326.

Touw, A., 1983. The identity of Rhodobryum

spathulatum (Homsch.) Poes. Lindbergia 9:

151-152.

Van Zanten, B.O., 1973. Musci. In Marion

and Prince Edward Islands. Report on the

South African biological and geological ex-

pedition/ 1965-1966 . Cape Town: Balkema.

Wilczek, R. & Demaret, F., 1976. Les

especes beiges du “complexe Bryum bico-

lor" (Musci). Bull. Jard. Bot. Nat.

Belg.lBull. Nat. Plantentuin Belg. 46:

511-541.

296

PROVISIONAL KEY TO THE FAMILIES OF FASCICLE 2

The following key is provided to allow access to the families treated in this fascicle. This key

is a continuation of the family key in Fascicle 1 and should be used in connection with it. When

necessary, couplets are incorporated into the key that refer to taxa which will be treated in other

fascicles.

1 Plants pleurocarpic or of pleurocarpic habit (Fascicle 3 & 4)

1 Plants acrocarpic or appearing acrocarpic but branches arising from a creeping stem or rhi-

zome:

2 Plants with a creeping stem or rhizome:

3 Plants forming dense mats or cushions on rock or bark; leaf cells frequently papillose

ORTHOTRICHACEAE (Fascicle 3)

3 Plants scattered or forming large turfs on soil; leaf cells smooth:

4 Plants small to medium-sized; leaf margins entire, without differentiated marginal

cells; capsules immersed to short-exserted GIGASPERMACEAE (p. 299)

4 Plants large to robust; leaf margins denticulate to dentate, marginal cells differentia-

ted and forming distinct border; capsules long-exserted BRYACEAE (p. 335)

2 Plants acrocarpic, without creeping stem or rhizome:

5 Leaves ecostate or costa rudimentary:

6 Plants minute, on dense, persistent protonema; leaves few, very small, margins

dentate EPHEMERACEAE (p. 305)

6 Plants small; leaves numerous, larger, margins entire:

7 Leaves orbicular-apiculate; costa absent; capsule immersed

GIGASPERMACEAE (p.299)

7 Leaves ovate to elliptical, acute; costa absent to rudimentary; capsule exserted . . .

BARTRAMIACEAE (p. 407)

5 Leaves costate:

8 Plants minute, usually on persistent protonema; capsules cleistocarpic, small, ±

sessile and globose EPHEMERACEAE (p. 305)

8 Plants larger, without persistent protonema; capsules stegocarpic or, if cleistocar-

pic, capsule not sessile and globose:

9 Leaves strongly toothed:

10 Leaves narrow with small, rounded leaf cells; costa percurrent

RHIZOGONIACEAE (p. 401)

10 Leaves broader with larger, ± rhomboidal leaf cells; costa short- to long-ex-

current:

11 Marginal leaf cells differentiated, forming ± distinct border; marginal

teeth unicellular BRYACEAE (p. 335)

1 1 Marginal leaf cells not differentiated, marginal teeth generally multi-cellu-

lar SPLACHNACEAE (p. 331)

9 Leaves serrate to entire:

1 2 Leaf cells papillose or prorate:

1 3 Leaves distichous, short and concave EUSTICHIACEAE (p. 399)

13 Leaves many ranked, elongated and flat BARTRAMIACEAE (p. 407)

297

12 Leaf cells smooth:

14 Leaf with distinct border:

15 Leaves oblong to elliptical; apices rounded or emarginate

MNIACEAE (p. 395)

15 Leaf apices acute to acuminate or obtuse:

16 Alar cells differentiated, enlarged, in single row; border strong and

extending to leaf base BARTRAMIACEAE (p. 407)

1 6 Alar cells not differentiated; border not distinct in leaf base:

17 Upper laminal cells with pointed ends, regular, ± rhomboi-

dal BRYACEAE (p. 335)

17 Upper laminal cells mostly with flattened ends, not uniform, trun-

cate-rhomboidal FUNARIACEAE (p. 3 1 1 )

14 Leaves without distinct border:

18 Leaf cells rounded or angular, subquadrate to hexagonal:

19 Leaf margins serrate; laminal cells incrassate

AULACOMNIACEAE (p. 405)

19 Leaf margins entire:

20 Upper laminal cells ± irregular in size and shape; walls firm, ±

wavy; plants with rhizomes GIGASPERMACEAE (p. 299)

20 Upper laminal cells regular, hexagonal to subrectangular, thin-

walled FUNARIACEAE (p. 311)

18 Leaf cells distinctly longer than broad, mostly rectangular or rhomboi-

dal:

21 Upper laminal cells rectangular to linear-rectangular:

22 Leaves broad, widest at midleaf or above FUNARIACEAE (p. 311)

22 Leaves narrow, widest at base:

23 Leaves linear-subulate above ovate base; costa broad, flattened

in cross section; axillary tubers frequent BRYACEAE (p. 335)

23 Leaves ovate-lanceolate to linear-lanceolate; costa narrow,

rounded in cross section; tubers absent

BARTRAMIACEAE (p. 407)

21 Upper laminal cells rhomboidal to oblong-rhomboidal; truncate-rhom-

boidal or fusiform:

24 Upper laminal cells with pointed ends, regular, rhomboidal to ob-

long-rhomboidal or fusiform BRYACEAE (p. 335)

24 Upper laminal cells with ends generally flattened, not uniform,

truncate-rhomboidal to rectangular FUNARIACEAE (p. 311)

299

GIGASPERMACEAE

Plants small, scattered or loosely caespitose, glaucous green to light green or dark green;

terricolous. Branches erect, arising from a long, highly branched, aphyllous rhizome; in section

central strand weak or absent, cortical cells more or less uniform. Leaves appressed to erect

spreading, generally concave, larger above, orbicular to broadly elliptical or obovate; apex

rounded and abruptly apiculate or long-acuminate; margins plane, entire. Costa absent or, if

present, short- to long-excurrent; in section cells not strongly differentiated. Laminal cells small,

angular or quadrate to hexagonal, thin-walled, smooth; basal cells slightly larger, rectangular.

Gemmae occasionally present at apex of leafy stems, lenticular.

Autoicous or dioicous with androgametophytes smaller. Perichaetia terminal, leaves generally

distinct, larger. Seta very short or elongate; capsule immersed or exserted, cleistocarpic and

subspherical or stegocarpic, gymnostomous and cupulate to short-cylindrical; exothecial cells lax,

smaller and transversely rectangular at mouth; stomata present at base of urn; peristome absent;

operculum plano-convex, apiculate; calyptra very small; spores large, granulate.

Known primarily from the Southern Hemisphere, Gigaspermaceae contains four genera. Only the American genus

Lorentziella, recognized by its costate leaves and cleistocarpic capsules, is not represented in southern Africa.

Gigaspermum, with three species, is the most widely distributed genus and is found in Europe, Africa, Australia and

New Zealand. Chamaebryum and Oedipodiella were both described from southern Africa. A variety of a species of the

latter, O. australis (Wager & Dix.) Dix. var. catalaunica P. Varde, has been described from Europe.

Identification of plants with capsules or gemmae should not prove difficult; sterile plants, especially of Oedipodiella

may be more problematical; however, detection of the rather copious rhizome system will properly place specimens in this

family. It is interesting that in all three genera found in southern Africa the rhizome is light yellow to brownish, aphyllous

and with only short, scattered rhizoids.

1 Leaves ecostate 1. Gigaspermum

1 Leaves costate:

2 Costa excurrent as smooth, hyaline awn; capsule exserted 2. Chamaebryum

2 Costa very short-excurrent; capsule immersed 3. Oedipodiella

1. GIGASPERMUM

Gigaspermum Lindb. in Ofvers. K. VetenskAkad. Forh. 21; 599 (1865); Broth, in Natiirl.

PflFam. 10: 316 (1924); Sim, Bryo. S. Afr. 289 (1926); Sainsb., N. Zeal. Mosses 240 (1955).

Type species: G. repens (Hook.) Lindb.

Plants small and inconspicuous when sterile; terricolous. Branches erect, leafy, simple, scat-

tered along leafless rhizome. Leaves small, distant, rounded with abruptly apiculate apices, ecos-

tate. Laminal cells small, thin-walled.

Autoicous. Perichaetia large, leaves distinct. Capsule immersed, gymnostomous, cupulate;

operculum convex with short apiculus; calyptra small, campanulate; spores large.

A genus of 3 species, Gigaspermum is the most widely distributed genus in the family. The species found in southern

Africa, G. repens, is also known from Australia and New Zealand.

Gigaspermum repens (Hook.) Lindb. in

Ofvers. K. VetenskAkad. Forh. 21:599(1865);

Broth, in Natiirl. PflFam. 10: 316 (1924); Sim,

Bryo. S. Afr. 289 (1926); Scott & Stone, Moss.

S. Aust. 250 (1976). Type: Australia, Menzies

s.n., 1791 (BM).

Anictangium repens Hook., Musci Exot. 2: 106 (1819).

Leptangium repens (Hook.) Mitt, in J. Linn. Soc., Bot. 4:

79(1859).

Physcomitrium breutelii C. Miill . in Bot. Ztg 13: 749

(1855). Leptangium breutelii (C. Miill.) Jaeg. in Verh. St.

Gall, naturw. Ges. 1874 — 75: 155 (1876). Gigaspermum

breutelii (C. Mull.) Par., Ind. Bryol. 511 (1896). Type.

300

Gigaspermaceae

Cape, Saldanha Bay, Breutel s.n. (BM, lecto.!), vide Fife

in Bryologist 83: 476 (1981).

Plants small, light green to glaucous

green, scattered or loosely caespitose; terrico-

lous. Branches erect, 2-5 mm high, arising

from a light yellow, highly branched, long-

creeping, aphyllus rhizome; in section rhizome

round, central strand absent, cortical cells in

4-5 rows, larger, thin-walled, outer row nar-

rower, branches in section round, central strand

indistinct, cells small, thin-walled, cortical

cells in 3-4 rows, thin-walled. Leaves distant,

erect-spreading, concave; orbicular-apiculate,

0,5 -0,8 mm long; apiculus of several clear

cells; margins plane, entire. Costa absent. Lam-

inal cells angular, quadrate to subhexagonal,

smooth; basal cells short-rectangular.

Autoicous. Perigonia axillary. Perichaetia

terminal on erect branches, leaves highly differ-

entiated; elliptical, long-acuminate, to 2,5 mm

long; laminal cells rhomboidal to rectangular,

hyaline. Seta very short, 0,3-0, 5 mm long,

yellowish; capsule immersed, gymnostomous,

cupulate, 0,8- 1,0 mm long, brownish yellow,

mouth very broad; exothecial cells lax, quadrate

to hexagonal, smaller in base, at mouth with

single row of transversely rectangular cells; sto-

mata present at base of urn, phaneropore; peri-

stome absent; operculum plano-convex, apicu-

late, cells not twisted; calyptra small, campanu-

late; spores rounded to angular, 80-100 pm,

reddish brown, finely granulate. Fig. 84.

Fig. 84. — Gigaspermum repens: 1. habit, x 1,2.

habit, x 10; 3. stem in cross section, x 175; 4. leaves, x

40; 5. leaf, X 120; 6. leaf in cross section, x 175; 7.

perichaetial leaf, x 40; 8. capsule and detached operculum,

x 10. ( 1 -8, Garside 6566).

Gigaspermaceae

301

Although G. repens is the most widely distributed

species in the genus, the plants are rare in southern Africa.

The plants form small patches among other mosses on dry,

rocky soils in central South West Atrica/Namibia, western

and central Cape, Orange Free State and southern Lesotho.

Map 111.

Vouchers: Garside 6566; Magill 4669; Volk 5253.

The scattered, sterile stems with minute, distant leaves

are difficult to find; however the conspicuous perichaetial

leaves and short, immersed, broad-mouthed capsules are

very distinctive and will quickly identify the species.

2. CHAMAEBRYUM

Chamaebryum Ther. & Dix., J. Bot., Lond. 60: 106 (1922); Broth, in Natiirl. PflFam. 10: 315

(1924); Sim, Bryo. S. Afr. 289 (1926). Type species: C.pottioides Ther. & Dix.

Plants small, caespitose; terricolous. Branches erect, densely leaved, arising along a long,

branched rhizome. Leaves crowded, broad, piliferous. Costa well developed, excurrent. Lamina l

cells small, thin- walled.

Monoicous. Perichaetia inconspicuous, leaves slightly larger. Capsule exserted, gymnosto-

mous, mouth narrower than urn; spores large.

The monotypic genus Chamaebryum, is endemic to southern Africa.

Chamaebryum pottioides Ther. & Dix.

in J. Bot., Lond. 60: 106 (1922); Sim, Bryo. S.

Afr. 289 (1926). Lectotype: Cape, Cape Town,

Wager 633 (BM, lecto.!; PRE!), vide Fife in

Bryologist 83: 475 (1981).

Plants small, light green to grey-green,

caespitose; terricolous. Stems erect, 2—6 mm

high, arising from a brownish, highly branched,

long-creeping rhizome; in section rhizome

round, central strand absent, cortical cells in 6

rows, large, thin-walled, outer 2 rows smaller,

brownish; branches in section round, central

strand not well defined, of 8—10 smaller, thin-

walled cells. Leaves crowded above, appressed

dry, erect-spreading wet; obovate to orbicular,

0,8— 1,2 mm long; apex obtuse, cuspidate to

piliferous; base not differentiated; margins

plane, entire. Costa percurrent in lower leaves,

excurrent as short, hyaline awn in upper leaves,

to 0,5 mm long; in section subround, bulging

dorsally, cells in 3-4 rows, not strongly differ-

entiated. Laminal cells subquadrate to angular,

smooth; basal cells quadrate to very short-rect-

angular. Gemmae rare, produced at apex of

sterile plants, bulging lenticular with a pro-

nounced hyaline, distal apiculus, to 0,2 mm in

diameter, cells ± hexagonal.

Monoicous. Perigonia terminal. Perichae-

tia terminal on erect branches; leaves slightly

larger. Seta erect, 0,6— 1 ,8 mm long, yellowish

when wet, greyish when dry; capsule exserted,

gymnostomous, short cylindrical, 1,0 -1,3 mm

long, bulging and reddish yellow when wet,

irregularly striated to sulcate and greyish when

dry, mouth narrow; exothecial cells lax, sub-

hexagonal, 3-4 rows at mouth transversely

rectangular; stomata present at base of urn,

weakly cryptopore; peristome absent; opercu-

lum plano-convex with minute apiculus, re-

maining attached to columella when capsule

first opens; calyptra not seen; spores rounded,

50-55 pm, reddish brown, granulate. Fig. 85:

1-8.

Endemic to southern Africa, C. pottioides is restricted

to drier, rocky areas of southern South West Africa/Nami-

bia and the western, central, northern and southern Cape.

The species frequently forms large colonies in association

with Desmatodon. Goniomitrium, Ephemerum and Pleuri-

dium. Map 1 12.

Map 112. — • Chamaebryum pottioides

302

Gigaspermaceae

Gigaspermaceae

303

Vouchers: Goldblatt 2399b; Lavranos 15554a; Magill

& Schelpe 3871, 4053; Oliver, Tolken & Venter 575; Van

Zanten 7608360a.

The concave, piliferous leaves, rhizomatous habit and

elevated, narrow-mouthed capsules will place this species.

Young or sterile plants could be confused with Goniomi-

trium but it lacks a rhizome and the plants are dark green

and scattered, with shorter seta and broad-mouthed cap-

sules. Sterile colonies of C. pottioides have a greyish

colouration when dry and are frequently quite compact.

3. OEDIPODIELLA

Oedipodiella Dix. in J. Bot., Lond. 60: 105 (1922); Broth, in Natiirl. PflFam. 10: 315 (1924);

Sim, Bryo. S.Afr. 288 (1926). Type species: O. australis (Wager & Dix.) Dix.

Plants medium-sized, dark green; terricolous. Branches erect, simple, scattered along leaf-

less, branching rhizome. Leaves mostly obovate; apex rounded-apiculate; costa short-excurrent.

Laminal cells small, quadrate to hexagonal. Gemmae produced at apex, lenticular.

Perichaetia terminal, leaves Ungulate; costa

carpic, subglobose-rostrate; spores large.

The genus is known from southern Africa and Europe.

Oedipodiella australis (Wager & Dix.)

Dix. in J. Bot., Lond. 60: 105 (1922); Sim,

Bryo. S. Aff. 288 (1926). Syntypes: Natal,

Umkomaas, Wager s.n.; Transvaal, Waterval,

Wager s.n.; Pretoria, Wager s.n. (all PRE!).

Oedipodium australe Wager & Dix. in Trans. R. Soc. S.

Afr. 4:3(1914).

Plants small to medium-sized, scattered or

loosely caespitose, dark green; terricolous.

Branches erect, 2—5 mm long, arising from a

light yellow, branching, aphyllus rhizome; in

section rhizome round, central strand absent,

cortical cells in 5-6 rows, lax, thin-walled,

outer 1—2 rows smaller; branches in section

round, central strand not well defined, cortical

cells in 4-5 rows, thin-walled. Leaves crowded

above, weakly contorted when dry, wide-

spreading when wet; obovate to spathulate or

broadly elliptical, 2-4 mm long; apex rounded,

abruptly apiculate; base oblong; margins plane,

entire. Costa short-excurrent, ventral superfi-

cial cells smooth, dorsal superficial cells pro-

rate; in section rounded, bulging dorsally, guide

cells 2, small, ventral surface cells 2, small,

dorsal substereid band very small, of 4— 6 cells,

dorsal surface cells larger, thin-walled. Laminal

cells rounded, subquadrate to hexagonal,

ending below apex. Capsules subsessile, cleisto-

smooth; basal cells larger, short-rectangular.

Gemmae produced at stem apex on sterile and

fertile plants, lenticular, to 0,3— 0,6 mm in dia-

meter, cells quadrate to hexagonal or short-rect-

angular.

?Dioicous. Perichaetia terminal; leaves

spathulate to Ungulate, 3, 0-3, 5 mm long, apex

rounded-apiculate; costa ending well below

apex. Seta very short, 0,1 mm long; capsules

immersed, cleistocarpic, subglobose, rostrate,

urn horizontally depressed, 1 ,0 mm high by 1 ,5

mm wide, beak 0,0 mm long, erect to oblique;

exothecial cells lax, quadrate to hexagonal; sto-

mata present at base of urn; calyptra not seen;

spores round, 45-50 /xm, reddish brown,

granulate. Fig. 85: 9-19.

Endemic to southern Africa, O. australis is found on

soil in wooded kloofs and forests of the northern and central

Transvaal, southern Natal, Transkei and the eastern Cape.

Map 111.

Vouchers: Bailey & Jacot Guillarmod 75 — 105h

(COLO); Gordon-Gray 505 1 ; Magill 4977, 5003.

In general habit and colouration O. australis could be

mistaken for species of Hyophila or Weisiopsis; however on

closer examination the large, lens-shaped gemmae and

branching rhizome will separate Oedipodiella. The small,

immersed, horizontally depressed, subglobose capsules

with long beaks are also very distinctive, but are rare and

easily overlooked.

Fig. 85.— Chamaebryum pottioides (1 — 8): 1. habit, x 1; 2. habit, x 10; 3. stem in cross section, x 87; 4. leaves,

X 40; 5. leaf in cross section, X 175; 6. basal leaf cells (left side), x 175; 7. leaf apex showing upper laminal cells, x 175;

8. portion of capsule mouth showing cells and spore, x 175. Oedipodiella australis (9-19): 9. habit, x 1; 10. habit, x

10; 11. stem in cross section, x 87; 12. leaves, x 40; 13. leaf in cross section, x 175; 14. basal leaf cells (left side), x 175;

15. laminal cells at upper left margin, x 175; 16. leaf apex, x 175; 17. gemmae, side view, x 40; 18. gemmae, face view,

X 40; 19. sporophyte, X 5. (1-8, Oliver etal. 77; 9- 19, Sim PRE-CH9678).

305

EPHEMERACEAE

Plants minute, ephemeral, gregarious or scattered, frequently on persistent protonemal mat.

Stems very short. Leaves few, larger above, spreading; linear-lanceolate to oblong-subulate; mar-

gins entire or serrate. Costa present or absent, sometimes weak below and strong above, percurrent

to excurrent; in section cells not differentiated. Laminal cells rectangular to rhomboidal or fu-

siform, smooth or prorate.

Dioicous. Seta very short; capsules cleistocarpic, globose to ellipsoidal, short apiculate; calyp-

tra cucullate, smooth or rough; spores large.

The family contains only two genera, Ephemerum and Micromitrium, only the former occurs in southern Africa. The

genera are separated by the cleistocarpic capsules of Ephemerum and stegocarpic capsules of Micromitrium.

EPHEMERUM

Ephemerum Hampe in Flora, Jena 20: 285 (1837); Broth, in Natiirl. PflFam. 10: 317 (1924); Sim,

Bryo. S. Afr. 289 (1926); Grout, Moss FI. N. Amer. 2: 67 (1935); Smith, Moss FI. Brit. Irel. 346

(1978). Lectotype species: E. serratum (Hedw.) Hampe, fide Grout, Moss FI. N. Amer. 2: 67

(1935).

With characters of family.

The genus consists of c. 28 species. The species are found in most areas of the world, but are most common in

temperate regions. Concentrations of species occur in North America, Europe, North Africa, eastern South America,

southern Africa and Australia.

Both E. sessile and E. serratum are excluded from the Flora. None of the southern African species have spores that

even approach the size given for E. sessile, i.e. 60-80 gm, or most of the Northern Hemisphere species. The southern

African species have weakly ornate spores that range between 25-40 pm in diameter.

The specimen Sim (1926) put into E. serratum is E. capense. Although the two species are similar, the latter has much

smaller leaves that barely reach the mid-point of the capsule, in addition to its smaller spores.

1 Leaves very small, to 0,5 mm long; margins spinose; costa absent 1 . E. capense

1 Leaves larger, 1-2 mm long; margins entire to serrate; costa present, occasionally obsolete:

2 Leaf margins serrate to coarsely serrate; cells in upper leaf prorate; calyptra rough 2. E. namaquense

2 Leaf margins entire to serrulate; laminal cells smooth, calyptra smooth:

3 Leaf margins entire to serrulate, occasionally with a few teeth at apex; perichaetial leaves not distinct, costa

well defined 3. E.rehmannii

3 Leaf margins entire; perichaetial leaves distinct, costa very weak to absent 4. E. diversifolium

1. Ephemerum capense C. Mull, in

Flora, Jena 71: 12 (1888); Broth, in Natiirl.

PflFam. 10: 318 (1924). Type: Cape, Somerset

East, Mt Boschberg, MacOwan s.n., 1882.

Plants minute, acaulescent, scattered on

persistent protonema, light green; terricolous.

Leaves few, 1-5 at base of vaginula, wide-

spreading; linear to elliptical, 0,2-0, 4 mm

long, only a few cells wide; apex acute to acu-

minate; margins plane, spinose. Costa absent.

Laminal cells rectangular to rhomboidal, 2—6:

1, smooth; basal cells rectangular.

Capsules ± sessile, cleistocarpic, orbicu-

lar, 0,3 mm long, reddish yellow, vaginula

globose, 0,1 mm long; exothecial cells quadrate

to short-rectangular; stomata few, present at

base of urn, phaneropore; calyptra cucullate,

0,2 mm long, smooth; spores subround, 37-40

pm, yellowish brown, sharply papillose. Fig.

86:9-16.

Endemic to southern Africa, E. capense is presently

known only from the shrublands of the central and eastern

Cape. Map 113.

Voucher: Sim 7094.

This species was treated by Sim (1926) as E. serratum,

but the plants are considerably smaller than that species in

all respects. The leaves of E. capense are very short and just

reach the mid-point of the capsule. They are rarely visible

when the capsules are mature, giving an impression of

naked capsules scattered on the dense persistent protonema.

The spores are the largest (37-40 gm) of any southern

African species of Ephemerum, but they are still smaller

than the range reported for£. serratum (40-70 /urn).

Ephemeraceae

Ephemeraceae

307

2. Ephemerum namaquense Magill, sp.

nov., E. crassinervio (Schwaegr.) Hampe simi-

lis, sed marginibus foliorum valde serratis, sto-

matibus limitatis ad basim thecarum et sporis

minoribus.

Type: Cape, Namaqualand, along road to

Niewoudtville, 10 km NE of Vanrhynsdorp, on

clay near road, sandy flats with dwarf succulent

shrubs, Magill & Schelpe 3921 (MO, holo.;

PRE; BOL).

Plants minute, scattered or gregarious,

light green to yellow-green; terricolous. Stems

to 0,5 mm long. Leaves larger above, erect-

spreading; lanceolate to linear-lanceolate,

0,7- 1,3 mm long; margins plane, serrulate to

coarsely serrate above. Costa percurrent to just

excurrent, distinct in leaf base; in section

round, 4-5 cells thick, cells undifferentiated,

incrassate, round, surface cells bulging out-

ward. Laminal cells rectangular to rhomboidal,

4-6: 1, prorate above mid-leaf; basal cells rec-

tangular, smooth.

Dioicous. Androgametophyte small,

leaves ovate, margins serrate. Perichaetia ter-

minal, leaves larger, subulate. Seta 0,1 -0,2

mm long, yellowish; vaginula elliptical 0,2 mm

long, reddish yellow; capsule globose to short-

elliptical, short-rostrate, 0,3-0, 5 mm long,

yellowish, beak to 50 pm long; exothecial cells

quadrate to rectangular, or some subhexagonal;

stomata restricted to extreme base, phanero-

pore, reddish; calyptra large, cucullate,

0,2-0, 3 mm long, rough by prorate cells;

spores immature, round, 33 pm, yellowish,

weakly granulate. Fig. 86: 1—8.

Endemic to southern Africa, E. namaquense is found

in small patches on soil among other mosses in shrublands

of the western Cape. Map 113.

Vouchers: Magill & Schelpe 3909; Oliver Ill’S',

Schelpe 4915a.

This species is closely related to E. crassinervium

(Schwaegr.) Hampe through the prorate leaf cells and rough

calyptra but differs in more coarsely serrate leaf margins;

stomata restricted to extreme base of capsule, and smaller,

less ornate spores.

3. Ephemerum rehmannii (C. Miill.)

Broth, in Natiirl. PflFam. 10: 319 (1924); Sim,

Bryo. S. Afr. 291 (1926). Type: Orange Free

State, Bloemfontein, Rehmann s.n., 1875 (G!).

Ephemerella rehmannii C. Miill. in Flora, Jena 71: 12

(1888).

Ephemerella nervosa Dix. in Trans. R. Soc. S. Afr. 18:

254 (1930). Ephemerum nervosa (Dix.) Schelpe in Mem.

bot. Surv. S. Afr. 43: 7 (1979). Type: Cape, King Wil-

liam’s Town, Wager 809 (BM; PRE!).

Plants minute, scattered or in small

groups, light green to yellow-green; terricolous.

Stems 0,5 mm long; in section round, central

strand absent, cortical cells large, thin- walled.

Leaves larger above, weakly secund dry, erect

wet; lanceolate, acuminate to subulate,

1,0-2, 5 mm long; margins weakly incurved,

entire to serrulate or occasionally with a few

teeth at apex. Costa long- to short-excurrent,

awn flexuose, smooth or with a few small serra-

tions, weaker in leaf base; in section round,

3—4 cells thick, cells round, incrassate, not dif-

ferentiated or internal cells more strongly thick-

ened. Laminal cells rectangular to oblong-

rhomboidal or fusiform, to 10: 1; basal cells

rectangular.

Perichaetial leaves longer, shape similar.

Seta very short, yellowish; vaginula cylindrical,

0,2 mm long; capsule ellipsoidal, short apicu-

late, 0,5— 0,8 mm long, yellowish to brownish-

Fig. 86. — Ephemerum namaquense (1-8): 1. habit, x 1; 2. habit (sporophyte immature), x 35; 3. leaves, x 70; 4.

leaf base, x 175; 5. cells at upper leaf, dorsal surface, X 175; 6. laminal cells at upper margin, x 700; 7. sporophyte,

ruptured to show spores, x 70; 8. calyptra, x 70. E. capense (9- 16): 9. habit, x 1; id. habit, x 50; 11-13. leaves, x 70;

14. leaf, x 350; 15. vaginula and lower sporophyte showing stomata, x 350; 16. spores, x 350. E. rehmannii (17-23):

17. habit, x 1; 18. habit, x 12; 19. leaves, x 35; 20. leaf base, x 175; 21. leaf apex, x 175; 22. median leaf cells, x 500;

23. spores, x 350. (1-8, Oliver PRE-CH12902; 9- 16, Sim 7094; 17-23, Rehmann s.n., 1875).

308

Ephemeraceae

yellow; exothecial cells rectangular, thin-

walled; stomata present at base of urn, phanero-

pore; calyptra cucullate, 0, 3-0,4 mm long,

smooth; spores subround, 25—35 jum, reddish

brown, papillose to sharply papillose. Fig. 86:

17-23.

Endemic to southern Africa, E. rehmannii is found in

the dry shrublands or grasslands of Zululand, Orange Free

State and the southern, eastern and western Cape. Map 114.

Vouchers: Schelpe 7625; Sim 7097; Wager 1459.

Most of the southern African specimens referred to E.

sessile belong here. Ephemerum rehmannii can be most

easily separated from E. sessile by its spores only half as

large, leaves frequently toothed above and the excurrent

costa that forms a generally smooth, flexuose awn.

An unusual form of this species collected in Bain’s

Kloof ( Schelpe 6369) is more similar to E. sessile through

its lanceolate-subulate leaves that reach a length of 2,5 mm.

The spores are only 35 gm in diameter however, not reach-

ing the 60-80 /urn reported for£. sessile.

4. Ephemerum diversifolium Mitt, in

Harvey, Tries. Cap. 1: 63 (1859); Sim, Bryo. S.

Afr. 291 (1926). Type: Cape, Uitenhage,

Zwartkop’s River, Zeyher s.n. (NY, holo. !).

Plants minute, scattered on persistent pro-

tonema, light green to yellow-green; temco-

lous. Stems to 0,1 mm long. Leaves larger

FIG. 87. — Ephemerum diversifolium: 1. habit, x 1;

2. habit, X 35; 3. leaf, x 70; 4. leaf in cross section, x

350; 5. leaf base, x 175; 6. leaf apex, x 175; 7. median

leaf cells, x 700; 8. sporophyte with calyptra and perichae-

tial leaf, X 70; 9. portion of capsule base showing stomata

and spores, x 175; 10. spores, x 350. (1 - 10, Harvey s.n.)

Ephemeraceae

309

above, weakly twisted dry, erect-spreading wet;

elliptical, acuminate to subulate, 0,8 — 1,2 mm

long; margins plane, entire. Costa excurrent to

long-excurrent, to 0,4 mm long, distinct in leaf

base; in section bulging dorsally, to 4 cells

thick, internal cells undifferentiated, surface

cells slightly smaller than laminal cells, incras-

sate. Laminal cells oblong-rhomboidal to fu-

siform, 8: 1 , smooth; basal cells rectangular.

Dioicous. Androgametophytes small,

gemmate, 0,25 mm tall. Pericnaetia terminal,

leaves narrow-elliptical to ligulate, 1,3 mm

long; costa very weak or absent in inner leaves;

cells irregularly fusiform. Seta very short, 0,05

mm long, yellowish; vaginula narrow, 0,15 mm

long; capsule ovate-apiculate, 0,7- 1,0 mm

long; exothecial cells rectangular to rhomboi-

dal, but with areas of shorter cells; stomata pre-

sent at extreme base of urn, phaneropore; calyp-

tra cucullate, 0,4 mm long, smooth; spores

subround, 25-30 gm, brownish yellow, granu-

late. Fig. 87.

A very distinct species known only from the type local-

ity in the shrublands of the southern Cape. Map 1 14.

Voucher: Type only.

The small plants are almost buried in the copious mass

of protonema which frequently sends up larger filaments

(stolons of Mitten) that are profusely branched at the tip.

The plants are probably most distinctive because of their

highly differentiated perichaetial leaves. When examined in

Hoyer’s solution, the narrowly elliptical to ligulate leaves

show the costa to be obsolete to absent. The costa in the

vegetative leaves is excurrent, frequently to a length of 0,4

mm and quite distinct throughout the leaf.

Insufficiently Known Species

Ephemerum piliferum Shaw in Cape Monthly Mag. 17:

314 (1878). Type: Cape, Oudeberg, Graaff-Reinet, MacLea

s.n., 1872. As suggested by Sim (1926) the description is

not adequate to place the species and type material has not

been located.

311

FUNARIACEAE

Plants annual or biennial, small to medium-sized, generally light green; terricolous. Stems

erect, simple; central strand generally present. Leaves more or less lax, mostly crowded above,

erect, broad and concave; margins plane to broadly incurved, entire or often serrate, rarely bor-

dered. Costa strong, percurrent to excurrent; in section rounded, bulging dorsally. Laminal cells

large, wide, smooth, rectangular to truncate-rhomboidal or occasionally hexagonal, thin-walled;

marginal cells generally narrower, rarely forming a distinct border; basal cells generally larger,

rectangular.

Monoicous or rarely dioicous. Perichaetia terminal, leaves not strongly differentiated. Cap-

sules immersed or long exserted, stegocarpic or occasionally cleistocarpic, erect to cemuous or

pendent; globose to pyriform, symmetric or curved, frequently with well defined neck; annulus

present, usually well developed, occasionally revoluble; peristome absent, single or double, teeth

16, strongly trabeculate, segments opposite teeth, cilia absent; operculum plano-convex to conic,

often apiculate; calyptra smooth, generally cucullate-inflated and rostrate or smaller, mitrate;

spores small, granulate.

The family Funariaceae consists of 16 genera, some large and very widespread in distribution ( Funaria , Physcomi-

trium), but most very small and quite restricted. The six genera present in southern Africa exhibit the full range of capsule

morphology expressed by the family; i.e. cleistocarpic or stegocarpic and gymnostomous to peristomate. The peristomate

capsules also exhibit a complete gradation in peristome development, from rudimentary to well developed, single or double.

The broad, concave leaves with large, lax leaf cells, generally with flattened ends, should place most sterile speci-

mens. The capsules are very distinctive and will easily identify the family and genera. Some southern African taxa of

Bryaceae bear a slight macroscopic resemblance to Funaria. The former are easily separated by peristome morphology, and

their upper laminal cells being evenly rhomboidal with pointed ends.

1 Plants small, stems very short; capsules immersed to emergent:

2 Capsule stegocarpic, mouth broad:

3 Upper leaves broadly obovate, margins entire; calyptra large, 8-plicate

1 . Goniomitrium

3 Upper leaves narrowly spathulate, margins strongly serrate; calyptra small, smooth:

4 Capsules immersed, small, hemispherical, 0,5-0, 6 mm long; spores papillose;

growing on alluvial mud and river banks in Zululand 2. Micropoma

4 Capsules immersed to emergent, cupulate, 1,0-1, 5 mm long; spores spinose;

growing on soil in semi-aria shrublands of the eastern Cape 5. Physcomitrium

2 Capsule cleistocarpic; calyptra minute, smooth:

5 Plants bulbiform; perichaetial leaves large, strongly concave, completely enclosing

sporophyte; capsule pyriform, pendent 3. Cygnicollum

5 Plants with erect to spreading leaves; perichaetial leaves spreading, exposing sporo-

phyte; capsule elliptical, erect 4. Physcomitrellopsis

1 Plants larger, stems elongated; capsule exserted on long seta:

6 Capsules cupulate, neck very short; exothecial cells quadrate to hexagonal; leaves with

distinct border of narrower, thin- walled cells 5. Physcomitrium

6 Capsules cylindrical to pyriform, neck long; exothecial cells rectangular; leaves without

distinct border or if present marginal cells incrassate 6. Funaria

1. GONIOMITRIUM

Goniomitrium Wils. in Hooker, J. Bot., Lond. 5: 142 (1846); Broth, in Natiirl. PflFam. 10: 324

(1924); Sim, Bryo. S. Afr. 292 (1926); Stone in J. Bryol. 11: 491 (1981). Type species: not

designated.

Rehmanniella C. Mull . in Bot. Zbl. 7: 347 (1881). Type species: R. africana C. Mull.

312

FUNARIACEAE

Funariaceae

313

Plants small, gregarious; terricolous. Stems short, simple; central strand present. Leaves

spreading wet, obovate-cuspidate; margins plane, entire. Costa short-excurrent. Laminal cells

hexagonal to rhomboidal, thin- walled.

Paroicous or dioicous. Seta short; capsule emergent, stegocarpic, gymnostomous, cupulate

with broad mouth; operculum plano-convex, apiculate; calyptra mitrate, 8-plicate; spores large.

The four species of Goniomitrium are equally divided between Africa and Australia. Each of the species is restricted in

its distribution and distinctive morphologically.

Goniomitrium africanum (C. Mull.)

Broth, in Natiirl PflFam. 1: 521 (1903); Sim,

Bryo. S. Afr. 292 (1926). Type: Orange Free

State, near Bloemfontein, Rehmann 171 (NH!).

Rehmanniella africana C. Mull, in Bot. Zbl. 7: 347

(1881).

Plants small, scattered or gregarious, dark

green; terricolous. Stems 1-2 mm high, simple,

radiculose below; in section round, central

strand weak, cells thin-walled, inner cortical

cells in 2-4 rows, outer cortical cells in 2 rows,

smaller, reddish brown. Leaves crowded and

larger above, infolded and appressed dry, erect-

spreading wet; broadly elliptical to obovate or

subspathulate, 1,5-2, 5 mm long; apex apiculate

to cuspidate; base oblong; margins plane, en-

tire. Costa short-excurrent; in section subround,

lamina attached ventrally, guide cells 2, ex-

posed ventrally, dorsal stereid band weak, 2

cells thick, occasionally substereids, dorsal sur-

face cells generally strongly thickened, smaller

than guide cells. Upper laminal cells hexagonal

to rhomboidal or subrectangular, thin-walled;

marginally quadrate to rectangular; basal cells

quadrate.

Paroicous. Perichaetial leaves slightly

larger, not distinct. Seta variable in length,

0, 5-2,0 mm long; capsule emergent to ex-

serted, erect, stegocarpic, cupulate, with broad

mouth, 1 mm long, irregularly sulcate when

dry, reddish yellow; exothecial cells quadrate to

rectangular or angular, thin-walled, with 1-2

rows of quadrate cells at mouth and 5-6 rows of

transversely rectangular cells just below; sto-

mata present at base of urn, phaneropore; peri-

stome absent; operculum plano-convex,

0,7-0, 8 mm in diameter, cells not twisted; ca-

lyptra mitrate, 1,5 mm long, smooth but 8-

sided because of longitudinal plications; spores

Map 115. — • Goniomitrium africanum

▲ Micropoma niloticum

rounded, 50-75 gm, yellowish, granulate with

a weak reticulate pattern. Fig. 88: 1-10.

Endemic to southern Africa, G. africanum occurs on

open soil, in dry rocky regions of southern South West

Africa/Namibia, the western, northern and central Cape

Province, and infrequently in the Orange Free State, Leso-

tho and central Transvaal. The species forms large patches

after the winter rains, but is only infrequently collected.

Map 115.

Vouchers: Magill & Schelpe 3834a, 4038; Oliver

6299, 7255; Smook & Harding 695; Volk 5298.

When sporophytes are present the large-mouthed cap-

sule and spreading, dark green, cuspidate leaves will place

the species. The large, unusual, 8-sided, plicate calyptra is

also very distinctive.

The seta length in different specimens is quite variable.

A recent study indicated that the variation was not signifi-

cant and found no supporting characters on the specimens

with long setae (Schelpe, pers. comm.).

In his recent generic revision of Funariaceae, Fife

(1985) reduced G. africanum to a subspecies of G. acumi-

natum Hook. & Wils., an Australian species.

FlG. 88. — Goniomitrium africanum ( 1- 10): 1. habit, X 1;2. habit, x 10; 3. plants, x 2; 4. stem in cross section, x

200; 5. leaves, x 40; 6. leaf in cross section, x 175; 7. basal leaf cells (right side), x 175; 8. leaf apex, x 175; 9.

sporophyte, x 10; 10. calyptra, x 10. Micropoma niloticum ( 1 1-19): 11. habit, x 1; 12. habit, x 10; 13. stem in cross

section, x 200; 14. leaves, x 40; 15. leaf in cross section, x 350; 16. basal leaf cells (left side), x 175; 17. leaf apex, x

175; 18. sporophyte, operculum detached, x 20; 19. spores, x 560. ( 1 - 10, Oliver 6299; 1 1 - 19, Junod 323).

314

Funariaceae

2. MICROPOMA

Micropoma Lindb. in Notis. Saellsk. F. FI. Fenn. Foerh. 1 1: 56 (1871); Broth, in Natiirl. PflFam.

10: 322 (1924); Sim, Bryo. S. Afr. 291 (1926). Type species: M. niloticum (Delile) Lindb.

Plants small, gregarious; terricolous. Stems short, simple; central strand present. Leaves

narrow, spathulate; margins serrate. Costa percurrent to subpercurrent. Laminal cells lax, rhom-

boidal to rectangular.

Autoicous. Seta short; capsule immersed, stegocarpic, hemispherical with very broad mouth,

gymnostomous; operculum convex-apiculate; calyptra mitrate, smooth; spores large.

The genus contains two species, M. niloticum from southern and eastern Africa as well as the Middle East, and M.

bukobense from central Africa. Both species were recently placed in Physcomitrium subgen. Cryptopyxis by Fife (1982).

Micropoma niloticum (Delile) Lindb. in

Broth, in Natiirl. PflFam. 1: 518 (1903); 10:

322 (1924); Sim, Bryo. S. Afr. 292 (1926); El-

Saadawy in Proc. Egypt. Acad. Sci. 25: 217

(1972). Type: Egypt, Delile s.n.

Gymnostomum niloticum Delile, FI. Aegypt. 2: 43

(1813). Physcomitrium niloticum (Delile) C. Mull, in Bot.

Ztg 16: 154(1858).

Plants small, scattered, gregarious, light

green, ± shiny; terricolous. Stems (0,5 — )2 — 3

mm long, occasionally elongated with a few

branches; in section round, central strand col-

lapsed, dark brown, cortical cells in 3-4 rows,

little differentiated or outer row ± larger.

Leaves weakly contorted dry, spreading wet;

narrowly spathulate, 1 — 2( — 3) mm long, apex

abruptly acuminate; base oblong; margins

plane, strongly serrate above base by projecting

cell ends. Costa weak, ending below apex or

extending into acumen; in section bulging dor-

sally, guide cells 2, ventral cells 2, in single

layer, similar to guide cells, dorsal stereid band

small, consisting of 2-4 cells, dorsal surface

cells smaller than guide cells, thickened. Lami-

nal cells lax, rectangular to rhomboidal, mostly

2-3: 1; basal cells rectangular, thin-walled.

Perichaetial leaves slightly larger, not dis-

tinct. Seta erect, 0,4-0, 6 mm long, yellowish

to yellow-brown; capsule immersed, urn hemis-

herical, 0,5 -0,6 mm long, smooth, yellow-

rown, neck absent; exothecial cells lax,

transversely rectangular to rhomboidal, 4 rows

at mouth abruptly narrower and transversely

rectangular; stomata present at extreme base of

urn, phaneropore; annulus absent; peristome

absent; operculum convex-apiculate, cells not

twisted; spores subround to angular, 30-37

/xm, light brown, papillose to strongly papil-

lose. Fig. 88: 11-19.

The species grows on alluvial mud and river banks in

the Gaza Strip, Egypt, Zimbabwe, Mozambique and South

Africa. Although M. niloticum has been known from

Maputo for some time (Sim, 1926), it has only recently

been discovered in Zululand, mixed with Ephemerum

rehmannii. Map 115.

Voucher: Taylor 478.

The species is most easily recognized by its narrow-

spathulate leaves and very broad-mouthed, immersed,

hemispherical capsules. Sterile stems are frequently

elongated with distant leaves.

3. CYGNICOLLUM

Cygnicollum Fife & Magill in Bryologist 85: 99 (1982). Type species: C. immersum Fife &

Magill.

Plants very small, bulbiform, light green; terricolous. Stems simple, central strand present.

Leaves obovate-acuminate, concave; margins crenulate above. Costa subpercurrent to short-excur-

rent. Upper laminal cells rhomboidal to oblong-rhomboidal, thin-walled; marginal cells narrower.

Perichaetia terminal, leaves large, conspicuous, strongly concave, enclosing sporophyte. Seta

short, erect; capsule cleistocarpic, pyriform, pendent, neck cygneous; calyptra small, campanulate;

spores irregularly papillose.

A very distinct genus recognized by the bulbiform habit and completely enclosed, pendent, cleistocarpic capsules. The

taxon seems, through most of its characters, properly placed in Funariaceae. However, it does not appear to be closely

related to any of the other genera of the family.

Funariaceae

315

Cygnicollum immersum Fife & Magill in

Bryologist 85: 99 (1982). Type: Cape, upper

slopes of Vanrhyns Pass, 40 km NE of Van-

rhynsdorp along road to Nieuwoudtville, Magill

& Schelpe 3952 (PRE, holo.; BUF; MICH;

MO).

Plants minute to small, bulbiform, soli-

tary, light green; terricolous. Stems to 1 mm

high, simple; in section round, central strand

small, inner cortical cells in 2 rows, large, thin-

walled, outer cortical cells in single row, small,

incrassate, reddish brown, epidermal cells

larger, thin-walled. Leaves larger above, erect-

spreading wet, little altered dry, lower leaves

obovate-apiculate, to 1,5 mm long; upper

leaves oblong-obovate, acuminate, 1,5 — 3,0

mm long; margins plane, entire below, bluntly

serrate above. Costa ending below apex in

lower leaves to short-excurrent in upper leaves;

in section rounded, with a large central stereid

band 2-4 cells thick and enclosing 1 -2 larger,

irregular cells or gaps, ventral cells in single

row, large, firm- walled, dorsal surface cells

similar to ventral cells, thickened. Upper lami-

nal cells rhomboidal to oblong-rhomboidal,

thin-walled; upper marginal cells narrower,

long-rectangular, but not forming distinct

border, upper ends projecting as marginal teeth;

basal cells lax, rectangular.

Autoicous. Perigonial branch short, at

base of main stem, gemmate. Perichaetia termi-

nal, leaves large, conspicuous, strongly con-

cave, completely enclosing capsule. Seta short,

erect, 0,7- 1,0 mm long , yellowish; capsule

broadly pyriform, cleistocarpic, pendent

because of cygneous neck, 1,0 — 1,3 mm long,

yellowish brown; exothecial cells quadrate to

rectangular or occasionally rhomboidal, thin-

walled; stomata present on neck, phaneropore;

calyptra small, narrowly campanulate, 0,6 mm

long, smooth; spores subreniform, 25 -35 /xm,

yellowish, irregularly papillose. Fig. 89.

Known only from soil in small open areas in shrub-

lands of the upper Vanrhyn’s Pass. Tne specimens were

collected on the south face of the escarpment at c. 800 m.

Fig. 89. — Cygnicollum immersum: 1. habit, x 1;2.

habit, x 10; 3. leaf, x 35; 4. leaf in cross section, x 350;

5. basal leaf cells (right side), x 175; 6. laminal cells at

upper left margin, x 350; 7. apical leaf cells, x 175; 8.

sporophyte, x 35; 9. calyptra, x 35; 10. stomatal appara-

tus, x 700; 11. spore, x 500. (1-11, Magill & Schelpe

3952).

316

Funariaceae

Funariaceae

317

The collection site in the north western Cape receives a high

rainfall for this region. Map 1 16.

Voucher: Magill & Schelpe 3937.

The completely enclosed sporophyte is very unique.

The seta is snort and erect but the neck of the pyriform

capsule is cygneous resulting in a pendent urn.

The cleistocarpic capsules and large, papillose spores

are known in Funariaceae, however the calyptra of Cygni-

collum is rather small for the family. Leaf shape and areola-

tion also confirm placement in Funariaceae.

Map 116. — •Cvgnicollum immersum

▲ Phy scomit rellopsis africana

4. PHYSCOMITRELLOPSIS

Physcomitrellopsis Broth. & Wager ex Dix. in J. Bot., Lond. 60: 107 (1922); Broth, in Natiirl.

PflFam. 10: 321 (1924); Sim, Bryo. S. Afr. 291 (1926). Type species: P. africana Wager & Broth.

Plants small, gregarious, green; terricolous. Stems erect, unbranched; central strand present.

Leaves spreading wet; spathulate-acuminate; margins plane, entire. Costa ending below apex.

Laminal cells rectangular, becoming rhomboidal above, smooth.

Perichaetia terminal, leaves larger, otherwise little differentiated. Capsule exposed, subses-

sile, erect, cleistocarpic; calyptra large, enclosing capsule, mitrate-rostrate; spores weakly papil-

lose.

Endemic to southern Africa, Physcomitrellopsis presently contains a single species found in the southeastern part of the

Flora area. The only other species described in the genus, Physcomitrellopsis indica Dix., was recently transferred to

Physcomitrium by Gangulee (1974).

Physcomitrellopsis africana Wager &

Broth, ex Dix. in J. Bot., Lond. 60: 107 (1922);

Sim, Bryo. S. Afr. 291 (1926). Type: Natal,

Wager s.n. (H, holo.; BM).

Plants small, gregarious, green; terrico-

lous. Stems 1,0— 1,5 mm high, simple; in sec-

tion round, central strand small, inner cortical

cells in 2-4 rows, thin-walled, outer cortical

cells in 2-3 rows, slightly thickened. Leaves

crowded above, weakly contorted dry, spread-

ing wet; spathulate-acuminate, 4-5 mm long;

base oblong; margins plane, entire below.

strongly dentate above base. Costa subpercur-

rent; in section subrounded, guide cells 2—3,

small, ventral cells in single row, similar to

guide cells, dorsal stereid band small, group of

6-8 cells, frequently with gap below guide

cells, dorsal surface cells ± larger, thin- walled.

Laminal cells rectangular to oblong-rhomboi-

dal; upper marginal cells bulging; basal cells

rectangular.

Autoicous. Perichaetia terminal, leaves

large, spathulate-acuminate, widespreading

Fig. 90. — Physcomitrellopsis africana (1 — 10): 1. habit, x 1; 2. habit, x 10; 3. portion of stem in cross section, x

140; 4. leaf, x 35; 5. leaf in cross section, x 350; 6. basal leaf cells at right margin, x 175; 7. median leaf cells, x 175; 8.

laminal cells at upper right margin, x 175,9. leaf apex, x 175; 10. spore, x 500. Physcomitrium spathulatum (1 1-21):

11. habit, x 1; 12. habit (var. spathulatum), x 10; 13. habit (var. sessile), x 10; 14. stem in cross section, x 140; 15.

leaf, X 35; 16. leaf in cross section, x 350; 17. basal leaf cells at right margin, x 175; 18. laminal cells at right margin, x

350; 19. leaf apex, X 175; 20. part of capsule mouth showing cells, x 245; 21. spore, x 500. ( 1 — 10, Nicholson 126B;

11 — 1 2 & 14-21, Edwards 827; 13, MacLea sub Rehmann 520).

