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
587
Forsstroemia producta ( Hornsch .) Par., Ind. Bryol. 498 (1896); Broth, in Natiirl. PflFam., edn 2, 11: 88 (1925); Sim, Bryo. S. Afr. 359 (1926); Stark, J. Hattori Bot. Lab. 63: 163 (1987). Neotype: South Africa, Cape Prov., Katberg Forest Reserve, Crosby & Crosby 7974 (MO), vide Stark (1987).
Pterogonium productum Hornsch. in Linnaea 15: 138 (1841). Neckera producta (Hornsch.) C. Miill., Syn. muse, frond. 2: 94 (1850). Lasia producta (Hornsch.) Jaeg. in Ber. Thatigk. St. Gallischen Naturwiss. Ges. 1875-1876: 204 (1877).
Plants medium-sized, forming tufts or rarely scattered and gregarious, green to yellow-green; corticolous to saxicolous. Primary stems creep- ing; secondary stem erect, 15-40 mm long, branching ± regular in a single plane, pinnate; in section oval, central strand absent, inner cortical cells thin-walled, hyaline, in 4—6 rows, outer cor- tical cells thick-walled, yellow, in 2 or 3 rows; paraphyllia absent; pseudoparaphyllia narrowly foliose. Leaves crowded, widespreading wet, appressed dry; stem leaves ovate, 1.0-1. 8 mm long; acute to acuminate; occasionally twisted at apex; rounded at base; margins plane, entire to weakly serrulate; branch leaves smaller but sim- ilar in other respects to stem leaves. Costa single, extending to midleaf or above, smooth; in section elliptical, consisting of three rows of undifferen- tiated cells. Upper laminal cells rhombic to quadrate, homogeneous, 11-20 pm long, 6-10 pm wide, walls thickened, smooth; basal cells not strongly differentiated, rectangular to fusiform, greenish, walls thickened, smooth; alar cells forming distinct groups, quadrate to trans- versely rectangular, green, walls thickened.
Perigonia gemmate. Perichaetia strongly differentiated; leaves lanceolate-acuminate, 2-3 mm long, leaf cells linear and ± sigmoid. Seta 1-3 mm long, yellow, smooth. Capsule emer-
gent to shortly exserted, ± erect, short-cylindri- cal, 1 .0-1.5 mm long, smooth, yellow-brown; exothecial cells irregular-rectangular, walls thin, cells at mouth darker, quadrate to transversely rectangular; annulus absent; neck cells not dif- ferentiated; stomata absent. Peristome double, light yellow; exostome teeth fragile and irregu- lar, linear, inflexed and closed wet, sigmoid and incurved dry; ± smooth cleft and perforated, with median zigzag line, 100-110 pm high; endostome segments rudimentary or sometimes absent with age. Operculum conic-rostrate, erect to curved. Calvptra cucullate, 3 mm long, weak- ly hairy. Spores rounded, 12-20 pm, granulate, brownish. Fig. 163: 1-11.
Forsstroemia producta is a widespread species found on trees and boulders in eastern North America and Mexico, central South America, eastern and southern Africa, eastern Australia, Tasmania, Korea and China. In south- ern Africa the species is generally collected on trees and saplings in forests of the northern and eastern Transvaal areas, KwaZulu-Natal, and
Fig. 163. — Forsstroemia producta (1-11): 1. habit (dry), x 1.5; 2. stem in cross section (cells partly shown), x 175; 3. leaf, x 32; 4. leaf in cross section, x 175; 5. basal leaf cells, x 160; 6. upper laminal cells, x 350; 7. leaf apex, x 160; 8. perichaetial leaf, x 32; 9. part of capsule mouth with peristome teeth, x 175; 10. calyptra, x 32; 11. spore, x 700. Leptodon smithii (12-24): 12. habit (dry), x 1; 13. habit (wet), x 1; 14. part of stem in cross section, x 175; 15. leaves, x 32; 16. leaf in proximal cross section, x 175; 17. leaf in distal cross section, x 175; 18. basal leaf cells, x 160; 19. cells at leaf apex, x 160; 20. paraphyllia, x 160; 21. perichaetial leaf, x 35; 22. part of capsule mouth with peristome (papillae partly shown), x 175; 23. operculum, x 35; 24. spore, x 700. (1-11, Crosby 7915; 12 & 13, Crosby 7925; 14—21, Crosby 8054; 22-24, Smook 3957.)
588
Leptodontaceae
the eastern, southern, central and southwestern Cape regions. Map 228.
Vouchers: Crosby & Crosby 7515, 7974; Magill 3227, 6781; Smook 3960.
Plants of F. producta have a long-creeping primary stem from which erect secondary stems arise. The branched secondary stems grow out perpendicularly from the substrate and are quite distinctive when growing alone
on small saplings. The plants generally have many obvious perichaetia along the erect stems and frequently several sporophytes per stem. The leaves have a single costa that extends to midleaf or beyond and this should separate F. producta from all similar taxa except perhaps Cryphaea exigua. The im- mersed capsules, narrower leaves and toothed leaf margins of the latter separate it from F. producta.
2. LEPTODON
Leptodon Mohr, Observ. hot. 27 (1803), nom. cons.; Broth, in Natiirl. PflFam., edn 2, 11: 179 (1925); Sim, Bryo. S. Afr. 398 (1926); Sainsb., N. Zeal, mosses 360 (1955). Type species: L. smithii (Hedw.) Web. & Mohr.
Plants medium-sized, in loose tufts, green; saxicolous or corticolous. Stems strongly curved when dry, bipinnately branched; in section central strand absent; paraphyllia numerous. Leaves oval; obtuse; weakly decurrent at base; margins plane, entire. Laminal cells short, weakly thick- ened, smooth; basal and alar cells not strongly differentiated.
Dioicous. Seta short. Capsule just exserted, um short ellipsoidal. Peristome double; exostome teeth whitish, papillose; endostomes rudimentary. Operculum curved-rostrate. Calyptra conical, hairy.
Traditionally placed in the Neckeraceae, Leptodon is treated in Leptodontaceae on the basis of its habit; whitish, double peristome; split, conical calyptra; absence of a central strand in the stem; and foliaceous pseudoparaphyllia. The placement of the family in a linear sequence may seem remote from it former placement, but this family should be viewed as an intermediate between Leucodontaceae/Cryphaeaceae on the one hand and Neckeraceae on the other.
Leptodon smithii (Hedw.) Web. & Mohr , Ind. Mus. PI. Crypt. 2 (1803); Broth, in Natiirl. PflFam., edn 2, 11: 180 (1925); Sim, Bryo. S. Afr. 398 (1926); Smith, Moss FI. Brit. Irel. 504 (1978). Type: United Kingdom.
Hypnum smithii Dicks, ex Hedw., Sp. muse, frond. 264 (1801). Neckera smithii (Hedw.) C. Mull., Syn. muse, frond. 2: 118 (1850).
Plants medium-sized, gregarious or forming loose tufts, dark green to green or yellow-green; corticolous or saxicolous. Primary stems creep- ing; secondary stem erect, 20-50 mm long, branching regular above stipe, dense, twice-pin- nate in single plane; in section oval, central strand absent, inner cortical cells thin-walled, hyaline, in 5 or 6 rows, outer cortical cells smaller, yellow, thick-walled, in 3-5 rows; axil-
lary hairs numerous, to 9 per axil, 3 cells long, basal cell brownish; paraphyllia numerous, branched, multicellular but not foliose, smooth; pseudoparaphyllia foliose. Leaves evenly spaced, spreading wet, weakly crisped dry; stem leaves ovate to elliptical, 1-2 mm long; obtuse to acute; rounded and narrowed to inser- tion, weakly decurrent; margins plane, entire; branch leaves smaller but similar in other respects to stem leaves, 0. 5-1.0 mm long. Costa single, extending to midleaf or ending below apex, ventral and dorsal surface smooth, cells elongate; in section elliptical and bulging dor- sally, cells undifferentiated, in three rows. Upper laminal cells quadrate to rectangular or rhombic to transversely rectangular, somewhat heterogeneous, 15-32 pm long, 15 pm wide, walls weakly thickened, smooth; basal cells rec-
Leptodontaceae
589
tangular, to 40 pm long, 10-15 pm wide, green- ish, walls incrassate, ± wavy, smooth; alar cells quadrate, yellowish, walls ± incrassate.
Perigonia gemmate; perigonial leaves broad- ly ovate-acuminate, 1.0-1. 2 mm long. Perichaetia strongly differentiated; perichaetial leaves lanceolate-subulate, 3. 0-3. 5 mm long, leaf cells linear and somewhat sigmoid, rectan- gular at margins, thickened. Seta 0.5-2. 0 mm long, yellow, smooth; vaginula sometimes as long as seta, hairy, 0.6 mm long. Capsule exsert- ed, erect, short ellipsoid, 1.2-1. 6 mm long, smooth, brownish; neck not differentiated; exothecial cells rectangular, somewhat irregular, walls weakly thickened, cells at mouth quadrate to transversely rectangular; annulus absent; stomata absent. Peristome double, whitish to yellowish; exostome teeth narrowly triangular, inflexed dry, erect wet, weakly papillose, with median zigzag line, 250-280 pm high; endo- stome segments and cilia absent; basal mem- brane low, rudimentary. Operculum curved-ros- trate, 0.5 mm long. Calyptra campanulate- mitrate, 2. 0-2. 5 mm long, hairy. Spores rounded, 15-20 pm, granulate, brown. Fig. 163: 12-24.
Leptodon smithii is found on bark and rock in woodlands and forests of Europe, southwest- ern Asia, Macaronesia, northern, eastern and southern Africa, southern South America and the Juan Fernandez islands, Australia and New Zealand. In the Flora area, the species is found in the northern, central and eastern Transvaal
Map 229. — Leptodon smithii
areas, Zululand, KwaZulu-Natal, eastern Free State and the eastern, southern, central, south- western, and northwestern Cape regions. Map 229.
Vouchers: Crosby & Crosby 8053; Ester- huysen 21628; Goldblatt 2114a; Magill 3375; Van Rooy & Perold 3812.
The pronounced curving of the stems and branches when dry help to identify the species in the field. When wet, the stems are erect from the substrate and branches spread outward in a single plane. The stem is covered with numer- ous paraphyllia, a character which separates L. smithii from mosses with similar habits, e.g. Forsstroemia and Pterogonium.
590
NECKERACEAE
The family Neckeraceae contains 12 genera found in forests and woodlands in temperate and tropical areas. Only one genus is found in the Flora area.
NECKERA
Neckera Hedw., Sp. muse, frond. 200 (1801); Broth, in Natiirl. PflFam., edn 2, 11: 184 (1925); Sim, Bryo. S. Afr. 400 (1926). Lectotype: N. permata Hedw.
Plants small to large, creeping, green to yellow green or golden; corticolous or saxicolous. Stems appressed to substrate, densely leaved, flattened; central strand absent; paraphyllia filamentous or absent; pseudoparaphyllia filamentous or narrowly foliose. Leaves strongly complanate, undulate, oblong-acute; margins plane, entire to serrulate. Costa short and double. Laminal cells linear, smooth; basal cells not distinct; alar cells in small groups.
Autoicous. Perigonia and perichaetia along stems or branches. Seta short or long, smooth. Capsule immersed, erect, oblong. Peristome double, incomplete; exostome teeth narrow, striate at base; endostome segments well developed above low basal membrane, cilia absent. Operculum rostrate. Calyptra cucullate, smooth. Spores small, round.
Neckera , a genus of about 90 species, is found in temperate and tropical forests. Plants can be recognized by their flattened appearance and undulate leaves.
Neckera valentiniana Besch. in Ann. Sci. Nat. Bot. 6, 10: 273 (1880); Broth, in Natiirl. PflFam., edn 2, II: 185 (1925); Sim, Bryo. S. Afr. 400 (1926). Type: Reunion, Paves Saint- Leu, Valentin s.n., 1876.
Plants medium-sized to large, forming mats, green to yellow-green; corticolous or saxi- colous. Stems flattened, 40-60 mm long, branching irregular, ± scattered; in section oval, central strand absent, inner cortical cells thin- walled, hyaline, in 6-8 rows, outer cortical cells thick-walled, yellow-brown, in 3 or 4 rows. Leaves evenly spaced, complanate, weakly crisped at apex, undulate, somewhat falcate; oblong-ovate, 2-3 mm long; acute; rounded at base; margins plane, entire to serrulate above. Costa short and double, smooth. Upper laminal cells linear-fusiform, homogeneous, 40-75 pm long, 3-6 pm wide, walls thin, smooth; basal cells not strongly differentiated, greenish, walls thickened, smooth; alar cells in small groups, quadrate, yellowish, walls thickened.
Perigonia gemmate; perigonial leaves ovate- acuminate, 1.0-1. 5 mm long. Perichaetia strong- ly differentiated; perichaetial leaves elliptical- acuminate, spreading at tips, to 4 mm long, leaf cells linear, thin-walled. Seta 0.5- 1.0 mm long, yellow-brown, smooth. Capsule exserted, erect, short-cylindrical, 1 .5 mm long, smooth, yellow-brown; exothecial cells ± irregular, quadrate to rectangular, walls weakly thick- ened; cells at mouth smaller, quadrate, thick- ened; stomata phaneropore, on lower urn. Peristome double and incomplete, yellow; exostome teeth linear, reflexed wet, inflexed dry, with median zigzag line, smooth above, weakly striate at base, 500 pm high; endostome segments linear above very low basal mem- brane, as long as teeth, smooth; cilia absent. Operculum obliquely rostrate. Calyptra cucul- late, smooth. Spores rounded, 18-25 pm, gran- ulate, green. Fig. 164.
Neckera valentiniana forms large mats on trees and rocks in forests of southern Africa and
Neckeraceae
591
the East African islands. In the Flora area the species is found in wet forests of the northern, central and eastern Transvaal areas, Zululand, KwaZulu-Natal and the eastern, southern and southwestern Cape regions. Map 230.
Voucher: Esterhuysen 25043; Magill 5725, 6264; Oliver 6787; Von Breitenbach 221; Van Rooy 1062.
