FLORA OF SOUTHERN AFRICA

BRYOPHYTA

Editor O.A. Leistner

Part 1 Musci

Fascicle 3

Erpodiaceae — Hookeriaceae

by Robert E. Magill & Jacques van Rooy

Digitized by the Internet Archive

in 2016

https://archive.org/details/floraofsoutherna13unse_0

FLORA OF SOUTHERN AFRICA

which deals with the territories of

SOUTH AFRICA, LESOTHO, SWAZILAND, NAMIBIA AND BOTSWANA

BRYOPHYTA

PART 1 MUSCI

Fascicle 3 Erpodiaceae — Hookeriaceae

Robert E. Magill & Jacques van Rooy

with drawings by

Gillian Condy

Scientific editor: O.A. Leistner

Technical editor: E. du Plessis

by

NATIONAL

Botanical

INSTITUTE

Pretoria

1998

Editorial Board

B.J. Huntley National Botanical Institute, Cape Town, RSA

R.B. Nordenstam Natural History Museum, Stockholm, Sweden

R.M. Polhill Royal Botanic Gardens, Kew, UK

J.J.A. van der Walt Paradyskloof, Stellenbosch, RSA

Typesetting and page layout by S.S. Brink, NBI, Pretoria

Reproduction by 4 Images, P.O, Box 34059, Glenstantia, 0010 Pretoria

Printed by Promedia, P.O. Box 255, Silverton, 0127 Pretoria

©, published by and obtainable from the

National Botanical Institute, Private Bag X101, Pretoria, 0001 South Africa

ISBN 1-874907-33-1

CONTENTS

Page

New taxa and new combinations published in Part 1, Fascicle 3 iv

Geographical regions referred to in this Fascicle v

Introduction to Fascicle 3 vii

Conspectus of classification 445

Literature cited in Fascicle 3 446

Provisional key to the families of Fascicle 3 449

ERPODIACEAE 451

RHACHITHECIACEAE 459

PT Y CHOMITRIACE AE 462

ORTHOTRICHACEAE 476

RHABDOWEISIACEAE 527

RACOPILACEAE 531

FONTINALACEAE 534

WARDIACEAE 538

HEDWIGIACEAE 541

CRYPHAEACEAE 550

LEUCODONTACEAE 553

PRIONODONTACEAE 558

TRACHYPODACEAE 560

PTEROBRYACEAE 565

METEORIACEAE 575

LEPTODONTACEAE 585

NECKERACEAE 590

THAMNOBRYACEAE 592

HOOKERIACEAE 601

Index to Fascicle 3 619

Appendix:

Plan of Flora of southern Africa A- 1

FSA contributions in Bothalia A-3

Alphabetical arrangement of taxa listed as ‘published’ in Plan of Flora of southern Africa A-4

NEW TAXA AND NEW COMBINATIONS PUBLISHED IN PART 1

FASCICLE 3

Distichophyllum mniifolium (Hornsch.) Sim var. taylorii (Sim) Magill, stat. nov., p. 614

Erpodium coronatum (Hook. & Wils.) Mitt, subsp. transvaaliense (Broth. & Wager) Magill , stat.

nov., p. 453

Pterobryopsis acutifolium (Brid.) Magill , comb, nov., p. 569

Pterobryopsis hoehnelii (C. Mull.) Magill , comb, nov., p. 567

Ptychomitriopsis aloinoides Magill , sp. nov., p. 463

Date of publication: January, 1998.

GEOGRAPHICAL REGIONS REFERRED TO IN THIS FASCICLE

B — Botswana

CC — central Cape

CE — eastern Cape

CN — northern Cape

CNW — northwestern Cape

CS — southern Cape

CSW — southwestern Cape

FS — Free State

KZN — KwaZulu-Natal

L — Lesotho

NAM — Namibia

S — Swaziland

TC — central Transvaal

TE — eastern Transvaal

TN — northern Transvaal

TS — southern Transvaal

Z — Zululand

v

INTRODUCTION TO FASCICLE 3

The following text constitutes Fascicle 3 of Part 1 of Volume Bryophyta in the Flora of south-

ern Africa Cryptogam series. This fascicle includes the families Erpodiaceae to Hookeriaceae (see

Conspectus of classification, p. 445).

Several inadequacies of the traditional classification system used for this Flora (see Fascicle 1,

p. 13) are evident in this fascicle. The families treated in Fascicle 3 should contain diplolepidous,

pleurocarpic mosses. Recent research on peristome development, including some members of the

Orthotrichales, indicates that the Ptychomitriaceae (Edwards 1979) would be better placed near the

Grimmiaceae (Fascicle 1) and that Rhabdoweisia of the Rhabdoweisiaceae is a member of the

Dicranaceae (Fascicle 1).

Amphidium, a genus traditionally placed close to Rhabdoweisia , but lacking a peristome, has

been included in Orthotrichaceae based on early development of the capsule (Lewinsky 1976). The

Orthotrichaceae appears to be a transition group containing both acrocarpic and pleurocarpic gen-

era and, when present, diplolepidous peristomes.

Two genera scheduled to appear in Fascicle 3 will be treated in Fascicle 4. These are Rigodium

(previously Lembophyllaceae now placed in Rigodiaceae), which is better placed in the Thuidiales

near Thuidiaceae, and Catagonium (previously Phyllogoniaceae now placed in Catagoniaceae),

recently revised by Lin (1983) and repositioned in the Hypnobryales near Plagiotheciaceae. The

Conspectus of classification has been modified to reflect these changes.

The notes on study and identification, and the Glossary included in Fascicle 1 , also apply here.

A more extensive list of terms used in bryology is available in Glossarium polyglottum bryologiae

(Magill 1990).

Two important characteristics are exhibited by many of the taxa treated in Fascicle 3. First, the

diplolepidous peristomes are generally incomplete, having endostome segments and cilia fre-

quently reduced or occasionally lacking. The exostome teeth are also commonly modified in a vari-

ety of ways resulting in a peristome that is quite distinct from the diplolepidous peristomes of the

Bryales treated in Fascicle 2 or those of the Thuidiales and Hypnobryales of Fascicle 4.

Second, many of the mosses treated here share a distinctive growth form. These plants have thin,

creeping, primary stems growing appressed to the substrate. Secondary stems or branches arise

more or less at right angles from these creeping stems. These innovations are said to be erect, a

term used here to indicate an orientation perpendicular to the substrate or creeping stem. In some

plants these secondary stems or branches become very long and pendent, forming large hanging

masses from trees and rocks.

Some of the geographical regions referred to in the previous two fascicles (Magill 1981, 1987),

partly demarcated by former provincial boundaries, have been changed in this fascicle to reflect

recent constitutional developments in southern Africa. The former western Transvaal region now

forms part of the northern Cape region, and the former Transkei region has been added to the east-

ern Cape region (see map on p. v). The name of the South West Africa/Namibia region has been

changed to Namibia, that of the Orange Free State region to Free State, and that of Natal to

KwaZulu-Natal. The name Zululand is retained for the northern part of KwaZulu-Natal Province.

The ‘Cape’ and Transvaal' regions referred to in this fascicle no longer coincide with the former

Cape and Transvaal provinces of South Africa.

The illustrations were prepared by Ms Gillian Condy; for technique and procedure, see Introduc-

tion to Fascicle 1, p. xv. Research on this fascicle was partly supported by grants to the Missouri

Botanical Garden from the National Science Foundation and the National Geographic Society.

vii

445

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

Grimmiaceae

Fascicle 2:

ORDER FUNARIALES

Family Gigaspermaceae

Ephemeraceae

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

Pterobryaceae

Meteoriaceae

Leptodontaceae

Neckeraceae

Thamnobryaceae

ORDER HOOKERIALES

Family Hookeriaceae

Fascicle 4\

ORDER THUIDIALES

Family Fabroniaceae

Leskeaceae

Thuidiaceae

Rigodiaceae

ORDER HYPNOBRYALES

Family Amblystegiaceae

Brachytheciaceae

Entodontaceae

Plagiotheciaceae

Catagoniaceae

Sematophyllaceae

Hypnaceae

Hylocomiaceae

CLASS POLYTRICHOPSIDA

ORDER POLYTRICHALES

Family Polytrichaceae

446

LITERATURE CITED IN FASCICLE 3

Literature references given outside formal literature treatments

ALLEN, B.H. 1983. On the costa of Fontinalis

(Musci). Lindbergia 9: 37 — 40.

ALLEN, B.H. 1987. A systematic account of

Pulchrinodus inflatus (Musci: Pterobry-

aceae), genus novum. New Zealand Journal

of Botany 25: 335-342.

ALLEN, B.H. & CROSBY, M.R. 1986. Revi-

sion of the genus Squamidium (Musci: Me-

teoriaceae). Journal of the Hatton Botanical

Laboratory 61: 423^476.

ANDERSON, L.E. & CRUM, H. 1959. Cyto-

taxonomic studies on mosses of the Canadian

Rocky Mountains. Bulletin of the National

Museum of Canada 160: 1-89.

ARZENI, C.B. 1954. The Pterobryaceae of the

southern United States, Mexico, Central

America, and the West Indies. American Mid-

land Naturalist 52: 1-67.

BIZOT, M. & POCS, T. 1974. East African

bryophytes. Acta Academiae Paedagogicae

Agriensis, N. Ser. 12: 383-449.

BUCK, W.R. 1980. Animadversions on Pteri-

gynandrum with special commentary on

Forsstroemia and Leptopterigynandrum.

The Bryologist 83: 451-465.

DE SLOOVER, J.L. 1983. Note de bryologie

africaine XII. — Porotrichum et Porotham-

nium. Bulletin du Jardin Botanique National

de Belgique 53: 97-152.

DIXON, H.N. 1920. New and interesting South

African mosses. Transactions of the Royal

Society of South Africa 8: 179-224.

DIXON, H.N. 1927. Studies in the bryology of

New Zealand V. New Zealand Institute,

Bulletin 3: 239-298.

DIXON, H.N. 1931. Ptychomitriopsis Dix. gen.

nov. Ptychomitriacearum. Journal of Botany,

British and Foreign 69: 284—285.

EDWARDS, S.R. 1979. Taxonomic implica-

tions of cell patterns in haplolepidous moss

peristomes. In G.C.S. Clark & J.G. Duckett,

Bryophyte systematics. Systematic Associa-

tion Special Volume 14: 315-346. Syste-

matics Association, London.

ENROTH, J. 1991. Thamnobryum capense

comb. nov. (Neckeraceae, Musci). Novon 1:

110.

LEWINSKY, J. 1976. On the systematic posi-

tion of Amphidium Schimp. Lindbergia 3:

227-231.

LEWINSKY, J. 1978. The genus Orthotrichum

Hedw. (Musci) in Africa south of the Tropic

of Cancer. Botanisk Tidsskrift 72: 61-85.

LEWINSKY, J. & VAN ROOY, J. 1990. New

species and a new record of Orthotrichum

from southern Africa: O. incurvomarginatum

sp. nov., O. armatum sp. nov., O. oreophilum

sp. nov. and O. firmum Vent. Journal of

Bryology 16,1: 67-78.

LIN, S.-H. 1983. A taxonomic revision of Phyl-

logoniaceae (Bryopsida). Part 1. Journal of

Taiwan Museum 36,2: 37-86.

MAGILL, R.E. 1981. Flora of southern Africa.

Bryophyta 1,1: 1-291.

MAGILL, R.E. 1987. Flora of southern Africa.

Bryophyta 1,2: 293^443.

MAGILL, R.E. (Ed.) 1990. Glossarium poly-

glottum bryologiae: a multilingual glossary

for bryology. Monographs in Systematic

Botany from the Missouri Botanical Garden

33, [x] + 297 pp. Missouri Botanical Garden,

St Louis.

MAGILL, R.E. & SCHELPE, E.A. 1979. The

bryophytes of southern Africa. An annotated

checklist. Memoirs of the Botanical Survey

of South Africa 43: 1-39.

MALTA, N. 1926. Die Gattung Zygodon Hook,

et Tayl. Eine monographische Studie. RIGA.

MANUEL, M.G. 1973. Studies in Cryphaea-

ceae I. A revision of the genus Cryphaea in

North America north of Mexico. The

Bryologist 76: 144—162.

447

MANUEL, M.G. 1974. A revised classification

of the Leucodontaceae and a revision of the

subfamily Alsioideae. The Bryologist 77:

531-550.

MITTEN, G. 1869. Musci austro-americani.

Journal of the Linnean Society 12: 1-659.

Reprinted in: Monographs in Systematics

Botany from the Missouri Botanical Garden

7 (1982).

MULLER, C. 1899. Contributiones ad Bryol-

ogiam austro-afram. Hedwigia 38: 52-155.

SHAW, J. 1986. Peristome structure in the

Orthotrichaceae. Journal of the Hattori Bot-

anical Laboratory 60: 119-136.

SIM, T.R. 1926. The bryophyta of South Africa.

Transactions of the Royal Society of South

Africa 15: 1-475.

SMITH, A.J.E. 1978. The moss flora of Britain

& Ireland , 706 pp. Cambridge University

Press, Cambridge.

STARK, L.R. 1987. A taxonomic monograph of

Forsstroemia Lindb. (Bryopsida: Leptodont-

aceae). Journal of the Hattori Botanical

Laboratory 63: 133-218.

TIXIER, P. 1989. Bryophyta exotica — 8. Recol-

tes de J.-P. Brunei au Togo (1983-1985).

Candollea 44: 493-511.

VAN ZANTEN, B.O. 1959. Trachypodaceae: a

critical revision. Blumea 9: All-515.

VITT, D.H. 1971. The infrageneric evolution,

phylogeny, and taxonomy of the genus Or-

thotrichum (Musci) in North America. Nova

Hedwigia 21: 683-711.

VITT, D.H. 1980a. The genus Macrocoma I.

Typification of names and taxonomy of the

species. The Bryologist 83,4: 405-436.

VITT. D.H. 1980b. The nomenclature and taxon-

omy of Macrocoma lycopodioides (Schwaegr.)

Vitt. Journal of Bryology 11: 219-229.

VITT, D.H. 1980c. The genus Macrocoma II.

Geographical variation in the Macrocoma

tenue-M. sullivantii species complex. The

Bryologist 83,4: 437-450.

WELCH, W.H. 1960. A monograph of the Fon-

tinalaceae , [vii] 4 357 pp. Martinus Nijhoff,

The Hague.

WHITTEMORE, A. & ALLEN. B. 1989. The

systematic position of Adelothecium Mitt,

and the familial classification of the Hook-

eriales (Musci). The Bryologist 92: 261-272.

Abbreviations and full titles of books quoted in this work*

Bot. Taschenb. = Botanisches Taschenbuch

Bryo. S. Afr. = The Bryophyta of South Africa

Bryol. austr. excurs. = Bryologia austriaca ex-

cursoria

Bryol. Eur. = Bryologia europaea

Bryol. Univ. = Bryologia universa

Cat. Afr. PI. = Catalogue of the African plants

Consp. regn. veg. = Conspectus regni veget-

abilis

Coroll, bryol. eur. = Corollarium bryologiae

europaeae

Engl. bot. = English botany

Enum. Bryin. exot. = Enumeratio Bryinearum

exoticarum

* Abbreviations of journal titles follow Lawrence, G.H.M.,

Buchheim, A.F.G.. Daniels, G.S. & Dolezal. H. (eds) 1968.

B-P-H, Botanico-Periodicum-Huntianum. Hunt Botanical

Library, Pittsburgh.

FI. bras. = Flora brasiliensis, Musci

FI. nov. zel. = Flora novae-zelandiae

Hist. phys. Madagascar, mousses = Histoire

physique, naturelle et politique de

Madagascar

Horae phys. berol. = Horae physicae berolinen-

ses

Icon. muse. = leones muscorum

Index bryol. = Index bryologicus

Index bryol. suppl. = Index bryologicus supple-

mentum

Index mus. pi. crypt. = Index musei plantarum

cryptogamarum

Monograph Fontinalaceae = A monograph of

the Fontinalaceae

Moss E.N. Amer. = Mosses of Eastern North

America

448

Moss FI. Brit. Irel. = The Moss Flora of Britain

and Ireland

Moss FI. Fenn. = Moss Flora of Fennoscandia

Moss FI. N. Amer. = Moss Flora of North

America north of Mexico

Moss FI. Pacific Northwest = Moss Flora of the

Pacific Northwest

Moss. E. India = Mosses of Eastern India and

adjacent regions

Moss. S. Austr. = Mosses of South Australia

Muse. Buitenzorg = Die Musci der Flora von

Buitenzorg

Muse, frond, ined. archip. ind. = Musci frondosi

inediti archipelagi indici

Muscol. brit. = Muscologia britannica

Muscol. recent. = Muscologia recentiorum

Muscol. recent, suppl. = Muscologia recentio-

rum supplementum

Muscol. recent, suppl. = Muscologia recentio-

rum supplementum seu species musco-

rum

N. Zeal. Mosses = A handbook of New Zealand

mosses

Natiirl. PflFam. = Die Natiirlichen Pflanzen-

familien

Observ. bot. = Observationes botanicae

Prodr. aetheogam. = Prodrome des cinquieme

families de FAetheogamie

Prodr. fl. bryol. Madagascar = Prodrome de la

flore bryologique de Madagascar

Revis. gen. pi. = Revisio generum plantarum

Skand. Bladmossfl. = Skandinaviens Bladmoss-

flora

Sp. muse, frond. = Species muscorum frondoso-

rum

Sp. muse, frond, suppl. = Species muscorum

frondosorum supplementum

Stud, handb. Brit, mosses = The student's hand-

book of British mosses

Syn. muse. eur. = Synopsis muscorum europae-

orum

Syn. muse, frond. = Synopsis muscorum fron-

dosorum omnium hucusque cognitorum

Syn. pi. = Synopsis plantarum

Tab. Calyptr. operc. = Tabula exhibens calyp-

tratarum operculatarum

449

PROVISIONAL KEY TO THE FAMILIES OF FASCICLE 3

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

a continuation of the family keys in Fascicles 1 and 2 and should be used in conjunction with them.

Where necessary, couplets are incorporated into the key that refer to taxa that have been dealt with

or will be treated in other fascicles.

1 Plants acrocarpic or appearing acrocarpic:

2 Calyptra cucullate:

3 Seta elongate; capsule long-exserted; gemmae occasionally present:

4 Gemmae present on stems and rhizomes ORTHOTRICHACEAE (p. 476)

4 Gemmae not present on stems (Fascicles 1 & 2)

3 Seta short; capsule immersed, emergent or short-exserted; gemmae absent:

5 Leaf cells papillose or striolate-papillose; peristome absent

ORTHOTRICHACEAE (p. 476)

5 Leaf cells smooth; peristome present:

6 Corticolous; perichaetial leaves long-sheathing, covering the seta; peristome teeth in

8 pairs RHACHITHECIACEAE (p. 459)

6 Terricolous; perichaetial leaves scarcely differentiated; peristome teeth 16, fused at

base, distant above, fugacious RHABDOWEISIACEAE (p. 527)

2 Calyptra mitrate:

7 Leaves erect, appressed, contorted or crisped when dry, muticous, dull; lamina and mar-

gins generally unistratose; capsules immersed or short-exserted

ORTHOTRICHACEAE (p. 476)

7 Leaves incurled when dry, glossy; lamina or margins frequently bistratose; cells smooth

or mammillose:

8 Leaves with long, hyaline awn, or if muticous then capsules immersed

GRIMMIACEAE (Fascicle 1)

8 Leaves muticous, capsules exserted PTYCHOMITRIACEAE (p. 462)

1 Plants pleurocarpic or of pleurocarpic habit, stems and branches frequently erect from a

creeping stem or rhizome:

9 Leaves ecostate or apparently so:

10 Plants aquatic or in splash zones:

11 Plants floating in streams, stems very long FONTINALACEAE (p. 534)

1 1 Plants forming tufts on rocks in splash zone, stems shorter WARDIACEAE (p. 538)

10 Plants of drier habitats, corticolous or saxicolous:

12 Plants large, branches ± erect; leaves widespreading when wet:

13 Leaf cells smooth, rhomboidal to elliptical LEUCODONTACEAE (p. 553)

13 Leaf cells papillose or granulose, quadrate to rectangular or fusiform

HEDWIGIACEAE (p .54 1 )

12 Plants small, stems and branches appressed to substrate; leaves complanate; leaf

cells hexagonal ERPODIACEAE (p. 451)

9 Leaves costate:

14 Stems distinctly heterophyllous:

15 Stems creeping along substrate, dark green; leaf margins unbordered . . v

RACOPILACEAE (p. 531)

15 Stems erect, green to glaucous green; leaf margins bordered by elongated cells ....

HOOKERIACEAE (p. 601)

14 Stems ± homophyllous:

16 Plants aquatic or in splash zones WARDIACEAE (p. 538)

450

16 Plants of drier habitats:

17 Stems distinctly dendroid; leaf cells smooth:

18 Stems incurved when dry; leaf apices obtuse LEPTODONTACEAE (p. 585)

18 Stems ± erect when dry; leaf apices acute to acuminate:

19 Leaves ± flat, leaf cells rhomboid, pitted THAMNOBRYACEAE (p. 592)

19 Leaves noticeably concave with flattened apices:

20 Leaf cells elongate, walls pitted PTEROBRYACEAE (p. 565)

20 Leaf cells rounded, walls smooth THAMNOBRYACEAE (p. 592)

17 Stems erect, pendent or creeping, if branched, not distinctly dendroid; leaf cells

smooth or papillose:

21 Stems and branches ± erect, perpendicular to substrate:

22 Alar cells not differentiated; costa single; calyptra initi ate, large, frequently cover-

ing the capsule; capsule exserted ORTHOTRICHACEAE (p. 476)

22 Alar cells differentiated; costa absent, single or double; calyptra cucullate or

conical and split up one side, not covering the capsule; capsule immersed or

exserted:

23 Leaf cells papillose PRIONODONTACEAE (p. 558)

23 Leaf cells smooth:

24 Costae double:

25 Leaf margins serrate; alar cells numerous, quadrate or transversely rectangu-

lar, green, walls smooth LEUCODONTACEAE (p. 553)

25 Leaf margins entire; alar cells in small group, thickened, pitted

PTEROBRYACEAE (p. 565)

24 Costae single:

26 Erect stems unbranched; leaves squarrose PTEROBRYACEAE (p. 565)

26 Erect stems irregularly branched; leaves erect to spreading:

27 Leaf margins ± entire; capsules exserted LEPTODONTACEAE (p. 585)

27 Leaf margins serrate above; capsules immersed . . CRYPHAEACEAE (p. 550)

21 Stems and branches pendent or ± creeping along substrate:

28 Plants with long pendent stems and branches:

29 Leaf cells smooth:

30 Branch leaves spirally ranked PTEROBRYACEAE (p. 565)

30 Branch leaves not in distinct rows METEORIACEAE (p. 575)

29 Leaf cells papillose:

31 Papillae seriate, or if papillae single then leaf margins strongly serrate

. TRACHYPODACEAE (p. 560)

31 Papillae numerous, scattered over cell lumens, or if papillae single then leaf

margins serrulate METEORIACEAE (p. 575)

28 Plants creeping over substrate:

32 Leaf cells papillose ORTHOTRICHACEAE (p. 476)

32 Leaf cells smooth:

33 Leaf margins distinctly bordered by elongated cells . . . HOOKERIACEAE (p. 601 )

33 Leaf margins not bordered:

34 Leaves complanate and undulate; capsules immersed

NECKERACEAE (p. 590)

34 Leaves appressed or spreading, flat; capsules exserted:

35 Costae strong and double, to midleaf or above . . . HOOKERIACEAE (p. 601 )

35 Costae single:

36 Stems heterophyllous; leaves with short awn .... RACOPILACEAE (p. 53 1 )

36 Stems homophyllous; leaves muticous .... ORTHOTRICHACEAE (p. 476)

451

ERPODIACEAE

Plants small, stringy, in loose wefts; corticolous or saxicolous. Stems branched; central strand

weak or absent; paraphyllia and pseudoparaphyllia absent. Leaves crowded or somewhat distant,

concave, clasping the stem or erect, ovate to elliptical, apex acute to rounded, muticous or with

long hyaline awn. Laminal cells uniform, quadrate to rectangular, smooth or papillose, weakly

thickened, alar cells not differentiated.

Autoicous. Perichaetia on short branches. Seta short or absent. Capsule immersed or just emer-

gent, cylindrical. Peristome absent or rudimentary. Operculum convex, beaked. Ccilyptra campanu-

late, sometimes twisted and clasping the seta, ribbed, naked. Spores large, granulate.

A family of five genera found in temperate regions of the world; two genera are represented in

southern Africa in dry woodland situations.

These tiny plants generally grow as scattered stems or loose groups on the bark of trees or rocks.

Macroscopically the plants resemble some leafy liverworts and can be easily overlooked.

Leaves uniform in size and shape; capsule immersed to emergent; calyptra not twisted ....

1. Erpodium

Leaves dimorphic, those facing substrate narrower and somewhat smaller; capsule exserted;

calyptra spirally twisted 2. Aulaeopilum

1. ERPODIUM

Erpodium (Brid.) C. Mull, in Bot. Zeitung (Berlin) 1: 11 A (1843); Broth, in Natiirl. PflFam., edn

2, 11: 2 (1925); Sim, Bryo. S. Afr. 347 (1926); Crum in Nova Hedwigia 23: 205 (1972). Type

species: E. domingense (Spreng.) C. Mull.

Plants small, slender; terricolous or saxicolous. Stems creeping, branched; central strand small

or absent. Leaves crowded, appressed and flattened against stem, ovate to elliptical, muticous or

awned; ecostate. Laminal cells small, isodiametric, smooth or papillose, walls weakly thickened.

Autoicous. Perigonia on branches, gemmate. Perichaetia on short branches along stem, obvious;

perichaetial leaves sheathing. Capsule immersed or emergent; annulus frequently well developed.

Peristome absent or rudimentary. Calyptra campanulate or mitrate, ribbed. Spores large, granulate.

A genus of 17 species, Erpodium is known from temperate and subtropical regions primarily in

the southern hemisphere. Four species are recognized in the Flora area.

1 Leaves with hyaline awn:

2 Leaf cells papillose 1 . E. beccarii

2 Leaf cells smooth 2. E. coronation subsp. transvaaliense

1 Leaves muticous:

3 Leaves broad, elliptical to orbicular; apex obtuse to rounded, cuspidate 3. E. grossirete

3 Leaves ovate to ovate-lanceolate; apex acute to obtuse 4. E. distichum

452

Erpodiaceae

Erpodiaceae

453

1. Erpodium beccarii C. Mm//, in Nuovo

Giom. Bot. Ital. 4: 18 (1872); Crum in Nova

Hedwigia23: 211 (1972). Type: Bogos, Abyssinia,

O. Beccari s.n.

Aulacopilum beccarii (C. Mull.) Mitt, in J. Linn. Soc.,

Bot. 13: 308 (1873).

Erpodium hanningtonii Mitt, in J. Linn. Soc., Bot. 22: 313

(1886); Sim, Bryo. S. Afr. 347 (1926).

Erpodium joannis-meyeri C. Mull, in Flora 73: 486

(1890).

Erpodium menyharthii C. Mull, in Verh. Zool. Bot. Ges.

Wien. 43: 13, 14 (1893).

Plants small and slender, creeping or in

wefts, dark green to grey-green; corticolous.

Stems to 20 mm long, branches few, scattered.

Leaves evenly spaced, erect-spreading wet, ap-

pressed dry; broadly ovate to elliptical, 1.0- 1.2

mm long; obtuse; awns hyaline, papillose, to

0.5 mm long; rounded at base; margins plane,

entire; ecostate. Upper laminal cells hexagonal

to subhexagonal, 10-18 pm long, walls weakly

thickened, homogeneous, papillose on both sur-

faces, papillae C-shaped, centred over lumen;

basal cells not strongly differentiated, quadrate

to rectangular, 25-37 pm long; alar cells not

strongly differentiated, transversely rectangular,

hyaline, walls thin.

Perigonial leaves ovate-acute, 0. 3-0.4 mm

long. Perichaetia strongly differentiated, hya-

line; perichaetial leaves with base sheathing,

ovate-acuminate, 1 .5-1.8 mm long, leaf cells

irregularly rectangular to quadrate at margins,

elongate fusiform to rhomboidal, alar cells

quadrate, not well defined. Seta 0. 1-0.3 mm

long, brown, smooth. Capsule immersed, erect,

short-cylindrical to broadly ovoid, 1 mm long,

smooth, yellow-brown, gymnostomous; exothe-

cial cells rectangular, walls thin, cells at mouth

not differentiated, annulus persistent, thick-

walled, in 3 or 4 rows, neck cells not differenti-

ated; stomata on lower urn, phaneropore. Oper-

culum convex-rostrate, beak curved or bent.

Calyptra campanulate, lobed and ribbed, serrate

on ribs. Spores rounded, 30—4-0 pm, weakly

papillose, brown. Fig. 126: 12-18.

Found on bark in woodlands of Central Ame-

rica, temperate South America, and eastern and

southern Africa and Madagascar. Erpodium bec-

carii is rarely collected in Namibia, KwaZulu-

Natal, Botswana and the Free State, but is more

common in woodland communities of the

Transvaal regions and Zimbabwe. Map 177.

Vouchers: Magill 3631, 4957, 5025; Vahr-

meijer 121B, 13151.

This species is most easily identified by its

long, hyaline awns that strongly contrast with

the dark green colour of the leaves. The papil-

lose cells of the awn and leaf lamina separate

this species from E. coronatum subsp. trans-

vaaliense.

2. Erpodium coronatum {Hook. & Wilson)

Mitt, subsp. transvaaliense {Broth. & Wager)

Fig. 126. — Erpodium coronatum subsp. transvaaliense (1-11): 1. habit (dry), x 1; 2. habit (wet), x 10; 3. stem in cross

section, x 175; 4. leaf, x 35; 5. basal leaf cells, x 122; 6. upper laminal cells, x 245; 7. cells at leaf apex, x 122; 8. capsule

with attached perichaetial leaf, x 25; 9. part of capsule mouth showing persistent annulus and peristome, x 175; 10. calyp-

tra, x 50; 11. spore, x 495. E. beccarii (12-18): 12. habit (dry), x 1; 13. habit (wet), x 10; 14. leaf, x 35; 15. upper laminal

cells, x 245; 16. perichaetial leaf, x 25; 17. part of capsule mouth showing exothecial cells and capsule mouth, x 175; 18.

spore, x 495. E. grossirete (19-24): 19. habit (dry), x 1; 20. habit (wet), x 10; 21. leaf, x 35; 22. upper laminal cells, x 245;

23. leaf apex, x 122; 24. perichaetial leaf, x 35. (1-3, 8 & 9, Smook 3181\ 4—7, 10 & 11, Magill 3039; 12, Magill 4958; 13,

Magill 3629; 14 & 15, Magill 3633; 16-18, Van Vuuren 1703; 19-24, Vahrmeijer 121a.)

454

Erpodiaceae

Magill, stat. nov. Type: Transvaal, Wolhuter,

Wager 189 (BM, holo.; PRE, GRA, iso.).

Erpodium transvaaliense Broth. & Wager in Dix. in

Trans. Roy. Soc. South Africa 8: 208 (1920); Broth, in

Natiirl. PflFam., edn 2, 1 1 : 3 (1925); Sim. Bryo. S. Afr. 347

(1926).

Plants small, creeping or in wefts, dark green

to brownish green; corticolous. Stems 10-20

mm long, branches few, scattered; in section

round, central strand small, inner cortical cells

thick-walled, hyaline, in 2 or 3 rows, outer cor-

tical cells thick-walled, brown, in 2 rows.

Leaves crowded, erect-spreading wet, appres-

sed dry; ovate to oblong, 0.8-1. 2 mm long;

acute to acuminate, aristate, awns hyaline,

smooth, length variable; rounded at base; mar-

gins plane, entire; ecostate. Upper laminal

cells irregular, walls thickened, homogeneous,

smooth on both surfaces; basal cells not strong-

ly differentiated, walls thin, papillae scattered,

hyaline-green, walls thin; alar cells not differen-

tiated.

Perichaetia green; leaves sheathing below,

oblong-cuspidate, 1. 8-2.0 mm long, leaf cells

quadrate to rectangular at margins, elongate to

rhomboidal at centre of leaf, alar cells quadrate,

not well defined. Seta 1 mm long, yellow,

smooth. Capsule immersed, erect, broadly ovoid,

1 .8 mm long, smooth, yellow to brown; exothe-

cial cells rectangular, walls thin, wavy, cells at

mouth not differentiated; annulus persistent, well

developed, thick-walled, 3 or 4 rows high; neck

cells not differentiated; stomata on lower urn,

phaneropore. Peristome single, hyaline; exos-

tome teeth irregular, erect, papillose, 150 pm

high. Operculum convex, long-rostrate, 0.5 mm

long. Calyptra mitrate, lobed and ribbed, 1 mm

long, naked. Spores rounded, 35-40 pm, weakly

papillose, light brown. Fig. 126: 1-11.

Endemic to southern Africa, E. coronatum

subsp. transvaaliense is the most commonly

collected Erpodium in the Flora area. The

plants are very small and difficult to locate in

crevices of tree bark in the northern and central

Transvaal regions, northern Cape area,

KwaZulu-Natal and northern Namibia. Map

178.

Map 178. — • Erpodium coronatum subsp. transvaaliense

♦ Erpodium grossirete

Vouchers: Magill 3039, 3646, 6391; Perold

115; Smook 1822.

The long, smooth, hyaline hairpoint and

smooth leaf cells separate this taxon from the

other southern African species. It differs from

the more widespread E. coronatum (Hook. &

Wils.) Mitt, in its well developed awn and

weakly ribbed calyptra. Although locally dis-

tinct, this taxon is best treated as a subspecies of

E. coronatum.

3. Erpodium grossirete C. Mull, in Verh.

Zool. Bot. Ges. Wien 43: 13 (1893); Broth, in

Natiirl. PflFam., edn 2, 11: 3 (1925). Type:

Zambesia, Boroma, Menyharth s.n ., Aug. 1890.

Plants small, creeping, compact, dark green;

corticolous. Stems julaceous, 10 mm long,

branching irregular, ± dense and erect from

creeping stem; in section round, central strand

absent, inner cortical cells thin-walled, hyaline,

in 3 rows, outer cortical cells weakly thickened,

in single row. Leaves ± crowded, erect-appressed

wet, appressed dry, weakly concave; stem

leaves broadly elliptical to ± orbicular, 0.6- 1.0

mm long; obtuse to rounded or truncate,

mucronate to cuspidate; abruptly rounded at

base; margins plane, entire, bordered; ecostate.

Upper laminal cells irregularly quadrate to

transversely hexagonal, 12-25 pm long, 12-31

Erpodiaceae

455

pm wide, walls ± thickened, homogeneous,

smooth on both surfaces; basal cells similar to

upper cells, 12-25 pm long, 15-31 pm wide,

hyaline, smooth; alar cells similar to basal cells

but more strongly transversely rectangular, hya-

line-green, walls thin.

Perigonial leaves ovate-acute. Perichaetia

green but quickly becoming hyaline; perichae-

tial leaves oblong-acuminate, strongly sheath-

ing, 2.0 mm long, leaf cells rectangular-oblong

or fusiform, shorter at margins, somewhat

thickened, basal and alar cells not well defined.

Seta very short. Capsule immersed, erect, cylin-

drical, 1.0-1. 6 mm long, smooth, yellow, gym-

nostomous; exothecial cells ± irregular, walls

thin, cells at mouth not differentiated; annulus

persistent, thick-walled, in 1 or 2 rows; neck

cells not differentiated; stomata not seen.

Operculum and calyptra not seen. Spores

rounded, 37 pm, granulate, brownish. Fig. 126:

19-24.

Endemic to southern Africa, E. grossirete is

rarely collected from bark of trees along

streams and in swamp forests in Malawi, north-

eastern Zimbabwe, Mozambique and the east-

ern Caprivi Strip of Namibia. Map 178.

Vouchers: Magadza 162; Vahrmeijer 121A.

Sim (1926) was incorrect in his interpreta-

tion of this species; all the specimens Sim

examined were E. distichum. These two species

can be easily separated on the basis of plant

habit, leaf shape, and leaf cell shape and size,

although the two species are similar in many

sporophytic characteristics. The rather large

cylindrical capsules and very short setae clearly

separate these two species into a distinct group

within the genus.

4. Erpodium distichum Wager & Dix. in

Trans. Roy. Soc. South Africa 8: 208 (1920);

Broth, in Natiirl. PflFam., edn 2, 11: 3 (1925).

Syntypes: Transvaal, Barberton, Wager 279\

Natal, Pietermaritzburg, Wager 226.

Plants small, slender, creeping, dark green;

corticolous. Stems up to 10 mm long, branches

few, scattered; in section round, central strand

absent, inner cortical cells thin-walled, hyaline,

in 3 rows, outer cortical cells hyaline, in single

row. Leaves evenly spaced, erect-spreading wet,

appressed dry; ovate to ovate-lanceolate, 0.8-

1 .2 mm long; acute to obtuse; weakly rounded

at base; margins plane, entire; ecostate. Upper

laminal cells hexagonal-fusiform, marginal

cells smaller, transversely rectangular, 25—41

pm long, 12-18 pm wide, walls thin to slightly

thickened at comers, homogeneous, smooth on

both surfaces; basal cells rectangular centrally,

transversely rectangular at margins, 12-15 pm

long, 30-40 pm wide, hyaline; alar cells

quadrate to transversely rectangular, 12 x 18

pm, hyaline-green, walls thin.

Perichaetia obvious, green; perichaetial

leaves oblong-acute, sheathing 1.5-1. 8 mm

long, leaf cells not strongly differentiated from

vegetative leaf cells. Seta very short. Capsule

Fig. 127.— Erpodium distichum: 1. habit (dry), x 5; 2.

habit (wet), x 15; 3. leaf, x 35; 4. basal leaf cells (right

side), x 175; 5. leaf apex, x 175; 6. perichaetial leaf, x 35;

7. spore, x 495. ( 1 , Wager PRE-CH 11 71 7; 2, Wager PRE-

CH 11944; 3-7, Wager PRE-CH 1059.)

456

Erpodiaceae

Map 179. — • Aulacopilum trichophyllum

♦ Erpodium distichum

erect, cylindrical, 1.5 mm long, smooth; exothe-

cial cells shortly rectangular, walls thin, cells at

mouth smaller, quadrate, in 3 or 4 rows; annu-

lus absent; neck cells not differentiated; stoma-

ta not seen; gymnostomous. Operculum and

calyptra not seen. Spores rounded, 35-38 pm,

granulate, light brown. Fig. 127.

Endemic to southern Africa, E. distichum is

rarely collected on trees in Zimbabwe, Mozam-

bique and the eastern Transvaal region. No

recent specimens have been seen and all the

South African specimens appear to be from a

single large collection made by Wager at

Barberton in 1914, although 'Maritzburg’ is also

cited on one of the syntypes. Map 179.

Vouchers; Eyles 2702; Sim 8988.

An unusual, narrow-leaved, muticous

species that is distinct gametophytically from

the other southern African species. The

extremely short seta clearly indicates a relation-

ship to E. grossirete (see p. 454). It is unclear

why the species has not been recollected, unless

it is restricted to a very small area of the eastern

Transvaal region.

2. AULACOPILUM

Aulacopilum Wilson in London J. Bot. 7: 90 (1848); Broth, in Natiirl. PflFam., edn 2, 11:4 (1925);

Sim, Bryo. S. Afr. 346 (1926); Crum in Nova Hedwigia 23: 218 (1972). Type species: A. glaucum

Wilson.

Plants small, slender; corticolous or saxicolous. Stems creeping, branched; central strand absent.

Leaves weakly concave, appressed, elliptical to ovate; apiculate or with short or long hyaline awn;

ecostate. Laminal cells isodiametric, multipapillose, walls weakly thickened.

Autoicous. Perigonia on stem, gemmate. Perichaetia on short branches; perichaetial leaves sheath-

ing. Capsule exserted, annulus not strongly differentiated. Peristome absent. Calyptra spirally

twisted, ribbed, clasping seta. Spores large, granulate.

Gametophytically Aulacopilum is similar in many respects to Erpodium ; however, the dimor-

phic leaves, (those facing the substrate are smaller and narrower) and the twisted calyptra are

important differences. The operculum remains attached to the columella and is retained by the

twisted and clasping calyptra, thus retarding spore discharge.

Aulacopilum trichophyllum Angstr. in C.

Midi in Bot. Zeitung (Berlin) 20: 393 (1862);

Sim, Bryo. S. Afr. 346 (1926). Type: South

Africa, Wahlberg s.n.

Aulacopilum incanum Mitt, in J. Linn. Soc., Bot. 13:

308 (1873).

Plants small, slender, in wefts, creeping, dark

green or sometimes glaucous; saxicolous or

corticolous. Stems flattened, up to 10 mm long,

branching ± regular; in section round, central

strand absent, inner cortical cells large, thin-

walled, hyaline, outer cortical cells smaller,

Erpodiaceae

457

thick-walled, yellow. Leaves dimorphic, evenly

spaced, ± appressed wet, appressed dry; upper

leaves elliptical to ovate, 0.6-1. 0 mm long;

acute, apiculate or short-awned; rounded at

base; margins plane, entire, ecostate; lower

leaves somewhat smaller and narrowly ovate-

acuminate. Upper laminal cells rounded hex-

agonal, 12-16 pm long, walls weakly thick-

ened, homogeneous, papillose on both surfaces,

papillae dense, centred over lumens; basal cells

rectangular only at central part of insertion,

12-16 pm long, 2^1 pm wide, hyaline; alar

cells not forming distinct groups.

Perigonial leaves ovate-cuspidate, 0.4 mm

long. Perichaetia green; perichaetial leaves

oblong-acuminate, sheathing, larger than stem

leaves but not strongly differentiated, 1.0- 1.2

mm long, leaf cells rectangular to rhombic. Seta

up to 1 mm long, yellowish, smooth. Capsule

exserted, erect, short-cylindrical, 0.5 mm long,

smooth, yellow-brown; exothecial cells

quadrate to rectangular, walls weakly thick-

ened, cells at mouth transversely rectangular in

1 or 2 rows; annulus weakly differentiated;

stomata on neck, phaneropore; gymnostomous.

Operculum convex-rostrate, cells not twisted.

Calyptra mitrate, large, completely covering

capsule and clasping seta, up to 1 mm long,

twisted and slit down one side, plicate, naked.

Spores rounded, 22-26 pm, granulate, yellow-

brown. Fig. 128.

These plants grow on trees or boulders in the

northern, eastern and central Transvaal areas,

Zululand and the eastern Cape region. The species

has also been reported from Uganda, Malawi and

Zimbabwe. The small, loose patches of dark green

threads are easily overlooked in open woodland

and coastal forest habitats. Map 179.

Vouchers: Linder 1232; Magill 4982, 6368;

Oliver 7354; Smook 1466; Wager 319.

Fig 128. — Aulacopilum trichophyllum: 1. habit (dry),

x 3; 2. habit (wet), x 5; 3. stem in cross section, x 350; 4.

dimorphic leaves, x 70; 5. basal leaf cells (left side), x 175;

6. upper laminal cells at right margin, x 700; 7. leaf apex, x

175; 8. perichaetial leaf, x 70; 9. habit showing capsule

with operculum, x 25; 10. calyptra, x 35; 11. spore, x 700.

(1 & 2, Magill 4982: 3-5, 8, 1 1 & 12, Junod 12: 6,1,9 &

10, Magill 3210.)

458

Erpodiaceae

The southern African plants are generally

dark green and do not exhibit the strong glau-

cous coloration of the Australian species A.

glaucum to which they are most closely related.

They also have a much stronger awn than the

Australian species.

459

RHACHITHECIACEAE

Plants minute to small, gregarious or caespitose, yellowish green; corticolous. Stems erect, cen-

tral strand of hydroids present. Leaves larger above, erect-incurved when dry, erect-spreading when

wet, oblong-spathulate; costa ending well below apex. Cells irregularly hexagonal, rectangular

below, thin-walled, smooth. Gemmae oblong, axillary.

Autoicous. Perigonia gemmate. Perichaetia terminal; leaves long, sheathing, covering seta.

Seta thick, straight or curved above, vaginula long. Capsule erect to inclined, ovate-cylindrical,

strongly 8-ribbed; stomata phaneropore; annulus revolvable. Peristome single, teeth in 8 pairs, tra-

beculate, smooth. Operculum conic-apiculate, oblique. Calyptra cucullate, widely split. Spores

rounded to elliptical.

RHACHITHECIUM

Rhachithecium Broth, ex Le Jolis in Mem. Soc. Sci. Nat. Cherbourg 29: 308 (1895); Sim, Bryo.

S. Afr. 272 (1926); Gangulee, Moss. E. India 5: 1158 (1976). Type species: not designated.

A genus of three species found in central and southern America, Africa, Madagascar, India, Sri

Lanka, southern and southeastern Asia and Japan. Rhachithecium perpusillum is the most wide-

spread of the three species and the only one known from the southern hemisphere.

Rhachithecium perpusillum (Thwait. &

Mitt.) Broth, in Natiirl. PflFam. 1,3: 1199 (1909);

Gangulee, Moss. E. India 5: 1159 (1976). Type:

Sri Lanka.

Zygodon perpusillus Thwait. & Mitt, in J. Linn. Soc.,

Bot. 13: 303 (1873).

Hypnodon transvaalensis C. Mull, in Hedwigia 38: 126

(1899). Rhachithecium transvaalense (C. Mull.) Broth, in

Natiirl. PflFam. 1,3: 1199 (1909); Sim, Bryo. S. Afr. 272

(1926); Crum in Bryologist 59: 32 (1956). Type: Transvaal,

near Utombi between Kook and Sand Rivers, Aug. 1884,

Wilms s.n. (PRE! ).

Plants minute to small, gregarious or caespi-

tose, yellowish green above, yellowish brown

below; corticolous. Stems erect, to 3.5 mm tall,

branching by subperichaetial innovations; rhi-

zoids brownish, smooth; in section round, cen-

tral strand of hydroids present, cortical cells in

2-4 rows, thin-walled, epidermis not differenti-

ated. Leaves ± crowded, larger above, erect-

incurved when dry, erect-spreading when wet,

keeled; oblong to oblong-spathulate to spathu-

late, (0.7—) 1 .0—1 ,6(— 1 .8) mm long; apex suba-

cute, mucronate or apiculate; margins plane,

entire below, frequently crenulate to serrulate

above, frequently flexuose. Costa ending well

below apex, ventral superficial cells rectangu-

lar, dorsal superficial cells linear; in section

crescent-shaped, ventral surface flat, laminal

insertion ventral, 2 guide cells exposed ventral-

ly, dorsal stereids in 2 or 3 rows, dorsal surface

cells not differentiated. Upper laminal cells

irregularly hexagonal, ± thin-walled, flat to

bulging, 12-29 pm, smooth; basal cells yellow-

ish, smooth, rectangular, thin-walled. Gemmae

oblong, 3-5 cells long, 1 or 2 cells wide,

80-130 pm long, 30-50 pm wide, axillary on

long stalks, yellowish or hyaline.

Map 180. — ♦ Rhachithecium perpusillum

• Ptychomitriopsis aloinoides

■ Ptychomitriopsis africana

460

Rhachitheciaceae

Autoicous. Perigonia on short branches,

gemmate; inner leaves concave, broadly ovate ±

apiculate, 0.2-0. 5 mm long. Perichaetia termi-

nal; leaves yellowish or hyaline, sheathing, cov-

ering seta, 1. 2-2.4 mm long, apex acute or

acuminate, cells irregularly hexagonal or rhom-

boidal above, rhomboidal to rectangular below,

thin-walled. Seta thick, 1.0-1. 5 mm long, yel-

lowish to brown, twisted anticlockwise above,

straight to curved above, vaginula long. Cap-

sule erect to inclined, ovate-cylindrical, con-

tracted below mouth when dry, 0.5-0. 8 mm

long, brownish, strongly 8-ribbed, neck short;

exothecial cells irregularly rectangular to

quadrate, shorter at mouth, thin-walled, cells of

ribs inflated, brownish or orange; stomata pre-

sent on neck, phaneropore; annulus revolvable.

Peristome teeth 16, in 8 pairs, oblong-lanceo-

late, brownish or orange, recurved when dry,

incurved when wet, trabeculate, smooth. Oper-

culum shortly conic-apiculate, apiculus oblique,

0.2-0. 3 mm long. Calyptra 0.4-0.6 mm long,

naked, cells prorate distally, yellowish brown.

Spores ( 1 8.5— )22.0— 3 1 ,0(— 35.0) pm, minutely

granulate to reticulate, brownish. Fig. 129.

The species is known from southern North

America, Mexico, South America, sub-Saharan

Africa, Madagascar, Sri Lanka, India and China.

In southern Africa R. perpusillum is rarely col-

lected on bark in woodlands and forests of the

northern and eastern Transvaal regions. Map 180.

Vouchers: Kluge 1042, 1048; Magill 3607,

6502, 6504.

The small plants, growing on b.ark of trees,

can easily be overlooked in the field. The spe-

cies can be recognized by the long-sheathing

perichaetial leaves, covering the long vaginula

and relatively thick seta and reaching the base

Fig. 129. — Rhachithecium perpusillum: 1. habit (dry),

x 10; 2. habit (wet), x 10; 3. stem in cross section, x 175; 4

& 5. leaves, x 35; 6. leaf in cross section, x 175; 7. cells at

leaf base (right side), x 175; 8. upper lamina! cells, x 350;

9. leaf apex, x 175; 10. axillary gemma, x 250; 11.

perichaetial leaf, x 35; 12. part of capsule wall with stoma,

x 350; 13. part of capsule mouth with incurved peristome

tooth, x 175; 14. calyptra, x 70; 15. spore, x 700. (1,2, 4—11

& 14, Kluge 1048 ; 3, Kluge 1042 ; 12, Magill 3607\ 13,

Magill 6502.)

Rhachitheciaceae

461

of the strongly 8-ribbed capsule. The oblong-

spathulate leaves with irregular hexagonal,

smooth upper laminal cells and the single peri-

stome with teeth smooth, recurved when dry and

incurved when wet also help to identify R. per-

pusillum.

462

PTY CHOMITRIACEAE

Plants small to medium-sized, loosely or densely caespitose, dark green to blackish green,

glossy above, brownish and dull below; saxicolous or rarely terricolous. Stems erect, branching

irregular; in section round, central strand small, inner cortical cells in 4 or 5 rows, thin-walled, yel-

lowish, outer cortical cells in 2-4 rows, somewhat thickened, reddish. Leaves erect-spreading with

plane to erect or rarely involute margins wet, incurled with involute margins dry; margins and/or

lamina generally bistratose, occasionally unistratose or multistratose. Costa smooth dorsally,

smooth or mammillose ventrally; in section with guide cells and dorsal and ventral stereid bands.

Laminal cells rounded-quadrate to short-rectangular or transversely rectangular especially near

margins, smooth, ± thickened; alar cells not differentiated.

Autoicous. Perichaetia terminal becoming lateral through innovation, occasionally polyseta-

ceous; perichaetial leaves undifferentiated or somewhat narrower. Seta erect, short or long. Capsule

erect, symmetrical, ovoid to short-cylindrical; exothecial cells irregularly rectangular, thin-walled;

stomata present at base of urn; annulus present. Peristome single, teeth 16, narrow-triangular, cleft

or perforated, ± fragile, ornately papillose. Operculum long-beaked. Calyptra mitrate, deeply

lobed, ± plicate, naked. Spores small.

A small family with four genera, two of which occur in southern Africa. Ptychomitrium is a

large, widespread genus found on rocks, generally in open, upland sites. Ptychomitriopsis is

endemic to southern Africa and is found on rocks in dry grasslands.

In view of the evidence presented by Edwards (1979) on peristome development, this family

and most of its genera are incorrectly placed in the Diplolepidae and belong to the Haplolepidae

near Grimmiaceae.