318

Funariaceae

wet. Seta 0,5 -0,7 mm long, yellowish; capsule

exposed, erect, cleistocarpic, globose to ellipti-

cal, 1,0- 1,2 mm long, smooth, reddish yel-

low; exothecial cells angular, subquadrate to

hexagonal; stomata present at base of urn,

phaneropore; calyptra swollen, completely en-

closing capsule, mitrate-rostrate, 1,3 mm long;

spores immature, rounded, 30 p m, hyaline,

weakly papillose with distinct tetrad scar. Fig.

90: 1-10.

Endemic to southern Africa, P. africana has been col-

lected twice in fairly large colonies at the edge of coastal

forests in Natal and Transkei. The exact type locality was

not given and the site has not been precisely located. Map.

1 16.

Voucher: Nicholas 126B.

The lax, spathulate-acuminate leaves with dentate mar-

gins are fairly distinct and should help to place sterile speci-

mens of P. africana. The short seta and exposed, globose to

elliptical, cleistocarpic capsule and large calyptra are very

distinctive; however also very small and easily overlooked

on dried specimens.

5. PHY SCOMITRIUM

Physcomitrium (Brid.) Fuernr. in Flora, Jena 13: 9 (1829); Broth, in Naturl. PflFam. 10: 322

(1924); Sim, Bryo. S. Afr. 292 (1926). Type species: P. sphaericum (Ludw.) Fuernr., vide Grout,

MossFl.N. Amer. 2:74(1935).

Gymnostomum subgen. Physcomitrium Brid., Bryol. Univ. 1: 97 (1826).

Plants mostly small, gregarious to loosely caespitose; terricolous. Stems unbranched; central

strand present. Leaves crowded above, obovate-acuminate; margins bordered, serrate above. Costa

subpercurrent to percurrent. Laminal cells rectangular to rhomboidal, smooth; narrower at margin.

Autoicous. Seta short or long; capsule cupulate, gymnostomous, mouth very broad; exothe-

cial cells subquadrate to hexagonal; operculum plano-convex; calyptra mitrate, smooth; spores

spinose.

Only a single species of Physcomitrium is known from southern Africa. The widespread genus contains approximately

83 species found on every continent except Antarctica.

The genus is most easily recognized by its cup-shaped, broad-mouthed capsules with very short necks. The exothecial

cells are shorter (quadrate to hexagonal) than those found in Funaria (rectangular to rhomboidal) and the peristome is

absent.

Physcomitrium spathulatum (Hornsch.)

C. Miill. in Linnaea 18: 695 (1845); Sim, Bryo.

S. Afr. 293 (1926). Type: Cape, between Kei

and Bashee Rivers, Drege s.n., 1832.

Gymnostomum spathulatum Hornsch. in Linnaea 15:

115(1841).

Physcomitrium spathulatum var. brevicollum C. Miill.,

Syn. Muse. 1: 118 (1849). Syntype: Cape, Pappe s.n.;

Philippstown, Ecklon s.n.

Physcomitrium brachypodium C. Mull, in Hedwigia 38:

59 (1899); Broth, in Naturl. PflFam. 10: 324 (1924). Type:

Cape, Groot Visch River, MacOwan s.n., June 1877.

Plants small to medium-sized, gregarious,

light green; terricolous. Stems 2-6 mm tall,

simple; in section round, central strand a small

group of thin-walled cells, inner cortical cells in

2—4 rows, large, thin- walled, outer cortical

cells in 2 rows, smaller, weakly thickened.

Leaves crowded above to rosulate, contorted

dry, erect-spreading wet, lamina weakly undu-

late; obovate to subspathulate or occasionally

elliptical, 2, 5-4, 5 mm long; apex acute to acu-

minate; margins weakly bordered, plane, ser-

rate above. Costa ending below apex to just

excurrent; in section subround, guide cells 2,

small, ventral cells 2, in single row, larger than

uide cells, weakly thickened, dorsal stereid

and weak, 2-4 cells, frequently with gap be-

low guide cells, dorsal surface cells similar to

ventral cells. Upper laminal cells rectangular to

oblong-hexagonal; marginal cells narrower,

long-rectangular to linear, forming distinct

border, thin-walled, upper ends of cells project-

ing as marginal serrations; basal cells rectangu-

lar.

Autoicous. Perichaetia terminal, leaves

not distinct. Seta very short, 0,3 -0,4 mm long

or elongated (3— )5— 8 mm long, reddish yel-

low; capsules, erect, symmetrical, stegocarpic,

cupulate or rarely very short-cylindrical,

1 ,0- 1 ,5 mm long, yellowish red, mouth broad,

neck very small, sulcate when dry; exothecial

cells subquadrate to hexagonal, thickened, but

thin-walled in base, with 4—6 rows of transver-

sely rectangular cells at mouth; stomata present

on neck, phaneropore; peristome absent; oper-

culum plano-convex, abruptly rostrate; calyptra

Funariaceae

319

Map 117. — • Physcomitrium spathulatum var. spathu-

latum

A Physcomitrium spathulatum var. sessile

mitrate, 1,5 mm long, smooth; spores round,

22-30(-35) /im, reddish brown, spinose, with

spines up to 3 /urn long. Fig. 90: 11-21.

1 Seta 3—8 mm long var. spathulatum

1 Seta 0,3-0, 4 mm long var. sessile

var. spathulatum.

Seta 3—8 mm long; capsule exserted;

spores round, 22—30 fj.m, spinose.

Reported from southern Africa, Zimbabwe and Mo-

zambique; in the Flora area the variety is infrequently col-

lected in moist situations in South West Africa/Namibia, the

central, eastern and southern Cape, Transkei, Natal and the

central and southern Transvaal. Map 117.

Vouchers: Edwards 827; Lambert 17a; Volk 2226;

Wells 54.

The short, erect, broad-mouthed, gymnostomous cap-

sule exserted on a long seta should separate this variety

from other taxa in Funariaceae. In addition the mitrate ca-

lyptra and bordered leaves are useful in identifying the va-

riety.

var. sessile (Shaw) Magill, stat. nov. Lec-

totype: Cape, near Graaff-Reinet, MacLea, sub

Rehmann 520 (PRE!; selected here; BM; BOL;

NH).

Physcomitrium sessile Shaw in Cape Monthly Mag. 17:

316(1878).

Seta 0,3 -0,4 mm long; capsules im-

mersed to emergent; spores round, 30 - 35 jam,

coarsely spinose.

Endemic to southern Africa, P. spathulatum var. ses-

sile is only rarely collected in the semi-arid grasslands of the

eastern parts of the central Cape Province. Map 117.

Voucher: Shaw s.n., 1867.

This variety is similar in all vegetative respects and

most sporophyte characters to the typical variety. The dif-

ferences are the very short seta that results in immersed to

emergent capsules, and the somewhat larger and more

coarsely spinose spores.

Shaw (1878) described the costa as excurrent. While

this is correct for some of the upper or perichaetial leaves,

which have costae very short-excurrent, the costa ends well

below the apex in most of the lower leaves.

This variety could be confused with Physcomitrellop-

sis africana ; however its capsules are much smaller and

cleistocarpic and its leaves are strongly toothed.

6. FUN ARIA

Funaria Hedw., Sp. Muse. 172 (1801); Broth, in Natiirl. PflFam. 10: 325 (1924); Scott & Stone,

Moss. S. Aust. 252 (1976); Smith, Moss FI. Brit. Irel. 338 (1978). Lectotype species: F.

hygrometrica Hedw., vide Britt. inN.L. Britt., FI. Bermuda 441 (1916).

Entosthodon Schwaegr., Sp. Muse. Suppl. 2: 44 (1823); Sim, Bryo. S. Afr. 293 (1926); Grout, Moss FI. N. Amer. 2: 78

(1935). Type species: E. templetonii (J.E. Sm.) Schwaegr.

Plants small to medium-sized, gregarious or loosely caespitose; terricolous. Stems erect,

simple; in section round, central strand weak, inner cortical cells in 2-4 rows, large, thin-walled,

outer cortical cells in 1-2 rows, smaller, incrassate, reddish yellow, epidermal cells large, thin-

walled, frequently collapsed. Leaves larger above, in a rosette, weakly contorted dry, patent to

widespreading wet; elliptical to obovate or subspathulate; apex acute to acuminate, occasionally

apiculate to cuspidate; base mostly oblong; margins plane or occasionally broadly incurved, entire

to crenulate or serrate, infrequently bordered. Costa ending below apex to percurrent, occasionally

excurrent. Laminal cells rectangular, lax, frequently rhomboidal near apex; marginal cells gener-

ally narrower, infrequently thickened.

Autoicous. Perichaetia terminal; leaves weakly differentiated. Seta elongate; capsule stego-

carpic, erect to pendent, symmetrical to asymmetrical, mostly pyriform, occasionally subcylindri-

cal, neck generally well developed, sulcate; exothecial cells rectangular; stomata present on neck;

320

Funariaceae

peristome single, double or absent; operculum plano-convex, apiculate; calyptra inflated-cucullate,

rostrate, beak erect to oblique; spores papillose to granulate.

A large genus of approximately 220 species, Funaria is found on every continent except Antarctica. In southern Africa

the genus is not common, but specimens are generally found in moist open areas in the eastern and southern parts of the

Flora area. Perhaps the most widely recognized moss, F. hygrometrica, is practically cosmopolitan in its distribution and

occasionally even considered a ‘weed’ in commercial greenhouses.

In his treatment of Funariaceae, Sim (1926) recognized nine species under Entosthodon, based on peristome devel-

opment. Although this approach has been followed by many authors, the separation of Entosthodon and Funaria, even in

southern Africa, is not altogether clear. Since an apparent gradation in peristome reduction is evident, I prefer to follow

Brotherus (1924) in uniting all of the species under Funaria. In southern Africa the genus is most closely related to

Physcomitrium; for differences see notes under that genus.

The southern African species of Funaria can be divided into two subgenera. The subgenus Funaria is identified by a

double peristome, with the endostome well developed to very rudimentary, and the cells of the operculum frequently

twisted counterclockwise when viewed from above. Three of the species found in the Flora area are included in this

subgenus, i.e. F. hygrometrica, F. rhomboidea and F. spathulata. The remaining species fall within the subgenus Entos-

thodon (Schwaegr. j Lindb. This subgenus is recognized by its capsules being gymnostomous or having a single peristome

which may be well developed or rudimentary . The cells of the operculum are not twisted or only very slightly so.

Since the peristomes of many of the species in the latter subgenus are fragile and lost soon after the capsule opens, their

presence or absence is not used as a major character in the key. Only three of the southern African species appear to be truly

gymnostomous. They are F . clavata, F. longicollis and F . urceolata.

1 Costa percurTent to excurrent in upper leaves:

2 Leaves apiculate; capsules erect to slightly inclined:

3 Leaves obovate to subspathulate; margins crenulate with a few strong teeth near apex 5. F. longicollis

3 Leaves elliptical or occasionally obovate; margins entire 4. F. rottleri

2 Leaves short-acuminate; capsules inclined to horizontal or pendent:

4 Leaves narrowly obovate; capsules inclined, symmetrical; peristome absent 6. F. urceolata

4 Leaves broadly elliptical to obovate; capsules inclined to pendent, asymmetrical; peristome double

10. F. hygrometrica

1 Costa ending below apex in upper leaves:

5 Leaf margins serrulate to crenulate or occasionally dentate:

6 Capsule asymmetrical, arcuate, if erect and straight then mouth obilque 9. F. spathulata

6 Capsules erect, symmetrical, pyriform or cylindrical above well defined neck:

7 Leaves bordered by 1-2 rows of narrow, incrassate cells; margins serrulate to serrate 1. F. limbata

7 Marginal cells bulging, slightly larger than laminal cells; margins crenulate, frequently strongly so

8. F. rhomboidea

5 Leaf margins entire:

8 Leaves bordered by 1-2 rows of narrow, incrassate cells 1 . F. limbata

8 Leaves not bordered; marginal cells occasionally narrower than laminal cells:

9 Leaf apices acute to obtuse; leaves broadly elliptical to obovate or occasionally ovate 2. F. succuleata

9 Leaf apices mucronate to apiculate or piliferous; leaves elliptical to obovate:

10 Capsules erect, pyriform, 1,2-1, 5 mm long; peristome present, fragile 3. F. bergiana

10 Capsules horizontal, clavate, to 4 mm long; peristome absent 7. F. clavata

1. Funaria limbata (C. Mull.) Broth, in

Natiirl. PfIFam. 1: 524 (1903); 10: 328 (1924).

Type: Cape, Zitskamma, Breutel s.n. (BM!).

Entosthodon limbatus C. Mull, in Bot. Ztg 16’ 155

(1858). 1 * * * 5 6 * * 9 10

Entosthodon marginatus C. Miill., Syn. Muse. 1: 125

(1848), Sim, Bryo. S. Afr. 297 (1926). Funaria marginata

(C. Miill.) Broth., horn, illeg. , in Natiirl. PfIFam. 1: 525

(1903), non Kindberg (1883). Type: Cape, Swellendam,

Ecklon s.n.

Entosthodon ampliretis C. Mull, in Hedwigia 38: 60

(1899). Funaria ampliretis (C. Mull.) Broth, in Natiirl.

PfIFam. 1: 525 (1903); 10: 329 (1924). Type: Natal, Um-

geni above Pietermaritzburg, Rehmann 174 (G !).

Entosthodon gracilescens C. Miill., horn, illeg., in Hed-

wigia 38: 59 (1899), non Schwaegr. ex C. Mull. (1858).

Type: Transvaal, near Lydenburg, Wilms s.n., Feb. 1883

(G, holo. !).

Entosthodon micropyxis C. Miill. in Hedwigia 38: 60

(1899); Funaria micropyxis (C. Miill.) Broth, in Natiirl.

Funariaceae

321

PflFam. 1: 524 (1903); 10: 328 (1924). Type: Cape, near

Blanco, Rehmann 175 (BM!; G!).

Physcomitrium leptolimbatum C. Mull, in Hedwigia 38:

59 (1899); Broth, in Natiirl. PflFam. 1: 520 (1903). Type:

Transvaal, near Lydenburg, Wilms s.n., Feb. 1888 (G,

holo.!).

Entosthodon marginatus var. obtusatus Sim, Bryo. S.

Afr. 297 (1926). Funaria ampliretis var. obtusata (Sim)

Wijk & Marg. in Taxon 9: 189 (1960). Type: Natal, Um-

hwati, New Hanover, Sim 8585 (PRE, holo. f).

Plants small, scattered, light green to yel-

low-green; terricolous. Stems 1—2 mm long.

Leaves larger and crowded above, contorted

dry, widespreading wet; elliptical to obovate,

2-3(-^) mm long; apex acute to obtuse or oc-

casionally rounded; base not differentiated;

margins plane, entire to weakly serrate above,

bordered by 1-2 rows of elongated, incrassate

cells. Costa ending just below the apex; in sec-

tion round, guide cells 2, small, thin-walled,

ventral cells in single row, 2, small, thin-

walled, dorsal stereid band small, 1-2 cells

thick, reddish yellow, dorsal surface cells in-

crassate. Laminal cells short, oblong-hexagonal

to rectangular or occasionally quadrate, thin-

walled; marginal cells narrower, elongate, in-

crassate; basal cells rectangular, thin- walled.

Seta 6-11 mm long, reddish yellow; cap-

sule erect, elliptical to short-cylindrical, 1-2

mm long, neck differentiated, urn frequently

constricted under mouth when dry; exothecial

cells rectangular, thickened, becoming shorter

to quadrate in neck, with 4-8 rows of transver-

sely rectangular to quadrate cells at mouth,

thickened; stomata on neck, phaneropore;

peristome single, irregular, teeth ± oblique,

triangular, 175 p.m high, irregularly perforated

and absent with age, striate below, granulate

above, reddish yellow; operculum plano-con-

vex, short-apiculate, cells weakly twisted,

counter clockwise; spores subround, 20—25

/im, light brown, vermiculate. Fig. 91: 1-13.

The species is known from eastern and southern Af-

rica. In the Flora area the species is infrequently collected in

the northern, eastern, central and southern Transvaal, Na-

tal, Transkei and the southern and southwestern Cape. Map

118.

Vouchers: Cholnoky 51, 417a; Crosby & Crosby 8091;

Gordon-Gray PRE-CH12820; Oliver 7308a; Rankin 54;

Russell 2562.

The species is distinguished from the other southern

African species of Funaria by its distinct border of narrow,

elongate and thickened marginal cells. Although the border

is occasionally weak, the cells are always more strongly

differentiated than in the other species. Sterile specimens

could be confused with Physcomitrium which also has bor-

dered leaves. The marginal cells of P. spathulatum, how-

ever, are not thickened.

Several specimens have been seen with broadly ellipti-

cal leaves and rounded to obtuse apices, but agree in other

respects to F. limbata. Although these specimens have been

recognized as a variety [F. ampliretis var. obtusata (Sim)

Wijk & Marg.] they are here considered to be just within the

normal variation of this taxon.

2. Funaria succuleata (Wager & Wright)

Magill, comb. nov. Type: Natal, Rydal Mount,

Wager s.n. PRE-CH12075 (PRE, holo. !; BM).

Physcomitrium succuleatum Wager & Wright in Trans.

R. Soc. S. Afr. 4: 3 (1914); Broth, in Natiirl. PflFam. 10:

324(1924).

Entosthodon cavifolius Mitt, in Harv., Thes. Cap. 1: 64

(1859); non Funaria cavifolia Card. & Broth. (1923).

Funaria harveyana Magill in Mem. bot. Surv. S. Afr. 43: 7

(1979). Type: Cape, near Cape Town, Harvey s.n. (NY!).

Plants small, gregarious, light green; terri-

colous. Stems 2—4 mm long. Leaves crowded

above, appressed, little contorted dry, erect-

spreading wet; broadly elliptical to obovate or

occasionally ovate, 1, 5-2,0 mm long, con-

cave; apex acute to obtuse; base scarcely dif-

ferentiated, margins plane to broadly incurved

above, entire. Costa ending below apex; in sec-

tion guide cells 2, exposed ventrally, dorsal

stereid band small, 2 cells thick, dorsal surface

cells similar in size to guide cells, weakly

thickened. Laminal cells rectangular, thin-

walled, rhomboidal or occasionally rhombic

near apex; marginal cells narrower and bulging

but not forming distinct border; basal cells rec-

tangular.

322

Funariaceae

Funariaceae

323

Map 119. — • Funaria succuleata

A Funaria bergiana

Seta to 4 mm long, reddish yellow; capsule

erect, pyriform, 1 ,2- 1 ,5 mm long, urn subglo-

bose, neck very short; exothecial cells rectangu-

lar, thickened, with 2-3 rows of quadrate cells

at mouth; peristome single, rudimentary and

often missing on deoperculate capsules, teeth

oblong, irregular, striate; operculum plano-con-

vex, cells not twisted; spores round, 25 pm,

reddish brown, weakly papillose. Fig. 91:

14-20.

Endemic to southern Africa, F. succuleata is found on

soil of the southwestern Cape, Lesotho, Orange Free State,

Natal and eastern Transvaal. Map 1 19.

Vouchers: Magill 4658; Tolken 5739a.

The species is recognized by its broadly elliptical to

obovate leaves; entire, unbordered margins and costa that

ends well below the apex. The leaves are generally concave

and occasionally have broadly incuved margins that may

obscure the typical leaf shape.

The new combination is provided since it does not

appear that Brotherus ever made the combination indicated

in Index Muscorum 2: 347 (1962). Brotherus frequently

made a combination by citing in the genus, his new combi-

nation followed by the author and genus of the basionym. In

this case, Natiirl. PflFam. 10: 324 (1924), he only stated his

opinion “ Ph. succuleatum Wager & Wright, Siidafr.,

gehort zu Funaria (Entosthodon) ” at the end of his treat-

ment of Physcomitrium.

The type at PRE is unfortunately mixed, because of

Wager’s practice of combining all subsequent collections of

a species in a single packet. Tne specimen at BM, however,

must have been sent soon after its collection, and hopefully

is not a mixture.

3. Funaria bergiana (Hornsch.) Broth, in

Natiirl. PflFam. 1: 524 (1903); 10: 328 (1924).

Type: Cape, “Montis Leonis”, Bergius s.n.,

15 Sept. 1816. (BM!).

Weissia bergiana Hornsch., Hoit. Phys. Berol. 59

(1820). Entosthodon bergii Bruch & Schimp., nom. illeg. ,

in B.S.G., Bryol. Eur. 3: 256 (1841). Physcomitrium ber-

gianus (Hornsch.) C. Mull, in Linnaea 18: 696 (1844).

Entosthodon bergianus (Hornsch.) C. Mull., Syn. Muse. 1:

126 (1848); Sim, Bryo. S. Afr. 296 (1926).

Funaria gymnostoma Dix. in S. Afr. J. Sci. 18: 318

(1922). Type: Natal, Goodoo, Wager s.n., Jan. 1918, PRE-

CHI 1990 (BM, holo., PRE!).

Plants small, gregarious, yellow-green to

light green; terricolous. Stems 2-3 mm long.

Leaves crowded above, weakly contorted and

appressed dry, widespreading wet; elliptical to

obovate, (1,5— )2, 0—2, 5 mm long; apex mu-

cronate to apiculate, apiculus to 0,5 mm long,

with linear, thickened cells, rarely absent on

some leaves; base scarcely differentiated to ob-

long; margins plane, entire. Costa ending be-

low apex; in section round, guide cells 2, ex-

posed ventrally, dorsal stereia band weak, 2—3

cells thick, cells reddish, dorsal surface cells

similar to guide cells. Laminal cells rectangu-

lar, rhomboidal and hexagonal; basal cells rec-

tangular.

Seta 4-6 mm long, yellowish; capsule

erect, pyriform, 1,2— 1,5 mm long, yellowish

brown, neck differentiated; exothecial cells rec-

tangular, incrassate, sometimes strongly so,

with 4-6 rows of quadrate to transversely rec-

tangular cells at mouth; stomata present on

neck; peristome single, fragile, teeth narrowly

triangular, 0,17 mm long, striate below, papil-

lose at apex, reddish yellow; operculum plano-

convex, cells not twisted; spores rounded,

25—30 pm, yellowish brown, weakly papil-

lose. Fig. 91: 21-27.

Fig. 91. — Funaria limbata (1-13): 1. habit, x 1; 2. habit, x 10; 3. part of stem in cross section, x 245; 4. leaf, x

35; 5. leaf in cross section, x 175; 6. basal leaf cells (right side), x 175; 7. blunt leaf apex, x 175; 8. acute leaf apex, x

175; 9. laminal cells, x 350; 10. capsule mouth and peristome teeth seen from above, x 87; 11. part of capsule mouth

showing cells, x 87; 12. capsule, dry, x 10; 13. spore, x 700. F. succuleata (14-20): 14. habit, x 1; 15. habit, x 10; 16.

leaf, x 35; 17. basal leaf cells (right side), x 175; 18. leaf apex, x 175; 19. laminal cells, x 350; 20. part of capsule mouth

showing cells and peristome tooth, x 245. F. bergiana (21-27): 21. habit, x 1; 22. habit, x 10, 23. leaf, x 35; 24. basal

leaf cells (right side), x 175; 25. laminal cells, x 350; 26. leaf apex, x 175; 27. part of capsule mouth showing cells and

peristome tooth, X 245. (1-6 & 8- 13, Thomas PRE-CH2961; 7, Sim 8585; 14-20, Wager s.n., PRE-CH12075; 21-26,

Anderson 6; 27, Volk 95%l).

324

Funariaceae

Funariaceae

325

Endemic to southern Africa, F. bergiana has been

collected from isolated communities in central South West

Africa/Namibia, the western and southern Cape, Natal and

eastern Transvaal. Map 119.

Vouchers: Anderson 6; Garside 6589; Magill 3135;

Schelpe 7772.

Funaria bergiana is recognized by its apiculate or

muticous leaves with costa ending below the apex and its

erect, symmetrical capsules. Bruch and Schimper (1841)

described and illustrated the costa of the lower stem leaves

as ‘costa excurrente mucronatis’. I have not seen leaves

with the costa excurrent, however the costa is frequently

longer, even to subpercurrent in the lower leaves. Cells of

the apiculus are elongated and generally incrassate and

could give the impression of an excurrent costa.

Remnants of a peristome were found on the only cap-

sule seen of F. gymnostoma; this and its leaf shape indicate

its relationship to F. bergiana.

4. Funaria rottleri (Schwaegr.) Broth, in

Natiirl. PflFam. 1: 525 (1903); 10: 329 (1924).

Type: Tranquebar, Rottler s.n. (G, holo. !).

Gymnostomum rottleri Schwaegr., Sp. Muse. Suppl. 1:

24 (1811). Physcomitrium rottleri (Schwaegr.) Hampe ex

C. Mull, in Linnaea 18: 696 (1845). Entosthodon rottleri

(Schwaegr.) C. Mull., Syn. Muse. 1: 121 (1848); Sim,

Bryo. S. Afr. 294(1926).

lEntosthodon campy lopodioides C. Mull, in Hedwigia

38: 60 (1899); Dix. in Trans. R. Soc. S. Afr. 8: 199(1920);

Sim, Bryo. S. Afr. 294 (1926). Type: Orange Free State,

Taaibosch Kranz, Rhenoster River, Rehmann s.n., 1875

(G!).

Plants small, scattered, light green to yel-

low-green; terricolous. Stems 2-5 mm long.

Leaves crowded above, in a rosette, weakly

contorted dry, widespreading wet; elliptical to

obovate, 2-4 mm long; apex acute to apiculate;

margins plane, entire or marginal cells bulging

above. Costa short-excurrent, mucro to 0,5 mm

long; in section guide cells 2, thin-walled, ven-

tral surface cells 2, thin-walled, dorsal stereid

band 2-6 cells thick, reddish, dorsal surface

cells smaller than guide cells, incrassate. Lami-

nal cells short-rectangular, thin-walled, becom-

ing rhomboidal near apex; basal cells rectangu-

lar.

Seta 5—10 mm long, reddish brown; cap-

sule erect to inclined, cylindrical to obconical,

3-4 mm long, reddish brown, mouth very

broad, urn constricted below mouth when dry,

neck differentiated; exothecial cells oblong-

hexagonal, incrassate, with 4-6 rows of

transversely rectangular cells at mouth; peri-

stome very rudimentary or absent; operculum

plano-convex, cells not twisted; spores subround

20—25 pm, brownish, papillose. Fig. 92: 1—8.

Reported from Asia and Africa, F. rottleri is quite

widespread in the eastern and southern parts of the Flora

area. The species grows on soil in open grasslands and

shrublands of the western, central and eastern Transvaal,

Orange Free State, Lesotho, Natal, Transkei and the east-

ern, southern, central, northern and southwestern Cape.

Map 120.

Vouchers: Ellis 3103; Hardy et al. 5339a; Jacot Guil-

larmod 8201; Magill 4549, 4757; Vahrmeijer PRE-

CH12690; Van Rooy 509; Von Breitenbach 96.

Most similar to F. longicollis, but differing in smaller

leaves with entire margins and longer, ± inclined capsules;

see note under that species.

5. Funaria longicollis Dix. in S. Afr. J.

Sci. 18: 318 (1922); Broth, in Natiirl. PflFam.

10: 327 (1924); 1 1: 530 (1925). Syntypes: Zim-

babwe, Zimbabwe Ruins, Sim 8735, 8796,

8797; Khami Ruins, Sim 8842 (BM; PRE!).

Entosthodon dixonii Sim, Bryo. S. Afr. 296 (1926), non

E. longicollis Mitt. (1869).

Fig. 92. — Funaria rottleri (1-8): 1. habit, X 1; 2. habit, x 10; 3. leaf, x 35; 4. leaf in cross section, x 200; 5. basal

leaf cells (left side), x 175; 6. laminal cells, x 350; 7. leaf apex, x 175; 8. part of capsule mouth showing cells, x 175.

F. longicollis (9-14): 9. habit, x 1; 10. habit, x 10; 11. leaf, x 35; 12. laminal cells at upper margin, x 350; 13. leaf

apex, x 175; 14. part of capsule mouth showing cells, x 350. F. urceolata ( 15-22): 15. habit, x 1; 16. habit, x 10; 17.

leaf, x 35; 18. laminal cells, x 350. 19. leaf apex, x 175; 20. capsule, dry, x 10; 21. exothecial cells, x 350; 22. part of

capsule mouth showing cells, x 350. F. clavata (23-28): 23. habit, x 1; 24. habit, x 10; 25. leaf, X 35; 26. basal leaf

cells, x 175; 27. leaf apex, x 175; 28. laminal cells, x 600. (1-8, Magill 4213; 9-14, Sim 8735; 15 — 22, Rooper s.n.;

23-28, Magill 4003).

326

Funariaceae

Plants small to medium-sized, scattered,

green to dark green; terricolous. Stems 2-5 mm

long. Leaves larger and crowded above, con-

torted dry, widespreading wet; obovate to

subspathulate, 3-4 (-5) mm long; apex acute to

obtuse, apiculate; base oblong; margins plane,

entire below, crenulate above, frequently irre-

gularly so with some strong teeth near apex.

Costa short excurrent, mucro to 0,2 mm long;

in section round, guide cells 2, small, thin-

walled, ventral cells in single row, slightly

larger than guide cells, thin- walled, dorsal ster-

eid band 2-3 cells thick, reddish, dorsal surface

cells larger than guide cells, thin-walled. Lami-

nal cells rectangular, becoming rhomboidal

above; marginal cells bulging, thin-walled; ba-

sal cells short-rectangular.

Seta 5-7 mm long, reddish yellow; capsule

erect, narrowly pyriform, 2, 0-2, 5 mm long,

red-brown, urn cylindrical, constricted below

mouth when dry, neck differentiated, sulcate;

exothecial cells rectangular, incrassate, with

8-10 rows of transversely rectangular cells at

capsule mouth; stomata present on neck, phane-

ropore; peristome absent, occasionally with

traces of rudimentary membrane; calyptra

plano-convex, cells very slightly twisted;

spores rounded, 25-30 pm, yellowish, sparsely

papillose with distinct tetrad scar (probably im-

mature). Fig. 92: 9-14.

The species has been collected only a few times in

southern Zimbabwe and in the northern, eastern and central

Transvaal. Map 121.

Vouchers: Magill 5018; Van Rooy 613, 621.

Similar in many respects to F. rottleri, but differing in

larger leaves with crenulate margins and smaller, erect cap-

sule. The spores of the two species also differ in size and F.

lon^icollis lacks a peristome. The peristome of F. rottleri is

rudimentary and fragile and missing on older capsules.

6. Funaria urceolata (Mitt.) Magill in

Mem. bot. Surv. S. Afr. 43: 7 (1979). Type:

Cape, East London, Rooper s.n. (NY, holo. !).

Entosthodon urceolatus Mitt, in Harvey, Thes. Cap. 1:63

(1859).

Funaria rufinervis Dix. in Trans. R. Soc. S. Afr. 18: 254

(1930). Type: Natal, National Park, Wager 758 (BM,

holo.!; PRE!).

Plants small, scattered, light green; terri-

colous. Stems 3-5 mm long. Leaves crowded

and larger above, weakly contorted dry, wide-

spreading wet; Ungulate to narrowly obovate,

3, 0-3 ,5 mm long; apex short-acuminate; base

oblong; margins plane, entire. Costa short ex-

current, mucro 0,2-0, 3 mm long; in section

round, guide cells 2, weakly thickened, ventral

cells in single row, thin- walled, dorsal stereid

band weak, 2 cells thick, dorsal surface cells

similar to ventral cells, thin-walled. Laminal

cells irregularly short-rectangular, thin-walled;

basal cells rectangular, thin-walled.

Seta 4-5 mm long, reddish yellow; capsule

inclined, narrow-pynform, 2-3 mm long,

mouth wide, urn constricted below mouth when

dry, neck differentiated, sulcate; exothecial

cells rectangular to rhomboidal, incrassate,

short rectangular to quadrate or hexagonal in

neck, thin- walled, with 4 rows of transversely

rectangular cells at mouth and 1-2 rows of

larger, ± quadrate cells just below; stomata

present on neck, subphaneropore; peristome ab-

sent; calyptra not seen, described as short-

conic, subplano-convex; spores subround,

27-30 pm, brownish, vermiculate to bluntly

papillose. Fig. 92: 15-22.

Endemic to southern Africa, F. urceolata is known

from the mountains of Natal and the southern Cape. Map

121.

Vouchers: Sim 10126; Wager 758.

The species is recognized by its narrow leaves with

short-excurrent costa, and narrowly pyriform, inclined,

gymnostomous capsules.

7. Funaria clavata (Mitt.) Magill, comb,

nov. Type: Cape of Good Hope, Menzies s.n.

(NY, holo.!).

Entosthodon clavata Mitt, in Harvey, Thes. Cap 1:63

(1859).

Funariaceae

327

Plants medium-sized, scattered, yellow-

green; terricolous. Stems 2-4 mm long. Leaves

crowded and larger above, appressed dry, wide-

spreading wet; elliptical to broadly elliptical or

obovate, 2, 0-3, 2 mm long; apex abruptly or

gradually apiculate to piliferous; base not dif-

ferentiated; margins plane, entire. Costa ending

below apex; in section bulging dorsally, guide

cells 3, thin- walled, ventral cells in single row,

larger than guide cells, thin-walled, dorsal ster-

eid band small, 1-2 cells thick, dorsal surface

cells small, incrassate. Laminal cells rectangu-

lar, thin-walled; basal cells larger, rectangular.

Seta 6 mm long, red-brown; capsule hori-

zontal, clavate, 4 mm long, red-brown, neck

not strongly differentiated, nearly smooth;

exothecial cells rectangular to rhomboidal,

strongly incrassate, in base shorter, quadrate to

hexagonal, thin-walled, with 4 rows of

transversely rectangular cells at capsule mouth;

stomata present at base of neck; peristome ab-

sent; operculum flat, cells not twisted; spores

rounded, 25 pm, brownish yellow, weakly pa-

pillose with tetrad scar (probably immature).

Fig 92: 23-28.

Endemic to southern Africa; the species is collected on

soil in the shrublands of the western Cape. Map 121 .

Vouchers: Magill <4 Schelpe 3845, 4003; Perold 481;

Schelpe 7716; V.d. Westhuizen 40.

Funaria clavata is a distinctive species, identified by

its broadly elliptical-apiculate leaves with costa ending be-

low the apex and its horizontal, gymnostomous, clavate

capsules.

8. Funaria rhomboidea Shaw in Cape

Monthly Mag. 17: 315 (1878). Type: Cape,

near Graaff-Reinet, McLea s.n., sub Rehmann

523. (NH!).

Entosthodon schinzii Geheeb in Bull. Herb. Boissier 4:

411 (1896). Funaria schinzii (Geheeb) Broth, in Natiirl.

PflFam. 1: 524 (1903). Type: South West Africa/Namibia,

Comagas, Schinz s.n., 24 Apr. 1885 (Z!).

Entosthodon rivale Geheeb in Bull. Herb. Boissier 4: 41 1

(1896). Physcomitrium rivale (Geheeb) Broth, in Natiirl.

PflFam. 1: 519 (1903). Type: South West Africa/Namibia,

Schinz s.n., Apr. 1885 (Z!).

Plants small, scattered, yellowish green;

terricolous. Stems 2-3 mm long. Leaves

crowded above, in a rosette, weakly contorted

dry, widespreading wet; elliptical to obovate,

2, 5-3, 5 (-4,5) mm long; apex acute to obtuse-

apiculate, occasionally very short acuminate;

base oblong; margins plane, crenulate to bluntly

toothed above base, occasionally strongly so.

Costa ending well below apex; in section

rounded, guide cells 2, thin- walled, ventral

cells 2, in single layer, thin- walled, dorsal ster-

eid band weak, 2-3 cells thick, dorsal surface

cells thin-walled. Laminal cells rectangular to

rhomboidal, thin-walled, becoming somewhat

irregular to rhombic at apex; marginal cells

slightly larger, bulging at distal ends to form

crenulate margin; basal cells rectangular.

Seta 6-7 mm long, reddish yellow; cap-

sules erect, pyriform, 1 ,5—2,0 mm long, red-

dish brown, urn short cylindrical or occasion-

ally inflated and globose, neck strongly sulcate,

collapsed wet or dry; exothecial cells rhom-

boidal, thickened, with 2-4 rows of transversely

rectangular cells at mouth, stomata present on

neck; peristome double, exostome teeth irregu-

lar, perforated, red-yellow, 0,10-0,15 mm

long, vertically striate below, papillose above,

endostome rudimentary, attached to teeth, yel-

lowish, smooth; operculum short-conic, cells

twisted counter-clockwise; spores round, 25-35

pm, brownish, weakly papillose. Fig. 93:1-8.

Endemic to southern Africa, F. rhomboidea is found in

dry grassland communities of central South West Africa/

Namibia, the central Cape, Orange Free State and northern

Natal. Map 122.

Vouchers: Potts 4; Tolken 5737; Volk 481, 5374,

12458.

Although considered a nomen nudum by Sim ( 1926) and

the authors of Index Muscorum (1962), Shaw (1878) pro-

vided a short discretion in English when he published F.

rhomboidea .

Map 122. — 9 Funaria rhomboidea

▲ Funaria spathulata

328

Funariaceae

Funariaceae

329

This species is similar to F. limbata, but differs in the

absence of a well defined border. The marginal cells in F.

rhomboidea are slightly larger with the upper ends of the

cells bulging outward resulting in a crenulate margin. It is

also similar to F. spathulata differing primarily in capsule

shape and size.

The type specimen of Entosthodon schinzii Geheeb

agrees well with Funaria rhomboidea. Only fragments of

the peristome were found in the mouth of the capsule exam-

ined, however their structure seems to indicate a peristome

similar to that of F. rhomboidea.

The type of Entosthodon rivale Geheeb contains slightly

larger plants, but otherwise agrees gametophytically with F.

rhomboidea. All of the sporophytes seen were immature, so

the original reference to a gymnostomous capsule could not

be substantiated. The capsule shape and exothecial cells

clearly indicate Funaria as the proper genus for the species.

9. Funaria spathulata Schimp. ex C. Mull.

in Hedwigia 38: 61 (1899); Broth, in Natiirl.

PflFam. 10: 330 (1924). Type: Cape, Groen-

kloof, Breutel s.n. (BM, holo.!; G!).

Entosthodon spathulatus (C. Mull.) Sim, Bryo. S. Afr.

298 (1926).

Funaria dieterlenii Th6r. in Bull. Mus. Hist, nat., Paris

30: 240 (1924); Broth, in Naturl. PflFam. 11: 530 (1925).

Type: Lesotho, Leribe, Dieterlen s.n. (PC, holo. !).

Plants small to medium-sized, scattered or

gregarious, light green to yellow-green; terrico-

lous. Stems 2-4 mm long. Leaves larger and

crowded above, spirally contorted dry, wide-

spreading wet; broadly elliptical to obovate or

subspathulate, (l,5-)2,0-3,5 mm long; apex

acute to short-acuminate; base oblong or scar-

cely differentiated; margins plane, crenulate to

denticulate. Costa ending below apex to almost

percurrent, in section round, guide cells 2, thin-

walled, ventral cells in single layer, 2, thin-

walled, dorsal stereid band small and weak, 2

cells thick, dorsal surface cells similar to guide

cells, weakly thickened. Laminal cells rectang-

ular or rhomboidal at apex; basal cells rectangu-

lar.

Seta (5— )8— 10 mm long, yellowish brown;

capsule asymmetrical, erect to inclined with ob-

lique mouth to arcuate with mouth vertical,

1,5-2, 5 mm long, yellow-brown, neck short,

well defined; exothecial cells on convex surface

rectangular, thickened, but irregular and shorter

on concave surface, on neck short-quadrate to

angular, thin-walled; stomata present on neck,

phaneropore; peristome double, exostome teeth

narrowly triangular, perforated, 0,3 mm long,

occasionally somewhat irregular, not twisted,

coarsely striated, reddish yellow, endostome

rudimentary, weakly papillose, yellow; opercu-

lum plano-convex, cells not twisted; spores

rounded, 23-32 pm, red-brown, strongly papil-

lose to vermiculate. Fig. 93: 9-15.

Endemic to southern Africa, F . spathulata is collected

in locally moist areas of the western and southern Cape,

Natal and Lesotho. Map 122.

Vouchers: Magill 4226, 4622; Oliver et al. 647; Schelpe

4917a;5c/iwt/7;81 1 1.

This species is macroscopically similar to F. hygrome-

trica, but distinct in its costa consistently ending below the

apex, its more erect capsules, rudimentary endostome and

cells of the operculum not twisted. There is also a resem-

blance to F. rhomboidea. however its capsules differ in size

and shape and the twisted cells of the operculum.

10. Funaria hygrometrica Hedw., Sp.

Muse. 172 (1801); Broth, in Naturl. PflFam.

10: 331 (1924); Sim, Bryo. S. Afr. 298 (1926);

Crum, Moss. Great Lakes Forest 135 (1973);

Scott & Stone, Moss. S. Aust. 254 (1976);

Smith, Moss FI. Brit. Irel. 340 (1978). Type:

Europe.

Funaria plagiostoma C. Mull, in Bot. Ztg 13: 748

(1855). Entosthodon plagiostomus (C. Mull.) Sim, Bryo.

S. AfT. 297 (1926). Syntypes: Cape, prope Swellendam,

Mundt; Pappe.

Funaria gracilescens Schimp. ex C. Miill. in Bot. Ztg

16: 154 (1858); Broth, in Naturl. PflFam. 10: 332 (1924).

Type: Cape, Zitzkamma, Breutel s.n. (BM, holo.!).

Funaria lonchopelma C. Miill. in Hedwigia 38: 61

(1899). Type: Cape, Montagu Pass, Rehmann 181 (BM!).

Plants medium-sized, gregarious, light

green; terricolous. Stems 3-8 mm long. Leaves

imbricate in bulb-like cluster, weakly contorted

dry; broadly elliptical to obovate, 2-4 mm long,

concave; apex acute to short acuminate; mar-

gins plane to broadly inrolled above, entire to

FIG. 93. — Funaria rhomboidea (1-8): 1. habit, x 1,2. habit, x 10; 3. leaf, x 35; 4. basal leaf cells at left margin,

x 175; 5. laminal cells, x 350; 6. leaf apex, x 175; 7. part of capsule mouth showing cells and peristome tooth, x 350; 8.

capsule, dry, x 10. F. spathulata (9- 15): 9. habit, x 1; 10. habit, x 10, 11. leaf, x 35; 12. basal leaf cells (left side), x

175; 13. laminal cells, x 350; 14. leaf apex, x 175; 15. part of capsule mouth showing cells, peristome tooth and spore

tetrads, x 175. F. hygrometrica (16— 25): 16. habit, x 1; 17. habit, x 10; 18. leaf, x 35; 19. basal leaf cells (right side),

x 175; 20. laminal cells, x 350; 21. leaf apex, x 175; 22. operculum (split and flattened by coverslip), x 70; 23. part of

capsule mouth showing cells and peristome, x 175; 24. capsules, dry and operculate on left, wet and deoperculate on right,

x 10; 25. calyptra, x 10. (1-8, Rehmann 523; 9- 15, Dieterlen 686; 16-21 & 23-25, Garside 6628; 2, Wood 282).

330

Funariaceae

weakly serrate. Costa ending just below apex to

short excurrent; in section round, guide cells 4,

ventral cells in single row of 2-3 cells, larger

than guide cells, thin- walled, dorsal stereid

band small, 2-4 cells thick, dorsal surface cells

similar in size to guide cells, incrassate. Lami-

nal cells inflated, subhexagonal to short rec-

tangular, somewhat narrower at margin; basal

cells oblong.

Seta 20-50 mm long, yellowish; capsules

horizontal to pendent, asymmetrical, arcuate,

subpyriform, to 3 mm long, reddish brown, sul-

cate, mouth oblique, neck differentiated, sul-

cate when dry; exothecial cells rectangular,

thin- walled, irregular and shorter on concave

surface, at mouth with 4-8 rows of transversely

rectangular cells, neck cells quadrate to an-

gular; stomata present on neck; annulus well

developed, revoluble; peristome double, exo-

stome teeth sigmoid, triangular-twisted, joined

at apex to an ephemeral, cancellate disk, ventral

surface papillose-striate below, papillose

above, dorsal surface strongly trabeculate, en-

dostome segments lanceolate, papillose, cilia

absent; operculum convex, cells twisted coun-

ter-clockwise; spores round, 14-17 pm, yel-

low-brown, smooth. Fig. 93: 16-25.

Almost cosmopolitan in its distribution F. hygromet-

rica is the most frequently collected and widespread species

of Funaria in southern Africa. Map 123.

Vouchers: Crosby & Crosby 7585; De Winter 9378a;

Esterhuysen 18578; Gar side 6545; Hardy 5003; Lambert 8;

Magill 3431, 4619, 6286; Perold 5; Phelan et al. 60; Pie-

naar 15; Smook 887; Van Rooy 1 , 453; Wells 70.

The plants of F. hygrometrica are generally the largest of

any of the species present in southern Africa. The asymme-

trical, horizontal to pendulous, sulcate capsules are very

distinctive, and with their broad oblique mouths, strongly

developed, revoluble annulus and double peristome will

quickly place the species.

SPLACHNACEAE

331

Plants medium-sized to large, forming loose tufts, dark green; terricolous. Stems erect, radi-

culose below; central strand present. Leaves lingulate to obovate; margins frequently serrate to

dentate above. Costa strong, generally excurrent; in section with guide cells and stereid bands.

Laminal cells hexagonal to oblong-rhomboidal, smooth.

Monoicous. Perichaetia terminal. Seta erect; capsule symmetrical, urn cylindrical, neck fre-

quently differentiated; peristome single, teeth 16, usually fused in pairs; operculum conic; calyptra

campanulate, rough or hairy; spores small.

Primarily a Northern Hemisphere family containing 7 genera, Splachnaceae is generally collected on decaying organic

matter. Only a single genus, Tayloria, is known from southern Africa.

TAYLORIA

Tayloria Hook, in J1 Sci. Arts, Lond. 2: 144 (1816); Broth, in Natiirl. PflFam. 10: 336 (1924);

Sayre in Grout, Moss FI. N. Amer. 2: 93 (1935); Sainsb., N. Zeal. Mosses 249 (1955); Gangulee,

Moss. E. India 4: 874 (1974); Smith, Moss FI. Brit. Irel. 352 (1978). Type species: T. splach-

noides (Schwaegr.) Hook.

With characters of family.

The genus Tayloria contains approximately 60 species that are rather evenly distributed. The largest concentration of

species is found in northwestern South America. Seven species are known from the African mainland and until recently,

only from the tropics above 15 ° S latitude. Tayloria ortnodonta (P. Beauv.) Demar. has been reported from Zimbabwe

(Magill & Schelpe 1979) and T. isleana (Besch.) Broth, is now known from southern Africa, its only locality outside

Reunion. The genus bears some vegetative resemblance to members of Mniaceae, but the sporophytes are very distinctive.

1 Leaves toothed in upper third, upper marginal cells and dentations more strongly thickened than laminal cells; calyp-

tra sparsely hairy 1. T. isleana

1 Leaves toothed in upper half; upper marginal cells not differentially thickened; calyptra papillose 2.T. orthodonta

1. Tayloria isleana (Besch.) Broth.,

Natiirl. PflFam. 1 (3): 503 (1903); 10: 339

(1924); Koponen & Weber in Ann. Bot. Fenn.

9: 131 (1972). Type: Reunion, G. del’Isle, Le-

pervanche 410, 1877 (BM!).

Orthodon isleanus Besch., Ann. Sci. Nat. Bot. Ser. 6, 9'

373(1880).

Plant medium-sized to large, loosely caes-

pitose, dark green to yellow-green; terricolous.

Stem erect, 8—12 mm long, simple; in section

round, central strand large, inner cortical cells

large, in 4—5 rows, thin-walled, outer cortical

cells smaller, in 2—3 rows, thick-walled.

Leaves larger above, when wet widespreading,

when dry erect-appressed; obovate, 2, 5 -3,0

mm long, apex obtuse, cuspidate, base oblong,

margins plane, entire below, strongly and

sharply dentate or spinose above, teeth multi-

cellular, directed forward, to 0,2 mm long, api-

cal cell elongated. Costa strong, short excur-

rent, awn 0,3 -0,6 mm long, smooth; in section

round, bulging dorsally, guide cells 2, exposed

ventrallv, ventral stereid band absent, dorsal

stereid band weak, 2-3 cells thick, dorsal sur-

face cells larger than guide cells, thick- walled.

Upper laminal cells somewhat variable in size

and shape, mostly rectangular-rhomboidal,

45—100 pm long, 30-38 pm wide, walls thin,

smooth; marginal cells and cells of teeth

somewhat thickened; basal cells rectangular.

Autoicous. Perichaetia at apex, perichae-

tial leaves similar to vegetative leaves, 2, 0-2, 5

mm long. Seta 6-7 mm long, yellowish; cap-

sule ovoid, 1, 5-2,0 mm long, constricted be-

low mouth when dry, reddish yellow; exothe-

cial cells rounded quadrate to rectangular or

angular; stomata on lower urn, subphaneropore;

peristome single, light yellow, teeth 16 in 8

332

Splachnaceae

Fig. 94. — Tayloria isleana (1-10): 1. habit, x 1; 2.

habit, x 10; 3. portion of stem in cross section, x 175; 4 &

5. leaves, x 35; 6. leaf in cross section, x 175; 7. basal leaf

cells at right margin, x 175; 8. upper laminal cells, X 400;

9. leaf apex, x 175; 10. peristome tooth and part of capsule

mouth, x 160. T. orthoaonta (11-16): 11. habit, X 1; 12.

habit, x 5; 13. leaf, X 17; 14. leaf in cross section, x 175;

15. basal leaf cells (left side), x 70; 16. leaf apex, x 175.

(1-10, Reid 721 1 ; 1 1 - 16, Rehmann 559).

Splachnaceae 333

MAP 124. — • Orthodontium lineare

A Tayloria isleana

♦Tayloria orthodonta

Eairs, triangular, weakly papillose, 0,45 pm

igh; operculum conic; calyptra campanulate,

3,0 — 3,5 mm long, hairy; spores rounded,

10-14 p m, more or less smooth, light yellow.

Fig. 94: 1-10.

The species was previously known only from Reunion.

The southern African specimen was collected in Karkloof,

northwest of Pietermartizburg, Natal in 1915, but was only

recently discovered in a mixed collection. Map 124.

Voucher: Reid 721 la.

The species can be indentified by its obovate-cuspidate

leaves with strongly toothed upper margins and leaf cells

that are large, thin-walled and smooth. Tayloria orthodonta

differs in having a shorter apiculus and papillose calyptra.