The large green mats and stems with com- planate, undulate leaves quickly identify this species in the Flora area. The plants are game- tophytically similar to the widespread species N. pennata Hedw., but differ in the well devel- oped endostomal segments produced by N. valentiniana. In addition, the numerous para- phyllia produced by N. valentiniana , while not unique, are generally absent in N. pennata.
Fig. 164. — Neckera valentiniana: 1. habit (dry, undu- lations partly shown), x 1.5; 2. part of secondary stem with branch and sporophyte (wet, undulations partly shown), x 3; 3. leaf, x 35; 4. basal leaf cells (left side), x 175; 5. upper laminal cells at right margin, x 50; 6. leaf apex, x 175; 7. paraphyllium, x 175; 8. perichaetial leaf, x 35; 9. part of capsule mouth with peristome, x 87; 10. spore, x 700. (1, Magill 6264\ 2, Von Breitenbach 221', 3-10, Van Rooy 1277.)
592
THAMNOBRYACEAE
Plants small to large or occasionally robust, dendroid, branches occasionally attenuate, dark green to yellow green or golden; corticolous or saxicolous. Primary stems long-creeping, appressed to substrate, naked; secondary stems erect and perpendicular to substrate; in section central strand small or absent; paraphyllia absent; pseudoparaphyllia narrowly foliose. Leaves concave, ovate to oblong; acute; margins plane, dentate. Costa strong and single. Laminal cells elongate or some- times rounded, smooth, sometimes pitted; basal cells distinct; alar cells in small groups and not strongly differentiated.
Dioicous. Perigonia and perichaetia along stems or branches. Seta long, smooth. Capsule exserted, erect. Peristome double, incomplete; exostome teeth narrow, striate below; endostome segments well developed above basal membrane, cilia generally absent. Operculum rostrate. Calyptra cucullate, smooth or with a few hairs. Spores small, round.
The recently segregated family Thamnobryaceae contains six genera found in forests and wood- lands of tropical and subtropical regions. Three genera are found in the Flora area.
1 Upper laminal cells irregularly rounded-quadrate 1. Pinnatella
1 Upper laminal cells rhombic to rhomboidal or fusiform:
2 Peristome complete, cilia present; leaves with marginal teeth made up of more than one
cell 2. Porothamnium
2 Peristome incomplete, cilia absent; leaves with marginal teeth or serrations made up of
single projecting cells 3. Porotrichum
1. PINNATELLA
Pinnatella Fleisch. in Hedwigia 45: 79 (1906); Broth, in Natiirl. PflFam., edn 2, 11: 195 (1925); Potier de la Varde in Mem. Soc. Sci. Nat. Cherbourg 42: 151 (1936); Enroth in Acta Bot. Fennica 151: 12 (1994). Lectotype species: P kuehliana (Bosch & Sande Lac.) Fleisch.
Plants small, dendroid, green; saxicolous. Secondary stems erect, branched above leafy stipe; branching mostly complanate. Leaves concave, somewhat plicate when dry, ovate; obtuse; strong- ly costate; branch leaves smaller. Leaf cells rounded-quadrate, somewhat irregular, thickened; alar cells not differentiated.
Dioicous. Seta short. Capsule exserted. Peristome double.
A genus of 15 species found in the wet tropical and subtropical forests of Africa, America, Asia and Australia. A single species is known from continental Africa.
Pinnatella minuta (Mitt.) Broth, in Natiirl. PflFam. 1,3: 857 (1906); Enroth in Acta Bot. Fennica 151: 12 (1994). Flolotype: Cuba, Wright s.n. (NY).
Porotrichum oblongifrondeum Broth, in Bot. Jahrb. Syst. 20: 200 (1894). Pinnatella oblongifrondea (Broth.) Broth, in Natiirl. PflFam. 1,3: 857 (1906). Lectotype: Tanzania, Usambara, Kwa Mshusa-Station, trockene Hochwalder bei Msingo, Holst 9193a (H).
Plants small to medium-sized, scattered or forming loose colonies, green; saxicolous or corticolous. Primary stems creeping; secondary stems erect, 8-12 mm long, dendroid, branch- ing irregular and ± complanate above stipe; in section oval, central strand absent, inner cortical cells thick-walled, yellow, in 6-8 rows, outer cortical cells somewhat smaller, thick-walled, yellow-brown; paraphyllia absent; pseudo-
Thamnobryaceae
593
paraphyllia foliose. Leaves evenly spaced, widespreading wet, somewhat crisped and incurved dry, concave; stem leaves ovate to broadly ovate, 1 mm long; obtuse, rounded- obtuse or acute; rounded to cordate at base; margins plane, entire below, weakly serrulate at apex; branch leaves ovate or ovate-ellipti- cal, 0.4-0. 6 mm long; obtuse to rounded or broadly acute; rounded at base; margins plane or erect, entire below, weakly serrulate at apex. Costa single, subpercurrent, smooth, frequent- ly spurred above; in section bulging dorsally, consisting of 3 or 4 rows of undifferentiated, thickened cells. Upper laminal cells irregular- ly rounded-quadrate, homogeneous, 6-12 pm long, walls thickened, smooth on both sur- faces; basal cells oblong, forming distinct group, 30-38 pm long, 5-7 pm wide, walls thickened, smooth; basal marginal cells small, quadrate, in 1 or 2 rows; alar cells not differ- entiated.
Sporophvte not known from the study area. Fig. 165: 1-8.
Pinnatella minuta is the only representative of the genus in the Neotropics, sub-Saharan Africa and the East African islands, and is also known from a single locality in the Nilgiri Mountains of southern India. In the Flora area the species is rare, having been collected only
Map 231. — ♦ Pinnatella minuta
• Porotrichum usagarum
twice in KwaZulu-Natal. Both of the South African specimens were sterile and appear to have been collected on rock, although this is not indicated on the labels. Map 231.
Vouchers: Sim 10327 , 10333.
The plants are most easily identified by their small dendroid habit, ovate leaves with obtuse apices, strong, single costa and short leaf cells. The plants are considerably smaller than all of the Porotrichum species, which share a similar habit.
2. POROTHAMNIUM
Porothamnium Fleisch., Muse. Buitenzorg 3: 927 (1908); Broth, in Natiirl. PflFam., edn 2, 11: 198 (1925); De Sloover in Bull. Jard. Bot. Belg. 53: 97-152 (1983). Type species: not selected.
Plants large, dendroid, in loose colonies; corticolous or saxicolous. Primary stems long-creep- ing, appressed to substrate; secondary stems erect, pinnately branched above stipe. Leaves ovate to broadly ovate-elliptical; apex acute to acuminate; margins plain, dentate. Costa single, extending to upper leaf, frequently ending in dorsal spine.
Dioicous. Capsule exserted, erect. Peristome double, complete; exostome teeth broad, striate; endostome segments broad, cleft and perforated along keel; cilia 3, as long as segments. Operculum rostrate. Calyptra cucullate, smooth. Spores small, granulate.
A genus of about 30 species found in wet forests of the Americas, Asia and Africa. Sim (1926) placed the species of Porotrichum in this genus but recent research by De Sloover (1983) has indi- cated that differences in peristome structure separate the two genera. In the Flora area the larger size and strong marginal leaf dentation of Porothamnium also separate it from Porotrichum.
Thamnobryaceae
595
Porothamnium stipitatum (Mitt.) Touw ex De Sloover in Bull. Jard. Bot. Belg. 53: 134 (1983). Type: Fernando Po, Mann s.n.
Trachyloma stipitatum Mitt, in J. Linn. Soc., Bot. 7: 156 (1864). .
Hypnum hildebrandtii C. Mull, in Linnaea 40: 287 (1876). Thamnium hildebrandtii (C. Mlill.) Jaeg. in Ber. Thatigk. St. Gallischen Naturwiss. Ges. 1877-1878: 467 (1880). Porothamnium hildebrandtii (C. Mlill.) Fleisch., Muse. Buitenzorg 3: 926 (1908). Type: Comoros, Johanna, Hildebrandt s.n., 1828 (BM, PC).
Neckera pterops Rehm. in Geh., Rev. Bryol. Lichenol. 5: 70 (1878), nom. nud. Porotrichurn pterops Rehm. ex Par., Index Bryol. suppl. 283 (1900), nom. nud. Type: not given, but Rehmann 329, fide Muller in Hedwigia 38: 130 (1899).
Thamnium afrum C. Mlill. in Hedwigia 38: 129 (1899). Type: South Africa, Natal, Inanda, Wood s.n. (Hb. Mac- Owan).
Plants large, frequently covering extensive areas, dark green; saxicolous or corticolous, rarely terricolous. Primary stems creeping; sec- ondary stems erect, to 1 20 mm long, stipe to 60 mm long, branches dense above stipe, branch- ing simple to pinnate; in section round, central strand small and dense in erect stems, inner cor- tical cells small, thin-walled, yellowish, outer cortical cells smaller, ± thick-walled, brown. Leaves flattened, spreading, weakly crisped when dry; stem leaves broadly ovate to oblong- ovate, ( 1 .0—) 1 .5— 2.5(— 3.0) mm long; acute to broadly acute; narrowed to insertion; margins plane, dentate, teeth multicellular; branch leaves similar to stem leaves but somewhat smaller. Costa single, extending to midleaf or above, smooth but terminating in a single dorsal tooth; in section bulging dorsally, in 3 rows of thickened, undifferentiated cells. Upper laminal cells irregularly rhomboidal, somewhat sig- moid, homogeneous, 35-50 pm long, 8-12 pm wide, walls weakly thickened, smooth; basal cells rectangular to rhomboidal, 40-75 pm
long, 8-12 pm wide, walls weakly thickened, smooth; alar cells not differentiated.
Perigonia axillary on branches, gemmate; perigonial leaves ovate, cuspidate, 1.0 mm long. Perichaetia along branches, obvious, green; perichaetial leaves oblong-acuminate, sheathing below, spreading at tips, to 2.5 mm long, leaf cells rhomboidal, thickened. Seta 15-25 mm long, red-brown, smooth. Capsule long-exserted, erect to inclined, ovoid, 2-3 mm long, smooth, red-brown, urn cylindrical, neck not differentiated; exothecial cells quadrate to rectangular or hexagonal, walls thickened, cells at mouth quadrate, strongly thickened, darker. Peristome double, yellow-brown; exostome teeth triangular, inflexed wet, erect dry, striate with median zigzag line, to 500 pm high; endostome segments above high basal mem- branes lanceolate and perforated or broad and cleft with diverging tips, shorter than teeth, weakly granulate; cilia present, linear, shorter than segments, weakly granulate. Operculum rostrate, 1 mm long. Calyptra cucullate, 2 mm long, smooth. Spores rounded, 9-15 pm, weak- ly granulate, brownish. Fig. 165: 9-17.
This species is found in moist forests of northern South America, sub-Saharan Africa, and the East and West African islands. In the Flora area P. stipitatum is found on rocks and bark in the forests of the northern and eastern Transvaal areas, Swaziland, Zululand, Kwa- Zulu-Natal and the eastern, southern and south- western Cape regions. Map 232.
Vouchers: Brenan M2845, Crosby & Crosby 9207; Magill 5127, 6023; Van Rooy 1783.
Gametophytically the relatively large size, dark green colour, and strongly dentate leaf margins identify this species. The leaf denta- tions are large enough to be seen easily with a hand lens and are made up of 2—4 cells each.
Fig. 165. — Pinnatella minuta (1-8): 1. habit (dry), x 1; 2. habit (wet), x 5; 3. part of stem in cross section, x 175; 4. stem leaf, x 35; 5. branch leaf, x 35; 6. stem leaf base (right side), x 175; 7. upper laminal cells of stem leaf, x 700; 8. stem leaf apex, x 175. Porothamnium stipitatum (9-17): 9. habit (dry), x 1; 10. part of stem with branch and sporophytes (wet), x 1.2; 11. part of stem in cross section, x 175; 12. stem leaf, x 35; 13. branch leaf, x 35; 14. upper laminal cells of stem leaf at left margin, x 350; 15. perichaetial leaf, x 18; 16. part of capsule mouth with peristome (papillae partly shown), x 87; 17. spore, x 700. (1-8, Sim 10327\ 9, Brenan 335P, 10, Sim PRE-CH9914: 1 1-17, Sim PRE-CH9901.)
596
Thamnobryaceae
The species is separated from Porotrichwn by its perfect peristome with striate teeth, broad segments and well-developed cilia.
Insufficiently known species
Porothamnium capense (Broth. & Dix.) Sim, Bryo. S. Afr. 404 (1926). Thamnium capense Broth. & Dix. in Bull. Torrey Bot. Club 43: 71 (1916). Type: In packing from Cape Town, 710. Communicated by G. Webster (BM!). The specimen is sterile and consists of a single, weakly branched creeping stem with leaves sharply serrate throughout and rounded-hexag- onal leaf cells and a tooth at the end of the costa. De Sloover (1983) suggested Thamnobryum Nieuwl. as a possible placement; Enroth (1991) subsequently made that combination — T. ca- pense (Broth. & Dix.) Enroth. The stem seemed more reminiscent of Ectropothecium Mitt., but since the plants have not been rediscovered and the collection site is unknown, the species is not formally treated here.
3. POROTRICHUM
Porotrichum (Brid.) Hampe in Linnaea 32: 154 (1863); Broth, in Natlirl. PflFam., edn 2, 11: 196 (1925); Bartram in Fieldiana, Bot. 25: 284 (1949); De Sloover in Bull. Jard. Bot. Belg. 53: 97-152 (1983). Lectotype species: P. longirostre (Hook.) Mitt.
Plants large, in loose colonies; corticolous, saxicolous or terricolous. Primary stems long-creep- ing, appressed to substrate; secondary stems erect, pinnately branched above long stipe. Leaves ovate to elliptical; apex acute to acuminate; margins plain, serrate above, entire below. Costa sin- gle, extending to upper leaf, frequently ending in dorsal spine.
Dioicous. Capsule exserted, erect. Peristome double; exostome teeth narrow, papillose to striate below; endostome segments narrow, perforated, as long as teeth, cilia absent. Operculum rostrate. Calyptra cucullate. Spores papillose.