Leaf lamina unistratose, or if bistratose then leaf margins broadly involute when wet; lami-

nal cells mostly larger than 10 pm 1. Ptychomitriopsis

Leaf lamina bistratose, or only margin bistratose, leaf margin plane to erect when wet; lami-

nal cells mostly smaller than 10 pm 2. Ptychomitrium

1 PTYCHOMITRIOPSIS

Ptychomitriopsis Dix. in J. Bot. 69: 284 (1931). Type species: P. africana Dix.

Plants small; saxicolous. Stems erect. Leaves oblong to narrowly elliptical; obtuse to rounded;

margins plane to broadly involute, entire. Costa subpercurrent or ending below apex, ventral super-

ficial cells similar to laminal cells, smooth, dorsal superficial cells long-rectangular, thickened,

smooth. Laminal cells small, weakly thickened.

Autoicous. Seta short, 2-3 mm long. Capsule cylindrical to ellipsoidal. Peristome teeth 16, cleft

and perforated. Spores small, essentially smooth to weakly papillose.

Ptychomitriopsis contains only two species, both endemic to southern Africa. The genus has

been rarely collected, perhaps because ol its small stature and occurrence in dry grassland habitats.

Macroscopically the genus is easily contused with small plants of Ptychomitrium. The genus is

separated from Ptychomitrium by smaller overall size, larger laminal cells, shorter basal cells, and

differences in leaf shape and anatomy.

Ptychomitriaceae

463

Leaves bistratose, upper margins broadly involute

Leaves unistratose, upper margins plane

1. Ptychomitriopsis aloinoides MagiiL sp.

nov., bene distinctci a speciebus aliis

Ptychomitriopsidis Dix., foliis bistratosis et

marginibus late involutis. Type: (Orange) Free

State, north slope of Wonderkop between

Rosendal and Marquard, 2827DA, on sandstone

boulder, 1800 m. Van Rooy 439 (MO, holo.!;

BM, NY, PRE).

Plants small, loosely caespitose, green to yel-

low-green; saxicolous. Stems 2-4 mm tall,

rarely branched; in section round, central strand

small, inner cortical cells in 2 or 3 rows, large,

thin-walled, outer cortical cells in 1 or 2 rows,

smaller, thin-walled, brownish. Leaves ± crowd-

ed, erect-spreading wet, incurved and appressed

dry; bistratose above base; oblong to lingulate,

1.0-1. 5 mm long; obtuse to rounded; base

scarcely differentiated; margins entire, plane to

erect below, broadly involute above. Costa end-

ing below apex to subpercurrent; in section

elliptical, guide cells 4, distinct, ventral stereid

band small, 2 cells thick, ventral surface cells

similar to ventral laminal cells, forming a con-

tinuous ventral layer, dorsal stereid band strong,

(2)3 or 4 cells thick, dorsal surface cells small,

incrassate but more strongly thickened on outer

wall. Upper laminal cells rounded-quadrate to

transversely rectangular (7-) 1 1-14 pm, smooth;

basal cells somewhat larger, short-rectangular,

thin-walled, hyaline.

Autoicous. Perichaetia terminal, leaves not

differentiated. Seta 2.0-3. 5 mm long, yellowish to

brownish. Capsule ellipsoidal, 1.0-1. 2 mm long,

brownish; annulus present, red. Peristome teeth

0.14 mm high, cleft and perforated, reddish yel-

low. Operculum rostrate, 0.7 mm long. Calyptra

mitrate, 1 .0—1.5 mm long. Spores rounded to ±

angular, 9-13 pm, essentially smooth, light green

to yellowish. Fig. 130: 13-18.

Endemic to southern Africa, P. aloinoides is

found on sandstone rocks or rock crevices in

grassland or shrubland of the Free State, north-

1 . P. aloinoides

. 2. P africana

em and northwestern Cape regions and Nami-

bia. Map 180.

Vouchers: Barnard CH-5762; Magill 6458;

Volk 5349.

This species differs from P. africana in its

broadly involute upper leaf margins, bistratose

lamina and slightly larger spores. The species is

separated from Ptychomitrium by its small

stature, larger leaf cells and leaf shape.

2. Ptychomitriopsis africana Dix. in J.

Bot. 69: 284 (1931). Type: Transvaal, Sout-

pansberg, Lake Fundudzi, Wager 112 (BM,

holo.!; PRE).

Ptychomitrium africanum (Dix.) Churchill in Funk &

Brooks, Advances in cladistics 143 (1981).

Ptychomitrium godfreyi Robinson in Bryologist 62: 225

(1960). Type: Natal, 25 miles NE of Ladysmith, 20 Feb.

1953, Godfrey (Duke, holo.!; NY. PRE, US).

Plants small, loosely to densely caespitose, .

green to yellow-green; saxicolous. Stems 3-5

mm tall, sparsely branched; in section round,

central strand large, inner cortical cells in 2 or 3

rows, large, thin-walled, outer cortical cells in

1(2) row(s), ± smaller, thin-walled, brownish.

Leaves erect-spreading wet, incurved and ±

crisped dry; unistratose; narrowly elliptical to

oblong, 2. 0-2. 5 mm long; obtuse to rounded

and short-apiculate; tapering to base; margins

plane, entire. Costa subpercurrent; in section

elliptical, guide cells 4, small, ventral stereid

band small, 2 cells thick, ventral surface cells

large, similar to laminal cells, thin-walled, dor-

sal stereid band 2 or 3 cells thick, dorsal surface

cells small, thickened or occasionally subster-

eids. Laminal ceils subquadrate to + transverse-

ly rectangular, (8—) 11—14 pm, firm-walled; basal

cells hyaline, short-rectangular, 2:1 to 3:1, thin-

walled.

Perichaetia terminal, leaves undifferentiated.

Seta 2-3 mm long, yellowish. Capsule cylindri-

464

Ptychomitriaceae

Ptychomitriaceae

465

cal, 1.0-1. 3 mm long, brownish with red mouth.

Peristome teeth lanceolate, cleft, papillose, red-

dish. Operculum rostrate, 0.7 mm long. Calyptra

mitrate, 0.8 mm long. Spores rounded, 7-8 pm,

yellowish, weakly papillose. Fig. 130: 1-12.

Ptychomitriopsis africana is endemic to

southern Africa. It is found on rock in grassland

in KwaZulu-Natal and the northern Transvaal

region. Map 180.

Vouchers: types only.

This species is identified by its small size,

narrowly elliptical leaves with rounded-obtuse

apices, and unistratose leaf lamina. As indicated

by Dixon (1931), the leaf cells of this plant are

‘large, very distinct and pellucid’, clearly sepa-

rating it from other southern African members

of this family.

2 PTY CHOMITRIUM

Ptychomitrium Fuernr. in Flora 2: 19 (1829), nom. cons.; Broth, in Natiirl. PflFam., edn 2, 1 1 :

8 (1925); Sim, Bryo. S. Afr. 213 (1926); Catcheside, Moss. S. Austr. 207 (1980); Crum &

Anderson, Moss. E.N. Amer. 2: 666 (1981). Type species: P. polyphyllum (Sw.) B.S.G.

Brachysteleum Reichenb., Consp. regn. veg. 1: 34 (1828), nom. rejec.; C. Mull., Syn. muse, frond 1: 766 (1849). Type

species: B. crispatum (Hedw.) Homsch.

Notarisia Hampe in Linnaea 11: 379 (1837). Type species: N. virginica Hampe.

Plants medium-sized; saxicolous or terricolous. Leaves tightly incurled dry, widespreading wet;

linear-lanceolate to ovate-lanceolate or ligulate above ovate to oblong base; margins plane to erect,

bistratose, or complete lamina bistratose, occasionally multistratose juxtacostally. Costa percur-

rent to subpercurrent; ventral superficial cells smooth to mammillose, similar to laminal cells; dor-

sal superficial cells smooth, rectangular, thickened. Laminal cells firm-walled to thickened.

Autoicous. Seta short or long. Capsule ovoid to short-cylindrical. Peristome teeth ± fragile,

mostly cleft and perforated. Spores granulate to weakly papillose.

Ptychomitrium contains 74 species found on rock or soil in rock crevices in temperate regions

of both hemispheres. The plants are generally fully exposed and have a distinctive dark green but

glossy appearance.

In southern Africa, the genus is most frequently found around rock outcrops in grassland or

shrubland of the eastern and southern Flora area.

1 Leaf margins bistratose, lamina unistratose or with bistratose patches 1 . P suberispatum

1 Leaf lamina bistratose, occasionally multistratose juxtacostally:

2 Laminal cells strongly mammillose:

3 Ventral laminal cells mammillose; dorsal laminal cells ± flat or occasionally papillose

marginally, mostly smaller than ventral cells; eastern Transvaal region, Swaziland,

Zululand 2. P. exaratifolium

Fig. 130. — Ptychomitriopsis africana (1-12): 1. habit (dry), x 5; 2. habit (wet), x 10: 3. stem in cross section, x 175;

4. leaf, x 35; 5. leaf in cross section, x 175; 6. basal leaf cells, x 350; 7. upper laminal cells, x 700; 8. leaf apex, x 175; 9.

part of capsule mouth with peristome teeth, x 175; 10. operculum, x 70; 11. calyptra, x 35; 12. Spores, x 700. P. aloinoides

(13-18): 13. habit (dry), x 5; 14. habit (wet), x 10; 15. leaf, x 35; 16. leaf in cross section, x 175; 17. part of peristome, x

350; 18. spores, x 700. Ptychomitrium suberispatum (19-23): 19. habit (dry), x 1; 20. habit (wet), x 5; 21. leaf, x 35; 22.

leaf in cross section, x 175; 23. leaf apex, x 175. (1-12, Wager 112\ 13 & 14, Giess 15385 ; 15-17, Russell 3932\ 18, Volk

5349\ 19 & 22, Schelpe 7583\ 20, Magill 5675; 21, Van Rooy 455\ 23, Cholnoky 4.)

466

PTYCH0M1TRIACEAE

3 Ventral and dorsal cells mammillose, similar in size and shape; Drakensberg of Lesotho

and western KwaZulu-Natal 3. P. diexaratum

2 Laminal cells smooth or bulging:

4 Leaf margins revolute in base 4. P depression

4 Leaf margins plane to ± reflexed in base:

5 Leaves abruptly ligulate above distinct obovate base 5. P. eurybasis

5 Leaves linear-lanceolate to ovate- or oblong-lanceolate, base not pronounced:

6 Leaf apex distinctly cucullate; leaves 2-3 mm long 6. P. cucullatifolium

6 Leaf apex acute (or rarely subcucullate); leaves 3-5 mm long:

7 Laminal cells 1 1-15 pm long; basal leaf cells short-rectangular, very thin-walled;

costa broad, in transverse section elliptical, ventral surface cells large, thin-

walled, distinct 7. P. crassinervium

7 Laminal cells less than 10 pm long; basal leaf cells rectangular to long-rectangu-

lar, ± thickened; costa narrow, in transverse section rounded, ventral surface

cells differentiated but generally not

1 . Ptychomitrium subcrispatum Ther. & P.

Varde in Revue Gen. Bot. 30: 65 (1918); Broth,

in Natiirl. PflFam., edn 2, 11:9 (1925); Sim,

Bryo. S. Afr. 216 (1926); De Sloover in Bull.

Jard. Bot. Belg. 46: 437 (1976). Type: Natal,

Van Reenen, 5000 ft, Wager no. 2 (PC, holo.!).

Glyphomitrium marginatum Wager & Dix. in Trans. Roy.

Soc. South Africa 8: 196 ( 1920). Ptychomitrium marginatum

(Wager & Dix.) Dix. in S. African J. Sci. 18: 315 (1922);

Broth, in Natiirl. PflFam., edn 2, 11:9 (1925); Sim, Bryo. S.

Afr. 216 (1926). Syntypes: Transvaal, Kaapsche Hoop,

Wager 298 (BM!, PRE!); Cape, Hogsback, Tjumie, D.B. &

M. Henderson 190 (BM!); D. Henderson 352b (BM!).

Plants medium-sized, loosely caespitose,

dark green; saxicolous or terricolous. Stems

1 0— 25(— 40) mm tall, irregularly branched.

Leaves incurved or appressed, with crisped apex

dry, erect-spreading wet; linear-lanceolate to

ovate- or oblong-lanceolate, (2.5— )3.5— 4.5(— 5.0)

mm long; acute to subcucullate; margins bi-

stratose, upper lamina unistratose, frequently

with bistratose patches. Costa percurrent, dorsal

and ventral superficial cells smooth; in section

bulging dorsally, guide cells 6, distinct, ventral

stereid band 3 or 4 cells thick, ventral surface

cells large and thin-walled, becoming smaller

and thickened distally, dorsal stereid band 4 or 5

cells thick, dorsal surface cells small and thick-

ened, not differentiated distally. Upper laminal

cells rounded-quadrate, becoming transversely

rectangular towards margins, 8.0-11.5 pm, ±

heterogeneous; basal cells short-rectangular to

rectangular, thin-walled, yellowish.

as distinct 8. P. crispatwn

Seta variable, 1 .0— 3.5(— 1 0.0) mm long, yel-

low to brown. Capsule short-cylindrical, 1.2-

1.5 mm, brownish. Peristome teeth cleft and

perforated, ornately papillose. Operculum ros-

trate. Calyptra mitrate, 1 mm long, long-

beaked. Spores round, 14-16 pm, brownish,

granulate. Fig. 130: 19-23.

This species is known from southern Africa,

Zimbabwe, Reunion and Macaronesia. In the

Flora area it is found on soil and rock in grass-

land and shrubland communities of the south-

western, southern and eastern Cape regions,

Free State, Lesotho, KwaZulu-Natal, Zululand,

Swaziland and the central, eastern and northern

Transvaal regions. Map 181.

Map 1 8 1 . — Ptychomitrium subcrispatum

Ptychomitriaceae

467

Vouchers: Hilliard & Burtt 14085; Magill

4143, 6318; Van Rooy 455; Schelpe 7583.

The species is most easily identified by its

bistratose leaf margins and mostly unistratose

lamina. Occasionally the lamina has bistratose

patches but these are generally small; see note

under P. crispatum (p. 475).

The name P. marginatum Dix. was used to

describe a variant of this species with long setae

and slight differences in basal leaf cell develop-

ment. Some specimens have been examined

with very long setae (up to 10 mm), and cap-

sules at the upper end of the size range of P

subcrispatum (1.5 mm); however, these speci-

mens agree in all other respects with this

species. Seta length has not proven to be a reli-

able character for the separation of species and

indeed the specimens examined show a great

deal of variation in this character.

2. Ptychomitrium exaratifolium Robin-

son in Bryologist 62: 227 (1963). Type:

Transvaal, Kruger National Park, vicinity of

Pretorius Kop, 15 Jan. 1953, R.K. Godfrey s.n.

(DUKE, holo.; NY!, US).

Plants small to medium-sized, caespitose,

dark green to yellow-green; saxicolous. Stems

5-10 mm tall, infrequently branched. Leaves

circinate-crisped dry, erect-spreading wet;

lanceolate from a broad obovate base, 3^4 mm

long; apex weakly cucullate; sheathing at base;

lamina and margins bistratose above base.

Costa subpercurrent to percurrent, ventral

superficial cells mammillose, dorsal superficial

cells smooth; in section elliptical, guide cells 6,

incrassate, ventral stereid band 1 or 2 cells

thick, cells sometimes substereid, ventral sur-

face cells mammillose, dorsal stereid band 2-4

cells thick, dorsal surface cells incrassate.

Upper laminal cells subquadrate, 5-10 pm,

thickened, ventral cells mammillose, dorsal

cells almost always shorter than ventral cells,

smooth or infrequently papillose or weakly

mammillose near margin; basal cells rectangu-

lar below, becoming quadrate at shoulders, thin-

walled.

Map 1 82. — ♦ Ptychomitrium exaratifolium

• Ptychomitrium diexaratum

Seta 2. 0-3. 5 mm long, reddish yellow.

Capsule ovoid, 1 ,0(— 1.5) mm long. Peristome

teeth lanceolate, cleft above, papillose, reddish.

Operculum rostrate, 0.7 mm long. Calyptra

mitrate, 1.8 mm long. Spores rounded to ellip-

soid, 20-30 pm, yellow-brown, granulate. Fig.

131: 1-10.

Endemic to southern Africa, P exaratifolium

is found on rock, especially dolerite, in grass-

land of the eastern Transvaal region, Swaziland

and Zululand. Map 182.

Vouchers: Magill 3578, 5480; Van Vuuren

1697, 1796.

This species is identified by its strongly mam-

millose ventral leaf cells, generally shorter and

smooth dorsal leaf cells and obovate leaf base.

Occasionally the dorsal leaf cells are almost as

high as the ventral cells and sometimes papillose

to weakly mammillose towards the leaf margins.

These specimens could be confused with P.

diexaratum , although the dorsal cells are never as

regularly mammillose (see p. 469). P. exaratifoli-

um also shows some resemblance to P. eurybasis,

especially, in leaf shape; however, its leaf cells

are flat and not mammillose. From illustrations

the Mexican species P. standleyi Crum looks

very close to P. exaratifolium. A few other

species are apparently disjunct between Mexico

and South Africa, e.g. Syntrichia chisosa

(Magill, Delgad. & L.R. Stark) R.H. Zander.

468

Ptychomitriaceae

Ptychomitriaceae

469

3. Ptychomitrium diexaratum Magill in

Magill & Schelpe in Mem. bot. Surv. S. Afr. 43:

3 (1979). Type: Lesotho, Sehlabathebe National

Park, in crevices of wet sandstone cliffs, grass-

land, 2400 m, Magill 4308 (PRE, holo.!; H, L,

MO, NY).

Plants small to medium-sized, caespitose,

dark green to yellow-green, brownish below;

saxicolous. Stems (5—) 1 0—1 5(— 20) mm tall,

occasionally branching above. Leaves crowded,

± contorted to circinate-incurved dry, erect-

spreading to widespreading wet; lanceolate to

occasionally acuminate above oblong or ellipti-

cal base, (2.0— )3.5^4.5 mm long; bistratose

above base, occasionally tristratose juxtacostal-

ly; apex acute to subcucullate; not narrowing to

insertion; margins plane or occasionally re-

flexed in base. Costa percurrent to subpercur-

rent, ventral superficial cells mammillose, dor-

sal superficial cells smooth; in section bulging

dorsally, guide cells 6-8, ventral stereid band

1(2) cell(s) thick, ventral surface cells similar to

ventral laminal cells, mammillose, dorsal

stereid band 2 or 3 cells thick, dorsal surface

cells substereids with conspicuously thickened

outer walls. Upper laminal cells quadrate to

rounded-quadrate, 6-10 pm, thickened, mam-

millose on dorsal and ventral leaf surfaces;

basal cells rectangular to long-rectangular, yel-

lowish to hyaline, thin-walled.

Seta 2-5 mm long, yellow-brown. Capsule

short, elliptical to ovoid, (1.0-)1. 5-2.0 mm

long, yellowish brown. Peristome teeth fragile,

narrowly triangular, cleft and perforated,

ornately papillose, orange-yellow. Operculum

rostrate, 1 mm long. Calyptra mitrate, 2. 0-2. 5

mm long. Spores rounded to angular, 12-18

pm, green to brown, verrucate. Fig. 131: 11-16.

Endemic to southern Africa, P. diexaratum is

found on rock in high grassland communities of

Lesotho and western KwaZulu-Natal. Map 182.

Vouchers: Hilliard & Burtt 10409; Jacot Guil-

larmod 6098, 6102; Magill 4199, 4270, 4695.

This species is very closely related to P

exaratifolium and is unlikely to be confused

with any other species. Ptychomitrium diexara-

tum is identified by the similar shape and size of

its dorsal and ventral leaf cells. Both leaf sur-

faces are strongly and evenly mammillose. The

weakly differentiated oblong to elliptical leaf

base and smaller spores are also useful charac-

ters. In addition, the leaves of this species are

occasionally tristratose juxtacostally. A few

leaves with somewhat smaller dorsal leaf cells

that are weakly mammillose have been exam-

ined, but conform in other respects to P.

diexaratum.

The two species are also separated spatially,

with P. diexaratum confined to upper elevation

grassland sites in the southern Drakensberg and

P. exaratifolium found in the lowveld of the

eastern Transvaal region, Swaziland and

Zululand.

4. Ptychomitrium depressum (C. Midi.)

Par., Ind. bryol. suppl. 1: 289 (1900); Broth, in

Natiirl. PflFam., edn 2, 11:9 (1925); Sim, Bryo.

S. Afr. 214 (1926). Type: Natal, Jammerlappen,

J. Dittrich s.n., 1898.

Brachysteleum depressum C. Mull, in Hedwigia 38: 153

(1899). Glyphomitrium depressum (C. Mull.) Broth, in

Natiirl. PflFam. 1,3: 441 (1902); Dix. in Trans. Roy. Soc.

South Africa 8: 196 (1920).

Plants medium-sized, caespitose, green;

saxicolous. Stems (5—) 1 0—25 mm high, little

branched. Leaves crowded, incurved and ±

crisped dry, erect- to widespreading wet; nar-

rowly acuminate above oblong base, 4—5 mm

Fig. 131. — Ptychomitrium exaratifolium (1-10): 1. habit (dry), x 1; 2. habit (wet), x 3; 3. stem in cross section, x 175;

4. leaf, x 35; 5. leaf in cross section, x 175; 6. basal leaf cells, x 350; 7. part of capsule mouth with peristome teeth, x 175;

8. operculum, x 35; 9. calyptra, x 35; 10. spore, x 700. P. diexaratum (11-16): 11. habit (dry), x 1; 12. habit (wet), x 3; 13.

leaf, x 35; 14. leaf in cross section, x 175; 15. upper laminal cells, x 700; 16. leaf apex, x 175. P. depressum (17-21): 17.

habit (dry), x 1; 18. habit (wet), x 3; 19. leaf, x 35; 20. leaf in cross section, x 175; 21. basal leaf margin in cross section, x

175. (1, 2, 7-10 & 12, Van Vuuren 1796 ; 3-5, Magill 5480', 6, Magill 4780; 11, Magill 4695; 13 & 15, Jacot Guillarmod

6102; 14, Magill 4308; 16, Magill 4201; 17 & 18, Pegler 1236; 19, Schelpe 7568; 20, Van Zanten 7609663; 21. Jacot

Guillarmod PRE-CH12844.)

470

Ptychomitriaceae

Ptychomitriaceae

471

long; acute; bistratose and frequently tristratose

at margins; base weakly sheathing; margins

plane above, spirally revolute in base. Costa

percurrent, smooth dorsally and ventrally; in

section bulging dorsally, guide cells 6, distinct,

ventral stereid band 2 or 3 cells thick, ventral

surface cells smaller than laminal cells, thick-

ened, dorsal stereid band 3 cells thick, dorsal

surface cells small, incrassate. Upper laminal

cells small, variable in size and shape, rounded-

quadrate to hexagonal or transversely short-rec-

tangular, 4-6 pm, incrassate; basal cells long-

rectangular to linear juxtacostally, abruptly

short-rectangular to quadrate at margins, yel-

lowish, thickened.

Seta 2-3 mm long, yellow-brown. Capsule

elliptical, 1 .5 mm long, yellow-brown, mouth

reddish. Peristome teeth narrowly triangular,

variably perforated. Operculum long-rostrate, 1

mm long. Calyptra mitrate, lobed below, 2 mm

long. Spores rounded, ( 1 2— ) 14(— 1 6) pm, yel-

lowish, granulate. Fig. 131: 17-21.

Endemic to southern Africa, P. depressum is

found in grassland, and open woodland and for-

est communities of the eastern Transvaal area,

Swaziland, Zululand, KwaZulu-Natal, the Free

State, and eastern and southwestern Cape

regions. Map 183.

Vouchers: Jacot Guillarmod 6143; Kemp

897; Magill 3522; Schelpe 7568; Taylor 446;

Van Zanten 7609663.

Closely related to P. crispatum, but the spi-

rally revolute basal leaf margins of P. depres-

sum easily separate the two species. As indicat-

ed by Dixon (1920), the revolute basal leaf mar-

gins of P. depressum are unique in the family.

5. Ptychomitrium eurybasis Dix. in S.

African J. Sci. 18: 316 (1922); Broth, in Natiirl.

PflFam., edn 2, 11:9 (1925); Sim, Bryo. S. Afr.

215 (1926). Syntypes: Zimbabwe, Macheke,

Eyles 1994 ; Matopos, Sim 8851 ; Zimbabwe, Sim

8808 (BM!, PRE!).

Plants small to medium-sized, caespitose,

dark green to blackish green; saxicolous. Stems

8-12 mm tall, irregularly and infrequently

branched. Leaves crowded, incurved and ± con-

torted above dry, erect-spreading wet; abruptly

ligulate above obovate base, 3-4 mm long;

acute; bistratose; base appressed, ± sheathing;

margins frequently multistratose. Costa percur-

rent to subpercurrent, smooth dorsally and ven-

trally; in section elliptical, guide cells 6, occa-

sionally with auxiliary cells, ventral stereid

band ± irregular, 1(2) cell(s) thick, cells fre-

quently not thickened, ventral surface cells

smaller than laminal cells, thin-walled, dorsal

stereid band 2 cells thick, dorsal surface cells

Fig. 132. — Ptychomitrium eurybasis (1-5): 1. habit (dry), x 1; 2. habit (wet), x 5; 3. leaf, x 35: 4. leaf in cross sec-

tion, x 175; 5. spore, x 700. P. cucullatifolium (6-12): 6. habit (dry), x 1; 7. habit (wet), x 5; 8 & 9. leaves, x 35: 10. leaf

in cross section, x 175; 11. upper laminal cells, x 700; 12. leaf apex (cells partly shown), x 350. P. crassinervium (13-18):

13. habit (dry), x 1; 14. habit (wet), x 5; 15. leaf, x 32; 16. leaf in cross section, x 175; 17. basal leaf cells, x 320; 18. upper

laminal cells, x 350. P. crispatum (19-24): 19. habit (dry), x 1; 20. habit (wet), x 5; 21. leaf, x 35; 22. leaf in cross sec-

tion, x 175; 23. basal leaf cells, x 350; 24. upper laminal cells, x 350. (1-5, Junod 7; 6, Van Rooy 1476; 7, Hilliard & Burn

13144; 8, Magill 5897; 9, Dealt & Killick 101 ; 10, Schelpe 2119; 11, Hilliard & Burn 10463; 12, Oliver 3231 ; 13, Smook

3585; 14, Cholnoky 617; 15 & 17, Magill 3970; 16, Van der Westhuizen & Deetlefs 27; 18, Esterhuysen 15655; 19,

Anderson PRE-CH13296; 20, Magill 5991; 21-23, Van Zanten 7609895; 24, Garside 6579.)

472

Ptychomitriaceae

not clearly defined, small, outer walls more

strongly thickened. Upper lamina I cells round-

ed-quadrate, occasionally short-rectangular or

transversely rectangular, 7-12 |am, larger juxta-

costally; basal cells rectangular, thin-walled,

yellowish, becoming shorter above.

Seta 2— 3(— 5 ) mm long, yellow-brown.

Capsule narrowly elliptical, 1.0- 1.8 mm long,

yellow- to red-brown, mouth red. Peristome

teeth fragile, perforated. Operculum rostrate, 1

mm long. Calyptra mitrate, 2. 0-2.4 mm long.

Spores rounded, variable in size, 38^40 or

50-52 pm, green, granulate. Fig. 132: 1-5.

This species is known from Malawi,

Zimbabwe and South Africa. In the Flora area a

few specimens have been collected in grassland

communities of the northern, eastern, and cen-

tral Transvaal regions, Swaziland, Free State

and the eastern Cape. Map 184.

Vouchers: Magill 3138, 6604; Retief 661;

Pegler 2151.

Ptychomitrium eurybasis is identified by its

distinctly obovate leaf base, long, ligulate blade

and flat, smooth, non-mammillate leaf cells.

The spores of this species are much larger than

those of the other southern African species. The

two fertile specimens examined from the Flora

area exhibited considerable differences in spore

size, one ranging from 38-40 pm, the other

from 50-52 pm. Otherwise the two specimens

were quite similar.

6. Ptychomitrium cucullatifolium (C.

Miill.) Jaeg. in Ber. Thatigk. St. Gallischen

Naturwiss. Ges. 1872-1873: 104 (1874); Broth,

in Natiirl. PflFam., edn 2, 11: 9 (1925); Sim,

Bryo. S. Afr. 215 (1926). Type: Cape, Witbergen,

Drege s.n., Jan. 1833 (BM!, G!, NY!).

Ptychomitrium crispatum var. brachycarpwn Homsch.

in Linnaea 15: 126 (1841). Brachysteleum cucullatifolium

C. Miill., Syn. muse, frond. 1: 769 (1849). Glyphomitrium

cucullatifolium (C. Mull.) Broth, in Natiirl. PflFam. 1,3: 441

(1902); Dix. in Trans. Roy. Soc. South Africa 8: 196 ( 1920).

Brachysteleum obtusatum C. Miill. in Hedwigia 38: 122

(1899). Ptychomitrium obtusatum (C. Miill.) Par., Ind.

bryol. suppl. 1: 289 (1900); Broth, in Natiirl. PflFam., edn

2, 11: 9 (1925); Robinson in Bryologist 62: 229 (1963).

Glyphomitrium obtusatum (C. Miill.) Broth, in Natiirl.

PflFam. 1,3: 441 (1902). Type: Transvaal, Middelburg to

Lydenburg, Wilms s.n. (G, holo.!).

Plants medium-sized, caespitose, dark green

to blackish green; terricolous or saxicolous.

Stems (5-) 10-20 mm high, occasionally

branching above. Leaves weakly appressed and

curved dry, erect-spreading wet; linear-lanceo-

late to ovate-lanceolate, (2.0— )2.5— 3.0 mm

long; bistratose above base; apex cucullate;

margins occasionally multistratose. Costa per-

current to subpercurrent, smooth dorsally and

ventrally; in section bulging dorsally, guide

cells 8, distinct, ventral stereid band 2 cells

thick, ventral surface cells distinct, slightly

smaller than laminal cells, dorsal stereid band 3

cells thick, dorsal surface cells small, outer wall

generally more strongly thickened. Upper lami-

nal cells rounded-quadrate or ± transversely

rectangular, 5— 7(— 1 0) pm, thickened; basal cells

distinct, rectangular, thin-walled, yellowish.

Seta 2-3 mm long, yellow-brown. Capsule

short-cylindrical, 1.2 mm long. Peristome teeth

narrowly triangular, occasionally irregular,

cleft, ornately papillose, yellowish. Operculum

rostrate, 0.5 mm long. Calyptra mitrate, 2 mm

long. Spores rounded, 11-14 pm, yellow-

brown, granulate. Fig. 132: 6-12.

This species is known from Zimbabwe and

southern Africa. In southern Africa over 95% of

Ptychomitriaceae

473

Map 185. — • Ptychomitrium cucullatifolium

♦ Ptychomitrium crassinervium

the specimens have been collected on soil and

rock in grassland in Lesotho, KwaZulu-Natal

and the Free State, the remaining specimens in

the eastern and central Cape regions (see note

below). Map 185.

Vouchers: Herman 606; Hilliard & Burtt

10463, 13615; Magill 4129, 4559, 5765; Smook

1117; Schelpe 2119; Van Rooy 412; Van Zinderen

Bakker 448.

Ptychomitrium cucullatifolium is very close-

ly related to P. crispatum. Numerous specimens

of P. crispatum are ± intermediate and difficult

to place in one or the other species. However,

almost all of these have been low-altitude col-

lections with a subcucullate leaf apex and have

here been placed in P. crispatum. Despite this

apparent overlap, P. cucullatifolium has been

maintained at the species level because these

specimens have leaves consistently cucullate,

leaf cells incrassate and are found predominant-

ly in the southern Drakensberg.

The type specimen and a few recent collec-

tions have come from the Cape region. Since the

holotype could not be examined (B), it is possi-

ble that the original gathering was a subcucullate

form of P. crispatum and the Drakensberg plants

represent an undescribed species. A specimen at

Geneva (G!) marked ‘Prom. Bon. Spei, Drege’,

however, conforms well with the concept of P

cucullatifolium presented here and has been

accepted as part of the original gathering.

Ptychomitrium obtusatum has not been main-

tained although preliminary examination of the

type showed an interesting difference in leaf

shape, i.e., broader and less acuminate. Speci-

mens referable to P. obtusatum appear to occur

in isolated populations within the rather restrict-

ed range of P. cucullatifolium. Leaves of several

specimens of P. cucullatifolium approach those

of P. obtusatum and since there is little or no

obvious difference in cell size, leaf length, spore

or capsule characters, P. obtusatum is here

regarded as a ‘race’ of P. cucullatifolium.

7. Ptychomitrium crassinervium (C.

Midi) Schimp. ex Par., Ind. bryol. suppl. 1: 289

(1900); Broth, in Natiirl. PflFam., edn 2, 11: 9

(1925); Sim, Bryo. S. Afr. 215 (1926). Type:

Cape, Groenekloof, Breutel s.n ., 1862 (BM!).

Brachysteleum crassinervium C. Miill. in Hedwigia 38:

121 (1899). Glyphomitrium crassinervium (C. Mull.) Broth,

in Natiirl. PflFam. 1,3: 441 (1902); Dixon in Trans. Roy.

Soc. South Africa 8: 196 (1920).

Plants small to medium-sized, caespitose,

dark green to blackish green; terricolous. Stems

5-15 mm tall, little-branched. Leaves incurved

and contorted above dry, erect-spreading wet;

linear-lanceolate to ovate-lanceolate, (2.0-)

3. 5-4. 5 mm long; acute; bistratose; base weak-

ly differentiated. Costa percurrent, wide below,

smooth dorsally and ventrally; in section nar-

rowly elliptical below, bulging dorsally above,

guide cells 8-10, distinct, frequently with auxil-

iary cells, ventral stereid band 2 or 3 cells thick,

ventral surface cells similar to ventral laminal

cells, dorsal stereid band 4-6 cells thick, dorsal

surface cells small and thickened proximally,

similar to dorsal laminal cells distally. Upper

laminal cells rounded-quadrate, 11-15 pm,

larger juxtacostally, walls thickened, frequently

somewhat irregularly so and producing ± wavy

walls; ventral row ± larger than dorsal in sec-

tion; basal cells short-rectangular, yellowish,

thin-walled or weakly thickened.

Seta (2— )4 — 5 mm long, yellow-brown. Cap-

sule short-elliptical, 1.5 mm long, brownish.

474

Ptychomitriaceae

Peristome teeth linear, cleft and perforated, red-

yellow, ornately papillose. Operculum rostrate,

1 mm long. Calyptra mitrate, 2 mm long.

Spores rounded, 30-34 pm, greenish to brown-

ish, weakly papillose. Fig. 132: 13-18.

Endemic to southern Africa, P. crassinervi-

um is found on rocky soil of road cuttings, steep

rocky slopes and clay banks in the western and

southwestern Cape regions, Zimbabwe and

Zambia. Map 185.

Vouchers: Cholnoky 617; Esterhuysen 15655;

Magill 3875, 4001; Oliver 7253, 7274.

Ptychomitrium crassinervium can be con-

fused with P. crispatum, but the following set of

characters should separate the two species. The

leaf cells of P. crassinervium are longer than 10

pm, the basal leaf cells are short-rectangular and

little-thickened, and the costa is broader below

and elliptical in transverse section, with surface

cells large, thin-walled and distinct. In contrast,

P crispatum has leaf cells shorter than 10 pm,

basal leaf cells long-rectangular, and a narrow

costa which is round in transverse section with

less clearly differentiated surface cells.

8. Ptychomitrium crispatum (Hedw.).

Jaeg. in Ber. Thatigk. St. Gallischen Naturwiss.

Ges. 1872-1873: 103 (1874); Broth, in Natiirl.

PflFam., edn 2, 11:9 (1925); Sim, Bryo. S. Afr.

214 ( 1 926). Type: Caput bonae spei. Thunberg s.n.

Encalypta crispata Hedw., Sp. muse, frond. 61 (1801).

Glyphomitrium crispatum (Hedw.) Brid. Muscol. recent,

suppl. 4: 30 (1818). Orthotrichum crispatum (Hedw.) Hook. &

Grev. in Edinburgh J. Sci. 1: 115 (1824). Brachypodium

crispatum (Hedw.) Brid., Bryol. univ. 1: 147 (1826). Notarisia

capensis Hampe in Linnaea 11: 379 (1837), nom. illeg.

Plants medium-sized, caespitose, dark green,

brownish below; saxicolous. Stems 1 0—20 mm

long, occasionally branching above. Leaves

incurved dry, widespreading wet; linear-lanceo-

late above short-oblong to elliptical base,

(2.5-)3.0-5.0 mm long; bistratose; apex acute

to subcucullate; lamina occasionally reflexed

above a weakly appressed base. Costa percur-

rent, narrow; ventral superficial cells smooth,

similar to ventral laminal cells; dorsal super-

ficial cells smooth, rectangular, thickened; in

section bulging dorsally (half-round through-

out), guide cells 6, small, ventral stereid band 2

or 3 cells thick, ventral surface cells similar to

laminal cells, dorsal stereid band (3— )5 or 6

cells thick, dorsal surface cells generally small,

thickened, not clearly differentiated but becom-

ing more pronounced distally. Upper laminal

cells ± rounded-quadrate, somewhat variable,

5-8 pm, thickened; basal cells rectangular to

rhomboidal, yellowish, thin-walled.

Seta (4 — )5— 8 mm long, yellowish. Capsule

short-cylindrical, 1. 5-2.0 mm long, yellow-

brown, mouth red. Peristome teeth narrowly trian-

gular, perforated, reddish yellow, ornately papil-

lose. Operculum rostrate, 1 mm long. Calyptra

mitrate, 2 mm long. Spores rounded, 12-16 pm,

brownish, bluntly papillose. Fig. 132: 19-24.

The most widespread southern African

species, P. crispatum has also been reported

from eastern Africa, Madagascar, southern

South America and the Juan Fernandez islands.

In the Flora area P crispatum is found on rock,

in rock crevices or shallow soil over rock in

grassland or shrubland communities of the west-

ern, southwestern, central, southern and eastern

Cape regions, the Free State, KwaZulu-Natal,

Zululand, Swaziland and the central, eastern and

northern Transvaal areas. Isolated specimens

have also been collected in the northern Cape

region and southern Lesotho. Map 1 86.

Map 186. — Ptychomitrium crispatum

Ptychomitriaceae

475

Vouchers: Bayliss 8200; Ellis 3108; Garside

6712; Hardy 5421a; Magill 3657, 5991, 6111;

Schelpe 7653; Van Zanten 760895.

Variations in this rather widespread and

somewhat plastic species reflect its relationship

to P. cucullatifolium (p. 472), P. depression (p.

469), and P. crassinervium (p. 473).

Another form of variation seen in specimens

of P. crispatum involves irregularity in the bi-

stratosity of the upper leaf lamina. Occasionally

unistratose patches are present, but these are

rarely as pronounced or extensive as in the

leaves of P subcrispatum. Specimens with mul-

tistratose margins are also occasionally encoun-

tered. These specimens can be confused with P.

subcrispatum , especially if the leaves are not

sectioned.

Insufficiently known species

Brachysteleum convolutifolium Shaw in

Cape Monthly Mag. 17: 379 (1878). Syntypes:

Cape, Graaff-Reinet, McLea s.n.; Bolus s.n.;

Shaw s.n. Type material has not been seen and

was apparently destroyed after Shaw’s death.

We would agree with Sim (1926) that the

description is insufficient to place this species

properly.

476

ORTHOTRICHACEAE

Plants small to large, forming mats or caespitose, variously green to brown; saxicolous, corti-

colous or terricolous. Primary stem prostrate or erect, variously branched; secondary stem hori-

zontal or erect, frequently branching by subperichaetial innovations; in section subpentagonal to

round, central strand absent or weak. Leaves appressed, erect, or variously twisted when dry, erect-

spreading to squarrose when wet; variously ovate, lanceolate or oblong, rarely fragile above, occa-

sionally rugulose to rugose; unistratose or bistratose or irregularly multistratose; apex variable;

base rarely differentiated; margin plane, recurved or rarely incurved, entire, crenulate, denticulate

or serrate above, rarely tuberculate below. Costa single, occasionally papillose or toothed dorsally.

Upper laminal cells generally small, rounded, incrassate, flat to bulging, smooth, mammillose or

papillose; basal cells generally rectangular to quadrate, smooth, papillose or tuberculate; alar cells

not differentiated. Gemmae present or absent, filiform or clavate to subround.

Autoicous, dioicous or synoicous, occasionally pseudautoicous. Perigonia terminal, lateral or

axillary. Perichaetia terminal on stems or branches. Seta short or long. Capsule immersed, emer-

gent or exserted, frequently ribbed. Peristome absent, single or double, parts occasionally fused or

reduced; exostome teeth 8, in 8 pairs or 16, outer plate thick, inner plate thin; endostome segments

8 or 16, alternating with teeth, not or weakly keeled, basal membrane low or absent, cilia absent.

Operculum mostly conic-rostrate. Calyptra cucullate, mitrate or campanulate, frequently large,

occasionally plicate, entire, lobed or lacerate below, naked or hairy. Spores granulate or papillose,

isosporous or anisosporous.

A large and diverse family with ± 550 species in 14 genera of which 9 genera and 31 species

occur in southern Africa. Members of the Orthotrichaceae are predominantly xerophytic and adapt-

ed to habitats on trees and rocks.

The family is characterized by leaves with small, incrassate and mostly papillose upper laminal

cells, larger basal cells, no differentiated alar cells, a large calyptra and the generally diplolepidous

peristome. The placement of genera with haplolepidous peristomes in the Orthotrichaceae is uncer-

tain (Shaw 1986). The orthotrichaceous peristome is characterized by an exostome with the outer

plate thicker than the inner plate and by the endostome segments generally alternating with the

exostome teeth, segments not or weakly keeled, basal membrane and cilia absent, and frequently

with a reduced number of inner peristomial layer divisions. Reduction of the peristome is fre-

quently found in the family.

The Orthotrichaceae can be divided into four subfamilies of which three (Zygodontoideae,

Orthotrichoideae and Macromitrioideae) occur in southern Africa.

Key to subfamilies and genera of the Orthotrichaceae

1 Primary stem erect, simple or sparsely branched; sporophytes produced on primary stem

and branches; gemmae occasionally present; calyptra cucullate or mitrate:

2 Calyptra cucullate, naked or sparsely hairy (subfamily Zygodontoideae, p. 477):

3 Cells smooth or papillose; costa in cross section with median guide cells absent, ven-

tral cells frequently larger; gemmae frequently present; capsule long-exserted; peri-

stome absent, single or double 1 . Zygodon

3 Cells papillose to papillose-striolate; costa in cross section with median guide cells present;

gemmae absent; capsule immersed or shortly exserted; peristome absent . . 2. Amphidium

2 Calyptra mitrate, hairy (subfamily Orthotrichoideae, p. 490):

4 Leaf base distinct, bordered 5. Ulota

4 Leaf base indistinct, not bordered:

Orthotrichaceae

477

5 Leaves crispate dry; perichaetial leaves hyaline below 4. Stoneobryum

5 Leaves erect to appressed or occasionally flexuose dry; perichaetial leaves chloro-

phyllous 3. Orthotrichum

1 Primary stem prostrate, with numerous horizontal or erect branches; sporophytes produced

on secondary stem and branches; gemmae absent; calyptra mitrate (subfamily Macro-

mitrioideae, p. 506):

6 Branch leaves secund, base decurrent, cells tuberculate 9. Cardotiella

6 Branch leaves appressed, flexuose, twisted or inrolled, base not decurrent:

7 Branches slender, widely spaced; branch leaves appressed dry; basal cells scarcely dif-

ferentiated, short 6. Macrocoma

7 Branches with a bushy appearance, crowded; branch leaves flexuose to variously twist-

ed or inrolled dry; basal cells differentiated, elongate:

8 Leaves flexuose, twisted to contorted or inrolled dry; calyptra plicate, naked or sparse-

ly hairy, laciniate or lacerate below 7. Macromitrium

8 Leaves twisted to spirally twisted around stem dry; calyptra smooth, naked, lobed

below 8. Schlotheimia

Subfamily ZYGODONTOIDEAE

Plants small to large; saxicolous, terricolous or corticolous. Primary stems erect, simple or

sparsely branched. Leaves appressed, twisted or crisped when dry; narrowly lanceolate or oblong

to elliptical; base scarcely differentiated. Upper laminal cells rounded or rounded-hexagonal,

incrassate, smooth or papillose or striolate-papillose; basal cells mostly rectangular. Gemmae fre-

quently present on stems or rhizoids.

Perichaetia terminal on primary stem and branches. Dwarf male plants absent. Capsule

immersed, short-exserted or long-exserted; ribbed. Stomata phaneropore. Peristome absent or sin-

gle or double. Calyptra cucullate, naked or sparsely hairy. Isosporous.

1. ZYGODON

Zygodon Hook. & Tayl., Muscol. brit. 70 (1818); Broth, in Natiirl. PflFam., edn 2, 11: 11

(1925); Sim, Bryo. S. Afr. 268 (1926); Malta, Die Gattung Zygodon 1-184 (1926); Sainsb., N. Zeal,

mosses 198 (1955); Scott & Stone, Moss. S. Austr. 245 (1976); Gangulee, Moss. E. India 5: 1153

(1976); Smith, Moss FI. Brit. Irel. 470 (1978); Crum & Anderson, Moss. E.N. Amer. 2: 679 (1981).

Type species: Z. conoideus (Dicks.) Elook. & Tayl.

Plants small to large, loosely caespitose to caespitose; corticolous or saxicolous to terricolous.

Stems erect, in section pentagonal to round, epidermis not differentiated. Leaves ± crowded, erect

to appressed or twisted when dry, erect-spreading to squarrose and frequently reflexed to recurved

when wet, occasionally in 5 indistinct rows, frequently rugulose; variously lanceolate or oblong to

elliptical; apex acute, acuminate, rounded-acute to obtuse or occasionally mucronate or apiculate,

frequently toothed; base scarcely differentiated, decurrent; margins plane or undulate, entire or

crenulate or serrate. Costa ending below apex, percurrent, subpercurrent or merging with elongat-

ed apical cells to become mucronate or apiculate. Upper laminal cells small, irregularly rounded-

hexagonal, incrassate, smooth or papillose; basal cells quadrate to rectangular or rhomboidal,

smooth or papillose. Gemmae frequently produced on stem or rhizoids, uniseriate or multicellular,

generally reddish brown.

478

Orthotrichaceae

Dioicous, autoicous or synoicous. Inflorescence terminal, overgrown by innovations,

perichaetial leaves scarcely differentiated, perigonial leaves and leaves of synoicous inflorescence

differentiated. Seta long, twisted anticlockwise above. Capsule erect, elliptical, oblong-cylindrical

or narrowly pyriform, 8-ribbed or plicate; mouth occasionally 3- or 4-sulcate, infrequently strong-

ly thickened; neck differentiated; stomata on neck, phaneropore. Peristome absent, single or dou-

ble. Operculum short- to long-conic-rostrate, straight to oblique. Calyptra cucullate, naked or

sparsely hairy. Spores round, ± granulate, brownish.

A genus of ± 90 species found mostly in tropical to temperate regions of the southern hemi-

sphere. Tropical South America is the major centre of described species. In southern Africa most

species are found on stems and branches of trees or on soil and rock in montane forests and wood-

lands of northern and eastern Transvaal, KwaZulu-Natal, and the southern and eastern Cape

regions. Two species occur in the Fynbos Biome of the southwestern Cape.

Zygodon is recognized by the tall plants with twisted to secund or erect to appressed leaves

when dry, small and frequently papillose leaf cells, long-exserted, ribbed or plicate capsule, cucul-

late calyptra, and numerous gemmae produced on rhizoids and in leaf axils.

1 Leaf cells smooth, margin entire; autoicous 1. Z. dixonii

1 Leaf cells papillose or smooth, margin serrate; dioicous or synoicous;

2 Leaf margin serrate; upper costa frequently with multicellular teeth dorsally; peristome

double; southwestern Cape region 2. Z. runcinatus

2 Leaf margin entire, crenulate or rarely with a single tooth above; costa smooth; peristome

single or absent; Transvaal regions, KwaZulu-Natal, eastern, southern and south-

western Cape regions:

3 Gemmae with transverse and longitudinal septa 6. Z. erosus

3 Gemmae with transverse septa only:

4 Leaf apex rounded-acute to rounded-obtuse, costa merging with apical cells to

become inucronate; capsule mouth sulcate when dry; peristome absent; calyptra

hairy , 5. Z trichomitrius

4 Leaf apex acute to acuminate, costa ending below apex; capsule mouth thickened,

erect when dry; peristome single, endostome segments fragile; calyptra naked:

5 Plants dioicous; Transvaal regions, KwaZulu-Natal and eastern Cape ... 3. Z. intermedius

5 Plants synoicous; southwestern Cape region 4. Z. leptobolax

1 . Zygodon dixonii Sim, Bryo. S. Afr. 271

(1926). Type: Natal, below Cathkin Peak, Sim

10004 (PRE, holo. !).

Plants small, caespitose, yellowish green

above, reddish brown below; saxicolous. Stems

up to 8 mm tall, branching by subperichaetial

innovations, tomentose below; rhizoids smooth

to coarsely papillose, reddish brown or pale

brown; in section round, cortical cells in 10

rows, inner cells thin-walled, gradually smaller

and incrassate towards outer cells, outer cells

bulging. Leaves ± equal in size to larger above,

twisted when dry, erect-spreading when wet;

narrowly oblong, 1 .0-1 .6 mm long, ventral sur-

face keeled; apex inucronate or apiculate; base

scarcely differentiated, margins decurrent; mar-

gins plane or occasionally recurved below,

entire, unistratose. Costa subpefcurrent to per-

current; ventral and dorsal superficial cells rec-

tangular to linear, incrassate; in section round,

bulging dorsally, lamina ventrally inserted, not

differentiated or 2 guide cells exposed ventral-

ly, ventral stereid band absent, dorsal stereids in

2 or 3 rows, dorsal surface cells not differenti-

ated, rough. Upper laminal cells rounded-

hexagonal, flat, smooth, ( 8—) 1 0— 1 2(— 17) pm;

basal cells large, reaching higher along costa,

quadrate to rectangular, thin-walled to incras-

sate, smooth. Gemmae absent.

Orthotrichaceae

479

Autoicous. Perigonia subperichaetial on short

branches, gemmate, leaves ovate. Perichaetia

terminal, leaves not differentiated. Seta 3.0-4. 5

mm long, brownish. Capsule exserted, erect,

elliptical or narrowly pyriform, brownish, 8-

ribbed, urn 0. 7-1.0 mm long, neck 0.5 mm long;

exothecial cells rectangular to quadrate, thin-

walled, longitudinal walls of rib cells incrassate,

smaller and rounded at mouth, incrassate; annu-

lus deciduous. Peristome double; exostome teeth

in 8 pairs, recurved when dry, narrowly lanceo-

late, fused below, perforated above, 230-310 pm

long, densely papillose to striolate-papillose, yel-

lowish brown; endostome segments alternating

with teeth pairs, shorter than teeth, erect when

dry, linear-lanceolate, papillose-striolate, hya-

line, basal membrane low. Operculum 0.5 mm

long, oblique. Calyptra 1.3 mm long, naked,

cells prorate above. Spores 11-15 pm. Fig. 133:

1-13.

Known only from the type locality in the

Drakensberg of KwaZulu-Natal. Map 187.

Voucher: type only.

The small plants and smooth leaf cells sepa-

rate Z. dixonii from other Zygodon species in

southern Africa.

2. Zygodon runcinatus C. Mull, in Hed-

wigia 38: 1 14 (1899); Broth, in Natiirl. PflFam.,

edn 2, 11: 15 (1925); Sim, Bryo. S. Afr. 268

(1926); Malta, Die Gattung Zygodon 109 (1926).