2. Tayloria orthodonta (P. Beauv.) Wijk

& Marg. in Taxon 17: 467 (1968).

Bryum orthodontum P. Beauv., Prodr. Cinq. Six. Fam.

Aetheogam. 48 (1805).

Plant medium-sized to large, caespitose,

yellow-green; terricolous. Stem erect, to 15 mm

long, simple; in section round, central strand

large, inner cortical cells large, in 4-6 rows,

thin-walled, outer cortical cells in 2-3 rows,

thick-walled. Leaves when wet widespreading,

when dry appressed, stem leaves oblong to ob-

ovate, 4, 0-4, 5 mm long, apex subacute to ob-

tuse, awned; base oblong; margins plane, entire

below, dentate in upper half, teeth multicellu-

lar, apical cell not differentiated. Costa strong,

short excurrent, awn 0,2-0, 3 mm long,

smooth; in section bulging dorsally, guide cells

large, 2-3, thickened, ventral stereid band ab-

sent, ventral surface cells large, thick-walled,

dorsal stereid band strong, 6-8 cells thick,

dosal surface cells large, thick-walled. Upper

laminal cells rectangular to rectangular-hexago-

nal, walls thickened, marginal cells thm-

walled, smooth, basal cells rectangular, walls

thin, smooth.

Autoicous. Perichaetia at apex, perichae-

tial leaves similar to vegetative leaves. Fig. 94:

11-16.

The species is known from Guinea through equatorial

Africa to Malawi and Reunion. This is the first report of the

species to South Africa. The specimen was collected by

Rehmann in woods at Houtbosch in the northern Transvaal.

Map 124.

Voucher: Rehmann 559.

The southern African specimen has the remnants of a

seta but other parts of the sporophyte are missing. The

species is similar to T. isleana with the important difference

of a papillose rather than hairy calyptra. Tne upper marginal

cells in this species are not thickened and the apical cells of

the marginal teeth are not elongated.

BRYACEAE

335

by J. van Rooy and R. E. Magell

Plants small to robust, solitary to caespitose or forming dense cushions, generally yellow-

green to dark green or reddish green, occasionally glossy or with metallic lustre; terricolous,

saxicolous, corticolous or humicolous. Stems erect, infrequently from rhizomes, simple or fre-

quently branched by subperichaetial innovations, in section with central strand; frequently tomen-

tose; tubers occasionally present. Leaves variable in size and shape, equidistant to crowded above

or in a rosette, mostly smaller below, imbricate or appressed to erect- spreading, straight or va-

riously twisted when dry; margins plane to recurved, entire to variably dentate, border present or

absent. Costa ending below apex, percurrent or excurrent as long or short awn; in section bulging

dorsally, ventral stereid band absent, generally with dorsal stereid band. Laminal cells mostly

rhomboidal, smooth; basal cells quadrate to rectangular. Propagulae, gemmae or filamentous

gemmae infrequently produced.

Dioicous, variably monoicous or heteroicous. Perichaetia terminal or quickly lateral through

innovations, leaves not strongly differentiated. Seta elongate, erect; capsule erect to pendulous,

generally pyriform, straight or rarely curved, neck differentiated; stomata on neck; annulus fre-

quently differentiated; peristome double or rarely single, exostome teeth 16, endostome segments

keeled, alternating with teeth, cilia present or absent, basal membrane low or high; operculum

convex, apiculate, mammillate or rostellate; calyptra cucullate, smooth; spores round, smooth to

papillose.

Nine of the twenty genera recognized in Bryaceae are known from southern Africa. Members of the family are easily

recognized by their sporophytic characters. The seta is long with a relatively large capsule, mostly inclined to pendulous and

generally pyriform in shape with the neck tapering to the seta. The peristome is generally double and frequently well

developed. The endostome frequently has a high basal membrane with 16 segments alternating with the exostome teeth and

with intermediate cilia varying in number. The peristome is variously reduced in some of the genera. Infrageneric variation

in peristome development is common in the Bryaceae and Rhodobryum is the only non-monotypic genus in the Flora area in

which the peristome is uniformly well developed. Sterile plants can be recognized by the leaves frequently crowded towards

the stem apex with the lower leaves distant and smaller, the smooth, rhomboidal laminal cells and the strong costae.

1 Leaf cells elongate, mostly 4 times or more as long as broad (>4:1); leaves generally

narrow:

2 Leaves imbricate, equidistant; stems frequently julaceous 7. Anomobryum

2 Leaves appressed to erect-spreading, crowded above, distant below, rarely equidistant;

stems not julaceous:

3 Leaves setaceous or linear:

4 Costa broad; exostome teeth as long as segments; endostome cilia

present 6. Leptobryum

4 Costa narrower; exostome teeth shorter than segments; endostome cilia

absent 1 . Orthodontium

3 Leaves ovate to lanceolate:

5 Sporophytes quickly lateral; peristome double, frequently appearing single,

exostome teeth rudimentary, short 2. Mielichhoferia

5 Sporophytes terminal; peristome double, exostome teeth long:

6 Costa percurrent or ending below apex 4. Pohlia

6 Costa excurrent 8. Bryum

1 Leaf cells shorter, mostly less than 4 times as long as broad (<4: 1); leaves generally broad:

7 Leaves imbricate, equidistant; costa percurrent; upper laminal cell walls thickened to

incrassate 7. Anomobryum

336

Bryaceae

7 Leaves appressed to erect-spreading or variously twisted, crowded above, distant

below; if imbricate and equidistant then costa excurrent or upper laminal cells thin-

walled:

8 Plants mostly corticolous, rarely terricolous; capsules erect to horizontal, mouth small:

9 Perichaetium a short, basal bud or quickly lateral through innovation; peristome

double, frequently appearing single, exostome teeth rudimentary,

short 2. Mielichhoferia

9 Perichaetium terminal but overgrown by subperichaetial innovation; peristome

double, exostome teeth ± long 3. Brachymenium

8 Plants mostly terricolous, rarely corticolous; capsules inclined to pendulous, mouth

large:

10 Stems frequently erect from rhizomes; upper leaves crowded in a conspicuous

rosette with lower leaves reduced; polysetaceous 9. Rnodobryum

10 Rhizomes absent; leaves equidistant and about equal in size, comose or crowded

in rosettes with lower leaves smaller; unisetaceous:

1 1 Plants small, reddish hyaline; leaves broadly ovate; exostome teeth shorter than

endostome segments 5. Plagiobryum

1 1 Plants small to robust, whitish or silvery green or variously green; leaves ovate

to lanceolate or oblong to obovate or spathulate; exostome not shorter than

segments 8. Bryum

1. ORTHODONTIUM

Orthodontium Schwaegr., Sp. Muse. Suppl. 2 (2): 123 (1827); Broth, in Natiirl. PflFam. 10: 349

(1924); Sim, Bryo. S. Afr. 317 (1926); Meijer in Act. bot. need. 1: 5-80 (1952); Sainsb., N. Zeal.

Mosses 254 (1955); Scott & Stone, Moss. S. Aust. 296 (1976); Smith, Moss FI. Brit. Irel. 361

(1978); Catcheside, Moss. South Austr. 243 (1980). Type species: O. lineare Schwaegr.

Plants small, loosely caespitose; terricolous or corticolous. Stems erect, little branched; in

section round, central strand oi collapsed cells present, cortical cells in 3-4 rows. Leaves erect-

spreading or flexuose, linear-lanceolate. Costa generally ending below apex; in section with guide

cells exposed ventrally. Upper laminal cells long-rhomboidal, basal cells frequently bulging.

Autoicous, synoicous or heteroicous. Capsule erect or inclined, ovate or pyriform; stomata

phaneropore; annulus apparently absent; peristome double, exostome teeth generally shorter than

segments, cilia absent, basal membrane low; operculum rostellate; calyptra small, cucullate; spores

round.

Orthodontium is a world-wide genus of 14 species. Vegetatively the genus may be confused with members of

Dicranaceae but the capsule and peristome characters will place it in Bryaceae. Sterile specimens are identified by size, leaf

shape and areolation.

Orthodontium lineare Schwaegr., Sp.

Muse. Suppl. 2 (2): 188 (1827); Broth, in

Natiirl. PflFam. 10: 350 (1924); Sim, Bryo. S.

Afr. 317 (1926); Meijer in Act. bot. neerl. 1: 28

(1952); Sainsb., N. Zeal. Mosses 255 (1955);

Scott & Stone, Moss. S. Aust. 298 (1976);

Smith, Moss FI. Brit. Irel. 361 (1978); Catche-

side, Moss. South Austr. 245 (1980). Type:

Cape, Menzies s.n. (G, holo.).

Plants small, loosely caespitose, yellow-

green or green above, brownish below; terrico-

lous or corticolous. Stems 1-5 mm tall, irregu-

larly branched, yellowish brown to reddish

brown; rhizoids smooth, brownish or reddish.

Bryaceae

337

Leaves crowded, larger above, erect-spreading

or flexuose wet or dry, linear-lanceolate or lin-

ear, (1 — )2— 4(— 6) mm long; apex acuminate to

setaceous, frequently reddish below; margins

plane, entire to denticulate above. Costa gener-

ally weak, ending below apex to subpercurrent;

in section round to subround, guide cells

exposed ventrally, dorsal stereid band strong,

stereids in 3—4 rows, dorsal surface cells in 1

row, incrassate. Upper laminal cells rhomboi-

dal to linear-rhomboidal, (80— ) 100 -150

(-200) n m long, (7, 5— )10, 0-18,75 pun wide;

basal cells larger, frequently reddish, bulging,

rhomboidal or subrectangular.

Autoicous, synoicous or heteroicous. Peri-

chaetial leaves linear-lanceolate. Seta 10-13

mm long, yellow or reddish, slightly twisted;

capsule erect or inclined, ovate, cylindrical or

pyriform, urn 1,0 -1,5 mm long, yellowish to

reddish brown, neck wrinkled dry, 0,2-0, 6

mm long; exothecial cells irregularly rectangu-

lar, thin-walled, shorter and incrassate above;

stomata at base of urn and on neck, phanero-

pore; annulus apparently absent; peristome yel-

lowish, exostome teeth distant, trabeculate ven-

trally, linear, to 200 pun long, minutely papil-

lose, endostome segments generally longer, al-

ternating with teeth, linear, nodose, cleft

below, extending into basal membrane, cilia

absent, basal membrane low, minutely papil-

lose; operculum conical, rostellate; calyptra

small, cucullate, ephemeral; spores 22-26 /u,m,

brownish. Fig. 95.

This species is known from Australia, New Zealand,

Europe and southern Africa. In the Flora area it is rarely

collected on soil or bark in the southwestern Cape and Zulu-

land. Map 124.

Vouchers: Magill 5386; Magill & Schelpe 4102; Pil-

lans 4271; Thorne PRE-CH3643.

Orthodontium lineare can be distinguished from other

taxa in Bryaceae by the erect to inclined capsule, the peri-

stome with teeth shorter than the nodose segments and the

low basal membrane without cilia. The linear leaves with

long-rhomboidal cells and the smooth rhizoids are also dis-

tinctive.

FIG. 95. — Orthodontium lineare: 1. habit, x 1; 2.

habit, x 3; 3. stem in cross section, x 175; 4. leaf, x 17; 5.

leaf in cross section, x 175; 6. basal leaf cells (right side),

X 175; 7. laminal cells at right margin, x 175; 8. leaf apex,

x 175; 9. capsule, x 15; 10. stomatal apparatus, x 350;

11. part of capsule mouth showing cells and peristome, x

175; 12. spore, X 700. (1-3, 5 & 7-12, Thorne PRE-

CH3643; 4 & 6, Magill 4102).

338

Bryaceae

2. MIELICHHOFERIA

Mielichhoferia Nees & Hornsch., Bryol. Germ. 2: 179 (1831); Broth, in Natiirl. PflFam. 10: 350

(1924); Sim, Bryo. S. Afr. 314 (1926); Andrews in Grout, Moss FI. N. Amer. 2: 184 (1935);

Sainsb., N. Zeal. Mosses 252 (1955); Gangulee, Moss. E. India 4: 892 (1974); Scott & Stone,

Moss. S. Austr. 295 (1976); Smith, Moss FI. Brit. Irel. 359 (1978). Type species: not designated.

Schizymenium Harv. in Hooker’s Icon. PI. 3: 202(1840). Type species: S. bryoides Harv.

Plants small to medium-sized, densely caespitose, light green to yellow-green, occasionally

glossy; terricolous. Stems erect, generally branched; in section with central strand. Leaves ovate to

lanceolate, not distinctly larger above; margins unbordered, plane, entire to denticulate. Costa

ending below apex to percurrent or occasionally very short excurrent; in section with dorsal stereid

or substereid band. Upper laminal cells elongate rhomboidal to linear-rhomboidal; shorter and

broader toward base, rectangular.

Dioicous. Perichaetia quickly lateral through elongation of subperichaetial innovation, leaves

weakly differentiated from vegetative leaves. Seta elongate, erect; capsule nearly erect to inclined,

pyriform, neck short; peristome double, frequently appearing single, exostome teeth rudimentary,

snort, blunt, endostome fragile, segments 16, linear, on a low basal membrane, occasionally

rudimentary, cilia absent; operculum convex; calyptra small, deciduous; spores rounded, finely

papillose.

Mielichhoferia contains c. 126 species, primarily of Southern Hemisphere distribution. The major centre of described

species is found in South America.

Shaw and Crum (1984) found that Mielichhoferia is synonymous with and has priority over Haplodontium Hampe, and

species traditionally included in Haplodontium have been transferred to Mielichhoferia. Several species traditionally

included in Mielichhoferia have been transferred to Schizymenium by Shaw (1985). Southern African species are treated

here as Mielichhoferia until their positions in the subfamily Mielichhoferioideae are determined.

Mielichhoferia is similar in many respects to Pohlia, but the genera can be separated on a combination of mostly

sporophytic characters. These characters can be summarized as follows: in Mielichhoferia, the perichaetia are produced on a

short, basal bud or quickly lateral through elongation of a subperichaetial innovation; because the plants are densely packed

the perichaetia are buried in the dense tuft. The peristome is double but appears single as the exostome teeth are rudimentary

and rarely visible above the annulus. The basal membrane and 16 linear segments of the endostome are fragile and quickly

lost on deoperculate capsules. A result of the weak and non-functional condition of the peristome is the rapid loss of spores

from the erect to inclined capsule.

In Pohlia the perichaetia are terminal on an elongated shoot and branching of this shoot is rare. As a result the

perichaetia are exposed and not overgrown by the plant. The peristome is double, consisting of 16 exostome teeth and an

endostome with basal membrane, broad segments and frequently cilia. The peristome is persistent and evident even on very

old capsules. This complete, functional peristome retains spores in the inclined to pendent capsules until suitable conditions

for dispersal exist.

1 Leaves lanceolate to oval-acuminate ,1—2 mm long l.M. bryoides

1 Leaves broadly elliptical or ovate, concave, 0,4— 1,0 mm long 2. M. subnuda

1. Mielichhoferia bryoides (Harv.) Wijk

& Marg. in Taxon 11: 221 (1962); Scott &

Stone, Moss. S. Austr. 295 (1976); Catcheside,

Moss. South Austr. 242 (1980). Type: Cape,

Harvey s.n. (BM, holo. !).

Schizymenium bryoides Harv. in Hooker’s Icon. PI. 3:

202(1840).

Mielichhoferia ecklonii Hornsch. in Linnaea 15: 118

(1841); Broth, in Natiirl. PflFam. 10: 352 (1924); Sim,

Biyo. S. Afr. 316 (1926); Sainsb., N. Zeal. Mosses 254

(1955). Syntypes: Cape, Zeyher s.n.; Lowenriicken, Ecklon

s.n., 24 Aug. 1824; Hottentots Hollands Mountains, Ecklon

s.n., 24 Oct. 1824 (BM!).

? Mielichhoferia rehmannii C. Mull, in Hedwigia 38: 64

(1899); Dix. in Trans. R. Soc. S. Afr. 8: 199 (1920); Broth,

in Natiirl. PflFam. 10: 353 (1924). Type: Cape, Devil’s

Peak, Rehmann 313.

Mielichhoferia transvaaliensis C. Miill. in Hedwigia 38:

64 (1899); Broth, in Natiirl. PflFam. 10: 351 (1924); Sim,

Bryo. S. Afr. 315 (1926). Type: Transvaal, Duivels

Krackler, near Lydenburg, Wilms s.n., Apr. 1887 (G,

holo.!).

Plants small to medium-sized, loosely

caespitose, light green, somewhat glossy; terri-

colous or saxicolous. Stems (2 — )5 — 10 mm

high, infrequently branched; in section angular,

Bryaceae

339

central strand small, inner cortex in 3 rows,

cells large, thin- walled, outer cortical cells in 2

rows, cells of inner row small, incrassate, outer

row stereids, reddish brown. Leaves appressed

to erect dry, erect- spreading wet; lanceolate to

ovate- or oval-acuminate, rarely ovate-acute,

(0,7 — )1 ,0— 1 ,8(— 2,0) mm long; margins

plane, entire below, weakly serrate above.

Costa ending below apex to subpercurrent or

infrequently percurrent; in section rounded,

bulging dorsally, guide cells 2, incrassate, ven-

tral cells in single row, 2-4, incrassate, dorsal

stereid band 2-3 cells thick, frequently with

hydroids below guide cells, dorsal surface cells

small, incrassate. Upper laminal cells mostly

6—15: 1, elongate-rnomboidal to linear-rhom-

boidal (50-)75-150 /urn long, 7-10 pm

wide; basal cells weakly differentiated, gener-

ally shorter, rectangular, occasionally quadrate.

Paroicous or synoicous. Perichaetia on

short, lateral branch, leaves slightly shorter,

triangular. Seta 8-10 (-15) mm long, yellow-

brown; capsules erect to inclined or infre-

quently pendent, ± asymmetrical, weakly

curved, subpyriform, (2,0— )2, 5— 3,0 mm

long, yellow-brown to dark brown, neck

weakly differentiated wet, collapsed dry,

0,5— 1,0 mm long; exothecial cells ± irregular,

rectangular to angular, 1—2 rows at mouth

smaller, quadrate; stomata present on neck,

phaneropore; annulus differentiated; peristome

double, frequently appearing single, exostome

teeth rudimentary, snort ana blunt, rarely visi-

ble above annulus, perforated, ± smooth, yel-

lowish, endostome with low basal membrane,

segments 16, linear, 170-200 pm high, appen-

diculate, frequently anastomosing, ± smooth to

variously papillose, cilia absent; operculum

convex; spores round, 18-22 pm, weakly pap-

illose, light brown. Fig. 96: 1-11.

The species is reported from eastern and southern Af-

rica, Australia, New Zealand and several subantarctic is-

lands. In the Flora area the species is found in shrublands,

grasslands and forest margins of the southwestern, south-

ern and eastern Cape, Natal, Lesotho and the eastern, cen-

tral and northern Transvaal. Map 125.

Vouchers: Cholnoky 940; Crosby & Crosby 9103; Es-

terhuysen 15697; Magill 4298, 4494; Oliver 6730; Van

Rooy 1005.

A lot of variation in plant size and leaf size and length

is accommodated in this circumscription of M. bryoides,

and it may be more correct to consider this a species com-

plex. For example the type of M. bryoides has narrower

leaves (lanceolate) than the types of M. transvaaliensis or

M. ecklonii (± ovate-acuminate). The leaves of M.

bryoides are also longer (to 2 mm) than those of either M.

transvaaliensis (±1,5 mm) or M. ecklonii (0,8- 1,2 mm).

Although the extremes appear distinct, the specimens are

treated as a single species since there is almost a continual

gradation in leaf size and shape, and additional separating

characters were not found.

Van Zanten (1973) included M. ecklonii in the syno-

nymy of M. campylocarpa (Hook. & Amott) Mitt. After

examining several specimens identified by Van Zanten as

M. campylocarpa from Marion Island, as well as several

other species of Mielichhoferia from the subantarctic is-

lands, it was decided not to refer the southern African speci-

mens of M. campylocarpa at this time. The most significant

differences between Marion Island and southern African

specimens are: (1) spore size, (2) leaf width in base, and (3)

capsule stature.

Variation in height of the endostome basal membrane

was found to be quite variable, vide Sim (1926) and Van

Zanten (1973), so the separation of M. transvaaliensis,

based primarily on that character, has not been maintained.

A small group of specimens from Natal and Lesotho

( Cholnoky 91; Dieterlen 790; Magill 4343) differ in having

distinctly quadrate basal cells on most leaves. Although

other specimens have weakly differentiated to rectangular

basal cells, some leaves (especially lower ones) occasio-

nally have quadrate cells. The specimens have therefore,

provisionally been placed here.

2. Mielichhoferia subnuda Sim , Bryo. S.

Afr. 315 (1926). Lectotype: Natal, Mooi River,

Sim 10197 (PRE!, selected here).

Mielichhoferia cholnoky i P. Varde in Revue Bryol.

lichen. 24: 31 (1955). Type: Natal, Drakensberg, Fairy

Glen, Cholnoky 18 (PC, holo.!).

Plants small, loosely caespitose, yellow-

green to light green or brownish; terricolous.

Stems 2-5 mm tall, infrequently branched; in

section round, central strand very small, inner

cortex in 2-3 rows, cells thin- walled, outer

cortex in 1-2 rows, cells with inner walls

340

Bryaceae

Fig. 96. — Mielicbhoferia bryoides (1-11): 1. habit,

x 1; 2. habit, X 8; 3. stem in cross section, x 175; 4-6.

variation in leaf shape, x 50; 7. leaf in cross section, x

350; 8. basal leaf cells (left side), x 175; 9. upper laminal

cells, X 700; 10. leaf apex, x 175; 11. part of capsule

mouth showing cells, annulus and peristome, x 150.

M. subnuda (12-20): 12. habit, x 1; 13. habit, x 8;

14-16. leaves, x 45; 17. basal leaf cells (right side), x

700; 18. upper laminal cells, x 700; 19. propagulum, x

175; 20. part of capsule mouth showing cells, annulus and

peristome, X 150. (1-3 & 6-10, Harvey s.n. (type M.

bryoides); 4, Ecklon s.n. (type M. ecklonii); 5 & 1 1 , Wilms

s.n. (type M. transvaaliensis ); 12-20, Sim 10197).

Bryaceae

341

strongly thickened and reddish. Leaves ap-

pressea dry, patent wet, weakly concave;

broadly elliptical, ovate or shortly oblong-

acute, (0,3 — )0,6 — 1 ,0 mm long; apex sub-

acute, mucronate; margins plane or occasion-

ally recurved below, entire. Costa ending be-

low apex to mucronate or rarely cuspidate; in

section rounded, bulging dorsally, guide cells

not differentiated, ventral cells in single row,

2-4, cells large, thin-walled, dorsal stereid

band 3-4 cells thick, occasionally substereids,

dorsal surface cells incrassate. Upper laminal

cells mostly 3-6: 1, rhomboidal to elongate-

rhomboidal, thin-walled, 37-83 p m long,

9-18 pm wide; basal cells quadrate to short

rectangular, not differentiated in some leaves.

Propagulae bud-like, axillary, 190-300 ;u,m

long.

?Dioicous. Perichaetia lateral through sub-

perichaetial innovation, leaves ovate-lanceolate

to lanceolate. Seta erect, 5-9 mm long, yel-

lowish, red or red-brown; capsules ± erect to

inclined, shortly pyriform or subglobose,

(0,6— )1 ,0- 1 ,5 mm long, yellow-brown, red or

red-brown, neck short, to 0,5 mm; exothecial

cells quadrate to rectangular, rhomboidal or an-

gular, incrassate, 2—3 rows at mouth smaller,

quadrate; stomata numerous on neck and lower

urn, phaneropore; annulus differentiated; peri-

stome double, frequently appearing single, exo-

stome teeth rudimentary, snort and blunt, rarely

visible above annulus, endostome yellowish,

rudimentary as short, smooth basal membrane

at mouth, fragile, quickly broken away, occa-

sionally with short, irregular segments; opercu-

lum convex; spores round, 16-20 pm, mi-

nutely papillose, yellow-brown. Fig. 96:

12-20.

Endemic to southern Africa, the species is collected in

grasslands and shrublands, frequently at high altitude in the

eastern Cape, Lesotho, Orange Free State and Natal. Map

126.

Vouchers: Jacot Guillarmod 284; Magill 4293, 4319;

Rennie & Lambert moss no. 56; Schelpe 7554; Van Rooy

1366.

Mielichhoferia cholnokyi does not differ from this

species. It is based on characters that Sim (1926) did not

fully describe for M. subnuda, e.g. excurrent costa, pyri-

form capsule, operculum shape.

The specimen, Sim 10197, is selected as lectotype be-

cause it conforms with the protologue and is marked ‘type’

in Sim’s hand, on the specimen.

One of the ‘paralectotypes’ of Bryum argenteum var.

rotundifolium (Sim 10232) belongs here (see note under

B. cellulare).

3. BRACHYMENIUM

Brachymenium Schwaegr., Sp. Muse. Suppl. 2: 131 (1824); Broth, in Natiirl. PflFam. 10: 365

(1924); Sim, Bryo. S. Afr. 317 (1926); Andrews in Grout, Moss FI. N. Amer. 2: 207 (1935); Ochi

in J. Fac. Educ. Tottori Univ. 23: 3 (1972); Gangulee, Moss. E. India 928 (1974); Scott & Stone,

Moss. S. Austr. 290 (1976). Lectotype species: B. nepalense Hook, in Schwaegr., fide Ochi

(1972).

Plants small to large, forming dense cushions or turfs, occasionally gregarious, light green

and glossy or yellow-green to green or brownish green; terricolous, saxicolous or corticolous.

Stems erect, sparsely branched, occasionally lower stem densely matted with red-brown tomentum;

in section round, central strand present, inner cortical cells large, thin-walled, outer cortical cells in

1-2 rows, smaller, weakly thickened. Leaves ovate to elliptical or occasionally obovate, equally

spaced or crowded above; margins infrequently weakly bordered, plane to erect, entire or weakly

serrate near apex. Costa percurrent to excurrent as short or long awn; in section with weak dorsal

342

Bryaceae

stereid band. Upper laminal cells firm, rhomboidal to oblong-rhomboidal or sometimes lax, linear-

rhomboidal to fusiform; apical cells occasionally differentiated, hyaline; basal cells quadrate to

rectangular, frequently distinct. Propagulae infrequent, bud-like, axillary.

Dioicous or autoicous. Perichaetia terminal but overgrown by subperichaetial innovations.

Seta elongate, erect; capsules erect to inclined or horizontal, pyriform to cylindrical-pyriform or

clavate, neck differentiated; stomata present on neck, phaneropore; annulus differentiated;

Eeristome double, exostome teeth 16, narrowly triangular, endostome variable, consisting of a

igh basal membrane, segments rudimentary or well developed, cilia rudimentary or absent;

operculum low, conical; calyptra cucullate, smooth, yellowish; spores large or small, greenish.

The genus Brachymenium contains c. 170 species found almost entirely in the tropics and temperate regions of the

Southern Hemisphere. Six species occur in southern Africa; three are endemic to Africa, two are found in Africa and Asia,

and the last also occurs in Australia and South America.

Vegetatively the genus is similar to Bryum, however the small-mouthed, narrow capsule with incomplete endostome

will identify Brachymenium. Most southern African specimens are recognizable on vegetative characters alone, but pro-

blems may be encountered in separating sterile specimens of B. dicranoides, B. leptophyllum and 6. nepalense from similar

Bryum species.

1 Plants small, forming loose turfs on soil or rock, stems sparsely tomentose; spores less than 20 pm in diameter:

2 Plants dull, dark green; upper laminal cells firm, rhomboidal to oblong-rhomboidal, mostly 25-40 /xm

long 1. B. dicranoides

2 Plants glossy, light green; upper laminal cells lax, linear-rhomboidal to fusiform, mostly 75 — 150 /xm

long 2. B. acuminatum

1 Plants larger, mostly corticolous; spores larger than 20 pm in diameter:

3 Plants forming dense cushions, strongly tomentose below; leaves without obvious border, costa generally long-

excurrent as weakly serrate, hyaline awn:

4 Leaf apex hyaline, serrate; spores 40-50 /xm in diameter 3. B. pulchrum

4 Leaf apex chlorophyllous, entire to weakly serrate; spores ± 20 /xm in diameter 4. B. angolense

3 Plants loosely caespitose or gregarious, sparsely tomentose below; marginal cells narrower and ± thickened

forming weak border 1-3 cells wide; costa short excurrent, smooth:

5 Leaf margins revolute from base to near apex; border frequently well defined at apex; endostome segments

absent or rudimentary; spores ± 25 /xm in diameter 5.6. nepalense

5 Leaf margins only reflexed; border weakly differentiated above; endostome segments well developed;

spores 35-37 pm in diameter 6. 8. leptophyllum

1. Brachymenium dicranoides (Horn-

sch.) Jaeg. in Verh. St. Gall, naturw. Ges.

1873-74: 113 (1875); Broth, in Naturl.

PflFam. 10: 366 (1924); Sim, Bryo. S. Afr. 320

(1926); Ochi in J. Fac. Educ. Tottori Univ. 23:

6 (1972). Type: Cape, Table Mountain, Ecklon

s.n. (BM!;H).

Bryum dicranoides Homsch. inLinnaea 15: 134(1841).

Bryum liliputanum C. Miill. in Hedwigia 38: 66 (1899).

Brachymenium liliputanum (C. Miill.) Broth, in Naturl.

PflFam. 10: 366 (1924). Type: Cape, Cape Town, Reh-

mann 241 (PRE!).

? Bryum neesii C. Mull, in Hedwigia 38: 66 (1899).

Brachymenium neesii (C. Miill.) Par., Ind. Bryol. Suppl.

39 (1900); Broth, in Naturl. PflFam. 10: 366 (1924). Type:

Cape, Gnadenthal, Breutel s.n.

Plants small, forming turfs, yellowish

green to brownish green; terricolous or occa-

sionally saxicolous. Stem 2-4(-7) mm long,

sparsely radiculose below; in section cortical

cells in 2-3 rows. Leaves small, ± concave,

appressed to erect dry, erect-spreading wet;

ovate to ovate-lanceolate or elliptical,

(0,3-)0,5-0,8 (-1,0) mm long; apex acute to

acuminate, frequently cuspidate; margins gen-

erally plane, entire. Costa short- to long-excur-

rent, yellow to yellow-brown or brown, awn

smooth, 0,1 -0,3 mm long; in section bulging

dorsally, ventral cells in single layer, large,

thin-walled, dorsal stereid band 2-4 cells

thick, dorsal surface cells smaller than ventral

cells, incrassate. Upper laminal cells mostly

3-5: 1, rhomboidal to oblong-rhomboidal,

(20-)25-40(-65) /xm long, (5— )8— 10(- 13)

/xm wide, occasionally incrassate; marginal

cells frequently rectangular, not forming dis-

tinct border; basal cells weakly differentiated,

quadrate to rectangular. Propagulae infrequent,

bud-like, axillary.

Bryaceae

343

Perichaetial leaves ovate-lanceolate to

lanceolate, costa excurrent. Seta 4-15 mm

long, red-yellow, reddish brown or yellow-

brown; capsule suberect to horizontal, cylindri-

cal-pyriform to pyriform, (0,6-)0,8-l,6 mm

long, yellowish brown or reddish brown, mouth

small, neck wrinkled dry; exothecial cells irre-

gular to subrectangular, incrassate, smaller at

mouth, transversely rectangular; peristome

teeth linear, 250-400 /xm long, hyaline above,

yellow to yellow-brown below, finely papil-

lose, endostome fragile, basal membrane /

height of teeth, segments linear, narrowly per-

forated, just shorter than teeth, cilia absent;

operculum conical; spores round, 10-15 /um,

yellow-brown with age, weakly papillose. Fig.

97:1-11.

This species is known from grasslands and shrublands

of eastern and southern Africa. In the flora area it is infre-

quently collected in the southwestern Cape. Map 127.

Vouchers: Gar side 6272, 6548; Magill 6235.

This species is very similar to Bryum bicolor and

sterile specimens of these two species are difficult to

separate. In general, B. dicranoides is a smaller plant with

shorter, narrower leaves and smaller laminal cells. The leaf

margins are also plane, while in Bryum bicolor they tend to

be re flexed or recurved below. Both species produce axil-

lary, bud-like propagulae, but tubers have only been found

on Bryum specimens. Specimens with sporophytes can be

easily separated on capsule shape and orientation, and peri-

stome architecture.

This species may also be confused with Anomobryum

drakensbergense but differences in habit, leaf shape and

areolation and sporophytic characters will separate the two

species.

Southern African specimens treated by Ochi (1972) as

B. dicranoides and B exile (Doz. & Molk.) Bosch. & Lac.

Map 127. — • Brachymenium dicranoides

▲ Brachymenium angolense

are included here. Differences in awn length and leaf shape

are considered normal variation for the African species.

Examination of specimens of B. exile from India confirms

Gangulee’s (1974) circumscription of that species and in the

absence of any southern African specimens it is excluded

from the Flora.

2. Brachymenium acuminatum Harv. in

Hooker’s Icon. PI. 1: 19 (1836); Broth, in

Natiirl. PflFam. 10: 367 (1924); Ochi in J. Fac.

Educ. Tottori Univ. 23: 4 (1972); Gangulee,

Moss. E. India 4: 950 (1974). Type: Nepal,

Wallich s.n. (BM!).

Brachymenium borgenianum Hampe in Linnaea 38: 211

(1874); Broth, in Naturl. PflFam. 10: 367 (1924); Sim,

Bryo. S. Afr. 320 (1926). Type: Madagascar, Borgen subl.

Mielichhoferia squarrulosa C. Mull, in Hedwigia 38: 64

(1899). Type: Cape, Cape Town, RehmannlW (BM!).

Bryum pallidojulaceum C. Miill. in Hedwigia 38: 67

(1899); Brachymenium pallidojulaceum (C. Miill.) Par.,

Ind. Bryol. Suppl. 39 (1900); Broth, in Naturl. PflFam. 10:

367 (1924). Type: Transvaal, Lake Chriss, Wilms s.n.,

Apr. 1885 (G, holo.!).

Brachymenium lonchopus P. Varde in Arch. Bot. 1: 57

(1928). Type: Rep. Congo, Oubangui, Bambari, Tisserant

s.n. (PC, nolo.).

Plants small, forming loose turfs, light

freen to yellowish green, glossy; terricolous.

terns (2— )5— 10 mm long, sparsely radiculose

below; in section cortical cells in 2-4 rows.

Leaves erect- appressed, little altered dry, con-

cave; ovate to ovate-acuminate or ovate-lanceo-

late, 0,5 -0,8 mm long; apex acute to acumi-

nate; base flat, not concave; margins erect, en-

tire. Costa short-excurrent, mucro 0,1 -0,2 mm

long; in section subround, ventral cells 2, large,

thin- walled, dorsal substereid band 2 cells

thick, dorsal surface cells undifferentiated. Up-

per laminal cells mostly 6—10: 1, fusiform to

linear-rhomboidal, ± sinuolate, (50— )75 — 150

/xm long, 12-18 /xm wide, dorsal surface

weakly thickened; basal cells generally distinct,

quadrate to short-rectangular, 1-2: 1, thin-

walled.

Perichaetial leaves triangular, ± 1 mm

long. Seta (6— )10— 15(— 22) mm long, yellow-

ish; capsule suberect to inclined or almost hori-

zontal, clavate to cylindrical-pyriform or occa-

sionally smaller and pyriform, 2, 5-3, 5 mm

long, yellowish, neck sulcate, to 1 mm long;

exothecial cells subrectangular, walls ± wavy,

1-2 rows of transversely rectangular cells at

mouth; peristome teeth 250—300 /xm long, en-

dostome with basal membrane 175-200 /xm

high, segments rudimentary; spores round,

16-18 /xm, green, weakly papillose. Fig. 97:

12-21.

344

Bryaceae

FIG. 97. — Brachymenium dicranoides (1-11): 1.

habit, x 1; 2. habit, x 5; 3. stem in cross section, x 175; 4.

rhizoid, X 350; 5. leaves, x 70; 6. leaf in proximal cross

section, x 350; 7. basal leaf cells (right side), x 175; 8.

leaf apex (left side), x 175; 9. perichaetial leaf, x 70; 10.

part of capsule mouth showing cells and peristome, x 175;

11. spores, x 700. B. acuminatum (12-21): 12. habit, x

1; 13. habit, x 5; 14 & 15. leaves, x 70; 16. leaf in proxi-

mal cross section, x 350; 17. basal leaf cells (left side), x

175; 18. leaf apex, x 350; 19. perichaetial leaf, x 70; 20.

sporophyte, x 5; 21. part of capsule mouth showing cells,

peristome and spores, x 175. (1 -3 & 6, Magill 6235; 4, 5,

7-9 & 11, Garside 6272; 10, Magill 6548; 12, 13, 18, 20

& 21, Garside 6608; 14, 15 & 17, Brenan M2786; 16,

Brenan M3254; 19, Wager PRE-CHI 161).

Bryaceae

345

Map 128. — • Brachymenium acuminatum

Brachymenium acuminatum has been reported from

eastern and southern Africa, South America, India, south-

east Asia and Australia. In southern Africa the species is

found in grasslands and woodlands of the northern, central,

southern and eastern Transvaal, Swaziland, Natal and Leso-

tho. A few specimens have also been collected in the north-

western and southwestern Cape, eastern Orange Free State,

central South West Africa/Namibia and Botswana. Map

128.

Vouchers: Brenan M2786; Cholnoky 315, 574; Gar-

side 6608; Magill 3424, 3832; Oliver 7162; Pearson 9849;

Van Rooy 995.

The glossy, light green, concave leaves, with lax,

elongate upper leaf cells and generally strongly differen-

tiated basal cells will separate B. acuminatum from other

species of the genus.

Although the leaf is generally concave, above the base,

when cut in cross-section the entire lamina is broadly re-

curved. This is apparently due to a slight thickening of the

dorsal cell facies.

3. Brachymenium pulchrum Hook, in

Hooker, bot. Misc. 1: 136 (1829); Broth, in

Naturl. PflFam. 10: 367 (1924); Sim, Bryo. S.

Afr. 318 (1926); Ochi in J. Fac. Educ. Tottori

Univ. 23: 12 (1972). Type: Cape, Swellendam,

Mundt s.n. (BM).

Brachymenium julaceum Homsch. in Linnaea 15: 133

(1841); Broth, in Naturl. PflFam. 10: 367 (1924). Bryum

koratranum C. Mull., Syn. Muse. 1: 324 (1848). Brachv-

menium koratranum (C. Miill.) C. Mull, in Rehmann,

Musci Austr. Afr. no. 218 (1875) nom. illeg. ; Dixon in

Trans. R. Soc. S. Afr. 8: 200 (1920). Type: Cape, Outni-

qualand, Koratra, Drige s.n.

Plants medium-sized to large, forming

dense cushions, light green to yellow-green, ±

glossy, red-brown below; corticolous. Stems

(5— )10— 35 mm long, irregularly branched, ±

julaceous, lower stem densely matted with red-

brown tomentum; in section cortical cells in

6—8 rows. Leaves crowded, weakly concave,

appressed with reflexed awns wet or dry; ovate

to oval, 0,8— 1,2 mm long; apex acute, hyaline;

margins plane to erect, serrate at apex. Costa

short- to long-excurrent, awn hyaline, sparsely

toothed, (0,2 — )0,5 — 1 ,0 mm long; in section

round, ventral cells 2, in single layer, exposed

walls very thin, inner walls thickened, dorsal

stereid band 2-3 cells thick, dorsal surface

cells undifferentiated or occasionally subste-

reids. Upper laminal cells mostly 3-5: 1,

oblong-rnomboidal to fusiform, ± thickened,

somewhat flexuose, (75-) 87- 100 (- 125) pm

long, 20-25 gm wide; marginal cells above

mid-leaf becoming narrowly fusiform, hyaline

and strongly thickened, several cells wide at

apex; basal cells quadrate, forming distinct

group.

Perichaetial leaves ovate-lanceolate, to 2

mm long. Seta 10-25 mm long, brownish; cap-

sule erect to inclined, pyriform to cylindrical-

pyriform, 3-4 mm long, yellow-brown, neck

differentiated, 1,0- 1,5 mm long; exothecial

cells rectangular to rhomboidal, thin-walled,

1—2 rows at mouth smaller, quadrate; peri-

stome teeth linear, 300 pm high, strongly re-

flexed against capsule wall when dry, endo-

stome consisting of a high basal membrane, to

200 pm high; operculum low-conic; calyptra

cucullate, smooth, 1,7 mm long, yellowish;

spores round, 40-50 pm, green, essentially

smooth. Fig. 98: 1-8.

This species is known from eastern and southern Af-

rica, Madagascar and India. In southern Africa it is found

on bark in woodlands and forests of the northern and eastern

Transvaal, Swaziland, Zululand, Natal, Transkei and the

eastern and southern Cape. Map 129.

Map 129. — •Brachymenium pulchrum

346

Bryaceae

12

Bryaceae

347

Vouchers: Crosby & Crosby 7656, 7827; De Winter

9378; Hilliard & Burtt 10083; Kemp 851; Kluge 1038;

Magill 3747, 5500, 5993; Rankin 25; Schelpe 7553, 7867;

Smook 1419a; Van Rooy 274, 2312; Von Breitenbach 111,

403.

Brachymenium pulchrum forms large, compact cush-

ions on tree trunks and branches. The cushions are held

together by a copious growth of red-brown tomentum. Ex-

posed leaves at the top of stems are generally ± glossy with

costa excurrent as a hyaline awn. The generally strongly

differentiated apical leaf cells will also help to identify the

species.

The closely related species, B. angolense could be

confused with B. pulchrum, however it is somewhat smal-

ler, and lacks the differentiated apical leaf cells (see note

there).

4. Brachymenium angolense ( Welw . &

Duby) Jaeg. in Verh. St Gall, naturw. Ges.

1873—74: 117 (1875); Broth, in Natiirl.

PflFam. 10: 368 (1924); Ochi in J. Fac. Educ.

Tottori Univ. 23: 11 (1972). Type: Angola,

Huilla, Welwitsch 26 (G; H).

Bryum angolense Welw. & Duby in Mem. Soc. Phys.

Hist. nat. Geneve 21: 219 (1870).

Bryum campylotrichum C. Mull, in Hedwigia 38: 65

(1899). Brachymenium campylotrichum (C. Mull.) Broth,

in Natiirl. PflFam. 1: 558 (1903); 10: 367 (1924); Sim,

Bryo. S. Afr. 318 (1926). Type: Transvaal, Drakensberg,

Wilms s.n., 1884 (G).

Plants small to medium-sized, forming

cushions, light green to yellow-green, ±

glossy; corticolous. Stems (2 — )5 — 10 mm long,

densely matted below with red-brown tomen-

tum; in section cortical cells in 4-5 rows.

Leaves crowded, weakly concave, appressed

with flexuose awns wet or dry; ovate to oval,

0,7- 1,0(1, 4) mm long; apex acute, chloro-

phyllous; margins plane, entire to weakly ser-

rate at apex. Costa long-excurrent, awn hya-

line, weakly toothed, 0,6-0, 8 mm long; in sec-

tion bulging dorsally, ventral cells 2, in single

layer, large, thin- walled, dorsal stereid band

4-5 cells thick, frequently with hydroids below

ventral cells, dorsal surface cells substereids or

incrassate. Upper laminal cells mostly 3-4: 1,

oblong-hexagonal to oblong-rhomboidal, thin-

walled, 50— 62(— 75) pm Tong, 15 — 20(— 25)

pm wide; marginal cells smaller, rectangular to

quadrate above, weakly thickened, chlorophyl-

lous; basal cells moderately differentiated, rec-

tangular to quadrate. Propagulae bud-like, axil-

lary or on tomentum.

Sporophyte not known from southern Af-

rica. Fig. 98: 9-15.

This species is known from woodlands and forests of

eastern, central and southern Africa. In the Flora area B.

angolense has been infrequently collected in dry forests of

the northern and eastern Transvaal, Natal and the Caprivi

Strip of South West Africa/Namibia. Map 127.

Vouchers: Magill 3158; Symons 10198; Vahrmeijer

119; Wager 253.

This species is very closely related to B. pulchrum.

Because sporophytes are not known from the Flora area the

two species are separated by size, development of apical

leaf cells and upper laminal cell length. Spore size has also

been reported (Ochi, 1972) useful in separating the species.

5. Brachymenium nepalense Hook, in

Schwaegr., Sp. Muse. Suppl. 2: 131 (1824);

Broth, in Natiirl. PflFam. 10: 369 (1924); Ochi

in J. Fac. Educ. Tottori Univ. 23: 32 (1972);

Gangulee, Moss. E. India 4: 937 (1974). Type:

Nepal, Hooker s.n.

Plants medium-sized, loosely caespitose or

fregarious, green to yellow-green; corticolous.

terns 5-10 mm long, sparsely radiculose be-

low; in section cortical cells in 4—6 rows.

Leaves crowded and larger above, ± comose,

spirally twisted around stem when dry, erect-

spreading wet; oblong to elliptical or obovate,

1,5-2, 2 mm long; apex obtuse, cuspidate; mar-

gins narrowly revolute from base to near apex,

weakly serrate at apex. Costa short -excurrent;

in section round, guide cells 2, small, ventral

cells 2, in single layer, similar to guide cells,

dorsal stereid band strong, 4 cells thick, with

hydroids present below guide cells, dorsal sur-

face cells similar to ventral surface cells, fre-

quently thickened. Upper laminal cells mostly

Fig. 98. — Brachymenium pulchrum (1-8): 1. habit, x 1; 2. habit, x 3,5; 3. leaf, x 35; 4. part of leaf in cross

section, x 350; 5. basal leaf cells (left side), x 70; 6. upper laminal cells (left side), x 175; 7. capsule, x 8,5; 8. part of

capsule mouth showing cells and peristome, x 88. B. angolense (9- 15): 9. habit, x 1; 10. habit, x 3,5; 11. leaf, x 35;

12. leaf in cross section, x 175; 13. basal leaf cells (left side), x 175; 14. upper laminal cells, x 175; 15. propagula, x 70.

B. nepalense ( 16-22): 16. habit, x 1; 17. habit, x 3,5; 18. leaf, x 18; 19. leaf in cross section, x 175; 20. basal leaf cells

(right side), X 175; 21. upper laminal cells (left side), x 175; 22. part of capsule mouth showing cells, peristome and spore,

X 117. B. leptophyllum (23-30): 23. habit, x 1; 24. habit, x 3,5; 25. leaf, x 35; 26. leaf in cross section, x 175; 27.

basal leaf cells (left side), x 175; 28. upper laminal cells (left side), x 175; 29. capsule, x 5; 30. part of capsule mouth

showing cells, peristome and spore, x 88. (1, 2, 4 & 7, Kemp 851; 3, Von Breitenbach 183; 5, Schelpe 7934; 6, Kemp

1514; 8, Smook 2604; 9, 10, 12 & 15, Magill 3300; 11, 13 & 14, Magill 3157; 16 & 18, Crosby & Crosby 7657; 17 & 22,

Von Breitenbach 143; 19, Rehmann 560; 20 & 21 , Magill 4813; 23, 25, 26, 27 & 29, Kluge 1041A; 25, Sim 10196; 28 &

30, Sim 8823).

348

Bryaceae

Map 130. — • Brachymenium nepalense

▲ Brachymenium leptophyllum

2-3: 1, oblong-hexagonal, 37-50 pm long,

15-20 pm wide, thin- walled, weaklv pitted;

marginal cells elongate and thickened, in 2-3

rows forming weak border, generally not visi-

ble because of revolute margins; basal cells

quadrate to rectangular, 1-2: 1, thin-walled,

faintly reddish.

Perichaetial leaves triangular, smaller than

vegetative leaves. Seta 20 mm long, reddish

yellow; capsule erect, cylindrical-pyriform,

2, 5-4,0 mm long, yellow-brown, mouth red-

dish, neck sulcate, 1,0-1, 5 mm long; exo-

thecial cells rectangular, weakly thickened,

smaller at mouth; peristome teeth narrowly tri-

angular, ± 300 /xm high; endostome with high

basal membrane, segments absent; operculum

conical; spores round, 25 /Am, green, papillose.

Fig. 98: 16-22.

The species has been reported from Japan to southeast

Asia, New Guinea, India, central and southern Africa, Ma-

dagascar and Mauritius. In the flora area the species has

been rarely collected in the forests of the eastern and north-

ern Transvaal and Natal. Map 130.

Vouchers: Crosby & Crosby 7657; Magill 4813; Von

Breitenbach 1410.

This species is near the east African species B. rigidum

Broth. & Par. but differs in having the segments of the

endostome rudimentary or absent and more narrowly revo-

lute leaf margins. It is also related to B. leptophyllum, the

two species differ primarily in the curvature of the leaf

margin and strength of the border.

Sterile specimens of B. nepalense could be mistaken

for larger species of Bryum, for example Bryum torques-

cens, but specimens can usually be separated on leaf shape

and curvature of the leaf margins.

6. Brachy menium leptophyllum (C.

Mull.) Jaeg. in Verh. St Gall, naturw. Ges.

1873-74: 111 (1875); Broth. inNaturl. PflFam.

10: 369 (1924); Ochi in J. Fac. Educ. Tottori

Univ. 23: 20 (1972). Lectotype: Ethiopia,

Schimper 451 (H), fide Ochi (1972).

Bryum leptophyllum Bruch & Schimp. ex C. Miill., Syn.

Muse. 1:273(1848).

Brachymenium variable Dix. in Smithson, misc. Colins

69 (8): 2 (1918); Broth, in Natiirl. PflFam. 10: 369 (1924);

Sim, Bryo. S. Afr. 319 (1926). Type: Uganda, Namo-

nyungi, Diimmer 2577 (PRE!).

Plants medium-sized, loosely caespitose or

gregarious, green, ± glossy; corticolous or

occasionally terricolous. Stems 2-5 mm long,

radiculose below; in section cortical cells in 4-6

rows. Leaves appressed and spirally twisted

around stem dry, spreading wet; elliptical,

1,0-1 ,8 mm long, apex acute, fragile, mucro-

nate; margins rerlexed from base to near apex,

entire to weakly serrate at apex. Costa short-

excurrent, mucro smooth, 0, 1-0,5 mm long; in

section bulging dorsally, guide cells 2, ventral

surface cells 2, thin-walled, dorsal stereid band

3 cells thick, reddish, with hydroids below

guide cells, dorsal surface cells weakly differ-

entiated but larger and thin-walled laterally.

Upper laminal cells mostly 2-3: 1, oblong-hex-

agonal, (35-) 50-62 (-75) pm long, 16—20 pm

wide, thin-walled; marginal cells narrower and

slightly thickened, forming very weak border, 1

(-2) cells wide; basal cells short rectangular,

2-3: 1.