A genus of over 50 species found in the wet temperate to tropical areas of Central and South America, Africa and Asia. The separation of this genus from Porothamnium follows the treatment of De Sloover (1983) and is based on several easily observable gametophytic characters and sig- nificant differences in the peristomes of the two genera.
Gametophytically the African species placed in Porotrichum are smaller plants, dark green to yellow-green in colour, and have serrate to denticulate upper leaf margins. The differences exhib- ited by the peristomes of Porotrichum and Porothamnium are reminiscent of the differences between the Hookeriaceous and Daltoniaceous peristomes (Whittemore & Allen 1989).
The peristome teeth of the African Porotrichum species are narrow and papillose to papillose- striate below while the African Porothamnium species have broad teeth that are characterized as striate throughout. The endostome segments are narrow and perforated and the cilia are absent or rudimentary in these species of Porotrichum , while Porothamnium has broad, cleft or perforated segments and well-developed cilia.
Thamnobryaceae
597
1 Leaf cells short, rhomboidal; leaves broadly elliptical, apex acute, margins denticulate to
midleaf or below; plants green to dark green 1 . P. usagarum
1 Leaf cells longer, rhomboidal to fusiform; leaves ovate to ovate-elliptical, apex short acuminate, margins serrate above; plants green to yellow-green:
2 Costa disappearing in upper leaf, not ending in dorsal spine; common in moist forests . .
2. P. madagassum
2 Costa ending in short dorsal spine; rare 3. P. elongatum
1. Porotrichum usagarum Mitt, in J. Linn. Soc., Bot. 22: 315 (1886); Enroth in Ann. Bot. Fenn. 28: 197-200 (1991); Enroth & Hodgetts in J. Bryol. 19: 140 (1996). Lectotype: Tanzania, Usagara Mountains, Hannington s.n. (NY; H, S, isolecto.) fide Enroth (1991).
Porotrichum molliculum Broth, in Bot. Jahrb. Syst. 24: 257 (1897). Thamnium molliculum (Broth.) Kindb. in Hedwigia41: 220 (1902). Porothamnium molliculum (Broth.) Fleisch. in Broth, in Natiirl. PflFam., edn 2, 11: 199 (1925). Syntypes: Tanzania, Volkens s.n., 1941; Scott Elliott 260.
Porotrichum natalense C. Mull, in Hedwigia 38: 129 (1899). Porothamnium natalense (C. Mull.) Fleisch., Muse. Buitenzorg 3: 926 (1908). Thamnium natalense (C. Mull.) Kindb. in Hedwigia 41: 239 (1902). Type: Natal, Inanda, Rehmann 334 (PRE).
Thamnium penniforme (‘ pennaeforme ’) Kindb. var. brachyphyllum Dix. in Trans. Roy. Soc. South Africa 18: 257 (1929). Type: Transkei, Port St Johns, Wager 954 (PRE).
Plants medium-sized, in loose colonies, dark green and somewhat glossy; corticolous or saxi- colous. Primary stems long-creeping; sec- ondary stems erect, to 120 mm long, branching somewhat irregular above stipe; in section round, central strand absent, inner cortical cells thin-walled, hyaline, in 8-10 rows, outer corti- cal cells smaller, thick-walled, yellow-brown, in 8-12 rows. Leaves ± crowded, spreading wet, appressed dry; stem leaves ovate to ovate-ellip- tical, 1. 5-3.0 mm long; apex acute or acumi- nate, apiculate on larger, broader leaves; nar- rowed to insertion; margins plane, denticulate; branch leaves ovate to elliptical, 1.0- 1.5 mm long; acute; narrowed to insertion; margins plane, denticulate, cells not differentiated. Costa single, extending to midleaf or above, smooth, ending distally in short dorsal spine, ventral and dorsal surface smooth; in section elliptical, cells not strongly differentiated. Upper laminal cells rhombic to rhomboidal.
homogeneous, 12-22 pm long, 5-6 pm wide, walls somewhat thickened, smooth; basal cells rectangular, 12-40 pm long, 5-6 pm wide, hya- line, walls thickened, smooth; alar cells not dif- ferentiated.
Perigonia on stems and branches, gemmate; perigonial leaves ovate. Perichaetia along stem and branches, green; perichaetial leaves oblong- acuminate, 1.2-1. 8 mm long. Seta 10-15 mm long, yellow-brown, smooth. Capsule exserted, erect, ellipsoid, 2.0-2.5 mm long, smooth, brown. Peristome double, brown; exostome teeth linear, papillose, 600-800 pm high; endostome seg- ments linear above low basal membrane, as long as teeth, papillose, cilia absent. Operculum long- rostrate, 1.5 mm long. Calyptra cucullate, smooth or with a few hairs. Spores rounded, 12-15 pm, granulate, brown. Fig. 166: 10-14.
This species is known from moist forests of central, eastern and southern Africa, Madagas- car and the East African islands. In the Flora area P. usagarum is infrequently collected in the forests of Zululand and KwaZulu-Natal, and has rarely been found in the northern and eastern Transvaal areas and the eastern Cape region. Map 231.
Vouchers: Brenan M3356; Crosby & Crosby 7574; Magill 4908; Van Rooy 158. ’
The plants are generally smaller, darker in colour and have shorter leaves than the other species of Porotrichum in the Flora area. In addition the leaf cells are shorter and broader especially at the apex and the margins are more strongly denticulate. This species could be con- fused with very small plants of Porothamnium stipitatum , but that species has marginal teeth on the leaf blade which are composed of 2^1 cells each.
Thamnobryaceae
599
2. Porotrichum madagassum Kiaer ex Besch. in Ann. Sci. Nat. Bot. 6, 10: 332 (1880). Type: Madagascar, Borgen s.n., 1875.
Porotrichum penniforme ('pennaeforme') C. Mull, in Hedwigia 38: 127 (1899). Porothamnium penniforme (‘ pen- naeforme') (C. Miill.) Fleisch., Muse. Buitenzorg 3: 926 (1908). Syntypes: South Africa, Cape Prov., Oudebosch, Breutel s.n.; Blanco, Rehmann s.n., Oct. 1875; Somerset East, Boschberg, MacOwan s.n., Nov. 1873 (PRE).
Plants medium-sized to large, in loose colonies, green to yellow-green; saxicolous or sometimes corticolous. Primary stems long- creeping; secondary stems erect, 60-100 mm long, branching irregular above stipe, ± den- droid; in section round to oval, central strand small, inner cortical cells thin-walled, in 12-14 rows, outer cortical cells thick-walled, red- brown, in 8-10 rows. Leaves somewhat crowd- ed, flattened, erect-appressed; stem leaves broad- ly ovate to elliptical, 1. 5-2.5 mm long; acute to short-acuminate; narrowed to insertion; margins plane, serrate above, entire below; branch leaves ovate to elliptical, somewhat smaller. Costa sin- gle, extending to midleaf or above, smooth, not ending in spine; in section elliptical, of 2 or 3 rows of undifferentiated cells. Upper laminal cells rhomboidal to somewhat fusiform, homo- geneous, 20-40 pm long, 5-6 pm wide, walls weakly thickened, smooth; basal cells ± oblong, not strongly differentiated, 35-65 pm long, 6-8 pm wide, greenish, walls thickened and pitted, smooth; alar cells not differentiated.
Perigonia gemmate. Perichaetial leaves oblong-acuminate, to 3 mm long, leaf cells oblong, thickened and pitted. Seta 10-15 mm long, red-brown, smooth. Capsule exserted, erect, ovoid, 1. 5-2.0 mm long, smooth, red- brown; exothecial cells ± hexagonal, walls thin; cells at mouth rounded; annulus present, differ- entiated; stomata phaneropore on neck. Peri- stome double, yellow; exostome teeth linear,
recurved dry, inflexed wet, papillose-striate, with median zigzag line or perforated along midline, 700-800 pm high; endostome segments linear and perforated above low basal membrane, as long as teeth, papillose; cilia absent. Operculum obliquely rostrate, 1 mm long. Calyptra cucul- late, ± smooth. Spores rounded, 18-20 pm, papillose, brownish. Fig. 166: 1-9.
Porotrichum madagassum is frequently found in moist forests of central, eastern and southern Africa, Madagascar and the East African islands. In southern Africa the species is found in the forests of the northern and eastern Transvaal areas, Swaziland, Zululand, KwaZulu- Natal and the eastern, southern and southwest- ern Cape regions. Map 233.
Vouchers: Crosby & Crosby 8034; Magill 7002; Oliver 6807; Von Breitenbach 174.
In addition to being the most commonly col- lected species of Porotrichum in the Flora area, P. madagassum is recognized by its yellow- green colour, ± flaccid appearance and leaves that appear more delicate than other related
Fig. 166. — Porotrichum madagassum (1-9): 1. habit (wet), x 1; 2. part of stem in cross section, x 175; 3. stem leaf, x 35; 4. branch leaf, x 35; 5. basal cells of stem leaf (left side), x 175; 6. upper laminal cells of stem leaf at left margin, x 350; 7. perichaetial leaf, x 35; 8. part of capsule mouth with peristome, x 70; 9. spore, x 700. P. usagarum (10-14): 10. habit (dry), x 1; 11. branch leaf, x 35; 12. stem leaf, x 35; 13. upper laminal cells of stem leaf at left margin, x 350; 14. stem leaf apex, x 175. P. elongatum (15-19): 15. habit (dry), x 1 ; 16. stem leaf, x 35; 17. branch leaf, x 35; 18. upper lami- nal cells of stem leaf showing dorsal spine of costa, x 700; 19. stem leaf apex, x 175.(1, Van Rooy 1622; 2, 4 & 6, Van Rooy 1426; 3 & 5, Von Breitenbach 153; 7, Wood 2651; 8 & 9, Sun 7282; 10-14, Van Rooy 1884; 15, 16, 18 & 19, Van der Schijff 4956; 17, Sim PRE-CH9906.)
600
Thamnobryaceae
species. The species is quite variable and indi- vidual specimens have been confused with each of the other species. It is perhaps most difficult to separate from P. elongation (see note below), but that species has the costa ending in a dorsal spine. Specimens with shorter leaf cells have been confused with P. usagarum , but that species is always smaller, darker green and has shorter, broader leaves. The very strong teeth on the leaf margins of Porothamnium stipitatum separate it from P. madagassum which has only a weakly serrate apex.
3. Porotrichum elongatum (Welw. & Dub.) Gepp in Hiem, Cat. Afr. PI. 2,2: 294 (1901); Broth, in Natiirl. PflFam., edn 2, 11: 197 (1925); De Sloover in Bull. Jard. Bot. Belg. 53: 101 (1983). Type: Angola, Golungo-Alto region, Welwitsch s.n.
Homalia elongata Welw. & Duby in Mem. Soc. Phys. Geneve 2 1 : 429 (1871).
Porotrichum comorense Hampe ex C. Miill. in Linnaea 40: 270 (1876); Broth, in Natiirl. PflFam., edn 2, 11: 197 ( 1925). Porothamnium comorense (C. Miill.) Sim, Bryo. S. Afr. 402 (1926). Type: Comoros, Irhamia, Hildebrandt s.n., 1834 (BM, PC).
Plants medium-sized, forming loose colonies, green to yellow-green; corticolous. Primary stems creeping; secondary stems erect, 30-120 mm long, branching irregular above stipe, pinnate or lower branches irregularly once-pinnate; in sec- tion round to oval, central strand small, inner cor- tical cells thin-walled, hyaline, in 10-12 rows, outer cortical cells thick-walled, yellow, in 4—6 rows. Leaves spreading wet, appressed dry; stem leaves broadly ovate to elliptical, 1.0-1. 5 mm long; short acuminate; rounded at base; margins plane to erect above, serrulate above; branch leaves similar to stem leaves but smaller and more elliptical, 0.5-1. 0 mm long; acute to acuminate; rounded at base; margins plane, serrulate above. Costa smooth but ending below apex in a short dorsal spine; in section elliptical. Upper laminal cells rhomboidal to fusiform, homogeneous, 20-40 pm long, 5-7 pm wide, walls thin, smooth; basal cells fusiform and longer but not forming distinct groups, greenish, walls ± thickened, smooth; alar cells not forming distinct groups.
Perigonia and perichaetia not seen. Sporo- phyte not found in southern Africa but described as: Seta erect, to 14 mm long. Capsule erect, elliptical, 1.5-1. 8 mm long. Peristome teeth 700-750 pm, papillose; endostome segments almost as long as teeth, cilia absent. Operculum conic-rostrate, 1.0- 1.5 mm long. Calyptra cucullate, smooth, 2.0-2. 5 mm long. Spores round, 10-15 pm, finely papillose, light yellow. Fig. 166: 15-19.
Porotrichum elongatum is found in forests of sub-Saharan Africa, Madagascar and the East and West African islands. In the Flora area the species is extremely rare and currently known from a few specimens collected in KwaZulu- Natal and the northern and eastern Transvaal regions. Map 234.
Vouchers: Loocke PRE-CH11205; J. Sim PRE-CH9906; Van der Schijff4956.
Only a few South African specimens appear to fit within the concept of this species. The col- lections are very similar to P. madagassum but differ in having the costa end in a short dorsal spine. All of the specimens seen from southern Africa are poor or mixed and may indicate that these specimens represent random introductions and not an established species. The prorate leaf cells described for this species were not found on the South African specimens. It is also pos- sible that these specimens represent an aberrant form of P. madagassum.
601
HOOKERIACEAE
Plants small to large, in tufts or mats, dark green to yellow-green; corticolous, terricolous or saxi- colous. Stems generally creeping, short; central strand present or absent; paraphyllia absent; pseudopa- raphyllia absent. Leaves large, frequently appressed and flattened; margins frequently bordered, toothed or entire. Costa generally strong, double, extending to above midleaf, sometimes single or short and double. Laminal cells large, firm-walled, frequently pitted; alar cells not differentiated.