Isosyntypes: Cape, Table Mountain, Rehmann

150 ; Devil’s Peak, Rehmann 150b (both PRE!).

Plants medium-sized, caespitose, yellowish

green or olivaceous above, brown to dark

brown below; saxicolous to terricolous. Stems

to 40 mm tall, branching dichotomously or by

subperichaetial or subperigonial innovations,

sparsely tomentose below; rhizoids smooth to

coarsely papillose, red-brown or red; in section

pentagonal to subround, inner cortex 7-14 cells

across, thin-walled to incrassate, smaller

towards outside, outer cortical cells smaller, in

1-4 rows, substereids or stereids. Leaves ±

crowded, ± equal in size, erect to appressed

when dry, widespreading to squarrose-recurved

Map 187. — ♦ Zygodon dixonii

• Zygodon runcinatus

from ± sheathing bases when wet, in 5 indistinct

rows, spiralled around stem; ovate-lanceolate to

lanceolate or oblong-lanceolate, 1 .4— 3.0(— 3.6)

mm long; ventral surface keeled; apex acute to

acuminate, toothed; base ovate to oblong, ±

sheathing, margins decurrent; margins plane,

serrate to strongly serrate in upper half to two-

thirds, teeth frequently multicellular, unistratose

with bistratose patches, thickened. Costa end-

ing below apex to subpercurrent, ventral and

dorsal superficial cells rounded to elongate,

incrassate; in section round, bulging dorsally,

lamina ventrally inserted, not differentiated or

2^4 ventral guide cells larger, ventral stereid

band absent, dorsal cells in 1-3 rows, incrassate

to stereids, with dorsal teeth distally, teeth mul-

ticellular. Upper laminal cells small, irregularly

rounded-hexagonal, flat, (7.0-)8.5-12.5(-14.0)

pm, smooth to papillose, 1^1 papillae scattered

over lumen, small, low; basal cells rhomboidal

to rectangular, thin-walled to incrassate, smooth

or walls papillose. Gemmae axillary, sub-

clavate, uniseriate, with transverse septa.

Dioicous. Perigonia terminal, leaves broadly

ovate to orbicular. Perichaetia terminal; inner

leaves oblong-acuminate or ovate-acuminate to

lanceolate, 1.2-2.5(-2.8) mm long; apex acumi-

nate; margins plane, entire to crenulate above;

costa frequently weak, ending below apex; lami-

480

Orthotrichaceae

Orthotrichaceae

481

nal cells rounded to vermiculate above, vermicu-

late below, occasionally papillose. Seta 3-5 mm

long, yellowish or brownish. Capsule exserted,

erect, elliptical or narrowly pyriform, 8-ribbed,

urn 1.0-1. 8 mm long, neck 0.3-0.6 mm long;

exothecial cells rectangular to quadrate, irregu-

larly incrassate, smaller at mouth, rounded-

quadrate, cells of ribs differentiated; annulus

deciduous. Peristome double, recurved when

dry; exostome teeth in 8 pairs, narrowly triangu-

lar or lanceolate, fused below, perforated above,

180-300 pm long, densely papillose above, stri-

olate-papillose below, yellowish or yellowish

brown; endostome segments alternating with

teeth pairs, linear, minutely papillose or smooth,

pale yellow or hyaline, basal membrane low or

absent. Operculum 0.5 mm long, oblique. Calyp-

tra 2.5-3. 0 mm long, smooth, naked. Spores

15.0-22.5 pm. Fig. 133: 14-26.

The species is known from the Fynbos

Biome of the southwestern Cape region and

from Tanzania. Map 187.

Vouchers: Barnard PRE-CH3405, PRE-

CH6032; Esterhuysen 16594; Garside 6619;

Magill & Schelpe 3967, 3998; Schelpe 4969;

Thorne PRE-CH6416.

Zygodon runcinatus is easily recognized by

the strongly serrate leaf margins and the multi-

cellular teeth on the dorsal side of the upper

costa.

3. Zygodon intermedius B.S.G. , Bryol.

eur. 3: 41 (1838); Broth, in Naturl. PflFam., edn

2, 11: 15 (1925); Malta, Die Gattung Zygodon

75 (1926); Lewinsky in Lindbergia 15: 131

(1990). Type: New Zealand, Dusky Sound,

Menzies 66 (BM, holo.).

Zygodon transvaalensis Rehm. ex Sim, Bryo. S. Afr.

271 ( 1926). Type: Transvaal, in mont. Lechlaba ad arborum

truncos, Rehmatm 500 (PRE, iso.!).

Plants small to medium-sized, loosely caes-

pitose to caespitose, yellowish green to brown;

terricolous or corticolous. Stems up to 25 mm

tall, branching by innovations, tomentose

below; rhizoids smooth to papillose or granu-

lose, reddish brown or reddish; in section sub-

pentagonal or round, central strand absent, inner

cortex 5-10 cells across, thin-walled to incras-

sate, outer cortical cells smaller, in 1-3 rows,

incrassate or with substereids to stereids, outer

cells frequently rough. Leaves ± crowded, ±

equal in size or larger above, twisted or secund

and ± appressed when dry, erect-spreading to

squarrose and recurved when wet; narrowly

oblong-lanceolate to lanceolate or occasionally

oblong to elliptical, 0.6-2. 5 mm long, ventral

surface keeled to broadly keeled; apex acute to

acuminate, occasionally apiculate, with a single

clear tooth; base scarcely differentiated, decur-

rent; margins ± undulate above, rarely recurved

below, entire or rarely with a single tooth above,

entire to crenulate below, unistratose. Costa

ending below apex; ventral and dorsal superfi-

cial cells long-rectangular to linear; in section

subround to round, bulging dorsally, guide cells

exposed ventrally, dorsal stereid band 1-3 cells

thick, dorsal surface cells not differentiated, ±

rough. Upper laminal cells irregularly rounded-

hexagonal, flat, 7.5-15.0 pm, papillose to

densely papillose, papillae 4-7, scattered over

lumen, low and blunt; basal cells short-rectan-

gular to quadrate, thin-walled to incrassate or

transverse walls incrassate, smooth or with a

single papilla over lumen or walls papillose.

Gemmae produced on stem, filiform or clavate,

uniseriate, with transverse septa, (2)3-5 cells

long, reddish brown.

Dioicous. Perigonia and perichaetia termi-

nal, quickly overgrown by innovations; perigo-

nial leaves shortly ovate-apiculate or ovate -

Fig. 133. — Zygodon dixonii (1-13): 1. habit, (dry), x 5; 2. habit (wet), x 7; 3. stem in cross section, x 175; 4. leaf, x

35; 5. leaf in cross section, x 175; 6. basal leaf cells (right side), x 175; 7. upper laminal cells, x 350; 8. leaf apex, x 175;

9. stoma of capsule wall, x 350; 10. part of capsule mouth with peristome, x 175; 11. operculum, x<70; 12. calyptra, x 35;

13. spores, x 700. Z. runcinatus (14—26): 14. habit (dry), x 3; 15. habit (wet), x 4; 16. stem in cross section, x 175; 17.

leaf, x 35; 18. leaf in cross section, x 175; 19. basal leaf cells (right side), x 175; 20. laminal cells at left margin, x 700; 21.

cells at leaf apex, x 175; 22. gemma, x 350; 23. perichaetial leaf, x 35; 24. part of capsule mouth with peristome, x 175;

25. operculum, x 70; 26. spore, x 700. (14, 20, 23, 24 & 26, Barnard 50331; 15, Esterhuysen 16594\ 16 & 18, Barnard

44154', 17 & 21, Thorne 6416; 19, Garside 6619; 22, Rehmann 150; 25, Sim 9398.)

482

Orthotrichaceae

Fig. 134. — Zygodon trichomitrius (1-11): 1. habit (dry), x 2;

2. habit (wet), x 3; 3. leaf, x 35; 4. leaf cross section, x 175; 5. basal

leaf cells, x 350; 6. upper laminal cells, x 350; 7. cells at leaf apex, x

350; 8. gemma, x 350; 9. part of capsule mouth, x 175; 10. calyptra

x 18; 11. spore, x 700. Z. erosus (12-23): 12. habit (dry), x 1; 13.

habit (wet), x 2; 14. leaf, x 35; 15. leaf cross section, x 175; 16. basal

leaf cells, x 350; 17. upper laminal cells, x 350; 18. leaf apex, x 350;

19. gemma x 175; 20. filamentous gemmae at leaf base, x 175; 21.

part of capsule mouth, x 175; 22. calyptra x 18; 23. spore, x 700. Z.

intermedins (24-34): 24. habit (dry ), x 1 ; 25. habit (wet), x 3; 26 &

27. leaves, x 35; 28. leaf cross section, x 175; 29. basal leaf cells, x

350; 30. upper laminal cells, x 700; 31. leaf apex, x 350; 32. gemma

x 350; 33. penchaetial leaf, x 35; 34. part of capsule mouth, x 175;

35. calyptra x 35. (12, 18, 21-23, Sim 10045: 13, Sim 10070: 14,

19 & 20, Stirton 8991: 15, Magill 5508: 16, Magill5527: 17, Van

Roox 1629: 24, 29 & 32, Van Rooy 1508: 25 & 35, Magill 5667: 26, Sim 10030: 27, Sim 10101: 28, Jacot Guillarmod PRE-CH12583: 30,

Sim PRE-CH7579: 31 & 34. Magill 5662: 33, Magill 5684.)

Orthotrichaceae

483

acuminate; perichaetial leaves scarcely differ-

entiated. Seta 5-10 mm long, brownish.

Capsule oblong-cylindrical or narrowly pyri-

form, 1.5-2. 2 mm long, brownish, 8-ribbed,

neck differentiated; exothecial cells quadrate to

short-rectangular, incrassate, smaller at mouth;

annulus apparently absent. Peristome fragile,

exostome teeth rudimentary; endostome seg-

ments distant, ± linear, irregular in outline,

85-105 pm long, yellowish, essentially smooth,

basal membrane absent. Operculum 0.8 mm

long, ± oblique. Calyptra 2.0-2. 5 mm long,

smooth. Spores 1 7—25 pm. Fig. 134: 24-34.

Zygodon intermedins frequently grows on

bark of trees but has also been found on soil.

This pantropical species has been reported from

Australasia, tropical Asia, South America, the

Juan Fernandez islands, sub-Saharan Africa and

the Mascarenes. In the Flora area it is known

from montane forests in the northern and east-

ern Transvaal regions, KwaZulu-Natal, the east-

ern Cape region, and from cliffs at Sani Pass,

KwaZulu-Natal. Map 188.

Vouchers: Crosby 7927; Magill 4471, 5662,

5684, 7323; Sim 10101; Van Rooy 1505.

The dioicous sexual condition and geograph-

ical distribution separate Z. intermedius from

the very similar Z. leptobolax (see note below).

4. Zygodon leptobolax C. Mull, in Hed-

wigia 38: 113 (1899); Broth, in Natiirl. PflFam.,

edn 2, 11: 14 (1925); Sim, Bryo. S. Afr. 270

(1926); Malta, Die Gattung Zygodon 130

(1926). Type: Cape, Cape Town, Rondebosch,

Rehmann 499 (PRE, iso.!).

Plants small to medium-sized, loosely caes-

pitose, yellowish brown to brown; corticolous.

Stems up to 11 mm tall, branching by innova-

tions, tomentose below; rhizoids smooth to papil-

lose, reddish brown; in section subpentagonal to

round, central strand absent, inner cortex 3-6

cells across, thin-walled or incrassate towards

outside, outer cortical cells smaller, in 1-3 rows,

consisting of substereids, outer cells occasional-

ly rough. Leaves ± crowded, slightly larger

above, twisted to secund and ± appressed when

dry, erect-spreading when wet; oblong-lanceo-

late to lanceolate, 0.7-2.0 mm long; ventral sur-

face broadly keeled; apex acute to acuminate,

with a single clear tooth above; base scarcely dif-

ferentiated, decurrent; margins ± undulate above,

plane or rarely recurved below, entire or rarely

with a single blunt tooth above, entire to crenu-

late below, unistratose. Costa ending below

apex; ventral and dorsal superficial cells long-

rectangular to linear; in section subround,

bulging dorsally, guide cells exposed ventrally,

dorsal stereid band 1-3 cells thick, dorsal surface

cells ± rough. Upper laminal cells irregularly

rounded-hexagonal, 1 1— 17(— 20) pm, papillose to

densely papillose, frequently obscured by papil-

lae, 4-7 papillae scattered over lumen, low and

blunt; basal cells short-rectangular to quadrate,

incrassate, smooth or with single papilla over

lumen or walls papillose. Gemmae produced on

stem, subclavate, uniseriate with transverse

septa, 3 or 4 cells long, reddish brown.

Synoicous. Inflorescence terminal; leaves

broadly ovate-acuminate to oblong-acuminate,

0.5-1. 3 mm long. Seta 4.5-10.0 mm long, red-

dish brown. Capsule oblong-cylindrical or nar-

rowly pyriform, 1-2 mm long, 8-ribbed, neck

differentiated; exothecial cells quadrate to rec-

tangular, longitudinal walls incrassate along

ribs, smaller and incrassate at mouth; annulus

apparently absent. Peristome fragile, exostome

absent; endostome segments short, distant, yel-

lowish, basal membrane absent. Operculum

0. 6-0.8 mm long. Calyptra 1. 8-2.0 mm long,

smooth. Spores 17-28 pm. Fig. 135.

This is another species of Zygodon restricted

to the Fynbos Biome of the southwestern Cape

region where it was collected from bark on the

slopes of Table Mountain. Map 188.

Vouchers: Rehmann 148; Sim 9150.

In the absence of the synoicous inflorescence

Z. leptobolax is practically indistinguishable

from similar specimens of Z. intermedius. The

two species are characterized by the falcate

stems, twisted to secund and frequently ap-

pressed leaves, acute to acuminate leaf apex

with a single clear tooth above, wavy leaf mar-

gins, costa ending below the apex, uniseriate

484

Orthotrichaceae

Fig 135. — Zygodon leptobolax: 1. habit (dry), x 3; 2.

habit (wet), x 5; 3. leaf, x 35; 4. leaf apex, x 350; 5. gemma,

x 350; 6. part of synoicous inflorescence, x 70; 7. leaf of

synoicous inflorescence, x 35. (1, 2, 4 & 7, Rehmann 499 ;

3, Rehmann 148\ 5 & 6, Sim 9150.)

gemmae, thickened capsule mouth, and the sin-

gle peristome with endostome segments distant

and reduced.

5. Zygodon trichomitrius Hook. & Wilson

in London J. Bot. 5: 143 (1846); Broth, in Natiirl.

PflFam., edn 2, 11; 14 (1925); Sim, Bryo. S. Afr.

269 (1926); Malta, Die Gattung Zygodon 62

(1926). Type: Cape, on trees in the forest of

Grootvadersbosch, Swellendam district, Zeyher

s.n. (BM-Hook., lecto.!, selected here).

Plants medium-sized to large, loosely caespi-

tose to caespitose, yellowish green or light

green to green above, olivaceous or yellowish

brown to brown below; corticolous. Stems fre-

quently falcate above, up to 40 mm tall, branch-

ing by subperichaetial or subperigonial innova-

tions, sparsely tomentose to tomentose below;

rhizoids smooth to coarsely papillose, reddish

Map 188. — • Zygodon intermedius

♦ Zygodon leptobolax

brown to red; in section subpentagonal or sub-

round, inner cortex 7-11 cells across, thin-

walled, incrassate towards outside, outer corti-

cal cells smaller, in 2-A rows, consisting of sub-

stereids or stereids. Leaves ± crowded, ± equal

in size to larger above, twisted or contorted

when dry, widespreading to squarrose and

recurved when wet, infrequently rugulose, in 5

indistinct rows, spiralled around stem; narrowly

oblong-lanceolate or lanceolate, ( 1 .7— )2. 0-3.4

(-3.7) mm long; ventral surface keeled; apex

acute to rounded-acute to rounded-obtuse,

mucronate, cells of mucro fusiform, smooth,

incrassate, with single tooth; base scarcely dif-

ferentiated, ± sheathing, margins decurrent;

margins plane, generally crenulate, unistratose.

Costa merging with elongated cells of mucro,

ventral and dorsal superficial cells long-rectan-

gular to linear; in section subround, bulging

dorsally, lamina ventrally inserted, 2-\ guide

cells exposed ventrally, ventral stereid band

absent, dorsal stereid band 1-3 cells thick, dor-

sal surface cells not differentiated, rough.

Upper laminal cells irregularly rounded-hexag-

onal, flat, 7.5-15 pm, papillae low, blunt, scat-

tered; basal cells rhomboidal to rectangular,

thin-walled to incrassate, smooth or with single

papilla over lumen or papillose along walls.

Gemmae numerous, on stem, subclavate, unise-

riate with transverse septa, 3-5 cells long, red-

dish brown.

Orthotrichaceae

485

Dioicous. Perigonia terminal, quickly over-

grown by subperigonial innovation; leaves

broadly ovate, ovate-apiculate or shortly ovate-

acuminate. Perichaetia terminal; leaves scarce-

ly differentiated, inner leaves occasionally lin-

ear from a lanceolate base, shorter, 1.2-2. 8 mm

long, apex acuminate. Seta 8.5-11.0 mm long,

brownish. Capsule narrowly pyriform, brown-

ish, plicate, mouth 3- or 4-sulcate when dry,

orange to brown, urn 1.4— 2.0 mm long, neck

0.5-0.8 mm long; exothecial cells irregularly

quadrate to rectangular, smaller at mouth, lon-

gitudinal walls incrassate above; annulus appar-

ently absent. Peristome absent. Operculum

1.0-1. 5 mm long, straight to oblique. Calyptra

2.6-3. 0 mm long, sparsely hairy, hairs uniseri-

ate, ascending, smooth to crenulate, cells occa-

sionally prorate distally above. Spores 17-20

pm. Fig. 134: 1-11.

The typical variety of Z. trichomitrius is

endemic to southern Africa. The var. mild-

braedii (Broth.) Malta is found in central and

eastern Africa (Malta 1926). In the Flora area

the species is found on trunks, branches and

exposed roots of trees and rarely on rock in

forests of the northern and eastern Transvaal

regions, KwaZulu-Natal and the eastern,

southern and southwestern Cape regions. Map

189.

Vouchers: Crosby & Crosby 7634; Magill

3749, 5956, 6217; Schelpe 7869a; Van Rooy

2285, 2296.

The recurved leaves, relatively broad above

with rounded-acute to rounded-obtuse apices and

mucronate costae, the uniseriate gemmae, and

gymnostomous capsule with sulcate mouth place

plants in this species. For differences between

Z. trichomitrius and Z. erosus , see p. 486.

6. Zygodon erosus Mitt, in J. Linn. Soc.,

Bot. 22: 305 (1886); Broth, in Natiirl. PflFam.,

edn 2, 11: 15 (1925); Malta, Die Gattung Zygo-

don 63 (1926). Type: Tanzania, Kilimanjaro,

Hannington s.n. (NY, holo.!).

Zygodon africanus Sim, Bryo. S. Afr. 270 ( 1926). Type:

Transvaal, MacMac, MacLea sub Rehmann 497 (PRE! ).

Plants medium-sized to large, caespitose,

yellowish green to green above, yellowish

brown to brown below; corticolous. Stems up

to 50 mm tall, branching dichotomously or by

subperichaetial or subperigonial innovations,

tomentose below; rhizoids smooth to coarsely

papillose, reddish brown to red; in section

weakly pentagonal to subround, inner cortex

6-12 cells across, thin- walled, outer cortical

cells smaller, in 2-4 rows, incrassate or con-

sisting of stereids. Leaves crowded, ± equal in

size to larger above, twisted or contorted when

dry, erect-spreading to squarrose and reflexed

to recurved when wet, frequently rugulose,

frequently in 5 indistinct rows, spiralled

around stem; narrowly ovate-lanceolate to

oblong-lanceolate or linear-lanceolate, 1.9-3. 6

mm long; ventral surface keeled; apex acumi-

nate, acute or occasionally obtuse, apiculate,

cells of apiculus elongate, smooth, incrassate,

frequently with single tooth; base scarcely dif-

ferentiated, ± sheathing, margins decurrent;

margins plane, generally crenulate, unistratose.

Costa ending below apex or merging with

elongated cells of apiculus; ventral and dorsal

superficial cells long-rectangular to linear; in

section subround, bulging dorsally, lamina

ventrally inserted, 2 guide cells exposed ven-

trally, ventral stereid band absent, dorsal

stereid band 1-4 cells thick, dorsal surface

cells not differentiated, rough. Upper laminal

cells irregularly rounded-hexagonal, ± flat.

Map 189. — Zygodon trichomitrius

486

Orthotrichaceae

8.5— 1 2.5(— 15.0) (am, papillae low, scattered;

basal cells rectangular, thin-walled to incras-

sate, smooth or walls papillose, occasionally

reaching higher along costa. Gemmae numer-

ous, on stem and rhizoids, clavate to subround,

with transverse and longitudinal septa, reddish

brown.

Dioicous. Perigonia terminal, quickly lateral

by subperigonial innovation; leaves ovate or

shortly ovate-acuminate. Perichaetia terminal,

leaves scarcely differentiated. Seta 7-14 mm

long, yellowish or brownish. Capsule cylindri-

cal or narrowly pyriform, yellowish or brown,

plicate, mouth sulcate dry, reddish brown, urn

1.5- 1. 8 mm long, neck 0.5 mm long; exothecial

cells irregularly quadrate to rectangular, smaller

at mouth, longitudinal walls incrassate above;

annulus apparently absent. Peristome absent.

Operculum 1 mm long, oblique. Calyptra 2.4

mm long, hairy, hairs ascending, uniseriate,

smooth, cells smooth. Spores 16-20 pm, papil-

lose. Fig. 134: 12-23.

Zygodon erosus is known from India, and

central, eastern and southern Africa. In southern

Africa the species occurs on bark or infrequently

on forest litter in montane forests of the eastern

Transvaal region and KwaZulu-Natal. Map 190.

Vouchers: Esterhuysen 20216; Magill 5508 ,

5527, 5616; Sim 10045, 10070; Stirton 8991;

Van Rooy 1629.

Plants of this species are easily identified by

the numerous clavate to subround gemmae,

with transverse and longitudinal septa, on the

stems and rhizoids. The narrower leaves with

acute to acuminate, apiculate apices and the

gemmae characters separate Z. erosus from the

closely related Z. trichomitrius.

Insufficiently known species

Zygodon cernuus C. Miill. in Hedwigia 38:

114 (1899); Broth, in Natiirl. PflFam., edn 2, 11:

14 (1925); Malta, Die Gattung Zygodon 56

(1926). Type: Cape, Somerset East, Mt Bosch-

berg, Jan. 1878, MacOwan s.n. Sim (1926) did

not see the type and it could not be located for

this treatment.

Zygodon perreflexus C. Miill. in Hedwigia

38: 115 (1899); Broth, in Natiirl. PflFam., edn

2, 11: 14 (1925); Malta, Die Gattung Zygodon

134 (1926). Type: Cape, Cape Town, Clare-

mont, Oct. 1876, Rehmann 297. The type could

not be located. The specimen Rehmann 148

(PC!, BM!) collected at Rondebosch and origi-

nally named as Z. perreflexus, is Z. leptobolax.

2. AMPHIDIUM

Amphidium Schimp. nom. cons., Coroll, bryol. eur. 39 (1856); Broth, in Natiirl. PflFam., edn 2,

10: 192 (1924); Nyholm, Moss FI. Fenn. 309 (1954); Sainsb., N. Zeal. Mosses 197 (1955);

Gangulee, Moss. E. India 5: 1150 (1976); Crum & Anderson, Moss. E.N. Amer. 2: 683 (1981); Van

Rooy in Lindbergia 17: 59 (1992). Type species: A. lapponicum (Hedw.) Schimp.

Plants small to medium-sized, densely caespitose; saxicolous to terricolous. Stems erect, papil-

lose. Leaves ± crowded, erect and crisped when dry, erect-spreading and flexuose when wet,

keeled, unistratose; narrowly oblong-lanceolate or linear-lanceolate; apex acute to acuminate; base

Orthotrichaceae

487

scarcely differentiated, sheathing; margins entire or irregularly serrate. Costa ending below apex;

in section with median guide cells. Upper laminal cells rounded, incrassate, papillose on dorsal and

ventral surface, occasionally striolate; basal cells rectangular, smooth to striolate.

Autoicous. Perigonia terminal on short, subperichaetial branches; perichaetia terminal, leaves

weakly or strongly differentiated. Seta short. Capsule erect to inclined, immersed or short-exsert-

ed, short-pyriform or urceolate, 8-ribbed; stomata phaneropore; annulus and peristome absent.

Operculum conic-apiculate or conic-rostellate. Calyptra cucullate, naked, cells generally papillose.

Spores subround to subtriangular, papillose, brownish.

Amphidium contains 12 species and is found on every continent except Antarctica. Five species

are known from Africa.

The genus is recognized by its narrow leaves; median guide cells of the costa; rounded and

papillose to striolate-papillose upper laminal cells; immersed to shortly exserted, 8-ribbed, gym-

nostomous capsule; and cucullate, naked calyptra.

The similarities between Amphidium and Orthotrichum in the development of the young cap-

sules and the papillose calyptrae (Lewinsky 1976), and other morphological characters indicate a

position in the Orthotrichaceae rather than in Dicranaceae or Rhabdoweisiaceae.

Leaf margin irregularly serrate above; perichaetial leaves not or scarcely differentiated, linear-

lanceolate, 2. 5-3. 8 mm long LA. tortuosum

Leaf margin entire; perichaetial leaves differentiated, abruptly apiculate or acuminate above an

oval or oblong base, 1.2-2.0 mm long 2. A. lapponicum

1. Amphidium tortuosum (Hornsch.)

Cufodontis in Oesterr. Bot. Z. 98: 221 (1951);

Van Rooy in Lindbergia 17: 59 (1992). Type:

Cape, ‘Bei dem Wasserfalle auf der ostlichen

Seite des Teufelsberges, 3te Hohe’, Ecklon (not

located).

Syrrhopodon tortuosus Homsch. inLinnaea 15: 117 (1841).

Zygodon cyathicarpus Mont, in Ann. Sci. Nat. Bot. 3,4:

106 (1845). Amphoridium cyathicarpum (Mont.) Jaeg., Ber.

Thatigk. St. Gallischen Naturwiss. Ges. 1872-1873: 108

(1874). Amphidium cyathicarpum (Mont.) Broth, in Natiirl.

PflFam. 1,3: 460 (1902); Sim, Bryo. S. Afr. 267 (1926); vide

Robinson in Smithsonian Contr. Bot. 27: 20 (1975). Type:

Chile.

Plants small, yellow-green to green above,

yellow-brown to brown below; saxicolous to

terricolous. Stems up to 28 mm tall, branching

by subperichaetial innovations, scarcely tomen-

tose below; rhizoids red or reddish brown,

smooth to papillose; in section subround to sub-

triangular, inner cortex 4-8 cells across, thin-

walled, outer cortical cells smaller, in 1 or 2

rows, rarely absent on one side, incrassate,

minutely papillose. Leaves crowded, ± equal in

size, erect and crisped when dry, erect-spread-

ing and flexuose when wet; linear-lanceolate to

linear, 2.CM-.8 mm long; apex acute to acumi-

nate, with a single apical tooth; margins plane,

frequently recurved on one side below, irregu-

larly serrate, frequently denticulate at shoul-

ders, rarely decurrent. Costa ending below

apex; ventral and dorsal superficial cells nar-

rowly rectangular; in section crescent-shaped to

subround, bulging dorsally, ventrally flat, lami-

nal insertion ventral, guide cells in one layer

medially, ventral substereid or stereids in one

layer, papillose, dorsal stereids in 1 or 2 rows,

surface rough. Upper laminal cells rounded-

quadrate or oval, incrassate, bulging ventrally,

papillose, striolate, 6-14 pm, papillae low,

blunt, scattered over dorsal and ventral surface;

basal cells rectangular, thin-walled, smooth to

striolate.

Autoicous. Perigonia terminal on short, sub-

perichaetial branches; inner leaves ovate to

ovate-apiculate to ovate-acuminate, margins

irregularly crenulate. Perichaetia terminal, fre-

quently overgrown by subperichaetial innova-

tions; leaves scarcely differentiated, linear-

lanceolate, 2.5-3. 8 mm long, sheathing below.

488

Orthotrichaceae

Fig 136. — Amphidium tortuosum (1-12): 1. habit

(dry), x 1; 2. habit (wet), x 5; 3. part of stem in cross sec-

tion, x 350; 4. leaves, x 35; 5. leaf cross section, x 350;

6. basal leaf cells, papillae partly shown, x 350; 7. upper

laminal cells, x 700; 8. leaf apex, x 350; 9. upper part of

plant, x 18; 10. part of capsule mouth, x 175; 11. opercu-

lum, x 70; 12. spore, x 700. A. lapponicum (13-25): 13.

habit (dry), x 1; 14. habit (wet), x 5; 15. part of stem in

cross section, x 350; 16. part of rhizoid, x 350; 17.

leaves, x 28; 18. leaf cross section, x 175; 19. upper lam-

inal cells, x 700; 20. leaf apex, x 350; 21. perichaetial

leaf, x 28; 22. upper part of plant, x 28; 23. calyptra, x

55; 24. operculum, x 55; 25. spore, x 700. (1, 4, 6 & 8,

Magill 7141 ; 2, Esterhuysen 26177: 3 & 10, Esterhuysen

35932; 5, Magill 7088: 7, 9 & II. Magill 7159: 12,

Magill 7150, 13. 14, 16 & 20, Van Rooy 1304: 15, 17, 21, 23 & 24, Van Rooy 1316 ; 18, 19 & 22, Van Rooy 1308: 25, Van

Rooy 3081 .)

Orthotrichaceae

489

base oblong or oval. Seta 0.9-1. 5 mm long, fre-

quently curved, yellowish. Capsule erect to

inclined, immersed, urceolate, brownish, urn

0.3-0. 5 mm long, neck 0.2-0. 3 mm long;

exothecial cells irregularly rhomboidal to rec-

tangular, ± thin-walled, in 2 or 3 transverse rows

at mouth, smaller at mouth; stomata present on

neck. Operculum conic-apiculate, oblique.

Calyptra 0. 5-0.6 mm long, cells smooth to

papillose. Spores 12-22 pm. Fig. 136: 1-12.

Primarily a southern hemisphere species,

Amphidium tortuosum is known from Mexico,

South America, Juan Fernandez islands. South

Georgia, Hawaii, Australasia, Indonesia and

sub-Saharan Africa. In southern Africa the

species is rarely collected from rock crevices

on the mountains of KwaZulu-Natal and

Lesotho and on Table Mountain in the south-

western Cape region. Map 191.

Vouchers: Esterhuysen 26177 , 35932;

Magill 7141, 7150, 7159; Van Zanten 7609902.

The plants are recognized by their crisped,

linear-lanceolate to linear leaves with irregular-

ly serrate margins and striolate-papillose cells,

scarcely differentiated perichaetial leaves and

immersed capsules.

2. Amphidium lapponicum ( Hedw .)

Schimp., Coroll, bryol. eur. 39 (1856); Broth, in

Natiirl. PflFam., edn 2, 10: 193 (1924); Nyholm,

Moss FI. Fenn. 310 (1954); Crum & Anderson,

Moss. E.N. Amer. 2: 685 (1981); Van Rooy in

Lindbergia 17: 62 (1992). Type: Hedwigia lap-

ponica Hedw. St. Cr. v. 3. p. t„ nomine Gymnost.,

Exempl. a Cl. Swartz. (G, holo.!).

Anictangium lapponicum Hedw., Sp. muse, frond. 40

(1801). Zygodon lapponicus (Hedw.) B.S.G., Bryol. eur. 3:

38 (1838). Amphoridium lapponicum (Hedw.) Schimp. in

Syn. muse. eur. 1: 247 (1860).

Plants small to medium-sized, yellowish

green to green above, brown below; saxi-

colous. Stems up to 38 mm tall, branching by

subperichaetial innovations, scarcely tomen-

tose below; rhizoids red-brown, smooth to

papillose; in section subround to subtriangular,

inner cortex 5-10 cells across, thin-walled or ±

incrassate, outer cortical cells smaller, in 1 or 2

rows, incrassate, papillose. Leaves ± crowded,

± equal in size, erect and crisped when dry,

erect-spreading and flexuose when wet; nar-

rowly oblong-lanceolate to linear-lanceolate,

2.0-3. 3 mm long; apex acute to acuminate, fre-

quently with a single apical tooth; margins

plane above, recurved below, entire, rarely

decurrent. Costa ending below apex; ventral

and dorsal superficial cells narrowly rectangu-

lar; in section subround or crescent-shaped,

bulging dorsally, ventrally flat, laminal inser-

tion ventral, guide cells in one layer medially,

ventral substereids or stereids in one layer,

papillose, dorsal substereids or stereids in 1 or

2 rows, papillose. Upper laminal cells irregu-

larly rounded, incrassate, ± flat, papillose,

weakly striolate, 8.7-15.0 pm, papillae ± low,

blunt, scattered over dorsal and ventral surface;

basal cells rectangular, thin-walled to incras-

sate, smooth to striolate.

Autoicous. Perigonia terminal on short, sub-

perichaetial branches; inner leaves ovate-apicu-

late or ovate-acuminate. Perichaetia terminal,

frequently overgrown by subperichaetial innova-

tions; leaves differentiated, sheathing, oval or

oblong, 1. 2-2.0 mm long, apex abruptly apicu-

late or acuminate, costa ending below apex,

upper laminal cells irregularly rectangular or

rhomboidal to vermiculate, incrassate, striolate.

Seta short, 0.4-0.6 mm long, yellowish to brown.

490

Orthotrichaceae

Capsule erect, immersed to shortly exserted,

shortly pyriform or urceolate, brownish, urn

0.5-0. 7 mm long, neck 0.3-0. 6 mm long;

exothecial cells irregularly rhomboidal to rectan-

gular, incrassate, in 2-5 transverse rows at

mouth, smaller at mouth; stomata present at base

of urn and on neck. Operculum conic-rostellate,

beak turned to one side. Calyptra 1 mm long,

cells papillose. Spores 11-14 pm. Fig. 136;

13-25.

Amphidium lapponicum is widely distributed

in temperate to arctic regions of the northern

hemisphere, disjunct in South Africa and prob-

ably New Zealand, and also known from

Macaronesia and northern Africa. In southern

Africa the species is known from wet basalt

rock crevices in the Drakensberg Mountains of

KwaZulu-Natal and the Maluti Mountains in

Lesotho. Map 192.

Vouchers: Esterhuysen 25177a; Van Rooy

1304, 1308, 1316, 3081, 3104.

Subfamily ORTHOTRICHOIDEAE

Plants small to medium-sized; saxicolous, terricolous or corticolous. Primary stems erect, sim-

ple or sparsely branched. Leaves appressed, erect or crisped when dry; ovate-lanceolate, oblong to

oblong-lanceolate or linear-lanceolate above an oval or ovate base; base rarely strongly differenti-

ated. Upper laminal cells rounded-hexagonal or irregularly rounded, incrassate, smooth or papil-

lose; basal cells mostly rectangular. Gemmae absent or present, on leaf lamina or rarely on rhizoids.

Perichaetia terminal on primary stems and branches. Dwarf male plants absent. Capsule

immersed, emergent or short-exserted, frequently ribbed. Stomata phaneropore or cryptopore.

Peristome double. Calyptra mitrate, hairy. Isosporous.

3. ORTHOTRICHUM

Orthotrichum Hedw., Sp. muse, frond. 162 (1801); Broth, in Natiirl. PflFam., edn 2, 11: 17

(1925); Sim, Bryo. S. Afr. 273 (1926); Nyholm, Moss FI. Fenn. 321 (1954); Vitt in Nova Hedwigia

21 : 683-7 II (1971); Smith, Moss FI. Brit. Irel. 473 (1978); Lewinsky in Lindbergia 4: 68 (1978);

Crum & Anderson, Moss. E.N. Amer. 2: 687 (1981). Lectotype species: O. anomalum Hedw., vide

Grout, Moss FI. N. Amer. 2: 106 (1935).

Plants small to medium-sized, loosely caespitose to caespitose; corticolous or saxicolous. Stems

erect, simple or sparsely branched by subperichaetial innovations, scarcely tomentose below; rhi-

zoids smooth, reddish brown. Leaves erect to appressed when dry, occasionally flexuose, erect-

spreading to spreading or recurved below when wet; unistratose or rarely bistratose; generally

Map 192. — • Amphidium lapponicum

♦ Orthotrichum firmum

The species is distinguished from Am-

phidium tortuosum by its broader leaves with

entire margins and larger papillae on the cells,

differentiated perichaetial leaves, and immersed

to short-exserted capsules.

Orthotrichaceae

491

ovate-lanceolate; margins incurved or recurved to revolute. Costa with cells scarcely differentiat-

ed in section. Upper laminal cells mostly rounded-hexagonal, frequently incrassate, flat to bulging,

smooth or papillose; basal cells rectangular to quadrate, smooth or papillose. Gemmae present or

absent, filiform, uniseriate, on leaf lamina or rarely terminal on rhizoid.

Autoicous. Perigonia gemmate. Perichaetia terminal; leaves scarcely differentiated, inner

leaves smaller. Seta brownish. Capsule immersed, emergent or exserted, ovate-cylindrical to cylin-

drical, smooth or frequently 8-ribbed, brownish, neck short; stomata phaneropore or cryptopore.

Peristome double, exostome teeth 16 or in 8 pairs; endostome processes absent, 8 or 16, alternat-

ing with teeth, stout or slender, 1 or 2 cells wide, basal membrane low or absent. Operculum conic-

rostellate or conic-apiculate. Calyptra mitrate, plicate, frequently lobed below, hairy. Spores sub-

round to round, brownish, papillose or granulate.

The genus contains ± 200 species and is mainly temperate in distribution. South America is the

major centre of described species, followed by North America and Europe. Of the eight species pre-

sent in southern Africa, four are endemic, two are also known from eastern Africa and two are

widespread.

Orthotrichum is characterized by erect stems with erect to appressed or occasionally flexuose

leaves and mitrate, hairy and usually plicate calyptrae. The southern African species with crypto-

pore stomata belong to section Diaphanum Vent, (subgenus Orthotrichum), O. rupestre belongs to

section Rupestria Schimp. (subgenus Phaneroporum Delogne) and the remaining phaneropore

species belong to section Leiocarpa Mol. (subgenus Phaneroporum). The adaptation to xeric habi-

tats on rock and tree trunks is the major evolutionary trend in the genus (Vitt 1971).

1 Leaf margins incurved 3.0. incurvomarginatum

1 Leaf margins recurved to revolute:

2 Stomata phaneropore:

3 Capsule exserted:

4 Exostome teeth 16; endostome segments 16 1. O.firmum

4 Exostome teeth in 8 pairs; endostome segments 8 2.0. oreophilum

3 Capsule immersed to emergent:

5 Leaf apex acute to acuminate; exostome teeth 16; endostome segments absent or 16,

narrow 5. O. rupestre

5 Leaf apex acuminate to mostly aristate; exostome teeth in 8 pairs; endostome segments

8, broad ; 4. O. armatum

2 Stomata cryptopore:

6 Leaf apex aristate, cells elongate; costa merging with apical cells 8.0. diaphanum

6 Leaf apex rounded-obtuse, acute, acuminate or apiculate, cells not or scarcely differen-

tiated, rounded; costa ending below apex or subpercurrent:

7 Leaves mostly oblong-lanceolate or elliptic; apex apiculate; margins recurved at

midleaf, frequently plane above and below 1.0. transvaalense

1 Leaves narrowly ovate-lanceolate or lanceolate; apex variable, rounded-obtuse, acute

or acuminate, frequently channelled; margins broadly recurved to revolute

6. O. subexsertum

492

Orthotrichaceae

Orthotrichaceae

493

1. Orthotrichum firmum Vent, in Nuovo

Giorn. Bot. Ital. 4: 15 (1872); Broth, in Natiirl.

PflFam., edn 2, 1 1 : 20 ( 1925); Lewinsky & Van

Rooy in J. Bryol. 16: 76 (1990). Type; Bogos

Abyssiniae circa Keren, Beccari s.n. fide

Lewinsky in Bot. Tidsskr. 72: 65 (1978).

Plants small to medium-sized, caespitose,

yellowish green or olivaceous above, green-

brown below; corticolous. Stems 15-25 mm tall,

scarcely tomentose below; in section subround,

central strand absent, inner cortex 7-13 cells

across, thin-walled to incrassate, outer cortical

cells smaller, in 2-4 rows, incrassate to stereids,

epidermis not differentiated. Leaves ± crowded,

erect when dry, spreading when wet; ovate-

lanceolate to lanceolate, (2.2— )2.8— 3.5 (-5.0)

mm long; ventral surface keeled; apex acute to

acuminate, frequently channelled, 0. 1— 0.2(— 0.3 )

mm long; margins recurved, entire, flexuose;

unistratose. Costa ending below apex or extend-

ing into channelled apex; superficial cells round-

ed above, linear below, flat to bulging, smooth;

in section crescent-shaped to subround, bulging

dorsally, cells scarcely differentiated, incrassate.

Upper laminal cells rounded-hexagonal to oval,

incrassate, slightly bulging, 11-20 pm long,

papillae low, blunt, 1-3 over lumen, mostly sim-

ple; basal cells smooth or walls papillose, rec-

tangular, quadrate towards margin, thin-walled

to incrassate.

Gonioautoicous. Perigonial leaves ovate.

Perichaetia terminal; leaves scarcely differenti-

ated, inner leaves smaller. Seta 24-3.8 mm

long, occasionally polysetaceous, twisted anti-

clockwise above. Capsule exserted, oblong-

cylindrical, 1.5-2. 5 mm long, 8-ribbed above.

neck weakly differentiated; exothecial cells

irregularly rectangular, longitudinal walls

incrassate, shorter at mouth; stomata at base of

urn and on neck, phaneropore. Peristome dou-

ble; exostome teeth 16, recurved when dry,

oblong, blunt, 300-350 pm long, yellowish

brown, densely papillose; basal membrane

absent, processes 16, alternating with teeth,

200-310 pm long, 2 cells wide, cell divisions

prominent below, papillose. Operculum 0.5 mm

long, conic-rostellate, yellowish with reddish

brown rim. Calvptra 3. 0-3. 5 mm long; hairs

ascending, papillose. Spores 20-35 pm, papil-

lose. Fig. 137: 15-22.

Known from eastern and central Africa,

India, and recently reported from southern

Africa. The species is rarely collected in mon-

tane forests and wooded areas of the northern

Transvaal area, KwaZulu-Natal and the eastern

Cape region. Map 192.

Vouchers: Magill 3710, 5686a, 5691.

The recurved leaf margins, channelled apex,

long-exserted capsule and 16 exo- and endo-

stome teeth separate O. firmum from other

phaneropore species in the Flora area.

2. Orthotrichum oreophilum Lewinsky &

Van Rooy in J. Bryol. 16: 74 (1990). Type: Le-

sotho, Kotisephola Pass, Van Rooy 3423 (PRE,

holo.!).

Plants small to medium-sized, loosely caespi-

tose to caespitose, olivaceous to dark green

above, dark green to brownish below; saxi-

colous. Stems 10-28 mm tall; in section subpen-

Fig. 137. — Orthotrichum incurvomarginatum ( 1-14): 1. habit (dry), x 1.4; 2. habit (wet), x 5; 3. part of stem in cross

section, x 175; 4. leaf, x 18; 5. leaf in cross section, x 175; 6. basal leaf cells, x 350; 7. upper laminal cells, x 350; 8. cells

at leaf apex, x 175; 9. perichaetial leaf, x 18; 10. part of capsule wall with stoma, x 350; 11. part of capsule mouth with

peristome (papillae partly shown), x 175; 12. operculum, x 88; 13. calyptra, x 18; 14. spore, x 700. O. firmum (15-22):

15. habit (dry), x 1.4; 16. habit (wet), x 5; 17. leaf, x 18; 18. leaf in cross section, x 175; 19. basal leaf cells (right side), x

350; 20. leaf apex, x 175; 21. part of capsule mouth with peristome (papillae partly shown), x 175; 22. spore, x 700. O.

oreophilum (23-30): 23. habit (dry), x 1 ; 24. habit (wet), x 5; 25. leaf, x 25; 26. leaf in cross section, x 175; 27. upper lam-

inal cells (left side), x 350; 28. perichaetial leaf, x 50; 29. part of capsule mouth with peristome (papillae partly shown), x

175; 30. spore, x 700. ( 1, 2 & 4, Magill & Schelpe 3975: 3 & 9, Magill & Schelpe 4022: 5-7, Magill & Schelpe 3943: 8,

10 & 13, Schelpe 4968 ; 11, Magill & Schelpe 4012: 12, Barnard 49647- 14, Van der Westhuizen PRE-CH13542: 15, 16, 18,

19 & 22, Magill 5691: 17, Magill 3710: 20, Sim PRE-CH5736: 21, Oliver 6775: 23, 24, 28 & 30. Van Rooy 3423: 25, Van

Rooy 3376: 26, Van Rooy 3033: 27, Van Rooy 3078: 29, Van Rooy 3075.)

494

Orthotrichaceae

tagonal to subround, central strand absent, inner

cortex 6-8 cells across, incrassate, outer cortical

cells smaller, in 1-3 rows, substereids to ster-

eids, epidermis not differentiated. Leaves ±

crowded above, erect to erect-spreading, fre-

quently flexuose when dry, spreading when wet;

unistratose; ovate-lanceolate to narrowly lanceo-

late, ( 1 .4 — )2.0— 3.0(— 3.7) mm long; ventral sur-

face keeled; apex acuminate; margins irregular-

ly recurved, entire, unistratose. Costa ending

below apex; superficial cells rounded above to

linear below, bulging dorsally, papillose; in sec-

tion crescent-shaped to subround, bulging dor-

sally, cells scarcely differentiated, incrassate to

substereids. Upper laminal cells irregularly

rounded-hexagonal to oval, incrassate, flat to

slightly bulging, 10-20 pm, papillae bifid; basal

cells yellowish to orange or reddish brown

below, rectangular to quadrate at margin, incras-

sate, walls occasionally papillose, frequently pit-

ted, cells occasionally inflated at comers.

Gonioautoicous. Perigonial leaves ovate.

Perichaetia terminal; inner leaves narrowly

subtriangular, as long as stem leaves or inner

leaves smaller. Seta 1.2-1. 8 mm long, straight

or slightly twisted anticlockwise above. Capsule

shortly exserted, oblong-cylindrical, 1. 6-2.0

mm long, smooth, neck weakly differentiated;

exothecial cells irregularly rectangular to

quadrate above, longitudinal walls strongly

incrassate; stomata phaneropore, on um. Peri-

stome double, exostome teeth in 8 pairs, trian-

gular, cleft down middle, perforated above,

140-260 pm long, pale yellow, papillose, fre-

quently with vermiculate lines; basal membrane

absent, processes 8, alternating with teeth pairs,

narrowly oblong or linear, frequently 2 cells

wide, 125-210 pm long, median lines conspic-

uous, ± papillose. Operculum conic-rostellate,

0. 3-0.5 mm long, yellowish with reddish brown

rim. Calyptra 2. 0-2. 3 mm long, hairs inultiseri-

ate, papillose. Spores 15-25 pm, granulate. Fig.

137: 23-30.

Endemic to the mountains of Lesotho, O. oreo-

philum has been collected only above 3000 m.

Map 193.

Vouchers: Magill 4569; Van Rooy 3015,

3033, 3376.

Map 193. — ♦ Orthotrichum oreophilum

• Orthotrichum incurvomarginatum

Growing on rock like O. rupestre , this pha-

neropore species can be recognized by the erect

to erect-spreading, flexuose leaves when dry,

generally acuminate leaf apex, exserted cap-

sule, 8 exostome teeth pairs, 8 endostome pro-

cesses and strongly incrassate exothecial cells.

3. Orthotrichum incurvomarginatum

Lewinsky & Van Rooy in J. Bryol. 16: 67

(1990). Type: Cape, upper slopes of Mount

Synnott, 7 km N of Clanwilliam, Magill &

Schelpe 4012 (PRE, holo.!).

Plants small to medium-sized, caespitose,

yellowish green or olivaceous above, yellowish

brown or brown-green below; corticolous.

Stems 8-26 mm tall, scarcely tomentose below;

in section subround to subpentagonal, central

strand absent, inner cortex 11-14 cells across,

thin-walled to incrassate, outer cortical cells

smaller, in 1-4 rows, incrassate to stereids, epi-

dermis not differentiated, outer surface rough.

Leaves ± crowded, erect when dry, spreading

when wet; ovate-lanceolate to lanceolate,

2.4 — 3.7( — 4.3 ) mm long; ventral surface keeled;

apex narrowly acuminate to subulate, 0. 1-0.4

(-0.7) mm long, cells rounded to elongate; mar-

gins incurved, entire; unistratose. Costa weak

above, extending into acumen; ventral and dor-

sal superficial cells rounded above, linear

Orthotrichaceae

495

below, flat to bulging, smooth to weakly papil-

lose; in section crescent-shaped to subround,

bulging dorsally, cells scarcely differentiated,

incrassate. Upper laminal cells rounded-hexag-

onal to oval, incrassate, slightly bulging,

10.0-1 8. 7(— 33.0) pm long, papillae low, blunt,

1-3 over lumen, simple or forked; basal cells

smooth or papillose, rectangular, quadrate

towards margin, generally thin-walled.

Gonioautoicous. Perigonial leaves ovate.

Perichaetia terminal; leaves narrowly oblong or

lanceolate, 2. 0-3. 6 mm long, apex acuminate to

subulate, 0.2-0.6(-1.0) mm long, costa extend-

ing into acumen. Seta short, (0.3— )0.5— 0.8(— 1 .3)

mm long. Capsule immersed to emergent,

oblong-cylindrical, 1.9-2.3(-3.0) mm long, 8-

ribbed, neck differentiated; exothecial cells rec-

tangular, longitudinal walls incrassate, shorter

at mouth; stomata at base of urn and on neck,

phaneropore. Peristome double; exostome teeth

in 8 pairs, triangular, 160-275 pm long, dense-

ly papillose; basal membrane absent, processes

8, alternating with teeth pairs, 175-240 pm

long, 2 cells wide, coarsely papillose, cell divi-

sions prominent below. Operculum 0.3 mm

long, conic-apiculate, yellowish with red or red-

brown rim. Calyptra 2. 5-3.0 mm long; hairs

ascending, multi- to uniseriate, crenulate to den-

ticulate, smooth to papillose. Spores 16-25 pm,

papillose. Fig. 137: 1-14.

Endemic to southern Africa, O. incurvomar-

ginatum is found on indigenous and exotic trees

and shrubs in the winter rainfall regions of the

southwestern and northwestern Cape. Map 193.

Vouchers: Barnard PRE-CH3114; Magill &

Schelpe 3943, 4025; Schelpe 4968.

The incurved leaf margins distinguish O.

incurvomarginatum from other species of the

genus in the Flora area.

4. Orthotrichum armatum Lewinsky &

Van Rooy in J. Bryol. 16: 69 (1990). Type:

Cape, Hogsback, Oliver 6775 (C, holo.!).