Perichaetial leaves ± triangular, smaller

than vegetative leaves. Seta 8-14 mm long, yel-

low-brown; capsule suberect to horizontal, py-

riform to clavate, 2,5-3 ,0 mm long, yellowish

to brownish, neck sunken, to 1 mm long;

exothecial cells irregular with wavy walls, at

mouth regular and quadrate; peristome teeth

narrowly lanceolate, 350 pm high, endostome

with high basal membrane, segments as high as

teeth, cilia rudimentary; operculum short-coni-

cal, reddish; spores round, 33-37 pm, brown-

ish, papillose. Fig. 98: 23-30.

Brachymenium leptophyllum is known from central,

eastern and southern Africa. In the Flora area the species

has been infrequently collected in forests of Natal and the

eastern Transvaal. Map 130.

Vouchers: Kluge 1041a; Sim 10196; Smook 1414.

The species is related to B. rigidum and B. nepalense

but is distinct in its smaller size, weak leaf border and

entire, reflexed margins and frequently eroded leaf apex.

When sterile the species could be mistaken for species of

Bryum, especially Bryum capillare and B. torquescens.

Bryaceae

4. POHLIA

349

Pohlia Hedw., Sp. Muse. 171 (1801); Broth, in Natiirl. PflFam. 1: 546 (1903); Andrews in Grout,

Moss FI. N. Amer. 2: 188 (1935); Sainsb., N. Zeal. Mosses 257 (1955); Gangulee, Moss. E. India

4: 906 (1974); Scott & Stone, Moss. S. Austr. 299 (1976); Smith, Moss FI. Brit. Irel. 363 (1978).

Type species: P. elongata Hedw.

Webera Hedw., Sp. Muse. 168 (1801), nom. illeg. ; Broth, in Natiirl. PflFam. 10: 357 (1924); Sim, Bryo. S. Afr. 321

(1926). Type species: not designated.

Plants small to large, caespitose, occasionally gregarious, green to yellow-green or light

green, occasionally with metallic lustre; terricolous. Stems erect, generally simple, tomentum

sparse; in section angular, central strand large, inner cortical cells in 3-5 rows, large and thin-

walled, slightly smaller and weakly thickened towards outside, outer cortical cells in 1-2 rows,

stereids or substereids, reddish. Leaves ovate to ovate-lanceolate, becoming longer and narrower

toward apex, forming comal tuft; margins unbordered, plane to recurved, entire to denticulate.

Costa percurrent to ending below apex; in section dorsal stereid band weak. Upper laminal cells

elongate-rhomboidal to fusiform or linear-rhomboidal, smooth, becoming slightly broader and

shorter toward base. Propagulae infrequent, axillary.

Dioicous or variably monoicous. Perichaetia terminal on main stem, leaves similar to leaves

of comal tuft. Seta elongate, flexuose; capsule nearly erect to inclined or horizontal, infrequently

pendulous, pyriform to cylindrical-pyriform, neck long or short; peristome double, exostome teeth

16, endostome segments on basal membrane, cilia present, rudimentary or occasionally absent;

operculum short-conical; calyptra small, deciduous; spores subround, finely papillose.

A genus of c. 115 species, Pohlia is found throughout the world, but its species are concentrated in the Northern

Hemisphere. Three of the species found in southern Africa are very widespread in their distributions and have been reported

from every continent. The fourth, P. baronii is presently known only from southern Africa and Madagascar; however, it is

closely related to the widespread Northern Hemisphere, gemmae-producing, species complex of Section Pohliella.

Pohlia is similar in many respects to Mielichhoferia and specimens have been frequently confused, see note under

Mielichhoferia.

1 Leaf base decurrent; propagulae present in leaf axils 1 P baronii

1 Leaf base not decurrent; propagulae absent or rarely in small numbers:

2 Leaves below comul tufts broadly lanceolate; costa generally ending well below apex; plants with metallic lus-

tre 2. P. cruda

2 Leaves below comal tufts narrower, ovate-lanceolate to lanceolate; costa percurrent; plants without distinct metal-

lic lustre:

3 Capsules ± curved, inclined to horizontal, cylindrical-pyriform, neck as long as um; cilia rudimentary or ab-

sent 3. P. elongata

3 Capsules symmetrical, pendent, pyriform, neck short; cilia well developed 4 . P. nutans

1. Pohlia baronii Wijk & Marg. in Taxon

10: 25 (1961). Type: Madagascar, Betsileo,

Besson s.n., Musci Madagasc. no. 30 (PC!).

Webera decurrens Ren. & Card, in Revue Bot. 9: 396

(1891). Webera annotina (Hedw.) Bruch var. decurrens

(Ren. & Card.) Ren. & Card, in Bull. Soc. r. Bot. Belg. 32:

104 (1894); Ren., Prod. FI. Bryol. Madagascar 163 (1897).

Plants small to medium-sized, scattered

and frequently gregarious, yellow-green to light

green, but with obvious red stem and lower

costae; terricolous. Stems 5-20 mm long, ±

elongated, reddish. Leaves ± distant, exposing

red stem, incurved dry, erect-spreading wet;

ovate-acuminate to ovate-lanceolate, (0,7-)

1 ,0- 1 ,4 mm long, occasionally longer and nar-

rower above; apex acuminate; margins decur-

rent, ± plane, entire or serrulate at apex. Costa

percurrent to subpercurrent; in section rounded,

Bulging dorsally, guide cells 2, slightly thick-

ened, ventral cells in single layer of 2-4 cells,

dorsal stereid band small, of 4— 6 cells, red-

dish, dorsal surface cells similar to ventral

cells, large, weakly thickened. Upper laminal

cells mostly 5-9: 1, elongate-rhomboidal to

fusiform, 50-87x10 /xm; basal cells ± rec-

tangular. Propagulae multicellular, gemmate

350

Bryaceae

Fig. 99. — Pohlia baronii: 1. habit, X 1; 2. habit, x

3; 3. leaves, x 70; 4. leaf in cross section, x 175; 5. basal

leaf cells (left side), x 175; 6. leaf cells at upper right

margin, x 700; 7. leaf apex, x 175; 8 & 9. gemmae show-

ing variation in shape, X 145.(1-8, Vorster 892; 9, Magill

4285).

but variable in shape, ovoid to elongate-

vermicular, 180-300 /u,m long, yellow-green

to brownish, ovoid forms with several, short,

incurved leaf primordia, elongated forms fre-

quently twisted, with 1-2, frequently bent,

longer leaf primordia.

Sporophyte unknown. Fig. 99.

Pohlia baronii is presently known from Madagascar

and southern Africa. In the Flora area it is infrequently

collected in grasslands and open shrublands of Natal, Leso-

tho and the eastern, southern and central Transvaal. Map

131.

Vouchers: Cholnoky 873; Sim 10231; Stirton 6759;

Vorster 892, 1795.

Map 131 . — • Pohlia baronii

The plants are recognized by their somewhat elongated

stem and well-spaced leaves. The leaf bases are decurrent

and the decurrencies are generally obvious on detached

leaves.

The production of elongated bud-like propagulae in

leaf axils clearly indicate the relationship of P. baronii to

the propagulae-producing species of section Pohliella of the

Northern Hemisphere. The plants are, in most respects,

similar to Pohlia proligera, differing only in shorter,

broader leaves and fewer propagulae. It seems probable that

P. baronii is only a Southern Hemisphere race of P. proli-

gera.

2. Pohlia cruda (Hedw.) Lindb., Musci

Scand. 18 (1879); Broth, in Naturl. PflFam. 1:

548 (1903); Andrews in Grout, Moss FI. N.

Amer. 2: 192 (1935); Flowers, Moss. Utah 343

(1973); Clarke in Br. Antarct. Surv. Bull. 37:

65 (1973); Smith, Moss FI. Brit. Irel. 367

(1978). Type: Europe.

Mnium crudum Hedw., Sp. Muse. 189 (1801). Webera

cruda (Hedw.) Fuemr. in Flora, Jena 12: 35 (1829); Broth,

in Naturl. PflFam. 10: 359 (1924).

Webera depauperata Sim, Bryo. S. Afr. 323 (1926).

Pohlia depauperata (Sim) Schelpe in Mem. bot. Surv. S.

Afr. 43: 7 (1979). Type: Natal, Giant’s Castle, Symons sub

Sim 10208 (PRE, holo. !).

Plants large, loosely caespitose, yellow-

green to hyaline green or occasionally reddish

green, with distinct metallic lustre; terricolous

or saxicolous. Stems (5—) 10 — 30 mm long,

dark red, infrequently branched, tomentum

sparse, red-brown. Leaves larger above, form-

ing comal tuft, erect-spreading wet, little al-

tered dry; broadly lanceolate to elliptical,

(1,2-) 1 ,8-3,5 mm long, abruptly narrower at

stem apex, spreading; margins ± plane, mi-

nutely serrate to denticulate near apex. Costa

Bryaceae

351

ending below apex or occasionally ± percurrent

on some leaves; in section elliptical, guide cells

2, incrassate, occasionally with 1-2 ventral

stereids, ventral surface cells slightly smaller

than guide cells, weakly thickened, dorsal ster-

eid band small, of 3-4 cells, generally with

hydroids below guide cells, dorsal surface cells

similar to ventral surface cells but more

strongly thickened. Upper laminal cells mostly

8-14: 1, fusiform to linear-rhomboidal, ver-

micular, 80—150x10—12 /xm; basal cells

long-rectangular.

Dioicous, autoicous orparoicous, but spo-

rophytes infrequent. Penchaetia terminal,

leaves linear, to 3 mm long. Seta 10-15 mm

long, ± flexuose, yellow-brown; capsule in-

clined to horizontal, elliptical, 2,0 — 2,5 mm

long, yellow-brown, neck short, inconspicuous;

exothecial cells rectangular to rhomboidal, 2—4

rows at mouth smaller and 1 — 2 transversely

rectangular; stomata at base of urn and on neck,

phaneropore; annulus differentiated; peristome

double, exostome teeth narrowly triangular, to

0,5 mm long, finely papillose, endostome with

basal membrane to A height of teeth, segments

fragile, ± irregular and cleft, not as high as

teeth, cilia 2-3; operculum conic-apiculate;

spores rounded, 25-30 /xm, finely papillose,

light brown. Fig. 100: 1-10.

A widespread species, P. cruda is known from North,

Central and South America, Europe, Asia, southern Africa,

Australia, New Zealand, Antarctica and Oceania. In the

Flora area, the species is restricted to the high grass-heath-

lands in and around Lesotho. Map 132.

Vouchers: Esterhuysen 21622; Magill 4189, 4402,

4592; Van Rooy 1323.

Map 132. — 9 Pohlia cruda

▲ Pohlia nutans

Pohlia cruda is unlikely to be confused with other

species because of a distinct metallic lustre, light to hyaline

green leaves and the dark red stem and lower costa. In

addition the broad stem leaves are distinct from the longer,

narrower comal leaves at the apex.

Sim’s (1926) observations on the peristome of P. de-

pauperata could not be verified since all available capsules

nad been dissected. The type specimen agrees in all other

characters and in distribution to southern African specimens

of P. cruda with complete peristomes.

3. Pohlia elongata Hedw., Sp. Muse. 171

(1801); Broth, in Natiirl. PflFam. 1: 547

(1903); Andrews in Grout, Moss FI. N. Amer.

2: 193 (1935); Flowers, Moss. Utah 346

(1973); Smith, Moss FI. Brit. Irel. 364 (1978).

Type: Germany.

Webera elongata (Hedw.) Schwaegr., Spec. PI. edn 4,5:

48 (1832); Broth, in Natiirl. PflFam. 10: 358 (1924).

Bryum leptoblepharon C. Mull., Syn. Muse. 1: 337

(1848). Pohlia leptoblepharon (C. Mull.) Broth, in Naturl.

PflFam. 1: 547 (1903). Webera leptoblepharon (C. Mull.)

Jaeg. in Verb. St Gall, naturw. Ges. 1873-74: 138 (1875);

Broth, in Naturl. PflFam. 10: 359 (1924); Sim, Bryo. S.

Afr. 324 (1926). Syntypes: Cape, Zwellendam, Ecklon

s.n., Oct. 1826; Grootvaders, Ecklon s.n., Oct. 1826 (both

BM!).

Bryum mielichhoferiacea C. Mull, in Hedwigia 38: 75

(1899). Webera mielichhoferia (C. Mull.) Par., Ind. Bryol.

Suppl. 328 (1900); Broth, in Naturl. PflFam. 10: 359

(1924); Sim, Bryo. S. Afr. 322 (1926). Pohlia mielichhofe-

ria (C. Miill.) Broth, in Naturl. PflFam. 1: 547 (1903).

Type: Cape, Table Mountain, Rehmann s.n., Nov. 1875

(cf. no. 222, BM!; G!; vide Dix. & Gepp, 1923).

Plants small to medium-sized, loosely

caespitose to gregarious, green to yellow-green;

terricolous. Stems 5— 10(— 20) mm long, red-

dish to reddish yellow, generally simple,

sparsely tomentose below. Leaves longer and

narrower above, forming comose tuft, appres-

sed dry, erect-spreading wet; ovate-lanceolate,

1-2 mm long, becoming linear-lanceolate,

1 ,7—3,5 mm long at apex, forming distinct tuft;

apex acuminate; margins generally plane but

reflexed medially in upper leaves, entire below,

generally denticulate at apex. Costa subpercur-

rent to percurrent; in section rounded, bulging

dorsally, guide cells 2—4, occasionally with

1-2 ventral stereid cells, ventral surface cells

large and weakly thickened but smaller than

guide cells, dorsal stereid band 3-4 cells thick,

frequently with hydroids below central guide

cells, dorsal surface cells similar to ventral sur-

face cells but more strongly thickened. Upper

lamina cells mostly 4—10: 1, oblong-rhomboi-

dal to linear, 37 — 100( — 175) x 8-12 p.m,

weakly vermicular; basal cells weakly differen-

tiated, linear to long-rectangular at insertion,

walls straight.

Bryaceae

353

Autoicous. Perichaetia terminal, leaves

ovate-acuminate, to 2 mm long. Seta erect,

10—30 mm long, yellow-brown; capsule gener-

ally ± asymmetrical, curved, inclined to hori-

zontal, cylindrical-pyriform, 4-5 mm long,

smooth, yellow-brown, neck ± distinct, mostly

as long as urn; exothecial cells elongate-rhom-

boidal, weakly thickened, 2-3 rows at capsule

mouth smaller, quadrate; stomata present on

neck, phaneropore; annulus differentiated; peri-

stome double, yellowish, exostome teeth nar-

rowly triangular, finely papillose, endostome

with low basal membrane, to 'A height of teeth,

segments broad, as high as teeth, cilia rudimen-

tary to absent; operculum conic-apiculate;

spores rounded, 18-25 /xm, weakly papillose,

light brown. Fig. 100: 11 — 19.

A widespread species found in North America,

Europe, Asia, Africa and Kerguelen Island. In the Flora

area, P. elongata is collected in grasslands and open shrub-

lands of the southwestern, southern and eastern Cape,

Natal, Lesotho and the eastern, central and northern

Transvaal. Map 133.

Vouchers: Cholnoky 914; Edwards 582; Magill 3780,

4479, 4902, 6065; Venter 4534.

Sporophytes are common on specimens of P. elongata

and are important in separating specimens from other spec-

ies of Pohlia, especially P. nutans. The long, inclined cap-

sules, with the neck as long as the urn, will place specimens

in P. elongata. The cilia of the endostome are variable, but

never as well developed as in P. nutans; see note under that

species.

4. Pohlia nutans (Hedw.) Lindb., Musci

Scand. 18 (1879); Broth, in Naturl. PflFam. 1:

548 (1903); Andrews in Grout, Moss FI. N.

Amer. 2: 194 (1935); Flowers, Moss. Utah 344

(1973); Clarke in Br. Antarct. Surv. Bull. 37:

69 (1973); Smith, Moss FI. Brit. Irel. 367

(1978). Type: Germany.

Webera nutans Hedw., Sp. Muse. 168 (1801); Broth, in

Naturl. PflFam. 10: 360 (1924); Sim, Bryo. S. Afr. 322

(1926).

IBryum ecklonianum C. Mull, in Bot. Ztg 13: 752

(1855); Dixon in Trans. R. Soc. S. Afr. 8: 200 (1920);

Pohlia ecklonianum (C. Mull.) Jaeg. in Verh. St Gall,

naturw. Ges. 1873—74: 138 (1875). Type: Cape, Klein-

riviersberg, Caledon district, Ecklon s.n., Nov. 1825.

Bryum afronutans C. Miill. in Hedwigia 38: 76 (1899).

Webera afronutans (C. Miill.) Par., Ind. Bryol. Suppl. 327

(1900). Type: Cape, Montagu Pass, Rehmann s.n., Oct.

1875 (cf. no. 221, BM!, vide Dixon & Gepp, 1923).

Bryum humidulum Sull. & Lesq. in Proc. Am. Acad.

Arts Sci. 4: 278 (1959). Type: Cape, nr Simonstown, C.

Wright s.n. , 29 Sept. 1853 (FH, holo. !).

Plants small to medium-sized, loosely

caespitose, green to yellow-green; terricolous.

Stems 10—30 mm high, reddish, mostly simple,

sparsely tomentose below. Leaves larger above,

forming comose tuft, erect to erect-spreading

wet or dry, below ovate-lanceolate 1 ,3—2,5

mm long, above linear-lanceolate, 2-3 mm

long; apex acuminate; margins plane to nar-

rowly recurved, entire to denticulate at apex.

Costa percurrent to subpercurrent; in section

bulging dorsally, guide cells 2, incrassate,

occasionally with 1—2 ventral stereids, ventral

surface cells similar to guide cells, dorsal ster-

eid band weak, only a few cells and hy droids

below guide cells, dorsal surface cells similar to

ventral cells, incrassate. Upper laminal cells

mostly 5-8: 1, linear to elongate-hexagonal,

50-75 (-100) x 10-12 /xm, frequently in-

crassate; basal cells rectangular.

Paroicous. Antheridia in axils of comal

leaves; perichaetia terminal, leaves similar to

FIG. 100. — Pohlia cruda ( 1 — 10): 1. habit, x 1; 2. habit, x 3; 3. part of stem in cross section, x 175; 4. leaves, x 35;

5. leaf from stem apex, x 35; 6. leaf in cross section, x 175; 7. basal leaf cells (right side), x 175; 8. laminal cells, x 700;

9. leaf apex, x 175; 10. part of capsule mouth showing cells, peristome and spores, x 60. P. elongata (11-19): 11. habit,

x 1; 12. habit, x 3; 13 & 14. leaves, x 35; 15. leaf in cross section, x 175; 16. basal leaf cells (left side), x 175; 17.

laminal cells, x 700; 18. leaf apex, x 175; 19. part of capsule mouth showing cells and peristome, x 175. P. nutans

(20— 28): 20. habit, x 1; 21. habit, x 3; 22. leaves, x 35; 23. leaf in proximal cross section, x 175; 24. leaf in distal cross

section, x 175; 25. basal leaf cells (left side), x 175; 26. laminal cells, x 700; 27. leaf apex, x 175; 28. part of capsule

mouth showing cells and peristome, X 100. (1-10, Esterhuysen 21621; 1 1 - 19, Edwards 582; 20— 28, Thorne 50374).

354

Bryaceae

comal leaves. Seta flexuose, 10—30 mm long,

brownish; capsule subhorizontal to pendent,

pyriform, 2 mm long, neck short; exothecial

cells irregularly rectangular, thin-walled; 1—2

rows at mouth darker, transversely rectangular;

stomata present on neck, phaneropore; annulus

differentiated; peristome double, yellowish,

exostome teeth narrowly triangular, finely pap-

illose, endostome with high basal membrane,

segments broad, as high as teeth, cilia well de-

veloped, 2-3, elongate; operculum convex-

apiculate; spores rounded, 18—25 /urn, weakly

papillose, brownish. Fig. 100: 20— 28.

A very widespread species, P. nutans is found in North

and South America, Europe, Asia, Africa, Australia, New

Zealand, Antarctica and Oceania. In southern Africa the

species is known from the central, southern and south-

western Cape. Map 132.

Vouchers: Thorne 50374; Wager 439.

A strict definition of P. nutans is used in the Flora.

Plants placed in the species have short-pyriform capsules

that are generally subhorizontal to pendent; the neck of the

capsule is shorter than the urn, and the endostome cilia are

well developed and as long as segments.

Variation in leaf shape and size, leaf cell length and

width, and stature of plants makes separation of sterile

specimens of P. nutans and P. elongata rather arbitrary.

Insufficiently Known Species

Pohlia philonotula (C. Mull.) Broth, in Natiirl.

PflFam. 1: 552 (1903). Basionym: Bryum philonotula C.

Mull, in Hedwigia 38: 76 (1899). Type: Transvaal, Kuilen

near Lydenburg, Wilms s.n., Feb. 1888, Herb. Jack. The

type has not been located. The species was described sterile

and generic placement is uncertain; see note in Sim (1926).

Pohlia pseudophilonotula (C. Mull.) Broth, in Natiirl.

PflFam. 1: 552 (1903). Basionym: Bryum pseudophilono-

tula C. Mull, in Hedwigia 38: 76 (1899). Type: Transvaal,

Lake Chriss, Wilms s.n., Apr. 1885, Herb. Jack. The type

has not been located. The species was described sterile and

generic placement is uncertain; see note in Sim (1926).

Pohlia macleai (Sim) Schelpe in Mem. bot. Surv. S.

Afr. 43: 7 (1979). Webera macleai Sim, Bryo. S. Air. 323

(1926). Type: Cape, Rhenosterberg, MacLea sub Rehmann

548 (BM; BOL; NH!; PRE!). Sim’s (1926) peristome ob-

servations could not be verified. The specimen at PRE

shows only fragments of an endostome remaining and the

NH specimen is sterile. In addition, gametophytically, the

specimens resemble a large specimen of Mielichlwferia

bryoides and the sporophytes of the PRE specimen are from

short basal buds. Dixon in Sim (1926) reported that the

‘London’ specimen is a tall form of P. elongata with the

habit of Mniobryum albicans. It is possible that the collec-

tion (Rehmann 548) was mixed and more duplicates must be

examined to clarify the problem.

5. PLAGIOBRYUM

Plagiobryum Lindb. in Ofvers. K. VetenskAkad. Forh. 19: 606 (1863); Broth, in Natiirl. PflFam.

10: 372 (1924); Andrews in Grout, Moss FI. N. Amer. 2: 209 (1935); Gangulee, Moss. E. India 4:

952 (1974); Smith, Moss FI. Brit. Irel. 381 (1978). Type species: P. zierii (Hedw.) Lindb.; fide

Andrew in Grout, Moss FI. N. Amer. 2: 209 (1935).

Plants mostly small, gregarious; terricolous. Stems erect, little branched; central strand small.

Leaves imbricate, broadly ovate, concave. Costa generally ending below apex. Laminal cells lax,

thin- walled.

Dioicous. Seta short, curved; capsule clavate; peristome double, exostome teeth shorter than

segments, cilia rudimentary; operculum conic; spores large.

A genus of seven species found at high altitudes or latitudes of Europe, North America, Asia, Africa and New

Zealand. This is the first report of the genus from southern Africa, where it is restricted to the afro-alpine region of the

Drakensberg and the high grass-heathland of Lesotho.

The genus could be confused with the hyaline or silvery species of Bryum, but the distinct capsule and peristome will

easily separate the two genera. Sterile specimens are most easily identified by size, leaf shape and colouration, lax leaf cells

and habitat.

Plagiobryum zierii (Hedw.) Lindb. in

Ofvers. K. VetenskAkad. Forh. 19: 606 (1863);

Broth, in Natiirl. PflFam. 10: 373 (1924);

Andrews in Grout, Moss FI. N. Amer. 2: 210

(1935); Nyholm, Moss FI. Fenn. 2: 212 (1954);

Gangulee, Moss. E. India 4: 953 (1974); Smith,

Moss FI. Brit. Irel. 381 (1978). Type: Europe.

Bryum zierii Hedw., Sp. Muse. 182 (1801). Pohlia zierii

(Hedw.) Schwaegr., Spec. PI. edn 5, 5: 76 (1830).

Plants small to medium-sized, gregarious

or loosely tufted, light green to hyaline with

faint reddish tint; terricolous. Stems 4—10 mm

high, julaceous, infrequently branched, sparsely

Bryaceae

355

radiculose below; in section rounded, inner cor-

tical cells in 2-3 rows, thin-walled, hyaline,

outer cortical cells in 2 rows, weakly thickened,

reddish. Leaves imbricate wet or dry, concave;

broadly ovate, 0,6- 1 ,0 mm long, 0,5-0, 7 mm

wide; apex acute, hyaline; margins plane to

erect, entire. Costa reddish, ending below apex

to subpercurrent; in section bulging dorsally,

ventral cells 2—3, in single layer, large, thin-

walled, dorsal stereid band 3-4 cells thick, oc-

casionally cells only weakly thickened, dorsal

surface cells 3-4, slightly smaller then ventral

cells. Upper laminal cells rhomboidal to ob-

long-rhomboidal, 2-4: 1, hyaline, lax, thin-

walled but firmer below; upper marginal cells

short-rectangular, in 1-2 rows, not forming

distinct border; basal cells rectangular, 2-3: 1,

thin- walled, reddish.

Dioicous. Male plants similar, perigonia

terminal or lateral through innovation.

Perichaetia terminal, frequently overgrown by

subperichaetial innovations; leaves triangular to

elliptical, 1,2— 1,5 mm long, reddish to

yellowish. Seta 6-10 mm long, stout, ±

curved, yellow-brown; capsules horizontal to

pendulous, clavate or gibbous, to 5 mm long,

mouth oblique, neck differentiated, 1-2 mm

long; exothecial cells irregularly rectangular,

thickened, smaller and quadrate at mouth; sto-

mata cryptopore, numerous on neck; annulus

differentiated; peristome double, exostome

teeth triangular, 0,3 mm long, weakly papil-

lose, shorter than endostome, endostome seg-

ments narrow above high basal membrane, yel-

lowish, weakly papillose, cilia rudimentary;

operculum conical; spores large, rounded,

6. LEPTOBRYUM

Leptobryum (B.S.G.) Wils., Biyol. Brit. 219 (1855); Broth, in Naturl. PflFam. 10: 373 (1924);

Sim, Biyo. S. Afr. 324 (1926); Gangulee, Moss. E. India 4: 896 (1974); Catcheside, Moss. South

Austr. 246 (1980); Crum & Anderson, Moss. E. N. Amer. 1: 536 (1981). Type species: L.

pyriforme (Hedw.) Wils.; fide Andrews in Grout, Moss FI. N. Amer. 2: 187 (1935).

Plants mostly small, loosely caespitose; terricolous or corticolous. Stems erect, rarely branch-

ing; in section round, central strand large, cells thin-walled, inner cortical cells in 3-4 rows, outer

cortical cells in 1 —2 rows, smaller and incrassate; tubers subterranean or axillary on short rhizoids,

numerous, ovoid, cells large and few. Leaves distant below, comose and larger above, subulate or

setaceous above an ovate base. Costa broad, frequently filling subula, mostly subpercurrent; in

section crescent-shaped, guide cells large. Laminal cells linear.

Synoicous. Seta long; capsule inclined to pendulous, pyriform; stomata phaneropore; annulus

differentiated; peristome double, exostome teeth generally as long as segments, cilia 2-3, basal

membrane high; operculum conic, mammillate; calyptra cucullate; spores round.

A cosmopolitan genus of 6 species. The long, narrow leaves resemble those of Dicranella and some other genera of

the Dicranaceae but the inclined to pendulous, pyriform capsule and peristome characters place plants in Bryaceae.

Map 134. — • Plagiobryum zierii

A Leptobryum pyriforme

30 -40 pm, yellow-brown, papillose. Fig. 101:

1-11.

New to southern Africa, P. zierii is also known from

Europe, Asia, India, northern Africa and North America. In

the Flora area the species is found on soil of cliff faces or

rock crevices in the grass-heathlands of Lesotho and the

Drakensberg escarpment of Natal and at high altitudes in

Natal. Map 134.

Vouchers: Dealt & Killick 98a; Esterhuysen 26172;

Magill 4454 , 4498 , 4506a; McVean 269124; Van Rooy

1325.

Specimens with sporophytes are distinctive and easily

recognised. Sterile specimens may be macroscopically con-

fused with specimens of Bryum cellulare or perhaps larger

specimens of B. argenteum. The shorter, acute apex will

separate this species from the east African species, Plagio-

bruym piliferum P. Varde.

356

Bryaceae

Bryaceae

357

Leptobryum pyriforme (Hedw.) Wils.,

Bryol. Brit. 219 (1855); Broth, in Naturl.

PfCFam. 10: 374 (1924); Sim, Bryo. S. Afr. 325

(1926); Gangulee, Moss. E. India 4: 896

(1974); Catcneside, Moss. South Austr. 246

(1980); Crum & Anderson, Moss. E. N. Amer.

1: 536 (1981). Type: Europe.

Webera pyriformis Hedw., Sp. Muse. 169 (1801).

Plants small to medium-sized, loosely

caespitose, green to yellow-green; terricolous

or corticolous. Stems to 25 mm tall, rarely

branching, brownish or reddish-brown or

purple, occasionally radiculose below; rhizoids

papillose, brownish to red or purple; tubers

ovoid, numerous, subterranean or frequently

axillary on short rhizoids, 120-160 /im long,

cells large and few, brown or reddish brown.

Leaves distant below, comose and larger above,

erect-spreading to widely spreading, frequently

flexuose; linear-lanceolate or ovate- subulate,

0, 5-3,0 mm long below, 3, 5-8,0 mm long

above; apex setaceous or subulate; base

generally ovate, frequently sheathing; margins

plane, entire or denticulate, often bistratose

above; border absent. Costa broad, frequently

filling subula, subpercurrent to percurrent or

aristate, ventral and dorsal superficial cells

smooth to rough; in section crescent-shaped,

laminal insertion medially, guide cells large,

ventral stereid band absent, ventral cells in 1 — 2

rows, dorsal cells in 1—2 rows with scattered

stereids or substereids. Upper laminal cells

linear-rhomboidal or linear-rectangular, thin-

walled, 75—125 ixm long, 5-9 /xm wide,

mostly 10-15: 1; basal cells irregularly linear-

rectangular, frequently brownish or reddish-

brown below.

Synoicous. Perichaetia terminal, leaves

ovate-subulate. Seta 12—25 mm long, yellow-

brown or reddish brown; capsule inclined to

pendulous, pyriform, yellowish brown or

orange, frequently shiny; urn 0, 7-0,9 mm

long, neck generally 1 mm long, sulcate;

exothecial cells irregularly rectangular to quad-

rate, thin-walled, 3-8 rows at mouth smaller;

peristome double, yellowish, exostome teeth,

oblong-acuminate, trabeculate, finely papil-

lose, endostome hyaline or yellowish, segments

keeled, perforated, cilia 2—3, appendiculate,

basal membrane high, minutely papillose; oper-

culum conic, mammillate; calyptra cucullate;

spores round, 12,5 /xm, pale, smooth or finely

papilose. Fig. 101: 12-22.

This species has a cosmopolitan distribution. In

southern Africa Leptobryum pyriforme is infrequently col-

lected on bark or soil in the southern Cape and Natal. Map

134.

Vouchers: Edwards PRE-CH 10396; Schoeman 18;

Sim 10205; Wager 279, 435.

Leptobryum pyriforme will be recognized by the long

subulate leaves above ovate bases, the broad costa with

large guide cells in section, the linear-rectangular laminal

cells, the ovoid tubers with large cells and the wide-

mouthed capsule.

7. ANOMOBRYUM

Anomobryum Schimp. in Syn. Muse. 382 (1860); Broth, in Naturl. PflFam. 10: 371 (1924); Sim,

Bryo. S. Afr. 325 (1926); Gangulee, Moss. E. India 4: 954 (1974); Smith, Moss FI. Brit. Irel. 383

(1978). Type species: A.julaceum (Brid.) Schimp.

Plants small to medium-sized, caespitose, mostly yellowish green, frequently glossy; terricol-

ous or saxicolous. Stems erect, frequently julaceous, branched; in section with central strand;

rhizoids densely papillose. Leaves evenly spaced along stem, about equal in size, imbricate,

concave, mostly ovate; margins generally plane, entire to denticulate above; border absent. Costa

ending below apex to mucronate. Upper laminal cells rhomboidal to linear- vermicular, frequently

incrassate; basal cells broader, quadrate to rectangular.

Fig. 101— Plagiobryum zierii (1-11): 1. habit, X 1;2. habit, x 3,5; 3. leaves, x 70; 4. leaf in cross section, x

175; 5. basal leaf cells (right side), X 175; 6. laminal cells, x 600; 7. leaf apex, x 175; 8. perichaetial leaf, x 70; 9.

sporophyte,. x 3,5; 10. part of capsule mouth showing cells, annulus and peristome, x 140; 11. spore, x 350. Lepto-

bryum pyriforme ( 12—22): 12. habit, x 1; 13. habit, x 3,5; 14. stem in cross section, x 175; 15. leaf, x 35; 16. leaf in

proximal cross section, x 260; 17. leaf in distal cross section, x 260; 18. basal leaf cells (right side), x 175; 19. upper leaf

lamina, x 175; 20. tuber, x 175; 21. sporophyte, x 3,5; 22. part of capsule mouth showing cells and peristome, x 70.

(1-7, Magill 4506a; 8-11, Dealt & Killick 98a; 12-15, 21 & 22, Edwards PRE-CH10396; 16-18, Wager 279; 19 & 20,

Sim 10205).

358

Bryaceae

Bryaceae

359

Dioicous. Perichaetia terminal but quicklv overgrown bv elongation of subperichaetial inno-

vations. Seta elongate; capsule inclined to pendulous, generally pyriform, neck wrinkled when dry;

stomata phaneropore; annulus differentiated; peristome double, exostome teeth trabeculate, endo-

stome segments keeled, broadly perforated, cilia rudimentary or 2-3, basal membrane high; oper-

culum conic, blunt to apiculate; calyptra cucullate; spores round.

The c. 60 species of Anomobryum occur in tropical and temperate regions world- wide. The sporophytic and many

gametophytic characters of Anomobryum are similar to those of Bryum and the genus has occasionally been placed in

synonymy with Bryum or even Pohlia. Anomobryum is recognized here as a separate genus on the basis of its distinct habit

and leaf areolation.

1 Costa ending below apex; margins denticulate above; upper laminal cells linear-rhomboidal to linear- vermicular,

50-140 fim long 1. A.filiforme

1 Costa percurrent; margins entire; upper laminal cells rhomboidal, occasionally vermicular, 35— 63(— 85) /xm long

2. A. drakensbergense

1 . Anomobryum filiforme (Dicks.) Solms

in Rabenh., Bryoth. Eur. 25: 331 (1873);

Broth, in Natiirl. PflFam. 10: 372 (1924);

Gangulee, Moss. E. India 4: 958 (1974); Smith,

Moss FI. Brit. Irel. 383 (1978). Type: Europe.

Bryum filiforme Dicks., PI. Crypt. Brit. 4: 16 (1801).

Bryum promontorii C. Mull, in Hedwigia 38: 69 (1899),

fide Ochi in J. Fac. Educ. Tottori Univ. 23: 38 (1972).

Anomobryum promontorii (C. Mull.) Dix. in Trans. R. Soc.

S. Afr. 8: 201 (1920); Broth, in Natiirl. PflFam. 10: 372

(1924); Sim, Bryo. S. Afr. 325 (1926). Syntypes: Orange

Free State, Witteberg Mtn above Kadziberg. Rehmann 214

(540) (PRE!); Transvaal, Lydenburg, Apr. 1887, Wilms in

Herb. Jack.

Plants small to medium-sized, caespitose,

olivaceous or green or mostly yellowish green

to yellow above, olivaceous or brownish to red-

dish brown below, generally glossy; terricolous

or saxicolous. Stems to 37 mm tall, julaceous,

brown to red-brown, branching by forks or sub-

perichaetial innovations, tomentose below; in

section round, central strand of thin-walled

cells present, inner cortex 2-3 cells wide, thin-

walled, outer cortex 1-2 cells wide, incrassate;

rhizoids densely papillose, brownish to red-

brown. Leaves evenly spaced along stem, about

equal in size, imbricate, concave, ovate or

oblong-acute, (0,5— )0,9— 1 ,6 mm long; apex

acute or obtuse, apiculate, margins plane

or occasionally recurved below, denticulate

above; border absent. Costa ending below

apex, frequently red below; in section ovoid,

ventrally concave, laminal insertion median,

guide cells frequently exposed ventrallv, dorsal

stereid band strong, dorsal surface cells incras-

sate. Upper laminal cells linear-rhomboidal

or linear-vermicular, incrassate, occasionally

itted, (37— )50— 140 pm long, 7-12 ju,m wide;

asal laminal cells frequently reddish, irregu-

larly rectangular.

Dioicous. Perichaetia terminal but quickly

overgrown by innovations, leaves ovate-lanceo-

late or lanceolate. Seta 13-20 mm long, yellow-

ish red or reddish brown; capsule inclined to

pendulous, generally pyriform, yellowish

brown or reddish brown; urn 0, 8-1,1 mm long,

contracted below mouth when dry, neck

0,4-1 ,0 mm long, wrinkled when dry; exothe-

cial cells irregularly rectangular to quadrate,

incrassate, smaller at mouth; stomata present on

neck, phaneropore; peristome teeth yellow to

yellow-brown, orange below, hyaline above,

narrowly oblong-acuminate, trabeculate, finely

papillose, endostome segments keeled, broadly

perforated, cilia 2-3, appendiculate, basal

membrane high, hyaline to yellowish above,

yellow below, finely papillose; operculum

conic, apiculate; calyptra cucullate; spores

13—16 /xm, finely papillose. Fig. 102: 1-11.

This widely distributed species is known from Europe,

Asia, Oceania, Iceland, North and South America and Af-

rica. In southern Africa A. filiforme is frequently collected

Fig. 102. — Anomobryum filiforme (1-11): 1. habit, x 1 ; 2. habit, x 1; 3. habit, x 10; 4. stem in cross section, x

175; 5. leaves, x 40; 6. leaf in proximal cross section, x 420; 7. leaf in distal cross section, x 420; 8. basal leaf cells (left

side), x 175; 9. leaf apex, x 175; 10. part of capsule mouth showing cells and peristome, x 175; 11. spore, x 700.

A. drakensbergense (12-25): 12. habit, x 1; 13. habit, x 3; 14. stem in cross section, x 175; 15. rhizoid, x 350; 16.

leaves, x 35; 17. leaf in proximal cross section, x 350; 18. basal leaf cells (left side), x 175; 19. upper laminal cells, x

350; 20. leaf apex (left side), x 175; 21. perichaetial leaf, x 35; 22. sporophyte, x 5; 23. part of capsule wall showing cells

and stoma, x 350; 24. part of capsule mouth showing cells and peristome, x 175; 25. spore, x 700. (1 & 4-9, Rehmann

214; 2, 3, 10 & 1 1 , Cholnoky 442; 12, 15, 17 & 21 -24, Esterhuysen 34594; 13, 16 & 18-20, Smook 1095; 14, Van Rooy

21).

360

Bryaceae

along the Drakensberg of Natal, Lesotho and Transvaal and

also in the eastern Cape, Natal Midlands, Zululand, eastern

Orange Free State, southern, central, eastern and northern

Transvaal. Map 135.

Vouchers: Cholnoky 127, 461; Hardy 3942; Magill

4328; Oliver 6752; VanRooyU , 341, 1493, 1632.

This species can be recognized by the julaceous, yel-

lowish green, glossy plants and the evenly spaced leaves

with costae ending below the apex, the denticulate upper

margins and linear-vermicular, incrassate upper laminal

cells.

2. Anomobryum drakensbergense Van

Rooy in Bothalia 16: 42 (1986). Type: Natal,

Organ Pipes Pass, Esterhuysen 34594 (PRE,

holo.!; MO; BOL).

Plants small to medium-sized, caespitose,

yellowish green to brownish above, yellow-

brown to brown below; saxicolous or terrico-

lous. Stems to 20 mm tall, branching by forks or

subperichaetial innovations, frequently tomen-

tose below, yellowish green or reddish brown to

brown; in section round, central strand of thin-

walled cells present, inner cortical cells in 2-4

rows, thin-walled, outer cortical cells 1-2 rows,

thin-walled to incrassate. Leaves ± equidistant,

about equal in size or subperichaetial leaves

larger, imbricate, freauently concave, erect

when dry, erect-spreading when wet; shortly

oblong-acute or ovate to ovate-lanceolate,

(0,5— )0,6— 1 ,5 mm long; apex acute; margins

plane or rarely recurved, entire; border absent.

Costa percurrent or occasionally mucronate,

generally yellow, frequently reddish below; in

section subround to round, laminal insertion

ventral, ventral surface cells present, guide

cells incrassate, dorsal stereid band strong, dor-

sal surface cells incrassate. Upper laminal cells

short-rhomboidal to linear-rnomboidal, occa-

sionally vermicular, frequently incrassate,

(25— )35 — 63(— 85) pm long, 7-19 pm wide; ba-

sal laminal cells frequently reddish, quadrate.

Dioicous. Perichaetia terminal but quickly

overgrown by innovations; leaves ovate-lanceo-

late to lanceolate, upper laminal cells vermi-

cular. Seta 10-16 mm long, yellowish red or

reddish brown; capsule pyriform, inclined to

horizontal, yellowish to reddish or brown, fre-

quently contracted below mouth when dry, urn

1,0—1 ,8 mm long, neck 0,5-1, 5 mm long,

wrinkled when dry; exothecial cells irregularly

rectangular to quadrate, incrassate, smaller at

mouth; stomata present on neck, phaneropore;

annulus differentiated; peristome teeth narrowly

oblong-acuminate, frequently blunt and irregu-

lar in outline, 190-300 pm long, bordered, tra-

beculate, yellowish to reddish, frequently hya-

line above, minutely papillose, endostome seg-

ments broad below, tapering above, keeled,

broadly perforated, yellowish to hyaline, cilia

rudimentary, basal membrane high, yellow, mi-

nutely papillose; operculum conic, blunt to mu-

cronate; calyptra cucullate; spores 12-18 fim,

granulose. Fig. 102: 12-25.

The species is known from the Drakensberg of Natal

and mountains of Lesotho where it grows on soil in rock

crevices, from 2 100-3 050 m. Map 136.

Vouchers: Esterhuysen 21623, 35941; Magill 4142a,

4604, 4705; Smook 1089, 1095, 1096a; Van Rooy 21, 1087,

1090.

The species will be recognized by its habit, branching

patterns of the stem, leaf shape and areolation. A.

drakensbergense can be distinguished from A. filiforme by

the percurrent costa, entire leaf margins, shorter and wider

upper laminal cells and reduced peristome.

Map 136. — •Anomobryum drakensbergense

Bryaceae

8. BRYUM

361

Bryum Hedw., Sp. Muse. 178 (1801); Broth, in Natiirl. PflFam. 10: 374 (1924); Sim, Bryo. S.

Afr. 326 (1926); Andrews in Grout, Moss FI. N. Amer. 2: 211 (1935); Sainsb., N. Zeal. Mosses

265 (1955); Gangulee, Moss. E. India 4: 961 (1974); Smith, Moss FI. Brit. Irel. 383 (1978);

Catcheside, Moss. South Austr. 250 (1980); Crum & Anderson, Moss. E. N. Amer. 1: 538 (1981).

Lectotype species: B.argenteum Hedw.; fide Britton, FI. Bermuda 442 (1916).

Plants small to robust, gregarious or caespitose, whitish to silvery green or yellowish green to

green or reddish green to red, occasionally glossy; terricolous to saxicolous, corticolous or humico-

lous. Stems erect, frequently branching by subperichaetial innovations, frequently tomentose be-

low; tubers occasionally produced, on long or short rhizoids, subterranean or axillary; in section

round, central strand present, inner cortical cells in 2-5 rows, thin-walled, outer cortical cells in

1-2 rows, smaller, frequently incrassate or stereids. Leaves orbicular, ovate to lanceolate, ellipti-

cal, oblong to obovate or spathulate, equidistant to crowded above, about equal in size or larger

above; margins plane to recurved, entire to denticulate or dentate above, border present or absent.

Costa ending below apex or percurrent to excurrent as short or long awn; in section bulging

dorsally, dorsal stereid band weak to strong. Upper laminal cells hexagonal, rhomboidal or linear-

rhomboidal, occasionally vermicular; basal cells quadrate to rectangular. Propagulae infrequent,

bud-like, axillary; filamentous gemmae rare, axillai^ or on leaf lamina.

Dioicous, synoicous or autoicous. Perichaetia terminal but frequently overgrown by subperi-

chaetial innovations. Seta elongate, erect; capsules inclined to pendulous, ovoid or oblong-cylin-

drical to pyriform or clavate, straight or curved, neck differentiated; stomata on neck, phaneropore;

annulus differentiated; peristome double, exostome teeth 16, narrowly oblong-acuminate, fre-

quently bordered, trabeculate at back, papillose or rarely striolate below, endostome segments

perforated, cilia absent or 2-4, nodose to appendiculate, basal membrane low or high; operculum

conic, blunt to apiculate or mammillate; calyptra cucullate; spores round.

Bryum is found throughout the world and is one of the largest moss genera with c. 800 species. South America is the

major centre of described species, followed by tropical Africa and Europe. Sterile plants can usually be recognized by the

subperichaetial branching, the relatively broad leaves crowded towards the stem apex with the lower leaves smaller, the

strong costae and the typical smooth, broad, hexagonal to rhomboidal laminal cells. Fertile plants are recognized by their

inclined to pendulous, narrowly pyriform capsules with double, generally well developed peristomes. Some species of

Bryum may be mistaken for Brachymenium, Plagiobryum or Rhodooryum; see notes under those genera.

The genus is difficult to treat taxonomically and the positive results obtained by regional studies and studies on critical

groups (Crundwell & Nyholm, 1964; Ochi, 1970, 1972; Syed, 1973; Smith & Whitehouse, 1978; Mohamed, 1979)

underline the need for more research and a monographic study of Bryum. Many species have recently been reduced to

synonymy while other species with broad morphological variation have been shown to consist of aggregates or complexes

of several species or infraspecific taxa.

The presence or absence, position, size, colour and shape of propagulae, gemmae and rhizoidal gemmae (tubers) have

proved to oe important taxonomic characters. Because of their size and colour, subterranean tubers are difficult to locate and

can easily be overlooked. Care should be taken when studying mixed collections of Bryum since it is often difficult to prove

to which plant the tubers belong. The costal anatomy is also a useful taxonomic character in some instances. Cross-sections

should be taken from the lower third of the leaf as the costa is occasionally weak above.

In addition to the key, species descriptions and illustrations, familiarity with the species is often required to identify

plants. The variability expressed by most of the species makes identification of some specimens, especially sterile ones,

very difficult.

1 Leaf border absent or laminal cells gradually narrowed towards margin:

2 Plants whitish or silvery green; upper laminal cells hyaline 2. B. argenteum

2 Plants variously green, not whitish; laminal cells chlorophyllose:

3 Axillary tubers present, cells protuberant 6. B. nitens

3 Axillary tubers absent:

4 Leaf margins recurved to revolute above:

5 Costa long excurrent 11 .6. caespiticium

5 Costa percurrent to short excurTent 7.6. alpinum

362

Bryaceae

4 Leaf margins plane above:

6 Leaves generally equidistant and about equal in size, occasionally larger above; plants frequently shiny:

7 Plants small; leaves <1 mm long, cells lax 1 B. cellulare

7 Plants medium-sized to large; leaves >1 mm long, cells firm-walled to incrassate 7. 8. alpinum

6 Leaves more crowded towards stem apex or comose, larger above; plants not distinctly shiny:

8 Costa ending below apex to percurrent 7.8. alpinum

8 Costa excurrent:

9 Plants small to medium-sized; leaves erect-spreading when wet, not in rosettes, generally <2 mm

long, apex acute to acuminate:

10 Comal leaves appressed dry, broadly ovate or subtriangular 4.6. radiculosum

10 Comal leaves erect and occasionally twisted dry, ovate-lanceolate, lanceolate, oblong-lanceolate

or elliptical:

1 1 Tubers brownish; leaves ovate to ovate-lanceolate, costa generally long excurrent 3. 6. bicolor

1 1 Tubers reddish; leaves lanceolate, oblong-lanceolate or elliptical, costa mucronate to short ex-

current 5.6. microerythrocarpum

9 Plants medium-sized to large; leaves frequently in rosettes, generally >2 mm long, apex acute to obtuse:

12 Upper laminal cells 25-36 pm long; leaf apex generally obtuse 14.6. viridescens

12 Upper laminal cells 40- 100 /am long; leaf apex generally acute 15.6. canariense

1 Leaf border 1- 12 cells wide, occasionally weak:

13 Leaf margins recurved to re volute above, apex acuminate 11.6. caespiticium

13 Leaf margins plane above, apex generally acute:

14 Leaf border bi- to multistratose:

15 Leaves ovate, ovate-lanceolate or lanceolate, 1 ,2-2,5 mm long; border indistinct, unistratose to bistratose;

margins generally plane 10.8. turbinatum

15 Leaves elliptical to obovate, 2, 4-3, 7 mm long; border distinct, generally multistratose; margins recurved

below 13. 8. donianum

14 Leaf border unistratose:

16 Leaves frequently decurrent 12.8. pseudotriquetrum

16 Leaves not or rarely decurrent:

17 Plants small to medium-sized; leaves spirally twisted around stem; tubers frequently present:

18 Tubers brown; leaves 1,0-1, 8 mm long, border inconspicuous, 1 — 2( — 3) cells wide; mostly ste-

rile 8.8. capillare

18 Tubers red; leaves 2-4 mm long, border conspicuous, 2-3 cells wide; frequently fruiting

9.8. torque see ns

17 Plants medium-sized to robust; leaves not spirally twisted around stem, variously twisted; tubers occasion-

ally persent:

19 Costa in section subround to round, dorsal stereid band well defined:

20 Leaves ovate, ovate-lanceolate or lanceolate; margins generally plane; border indistinct

10. 8. turbinatum

20 Leaves elliptical, oblong, oblong-spathulate or spathulate; margins recurved to revolute below;

border distinct:

21 Upper laminal cells long-rhomboidal, generally 50-80 /am long:

22 Laminal cells gradually narrowed towards border, border 4-8 cells wide; tubers large, mostly

350-550 pm 16.8. erythrocaulon

22 Laminal cells abruptly narrowed at border, border 2-3 cells wide; tubers smaller, mostly

170-280 /xm 9. 8. torque scens

21 Upper laminal cells short-rhomboidal, generally 30—55 /am long:

23 Axillary filamentous gemmae frequently present; leaves generally flat, border 2-6 cells wide

18.8. andicola

23 Filamentous gemmae absent; leaves frequently concave, border 4-12 cells wide

17.8. perlimbatum

19 Costa in section subrectangular, dorsal stereid band not well defined or absent, substereids frequently

present:

Bryaceae

363

24 Leaf margin denticulate to dentate, border conspicuous, 2-5 cells wide; filamentous gemmae

absent 19. B. pycnophyllum

24 Leaf margin strongly dentate to spinose, border inconspicuous, 1-3 cells wide; filamentous

gemmae frequently present on leaf lamina; 20. B. aubertii

1. Bryum cellulare Hook, in Schwaegr.,

Sp. Muse. Suppl. 3(1): 214a (1827); Broth, in

Natiirl. PflFam. 10: 384 (1824); Baitram,

Mosses of the Phillipines 138 (1939); Ochi in J.