Perichaetia lateral along stems, generally small. Seta mostly short, smooth or rough above, some- times throughout. Capsule small, inclined, dark brown. Peristome double, well developed; exostome teeth striate or papillose below, frequently strongly grooved; endostomes generally lacking cilia. Operculum rostrate. Calyptra cucullate, smooth to rough or hairy. Spores small, weakly ornamented.
The family is widespread in the tropics and warm, moist temperate regions. It contains 32 gen- era; only 8 are presently known from the Flora area.
1 Costa ending in lower leaf, short and double or single and bifurcate 1. Calyptrochaeta
1 Costa extending above midleaf, single or double:
2 Costa double:
3 Leaf cells variably papillose; leaves frequently blunt or rounded at apex 2. Callicostella
3 Leaf cells smooth; leaves acute:
4 Leaves bordered, margins entire or serrate above; leaf cells subquadrate to hexagonal
3. Cyclodictyon
4 Leaves not bordered, margins serrate above; leaf cells rhomboidal:
5 Leaves coarsely and bluntly serrate above by inflated cells that are frequently two-
pointed; seta smooth; capsule inclined to nodding 4. Hookeriopsis
5 Leaves sharply serrate above by narrow, single-pointed cells; seta scabrous; capsule
± erect 5. Lepidopilidium
2 Costa single:
6 Stems simple or weakly branched, homophyllous; leaf margins entire ... 6. Distichophyllum 6 Stems highly branched, heterophyllous; leaf margins serrate:
7 Plants flabellate; amphigastria broadly ovate, abruptly cuspidate 7. Hypopterygium
7 Plants pinnate; amphigastria lanceolate 8. Lopidium
1. CALYPTROCHAETA
Calyptrochaeta Desv. in Mem. Soc. Linn. Paris 3: 226 (1825). Type species: C. cristata (Hedw.) Desv.
Eriopus Brid., Bryol. univ. 2: 788 (1827); Sim, Bryo. S. Afr. 442 (1926); Broth, in Natilrl. PflFam., edn 2, 11: 232 (1925). Type species: E. cristata (Hedw.) Brid.
Plants medium-sized, in tufts; saxicolous or terricolous. Stems erect, sparsely branched; central strand present. Leaves flattened in single plane, larger above; elliptical to broadly spathulate; rounded-apiculate; margins bordered, plane, serrate. Costa short and' double. Laminal cells large, hexagonal, thin-walled, pitted.
602
Hookeriaceae
Hookeriaceae
603
Dioicous. Seta roughly papillose throughout. Capsule inclined. Peristome double, complete; cilia present. Operculum rostrate. Calyptra cucullate. Spores small.
This genus includes 15 species found mostly in the southern hemisphere. A single species is known from eastern and southern Africa and Madagascar. Some specimens of Calyptrochaeta could be confused with Plagiomnium Koponen; however, that genus has leaves with a long, single costa and a truncate apex.
Calyptrochaeta asplenioides {Brid.) Cros- by in Rev. Bryol. Lichenol. 42: 712 (1976). Type: in Insula Barbonia habitat, Bory St Vincent s.n. (BM).
Pterygopltyllum asplenioides Brid., Muscol. recent, suppl. 4: 151 (1818). Eriopus asplenioides (Brid.) Besch. in Ann. Sci. Nat. Bot. 6, 10: 281 (1880); Broth, in Natiirl. PflFam., edn 2, 11:233(1925).
Hookeria mniacea C. Mull, in Bot. Zeitung (Berlin) 17: 247 (1859). Eriopus mniaceus (C. Mull.) Broth, in Natiirl. PflFam. 1,3: 931 (1907); Sim, Bryo. S. Afr. 442 (1926). Type: Prom, bonae spei, Soutkloof, Breutei s.n.
Plants medium-sized to large, forming turfs, light green to dark green; saxicolous or terri- colous. Stems suberect, 10-30 mm long, branching sparse, irregular; in section round, central strand small, inner cortical cells thin- walled, hyaline, in 3 or 4 rows, outer cortical cells somewhat thick-walled, yellow to red, in 3 rows. Leaves evenly spaced, larger above, spreading wet, appressed dry, ± flattened into one plane; somewhat asymmetrical, broadly ovate to elliptical or broadly spathulate, largest leaves 3-6 mm long; rounded and apiculate; strongly narrowed to insertion; margins plane, serrate to denticulate above midleaf; bordered by elongated, thickened cells, unistratose. Costa short and double or single and bifurcate, ending in lower leaf, smooth; in section ellipti-
cal, cells small, thickened. Upper laminal cells homogeneous, hexagonal to rhomboidal, 100-112 pm long, 50-56 pm wide, walls thick- ened, pitted, smooth; basal cells rhomboidal, greenish, walls thickened, pitted, smooth; alar cells not differentiated.
Dioicous. Perigonia not seen. Perichaetia small, green; perichaetial leaves ovate-acumi- nate but with some leaves truncate, 1-2 mm long, leaf cells rectangular to rhombic or fusiform. Seta to 10 mm long, yellow-brown, roughly papillose throughout. Capsule exsert- ed, inclined to nodding, weakly pyriform, 1.8-2. 5 mm long, smooth, red brown; urn ovoid; neck differentiated, shorter than urn, curved; exothecial cells rounded to quadrate, walls thickened, collenchymatous, cells at mouth quadrate, neck cells irregular, quadrate to rectangular, stomata not seen. Peristome double, yellow; exostome teeth triangular, reflexed dry, inflexed wet, striate below with obvious zigzag line, papillose above, 400 pm high; endostome segments lanceolate above high basal membrane, keeled, as long as teeth, weakly granulate; cilia single, broad, almost as long as segments, weakly granulate. Operculum convex-rostrate, 0.5 mm long. Calyptra cucul- late, rough when young. Spores rounded, 12-17 pm, weakly granulate, yellow-brown. Fig. 167: 1-11.
Fig. 167. — Calyptrochaeta asplenioides ( 1-1 1): 1. habit (dry), x 1; 2. habit (wet), x 3; 3. part of stem in cross section, x 87; 4. leaf, x 18; 5. part of leaf in cross section, x 175; 6. basal leaf cells (right side), x 70; 7. cells at leaf apex, x 70; 8. perichaetial leaf, x 18; 9. seta in cross section, x 87; 10. part of capsule mouth with peristome, x 175; 11. spore, x 700. Callicostella tristis (12-24): 12. habit (dry), x 1; 13. habit (wet), x 3; 14. part of stem in cross section, x 175; 15. leaf, x 35; 16. leaf in cross section, x 175; 17. basal leaf cells (right side), x 175; 18. upper laminal cells at right margin, x 350; 19. leaf apex, x 175; 20. perichaetial leaf, x 35; 21. part of capsule mouth with peristome, x 175; 22. operculum, x 35; 23. calyp- tra, x 28; 24. spore, x 700. ( 1 , 2 & 5, Crosby 8606 ; 3, Magiil 6006; 4, 6 & 7, Jacot Guillarmod PRE-CHI 2571- 8-11, Crosby 7389; 12 & 13, Van Rooy 166; 14-16, 18-22 & 24, Van Rooy 977; 17, Wager PRE-CHI 1624; 23. Taylor PRE-CH12635.)
604
Hookeriaceae
Map 235. — ♦ Calyptrochaeta asplenioides • Callicostella tristis
Calyptrochaeta asplenioides is known from Tanzania, Madagascar, the East African islands and South Africa. In the Flora area it is restrict- ed to indigenous forests of the southern and southwestern Cape regions. The relatively large plants are found on rock or humus near streams or on rock overhangs where water is plentiful. Map 235.
Vouchers: Crosby & Crosby 8606; Ester - huysen 26687; Magill 6006.
Within the family the species is recognized by its large bordered leaves, short, mostly dou- ble costa, and large hexagonal leaf cells. Sporophytes are rare but when present the papil- lae on the seta are obvious with a hand lens.
2. CALLICOSTELLA
Callicostella (C. Mull.) Mitt, in J. Linn. Soc., Bot. Suppl. 1: 136 (1859), nom. cons.; Sim, Bryo. S. Afr. 444 (1926); Broth, in Natiirl. PflFam., edn 2, 11: 238 (1925). Type species: C. papillata (Mont.) Mitt.
Schizomitrium B.S.G., Bryol. eur. 5: 59 (1851), nom. rejic. Type species: not designated.
Plants small, in mats; terricolous, corticolous or saxicolous. Stems creeping; central strand absent. Leaves complanate, broadly oblong to lingulate; truncate; margin serrate, not bordered. Costa double, extending to upper leaf. Alar cells not differentiated.
Seta elongate, ± smooth. Capsule horizontal, dark red. Peristome double, incomplete; cilia absent. Operculum beaked. Calyptra mitriform. Spores small, green.
A genus of over 100 species found in tropical and subtropical forests, mostly of the southern hemisphere.
Callicostella tristis (C. Mull.) Broth, in Natiirl. PflFam. 1,3: 938 (1907); Sim, Bryo. S. Afr. 444 (1926); Broth, in Natiirl. PflFam., edn 2, 11: 239 (1925). Type: South Africa, Natal, Inanda, Rehmann s.n. (BM, NH, in PRE Reh- mann specimen numbered as 624).
Hookeria tristis C. Mull., Hedwigia 38: 130 (1899). Schizomitrium triste (C. Mull.) Ochyra in Ochyra & Poes, Acta Bot. Acad. Sci. Hung. 28: 382 (1982).
Callicostella applanata Broth. & Bryhn in Bryhn, Forh. Vidensk.-Selsk. Kristiania 4: 19(1911 ); Sim in Bryo. S. Afr. 445 (1926); Broth, in Natiirl. PflFam., edn 2, 11: 239 (1925). Schizomitrium applanatum (Broth. & Bryhn) Ochyra in Ochyra & Poes, Acta Bot. Acad. Sci. Hung. 28:
381 (1982). Type: South Africa, Zululand, Eshowe, H. Bryhn. Nov. 1908 (H-BR).
Plants small, forming mats, grey-green to yellow-green; terricolous, humicolous, saxi- colous or corticolous. Stems creeping, 15-30 mm long, branches irregular, scattered; in sec- tion round or oval, central strand absent, inner cortical cells thin-walled, in 2 or 3 rows, outer cortical cells smaller, ± thick-walled, red- brown. Leaves evenly spaced, spreading wet, ± crisped and curved downward dry; broadly ovate to oblong, 1-2 mm long; apex variable, truncate or rounded to obtuse or almost acute;
Hookeriaceae
605
rounded and narrowed to insertion; margins plane, serrate, cells not differentiated. Costa double, ending below apex, with a single dorsal tooth at each tip, smooth or ridged dorsally; ventral surface smooth, cells elongate; dorsal surface toothed above, bulging or ridged, cells elongate; in section bulging dorsally. Upper laminal cells irregularly shaped, homogeneous, 12-20 pm long, 5-8 pm wide, walls weakly thickened, papillose above dorsally but some papillae on ventral surface; papillae single, low, centred over lumen; basal cells almost rectan- gular, forming distinct group, 25-37 pm long, 8-15 pm wide, hyaline, smooth; alar cells not forming distinct groups.
Dioicous? Perichaetia small, green; peri- chaetial leaves ovate to oblong, 1 mm long; acute to acuminate; leaf cells ± rectangular. Seta 10-15 mm long, red-yellow, ± smooth. Capsule exserted, horizontal, ovoid, 1 mm long, smooth, dark red; urn ovoid; neck tapering, shorter than urn; exothecial cells short rectangular, walls collenchymatous, cells at mouth transversely rectangular, neck cells rectangular; stomata on neck cryptopore, formed of 4 cells, some not functional. Peristome double, red; exostome teeth triangular, inflexed at tips dry, erect appressed wet, striate and deeply furrowed below, papillose above, 200 pm high; endo- stome segments lanceolate and keeled above high basal membrane, as long as teeth, weakly
granulate; cilia absent. Operculum rostrate, when young body yellow, beak dark red, 0.5 mm long. Calyptra large, mitrate, 2 mm long, rough above. Spores rounded, 10-13 pm, weak- ly papillose, green. Fig. 167: 12-24.
Reported from central, western and southern Africa, Callicostella tristis is found on rocks, humus or soil in forests of the northern, central and eastern Transvaal areas, Swaziland, Zululand, KwaZulu-Natal, and the eastern Cape region. Eleven other species have been recorded from east- ern Africa, several related to C. tristis. Map 235.
Vouchers: Bernard 9076-A; Edwards 13; Magill 5337; Van Rooy 166 , 1886.
The leaves of this species are complanate and become somewhat crisped and down- turned when dry. The unbordered leaves with strong, double costae and papillose upper lami- nal cells place the plants in the Hookeriaceae. The papillae are variable and leaf cells must fre- quently be cleared to detect them.
Callicostella tristis is similar to the wide- spread tropical American species C. pallida (Homsch.) Angstr., but differs mostly in a weaker ornamentation of its seta and variable leaf cell papillae. The characters used to differ- entiate C. applanata are quite variable even on the type specimen; this species is therefore placed in the synonymy of C. tristis.
3. QYCLODICTYON
Cyclodictyon Mitt, in J. Linn. Soc., Bot. 7: 163 (1864); Sim, Bryo. S. Afr. 442 (1926); Broth, in Natiirl. PflFam., edn 2, 11: 236 (1925). Type species: C. pteridioides R Beauv.
Plants small to medium-sized, in small mats, green to yellow green; humicolous or saxicolous. Stems creeping; central strand absent. Leaves flattened, broadly ovate to elliptical; margins bor- dered, entire to serrate. Laminal cells large, ± hexagonal, thin-walled; basal cells rectangular; alar cells not differentiated.
Seta elongate, smooth. Capsule horizontal, blackish. Peristome double, complete; cilia single, rudimentary. Operculum rostrate. Calyptra mitrate, rough. Spores small, green.
A tropical to subtropical genus of about 100 species, most of which are found in the Americas. About 20 taxa have been reported or described from Africa, and one or two species are known from Europe, Australia and Pacific islands.