Plants medium-sized, mixed with other

bryophytes, olivaceous or yellowish green

above, greenish brown to brown below; corti-

colous. Stems 10-15 mm tall; in section sub-

pentagonal to subround, central strand absent,

inner cortex 12-14 cells across, incrassate,

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

stereids to stereids, epidermis not differentiated,

outer surface rough. Leaves ± crowded, erect

when dry, spreading when wet; unistratose;

ovate-lanceolate, 2. 5^4.0 mm long; ventral sur-

face keeled; apex narrowly acuminate to aris-

tate, 0.3-0. 6 mm long; margins recurved for

lower ± three-quarters, plane above, entire to

crenulate, unistratose. Costa extending into

arista; superficial cells rounded to oval above,

linear below, ± flat, smooth; in section crescent-

shaped to subround, bulging dorsally, cells

scarcely differentiated, incrassate. Upper lami-

nal cells shortly rounded-rhomboidal, oval or

irregular in shape, flat to weakly bulging,

( 1 OX)—) 1 5.0— 27.5(— 35 .0) pm long, 7.5-13.8 pm

wide, smooth to papillose, papillae low, blunt,

mostly simple; basal cells yellowish to yellow-

ish brown below, walls papillose, frequently pit-

ted, rhomboidal to rectangular, quadrate at mar-

gin, incrassate.

Gonioautoicous. Perigonial leaves ovate.

Perichaetia terminal; leaves ovate-lanceolate or

lanceolate, apex aristate to subulate, inner leaves

smaller. Seta short, 0.8-1. 5 mm long. Capsule

immersed to emergent, ovate-cylindrical, 1. 5-2.2

mm long, 8-ribbed in upper half when dry, neck

weakly differentiated; exothecial cells irregular-

ly rectangular, thin-walled to incrassate, smaller

at mouth, incrassate; stomata phaneropore, on

urn. Peristome double; exostome teeth in 8

pairs, reflexed when dry, cleft down middle, tri-

angular, 280-330 pm long, yellowish brown,

minutely striolate-papillose; basal membrane

absent, processes 8, alternating with tooth pairs,

broad, stout, oblong-lanceolate or triangular,

blunt, 2 cells wide, dividing lines conspicuous,

papillose with ridges along walls. Operculum

conic-apiculate, 0.4 mm long. Calyptra 2.2-3.0

mm long, weakly plicate, hairy. Spores 25-34

(-53) pm, papillose. Fig. 138: 1-5.

This endemic is known only from Hogsback

in the eastern Cape region. Map 194.

Voucher: type only.

496

Orthotrichaceae

Orthotrichum armatum is most easily sepa-

rated from other phaneropore species in south-

ern Africa by the narrowly acuminate to aristate

leaf apex and recurved margins. The species is

also characterized by the crenulate leaf margins,

immersed to emergent capsule, 8 pairs of exo-

stome teeth and 8, broad, stout, blunt endo-

stome processes.

5. Orthotrichum rupestre Schleich. ex

Schwaegr., Sp. muse, frond, suppl. 1,2: 27

(1816); Broth, in Natiirl. PflFam., edn 2, 11: 17

(1925); Nyholm, Moss FI. Fenn. 324 (1954);

Lawton, Moss FI. Pacific Northwest 226 ( 1971);

Crum & Anderson, Moss. E.N. Amer. 2: 696

(1981). Lectotype: Austria, Schwagrichen s.n.

(G) fide Lewinsky in Lindbergia 9: 56 (1983).

Orthotrichum macleanum Shaw in Cape Monthly Mag.

17: 378 ( 1878). Isotypes: Cape, Graaff-Reinet, MacLea sub

Rehmann 514 (BM!; H!; PRE!).

Plants small to medium-sized, caespitose, oli-

vaceous above, dark green or greenish brown

below; saxicolous or rarely corticolous. Stems

5 — 40 mm tall, tomentose below; in section sub-

round to subpentagonal, central strand absent,

inner cortex 9-20 cells across, thin-walled to

incrassate, outer cortical cells smaller, in 2-A

rows, substereids or stereids, epidermis not dif-

ferentiated, outer surface rough. Leaves ± crowd-

ed, erect-appressed when dry, erect-spreading to

spreading or recurved when wet, unistratose or

bistratose; ovate-lanceolate or lanceolate,

(1.7-)2.3-3.8(-^L2) mm long; ventral surface

keeled; apex acute to acuminate; margins

recurved to revolute, entire, unistratose to tris-

tratose. Costa subpercurrent; superficial cells

rounded to linear, flat to slightly bulging, smooth

to papillose; in section crescent-shaped to sub-

round, bulging dorsally, cells scarcely differenti-

Fig. 138. — Orthotrichum armatum (1-5): 1. habit

(dry), x 3; 2. leaf, x 17; 3. leaf in cross section, x 175; 4.

upper laminal cells, x 350; 5. cells at leaf apex, x 175. O.

rupestre (6-13): 6. habit (dry), x 1; 7. habit (wet), x 3; 8.

leaf, x 18; 9. leaf in cross section, x 175; 10. basal leaf cells,

x 350; 11. upper laminal cells, x 350; 12. part of capsule

mouth with peristome teeth (papillae partly shown), x 175;

13. spore, x 700. (1-5, Oliver 6775\ 6 & 7, Magill 5886: 8,

1 1-13, Rehmann 514 ; 9, Magill 5887a ; 10, Magill 5913.)

Orthotrichaceae

497

Map 194. — ■ Orthotrichum armatum

• Orthotrichum rupestre

♦ Orthotrichum transvaalense

ated, incrassate to substereids. Upper laminal

cells irregularly rounded-hexagonal, incrassate,

flat or slightly bulging, 8.7-15.0 (-22.5) |jm

long, papillae low, blunt, 1-3 over lumen, simple

or bifid; basal cells yellow to orange below,

smooth, walls papillose or 1 papilla over lumen,

rectangular to quadrate at margin, thin-walled to

incrassate, occasionally pitted.

Gonioautoicous. Perigonial leaves ovate.

Perichaetia terminal; leaves ovate-acuminate to

oblong-acuminate or lanceolate, ( 1 .8— )3. 2—4.5

mm long. Seta short, 0.4—1 .2 mm long, 1 or 2 per

perichaetium. Capsule immersed to emergent,

1. 4-2.4 mm long, 8-ribbed above; urn ovate-

cylindrical to oblong-cylindrical, neck contract-

ed; exothecial cells irregularly rectangular to

quadrate, shorter above, incrassate; stomata

phaneropore, on urn. Peristome single or infre-

quently double, preperistome occasionally pre-

sent; exostome teeth 16, narrowly triangular, per-

forated above, 160-275 pm long, yellowish,

coarsely papillose to reticulate; basal membrane

absent, processes absent or 16, alternating with

exostome teeth, narrow, pale. Operculum 0.4— 0.6

mm long, conic-rostellate, yellowish brown with

reddish brown rim. Calyptra 0. 8-1.0 mm long,

hairy or almost smooth with multicellular out-

growths. Spores 15.0-27.5 pm, granulate. Fig.

138:6-13.

This widespread species has been reported

from all continents. In Africa O. rupestre is

known from northern and eastern Africa and in

the Flora area it is infrequently collected at high

altitudes in Lesotho and the central and south-

western Cape regions. The plants grow on rock

and rarely on tree trunks in southern Africa.

Map 194.

Vouchers: Barnard PRE-CH6220; Ester-

huysen 24359; Magill 5886, 5891, 5913; Van

Rooy 2996.

This variable species is characterized by

phaneropore stomata, immersed to emergent

capsules with 16 exostome teeth, and leaves

with acute to acuminate apices and recurved to

revolute margins. African plants generally lack

an endostome (Lewinsky 1978) but endostome

processes were observed in some specimens

from Lesotho (Van Rooy 2996). Exostome teeth

ornamentation varies from papillose to coarsely

papillose to reticulate. One of the Lesotho speci-

mens (Van Rooy 2996) differs from other south-

ern African material in the bistratose leaves and

the almost naked calyptra with multicellular,

leaf-like outgrowths.

6. Orthotrichum subexsertum Schimp. ex

C. Mull, in Bot. Zeitung (Berlin) 16: 164

(1858); Broth, in Natiirl. PflFam., edn 2, 11: 21

(1925); Lewinsky in Bot. Tidsskr. 72: 80

(1978). Type: Cape, Genadendal, no collector

given (BM!).

Orthotrichum pseudotenellum sensu Sim, Bryo. S. Afr.

275 (1926).

Plants small to medium-sized, caespitose,

olivaceous or yellowish green above, greenish

brown or yellowish brown to brown below; cor-

ticolous or rarely saxicolous. Stems 5-18 mm

tall, tomentose below; in section subround to

subpentagonal, central strand absent, inner cor-

tex 7-15 cells across, thin-walled to incrassate,

central cell walls occasionally collapsed, outer

cortical cells smaller, in 1-3 rows, incrassate or

substereids, epidermis not differentiated, outer

surface ± rough. Leaves erect to appressed

when dry, occasionally flexuose, spreading

when wet; unistratose, rarely with bistratose

Orthotrichaceae

499

patches; ovate-lanceolate to lanceolate, (1.7-)

1 .9— 3.5(— 3.7) mm long; ventral surface keeled;

apex acuminate, acute or rounded-obtuse, fre-

quently channelled; margins broadly recurved

to revolute, entire to crenulate below, serrulate

to serrate at apex, unistratose. Costa ending be-

low apex to subpercurrent; superficial cells

rounded above, elongate below, flat to slightly

bulging, smooth to weakly papillose; in section

crescent-shaped, subround or subquadrate,

bulging dorsally, cells scarcely differentiated,

incrassate. Upper laminal cells rounded-hex-

agonal, incrassate, flat to bulging, 10-20 pm,

smooth to papillose, papillae low, blunt; basal

cells rectangular to quadrate, thin-walled to

incrassate, ± flat, smooth or walls papillose,

occasionally pitted. Gemmae fusiform, on leaf

lamina or rarely terminal on rhizoid, uniseriate,

up to 17 cells long, reddish brown.

Autoicous. Perigonia gemmate, on short or

long branches; inner leaves ovate. Perichaetia

terminal; leaves scarcely differentiated or inner

leaves smaller. Seta (0.25-)0. 6-1.1 mm long,

straight or twisted anticlockwise above.

Capsule emergent, ± cylindrical, 1.5-2. 8 mm

long, contracted below mouth when dry, 8-

ribbed, neck differentiated; exothecial cells

quadrate to rectangular, thin-walled or longitu-

dinal walls incrassate, smaller and incrassate

towards mouth, differentiated along ribs; sto-

mata cryptopore, on base of urn and neck.

Peristome double; exostome teeth in 8 pairs,

reflexed when dry, cleft down middle, frequent-

ly perforated, triangular, ( 160— )210— 330 pm

long, brownish, densely papillose; basal mem-

brane low, processes 8, alternating with teeth

pairs, linear from broad base, 140-230 pm long,

smooth or weakly papillose, striolate below.

Map 195. — Orthotrichum subexsertum

pale yellow. Operculum conic-apiculate, 0.2-

0.5 mm long. Calyptra 1.8-2. 7 mm long, hairs

uniseriate above, bi- to multiseriate below.

Spores ( 12.5— )17.5—20.0(— 22.5) pm, granulate.

Fig. 139: 1-14.

Restricted to Africa, the species is known

from Kenya, Rwanda and South Africa.

Orthotrichum subexsertum is found on indige-

nous or exotic trees and shrubs and rarely on

rock. In the Flora area most specimens have

been collected in the southern and southwestern

Cape regions but the species is also known from

the eastern Cape, KwaZulu-Natal and Zululand.

The species is often mixed with O. diaphanum.

Map 195.

Vouchers: Barnard PRE-CH3404; Magill

5931a; Magill & Schelpe 4026; Sim 9498;

Thome PRE-CH6420; Van Rooy 799.

The ovate-lanceolate to lanceolate leaves

with broadly recurved to revolute margins.

Fig. 139 — Orthotrichum subexsertum (1-14): 1. habit (dry), x 3; 2. habit (wet), x 5; 3. part of stem in cross section,

x 175; 4. leaf, x 35; 5. leaf in cross section, x 175; 6. basal leaf cells, x 350; 7. upper laminal cells, x 350; 8. cells at leaf

apex, x 175; 9. gemma, x 175; 10. part of capsule wall with stoma, x 350; 11. part of capsule mouth with peristome, x 175;

12. operculum, x 70; 13. calyptra, x 18; 14. spore, x 700. O. transvaalense (15-22): 15. habit (dry), x 1.4; 16. habit (wet),

x 5; 17. leaf, x 30; 18. leaf in cross section, x 175; 19. upper laminal cells, x 350; 20. leaf apex, x 175; 21. gemma, x 175;

22. part of capsule mouth with peristome, x 175. O. diaphanum (23-30): 23. habit (dry), x 3; 24. habit (wet), x 5; 25. leaf,

x 35; 26. leaf in cross section, x 175; 27. basal leaf cells (left side), x 350; 28. leaf apex, x 175; 29. gemma, x 175; 30. part

of capsule mouth with peristome, x 175. ( 1, 2 & 12, Sim 9498; 3, Barnard 4965; 4, Wager PRE-CH 12070 ; 5, 7, 11 & 13,

Thome PRE-CH6420; 6, Magill 4033; 8, Magill 4026; 9, Van Rooy 1499a; 10, Barnard 49634 ; 14, Van Rooy 799 ; 15 & 16,

Magill 5051 ; 17, Van Rooy 594 ; 18, Magill 5061 ; 19-22, Smook 6416 ; 23 & 24, Van Rooy 2371; 25 & 28, Van Rooy 553;

26, Magill 4032; 27, Magill 6127; 29, Schelpe 7644; 30, Van Rooy 2368.)

500

Orthotrichaceae

emergent capsule, contracted below the mouth

and 8-ribbed when dry, double peristome with 8

pairs of exostome teeth and 8 narrow endo-

stome segments, and the fusiform gemmae

define this cryptopore species. The variable leaf

apex can be acuminate, acute or rounded-obtuse

and is frequently channelled and toothed.

7. Orthotrichum transvaalense Sim ,

Bryo. S. Afr. 276 (1926); Lewinsky in Bot.

Tidsskr. 72: 79 (1978). Type: Transvaal, Mont.

Lechlaba, Houtbosch, Rehmann 517 (PRE!).

Plants small, loosely caespitose to caespitose,

green or yellowish green above, brown-green or

yellowish brown to brown below; corticolous.

Stems 3-13 mm tall; in section subround, central

strand absent or cell walls collapsed, inner cor-

tex 6-10 cells across, thin-walled, outer cortical

cells weakly differentiated, smaller, in 1 or 2

rows, firm-walled to incrassate, epidermis not

differentiated. Leaves larger above, erect to

appressed when dry, erect-spreading when wet,

unistratose, concave; oblong-lanceolate, lanceo-

late or elliptic, (1.4—) 1.7-2.8(-4.0) mm long;

ventral surface keeled; apex acute or apiculate,

60-220 pm long, cells not or weakly differenti-

ated, rounded, incrassate, occasionally chan-

nelled; margins recurved, plane above, frequent-

ly plane below, entire, occasionally denticulate

at apex, unistratose. Costa ending below apex;

superficial cells rounded above, rectangular to

linear below; in section crescent-shaped, cells

scarcely differentiated, thin-walled to incrassate.

Upper laminal cells rounded-hexagonal, thin-

walled to incrassate, weakly bulging to bulging,

( 1 5.0—) 1 7.5— 22.5(— 27.5) pm, smooth or weakly

papillose; basal cells smooth or walls papillose,

rectangular to quadrate, thin-walled or occasion-

ally incrassate. Gemmae fusiform, on lamina, up

to 13 cells long, brownish or reddish brown,

smooth.

Autoicous. Perigonia gemmate, on short or

numerous on long branches; inner leaves ovate.

Perichaetia terminal; leaves scarcely differenti-

ated, as long as stem leaves or inner leaves

smaller. Seta short, 0.5-0. 7 mm long, straight.

Capsule emergent, ovate-cylindrical, 1. 2-2.1

mm long, 8-ribbed, neck short; exothecial cells

rectangular, thin-walled, weakly differentiated

along ribs, rounded at mouth, incrassate; sto-

mata cryptopore, on base of urn. Peristome dou-

ble; exostome teeth loosely in 8 pairs, triangu-

lar, cleft almost entire length, perforated,

160-320 pm long, yellowish brown, recurved

when dry, densely papillose; basal membrane

absent, processes 8 or occasionally 16, linear, 1

cell wide, 130-260 pm long, yellowish or hya-

line, smooth or weakly papillose. Operculum

short conic-apiculate. Calyptra 1.6-2. 3 mm

long, smooth to scabrous above, lobed below,

hairs uniseriate above, multiseriate below, den-

ticulate. Spores 17-23 pm, papillose to granu-

late. Fig. 139: 15-22.

This endemic is rarely collected from indige-

nous and exotic tree trunks in the Free State,

KwaZulu-Natal and the northern Transvaal

region. It is frequently found mixed with O.

diaphanum, and species of Fabronia Raddi and

Syntrichia Brid. Map 194.

Vouchers: Magill 5051; Smook 6416; Van

Rooy 594.

The species is closely related to O. dia-

phanum but can be distinguished by the wider,

concave leaves, generally appressed when dry;

apiculate leaf apex consisting of scarcely differ-

entiated, rounded cells; the costa ending below

the apex; and the exostome teeth loosely in 8

pairs.

8. Orthotrichum diaphanum Schrad. ex

Brid., Muscol. recent. 2,2: 29 (1801); Broth, in

Natiirl. PflFam., edn 2, 11: 21 (1925); Nyholm,

Moss FI. Fenn. 345 (1954); Lewinsky in Bot.

Tidsskr. 72: 77 (1978); Smith, Moss FI. Brit.

Irel. 486 (1978); Crum & Anderson, Moss. E.N.

Amer. 2: 696 (1981). Type: Europe.

Orthotrichum glaucum Spreng., Syst. veg. 4,2: 323

(1827); Sim. Bryo. S. Afr. 275 (1926). Type: Cape, Cape

Town, Table Mountain, Ecklon s.n. (BM!).

Orthotrichum pseudotenellum Hampe ex C. Mull, in

Bot. Zeitung (Berlin) 17: 230 (1859); Broth, in Natiirl.

PflFam., edn 2. 1 1 : 22 (1925). Type: Cape, Genadendal,

Breutel s.n. (BM!).

Orthotrichum piliferum Sim, Bryo. S. Afr. 274 (1926).

Lectotype: Cape, Uitenhage, Sim 9001 (PRE!) vide

Lewinsky in Bot. Tidsskr. 72: 77 (1978).

Orthotrichaceae

501

Plants small, loosely caespitose to caespi-

tose, green or yellowish green above, yellowish

brown to brown below; corticolous. Stems up to

13 mm tall, occasionally forked, scarcely

tomentose below; in section subround to sub-

pentagonal, central strand absent or cell walls

collapsed, inner cortex 4—11 cells across, thin-

walled to incrassate, outer cortical cells weakly

differentiated, smaller, in 1 or 2 rows, incras-

sate, epidermis not differentiated. Leaves larger

above, erect, frequently flexuose when dry,

erect-spreading to spreading or reflexed below

when wet, unistratose; ovate-lanceolate or

oblong-lanceolate to lanceolate, (1.5— )1. 8-2.8

(-3.0) mm long; ventral surface keeled; apex

gradually or abruptly aristate, 0. 1— 0.5(— 1 .0)

mm long, cells elongate, incrassate, yellowish

or hyaline, occasionally channelled; margins

broadly recurved to revolute, frequently plane

above and below, entire, crenulate to denticulate

at apex, unistratose. Costa merging with apical

cells, occasionally ending below apex in lower

leaves; superficial cells rounded above, rectan-

gular to linear below, flat to bulging dorsally,

smooth or weakly papillose; in section crescent-

shaped to subround, bulging dorsally, cells

scarcely differentiated. Upper laminal cells

rounded-hexagonal, thin-walled or occasionally

incrassate, thickened at the corners, bulging,

1 5— 25(— 30) pm, smooth or weakly papillose;

basal cells rectangular to quadrate, smooth or

walls papillose, thin-walled or occasionally

incrassate. Gemmae fusiform, on lamina, up to

19 cells long, reddish brown, smooth.

Autoicous. Perigonia gemmate, on short or

numerous on long branches, occasionally

appearing to be on separate plants. Perichaetia

terminal; leaves scarcely differentiated, as long

as stem leaves or inner leaves smaller. Seta

short, 0.3-0. 8 mm long, ± straight. Capsule

emergent, ovate-cylindrical, 1. 2-2.1 mm long,

weakly 8-ribbed, neck short; exothecial cells

rectangular, thin-walled, weakly differentiated

along ribs, smaller towards mouth, incrassate;

stomata cryptopore, on base of urn. Peristome

double; exostome teeth 16, narrowly subtrian-

gular, frequently perforated above, 130-260 pm

long, yellowish or brownish, recurved in old

capsules, densely papillose to striolate-papil-

lose; basal membrane absent, processes 16, lin-

ear, 150-190 pm long, yellowish or hyaline,

smooth to spiculose. Operculum conic-apicu-

late, 0.2-0.4 mm long, yellowish brown to

brown. Calyptra 1.5-2. 3 mm long, smooth to

scabrous, hairs uniseriate above to multiseriate

below, denticulate. Spores 13-23 pm, granulate.

Fig. 139: 23-30.

Orthotrichum diaphanum is known from

North and South America, Europe, the western

part of temperate Asia, Macaronesia, Hawaii

and northern, eastern and southern Africa. In the

Flora area the species is most often collected in

the eastern Free State and the eastern, central,

southern, southwestern and northwestern Cape

regions but it is also known from KwaZulu-

Natal and the southern Transvaal region. Plants

grow on bark of indigenous and exotic trees and

shrubs, and is frequently collected with O.

subexsertum and species of Fabronia Raddi and

Syntrichia Brid. Map 196.

Vouchers: Barnard PRE-CH8999 ; Magill

5848, 5914; Magill & Schelpe 4032; Van Roov

393, 2348, 2371, 2684.

This cryptopore species is recognized by the

yellowish or hyaline awn formed by the elon-

gated cells of the leaf apex. Also see note under

O. transvaalense (p. 500).

502

Orthotrichaceae

Insufficiently known species

Orthotrichum afrofastigiatum C. Mull, in

Hedwigia 38: 113 (1899); Broth, in Natiirl.

PflFam., edn 2, 11: 17 (1925); Sim, Bryo. S.

Afr. 273 (1926). Type: Eastern Cape, Mac-

Owan s.n. (H!). The name has been misap-

plied by Lewinsky (1978) and is regarded as

a nomen dubium (Lewinsky & Van Rooy

1990).

4. STONEOBRYUM

Stoneobryum Norris & Robinson in Bryologist 84: 96 (1981). Type species: S. bunyaense Norris

& Robinson.

Plants small; corticolous. Stem erect, branching by subperichaetial innovations. Leaves larger

above, crispate dry, oblong to oblong-lingulate, base occasionally sheathing. Costa in section

with cells scarcely differentiated. Upper laminal cells rounded-hexagonal, incrassate, smooth;

basal cells rectangular, thin-walled to incrassate, frequently papillose. Gemmae absent.

Autoicous. Perigonia lateral, gemmate. Perichaetia terminal; leaves highly differentiated, hya-

line, erect, enveloping capsule. Seta very short. Capsule immersed, ribbed; stomata cryptopore;

annulus absent. Peristome double; exostome teeth in 8 pairs, triangular, minutely papillose to stri-

olate-papillose; endostome processes narrow, keeled, alternating with teeth pairs, shorter than

teeth, minutely striolate-papillose, basal membrane absent. Operculum short-conical. Calyptra

mitrate, hairy. Spores round, minutely papillose, brownish.

The genus contains two species: Stoneobryum bunyaense in Australia and S. mirum in South

Africa. The crispate leaves, the highly differentiated, hyaline perichaetial leaves enveloping the

almost sessile capsule, the cryptoporous stomata, and the shortly mitrate, hairy calyptra separate

Stoneobryum from other genera in Orthotrichoideae.

Stoneobryum mirum (Lewinsky) Norris

& Robinson in Bryologist 84: 98 (1981). Type:

Natal, Scheepers Nek, Arcadia, Sim 10104

(PRE, holo.!).

Orthotrichum mirum Lewinsky in Bot. Tidsskr. 72: 73

(1978).

Orthotrichum afrofastigiatum sensu Sim, Bryo. S. Afr.

273 (1926).

Plants small, caespitose, yellowish green or

olivaceous above, brownish below; corticolous.

Stem up to 15 mm tall, tomentose below; rhi-

zoids reddish brown, smooth; in section with

inner cortex 5-15 cells across, thin-walled to

incrassate, outer cortical cells smaller, in 1-3(4)

rows, incrassate or consisting of substereids or

stereids. Leaves ± crowded, larger above,

crispate dry, erect-spreading to widespreading

wet, unistratose; oblong to oblong-lingulate,

1. 8-3.0 mm long; ventral surface keeled; apex

acute or occasionally acuminate or cuspidate;

base occasionally sheathing; margins plane to

recurved below, entire. Costa ending below

apex, percurrent or infrequently mucronate; in

section round, bulging dorsally, ventrally flat,

laminal insertion ventral, ventral stereid band

absent, 2^1 ventral cells larger, dorsal stereids

or substereids present. Upper laminal cells

rounded-hexagonal, incrassate, essentially flat,

1 1-19 pm, smooth; basal cells rectangular, thin-

walled to incrassate, infrequently papillose.

Autoicous. Perigonia lateral, gemmate,

leaves ovate. Perichaetia terminal, frequently

overgrown by subperichaetial innovations;

inner leaves hyaline, erect, enveloping cap-

sule, broadly oblong, concave, apex rounded,

margin plane, entire below, toothed at apex,

costa ending below apex, upper laminal cells

rhomboidal, hyaline, thin-walled. Seta very

short. Capsule cylindrical, 1-2 mm long,

abruptly narrowed to a short neck, ribbed;

exothecial cells rectangular to quadrate, thin-

walled to incrassate, smaller at mouth; stoma-

Orthotrichaceae

503

Map 197. — • Stoneobryum mi rum

♦ Ulota ecklonii

ta present at base of urn. Peristome double;

teeth 16, in 8 pairs, triangular, 240-300 pm

long, brownish or yellowish, minutely papil-

lose to striolate-papillose; endostome process-

es narrow, keeled, alternating with teeth pairs,

shorter than teeth, minutely striolate-papillose,

pale yellow, basal membrane absent. Opercu-

lum short-conical. Calyptra 0. 8-1.0 mm long,

hairs ascending, uniseriate, smooth. Spores 12-22

pm. Fig. 140.

Stoneobryum mirum is endemic to southern

Africa and is found on trees in the Midlands and

Drakensberg foothills of KwaZulu-Natal. Map

197.

Vouchers: Hilliard & Burtt 13298A; Magill

5681; Sim 10102, 10103, 10109, PRE-CH7751.

This species is most easily recognized by its

highly differentiated, hyaline perichaetial

leaves, enveloping the almost sessile capsule.

Fig. 140. — Stoneobryum mirum: 1. habit (dry), x 1;

2. habit (wet), x 5; 3. stem in cross section (cells partly

shown), x 130; 4. leaf, x 18; 5. leaf in cross section, x 175;

6. basal leaf cells (right side), x 175; 7. upper laminal cells

at right margin, x 700; 8. leaf apex, x 175; 9. perichaetial

leaf, x 26; 10. perichaetial leaf cells, x 175; 11. part of

capsule wall with stoma, x 350; 12. part of capsule mouth

with peristome (papillae partly shown), x 175; 13. calyp-

tra, x 35; 14. spore, x 700. (1-3, 6, 7, 9, 11 & 13, Magill

5681, 4 & 14, Sim 10109\ 5, 8, 10 & 12, Sim 10102.)

504

Orthotrichaceae

5. ULOTA

Ulota Mohr in Ann. Bot. (Konig & Sims) 2: 540 (1806); Broth, in Natiirl. PflFam., edn 2, 1 1: 24

(1925); Sim, Bryo. S. Afr. 272 (1926); Nyholm, Moss FI. Fenn. 315 (1954); Sainsb., N. Zeal, moss-

es 217 (1955); Lawton, Moss FI. Pacific Northwest 228 (1971); Smith, Moss FI. Brit. Irel. 486

(1987); Crum & Anderson, Moss. E.N. Amer. 2: 720 (1981). Lectotype species: U. crispa (Hedw.)

Brid., vide Gangulee, Moss. E. India 5: 1167 (1976).

Plants small, caespitose; terricolous. Stems erect, central strand absent. Leaves crowded, crisped

when dry, erect-spreading when wet; linear-lanceolate above an oval or ovate base; apex acute to

acuminate; base sheathing; margins generally plane, entire. Costa ending below apex. Upper lami-

naI cells irregularly rounded, incrassate, smooth to weakly papillose; basal cells rectangular or

rhomboidal to vermicular, incrassate, bordered by rectangular to quadrate cells in 4—12 rows, trans-

verse walls incrassate, longitudinal walls thin.

Autoicous. Perigonia terminal on short branches. Perichaetia terminal, leaves weakly differen-

tiated. Seta twisted anticlockwise above. Capsule shortly exserted, oval, contracted below mouth

when dry, ribbed, neck gradually narrowed to seta; stomata present in lower half of urn, phanero-

pore; annulus absent. Peristome double; teeth 16, in 8 pairs; endostome segments linear, alternat-

ing with teeth pairs. Operculum conic-rostellate. Calyptra campanulate, lobed below, hairy. Spores

round, granulose, brownish.

A genus of 50-60 species found mostly as epiphytes in moist temperate forests of the world. In

southern Africa Ulota is most easily recognized by the distinct border of the differentiated leaf

base.

Ulota ecklonii (Hornsch.) Jaeg., Ber.

Thatigk. St. Gallischen Naturwiss. Ges. 1 872—

1873: 163 (1874); Broth, in Natiirl. PflFam.,

edn 2, 11: 25 (1925); Sim, Bryo. S. Afr. 273

(1926). Type: Cape, Table Mountain, Ecklon

s.n. (BM!).

Orthotrichum ecklonii Hornsch. in Linnaea 15: 129

(1841).

Plants small, caespitose, yellow-green

above, brown to reddish brown below; terri-

colous. Stems up to 18 mm tall, branching by

subperichaetial innovations, tomentose below;

rhizoids red-brown to brown, smooth; in sec-

tion round, with inner cortex 4-8 cells across,

incrassate, outer cortical cells smaller, in 1-3

rows, incrassate, outer surface rough. Leaves

crowded, larger above, crisped when dry, erect-

spreading when wet; linear-lanceolate above an

oval or ovate base, ( 1 .3—) 1 .5— 3.0(— 3.7) mm

long; ventral surface keeled; apex acute to

acuminate; base sheathing; margins plane,

occasionally narrowly recurved in midleaf on

one side, entire. Costa ending below apex; ven-

tral superficial cells rectangular to linear; dor-

sal superficial cells oval above, narrowly rec-

tangular to linear below; in section subround to

crescent-shaped, bulging dorsally, ventrally

flat, laminal insertion ventral, cells not differ-

entiated, incrassate to stereids. Upper laminal

cells irregularly rounded, incrassate, ± flat,

smooth to weakly papillose, 10.5-16.0 (-20.0)

pm, papillae low, simple, blunt; basal cells rec-

tangular or rhomboidal to vermicular, incras-

sate, smooth, occasionally pitted, border dis-

tinct, cells in 4-12 rows, short-rectangular to

quadrate, transverse walls incrassate, longitudi-

nal walls thin.

Autoicous. Perigonia terminal on short

branches, inner leaves ovate. Perichaetia ter-

Orthotrichaceae

505

minal, leaves weakly differentiated. Seta

2. 0-2. 5 mm long, brownish, twisted anticlock-

wise above. Capsule shortly exserted, oval,

contracted below mouth when dry, ribbed,

brownish, urn 1.0-1. 2 mm long; neck gradual-

ly narrowed to seta, ± 0.5 mm long; exothecial

cells irregularly rectangular or quadrate,

smaller at mouth, irregularly rounded or oval,

incrassate; stomata present in lower half of

urn; annulus absent. Peristome reflexed when

dry; exostome teeth 16, in 8 pairs, fused below,

perforated above, lanceolate, 270-300 pm,

yellowish brown, minutely papillose to stri-

olate-papillose; endostome segments fragile,

alternating with teeth pairs, linear, hyaline or

yellowish, essentially smooth, basal mem-

brane absent. Operculum 0.5 mm long.

Calyptra 2. 0-2. 4 mm long, lobed below, hairs

ascending, multiseriate, denticulate. Spores

23-35 pm. Fig. 141.

Endemic to southern Africa, Ulota ecklonii

is known from Table Mountain and the

Hottentots Holland Mountains in the Fynbos

Biome of the southwestern Cape region. Map

197.

Vouchers: Thorne PRE-CH3638; Van Zanten

7608185.

The linear-lanceolate leaves above an ovate,

bordered base; rounded, smooth to weakly

papillose laminal cells; shortly exserted, ribbed

capsule with a double peristome; and hairy

calyptra place plants in this species.

Fig. 141. — Ulota ecklonii: 1. habit (dry), x 1; 2. habit

(wet), x 5; 3. stem in cross section, x 175; 4. leaf, x 25;

5. leaf in cross section, x 175; 6. basal leaf cells (left

side), x 350; 7. upper laminal cells at left margin, x 700;

8. leaf apex, x 175; 9. perigonial leaf, x 35; 10.

perichaetial leaf, x 35; 11. capsule, x 18; 12. part of cap-

sule mouth with peristome, x 122; 13. operculum, x 50;

14. calyptra, x 25; 15. spore, x 490. (1 & 2, Thorne PRE-

CH3638-, 3, 4, 7, 8 & 10-14, Thorne SAMH50485\ 5, 6, 9

& 15, Ecklon s.n.)

506

Orthotrichaceae

Subfamily Macromitrioideae

Plants medium-sized to large; saxicolous or corticolous. Primary stems prostrate, with numer-

ous erect or ascending secondary stems and branches. Leaves dimorphic or stem leaves scarcely

differentiated, appressed or variously twisted to spirally twisted or secund when dry, ovate or var-

iously lanceolate or oblong, base scarcely differentiated. Upper laminal cells rounded to rounded-

quadrate or rounded-hexagonal, incrassate, smooth or papillose; basal cells scarcely differentiated

or rectangular. Gemmae absent.

Perichaetia terminal on secondary stems and branches. Dwarf male plants frequently present in

some genera. Capsule exserted, ribbed. Stomata phaneropore. Peristome single or double. Calyptra

mitrate, naked or hairy. Isosporous or anisosporous.

6. MACROCOMA

Macrocoma ( Homsch. ex C. Mull.) Grout in Bryologist 47: 4 (1944); Vitt in Rev. Bryol. Lichenol.

39: 207 (1973); Vitt in Bryologist 83: 407 (1980a); Van Rooy & Van Wyk in Bryologist 95: 206

(1992). Type species: M.fUifortne (Hook. & Grev.) Grout.

Macromitrium sect. Macrocoma Homsch. ex C. Mull, in Bot. Zeitung (Berlin) 3: 522 (1845); Sim, Bryo. S. Afr. 278

( 1926); Catcheside, Moss. S. Austr. 211 ( 1980). Macromitrium subgen. Macrocoma (Homsch. ex C. Mull.) Broth, in Natiirl.

PflFam. 1,3: 477 (1902); Gangulee, Moss. E. India 5: 1171 (1976).

Plants medium-sized, slender, forming mats; corticolous or occasionally saxicolous. Primary

stems creeping, irregularly or subpinnately branched; secondary stems ascending to erect, widely

spaced. Leaves appressed to spirally appressed or rarely erect when dry, spreading to squarrose

when wet; generally ovate-acuminate, lanceolate or lanceolate-ligulate; frequently excavate below,

unistratose or bistratose; apex flat or occasionally cucullate, occasionally fragile; stem leaves

scarcely differentiated. Upper laminal cells oval or rounded-quadrate, smooth above to papillose

below, incrassate, frequently with intercellular spaces; basal cells scarcely differentiated.

Autoicous or dioicous. Perigonia terminal or lateral, leaves broadly ovate-apiculate. Perichaetia

terminal, leaves mostly oblong. Seta straight or twisted counterclockwise, ochrea frequently pre-

sent. Capsule exserted, elliptic or oblong-cylindrical, frequently ribbed; stomata present at base of

urn, phaneropore. Peristome double, exostome parts fused, endostome parts fused. Operculum

conic-subulate. Calyptra mitrate, covering capsule, naked or densely hairy, generally yellow-

brown. Spores round, brownish, granulose; isosporous.

Of the 11 species (one with two subspecies) recognized by Vitt (1980a), four occur in Africa,

three are known from southern Africa and two species are endemic to the Flora area. The genus is

characterized by creeping, slender stems; widely spaced branches with crowded, appressed leaves;

uniformly rounded to oval leaf cells; a large, mitrate, hairy calyptra covering the entire capsule; and

the presence of an ochrea (a collar-like sheath around the base of the seta).

1 Branch leaf apex fragile; leaf cells generally rounded-quadrate, frequently obscure

2. M. lycopodioides

1 Branch leaf apex intact; leaf cells generally oval, clear:

2 Calyptra densely hairy; Cape regions to the northern Transvaal region . . . 3. M. tenue subsp. tenue

2 Calyptra naked or with a few hairs; southwestern Cape 1. M. pulchella

Orthotrichaceae

507

1. Macrocoma pulchella ( Hornsch .) Vitt in

Rev. Bryol. Lichenol. 39: 219 (1973); Van Rooy

& Van Wyk in Bryologist 95: 206 (1992). Type:

Cape, Bergius s.n. (BM, lecto.!) vide Vitt in

Bryologist 83: 425 (1980a).

Schlotheimia pulchella Hornsch., Horae phys. berol. 61

(1820). Macromitrium pulchellum (Hornsch.) Brid. in

Bryol. univ. 1: 313 (1826); Broth, in Natiirl. PflFam., edn 2,

11: 30 (1925); Sim, Bryo. S. Afr. 279 (1926).

Plants yellow-green or olivaceous above,

brown below; corticolous. Primary stem up to

50 mm long, branching irregular; secondary

stem in section with inner cortex 5 or 6 cells

across, incrassate, outer cortical cells in 1 or 2

rows, consisting of substereids or stereids, outer

surface rough. Stem leaves crowded to some-

what distant, frequently reflexed; ovate-acumi-

nate, lanceolate or lanceolate-ligulate, 0.8-1. 1

mm long. Branch leaves crowded, ± equal in

size, appressed to spirally appressed or erect

when dry, occasionally twisted, erect-spreading

to spreading when wet; occasionally reflexed;

lanceolate or oblong, 0.8-1. 8 mm long, ±

keeled, unistratose, occasionally with bistratose

patches; apex acuminate, acute, rounded acute to

obtuse, rarely cucullate; margins plane above,

plane or recurved below, entire. Costa ending

below apex to percurrent or rarely mucronate;

ventral superficial cells linear, smooth; dorsal

superficial cells oval above, linear below,

smooth to bulging; in section crescent-shaped to

subround, flat ventrally, bulging dorsally, guide

cells absent, 2^1 ventral substereids or stereids

present, dorsal surface cells incrassate. Upper

laminal cells oval above, oval to rounded

quadrate below, incrassate, flat to bulging above,

bulging or mammillose below, occasionally with

intercellular spaces, 7.5-14.0 pm; basal cells

frequently longer and narrower towards costa.

Dioicous. Perigonia terminal on branches or

branchlets, leaves broadly ovate to ovate-

acuminate. Perichaetia terminal on branches or

branchlets; leaves oblong, oblong-acuminate,

oblong-lanceolate or oblong-ligulate, 1.0- 1.8

mm long. Seta 3. 2-8.0 mm long, yellowish or

reddish to brown, ochrea weakly developed or

absent. Capsule elliptic or oblong-cylindrical;

urn 0.8-1. 4 mm long, yellowish brown to

Map 198. — ♦ Macrocoma pulchella

• Macromitrium levatum

■ Macromitrium richardii

brown, ribbed; neck 0.2-0. 8 mm long; exothe-

cial cells irregularly rectangular to quadrate,

smaller at mouth, incrassate. Peristome double;

outer layer 7-10 cells high, irregularly split,

striolate-papillose, yellowish brown; inner

membrane frequently plicate, outer surface

smooth, inner surface coarsely papillose, hya-

line. Operculum 0.5-0. 8 mm long. Calyptra

1.5-1. 8 mm long, naked or with a few hairs,

yellow below, red-brown above. Spores 23-28

pm. Fig. 142: 1-16.

The species is endemic to South Africa and

known only from a few collection sites in the

Fynbos Biome of the southwestern Cape region.

Map 198.

Vouchers: Barnard SAM no. 49635; Reh-

mann 164; Wood PRE-CH3401.

Macrocoma pulchella is most easily recog-

nized by the almost naked calyptra and the peri-

stome of two well-developed membranes.

2. Macrocoma lycopodioides (Schwaegr.)

Vitt in Rev. Bryol. Lichenol. 39: 219 (1973);

Van Rooy & Van Wyk in Bryologist 95: 207

(1992). Type: Cape, Knysna, near Kruisvallei,

at Mantis Station, Burchell 5144-7 , upper left

plant (G, lecto.!) vide Vitt in Bryologist 83: 423

(1980a).

508

Orthotrichaceae

Orthotrichaceae

509

Macromitrium lycopodioides Schwaegr. in Sp. muse,

frond, suppl. 2,2: 141 (1827); Broth, in Natiirl. PflFam., edn

2, 11: 30 (1925); Sim, Bryo. S. Afr. 279 (1926). Macro-

mitrium tenue var. lycopodioides (Schwaegr.) C. Mull, in

Bot. Zeitung (Berlin) 3: 522 (1845).

Plants yellowish green or olivaceous above,

brown below; corticolous or occasionally saxi-

colous. Primary stem up to 90 mm long, branch-

es irregular, young stem often subpinnately

branched; secondary stem in section with inner

cortex 5-9 cells across, incrassate, outer cortical

cells in 1-4 rows, consisting of substereids or

stereids, outer surface rough. Stem leaves crowded

to somewhat distant, frequently reflexed; ovate-

acuminate or lanceolate, 0. 4-1.1 mm long.

Branch leaves crowded, ± equal in size, appressed

to spirally appressed when dry, spreading to wide-

ly spreading when wet, occasionally reflexed;

lanceolate, 0.5- 1.0 mm long, keeled to narrowly

keeled above, broadly keeled below; bistratose or

occasionally unistratose with bistratose patches or

unistratose; apex acuminate to acute, occasionally

cucullate, fragile; margins plane above, plane to

recurved below, entire above, entire or occasional-

ly crenulate to denticulate below. Costa ending

below apex to percurrent; ventral superficial cells

rectangular or linear; dorsal superficial cells oval

or rounded-quadrate above, linear below, smooth

to bulging; in section crescent-shaped to sub-

round, flat ventrally, bulging dorsally, guide cells

absent, incrassate or consisting of 2 -4 ventral sub-

stereids or stereids, dorsal surface cells incrassate.

Upper laminal cells rounded-quadrate, frequently

obscure, homogeneous, incrassate, flat to bulging

above, frequently mammillose to papillose below,

intercellular spaces numerous, 7.5-14.0 pm; basal

cells frequently longer and narrower towards

costa.

Dioicous. Perigonia terminal on branches or

branchlets or lateral on short branchlets, leaves

broadly ovate-acuminate to ovate-apiculate.

Perichaetia terminal on branches or branchlets;

leaves oblong-acuminate or oblong-subulate,

1.8-2. 3 mm long. Seta 1. 8^f.O mm long, yel-

lowish or reddish brown to brown, ochrea fre-

quently present. Capsule elliptic or oblong-

cylindrical; urn 1.0-1. 6 mm long, yellowish

brown to brown, smooth, ribbed, neck up to 0.5

mm long; exothecial cells irregularly rounded

to rectangular, smaller at mouth, incrassate.

Peristome double; outer layer low, fused, pale

brown, coarsely papillose; inner membrane low,

outer surface smooth, inner surface coarsely

papillose, hyaline. Operculum 0.7-0. 8 mm

long. Calvptra 2. 2-2. 8 mm long, yellow to yel-

low-brown to brown, densely hairy. Spores

23-35 pm. Fig. 142: 17-24.

Endemic to the Flora area, M. lycopodioides

is known from the southwestern, southern and

eastern Cape regions, the Free State, KwaZulu-

Natal, Zululand, Swaziland, and the eastern,

central and northern Transvaal areas. Map 199.

Vouchers: Crosby & Crosby 7641, 8085;

Magill 3500, 5932, 5995; Smook 1846; Van

Rooy 2287; Von Breitenbach 488.

This species is easily distinguished from the

closely related M. tenue by its fragile leaf apex.

The obscure, rounded-quadrate laminal cells of

the branch leaves with numerous intercellular

spaces, and the oblong-subulate perichaetial

leaves also help to identify M. lycopodioides.

Vitt (1980b) suggests that the fragile leaf apex

and intercellular spaces are caused by the break-

down of components of the middle lamella.

Fig. 142. — Macrocoma pulchella (1-16): 1. habit (dry), x 1; 2. habit (wet), x 7; 3. stem in cross section, x 175; 4. stem

leaf, x 35; 5. branch leaf, x 35; 6. leaf in cross section, x 175; 7. basal leaf cells (right side), x 350; 8. upper laminal cells

(right side), x 350; 9. leaf apex, x 350; 10. perigonial leaf, x 35; 11. perichaetial leaf, x 35; 12. part of capsule wall with

stoma, x 350; 13. part of capsule mouth with peristome, x 175; 14. operculum, x 35; 15. calyptra, x 25; 16. spore, x 700.

M. lycopodioides (17-24): 17. habit (dry), x 1; 18. habit (wet), x 7; 19. stem leaf, x 35; 20. branch leaves, x 35; 21. leaf

in cross section, x 175; 22. basal leaf cells, x 350; 23. upper laminal cells, 350; 24. perichaetial leaf, x 35. M. tenue subsp.

tenue (25-32): 25. habit (dry), x 1 ; 26. habit (wet), x 5; 27. stem leaf, x 35; 28. branch leaf, x 35; 29. leaf in cross section,

x 175; 30. upper laminal cells, x 700; 31. leaf apex, x 350; 32. perichaetial leaf, x 35. (1. 2, 4 & 14, Barnard SAMH49635;

3,6,9, 11. 15 & 16, Wood PRE-CH340P, 5,7, 8, 12 & 13. Rehmann 164- 10, Wood SAMH50327; 17 & 18, Leighton PRE-

CHI 2797: 19 & 20, Von Breitenbach 488: 21. Magill 5932: 22, Magill 3500: 23, Van Rooy 2287: 24. Crosby 7641: 25,

Smook 1413: 26, Von Breitenbach 334a: 27 & 32, Von Breitenbach 198: 28 & 31, Oliver 7665: 29, Van Rooy 798: 30,

Hilliard & Burn 10438.)

510

Orthotrichaceae

3. Macrocoma tenue (Hook. & Grev.) Vitt

subsp. tenue in Rev. Bryol. Lichenol. 39: 217

(1973); Van Rooy & Van Wyk in Bryologist 95:

208 (1992). Type: Cape of Good Hope, Menzies

s.n. (BM, lecto.!) vide Vitt in Bryologist 83: 425

(1980a).

Orthotrichum tenue Hook. & Grev. in Edinburgh J. Sci.

1: 120 (1824). Macromitrium tenue (Hook. & Grev.) Brid.

in Bryol. uni v. 1 : 740 ( 1 826); Broth, in Natiirl. PflFam., edn

2, 11: 30 (1925); Sim, Bryo. S. Afr. 280 (1926); Scott &

Stone, Moss. S. Austr. 232 (1976); Catcheside, Moss. S.

Austr. 212 (1980).

Orthotrichum microphyllum Hook. & Grev. in

Edinburgh J. Sci. 1: 121 (1824). Macromitrium microphyl-

lum (Hook. & Grev.) Brid. in Bryol. univ. I: 737 (1826).

Type: Cape, Burchell s.n. (E, lecto.!) vide Vitt in Bryologist

83: 427 (1980a).

Maschalocarpus ecklonii Spreng. in Syst. veg. 4,2: 321

(1827). Type: Cape, Table Mountain, Ecklon s.n. (H, lecto.!)

vide Vitt in Bryologist 83: 428 (1980a).

Macromitrium dregei Homsch. in Linnaea 15: 131

(1841). Macromitrium tenue var. dregei (Homsch.) C. Mull,

in Bot. Zeitung (Berlin) 3: 522 (1845). Type: Cape, Drege

s.n. (BM, lecto.!) vide Vitt in Bryologist 83: 428 (1980a).

Macromitrium confusum Mitt, in J. Linn. Soc., Bot. 22:

305 (1886). Type: Cape, East London, Capt. Rooper s.n.

(NY, lecto.!) vide Vitt in Bryologist 83: 428 (1980a).

Macromitrium dawsoniaemitrium C. Mull, in Hedwigia

38: 116 (1899). Type: Cape, in sylvis Knysna, Rehmann

160 , (H, lecto.!) vide Vitt in Bryologist 83: 428 (1980a).

Leiomitrium capense Broth, in Denkschr. Kaiserl. Akad.

Wiss., Math.-Naturwiss. Kl. 88: 736 (1913). Coleochaetium

capense (Broth.) Broth, in Natiirl. PflFam., edn 2, 1 1 : 26

(1925). Type: Cape, slope of Table Mountain, 1909,

Brunnthaler s.n. (H, holo.!) vide Vitt in J. Hattori Bot. Lab.

49: 110(1981).

Plants yellowish green to dark green or oli-

vaceous above, dark green or brown below; cor-

ticolous, occasionally saxicolous. Primary stem

up to 150 mm long, branches irregular; sec-

ondary stem in section with inner cortex 6-10

cells across, incrassate, outer cortical cells in

2-4 rows, incrassate or consisting of substerei-

ds or stereids, outer surface rough. Stem leaves

crowded to somewhat distant, frequently re-

flexed; ovate-acuminate to lanceolate or lance-

olate-ligulate, 0.5-1. 7 mm long. Branch leaves

crowded, ± equal in size, appressed to spirally

appressed when dry, erect-spreading to widely

spreading when wet, occasionally reflexed, in-

Map 199. — Macrocoma lycopodioides

frequently falcate; lanceolate, lanceolate-ligulate

or infrequently lanceolate-subulate, 0.5- 1.6

mm long, frequently narrowly keeled above,

broadly keeled below, unistratose, frequently

with bistratose patches above; apex acuminate,

acute or rounded acute, occasionally cucullate;

margins plane above, plane to recurved below,

entire above, entire below or denticulate at base.

Costa ending below to percurrent or infrequent-

ly mucronate; ventral superficial cells narrowly

rectangular or linear; dorsal superficial cells

oval above, linear below, smooth to bulging; in

section crescent-shaped to subround, flat ven-

trally, bulging dorsally, guide cells absent, 2^1

ventral substereids, stereids or incrassate cells

present, dorsal surface cells incrassate. Upper

laminal cells oval to rounded, homogeneous,

incrassate, flat to bulging above, frequently

mammillose to papillose below, frequently

with intercellular spaces below, 6-18 pm; basal

cells frequently longer and narrower towards

costa.

Autoicous. Perigonia terminal on branches

or branchlets, leaves broadly ovate-apiculate.

Perichaetia terminal on branches or branchlets;

leaves oblong-ligulate, oblong-lanceolate or

lanceolate, infrequently falcate, 1.3-2. 3 mm

long. Seta 2. 5-7. 5 mm long, yellowish brown or

reddish brown to brown, ochrea frequently pre-

sent. Capsule elliptic or oblong-cylindrical; urn

1-2 mm long, yellowish brown or reddish

Orthotrichaceae

511

brown to brown, smooth to ribbed; neck 0.2-0.6

mm long; exothecial cells irregularly oval to

rectangular, smaller at mouth, incrassate.

Peristome double; outer layer 2-5 cells high,

fused, coarsely papillose, pale brown; inner

membrane low, outer surface smooth, inner sur-

face coarsely papillose, hyaline. Operculum

0.4- 1.0 mm long. Calyptra 2-\ mm long, yel-

low to yellowish brown to brown, densely hairy.

Spores ( 1 8— )27— 37( — 43) pm. Fig. 142: 25-32.