Fac. Educ. Tottori Univ. 24: 42 (1973); Gangu-

lee, Moss. E. India 4: 969 (1974). Type: Asia.

Bryum argenteum var. rotundifolium Sim, Bryo. S. Afr.

328 (1926). Type: Natal, Giants Castle, Symons sub Sim

10210 (PRE, lecto. ! , selected here).

Plants small, caespitose, pale green or red-

dish green above, reddish, yellowish or red-

brown to brown below, slightly glossy; saxico-

lous and terricolous. Stems 3-20 mm tall,

simple or branching by subperichaetial innova-

tions, red to brown, tomentose below; rhizoids

reddish brown, papillose, tubers 80-220 pm,

reddish brown. Leaves equidistant, about eaual

in size, erect when dry, erect to erect-spreading

when wet, concave; orbicular or ovate to ob-

long, (0,2-)0,4-0,8(-l,0) mm long; apex

acute to rounded; margins plane, entire; border

absent or rarely 1 cell wide. Costa weak, end-

ing well below apex to percurrent or rarely mu-

cronate, red or reddish brown; in section round,

ventral surface cells generally present, dorsal

cells incrassate to stereids, hydroids absent.

Upper laminal cells lax, hexagonal or rhomboi-

dal to subrectangular, (25 — )32— 100( — 138)

/urn long, (10 — )12 — 28( — 35) gm wide; basal

cells lax, subrectangular to rectangular or qua-

drate.

Dioicous. Perichaetia terminal; leaves ob-

long to oblong-lanceolate, 1,0- 1,8 mm long,

inner leaves smaller, triangular. Seta 7-12 mm

long, yellowish to reddish brown; capsule in-

clined to horizontal, clavate-cylindrical, urn 1 ,3

mm long, neck 0,6 mm long; exothecial cells

thin-walled, irregular in shape, smaller towards

mouth, incrassate at mouth; stomata phanero-

pore; annulus present; peristome double, exo-

stome teeth yellow to orange, hyaline above,

narrowly oblong-acuminate, 0,3 mm long,

striolate below, minutely papillose above, en-

dostome segments linear, narrowly perforated,

ellowish to orange, cilia absent, basal mem-

rane low; operculum conic, rostellate; calyptra

not seen; spores 20-28 pm, finely papillose.

Fig. 103: 1-13.

Bryum cellulare is known from Australia, Oceania,

Japan, southern Asia, India, southern Europe and northern,

central and southern Africa. In the Flora area the species is

infrequently collected in Lesotho, eastern Orange Free

State, Natal, Zululand, southern, central and eastern

Transvaal and South West Africa/Namibia. Map 137.

Vouchers: Cholnoky 163a, 169; Farkas 14b; Oliver

7070; Perold 39, 367; Phelan, Smook & Taylor 57; Van

Rooy 442.

A group of specimens from Transvaal and Zululand

( Cholnoky 163a; Perold 39; Retief et al. 1 138; Wager 243,

1426) including the only fruiting specimen ( Farkas 14b),

have costae ending well below the leaf apices and resemble

B. pocsii Bizot. In the original description of B. pocsii, it is

distinguished from B. cellulare on the basis of the shorter

costa. Variation in the length of the costa, from ending well

below the apex to percurrent, was observed in some speci-

mens ( Cholnoky 169). A narrow, indistinct leaf border is

occasionally produced by southern African plants in this

group. B. pocsii probably represents an undescribed variety

of B. cellulare.

The majority of southern African specimens belong to

a second group with smaller, rounder leaves, smaller lami-

nal cells and percurrent costae. Plants in this group gener-

ally lack leaf borders. Some specimens collected in Lesotho

( Magill 4157, 4218, 4649) nave short, round leaves but

agree in other characters with plants in this second group.

Tne type of B. argenteum var. rotundifolium also belongs to

this group. The specimen, Symons sub Sim 10210 is se-

lected as lectotype because it closely matches the protolo-

gue and the word ‘type’ appears on the original label in

Sim’s handwriting. No fruiting material has been seen but

tubers are occasionally found in specimens belonging to this

group.

Sterile plants of B. cellulare can be recognized by the

evenly spaced, reddish green, glossy, concave leaves with

lax laminal cells, entire and plane margins without a distinct

Bryaceae

365

leaf border and distinctly larger perichaetial leaves. Fruiting

specimens of B. cellulare are easily separated from other

taxa in Bryaceae. Gametophytically the species resembles

B. nitens but the shorter leaves and lamrnal cells, glossy

plants and the absence of axillary tubers with protuberant

cells will distinguish it from that species. B. cellulare may

also be confused with Plagiobryum zierii, but that species

has hyaline leaf apices.

2. Bryum argenteum Hedw., Sp. Muse.

181 (1801); Lawton, Moss FI. Pacific North-

west 165 (1971); Ochi in J. Fac. Educ. Tottori

Univ. 23: 45 (1972); Scott & Stone, Moss. S.

Austr. 273 (1976); Crum & Anderson, Moss.

E. N. Amer. 1: 570 (1981). Type: Europe.

Bryum stellipilum C. Mull, in Hedwigia 38: 66 (1899).

Type: Transvaal, Spitzkop prope Lydenburg, Wilms s.n.,

Feb. & Apr. 1887 (G!).

Bryum capensiargenteum C. Mull, in Hedwigia 38: 67

(1899), fide Ochi in J. Fac. Educ. Tottori Univ. 23: 46

(1972). Type: Cape, Boschberg prope Somerset East, Mac-

Owan s.n. (L, iso.).

Bryum oranicum C. Mull, in Hedwigia 38: 68 (1899).

B. argenteum var. australe Sim, Bryo. S. Afr. 328 (1926).

Type: Orange Free State, Rehmann 260 (PRE, iso. !).

Bryum argenteum var. proliferum Sim, Bryo. S. Afr.

328 (1926). Type: Sim s.n.

Bryum argenteum var. viride Sim, Bryo. S. Afr. 328

(1926). Type: none cited.

Plants small to medium-sized, forming

turfs or cushions, whitish or silvery green to

green above, green below; terricolous or saxi-

colous. Stems julaceous, ( 1,0 -)5 ,0-25,0 mm

tall, branching below, yellowish green or red to

red-brown, frequently radiculose below; rhi-

zoids red-brown, coarsely papillose, tubers ab-

sent. Leaves crowded, occasionally distant be-

low, imbricate wet or dry, concave; broadly

ovate to oval or broadly oblong to obovate,

0,5- 1,5 mm long; apex hyaline, apiculate to

acuminate; margins plane, entire; border ab-

sent. Costa ending below apex to aristate,

weak, frequently reddish below, awn to 0,8 mm

long, hyaline, frequently denticulate; in section

subround, ventral surface cells 2-3, dorsal

stereid band present, stereids in 1—2 rows, dor-

sal surface cells incrassate, guide cells absent,

hydroids few. Upper laminal cells hyaline,

rnomboidal, occasionally flexuose, frequently

thickened at the comers, (25— )37— 75( — 100)

/xm long, 10-17 pm wide, narrowly rectangu-

lar at margin; basal cells frequently reddish,

uadrate to rectangular, extending up margin in

-4 rows. Propagulae occasionally present in

leaf axils.

Dioicous. Perichaetia terminal; leaves ob-

long-acuminate to triangular, 1,0- 1,5 mm

long. Seta 7— 12(— 20) mm long, yellowish or

red to reddish brown; capsule pendulous, ovoid

or pyriform, 1-2 mm long, yellowish to red-

dish yellow or dark red to reddish brown, con-

tracted below mouth when dry, neck short,

occasionally larger than um, frequently

abruptly narrowed towards seta, wrinkleo when

dry; exothecial cells irregularly rectangular, in-

crassate, shorter towards mouth, 2-4 rows at

mouth transversely elongated; peristome dou-

ble, exostome teeth yellowish to dark red or

reddish brown, apex hyaline, narrowly oblong-

acuminate, 0,2-0, 4 mm long, finely papillose,

endostome segments keeled, perforated, cilia

2-3, basal membrane high, minutely papillose,

yellowish; operculum low-conic, blunt to api-

culate; spores 10-15 pm, finely papillose. Fig.

103: 14-20.

Bryum argenteum is cosmopolitan in distribution and

common throughout the Flora area. It is frequently found in

disturbed places and even cities where it grows along paths,

tarred roads, on walls, roofs and other concrete structures.

Map 138.

Vouchers: Kemp 1449; Killick 4215; Kluge 1039; Ma-

gill 3147, 4315, 4920; Smook 1086; Smook & Phelan 671;

Van Rooy 2, 487, 703, 1375; Volk 01 162.

Bryum argenteum is most easily identified by the whit-

ish or silvery green appearance of the julaceous stems and

the broadly ovate, concave leaves with hyaline upper lami-

nal cells.

Considerable variation in plant and leaf size, leaf

shape, acumen length and nerve length is present in south-

ern African plants. For example, the nerve may end below

the apex, extend into the acumen or become long-excurrent

as a toothed awn. A continual gradation in nerve length was

Fig. 103. — Bryum cellulare (1-13): 1. habit, x 1; 2. habit, x 2,5; 3. stem in cross section, x 70; 4. leaves, x 35; 5.

leaf in proximal cross section, x 175; 6. basal leaf cells, x 175; 7. leaf apex, x 175; 8. rhizoid with tuber, x 70; 9.

perichaetial leaf, X 35; 10. sporophyte, X 2,5; 11. part of capsule wall showing cells and stoma, x 175; 12. part of capsule

mouth showing cells and peristome, x 175; 13. spore, x 700. B. argenteum (14—20): 14. habit, x 1; 15. habit, x 2,5; 16.

leaves, x 35; 17. leaf in proximal cross section, x 175; 18. basal leaf cells, x 70; 19. leaf apex, x 175; 20. propagula, x

70. B. bicolor (21-28): 21. habit, x 1; 22. habit, x 2,5; 23. leaf, x 35; 24. leaf in proximal cross section, x 175; 25. basal

leaf cells (left side), X 175; 26. leaf apex, X 175; 27. propagula, X70; 28. rhizoid with tuber, x 70. (1 & 2 0/;ver7070; 3,

4 & 9, Cholnoky 169; 4, 6 & 8, Anderson 15; 5, Perold 367; 7, Schelpe 2088a; 10-13 Farkas 14b; 14, Van Rooy 487; 15,

Anderson PRE-CH13277; 16, Hardy 5340 & Tolken 5587; 17, Oliver 7351; 18, Perold 58, 19, Magill 4315; 20, Filter

PRE-CH12602; 21 & 22, DeWinter 9300; 23-26, Van Rooy 304; 27 , Magill 6325; 28, VanRooyllS.)

366

Bryaceae

observed in southern African plants and the length of the

nerve frequently varies among plants in a population or

even leaves on the same plant.

Plants of B. argenteum with excurrent costae are often

treated as the var. lanatum (P. Beauv.) Hampe and many

southern African specimens were also referred to B. arach-

noideum C. Miill. The varieties described by Sim (1926)

and treated here as synonyms represent different races of B.

argenteum in southern Africa.

Plants with a rounded leaf apex, costa ending well

below the apex and the apiculus short or absent (Van Zinde-

ren Bakker 453; Magill 7065, 7145; Perold 1; Van Rooy 5)

resemble the Angolan B. albopulvinatum C. Mull. These

plants conform well in other characters to B. argenteum and

as the leaf shape and development of the apiculus are vari-

able among southern African plants, they are treated here

provisionally. More specimens from the rest of Africa

should be studied to properly assess the relationship be-

tween B. albopulvinatum and B. argenteum.

3. Bryum bicolor Dicks., PI. Crypt. Brit.

4: 16 (1801); Broth, in Naturl. PflFam. 10: 393

(1924); Ochi in J. Fac. Educ. Tottori Univ. 23:

51 (1972); Flowers, Mosses: Utah and the West

378 (1973); Crum & Anderson, Moss. E. N.

Amer. 1: 564 (1981). Type: Europe.

Bryum rigidicuspis Dix. in S. Afr. J. Sci. 18: 320 (1922);

Broth, in Naturl. PflFam. 11: 530 (1925); Sim, Bryo. S.

Afr. 330 (1926). Syntypes: Natal, Van Reenen Pass, Wager

74; Zimbabwe, Zimbabwe, Sim 8790; Khami Ruins, Sim

8839, 8867 (all PRE!).

Plants small, loosely caespitose to caespi-

tose, yellowish green to green above, brownish

below; terricolous. Stems 3—15 mm tall, simple

or branching by subperichaetial innovations,

occasionally tomentose below; rhizoids reddish

to reddish brown, papillose, tubers 100-360

gm, brown. Leaves comose, erect to erect-

spreading when dry, erect-spreading when wet,

slightly concave; ovate to ovate-lanceolate or

lanceolate, (0,3— )0,5 — 1 ,0(— 1 ,2) mm long;

apex acuminate; margins plane to recurved be-

low, entire to denticulate above; border absent.

Costa strongly aristate, awn 80-340(-480)

/Am long, smooth to denticulate above, yellow-

ish green or reddish; in section round, laminal

insertion ventral, ventral surface cells present,

dorsal stereid band strong, dorsal surface cells

incrassate, hydroids present between guide cells

and stereid band. Upper laminal cells rhomboi-

dal, thin-walled to incrassate, 22—68 /Am long,

7-15 /Am wide; basal cells rectangular to qua-

drate, occasionally reddish below. Propagulae

bud-like, axillary, 180-600 /Am long.

Dioicous. Perichaetia terminal, leaves lan-

ceolate to triangular. Seta 6 — 15( — 22) mm

long, yellowish or red to reddish brown; cap-

sule horizontal to pendulous, ovoid or shortly

oblong-cylindrical, 0,8-1 ,8 mm long, dark red

or reddish brown when mature, slightly con-

tracted below mouth when dry, neck short, fre-

quently broadly rounded to seta, wrinkled when

dry; exothecial cells irregular in shape, incras-

sate, smaller towards mouth; peristome double,

exostome teeth yellow to orange, endostome

segments tapering above, widely perforated, ci-

lia 2—3, appendiculate, basal membrane high,

finely papillose, yellowish; operculum mam-

millate; spores 8-10 /Am, minutely papillose.

Fig. 103:21-28.

The species is almost cosmopolitan in distribution. In

southern Africa B. bicolor is collected on soil in the north-

western, southwestern, southern, eastern, central and north-

ern Cape, Transkei, eastern Orange Free State, Natal, Zulu-

land, southern, central, eastern and northern Transvaal,

South West Africa/Namibia and Botswana. The species is

frequently collected in disturbed places. Map 139.

Bryaceae

367

Vouchers: Anderson PRE-CHI 2832; De Winter 9300;

Garside 6230; Hoffman 44; Smook 895, 2271; Taylor 475;

VanRooy 304, 398, 614, 1985, 2189.

Bryum bicolor is recognized by the ovate to ovate- lan-

ceolate leaves with yellowish, strongly aristate costae, ab-

sence of a leaf border, axillary propagulae and short, thick

capsules with short, rounded necks. Brownish tubers are

frequently produced although specimens with reddish (Re-

tie/ 1239a) and yellowish tubers with larger cells (Cholnoky

706, 708) were also found.

Wilczek & Demaret (1976) and Smith & Whitehouse

(1978) described several species from the B. bicolor com-

lex in Europe. Southern African plants are treated as B.

icolor until a world-wide study can properly assess the

relationships within this complex.

Bryum rigidicuspis has rounded leaf apices and longer

setae but agrees in other characters with B. bicolor. Sterile

plants of B. rigidicuspis may be confused with B. capillare

but the spirally twisted leaves, rounded leaf apices and leaf

areolation will identify B. capillare.

In the absence of tubers or propagulae, immature spec-

imens of B. bicolor may be mistaken for sterile Archidium

Brid. plants, especially A. acanthophyllum Snider. B. bico-

lor can also be confused with sterile plants of small Brachy-

menium species, e.g. B. dicranoides which is a smaller

lant with smaller leaves (see note under that species),

pecimens with broadly ovate leaves and shorter costae

(Garside 6230, 6277) resemble Bryum radiculosum but are

treated here provisionally. For differences between B. bico-

lor and B. radiculosum see note under that species.

4. Bryum radiculosum Brid., Sp. Muse.

3: 18 (1817); Crundwell & Nyholm in Trans.

Brit, biyol. Soc. 4 (4): 603 (1964); Ochi in J.

Fac. Educ. Tottori Univ. 23: 54 (1972). Type:

Italy, Rome, 1806, Herb. Brid. (B, holo. !).

?Bryum subdecursivum C. Mull, in Hedwigia 38; 74

(1899); Broth, in Naturl. PflFam. 10: 394 (1924). Type:

Cape, Cape Town, Rehmann 254 (BM, iso. !).

Bryum subcavifolium Dix. in Sim, Bryo. S. Afr. 332

(1926), fide Ochi in J. Fac. Educ. Tottori Univ. 23: 54

(1972). Type: Cape, Camps Bay, Rehmann 255 (BM,

holo.).

Plants small, loosely caespitose to caespi-

tose, yellowish green or reddish green above,

reddish brown to brown below; terricolous.

Stems 2—10 mm tall, simple or branching by

subperichaetial innovations, occasionally to-

mentose below; rhizoids red-brown or brown-

ish, papillose, tubers 120—240 gm, pale brown

to reddish brown. Leaves comose, smaller be-

low, imbricate above when dry, erect to erect-

spreading wet, concave; broadly ovate to

triangular, 0,5- 1,1 mm long; apex acute to

acuminate; margins plane to frequently re-

curved, entire to denticulate above; border ab-

sent. Costa strong, mucronate to shortly aris-

tate, yellowish to red, occasionally hyaline

above, awn 0,1 -0,3 mm long; in section sub-

round to round, laminal insertion ventral, ven-

tral surface cells present, dorsal stereid band

strong, stereids in 2-4 rows, dorsal surface

cells incrassate, hydroids present between guide

cells and stereid band. Upper laminal cells

rhomboidal, thin-walled to incrassate, (25—)

35-63 /zm long, 10-23 /zm wide; basal cells

quadrate to short-rectangular, frequently thick-

ened at the comers, generally pitted. Gemmae

occasionally present, ovoid, stalked, axillary,

freen to yellowish green or reddish brown to

rown, 80-200 /zm long.

Dioicous. Perichaetia terminal, leaves ob-

long-lanceolate to triangular. Seta 15-18 mm

long, red to reddish brown; capsule pendulous,

ovoid, 1,0— 1,8 mm long, red, neck short,

broadly rounded to seta, wrinkled when dry;

exothecial cells irregular in shape, incrassate,

shorter towards mouth; 2—3 rows at mouth

transversely elongated; peristome double, exo-

stome teeth oblong-acuminate, reddish or

orange, hyaline above, finely papillose, endo-

stome segments tapering above, widely perfo-

rated, cilia 2-3, appendiculate, basal mem-

brane high, yellowish, finely papillose; opercu-

lum convex, mammillate; spores 12-15 /zm,

smooth to weakly papillose. Fig. 104: 1-9.

This species is known from Europe, southeast Asia,

Japan, Oceania, Australia, North America and northern and

southern Africa. In southern Africa B. radiculosum is occa-

sionally collected in the central, southern, southwestern and

northwestern Cape, Transkei and South West Africa/Nami-

bia. Map 140.

Vouchers: Magill 3964, 4023a; Magill & Schelpe

3922; Oliver 345; Perold 462; Schelpe 7739, 7774; Stirton

9097 ; Van Rooy 717.

368

Bryaceae

/,/* v l|

wu & mil

FlG. 104. — Bryum radiculosum (1-9): 1. habit, x 1;

2. habit, x 2,5; 3. leaf, x 35; 4. leaf in proximal cross

section, x 88; 5. basal leaf cells (right side), X 175; 6.

upper laminal cells (left side), x 175; 7. gemmae, x 70; 8.

tuber, x 70; 9. sporophyte x 5. B. microerythrocarpum

(10-18): 10. habit, x 1; 11. habit, x 3,5; 12. leaf, X 35;

13. leaf in proximal cross section, x 175; 14. basal leaf

cells (left side), x 175; 15. upper laminal cells at left mar-

gin, x 175; 16. tuber, X 35; 17. perichaetial leaf, x 35; 18.

sporophyte, x 3,5. B. turbinatum (19-26): 19. habit, x 1;

20. habit, x 3,5; 21. leaf, x 35; 22. leaf in distal cross

section showing multicellular knob at margin, x 175; 23.

leaf in proximal cross section, x 175; 24. basal leaf cells

(leftside), x 175; 25. upper laminal cells (left side), x 175;

26. capsule, x 7. (1, 2, 4 & 5, Oliver 345; 3 & 6, Magill

3922, 7, Magill 4023; 8, Schelpe 7774; 9, Wager 58; 10,

11, 15, 17 & 18, Sim 3894; 12, 14 & 16, Sim 3916; 13, Van

Rooy 628; 19, 20 & 26, Van Rooy 2623; 21 & 24, Wager

s.n. (GRA sheet 188); 23, Magill 4229; 25, Potts 2).

Bryaceae

369

Bryum radiculosum can be mistaken for B. bicolor but

the broader leaves, appressed towards the stem apex when

dry, with stronger recurved margins and shorter excurrent

costae and the axillary gemmae will usually separate B.

radiculosum. This species is generally found in the southern

and western Cape while B. bicolor ranges northward into

the Transvaal. Propagulae similar to those of B. bicolor

were found in some specimens ( Magill 3953, 3962) but the

plants conform well in other characters to B. radiculosum.

A few specimens resembling B. bicolor ( Schelpe 7799,

7819; Oliver 364b; Anderson 21) were difficult to place but

are treated here provisionally. The specimen Rehmann 252

with lanceolate leaves and longer excurrent costae is treated

here because of the axillary gemmae.

The type specimen of B. subcavifolium was not avail-

able to Sim (1926) and has not been located for this study.

The original description by Dixon in Sim (1926) agrees well

with B. radiculosum and we therefore follow Ochi (1972) in

reducing it to synonymy. Ochi (1972) referred B. subdecur-

sivum to Brachymenium exile (Doz. & Molk.) Bosch. &

Lac. The type specimen in BM is sterile and is treated here

on vegetative characters alone. Bryum subdecursivum is not

synonymous with B. argenteum as suggested by Sim

(1926).

In their treatment of the European species of B. eryth-

rocarpum complex Crundwell & Nyholm (1964) did not

report stalked, axillaiy gemmae for this species and no gem-

mae could be found in the type specimen. Additional sepa-

rating characters were not found in southern African plants

bearing gemmae.

5. Bryum microerythrocarpum C. Mull.

& Kindb. in Macoun, Cat. Can. PI. 6: 124

(1892); Broth, in Natiirl. PflFam. 10: 394

(1924); Crundwell & Nyholm in Trans. Br.

bryol. Soc. 4: 622 (1964); Lawton, Moss FI.

Pacific Northwest 170 (1971); Ochi in J. Fac.

Educ. Tottori Univ. 23: 54 (1972); Smith, Moss

FI. Brit. Irel. 425 (1978). Type: Canada.

Bryum zuluense Broth. & Bryhn, Forh. Vid. Selsk.

Christiania 1911: 14 (1911); Broth, in Natiirl. PflFam. 10:

394 (1924). Type: Zululand, Eshowe, Jan. 1909, Bryhn

s.n.(Hl).

Plants small to medium-sized, loosely

caespitose to caespitose, green above, yellow-

ish brown or reddish brown below; terricolous

or humicolous. Stems 3-15 mm tall, simple or

branching by subperichaetial innovations, to-

mentose below; rhizoids reddish or reddish

brown, papillose, tubers (80-)200-300 /urn, red

or red-brown. Leaves larger above, erect and

slightly twisted dry, erect-spreading wet; lan-

ceolate, oblong-lanceolate or elliptical, (0,8-)

1, 0-2,0 (-2,2) mm long; apex acuminate; mar-

gins plane to recurved below, denticulate

above; border generally absent. Costa mucro-

nate to short-excurrent, occasionally reddish

below, awn 0,1 -0,3 mm long; in section sub-

round, lamina inserted ventrally, ventral sur-

face cells present, dorsal stereid band in 1-3

rows, dorsal surface cells incrassate, hydroids

forming large gap between guide cells and ster-

eid band. Upper laminal cells rhomboidal, thin-

walled or occasionally incrassate, frequently

narrower and incrassate towards margin, 40-68

(-98) fim long, 10-18 pcm wide; basal cells

rectangular to quadrate, thin- walled or incras-

sate, occasionally pitted. Propagulae not seen.

Dioicous. Perichaetia terminal, leaves lan-

ceolate to narrowly triangular, margins re-

curved. Seta 12-28 mm long, yellowish to red;

capsule pendulous, oblong-cylindrical or nar-

rowly pyriform, yellowish brown, reddish

brown to red, urn 1,0-2, 2 mm long, neck

wrinkled when dry, frequently curved, 0,8- 1 ,2

mm long; exothecial cells irregular in shape,

incrassate, smaller towards mouth, 1-3 rows at

mouth transversely elongated; peristome dou-

ble, exostome teeth narrowly oblong-acumi-

nate, bordered, yellow, orange or red-brown

below, hyaline above, minutely papillose, en-

dostome segments tapered above, broadly per-

forated, cilia 2-3, nodose to appendiculate, ba-

sal membrane high, yellowish, minutely papil-

lose; operculum conical, mammillate; calyptra

cucullate; spores 11-14 /urn, smooth to mi-

nutely granulose. Fig. 104: 10-18.

The species is found in northern America, Europe,

New Zealand and southern Africa. In the Flora area B.

microerythrocarpum is infrequently collected in the south-

western Cape, Natal and central Transvaal. Map 141 .

Vouchers: Sim PRE-CH3894, 3916; Van Rooy 628.

In comparison with Bryum bicolor and B. radiculo-

sum, this species has longer and narrower capsules. Sterile

specimens are sometimes difficult to distinguish from B.

Map 141. — # Bryum microerythrocarpum

▲ Bryum caespiticium

370

Bryaceae

hm » > »7

FIG. 105. — Bryum alpinum (1-10): 1. habit, x 1; 2.

habit, X 2,5; 3 & 4. leaves, x 35; 5. leaf in proximal cross

section, X 175; 6. basal leaf cells, x 175; 7 & 8. leaf

apices, x 175; 9. tuber, X 50; 10. sporophyte, x 2,5.

B. nitens (11-17): 11. habit, X 1; 12. habit, x 2,5; 13.

leaf, x 35; 14. leaf in proximal cross section, X 175; 15.

basal leaf cells (right side), x 175; 16. leaf apex, x 175;

17. axillary tuber, x 175. (I, Schmitz 781 1; 2, Rankin 126;

3, Cholnoky 464; 4, Jacot Guillarmod 6055; 5, Van Rooy

1116; 6, Magill 4149; 7, Magill 4178; 8, Wager 1510; 9,

Smook 1224; 10, Esterhuysen 15530; 11 & 12, Van Rooy

624; 13, Smook 1821; 14, Stirton 6763; 15, Perold 22; 16,

Sim 8156; 17, Stirton 6764).

Bryaceae

371

bicolor. The lanceolate leaves with short-excurrent costae

and the reddish tubers will help to identify specimens of B.

microerythrocarpum .

6. Bryum nitens Hook., Icon. PI. 1: 19

(1836); Broth, in Natiirl. PflFam. 10: 395

(1924); Ochi in J. Fac. Educ. Tottori Univ. 23:

62 (1972). Type: India.

Plants medium-sized, loosely caespitose to

caespitose, green to yellowish green or reddish

green above, green or yellowish brown to

brown below; terricolous. Stems 5-35 mm tall,

simple or branching by innovations, red to red-

dish brown, scarcely tomentose below; rhizoids

yellowish brown or reddish brown, papillose,

tubers axillary or on long rhizoids, 80-220

pm, cells generally protuberant, yellowish

brown or reddish brown. Leaves distant below,

crowded above to comose, concave, erect to

erect- spreading dry, erect-spreading to wide-

spreading wet; oblong-lanceolate, 1, 2-2,2 mm

long; apex acuminate; margins plane, entire to

denticulate at apex; border absent. Costa weak,

mucronate or occasionally percurrent, occasio-

nally reddish; in section subround to round,

lammal insertion ventral, guide cells occasio-

nally absent, ventral stereid band absent, dorsal

stereid band small, dorsal surface cells incras-

sate. Upper laminal cells long-rhomboidal,

thin-walled, (55— )68— 103(— 125) pm long,

12-18 pm wide; basal cells abruptly broader

below, subrectangular to rectangular or qua-

drate. Propagulae occasionally present in leaf

axils.

Dioicous. Sporophytes not seen. Fig. 105:

11-17.

Bryum nitens is known from India, southern and south-

eastern Asia, Japan, Oceania, Australia, Indian Ocean Is-

lands, Madagascar and Africa. In southern Africa the spe-

cies is occasionally collected on banks of rivers, streams

and disturbed water courses in the northern, central and

eastern Cape, Transkei, Natal, Zululand, central, southern,

eastern and northern Transvaal and South West Africa/

Namibia. Map 142.

Vouchers: Cholnoky 165; De Winter & Giess 6793a;

Perold 22, 369; Smook 1821; Stirton 6764; Van Rooy 624,

957, 1709.

Bryum nitens will be recognized by the distantly

spaced leaves on the lower stem, the concave leaves with

weak costae, long-rhomboidal laminal cells, abruptly

broader basal cells and plane margins. The presence of

numerous axillary tubers with protuberant cells will also

help to identify plants.

Some specimens ( Herman 90 and Van Rooy 741) may

be confused with B. cellulare because of their shorter leaves

Map 142. — • Bryum nitens

▲ Bryum turbinatum

and laminal cells but they agree well in other characters

with B. nitens (See note underB. cellulare).

7. Bryum alpinum Huds. ex With., Syst.

Arr. Brit. PI. edn 4, 3: 824 (1801); Broth, in

Natiirl. PflFam. 10: 396 (1924); Ochi in J. Fac.

Educ. Tottori Univ. 23: 59 (1972); Smith, Moss

FI. Brit. Irel. 414 (1978). Type: Europe.

Bryum afroalpinum Rehm. ex C. Mull, in Hedwigia 38:

73 (1899); Broth, in Natiirl. PflFam. 10: 396 (1924). Syn-

types: Orange Free State, Kadziberg, Rehmann 248, 248B

(PRE!).

Bryum wilmsii C. Mull, in Hedwigia 38: 74 (1899);

Broth, in Natiirl. PflFam. 10: 397 (1924), fide Sim, Bryo.

S. Afr. 332 (1926). Type: Transvaal, Spitzkop, Apr. 1887,

Wilms in Herb. Jack.

Webera revoluta Sim, Bryo. S. Afr. 324 (1926), non

Pohlia revoluta Card. (1909). Pohlia simii Schelpe in

Mem. bot. Surv. S. Afr. 43: 7 (1979). Type: Transvaal,

Houtbosch, Rehmann 566 (PRE, holo. !).

Plants medium-sized to large, caespitose,

red to reddish green or green to yellowish green

above, yellowish brown or reddish brown to

brown below, frequently shiny; terricolous or

saxicolous. Stems 5-55 mm tall, simple or

branching by innovations, occasionally tomen-

tose below; rhizoids red-brown, coarsely papil-

lose, tubers rare, 140 - 300 (- 1000) pm, red to

red-brown. Leaves crowded or occasionally dis-

tant below, generally equidistant, about equal

in size, appressed to erect when dry, erect to

erect-spreading when wet, lanceolate to oblong-

lanceolate, ovate-lanceolate or oblong to ellipti-

cal, frequently concave, 1-3 mm long; apex

acuminate to rounded-obtuse, occasionally

cucullate; margins plane to revolute, entire or

372

Bryaceae

crenulate to denticulate at apex; border absent.

Costa ending below apex to short-excurrent,

reddish or yellowish to brown; in section sub-

round to round, laminal insertion ventral, ven-

tral surface cells incrassate, dorsal stereid band

strong, stereids in 2-4 rows, dorsal surface

cells incrassate or substereids, hydroids present

between stereid band and guide cells. Upper

laminal cells short-rhomboidal to linear-rhom-

boidal or vermicular, occasionally pitted, fre-

quently incrassate, (32 — )37 — 63( — 98) p m

long, (7— )10 — 15( — 20) pm wide; basal cells

rectangular to quadrate, frequently orange or

yellow to brown, frequently pitted, occasion-

ally bulging below, walls frequently thickened.

Dioicous, sporophytes rare. Perichaetia

terminal; leaves lanceolate or triangular,

1,0- 1,4 mm long, cells linear-rhomboidal to

vermicular. Seta 13-40 mm long, red to red-

dish brown; capsule pendulous, clavate or pyri-

form, red to reddish brown, urn 1 - 2 mm long,

neck frequently wrinkled when dry, 0,5 — 1,2

mm long; exothecial cells irregular in shape,

incrassate, smaller towards mouth; peristome

double, exostome teeth narrowly oblong-acu-

minate, yellowish brown below, hyaline above,

minutely papillose, endostome segments ta-

pered above, broadly perforated, cilia 2-3, no-

dose to appendiculate, basal membrane high,

yellowish, minutely papillose; operculum mam-

millate; calyptra cucullate; spores 17-23 pm,

minutely papillose. Fig. 105: 1-10.

Bryum alpinum has a cosmopolitan distribution. In

southern Africa this species is frequently collected on wet

soil and rock in the mountains of Lesotho, Natal and Orange

Free State and occasionally in the southwestern, northwes-

tern, central and eastern Cape, Transkei and southern, cen-

tral, eastern and northern Transvaal. Map 143.

Vouchers: Cholnoky 464; Dealt & Killick 98, 129,

142; Esterhuysen 15530; Magill 4139; Russell 3796; Van

Rooy 11, 318, 674, 1108, 1472.

A broad species concept is employed here to accom-

modate the wide range of variation in leaf morphology dis-

played by southern African plants. It may be more correct to

consider this a species complex. This concept of B. alpinum

is likely to include the North American B. miniatum Lesq.

and the European B. muehlenbeckii B.S.G., B. mildeanum

Jur. and B. gemmiparum De Not.

Plants with lanceolate leaves, acuminate leaf apices,

revolute leaf margins, linear-rhomboidal to vermicular lam-

inal cells, thick cell walls and short excurrent costae, are

frequently red to reddish green in colour. Plants with

broader, oblong-lanceolate or elliptical leaves, rounded-

obtuse leaf apices, generally plane leaf margins, rhomboi-

dal to short-rhomboidal laminal cells with thinner walls and

costae ending below the leaf apices, are green to yellowish

green in colour. Intermediates are frequently found and

variation sometimes occur among leaves of the same plant.

Narrow leaved plants may resemble species of Pohlia

but B. alpinum will be recognized by its mucronate to short

excurrent costa and the crowded leaves which are more or

less evenly spaced along the stem. Plants with narrow

leaves, short-excurrent costae and long-rhomboidal or ver-

micular, incrassate upper laminal cells (Wager PRE-

CH106, 483), resembling the east African B. stenophyllum

Dix. (pers. comm. Ochi, 1985), are treated here provision-

ally.

8. Bryum capillaretfedw., Sp. Muse. 182

(1801); Syed in J. Bryol. 7: 269 (1973); Smith,

Moss FI. Brit. Irel. 401 (1978); Catcheside,

Moss. South Austr. 257 (1980). Type: Europe.

Plants small to medium-sized, caespitose,

green or yellowish green or rarely reddish green

above, brownish or occasionally reddish brown

below; terricolous or saxicolous. Stems

1 — 7( — 15) mm tall, simple or branching by in-

novations, occasionally tomentose below; rhi-

zoids yellowish brown to reddish brown, papil-

lose, tubers abundant, (180-)230-300 (-450)

pm, yellowish brown to brown. Leaves evenly

spaced along stem or crowded above, larger

above, spirally twisted around stem when dry,

erect and slightly bent towards direction of

twisting when wet; ovate to oblong or occasion-

ally obovate-spathulate; (0,8 — )1,0— 1 ,6 mm

long; apex acute to obtuse; margins plane

above, plane to recurved below, entire to denti-

culate above; border generally inconspicuous,

occasionally absent, cells in 1 — 2( — 3) rows,

sinuate. Costa cuspidate to aristate, awn (0,2-)

Bryaceae

373

0,4-0,6(-0,7) mm long, generally smooth,

mostly yellowish, infrequently reddish below;

in section subround to round, laminal insertion

ventral, ventral surface cells differentiated, dor-

sal stereid band strong, stereids in 2-4 rows,

dorsal surface cells incrassate, hydroids present

between stereid band and guide cells. Upper

laminal cells rhomboidal to hexagonal, thin-

walled, (26— )35— 50(— 63) pm long, (12—)

15 — 19( — 23) pm wide; basal cells quadrate to

rectangular, infrequently pitted, infrequently

reddish below. Filamentous gemmae absent.

Dioicous or synoicous. Perichaetia termi-

nal, leaves narrowly lanceolate. Seta 15-25

mm long, yellowish above, reddish below; cap-

sule pendulous, yellowish or brown, cylindrical

to pyriform, urn 1,0- 1,5 mm long, contracted

below mouth when dry, neck wrinkled when

dry, 0,9- 1,4 mm long; exothecial cells irregu-

larly rectangular, incrassate, smaller at mouth;

peristome double, exostome teeth oblong-acu-

minate, finely papillose, endostome segments

tapered above, broadly perforated, cilia 2-3,

appendiculate, basal membrane high, minutely

papillose; operculum conic, apiculate; calyptra

not seen; spores small, 11 — 15 pm, pale yel-

low, smooth to granulose. Fig. 106: 1-8.

Bryum capillare is almost cosmopolitan and is found

throughout the drier northern regions of the Flora area. It is

frequently collected in Swaziland and Transvaal and occa-

sionally in Transkei, Natal, Zululand, northern and eastern

Cape, Botswana and South West Africa/Namibia. Map 144.

Vouchers: Ellis 3114; Magill 3642, 4929; Relief &

Herman 17a; Smook 1461, 2714a; Van Rooy 744, 928,

1909; Van Vuuren 1725.

Bryum capillare can be recognized when dry by leaves

that are spirally twisted around the stem. The leaves are

mostly ovate to oblong in shape, with rounded apices,

inconspicuous and narrow borders, smooth to slightly denti-

culate margins and strongly excurrent costae forming

smooth awns. Brown tubers are produced in abundance and

sporophytes are rare. In some specimens (Magill 5074, Rus-

sell 2669, Van Rooy 709) the leaf border is weak to absent

but the plants conform well in other characters to B. cap-

illare.

The only fruiting specimen collected so far ( Schelpe

2135), differs from sterile plants of B. capillare in being

lax, with reddish green, oblong^spathulate leaves, distantly

spaced along the red stem. The specimen is placed in

B. capillare on the basis of the leaf and upper laminal cell

length, the inconspicuous and narrow leaf borders, the en-

tire to slightly denticulate leaf margins and the rounded leaf

apices. No tubers could be found in the specimen.

For differences between B. capillare and B. torque s-

cens see note under that species.

9. Bryum torquescens Bruch ex De Not. ,

Syll. no. 163 (1838); Bruch, Bryol. Eur. 4: 49

(1839); Sim, Bryo. S. Afr. 334 (1926) p.p.;

Syed in J. Bryol. 7: 307 (1973); Smith, Moss

FI. Brit. Irel. 406 (1978); Catcheside, Moss.

South Austr. 257 (1980). Type: Italy.

Bryum decursivum C. Mull, in Hedwigia 38: 70 (1899),

fide Syed in J. Bryol. 7: 308 (1973). Type: Cape, Cape

Town, Rehmann 246 (BM).

Bryum lonchopyxis C. Miill. in Hedwigia 38: 72 (1899).

Type: Cape, Grahamstown, MacOwan s.n., June 1874

(GRA!).

Bryum porphyreothrix C. Miill. in Hedwigia 38: 70

(1899), fide Syed in J. Bryol. 7: 308 (1973). Type: Cape,

Montagu Pass, Oct. 1875, Rehmann sub 103.

Bryum torquescentulum C. Miill. in Hedwigia 38: 71

(1899) fide Syed in J. Bryol. 7: 308 (1973). Type: Cape,

Port Elizabeth, Ecklon s.n., Feb. 1830.

Bryum torquescentulum var. nutans C. Mull, in Hedwi-

gia 38: 71 (1899). Type: Cape, Groenekloof, Breutel s.n.

Bryum aterrimum C. Miill. ex Sim, Bryo. S. Afr. 334

(1926). Type: Cape, between Knysna and Belvedere, Reh-

mann 235 (PRE!).

?Bryum acuminatum Sim, Bryo. S. Afr. 336 (1926), non

(Hopp. & Homsch.) B.S.G., Bryol. Eur. 4: 91 (1839).

Bryum simii Schelpe in Mem. bot. Surv. S. Afr. 43: 7

(1979). Type: Natal, Cathkin, Owen 15 (PRE, holo. !).

Plants medium-sized, caespitose, green or

reddish green above, brown to red-brown be-

low; corticolous or saxicolous. Stems 1-15

(-20) mm tall, branching by subperichaetial

innovations, innovations to 10 mm long, to-

mentose below; rhizoids red-brown, coarsely

papillose, tubers (140— ) 170 -280 pm, red.

Leaves crowded above to comose, larger above,

twisted to spirally twisted around stem when

374

Bryaceae

FIG. 106. — Bryum capillare (1-8): 1. habit, X 1; 2. habit (dry), x 7; 3. habit (wet), x 10; 4. leaf, X 35; 5. leaf in

proximal cross section, x 175; 6. basal leaf cells (right side), X 175; 7. leaf apex, x 175; 8. tuber, x 70. B. torquescens

(9-15): 9. habit, x 1 ; 10. habit (wet), x 3,5; 11. rhizoid, x 350; 12. leaf, x 30; 13. leaf in proximal cross section, x 175;

14. laminal cells at upper right margin, x 175; 15. rhizoid with tuber, x 88. (1-3 & 8 , Relief & Herman 17a; 4, Ellis 31 14;

5, Magill 4929; 6 & 7, Van Vuuren 1728; 9-1 1 & 14, Crosby & Crosby 8173; 12, Magill 3924; 13, Cholnoky 301; 15, Van

Rooy 821).

Bryaceae

375

dry, patent when wet; elliptical to oblong-spa-

thulate or oblong, (1 ,5 — )2,0 — 3,0( — 3,5) mm

long; apex acute to acuminate; margins plane

above, plane to recurved below, denticulate

above; border ( 1 — )2— 3 cells wide, incrassate.

Costa mucronate to aristate, awn (0,2-)

0,4-0,6(-0,8) mm long, denticulate above,

frequently reddish; in section round to sub-

round, laminal insertion ventral, ventral sur-

face cells present, dorsal stereid band strong,

dorsal surface cells incrassate, hydroids present

between guide cells and stereid band. Upper

laminal cells rhomboidal to hexagonal, (45—)

50 — 75( — 88) pm long, (15 — )16 — 22( — 25) pm

wide; basal laminal cells rectangular to quad-

rate, occasionally pitted, frequently reddish be-

low. Filamentous gemmae absent.

Synoicous, autoicous or dioicous, fre-

quently fruiting. Perichaetia terminal, leaves

lanceolate to triangular. Seta (15 — )20 — 30

(-35) mm long, yellowish or reddish brown;

capsule cemuous to pendulous, cylindrical, red

or reddish brown, slightly contracted below

mouth when dry, urn 1,4— 2,7 mm long, neck

occasionally wrinkled when dry; exothecial

cells shorter above, in longitudinal rows, at

mouth in 1-2 transverse rows; peristome dou-

ble, exostome teeth oblong-acuminate, gener-

ally orange, finely papillose, endostome seg-

ments frequently broad below, abruptly nar-

rowed above, broadly perforated, cilia 3, ap-

pendiculate, basal membrane high, finely

papillose; operculum hemispherical, apiculate;

calyptra cucullate; spores small, 12—14 pm,

pale yellow, smooth to minutely papillose. Fig.

106:9-15.

Bryum torquescens is known from Europe, the Middle

East, Asia, North and South America, Australia, New Zea-

land and Africa. In southern Africa this species occurs

mostly in the Fynbos Biome but also in forests, woodlands

and more arid regions of the southwestern, northwestern,

southern and eastern Cape. This species is also infrequently

collected in the central Cape, eastern Orange Free State,

Natal, Zululand and central, eastern and northern

Transvaal. B. torquescens occasionally grows on bark in

southern Africa but is mostly found on soil or rock. Map

145.

Vouchers: Cholnoky 125, 301; Crosby <4 Crosby

8173; Magill 3677, 6085, 6149, 6280, 6301; Magill &

Schelpe 3947 ; Van Rooy 821.

Bryum torquescens has been treated as a synonym or a

variety of B. capillare by many authors. It can be distin-

guished by the reddish green leaves, mostly oblong-spathu-

late in shape, with conspicuous borders and denticulate

margins and awn. This species also has longer leaves and

upper laminal cells than B. capillare. Numerous red tubers

are generally present on the rhizoids and sporophytes are

frequently produced while B. capillare bears brownish tub-

ers and is mostly sterile. B. torquescens has a southern

distribution in the Flora area while B. capillare occurs in the

drier northern regions.

Some specimens from Natal, the eastern Orange Free

State and Transvaal (Filter PRE-CH 1 3407 ; Sim 3928 , PRE-

CH9500; Van Rooy 575, 1539) consist of smaller plants

with softer leaves, smaller upper laminal cells, weaker leaf

borders, denticulation and costae, and red tubers with pro-

tuberant cells. The type of B. simii ( Owen 15) also has

softer leaves, narrow leaf borders, weak denticulation and

shorter upper laminal cells, but the tubers are reddish brown

without protuberant cells. The specimens are treated here

provisionally.

10. Bryum turbinatum (Hedw.) Turn.,

Muse. Hib. 127 (1804); Broth, in Natiirl.

PflFam. 10: 386 (1924); Lawton, Moss FI.

Pacific Northwest 174 (1971); Flowers, Moss.

Utah 368 (1973); Smith, Moss FI. Brit. Irel.

395 (1978); Crum & Anderson, Moss. E.N.

Amer. 1: 551 (1981). Type: Europe.

Mnium turbinatum Hedw.,Sp. Muse. 191 (1801).

Bryum radicale Rehm. ex Dix. in Sim, Bryo. S. Afr. 333

(1926). Type: Orange Free State, Bethlehem, Rehmann 245

(BM, holo.!).

Plants medium-sized, loosely caespitose or

caespitose, green, yellow-green or reddish

Seen above, brown and frequently shiny be-

w; terricolous. Stems 4-45 mm tall, branch-

ing by subperichaetial innovations, red or red-

brown, frequently tomentose below; rhizoids

red-brown, papillose. Leaves crowded above,

frequently distant below, erect and slightly

twisted dry, erect-spreading to widespreading

wet, generally concave; ovate, ovate-lanceolate

376 Bryaceae

Fig. 107. — Bryum caespiticium (1-7): 1. habit, x 1;

2. habit, x 2,5; 3. leaf, x 18; 4. leaf in proximal cross

section, X 88; 5. basal leaf cells (left side), x 120; 6. leaf

apex, x 120; 7. sporophyte, x 2,5. B. pseudotriquetrum

(8—14): 8. habit, x 1; 9. habit, x 2,5; 10. leaf, x 18; 11.

leaf in cross section, x 175; 12. basal leaf cells (right side),

x 175; 13. laminal cells at upper right margin, x 175; 14.

sporophyte, x 2,5. B. donianum ( 15—20): 15. habit, X 1;

16. habit, x 2,5; 17. leaf, x 35; 18. leaf in proximal cross

section, x 88; 19. basal leaf cells (left side), x 70; 20. leaf

apex, x 175. (1-3, 5 & 7, Russell 3818; 4, Magill 421 1; 6,

Meyer 1046a; 8 & 9, Magill 5548; 10, 11 & 13, Magill

5646; 12, Crosbv & Crosby 7832; 14, Magill 4593; 15, 16

& 1 8-20 , Gar side 665 1 ; 17, Magill 6286).

Bryaceae

377

or lanceolate, (0,8— )1 ,2—2,5 mm long; apex

acuminate; margins rarely decurrent, plane

above, plane or narrowly recurved below, en-

tire or denticulate above; border indistinct, 1-4

cells wide, cells linear- rhomboidal to linear,

incrassate, unistratose to bistratose. Costa

mucronate, cuspidate or short excurrent, awn

30-260 fim long; in section round to subround,

lamina inserted ventrally, ventral surface cells

differentiated, dorsal stereid band strong, ste-

reids in 2-4 rows, dorsal surface cells mcras-

sate, hy droids present between stereid band and

guide cells. Upper laminal cells rhomboidal,

generally thin-walled, 42-88 pm long, 15-28

pm wide; basal cells quadrate to rectangular,

frequently bulging, occasionally reddish below.

Propagulae occasionally present, bud-like, axil-

lary.

Dioicous. Perichaetia terminal, leaves lan-

ceolate or narrowly triangular, margins

recurved below, border indistinct. Seta 15-35

mm long, yellowish brown or reddish brown;

capsule pendulous, shortly pyriform, frequently

contracted below mouth when dry, yellowish

brown to red-brown, urn 0,6—1 ,4 mm long,

neck 0, 8-2,0 mm long, wrinkled when dry;

exothecial cells irregular in shape, incrassate,

smaller towards mouth; peristome double, exo-

stome teeth narrowly oblong-acuminate, yellow

below, hyaline above, minutely papillose, en-

dostome segments tapered above, broadly per-

forated, cilia 2-3, appendiculate, basal mem-

brane high, yellowish, minutely papillose;

operculum conical; calyptra not seen; spores

15-23 pm, pale, minutely granulose. Fig. 104:

19-26.

The species is known from North America, Europe,

Asia, Japan, North and South Africa. In southern Africa, B.

turbinatum is infrequently collected in the central and east-

ern Cape, Orange Free State, Lesotho and Natal. Map 142.

Vouchers: Liebenberg PRE-CH7647; Lyle PRE-

CH9553; Magill 4229, 5907; Potts 2; Russell 3781; Van

Rooy 2623, 2628.