Hookeriaceae
607
Cyclodictyon vallis-gratiae (Hampe) O. Kuntze, Revis. gen. pi. 2: 835 (1891); Sim, Bryo. S. Afr. 443 (1926); Broth, in Natiirl. PflFam., edn 2, 11: 237 (1925). Type: Prom, bon. spei, Genadendal, s.n. & s.l. (BM herb. Hampe).
Hookeria vallis-gratiae Hampe ex C. Mull, in Bot. Zeitung (Berlin) 16: 169 (1858).
Hookeria breuteliana Hampe in C. Miill. in Bot. Zeitung (Berlin) 17: 247 ( 1859). Cyclodictyon breutelianum (Hampe) O. Kuntze, Revis. gen. pi. 2: 835 (1891 ); Broth, in Natiirl. PflFam., edn 2, 11: 236 ( 1925). Cyclodictyon vallis- gratiae fo. breuteliana Demar. & P. Varde in Bull. Jard. Bot. Etat 21: 45 (1951). Type: Prom. bon. spei, Oude Bosch inter H. vallis-gratiae, Breutel s.n. (BM).
Hookeria breutelii Hampe ex Kindb., Enum. Bryin. exot. 16 (1888), nom. illeg. inch spec, prior.
Plants small to medium-sized, forming mats, green; humicolous or saxicolous. Stems creep- ing, 10-30 mm long, branching sparse, irregu- lar; in section oval, central strand absent, inner cortical cells large, thin-walled, outer cortical cells larger, thin-walled. Leaves somewhat dis- tant, spreading wet, spreading and weakly crisped dry; asymmetrical, oblong to lingulate, 1.2-2. 5 mm long; apex rounded and apiculate; narrowed to insertion; margins plane, serrate above, entire below, weakly bordered by elon- gated cells, unistratose. Costa double, ending below apex, smooth but each ending in a dorsal tooth; in section bulging dorsally, consisting of a small group of weakly thickened cells. Upper latninal cells quadrate to hexagonal, homoge- neous, 25-37 pm long, walls thin, smooth on both surfaces; basal cells forming distinct group, rectangular to rhomboidal, 50-75 pm long, 25 pm wide, greenish, walls thin, smooth; alar cells not differentiated.
Autoicous. Perigonia on stem, gemmate; perigonial leaves lanceolate. Perichaetia along
stem, small, green; perichaetial leaves ovate- acuminate, 1-1.5 mm long, leaf cells large, thin- walled. Seta 14—18 mm long, red-brown, smooth. Capsule exserted, horizontal, ellipsoid, 1-2 mm long, smooth, red-brown, urn ellipsoidal; neck tapering, shorter than urn; exothecial cells rectan- gular, walls firm, cells at mouth smaller, quadrate to rectangular, neck cells rectangular; stomata phaneropore, on neck, consisting of 3-6 cells, some not functional. Peristome double, red-yel- low; exostome teeth lanceolate, inflexed at tip dry, incurved appressed wet, striate below with a deep but broad median furrow, coarsely papillose by a few scattered papillae above, 450 pm high; endostome segments lanceolate, keeled above basal membrane, as long as teeth, papillose; cilia single, rudimentary, granulate. Operculum ros- trate, 1.0-1. 2 mm long. Calyptra mitrate, 1.5-1. 8 mm long, rough. Spores rounded, 8-12 pm, granu- late, green. Fig. 168: 1-13.
This species is found in moist forests of east- ern, western and southern Africa and the East and West African islands. The plants are rather
Fig. 168. — Cyclodictyon vallis-gratiae (1-13): 1. habit (dry), x 1: 2. distal part of stem (wet), x 5; 3. part of stem in cross section, x 175; 4. leaf, x 32; 5. part of leaf in cross section, x 175; 6. leaf base (left side), x 87; 7. leaf apex, x 87; 8. perichaetial leaf, x 35; 9. distal part of sporophyte, x 5; 10. part of capsule mouth with peristome (papillae partly shown), x 122; 11. operculum, x 25; 12. calyptra, x 25; 13. spore, x 700. Hookeriopsis pappeana (14-25): 14. habit (dry), x 1; 15. distal part of stem with sporophyte (wet), x 3; 16. part of stem in cross section, x 175; 17. leaf, x 35; 18. part of leaf in cross section, x 175; 19. basal leaf cells (right side), x 175; 20. cells at leaf apex, x 175; 21. perichaetial leaf, x 35; 22. part of capsule mouth with peristome, x 122; 23. operculum, x 25; 24. calyptra, x 25; 25. spore, x 700. (1, 2, 4, 5 & 9, Van Rooy 1I94\ 3, 6 & 7, Mogg 12558 ; 8 & 10-13, Wager 15; 14, Van Rooy 1845 ; 15 & 21-25, Van Rooy 873; 16-18 & 20, Crosby 7899; 19, Sim 7232.)
608
Hookeriaceae
rare in the Flora area but have been found on decaying wood or on rock in kloof forests of the northern, eastern, central and southern Trans- vaal areas, Zululand, KwaZulu-Natal and the eastern, southern and southwestern Cape regions. Map 236.
Vouchers: Arnell, 1973; Crosby & Crosby 7558, 7521; Wager 15.
There are 11 species of Cyclodictyon in eastern Africa, most of which are closely
related to C. vallis-gratiae. The southern African species is known by its broad, bor- dered leaves that have very large hexagonal leaf cells and a double costa reaching well above midleaf.
Sim’s (1926) reference to C. laete-virens Mitt, in southern Africa was in error. The two speci- mens cited by Sim were both Hookeriopsis pap- peana', one of them was mixed with C. vallis- gratiae.
4. HOOKERIOPSIS
Hookeriopsis (Besch.) Jaeg. in Ber. Thatigk. St. Gallischen Naturwiss. Ges. 1875-1876: 358 (1877); Sim, Bryo. S. Afr. 444 (1926); Broth, in Natiirl. PflFam., edn 2, 11: 240 (1925); Gangulee, Moss. E. India 7: 1512 (1977). Type species: not designated.
Plants medium-sized to large, forming loose mats, dark green to yellow-green; saxicolous, cor- ticolous or terricolous. Stems creeping; central strand absent. Leaves complanate, somewhat asym- metrical, elliptical; short acuminate to acute; margins plane, dentate above by enlarged marginal cells. Costa double, extending to near midleaf or just above. Laminal cells rhomboidal to elongate- rhomboidal, weakly thickened and pitted; alar cells not differentiated.
Autoicous. Seta elongate, smooth or rough above. Capsule inclined to horizontal or nodding. Peristome double, incomplete, cilia absent. Operculum rostrate. Calyptra mitrate. Spores small.
A genus of over 120 species found in Central and South America, Africa, India and Asia. Ten species have been reported from Africa. The single species reported from the Flora area is also found in east Africa.
Hookeriopsis pappeana (Hampe) Jaeg. in Ber. Thatigk. St. Gallischen Naturwiss. Ges. 1875-1876: 360 (1877); Sim, Bryo. S. Afr. 444 (1926); Broth, in Natiirl. PflFam., edn 2, 11: 243 (1925). Type: South Africa, Cape Prov., Zwellendam, Pappe s.n. (BM, iso.).
Hookeria pappeana Hampe, Icon. Muse. 2 (1844).
Plants medium-sized, forming loose mats, dark green to yellow green, occasionally tinged with dark red; terricolous, saxicolous or corti- colous. Stems creeping, 10-30 mm long, branching sparse, irregular; in section round, central strand absent, inner cortical cells thin- walled, hyaline, in 3 or 4 rows, outer cortical cells smaller, weakly thick-walled, yellow- brown, in 2 or 3 rows, epidermis fragile, of enlarged thin-walled cells. Leaves crowded, widespreading wet, spreading and crisped dry.
± falcate and somewhat asymmetrical; oblong to elliptical, 1.5-2. 5 mm long; apex shortly acuminate; rounded to base; margins plane, entire below, dentate by enlarged, frequently double-pointed marginal cells; not bordered. Costa double, extending to midleaf, smooth; in section a small group of smaller, weakly thick- ened cells, bulging dorsally. Upper laminal cells rhomboidal to subfusiform, homogeneous, (37)50-75 pm long, 12-18 pm wide, walls thickened, weakly pitted, smooth; basal cells not strongly differentiated, oblong to rhom- boidal, 50-75 pm long, 12-18 pm wide, walls thickened, weakly pitted, smooth; alar cells not differentiated.
Autoicous. Perigonia on stem, gemmate; perigonial leaves ovate-cuspidate, to 1 mm long. Perichaetia along stem, not strongly dif- ferentiated; perichaetial leaves lanceolate to
Hookeriaceae
609
ovate-acuminate, to 2 mm long, leaf cells linear, somewhat sigmoid. Seta 10-15 mm long, dark red-brown, smooth. Capsule exserted, inclined to drooping, short-cylindrical, 1.0-1. 5 mm long, smooth, dark red-brown; urn cylindrical; neck shorter than urn; exothecial cells shortly rectangular, walls weakly collenchymatous, cells at mouth quadrate to transversely rectan- gular, neck cells quadrate; stomata on neck, phaneropore. Peristome double, yellow-brown; exostome teeth lanceolate, erect with inflexed tips dry, erect appressed wet, striate below with deep furrow, papillose above, 500 pm high; endostome segments lanceolate and keeled above short basal membrane, as long as teeth, granulate, cilia absent. Operculum rostrate, 0.5 mm long. Calyptra mitrate, with a few hairs, especially when young, to 2 mm long. Spores rounded, 12-14 pm, weakly granulate, green- ish. Fig. 168: 14-25.
Hookeriopsis pappeana is known from Tan- zania, Uganda, Kenya, Zaire, Zimbabwe and South Africa and has also been reported from China. In the Flora area the species has been collected on soil, rocks and bark at the base of trees, in forests of the northern and eastern Transvaal areas, Zululand, KwaZulu-Natal and the eastern, central, southern and southwestern Cape regions. Map 237.
Vouchers: Crosby & Crosby 8063; Magill 5148, 6203; Smook 1358; Van Rooy 2234.
Hookeriopsis pappeana is identified by its
Map 237. — Hookeriopsis pappeana
unbordered leaves, double costa and dentation of the upper leaf margins formed by enlarged cells. The species closely resembles Lepidopili- dium hanningtonii ; the two can be separated only under the microscope. The most pro- nounced vegetative difference is seen in the leaf margins of the two species. The upper leaf mar- gin of H. pappeana is dentate by enlarged mar- ginal cells that project out from the leaf margin. These cells are frequently large enough to be visible under a dissecting scope, although this is not always so. In addition the blunt tips of these cells generally have two papilla-like points. In contrast, L. hanningtonii is serrate by narrow, sharp-pointed cells. See p. 612 for further dif- ferences.
5. LEPIDOPILIDIUM
Lepidopilidium (C. Miill.) Broth, in Hedwigia 39: 273 (1900); Broth, in Natiirl. PflFam., edn 2, 11: 243 (1925). Type species: not designated.
Plants medium-sized, forming loose mats, green; corticolous. Stems creeping; central strand absent. Leaves flattened, lateral leaves widespreading, weakly falcate; oblong-acuminate; margins serrate, unbordered. Costa double, ending near midleaf. Laminal cells rhomboidal, thin-walled, smooth, extending to base of leaf; alar cells not differentiated. Gemmae produced in tufts along stem, cylindrical.
Seta short, papillose. Capsule erect. Peristome double, incomplete; cilia absent. Operculum ros- trate. Calyptra mitrate. Spores large for the family.
18
Hookeriaceae
611
The 29 species of Lepidopilidium are found in tropical America (14), Africa (5), the East African islands (9) and Asia (1). The genus is related to Hookeriopsis, but is distinct in its scabrous seta and erect capsule.
Lepidopilidium hanningtonii (Mitt.) Broth, in Natiirl. PflFam. 1,3: 944 (1907); Broth, in Natiirl. PflFam., edn 2, 11: 244 (1925). Type: Tanzania, Usagara Mts, Hannington s.n., Oct. 1883 (BM, holo.).
Lepidopilum hanningtonii Mitt, in J. Linn. Soc., Bot. 22: 309 (1886).
Plants medium-sized to large, forming mats, dark green to yellow-green or sometimes tinted red-brown; corticolous. Stems creeping, 20-40 mm long, branching irregular; in section round, central strand absent, inner cortical cells thin- walled, hyaline, in 4 rows, outer cortical cells smaller, thick-walled, yellow, in 3 rows, epider- mis of enlarged thin-walled cells fragile. Leaves somewhat distant but evenly spaced, wide- spreading wet, spreading and narrowed dry, somewhat falcate; oblong to elliptical, 2. 0-3. 2 mm long; acuminate; narrowed to insertion; margins plane, serrate, not bordered. Costa short and double or double and extending to midleaf, smooth; in section a small group of cells bulging dorsally. Upper laminal cells rhomboidal, homogeneous, 66-112 pm long, 15-22 pm wide, walls thin to weakly thickened, smooth; basal cells quadrate to short rectangu- lar at attachment, 15-20 pm long, hyaline, walls thin, smooth; alar cells not differentiated. Gemmae on stem, in small tufts, cylindrical, to 250 pm, green, smooth.
Autoicous. Perichaetia not obvious; peri- chaetial leaves lanceolate-acuminate, 1. 5-2.2 mm long, leaf cells fusiform. Seta 2-6 mm
Map 238. — • Lepidopilidium hanningtonii
♦ Distichophyllum mniifolium var. taylorii
long, brown, papillose. Capsule exserted, ± erect, ellipsoid, 1 mm long, smooth, yellow- green, mouth red; urn ellipsoidal; neck shorter than urn; exothecial cells quadrate to rectangu- lar, walls weakly collenchymatous, cells at mouth quadrate to transversely rectangular, neck cells quadrate; stomata on neck, phanero- pore. Peristome double, red-brown; exostome teeth lanceolate, inflexed dry, erect appressed wet, striate with broad median furrow below, essentially smooth above, 500 pm high; endo- stome segments lanceolate and keeled above low basal membrane, as long as teeth, granu- late; cilia absent. Operculum rostrate, 0.5 mm long. Calyptra mitrate, to 2 mm long, smooth. Spores rounded, 22-25 pm, granulate, light brown. Fig. 169: 1-13.