This subspecies is known from central, east-

ern and southern Africa, Reunion, Madagascar,

St. Helena, Australia, Tasmania and New Zea-

land. In the Flora area it is found on trees and

occasionally rocks in the Fynbos Biome of the

southern and southwestern Cape and montane

forest and grassland of the central and eastern

Cape regions, the Free State, Lesotho, KwaZulu-

Natal, Zululand, Swaziland and the eastern, cen-

tral and northern Transvaal areas. Map 200.

Vouchers: Hilliard & Burtt 14087; Kemp

855; Magill 4557, 5701; Smook 1672; Stirton

8132; Van Rooy 801, 1414, 1895; Von Breiten-

bach 198.

Macrocoma tenue is recognized by the

densely hairy calyptra, slender branches with

appressed leaves, intact leaf apex, small laminal

cells which are oval with incrassate walls, and

the low exostome and endostome membranes.

The autoicous sexual condition of M. tenue

subsp. tenue also helps to distinguish it from M.

pulchella and M. lycopodioides. Vitt (1980c)

has shown that M. tenue can be divided into two

subspecies and southern African plants belong

to subspecies tenue.

7. MACROMITRIUM

Macromitrium Brid., Muscol. recent, suppl. 4: 132 (1818); Broth, in Natiirl. PflFam., edn 2, 11;

28 (1925); Sim, Bryo. S. Afr. 278 (1926) p.p.; Van Rooy & Van Wyk in Bryologist 95: 208 (1992).

Type species: M. pallidum (R Beauv.) Wijk & Marg. vide Vitt in J. Hattori Bot. Lab. 54: 5 (1983).

Plants medium-sized to large, forming mats; corticolous or saxicolous. Primary stems creeping;

secondary stems erect, crowded, bushy. Leaves secund or variously twisted when dry; narrowly

oblong-lanceolate or ligulate; unistratose, occasionally with bistratose patches or irregularly uni-

stratose to multistratose; apex obtuse, acute or acuminate, infrequently fragile; margins plane or

recurved on one side, entire or crenulate or denticulate, prorate or tuberculate; stem leaves small-

er, generally lanceolate. Costa strong, single. Upper laminal cells rounded, incrassate, flat to mam-

millose, smooth or multipapillose; basal laminal cells rectangular, occasionally sinuate, longitudi-

nal walls strongly incrassate, frequently tuberculate.

Autoicous or pseudautoicous, dwarf male plants axillary on secondary stem or on leaf lamina.

Perichaetia terminal, overgrown by subperichaetial innovations; leaves oblong-lanceolate or

oblong-subulate, unistratose. Seta occasionally twisted clockwise above. Capsule exserted, erect,

ovate-cylindrical, smooth or ribbed; stomata present on neck, phaneropore. Peristome single or

double; exostome teeth 16, separate or fused, segments and cilia absent, basal membrane absent or

512

Orthotrichaceae

high. Operculum conic-rostrate. Calyptra large, mitrate, laciniate to lacerate below, plicate, naked

or with a few hairs. Spores round, brownish, anisosporous or isosporous.

A large tropical to temperate genus of approximately 370 species, most of them occurring in

Oceania, southeastern Asia, Australasia and tropical South America. The genus is absent from

Europe and large parts of Asia and North America.

Members of the genus can be recognized by: 1. creeping primary stem with numerous erect

bushy branches; 2. differentiated stem leaves; 3. branch leaves that are variously twisted when dry;

4. small, rounded upper laminal cells; 5. differentiated perichaetial leaves; 6. erect, frequently pli-

cate capsules; 7. reduced peristomes; 8. sexual condition that is frequently pseudautoicous and

plants that are frequently anisosporous; and 9. large, mitrate, plicate, laciniate to lacerate calyptrae.

1 Branch leaves 2.5-5 .0 mm long; margins irregularly denticulate towards apex; peristome

double 1 . M. levatum

1 Branch leaves 1. 0-3.0 mm long; margins entire, weakly prorate or crenulate-papillose

towards apex; peristome single:

2 Branch leaves with upper laminae irregularly unistratose to multistratose; apex fragile . .

5. M. serpens

2 Branch leaves with upper laminae unistratose, occasionally with bistratose patches; apex

intact:

3 Branch leaf apex acute to acuminate; basal cells reaching midleaf; upper cells flat to

bulging, smooth 2. M. macropelma

3 Branch leaf apex broadly acute to rounded-obtuse; basal cells restricted to lower quar-

ter to third of leaf; upper cells strongly bulging, multipapillose:

4 Branch leaves frequently with bistratose patches; perichaetial leaves longer than

branch leaves; seta 1.5-3. 5 mm long; pseudautoicous, anisosporous .... 4. M. lebomboense

4 Branch leaves unistratose; perichaetial leaves shorter or as long as branch leaves; seta

4.5-12.0 mm long; autoicous, isosporous 3. M. richardii

1. Macromitrium levatum Mitt, in J. Linn.

Soc., Bot. 7: 152 (1863); Broth, in Natiirl.

PflFam., edn 2, 11: 43 (1925); Van Rooy & Van

Wyk in Bryologist 95: 208 (1992). Type:

Cameroon, Cameroon Mountain, 8 000-10 000 ft,

on trees and rocks, Mann s.n. (NY-Mitt., holo.!).

Macromitrium mannii sensu Sim, Bryo. S. Afr. 281

(1926).

Plants medium-sized to large, forming mats,

green, yellowish green, brown-green, or yel-

low-brown above, brown below; saxicolous or

corticolous. Primary stem up to 140 mm long,

branching regularly, tomentose below; rhizoids

smooth, red-brown; secondary stem erect, 2^40

mm tall, branching by subperichaetial innova-

tions; in section inner cortex 10-16 cells across,

incrassate, outer cortical cells smaller, in 2-5

rows, incrassate, substereids or stereids, outer

surface rough. Stem leaves crowded to ± distant.

fragile on old stems; ovate-acuminate, ovate-

lanceolate or ovate-subulate, 1. 0-2.4 mm long.

Branch leaves crowded, erect, spirally twisted

to contorted when dry, erect-spreading, fre-

quently reflexed, flexuose when wet, rugose,

unistratose; narrowly lanceolate, narrowly

oblong-lanceolate or narrowly oblong-subulate,

2. 1-4. 8 mm long, keeled; apex acute to acumi-

nate; margins plane, occasionally reflexed

below, irregularly denticulate above, plane or

prorate to tuberculate below. Costa ending

below apex to mucronate; ventral and dorsal

superficial cells linear, smooth; in section cres-

cent-shaped to subround, bulging dorsally,

guide cells incrassate or substereids, dorsal

stereids present. Upper laminal cells rounded or

rounded-hexagonal, incrassate, bulging, fre-

quently in conspicuous longitudinal rows,

7.5-18.0 pm, mammillose to strongly mammil-

lose; basal cells long-rectangular, longitudinal

Orthotrichaceae

513

walls irregularly incrassate, pitted, tuberculate,

marginal cells frequently thinner walled, inflat-

ed, smooth or prorate to tuberculate.

Pseudautoicous; dwarf male plants rare, on

branch leaf lamina. Perichaetia terminal, fre-

quently overgrown by subperichaetial innova-

tions; leaves oblong-lanceolate or oblong-subu-

late, 3.2— 4.4 mm long. Seta 4.5-14.0 mm long,

yellowish brown or reddish brown to brown.

Capsule reddish brown to brown, ribbed; urn

ovate-cylindrical, 1.0-1. 6 mm long; neck to 0.5

mm long; exothecial cells irregular in shape,

smaller at mouth, incrassate; annulus decidu-

ous, cells subrectangular. Peristome double,

erect when dry; exostome teeth fused, blunt,

frequently fragile above, 170-225 pm long, yel-

low-brown to brown, densely papillose-striolate;

basal membrane high, hyaline, papillose, seg-

ments and cilia absent. Operculum 1 .0—1 .2 mm

long. Calyptra 3. 0-3. 6 mm long, lacerate to

base of rostrum, essentially naked, yellow to

yellowish brown or reddish brown. Spores

17.0-36.5 pm, minutely papillose; aniso-

sporous. Fig. 143: 1-13.

Macromitrium levatum is widespread in

Africa south of the Sahara, and is also found on

the Comoros and Madagascar. In the Flora area

it is found on stems and branches of trees and

shrubs and on rock in montane forests and

wooded areas of the eastern Cape, KwaZulu-

Natal and the eastern and northern Transvaal

regions. Map 198.

Vouchers: Crosby & Crosby 9216: Ester-

huysen 20218; Magill 5512, 6566: Rankin 148;

Smook & Phelan 876; Van Rooy 1417, 1604,

2299.

The species is most easily identified by the

large plants with long, spirally twisted and con-

torted branch leaves. The unistratose branch

leaves with denticulate upper margins, bulging

upper laminal cells arranged in conspicuous

longitudinal rows, tuberculate basal laminal

cells, calyptra that is lacerated to the base of the

rostrum, and the double peristome also help to

place plants of M. levatum.

Tixier (1989) placed this species in syn-

onymy under the Asian Macromitrium sulcatum

(Hook.) Brid. but until the type specimens are

compared, M. levatum is here maintained as a

distinct species.

2. Macromitrium macropelma C. Miill.

in Bot. Zeitung (Berlin) 14: 420 (1856); Broth,

in Natiirl. PflFam., edn 2, 11: 34 (1925); Sim,

Bryo. S. Afr. 283 (1926). Lectotype: Cape,

Grootvadersbosch, Ecklon (PRE!) fide Van

Rooy & Van Wyk in Bryologist 95: 210 (1992).

Plants medium-sized to large, forming mats,

yellow-green above, brown below; corticolous.

Primary stem up to 60 mm long, branching

regularly, tomentose below; rhizoids smooth,

reddish brown; secondary stem 4—14 mm tall,

branching by subperichaetial innovations; in

section inner cortex 10-12 cells across, incras-

sate, outer cortical cells smaller, in 1-3 rows,

stereids, outer surface rough. Stem leaves ± dis-

tant, ovate-acuminate or ovate-subulate or

shortly oblong-subulate, 1.0-1. 7 mm long.

Branch leaves crowded, erect-twisted to spiral-

ly twisted when dry, erect to erect-spreading

when wet, ± rugose, unistratose; narrowly

lanceolate, narrowly oblong-lanceolate or nar-

rowly oblong-subulate, 1.5-2. 3 mm long,

keeled; apex acute to acuminate; margins plane,

occasionally recurved on one side of leaf, entire

to weakly prorate. Costa ending below apex to

mucronate; dorsal and ventral superficial cells

linear, smooth; in section subround to round,

bulging dorsally, cells undifferentiated, stereids.

Upper laminal cells irregularly rounded to oval,

incrassate, flat to bulging, 7.5-18.0 pm,

smooth; basal cells long-rectangular to linear,

longitudinal walls strongly incrassate, smooth.

Pseudautoicous? Perigonia not seen. Peri-

chaetia terminal, frequently overgrown by sub-

perichaetial innovations; leaves lanceolate,

oblong-lanceolate, or oblong-acuminate, 2.2- 2.8

mm long. Seta 8-13 mm long, reddish brown to

brown. Capsule reddish brown to brown, mouth

darker, weakly ribbed above; urn ovate-cylindri-

cal, 1.0-1. 2 mm long; neck up to 0.4 mm long,

wrinkled when dry; exothecial cells irregularly

quadrate to rectangular, smaller at mouth, incras-

sate. Peristome single; exostome teeth 16, loose-

514

Orthotrichaceae

Fig. 143. — Macromitrium levatum (1-13): 1. habit

(dry), x 1; 2. habit (wet), x 5; 3. branch in cross section, x

175; 4. stem leaf, x 35; 5. branch leaf, x 35; 6. branch leaf

cross section, x 175; 7. basal cells of branch leaf, x 350; 8.

upper laminal cells of branch leaf, x 350; 9. branch leaf apex,

x 350; 10. perichaetial leaf, x 35; 11. sporophyte, x 5; 12. part

of capsule mouth, x 175; 13. spores, x 700. M. richardii

(14-25): 14. habit (dry), x 1; 15. habit (wet), x 5; 16. stem

leaf, x 35; 17. branch leaf, x 35; 18. branch leaf in cross sec-

tion, x 175; 19. basal cells of branch leaf, x 350; 20. upper

laminal cells of branch leaf, x 350; 21. cells of branch leaf

apex, x 350; 22. perichaetial leaf, x 35; 23. part of capsule

mouth with peristome, x 175; 24. calyptra, x 18; 25. spore, x

700. (1, Smook & Phelan 876; 2, Magill 5598 ; 3, Sim PRE-

CH9021; 4, Vorster 587; 5-7, Crosby 9216; 8, Vorster 500: 9,

Von Breitenbach 447; 10-13, Esterhuysen 20218; 14, 15, 17 & 23, Magill 6085; 16, 18 & 22, Magill 6074; 19-21, Wagener

PRE-CHI 3349; 24 & 25, Thorne SAMH49510.)

Orthotrichaceae

515

ly in 8 pairs, narrowly oblong, blunt, frequently

irregular in outline, (85-) 120-190 pm long, pale

brown, finely papillose. Operculum 0.8 mm

long. Calyptra mitrate, 2.2-2. 5 mm long, lacini-

ate below, essentially naked, yellowish brown.

Spores 25^-8 pm, minutely granulose; aniso-

sporous. Fig. 144: 1-14.

Endemic to southern Africa, Macromitrium

macropelma is found on trees or rock in a few

localities in the southern and southwestern

Cape. Map 201.

Vouchers: Barnard SAM no. 49291; Reh-

mann 167; Van Zanten 7609424.

The slender, unistratose branch leaves with

smooth, flat to bulging and strongly incrassate

upper laminal cells and smooth basal laminal

cells separate specimens of M. macropelma

from the other species of Macromitrium in the

Flora area. Dwarf male plants were not found

but the presence of anisomorphic spores sug-

gests a pseudautoicous sexual condition.

3. Macromitrium richardii Schwaegr.,

Sp. muse, frond, suppl. 2,2: 188 (1827); Broth,

in Natiirl. PflFam., edn 2, 11: 34 (1925); Crum

& Anderson, Moss. E.N. Amer. 2: 736 (1981);

Van Rooy in J. Bryol. 16: 213 (1990). Type:

Guiana, Richard s.n. (G, holo.).

Plants medium-sized, forming mats, oliva-

ceous or yellowish green above, reddish brown

to brown below; corticolous. Primary stem

tomentose below; rhizoids smooth, reddish

brown; secondary stems numerous, 2-10 mm

tall, branching by subperichaetial innovations;

in section inner cortex 9-11 cells across, incras-

sate, outer cortical cells smaller, in 1 or 2 rows,

consisting of substereids or stereids, outer sur-

face rough. Stem leaves secund, ovate-lanceo-

late or lanceolate, 0.75-1.4 mm long. Branch

leaves ± crowded, tightly inrolled, twisted when

dry, erect-spreading, incurved when wet, occa-

sionally rugulose, unistratose; narrowly lanceo-

late or ligulate, (1. 4-) 1.6-2. 8 mm long, ventral

surface keeled; apex acute, obtuse or

mucronate, weakly cucullate; margins plane or

recurved on one side, crenulate. Costa ending

Map 201. — ♦ Macromitrium macropelma

• Macromitrium lebomboense

below apex to subpercurrent or occasionally

mucronate, superficial cells linear; in section

round, bulging dorsally, consisting of subster-

eids or stereids. Upper laminal cells rounded-

hexagonal to rounded, incrassate, bulging,

(8.7— ) 1 0.0— 1 5.0 pm, smooth to multipapillose,

papillae low, blunt; basal cells smooth, rarely

papillose below, elliptical to rectangular, incras-

sate, frequently pitted.

Autoicous. Perigonia on short or long

branches. Perichaetia terminal; leaves oblong-

lanceolate or lanceolate, as long as branch

leaves. Seta 4.5-7.5(-12.0) mm long, yellow-

ish or reddish brown. Capsule ovate-cylindri-

cal, 1.2-1. 8 mm long, reddish brown, weakly

to strongly 8-ribbed; mouth contracted, 8-

ribbed; neck differentiated; exothecial cells

irregularly rounded-quadrate to rectangular,

incrassate, smaller at mouth. Peristome single,

reduced, inserted below mouth; teeth 16,

oblong, blunt, 55-150 pm long, yellowish

brown or pale yellow, striolate papillose.

Operculum 0.8 mm long. Calyptra naked or

sparsely hairy, lacerate below, 2. 0-2. 3 mm

long. Spores 21-34 pm, densely papillose;

isosporous. Fig. 143: 14-25.

This African-Neotropical disjunct is known

from North, Central and South America, and the

Caribbean. In Africa, Macromitrium richardii is

516

ORTHOTRICHACEAE

Orthotrichaceae

517

found on tree branches in lowland forests and

wooded areas of the southern Cape and

Zululand. Map 198.

Vouchers: Magill 5363, 6035a, 6074; Thome

SAM No. 49510; Wagener PRE-CHI 3349.

This species can be identified by the tightly

inrolled, ligulate-lanceolate branch leaves with

acute to obtuse apices, strongly bulging, gener-

ally multipapillose laminal cells, and the 8-

ribbed capsule with a contracted mouth.

Compared with the closely related M. lebom-

boense, this species has narrower branch leaves,

shorter perichaetial leaves and calyptra, a

longer seta, an autoicous sexual condition and

isomorphic spores.

4. Macromitrium lebomboense Van Rooy

in J. Bryol. 16: 209 (1990). Type: Natal, Le-

bombo Mountains, near Nambulugwana, Van

Rooy 227 (PRE, holo.!).

Plants medium-sized, forming mats, yellow-

ish green or greenish brown above to brown

below; corticolous or rarely saxicolous. Pri-

mary stem tomentose below; rhizoids smooth,

reddish brown to brown; secondary stems

numerous, 2-8 mm tall, branching by sub-

perichaetial innovations; in section inner cortex

9-13 cells across, incrassate, outer cortical cells

smaller, in \-4 rows, consisting of stereids,

outer surface rough. Stem leaves secund, lanceo-

late, 0.8-1. 5(-l. 8) mm long. Branch leaves

crowded, tightly inrolled, twisted when dry,

erect-spreading when wet, rugulose below,

unistratose or frequently with bistratose patch-

es; generally ligulate, (1 .2— )1 .6— 2.3(— 2.5) mm

long; ventral surface keeled; apex rounded-

obtuse; margins plane, frequently recurved on

one side. Costa ending below to percurrent;

superficial cells linear, bulging; in section sub-

round, bulging dorsally, consisting of stereids.

Upper laminal cells small, rounded-hexagonal

to rounded-quadrate, incrassate, bulging, 7.5-

10.0 pm, papillae low, blunt, scattered over

lumen; basal cells smooth, rarely papillose

below, rectangular, longitudinal walls incras-

sate, frequently sinuate.

Pseudautoicous; dwarf male plants axillary,

rarely on leaf base, 0.5-1 .2 mm long, branching

by subperigonial innovations. Perichaetia ter-

minal; leaves erect, longer than branch leaves,

broadly lanceolate or oblong-lanceolate, (2.0-)

2.2— 2.8(— 3. 1 ) mm long. Seta (1.7-)2.2-2.8

(-3.1) mm long, yellowish brown or reddish

brown. Capsule ovate-cylindrical, (1.2— )1.5— 1.8

(-2.0) mm long, yellowish brown or reddish

brown, weakly ribbed dry, mouth erect, neck

differentiated; exothecial cells irregularly qua-

drate to rectangular, incrassate, smaller at

mouth. Peristome single, inserted below mouth;

teeth 16, occasionally in pairs, narrowly oblong,

blunt, 155-250 pm long, yellowish, papillose-

striolate. Operculum 1.0-1. 3 mm long.

Calyptra 2. 5-3. 5 mm long, deeply plicate, lac-

erate below, naked or sparsely hairy. Spores

12.0-30.0(-36.0) pm, minutely papillose; aniso-

sporous. Fig. 144: 23-31.

Macromitrium lebomboense is endemic to

southern Africa and is found on tree trunks and

branches and rarely on rock in lowland forests

and wooded streams of Zululand, KwaZulu-Natal,

and the eastern Cape region. Map 201 .

Fig. 144. — Macromitrium macropelma (1-14): 1. habit (dry), x 1; 2. habit (wet), x 4; 3. part of branch in cross sec-

tion, x 175; 4. stem leaf, x 35; 5. branch leaf, x 35; 6. branch leaf in cross section, x 175; 7. basal cells of branch leaf, x

350; 8. upper laminal cells of branch leaf, x 350; 9. branch leaf apex, x 350; 10. perichaetial leaf, x 35; 11. part of capsule

wall with stoma, x 350; 12. part of capsule mouth with peristome teeth, x 175; 13. calyptra, x 18; 14. spore, x 700. M. ser-

pens (15-22): 15. habit (dry), x 1; 16. habit (wet), x 4; 17. stem leaf, x 35; 18. branch leaf, x 35; 19. branch leaf in cross

section, x 175; 20. basal cells of branch leaf, x 350; 21. dwarf male plant at base of branch leaf, x 35; 22. Spores (papillae

partly shown), x 700. M. lebomboense (23-31): 23. habit (dry), x 1; 24. habit (wet), x 5; 25. branch in cross section (cells

partly shown), x 130; 26. stem leaf, x 35; 27. branch leaf, x 35; 28. branch leaf in cross section, x 175; 29. upper laminal

cells of branch leaf at left margin, x 700; 30. perichaetial leaf, x 35; 31. Spores (papillae partly shown), x 700. (1-3 & 5-14,

Barnard SAMH49291 ; 4, Sim PRE-CH9017; 15, Schelpe 7886 ; 16, Van Zinderen Bakker 264 ; 17 & 20, Oliver 7056; 18,

PRE-CH10726 ; 19, Jacot Guillarmod 6164a; 21, Rehmann PRE-CH5717; 22, Rehmann 157; 23, 24 & 30, Van Rooy 227;

25, Bosman PRE-CH1576; 26, Wager PRE-CH12021; 27, Van Rooy 196; 29 & 31, Smook 1523.)

518

Orthotrichaceae

Vouchers: Bosnian PRE-CH1576; Linder

1191; Magill 5321; Smook 1523; Van Rooy 948,

1714.

Related to Macromitrium richardii (see p.

515), M. lebomboense can be recognized by the

perichaetial leaves that are longer than the

branch leaves, short seta, weakly ribbed capsule

with erect mouth, well-developed exostome of

16 teeth, deeply plicate, lacerate, naked to

sparsely hairy calyptra, and anisomorphic

spores with correlated pseudautoicous sexual

condition.

5. Macromitrium serpens (Hook. &

Grev.) Brid. in Bryol. univ. 1: 736 (1826);

Broth, in Natiirl. PflFam., edn 2, 11: 35 (1925);

Sim, Bryo. S. Afr. 282 (1926). Lectotype: Cape,

Burchell s.n. (E! ; BM, isolecto.!)/i7/e Van Rooy

& Van Wyk in Bryologist 95: 210 (1992).

Orthotrichum serpens Bruch ex Hook. & Grev. in

Edinburgh J. Sci. 1: 119 (1824).

Macromitrium tristratosum Dix. in Kongel. Norske

Vidensk. Selsk. Skr. (Trondheim) 1932, 4: 10 (1932).

Lectotype: Natal, Zululand, Eshowe, indigenous forest,

Hoeg 121 (BM!) fide Van Rooy & Van Wyk in Bryologist

95:210(1992).

Plants medium-sized to large, forming mats,

yellowish green, green or greenish brown above,

greenish brown to brown below; corticolous.

Primary stem up to 100 mm long, branching reg-

ularly, tomentose below; rhizoids smooth, red-

dish brown; secondary stem 2-10 mm tall,

branching by subperichaetial innovations; in sec-

tion inner cortex 10-15 cells across, incrassate,

outer cortical cells smaller, in 1-3 rows, consist-

ing of substereids to stereids, outer surface

rough. Stem leaves crowded to ± distant, fragile

on old stems; ovate-acuminate or ovate-subulate,

1.0-2. 3 mm long. Branch leaves crowded, erect,

curved to twisted, inrolled when dry, erect-

spreading, indexed when wet, ± rugose, irregu-

larly unistratose to multistratose; narrowly ovate-

lanceolate, narrowly lanceolate or narrowly

lanceolate-subulate, 1. 7-3.0 mm long, keeled;

apex acuminate, fragile; margins plane, entire to

crenulate-papillose above, entire below. Costa

ending below apex to percurrent, ventral and dor-

sal superficial cells linear, smooth; in section

subround to round, bulging dorsally, guide cells

frequently differentiated, incrassate, dorsal

stereids present. Upper laminal cells rounded,

incrassate, bulging, 8.7-15.0 pm, multipapillose,

papillae small, low; basal cells rectangular, lon-

gitudinal walls incrassate, straight to sinuate, fre-

quently unipapillose or tuberculate.

Pseudautoicous; dwarf male plants on sec-

ondary stem, leaves ovate or ovate-apiculate.

Perichaetia terminal, overgrown by subperi-

chaetial innovations; leaves oblong-lanceolate,

1.6-2. 7 mm long, unistratose. Seta 3. 5-6.0 mm

long, reddish brown. Capsule reddish brown,

mouth darker, smooth to weakly ribbed above;

urn ovate-cylindrical, 0.8-1. 4 mm long; neck

up to 0.4 mm long, wrinkled dry; exothecial

cells irregularly quadrate to rectangular, smaller

at mouth, incrassate. Peristome single; exos-

tome teeth 16, narrowly oblong, blunt, fre-

quently irregular in outline, ( 1 25—) 1 50—225 pm

long, pale yellow or yellowish brown, hyaline

above, erect to incurved dry, striolate-papillose.

Operculum 0. 8-1.0 mm long. Calyptra 2. 8-3. 2

mm long, lacerate below, plicate, essentially

naked, yellow to yellowish brown below, yel-

lowish brown or reddish brown above. Spores

16-38 pm, minutely papillose; anisoporous.

Fig. 144: 15-22.

Macromitrium serpens is known from east-

ern and southern Africa, Madagascar and the

Mascarenes. In the Flora area it is found on

Map 202. — Macromitrium serpens

Orthotrichaceae

519

stems and branches of trees and rarely on rock

in indigenous forests of the southern and eastern

Cape regions, KwaZulu-Natal Midlands and

Zululand. Map 202.

Vouchers: Crosby & Crosby 8060; Magill 5166,

6035; Oliver, Balsinhas & Vorster 7056; Russell

2689; Schelpe 7886; Van Rooy 888, 2066.

The species is recognized by its fragile leaf

apex, sinuate basal laminal cells and papillose,

irregularly unistratose to multistratose upper

leaf lamina. The structure of the upper leaf

lamina is unusual in that the cells are displaced

in dorsal and ventral ‘pillars’. The laminal cells

are also arranged in longitudinal rows, giving

the leaf a lamellate appearance.

8. SCHLOTHEIMIA

Schlotheimia Brid., Muscol. recent, suppl. 2: 16 (1812); Broth, in Natiirl. PflFam., edn 2, 11: 46

(1925); Sim, Bryo. S. Afr. 284 (1926); Bartram, Mosses of the Philippines 184 (1939); Sainsb., N.

Zeal, mosses 238 (1955); Gangulee, Moss. E. India 5: 1190 (1976); Crum & Anderson, Moss. E.N.

Amer. 2: 741 (1981); Van Rooy & Van Wyk in Bryologist 95: 210 (1992). Lectotype species: S.

torquata (Hedw.) Brid .fide E.G. Britton, Bahama Flora (1920).

Plants medium-sized to large, forming mats or cushions, greenish or olivaceous or brownish

green above, orange-brown or reddish brown below; saxicolous or corticolous. Primary stem pros-

trate; secondary stems erect, crowded, bushy. Leaves crowded, appressed and twisted to spirally

twisted around stem when dry, erect-spreading to squarrose when wet, rugose; oblong, oblong-lin-

gulate or oblong-lanceolate; apex acute, subacute, rounded-obtuse or cuspidate; stem leaves small-

er, shortly oblong or ovate to lanceolate. Costa ending below apex, mucronate or cuspidate. Upper

laminal cells rounded-hexagonal, incrassate, ± flat, smooth; basal cells rhomboidal to rectangular,

incrassate, pitted.

Pseudautoicous or dioicous, dwarf male plants axillary on secondary stems. Perigonia terminal.

Perichaetia terminal; leaves oblong, oblong-lanceolate or lanceolate. Seta twisted clockwise

above. Capsule exserted, erect, ovoid or oblong-cylindrical, weakly to strongly ribbed; stomata on

neck and base of urn, phaneropore. Peristome double, well-developed; exostome teeth 16, consist-

ing of 2 divisions; endostome segments 16-32, narrow, alternating with exostome teeth, shorter

than teeth, basal membrane low. Operculum conic-rostrate. Calyptra large, completely covering

capsule, campanulate, lobed below, naked. Spores round, brownish; isosporous or anisosporous.

A genus of ± 130 species mostly found in tropical and subtropical regions. The tropical forests

of South America are the major centre of described species, 28 species are known from Madagascar

and 19 from Africa. Schlotheimia is recognized by the rusty or chestnut-green and brown colour of

the plants; prostrate stems with erect, bushy branches; differentiated stem leaves; crowded, rugose

branch leaves that are twisted to spirally twisted around the branch when dry; the large, naked, bell-

shaped calyptra, lobed below and covering the whole capsule; and the well-developed peristome.

1 Stem leaf costa aristate; dioicous; isosporous 3. S. rufopallens

1 Stem leaf costa ending below the apex, mucronate or short-excurrent; pseudautoicous;

anisosporous:

2 Branch leaves broad towards apex; apex acute to rounded-obtuse; costa mucronate ....

1.5. ferruginea

2 Branch leaves gradually narrowed to the cuspidate apex; costa ending below the apex or

exserted as a short point 2. 5. percuspidata

520

Orthotrjchaceae

Orthotrichaceae

521

1. Schlotheimia ferruginea (Bruch ex

Hook. & Grev.) Brid. in Bryol. univ. 1: 743

(1826); Broth, in Natiirl. PflFam., edn 2, 11: 48

(1925); Sim, Bryo. S. Afr. 286 (1926). Lectotype:

Cape, Burchell s.n. (E!; BM, isolecto. ! ) fide Van

Rooy & Van Wyk in Bryologist 95: 212 (1992).

Orthotrichum ferrugineum Bruch ex Hook. & Grev. in

Edinburgh J. Sci. 1: 118 (1824 ). Macromitrium ferrugineum

(Bruch ex Hook. & Grev.) C. Mull, in Bot. Zeitung (Berlin)

3: 544 (1845).

Schlotheimia subventrosa Broth. & Bryhn in Forh.

Vidensk.-Selsk. Kristiania 4: 13 (1911). Type: Zululand,

Eshowe, Aug. 1909, H. Bryhn s.n. (H, holo.!).

Plants medium-sized to large, forming mats

or cushions, green to brownish green above,

brown, orange-brown or reddish brown below;

saxicolous or corticolous. Primary stem up to

120 mm long, branching regularly, frequently

tomentose; rhizoids essentially smooth, brown

to reddish brown; secondary stem up to 35 mm

tall, branching by subperichaetial innovations;

in section inner cortex 7-10 cells across, incras-

sate, outer cortical cells smaller, in 1-3 rows,

consisting of substereids to stereids, outer sur-

face occasionally rough. Stem leaves crowded

to ± distant; ovate to lanceolate or shortly

oblong, 0.5-1. 5 mm long; apex acute to obtuse;

margins frequently decurrent; costa mucronate

to short-excurrent, awn up to 260 pm long.

Branch leaves crowded, ± equal in size, ap-

pressed and twisted to spirally twisted around

stem when dry, erect-spreading when wet,

rugose; broadly oblong to oblong-lingulate, 1.0-

2.5 mm long; ventral surface keeled; apex

broadly acute to rounded obtuse; margins plane,

entire. Costa mucronate; in section round, lami-

na ventrally inserted, 2 ventral cells larger, con-

sisting of stereids, dorsal surface rough. Upper

laminal cells rounded-hexagonal to rounded,

incrassate, ± flat, 7.5-15.0 pm, smooth; basal

cells narrowly rhomboidal to narrowly rectan-

gular, incrassate, pitted.

Pseudautoicous; dwarf male plants axillary

on secondary stem. Perichaetia terminal, fre-

quently overgrown by subperichaetial innova-

tions; leaves broadly oblong to oval or oblong-

lanceolate, 1.8-3. 2 mm long. Seta 1. 4-4.5 mm

long, yellowish brown or reddish. Capsule oblong-

cylindrical, brown to red-brown, smooth to

weakly ribbed; urn 1 .4—2.5 mm long; neck 0.2-

0.4 mm long, frequently wrinkled dry; exothe-

cial cells irregular in shape, incrassate, smaller

and occasionally transversely elongate at

mouth. Peristome double; exostome teeth 16,

revolute dry, linear, consisting of 2 divisions,

300-455 pm long, brown, densely striolate-

papillose; endostome segments 16-32, outer

surface smooth, inner surface vertically striolate-

papillose, yellow to pale yellowish brown.

Operculum 1 mm long. Calyptra 3. 2-4.0 mm

long, yellow to brown or reddish brown. Spores

13-40 pm, granulate or minutely papillose;

anisosporous. Fig. 145: 1-14.

Schlotheimia ferruginea is the most fre-

quently collected species of the genus in the

Flora area. It is found on stems and branches of

trees and shrubs and on rock in forests and

wooded areas of the northern and eastern Trans-

vaal regions, Swaziland, Zululand, KwaZulu-

Natal and the eastern Cape region, and occa-

sionally in the southern Cape. This African

Fig. 145. — Schlotheimia ferruginea (1-14): 1. habit (dry), x 3; 2. habit (wet), x 5; 3. branch in cross section (cells part-

ly shown), x 175; 4. stem leaf, x 35; 5. stem leaf apex, x 175; 6. branch leaf, x 35; 7. branch leaf in cross section, x 175;

8. basal cells of branch leaf at right margin, x 700; 9. upper laminal cells of branch leaf, x 700; 10. branch leaf apex, x 175;

11. perichaetial leaf, x 35; 12. part of capsule mouth with peristome (papillae partly shown), x 175; 13. calyptra, x 12; 14.

spores (papillae partly shown), x 700. S. percuspidata ( 15—22): 15. habit (dry), x 1; 16. habit (wet), x 3; 17. stem leaf, x

35; 18. stem leaf apex, x 175; 19. branch leaf, x 35; 20. branch leaf apex, x 175; 21. perichaetial leaf, x 35; 22. Spores

(papillae partly shown), x 700. S. rufopallens (23-30): 23. habit (dry), x 5; 24. habit (wet), x 2; 25. stem leaf, x 35; 26.

stem leaf apex, x 175; 27. branch leaf, x 35; 28. branch leaf apex, x 175; 29. perichaetial leaf, x 35; 30. spore, x 700. (1, 2

& 10, Hilliard & Burtt 11841b; 3, Van Rooy 114; 4, Retief & Herman 42; 5 & 14, Wells 75; 6 & 13, Van Rooy 2161; 7, Von

Breitenbach 95; 8 & 9, Magill 5514; 11, Von Breitenbach 102; 12, Hoffman 41; 15 & 16, Stirton 9813; 17, Van Rooy 1645;

18 & 19, Magill 5510; 20, Von Breitenbach 466; 21, Magill 5216; 22, Smook 904; 23, Smook 6190; 24, Von Breitenbach

414; 25, Magill 6190; 26, Jacot Guillarmod PRE-CH13540; 27, Schelpe 7871; 28, Sim 9288; 29, Cooper 17; 30, Pillans

PRE-CH6261.)

522

Orthotrichaceae

endemic is also known from Tanzania, Zambia

and Zimbabwe. Map 203.

Vouchers: Crosby & Crosby 9201; Hilliard

& Burn 11841B; Kemp 1053; Magill 5514 ,

6576; Stirton 8690; Van Rooy 1188 , 1883,

2161; Von Breitenbach 121.

This species is identified by the relatively

broad branch leaf with subacute to rounded-

obtuse apex and mucronate costa, and the stem

leaf with acute to obtuse apex and mucronate to

short-excurrent costa.

2. Schlotheimia percuspidata C. Miill. in

Hedwigia 38: 117 (1899); Broth, in Natiirl.

PflFam., edn 2, 11: 48 (1925); Sim, Bryo. S.

Afr. 285 (1926); Van Rooy & Van Wyk in

Bryologist 95: 212 (1992). Type: Cape, Blanco,

Oct. 1875, Rehmann s.n.

Plants medium-sized to large, forming mats,

olivaceous or yellowish green to green above,

orange-brown or reddish brown to brown

below; corticolous. Primary stem up to 70 mm

long, branching regularly, tomentose; rhizoids

smooth to weakly papillose, orange-brown to

brown; secondary stem up to 38 mm tall,

branching by subperichaetial innovations; in

section inner cortex 5-8 cells across, incras-

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

consisting of stereids. Stem leaves crowded;

ovate to ovate-lanceolate, 0.6-1. 3 mm long;

apex acute or acuminate; margins decurrent;

costa ending below apex, extending into acu-

men or short-excurrent, awn up to 220 pm

long. Branch leaves crowded, ± equal in size,

appressed and twisted to spirally twisted

around stem when dry, erect-spreading to wide-

spreading when wet, rugose; oblong to oblong-

lanceolate, 1.5-2. 8 mm long; ventral surface

keeled; apex acute, subacute or cuspidate; mar-

gins plane, occasionally reflexed on one side

below, entire. Costa ending below apex or

extending into cuspidate point; in section sub-

round, lamina ventrally inserted, cells not dif-

ferentiated or 2 ventral cells larger, consisting

of stereids, dorsal surface smooth to rough.

Upper laminal cells rounded-hexagonal to

rounded, incrassate, ± flat, 8-13 pm, smooth;

basal cells rhomboidal to rectangular, incras-

sate, strongly pitted.

Pseudautoicous; dwarf male plant axillary on

secondary stem, branching by subperigonial

innovations. Perichaetia terminal, frequently

overgrown by subperichaetial innovations;

leaves oblong-lanceolate or lanceolate, 2. 3-3.0

mm long, apex acuminate or cuspidate, costa

ending below apex to cuspidate. Seta 3-5 mm

long, yellowish brown or red-brown. Capsule

ovoid or oblong-cylindrical, red-brown to

brown, smooth to weakly ribbed; urn 1.3-1. 8

mm long; neck 0.2-0. 5 mm long, wrinkled dry;

exothecial cells irregular in shape, incrassate,

smaller at mouth. Peristome double; exostome

teeth 16, revolute dry, linear, consisting of 2

divisions, 400^-70 pm long, brown, densely

striolate-papillose; endostome segments 16-32,

outer surface smooth, inner surface vertically

striolate-papillose, yellowish. Operculum 1 mm

long. Calyptra 3. 0-3. 5 mm long, yellowish

brown or reddish brown. Spores 12-35 pm,

granulate; anisosporous. Fig. 145: 15-22.

Schlotheimia percuspidata grows on bark of

trees and on rock, and is infrequently collected

in montane forests of the northern and eastern

Transvaal regions, Zululand and KwaZulu-

Natal. This African endemic is also known from

Tanzania, Malawi and Zimbabwe. Map 204.

Orthotrichaceae

523

Map 204. — Schlotheimia percuspidata

Vouchers: Filter 26; Magill 5216, 5510;

Smook 904; Van Rooy 1645; Von Breitenbach

172, 466.

Schlotheimia percuspidata is most easily

identified by the branch leaves that are gradual-

ly narrowed towards the cuspidate apices. The

stem leaf costa usually ends below the acute to

acuminate apex but sometimes also extends into

the acumen or becomes short-excurrent.

Several specimens from the eastern

Transvaal region ( Brenan M3 3 30; Crosby &

Crosby 7646, 7648, 7658, 13398; Stirton 9813;

Vorster 503) differ in the acute to rounded-

obtuse apices and mucronate costae of their

branch leaves. These specimens can, however,

be distinguished by the stem leaf costa which

ends below the acuminate apex.

3. Schlotheimia rufopallens C. Mull, in

Hedwigia 38: 117 (1899); Broth, in Natiirl.

PflFam., edn 2, 1 1 : 48 (1925). Lectotype: Cape,

Blanco, Rehmann 154 (BM!) fide Van Rooy &

Van Wyk in Bryologist 95: 212 (1992).

Schlotheimia exrugulosa C. Mull, in Hedwigia 38: 118,

(1899). Lectotype: Cape, Table Mountain, Rehmann 152

(BM!) fide Van Rooy & Van Wyk in Bryologist 95: 212

(1992).

Plants medium-sized, in mats, olivaceous

above, orange-brown, reddish brown or brown

below; corticolous. Primary stem to 70 mm

long, branching regularly, frequently tomen-

tose; rhizoids smooth to weakly papillose,

orange or reddish brown; secondary stem to 30

mm tall, branching by subperigonial and sub-

perichaetial innovations; in section inner cortex

5-9 cells across, incrassate, outer cortical cells

smaller, in 1-3 rows, consisting of stereids,

outer surface occasionally rough. Stem leaves

generally crowded; ovate-lanceolate to lanceo-

late, 0.5—1 ,5(— 1 .8) mm long; apex acute to

acuminate; margins slightly decurrent; costa

aristate, awn (120-)180-^100(^160) pm long.

Branch leaves crowded, ± equal in size, ap-

pressed and twisted to spirally twisted around

stem when dry, erect-spreading to squarrose

when wet, rugose; oblong, ( 1 .0—) 1 .3—2.5 mm

long; ventral surface keeled; apex acute, sub-

acute or rounded-obtuse; margins plane or

reflexed on one side below, entire, weakly pro-

rate or crenulate above. Costa mucronate to

cuspidate; in section subround, lamina ventrally

inserted, 2 ventral cells larger, consisting of

stereids, dorsal surface prorate. Upper laminal

cells rounded-hexagonal to rounded or oval,

incrassate, ± flat, 5-12 pm, smooth; basal cells

rhomboidal to rectangular, incrassate, strongly

pitted, infrequently papillose.

Dioicous. Perigonia terminal, leaves ovate.

Perichaetia terminal, frequently overgrown by

subperichaetial innovations; leaves broadly

oblong, oblong-lanceolate or lanceolate, 1.8-

2.8 mm long. Seta 3-9 mm long, yellowish

brown or reddish brown. Capsule oblong-cylin-

drical, yellowish brown, reddish brown or red

brown, ribbed, urn 1.0-1. 5 mm long, neck 0.5

mm long; exothecial cells irregular in shape,

incrassate, smaller towards mouth, 2-A rows at

mouth transversely elongated. Peristome dou-

ble; exostome teeth 16, revolute dry, linear, con-

sisting of 2 divisions, 280^410 pm long, yel-

low-brown, densely striolate-papillose; endo-

stome segments 16-32, outer surface smooth,

inner surface vertically striate-papillose, yel-

lowish or hyaline. Operculum 1 mm long.

Calyptra 2.0-3. 5 mm long, yellow-brown, red-

dish brown or brown. Spores 20-32 pm, granu-

late; isosporous. Fig. 145: 23-30.

524

Orthotrichaceae

Schlotheimia rufopallens is known only

from scattered localities in the southwestern,

southern and eastern Cape regions, KwaZulu-

Natal, Zululand, and the central and northern

Transvaal areas. Map 205.

Vouchers: Cooper 17; Magill 5185, 5335;

Schelpe 7871; Smook 1597, 1620; Von Breiten-

bach 344, 414.

Schlotheimia rufopallens can be recognized

by the aristate costae and acute to acuminate

apices of its stem leaves. This species has nar-

rower branch leaves and a longer excurrent

stem leaf costa than Schlotheimia ferruginea

and S. percuspidata. The dioicous sexual condi-

tion and isomorphic spores also help to identify

plants.

Insufficiently known species

Schlotheimia rufoaeruginosa C. Mull, in

Linnaea 39: 410 (1875); Broth, in Natiirl.

PflFam., edn 2, 11: 48 (1925); Sim, Bryo. S.

Afr. 284 (1926). Type: ‘Africa australis, Natal,

Drakensberg, in Polypodii incani rhizomate: M.

Lea 1874.’ The type could not be located.

Schlotheimia rufoglauca C. Mull, in

Hedwigia 38: 118 (1899); Broth, in Natiirl.

PflFam., edn 2, 11: 48 (1925). Type: ‘Prom,

bonae spei, Knysna district, in sylvis prope

Estemek, Nov. 1875, . . . Rehmann s.n.' The

type could not be located. Rehmann specimens

in H! and PRE!, originally named as S. rufo-

glauca but collected near Portland ( Rehmann

153), are S. rufopallens. Sim (1926) treated the

species as a synonym of S. rufoaeruginosa.

Schlotheimia ventrosa C. Mull., Syn. muse,

frond. 1: 756 (1849); Broth, in Natiirl. PflFam.,

edn 2, 11: 47 (1925). Type: ‘Prom. b. spei,

Olifantshoek in Districtu Uitenhagen: Ecklon.

Hb. Kunz.’ The original material in Herb.

Kunze (LZ) was probably destroyed and suit-

able type material could not be located. Sim

(1926) noted that this species (from descrip-

tions only) may be a synonym of S. grevilleana

Mitt. Magill & Schelpe (1979) noted that S.

subventrosa (treated here as a synonym of S.

ferruginea) is conspecific with S. ventrosa. A

specimen in BM named as S. ventrosa and col-

lected at the type locality by Pappe is S. ferru-

ginea.

9. CARDOTIELLA

Cardotiella Vitt in J. Hattori Bot. Lab. 49: 101 (1981); Van Rooy & Van Wyk in Bryologist 95: 212

(1992). Type species: C. subappendiculata (Broth.) Vitt.

Plants large, forming mats, green to yellowish green or yellowish brown above, brown below;

saxicolous or corticolous. Primary stems creeping, tomentose; secondary stems erect-curved, wide-

ly spaced. Leaves crowded, secund, in 4 or 5 rows, generally rugose, frequently falcate, ovate to

lanceolate; margins decurrent, irregularly denticulate above, denticulate to spinulose below; stem

leaves smaller. Laminal cells rounded to rounded-hexagonal, homogeneous, papillose to strongly

papillose, frequently obscure below; cells of decurrency irregularly rectangular, inflated, smooth,

frequently tuberculate at margin.

Orthotrichaceae

525

Dioicous. Perigonia lateral, gemmate. Perich^etia terminal on secondary stems, leaves ovate-

subulate to oblong-subulate. Seta short. Capsule exserted, elliptic, 8-ribbed; stomata phaneropore.

Peristome double; exostome teeth 16, fused below; endostome segments 8-16, alternating with

teeth. Operculum conic-rostrate. Calyptra mitrate, lobed at base, weakly plicate, hairy. Spores

round, minutely papillose, pale brown.

Cardotiella is a genus of six species; four are endemic on the East African islands of

Madagascar, Mauritius and Reunion, one species is known only from the Neotropics, and one is

endemic to South Africa. The genus is characterized by the erect-curved branches with secund,

rugose leaves; leaf decurrency of large, inflated and smooth cells; strongly unipapillose, homoge-

neous laminal cells; well-developed peristome, and the large mitrate calyptra.

Cardotiella secunda (C. Mull.) Vitt in J.

Hattori Bot. Lab. 49: 105 (1981); Van Rooy & Van

Wyk in Bryologist 95: 212 (1992). Type: Cape,

Olifantshoek District, Uitenhage, Pappe s.n.

Macromitrium secundum C. Mull, in Bot. Zeitung

(Berlin) 14: 420 (1856); Sim, Bryo. S. Afr. 281 (1926).

Coleochaetium secundum (C. Mull.) Broth, in Nattirl.

PflFam. 1,3: 475 (1902).

Macromitrium schlotheimiaeformis Par., Ind. bryol.

suppl. 1 : 241 (1900). Cardotiella schlotheimiaeformis (Par.)

Vitt in J. Hattori Bot. Lab. 49: 103 ( 1981). Lectotype: Cape,

Cape Town. Devil’s Peak, Rehmann 151 (NY; PRE, isolec-

td. !).

Plants large, forming mats, green to yellow-

ish green or yellowish brown above, brown

below; saxicolous or corticolous. Primary stem

creeping, up to 70 mm long, irregularly branch-

ed, frequently tomentose; rhizoids smooth, red-

dish brown; secondary stem erect-curved,

branching by subperichaetial innovations, occa-

sionally tomentose below; in section round,

central strand absent, inner cortex 8-12 cells

across, thin to thick-walled, outer cortical cells

in 2-&, rows, walls incrassate or consisting of

stereids, outer surface rough. Stem leaves weak-

ly rugose to rugose, occasionally falcate, fre-

quently reflexed; ovate, ovate-acuminate,

ovate-ligulate or ovate-lanceolate to lanceolate,

(0.8—) 1 .0—1 .7(— 2.0) mm long; apex acute to

acuminate; margins plane, entire to irregularly

denticulate above, denticulate to spinulose

below, decurrent; costa ending below apex to

subpercurrent or infrequently mucronate.

Branch leaves crowded, ± equal in size, secund,

in 4 or 5 rows, generally rugose, occasionally

falcate; ovate-lanceolate or oblong-lanceolate

to lanceolate to narrowly lanceolate, 1.0-2. 5

mm long; apex acuminate, acute or rarely

rounded-obtuse; margins plane, entire to irregu-

larly denticulate above, denticulate to spinulose

below, decurrent; cells of decurrency irregularly

rectangular, inflated, smooth, frequently tuber-

culate at margin; unistratose. Costa ending

below apex to subpercurrent or occasionally

mucronate; ventral superficial cells rounded or

short-rectangular above, linear below; dorsal

superficial cells rounded to oval above, linear

below; in cross section subround, flat ventrally,

bulging dorsally, 2 ventral surface cells incras-

sate or substereids, central cells incrassate or

consisting of stereids, dorsal surface cells

incrassate or consisting of substereids. Laminal

cells rounded-hexagonal or rounded-quadrate,

homogeneous, incrassate, frequently obscure,

7-10 pm, smooth to papillose above, papillose

to strongly papillose or spinulose below, papil-

lae single, conical; basal marginal cells occa-

sionally subquadrate, protruding.

Dioicous. Perigonia lateral, gemmate, leaves

broadly ovate-apiculate. Perichaetia terminal

on branches; leaves ovate-subulate to oblong-

subulate, 2.3-3. 2 mm long. Seta 1.0-1. 8 mm

long, yellowish brown or reddish brown.

Capsule elliptic; urn 1.2-1. 8 mm long; neck

0. 6-1.0 mm long; yellowish brown or reddish

brown to brown, 8-ribbed; exothecial cells

irregular rectangular to quadrate, smaller at

mouth, incrassate; stomata present at base of

urn and on neck; annulus weakly differentiated.

Peristome recurved when dry; exostome teeth

16, fused below to form 8 pairs, perforated

above, 0.4 mm long, yellow-brown, papillose

below, vertically striolate-papillose above;

526

Orthotrichaceae

Map 206. — • Cardotiella secunda

♦ Rhabdoweisia fugax

■ Rhabdoweisia crispata

endostome segments 8-16, alternating -with

teeth, shorter than exostome, narrow, hyaline,

outer surface smooth, inner surface vertically

striolate. Operculum 0. 8-1.0 mm long.

Calyptra 2.2-2. 5 mm long, yellow or yellowish

brown. Spores 15-23 pm. Fig. 146.

Endemic to southern Africa, Cardotiella

secunda is known from scattered localities in

forests of the southwestern, southern and east-

ern Cape, and Zululand. Map 206.

Vouchers: Magill 5476, 6047, 6269, 6307,

7640; Schelpe 7646; Van Rooy 944, 1714a; Van

Zinderen Bakker 249.

The erect-curved branches, secund, rugose

leaves with decurrent margins, strongly papil-

lose laminal cells and peristome characters

separate C. secunda from other members of the

subfamily.