Fruiting plants are easily recognized by the broadly

pyriform capsule, frequently contracted below the mouth

when dry. Vegetatively B. turbinatum can be distinguished

from the closely related B. pseudotriquetrum by the

indistinct, occasionally bistratose leaf borders, plane, rarely

decurrent leaf margins, thinner walled laminal cells and

broader leaf bases. The species may also be confused with

B. caespiticium but that species has a longer excurrent costa

and recurved to revolute leaf margins.

11. Bryum caespiticium Hedw. Sp.

Muse. 180 (1801); Broth, in Natiirl. PflFam.

10: 390 (1924); Lawton, Moss FI. Pacific

Northwest 167 (1971); Ochi in J. Fac. Educ.

Tottori Univ. 23: 87 (1972); Smith, Moss FI.

Brit. Irel. 412 (1978); Catcheside, Moss. South

Austr. 264 (1980); Crum & Anderson, Moss.

E. N. Amer. 1: 558 (1981). Type: Europe.

Plants small to medium-sized, caespitose,

yellowish green or reddish green above, brown-

ish below; terricolous. Stems 5-15 mm tall, fre-

quently branching by subperichaetial innova-

tions, densely tomentose below, red to reddish

brown; rhizoids red-brown, coarsely papillose,

tubers absent. Leaves frequently in successive

comal tufts, smaller below, frequently concave,

imbricate to erect, slightly twisted when dry,

erect to erect-spreading when wet; oblong-lan-

ceolate to ovate-lanceolate or ovate-oblong

(1 ,1— )1 ,5— 2,3(— 2,7) mm long; apex acuminate;

margins generally recurved to re volute, entire

to denticulate at apex; border indistinct. Costa

long-excurrent, awn 0,2-0, 8 mm long, entire to

denticulate above, reddish below; in section

subround to round, laminal insertion ventral,

ventral stereid band absent, ventral surface cells

present, dorsal stereid band strong, stereids in

3-5 rows, dorsal surface cells incrassate, hy-

droids present between guide cells and stereid

band. Upper laminal cells rhomboidal, infre-

quently pitted, (35— )40— 67(— 73) pm long,

(10— )12— 15(— 19) pm wide, narrower towards

margin; basal cells rectangular, frequently red-

dish below, pitted.

Dioicous. Perichaetia terminal, leaves lan-

ceolate to triangular. Seta 13-30 mm long, yel-

lowish to reddish to reddish brown; capsule

pendulous, oblong -cylindrical to clavate, yel-

lowish to yellowish brown or reddish brown,

frequently contracted below mouth when dry,

urn 1-2 mm long, neck wrinkled when dry,

0,5-1 ,2 mm long; exothecial cells irregular rec-

tangular, incrassate, smaller towards mouth,

1-3 rows at mouth transversely elongated; sto-

mata on neck, phaneropore; peristome double,

exostome teeth narrowly oblong-acuminate,

yellowish, hyaline above, finely papillose, en-

dostome segments tapering above, perforated,

cilia 2-3, appendiculate, basal membrane high,

yellowish or hyaline, smooth or minutely papil-

lose; operculum conic, apiculate; calyptra cu-

cullate; spores yellowish brown, 1 5—28 pm,

minutely papillose. Fig. 107: 1-7.

Bryum caespiticium is almost cosmopolitan in distribu-

tion. In southern Africa this species is infreauently collected

at high elevations in and around Lesotho. Map 141.

Vouchers: Magill 4211, 4460, 4527, 4574a; Meyer

1046b, Russell 3818; VanZinderen Bakker 452.

378

Bryaceae

The plants are recognized by their caespitose habit and

the oblong-acuminate leaves with reddish bases, recurved to

re volute margins and long excurrent costae. See note under

B. turbinatum.

12. Bryum pseudotriquetrum (Hedw.)

Gaertn., Meyer & Schreb. in Oek. Techn. FI.

Wetterau 3: 102 (1802); Broth, in Natiirl.

PflFam. 10: 387 (1924); Ochi in J. Fac. Educ.

Tottori Univ. 23: 82 (1972); Gangulee, Moss.

E. India 4: 991 (1974); Scott & Stone, Moss. S.

Austr. 282 (1976); Smith, Moss FI. Brit. Irel.

411 (1978); Crum & Anderson, Moss. E. N.

Amer. 1: 558 (1981). Type: Europe.

Mnium pseudotriquetrum Hedw., Sp. Muse. 190(1801).

Bryum decurrens C. Miill. in Bot. Ztg 13: 751 (1855);

Broth, in Natiirl. PflFam. 10: 386 (1924). Type: Cape,

Winterberg, Ecklon s.n., Jun. 1825 (BM!).

Bryum brachymeniaceum C. Miill. in Hedwigia 38: 71

(1899); Broth, in Natiirl. PflFam. 10: 398 (1924); Sim,

Bryo. S. Afr. 333 (1926). Type: Cape, Rondebosch, at tree

roots, Rehmann 224 (PRE!).

Bryum aulacomnioides C. Mull, in Hedwigia 38: 72

(1899); Broth, in Natiirl. PflFam. 10: 387 (1924); Sim,

Bryo. S. Afr. 331 (1926). Type: Cape, Boschberg, Somer-

set East, MacOwan s.n. , 1878 (GRA!).

Bryum aulacomnioides var. limbatum Sim, Bryo. S. Afr.

331 (1926). Syntypes: Cape, Cape Town, Sim 9534; Natal,

Rosetta Farm, Sim 10,235; Litte Berg, Owen 40; Aller-

thorpe, Sim 10,236; Edgehill, Sim 10,237; Giants Castle,

Symons s.n.; Impolweni Bridge, Sim 10,247; Giants Castle,

Sim 10,245; Mont Aux Sources, Sim 10,246; Transvaal,

Rosehaugh, Sim 8031 (all PRE!).

Plants medium-sized to large, caespitose,

freen to reddish green above, brownish or red-

rown below; terricolous or saxicolous. Stems

(3,0-)18-42 mm tall, frequently densely

tomentose below, red, branching by innova-

tions; rhizoids red-brown, coarsely papillose,

tubers not seen. Leaves crowded above, fre-

quently distant below, twisted to spirally

twisted around stem dry, erect-spreading wet,

elliptical or ovate-lanceolate or ovate, (0,8 — )

1 ,8— 2,5( — 4,2) mm long; apex acuminate;

margins decurrent, frequently red, plane above,

Elane to revolute below, denticulate at apex;

order strong, cells in (2 — )4 — 8 rows, linear,

incrassate, frequently red below. Costa percur-

rent to short-excurrent, yellowish or red; in sec-

tion subround to round, laminal insertion ven-

tral, ventral surface cells differentiated, dorsal

stereid band strong, stereids in 2-4 rows, dor-

sal surface cells incrassate, hydroids present

between stereid band and guide cells. Upper

laminal cells short-rhomboidal to hexagonal,

(22 — )32 — 45(— 68) pm long, ( 10 — )17 — 22

(-25) pm wide, pitted; basal cells rectangular,

frequently red below, pitted. Filamentous

gemmae absent.

Dioicous or synoicous, sporophytes infre-

quent. Perichaetia terminal, leaves lanceolate to

triangular, 0,7- 1,5 mm long, cells incrassate,

elongate towards apex, border weak. Seta

16-32 mm long, yellowish or red-brown; cap-

sule pendulous, straight, clavate, yellowish or

brownish or red-brown, urn 1,2-2, 8 mm long;

neck wrinkled when dry, 0,9 -2,0 mm long;

exothecial cells irregular rectangular, incras-

sate, shorter above, in 1-2 transverse rows at

mouth; peristome double, exostome teeth yel-

lowish or orange, minutely papillose, endo-

stome segments tapered above, broadly perfo-

rated, cilia 2-4, appendiculate, basal mem-

brane high, smooth to minutely papillose; oper-

culum conical, apiculate; calyptra not seen;

spores 12-21 pm, smooth to granulose. Fig.

107:8-14.

This cosmopolitan species occurs at higher altitudes in

the southwestern and eastern Cape, Natal, Lesotho, eastern

Orange Free State and southern, central and eastern

Transvaal. Map 146.

Vouchers: Arnold 1270; Deall & Killick 72, 106,

115a; Magill 4145, 4593, 5646; Smook 886; Van Rooy

466, 1378.

B. pseudotriquetrum is most easily identified by the

reddish green, generally decurrent and more or less dis-

tantly spaced leaves along the red stem. The leaves are

mostly ovate or elliptical in shape, with a strong border and

short-excurrent, reddish costa.

Ochi (1972) reported the specimen Rehmann 247 as B.

turbinatum but the Rehmann specimen in PRE belongs

here. For differences between B. pseudotriquetrum and B.

turbinatum see note under B. turbinatum.

Bryaceae

379

13. Bryum donianum Grev. in Trans.

Linn. Soc. Lond. 15: 345 (1827); Broth, in

Natiirl. PflFam. 10: 399 (1924); Ochi in J. Fac.

Educ. Tottori Univ. 23: 94 (1972); Smith, Moss

FI. Brit. Irel. 401 (1978). Type: Europe.

Plants medium-sized to large, caespitose,

green or reddish green above, brown to red-

brown below; terricolous. Stems 5-15 mm tall,

branching by subperichaetial innovations,

tomentose below; rhizoids red-brown, coarsely

papillose, tubers absent. Leaves comose, larger

above, erect and frequently twisted dry, erect-

spreading wet; elliptical or obovate or obovate-

lanceolate, 2,4-3, 7 mm long; apex acuminate;

margins plane above, recurved below, obscure-

ly denticulate above; border strong, cells in

2-4 rows, incrassate or substereids, bi- to mul-

tistratose, yellowish or occasionally reddish, in

section forming multicellular knob. Costa

shortly excurrent, yellowish or reddish; in sec-

tion subround to round, laminal insertion ven-

tral, ventral surface cells present, dorsal stereid

band strong, stereids in 3-5 rows, dorsal sur-

face cells incrassate, hydro ids present between

stereid band and guide cells. Upper laminal

cells rhomboidal, frequently pitted, in

conspicuous rows, (27— )42— 55(— 62) gm

long, (11 — )15 — 20( — 25) pm wide; basal cells

rectangular, pitted, pink to reddish below. Fila-

mentous gemmae absent.

Dioicous. Perichaetia terminal; leaves lan-

ceolate to triangular, costa aristate. Seta 20-38

mm long, yellowish to red or reddish brown;

capsule pendulous, straight or slightly curved,

cylindrical or pyriform, contracted below

mouth when dry, yellowish to red-brown, urn

2-3 mm long, neck wrinkled when dry, 1-3

mm long; exothecial cells irregular in shape,

incrassate, shorter above; peristome double,

exostome teeth oblong-acuminate, yellow to

orange, minutely papillose, endostome seg-

ments tapered above, broadly perforated, cilia

2-3, appendiculate, basal membrane high, mi-

nutely papillose, operculum mammillate; calyp-

tra not seen; spores round, 12-14 pm, pale

yellow, essentially smooth. Fig. 107: 15-20.

New to southern Africa, B. donianum is also known

from Europe, North Africa, the Middle East, eastern Asia

and Macaronesia. In the Flora area only three collections

have been made in the southwestern Cape. Map 147.

Vouchers: Garside 6651, 6654; Magill 6286.

This species could be confused with B. torquescens or

B. pseudotriquetrum but the strong, distinct, bi- to multi-

stratose border clearly separates it.

Map 147. — • Bryum viridescens

▲ Bryum donianum

14. Bryum viridescens Welw. & Dub.,

Mem. Soc. Phys. Hist. Nat. Geneve 21: 218

(1870); Broth, in Natiirl. PflFam. 10: 400

(1924); Ochi in J. Fac. Educ. Tottori Univ. 23:

96 (1972). Type: Angola, Huilla, Welwitsch 28

(BM).

Bryum polytrichoideum sensu Sim, Bryo. S. Afr. 338

(1926).

Plants medium-sized to large, solitary or

loosely caespitose, green above, reddish brown

or yellowish brown below; terricolous or corti-

colous. Stems 5-35 mm tall, frequently

branching by subperichaetial innovations, inno-

vations to 15 mm tall, frequently tomentose;

rhizoids reddish brown, papillose, tubers

(200— )260-600(-800) ^im, reddish brown or

yellowish brown. Leaves comose, frequently in

successive comal tufts, smaller below, ap-

pressed dry, erect-spreading and generally in a

rosette wet; broadly oblong to obovate or infre-

quently spathulate, (1,6 — )2,0— 3 ,0( —3,8) mm

long; apex obtuse or rarely acute; lamina fre-

quently indexed above; margins plane above,

recurved in lower M to 3/», entire to crenulate to

denticulate; border absent. Costa strong, aris-

tate, awn 180-360 pm long, smooth to denti-

culate above; in section subround to round, ven-

tral surface cells present, dorsal stereid band

strong, stereids in 3-5 rows, dorsal surface

cells incrassate to substereid, hydroids present

between guide cells and stereid band. Upper

laminal cells hexagonal, pitted, 17-25 /xm

long, (12 — )15 — 23 pm wide; basal cells rec-

tangular to quadrate, pitted.

380

Bryaceae

Fig. 108. — Bryum viridescens (1-12): 1. habit (dry),

x 2; 2. habit (wet), x2,5 ; 3. part of stem in cross section,

x 175; 4. rhizoid, x 175; 5. leaf, x 18; 6. leaf in proximal

cross section, x 70; 7. basal leaf cells (left side), x 70; 8.

leaf apex, x 175; 9. rhizoid with tuber, x 35; 10. perichae-

tial leaf, x 18; 11. part of capsule mouth showing cells and

peristome, x 70; 12. spore, x 700. B. canariense (13-19):

13. habit (dry), x 1; 14. habit (wet), x 2,5; 15. leaf, x 18;

16. leaf in proximal cross section, x 70; 17. basal leaf cells

(left side), x 175; 18. leaf apex (left side), x 175; 19. tuber

attached to rhizoid, x 175. (1 & 10-12, Wits Univ. PRE-

CH 12854; 2, Hilliard & Burn 9977; 3, Magill 3736; 4 & 6,

Von Breitenbach 145; 5, Stirton 9893; 7 & 8, Herman 76b;

9, Von Breitenbach 93; 13-18, Oliver 7238; 19, Magill

4091).

Bryaceae

381

Dioicous. Perichaetia terminal; leaves

narrowly oblong to triangular, 1—2 mm long,

costa aristate. Seta 18 — 35 mm long, reddish

brown or yellowish brown; capsule pendulous,

oblong-cylindrical or infrequently pyriform,

frequently contracted below mouth when dry,

yellowish brown or reddish brown, urn

1,5 -3,0 mm long, neck frequently wrinkled

when dry, 1,0— 1,2 mm long; exothecial cells

irregular in shape, incrassate, smaller above;

peristome double, exostome teeth narrowly ob-

long-acuminate, hyaline above, yellowish to

yellowish brown below, finely papillose, en-

dostome segments tapering above, broadly per-

forated, cilia 3—4, appendiculate, basal mem-

brane high, yellowish, minutely papillose;

operculum mammillate or rostellate; calyptra

not seen; spores 12—20 pm, papillose. Fig.

108:1-12.

The species is known from Angola and South Africa.

In the Flora area B. viridescens is collected on soil and bark

in forests, woodlands and grasslands of the central, eastern

and northern Transvaal and Swaziland and rarely in Natal

andTranskei. Map 147.

Vouchers: Crosby & Crosby 9211; Herman 8 1 ; Magill

3444, 3736, 4879; Smook 854a; Stirton 9893; Van Rooy

1382, 1878; Vorster 800.

The species is identified by the upper leaves arranged

in a rosette, red-brown tomentum, absence of a leaf border,

hexagonal laminal cells, inflexed upper leaf laminae and

obtuse leaf apices.

The consistently shorter laminal cells and obtuse leaf

apices will separate B. viridescens from the closely related

B. canariense. B. viridescens is known from Transvaal and

Swaziland and rarely from Natal and Transkei while B.

canariense is known from the Cape, Orange Free State,

Natal, Zululand and rarely from Transvaal.

Mohamed (1979) cited the specimen MacLea 558 in

BM under B. appressum Ren. & Card, but duplicates in

PRE and NH belong here. B. appressum is therefore ex-

cluded from the Flora area .

15. Bryum canariense Brid., Sp. Muse.

3: 29 (1817); Broth, in Natlirl. PflFam. 10: 400

(1924); Sim, Bryo. S. Afr. 338 (1926);

Andrews in Grout, Moss FI. N. Amer. 2: 239

(1940); Lawton, Moss FI. Pacific Northwest

167 (1971); Smith, Moss FI. Brit. Irel. 409

(1978). Type: Canary Islands, Tenerife Island,

Rudley s.n., Bory St. Vine. (B, holo. !).

Bryum mundtii C. Mull, in Bot. Ztg 17: 206 (1859);

Broth, in Naturl. PflFam. 10: 401 (1924); Sim, Bryo. S.

Afr. 337 (1926); Ochi in J. Fac. Educ. Tottori Univ. 23:

100 (1972). Syntypes: Cape: Grootvadersbosch, Mundt

s.n.; Addo, Ecklon s.n., 1829; Genadendal, Breutel s.n.;

Phillipstown ad Katrivier, Ecklon s.n.; Zwarte Hoogdene,

Ecklon s.n., May 1823; Krakamma, Ecklon s.n., July 1832;

Camps Bay, Ecklon s.n., Sept. 1824.

Bryum pervirens C. Mull, in Rabenh. Bryoth. Eur. 28: n.

1399 (1884). Type: Cape, Somerset East, Boschberg, no.

23 Hb. MacOwen (GRA!).

Bryum canariensiforme Dix. in Bull. Torrey bot. Club

43: 68 (1916); Broth, in Naturl. PflFam. 10: 400 (1924);

Sim, Bryo. S. Afr. 336 (1926). Type: Cape, Hobkirk 928,

July 25 1900.

Plants medium-sized to large, solitary or

loosely caespitose to caespitose, green to

ellowish green or olivaceous above, yellowish

rown to brown below; terricolous or saxi-

colous. Stems 3-22 mm tall, frequently

branching by subperichaetial innovations, inno-

vations to 14 mm long, occasionally tomentose

below; rhizoids red-brown, coarsely papillose;

tubers rare, 200-760 /urn, reddish brown or yel-

lowish brown. Leaves comose, frequently semi-

rosulate, frequently in successive comal tufts,

distant and smaller to reduced below, appressed

to erect or occasionally twisted dry, erect-

spreading to widely spreading wet; broadly

ovate to oblong, obovate-oblong or obovate,

(l,7-)2,0-4,0(-6,0) mm long; apex acute to

infrequently obtuse; lamina infrequently in-

flexed above; margins plane above, plane to

recurved in lower A to 2A, entire to crenulate or

denticulate above; border absent or indistinct.

Costa strong, aristate, awn 100-240 pm long,

smooth to occasionally denticulate above, occa-

sionally yellowish or reddish below; in section

subround to round, ventral surface cells pre-

sent, dorsal stereid band strong, stereids in 3-4

rows, dorsal surface cells incrassate to sub-

stereid, hydroids present between guide cells

and stereid band, upper laminal cells rhomboi-

dal, occasionally pitted, frequently incrassate,

frequently gradually narrowed towards margin,

(35 — >45 — 80( — 100) pm long, (11— )12— 20

(-22) pm wide; basal cells rectangular to qua-

drate , pitted , occasionally yellow below .

Dioicous, frequently with sporophytes.

Perichaetia terminal; leaves narrowly oblong-

lanceolate to triangular, 1—3 mm long, costa

aristate. Seta (13 — )18 — 35( — 42) mm long,

yellowish or reddish to reddish brown; capsule

pendulous, straight or slightly curved, obtong-

cylindrical or narrowly pyriform, yellowish

brown to reddish brown, urn 1,0-2, 8 mm

long, neck frequently wrinkled when dry,

0,6- 1,5 mm long; exothecial cells irregular in

shape, incrassate, smaller above; peristome

douole, exostome teeth narrowly oblong-acu-

minate, hyaline above, yellowish below, finely

papillose, endostome segments tapering above,

widely perforated, cilia 2-3, noaose to appen-

diculate, basal membrane high, minutely papil-

382

Bryaceae

Fig. 109. — Bryum perlimbatum (1-7): 1. habit, x

2,5; 2. leaf, x 18; 3. leaf in proximal cross section, x 70;

4. basal leaf cells (left side), x 70; 5. cells at upper right

margin, x 175; 6. tuber attached to rhizoid, x 70; 7. peri-

chaetial leaf, x 35. B. andicola (8-14): 8. habit (dry), X 1;

9. habit (wet), x 2,5; 10. leaf, x 18; 11. leaf in proximal

cross section, x 70; 12. cells at upper left margin, x 175;

13. tuber, x 35; 14. filamentous gemmae, X 175.

B. erythrocaulon (15—21): 15. habit (dry), x 1; 16. habit

(wet), x 3,5; 17. leaf, x 18; 18. leaf in proximal cross

section, x 70; 19. basal leaf cells (right side), x 70; 20.

cells at upper left margin, x 175; 21. tuber, x 70. (1-7,

Benove 37; 8 & 9, Van Rooy 1766; 10 & 14, Van Rooy

1424; 11-13, Van Rooy 1392; 15 & 16, Moll & Cunning-

ham 6009; 17, 19 & 20, DeKock9 ; 18, Cholnoky 307; 21,

Jacot Guillarmod 75-100).

Bryaceae 383

lose, yellowish; operculum mammillate or ros-

tellate; spores (12 — )15 — 20( — 23) /urn, finely

papillose. Fig. 108: 13-19.

The species is known from Europe, North America,

Atlantic Islands and northern and southern Africa. In south-

ern Africa B. canariense is frequently collected in the

southwestern, northwestern, southern, eastern and central

Cape, eastern Orange Free State, Transkei, Natal and Zulu-

land and rarely in eastern and northern Transvaal. Map 148.

Vouchers: Crosby & Crosby 9263; Esterhuysen

27184; Goldblatt 2123; Jacot Guillarmod 7808; Magill

4096, 6183; Oliver 7238; Smook 3480, 4054; Van Rooy

528, 939.

The plants are recognized by their leaves in successive

comal tufts and the absence of a well defined leaf border. In

the majority of southern African plants the laminal cells

gradually narrow and elongate towards the margin.

B. mundtii has longer leaves and laminal cells, longer

and narrower cells along the leaf margin and stronger denti-

culate leaf margins than the type of B. canariense. Al-

though the two extremes appear distinct, a continual grada-

tion in leaf size and shape, cell size, differentiation of

border cells and denticulation, from B. canariense to B.

mundtii, is evident in southern African plants. Additional

separating characters were not found. Differentiation of the

border cells even varies among leaves on the same plant

(Oliver 7239, Barnard SAM-46150).

Plants with narrow, incrassate border cells ( Schelpe

7649, Garside 6513, Van Rooy 774) can be mistaken for B.

erythrocaulon but the laminal cells of these specimens grad-

ually narrow towards the margin and the border is not as

well defined as in B. erythrocaulon.

Southern African plants of B. canariense are fre-

quently fruiting but tubers are rarely found.

For differences between B. canariense and B. virides-

cens see note under that species.

16. Bryum erythrocaulon (Schwaegr.)

Brid., Mant. Muse. 119 (1819); Mohamed in J.

Bryol. 10: 428 (1979). Type: Mauritius, Com-

merson s.n. (G!).

Mnium erythrocaulon Schwaegr., Sp. Muse. Suppl. 1:

127(1816).

Plants medium-sized to large, loosely

caespitose, green or reddish green above,

brown or rea-brown below; temcolous or on

charcoal. Stems to 20 mm tall, branching by

innovations, tomentose below; rhizoids reddish

brown, coarsely papillose, tubers large, (250—)

350-550(-750) gm, yellow-brown. Leaves

comose, frequently in successive comal tufts,

larger above, erect or slightly twisted dry,

erect-spreading and frequently rosulate wet; ob-

long to oblong-spathulate, 3, 0-4, 5 mm long;

apex acute; margins plane above, recurved in

lower / to %, dentate above; border 4-8 cells

wide, cells sinuate, incrassate. Costa mucro-

nate to shortly excurrent, denticulate above,

frequently orange below; in section subround to

round, laminal insertion ventral, ventral surface

cells present, dorsal stereid band strong, ster-

eids in 3-5 rows, dorsal surface cells incras-

sate, hy droids present between stereid band and

ide cells. Upper laminal cells long-rhom-

idal, frequently incrassate, pitted, (45-)

52— 73(— 83) pm long, 12-19 pm wide, grad-

ually narrowed towards border; basal cells rec-

tangular, pitted, reddish or orange below. Fila-

mentous gemmae absent.

Sporophytes not seen. Fig. 109: 15-21.

This species is known from Madagascar, Mauritius

and southern Africa. In the Flora area B. erythrocaulon is

infrequently collected in forests and woodlands of the

southern, southwestern and eastern Cape, Transkei, Natal,

Zululand and northern Transvaal. Map 149.

384

Bryaceae

Vouchers: Cholnoky 180b, 306, 307; De Kock 9; Lam-

bert 12; Moll & Cunningham 6009; Russell 2672.

The leaves of B. erythrocaulon frequently have a red-

dish green tinge, are rosulate when wet and oblong to ob-

long-spathulate in shape. The upper laminal cells are long-

rhomboidal, generally incrassate and gradually narrow to-

wards the wide, distinctive border.

17. Bryum perlimbatum Card, in Bull.

Herb. Boissier 2 (5): 1007 (1905); Ochi in J.

Fac. Educ. Tottori Univ. 23: 105 (1972);

Mohamed in J. Bryol. 10: 449 (1979). Type:

Falkland Islands, Dupemey Harbour, Skotts-

berg 236 (S).

Bryum herpetineuron Ther. in Bull. Mus. Hist. Nat.

Paris 34: 1 17 (1928). Type: Natal, Port Natal, Ultima Espe-

ranzo, Benove 37 (PC!).

Plants medium-sized, caespitose, green to

yellowish green above, reddish or reddish

brown below; terricolous. Stems 3-11 mm tall,

simple or branching by subperichaetial innova-

tions; rhizoids red-brown, densely papillose,

tubers 150-550 pm, reddish brown. Leaves

comose, larger above, erect and twisted dry,

rosulate wet; oval or broadly oblong, frequently

concave, 1, 5-4,0 mm long; apex acute to ob-

tuse; margins recurved to revolute in lower % to

/, smooth to denticulate above; border 4-12

cells wide, cells narrow, incrassate. Costa aris-

tate, awn to 0,3 mm long, frequently reddish

below; in section round, laminal insertion ven-

tral, ventral surface cells present, dorsal stereid

band strong, stereids in 4-7 rows, dorsal sur-

face cells incrassate, hy droids present between

stereid band and guide cells. Upper laminal

cells short-rhomboidal, pitted, (30- ) 40-53

(-60) pm long, (15-) 17 — 20 (-23) pm wide;

basal cells rectangular, pitted, reddish below.

Filamentous gemmae absent.

Dioicous. Sporophytes not seen. Fig. 109:

Bryum perlimbatum occurs in New Zealand, South

America, the Falkland Islands and southern Africa. Only

one collection, the type specimen of B. herpetineuron, is

known from the Flora area. Map 150.

Voucher: Type of synonym only.

This species can be recognized by the concave,

broadly ovate or oblong leaves with wide borders.

18. Bryum andicola Hook, in Kunth,

Syn. PI. Aequin. 1: 58 (1822); Mohamed in J.

Bryol. 10: 421 (1979). Type: Mexico.

Bryum pumiliroseum Dix. in Trans. R. Soc. S. Afr. 8:

203 (1920); Sim, Bryo. S. Afr. 336 (1926). Syntypes:

Cape, Hogsback, Tjumie, 1917, Henderson 326, 327, 337

(BOL!).

Plants medium-sized to large, loosely

caespitose, green to yellow-green above, brown

to reddish brown below; temcolous and cortico-

lous. Stems 3-18 (-30) mm tall, simple or

branching by innovations, tomentose below;

rhizoids red-brown, densely papillose, tubers

large, 525-1100 pm, reddish brown or yellow-

brown. Leaves comose, frequently in succes-

sive comal tufts, larger above, twisted dry, ro-

sulate above wet; elliptical to oblong or spathu-

late, (2,5-) 2, 8-3, 8 (-4,0) mm long; apex

acute to occasionally obtuse; margins recurved

to re volute in lower % to3/, serrulate to serrate

above; border 2—6 cells wide, cells sinuate,

incrassate. Costa mucronate to excurrent as

short awn, frequently denticulate above, awn to

0,3 mm long, reddish to reddish brown below;

in section generally round, dorsal stereid band

strong, stereids in 4-6 rows, dorsal surface

cells incrassate, hy droids present between ster-

eid band and guide cells. Upper laminal cells

short-rhomboiaal, frequently pitted, (25-)

33-45 (-50) pm long, (12-) 13-18 (-20)

pm wide; basal cells rectangular to occasionally

quadrate, pitted, frequently reddish to reddish

brown below. Filamentous gemmae present in

axils of comal leaves, highly branched, coar-

sely papillose, reddish brown to brown.

Dioicous, sporophytes infrequent. Peri-

chaetia terminal, leaves oblong to lanceolate or

triangular. Seta 15-20 mm long, yellowish to

Bryaceae

385

reddish brown; capsule horizontal, pyriform,

brown, slightly contracted below mouth when

dry, urn 1,1 — 1,5 mm long, neck wrinkled

when dry; exothecial cells irregularly rectangu-

lar, 4-7 rows at mouth smaller, quadrate, in-

crassate; peristome double, exostome teeth nar-

rowly oblong-acuminate, to 0,4 mm long,

finely papillose, endostome fragile, minutely

papillose; operculum and calyptra not seen;

spores round, 20—25 /xm, finely papillose,

light brown. Fig. 109: 8— 14.

Bryum andicola is known from North, Central and

South America and surrounding islands. East Africa and

southern Africa. In the Flora area the species has been col-

lected in forests and woodlands of the southwestern and

central Cape, Transkei, Natal, eastern Orange Free State,

Lesotho, eastern, southern, central and northern Transvaal

and Swaziland. Map 150.

Vouchers: Cholnoky 102; Magill 3536; Pienaar 33;

Raven 26126; Van Rooy 1424, 1589, 1627; Wager 1 155.

This species is most easily identified by the conspicu-

ous, reddish brown filamentous gemmae in the axils of the

comal leaves. The leaves which are twisted when dry and

rosulate when wet and the presence of large tubers will also

help to distinguish B. andicola.

19. Bryum pycnophyllum (Dix.) Moha-

med in J. Brvol. 10: 435 (1979). Syntypes:

Zimbabwe [Rhodesia], Zimbabwe, Sim 8737;

Umtali, Eyles 1725 (both PRE!).

Bryum truncorum (Brid.) Brid. var. pycnophyllum Dix.

inS.Afr.J.Sci. 18:321 (1922).

Bryum truncorum sensu Sim, Bryo. S. Afr. 337 (1926).

Plants medium-sized to robust, loosely

caespitose to caespitose, green to yellowish

green above, brownish below; terricolous or hu-

micolous. Stems 5-45 mm tall, simple or

branching by innovations, densely tomentose

below; rhizoids red-brown or red, coarsely pa-

pillose, tubers generally absent. Leaves evenly

spaced along stem or comose above, frequently

in successive comal tufts, larger above, twisted

dry, erect-spreading wet, elliptical to obovate,

(2,5—) 3, 0-6,0 mm long; apex acute; margins

plane above, recurved in lower lA to %, denti-

culate to dentate above; border 2-5 cells wide,

cells sinuose, incrassate, yellowish. Costa aris-

tate, awn to 0,7 mm long, entire to denticulate

above, yellowish; in section semi-circular or

subquadrate to subrectangular, ventral surface

cells present, dorsal substereid band weak to

strong, dorsal surface cells incrassate or subste-

reids, hydroids present between stereid band

and guide cells. Upper laminal cells short-

rhomboidal, frequently narrower and incrassate

towards border, pitted, (35-) 40-63 (-78)

pm long, (18 — ) 20—23 (—28) pm wide; basal

cells rectangular, pitted. Filamentous gemmae

absent.

Dioicous. Perichaetia terminal, leaves nar-

rowly oblong or triangular. Seta 15-30 mm

long, yellowish to reddish brown; capsule hori-

zontal, straight to curved, yellowish or red-

brown, urn 2,2— 3,0 mm long, neck wrinkled

dry; exothecial cells irregularly rectangular, in-

crassate; peristome double, exostome teeth ob-

long-acuminate, 43—58 pm long, apex hya-

line, finely papillose, endostome segments

keeled, perforated, cilia 2-4, appendiculate,

basal membrane high, finely papillose; opercu-

lum conic; spores 18—22 pm, pale yellow to

brown, weakly papillose. Fig. 110: 1-7.

B. pycnophyllum is known from Africa south of the

equator and has been collected in Tanzania, Zimbabwe and

South Africa. In the Flora area this species is frequently

collected in the southern, eastern, central and northern

Cape, Transkei, Lesotho, Orange Free State, Natal, Zulu-

land and in the southern, eastern, central and northern

Transvaal. Map 151.

Vouchers: Crosby & Crosby 150\',Magill 3520, 3807,

4885; Rankin 216; Smook 2312; Van Rooy 73, 558, 745,

1041, 1357, 1602.

When dry, this species is recognized by the twisted

leaves with yellowish borders and costae. The shape of the

costa is generally subrectangular in cross-section, con-

taining a large gap of collapsed hydroids and often weak

dorsal stereid band. Filamentous gemmae and tubers are

generally absent.

Tubers were found in two specimens, Smook 1411

from Natal and Vahrmeijer PRE-CHI 2682 from the Cape.

The reddish tubers, 150-250 /xm in diameter, resemble

those of B. torquescens and plants of B. pycnophyllum in

the two specimens might have been mixed with that species.

386

Bryaceae

Fig. 110. — Bryum pycnophyllum (1-7): 1. habit

(dry), x 2; 2. habit (wet), x 3,5; 3 & 4. leaves, x 18; 5.

leaf in proximal cross section, X 70; 6. basal leaf cells

(right side), x 175; 7. cells at upper right margin, x 175,.

B. aubertii (8-15): 8. habit (dry), x 2; 9. habit (wet), x

3,5; 10 & 11. leaves, X 12; 12. leaf in proximal cross

section, x 70; 13. basal leaf cells (right side), x 175; 14.

cells at upper right margin, x 175; 15. filamentous gem-

mae, X 1/5. (1, Pegler 1450; 2, Van Rooy 1702; 3, Van

Rooy 548; 4, 5 & 7, Smook 1724a; 6, Crosby & Crosby

7742; 8 & 9, Van Rooy 1004a; 10, Van Rooy 350; 11,

Schelpe 7580; 12 & 15, Smook 2549; 13 & 14, Ruch 2018).

Bryaceae

387

More tubers must be found on the rhizoids of B. pycnophyl-

lum before it can be reported as a tuber-bearing species.

Plants growing under more xeric conditions, at high

altitudes and dry regions of the eastern Orange Free State,

Lesotho, central and northern Cape (Van Rooy 548, 743;

Magill 4635, 4637, 5839; Smook 2838) are generally smal-

ler and have smaller, rounded leaves with weaker borders

and denticulations. These plants still fall well within the

wide range of variation in plant size and morphology dis-

played by B. pycnophyllum.

South African specimens of B. pycnophyllum were

generally referred to B. billardieri Schwaegr. but that

species bears large tubers, its leaves are not distinctly

twisted when dry and it is not known from Africa. Speci-

mens referred to the East African B. voeltzkowii Broth, are

large facies of B. pycnophyllum.

20. Bryum aubertii (Schwaegr.) Brid.,

Mant. Muse. 119 (1819); Ochi in J. Fac. Educ.

Tottori Univ. 23: 118 (1972). Type: Insula

Franciae, Aubert s.n. (G, holo.!).

Mnium aubertii Schwaegr., Sp. Muse. Suppl. 1 (2): 132

(1816). Rhodobryum aubertii (Schwaegr.) Trier. in Recueil

Publ. Soc. Havraise Etud. Div. 89 (2): 128 (1922);

Mohamedin J. Bryol. 11:692(1981).

Plants large to robust, loosely caespitose,

green or reddish green above, dark- green or

brownish below; terricolous or corticolous.

Stems 5-60 mm tall, branching by innovations,

frequently tomentose; rhizoids red-brown,

papillose, tubers absent. Leaves equidistant to

comose, successive comal tufts frequently pre-

sent, twisted dry, erect-spreading wet; ellipti-

cal, oblong-spathulate or oblong-acuminate,

3-6 mm long; apex acute to acuminate; mar-

gins plane above, plane to recurved below, den-

tate to weakly spinose in upper lA to 2A; border

absent or 1 —3 (—4) cells wide, generally incon-

spicuous. Costa short-excurrent or occasionally

ending below apex, yellowish or reddish, fre-

quently weak above; in section subrectangular,

ventral surface cells present, dorsal stereid band

absent, dorsal cells incrassate to substereid, hy-

droids present between stereid band and guide

cells. Upper laminal cells rhomboidal, fre-

uently pitted, (46-) 55-75 (-88) /urn long,

0-25 /urn wide; basal cells rectangular, pitted,

occasionally reddish. Filamentous gemmae fre-

quently produced on leaf laminae, branched,

yellowish brown or reddish brown, finely papil-

lose.

Dioicous. Perichaetia terminal; leaves lan-

ceolate to triangular, margins entire to dentate,

border absent, costa aristate. Sporophytes not

known. Fig. 110: 8-15.

Bryum aubertii is known from Madagascar, Reunion,

Mauritius, central and southern Africa. In the Flora area this

species is infrequently collected on soil, rock, humus or

bark in forests and woodlands of Transkei, Natal, Zululand,

eastern Orange Free State, Lesotho, southern, central and

northern Transvaal and on Table Mountain in the

southwestern Cape. Map 152.

Vouchers: Crosby & Crosby 7879; Magill 5712; Retief

733; Smook 1825, 2549; Van Rooy 121, 382, 1004a, 1665,

2226.

The species is recognized by the strongly toothed leaf

margins, the weak leaf borders and the presence of fila-

mentous gemmae on the leaf laminae. Southern African

plants in general are smaller and have stronger toothed leaf

margins than the type specimen. No filamentous gemmae

could be found in the type specimen.

A specimen from Natal (Van Rooy 1527) has entire to

weakly dentate leaf margins but agrees in all other charac-

ters with southern African specimens of B. aubertii.

Although B. aubertii resembles species of Rhodo-

bryum, southern African specimens are treated here because

of their habit.

Insufficiently Known Species

Bryum condensatum Hampe in C. Miill., Bot. Ztg 16:

155 (1858); Sim, Bryo. S. Afr. 329 (1926). Type: Cap,

Genadendal (probably collected by Breutel). The type has

not been seen. Ochi (1972) treated the speies provisionally

as a synonym of B. canariense without citing a type speci-

men and it is not clear whether he has seen the type or not.

Bryum laxogemmaceum C. Miill. in Hedwigia 38: 75

(1899); Broth, in Natiirl. PflFam. 10: 394 (1924). Typ:

Cape, Knysna, Estemek, Rehmann 253 (BM, iso.). The

type has not been seen. Sim (1926) considered the species

to be synonymous with B. erythrocarpum Schwaegr.

Crundwell & Nyholm (1964) noted that the isotype in Brit-

ish Museum is mostly Funaria hygrometrica and the

scanty, sterile Bryum material is indeterminable.

Bryum macleanum C. Miill. in Hedwigia 38: 74

(1899); Broth, in Natiirl. PflFam. 10: 394 (1924). Typ:

Transvaal, Lydenburg, 1891, MacLea in Herb. Jack. The

Map 152. — # Bryum aubertii

388

Bryaceae

type has not been seen. Sim (1926) treated the species as a

synonym of B. erythrocarpum and Crundwell & Nyholm

(1964) noted that the type specimen was probably destroyed

with Muller’s herbarium in Berlin.

Bryum pappeanum C. Miill. in Bot. Ztg 13: 752

(1885); Broth, in Natiirl. PflFam. 10: 399 (1924). Type:

Cape, Pappe s.n., 1838. The type has not been seen. Sim

(1926) suggested that this species is synonymous with B.

capillare, and Ochi (1972) also treated it as a synonym of

B. capillare, without having seen the type.

Bryum polytrichoideum C. Miill. in Bot. Ztg 17: 206

(1859). Type: Cape, Rondebosch, Pappe s.n. (BM, syn.).

The type has not been seen. Brotherus (1924) considered

the species to be closely related to B. canariense and Sim

(1926) confused it with B. viridescens without having seen

the type. Ochi (1972) after seeing the type, treated it as a

synonym of B. capillare.

Bryum prionotes Shaw in Cape Monthly Mag. 17: 319

(1878). Type: Cape, Katberg, s.l. Sim (1926) suggested

that this is a synonym of B. muehlenbeckii B.S.G. without

having seen the type. The Shaw collection was probably

destroyed sometime after his death and attempts to locate

specimens in southern African and European herbaria have

been unsuccessful.

Bryum transvaaloalpinum C. Miill. in Hedwigia 38: 73

(1899). Philonotis transvaaloalpinum (C. Miill.) Broth, in

Natiirl. PflFam. 1: 598 (1909). Type: Transvaal, between

Middelburg and Lydenburg, Dec. 1883, Wilms s.n. in

Herb. Jack. The type has not been seen. Judging by the

orginal description, it is likely that Sim (1926) is correct in

referring this species to Bryum alpinum, although he had

not seen the type.

9. RHODOBRYUM

Rhodobryum (Schimp.) Limpr., Laubm. Deutschl. 2: 444 (1892); Broth, in Natiirl. PflFam. 10:

402 (1924); Sim, Bryo. S. Afr. 339 (1926); Andrews in Grout, Moss FI. N. Amer. 2: 241 (1940);

Iwatsuki & Koponen in Acta Bot. Fenn. 96: 3 (1972); Gangulee, Moss. E. India 4: 1015 (1974);

Smith, Moss FI. Brit. Irel. 430 (1978); Crum & Anderson, Moss. E. N. Amer. 1: 575 (1981).

Type species: R. roseum (Hedw.) Limpr.

Plants large to robust, solitary or gregarious; terricolous or humicolous. Stems frequently erect

from rhizomes, to 70 mm tall, simple or branching by subperichaetial innovations, tomentose; in

section with central strand, inner cortical cells thin-walled, outer cortical cells smaller, incrassate.

Leaves loosely or tightly rosulate above, distant and reduced below, spathulate or obovate; margins

denticulate to dentate above, border 1-4 cells wide. Costa ending below apex to short excurrent; in

section with ventral cells in several rows, dorsal stereid or substereid band frequently small or

absent, rarely large, hydroids differentiated. Upper laminal cells rhomboidal, basal cells rectangu-

lar.

Dioicous, frequently poly setaceous. Perichaetia and perigonia terminal. Seta long; capsule

horizontal to pendulous, frequently curved, generally obiong-cylindrical, frequently contracted

below mouth when dry, neck frequently wrinkled when dry; stomata phaneropore; annulus present;

peristome double, exostome teeth 16, oblong-acuminate, trabeculate, hyaline above; endostome

segments broadly perforated, keeled, cilia appendiculate, basal membrane high; operculum conic,

frequently mammillate; calyptra cucullate; spores round, smooth to minutely papillose.

This genus contains approximately 48 species and is found from the temperate to the tropical zone on all the continents,

excluding Antarctica. Tropical and subtropical Africa and South America are the two major centres of diversity.

Rhodobryum has many characters in common with the larger species of Bryum, but all or a combination of the

following characters will usually place plants in this genus: 1 . plants robust; 2. stems erect from rhizomes; 3. upper leaves

arranged in distinct rosettes with lower leaves distant and reduced; 4. costa in cross-section with ventral cells in several rows

and dorsal stereid or substereid band small or absent; 5. more than one sporophyte produced from a single perichaetium.

The costal anatomy has proved to be an important taxonomic character. Cross-sections were taken from the lower third

of the leaf as the costa is frequently weak above.

The four species known from southern Africa are collected on humus-rich soil or litter in forests and dense, moist

woodlands.

1 Costa in cross-section with dorsal stereid band absent; leaf margins generally plane 3. R. commersonii

1 Costa in cross-section with dorsal stereid or substereid band; leaf margins recurved to revolute below:

Bryaceae

389

2 Costa in section with dorsal stereid band large; rhizomes absent 1 . R. keniae

2 Costa in section with dorsal stereid or substereid band small; stems frequently erect from rhizomes:

3 Leaves obovate-spathulate, undulate when wet; costa in section with substereid band 4.R. umbraculum

3 Leaves spathulate or oblong-spathulate, flat when wet; costa in section with stereid band 2.R. roseum

1. Rhodobryum keniae (C. Mull.) Broth.

in Naturl. PflFam. 10: 404 (1924). Type: Kenia

[Kenya], VonHohnel s.n., (H, iso.!).

Bryum keniae C. Mull, in Flora, Jena 73: 475 (1890);

Ochi in J. Fac. Educ. Tottori Univ. 23: 112 (1972); Bizot &

P6cs in Acta bot. hung. 25: 256 (1979).

Plants large to robust, gregarious, green or

yellowish green above, brown or reddish brown

below; terricolous or humicolous. Stems erect,

5—70 mm tall, frequently branching by subpe-

richaetial innovations, brown or reddish brown,

densely tomentose; rhizoids reddish brown,

densely papillose. Leaves in successive rosettes

through innovation, distant and reduced below,

erect and twisted dry, widespreading wet; spa-

thulate or oblong-spathulate, 6-11 mm long;

apex acute to obtuse; margins recurved to revo-

lute in lower % to %, denticulate to dentate

above; border 2— 3(— 4) cells wide. Costa cus-

pidate or short excurrent, smooth, frequently

yellowish or reddish below; in section round or

subround, ventrally flat or rarely concave, lami-

nal insertion ventral, ventral cells in 2 rows,

dorsal stereid band strong, stereids in 3-5

rows, dorsal surface cells incrassate, hydroids

present between stereid band and guide cells.

Upper laminal cells rhomboidal to short-rhom-

boidal, 62 — 85( — 100) /zm long, 22 — 30( — 35)

pm wide, pitted; basal cells rectangular, pitted,

frequently yellowish or reddish below. Fila-

mentous gemmae subperichaetial on stem, red-

dish brown, coarsely papillose.

Dioicous, infrequently poly setaceous. Pe-

richaetial, leaves triangular, apex acuminate,

margins serrulate, costa aristate. Seta (27—)

30— 35(— 48) mm long, yellowish or reddish

brown; capsule inclined to pendulous, yellow-

ish brown or reddish brown, oblong-cylindri-

cal, straight or slightly curved, slightly con-

tracted below mouth when dry, urn 3-5 mm

long, neck wrinkled when dry, 1-2 mm long;

exothecial cells irregular in shape, incrassate,

smaller at mouth; peristome teeth narrowly ob-

long-acuminate, yellowish brown below, hya-

line above. Finely papillose; endostome seg-

ments keeled, tapered above, broadly perfo-

rated, cilia 3, appendiculate, basal membrane

high, yellowish, finely papillose; operculum

conic, mammillate; calyptra not seen; spores

round, 12-18 pm, weakly papillose. Fig. Ill:

17-22.

Rhodobryum keniae occurs in submontane and mon-

tane forests of tropical and subtropical Africa. In southern

Africa this species is infrequently collected on soil or humus

in forests and dense woodlands of the eastern Cape, Natal,

Swaziland and eastern and northern Transvaal. Map 153.

Vouchers: Magill 3533; Manders 67; Obermeyer PRE-

CHI 1892; Stirton 1779; Van der Schijff 6184; Van Rooy

1222, 1586; Von Breitenbachlh.

This species is similar to those Bryum species with

larger plants in the absence of rhizomes, the costal anatomy

and the arrangement of leaves in successive rosettes. R.

keniae is occasionally poly setaceous, the upper leaves are

tightly rosulate and the lower leaves are distant and re-

duced. It has therefore provisionally been placed in Rhodo-

bryum rather than Bryum.

Small, yellowish brown or reddish brown, papillose,

subperichaetial filamentous gemmae are present on south-

ern African plants.

2. Rhodobryum roseum (Hedw.) Limpr.,

Laubm. Deutschl. 2: 445 (1892); Broth, in

Naturl. PflFam. 10: 404 (1924); Sim, Bryo. S.

Afr. 340 (1926); Andrews in Grout, Moss FI.

N. Amer. 2: 241 (1940); Nyholm, Moss FI.

Fenn. 259 (1954); Gangulee, Moss. E. India 4:

1016 (1974); Smith, Moss FI. Brit. Irel. 430

(1978); Crum & Anderson, Moss. E. N. Amer.

1: 575 (1981). Type: Europe.

Map 153. — # Rhodobryum keniae

Bryaceae

391

Mnium roseum Hedw., Sp. Muse. 194 (1801). Bryum

roseum (Hedw.) Gaertn., Meyer & Scherb. in Oek. Techn.

FI. Wetterau 3: 104 (1802); Ochi in J. Fac. Educ. Tottori

Univ. 23: 111 (1972).

Mnium spathulatum Homsch. in Linnaea 15: 135 (1841).

Rhodobryum spathulatum (Homsch.) Poes in Bizot & Poes

in Acta bot. hung. 25: 257 (1979); Koponen, Li & Zang in

Ann. Bot. Fenn. 19: 78 (1982). Type: Cape, Drege, L no.

910. 115-97 (L, lecto. !), vide Touw in Lindbergia 9: 151

(1983).

Bryum leucothrix C. Miill. in Hedwigia 38: 69 (1899).

Rhodobryum leucothrix (C. Miill.) Broth, in Natiirl.

PflFam. 10: 404 (1924). Type: Cape, Boschberg, Mac-

Owan 20 (GRA, iso. !).

Plants large to robust, solitary or grega-

rious, green above, reddish brown below; terri-

colous or humicolous. Stems erect from rhi-

zomes, 10-50 mm tall, simple or occasionally

branching by subperichaetial innovations, rea-

brown, tomentose; rhizoids reddish brown,

densely papillose. Leaves distant and reduced

below, rosulate above, erect and twisted dry,

spreading to squarrose wet; spathulate or ob-

long-spathulate, 7-14 mm long; apex acute or

occasionally obtuse; margins recurved to revo-

lute in lower / to %, denticulate to dentate

above; border weak, 1 — 2( — 3 ) cells wide.

Costa short excurrent or occasionally ending

below apex, tapering above; in section sub-

round or semi-crescent shaped, ventrally flat to

slightly concave, laminal insertion ventral, ven-

tral cells in 4-6 rows, dorsal stereid band

small, dorsal cells generally in 2 rows, hydroids

present between stereid band and ventral cells.

Upper laminal cells rhomboidal, (72—)

85— 1 12(— 137) pm long, (25-)30-35 pm

wide, frequently pitted; basal cells rectangular,

frequently pitted. Filamentous gemmae absent.