Fig. 169. — Lepidopilidium hanningtonii (1-13): 1. habit (dry), x 1; 2. habit (wet), x 5; 3. part of stem in cross sec- tion, x 175; 4. leaf, x 32; 5. part of leaf in cross section, x 175; 6. basal leaf cells (right side), x 175; 7. cells at leaf apex, x 175; 8. perichaetial leaf, x 35; 9. seta in cross section (cells partly shown), x 175; 10. part of capsule mouth showing cells and peristome, x 175; 11. operculum, x 25; 12. calyptra, x 25; 13. spore, x 700. Distichophyllum mniifolium var. mnii- folium (14—25): 14. habit (dry), x 1; 15. habit (wet), x 5; 16. stem in cross section (cells partly shown), x 175; 17. leaf, x 35; 18. part of leaf in cross section, x 175; 19. basal leaf cells (right side), x 175; 20. upper laminal cells at right margin, x 350; 21. leaf apex, x 175; 22. perichaetial leaf, x 35; 23. part of capsule mouth showing cells and peristome, x 175; 24. calyptra, x 58; 25. spores, x 700. (1 & 2, Van Rooy 2223; 3, 8, 9 & 11-13, Magill 5745; 4, Crosby 7786; 5-7 & 10, Kemp 1499 ; 14-19, 21 & 23, Duthie PRE-CH8220 ; 20, Russell 2534 ; 22 & 24, Jacot Guitlannod PRE-CH12569.)
612
Hookeriaceae
Endemic to Africa, L. hanningtonii is found in forests of eastern and southern Africa. In southern Africa the species is found on stems or trunks of trees and shrubs in natural forests in Swaziland, Zululand, KwaZulu-Natal and the eastern Cape region. Map 238.
Vouchers: Crosby & Crosby 7662; Magill 5109; Van Rooy 2223.
Lepidopilidium hanningtonii is recognized by its elongate and gently curved leaves, unbor- dered, serrate leaf margins and double costa that ends below midleaf. The shorter papillose seta
and erect capsule separate fertile specimens from Hookeriopsis pappeana. Sterile specimens are generally more difficult to separate. The leaves of L. hanningtonii are more sharply ser- rate and as a rule longer and narrower. They are also more curved (weakly falcate), especially when dry. As the leaves dry the cells collapse inward, narrowing the width of the leaf. This also gives the leaves a less crowded appearance on the stem. Each of these conditions also occurs in H. pappeana, but rarely to the same degree. Stems of H. pappeana appear fuller and leaves not as narrow, when the two species are compared side by side (see p. 609).
6 DISTICHOPHYLLUM
Distichophyllum Doz. & Molk., Muse, frond, ined. archip. ind. 4: 99 (1846). Type species: not des- ignated.
Plants medium-sized, green; corticolous. Stems erect to suberect; central strand absent. Leaves inserted around stem, bordered; Ungulate to spathulate; apiculate; margins plane, entire to serrulate. Costa single, ending below apex. Laminal cells large, thickened, smooth; border cells elongate, incrassate; basal cells large, rectangular; alar cells not differentiated.
Seta lateral. Capsule inclined. Peristome double, incomplete; cilia absent. Operculum rostrate. Calyptra mitrate. Spores small.
Most of the 118 species of Distichophyllum are found around the southern Pacific basin or India; only three species are known from Africa. Many of the other species have complanate, dimorphous leaves and a much shorter costa than the African species.
Distichophyllum mniifolium (Hornsch.) Sim, Bryo. S. Afr. 441 (1926). Type: South Africa, Cape, nr. Koratra, Dr'ege s.n., 5 Oct. 1831 (BM, iso.).
Hookeria mniifolia Hornsch. in Linnaea 15: 141 (1841). Mniadelphus homschuchii C. Mull., Syn. muse, frond. 2: 22 (1850), nom. illeg. inch spec, prior. Leskeodon mniifolius (Hornsch.) Biz. in Biz. & Poes in Acta Acad. Paedagog. Agriensis, n.s. 12: 436 (1974).
Mniadelphus homschuchii C. Mull, in Hedwigia 38: 130 (1899). Type: South Africa, Cape, Genadendal, Breutel s.n. (BM, iso.).
Plants medium-sized, forming turfs, green to yellow-green; corticolous. Stems ± erect, 5-12 mm long, branching sparse, irregular; in
section round, central strand absent, inner cor- tical cells large, thin-walled, 2 or 3 cells across, outer cortical cells smaller, thin-walled, red- brown. Leaves evenly spaced, larger above, spreading wet, appressed dry, somewhat crisped; Ungulate to spathulate, 1.2-2. 2 mm long; apiculate; not narrowed at base; margins plane, ± entire or weakly serrulate at apex; bor- dered by elongated cells. Costa single, ending below apex, smooth; ventral and dorsal sur- faces smooth, cells elongate; in section bulging dorsally, of 6-8 smaller, incrassate cells. Upper laminal cells rounded-quadrate to hexagonal, ± heterogeneous, (6— )8— 1 2(— 15) pm long, walls thickened, smooth; border cells thickened and
Hookeriaceae
613
pitted; basal cells strongly differentiated and forming distinct group, rectangular to oblong- hexagonal, (12-)20-55 pm long, 12-18 pm wide, hyaline, walls smooth; alar cells not dif- ferentiated.
Autoicous. Perichaetia not obvious; peri- chaetial leaves oblong-acuminate, 0.5-0. 8 mm long, leaf cells ± oblong. Seta 5-6 mm long, reddish, smooth. Capsule exserted, inclined, short-ellipsoidal, 1 .0—1.5 mm long, smooth, brown; neck shorter than urn; exothecial cells rounded to hexagonal, walls collenchymatous, cells at mouth transversely rectangular; neck cells rectangular; stomata on neck, phanero- pore, guard cells 2. Peristome double, yellow- ish; exostome teeth lanceolate, reflexed with inflexed tip dry, erect appressed wet, striate with deep median furrow below, papillose above, 200 pm high; endostome segments lanceolate and keeled above basal membrane, as long as teeth, papillose; cilia absent. Operculum rostrate. Calyptra mitrate, 1 mm long, fringed below with unicellular hairs. Spores rounded, 7-10 pm, smooth to granulate, brownish.
On a single sheet in the Natural History Museum (BM) there are four specimens filed under Mniadelphus hornschuchii. The top and bottom specimens from Genadendal are prob- ably isotypes of M. hornschuchii C. Mull, [horn, illeg.; Hedwigia 38: 130 (1899)]. The two specimens in the centre are probably iso- types of Hookeria mniifolia Hornsch. [Linnaea 15: 141 (1841)]. These latter specimens were incorrectly transferred to M. hornschuchii [nom. illeg.; Syn. muse, frond 2: 22 (1850)] by Muller.
Hookeria mniifolia was transferred to Distichophyllum by Sim (1926) and later to Leskeodon Broth, by Bizot & Poes (1974). The southern African specimens have hookeria- ceous peristomes like Distichophyllum, not the daltoniaceous peristome of Leskeodon, and therefore Sim’s placement is retained.
Map 239. — Distichophyllum mniifolium var. mniifolium
Endemic to Africa, D. mniifolium is rarely collected on wood in forests of the southern and southwestern Cape regions. It has also been reported from Tanzania by Bizot & Poes (1974). Two varieties are recognized:
Leaf cells small, irregular, 6-10 pm ....
var. mniifolium
Leaf cells larger, uniform, 10-16 pm . . .
var. taylorii
Distichophyllum mniifolium (Hornsch.) Sim var. mniifolium.
See species description and key. Fig. 169: 14-25.
Found on wood in forests of the southern and southwestern Cape regions, and reported from Tanzania. Map 239.
Vouchers: Duthie (PRE-CH8220); Jacot Guillarmod PRE-CH12569; Russell 2534.
The suberect plants, large leaf cells, single costa and strong leaf borders should place specimens of var. mniifolium. The strongly dif- ferentiated basal leaf cells are reminiscent of the cancellinae of Calymperaceae. The small num- ber of specimens examined seems to indicate that the variety is quite variable.
614
Hookeriaceae
Fig. 170. — Distichophyllum mniifolium var. taylorii: 1. habit (dry), x 1 ; 2. habit (wet), x 5; 3. leaf, x 32; 4. basal leaf cells (left side), x 175; 5. upper laminal cells at right margin, x 350; 6. cells at leaf apex, x \15XTaylor sub Sim 10281.)
Distichophyllum mniifolium (Hornsch.) Sim var. taylorii (Sim) Magill, stat. nov. Type: South Africa, Cape Prov., Wilderness, George, Taylor s.n. (sub Sim 10281, PRE, holo.!).
Distichophyllum taylorii Sim, Bryo. S. Afr. 441 (1926).
See species description and key. Fig. 170.
Endemic to the southern Cape area. Map 238.
Voucher: type only.
The specimen differs from the typical variety by the characters used in the key (cf. Sim 1926: 441), although this cell size dif- ference is magnified by the more regular cell shape of var. taylorii. The leaves of the type are short for the species, the border is some- what stronger and the upper margins are practically entire. Since each of these latter character states is also found in specimens of var. mniifolium, additional specimens of var. taylorii are needed to assess its status prop- erly.
7. H YPOPTERY GIUM
Hypopterygium Brid., Bryol. univ. 2: 709 (1827); Sim, Bryo. S. Afr. 445 (1926); Broth, in Natiirl. PflFam., edn 2, 11: 273 (1925). Type species: not designated.
Plants small to large, scattered, light green; terricolous, saxicolous or corticolous. Primary stems long-creeping; secondary stems erect, heterophyllous, flabellate above naked stipe. Leaves dimor- phic, in three ranks; lateral ranks larger, spreading; amphigastria small, rounded and apiculate; mar- gins generally bordered, plane, entire to dentate. Costa variable, single or forked, short or extend- ing to midleaf or above. Laminal cells rhomboidal; border cells elongate.
Autoicous. Seta elongate, smooth. Capsule horizontal. Peristome double, incomplete; cilia pre- sent. Operculum rostrate. Calyptra cucullate. Spores small.
A genus of 59 species found in tropical America, Africa and Asia. Ten species are known from Africa. H. laricinum and H. viridissimum C. Mull., the two most widely distributed species, are known from western and eastern Africa and Madagascar. The genus is sometimes placed with the other dimorphic-leaved genus, i.e. Lopidium, in a separate family, Hypopterygiaceae.
Hookeriaceae
615
Hypopterygium laricinum (Hook.) Brid., Bryol. univ. 2: 714 (1827); Sim, Bryo. S. Afr. 446 (1926); Broth, in Natiirl. PflFam., edn 2, 11: 275 (1925). Syntypes: Prom. bon. spei, D. Menzies 75 (BM); in jugis andian regione tem- perata, Humboldt & Bonpland s.n.
Hypnum laricinum Hook., Musci exot. 1: 35 (1818).
Plants small to large, scattered, gregarious, green to grey-green; terricolous, saxicolous or corticolous. Primary stems long-creeping, fre- quently tomentose; secondary stem erect, 20-40 mm long; branching regular, flabellate above stipe; in section round, central strand large, inner cortical cells thin-walled, large, hyaline, in 5 or 6 rows, outer cortical cells smaller, weakly thick-walled, in 2 or 3 rows, epidermis of enlarged thin-walled cells, hyaline. Leaves dimorphic, flattened, widespreading wet, spreading and weakly crisped dry; stem leaves asymmetrical, broadly ovate in lateral ranks, 1.8-2. 2 mm long; obtuse and apiculate or cuspi- date; rounded at base; margins plane, entire, unistratose, bordered by elongated cells, hya- line; amphigastria orbicular or broader than long, 1.0-1. 2 mm long; branch leaves asym- metrical, broadly ovate, 1.2-1. 5 mm long; obtuse and apiculate Or cuspidate; rounded at base; margins plane, serrate, unistratose, bor- dered by elongated cells, hyaline; branch amphigastria orbicular, to 1 mm long, margins entire. Costa variable, single, short or extending to midleaf, sometimes extending to apex, fre- quently forked, displaced proximally in lateral leaves; ventral and dorsal surfaces smooth, cells elongate; in section flattened, 3 rows of small, unthickened cells. Upper laminal cells rhom- boidal, homogeneous, 45-75 pm long, 20-30 pm wide, walls thin, weakly pitted, smooth; basal cells not differentiated or some rectangu- lar cells at attachment, walls thin, weakly pitted, smooth; alar cells not differentiated.
Autoicous. Perigonia on stem, gemmate; perigonial leaves ovate-acuminate, 1.5 mm long. Perichaetia obvious, green; perichaetial leaves ovate to oblong, 1. 5-2.0 mm long; acuminate; leaf cells fusiform, pitted. Seta 8-10 mm long, brown, smooth. Capsule exserted,
horizontal to nodding, short-cylindrical, 1.5 mm long, smooth, brown; neck shorter than urn; exothecial cells quadrate to hexagonal, walls thin, cells at mouth quadrate, neck cells qua- drate, annulus present; stomata raised, phanero- pore. Peristome double, yellow; exostome teeth fragile, oblong, weakly striate, with median zigzag line, 350 pm high (old and broken); endostome segments linear above low basal membrane, shorter than teeth, smooth; cilia 1-3, shorter than segments. Operculum long- rostrate, 1 mm long. Calyptra cucullate, smooth. Spores rounded, 10-15 pm, granulate, yellowish. Fig. 171: 1-13.
Hypopterygium laricinum is found in Central and South America, eastern and southern Africa, Gabon and the East and West African islands. In the Flora area, it is collected on soil, rocks or at the base of trees in forests of the northern and eastern Transvaal regions, Zulu- land, KwaZulu-Natal and the eastern, southern and southwestern Cape areas. Map 240.
Vouchers: Glen 3324; Magill 5105, 5451, 6212; Schelpe 7890; Van Rooy 2103.
This species is recognized by its erect, flabel- late secondary stems and dimorphic leaves. It could be confused with Lopidium penniforme which also has erect secondary stems, dimorphic leaves and grows in similar habitats; however, the amphigastria of L. penniforme are lanceolate
616
Hooke riaceae
Hookeriaceae
617
to ovate-acuminate while those of H. laricinum are orbicular. In addition, the secondary stems of L. penniforme are pinnately branched.