Fig 146. — Cardotiella secunda: 1. habit (dry), x 1; 2.

habit (wet), x 5; 3. branch in cross section (cells partly

shown), x 175; 4. stem leaf, x 35; 5. branch leaf, x 35; 6.

branch leaf in cross section, x 175; 7. basal cells of branch

leaf at left margin, x 350; 8. upper laminal cells of branch

leaf (left side), x 350; 9. branch leaf apex, x 350; 10. part of

capsule mouth with peristome (papillae partly shown), x

140; 11. calyptra, x 18; 12. spore, x 700. (1, 3, 5 & 11,

Schelpe 7640', 2, 1 0 & 12, Magill 6269 ; 4 & 6, Crosby 6269',

7, Taylor 6312\ 8, Van Zinderen Bakker 249', 9, Magill

5476.)

527

RHABDOWEISIACEAE

Plants small, caespitose; terricolous. Stems erect; in section round, central strand absent; rhi-

zoids smooth. Leaves crisped when dry, erect to erect-spreading when wet; linear-lanceolate, nar-

rowly oblong-lanceolate or narrowly ligulate; margins plane or recurved on one side below, entire,

crenulate or irregularly denticulate. Costa ending below apex, in section with median guide cells.

Upper laminal cells rounded-quadrate or rounded-hexagonal, incrassate, smooth; basal cells rec-

tangular, thin-walled, frequently hyaline.

Autoicous. Perichaetia terminal, leaves scarcely differentiated. Capsule exserted, erect, 8-

ribbed; stomata phaneropore; annulus absent. Peristome single, frequently fugacious, teeth 16, dis-

tant, narrow, smooth, basal membrane low. Operculum conic-rostrate, oblique. Calyptra cucullate,

smooth, naked. Spores subtriangular-globular; granulate, brownish.

Anderson & Crum (1959) proposed the family to contain the genera Rhabdoweisia B.S.G.,

Rhabdoweisiella Williams and Amphidium Schimp. Amphidium is closely related to some of the

diplolepidous genera in the Orthotrichaceae (Lewinsky 1976) and is treated there for the Flora (see

p. 486). However, the haplolepidous origin of the peristome in the Rhabdoweisiaceae indicates a

position in the Dicranales rather than the Orthotrichales.

RHABDOWEISIA

Rhabdoweisia B.S.G., Bryol. eur. 1: 97 (1846); Broth, in Natiirl. PflFam., edn 2, 10: 194 (1924);

Sim, Bryo. S.Afr. 151 (1926); Lawton in Bryologist 64: 141 (1961); Smith, Moss FI. Brit. Irel. 128

(1978); Crum & Anderson, Moss. E.N. Amer. 1: 176 (1981); Van Rooy in Bryologist 94: 409

(1991). Type species: R. fugax (Hedw.) B.S.G.

The genus contains approximately five species and occurs in Europe, Greenland, North and

South America, Asia and Africa. Rhabdoweisia is represented in Africa by four species of which

two occur in southern Africa.

Leaf margins entire to crenulate, apex generally acuminate, laminal cells in (4)5— 7(8) rows

on each side of costa at leaf middle; spores 1 1-17 pm 1 . R. fugax

Leaf margins irregularly denticulate, apex generally acute, laminal cells in (5— )7— 9( 1 0) rows

on each side of costa at leaf middle; spores 16-25 pm 2. R. crispata

1. Rhabdoweisia fugax (Hedw.) B.S.G. ,

Bryol. eur. 1: 98 (1846); Broth, in Natiirl.

PflFam. 1,3: 313 (1909); Sim, Bryo. S. Afr. 151

(1926); Nyholm, Moss FI. Fenn. 49 (1954);

Smith, Moss FI. Brit. Irel. 128 (1978); Van Rooy

in Bryologist 94: 409 (1991). Type: Germany;

Sudeten, Breutel, Musci frondosi 260 (NY,

neo.), vide Lawton in Bryologist 64: 144 ( 1961).

Weissia fugax Hedw., Sp. muse, frond. 64 (1801).

Plants yellowish green to brown; terricolous.

Stems up to 5 mm tall, branching by sub-

perichaetial innovations; rhizoids reddish

brown, smooth; in section with cortical cells

thin-walled, epidermis not differentiated. Leaves

larger above, crisped when dry, erect to erect-

spreading when wet, reflexed; linear-lanceolate,

1.0-1. 8 mm long; keeled; apex acuminate or

occasionally acute; base scarcely differentiated,

± sheathing in upper leaves; margins plane or

frequently recurved on one side below, entire to

crenulate at apex, frequently flexuose above.

Costa ending below apex to subpercurrent, ven-

tral superficial cells rectangular, dorsal superfi-

cial cells linear; in section crescent-shaped to

subround, ventral surface flat, lamina ventrally

inserted, guide cells in 1 layer, ventral stereid

band absent, ventral surface cells incrassate.

Rhabdoweisiaceae

Fig 147 — Rhabdoweisia fugax (1-13): 1. habit (dry), x

5; 2. habit (wet), x 10; 3. stem in cross section, x 350; 4.

leaves, x 35; 5. leaf in cross section, x 350; 6. basal leaf

cells, x 350; 7. upper laminal cells (left side), x 700; 8. leaf

apex, x 350; 9. capsule (dry), x 20; 10. capsule with oper-

culum (wet), x 50; 11. part of capsule wall with stoma, x

700; 12. part of capsule mouth with peristome teeth, x 350;

13. spores (papillae partly shown), x 700. R. crispata

(14-23): 14. habit (wet), x 10; 15. leaves, x 35; 16. leaf in

cross section, x 350; 17. basal leaf cells (left side), x 350;

18. upper laminal cells at right margin, x 350; 19. cells at

leaf apex, x 350; 20. capsule (dry), x 20; 21. capsule (wet),

x 50; 22. calyptra, x 35; 23. spore, x 700. (3-7, 9 & 11,

Wager PRE-CH535 ; 1, 2, 8, 10, 12 & 13, Wager 1072c ;

14-23, Van Rooy 3689.)

Rhabdoweisiaceae

529

bulging, dorsal stereid band small, dorsal sur-

face cells incrassate, bulging. Upper laminal

cells rounded-quadrate or rounded-hexagonal,

incrassate, bulging, 8.7-16.3 pm, smooth, in

(4)5-7(8) rows on each side of costa at leaf

middle, in 3-5(6) rows just below apex; basal

cells occasionally reaching higher along costa,

rectangular, thin-walled, smooth.

Autoicous. Perigonia terminal on short

branches, inner leaves ovate. Perichaetia termi-

nal, leaves scarcely differentiated. Seta 1. 8-2.4

mm long, yellowish to brown, twisted anti-

clockwise above. Capsule exserted, ovoid, 0.7

mm long, yellowish to brownish, weakly 8-

ribbed, neck short, mouth orange; exothecial

cells irregular rectangular, thin-walled, smaller

above, 2-4 rows at mouth transversely elongat-

ed, cells of ribs weakly differentiated, bulging;

stomata on neck, few, phaneropore. Peristome

single, inserted below mouth, teeth 16, distant,

filiform, 93-110 pm long, orange, smooth,

basal membrane low. Operculum 0.5 mm long.

Calyptra not seen. Spores 11-15 pm. Fig. 147:

1-13.

The species is known from Europe, the

Caucasus, eastern Siberia, Macaronesia, Sri

Lanka, China, the Neotropics, and southern

Africa. In the Flora area the species is known

from the Drakensberg of KwaZulu-Natal and

the eastern Transvaal region. Map 206.

Vouchers: MacLea sub Rehmann 501; Wager

PRE-CH11570.

Rhabdoweisia fugax is recognized by its lin-

ear-lanceolate leaves with acuminate apices,

entire to crenulate leaf margins, and the single

peristome with distant, filiform teeth abruptly

narrowed from a low basal membrane.

2. Rhabdoweisia crispata (Dicks.) Lindb.

in Act. Soc. Sci. Fenn. 10: 22 (1871); Smith,

Moss FI. Brit. Irel. 130 (1978); Crum &

Anderson, Moss. E.N. Amer. 1: 177 (1981).

Type: Herb. Dickson, sheet 32 no. 30 (BM,

lecto.!) fide Van Rooy in Bryologist 94: 409

(1991)."

Bryum crispatum Dicks., PI. crypt, brit. 4: 29 (1801).

Plants green above, light brown below; terri-

colous. Stems up to 5 mm tall, branching by

subperichaetial innovations, tomentose below;

rhizoids reddish brown, smooth; in section with

cortical cells thin-walled, outer 1 or 2 rows

smaller, epidermis not differentiated. Leaves

larger above, crisped when dry, erect to erect-

spreading when wet, recurved; narrowly

oblong-lanceolate, narrowly ligulate or linear-

lanceolate, 1 .0— 2.3(— 3 .2) mm long; keeled;

apex acute or occasionally acuminate; base

scarcely differentiated, ± sheathing; margins

plane or frequently recurved on one side below,

irregularly denticulate or serrulate, flexuose

above. Costa ending below apex, ventral super-

ficial cells rectangular, dorsal superficial cells

linear; in section crescent-shaped, ventrally flat,

lamina ventrally inserted, guide cells in 1 layer,

ventral stereid band absent, ventral surface cells

incrassate, dorsal stereid band small, dorsal sur-

face cells incrassate or substereids, bulging.

Upper laminal cells irregularly rounded-

quadrate or rounded-hexagonal, incrassate,

bulging, 1 1 .2— 17.5(— 20.0) pm, smooth, in (5-)

7-9(10) rows on each side of costa at leaf mid-

dle, in (4)5 or 6(7) rows just below apex; basal

cells rectangular, thin-walled.

Autoicous. Perigonia terminal on short

branches below perichaetia, inner leaves ovate.

Perichaetia terminal, leaves scarcely differenti-

ated. Seta 1.6-2. 7 mm long, yellowish brown,

twisted anticlockwise above. Capsule exserted,

ovoid, 0.4-0. 8 mm long, yellowish brown to

brown, weakly 8-ribbed, neck short; exothecial

cells irregular-rectangular, smaller at mouth,

cells of ribs weakly differentiated; stomata few,

on neck, phaneropore. Peristome single, insert-

ed below mouth, fugacious, teeth not seen,

basal membrane low, orange. Operculum 0.5-

0.7 mm long. Calyptra 0.8- 1.2 mm long,

smooth, naked. Spores 16.0-22. 5(— 25.0) pm.

Fig. 147: 14-23.

Rhabdoweisia crispata is rather widespread

in the temperate to arctic northern hemisphere

and has also been reported from Hawaii, the

Neotropics, Juan Fernandez islands, Lebanon,

Bhutan, Java, and North and South Africa. The

species has been collected recently (1987) on

530

Rhabdoweisiaceae

the Drakensberg escarpment at Sani Pass. Map

206.

Voucher: Van Rooy 3689.

This species is most easily separated from

Rhabdoweisia fngax by the shape of the peri-

stome teeth. The filiform teeth of R. fngax

abruptly narrow from the basal membrane while

the narrowly lanceolate teeth of R. crispata

taper evenly from the broad base. The only col-

lection of R. crispata known from southern

Africa, however, has immature capsules and

capsules with the teeth broken away.

The two species are more difficult to sepa-

rate in the absence of peristome teeth, but the

broader leaves with acute apices and irregularly

denticulate margins, and the larger spores will

usually distinguish R. crispata from R. fugax.

531

RACOPILACEAE

A small family containing only two genera, Racopilaceae is primarily of southern hemisphere

distribution. The genera, including Racopilum , the only representative in southern Africa, are rec-

ognized by their strongly dimorphic and ranked leaves which are tightly curled up and over the

stem when dry.

RACOPILUM

Racopilum R Beauv., Prodr. Aetheogam. 36 (1805); Sim, Bryo. S. Afr. 447 (1926); Broth, in

Natiirl. PflFam., edn 2, 11: 52 (1926); Catcheside, Moss. S. Austr. 291 (1980). Type species: R.

mnioides P. Beauv.

Plants small to large, in loose mats, dark green; mostly terricolous. Stems creeping, heterophyl-

lous; central strand small. Leaves 4-ranked, incurved dry, widely spreading wet, leaves in lateral

ranks larger than dorsal ranks; strongly cuspidate; margins weakly serrate above. Leaf cells rec-

tangular to quadrate, minutely mammillose, rectangular below; alar cells not differentiated.

Dioicous or autoicous. Perigonia and perichaetia along stem. Seta long, smooth. Capsule hori-

zontal, ribbed. Peristome double, complete. Operculum rostrate. Calyptra cucullate, somewhat

hairy. Spores small, green.

A genus with over 60 species, found in the Americas, Africa, Madagascar, India, southern Asia,

Australia, New Zealand and Oceania. The genus is easily identified by its dimorphous leaves which

spread widely when wet. The leaves are arranged in four rows. The two apparently lateral rows of

leaves are larger and broader than the two rows on the dorsal stem surface. Some species are

described as having dwarf male plants on the rhizoids of the larger female plants. However, the

southern African species has male and female plants of equal size; the males produce large perigo-

nia between the lateral leaves.

Racopilum capense C. Mull, in Hedwigia

38: 124 (1899); Sim, Bryo. S. Afr. 447 (1926);

Broth, in Natiirl. PflFam., edn 2, 11: 53 (1925).

Syntypes: Cape, Claremont, Rehmann, Oct.

1876; Touws River, Rehmann , Nov. 1875;

Somerset East, Boschberg, MacOwan s.n. (G!);

Natal, Inanda, Rehmann 297 (PRE!); Van

Reenen Pass, Rehmann 297b (PRE!).

Plants small to large, forming mats, dark

green; terricolous, humicolous, saxicolous or

corticolous. Stems creeping, up to 60 mm

long, branching regular, pinnate; in section

oval, central strand small, inner cortical cells

thin-walled, hyaline, in 5 or 6 rows, outer cor-

tical cells thick-walled, red-brown, in 2 or 3

rows; paraphyllia and pseudoparaphyllia

absent. Leaves evenly spaced, 4-ranked, wide-

spreading wet, incurved dry; lateral leaves

broadly ovate to elliptical, 1-2 mm long;

acute, cuspidate or short-awned; rounded at

base; margins plane, entire below, serrate

above; dorsal leaves ovate, 0.8-1. 5 mm long;

acute, cuspidate or short-awned; rounded at

base; margins plane, weakly serrate. Costa

single, short-excurrent, smooth, yellow, both

surfaces smooth; in section bulging dorsally,

guide cells thin-walled, ventral stereid band

absent, ventral surface cells large, thick-

walled, dorsal stereid band 2 cells thick, dor-

sal surface cells small, thick-walled. Upper

laminal cells hexagonal to rhomboidal,

becoming rectangular at margins in lateral

leaves, rhombic to quadrate in dorsal leaves,

7-12 pm long, walls weakly thickened, ±

homogeneous, minutely mammillose on both

surfaces; basal cells quadrate to short-rectan-

gular, 12-24 pm long, 12 pm wide, hyaline,

walls smooth, weakly thickened; alar cells not

differentiated.

532

Racopilaceae

Dioicous. Perigonia axillary on stem, gem-

mate; perigonial leaves orbicular-acuminate.

Perichaetia strongly differentiated; perichaetial

leaves oblong to ovate, apex long-cuspidate,

1-2 mm long with awn 1-2 mm long, leaf cells

fusiform, thin-walled and pitted. Seta 15-22

mm long, red-brown, smooth. Capsule exsert-

ed, horizontal to nodding, cylindrical to weakly

pyriform, 2. 5-3.0 mm long, ribbed, red-yellow,

urn cylindrical, neck not differentiated; exothe-

cial cells rectangular, walls firm, cells at mouth

quadrate; annulus differentiated; stomata pha-

neropore on lower urn. Peristome complete,

orange; exostome teeth linear from a broader

base, erect with inflexed tips dry, incurved wet;

striate below with median zigzag line, papillose

above, up to 700 pm high; endostome segments

keeled and perforated, as long as teeth, on high

basal membrane, granulate; cilia 3, linear, as

long as segments, granulate. Operculum coni-

cal and long-rostrate, 1.5 mm long. Calyptra

Fig. 148. — Racopilum capense: 1. habit (dry), x 1; 2.

dorsal view of branch, x 5; 3. stem in cross section (cells

partly shown), x 175; 4. dorsal leaf, x 32; 5. lateral leaf, x

32; 6. lateral leaf in cross section, x 175; 7. basal cells of

lateral leaf, x 350; 8. upper laminal cells of lateral leaf

(right side), x 350; 9. lateral leaf apex, x 175; 10. part of

capsule mouth with peristome, x 70. (1, 4 & 5, Crosby

7886- 2, Crosby 9141\ 3 & 6-10, Crosby 7519.)

Racopilaceae

533

not seen. Spores rounded, 12-16 |am, granulate,

green. Fig. 148.

In the Flora area, R. capense is found in for-

est and woodland sites, especially near streams,

in the northern, eastern and central Transvaal

regions, Swaziland, KwaZulu-Natal, Zululand,

and the eastern, southern and southwestern Cape

areas. It is also found throughout western, south-

eastern and eastern Africa, the East African

islands and the Arabian peninsula. Map 207.

Vouchers: Crosby & Crosby 9141; Magill

3487, 4783; Oliver 7 167 A; Van Rooy 258.

The single southern African species is easily

identified by its dark green, creeping stems with

dimorphic leaves. Racopilum capense is

dioicous, with male plants as large as the female

plants ( Kemp 1474). Its nearest relatives are

either monoicous, R. tomentosum (Hedw.) Brid.,

or produce dwarf males on rhizoids of female

plants, R. cuspidigerum (Schwaegr. ) Angstr.

534

FONTINALACEAE

A family of four genera found primarily in temperate regions, especially in the northern hemi-

sphere. In the Flora area, Fontinalaceae is represented by two species, Fontinalis antipyretica and

F. squamosa. The plants, found in streams of the southwestern Cape, are believed to have been

introduced with fish from Europe where both species are known to occur.

FONTINALIS

Fontinalis Hedw., Sp. muse, frond. 298 (1801); Broth, in Natiirl. PflFam., edn 2, 11: 57 (1925);

Sim, Bryo. S. Afr. 354 (1926); Welch, Monograph Fontinalaceae 18 (1960). Lectotype species: F.

antipyretica Hedw. cf. Myrin in Kongl. Vetensk. Acad. Handl. 1832: 273 ( 1832).

Plants long, in loose floating mats, dark green; aquatic and saxicolous. Stems elongated and

branched; central strand absent; paraphyllia absent; pseudoparaphyllia absent. Leaves erect to

spreading, frequently carinate or concave; margins plane, entire. Costa present as bistratose region

in extreme leaf base. Laminal cells elongate, fusiform, thin-walled; alar cells strongly differentiat-

ed, enlarged, quadrate.

Dioicous. Sporoplrytes not known from the Flora area but described as: Capsule immersed to

emergent, oval to cylindrical. Peristome double. Operculum long-conical. Spores medium-sized.

A genus of over 37 species found mostly in temperate regions of the northern hemisphere. The

plants generally grow submerged on rock in streams or pools, but are also found exposed when

water levels drop for short periods. The large mats fan out in still water and can cover extensive

areas when conditions are right. The unistratose leaves and lack of vascular tissue separate speci-

mens from aquatic vascular plants. Fontinalis can be confused with Wardia , another aquatic moss

that grows in the mountains of the southwestern Cape. A comparison of the two indicates that

Fontinalis has much longer stems which branch throughout, longer and narrower leaves which are

frequently keeled and an immersed to emergent capsule.

Stem leaves keeled or deeply channelled when wet (most obvious at stem or branch tips) . .

1. F. antipyretica var. gracilis

Stem leaves concave when wet, not keeled 2. F. squamosa

1. Fontinalis antipyretica Hedw. var. gra-

cilis (Lindb.) Schimp., Syn. muse. eur. 2: 552

(1876); Welch, Monograph Fontinalaceae 46

(1960); Smith, Moss FI. Brit. Irel. 495 (1980).

Type: Finland, Lindberg, 1868, fide Welch

(1960).

Fontinalis gracilis Lindb. in Hedwigia 6: 39 (1867);

Broth, in Natiirl. PflFam., edn 2, 1 1 : 58 (1925).

Fontinalis antipyretica sensu Sim, Bryo. S. Afr. 354

(1926).

Plants long and slender, in floating mats, dark

green to yellow green; aquatic and saxicolous.

Stems up to 300 mm long, branches numerous,

regular; in section round, central strand absent,

inner cortical cells thin-walled, hyaline, becom-

ing more thick-walled and yellowish towards

margin, in 6-8 rows, outer cortical cells thick-

walled, red, in single row; axillary hairs with

basal cells brown, 6 or 7 cells long, hyaline.

Leaves somewhat distant, spreading and keeled

wet, appressed or spreading dry; 3. 5-5.0 mm

long, 1.8-2. 2 mm wide; acute and frequently

somewhat cucullate, weakly reflexed at tip; nar-

rowed to insertion; margins plane, entire. Costa

restricted to a bistratose region at insertion.

Upper laminal cells linear, ± fusiform and sig-

moid, homogeneous, smooth, 80-190 pm long,

10-12 pm wide, walls thin; basal cells not dif-

ferentiated, yellowish, walls thin, smooth; alar

Fontinalaceae

535

cells strongly differentiated, forming distinct

groups, large, quadrate, hyaline, walls thin.

Sporophytes not known from the Flora area.

Fig. 149: 7-11.

Widely distributed with numerous local sub-

specific taxa, F. antipyretica is found on rock

and wood in cold water streams and ponds

throughout the northern hemisphere, including

North America, Europe and Asia. The variety

gracilis was introduced into streams of the

southwestern Cape and according to Sim

(1926), was first collected in 1919. Map 208.

Vouchers: Mogg 2837; Oliver 9052b; Pillans

9979; Wicht 11118.

This variety is believed to have been intro-

duced with fish from the northern hemisphere.

The specimens clearly exhibit the characters of

the European variety, F. antipyretica var. gra-

cilis. In a group that is thought to express a

great deal of environmental adaptation, this

population, which has remained uniform for ±

80 years, should provide interesting insights

into environmentally induced character modifi-

cation.

The South African specimens are uniform

and exhibit several interesting characters. For

example, the leaves are flattened when dry and

show a keel only when wet. Leaves are fre-

quently split along the keel, especially on

slides, thus masking the carinate condition. This

may cause some confusion with the other

species, F. squamosa , which has concave

leaves. In F. antipyretica var. gracilis the leaves

at the stem tips are distinctly keeled when wet,

giving the stem an angled appearance. The

leaves also appear ecostate, but have a bi-

stratose area at the base which represents a rudi-

mentary costa as described by Allen (1983).

2. Fontinalis squamosa Hedw., Sp. muse,

frond. 299 (1801); Broth, in Natiirl. PflFam.,

edn 2, 11: 60 (1925); Welch, Monograph Fon-

tinalaceae 99 (1960). Type: England.

Plants long and slender, forming floating

mats, green to dark green; aquatic and saxi-

colous. Stems up to 400 mm long, branching

irregular, somewhat fasciculate; in section

round to oval, central strand absent, inner cor-

tical cells thin-walled, large, hyaline, outer cor-

tical cells thick-walled and yellow, becoming

smaller and reddish toward margin; axillary

hairs with brown basal cells. Leaves evenly

spaced, spreading wet, appressed dry, concave;

stem leaves ovate-lanceolate, 3. 5-4. 5 mm

long; acute to obtuse, weakly decurrent at base;

margins plane to weakly inflexed, entire, weak-

ly bordered, marginal cells not differing in

shape but with a faint yellowish colour; branch

leaves similar to stem leaves although some-

what smaller, 3. 0-4.0 mm long. Costa absent

or a few bistratose areas at extreme base of

leaves. Upper laminal cells linear, somewhat

fusiform, homogeneous, 110-125 pm long,

8-10 pm wide, walls thin, smooth; basal cells

rectangular, forming distinct group at insertion,

reddish brownish, smooth; alar cells strongly

differentiated, enlarged and inflated, reddish,

walls thin.

Sporophyte not known from Flora area. Fig.

149: 1-6.

Found on rocks in streams in Europe, Asia

and northern and southern Africa, the species

was probably also introduced with fish import-

ed from Europe. At this time, the only collec-

tion is from below the main dam on the Eerste

536

Fontinalaceae

FIG. 149. — Fontinalis squamosa (1-6): 1. habit (dry), x

1; 2. part of stem in cross section, x 175; 3. stem leaf, x 35;

4. branch leaf, x 32; 5. basal leaf cells (left side), x 160; 6.

branch leaf apex, x 160. F. antipyretica var. gracilis (7-1 1 ):

7. habit (dry), x 1; 8. branch leaf, x 35; 9. branch leaf (side

view), x 35; 10. upper laminal cells, x 320; 11. branch leaf

apex, x 175. (1-6, Oliver 9052\ 7-9 & 11, Oliver PRE-

CH13598 ; 10, Mogg 2837.)

Fontinalaceae

537

River in Jonkershoek Valley. The collection was

mixed with F. antipyretica but it is unclear

whether they were growing intermixed. Map

209.

Voucher: Oliver 9052.

The rather unusual condition of the South

African specimens of F. antipyretica not show-

ing the keeled leaf condition, unless wet, has no

doubt masked the presence of F. squamosa in

South Africa. More collections of Fontinalis

may extend the range of this exotic taxon to

other areas where F. antipyretica is known to

occur.

Fontinalis squamosa is very similar to the

more widespread species F. dalecarlica B.S.G.

The two species differ in the development of

leaf marginal cells. The specimens of F.

squamosa show little or no change in cell size

toward the margins, but exhibit a faint yellow

tint on the 1-3 cells along the leaf margins. The

cells of F. dalecarlica become slightly narrower

toward the margins and do not exhibit a colour

change. F. dalecarlica is not found in the United

Kingdom, the apparent source of the introduced

fish.

538

WARDIACEAE

The family Wardiaceae, containing a single genus and species, is endemic to the southwestern

Cape. The plants are found on rock in mountain streams and are recognized by their blackish

colour, mostly ecostate leaves and enlarged alar cells. The alar cells are fragile and frequently erod-

ed away by stream action or left on the stem when leaves are removed.

WARDIA

Wardia Harv. & Hook, in Companion Bot. Mag. 2: 183 (1837); Broth, m Natiirl. PflFam., edn 2,

1 1; 55 (1925); Sim, Bryo. S. Afr. 353 (1926). Type species; W. hygrometrica Harv. & Hook.

Plants in loose mats, usually blackish green; aquatic and saxicolous. Stems branched above

stipe; central strand absent. Leaves erect to spreading wet, appressed dry; variable in size and

shape; acute to cuspidate; margins plane, entire. Costa highly variable, absent, present only in base,

discontinuous and present at base and apex only, or rarely strong and extending from base to apex;

in section cells thickened, not differentiated. Laminal cells linear-fusiform, thickened; alar cells

strongly differentiated, enlarged and inflated, hyaline to yellowish.

Dioicous or rarely autoicous. Seta short and thick. Capsule exserted, erect, systylous. Peristome

very short, exostome teeth truncate. Operculum curved-rostrate, attached to columella, persistent.

Calyptra small, cucullate. Spores large, finely granulate.

Wardia hygrometrica Harv. & Hook, in

Companion Bot. Mag. 2: 1 83 ( 1 837); Sim, Bryo.

S. Afr. 354 (1926); Broth, in Natiirl. PflFam.,

edn 2, 1 1: 56 (1925); Welch in Bryologist 46: 27

(1943). Type: Cape of Good Hope, Table

Mountain, Harvey s.n.

Fontinalis duthieae Dix. in Sim, Bryo. S. Afr. 354

(1926); Welch in Bryologist 50: 187 (1947). Syntypes:

South Africa, Platteklip Rock, Cape Town, Sim 9389; Table

Mt. Sim 9385 (PRE).

Plants small to large, forming mats, dark

green or yellow-green to blackish green; aquatic

and saxicolous, in splash zone or submerged.

Stems naked and black below when old,

suberect, 15-80 mm long, branching irregular

above stipe, subfastigiate; in section round to

oval, central strand absent, inner cortical cells

large, thin-walled, hyaline, in 5 or 6 rows, outer

cortical cells smaller, yellowish, thick-walled, in

5 or 6 rows, epidermis absent; axillary hairs not

seen; paraphyllia absent; pseudoparaphyllia

absent. Leaves evenly spaced, spreading to

erect-appressed wet, appressed dry, ± concave;

on stipe lanceolate to triangular, 1. 5-2.0 mm

long; acute; margins plane, entire, eroded; upper

stem and branch leaves variable; narrowly or

broadly elliptical or short and broadly oblong,

1.2-2. 2 mm long; acute to narrowly acuminate

or occasionally mucronate to apiculate; rounded

at base; margins plane, entire, cells not differen-

tiated. Costa variable, absent, present only at

base, present and discontinuous only at apex and

base, or single and strong throughout, smooth;

ventral and dorsal surface cells elongate; in sec-

tion flat below, round above, guide cells not dif-

ferentiated, in best development consisting of

3-6 rows of undifferentiated, thickened cells;

most leaves with bistratose region at extreme

base and thickened awn at apex. Upper laminal

cells linear-fusiform and somewhat sigmoid,

homogeneous, 57-89 pm long, 7-10 pm wide,

walls thickened or rarely thin, smooth; basal

cells linear to long-rectangular, 50-125 pm

long, 7-10 pm wide, brownish yellow, walls

incrassate, smooth; alar cells strongly differenti-

ated, forming distinct groups, enlarged and

inflated, hyaline, walls thin.

Dioicous or rarely autoicous. Perichaetia at

apex; perichaetial leaves numerous, broadly

ovate, apex acute, sheathing below. Seta 4-6

mm long, yellow-brown to blackish, twisted

clockwise when dry, smooth, thick. Capsule

Wardiaceae

539

exserted, erect, short-cylindrical or rarely

ovoid, 1 .0-2.2 mm long, smooth but somewhat

ribbed when young and dry, yellow-brown to

black; urn short-cylindrical; neck not differenti-

ated; exothecial cells irregular, rounded to

quadrate or hexagonal, walls incrassate, cells at

mouth quadrate, darker; annulus not differenti-

ated, neck cells hexagonal to rectangular, thick-

ened; stomata absent. Peristome rudimentary,

yellow to orange; exostome teeth irregular,

short-rectangular, truncate, erect, 40-50(-95)

jam high, smooth with irregular prostome devel-

opment; endostome absent. Operculum curved-

rostrate, attached to columella, persistent,

1.0- 1.3 mm long. Calyptra not seen. Spores

rounded, 25-31 (-37) pm, granulate, brown.

Fig. 150.

Endemic to southern Africa, W. hygrometri-

ca is found in mountain streams of the south-

western Cape. Most specimens have been col-

lected on Table Mountain, fertile ones mostly

between October and February. Map 210.

Vouchers: Bews 8498; Crosby & Crosby 8185;

Esterhuysen 24545; Thome PRE-CH3442.

Although presently restricted in distribution,

W. hygrometricci expresses a high degree of

gametophytic variability. Plants show variation

in stem length and firmness, as well as the stage

at which stipes develop. This variation in habit

appears to be related to the age of individual

plants, younger plants being small, flaccid and

without a differentiated stipe, while older plants

are long, stout and frequently have well devel-

oped stipes. Variation in leaf size and shape is

frequently considerable between individual col-

lections. Much of the gametophytic variation

expressed by this species could be a result of the

semi-aquatic nature of the plants.

Fig. 150. — Wardia hygrometrica: 1. habit (dry), x 1; 2.

habit (wet), x 3; 3. stem in cross section, x 175; 4. ecostate

leaf, x 35; 5. costate leaf, x 35; 6. part of ecostate leaf in

cross section, x 175; 7. part of costate leaf in cross section,

x 175; 8. basal leaf cells (right side), x 175; 9. cells at leaf

apex, x 175; 10. perichaetial leaf, x 35; 11. distal part of

sporophyte (dry), x 10; 12. calyptra, x 17; 13. spore, x 700.

(1, 2 & 11, Wager CHU690\ 3, 4, 6, 8 & 9, Van Zanten

7608227\ 5 & 7, no collector given, CH8648\ 10 & 13,

Harrison BOL 249 17; 12, Sim 9257.)

540

WARDIACEAE

Map 210. — ♦ Wardia hygrometrica

• Hedwigia ciliata

ed as discontinuous and completely lacking

from the middle of the leaf. More recently spec-

imens have been found ( PRE-CH8648 ) with

well developed costae, extending from base to

apex. This is the first report of specimens with

fully developed costae.

The sporophyte is also adapted to the aquatic

habitat as seen in the short, stout setae, and

absence of stomata. The operculum remains

attached to the columella after the capsule has

opened. At the point of attachment, inside the

base of the operculum, is a large plug of tissue

that is normally broken down during the final

stages of peristome development. This is in part

responsible for the short, truncate peristome

teeth that are just visible at the capsule mouth.

The presence of a costa in some specimens

of Wardia was first noted by Allen (1987) when

he discovered a small bistratose area at the

extreme base of some leaves. He further found

that the discoloured acumen in most leaves was

multistratose. The costa was therefore interpret-

The type of Fontinalis duthieae Dix. in Sim

was found to be a light-coloured, somewhat

attenuate specimen of W. hygrometrica. The

leaves illustrated by Sim are not outside the

range of those found on other specimens of W.

hygrometrica ; furthermore the laminal and alar

cells clearly indicate the relationship of his

specimens.

541

HEDWIGIACEAE

Plants medium to large, erect to creeping, grey-green or yellow-green to green, occasionally

with reddish tint; saxicolous, corticolous or terricolous. Stems ± julaceous, freely branched; in sec-

tion central strand absent or weak; paraphyllia absent; pseudoparaphyllia absent or filamentous.

Leaves appressed dry, widespreading wet, frequently with hyaline or reddish apex; broadly ovate

to elliptical; acute to acuminate or sometimes piliferous; margins recurved below, entire to serru-

late, frequently decurrent at base. Laminal cells quadrate to rectangular or sometimes fusiform,

generally incrassate and pitted, papillose or granulose; basal cells longer; alar cells quadrate, some-

times strongly differentiated.

Autoicous or dioicous. Perigonia terminal or axillary, gemmate. Perichaetia terminal on stems

or short branches, not strongly differentiated; perichaetial leaves imbricate, oblong, margins entire,

serrulate or ciliate. Seta short or long. Capsule immersed or exserted, cupulate to cylindrical, occa-

sionally weakly ribbed, gymnostomous. Operculum rostrate, beak curved. Calyptra cucullate.

Spores small to medium-sized, brown or greenish.

A small mostly southern hemisphere family of five genera, four of which occur in the Flora

area. The genera are generally recognized by their extensive, loose tufts formed over exposed rock

surfaces at higher elevations.

1 Leaf cells granulate, alar cells strongly differentiated 1 . Rhacocarpus

1 Leaf cells papillose, alar cells not strongly differentiated:

2 Leaf apex hyaline, at least at stem tips:

3 Plants grey-green, leaf cell papillae large, 1 or 2 per cell; perichaetial leaf margins ciliate;

capsule immersed 2. Hedwigia

3 Plants green to yellow-green; leaf cell papillae small, scattered over surface;

perichaetial leaves entire; capsule exserted 3. Braunia

2 Leaf apex green or yellow-green:

4 Leaves somewhat plicate when dry; capsule exserted 3. Braunia

4 Leaves not plicate when dry; capsule immersed 4. Hedwigidium

1. RHACOCARPUS

Rhacocarpus Lindb., in Ofvers. Forh. Kongl. Svenska Vetensk.-Akad. 19: 607 (1863); Sim, Bryo.

S. Afr. 352 (1926); Broth, in Natiirl. PflFam., edn 2, 11: 73 (1925); Sainsb., N. Zeal, mosses 334

(1955). Type species: R. humboldtii (Hook.) Lindb.

Harrisonia Spreng., Syst. veg. 4,1: 145 (1827), hom. illeg.

Plants large, creeping, yellow-green to grey-green, frequently tinted with red; saxicolous or ter-

ricolous. Stems frequently highly branched; central strand absent. Leaves ovate to oblong, fre-

quently with reddish hair-point; margins recurved below, serrulate above; ecostate. Laminal cells

fusiform, densely granulate, incrassate and pitted; marginal cells smooth; alar cells strongly differ-

entiated, enlarged, coloured.

Dioicous. Capsule exserted, weakly ribbed. Peristome absent. Operculum rostrate. Calyptra

cucullate, reddish. Spores medium-sized.

Rhacocarpus contains 22 species that are, with the exception of a single Central American

species, restricted to the southern hemisphere. Except for the gymnostomous capsules, the genus

542

Hedwigiaceae

Hedwigiaceae

543

has little in common with other genera of the family. The single southern African species is rather

restricted in its local distribution, but widely distributed throughout the hemisphere.

Rhacocarpus purpurascens (Brid.) Par .,

Index bryol. suppl. 292 (1900); Broth, in Natiirl.

PflFam., edn 2, 11: 75 (1925); De Sloover in

Bull. Jard. Bot. Belg. 43: 343 (1973); Scott &

Stone, Moss. S. Austr. 356 (1976). Type:

Reunion.

Hypnum purpurascens Brid., Muscol. recent, suppl. 2:

121 (1812).

Harrisonia breuteliana C. Mull, in Oesterr. Bot. Z. 47:

392 (1897). Rhacocarpus breutelianus (C. Mull.) Broth, in

Natiirl. PllFam. 1,3: 720 (1905). Type: South Africa,

Saldanha Bay, Breutel 1862.

Harrisonia ecklonianci C. Mull, in Oesterr. Bot. Z. 47:

398 (1897). Rhacocarpus ecklonianus (C. Mull.) Broth, in

Natiirl. PflFam. 1,3: 722 (1905); Sim, Bryo. S. Afr. 352

(1926). Syntype: South Africa, Cape Town, Table

Mountain, Rehmann 197, 314 (PRE).

Harrisonia gracillima C. Mull, in Oesterr. Bot. Z. 47:

391 (1897). Rhacocarpus gracillimus (C. Mull.) Broth, in

Natiirl. PflFam. 1,3: 720 (1905). Type: South Africa, Cape

Town, Table Mountain, Rehmann 312.

Harrisonia rehmanniana C. Mull, in Oesterr. Bot. Z. 47:

391 (1897). Rhacocarpus rehmannianus (C. Mull.) Wijk et

Marg. in Taxon 9: 52 (1960); Sim. Bryo. S. Afr. 353 (1926).

Syntypes: South Africa, Cape, Table Mountain, Rehmann

313; Ecklon s.n.; Montague Pass, Rehmann, Oct. 1875;

Saldanha Bay, Breutel s.n., 1862 (G?).

Plants medium to large, forming large, loose

tufts, yellow-green to grey-green, frequently

tinged with red, yellow at apex, brown to black

below; saxicolous or terricolous. Stems creep-

ing, to 100 mm long, branches numerous; in

section round, central strand absent, inner cor-

tical cells large, hyaline, in 3-5 rows; outer cor-

tical cells smaller, red-yellow, thick-walled, in

2 or 3 rows, axillary hairs few, 2 or 3 cells long,

basal cells brown; pseudoparaphyllia absent.

Leaves evenly spaced, widespreading wet,

appressed with reflexed tips dry; stem leaves

ovate to oblong, 1. 5-2.0 mm long; acute to

acuminate; apex frequently hair-pointed, awn

reddish, weakly narrowed to insertion; margins

recurved and reddish below, plane to broadly

inflexed above, serrulate above, faintly bor-

dered; ecostate. Upper laminal cells fusiform,

somewhat sigmoid, heterogeneous, 12-75 pm

long, 6-12 pm wide, walls incrassate and pit-

ted, densely granulate on both surfaces, border

cells in 2-16 rows, narrower and smooth,

thicker in section; basal cells somewhat longer

but not strongly differentiated, 60-100 pm

long, 8-12 pm wide, greenish yellow, reddish

across insertion, walls incrassate and pitted,

densely granulate; alar cells strongly differenti-

ated and forming distinct groups, quadrate to

short rectangular, reddish, smooth, thin-walled.

Perigonial leaves ovate-acute, 1 mm long.

Perichaetial leaves oblong-acuminate, imbri-

cate, 2. 5-3.0 mm long, margins weakly serru-

late; leaf cells rectangular, incrassate and pitted.

Seta 6 mm long, yellow-brown, smooth.

Capsule exserted, erect, ellipsoid, 2 mm long,

weakly ribbed, yellow-brown, gymnostomous;

neck shorter than urn; exothecial cells rounded

to quadrate, walls thin, cells at mouth quadrate

to transversely short-rectangular, neck cells

smaller, quadrate, thickened; stomata on neck,

numerous, phaneropore. Operculum conic-ros-

trate, beak curved, 0.5 mm long. Calvptra

cucullate, reddish, 1 mm long, smooth. Spores

not seen. Fig. 151: 1-11.

Rhacocarpus purpurascens is found in

Central and South America, the West Indies,

eastern and southern Africa, the East African

islands, Australia and New Zealand and some

southern temperate and subantarctic islands. In

southern Africa the plants are rather restricted in

Fig. 151. — Rhacocarpus purpurascens (1-11): 1. habit (dry), x I ; 2. habit (wet), x 5; 3. stem in cross section (cells

partly shown), x 175; 4 & 5. leaves, x 35; 6. leaf in cross section, x 175; 7. basal leaf cells (right side), x 175; 8. upper lam-

inal cells, x 350; 9. upper laminal cells at right margin, x 350; 10. leaf apex, x 175; 11. perichaetial leaf, x 32. Hedwigia

ciliata (12-21); 12. habit (dry), x 1; 13. habit (wet), x 5; 14. part of stem in cross section, x 175; 15. leaf, x 25; 16. leaf in

cross section, x 175; 17. basal leaf cells at right margin (papillae partly shown), x 175; 18. upper laminal cells, x 350; 19.

leaf apex, x 175; 20. perichaetial leaf, x 25; 21. spore, x 700. (1, 5-7, 9 & 11, Esterhuysen 15873; 2, Bews PRE-CH8665;

3, 4, 8 & 10, Magill 4361 ; 12 & 13, Hilliard & Burtt 11841a; 14-19, Magill 5674; 20 & 21, Magill 7362.)

544

Hedwigiaceae

distribution. The species is found on wet rock or

soil, frequently associated with Sphagnum L., in

mountains of the northern and eastern Transvaal

regions, Lesotho and the southern and south-

western Cape. Map 211.

Vouchers: Anderson 1204; Boucher 3658;

Esterhuysen 15873; Magill 4361, 6324; Magill

& Schelpe 4061.

This magnificent moss is easily recognized

by its distinctive coloration, densely granulate

cell ornamentation and pronounced alar regions.

The variations expressed by the plants, i.e. apex

angle and hair-point development, width of

smooth-celled leaf border, branching pattern and

plant size, do not seem to be correlated.

However, subspecific names may be required

when the species is examined on a worldwide

basis. For example, the leaf apex is quite vari-

able in southern Africa. It can be abruptly

acuminate with a rather broad acumen filled

with thickened and pitted laminal cells, or nar-

row and extending out into a short or long awn.

The fertile specimens were either too young

or too old and missing the capsule, and there-

fore spores were not found. They have been

described elsewhere as 20-22 pm in diameter

and finely papillose.

2. HEDWIGIA

Hedwigia P. Beauv. in Mag. Encycl. 5: 304 (1804), nom. cons.; Sim, Bryo. S. Afr. 348 (1926);

Broth, in Natiirl. PflFam., edn 2, 11: 67 (1925); Scott & Stone, Moss. S. Austr. 352 (1976); Smith,

Moss FI. Brit. Irel. 492 (1980); Crum & Anderson, Moss. E.N. Amer. 2: 744 (1981). Type species:

H. ciliata (Hedw.) P. Beauv.

Plants large, glaucous green; saxicolous. Stems erect or prostrate, irregularly branched; central

strand small. Leaves crowded, patent wet, broadly elliptical, concave, acuminate apex hyaline,

ecostate. Leaf cells irregularly rectangular, incrassate, papillose by single, simple or branched

papillae centred over lumens; papillae 2-6 and seriate on linear, interior basal cells.

Autoicous. Seta very short. Capsule immersed, cupulate. Peristome absent. Operculum convex.

Calyptra cucullate. Spores small, brown.

A genus generally considered to contain only one species, H. ciliata. The species does exhibit

much variability and this has resulted in the description of a large number of subspecific taxa. The

genus is rather widely distributed on exposed rock in temperate forest throughout the world.

Hedwigia ciliata (Hedw.) P. Beauv., Prodr.

aetheogam. 15 (1805); Smith, Moss FI. Brit.

Irel. 492 (1978); Scott & Stone, Moss. S. Austr.

354 (1976); Catcheside, Moss. S. Austr. 295

(1980). Type: North America.

Hedwigia albicans Lindb. in Ofvers. Forh. Kongl.

Svenska Vetensk.-Akad. 21: 421 ( 1864); Sim, Bryo. S. Afr.

349 (1926); Broth, in Natiirl. PflFam., edn 2, 11: 68 (1925).

Type: not given.

Hedwigia macowaniana C. Mull, in Flora 71: 415

( 1888). Type: South Africa, Cape, Somerset East, MacOwan

s.n.

Aniclangium ciliatum Hedw., Sp. muse, frond. 40 (1801).

Hedwigiaceae

545

Hedwigia macowaniana C. Mull, in Hedwigia 38: 122

(1899), hom. illeg. Hedwigia ciliata fo. macowaniana (C.

Mull.) Fleisch. in Hedwigia 61: 403 (1920). Type: South

Africa, Cape, Mt. Boschberg, MacOwan s.n., July 1877

(GRA, H).

Hedwigia macowani C. Mull, ex Dix. & Gepp in Bull.

Misc. Inform. 1923: 229 (1923), nom. illeg. Type: South

Africa, ‘Musci MacOwanianis No. 19’; sub Rehmann 596,

596b, 596c (PRE).

Plants large, forming extensive, loose tufts,

glaucous, grey-green to yellow-green; saxi-

colous. Stems julaceous, erect to pendent, to 70

mm long, branching irregular; in section round,

central strand not well defined, inner cortical

cells thin-walled, yellow, in 4—6 rows, outer cor-

tical cells thick-walled, smaller, red-brown, in 2

or 3 rows; pseudoparaphyllia filamentous.

Leaves crowded, patent wet, appressed with

weakly spreading hyaline tips dry, concave, fre-

quently secund in prostrate plants; broadly ellip-

tical to ovate, 2.0-2. 5 mm long; acuminate; apex

hyaline, spinose papillate to almost smooth;

weakly decurrent at base; margins plane to nar-

rowly reflexed below, plane above or narrowly

recurved at transition to hyaline acumen, entire;

ecostate. Upper laminal cells irregularly rectan-

gular, homogeneous, 10-25 pm long, 2-10 pm

wide, walls incrassate, pitted, papillose on both

surfaces; papillae single, centred over lumen,

simple or branched; basal cells rectangular

toward margins, linear in centre and strongly

differentiated, 30-150 pm long, 6-12 pm wide,

yellow to brownish, walls incrassate, pitted and

irregular, papillose, papillae 2-6, seriate and

centred over lumens; alar cells quadrate, brown-

ish, walls thickened, smooth.

Perigonial leaves ovate-acuminate, 1 mm

long. Perichaetial leaves imbricate; oblong to

elliptical, 2.5-3.5 mm long; acuminate; margins

with long, hyaline, ciliate hairs; leaf cells irre-

gularly rectangular to angular, incrassate above,

rectangular below, thickened, pitted. Seta up to 1

mm long, yellow, smooth. Capsule immersed,

erect, gymnostomous, oblate to cupulate, 1-2

mm long, smooth, yellow to yellow-brown; neck

shorter than um; exothecial cells ± rectangular to

angular, walls thickened, cells at mouth becom-

ing smaller, rounded and collenchymatous, in

10-15 rows, neck cells irregular rectangular,

thin-walled; stomata not seen. Operculum con-

vex and minutely beaked, 0.5 mm long. Calyptra

small, cucullate. Spores rounded to angular,

22-26 pm, verruculose, brown. Fig. 151: 12-21.

Widespread on rocks in forest or forest mar-

gins, frequently in exposed situations, H. cilia-

ta is known throughout the temperate to arctic

northern hemisphere. Central and South Ame-

rica, eastern and southern Africa and Mada-

gascar, Australia, Tasmania and New Zealand.

In southern Africa this species is uncommon in

the northern and eastern Transvaal regions,

KwaZulu-Natal, Lesotho and the eastern and

central Cape areas. Map 210.

Vouchers: Hilliard & Burtt 11841; Magill 6947,

7424; Van Rooy & Perold 3857.

Hedwigia ciliata is most easily identified by

its grey-green colour, large leaf cell papillae and

pronounced hyaline leaf apex. Specimens are

uncommon but usually cover large areas on

exposed boulders in forest openings or margins.

The plants are generally collected with sporo-

phytes, a condition that is not as common in

other members of the family. The cup-shaped

capsules are immersed, but their orange urns and

reddish mouths are generally apparent through

the large, ornately ciliate perichaetial leaves.

3. BRAUNIA

Braunia B.S.G. , Bryol. eur. 3: 159 (1846); Sim, Bryo. S. Afr. 350 (1926); Broth, in Natiirl. PflFam.,

edn 2, 11: 70 (1925). Type species: B. sciuroides (Bals. & De Not.) B.S.G.

Plants large, loosely caespitose; saxicolous or corticolous. Stems suberect, branched; central

strand present. Leaves widespreading wet, broadly elliptical, ± plicate; margins narrowly recurved;

apex frequently hyaline; ecostate. Laminal cells quadrate to rectangular, thickened, papillose; inte-

rior basal cells linear, seriate papillose.

Hedwigiaceae

547

Autoicous. Seta elongate, smooth. Capsule long-exserted. Peristome absent. Operculum ros-

trate, beak curved. Calyptra cucullate. Spores small, brown.

Braunia contains 23 species scattered in the Americas, Africa and Asia. One species, B. secun-

da, found throughout the range of the genus, is known from southern Africa.

Braunia secunda (Hook.) B.S.G., Bryol.

eur. 3: 161 (1846); Sim, Bryo. S. Afr. 350

(1926); Broth, in Natiirl. PflFam., edn 2, 11: 71

(1925); Grout, Moss FI. N. Amer. 2: 44 (1933).

Type: Mexico, Humboldt & Bonpland s.n.

Hedwigia secunda Hook., Musci exot. 1: 46 (1818).

Anictangium secundum (Hook.) Hook, in Kunth, Syn. pi. 1:

47 (1822).

Neckera diaphana C. Mull.. Syn. muse, frond. 2: 105

(1850). Braunia diaphana (C. Mull.) Jaeg. in Ber. Thatigk.

St. Gallischen Naturwiss. Ges. 1874—1875: 172 (1876).

Type: Prom. bon. spei, Pappe s.n.

Braunia secunda var. pinnata Sim, Bryo. S. Afr. 351

(1926). Type: South Africa, Natal, Giant’s Castle, 1915,

Symons s.n. sub Sim 10266 (PRE).

Plants large, forming extensive tufts, green

to yellow-green; saxicolous or corticolous.

Stems suberect, 20-70 mm long, branching

irregular; in section round, central strand

absent, inner cortical cells thick-walled, yellow,

in 5-7 rows, outer cortical cells smaller, thick-

walled, red-brown, in 2 or 3 rows; pseudopara-

phyllia absent. Leaves evenly spaced, patent to

widespreading wet, weakly appressed dry, pli-

cate; broadly elliptical to ovate-lanceolate,

2. 0-2. 5 mm long; acute to acuminate; apex fre-

quently hyaline and papillose, especially at

stem tips; base weakly decurrent; margins nar-

rowly recurved frequently throughout, entire

below, sometimes toothed at apex; ecostate.

Upper laminal cells quadrate to rectangular, fre-

quently irregular, homogeneous, 7-16 pm long,

6-8 pm wide, walls incrassate, papillose on

both surfaces, frequently almost smooth above;

papillae numerous, low, scattered; basal cells

quadrate to short rectangular marginally, linear

and forming distinct group internally, 60-100

pm long, 8-12 pm wide, yellowish, walls thick-

ened, weakly pitted, papillose, papillae low,

seriate; alar cells not strongly differentiated.