Dioicous, frequently poly setaceous. Peri-

chaetia terminal; leaves lanceolate or triangular,

margins entire to denticulate above, costa form-

ing denticulate awn. Seta 27-42 mm tall, yel-

lowish red or reddish brown; capsule pendu-

lous, yellowish brown, oblong-cylindrical,

slightly curved, slightly contracted below

mouth dry, urn 2, 5-5,0 mm long, neck short;

exothecial cells irregular in shape, incrassate,

smaller at mouth; peristome teeth narrowly ob-

long-acuminate, yellowish brown or orange,

hyaline above, finely papillose, endostome seg-

ments keeled, tapered above, broadly perfo-

rated, cilia 2-3, appendiculate, basal mem-

brane high, yellowish, minutely papillose;

operculum conic, mammillate; spores round,

17-23 pm, minutely papillose. Fig. Ill:

11-16.

This widely distributed species is known from Europe,

Asia, Japan, northern and central America and eastern and

southern Africa. In southern Africa it is infrequently col-

lected on soil or humus in forests of the eastern and central

Cape, Natal and eastern and northern Transvaal. Map 154.

Vouchers: Esterhuysen 20229a; Filter 1; Junod 4019;

Magill 5520; Rennie & Lambert moss no. 24; Van Rooy

1171, 1492; Young 3016.

Rhodobryum roseum can be recognized by its tightly

rosulate leaves, the spathulate or oblong-spathulate leaf

shape, recurved to revolute lower leaf margins and the cos-

tal anatomy.

Although Iwatsuki & Koponen (1972) and Koponen et

al. (1982) listed several characters to separate R. roseum

from R. spathulatum (=R. ontariense Kindb.), these cha-

racters are not reliable for separation of specimens in sou-

thern Africa.

The lectotype of Mnium spathulatum at L, selected by

Touw (1983), consists mostly of Bryum aubertii. However,

a plant affixed to the sheet above the original handwriting,

closely resembles Rhodobryum roseum. The original collec-

tion could have been mixed and we therefore follow Ochi

(1972) in including R. spathulatum in the synonymy of R.

roseum.

FlG. 111. — Rhodobryum umbraculum (1-10): 1. habit (dry), x 1;2. habit (wet), x 1 ; 3. stem in cross section, x

42; 4. leaf, x 5; 5. leaf in proximal cross section, x 175; 6. cells at upper right margin, x 175; 7. leaf apex, x 42; 8.

perichaetial leaf, x 5; 9. sporophyte, x 5; 10. part of capsule mouth showing cells, peristome and spores x 70. R. roseum

( 1 1—16): 11. habit (wet), x 1; 12. leaf, x 5; 13. leaf in proximal cross section, x 175; 14. cells at upper right margin, x

175; 15. leaf apex, x 42; 16. perichaetial leaf, x 5. R. keniae (17-22): 17. habit, x 1; 18. leaf, x 5; 19. leaf in proximal

cross section, x 175; 20. cells at upper right margin, x 175; 21. leaf apex, x 42; 22. perichaetial leaf, x 5. (1-3 & 6-10,

C rosby & Crosby 1905, 4 & 5, Kluge 1004; 1 1-16, Magill 5520; 17-21 , Magill 3537; 22, Obermeyer PRE-CHI 1893).

392

Bryaceae

Fig. 112. — Rhodobryum commersonii: 1. habit

(wet), x 1,5; 2. leaf, x 5; 3. costa in proximal cross sec-

tion, x 175; 4. basal leaf cells (right side), x 88; 5. cells at

upper left margin, x 175; 6. leaf apex, x 42; 7. perichaetial

leaf, x 5 . (1 & 7 ,Junod 13; 2—6, Smook & Phelan 664).

3. Rhodobryum commersonii (Schwaegr.)

Par., Ind. Bryol. 115 (1898); Broth, in Natiirl.

PflFam. 10: 404 (1924); Sim, Bryo. S. Afr. 340

(1926); Mohamed in J. Bryol. 11; 691 (1981).

Type: Reunion, Commerson H-89 (H, iso.).

Mnium commersonii Schwaegr., Sp. Muse. Suppl. 1 (2):

134 (1816). Bryum commersonii (Schwaegr.) Bna., Mant.

Muse. 1 19 (1819); Ochi in J. Fac. Educ. Tottori Univ. 23:

121 (1972).

Plants large to robust, gregarious, green

above, brownish below; terricolous or humico-

lous. Stems frequently erect from rhizomes, to

50 mm tall, simple or occasionally branching

by subperichaetial innovations, red-brown, to-

mentose below; rhizoids reddish brown, papil-

lose. Leaves crowded or loosely rosulate above,

distant and smaller below, erect dry, wide-

spreading to squarrose wet, undulate and

slightly twisted dry, flat to weakly undulate

above wet, obovate, 7-15 mm long; apex acute

to obtuse; margins plane above, plane to occa-

sionally recurved below, denticulate to dentate

above; border 1-3 cells wide. Costa subpercur-

rent to short excurrent, frequently reddish; in

section subrectangular, laminal insertion ven-

tral, ventral cells in 3-5 rows, dorsal stereid

band absent, dorsal cells thin-walled, occasio-

nally incrassate dorsally, hydroids present in

centre of costa. Upper laminal cells rhomboidal

to hexagonal, pitted, (40— )67— 100(— 135) pm

long, (30— )33— 45(— 58) pm wide; basal cells

rectangular, pitted, occasionally reddish below.

Filamentous gemmae absent.

Dioicous, infrequently poly setaceous.

Perichaetia terminal, leaves lanceolate or

triangular. Seta 30-57 mm long, yellowish or

reddish or reddish brown; capsule horizontal to

pendulous, curved, oblong-cylindrical, con-

tracted below mouth dry, brown or reddish

brown, urn 4, 5-6,0 mm long, neck wrinkled

dry, 1 ,2-2,0 mm long; exothecial cells irregular

in shape, smaller at mouth, incrassate; peri-

stome teeth oblong-acuminate or lanceolate,

yellowish brown below, hyaline above, trabe-

culate, finely papillose, endostome segments

keeled, broad below, tapered above, broadly

perforated, cilia 2, appendiculate, basal mem-

brane high, yellowisn, minutely papillose;

spores round, 14-18 pm, minutely papillose.

Fig. 112.

Rhodobryum commersonii occurs in India, Madagas-

car, Comoro Islands and eastern, central and southern Af-

rica. In the Flora area this species is collected on humus or

soil in forests and dense woodlands of the southern and

eastern Cape, Transkei, Natal, Zululand, Swaziland and

eastern and northern Transvaal. Map 155.

Vouchers; Kemp 1513; Knox 6; Under 1229; Oliver

7153; Russell 2555; Schelpe 7937; Smook 1533; Smook &

Phelan 664; Van Rooy 1573; Von Breitenbach 109.

Rhodobryum commersonii can be distinguished from

other species of Rhodobryum by its obovate leaves, mostly

plane leaf margins, subrectangular shape of the costa in

cross section and the absence of a dorsal stereid band in the

costa.

4. Rhodobryum umbraculum (Hook.)

Schimp. ex Par., Ind. Bryol. 1122 (1898);

Broth, in Natiirl. PflFam. 10: 404 (1924); Sim,

Bryo. S. Afr. 339 (1926). Type: Cape, Bur-

chell-Hooker 2507 (BM, lecto.), fide Ochi in J.

Fac. Educ. Tottori Univ. 23: 115 (1972).

Bryaceae

393

Bryum umbraculum Bruch ex Hook., Musci Exot. 2:

133 (1819); Ochi in J. Fac. Educ. Tottori Univ. 23: 115

(1972).

Plants large to robust, solitary or gregar-

ious, green or dark green above, reddish brown

below; terricolous or humicolous. Stems erect

from rhizomes, to 40 mm tall, simple or occa-

sionally branching by subperichaetial innova-

tions, red-brown, tomentose; rhizoids reddish

brown, papillose. Leaves distant and reduced

below, rosulate above, undulate, erect dry,

widely spreading to squarrose wet; broadly obo-

vate-spathulate, 7-15 mm long; apex generally

obtuse, occasionally apiculate; margins plane

above, recurved below, denticulate above;

border (2-)3^f cells wide, occasionally red-

dish. Costa ending below apex to apiculate,

frequently reddish; in section subround to semi-

crescent shaped, ventrally slightly concave,

laminal insertion ventral, ventral cells in 5-6

rows, dorsal substereid band weakly differen-

tiated, dorsal cells incrassate, reddish, hydroids

present between substereids and ventral

cells. Upper laminal cells rhomboidal,

(85— ) 100— 1 50(— 180) pm long, (35-)45-55

pm wide, pitted, gradually narrowed towards

border; basal cells rectangular, pitted. Fila-

mentous gemmae absent.

Dioicous, poly setaceous. Perichaetia ter-

minal; leaves oolong-acuminate or triangular,

reddish below; margins entire to denticulate;

border weak or absent; costa ending below

apex. Seta 27-32 mm long, reddish brown; cap-

sule horizontal to pendulous, brown or reddish

brown, straight or slightly curved, oblong-

cylindrical, contracted below mouth dry, urn

3, 0-3, 5 mm long, neck 1,0-1, 8 mm long,

wrinkled dry; exothecial cells irregular in

shape, incrassate, smaller at mouth, 1-2 rows at

mouth in transverse rows; peristome teeth nar-

rowly oblong-acuminate or lanceolate, yellow-

ish brown, hyaline above, trabeculate, finely

papillose, endostome segments keeled, broad

below, tapered above, broadly perforated, cilia

3, appendiculate, basal membrane high,

yellowish, minutely papillose; operculum

conic; spores round, 15-17 pm, essentially

smooth. Fig. 1 11: 1-10.

Rhodobiyum umbraculum is known from eastern and

southern Africa. In the Flora area this species is collected in

forests of the southern and eastern Cape, Transkei, Natal,

northeastern Orange Free State, Zululand and eastern and

northern Transvaal. Map 156.

Vouchers: Crosby & Crosby 7905, 8094; Jacot Guil-

larmod PRE-CH13168; Kemp 849; Magill 5217, 6011;

Oliver 6722; Van Rooyl6, 1251, 1420, 2232; Von Breiten-

bach 12.

Rhodobryum umbraculum is easily recognized by its

darkish green, undulate, broadly obovate-spatnulate leaves

and costal anatomy.

395

MNIACEAE

Plants medium-sized to large, forming loose mats, dark green to yellow-green; terricolous or

saxicolous. Stems erect, densely leaved above, tomentose below; central strand present; sometimes

producing long-creeping, arching or prostrate, plagiotropic stolons, leaves distant. Leaves of stem

crisped ary, spreading wet; oblong to elliptical or spathulate; stolon leaves shorter, rounded;

margins generally bordered, frequently dentate, sometimes doubly dentate. Costa strong, guide

cells present, with one or two stereid bands. Laminal cells generally isodiametric, smooth; mar-

ginal cells elongated.

Perichaetia terminal on erect stems, poly setaceous. Seta elongate; capsule erect to pendulous;

peristome generally complete; operculum rostrate; calyptra small, cucullate, generally naked;

spores medium-sized.

Only one of the ten genera recognized in Mniaceae is present in the Flora area. The family is primarily Northern

Hemisphere in its distribution with its major species concentration in Asia.

PLAGIOMNIUM

Plagiomnium Koponen in Ann. Bot. Fenn. 5: 145 (1968); Smith, Moss FI. Brit. Irel. 439 (1978).

Type: P. cuspidatum (Fledw.) Kop.

Fertile stems erect, radiculose, producing plagiotropic stolons. Leaves large, margin with

unistratose border, dentate, teeth single. Costa with dorsal stereid band only.

A widespread genus with approximately 25 species found on soil or rock in moist or shaded habitats.

Plagiomnium rhynchophorum (Hook.)

Kop. var. reidii (Dix.) Kop. in Ann. Bot. Fenn.

18: 109 (1981). Type: Natal, Town Bush, Pie-

termaritzburg, Reia sub Sim 7552 (BM, holo. !;

PRE!).

Mnium rostratum Schrad. var. reidii Dix. in Trans. Roy.

Soc. S. Afr. 8: 204 (1920); Sim, Bryo. S. Afr. 342 (1926).

Mnium ecklonii C. Mull, in Bot. Ztg 13: 749 (1855);

Broth, in Naturl. PflFam. 10: 415 (1924). Type: Cape,

Winterberg, Ecklon s.n., 1825.

Plants medium-sized to large, in loose

mats, dark green; terricolous or saxicolous.

Stems erect, to 40 mm high, radiculose, densely

leaved above; in section round, central strand

large, cortical cells in 8-10 rows, large, thin-

walled, becoming smaller and thickened toward

margin. Plagiotropic stolons arising from vari-

ous parts of erect stem, long creeping with

leaves distant, anchored to substrate by rhizoid

tufts; in section rounded, central strand small,

inner cortical cells in 3-5 rows, large, thin-

walled, outer cortical cells in 2 rows, smaller,

thickened. Stem leaves larger and crowded to-

ward apex, somewhat contorted dry, wide-

spreading wet, undulate; oblong to oblong-spa-

thulate or narrowly elliptical, 5-b mm long;

apex rounded to truncate or emarginate, mucro-

nate; base narrow, slightly decurrent; margins

plane, dentate to denticulate, occasionally en-

tire. Costa strong below, tapering to apex, short

excurrent as mucro; in proximal section ellipti-

cal, guide cells 4-6, large, thin- walled, ventral

cells in 4 rows, first row above guide cells

large, thin-walled, 3 outer rows smaller, incras-

sate, dorsal stereid band in small group of ± 12

cells in centre of costa, dorsal cells in 2-3

rows, larger, incrassate; in distal section

rounded, guide cells large, distinct, dorsal ster-

eid band absent, ventral and dorsal cells simi-

lar, each in 2 rows, thickened. Stolon leaves

distant, somewhat scattered along stolon; orbi-

cular to broadly elliptical, (l-)2-4 mm long;

apex rounded-mucronate; margins plane, entire

to serrate, strongly bordered; costa as in stem

leaves. Laminal cells rounded, collenchyma-

tous, quadrate to hexagonal or short-rectangular

to short-rhomboidal, mostly 1-2:1, smooth,

cell walls yellowish; extreme basal cells rec-

tangular; marginal cells strongly differentiated,

in 3—4 rows, elongate, ± vermiculate,

thickened, yellowish; marginal teeth of 1-3

shorter cells.

Mniaceae

397

Synoicous. Perichaetia terminal on erect

stems, polysetaceous; leaves similar to upper

stem leaves. Seta erect, 14 — 16( — 20) mm long,

yellowish above, reddish below, arcuate dis-

tally; capsule horizontal to pendulous, ovate to

cylindrical-ovate, 3, 5-4, 5 mm long, yellow-

ish; exothecial cells quadrate to subhexagonal,

2 rows at mouth transversely rectangular; sto-

mata scattered on lower urn, cryptopore; peri-

stome double, reddish yellow, exostome teeth

16, triangular, 0,5-0, 6 mm long, strongly tra-

beculate, endostome yellowish, finely papil-

lose, basal membrane to 0,15 mm high, seg-

ments alternating with teeth, weakly keeled,

perforated above, cilia 3, as long as segments;

operculum long rostrate; spores rounded,

25-27 pm, yellowish, essentially smooth. Fig.

113: 1-12.

A member of the widespread P. rostratum complex, P.

rhynchophorum is known from Asia, India and Afnca. The

variety P. rhynchophorum var. reidii is widespread in moist

forests, woodlands and grasslands in the southern parts of

the Cape Province, Transkei, Natal, eastern Orange Free

State, Lesotho and the escarpment of the eastern Transvaal.

Map 157.

Vouchers: Crosby & Crosby 7667; Jacot Guillarmod

6058; Lambert 4; Magill 4369, 5553; Van Rooy 476, 1444,

2826.

The dark green mats formed by plagiotropic stolons

with distant, rounded leaves should quickly place most

southern African specimens of Plagiomnium. Specimens

with erect, fertile stems almost always have associated pla-

giotropic stolons. Dried fertile stems of Plagiomnium bear

some resemblance to Atrichum P. Beauv., however in the

Flora area specimens can easily be separated on leaf shape

or presence of the ventral costal lamellae produced by Atri-

chum.

The southern African taxon has for some time been

referred to P. rostratum (Hook.) Kop. (= Mnium rostra-

tum) and a broader interpretation of that species (cf. Gangu-

lee, 1974) would certainly include the southern African

specimens. Koponen (1972) has indicated however, that the

P. rostratum complex can be divided. It was therefore

decided to treat the southern African specimens as a variety

of P. rhynchophorum at this time.

FIG. 113 — Plagiomnium rhynchophorum var. reidii (1- 12): 1. habit, x 1; 2. habit, x 2,5; 3. portion of stem in

cross section; x 170; 4. portion of plagiotropic stolon in cross section, x 170; 5. upper stem leaf, x 12; 6. stolon leaf, x 12;

7. proximal stem leaf in cross section, x 170; 8. distal cross section of stem leaf costa, x 170; 9. basal leaf cells (right

side), x 175; 10. cells at upper stem leaf margin, x 700; 11. stem leaf apex, x 175; 12. part of capsule mouth showing

peristome, and side view of exostome tooth at right, x 70. Eustichia longirostris (13-20): 13. habit, x 1; 14. habit, x 5;

15. stem cross section, x 175; 16. leaves, x 70; 17. leaf in cross section, x 175; 18. basal leaf cells, papillae partly shown

(right side), x 175; 19. upper laminal cells at left margin, papillae partly shown, x 700; 20. leaf apex, x 175. (1-12,

Rennie 938; 13—20, Cholnoky 932). ^

399

EUSTICHIACEAE

Plants slender, forming dense tufts, light green; terricolous or saxicolous. Stems frequently

elongate; central strand present. Leaves distichous, keeled, ovate-cuspidate; margins erose-denti-

culate. Costa short excurrent; in section cells weakly differentiated, mostly stereids. Laminal cells

small, papillose.

Seta elongate; capsule suberect; exostome absent, endostome of 16 segments, weakly per-

forated, vertically striolate; operculum rostrate; calyptra cucullate.

The family consists of a single genus, Eustichia, that occurs at high altitudes in Central and South America and Africa.

EUSTICHIA

Eustichia Mitt, in J. Linn. Soc., Bot. 12: 603 (1869); Broth, in Natiirl. PflFam. 10: 421 (1924);

Sim, Bryo. S. Afr. 287 (1926); Bartram in Fieldiana 25: 185 (1949). Type species: E. longirostris

(Brid.) Brid.

With characters of family.

The genus contains 6 species; all except E. longirostris, the most widely distributed species, are restricted to the

Americas. In southern Africa the genus is found at high altitude in the Drakensberg.

Eustichia longirostris (Brid.) Brid.,

Bryol. Univ. 2: 789 (1827); Broth, in Natiirl.

PflFam. 10: 421 (1924); Sim, Bryo. S. Afr. 287

(1926). Type: Bourbon.

Pterigynandrum longirostrum Brid., Mant. Muse. 131

(1819). Diplostichum longirostre (Brid.) Mont, in Annls

Sci. nat.,bot. ser. 3,4: 117(1845).

Diplostichum africanum C. Mull, in Hedwigia 38: 53

(1899). Eustichia africana (C. Mull.) Par., Ind. Bryol.

Supp. 152 (1900); Broth, in Natiirl. PflFam. 10: 421

(1924); Sim, Bryo. S. Afr. 287 (1926). Type: Orange Free

State, above Kadziberg, Rehmann 279 (PRE!).

Plants small but occasionally elongated,

loosely caespitose, light green; terricolous.

Stems up to 50 mm long, mostly buried,

exposed chlorophyllose region 10-20 mm long,

irregularly branched, radiculose below; in sec-

tion subround, central strand present, inner cor-

tical cells in 2-3 rows, large, thin-walled, outer

cortical cells in 2 rows, stereids reddish yellow.

Leaves complanate and distichous, condupli-

cate to carinate-concave, crowded to somewhat

distant, appressed but little altered dry, erect

wet; ovate-cuspidate; margins plane, erose-den-

ticulate by projecting cell ends, frequently with

adjoining cells projecting to form stronger

tooth. Costa short excurrent, reflexed; in sec-

tion cells not strongly differentiated, consisting

of a single or double row of stereids with dorsal

and ventral surface cells slightly larger and in-

crassate or more strongly thickened on outer

wall. Laminal cells subquadrate to short-rec-

tangular, thickened, mostly 1-2 : 1, papillose,

cells with 3-4 low, blunt papillae over dorsal

Map 158. — • Eustichia longirostris

A Pyrrhobryum vallis-gratiae

surface, ventral surface smooth; basal cells rec-

tangular.

Sporophyte not known from southern

Africa. Fig. 113: 13-20.

Collected at high altitudes in South America, southern

Africa and the East African Islands. In the Flora area the

species is known from subalpine grasslands of the Drakens-

berg of Lesotho, Natal and the Orange Free State. Map 158.

Vouchers: Cholnoky 932; Esterhuysen 26174; Magill

5513.

The slender, light green plants, with distichous, keeled

leaves, form dense tufts on soil among rocks. The stems are

frequently very long and the lower buried portions are

yellowish brown and held together by interwoven rhizoids.

Eustichia africana, described from the Kadziberg in

the northeastern Orange Free State is a depauperate form of

this species.

401

RHIZOGONIACEAE

Plants small to large, forming loose tufts, yellowish green to dark green; growing on various

substrates. Stems erect, simple or highly branched, occasionally with stipe-like lower portion and

densely leaved upper portion, matted with dense tomentum at base or throughout lower stem; in

section with central strand. Leaves distichous or spirally arranged, margins frequently thickened,

often prominently toothed, sometimes doubly so, base occasionally decurrent. Costa strong, often

spinose dorsally; in section with guide cells and dorsal and ventral stereid bands. Laminal cells

isodiametric, incrassate.

Autoicous or dioicous. Perichaetia on short lateral branches at base of stem or above, leaves

differentiated. Seta erect, generally very long; capsules erect to inclined, short-necked; peristome

double, complete; operculum conic to conic-rostrate; calyptra cucullate; spores small.

A family of 9 genera confined mostly to the Southern Hemisphere. The family is recognized by its isodiametric laminal

cells, generally differentiated and toothed leaf margins, strong costa, and basal or occasionally lateral perichaetia. The

peristome is generally double and complete, however two genera, Hymenodon Hook. f. & Wils. and Hymenodontopsis

Hen., lack exostome teeth. Only a single genus, Pyrrhobryum, is present in southern Africa.

PYRRHOBRYUM

Pyrrhobryum Mitt, in J. Linn. Soc., Bot. 10: 174 (1868), emend. Manuel, Cryptogamie 1: 68

(1980). Lectotype: P. spiniforme (Hedw.) Mitt., fide Manuel (1980).

Rhizogonium Brid. p.p., Bryol. Univ. 2: 644 (1827); Sim, Bryo. S. Afr. 342 (1926).

Manuel (1980) separated the genus Pyrrhobryum from Rhizogonium on the basis of spirally arranged leaves with

multistratose borders, doubly toothed leaf margins, toothed dorsal costa and capsules with distinct necks. Both of the

species in southern Africa belongs to the segregate genus of 28 species found primarily in the Southern Hemisphere. The

widespread species P. spiniforme extends northward into southern North America and Asia.

1 Stems branching above; leaves appressed when dry, ovate-lanceolate, to 2 mm long 1 . P. vallis-gratiae

1 Stems simple; leaves crisped when dry, linear-lanceolate, 4-8 mm long 2.P. spiniforme

1. Pyrrhobryum vallis-gratiae (Hampe)

Manuel in Cryptogamie 1: 70 (1980). Type:

Cape, Gnadenthal, Breutel s.n. (BM, holo. !).

Mnium vallis-gratiae Hampe in Bot. Ztg 17: 205 (1859).

Rhizogonium vallis-gratiae (Hampe) Hampe ex Jaeg. in

Verh. St. Gall, naturw. Ges. 1873-74: 225 (1875); Broth,

in Natiirl. PflFam. 10: 428 (1924); Sim, Bryo. S. Afr. 344

(1926).

Plants small to medium-sized, caespitose,

dark green or yellow-green with age; terrico-

lous, saxicolous or corticolous. Stems 6-30

mm long, branching above short stipe, tomen-

tose at base, stipe dark brown, naked or with

small, distant, scale leaves; in section angular,

central strand present, inner cortical cells in

3—4 rows, thickened, outer cortical cells ste-

roids, in 2 rows, reddish. Leaves appressed dry,

erect-spreading wet; ovate-lanceolate, 1,8 -2,0

mm long; apex acuminate; margins plane,

doubly toothed above mid-leaf, bistratose at

base, multistratose above; in section forming

small knob, 3—4 cells in diameter. Costa per-

current, ventral surface smooth, cells rounded-

quadrate, dorsal surface sparsely spinose above

mid-leaf, cells short-rectangular; in proximal

section elliptical, guide cells 6-8, thickened,

ventral stereid band 2 cells thick, dorsal stereid

band 3 cells thick, ventral and dorsal surface

cells similar, thickened; in distal section sub-

triangular, guide cells 4, thickened, ventral

stereid band 1—2 cells thick, dorsal stereid

band 2 cells thick, ventral and dorsal surface

402

Rhizogoniaceae

Fig. 114. — Pyrrhobryum vallis-gratiae (1-12): 1.

habit, x 1; 2. habit, x 2,5; 3. part of stem in cross section,

x 350; 4. leaf, x 35; 5. leaf in cross section, x 175; 6.

basal leaf cells (right side), x 175; 7. median leaf cells, X

700; 8. leaf apex, x 175; 9. cells at upper right margin, x

700; 10. perigonial leaf, x 35; 11. perichaetial leaf, x 35;

12. part of capsule mouth showing peristome, x 175. P.

spiniforme ( 13 — 23): 13. habit, X 1; 14. habit, x 2; 15.

part of stem in cross section, x 350; 16. leaf, x 27; 17. leaf

cross section, proximal, x 175; 18. leaf cross section, dis-

tal, x 175; 19. basal leaf cells (right side), x 175; 20.

laminal cells at upper right margin, x 700; 21. median leaf

cells, x 700; 22. leaf apex, x 175; 23. portion of peri-

stome, X 120. (1 - 12, Esterhuysen 26387; 13-23 , Magill

5234).

Rhizogoniaceae 403

cells similar, thickened. Laminal cells rounded,

subquadrate, incrassate, somewhat irregular in

size and shape, mostly 1:1, smooth; basal cells

subquadrate, thickened.

Dioicous. Plants similar; perigonia gem-

mate, on short, lateral branches; leaves oval,

abruptly subulate, 1 mm long, margins weakly

toothed above. Perichaetia lateral on short

branches; leaves oblong to elliptical, abruptly

subulate, 2,5— 3,0 mm long; margins witn

single teeth in subula; lower laminal cells rec-

tangular, somewhat sinuolate, rounded and sub-

quadrate above mid-leaf. Seta 15-20 mm long,

yellowish; capsule inclined to horizontal, ar-

cuate-clavate, 2 mm long, reddish yellow;

exothecial cells subhexagonal, incrassate; sto-

mata present at base of urn, phaneropore; annu-

lus present; peristome double, exostome teeth

triangular, 0,5 mm long, brownish yellow, with

zig-zag median line, striate below, papillose

above, endostome fragile, light yellow, weakly

papillose, basal membrane high, segments as

high as teeth, narrowly slit along keel, cilia

shorter, 1-2; operculum conic with very short

apiculus, cells not twisted; calyptra not seen;

spores round, 20-25 pm, green, spiculate.

Fig. 114: 1-12.

Endemic to southern Africa, P. vallis-gratiae is found

in montane or kloof forests of the southwestern and south-

ern Cape. Map 158.

Vouchers: Brenan M2750; Crosby & Crosby 8154;

Esterhuysen 19182; Magill & Schelpe 4059, 4074.

The species is recognized by its doubly toothed, mul-

tistratose leaf margins, spinose dorsal costal surface and

isodiametric laminal cells. In comparison with P. spini-

forme, the leaves of P. vallis-gratiae are much shorter and

appressed when dry, giving the plants a julaceous appear-

ance. In addition the perichaetia are borne on short lateral

branches on the upper part of the stem, while in P. spini-

forme the perichaetia are borne at the base of the stems.

2. Pyrrhobryum spiniforme (Hedw.)

Mitt, in J. Linn. Soc., Bot. 10: 174 (1868);

Manuel in Cryptogamie 1: 69 (1980). Type:

Jamaica.

Hypnum spiniforme Hedw. Sp. Muse. 236 (1801).

Mnium spiniforme (Hedw.) C. Mull., Syn. Muse. 1: 175

(1848). Rhizogonium spiniforme (Hedw.) Bruch ex Krauss

in Flora 29: 134 (1846); Broth, in Natiirl. PflFam. 10: 428

(1924); Sim, Bryo. S. Afr. 344 (1926); Andrews in Grout,

Moss FI. N. Amer. 2: 260 (1933); Gangulee, Moss. E.

India 4: 1068(1974).

Plants medium-sized, caespitose, dark

green to yellow-green; terricolous, saxicolous

or corticolous. Stems 10—50 mm long; gener-

ally simple, radiculose at base, lower stem

forming a short, naked, dark brown stipe, when

young with distant scale leaves; in section an-

gular, central strand large, inner cortical cells in

6-8 rows, incrassate, outer cortical cells in

2-3 rows, strongly thickened, reddish. Leaves

crisped dry, spreading wet; linear-lanceolate to

lanceolate, (3-)5— 8 mm long; apex acumi-

nate; margins plane, doubly toothed by clear,

sharp teeth above base, bistratose below, mul-

tistratose above, in section forming knob, 3-4

cells in diameter. Costa percurrent, ventral sur-

face smooth, cells quadrate to short- rectangu-

lar, dorsal surface with distant spines, cells

short-rectangular to quadrate; in proximal sec-

tion elliptical, guide cells small, 6-8, incras-

sate, ventral stereid band strong, 3-4 cells

thick, dorsal stereid band 2-3 cells thick, ven-

tral and dorsal surface cells slightly larger, in-

crassate; in distal section subtriangular, guide

cells 4—6, incrassate, ventral stereid band 2—3

cells thick, dorsal stereid band 2-3 cells thick,

ventral and dorsal surface cells slightly larger,

incrassate. Laminal cells somewhat variable in

size and shape, rounded, quadrate to subhexa-

gonal, mostly 1( — 2): 1 , thickened, smooth; ba-

sal cells quadrate.

Synoicous. Paraphyses numerous, fili-

form. Perichaetia on short lateral branch at base

of vegetative shoot; leaves obovate to elliptical,

abruptly constricted to short subula, 1,5 -2,0

mm long, margins serrate in subula, cells fu-

siform, mostly 10:1, incrassate. Seta erect to

curved, 10-50 mm long, reddish yellow; cap-

sule inclined to horizontal, clavate-arcuate, to 3

mm long, reddish yellow, gradually narrowing

to a short neck, exothecial cells short-rectangu-

lar, somewhat shorter at mouth, reddish; sto-

mata present on neck, phaneropore; annulus

present; peristome double, yellowish, exostome

teeth triangular, 0,6-0, 7 mm long, with weak

median zig-zag line, striate below, coarse lv pa-

pillose above, endostome fragile, light yellow,

weakly papillose, basal membrane high, seg-

ments as nigh as teeth, slit along keel, cilia

shorter, 1 or 2; operculum obliquely rostrate,

beak to 1 ,5 mm long; calyptra not seen; spores

round, 20-25 pm, green, spiculate. Fig. 114:

13-23.

404

Rhizogoniaceae

Although previously considered a widespread Sou-

thern Hemisphere species, recent authors (Scott & Stone,

1976; Sainsbuiy, 1955) have questioned its presence in

Australia and New Zealand. In southern Africa the species

is found in forests of the southwestern, southern and eastern

Cape, Transkei, Natal, Zululand and the eastern and nor-

thern Transvaal. Map 159.

Vouchers: Crosby & Crosby 13395; Magill 5234,

6040; Schelpe 7862; Van Rooy 829; Von Breitenbach 79.

This species is easily recognized by its stem that con-

sists of a lower, ± naked stipe-like portion with scattered

scale leaves and an unbranched, densely leaved upper por-

tion. The long, narrow, strongly toothed leaves are crisped

when dry, thus separating it from the other southern African

species. Pyrrhobryum spiniforme is closely related to the

droicous Australian species P. paramattense (C. Miill.) Ma-

nuel (cf. Scott & Stone, 1976, as Rhizogonium).

AULACOMNIACEAE

405

Plants medium-sized, in tufts, dark green to yellowish green, growing on a variety of sub-

strates. Stems erect, branched, tomentose; central strand present. Leaves larger above, ovate-lan-

ceolate, cuspidate; margins plane, irregularly toothed above. Costa short-excurrent; in section with

guide cells and two stereid bands. Laminal cells small, irregularly isodiametric, incrassate,

smooth.

Dioicous. Perichaetia terminal. Seta elongate; capsule erect, cylindrical, striate; peristome

double, exostome teeth papillose with median zig-zag line, endostome complete, segments keeled,

as long as teeth, cilia single, shorter than teeth; operculum conical; calyptra cucullate; spores

small.

The family contains two genera, Aulacomnium and Leptotheca-, only the latter is known from southern Africa. Two

very widely distributed species of Aulacomnium are known from eastern Africa, A. palustre (Hedw.) Schwaegr. and A.

turgidum (Wahlenb.) Schwaegr.; they may be separated from Leptotheca by their sharply papillose leaf cells and costa

ending below the leaf apex.

LEPTOTHECA

Leptotheca Schwaegr., Sp. Muse. Suppl. 2: 135 (1824); Broth, in Natiirl. PflFam. 10: 440

(1924); Sim, Bryo. S. Afr. 299 (1926); Sainsb., N. Zeal. Mosses 287 (1955). Type species:

L. gaudichaudii Schwaegr.

With characters of family.

The genus contains 4 species, two endemic to the Americas, one Australian and L. gaudichaudii which is known from

southern South America, southern Africa, Australia and New Zealand.

Leptotheca gaudichaudii Schwaegr., Sp.

Muse. Suppl. 2: 135 (1824); Broth, in Natiirl.

PflFam. 10: 440 (1924); Sim, Bryo. S. Afr. 300

(1926); Sainsb., N. Zeal. Mosses 287 (1955);

Scott & Stone, Moss. S. Aust. 306 (1976).

Type: Australia.

Aulacomnium gaudichaudii (Schwaegr.) Mitt, in Hook-

er’s J. Bot. Kew Gdn Misc. 8: 262 (1856).

Plants medium-sized, caespitose, light

green to dark green, glossy; terricolous, saxico-

lous or corticolous. Stems 10—30 mm long,

iiTegularly branched, radiculose below; in sec-

tion subround, central strand very small, inner

cortical cells large, in 3-4 rows, weakly thick-

ened, outer cortical cells stereids, in 2 rows,

reddish. Leaves appressed and ± curved dry,

erect-spreading wet; ovate-lanceolate, 2-4 mm

long; apex acute, cuspidate; margins plane, ser-

rate above mid-leaf. Costa short-excurrent; in

section subround, guide cells 4-6, strongly

thickened, ventral stereid band 1 cell thick,

ventral surface cells substereids, dorsal stereid

band 2 cells thick, dorsal surface cells subste-

reids. Laminal cells variable in size and shape,

rounded to subquadrate, incrassate, mostly 1:1,

smooth; basal cells slightly larger, rounded-rec-

tangular, thickened. Gemmae infrequently pro-

duced, axillary in upper leaves and at apex,

filamentous, 12-14 cells long, yellow-brown.

406

Aulacomniaceae

Sporophyte not known from southern

Africa. Fig. 115.

Leptotheca gaudichaudii is known from Patagonia,

Terra del Fuego, Falkland Islands, southern Australia, Tas-

mania, New Zealand and southern Africa. In the Flora area

the species is restricted to rock recesses and crevices in the

mountain fynbos of the southwestern and southern Cape.

Map 160.

Vouchers: Brenan M2770; Esterhuysen 20038; Magill

5933,6338.

The glossy plants with ovate-lanceolate, cuspidate

leaves and small, smooth and incrassate leaf cells will help

to place specimens of L. gaudichaudii. Plants with gemmae

are infrequently found in southern Africa, however when

present the copious production of the long, brown, fila-

mentous gemmae, in the axils of the upper leaves, is very

obvious.

Fig. 115. — Leptotheca gaudichaudii: 1. habit, x 1;

2. habit, x 10; 3. stem in cross section, x 280; 4. leaves, x

50; 5. leaf in cross section, x 220; 6. basal leaf cells (right

side), x 175; 7. median leaf cells, x 700; 8. leaf apex, x

175; 9. portion of stem showing axillary, filamentous gem-

mae, X 87. (1—9, Esterhuysen 30969a).

407

BARTRAMIACEAE

Plants small to large, occasionally robust, caespitose; terricolous or saxicolous. Stem erect,

generally branching by subperichaetial innovation; central strand present, epidermis differentiated,

cells large, thin-walled, fragile. Leaves lanceolate to ovate, occasionally broader, lamina infre-

quently bistratose; apex acuminate to subulate, or infrequently acute to obtuse or rounded; base

scarcely differentiated or strongly differentiated with lamina reflexed above; margins plane to

recurved, entire to crenulate or serrate, frequently doubly so. Costa percurrent to short-excurrent,

rarely ending just below apex. Laminal cells rectangular to linear, generally incrassate, papillose or

prorate, papillae centered over lumen or displaced toward end of cell; basal cells rectangular or

quadrate, thin-walled, generally smooth.

Autoicous or dioicous. Perichaetia terminal, frequently overgrown by subperichaetial

branches. Seta elongate, erect; capsule erect to inclined or horizontal, urn globose, smooth or

rarely with wart-like projections, rugulose to furrowed or sulcate dry; peristome absent or present,

single or double, endostome with segments and cilia or frequently rudimentary; operculum convex

to conic or rostrate; calyptra cucullate, quickly lost; spores reniform or rounded, papillose or

warty.

Eight of the ten genera of Bartramiaceae are known from southern Africa. Members of the family are recognized by the

generally dull appearance of the appressed or wide-spreading leaves, and common occurrence of reddish tomentum on the

stem; rectangular, generally incrassate leaf cells with papillae mostly displaced toward cell ends, and especially by then-

globose or rounded, frequently inclined capsules.

1 Stem in cross-section triangular; leaf cells minutely verruculose-striolate 1 . Plagiopus

1 Stems rounded to angular in cross-section, not triangular; leaf cells smooth, papillose or

prorate:

2 Vegetative leaves less than 2,5 mm long:

3 Costa absent or rudimentary 5. Quathlamha

3 Costa well developed, extending above apex:

4 Leaves appressed in 5 distinct rows, ± triangular; costa abruptly excurrent as short

awn; peristome single, teeth joined above 4. Conostomum

4 Leaves variously erect to spreading, not in distinct rows; costa occasionally excur-

rent, never abruptly so; peristome absent or double, teeth not joined above:

5 Leaves mostly 1,5— 2,5 mm long; stems with few subperichaetial innovations;

capsule sulcate dry, mouth broad, peristome absent 3. Bartramia

5 Leaves mostly less than 1 ,5 mm long; stems with numerous subperichaetial in-

novations:

6 Capsules erect, globose or warty, little altered dry, gymnostomous, mouth

small 6. Bartramidula

6 Capsules inclined or horizontal, globose to ellipsoidal, striate to sulcate dry,

mouth wide, peristome present 7. Philonotis

2 Vegetative leaves (2,5-) 3, 0-6,0 mm long:

7 Leaves with distinct border of narrow, elongated cells 7. Philonotis

7 Leaves without distinct border:

8 Plants generally large, occasionally robust; leaves plicate in base or plications ex-

tending to upper leaf 8. Breutelia

8 Plants smaller; leaves without well defined plications:

9 Upper lamina bistratose:

408

Bartramiaceae

Bartramiaceae

409

10 Leaf base differentiated, hyaline and sheathing; plants dull, glaucous; leaf

margin serrate by projecting cell ends 3. Bartramia

10 Leaf base not differentiated; plants green; leaf margins denticulate, teeth for-

med by differentiated marginal cells 2. Anacolia

9 Upper lamina unistratose:

11 Leaves ovate- to oval-acuminate, some weakly plicate; peristome present,

double 8. Breutelia

11 Leaves linear-lanceolate to ovate- or oblong-lanceolate, lamina not plicate;

peristome absent 3 . Bartramia

1. PLAGIOPUS

Plagiopus Brid., Bryol. Univ. 1: 596 (1826); Broth, in Naturl. PflFam. 10: 448 (1924); Flowers in

Grout, Moss FI. N. Amer. 2: 157 (1935); Gangulee, Moss. E. India 4: 1080 (1974); Smith, Moss

FI. Brit. Irel. 454 (1978). Type species: P. serratus Brid.

Plants medium-sized, caespitose; saxicolous. Stems erect, irregularly branched, tomentose

below; in section triangular, central strand present, epidermal cells distinct. Leaves narrowly

lanceolate; margins recurved, doubly serrate above. Costa percurrent. Laminal cells rectangular,

papillose with numerous low, short ridges; basal cells slightly differentiated.

Synoicous. Seta elongate; capsule globose, weakly striate dry; peristome double; operculum

plano-convex; calyptra cucullate; spores warty.

Plagiopus contains two species, P. crassinervius (Mitt.) Broth, which is endemic to New Zealand and P. oederi

(Brid.) Limpr. which is widespread in Asia, Europe and some parts of North and South America. This is the first report of

Plagiopus from Africa. The genus is restricted to the Afro-alpine regions of the Drakensberg and the high grass-heatnlands

of Lesotho.

The genus resembles Bartramia but is separated from it, and the other genera of Bartramiaceae, by the triangular shape

of the stem in cross-section and the unusual leaf cell ornamentation.

Plagiopus oederi (Brid.) Limpr., Laubm.

Deutschl. 2: 548 (1895); Broth, in Naturl.

PflFam. 10: 449 (1924); Flowers in Grout,

Moss FI. N. Amer. 2: 157 (1935); Gangulee,

Moss. E. India 4: 1081 (1974); Smith, Moss FI.

Brit. Irel. 454 (1978). Type: Europe.

Bartramia oederi Brid., Muse. Rec. 2: 135(1803).

Plants medium-sized, caespitose, light

green to yellow-green above, reddish brown be-

low; saxicolous. Stems erect, 20-30 mm tall,

irregularly branched above, lower stems cov-

ered with reddish tomentum, radicles papillose

with numerous large, high papillae; in section

distinctly three-sided, subtriangular, central

strand very small, inner cortical cells in 6-8

rows, large, thin- walled, reddish, outer cortical

cells in 2 rows, stereids, reddish yellow, epider-

mal cells differentiated, thin-walled, fragile,

quickly eroded. Leaves erect and weakly flexu-

ose dry, erect-spreading wet; narrowly lanceo-

late, 2-4 mm long; apex acuminate; margins

recurved from base to upper leaf, entire below,

sharply serrate above, frequently doubly ser-

rate. Costa percurrent, superficial cells long-

rectangular, papillose; in section bulging dor-

Fig. 1 16. — Plagiopus oederi (1-13): 1. habit, x 1; 2. habit, x 3; 3. stem in cross section, x 87; 4. leaves, x 35; 5.

leaf in cross section, x 350; 6. basal leaf cells (left side), x 175; 7. upper laminal cells, x 700; 8. marginal leaf cells (right

side), x 700; 9. leaf apex, X 175; 10. capsule, wet, x 6; 11. capsule, dry, x 6; 12. part of capsule mouth showing cells

and peristome, x 140; 13. spore, x 700. Anacolia breutelii (14-27): 14. habit, dry (upper plant var. squarrifolia, lower

plant var. breutelii), x 1; 1$. habit, dry (var. squarrifolia), x 2; 16. part of stem in cross section, x 175; 17. lower leaf,

X 35; 18. upper leaf, X 35; 19. leaf in proximal cross section, X 350; 20. leaf in distal cross section, X 350; 21. basal leaf

cells (left side), x 175; 22. marginal leaf cells (right side), x 700; 23. leaf apex, x 175; 24. capsule, dry, x 6; 25. capsule,

wet, x 6; 26. part of capsule mouth showing cells and peristome, X 140; 27. exothecial cells, x 700. [1 — 13, VanZinderen

Bakker 446; 14 (upper plant) & 15 ,Pillans 10032; 14 (lower plant) & 16-27, Breutel s.n).

410

Bartramiaceae

sally, subround, guide cells 2-4, large, thick-

ened, ventral cells in single layer, smaller than

uide cells, thickened, dorsal substereid band

-4 cells thick, dorsal surface cells not differ-

entiated. Upper laminal cells rectangular, 4-6:

1, thickened, minutely verruculose-striolate, as

numerous low, short ridges, occasionally a few

cells also prorate distally; basal cells slightly

longer rectangular, minutely verruculose-strio-

late; alar cells not differentiated.

Perichaetia terminal, leaves similar to

vegetative leaves; basal leaf cells long-rectang-

ular to linear, smooth, yellowish. Seta 8-12

mm long, reddish yellow; capsule weakly in-

clined, urn globose, 2,0-2, 5 mm long, weakly

striate dry, reddish yellow, mouth frequently

oblique; exothecial cells large, hexagonal to

rectangular or quadrate, thin- walled; stomata

present at base of urn, phaneropore to subpha-

neropore; peristome double, exostome teeth 16,

narrowly triangular, 0,3 mm high, papillose,

with median zig-zag line, trabeculae strong, en-

dostome segments 16, fragile above, finely pa-

pillose; operculum plano-convex; calyptra cu-

cullate; spores subround to ovoid, 15-27 gm,

warty, yellow-brown. Fig. 116: 1-13.

New to Africa, the species has recently been collected

on rock in the subalpine grasslands of Natal and northern

Map 161. — # Plagiopus oederi

A Anacolia breutelii var. squarrifolia

and eastern Lesotho. Plagiopus oederi is also known from

Europe, Asia, North America and Oceania. Map 161.

Vouchers: Esterhuysen 26178, 35929; Meyer 1040D;

Van Rooy 1302, 1340; Van Zinderen Bakker 446.

In addition to the stem being ± triangular in cross-

section, the very distinctive leaf cell ornamentation will

identify this species. The cell surfaces are covered by nu-

merous low ridges that give an impression of small papillae

that were stretched by elongation of the cells. In cross-sec-

tion the cells appear to be covered with numerous low, blunt

papillae.

2. ANACOLIA

Anacolia Schimp., Syn. Muse, europ. edn 2: 513 (1876), nom. cons.; Broth, in Natiirl. PflFam.

10: 449 (1924); Flowers in Grout, Moss FI. N. Amer. 2: 154 (1935); Flowers in Bull. Torrey bot.

Club 79: 161 (1952); Gangulee, Moss. E. India 4: 1083 (1974). Lectotype species: A. webbii

(Mont.) Schimp., fide Flowers in Bull. Torrey bot. Club 79: 161 (1952).

Plants medium-sized, caespitose; terricolous or saxicolous. Stems erect, sparsely branched,

tomentose below; central strand present, epidermis not differentiated. Leaves linear-lanceolate,

lamina bistratose above; base not strongly differentiated; margins denticulate. Costa short-excur-

rent. Laminal cells short-rectangular, prorate; basal cells long-rectangular, weakly differentiated.

Dioicous. Perichaetia becoming lateral through innovation. Seta elongate; capsule inclined,

subglobose wet, rugulose dry; peristome double, endostome rudimentary; operculum convex;

calyptra cucullate; spores warty.

The nine species of Anacolia are found primarily in the Americas and Africa, although they are also known from

Europe and parts of Asia.

In southern Africa, the genus is recognized by having stems without a differentiated epidermis, coarsely serrate or

denticulate leaf margins, bistratose laminas and undifferentiated leaf bases. In addition the capsules are rugulose or

wrinkled when dry and the southern African species has a double peristome. Anacolia breutelii could be mistaken for a

Bartramia but in addition to the characters listed above, it is a slightly larger plant with a coarser look and texture.

Bartramiaceae

411

Anacolia breuteiii (C. Miill.) Magill,

comb. nov. Type: Cape, Soutkloof, Breutel

s.n. (BM!;G!).

Bartramia breuteiii C. Mull, in Bot. Ztg 16: 162 (1858).

Breutelia breuteiii (C. Miill.) Broth, in Natiirl. PflFam. 10:

470(1924).

Bartramia afrostricta C. Miill. in Hedwigia 38: 94

(1899). Syntypes: Cape, Cape Town, Rehmann 203 & 204

(PRE!).

Plants medium-sized, loosely caespitose,

green to yellow-green or reddish above, brown-

ish to blackish brown below; terricolous or saxi-

colous. Stems erect, 10—30 mm tall, sparsely

branched, reddish tomentum on lower stem; in

section round, central strand present, inner cor-

tical cells in 6-8 rows, thin-walled, yellowish,

outer cortical cells in 2 rows, cells small, ster-

eids or substereids, reddish to dark red, epider-

mal cells not differentiated. Leaves closely set,

generally appressed but infrequently squarrose-

recurved dry, erect-spreading to squarrose wet;

linear-lanceolate, (2,0-)2,5-3,5(-4,5) mm

long, ventral surface flat over costa, lamina bi-

stratose juxtacostally and in irregular patches

just above base, completely bistratose in upper

leaf; apex subulate; margins reflexed to re-

curved, doubly denticulate in subula by large,

smooth, clear cells, distinct from laminal cells.

Costa short-excurrent as denticulate awn, occa-

sionally reddish below, ventral superficial cells

rectangular, prorate to sharply dentate in upper

subula; in section bulging dorsally, subtriangu-

lar, guide cells 6, small but distinct, ventral

stereid to substereid band 1-3 cells thick, ven-

tral surface cells smaller than guide cells, in-

crassate, prorate, dorsal stereid to substereid

band strong, 3-4 cells thick, dorsal surface

cells incrassate, prorate. Laminal cells rectang-

ular, 1, 5-3,0: 1, weakly thickened, strongly or

weakly prorate distally and/or proximally; basal

cells long-rectangular, to 10: 1, thin-walled,

smooth, yellowish; alar cells weakly differen-

tiated, quadrate, clear.

Perigonia terminal, gemmate. Perichaetia

terminal, becoming lateral through innovation;

leaves similar to vegetative leaves, elliptical-

subulate. Sporophytes rare. Seta 7 mm long,

yellowish; capsule inclined, urn subglobose, 2

mm long, rugulose dry, red-brown; exothecial

cells subhexagonal, thickened in comers; sto-

mata present at base of um, phaneropore; peri-

stome double, exostome teeth 16, narrowly

triangular, 0,3 mm long, papillose, reddish yel-

low, endostome rudimentary, irregular and very

short, attached to teeth, smooth; operculum not

seen; young calyptra cucullate, smooth; spores

rounded, 12-18 fim, warty, yellowish. Fig.

116:14-27.