A few plants have short (0. 2-1.0 mm), brown ‘rhizoids’ in small tufts on the stems and branch-
es. They are longer than the gemmae found on Lopidium, have somewhat longer cells, are weakly granulate and occasionally have short lateral buds. Although each of these characters points toward rhizoids rather than gemmae, their position on an erect stem seems unusual.
8. LOPIDIUM
Lopidium Hook.f. & Wilson, FI. nov.-zel. 2: 119 (1855); Broth, in Natiirl. PflFam., edn 2, 1 1: 271 (1925). Lectotype species: L. concinnum (Hook.) Wilson, fide Dixon (1927).
Plants medium-sized to large, scattered, light green; saxicolous or corticolous. Primary stems long-creeping, tomentose; secondary stems erect, heterophyllous, pinnately branched above naked stipe; central strand small or absent. Leaves dimorphic, in three ranks; lateral ranks larger, widely spreading; amphigastria smaller and narrower; margins plane, serrulate, bordered by elongated cells. Costa short-excurrent. Laminal cells rhomboidal.
Seta short, rough. Capsule small, ± erect; annulus present. Peristome double, incomplete; cilia absent. Operculum rostrate. Spores small.
Lopidium contains ±19 species found in the southern hemisphere. Four species are known from Africa or the East African islands, only one of which extends into the Flora area. The genus has been placed by some authors in the family Hypopterygiaceae.
Lopidium penniforme (Brid.) Fleisch., Muse. Buitenzorg 3: 1073 (1908); Broth, in Natiirl. PflFam., edn 2, 11: 271 (1925). Type: Prom. bon. spei, Thunberg s.n.
Hypnum penniforme ( ‘pennae forme') Thunb. ex Brid., Muscol. recent, suppl. 2: 96 (1812). Hypopterygium penni- forme Ppennaeforme') (Brid.) Brid., Bryol. univ. 2: 717 (1827); Sim, Bryo. S. Afr. 446 (1926).
Hypopterygium polythrix Dix. in Kongel. Norske Vidensk. Selsk. Skr. (Trondheim) 1932, 4: 15 (1932). Type: South Africa, Natal, Zululand, Eshowe, in indigenous for- est, 22 Aug. 1929, Hoeg 127 (BM, lecto.l, selected here).
Plants medium-sized to large, scattered and gregarious, glaucous green; saxicolous or cor- ticolous. Primary stems long-creeping; sec-
ondary stem erect, 10-35 mm long, branching regular, pinnate above stipe; in section round, central strand absent but with central cavity, inner cortical cells thin-walled, hyaline, in 8-12 rows, outer cortical cells larger, thick- walled, brown, in 3-6 rows, epidermis absent. Leaves in ranks, dimorphic, widespreading wet, contorted dry, asymmetrical; stem leaves ovate or panduriform, 1.8-2. 2 mm long; obtuse and cuspidate; rounded at base; margins plane, serrulate throughout but stronger above, unistratose, bordered by elongated hyaline cells; amphigastria broadly ovate, abruptly narrowed, 1.2-1. 6 mm long; branch leaves lanceolate to elliptical, 1.2-1. 5 mm long; acute
Fig. 171. — Hypopterygium laricinum (1-13): 1. habit (dry), x 1; 2. part of secondary stem and branches (wet), x 5; 3. stem in cross section, x 175; 4. leaf, x 35; 5. amphigastrium, x 35; 6. part of leaf in cross section, x 175; 7. basal leaf cells (right side), x 175; 8. cells at leaf apex, x 175; 9. perichaetial leaf, x 35; 10. part of capsule mouth showing cells and peristome, x 70; 11. operculum, x 18; 12. calyptra, x 18; 13. spore, x 700. Lopidium penniforme (14—25): 14. habit (dry), x 1; 15. part of secondary stem and branch (wet), x 10; 16. part of stem in cross section, x 175; 17. secondary stem leaf, x 35; 18. branch leaf, x 35; 19. branch amphigastrium, x 35; 20. secondary stem amphigastrium, x 35; 21. secondary stem leaf in cross section, x 175; 22. basal leaf cells (right side), x 175; 23. branch leaf cells at left margin, x 350; 24. cells at branch leaf apex, x 175; 25. gemmae, x 175. (1, 2 & 12, Lewis PRE-CH12210; 3 & 6, Berry 49; 4,5,1 & 8, Sim 9430; 9, Crosby 8041; 10, 11 & 13, Von Breitenbach 90; 14—25, Magill 6008.)
618
Hookeriaceae
and cuspidate; rounded at base; margins plane, serrulate throughout, somewhat stronger above, unistratose, bordered by elongated hyaline cells; amphigastria lanceolate to ovate-acumi- nate, 0.6- 1.0 mm long. Costa single, shortly excurrent, displaced proximally in lateral leaves; ventral and dorsal surfaces smooth, cells elongate; in section elliptical, cells in 3 rows, smaller, incrassate. Upper laminal cells rounded quadrate to rectangular or transverse- ly rectangular, homogeneous, 10-15 pm long, in amphigastria 5-7 pm long, walls incrassate to somewhat collenchymatous, smooth; basal cells not differentiated, walls thickened and pitted, smooth; alar cells not differentiated. Gemmae on stem and branches, in small groups, cylindrical, 200-400 pm, brown, papillose.
Sporophyte not seen; see Sim (1926: 447). Fig. 171; 14-25.
Known from Tanzania, Zimbabwe and South Africa, L. penniforme is found on the base of trees, roots and moist rocks in forests of the southern, southwestern and eastern Cape regions, KwaZulu-Natal and Zululand. Map 241.
Vouchers: Magill 6021; Russell 2684; Stirton 9663; Van Rooy 2125.
Lopidium penniforme can be identified by its erect, pinnate secondary stems with dimor- phic leaves and narrow amphigastria. Although it is similar to Hypopterygium laricinum , the two species are easily separated by the shape of their amphigastria and branching patterns (see p. 615).
619
INDEX TO FASCICLE 3*
Aerobryopsis Fleisch
capensis (C. Mtill.) Fleisch
Aerobryum capense (C. Mull.) C. Mull. .
var. rupestre C. Mull
Amphidium Schimp
cyathicarpum (Mont.) Broth
lapponicum (Hedw.) Schimp
tortuosum (Homsch.) Robinson
Amphoridium cyathicarpum (Mont.) Jaeg.
lapponicum (Hedw.) Schimp
Anictangium ciliatum Hedw
lapponicum Hedw
secundum (Hook.) Hook
Anomodon gracilis (Hedw.) Garov
Antitrichia brasiliensis Homsch
Aulacopilum Wilson
beccarii (C. Mull.) Mitt
glaucum Wilson
incanum Mitt
trichophyllum Angstr.
Brachypodium crispatum (Hedw.) Brid.
Brachysteleum Reichenb
convolutifolium Shaw
crassinervium C. Mull
crispatum (Hedw.) Homsch
cucullatifolium C. Mull
depressum C. Mull
obtusatum C. Mull
Braunia B.S.G
diaphana (C. Mull.) Jaeg
elliottii Broth
erosa C. Mull
macowaniana C. Mtill
maritima C. Mtill
peristomata Dix
sciuroides (Bals. & De Not.) B.S.G. . .
secunda (Hook.) B.S.G
var. pinnata Sim
Bryum crispatum Dicks
Callicostella (C. Mull.) Mitt
applanata Broth. & Bryhn
pallida (Homsch.) Angstr.
papillata (Mont.) Mitt
tristis (C. Mull.) Broth
Calyptothecium Mitt
acutifolium (Brid.) Broth
africanum Broth
bescherellei (Ren.) Broth
hoehnelii (C. Mtill.) Argent
planifrons (Ren. <& Card.) Argent ....
pterobryoides Argent
subacutifolium Broth
Calyptrochaeta Desv
asplenioides (Brid.) Crosby
cristata (Hedw.) Desv.
... 579 ... 581 ... 581 ... 581 ... 486 ... 487 486, 489 ... 487 ... 487 ... 489 ... 544 ... 489 ... 547 ... 556 ... 578 ... 456 ... 453 456, 458 ... 456 ... 456 ... 474 ... 465 ... 475 ... 473 ... 465 ... 472 ... 469 ... 472 ... 545 ... 547 ... 555 ... 548 ... 548 ... 548 ... 555 ... 545 ... 547 ... 547 ... 529 ... 604 ... 604 ... 605 ... 604 ... 604 ... 567 ... 569 ... 567 ... 567 ... 567 ... 567 ... 567 ... 569 ... 601 ... 603 ... 601
* Synonyms are in italics.
Cardotiella Vitt 524
schlotheimiaeformis ( Par.)Vitt 525
secunda (C. Midi.) Vitt 525
subappendiculata (Broth.) Vitt 524
Coleochaetium capense (Broth.) Broth 510
secundum (C. Mtill.) Broth 525
Cryphaea Mohr 550
dentata Mitt 550
exigua (C. Mull.) Jaeg 550
CRYPHAEACEAE 550
Cyclodictyon Mitt 605
breutelianum (Hampe) O. Kuntze 607
laete-virens Mitt 608
pteridioides P. Beauv. 605
vallis-gratiae (Hampe) O. Kuntze 607
fo. breuteliana Demar. & P. Varde 607
Dicranaceae 487
Distichophyllum Doz. & Molk 612
mniifolium (Homsch.) Sim 612
var. mniifolium 613
var. taylorii (Sim) Magill 614
taylorii Sim 614
Encalypta crispata Hedw 474
Eriopus Brid 601
asplenioides (Brid.) Besch 603
cristata (Hedw.) Brid 601
mniaceus (C. Mull.) Broth 603
ERPOD1ACEAE 451
Erpodium (Brid.) C. Mtill 451
beccarii C. Mtill 453
coronatum (Hook. & Wilson) Mitt, subsp. trans-
vaaliense (Broth. & Wager) Magill 453
distichum Wager & Dix 455
domingense (Spreng.) C. Mtill 451
grossirete C. Mtill 454
hanningtonii Mitt 453
joannis-meyeri C. Mtill 453
menyharthii C. Mtill 453
transvaaliense Broth. & Wager 454
Fabronia Raddi 500, 501
Floribundaria Fleisch 583
floribunda (Doz. & Molk.) Fleisch 583
FONTINALACEAE 534
Fontinalis Hedw 534
antipyretica Hedw. 534
var. gracilis (Lindb.) Schimp 534
antipyretica sensu Sim 534
dalecarlica B.S.G 537
duthieae Dix 538
gracilis Lindb 534
squamosa Hedw 535
Forsstroemia Lindb 585
dendroides Dix 556
producta (Homsch.) Par 587
trichomitria (Hedw.) Lindb 585
Glyphomitrium crassinervium (C. Mtill.) Broth 473
crispatum (Hedw.) Brid 474
cucullatifolium (C. Mtill.) Broth 472
620
depressum (C. Mull.) Broth
marginatum Wager & Dix
obtusatum (C. Mlill.) Broth
Grimmia omithopodioides Web. & Mohr
Gymnostomum imberbe Sm
Harrisonia Spreng
breuteliana C. Mull
eckloniana C. Mlill
gracillima C. Mlill
rehmanniana C. Mlill
Hedwigia P. Beauv
albicans Lindb
ciliata (Hedw.) P. Beauv.
fo. macowaniana (C. Mlill.) Fleisch
imberbis (Sm.) Sprue
integrifolia P. Beauv
lapponica Hedw.