Perigonial leaves ovate-acute, 1.0-1. 2 mm

long. Perichaetial leaves oblong-acuminate, 3

mm long; margins entire; leaf cells quadrate to

short rectangular above, papillose, rectangular

to linear below, smooth, incrassate and pitted.

Seta 10-15 mm long, yellow-brown, smooth.

Capsule exserted, ± erect, gymnostomous,

cylindrical, 2. 0-3. 5 mm long, smooth, yellow-

brown, neck shorter than urn; exothecial cells

rounded, quadrate to oblong, walls thickened,

cells at mouth darker, quadrate, thickened, neck

cells not differentiated; stomata numerous on

neck, phaneropore. Operculum short- or long-

beaked, curved, 1 mm long. Calyptra cucullate.

Spores angular, 22-27 pm, papillose, brownish

green. Fig. 152: 1-10.

Braunia secunda is known from the south-

western United States, the West Indies through

Central America to Bolivia and northern Argen-

tina, northern, southern and eastern Africa and

India. In the Flora area the species is found on

boulders and at the base of trees in open sites in

forests or shaded kloofs of the northern and

eastern Transvaal regions, Swaziland, Zululand,

KwaZulu-Natal, and the central, eastern, south-

ern and southwestern Cape. Map 212.

Vouchers: Abbott 7112; Esterhuysen 17678;

Hilliard & Burtt 17139; Magill 3485 , 6951;

Meyer 2516g.

Specimens of Braunia secunda are quite vari-

able but are generally easily placed when they

Fig. 152. — Braunia secunda (1-10): 1. habit (dry), x 1; 2. habit (wet), x 3; 3. part of stem in cross section, x 175; 4.

leaf, x 35; 5. leaf in cross section, x 175; 6. basal leaf cells (left side), x 175; 7. upper laminal cells, x 350; 8. leaf apex, x

175; 9. perichaetial leaf, x 35; 10. spore, x 700. Hedwigidium integrifolium (11-21): 11. habit (dry), x 1; 12. habit (wet),

x 3; 13. part of stem in cross section, x 175; 14. leaf, x 35; 15. leaf in cross section, x 175; 16. basal leaf cells (right side),

x 175; 17. upper laminal cells, x 350; 18. cells at leaf apex, x 175; 19. perichaetial leaf, x 32; 20. operculum, x 35; 21.

calyptra, x 35. (1-10, Sipman 20021\ 11 & 13-20, Van Rooy 2270\ 12, Scheepers 1228.)

548

Hedwigiaceae

Map 212. — Braunia secunda

form large, yellow-green carpets over exposed

boulders in forest openings. The ecostate leaves

generally have a hyaline apex, especially at the

stem tips, although never as pronounced as in

Hedwigia ciliata. The leaves are also generally

distinctly plicate when dry, a condition that sep-

arates this species from H. ciliata and

Hedwigidium integrifolium (see below).

Fertile specimens are uncommon, but are

easily separated from Hedwigidium and Hed-

wigia by their long-exserted, cylindrical cap-

sules. The dryness of the open rock habitat may

account in part for the rarity of sporophytes, an

unusual fact considering the autoicous nature of

the plants.

4. HEDWIGIDIUM

Hedwigidium B.S.G., Bryol. eur. 3: 155 (1846); Sim, Bryo. S. Afr. 349 (1926); Broth, in Natiirl.

PflFam., edn 2, 11: 69 (1925). Type species: H. imberbe (Sm.) B.S.G.

Plants generally large, forming extensive loose tufts, yellow-green; generally saxicolous. Stems

suberect to creeping, branches few; central strand absent. Leaves patent wet, appressed dry, ellip-

tical-acuminate, apex green; margins weakly recurved below, entire, decurrent; ecostate. Laminal

cells rectangular, incrassate, walls frequently wavy.

Autoicous. Seta short. Capsule immersed, cupulate. Peristome absent. Operculum conic-ros-

trate, beak curved. Calyptra cucullate, smooth. Spores brownish.

A genus with a single species found in Central and South America, Europe, southern Asia and

India, central and southern Africa, Australia and New Zealand. The species has been treated by sev-

eral authors (Smith 1978) as belonging to Hedwigia , to which it is related.

Hedwigidium integrifolium (P. Beauv.)

Dix. in C. Jens., Skand. Bladmossfl. 369 (1939).

Type: North America.

Hedwigia integrifolia P. Beauv., Prodr. aetheogam. 60

(1805); Smith, Moss FI. Brit. Irel. 493 (1978); Scott &

Stone, Moss. S. Austr. 355 (1976).

Gymnostomum imberbe Sm., Engl. bot. 32: 2237

(1811). Hedwigidium imberbe (Sm.) B.S.G., Bryol. eur. 3:

157 (1846); Sim, Bryo. S. Afr. 349 ( 1926). Hedwigia imber-

bis (Sm.) Sprue., Muse, pyren. 263 (1847). Type: Ireland,

Hutchins, 1809.

Braunia macowaniana C. Mull, in Hedwigia 38: 123

(1899). Syntypes: South Africa, Cape, Somerset East, Mt

Boschberg, MacOwan s.n.; Natal, Jammerlappen, Dittrich,

1898 (PC).

Braunia erosa C. Mull, in Hedwigia 38: 124 (1899).

Type: South Africa. Cape Prov., Cape Town, near Ronde-

bosch, 1875, Rehmann s.n. (BM).

Braunia maritima C. Mull, in Hedwigia 38: 124 (1899).

Hedwigidium maritimum (C. Mull.) Par., Index bryol. suppl.

179 (1900). Syntypes: South Africa, Cape Prov., Table

Mountain, Rehmann 306, 307\ Transvaal, Lydenburg, Wilms

s.n., 1887 (BM).

Plants medium to large, forming large tufts,

yellow-green to yellow-brown; saxicolous or

humicolous. Stems suberect to creeping, up to

50 mm long, branches few, irregular; in section

round, central strand absent, inner cortical cells

large, yellowish, in 3 or 4 rows, outer cortical

Hedwigiaceae

549

cells smaller, thick-walled, reddish; axillary

hairs few, 3 cells long, basal cell brown;

pseudoparaphyllia absent. Leaves crowded,

patent wet, appressed dry; elliptical to lanceo-

late, 1. 5-2.5 mm long; acuminate, decurrent at

base; margins plane or recurved to above

midleaf, entire; ecostate. Upper laminal cells

rectangular or quadrate, homogeneous, 8-18

pm long, 5-7 pm wide, walls incrassate, fre-

quently wavy, weakly papillose; papillae low,

scattered; basal cells strongly differentiated

internally, linear, 35-75 pm long, 5-8 pm wide,

reddish yellow, walls incrassate, pitted, smooth;

alar cells not strongly differentiated from lami-

nal cells, somewhat larger, quadrate to rectan-

gular, yellow-brown, walls thickened.

Perigonial leaves ovate-acuminate, 1 mm

long. Perichaetial leaves oblong; acute to

acuminate, 3^1 mm long; base sheathing; mar-

gins serrulate at midleaf; leaf cells quadrate to

short-rectangular and papillose above, linear and

smooth below. Seta 0.8- 1.0 mm long, brown,

smooth. Capsule immersed, erect, gymnosto-

mous, cupulate, 1.5 mm long, weakly ribbed,

yellow to brown; neck not differentiated;

exothecial cells rounded quadrate, walls thin,

cells at mouth darker, quadrate; stomata not

seen. Operculum conic-rostrate, beak curved,

0.5 mm long. Calyptra cucullate, 1 mm long,

smooth, yellow-brown. Spores rounded angular,

20-25 pm, granulate, brown. Fig. 152: 11-21.

A rather widespread species known from

Central and South America, Europe, western,

eastern and southern Africa and the Mas-

carenes, southern and southeastern Asia,

Australia and New Zealand. In the Flora area

H. integrifolium is found on exposed rock or

humus over rock in the northern and eastern

Transvaal region, Swaziland, KwaZulu-Natal,

Lesotho and the eastern, central, southern and

southwestern Cape. Map 213.

Vouchers: Esterhuysen 21233; Crosby & Crosby

8142; Magill 3518; Van Rooy 2270.

Vegetatively this species resembles Braunia

secunda , and some specimens are not complete-

ly distinguishable because of the variability

expressed by both species. The following char-

acters are useful for separating sterile speci-

mens of the two species. The leaves of H. inte-

grifolium are narrower, only rarely weakly pli-

cate when dry and have a yellow-green apex,

whereas the leaves of B. secunda are generally

broad, distinctly plicate when dry, and the apex

(at least of those leaves at the stem apex) is hya-

line. The leaf cells of H. integrifolium also seem

to be more regularly rectangular with wavy,

incrassate walls.

The sporophyte of H. integrifolium is com-

pletely immersed and could be easily over-

looked since the perichaetial leaves are not

strongly differentiated. The autoicous condition

should lead to a higher number of plants with

sporophytes than have been found on recent

collections.

The need for a separate genus for this species

has been questioned, since it shares a similar

habit and immersed gymnostomous, cupulate

capsules with Hedwigia. In addition, the mid-

leaf marginal serrulations on the perichaetial

leaves may suggest a relationship with the cili-

ate perichaetial leaves of Hedwigia. On the

other hand, important character differences ex-

hibited by Hedwigidium include the lack of a

central strand in the stems, lack of pseudopara-

phyllia, absence of a hyaline leaf apex and dif-

ferences in leaf cell papillae.

550

CRYPHAEACEAE

A family of eight genera found in forests of temperate and tropical regions. The family is rec-

ognized by its branched secondary stems, strongly costate leaves, generally with decurrent bases,

autoicous gametophytes, immersed capsules, and conical calyptrae. In the Flora area the family is

represented by a single genus, Cryphaea. The family as defined by Manuel (1973) is more homo-

geneous than before. However, close ties with genera in Leucodontaceae and Leptodontaceae exist,

without clear lines of separation.

CRYPHAEA

Cryphaea Mohr in Web., Tab. Calyptr. operc. 3 (1814); Broth, in Natiirl. PflFam., edn 2, 11: 77

(1925); Sim, Bryo. S. Afr. 355 (1926); Crum & Anders., Moss. E.N. Amer. 2: 747 (1981). Type

species: not designated.

Plants medium-sized, scattered or in very loose tufts, green to yellow-green; corticolous.

Secondary stems erect from creeping primary stem, loosely pinnate; in section central strand

absent. Leaves lanceolate, margins entire to serrate, frequently decurrent at base. Costa single,

strong to near apex. Lamina l cells short, weakly thickened, smooth; basal cells not strongly differ-

entiated; alar cells quadrate, in large groups.

Autoicous. Perichaetia numerous, along secondary stem. Seta very short. Capsule immersed,

cylindrical; stomata absent. Peristome single or double and incomplete; exostome teeth recurved

when wet. Operculum conical. Calyptra conical but split along one side. Spores large.

A genus of about 68 corticolous species found in forests of temperate and tropical regions. Most

species are found in Central America and northern South America. Eight species are known from

Africa and the African islands.

The calyptra has been used as one of the major characters separating Cryphaeaceae from

Leucodontaceae. The cryphaeaceous calyptra, described as conical or mitrate, has been regarded as

distinct from the cucullate calyptra of the Leucodontaceae and was the major reason Manuel (1974)

removed Forsstroemia from Cryphaeaceae. The southern African species of Cryphaea was found

to have a conical calyptra split along one side, a condition also found in other species of the genus.

Therefore the technical distinction between the split-conical calyptra of Cryphaea and the cucul-

late or hood-shaped calyptra of Leucodontaceae is not as clear as previously suggested.

The peristome movement of Cryphaea is opposite that generally encountered in mosses. While

most mosses have peristome teeth erect or reflexed when dry to facilitate spore dispersal by wind,

the peristome of Cryphaea is closed when dry and open when wet. The advantage to Cryphaea is

uncertain, but appears to be connected to the corticolous nature of the plants.

Cryphaea exigua (C. Mull.) Jaeg. in Ber.

Thatigk. St. Gallischen Naturwiss. Ges. 1 874 —

1 875 : 179 (1876); Broth, in Natiirl. PflFam.,

edn 2, 11: 79 (1925); Sim, Bryo. S. Afr. 356

(1926). Type: South Africa, Cape, Philipstown,

s.l. & s.n. (Herb. Gottschean).

Pilotrichum exiguum C. Mull., Syn. muse, frond. 2: 166

(1851).

Cryphaea dentata Mitt, in J. Linn. Soc., Bot. 22: 311

( 1 886). Type: South Africa, Natal, Umgoye Mt, Plant s.n. (NY).

Plants medium-sized, scattered or gregarious,

green to yellow-green; corticolous. Primary

stems creeping; secondary stems erect (perpen-

dicular to substrate), julaceous, up to 60 mm

long, branching regular, pinnate; in section

round, central strand absent, inner cortical cells

large, thin-walled, hyaline, in 4 or 5 rows, outer

cortical cells smaller, thick-walled, yellow, in 2

or 3 rows; axillary hairs short, basal cells differ-

entiated, brownish; paraphyllia absent; pseudo-

Cryphaeaceae

551

paraphyllia sparse, short and filamentous. Leaves

crowded, widespreading wet, loosely appressed

dry; ovate to lanceolate, 1.6-2. 6 mm long;

acuminate; rounded and narrowed to insertion,

decurrent; margins plane, almost entire to serrate

above. Costa single, ending in acumen or sub-

percurrent, smooth; ventral surface smooth, cells

elongate; dorsal surface smooth, cells elongate;

in section subround, bulging dorsally, 2 or 3 rows

of undifferentiated cells. Upper laminal cells

rhombic or rounded, homogeneous, 12-25 pm

long, 7-10 pm wide, walls weakly thickened,

smooth; basal cells not strongly differentiated,

linear, 30-40 pm long, 7-10 pm wide, walls

weakly thickened, smooth; alar cells forming

distinct groups, quadrate or transversely rectan-

gular, walls weakly thickened.

Perigonia on stem, gemmate; leaves ovate-

acuminate, to 1 mm long. Perichaetia strongly

differentiated; leaves sheathing, oblong, abrupt-

ly aristate, 2-3 mm long; leaf cells linear-

fusiform, weakly thickened. Seta 0.2 mm long,

whitish, smooth. Capsule immersed but visible

through perichaetial leaf awns, erect, cylindri-

cal, up to 1.5 mm long, smooth, brown; neck

not differentiated; exothecial cells somewhat

irregular, quadrate to short rectangular, walls

thin, deteriorating below with age, cells at

mouth darker, quadrate; neck cells not differen-

tiated; annulus present, vesiculose; stomata

absent. Peristome white to yellowish; exostome

teeth narrowly triangular, incurved and sigmoid

dry, recurved wet, papillose, 320-350 pm high;

endostome segments linear, almost as long as

teeth, papillose, basal membrane absent, cilia

absent. Operculum conical, erect, 0.4 mm long.

Calyptra campanulate and split along one side,

0.5-0. 7 mm long, smooth. Spores rounded,

22-31 pm, granulate, light brown. Fig. 153.

Fig. 153. — Cryphaea exigua: 1. habit (dry), x 1; 2.

upper part of secondary stem (wet), x 3; 3. stem in cross

section, x 175; 4. leaf, x 35; 5. leaf in cross section, x 175;

6. basal leaf cells, x 175; 7. upper laminal cells at right mar-

gin, x 350; 8. leaf apex, x 175; 9. perichaetial leaf, x 35; 10.

part of capsule mouth with peristome (papillae partly

shown), x 175; 11. operculum, x 35; 12. calyptra, x 35; 13.

spore, x 700. (1, 2, 5-7 & 11-13, Crosby 7931- 3, 4, 8 &

10, Magill 3758\ 9, B return 3270.)

552

Cryphaeaceae

Endemic to Africa, C. exigua is known from

eastern and southern Africa. In the Flora area it

is found on small trees in dry forests of the

northern Transvaal area, Swaziland, Zululand,

and the eastern, southern and central Cape

regions. Map 214.

Vouchers: Brenan 3270; Crosby & Crosby

7931 , 8066; Magill 3412, 3764.

Specimens of Cryphaea exigua can be iden-

tified by their habit, narrowly costate leaves

with serrate margins, and numerous immersed

capsules along erect stems. The leaf shape and

marginal serrations also separate sterile speci-

mens from mosses with similar growth forms,

like Forsstroemia or Leptodon (see p. 585).

Map 214. — Cryphaea exigua

553

LEUCODONTACEAE

Plants small to large, forming loose tufts, green to yellow-green or brownish; saxicolous or cor-

ticolous. Secondary stems erect, julaceous dry, variably branched; in section central strand present

or absent; paraphyllia absent; pseudoparaphyllia present or absent. Leaves appressed dry, wide-

spreading wet; broadly ovate to elliptical; acute to acuminate; margins plain, entire or serrate.

Costa absent, short and double, or single extending to above midleaf. Upper laminal cells short,

rhombic to fusiform, smooth or prorate; basal cells elongate; alar cells in large groups, quadrate to

transversely rectangular.

Dioicous. Perichaetia along stem, obvious. Seta short or long. Capsule generally exserted,

short-cylindrical. Peristome double; exostome teeth somewhat irregular, linear, whitish, papillose

or smooth; endostome rudimentary, frequently only basal membrane present. Operculum rostrate.

Calyptra cucullate, smooth or with a few hairs. Spores granulate.

Leucodontaceae is a family of eight genera found in woodlands and forests throughout the

Americas, Europe, Africa and parts of Asia. Much has been written in recent years about the rela-

tionships between Leucodontaceae and Cryphaeaceae, but consensus has not been reached as to the

proper division of genera between the families. The families are divided in the Flora on the basis

of calyptra development, capsule exsertion and costa type. Forsstroemia has been placed in both

families, but in this treatment it is moved to the Leptodontaceae (see p. 585).

Leaves ecostate; margins entire; leaf cell smooth 1. Leucodon

Leaves costate; upper leaf margins serrate; some upper leaf cells prorate on dorsal surface

2. Pterogonium

1. LEUCODON

Leucodon Schwaegr., Sp. muse, frond, suppl. 1,2: 1 (1816); Broth, in Natiirl. PflFam., edn 2, 11:

91 (1925); Sim, Bryo. S. Afr. 357 (1926); Gangulee, Moss. E. India 5: 1218 (1976). Type species:

L. sciuroides (Hedw.) Schwaegr.

Plants large, full or slender, in extensive loose tufts, green to yellow-green; saxicolous or corti-

colous. Stems little-branched, julaceous. Leaves generally appressed dry, widespreading wet; mar-

gins plain, entire. Costa variable or absent. Laminal cells short, thickened, usually smooth; basal

cells forming distinct group in lower leaf; alar cells quadrate, forming large distinct groups.

Perichaetia along stem, obvious. Seta long. Capsule exserted, short-cylindrical. Peristome dou-

ble. Operculum rostrate. Calyptra cucullate, smooth. Spores large.

Leucodon is a genus of 33 species found in the Americas, Europe, Africa and northern Asia. A

large number of species have been described from Asia, but the genus has not been reported from

the East Indies, Australia or New Zealand.

The southern African species is distinct from other members of the genus by its complete lack

of a central strand in the stem (see Manuel 1974). The absence of a costa and smooth rather than

prorate upper leaf cells are also notable characters in the southern African species, although these

conditions are found in other species as well.

Leucodon assimilis (C. Mull.) Jaeg. in Ber. 2, 11: 92 (1925); Sim, Bryo. S. Afr. 358 (1926).

Thatigk. St. Gallischen Naturwiss. Ges. 1 875- Syntypes: South Africa, Grootvadersbosch, Pappe

1876: 217 (1877); Broth, in Natiirl. PflFam., edn s.n.\ Adoi, Uitenhagen District, Ecklon s.n.

554

Leucodontaceae

Leucodontaceae

555

Neckera assimilis C. Miill., Syn. muse, frond. 2: 92

(1850).

Leucodon capensis Schimp. in Ren., Prodr. fl. bryol.

Madagascar 184 (1898). Syntypes: La Reunion, Cilaos,

Eudei, Madagascar, Ambatomanga, Talazac , 1894 (PC).

Braunia elliottii Broth, in Bot. Jahrb. Syst. 24: 253

(1897). Type: East Africa, Shire Highlands, Sotchi, Elliott s.n.

Braunia peristomata Dix. in S. African J. Sci. 18: 324

(1922). Syntypes: Zimbabwe, Zimbabwe Ruins, Sim 8750,

8778, 8793, 8809- Fort Victoria, Sim 8843 (BM, PRE).

Leucodon assimilis var. humilis Sim, Bryo. S. Afr. 358

(1926). Type: South Africa, Transvaal, Houtbosch, Reh-

mann 605 (PRE).

Plants medium to large, full or slender, form-

ing extensive, loose tufts, green to yellow-green

or brownish; corticolous or saxicolous. Primary

stems creeping; secondary stems ± erect, jula-

ceous, ± curved when dry, 40-100 mm long,

branching sparse, irregular; in section round,

central strand absent, inner cortical cells large,

thin-walled, hyaline, in 6-8 rows, outer cortical

cells smaller, thick-walled, yellowish red; para-

phyllia absent; pseudoparaphyllia absent.

Leaves crowded, ecostate, widespreading wet,

appressed or occasionally spreading dry; stem

leaves ovate to elliptical, 1.2-2. 5 mm long;

abruptly short-acuminate or acute; rounded and

narrowed to insertion; margins plane, entire.

Upper laminal cells rhombic or some fusiform,

homogeneous, 1 0— 25(— 34) pm long, 3-6 pm

wide, walls thickened, frequently strongly so,

smooth; basal cells forming distinct group near

centre of leaf, confined to lower leaf or occa-

sionally partly extending to above midleaf, lin-

ear, 50-90 pm long, 4—6 pm wide, yellowish,

walls incrassate and pitted, smooth; alar cells

forming distinct groups, quadrate to transverse-

ly rectangular, extending to midleaf, green,

walls weakly thickened.

Perigonia on stem, gemmate. Perichaetia

along stem, obvious; perichaetial leaves imbri-

cate, oblong to lanceolate 3-5 mm long, apex

acute, leaf cells linear. Seta 6-12 mm long, yel-

low-brown, smooth. Capsule exserted, erect,

short-cylindrical to ovoid, 2. 0-2. 5 mm long,

smooth, yellow-brown; neck not differentiated;

exothecial cells irregular, quadrate to rectangu-

lar, walls thin, cells at mouth darker, quadrate;

annulus absent; stomata on lower urn, phanero-

pore. Peristome double, whitish to yellowish

with age; exostome teeth irregular, lanceolate,

papillose, cleft and perforated, 200-250 pm

high; endostome segments absent, basal mem-

brane rudimentary, papillose, cilia absent. Oper-

culum curved-rostrate, 1 mm long. Calyptra

cucullate, 5 mm long, smooth. Spores rounded,

25-30 pm, granulate, brownish. Fig. 154: 1-12.

Found on rock and trees, Leucodon assimilis

is known from central, eastern and southern

Africa and the East African islands. In the Flora

area specimens were collected in forests of the

northern, central and eastern Transvaal regions,

Swaziland, Zululand, KwaZulu-Natal, eastern

Free State, and the eastern, southern and south-

western Cape. Map 215.

Fig. 154. — Leucodon assimilis (1-12): 1. habit (dry), x 1; 2. habit (wet), x 3; 3. part of stem in cross section, x 175; 4.

leaf, x 32; 5. leaf in cross section, x 175; 6. basal leaf cells (left side), x 175; 7. upper laminal cells, x 350; 8. cells at leaf

apex, x 175; 9. perichaetial leaf, x 35; 10. part of capsule mouth with peristome (papillae partly shown), x 175; 11. oper-

culum, x 32; 12. spore, x 700. Pterogonium gracile (13-22): 13. habit (dry), x 1; 14. habit (wet), x 3; 15. part of stem in

cross section, x 160; 16. stem leaf, x 32; 17. branch leaf, x 35; 18. branch leaf in proximal cross section, x 175; 19. branch

leaf in distal cross section, x 175; 20. basal cells of branch leaf, x 175; 21. upper laminal cells of branch leaf, x 320; 22.

branch leaf apex, x 160.(1 , Magill 3768\ 2-1 & 9-12, Magill 3701',%, Magill 3271\ 13& 14, \l-20, Magill 3216: 15, 16,

& 21, Esterhuysen 24670.)

556

Leucodontaceae

Vouchers: Crosby & Crosby 8036; Ellis

3091; Kemp 744; Magill 3701, 6511; Schelpe

7874.

This species is most easily recognized when

it forms large, loose tufts on boulders or tree

limbs. The julaceous stems are ± erect and have

few branches. The leaves are ecostate, but have

a pronounced group of elongated basal cells in

the lower centre of the leaf. Leucodon assimilis

is similar to the North American species, L.

julaceus (Hedw.) Sull. which differs by having

the upper laminal cells papillose on the dorsal

surface. Specimens of L. assimilis are frequent-

ly confused with Braunia secunda (p. 547).

However, that species has weakly plicate

leaves, generally hyaline and papillose leaf tips,

and gymnostomous capsules.

Specimens of the var. humilis described by

Sim have a distinctive habit, their leaves re-

maining widespreading when dry, rather than

becoming appressed. No other characters were

found that would support recognition of the

variety; both expressions are occasionally pre-

sent in single collections.

2. PTEROGONIUM

Pterogonium Sw. in Monthly Rev. 2, 34: 537 (1801); Broth, in Natiirl. PflFam., edn 2, 11: 98

(1925); Sim, Bryo. S. Afr. 360 (1926); Smith, Moss FI. Brit. Irel. 503 (1978). Type species: P.

gracile (Hedw.) Sm.

Plants medium-sized, forming loose tufts, yellow-green; saxicolous or corticolous. Secondary

stems erect from creeping primary stems, densely branched, stems and branches curved when dry;

central strand present. Leaves ovate; margins serrulate. Costa short and double. Laminal cells short,

thickened, sparsely prorate on upper dorsal surface; basal cells elongate; alar cells differentiated.

Sporophyte not known from Flora area but described as dioicous. Perichaetial leaves long-

sheathing. Capsule exserted, ellipsoidal. Peristome double; exostome teeth papillose-striate below;

endostome segments short above basal membrane, cilia absent. Operculum conical. Calyptra

cucullate, with a few hairs. Spores small, 14-20 pm.

A genus of three species, Pterogonium is found in the Americas, Europe and Africa. A single

species, P. gracile with several regional varieties, is known from throughout Africa and the African

islands. The genus is recognized by its distinct branching pattern and curved stems and branches

when dry.

Pterogonium gracile (Hedw.) Sm., Engl,

bot. 16: 1085 (1802); Dixon, Stud, handb. Brit,

mosses, edn 3: 404 (1924); Lawton, Moss FI.

Pacific Northwest 240 (1971). Type: England

and Scotland.

Pterigynandrum gracile Hedw., Sp. muse, frond. 80

(1801). Anomodon gracilis (Hedw.) Garov., Bryol. austr.

excurs. 46 (1840). Neckera gracilis (Hedw.) C. Mull., Syn.

muse, frond. 2: 97 (1850).

Grimmia ornithopodioides Web. & Mohr, Bot.

Taschenb. 148 (1807). Pterogonium ornithopodioides

(Web. & Mohr) Lindb. in Ofvers. Forh. Kongl. Svenska

Vetensk.-Akad. 20: 411 (1863); Broth, in Natiirl. PflFam.,

edn 2, 11: 99 (1925); Sim, Bryo. S. Afr. 360 (1926). Type:

Europe.

Pterogonium gracile var. capense C. Mull, ex Dix.,

Stud, handb. Brit, mosses, edn 3: 405 (1924).

Forsstroemia dendroides Dix. in Sim, Bryo. S. Afr. 360

(1926), nom. nud.

Plants small to medium-sized, forming loose

tufts, yellow-green to glaucous green; corti-

colous or saxicolous. Primary stems creeping,

naked; secondary stems erect, 15-35 mm long;

branching dense above stipe, subdendroid, fas-

ciculate to pinnate, rarely flagellate, distinctly

curved when dry; in section oval, central strand

small, inner cortical cells thin-walled, yellow, in

stipe 8-10 rows wide, 5 or 6 rows in branches,

outer cortical cells smaller, thick-walled, yel-

Leucodontaceae

557

low-brown, in stipe 6-8 rows wide, 3 or 4 rows

in branches; paraphyllia absent; pseudopara-

phyllia absent. Leaves crowded, erect-spreading

wet, appressed dry; leaves on stipe broadly

ovate, 1.0-1. 5 mm long; acute; rounded at base;

margins plane, weakly serrulate; branch leaves

ovate to elliptical, 1. 2-2.2 mm long; acute;

rounded to cordate at base; margins plane, ser-

rate. Costa short and double, occasionally broad

at insertion and with 2-4 short spikes; in section

flat, consisting of 3 rows of undifferentiated

cells. Upper laminal cells short-fusiform to

rhomboidal, somewhat sigmoid, homogeneous,

12-25 pm long, 7-8 pm wide, walls thickened,

with a few sharply prorate cells on upper dorsal

surface; basal cells oblong to linear juxtacostal-

ly, forming distinct groups, irregularly quadrate

to transversely rectangular, green, walls thick-

ened; alar cells in distinct group, quadrate.

Sporophyte not known in Flora area. Fig.

154: 13-22.

Pterogonium gracile, found in eastern Africa

and the East African islands, Macaronesia,

Europe, the Middle East and western North

America, forms large tufts on boulders or on the

base of trees. In southern Africa the species is

found in the northern and eastern Transvaal

areas, KwaZulu-Natal, and the eastern, southern

and southwestern Cape. Map 216.

Vouchers: Crosby & Crosby 9199; Ester-

huysen 19186; Magill 3287; Van Rooy 2280.

When dry the stems and branches are curved,

giving the plants a distinctive appearance. The

serrate upper leaves, short and double costa and

strongly differentiated alar cells help to identify

specimens of P. gracile.

The branch leaves have a few scattered pro-

rate cells on the upper dorsal surface. The papil-

lae are obvious, even at lower magnifications,

in micropreparations and should also aid in

identifications.

558

PRIONODONTACEAE

The family Prionodontaceae contains two genera that are found in tropical and south temperate

regions of the Americas, Africa and Asia. Only one of the genera, Prionodon , is known from forests

of western, eastern and southern Africa and Madagascar.

PRIONODON

Prionodon C. Mull, in Bot. Zeitung (Berlin) 2: 129 (1844); Broth, in Nattirl. PflFam., edn 2, 11:

112 (1925); Sim, Bryo. S. Afr. 356 (1926); Griffin in Rickia 6: 9 (1974). Type species: P. densus

(Hedw.) C. Midi.

Plants mostly robust, green, forming large, loose tufts; corticolous. Secondary stems ± erect,

simple or branched; central strand absent. Leaves fragile, ovate-lanceolate, coarsely toothed above.

Costa single, strong, extending to upper leaf. Laminal cells oval, incrassate, unipapillose; alar cells

differentiated.

Dioicous. Perichaetia along secondary stems. Sporophytes rare. Seta short. Capsule exserted,

erect. Peristome double. Operculum obliquely rostrate. Calyptra cucullate, smooth.

A genus of 26 species found in forests of tropical and south temperate regions. The plants form

large, loose tufts on trees. The genus is apparently represented in Africa by a single variable

species, P. densus.

Prionodon densus (Hedw.) C. Mull, in

Bot. Zeitung (Berlin) 2: 129 (1844); Broth, in

Naturl. PflFam., edn 2, 11: 1 14 (1925); Bartram

in Fieldiana, Bot. 25: 245 (1949). Type:

Jamaica.

Hypnum densum Sw. ex Hedw., Sp. muse, frond. 282

(1801).

Prionodon rehmannii Mitt, in J. Linn. Soc., Bot. 22: 31 1

(1886); Broth, in Naturl. PflFam., edn 2, 11: 114 (1925);

Sim, Bryo. S. Afr. 356 (1926). Syntypes: Kilimanjaro,

Hannington s.n. \ Transvaal, Rehmann s.n. (BM, PRE).

Plants large to robust, loosely tufted, yel-

low-green; corticolous. Primary stems creep-

ing, tomentose; secondary stems erect, (40-)

70-130 mm long, branches few, scattered; in

section oval, central strand absent, inner corti-

cal cells thin-walled, hyaline to yellowish, in

8-10 rows, outer cortical cells thick-walled,

yellow to yellow-brown, in 2-4 rows; axillary

hairs with basal cells brownish, 3 or 4 cells

high; paraphyllia absent; pseudoparaphyllia

folious, low and wide. Leaves crowded, erect to

spreading wet, spreading and crisped dry,

strongly plicate; ovate to lanceolate, 3. 5-5. 5

mm long; apex long-acuminate, fragile and

missing on older leaves; long-decurrent at

base; margins plane, dentate to spinose. Costa

single, percurrent to ending below apex,

smooth; ventral and dorsal surface cells elon-

gate; in section bulging dorsally, guide cells

somewhat larger, ventral cells small, thick-

walled, in 1 or 2 rows, dorsal cells thick-

walled, in 3 or 4 rows. Upper laminal cells

oval, quadrate or rhombic, ± homogeneous,

25-50 pm long, 15-22 pm wide, walls incras-

sate, weakly papillose on both surfaces; papil-

lae single, low, centred over lumen; marginal

cells frequently not papillose; basal cells rec-

tangular, 50-100 pm long, 12-20 pm wide,

hyaline, walls incrassate, pitted, smooth; alar

cells quadrate to hexagonal, forming distinct

groups, greenish, walls incrassate, nodulose.

Perigonia not seen. Perichaetia not strongly

differentiated, perichaetial leaves lanceolate-

acuminate when young. Capsule not known

from Africa. Fig. 155.

The extremely variable and usually sterile

species P. densus is thought to be the most

Prionodontaceae

559

Map 217. — Prionodon densus

commonly collected species of Prionodon

throughout its range. It is found in Central and

South America, eastern Africa and the East

Arican islands and is probably the only species

in Africa. In the Flora area it is found on trees

in forests of the northern and eastern Transvaal

regions, KwaZulu-Natal, and the eastern Cape.

Map 217.

Vouchers: Brenan 3343; Crosby & Crosby

9929; Jacot Guillarmod 6136, 8117.

This species is recognized by its large, little-

branched secondary stems and narrow, coarse-

ly toothed, plicate leaves. The leaf tips are

very fragile and absent on most older leaves.

The leaf cells are short, oval to rhombic and

unipapillose on both surfaces. Prionodon

could perhaps be confused with several genera

of the Pterobryaceae that have a similar habit,

but these genera all have smooth leaf cells and

filamentous pseudoparaphyllia.

Fig. 155. — Prionodon densus: 1. habit (dry), x 1; 2.

part of stem in cross section, x 175; 3. leaf, x 32; 4. leaf in

cross section, x 175; 5. leaf base (cells partly shown), x

160; 6. upper laminal cells, x 700; 7. upper laminal cells at

right margin, x 350; 8. leaf apex, x 175. (1, 3 & 5-8. Crosby

7531; 2 & 4, Knox 28.)

560

TRACHYPODACEAE

Plants small to large, forming mats and frequently with long-pendent branches, green to yellow-

green or yellow-brown, blackish below; corticolous or saxicolous. Stems long-creeping, irregular-

ly branched to pinnate; central strand weak or absent. Paraphyllia and pseudoparaphyllia absent.

Leaves appressed or spreading, rarely curved, sometimes longitudinally plicate; broadly ovate to

lanceolate; apex long- or short-acuminate, frequently crisped; margins plane, almost entire to

strongly toothed, rounded to weakly auriculate at base. Costa single, extending to above midleaf or

ending in acumen. Laminal cells homogeneous, linear to fusiform, rarely isodiametric, unipapillate

or seriately papillose over lumens, cell walls thickened; basal cells longer, thickened and pitted;

alar cells not strongly differentiated.

Dioicous. Perigonia on stems and branches, gemmate. Perichaetia lateral on stems and branch-

es. Seta erect, 2-50 mm long, smooth to strongly papillose. Capsule erect to inclined, mostly

oblong-cylindrical, up to 3 mm long. Peristome double; exostome teeth yellowish, lanceolate with

median zigzag line, striate below, papillose above; endostome segments short or as long as teeth

above high or low basal membrane, smooth or papillose, cilia present or absent. Operculum coni-

cal, obliquely rostrate. Calyptra cucullate or mitrate, smooth or hairy. Spores small, rounded, papil-

lose.

A family of six genera found primarily in southeast Asia. A few taxa extend as far north as Japan

or to the south as far as Australia, South Africa and southern Brazil. The family is best represent-

ed in the tropical forests of Asia but is also known from forests of India, Africa and the East African

islands. South America, Central America and the larger Caribbean Islands.

Leaves crisped at apex when dry; laminal cells with a single papilla over each cell

1 . Trachypodopsis

Leaves weakly plicate, not distinctly crisped at apex; laminal cells seriate papillose with sev-

eral low, blunt papillae over each cell 2. Trachypus

1. TRACHYPODOPSIS

Trachypodopsis Fleisch. in Hedwigia 45: 64 (1906); Broth, in Natiirl. PflFam., edn 2, 11: 120

(1925); Sim, Bryo. S. Afr. 397 (1926); Van Zanten in Blumea 9: 511 (1959). Lectotype species: T.

serrulata (P. Beauv.) Fleisch., vide Van Zanten (1959).

Plants large, forming mats with pendent branches, green to yellow-green or brownish, not black-

ened; corticolous or saxicolous. Primary stems long-creeping; secondary stems and branches fre-

quently pendent; central strand present or absent. Leaves appressed and horizontally spreading,

crisped at apex; margins toothed. Costa single and slender, extending to upper leaf. Laminal cells

linear-fusiform, incrassate and pitted, papillose with a single papilla over each lumen; alar cells not

differentiated.

Dioicous. Seta erect, short, papillose. Capsule exserted, erect, ovoid. Peristome double, cilia

absent. Operculum obliquely rostrate. Calyptra cucullate. Spores small, yellowish, papillose.

Trachypodopsis contains five species found primarily in Asia. The genus is also known from

Africa and Central America as different varieties of the widespread species T. serrulata. In Africa

the typical variety is found in forests of eastern, western and southern Africa and the East African

islands.

Trachypodaceae

561

Trachypodopsis serrulata (P. Beauv.)

Fleisch. in Hedwigia 45: 67 (1906); Broth, in

Natiirl. PflFam., edn 2, 11: 122 (1925); Sim,

Bryo. S. Afr. 397 (1926); Van Zanten in Blumea

9: 517 (1959). Type: Mascarene islands.

Bourbon, Bory St. Vincent (L).

Pilotrichum serrulatum P. Beauv., Prodr. aetheogam. 83

(1805). Neckera serrulata (P. Beauv.) Brid., Muscol. recent,

suppl. 2: 29 ( 1812). Papillaria serrulata (P. Beauv.) Jaeg. in

Ber. Thatigk. St. Gallischen Naturwiss. Ges. 1875-1876:

274(1877). Trachypus serrulatus ( P. Beauv.) Besch. in Ann.

Sci. Nat. Bot. 6, 10: 269 (1880).

Neckera macleana Rehm. in Par., Index bryol. 852

(1896), nom. nud.; Sim, Bryo. S. Afr. 397 (1926).

Plants large, forming mats or somewhat pen-

dent, green or yellow-green to brownish; saxi-

colous. Stems creeping or pendent, 20-100 mm

long, branching irregular, branches ± pinnate; in

section round, central strand small, inner corti-

cal cells somewhat thick-walled, hyaline, in 7 or

8 rows, outer cortical cells smaller, thick-

walled, yellow-brown, in 3 or 4 rows; axillary

hairs few, hyaline, 3 or 4 cells long; paraphyllia

absent; pseudoparaphyllia absent but bud loose

with erect, reduced leaves. Leaves evenly

spaced, complanate, spreading to horizontal

wet, appressed dry; oblong to broadly ovate,

2.0-3. 5 mm long; short- to long-acuminate;

apex contorted especially when dry; rounded or

weakly auriculate at base; margins plane, den-

ticulate to dentate, some teeth recurved, cells

not differentiated but somewhat longer. Costa

single, ending in acumen, smooth, surface cells

elongate, smooth; in section elliptical, guide

cells 2, large, thin-walled, dorsal and ventral

cells in single row, smaller, thickened. Upper

laminal cells linear to fusiform, homogeneous,

25-37 pm long, 3-7 pm wide, walls incrassate

and pitted, papillose, papillae single, centred

over lumen; basal cells oblong-hexagonal,

25-50 pm long, 7-10 pm wide, yellow, walls

incrassate and pitted, smooth; alar cells round-

ed, quadrate to rectangular, hyaline, walls ±

thickened and pitted.

Perigonia on stem and branches, gemmate;

leaves ovate-acuminate, up to 1 mm long.

Perichaetia and sporophyte not seen. Fig. 156:

1-8.

The typical variety of T. serrulata is found

infrequently in forests of eastern, western and

southern Africa, the East and West African

islands and India (Assam). In the Flora area it

forms mats on rocks and trees in forests of the

eastern Transvaal region. Map 218.

Vouchers: Crosby & Crosby 13374; Kluge

2003; Smook & Phelan 846; Raven 26218.

The species is easily recognized by its

appressed and horizontally spreading leaves

and crisped leaf apex when dry. In addition the

leaf cells have a single, small papilla centred

over the lumen. This feature is also known in

Aerobryopsis capensis , but the leaves of that

species are widely spreading and lack the

crisped apex of Trachypodopsis.

2. TRACHYPUS

Trachypus Reinw. & Homsch. in Nova Acta Phys.-Med. Acad. Caes. Leop. -Carol. Nat. Cur. 14:

708 (1829); Broth, in Natiirl. PflFam., edn 2, 11: 118 (1925); Van Zanten in Blumea 9: 481 (1959).

Type species: T. bicolor Reinw. & Homsch.

562

Trachypodaceae

Trachypodaceae

563

Plants medium to large, forming lax mats, green to yellow-green or brownish, frequently black-

ish below; saxicolous or corticolous. Primary stems long-creeping; secondary stems erect, ascend-

ing or sometimes pendent; central strand absent. Leaves erect and contorted dry, spreading wet,

lamina somewhat plicate, margins entire to serrate. Costa single, slender and extending to midleaf

or above. Laminal cells fusiform, incrassate, seriate-papillose with several low, blunt papillae that

cover lumen; alar cells not differentiated.

Dioicous. Perigonia and perichaetia along stems and branches; perichaetial leaves long-acumi-

nate above sheathing base. Seta elongate, erect, papillose. Capsule exserted, ovoid. Operculum

obliquely long-rostrate. Calyptra cucullate, hairy. Spores rounded, yellowish, papillose.

The genus Trachypus contains five species known primarily from India, southeast Asia and

islands of the Pacific. Two varieties of the most

known from Africa and tropical America.

Trachypus bicolor Reinw. & Hornsch. var.

viridulus (Mitt.) Zant. in Blumea 9: 499 (1959).

Type: Ecuador, near Quito, Jameson s.n.

Neckera viridula Mitt, in Hooker’s J. Bot. Kew Gard.

Misc. 3: 331 (1851). Papillaria viridula (Mitt.) Jaeg. in Ber.

Thatigk. St. Gallischen Naturwiss. Ges. 1875-1876: 267

(1877). Trachypus viridulus (Mitt.) Broth, in Natiirl.

PflFam. 1,3: 831 (1906); Broth, in Natiirl. PflFam., edn 2,

11: 119 (1925).

Papillaria natalensis Sim, Bryo. S. Afr. 392 (1926).

Type: South Africa, Natal, Pietermaritzburg, Town Bush,

Sim 8703 (PRE).

Plants medium to large, forming mats, green

to yellow-green, black below; terricolous or

saxicolous. Stems creeping, up to 50 mm long,

branches erect from creeping stem (perpendi-

cular to substrate), pinnate; in section round,

central strand absent, inner cortical cells weak-

ly thick-walled, hyaline, in 4 or 5 rows, outer

cortical cells smaller, thick-walled, yellowish,

in 3 or 4 rows; axillary hairs not seen; para-

phy Ilia absent; pseudoparaphyllia absent.

Leaves ± crowded, spreading wet, erect-

appressed and contorted dry, ± weakly plicate;

ovate to lanceolate, 2 mm long; long-acumi-

widely distributed species, T. bicolor , are also

nate; weakly auriculate at base; margins plane,

weakly serrulate. Costa single, extending to

midleaf or above; ventral surface cells elon-

gate, papillose; dorsal surface cells elongate,

smooth; in section oblong, bulging dorsally,

guide cells not differentiated, cells in 3 or 4

layers, dorsal cells smaller, strongly thickened.

Upper laminal cells broadly fusiform, homoge-

neous, 27-37 pm long, 6-9 pm wide, walls

thickened, papillose; papillae 8-10, low, seri-

ate, centred over lumen and completely cover-

ing cells; basal cells rectangular, 25-50 pm

long, 6-8 pm wide, hyaline, walls thickened

and pitted, smooth at insertion; alar cells not

differentiated.

Dioicous; sporophyte not known from Flora

area. Fig. 156: 9-16.

Trachypus bicolor var. viridulus is found in

forests of southern and southeastern Asia, India,

Sri Lanka, eastern South Africa, the East and

West African islands, Central America, northern

South America, and the West Indies. In the

Flora area the taxon is found on rocks and trees

Fig. 156. — Trachypodopsis serrulata ( 1-8): 1. habit (dry), x 1; 2. part of secondary stem, x 3; 3. part of stem in cross

section, x 175; 4. leaf, x 35; 5. leaf in cross section, x 175; 6. basal leaf cells (left side), x 175; 7. upper laminal cells at left

margin, x 350; 8. leaf apex, x 175. Trachypus bicolor var. viridulus (9-16): 9. habit (dry), x 3; 10. habit (wet), x 5; 11.

part of stem in cross section, x 175; 12. leaf, x 35; 13. leaf in cross section, x 175; 14. basal leaf cells (papillae partly

shown), x 200; 15. upper laminal cells at right margin (papillae partly shown), x 350; 16. cells at leaf apex (papillae partly

shown), x 350. (1-8, Kluge 2003 ; 9, 11-14 & 16, Rankin 157; 10 & 15, Hilliard & Burn 10271.)

564

Trachypodaceae

Map 219. — • Trachypus bicolor var. viridulus

♦ Jaegerina stolonifera

in forests of the eastern Transvaal region and

KwaZulu-Natal. Map 219.

Vouchers: Burrows & Burrows 5984; Hil-

liard & Burtt 10271; Rankin 157.

The blackish coloration of the older parts of

stems and branches helps to identify the taxon,

although a similar coloration is also frequently

found in members of the Meteoriaceae. The

low, blunt, seriate papillae on the leaf cells are

distinctive and help to identify this species.

Trachypus bicolor var. hispidus (C. Mull.)

Card, and T. appendiculatus (Ren. & Card.)

Broth, have been reported from eastern Africa

but they are not clearly separated from the

southern African taxon and many intermediate

forms exist.

565

PTEROBRYACEAE

Plants small to large, occasionally robust, mostly dendroid, branches occasionally pendent,

green to yellow-green or golden green; corticolous or saxicolous. Primary stems appressed to sub-

strate, long-creeping, naked or with scale leaves; secondary stems erect, perpendicular to substrate,

usually branched; in section central strand absent. Leaves erect and weakly crisped or squarrose

dry, spreading to squarrose wet, generally strongly concave; broadly ovate to elliptical; rounded to

cordate or auriculate at base; margins plane below, generally broadly inflexed above, entire to ser-

rate. Costa variable. Laminal cells short to long, rhomboidal to fusiform or linear and sigmoid, gen-

erally thickened and pitted; basal cells rectangular, in yellowish band across insertion, thickened

and pitted; alar cells generally differentiated, forming distinct group, usually quadrate, frequently

thickened and pitted. Gemmae frequently present, cylindrical, multicellular, green or brown.

Mostly dioicous. Perigonia and perichaetia along secondary stem and branches, frequently

large, green. Seta generally very short. Capsule mostly immersed. Peristome single or double,

prostome frequently present; endostome usually rudimentary. Operculum conical to rostrate.

Calyptra small, ephemeral, mostly smooth. Spores large, angular, papillose.

A family of 27 genera found in tropical and subtropical forests throughout the world. Seven gen-

era are known from Africa, three extending into forests of southern Africa.

1 Leaves squarrose wet or dry 1. Jaegerina

1 Leaves appressed to widely spreading:

2 Branches julaceous; leaves not noticeably seriate 2. Pterobryopsis

2 Branch leaves spirally seriate 3. Orthostichopsis

1. JAEGERINA

Jaegerina C. Mull, in Linnaea 40: 273 (1876); Broth, in Natiirl. PflFam., edn 2, 11: 138 (1925);

Argent in J. Bryol. 7: 367 (1973). Type species: J. stolonifera (C. Miill.) C. Mull.

Plants large, green to yellow-green; corticolous. Primary stems creeping, with scale leaves; sec-

ondary stem erect, little branched; in section round, central strand absent. Leaves squarrose, some-

what plicate; broadly ovate; acute to acuminate; margins plane, serrulate; base cordate. Costa sin-

gle, extending to upper leaf. Laminal cells short- to long-elliptical, thickened and pitted; basal and

alar cells yellow but otherwise not strongly differentiated. Gemmae axillary, cylindrical.

Dioicous. Seta very short. Capsule immersed. Peristome single, exostome teeth broad.

Operculum conical. Calyptra small. Spores large.

A small genus of about 13 species found in forests of the Americas, Africa and Asia. Two species

are known from Africa and an additional four species are found on the East African islands.

Jaegerina stolonifera (C. Miill.) C. Miill.

in Linnaea 40: 274 (1876); Argent in J. Bryol. 7:

369 (1973). Type: Comores, Anjouan, Peters ,

Oct. 1843.

Plants large, loose-tufted, yellow-green to

brownish; corticolous. Primary stems long-

creeping; secondary stems erect to subpendent,

20-50 mm long, branches few, rare; in section

566

PTEROBRYACEAE

oval, central strand absent, inner cortical cells

thin-walled, hyaline, in up to 9 rows, outer cor-

tical cells smaller, thick-walled, in 6-8 rows;

paraphyllia absent; pseudoparaphyllia filamen-

tous. Leaves evenly spaced, in ranks, squarrose

wet or dry, weakly rugose; stem leaves broadly

ovate or lanceolate, 2. 5-5. 5 mm long; narrowly

acuminate; rounded to cordate at base; margins

plane, serrulate. Costa single, ending in acu-

men, smooth; in section elliptical or flat, guide

cells not differentiated, ventral cells in a single

row, dorsal cells larger, in single row. Upper

laminal cells elliptical, homogeneous, 10—35

pm long, 4-7 pm wide, walls thickened and pit-

ted, smooth; basal cells linear, 50-75 pm long,

5-7 pm wide, yellowish to brownish, walls

thickened and pitted, smooth; alar cells not

strongly differentiated, short-rectangular, brown-

ish, walls thickened and pitted. Gemmae scat-

tered along stems, cylindrical to clavate, brown-

ish, smooth.

Sporophyte not known from the Flora area.

Fig. 157.

Apparently rare, this large species is only

rarely collected in forests of eastern and south-

ern Africa, the East African islands and India.

Only a single specimen has been seen from the

Flora area, collected in forests of the eastern

Cape. Map 219.

Voucher: Wager PRE-CH1456.

Jaegerina stolonifera is very distinctive,

both in its size and squarrose leaves. Since the

species has not been recollected, the southern

African specimen may represent a local chance

introduction from the East African islands.

Fig. 157. — Jaegerina stolonifera: 1. habit (dry), x 3; 2.

part of stem in cross section, x 175; 3. scale leaf of prima-

ry stem, x 35; 4. leaf of secondary stem, x 25; 5. part of leaf

in cross section, x 175; 6. basal leaf cells (right side), x 175;

7. upper laminal cells at left margin, x 350; 8. leaf apex, x

175. (Wager PRE-CH1456.)

Pterobryaceae

567

2. PTEROBRYOPSIS

Pterobryopsis Fleisch. in Hedwigia 45: 56 (1906); Broth, in Natiirl. PflFam., edn 2, 11: 140

(1925). Lectotype species: P. crassicaulis (C. Mull.) Fleisch., fide Arzeni (1954).