Considerable confusion has arisen around the use of

the name Bartramia substricta sensu Sim (1926) and Dixon

(1920). A careful examination of type specimens at BM and

G has confirmed Muller’s descriptions and label data for

Bartramia breuteiii C. Miill. and B. substricta Schimp. ex

C. Mull.

The problem seems to have been created through mis-

takes in interpretation and observation. The specimens of A.

breuteiii are macroscopically similar to the European spec-

ies Bartramia stricta Brid. These southern African speci-

mens were coupled through this superficial similarity to

Muller’s name B. substricta by Dixon and Sim, and the

name B. breuteiii was apparently overlooked. The state-

ment by Dixon (1920), “B. afrostricta C. M. is entirely

identical with B. substricta Schimp. ex C. Miill., as a com-

arison of Rehm. M. Austr.-Afr. 203, 204, 205, with

chimper’s type as Kew shows”, further complicated the

problem since the syntypes of the former are Anacolia and

the type of the latter is a Breutelia. Since my observations

are compatible with Muller’s descriptions, it seems unlikely

that Dixon made more than a casual examination of the

specimens involved.

In the generic configuration used here Bartramia breu-

teiii is treated under Anacolia, while B. substricta is

transferred to Breutelia. As a result of incorrect use of the

name Bartramia substricta by Sim and Dixon, all of the

specimens of Anacolia breuteiii that were examined had

been placed under Bartramia substricta; while specimens of

Breutelia substricta had been placed under a later name, i.e.

Breutelia afroscoparia.

Presently known only from southern Africa, the species

is infrequently collected on rock or soil over rock at higher

elevations in the mountains of the southwestern and north-

western Cape and the Drakensberg of Natal, Lesotho and

Orange Free State. Two varieties are recognized; their dis-

tributions are practically sympatric .

1 Leaves erect to appressed dry var. breuteiii

1 Leaves squarrose-recurved dry var. squarrifolia

var. breuteiii.

Leaves erect to appressed dry, erect-

spreading wet; mostly 2—4 mm long.

Endemic to southern Africa, the typical variety is fre-

quently collected in the mountains of Lesotho, eastern

Orange Free State, Natal and the southern, southwestern

and northwestern Cape. Map 162.

Vouchers: Esterhuysen 20204; Magill 4396, 4713,

6092; Oliver 7237; Van Rooy 389.

The taxon is vegetatively similar to A. abyssinica

(C. Mull.) Flow., but sporophytically it is more closely

related to A. menziesii (Turn.) Par. The presence of a peri-

stome excludes this taxon from A. abyssinica.

412

Bartramiaceae

Bartramia squarrifolia Sim, Bryo. S. Afr. 302 (1926).

Leaves squarrose-recurved dry, wide-

spreading to squarrose wet; mostly 3—5 mm

long.

Endemic to southern Africa, this variety is rarely col-

lected in the Drakensberg of western Natal and a few sites in

the mountains of the southwestern Cape. Map 161 .

Vouchers: Hilliard <£ Burtt 10115; Magill & Schelpe

3849; Pillans 10032.

Similar to the typical variety in all respects except for

the squarrose spreading leaves that give the plants a very

distinctive appearance. Macroscopically the plants resemble

Bartramia capensis but are easily separated from that spec-

ies by differences in leaf cells, serration of the leaf margins

and bistratose lamina.

var. squarrifolia (Sim) Magill, stat. nov.

Lectotype: Cape, Lion’s Rump, Pillans 4089

(PRE, lecto. selected here!; BM!; BOL!).

3. BARTRAMIA

Bartramia Hedw., Sd. Muse. 164 (1801), nom. cons.; Broth, in Natiirl. PflFam. 10: 451 (1924);

Sim, Bryo. S. Afr. 300 (1926); Flowers in Grout, Moss FI. N. Amer. 2: 157 (1935); Sainsb.,

N. Zeal. Mosses 300 (1955); Gangulee, Moss. E. India 4: 1086 (1974); Scott & Stone, Moss. S.

Austr. 322 (1976); Smith, Moss FI. Brit. Irel. 454 (1978); Catcheside, Moss. South Austr. 279

(1980). Lectotype species: B. halleriana Hedw., fide Flowers in Grout, Moss FI. N. Amer. 2: 158

(1935).

Plants small to medium-sized, caespitose; terricolous or saxicolous. Stems erect, branches

few, generally subperichaetial innovations, sparsely tomentose below; in section round, central

strand present, epidermal cells ± distinct. Leaves linear to lanceolate or subulate above broad,

sheathing base; lamina unistratose or bistratose; margins mostly serrate. Costa short- to long-

excurrent. Laminal cells rectangular to long-rectangular, ± incrassate, prorate or papillose; basal

cells scarcely differentiated or very strongly differentiated.

Dioicous, autoicous or synoicous. Perichaetia becoming lateral through innovation. Seta short

or long; capsules ± inclined, globose to ovoid wet, sulcate 017; peristome absent; operculum conic

or convex-apiculate; calyptra cucullate, small; spores warty.

The genus contains approximately 90 species and is found on every continent, including Antarctica. Major centres of

described species are in Central America and western South America.

Bartramia is separated from the other southern African genera of the family by its long, slender leaves and absence of a

peristome although peristomes are generally present in the genus. It is unlikely to be confused with any other genus except

Anacolia ; see note there.

1 Leaves with well defined, hyaline, sheathing base; lamina bistratose above base 4. B. hampeana

1 Leaves without strongly differentiated base; lamina unistratose:

2 Leaves ovate-acuminate; laminal cells short-rectangular, incrassate, ± flexuose; leaf base to 20 cells wide on each

side of costa 3. B. aristaria

Bartramiaceae

413

2 Leaves linear-lanceolate; laminal cells short- to long-rectangular, not flexuose; leaf base less than 15 cells wide on

each side of costa;

3 Upper laminal cells short-rectangular, ± smooth to weakly prorate; costa percurrent to short-excurrent; spores

25 — 32 pm in diameter 1 . B . compacta

3 Upper laminal cells long-rectangular throughout, with single high papilla over lumen or displaced towards

proximal end of cell; costa short- to long-excurrent; spores to 50 /am in diameter 2. B. capensis

1. Bartramia compacta Hornsch., Hort.

Phys. Berol. 63 (1820); Broth, in Natiirl.

PflFam. 10: 457 (1924); Sim, Bryo. S. Afr. 303

(1926). Type: Cape, Lion’s Mountain, Bergius

s.n. (BM!).

Bartramia kraussii B.S.G., Bryol. Eur. 4: 40 (1842).

Type; Cape, Cape Town, Krauss s.n. (BM!; MANCH!).

IBartramia subasperrima C. Miill. in Hedwigia 38: 95

(1899); Broth, in Natiirl. PflFam. 10; 457 (1924). Type;

Cape, Cape Town, Rehmann 213; vide note below.

Plants small, caespitose, yellow-green to

light green or glaucous green; temcolous.

Stems 3—10 (—15) mm high, radiculose below;

in section round, central strand large, inner cor-

tical cells in 3—4 rows, large, thin-walled,

outer cortical cells in 1-2 rows, smaller,

weakly thickened, yellow-brown, epidermis not

differentiated or outer walls not as strongly

thickened. Leaves erect dry, erect-spreading to

widespreading wet; linear-lanceolate, (2,0-)

2,5— 3,0 mm long, lamina unistratose; apex

acuminate; margins reflexed, doubly serrate

above base. Costa percurrent to short-excur-

rent, superficial cells rectangular, smooth to

prorate ventrally, dentate above dorsally; in

section subround to triangular, bulging dor-

sally, guide cells 4, larger than other cells, in-

crassate, ventral cells in 1—3 rows, smaller

than guide cells, incrassate, dorsal stereid band

3-5 rows thick, dorsal surface cells not differ-

entiated or substereids. Upper laminal cells rec-

tangular, mostly 4-6 (-8): 1, weakly thick-

ened, weakly prorate; basal cells rectangular,

10: 1 , thin-walled; alar cells quadrate, not form-

ing distinct group.

Synoicous. Perichaetia lateral through in-

novation, leaves not differentiated. Seta 7—10

mm long or infrequently short, 2 mm long, red-

dish yellow; capsule ± erect, urn globose and

smooth wet, broadly short-cylindrical and sul-

cate dry, 1,0- 1,2 mm long, yellowish with

dark red mouth; exothecial cells subhexagonal,

weakly thickened or thin-walled with slightly

thickened comers, 2-4 rows of cells at mouth

smaller, quadrate to transversely rectangular;

stomata present at base of um, phaneropore;

peristome absent, occasionally with fragile,

hyaline fringe at mouth; operculum convex-api-

culate, more strongly apiculate dry; calyptra not

seen; spores subround, 25—32 /um, warty, yel-

lowish brown. Fig. 117: 1-12.

Dixon & Gepp (1923) suggested that Muller (1899)

was in error citing Rehmann 213 as the type for B. subas-

perrima as it would be out of place in Rehmann 's exsiccate.

However, it appears that Muller was not citing a specimen

from the exsiccate (Coll. Muse, austro-africani No. or Coll.

Muse. A. A. No.) but rather a number from a packet sent to

Muller by Rehmann prior to the distribution of the exsiccate

(Coll. No. 213). Dixon & Gepp’s reference to Rehmann’s

exsiccate No. 191 is also a mistake since in their list and on

the specimen, the number for B. subasperrima is 199. Most

of the evidence at this time indicates that the Rehmann

exsiccate specimens may represent isotype material; see

also Codd & Gunn ( 1982).

Endemic to southern Africa, the species is collected on

soil in mountains of the Cape Province. Two varieties are

recognized:

1 Seta very short, to 2 mm long; capsules immersed

to emergent var. macowaniana

1 Seta longer, 7-10 mm long; capsules exserted

var. compacta

var. compacta.

Stems mostly 5-10 mm high. Seta 7-10

mm long, exserting capsule above plants.

This variety is infrequently collected in the mountains

of the southwestern, northwestern and eastern Cape. A

single specimen is also known from the northern Cape. Map

Vouchers: Garside 6546, 6680; Magill & Schelpe

4016.

The short, rigidly erect, unistratose leaves, without

strongly differentiated bases, and the gymnostomous cap-

sules that are strongly sulcate when dry will separate speci-

mens of B. compacta from most members of the family.

The typical variety is separated from B. compacta var.

macowaniana primarily by its longer seta, and from B.

capensis by its narrower, more consistently erect leaves,

smooth to only weakly prorate leaf cells and much smaller

spores; see note under that species.

var. macowaniana (C. Miill.) Magill,

stat. nov. Type: Cape, Somerset East, Bosch-

berg, MacOwan s.n., 1877 (NBG!;NH!).

414

Bartramiaceae

Bartramiaceae

415

Map 163. — • Bartramia compacta var. compacta

▲ Bartramia compacta var. macowaniana

Bartramia macowaniana C. Miill. in Hedwigia 38: 95

(1899); Broth, in Natiirl. PflFam. 10: 457 (1924); Sim,

Bryo. S. Afr. 303(1926).

Stems mostly less than 5 mm long. Seta to

2 mm long; capsule just emergent.

Known only from the type locality in the southeastern

comer of the central Cape Province. Map 163.

Voucher: J. Sim 10149.

Gametophytically this variety is indistinguishable from

the typical variety. It is provisionally maintained at the rank

of variety on the basis of its consistently short seta. This

results in emergent capsules which are ‘nestled’ in the

leaves along the stem and give the plants a rather distinctive

appearance. The geographical isolation of the population is

also interesting, nowever the plants have only been col-

lected twice, in 1877 and 1921.

2. Bartramia capensis (R. Br.) Wijk &

Marg. in Taxon 8: 71 (1959). Type: Cape,

Table Mtn, R. Brown s.n. (BM!; MANCH!).

Glyphocarpa capensis R. Br. in Trans. Linn. Soc. Lond.

12: 575 (1819). Gymnostomum capense (R. Br.) Hook.,

Musci Exot. 2: 165 (1819). Glyphocarpus capensis ( R. Br.)

Brid., Bryol. Univ. 2: 91 (1827).

Bartramia quadrata Hook., Musci Exot. 2: 132 (1820).

Glyphocarpus quadratus (Hook.) Brid., Bryol. Univ. 2: 92

(1827). Type: Cape, George, Auteniqualand, Burchell s.n.

(E, holo.; BM!).

Bartramia sericea Homsch., Hort. Phys. Berol. 63

(1820); Broth, in Natiirl. PflFam. 10: 457 (1924); Sim,

Bryo. S. Afr. 303 (1926). Glyphocarpus sericeus

(Homsch.) Jaeg. in Verh. St. Gall, naturw. Ges. 1873—74:

62 (1875) nom. illeg. Type: Cape, Devil’s Peak, Bergius

s.n., vide Wijk & Marg., Ind. Muse. 5: 334 (1969).

Bartramia inserta Sull. & Lesq. in Proc. Am. Acad. Arts

Sci. 4: 279 (1859); Broth, in Natiirl. PflFam. 10: 457

(1924). Glyphocarpus insertus (Sull. & Lesq.) Jaeg. in

Verh. St. Gall, naturw. Ges. 1873-74: 62 (1875). Type:

Cape, Simonstown, C. Wright s.n., 26 Oct. 1853 (FH,

holo.!).

Breutelia aristaria (C. Mull.) Broth, var. plumosa Sim in

Trans. R. Soc. S. Afr. 15: 312 (1926), vide Dix. in Trans.

R. Soc. S. Afr. 8: 206 (1920) and C. Miill. in Hedwigia 38:

93 (1899). Type: Natal, Inanda, Rehmann 185 (G!).

Plants small, caespitose, light green; terri-

colous. Stems 5-8 mm high, sparsely

branched, with reddish tomentum below; in

section round, central strand small, inner corti-

cal cells in 2-4 rows, large, thin- walled, outer

cortical cells in 1—2 rows, stereids, dark red,

epidermal cells fragile, thin- walled, quickly

eroded and stem becoming fluted. Leaves erect-

spreading to widespreading dry or wet; nar-

rowly lanceolate to linear-lanceolate, (1,2—)

1,5 (-2,2) mm long, lamina unistratose; apex

long-acuminate; margins ± plane, serrulate, fre-

quently by projecting ends of adjoining cells.

Costa excurrent to long-excurrent, superficial

cells rectangular, smooth below, weakly pro-

rate in upper leaf; in section rounded, bulging

dorsally, guide cells 4, larger than other cells,

incrassate, ventral stereid or substereid band

1- 2 cells thick, exposed, dorsal stereid band 3

cells thick, rarely substereids, surface cells not

differentiated. Laminal cells long-rectangular

throughout, thickened, 7-12: 1, papillae few

and somewhat scattered, over lumen or dis-

placed toward proximal end; basal cells not

strongly differentiated, mostly 10: 1, generally

less than 15 cells wide on each side of costa;

alar cells forming small group of quadrate cells,

2- 3 cells wide by ±4 cells high.

Autoicous. Perigonia terminal on branch-

es. Perichaetia terminal, leaves slightly shorter

Fig. 1 17.— Bartramia compacta ( 1 — 12): 1. habit (var. macowaniana), x 1 ; 2. habit (var. compacta), x 1 ; 3. habit

(var. compacta), x 5; 4. part of stem in cross section, x 350; 5. leaf, x 35; 6. leaf in cross section, x350; 7. basal leaf

cells (right side), x 175; 8. laminal cells, x 700; 9. leaf apex, x 175; 10. exothecial cells, x 350; 11. part of capsule

mouth snowing cells, x 350; 12. spore, x 550. B. capensis (13-22): 13. habit, x 1; 14. habit, x 5; 15. part of stem in

cross section, x 350; 16. leaf, x 35; 17. leaf in cross section, x 350; 18. basal leaf cells (right side), x 175; 19. laminal

cells at right margin, x 700; 20. leaf apex, x 175; 21. spore, x 550; 22. part of capsule mouth showing cells, x 350.

B. aristaria (23 - 29): 23. habit, x 1; 24. habit, x 5; 25. leaves, x 35; 26. leaf in cross section, x 350; 27. basal leaf cells

(right side), x 175; 28. laminal cells at right margin, x 700; 29. leaf apex, x 175. (1,7. Sim 10149; 2- 12, Garside 6589a;

13-22, Esterhuysen 17438; 23-29, Thorne 50359).

416

Bartramiaceae

and broader than vegetative leaves; ovate-acu-

minate, 1, 5-2,0 mm long. Seta 5-10 mm

long, reddish brown; capsule ± erect to in-

clined, urn globose and smooth wet, ±quadrate

and sulcate dry, 1,0— 1,2 mm long; exothecial

cells oblong-hexagonal, thin-walled, at mouth

with 4-5 rows of transversely rectangular cells;

stomata present at base of urn, phaneropore;

peristome absent, occasionally with whitish

fringe at mouth; operculum conic to convex-

apiculate; calyptra not seen; spores large, sub-

round, 50 /xm, warty, red-brown. Fig. 117: 13-

22.

Endemic to southern Africa, the species is rare in the

shrublands of the central, southern and southwestern Cape.

A few specimens have also been found in Natal. Map 164.

Vouchers: Stephens PRE-CH9365; Thomas PRE-

CH3393.

The relationship between B. capensis and B. compacta

appears to be close since these species share many charac-

ters. The main differences are leaf shape and leaf position

when dry, type and degree of leaf cell papillosity and to

some extent leaf cell development. In addition the spores of

B. capensis are nearly twice as large as those of B. com-

pacta.

Bartramia capensis is unlikely to be confused with any

other species, with the possible exception of Anacolia breu-

telii, however the bistratose upper lamina and strongly

toothed leaf margins will separate that species.

The description of B. inserta as dioicous by Sullivant

& Lesquereux (1859) [see also Sayre (1978)] is apparently

incorrect. The perigonia are indeed terminal, but on elon-

gated innovations clearly attached to stems with sporo-

hytes. The compact tufting of the plants may account for

ullivant’s observation, or perhaps some stems are initially

terminated by perigonia. Recent examination of several

specimens with sporophytes (including the type of B. in-

serta) clearly indicates that B. capensis is autoicous.

»•: • •

Map 164. — 0 Bartramia capensis

The relationship of B. capensis and B. aristaria is still

in question; see note under that species.

3. Bartramia aristaria C. Mull, in Hed-

wigia 38: 93 (1899). Syntypes: Cape, Table

Mountain, Rehmann s.n., Nov. 1875; Montagu

Pass, Rehmann 184 (PRE!); Natal, Inanda,

Rehmann 194 (PRE!); non Rehmann 185.

Breutelia aristaria (C. Miill.) Broth, in Natiirl. PflFam.

1: 657 (1904); 10: 471 (1924); Sim, Bryo. S. Afr. 312

(1926).

Plants small to medium-sized, loosely

caespitose, light green to yellow-green; terrico-

lous. Stems 5-10 mm high, sparsely branched,

tomentose below, reddish; in section round,

central strand small, inner cortical cells in 3-4

rows, slightly thickened toward margin, outer

cortical cells in 1-2 rows, stereids, reddish,

epidermal cells larger, fragile, thin-walled.

Leaves erect to spreading wet or dry; ovate-acu-

minate to ovate-lanceolate, 1 ,5 — 2,5(— 3,0) mm

long, lamina unistratose; apex acuminate; mar-

gins reflexed, serrate above, frequently bistra-

tose. Costa excurrent to long-excurrent, super-

ficial cells rectangular, weakly prorate above;

in section bulging dorsally, guide cells 2-4,

larger than other cells, ventral stereid band

(1 — )2 — 3 cells thick, exposed, dorsal stereid

band 3-8 cells thick, occasionally substereids,

surface cells not strongly differentiated. Lami-

nal cells short-rectangular, incrassate, ±

flexuose, 6-8:1, papillae generally over

lumen; basal cells weakly differentiated, fre-

quently papillose, generally 20 or more cells

wide on each side of costa; alar cells occasion-

ally distinct.

Autoicous. Perichaetia terminal, leaves ±

broader. Seta 5-10 mm long, reddish; capsule

inclined, urn globose wet, cylindrical-sulcate

dry, 2 mm long, red-brown; exothecial cells

oblong-hexagonal, thin-walled, 4-5 rows of

transversely rectangular cells at mouth; stomata

present at base of urn, phaneropore; peristome

absent; operculum convex-apiculate; spores

subround, to 50 pm, warty, brownish. Fig.

117:23-29.

Endemic to southern Africa, this species is collected

on soil and rock in grassland and shrublands of the southern

and southwestern Cape, Natal and central and eastern

Transvaal. Map 165.

Vouchers: Bews PRE-CH9700; Esterhuysen 17827;

P illans mb.

Bartramiaceae

417

This species resembles a small Breutelia and was

treated in that genus by Brotherus (1924) and Sim (1926);

however, as Sim noted, the plants are very small for that

genus and there is no indication of leaf plications.

The species seems to be related to Bartramia capensis

and many specimens were previously identified as that

species or one of its synonyms. The two species are similar

in several sporophytic and gametophytic characters, but

differ primarily in leaf shape, number of cells across leaf

base, and leaf cell length and shape. These similarities

strongly suggest the need for additional research into the

relationship between the species.

4. Bartramia hampeana C. Mull, in Bot.

Ztg 16: 162 (1858); Broth, in Natiirl. PflFam.

10: 453 (1924); Sim, Bryo. S. Afr. 301 (1926);

Catcheside, Moss. South Austr. 281 (1980).

Type: Cape, Gnadenthal, Breutel s.n. (BM!).

Bartramia asperrima C. Miill. in Hedwigia 38: 94

(1899), hom. illeg. , non (Britt.) C. Miill. (1897); Broth, in

Natiirl. PflFam. 10: 453 (1924). Type: Cape, Gnadenthal,

Breutel s.n. (BM!).

Bartramia penicillata C. Miill. in Hedwigia 38: 94

(1899); Broth, in Natiirl. PflFam. 10: 453 (1924). Type:

Somerset East, Boschberg, MacOwan s.n. , 1878 (GRA!).

Bartramia ramentosa C. Miill. in Hedwigia 38: 94

(1899); Broth, in Natiirl. PflFam. 10: 456 (1924). Syntypes:

Transvaal, Lydenburg, Wilms s.n., Apr. 1887 (G!); Cape,

Somerset East, Boschberg, MacOwan s.n.

Plants small to medium-sized, loosely

caespitose, yellow-green to glaucous green; ter-

ricolous or saxicolous. Stems (3-)8-30(-60)

mm high, irregularly branched; in section

round, central strand present, inner cortical

cells in 3-5 rows, large, thin-walled, outer cor-

tical cells in 1—2 rows, stereids, dark red or

occasionally weakly thickened and undifferen-

tiated, epidermal cells differentiated, slightly

larger, thin-walled, frequently broken away and

stem weakly fluted. Leaves generally compact,

generally appressed dry, erect to erect-spread-

ing or occasionally widespreading wet; linear

above oblong to obovate base, (l,5-)2,5-4,0

(—5,0) mm long, bistratose above base; apex

subulate, frequently fragile, ± flexuose above;

base sheathing, hyaline, abruptly narrowing to

subula, occasionally not as oistinct in smaller

plants; margins plane, distinctly serrate to den-

ticulate, umstratose. Costa percurrent to short-

excurrent as serrate awn, superficial cells long-

rectangular, prorate; in section elliptical, guide

cells 6-12, ventral stereid band 1-3 cells

thick, occasionally substereids, ventral surface

cells strongly incrassate, smaller than guide

cells, rounded but sharply bulging at ends (pro-

rate), dorsal stereid band 3-8 cells thick, dor-

sal surface cells strongly incrassate, rounded

but bulging outward at prorate cell ends, occa-

sionally not differentiated. Laminal cells rec-

tangular to long-rectangular, 2-8:1, weakly

thickened, sharply prorate at both ends; basal

cells highly differentiated, long-rectangular,

10—15:1, thin-walled, hyaline, smooth, ±

glossy.

Dioicous. Perichaetia lateral through inno-

vation, leaves not differentiated. Seta variable

in length, 4 — 10( — 15) mm long, yellowish;

capsule suberect to inclined, urn subglobose to

broadly short-cylindrical, 1,5-2, 2 mm long,

sulcate dry, occasionally weakly ribbed wet,

yellowish with reddish mouth; exothecial cells

subrhomboidal to hexagonal, thin-walled, 3—4

rows at mouth transversely rectangular; stomata

present at base of urn, phaneropore; peristome

absent, occasionally with hyaline fringe at

mouth; operculum conic to convex-apiculate;

calyptra not seen; spores subround, 30-37 /xm,

warty, reddish brown. Fig. 118.

Presently recognized only from southern Africa, B.

hampeana is now believed to be conspecific with the Aus-

tralian species B. hampei (C. Miill.) Catcheside. In south-

ern Africa, B. hampeana is found on soil and rock in shrub-

lands or grasslands of mountains in the northwestern, south-

western, central, southern and eastern Cape, Lesotho,

Natal, Orange Free State and the eastern and central

Transvaal. Map 166.

Vouchers: Esterhuysen 24410; Hilliard & Burn

12222, 125742; Magill 4155, 4699, 5788, 5906; Rankin 78;

Schelpe 7684; Smook 1723; Van Rooy 317; Van Zanten et

al. 76091011.

This is the only southern African species of Bartramia

with the highly differentiated leaf base of the ‘vaginella’

group. In most specimens the hyaline, sheathing base, that

abruptly constricts to a long, narrow subula, is obvious even

on leaves still attached to the stem. Some smaller specimens

418

Bartramiaceae

Map 166. — • Bartramia hampeana

tiated basal leaf cells, as well as other characters specific to

B. hampeana.

The seta of B. hampeana is quite variable in length,

although there appears to be a continual gradation in

lengths, unlike the situation in B. compacta. The sporo-

phytes are quickly overgrown by subperichaetial innova-

tions, thus even capsules with longer seta appear emergent

as the cushions enlarge. The few plants with very long seta

have short stems and almost no branches. These plants at

first appear distinct, but do not differ in other respects.

Fig. 118. — Bartramia hampeana: 1. habit, x 1; 2.

habit, x 4; 3. leaves, x 35; 4. leaf, x 35; 5. leaf in proxi-

mal cross section, x 350; 6. leaf in distal cross section, x

350; 7. basal leaf cells (right side), X 175; 8. basal leaf

cells, x 700; 9. laminal cells at upper right margin, x 700;

10. leaf apex, x 175; 11. part of capsule mouth showing

cells, X 350. (1-11, Barnard 50345).

do not have the pronounced shoulders of the constriction.

These specimens do, however, exhibit the strongly differen-

Insufficiently Known Species

Bartramia delagoae C. Miill. in Hedwigia 38; 92

(1899). Philonotis delagoae (C. Miill.) Par., Ind. Bryol.

Suppl. 266 (1900); Broth, in Natiirl. PflFam. 10: 462

(1924); Dix. in Trans. R. Soc. S. Afr. 8: 205 (1920). Type:

Transvaal, Omtombi, Delagoa Bay to Lydenburg, Wilms

s.n., Aug. 1884 (Herb. Jack). The type of this species has

not been located.

Bartramia spielhausii C. Miill. in Hedwigia 38: 91

(1899). Breutelia spielhausii (C. Miill.) Par., Ind. Bryol.

Suppl. 54 (1900); Broth, in Natiirl. PflFam. 10; 471 (1924).

Syntypes: Cape, Table Mountain, Spielhaus s.n. (Herb.

Bramer); Somerset East, Boschberg, MacOwan s.n., 1883.

A type has not been located. Dixon (1920) indicated that

this species (from description only) was not distinct from

Rehmann 185 (type of Breutelia aristaria var. plumosa

Sim — synonym of Bartramia capensis).

4. CONOSTOMUM

Conostomum Sw. in Web. & Mohr., Naturh. Reise Schwedens 122 (1804); Broth, in Natiirl.

PflFam. 10: 457 (1924); Sainsb., N. Zeal. Mosses 305 (1955); Smith, Moss FI. Brit. Irel. 458

(1978). Type species: C. arcticum Sw.

Plants small to medium-sized, densely caespitose; terricolous or saxicolous. Stems erect,

branched but with few subperichaetial innovations, densely tomentose below; in section rounded,

central strand present, epidermal cells differentiated. Leaves in 5 distinct rows, imbricate wet or

Bartramiaceae

419

dry; triangular to oblong- lanceolate; margins reflexed and serrate at apex; lamina bistratose juxta-

costally. Costa short-excurrent. Laminal cells rhomboidal to rectangular, smooth below, strongly

prorate near apex; basal cells rectangular, smooth.

Autoicous. Seta flexuose; capsule oval, sulcate dry; peristome single, teeth joined at apex;

operculum rostrate; spores with bulging plates, ± warty.

A genus of 15 species found at high altitude in mountainous regions; primarily in the Southern Hemisphere. This is the

first report of the genus for Africa. The specimens were collected in the alpine grass-heathlands of northern Lesotho and the

Drakensberg of Natal.

The genus should not be confused with any other genus of Bartramiaceae, although the prorate laminal cells, spore

ornamentation and capsule structure clearly indicate the relationship of Conostomum to other members of the family.

Conostomum pentastichum (Brid.)

Lindb. in Ofvers. K. VetenskAkad. Forh. 20:

392 (1863). Type: South America, Chile.

Bartramia pentasticha Brid., Muse. Rec. 2: 134 (1803).

Plants small to medium-sized, forming

dense cushions, yellow-brown to yellow-green,

brownish below; terricolous or saxicolous.

Stems 5-10 (-15) mm tall, irregularly

branched, tomentum dense on lower stem, red-

brown; in section subround to angular, central

strand small, inner cortical cells in 3 rows,

large, thin-walled, outer cortical cells in 2

rows, thickened or substereids, red-yellow, epi-

dermal cells large, thin-walled, fragile, stem

becoming fluted. Leaves crowded, erect-ap-

pressed in 5 distinct rows wet or dry; narrowly

triangular to oblong-lanceolate, 1 ,0—1 ,2 mm

long, lamina bistratose juxtacostally from base;

cells with strongly thickened adjoining walls,

outer walls thin, bulging, 4-8 rows at margin

unistratose, cells rounded, ± evenly thickened;

apex acute, toothed; margins plane below,

weakly reflexed above, entire, serrate near

apex. Costa short-excurrent as smooth awn,

0,2-0, 3 mm long, ventral superficial cells rec-

tangular, weakly prorate and ± thickened near

apex, dorsal superficial cells rectangular,

strongly prorate at distal ends and ± thickened

near apex; in section guide cells not well de-

fined, ± 6, substereids, ventral cells in single

layer continuous with upper layer of lamina,

cells enlarged, inner walls strongly thickened,

outer walls very thin- walled, dorsal substereid

band 2 cells thick, dorsal surface cells continu-

ous with lower layer of lamina, similar to ven-

tral surface cells. Upper laminal cells rhomboi-

dal to fusiform or narrow-rectangular, 7—10: 1,

becoming weakly thickened and distally prorate

above, slightly narrower toward margin; basal

cells rectangular, 2-5: 1, thin-walled, smooth.

Perigonia terminal on branches, leaves el-

liptical, yellowish; costa short-excurrent. Peri-

cnaetia terminal, leaves similar to vegetative

leaves or slightly broader; laminal cells narrow,

thin-walled. Seta 3-4 mm long, flexuose, yel-

low-brown; capsule suberect, urn oval-orbicu-

lar wet, elliptical and sulcate dry, 1,5 mm long,

red-brown; exothecial cells hexagonal to ob-

long-hexagonal, 2 rows smaller and quadrate at

mouth; stomata present at base of urn, phanero-

pore; peristome single, fragile, rea-yellow,

teeth 16, linear, smooth, joined at apex; opercu-

lum obliquely rostrate; calyptra not seen; spores

oval-reniform, 50-60 pm, reddish brown, with

low, bulging, ± hexagonal plates covering dis-

tal surface. Fig. 119.

New to southern Africa, C .pentastichum is found in

South America, Tasmania, New Zealand and a few subant-

arctic islands. In the Flora area the species is found on soil

and rock in the alpine grass-heathlands of northern Lesotho

and the Drakensberg of Natal. Map 167.

Vouchers: Magill 7130A; Van Zanten et al. 7609961,

7609977 c. fr.

The unusual rigid ranking of the leaves up the stem,

and shape of leaves and leaf cells should place specimens of

C. pentastichum. The southern African plants are at the

lower limits of measurements published for New Zealand

plants (Sainsbury, 1955) where stems reach 40 mm high

and the seta and capsules are 10-35 mm and 1,5-3, 5 mm

long, respectively.

Bartramiaceae

Bartramiaceae

5. QUATHLAMBA

421

Quathlamba Magill, gen. nov., plantae parvae, laxe caespitosae, subvirides; terricolae. Caules

infra perichaetia ramificantes. Folia ovata, appressa. Costa destituta vel rudimentalis . Cellulae

laminae leves.

Perichaetia terminalia; folia costata. Theca exserta globosa, orificio parvo, peristomio desti-

tute, operculo plane -convexo; calyptra parva cucullata; sporae verrucatae.

Type species: Q. debilicostata Magill.

Plants small, forming loose tufts, light green; terricolous. Stems branching by subperichaetial

innovations. Leaves ovate, appressed. Costa absent or rudimentary. Laminal cells smooth.

Perichaetia terminal, leaves costate. Capsule exserted, globose, mouth small; peristome ab-

sent; operculum plano-convex; calyptra small, cucullate; spores warty.

Quathlamba, barrier of uppointed spears, is the Zulu name for the Drakensberg, habitat of the genus.

Quathlamba debilicostata Magill, sp.

nov. bene distincta plantis parvis, foliis ovato-

acutis, 0,5-0, 7 mm longis, costis destitutis vel

rudimentalibus , cellulis laminae levibus, cap-

sulis globularibus, peristomio destitute et spo-

ris verrucatis.

Type: Lesotho, top of Sani Pass, on soil of

rock crevices along northern cliff face just E of

Mountain Lodge, 2860 m, Magill 4512 (MO,

holo.; PRE).

Plants small, in loose tufts or cushions,

light green above, tan below; terricolous. Stems

erect, 5-15 mm high, julaceous, branching by

subperichaetial innovations; in section round,

central strand small, cortical cells in 3—4 rows,

undifferentiated. Leaves crowded above, erect-

appressed wet or dry; ovate to elliptical or ob-

long-elliptical, 0,5-0, 7 mm long; apex acute,

very weakly concave; margins plane, entire.

Costa absent or rudimentary, when present con-

sisting of a few elongated or brownish cells

extending into lower 'A of leaf; in section with a

single dorsal and ventral cell, both smaller than

cells of lamina. Laminal cells irregular, rhom-

boidal to rectangular, 50-75 /u,m x 25 pm,

thin- walled or occasionally weakly thickened,

smooth; basal cells rectangular to short-rectang-

ular, 30—75 pm x 25 pm\ thin-walled.

Autoicous. Antheridia axillary in upper-

most leaves. Perichaetia terminal but quickly

overgrown by subperichaetial innovations;

leaves ovate-acuminate, 0,75-1,50 mm long;

costa present, weak to above midleaf; in section

with 2-3 cells ventrally and 3—4 cells dorsally;

laminal cells rectangular to long-rectangular,

weakly thickened. Seta 1, 5-2,0 mm long, light

brown; capsule exserted, erect, gymnostomous,

urn globose wet, sulcate dry, 1 ,8—2,2 mm long,

light brown, mouth small; exothecial cells an-

gular, ± subquadrate to rectangular, thin-

walled, 3-4 rows of transversely rectangular,

thickened cells at mouth, dark brown; stomata

in two rows at base of urn, phaneropore; peris-

tome absent; operculum plano-convex; calyptra

quickly lost, small, cucullate, 0,7 mm long,

dark brown, appearing leathery; spores rounded

but flattened to concave on proximal face, 50

pm, warty, yellow-brown. Fig. 120.

Endemic to southern Africa, the species is found on

soil in rock crevices of cliffs in the alpine heathland at Sani

Pass, Lesotho. Map 167.

Vouchers: Magill 4443, 4485, 7154a.

Gametophytically these plants are very unusual in the

family, but the globose urn with small mouth and warty

spores and two guard cells per stoma indicate its

relationship to other members of Bartramiaceae.

Fig. 1 19. — Conostomum pentastichum: 1. habit, x 1; 2. habit, x 5; 3. stem in cross section, x 350; 4. leaves, x

70; 5. leaf in cross section, x 175; 6. basal leaf cells (left side), x 175; 7. upper laminal cells at left margin (dorsal surface),

x 350; 8. upper laminal cells (dorsal surface), x 700; 9. leaf apex, x 700; 10. upper exothecial cells, x 175; 11. part of

capsule mouth, showing cells and broken peristome, x 175. (1-11, VanZanten 7609977).

Fig. 120. — Quathlamba debilicostata: 1. habit, x 1; 2. habit, x 10; 3. stem cross section, x 175; 4 & 5. leaves, x

35; 6. leaves in cross section, x 175; 7. basal leaf cells, x 175; 8. leaf apex, x 175; 9. perichaetial leaf, x 35; 10. part of

capsule mouth with spores, x 70; 11. spore, x 700. (1- 1 1, Magill 4512).

Bartramiaceae

Rg 121. — Bartramidula comosa (1-14): 1. habit, x

1; 2. habit, x 5; 3. part of stem in cross section, x 700; 4.

stem leaf, x 35; 5. subperichaetial leaf, x 35; 6. leaf in

cross section, x 700; 7. basal leaf cells (left side), x 175;

8. laminal cells at left margin, x 700; 9. leaf apex, x 175;

10. perichaetial leaf, x 35; 11. capsule, dry, x 10; 12.

capsule, wet, x 10; 13. exothecial cells, x 175; 14. spore,

7 x 450. B. globosa (15-27): 15. habit, x 1; 16. habit, x 5;

17. part of stem in cross section, x 700; 18. leaf, x 35; 19.

leaf in cross section, x 700; 20. basal leaf cells (right side),

x 175; 21. laminal cells at right margin, X 700; 22. leaf

apex, x 175; 23. perichaetial leaf, x 35; 24. capsule, wet,

x 10; 25. capsule, dry, x 10; 26. part of capsule mouth

showing cells, X 175; 27. spore, X 450. (1-14, Esterhuy-

sen PRE-CHI 2606; 15-27, Oliver 6793).

Bartramiaceae

6. BARTRAMIDULA

423

Bartramidula B.S.G., Bryol. Eur. 4: 55 (1846), nom. cons.; Broth, in Natiirl. PflFam. 10: 459

(1924); Sim, Bryo. S. Air. 304 (1926); Flowers in Grout, Moss FI. N. Amer. 2: 163 (1935);

Smith, Moss FI. Brit. Irel. 458 (1978); Catcheside, Moss. South Austr. 285 (1980). Lectotype

species: B. wilsonii B.S.G., fide Flowers in Grout, Moss FI. N. Amer. 2: 163 (1935).

Plants small, caespitose or in small groups; terricolous. Stems sparsely branched by subperi-

chaetial innovations; central strand large, epidermis differentiated. Leaves linear-lanceolate to

ovate-acuminate, margins serrate. Costa short-excurrent. Laminal cells narrowly rectangular, in-

crassate, prorate; basal cells rectangular, smooth.

Dioicous or synoicous. Perichaetia becoming lateral through innovation; leaves ovate-acumi-

nate or strongly differentiated, long-linear. Seta elongate; capsule erect, symmetrical, round,

smooth or warty, little altered dry; peristome absent; spores papillose or warty.

The 21 species of Bartramidula are evenly distributed in temperate and tropical regions of the world. The two southern

African species could easily be mistaken for species of Philonotis when sporophytes are absent.

1 Capsule wall warty; spore surface warty 1. B. comosa

1 Capsule wall smooth; spore surface papillose 2. B. globosa

1. Bartramidula comosa (Hampe & C.

Mull.) Broth, in Natiirl. PflFam. 1: 644 (1904);

10: 460 (1924); Sim, Bryo. S. Afr. 304 (1926).

Type: Cape, Houteniqua, Montagu Pass, Breu-

tel s.n. (BM!).

Bartramia comosa Hampe & C. Mull, in Bot. Ztg 17:

221 (1859), horn, illeg. , non Mitten (1859).

Plants small, loosely caespitose or in small

groups, yellowish green; terricolous. Stems

erect, to 5 mm high, usually with a few subperi-

chaetial branches; in section round, central

strand large, inner cortical cells in 2-3 rows,

large, thin- walled; outer cortical cells in 1-2

rows, smaller, incrassate, reddish, epidermal

cells thin-walled. Leaves somewhat crowded,

appressed to erect dry, erect wet; narrowly lan-

ceolate, 1 ,0—1 ,2(— 1 ,5) mm long; apex acumi-

nate to subulate; margins plane, serrate above

base. Costa short-excurrent, serrate; in section

rounded, bulging dorsally, guide cells 2, incras-

sate, ventral cells in single layer, similar to

guide cells, incrassate, dorsal stereid band

small, of 4-8 cells, dorsal surface cells similar

to ventral cells. Laminal cells linear-rectangu-

lar, 4—6: 1, incrassate, weakly prorate; basal

cells slightly wider, 2-4: 1, smootn.

Dioicous. Perichaetia terminal or lateral

through innovation; leaves strongly differen-

tiated, long-linear, 4-5 mm long; margins

weakly serrate, teeth distant; costa excurrent;

laminal cells linear, mostly smooth. Seta 3-6

mm long, yellowish to reddish yellow; capsule

erect, symmetrical, urn globose, warty, to 1

mm long, yellowish to brownish; exothecial

cells irregularly arranged, quadrate to rect-

angular or oblong-hexagonal or occasionally

transversely rectangular, warty protuberances

on capsule wall multicellular, 4-6 rows of

short, transversely rectangular cells at capsule

mouth; stomata present at base of urn, phanero-

pore; peristome absent; operculum plano-con-

vex; only juvenile calyptra seen, cucullate, ap-

parently quickly lost; spores subround, 35-42

fim, densely warty, brownish. Fig. 121: 1-14.

Endemic to southern Africa, B. comosa is collected on

soil in mountains of the southern and southwestern Cape.

Map 168.

Voucher: Esterhuysen 24360.

Vegetatively similar to B. globosa ; the narrower

leaves, differences in costal anatomy and more strongly

thickened leaf cells will help to separate sterile specimens of

B. comosa. Sporophytically this species is very distinctive

with its very long-linear perichaetial leaves, warty capsule

and large warty spores.

2. Bartramidula globosa (C. Mull.)

Broth, in Natiirl. PflFam. 1: 644 (1904); 10:

460 (1924); Sim, Bryo. S. Afr. 305 (1926).

Type: Cape, Gnadenthal, Breutel s.n., 1856

(BM!).

Bartramia globosa C. Miill. in Hedwigia 38: 90 (1899).

424

Bartramiaceae

Map 168. — • Bartramidula globosa

▲ Bartramidula comosa

Bartramia globosa var. tenuicaulis C. Mull, in Hedwigia

38: 90 (1899). Bartramia sordida C. Mull. , Gen. Muse. Fr.

335 (1900). Bartramidula globosa var. tenuicaulis (C.

Mull.) Sim, Bryo. S. Afr. 305 (1926). Type: not desig-

nated.

Plants small, loosely caespitose or forming

turfs, green to yellow-green; terricolous or saxi-

colous. Stems erect, (2-) 4-8 (-10) mm high,

branching by subperichaetial innovations,

weakly tomentose below; in section round, cen-

tral strand large, innner cortical cells in 2-4

rows, large and thin-walled, but outer row

smaller, outer cortical cells in 1-2 rows, ster-

eids or substereids, reddish, epidermal cells

medium-sized, thin- walled, frequently col-

lapsed or eroded producing fluted stem. Leaves

somewhat crowded, appressed to erect dry,

erect-spreading wet; narrowly lanceolate or

occasionally ovate-acuminate, 0,8-1, 4 mm

long; apex acuminate; margins plane to weakly

reflexed, sharply serrate above. Costa short ex-

current as serrate awn or occasionally only per-

current, filling acumen; in section rounded,

bulging dorsally, guide cells 2, incrassate, ven-

tral cells 2, in single layer, similar to guide

cells, weakly thickened, dorsal stereid band 1-2

cells thick, frequently with distinct gap below

guide cells, dorsal surface cells similar to ven-

tral cells. Laminal cells linear-rectangular,

8-10: 1 , incrassate, distinctly prorate in upper

leaf; basal cells in distinct group, rectangular,

2-4: 1, slightly bulging, weakly thickened.

Synoicous. Perichaetia terminal or lateral

through innovation; leaves ovate to short-

oblong, acuminate, to 2 mm long, laminal cells

rectangular. Seta 7—12 mm long, reddish yel-

low; capsule erect, symmetrical, urn globose,

1, 5-2,0 mm long, ± smooth, little altered to

wrinkled dry, mouth small; exothecial cells ob-

long-hexagonal, thin- walled, becoming qua-

drate above, 3-4 rows at mouth transversely

rectangular; stomata numerous at base of urn,

phaneropore; peristome absent, frequently with

white fringe in mouth; operculum plano-

convex; calyptra not seen; spores subround to

reniform, 37—42 /tm, weakly papillose, yellow-

brown. Fig. 121: 15-27.

Endemic to southern Africa, B. globosa is found on

soil and rock in the central, southwestern, southern and

eastern Cape, Natal, Lesotho, eastern Orange Free State

and the central and northern Transvaal. Map 168.

Vouchers: Magill 4449; Schelpe 7558; Van Rooy 467.

Sterile specimens are practically indistinguishable

from Philonotis dregeana. The erect, symmetrical, globose

capsules, with small mouths, will help to identify speci-

mens of B. globosa. In addition the capsules are inflated

and ± smooth wet or dry, a distincitive feature within the

family. The two southern African species of Bartramidula

are most easily identified by their sporophyte characters,

which will also separate the species from other members of

the family.

7. PHILONOTIS

Philonotis Brid., Bryol. Univ. 2: 15 (1827); Broth, in Naturl. PflFam. 10: 460 (1924); Sim, Bryo.

S. Afr. 305 (1926); Flowers in Grout, Moss FI. N. Amer. 2: 164 (1935); Sainsb., N. Zeal. Mosses

310 (1955); Gangulee, Moss. E. India 4: 1 108 (1974); Scott & Stone, Moss. S. Austr. 336(1976);

Smith, Moss FI. Brit. Irel. 459 (1978); Catcheside, Moss. South Austr. 288 (1980). Lectotype

species: P.fontana (Hedw.) Brid., fide Flowers in Grout, Moss FI. N. Amer. 2: 164(1935).

Plants small to medium-sized, rarely large, caespitose, green to yellowish green, occasionally

bluish green; terricolous or saxicolous. Stems erect, branching by numerous, generally whorled,

subperichaetial innovations; central strand present, epidermal cells large, thin-walled, fragile.

Bartramiaceae

425

Leaves ovate to lanceolate; apex acute to acuminate, occasionally obtuse to rounded; base scarcely

differentiated; margins plane to strongly recurved, entire to crenulate or doubly serrate. Costa

subpercurrent to short-excurrent. Laminal cells short-rectangular to linear, incrassate, smooth,

papillose or prorate; basal cells usually slightly larger, rectangular, thin-walled, generally smooth.

Autoicous or dioicous. Perigonia terminal, budlike or discoid with leaves abruptly narrowed

and reflexed above broad sheathing base. Perichaetia terminal, quickly overgrown by subperichae-

tial branches. Seta elongate; capsule inclined, globose to ovoid wet, sulcate dry; peristome double,

endostome with segments and cilia, occasionally rudimentary; operculum convex to conic; spores

rounded to reniform, papillose.

Philonotis is a widespread genus with just over 170 species. Specimens are commonly collected on shallow soil or

rock, especially in association with seepage areas, at the back of waterfalls or on marshy ground.

To accommodate the unusually large amount of gametophytic variation exhibited by specimens of Philonotis, a broad

approach to the circumscription of species has been adopted. This variation is frequently expressed in leaf size and shape,

leaf cell size and ornamentation (both between individual plants in a specimen and between fertile and sterile collections)

and plant size and degree of branching.

Although the situation has been rationalized at the species level, a number of problems still exist, e.g. separation of

sterile specimens of Philonotis dregeana and Bartramidula globosa; relationship between P. dregeana an aP. afrocapilla-

ris. Solution to the problem of infraspecific variation will require considerable research, including cultivation experiments.

For instance differences in appearance between smooth to weakly papillose specimens and strongly papillose forms are very

striking, but apparently a result of environmental differences.

1 Leaf cells papillose or prorate:

2 Leaf margins plane; leaves frequently keeled 4. P.falcata

2 Leaf margins re flexed below, recurved to revolute above:

3 Leaf cells short-rectangular, prorate; plants yellow-green 3. P. africana

3 Leaf cells quadrate to hexagonal, papilla over lumen; plants with glaucous bloom 5. P. scabrifolia

1 Leaf cells smooth or some cells weakly prorate:

4 Leaves with border of narrow, elongated cells 6. P. vagans

4 Leaves without border:

5 Leaf cells ± lax, rectangular to oblong-hexagonal; costa generally ending below apex 2.P. hastata

5 Leaf cells regularly narrow-rectangular, generally incrassate; costa short-excurrent 1. P. dregeana

1. Philonotis dregeana (C. Mull.) Jaeg.

in Verh. St Gall, naturw. Ges. 1873—74: 89

(1875); Broth, in Nattirl. PflFam. 10: 463

(1924). Type: Cape [Bartramia sp. coll. Dre-

geana III. A,e,9], Drege s.n. (G!).

Bartramia dregeana C. Mull, in Bot. Ztg 14: 419 (1856).

Bartramia androgyna Hampe in Bot. Ztg 28: 34 (1870).

Philonotis androgyna (Hampe) Jaeg. in Verh. St Gall, na-

turw. Ges. 1873-74: 89 (1875); Sim, Bryo. S. Afr. 306

(1926). Type: Natal, Umpumulo, Borgen s.n., 20 Mar.

1867 (BM, nolo.!).

Bartramia afrouncinata C. Miill. in Hedwigia 38: 91

(1899); Dix. in J. Bot., Lond. 62: 235 (1924); Sim, Bryo.

S. Afr. 307 (1926). Philonotis afrouncinata (C. Miill.)

Par., Ind. Bryol. Suppl. 265 (1900); Broth, in Natiirl.

PflFam. 10: 463 (1924). Type: Cape, MacOwan s.n., 1876

(?G!).

Bartramia afrouncinata var. gracilescens C. Miill. in

Hedwigia 38: 92 (1899). Philonotis afrofontana var. graci-

lescens (C. Miill.) Par., Ind. Bryol. Suppl. 265 (1900);

Dix. in Trans. R. Soc. S. Afr. 8: 205 (1920); Sim, Bryo. S.

Afr. 307 (1926). Bartramia gracilescens (C. Miill.) C.

Miill., Gen. Muse. Fr. 339 (1900). Type: Cape, Boschberg,

MacOwan s.n., 1874 (G!).

Bartramia pernana C. Miill. in Hedwigia 38: 92 (1899).

Philonotis pernana (C. Miill.) Par., Ind. Bryol. Suppl. 267

(1900). Type: Cape, prope Belweder, Rehmann 191, fide

Dix. & Gepp (1923).

Philonotis afrocapillaris Dix. ex Sim, Bryo. S. Afr. 309