Plants medium-sized to large, forming lax tufts, glossy, yellow-green to green; mostly terri-

colous. Primary stems long-creeping; secondary stems erect to short-pendent, irregularly to pin-

nately branched, densely leaved throughout; in section central strand absent. Leaves usually imbri-

cate, strongly concave; broadly ovate to oval; acute to abruptly acuminate; subcordate to weakly

auriculate at base. Costa absent, short and double, or single and extending to above midleaf. Upper

laminal cells linear-fusiform to rhomboidal, usually strongly thickened and pitted; basal cells

rhomboidal to rectangular, coloured across insertion; alar cells ± differentiated, smaller, quadrate.

Gemmae axillary, cylindrical.

Dioicous. Sporophytes rare. Seta short, smooth. Capsule immersed, ovoid. Peristome irregular,

smooth; prostome present. Operculum conic-rostrate. Calyptra small, ephemeral. Spores large, up

to 50 pm.

Pterobryopsis is a genus of 43 species centred in southeast Asia and India. A few species are

also known from the Americas, Africa and Australia. The generic distinctions between

Pterobryopsis and Calyptothecium Mitt, have faded as more species have been discovered or

moved into the genera. Although a completely satisfactory solution to the problem of separating

the genera will not be found until they are monographed, a more restrictive view of Calyptothecium

is adopted here and the southern African species are transferred to Pterobryopsis. When Mitten

(1869) described Calyptothecium , the two obvious characters he used were the distichous branch-

es and flattening of the leaves along the stem. These characters were also partly responsible for the

positioning of the genus in Neckeraceae. In Africa three species have their stem leaves complanate,

C. bescherellei (Ren.) Broth., C. planifrons (Ren. & Card.) Argent, C. pterobryoides Argent, and

are here considered to represent species of Calyptothecium. However, the two species that reach

southern Africa have ± julaceous stems and are treated in Pterobryopsis.

1 Costa absent or very short, single or double, restricted to lower third of leaf 1 . P. hoehnelii

1 Costa single, occasionally spurred, extending to above midleaf:

2 Leaves elliptical or oblong-ovate, upper margins plane and entire 2.P. acutifolium

2 Leaves broadly ovate to oval, upper margins inflexed and serrulate 3. P. rehmannii

1. Pterobryopsis hoehnelii (C. Mull.)

Magill, comb. nov. Type: Kenya, Mt Kenya,

Leikipia region, Hoehnel s.n.

Neckera hoehnelii C. Mull, in Flora 73: 489 (1890).

Calyptothecium hoehnelii (C. Mull.) Argent in J. Bryol. 7:

571 (1973).

Calyptothecium africanum Broth, in Bot. Jahrb. Syst.

20: 198 (1894). Syntypes: South Africa, Rehmann 332

(PRE); Usambara: Hochwaelder; Lugulua-Wald;

Muandara-Wald; Bulua-Wald; Hochwald, Holst 9044.

2636. 2621. 4312, 3291.

Plants large, somewhat scattered or forming

large turfs, yellow-green; corticolous or terri-

colous. Primary stems long-creeping; secondary

stem erect, 30-100 mm long, branches distichous

above stipe, pinnate; in section oval, central

strand absent, inner cortical cells thin-walled,

yellowish, in 5-7 rows, outer cortical cells thick-

walled, smaller, red-brown, in 3-5 rows; axillary

hairs very short, 2 or 3 cells long, apical cell larg-

er, brownish; paraphyllia absent; pseudopara-

phyllia filamentous, 8-12 cells long, 2 or 3 cells

wide at base. Leaves crowded, concave, spread-

ing wet, somewhat appressed to spreading dry;

stem leaves oblong to oblong-ovate, 2. 5-4.0 mm

long; acuminate; subcordate at base; margins

broadly incurved above, entire; branch leaves

similar to stem leaves although generally small-

Pterobryaceae

Pterobryaceae

569

er. Costa absent, short and double or short and

single, restricted to lower leaf, weak, smooth,

hyaline; in section elliptical consisting of a group

of 4 slightly smaller cells. Upper laminal cells

linear-fusiform, homogeneous, 75-120 pm long,

7-8 pm wide, walls incrassate and pitted,

smooth; basal cells ± rectangular, forming dis-

tinct group across insertion, 25-50 pm long,

6-12 pm wide, brownish, walls thickened and

pitted, smooth; alar cells not differentiated.

Gemmae in leaf axils, filiform, of 6-10 cells,

55-175 pm long, brownish green, smooth.

Dioicous. Perigonia gemmate, perigonial

leaves ovate-cuspidate. Perichaetia green; peri-

chaetial leaves ovate, long-acuminate, 4.5-5. 5

mm long, leaf cells linear, thickened and pitted.

Seta yellowish, smooth. Capsule immersed,

erect, ovoid, 1. 2-2.0 mm long, smooth, brown-

ish; neck not differentiated; exothecial cells ir-

regularly rectangular, walls thin, cells at mouth

quadrate and darker; stomata not seen. Peri-

stome single, yellow-red; prostome present;

exostome teeth irregular, linear-lanceolate,

smooth, to 400 pm high; endostome absent.

Operculum conic-rostrate, 0.5 mm long.

Calyptra small. Spores rounded, heterosporous,

25-48 pm, granulate, brown. Fig. 158: 1-13.

Found in forests of eastern, central and

southern Africa and the island of Bioko, P

hoehnelii is the most commonly encountered

species of Pterobryopsis in Africa. In southern

Africa, the species is found on trees, soil and

rock in forests of the eastern Transvaal region,

Zululand, KwaZulu-Natal, and the eastern and

southern Cape. Map 220.

Vouchers: Crosby & Crosby 13381a; Magill

5122, 6036; Schelpe 7518; Van Rooy 880.

The plants usually cover large areas on tree

trunks and branches, rocks or soil at the base of

trees. The large, ± dendroid plants are usually

recognized by their size, large concave leaves,

entire leaf margins, short or absent costa, and

smooth, strongly thickened and pitted leaf cells.

Some specimens have been seen with leaves

only weakly concave. However, most plants

have strongly concave leaves.

Sporophytes are frequently found on P.

hoehnelii although the immersed capsules, large

green perichaetial leaves and densely leaved

stems make them inconspicuous.

2. Pterobryopsis acutifolium (Brid.)

Magill , comb. nov. Type: Mascarene Isl., Re-

union, s.L, no. XI.

Neckera acutifolia Brid., Bryol. univ. 2: 757 (1827).

Calyptothecium acutifolium (Brid.) Broth, in Par., Index

bryol., edn 2, 1: 288 (1904); Argent in J. Bryol. 7: 571

(1973).

Calyptothecium subacutifolium Broth, in Bot. Jahrb.

Syst. 24: 254 (1897). Type: South Africa, Pondoland,

Egorawald, Beyrich 40.

Fig. 158.— Pterobryopsis hoehnelii (1-13): 1. habit (dry), x 1; 2. distal part of branch with sporophyte (wet), x 5; 3.

stem in cross section (cells partly shown), x 175; 4. leaf, x 25; 5. leaf in cross section, x 175; 6. basal leaf cells (left side),

x 175; 7. upper laminal cells, x 350; 8. leaf apex, x 122; 9. perichaetial leaf, x 25; 10. part of capsule mouth with peristome

teeth, x 122; 11. operculum, x 35; 12. calyptra, x 35; 13. spore, x 490. P. acutifolium (14-1 8): 14. habit (dry), x 1 ; 15. leaf,

x 35; 16. basal leaf cells (left side), x 175; 17. upper laminal cells at right margin, x 350; 18. leaf apex, x 175. P. rehman-

nii (19-23): 19. habit (dry), x 1; 20. leaf, x 35; 21. basal leaf cells (left side), x 175; 22. upper laminal cells at right mar-

gin, x 350; 23. leaf apex, x 175. (1, Brenan 3352\ 2 & 10-13, Sim 7125\ 3-5, 7 & 9, Sim 7014\ 6 & 8, Crosby 7792\ 14,

Sim PRE-CH9993\ 15 & 16, Magill 5583\ 17 & 18, Crosby 7843\ 19-23, Buchanan s.n.)

570

Pterobryaceae

Plants medium-sized, forming large turfs,

green to yellow-green; corticolous or saxi-

colous. Primary stems creeping; secondary

stem erect, 20-70 mm long, branching irregular,

distichous above stipe, pinnate; in section oval,

central strand absent, inner cortical cells large,

hyaline, thin-walled, in 3-5 rows, outer cortical

cells smaller, brownish, thick-walled, in 4-6

rows; axillary hairs not seen; paraphyllia

absent; pseudoparaphyllia filamentous. Leaves

crowded, erect- spreading wet, appressed dry,

somewhat crisped, concave; stem leaves ovate,

elliptical or oblong-ovate, 2. 6-3. 2 mm long;

acute; cordate to subauriculate at base; margins

plane, entire; branch leaves similar but smaller.

Costa single, slender, ending below apex,

smooth; in section bulging dorsally, 2 cells

thick. Upper laminal cells fusiform, homoge-

neous, 50-75 pm long, 6-8 pm wide, walls

thickened and pitted, smooth; basal cells rectan-

gular, much shorter and brownish at insertion,

walls incrassate and pitted; alar cells not differ-

entiated. Gemmae in leaf axils, few, filiform, of

4-8 cells, to 62 pm long, greenish, smooth.

Dioicous. Perigonia gemmate. Perichaetia

green. Sporophyte not seen but described as:

Seta very short. Capsule immersed, ovoid.

Peristome single, yellowish, teeth irregular.

Operculum conic-rostrate. Calyptra cucullate.

Spores rounded, variable, 18-40 pm, papillose.

Fig. 158: 14-18.

This species is known from forests through-

out Africa and the African islands. In the Flora

area P. acutifolia is infrequently collected on

trees or rocks in forests of the eastern Transvaal

area, KwaZulu-Natal and the eastern Cape

region. Map 221.

Vouchers: Crosby & Crosby 7843; Magill

3220, 5583; Sim 9993; Van Rooy 927 , 1548.

Pterobryopsis acutifolium is somewhat vari-

able in plant size and leaf shape. The plants can

be very short and compact or larger and full. The

leaves, which exhibit some variation in shape of

the lamina and apex, are occasionally crisped

when dry. The plants are identified by their large

size, single costa, entire leaf margins, and

smooth, strongly thickened and pitted leaf cells.

Map 221. — • Pterobryopsis acutifolium

♦ Pterobryopsis rehmannii

3. Pterobryopsis rehmannii Magill in

Magill & Schelpe in Mem. bot. Surv. S. Afr. 43:

5 (1979). Type: South Africa, Natal, Buchanan

s.n. (Rehmann Musci Austro-africani 615; NH,

holo.; BM).

Plants large, loose-tufted, green, glossy; cor-

ticolous. Primary stems long-creeping; sec-

ondary stem erect, up to 60 mm long, branches

few, pinnate. Leaves crowded throughout, erect

to imbricate wet or dry, concave; stem leaves

broadly ovate, 2. 0-2. 2 mm long; acute; weakly

decurrent at base; margins plane to inflexed

above, serrulate above; branch leaves elliptical

to oval, concave, 1.2-1. 8 mm long; acute; mar-

gins plane to inflexed above, serrulate above.

Costa single, ending below apex, slender,

smooth. Upper laminal cells rhomboidal, homo-

geneous, 55-60 pm long, walls thickened,

smooth; basal cells oblong, walls thickened,

smooth; alar cells forming distinct groups,

quadrate, walls thickened. Gemmae in leaf axils,

scattered, cylindrical, of 6-8 cells, green,

smooth.

Sporophyte not known. Fig. 158: 19-23.

Duplicate material of the specimen was dis-

tributed by Rehmann in the second part of his

Musci Austro-africani exsiccati. The specimen

Pterobryaceae

571

was collected in KwaZulu-Natal by Buchanan

between 1861 and 1874, but no further collec-

tion data are available. From substrate frag-

ments in the collection it appears that the speci-

men was collected from a tree. Map 221.

Voucher: type only.

The erect plants with short branches and im-

bricate, strongly concave, oval to broadly ovate

leaves with serrulate upper margins identify this

species. The plants are large and like other mem-

bers of the genus probably cover large areas of

trees and rocks in closed forests. It is therefore

curious that P. rehmannii has not been recollected.

3. ORTHOSTICHOPSIS

Orthostichopsis Broth, in Natiirl. PflFam. 1,3: 804 (1906); Argent in J. Bryol. 7: 597 (1973). Type

species: O. tetragonum (Hedw.) Broth.

Plants large, frequently long and slender, erect when young, becoming pendent with age, yel-

low-green to brownish; corticolous or occasionally saxicolous. Primary stems long-creeping,

naked or with scale leaves; secondary stems erect but stems and branches elongated and hanging

with age; in section central strand absent. Leaves generally arranged in distinct spiral rows, but

occasionally only branch leaves ranked; ovate to oval; acute to acuminate; margins broadly

inflexed above, entire to serrulate, cordate at base. Costa single, extending to above midleaf, occa-

sionally shorter, single or forked. Leaf cells oblong, thickened and pitted, smooth; alar cells form-

ing a distinct group.

Dioicous. Seta very short. Capsule immersed. Peristome double, endostome rudimentary.

Operculum short-rostrate. Calyptra hairy.

A genus of 20 species found in forests of Central and South America, Asia and Africa south of

the Sahara. Two species are found in the Flora area, both rare and restricted to forests of the

Transvaal regions, KwaZulu-Natal and the eastern Cape.

Leaves with a long, narrow acumen; plants frequently long-pendent, up to 150 mm long

1.0. subimbricata

Leaves with a short apiculus; plants usually short and erect, 10-40 mm long .... 2. O. pinnatella

1. Orthostichopsis subimbricata (Hampe)

Broth, in Natiirl. PflFam. 1,3: 805 (1906). Type:

Madagascar, Borchgrevink 13.

Neckera subimbricata Hampe in Linnaea 38: 216

(1874). Pilotrichella subimbricata (Hampe) Jaeg. in Ber.

Thatigk. St. Gallischen Naturwiss. Ges. 1875-1876: 255

(1877).

Neckera chrysoneura Hampe ex C. Mull, in Linnaea 40:

263 (1876). Orthostichopsis chrysoneura (C. Mull.) Broth,

in Natiirl. PflFam. 1,3: 805 (1906). Type: Comoro Island,

Johanna , Hildebrandt s.n., 1875.

Plants medium-sized to long and slender,

pendent, green to yellow-green; corticolous.

Primary stem long-creeping; secondary stem

long-pendent, up to 150 mm long; branching

irregular, subdistichous, up to 25 mm long; in

section round to oval, central strand absent,

inner cortical cells thin-walled, yellowish, in

4-6 rows, outer cortical cells smaller, thick-

walled, red-brown, in 3 or 4 rows; paraphyllia

absent; pseudoparaphyllia filamentous. Leaves

somewhat crowded, concave, erect to spreading

wet, ± appressed dry; stem leaves elliptical to

ovate, 1.9-2. 5 mm long; acuminate, hair-point-

ed; rounded to cordate at base; margins broadly

inflexed above, serrulate above; branch leaves

similar to stem leaves but somewhat smaller,

1.5-2. 2 mm long; subulate and hair-pointed;

cordate at base; margins broadly inflexed

above, serrulate above. Costa single or forked,

ending below apex, smooth; ventral surface

cells elongate; dorsal surface cells elongate; in

572

Pterobryaceae

Fig. 159. — Orthostichopsis subimbricata (1-9): 1.

habit (dry), x 1; 2. habit (wet), x 3; 3. stem in cross section

(cells partly shown), x 175; 4. stem leaf, x 35; 5. branch leaf,

x 35: 6. stem leaf in cross section, x 175; 7. basal cells of

branch leaf (right side), x 175; 8. upper laminal cells of stem

leaf at left margin, x 350; 9. stem leaf apex, x 175. O. pin-

natella (10-16): 10. habit (dry), x 1; 11. habit (wet), x 3; 12.

stem leaf, x 70; 13. branch leaf, x 70; 14. stem leaf in cross

section, x 175; 15. stem leaf basal cells, x 175; 16. branch

leaf apex, x 175.(1 & 3-9, Crosbv 13382\ 2, Van Rooy 1890\

10-16, Magill 5440.)

Pterobryaceae

573

section flat, consisting of 6-8 undifferentiated

cells. Upper laminal cells linear, weakly sig-

moid, homogeneous, 37-62 pm long, 3-6 |rm

wide, walls incrassate and pitted, smooth; basal

cells rectangular, 40-50 pm long, 6-8 pm wide,

yellowish, walls incrassate and pitted, smooth;

alar cells strongly differentiated, quadrate, yel-

low-brown, walls incrassate and pitted.

Dioicous? Perigonia not seen. Perichaetia

not strongly differentiated, green; perichaetial

leaves convolute, acuminate, 1.2 mm long.

Sporophyte not known from southern Africa but

capsule described as immersed. Fig. 159: 1-9.

The species is found in forests of eastern and

southern Africa and the East African islands. In

the Flora area, O. subimbricata is found in

dense forests of the northern and eastern

Transvaal regions and the eastern Cape area.

Map 222.

Vouchers: Crosby & Crosby 13375A, 13382;

Van Rooy 1890.

The plants could be easily confused with

Squamidium of the Meteoriaceae. However,

Orthostichopsis does not exhibit the blackening

of leaves and stems common in Squamidium. In

addition the costa in Orthostichopsis, although

somewhat variable, is always stronger than that

of Squamidium and the alar cells are not as pro-

nounced. The weakly seriate branch leaves of

this species resemble those of Pilotrichella, but

the differences in costa type and length as well

as capsule exsertion separate the taxa.

2. Orthostichopsis pinnatella (Broth.)

Broth, in Nattirl. PflFam. 1,3: 805 (1906). Type:

Tanzania (Usambara), Holst 9205 e.

Pilotrichella pinnatella Broth, in Bot. Jahrb. Syst. 20:

198 (1894).

Plants small, forming mats, green to yellow-

green; corticolous or saxicolous. Primary stems

long-creeping, flattened, with appressed scale

leaves; secondary stems erect, 10-40 mm long,

branching ± regular, distichous above stipe,

curved when dry, pinnate; in section oval, cen-

Map 222. — • Orthostichopsis subimbricata

♦ Orthostichopsis pinnatella

tral strand absent, inner cortical cells thin-

walled, yellowish, 3 or 4 cells wide, outer corti-

cal cells smaller, thick-walled, red-brown, 3 or

4 cells wide; paraphyllia absent; pseudopara-

phyllia filamentous. Leaves somewhat crowd-

ed, in ranks, erect wet, ± appressed dry, strong-

ly concave; stem leaves elliptical to ovate,

0.8-1. 1 mm long; apiculate; rounded to subau-

riculate at base; margins broadly inflexed above

to overlapping at apex, serrulate; branch leaves

similar but smaller than stem leaves, 0.5-0. 8

mm long. Costa single and extending to midleaf

or above, occasionally forked, sometimes short-

er and double in branch leaves; dorsal and ven-

tral surface cells elongate, smooth; in section

elliptical, a small group of 6-8 cells in lower

leaf. Upper laminal cells ± rhomboidal and sig-

moid, heterogeneous, 15-25 pm long, 5-8 pm

wide, walls weakly thickened, smooth; basal

cells rectangular, 20-28 pm long, 6-8 pm wide,

yellowish, walls thickened, smooth; alar cells

quadrate, forming distinct groups, yellow, walls

thickened.

Gametangia and sporophytes not known.

Fig. 159: 10-16.

Known only from the African mainland, O.

pinnatella is found in forests of Tanzania,

Malawi and South Africa. In southern Africa

574

Pterobryaceae

the plants grow in small clumps on trees in

forests of Zululand. Map 222.

Vouchers: Crosby & Crosby 7799; Magill

5440.

The rather small, dendroid plants are usually

quite distinctive; although when growing in large

mats, the strongly seriate, concave leaves resem-

ble those of Pilotrichella. Orthostichopsis pin-

natella can be separated from species of

Pilotrichella by its long, single costa, serrulate

upper leaf margins, shorter leaf cells, and dis-

tinct, although small group of alar cells. The

costa of O. pinnatella is somewhat variable, and

some leaves may have a very short and double

costa restricted to the lower leaf, but most branch

and stem leaves have a single, slender or occa-

sionally forked costa, extending to the upper leaf.

575

METEORIACEAE

Plants slender and creeping with long-pendent branches, green to yellow-green, frequently

becoming blackish on older parts of stem and branches; corticolous or saxicolous. Primary stems

appressed to substrate, highly branched; in section generally with weak central strand; paraphyllia

and pseudoparaphyllia absent. Leaves appressed or widespreading, frequently concave; ovate to

elliptical; acute to long-acuminate or piliferous; margins plane or incurved above, entire or weak-

ly serrate, rounded to base, frequently decurrent. Laminal cells short or long, thickened and fre-

quently pitted, generally papillose; basal cells distinct; alar cells mostly differentiated.

Dioicous or autoicous. Perichaetia frequently distinctive. Seta elongate. Capsule exserted or

immersed. Peristome double, well developed; exostome teeth 16, yellowish, somewhat striate

below, papillose above; endostome segments almost as high as teeth, thin, hyaline, above a fre-

quently high basal membrane, cilia present or absent. Operculum conic-rostrate. Calyptra cucullate

or mitrate. Spores small, weakly papillose.

1 Costa short and double or lacking 1 . Pilotrichella

1 Costa single, extending to above midleaf, frequently very fine:

2 Leaf cells smooth 2. Squamidium

2 Leaf cells papillose:

3 Cells with a single papilla 3. Aerobryopsis

3 Cells with several papillae over lumen:

4 Median leaf cells short-elliptic; leaf margins entire; branch leaves generally appressed

4. Papillaria

4 Median leaf cells elongate, ± fusiform; leaf margins serrate; branch leaves spreading

5. Floribundaria

1. PILOTRICHELLA

Pilotrichella (C. Mull.) Besch. in Mem. Soc. Sci. Nat. Cherbourg 16: 222 (1872); Sim, Bryo. S.

Afr. 395 (1926); Broth, in Nattirl. PflFam., edn 2, 11: 157 (1925). Type species: P. imbricata (P.

Beauv.) Besch.

Neckera subsect. Pilotrichella C. Mull., Syn. muse, frond. 2: 129 (1850).

Plants frequently in large pendent masses, dull green to yellow-green; saxicolous or corticolous.

Stems creeping or pendent; branches usually long- pendent. Leaves panduriform, deeply concave,

often in spiral rows; apiculate to piliform; margins entire or minutely serrate. Costa short and dou-

ble or absent. Laminal cells elongate, smooth; alar cells generally forming distinct group.

Perichaetia not obvious. Seta straight, smooth. Capsule exserted, ovoid. Peristome double.

Operculum conic-rostrate. Calyptra cucullate, pilose.

A genus of about 65 species found throughout Africa and Central and South America. The genus

is recognized by its small, strongly concave, frequently panduriform leaves with a sharp apex and

smooth leaf cells. The southern African species is uniform throughout its range.

Pilotrichella pandurifolia (C. Mull.) Jaeg.

in Ber. Thatigk. St. Gallischen Naturwiss. Ges.

1875-1876: 255 (1877); Broth, in Natiirl.

PflFam., edn 2, 11: 158 (1925); Sim, Bryo. S.

Afr. 395 (1926). Type: Cape, Zeyher s.n., 1823

(BM).

Neckera pandurifolia (' panduraefolia') C. Mull, in Bot.

Zeitung (Berlin) 13: 767 (1855).

576

Meteoriaceae

Meteoriaceae

577

Pilotrichella kuntzei C. Mull, in Hedwigia 38: 127

(1889). Type: Cape, Kingwilliamstown, Perie Forest,

Kuntze s.n., 1894.

Pilotrichella cuspidata Broth, in Bot. Jahrb. Syst. 24:

255 (1897). Syntypes: Pondoland, Beyrich 38, Bachmann 6.

Pilotrichella conferta Ren. & Card, in Bull. Soc. Roy.

Bot. Belg. 38,1: 24 (1900). Type: Lesotho, Vemet s.n. (PC).

Plants long and slender, forming dense pen-

dent masses, light green to yellow-brown or

dark green; saxicolous or corticolous. Primary

stems creeping; secondary stems erect to pen-

dent, 20-150 mm long, branches few, occasion-

ally erect and dendroid in new secondary

growth; in section round, central strand absent,

inner cortical cells in 5 rows hyaline, thick-

walled, outer cortical cells in 4 or 5 rows small-

er, thick-walled, yellow-brown. Leaves crowd-

ed, erect-spreading wet, spreading dry; stem

leaves obovate-cuspidate, 1. 5-2.0 mm long;

margins erect, entire; branch leaves generally

panduriform; obovate to oblong, 0.8-1. 5 mm

long; acute; cuspidate or sometimes mucronate;

± auriculate or rounded at base; margins plane

below, becoming erect to incurved above,

entire. Costa short and double or absent, ventral

and dorsal surfaces smooth; in section consist-

ing of 3 or 4 undifferentiated cells. Upper lami-

nal cells oblong-fusiform; 35-50 pm long,

walls weakly thickened, smooth; basal cells

similar to somewhat longer juxtacostally; alar

cells few quadrate, walls weakly thickened.

Dioicous. Perigonia on branches, gemmate.

Perichaetia along branches; perichaetial leaves

long-sheathing, 2.0-2.6 mm long, leaf cells lin-

ear. Seta 2-3 mm long, yellow-brown; vaginula

2 mm long, beset with hairs that reach base of

um. Capsule exserted, erect, ovoid, 1-2 mm

long, smooth, brown, neck weakly differentiat-

ed; exothecial cells subquadrate to subhexago-

nal, walls weakly thickened, cells at mouth

brownish; stomata few, phaneropore. Peristome

double, yellow; exostome teeth fragile, triangu-

lar, papillose, 0.4 mm high; endostome seg-

ments linear above very low basal membrane,

perforated, almost as long as teeth, smooth; cilia

rudimentary. Operculum conic-rostrate, to 1.5

mm long. Calyptra not seen. Spores rounded,

20-25 pm, papillose, green. Fig. 160: 1-11.

Pilotrichella pandurifolia is known from

forests of eastern and southern Africa. In the

Flora area the species is found on boulders,

rocks and trees in the forests of the northern,

central and eastern Transvaal areas, Swaziland,

Zululand, KwaZulu-Natal and the eastern and

southern Cape regions. Map 223.

Vouchers: Magill 3731, 5445, 6584; Smook

& Phelan 672; Stirton 9929; Von Breitenbach

353.

The 5 -ranked, strongly concave, panduri-

form branch leaves and short double costa

should place specimens of this species. The

Fig. 160. — Pilotrichella pandurifolia (1-11): 1. habit (dry), x 1; 2. part of secondary stem with branches (wet), x 3; 3.

part of stem in cross section, x 175; 4. stem leaf, x 35; 5. branch leaf, x 35; 6. basal cells of branch leaf, x 175; 7. upper

laminal cells of branch leaf at left margin, x 490; 8. cells at branch leaf apex, x 245; 9. perichaetial leaf, x 35; 10. part of

capsule mouth with peristome, x 122; 11. spore, x 700. Squamidium brasiliense (12-22): 12. distal part of secondary stem

with branches (dry), x 1; 13. part of secondary stem with branches (wet), x 3; 14. part of stem in cross section, x 175; 15.

stem leaf, x 32; 16. branch leaf, x 32; 17. basal cells of branch leaf, x 175; 18. upper laminal cells of branch leaf, x 320;

19. branch leaf apex, x 175; 20. perichaetial leaf, x 18; 21. part of capsule mouth with peristome, x 70; 22. spore, x 700.

(1-8, Burrows 5039; 9-11, Sim PRE-CHI6623; 12, Filter 23\ 13, 14, 20-22, Crosby 7902\ 15, 16 & 18, Crosby 13404', 17

& 19, Magill 5200.)

578

Meteoriaceae

above characters remain constant throughout

the growth stages of the plants. Juvenile plants

are frequently collected with stems short and

unbranched. These plants quickly develop into

an erect dendroid stage, somewhat like

Porothamnium. In later stages the stems elon-

gate up to 150 mm and have branches that fre-

quently exceed 50 mm. This last stage results in

the more typical habit, e.g. long-pendent stems

hanging from boulders and tree trunks.

Sporophytes are frequently found on the distal

branches.

2. SQUAMIDIUM

Squamidium (C. Mull.) Broth, in Natiirl. PflFam. 1,3: 807 (1906); Sim, Bryo. S. Afr. 390 (1926);

Allen & Crosby in J. Hattori Bot. Lab. 61: 431 (1986). Type species: S. lorentzii (C. Miill.) Broth.

Plants large and pendent, green to yellow-green with blackish tint on older parts of stems and

branches; terricolous or saxicolous. Primary stems long-creeping; secondary stems and branches

long and pendent. Leaves appressed, concave, abruptly cuspidate or piliferous; cordate and decur-

rent at base; margins plane, entire. Costa generally single, extending to upper leaf. Laminal cells

linear-fusiform, smooth; alar cells strongly differentiated.

Perichaetia differentiated. Seta to 1 mm long, smooth. Capsule immersed, ovoid. Peristome

double. Operculum convex. Calyptra cucullate, pilose.

A genus of seven species found in Central and South America, Africa, and the East African

islands. The genus might be confused with Orthostichopsis of the Pterobryaceae but Squamidium

has leaves with a stronger costa, decurrent leaf bases and no pseudoparaphyllia.

Squamidium brasiliense (Homsch.) Broth.

in Natiirl. PflFam. 1,3: 809 (1906); Allen &

Crosby in J. Hattori Bot. Lab. 61: 454 (1986).

Type: Brazil, near Mandioccam, Martius s.n.

(BM).

Antitrichia brasiliensis Homsch., FI. bras. 1,2: 52 (1840).

Meteorium biforme (Hampe) Besch. in Ann. Sci. Nat.

Bot. 6, 10: 269 (1880); Pilotrichella biformis (Hampe) C.

Mull, ex Geh., Abh. Naturwiss. Vereine Bremen 7: 209

(1881); Squamidium biforme (Hampe) Broth, in Natiirl.

PflFam. 1,3: 809 (1906). Type: Madagascar, Alamazan-

troskoven, Borgen s.n. (BM).

Pilotrichella rehmannii C. Miill. in Geh., Rev. Bryol.

Lichgnol. 5: 70 (1878); Meteorium rehmannii C. Miill.,

Hedwigia 38: 127 (1899); Squamidium rehmannii (C.

Miill.) Broth., Natiirl. PflFam. 1,3: 809 (1906); Sim, Bryo.

S. Afr. 391 (1926). Lectotype: Cape, Montagu Pass, near

Blanco, Rehmann 323 (BM), vide Allen & Crosby, 1986.

Plants medium-sized to large, pendent, green

to yellow-green turning blackish brown to black

on older stems and branches; corticolous or saxi-

colous. Primary stems creeping; secondary

stems pendent, julaceous, to 200 mm long,

branching regular, in widely spaced groups of

2-5 branches; in section round, central strand

variable, small or absent, inner cortical cells

hyaline, thin-walled, in 6 rows, outer cortical

cells yellow, thick-walled in 4 or 5 rows. Leaves

± crowded and concave, ± erect wet, appressed

dry; stem leaves ovate-lanceolate to oblong, 3.5

mm long; abruptly setaceous; subcordate and

decurrent at base; margins plane, entire, cells not

differentiated; branch leaves broadly ovate to

elliptical or oblong; abruptly cuspidate to seta-

ceous; subcordate and decurrent at base; mar-

gins plane, entire. Costa variable, mostly single,

extending to upper leaf, on stem leaves extend-

ing to midleaf, percurrent or short and double on

branch leaves; ventral and dorsal surface

smooth; in section flat, guide cells not differen-

tiated, ventral and dorsal surface cells thick-

walled. Upper laminal cells linear-fusiform,

walls weakly thickened, homogeneous, smooth;

basal cells rectangular, hyaline, walls thickened

and pitted, smooth; alar cells strongly differenti-

ated, quadrate, yellowish, walls thickened.

Dioicous. Perigonia on stems and branches,

gemmate. Perichaetia along stem; perichaetial

Meteoriaceae

579

leaves oblong-acuminate, base sheathing, 4 mm

long, leaf cells linear, fusiform. Seta 0. 5-1.0

mm long, brown, smooth. Capsule immersed,

ovoid, 2 mm long, smooth, black-green, neck

not differentiated; exothecial cells subquadrate

broadly rectangular, walls thickened, cells at

mouth quadrate, neck cells smaller; stomata

phaneropore. Peristome double, yellow-brown;

exostome teeth long-linear, smooth, papillose,

0.7 mm high; endostome segments linear above

low basal membrane, perforated, shorter than

teeth, smooth; cilia absent. Operculum ± con-

vex, 0.7 mm long. Calyptra cucullate, 1 mm

long, pilose. Spores rounded, 25-35 pm, granu-

late, yellow. Fig. 160: 12-22.

This species is known from South America,

eastern and southern Africa and the East African

islands. In the Flora area it is found on trees and

boulders in forests of the northern, central and

eastern Transvaal areas, Swaziland, Zululand,

KwaZulu-Natal and the eastern and southern

Cape regions. Map 224.

Vouchers: Crosby & Crosby 7902; Filter 18;

Leighton 3371a; Magill 5204, 6569; Pienaar 11.

Squamidium brasiliense is recognized by its

long, narrow, pendent stems with ± grouped,

julaceous branches. The branch leaves are

erect-appressed and frequently strongly con-

cave, making the branches appear fuller and

much larger than the stem. The plants are green

to yellow-green above but take on a shiny

blackish or brownish black tint on older parts of

stem or branches. This is also observed in other

members of the family, e.g. Papillaria, although

the plants are dull in appearance. Squamidium

brasiliense might be confused with Ortho-

stichopsis of the Pterobryaceae which is recog-

nized by its leaves in 5 -ranked spirals and fila-

mentous pseudoparaphyllia.

3. AEROBRYOPSIS

Aerobryopsis Fleisch. in Hedwigia 44: 304 (1905); Broth, in Natiirl. PflFam., edn 2, 11: 165

(1925); Sim, Bryo. S. Afr. 394 (1926); Noguchi in J. Hattori Bot. Lab. 41: 294 (1976); Gangulee,

Moss. E. India 5: 1315 (1976). Type species: not designated.

Plants large, yellow-green, shiny, with blackish tint on older parts; mostly corticolous. Primary

stems creeping; secondary stems and branches pendent. Leaves long-acuminate; cordate at base;

margins plane, entire to serrulate. Costa single. Laminal cells linear-fusiform with single papilla

over each lumen; alar cells not well defined.

Seta elongate, rough above. Capsule exserted. Peristome double. Operculum conic-rostrate.

Calyptra cucullate, smooth or with a few hairs.

A genus of ± 25 species found mostly in Asia, three species are known from Central and South

America, and one from Africa. Aerobryopsis is a lowland forest genus most easily recognized by

its widely spreading, glossy leaves.

Meteoriaceae

581

Aerobryopsis capensis (C. Mull.) Fleisch.

in Hedwigia 44: 306 (1905); Broth, in Natiirl.

PflFam., edn 2, 11: 164 (1925); Sim, Bryo. S.

Afr. 394 (1926). Type: Cape, Genadendal,

Breutel s.n. (BM).

Neckera capensis C. MU11. in Bot. Zeitung (Berlin) 16:

165 (1858); Aerobryum capense (C. Mull.) C. Mull, in

Linnaea 40: 262 (1876).

Aerobryum capense var. rupestre C. Mull, in Par., Index

bryol. 9 (1894), nom. nud.; Sim, Bryo. S. Afr. 394 (1926).

Plants medium-sized, pendent, yellow-

green, glossy; corticolous or rarely terricolous.

Primary stems flattened against substrate; sec-

ondary stems pendent, 200 mm long, branches

irregular; in section round, central strand small,

inner cortical cells in 5 or 6 rows, thin-walled

towards margin, outer cortical cells in 6 rows,

smaller, thick-walled. Leaves somewhat distant,

widespreading to reflexed, wavy to weakly pli-

cate; stem leaves ovate, 2.0-3. 0 mm long;

acuminate, acumens generally long and slender;

cordate at base; margins plane, serrulate. Costa

single, ending in acumen, ventral and dorsal

surfaces smooth; in section elliptical, cells in 3

equal rows, weakly thickened. Upper laminal

cells linear-fusiform, 50-76 pm long, walls

thickened, pitted, papillose, papillae single,

centred over cell lumens; basal cells rectangu-

lar, hyaline, walls more strongly thickened and

pitted, yellowish, smooth; alar cells not differ-

entiated.

Autoicous. Perigonia on branches, gem-

mate. Perichaetia along branches; perichaetial

leaves acuminate with base sheathing, leaf cells

linear-fusiform, thickened, pitted and smooth.

Seta 15-20 mm long, yellow-brown. Capsule

short-exserted, erect, short-cylindrical, 1. 5-2.0

mm long, brown; exothecial cells subquadrate

to rectangular, walls frequently collenchyma-

tous; stomata few, phaneropore. Peristome dou-

Map 225. — Aerobryopsis capensis

ble, yellow; exostome teeth broadly lanceolate,

striate with median zigzag line, 700-800 pm

high; endostome segments broadly keeled

above high basal membrane, as long as teeth, ±

smooth; cilia single, linear, short, papillose.

Operculum rostrate. Calyptra cucullate, 1 mm

long, smooth. Spores rounded, 12-17, granu-

late, brown. Fig. 161: 1-7.

The species is known from eastern and

southern Africa. In the Flora area A. capensis is

found in forests of the northern and eastern

Transvaal areas, Zululand, KwaZulu-Natal, and

the eastern, southern, central and southwestern

Cape regions. Map 225.

Vouchers: Magill 5557, 6022; Oliver 7179;

Rankin 143; Russell 2692b; Van Rooy 769a.

The glossy, yellow-green plants with widely

spreading leaves when wet or dry help to iden-

tify A. capensis. The single, small papilla cen-

tred over the broadest part of the linear-fusiform

leaf cells is distinctive in the family, but is also

seen in Trachypodopsis serrulata.

Fig. 161. — Aerobryopsis capensis (1-7): 1. habit (dry), x 1; 2. distal part of secondary stem (wet), x 3; 3. part of stem

in cross section, x 175; 4. leaf, x 35; 5. basal leaf cells, x 175; 6. upper laminal cells at left margin, x 350; 7. cells at leaf

apex, x 175. Papillaria africana (8-20): 8. habit (dry), x 1; 9. distal part of secondary stem with branch and sporophyte

(wet), x 3; 10. part of stem in cross section, x 175; 11. leaf, x 35; 12. part of leaf in cross section, x 175; 13. basal leaf cells,

x 175; 14. upper laminal cells at left margin, x 350; 15. cells at leaf apex, x 175; 16. perichaetial leaf, x 25; 17. part of cap-

sule mouth with peristome, x 175; 18. operculum, x 35; 19. calyptra, x 18; 20. spore, x 700. (1, Van Rooy 769a; 2, Filter

PRE-CHI 3387; 3 & 4, Crosby 7908 ; 5, Von Breitenbach 8; 6, Britten 2726b; 7, Von Breitenbach 317; 8, 11, 12 & 14, Van

Rooy 1951; 9, 16, 17 & 20, Arnold 1331; 10, Crosby 9240; 13 & 15, Magill 3529; 18, Sim 7231; 19, Crosby 9198.)

582

Meteoriaceae

4 PAPILLARIA

Papillaria (C. Mull.) C. Mull, in Ofvers. Forh. Kongl. Svenska Vetensk.-Akad. 33: 34 (1876) nom.

cons.; Broth, in Natiirl. PflFam., edn 2, 11: 161 (1925); Sim, Bryo. S. Afr. 391 (1926); Noguchi in

J. Hattori Bot. Lab. 41: 237 (1976); Gangulee, Moss. E. India 5: 1283 (1976). Type species: P.

nigrescens (Hedw.) Jaeg.

Neckera subsect. Papillaria C. Miill., Syn. muse, frond. 2: 134 ( 1850).

Plants hanging, slender, dull, green to yellow-brown, frequently with blackish tint on older parts;

corticolous. Primary stems creeping; secondary stems and branches pendent. Leaves lanceolate; cor-

date to auriculate at base; margins plane, entire to serrulate. Costa single, extending to midleaf or

above. Laminal cells short or long, multipapillose; alar cells not strongly differentiated.

Dioicous. Seta elongate, smooth. Capsule exserted. Peristome double. Operculum conic-ros-

trate. Calyptra cucullate, pilose.

A genus of ± 78 species found in Central and South America, Africa and Madagascar, India,

Asia, Australia and New Zealand, and some Pacific islands. The six species known from Africa are

recognized by their slender, pendent habit, dull yellowish to yellow-brown colour with blackish tint

on older stems and leaves, frequently auriculate leaves, and multipapillose leaf cells.

Papillaria africana (C. Miill.) Jaeg. in

Ber. Thatigk. St. Gallischen Naturwiss. Ges.

1875-1876: 272 (1877); Broth, in Natiirl.

PflFam., edn 2, 11: 163 (1925); Sim, Bryo. S.

Afr. 392 (1926). Type: Cape, Olifantshoek,

Ecklon s.n. (BM).

Neckera africana C. Mull., Syn. muse, frond. 2: 137

(1850). Meteorium africamnn (C. Miill.) Mitt., J. Linn.

Soc., Bot. 22: 314 (1886).

Plants slender, creeping along substrate with

pendent branches, green to yellow-green, dull;

saxicolous or corticolous. Primary stems creep-

ing; secondary stems pendent, to 400 mm long,

branching irregular; in section round, central

strand very small, inner cortical cells in 7 or 8

rows, weakly thickened, outer cortical cells in 4

or 5 rows smaller, thick-walled, yellow-green.

Leaves erect-spreading wet, appressed dry; ovate

to oval or oblong, ( 1 .4 — ) 1 .7— 2.0(— 2.5) mm;

abruptly long-acuminate, frequently extending

out as a long smooth awn; cordate or ± subauri-

culate and decurrent at base; margins plane,

entire. Costa ending in acumen, ventral and dor-

sal surfaces smooth; in section bulging dorsally,

to 3 cells thick, guide cells not differentiated,

dorsal surface cells thick-walled or not differen-

tiated. Upper laminal cells elliptical to rounded

rhombic, elongate-fusiform at apex, 10-12 pm

long, walls weakly thickened, papillose; papillae

small, scattered over lumen; apical and occasion-

ally marginal cells smooth; basal cells quadrate

to short rectangular, hyaline, smooth becoming

elongate-fusiform, strongly thickened and pitted

juxtacostally; alar cells small, quadrate.

Dioicous. Perigonia on branches, gemmate.

Perichaetia on distal branches; perichaetial

leaves oblong, 2.2 mm long, leaf cells fusiform,

thickened. Seta 2.5 mm long, brown; vaginula

long with numerous long paraphyses reaching

base of capsule. Capsule exserted, erect, ovoid,

1. 5-2.0 mm long, smooth, green-brown to

brown; exothecial cells quadrate to short rectan-

gular, walls thin, cells at mouth slightly larger,

darker; stomata numerous, phaneropore. Peri-

stome yellow; exostome teeth lanceolate, papil-

lose, perforated, 0.5 mm high; endostome seg-

ments linear above basal membrane, perforated,

shorter than teeth, almost smooth; cilia absent.

Operculum conic-rostrate, 1 .0 mm long. Calyp-

tra cucullate, 2 mm long, hairy. Spores rounded,

20-25 pm, papillose, green. Fig. 161: 8-20.

Found throughout eastern and southern

Africa and the East African islands, P. africana

grows hanging from branches of trees and bush-

es or less frequently on boulders in forests and

closed wooded areas. In southern Africa the

species is known from the northern and eastern

Meteoriaceae

583

Transvaal areas, Swaziland, Zululand, Kwa-

Zulu-Natal and the eastern, southern and south-

western Cape regions. Map 226.

Vouchers: Brenan 2795, 2841 ; Crosby 9198;

Magill 3673, 5201; Oliver 7060; Rankin 44;

Van Rooy 234.

These plants are generally recognized by

their slender, hanging branches and dull appear-

ance. They might be confused with other plants

with similar habit, e.g. Trachypns, Squamidium

or Floribundaria , but leaf shape, laminal cells

with scattered papillae, and differentiated basal

juxtacostal cells should identify specimens of P.

africana.

Map 226. — Papillaria africana

5 FLORIBUNDARIA

Floribundaria Fleisch. in Hedwigia 44: 301 (1905); Broth, in Natiirl. PflFam., edn 2, 11: 169

(1925); Sim, Bryo. S. Afr. 393 (1926); Noguchi in J. Hattori Bot. Lab. 41: 270 (1976); Gangulee,

Moss. E. India 5: 1299 (1976). Type species: not designated.

Plants small, dull, pendent; corticolous. Stems regularly branched. Leaves small, ovate-acumi-

nate, rounded at insertion; margins plane, entire to serrulate. Costa single, extending to midleaf.

Laminal cells linear, ± seriate-papillose; alar cells not distinct.

Seta elongate, smooth. Capsule exserted. Peristome double. Operculum conic-rostrate. Calyptra

cucullate, sparsely hairy. Spores rounded, papillose.

Floribundaria contains ± 34 species found in moist forests throughout the tropics. Two species

are known from Africa, both recognized by their small, delicate habit and seriate leaf cell papillae.

Floribundaria floribunda (Doz. & Molk.)

Fleisch. in Hedwigia 44: 302 (1905); Broth, in

Natiirl. PflFam., edn 2, 11: 170 (1925); Sim,

Bryo. S. Afr. 393 (1926). Type: ‘Java. Sumatra'

(L, holo.) fide Streimann in J. Hattori Bot. Lab.

69: 277-312 (1991).

Leskea floribunda Doz. & Molk. in Ann. Sci. Nat. Bot.

3, 2: 310 (1844); Meteorium floribundum (Doz. & Molk.)

Doz. & Molk., Muse, frond, ined. archip. ind. 6: 162. 53

(1848); Neckera floribunda (Doz. & Molk.) C. Mull, in

Linnaea 36: 9 ( 1869); Papillaria floribunda (Doz. & Molk.)

C. Mull, in Linnaea 40: 267 (1876).

Plants small, creeping over substrate or

forming thick masses, yellow-green to light

green; saxicolous or corticolous. Stems creep-

ing, 1CM-0 mm long, branching irregular; in

section round, central strand small, inner corti-

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

cells in 3 or 4 rows, smaller, thick-walled, yel-

low-brown. Leaves distant, widespreading wet,

reflexed and squarrose dry; stem leaves broadly

ovate to ovate-lanceolate, 0.9- 1.5 mm long;

acuminate; rounded at base; margins plane, ser-

rulate; branch leaves lanceolate to linear-lanceo-

late, 0.7-1. 2 mm long; acuminate; rounded at

base; margins plane, serrulate. Costa single,

weak, extending to midleaf or beyond but

always ending below apex, occasionally absent;

in section consisting of an undifferentiated

group of 4-6 cells. Upper laminal cells oblong-

fusiform, 25-38 pm long, walls weakly thick-

ened, papillose; papillae small, subseriate; basal

584

Meteoriaceae

cells not differentiated, somewhat broader; alar

cells and cells along insertion smooth.

Sporophyte not seen. Fig. 162.

Pantropical in distribution, Floribundaria

floribunda is known from Africa and the East

African islands, India, southern and southeast-

ern Asia, Japan, Oceania, Australia, and South

America. In southern Africa, the species is

found in dense shade on rocks, tree trunks, and

limbs of saplings in forests of the northern and

eastern Transvaal areas and KwaZulu-Natal.

Map 227.

Vouchers: Cholnoky 274; Crosby & Crosby

7893; Pienaar 25; Von Breitenbach 222.

The fine, slender plants with dull, wide-

spreading, distant leaves should place speci-

mens of F. floribunda. Sporophytes have not

been found in the Flora area, but they are borne

on short lateral shoots; the seta is ± 4 mm long

and carry the short ovoid capsule above the

leaves; the peristome is well developed and the

operculum is conic-rostrate.

Fig. 162. — Floribundaria floribunda: 1. habit (dry), x

I ; 2. habit (wet), x 3; 3. part of stem in cross section, x 175;

4. stem leaf, x 35; 5. branch leaf, x 35; 6. basal cells of stem

leaf (papillae partly shown), x 175; 7. upper laminal cells of

stem leaf (papillae partly shown), x 350; 8. stem leaf apex

(papillae partly shown), x 175. (1 & 3-7, Crosby 7540: 2,

Brenan 3237.)

585

LEPTODONTACEAE

Plants medium-sized, forming loose tufts, dark green to yellow-green; corticolous or saxicolous.

Secondary stems erect, sometimes incurved when dry, generally pinnately branched; branches in

single plane; in section central strand absent; paraphyllia present or absent; pseudoparaphyllia pre-

sent. Leaves appressed dry, widespreading wet; broadly ovate; apex rounded or acute to acuminate;

margins plane, entire. Costa strong, single, extending to midleaf or above. Upper laminal cells

short, rhombic; alar cells quadrate to transversely rectangular, frequently in large groups.

Autoicous or dioicous. Perigonia and perichaetia along erect stems and branches. Seta short.

Capsule exserted, short-cylindrical. Peristome double; exostome teeth somewhat irregular, smooth,

whitish; endostomes rudimentary. Operculum rostrate. Calyptra cucullate, somewhat hairy. Spores

small, granulate.

The family Leptodontaceae presently contains only four genera, two of which occur in the Flora

area. Forsstroemia and Leptodon are widely, although sporadically, distributed throughout the

world and appear to be of southern hemisphere origin. Both genera are represented in Africa by sin-

gle species.

Leaf apex acuminate; stems without paraphyllia; plants erect dry 1. Forsstroemia

Leaf apex obtuse; stems with numerous paraphyllia; plants incurved dry 2. Leptodon

1. FORSSTROEMIA

Forsstroemia Lindb. in Ofvers. Forh. Kongl. Svenska Vetensk.-Akad. 19: 605 (1863); Broth, in

Natiirl. PflFam., edn 2, 11: 87 (1925); Sim, Bryo. S. Afr. 359 (1926); Stark, J. Hattori Bot. Lab. 63:

145 (1987). Type species: F. trichomitria (Fledw.) Lindb.

Plants single or in small tufts; corticolous or saxicolous. Stems subdendroid, erect wet or dry;

pinnately branched. Leaves ovate; acute to acuminate; margins plane, entire to serrulate. Costa sin-

gle, extending to midleaf. Laminal cells rhombic, smooth; alar cells in distinct groups.

Autoicous. Seta short. Capsule emergent to exserted. Peristome double but with endostomes

rudimentary or sometimes absent. Operculum conic-rostrate. Calyptra cucullate.

A recent revision of Forsstroemia (Stark 1987) included all African specimens in F. producta ,

the most widely distributed species. Forsstroemia has for some time been treated in the family

Cryphaeaceae, but was moved to Leucodontaceae by Manuel (1974), primarily on the basis of a

cucullate calyptra. He discounted other differences, such as the presence of pseudoparaphyllia in

Forsstroemia, and created a new subfamily, Forsstroemioideae, for Forsstroemia and two other

discordant genera of the Leucodontaceae, Pseudocryphaea and Leucodontopsis. Although the sub-

family provides a better placement for Forsstroemia, it is also discordant within Leucodontaceae

because of the production of pseudoparaphyllia.

Buck (1980) subsequently moved the three genera and Leptodon to Leptodontaceae citing simi-

larities in several gametophytic and sporophytic characters. Leptodon, however, produces numer-

ous paraphyllia along its stem, while Forsstroemia , Pseudocryphaea Britt, and Leucodontopsis

Ren. & Card, lack paraphyllia. A relationship between Forsstroemia and Leptodon undoubtedly

exists. However, a firmer circumscription of Leptodontaceae is needed for a proper assessment of

other genera that belong here.

586

Leptodontaceae

Leptodontaceae