macowani C. Mlill. ex Dix. & Gepp
macowaniana C. Mlill 544,
secunda Hook
HEDWIGIACEAE
Hedwigidium B.S.G
imberbe (Sm.) B.S.G
integrifolium (P. Beauv.) Dix
maritimum (C. Mlill.) Par
Homalia elongata Welw. & Duby
Hookeria breuteliana Hampe
breutelii Hampe ex Kindb
mniacea C. Mull
mniifolia Homsch
pappeana Hampe
tristis C. Mlill
vallis-gratiae Hampe ex C. Mlill
HOOKER1ACEAE
Hookeriopsis (Besch.) Jaeg
pappeana (Hampe) Jaeg
Hypnodon transvaalensis C. Mlill
Hypnum densum Sw. ex Hedw
hildebrandtii C. Mull
laricinum Hook
penniforme Thunb. ex Brid
purpurascens Brid
smithii Dicks, ex Hedw
Hypopterygiaceae
Hypopterygium Brid
laricinum (Hook.) Brid
penniforme (Brid.) Brid
polythrix Dix
viridissimum C. A lull
Jaegerina C. Miill
stolonifera (C. Miill.) C. Miill
Lasia producta (Hornsch.) Jaeg
Leiomitrium capense Broth
Lepidopilidium (C. Miill.) Broth
hanningtonii (Mitt.) Broth
Lepidopilum hanningtonii Mitt
Leptodon Mohr
smithii (Hedw.) Web. & Mohr
LEPTODONTACEAE
Leskea floribunda Doz. & Molk
Leskeodon Broth
mniifolius (Homsch.) Biz 612
Leucodon Schwaegr 553
assimilis (C. Miill.) Jaeg 553
var. humilis Sim 555
capensis Schimp 555
julaceus (Hedw.) Sull 556
sciuroides (Hedw.) Schwaegr. 553
LEUCODONTACEAE 553
Leucodontopsis Ren. & Card. 585
Lopidium Hook. f. & Wilson 617
concinnum (Hook.) Wilson 617
penniforme (Brid.) Fleisch 617
Macrocoma (Homsch. ex C. Miill.) Grout 506
filiforme (Hook. & Grev.) Grout 506
lycopodioides (Schwaegr.) Vitt 507
pulchella (Homsch.) Vitt 507
tenue (Hook. & Grev.) Vitt subsp. tenue 510
Macromitrium Brid 511
sect. Macrocoma Homsch. ex C. Miill 506
subgen. Macrocoma (Homsch. ex C. Mlill.) Broth. . 506
confusum Mitt 510
dawsoniaemitrium C. Miill 510
dregei Homsch 510
ferrugineum (Bruch ex Hook. & Grev.) C. Miill. . . 521
lebomboense Van Rooy 517
levatum Mitt 512
lycopodioides Schwaegr 509
macropelma C. Miill 513
mannii sensu Sim 512
microphyllum (Hook. & Grev.) Brid 510
pallidum (P. Beauv.) Wijk & Marg 511
pulchellum (Homsch.) Brid 507
richardii Schwaegr 515
schlotheimiaeformis Par 525
secundum C. Miill 525
serpens (Hook. & Grev.) Brid 518
sulcatum ( Hook. ) Brid 513
tenue (Hook. & Grev.) Brid 510
var. dregei (Homsch.) C. Miill 510
var. lycopodioides (Schwaegr.) C. Mlill 509
tristratosum Dix 518
Maschalocarpus ecklonii Spreng 510
METEORIACEAE 575
Meteorium africanum (C. Miill.) Mitt 582
biforme (Hampe) Besch 578
floribundum (Doz. & Molk.) Doz. & Molk 583
rehmannii C. Miill 578
Mniadelphus homschuchii C. Miill 612
Neckera Hedw 590
subsect. Papillaria C. Mlill 582
subsect. Pilotrichella C. Mull 575
acutifolia Brid 569
africana C. Miill 582
assimilis C. Miill 555
capensis C. Miill 581
chrysoneura Hampe ex C. Miill 571
diaphana C. Miill 547
floribunda (Doz. & Molk.) C. Miill 583
gracilis (Hedw.) C. Miill 556
hoehnelii C. Miill 567
macleana Rehm 561
469
466
472
556
548
541
543
543
543
543
544
544
544
545
548
548
489
545
545
547
541
548
548
548
548
600
607
607
603
612
608
604
607
601
608
608
459
558
595
615
617
543
588
614
614
615
617
617
614
565
565
587
510
609
611
611
588
588
585
583
613
621
pandurifolia C. Mtill
pennata Hedw. 590,
producta (Homsch.) C. Mtill
pterops Rehm
serrulata (P. Beauv.) Brid
smithii (Hedw.) C. Mtill
subimbricata Hampe
valentiniana Besch
viridula Mitt
NECKERACEAE
Notarisia Hampe
capensis Hampe
virginica Hampe
Orthostichopsis Broth
chrysoneura (C. Mtill.) Broth
pinnatella (Broth.) Broth
subimbricata (Hampe) Broth
tetragonum (Hedw.) Broth
ORTHOTRICHACEAE
subfamily MACROMITRIOIDEAE
subfamily ORTHOTRICHOIDEAE
subfamily ZY GODONTOIDEAE
ORTHOTRICHUM Hedw
subgenus Orthotrichum
section Diaphanum Vent.
subgenus Phaneroporum Delogne
section Leiocarpa Mol
section Rupestria Schimp
afrofastigiatum C. Mull
afrofastigiatum sensu Sim
anomalum Hedw.
armatum Lewinsky & Van Rooy
crispatum (Hedw.) Hook. & Grev
diaphanum Schrad. ex Brid
ecklonii Homsch
ferrugineum Bruch ex Hook. & Grev
firmum Vent
glaucum Spreng
incurvomarginatum Lewinsky & Van Rooy
macleanum Shaw
microphyllum Hook. & Grev
mirum Lewinsky
oreophilum Lewinsky & Van Rooy
piliferum Sim
pseudotenellum Hamp. ex C. Mtill
pseudotenellum sensu Sim
rupestre Schleich. ex Schwaegr
serpens Bruch ex Hook. & Grev
subexsertum Schimp. ex C. Miill
tenue Hook. & Grev
transvaalense Sim
Papillaria (C. Miill.) C. Miill
africana (C. Miill.) Jaeg
floribunda (Doz. & Molk.) C. Mtill
natalensis Sim
nigrescens (Hedw.) Jaeg
serrulata (P. Beauv.) Jaeg
viridula (Mitt.) Jaeg
PiLOTRiCHELLA (C. Mull.) Besch
biformis (Hampe) C. Miill
conferta Ren. & Card
cuspidata Broth 577
imbricata (P. Beauv.) Besch 575
kuntzei C. Mtill 577
pandurifolia (C. Miill.) Jaeg 575
pinnatella Broth 573
rehmannii C. Mtill 578
subimbricata (Hampe) Jaeg 571
Pilotrichum exiguum C. Mtill 550
serrulatum P. Beauv 561
Pinnatella Fleisch 592
kuehliana (Bosch & Sande Lac.) Fleisch 592
minuta (Mitt.) Broth 592
oblongifrondea (Broth.) Broth 592
Plagiomnium Koponen 603
Porothamnium Fleisch 593
capense (Broth. & Dix.) Sim 596
comorense (C. Miill.) Sim 600
hildebrandtii (C. Miill.) Fleisch 595
molliculum (Broth.) Fleisch 597
natalense (C. Miill.) Fleisch 597
penniforme (C. Miill.) Fleisch 599
stipitatum (Mitt.) Touw ex De Sloover 595
Porotrichum (Brid.) Hampe 596
comorense Hampe ex C. Miill 600
elongatum (Welw. & Dub.) Gepp 600
longirostre (Hook.) Mitt 596
madagassum Kiaer ex Besch 599
molliculum Broth 597
natalense C. Mtill 597
oblongifrondeum Broth 592
penniforme C. Miill 599
pterops Rehm. ex Par 595
usugarum Mitt 597
Prionodon C. Miill 558
densus (Hedw.) C. Miill 558
rehmannii Mitt 558
PRIONODONTACEAE 558
Pseudocryphaea Britt 585
Pterigynandrum gracile Hedw 556
PTEROBRYACEAE 565
Pterobryopsis Fleisch 567
acutifolium (Brid.) Magill 569
crassicaulis (C. Miill.) Fleisch 567
hoehnelii (C. Miill.) Magill 567
rehmannii Magill 570
Pterogonium Sw 556
gracile (Hedw.) Sm 556
var. capense C. Mtill. ex Dix 556
omithopodioides (Web. & Mohr) Lindb 556
productum Homsch 587
Pterygophyllum asplenioides Brid 603
PT Y CHOMITRI ACEAE 462
Ptychomitriopsis Dix 462
africana Dix 462, 463
aloinoides Magill 463
Ptychomitrium Fuemr 465
africanum (Dix.) Churchill 463
crassinervium (C. Miill.) Schimp. ex Par 473
crispatum (Hedw.). Jaeg 474
crispatum var. brachycarpum Homsch 472
cucullatifolium (C. Miill.) Jaeg 472
575
591
587
595
561
588
571
590
563
590
465
474
465
571
571
573
571
571
476
506
490
477
490
491
491
491
491
491
502
502
490
495
474
500
504
521
493
500
494
496
510
502
493
500
500
497
496
518
497
510
500
582
582
583
563
582
561
563
575
578
577
622
depressum (C. Mull.) Par
diexaratum Magill
eurybasis Dix
exaratifolium Robinson
godfreyi Robinson
marginatum (Wager & Dix.) Dix
obtusatum (C. Miill.) Par
polyphyllum (Sw.) B.S.G
standleyi Crum
subcrispatum Ther. & P. Varde
RACOPILACEAE
Racopilum P. Beauv
capense C. Miill.
cuspidigerum (Schwaegr.) Angstr.
mnioides P. Beauv.
tomentosum (Hedw.) Brid.
Rhabdoweisia B.S.G
crispata (Dicks.) Lindb
fugax (Hedw.) B.S.G
RHABDOWEISIACEAE
Rhabdoweisiella Williams
RHACHITHECIACEAE
Rhachithecium Broth, ex Le Jolis
perpusillum (Thwait. & Mitt.) Broth. . . .
transvaalense (C. Miill.) Broth
Rhacocarpus Lindb
breutelianus (C. Miill.) Broth
ecklonianus (C. Miill.) Broth
gracillimus (C. Miill.) Broth
humboldtii (Hook.) Lindb
purpurascens (Brid.) Par
rehmannianus (C. Miill.) Wijk et Marg. .
Schizomitrium B.S.G
applanatum (Broth. & Bryhn) Ochyra . .
triste (C. Miill.) Ochyra
Schlotheimia Brid
exrugulosa C. Miill
ferruginea (Bruch ex Hook. & Grev.) Brid.
grevilleana Mitt.
percuspidata C. Miill
pulchella Homsch
rufoaeruginosa C. Miill.
rufoglauca C. Miill
rufopallens C. Miill
subventrosa Broth. & Bryhn
torquata (Hedw.) Brid.
ventrosa C. Miill
Squamidium (C. Miill.) Broth
biforme (Hampe) Broth
brasiliense (Homsch.) Broth
lorentzii (C. Miill.) Broth 578
rehmannii (C. Miill.) Broth 578
Stoneobryum Norris & Robinson 502
bunyaense Norris & Robinson 502
mirum (Lewinsky) Norris & Robinson 502
Syntrichia Brid. 500,501
chisosa (Magill, Delgad. & L.R. Stark) R.H. Zander 467
Syrrhopodon tortuosus Homsch 487
Thamnium afrum C. Miill 595
capense Broth. & Dix 596
hildebrandtii (C. Miill.) Jaeg 595
molliculum (Broth.) Kindb 597
natalense (C. Miill.) Kindb 597
penniforme Kindb. var. brachyphyllum Dix 597
THAMNOBRYACEAE 592
Trachyloma stipitatum Mitt 595
TRACHYPODACEAE 560
Trachypodopsis Fleisch 560
serrulata (P. Beauv.) Fleisch 560, 561
Trachypus Reinw. & Homsch 561
appendiculatus (Ren. & Card.) Broth 564
bicolor Reinw. & Homsch 561
var. hispidus (C. Miill.) Card. 564
var. viridulus (Mitt.) Zant 563
serrulatus (P. Beauv.) Besch 561
viridulus (Mitt.) Broth 563
Ulota Mohr 504
crispa (Hedw.) Brid. 504
ecklonii (Homsch.) Jaeg 504
Wardia Harv. c£ Hook 538
hygrometrica Harv. & Hook 538
WARDIACEAE 538
Weissia fugax Hedw 527
Zygodon Hook. & Tayl 477
africanus Sim 485
cemuus C. Miill 486
conoideus (Dicks.) Hook. & Tayl 477
cyathicarpus Mont 487
dixonii Sim 478
erosus Mitt 485
intermedius B.S.G 481
lapponicus (Hedw.) B.S.G 489
leptobolax C. Miill 483
perpusillus Thwait. & Mitt 459
perreflexus C. Miill 486
runcinatus C. Miill 479
transvaalensis Rehm. ex Sim 481
trichomitrius Hook. & Wilson 484
var. mildbraedii (Broth.) Malta 485
469
469
471
467
463
466
472
465
467
466
531
531
531
533
531
533
527
529
527
527
527
459
459
459
459
541
543
543
543
541
543
543
604
604
604
519
523
521
524
522
507
524
524
523
521
519
524
578
578
578
A-l
APPENDIX
PLAN OF FLORA OF SOUTHERN AFRICA
Cryptogam volumes will in future not be numbered, but will be known by the name of the group they cover. The num- ber assigned to the volume on Charophyta therefore becomes redundant. Occasional contributions to the Flora are pub- lished in Bothalia under the title FSA contributions.
Exotic families are marked with an asterisk.
Published volumes and parts are shown in bold.
INTRODUCTORY VOLUMES
The genera of southern African flowering plants Vol. 1: Dicotyledons (1975)
Vol. 2: Monocotyledons (1976)
Botanical exploration of southern Africa (1981)
CRYPTOGAM VOLUMES
Charophyta (as Vol. 9 in 1978)
Bryophyta: Part 1: Musci: Fascicle 1: Sphagnaceae-Grimmiaceae (1981)
Fascicle 2: Gigaspermaceae-Bartramiaceae (1987) Fascicle 3: Erpodiaceae-Hookeriaceae (1998) Fascicle 4: Fabroniaceae-Polytrichaceae
Hepatophyta
Anthocerotophyta
Pteridophyta (1986)
FLOWERING PLANTS VOLUMES
Vol. 1 : Stangeriaceae, Zamiaceae, Podocarpaceae, Pinaceae*, Cupressaceae, Welwitschiaceae, Typhaceae, Zoster- aceae, Potamogetonaceae, Ruppiaceae, Zannichelliaceae, Najadaceae, Aponogetonaceae, Juncaginaceae, Alismataceae, Hydrocharitaceae (1966)
Vol. 2 : Poaceae
Vol. 3 : Cyperaceae, Arecaceae, Araceae, Lemnaceae, Flagellariaceae Vol. 4 : Part 1 : Restionaceae
Part 2: Xyridaceae, Eriocaulaceae, Commelinaceae, Pontederiaceae, Juncaceae (1985)
Vol. 5 : Part 1: Colchicaceae, Eriospermaceae, Asphodelaceae (Chortolirion, 1995 in Bothalia 25: 31-33; Poellnitzia, 1995 in Bothalia 25: 35-36)
Part 2: Alliaceae, Liliaceae*, Hyacinthaceae, Agavaceae (1996 in Bothalia 26: 31-35)
Part 3: Dracaenaceae, Asparagaceae, Luzuriagaceae, Smilacaceae ( 1992)
Vol. 6 : Haemodoraceae, Amaryllidaceae, Hypoxidaceae, Tecophilaeaceae, Velloziaceae, Dioscoreaceae Vol. 7 : Iridaceae: Part 1 : Nivenioideae, Iridoideae Part 2: Ixioideae: Fascicle 1
Fascicle 2: Syringodea, Romulea (1983)
Vol. 8 : Musaceae, Strelitziaceae, Zingiberaceae, Cannaceae*, Burmanniaceae, Orchidaceae (Holothrix, 1996 in Bothalia 26: 125-140)
Vol. 9 : Casuarinaceae*, Piperaceae, Salicaceae, Myricaceae, Fagaceae*, Ulmaceae, Moraceae, Cannabaceae*, Urticaceae, Proteaceae
A-2
Vol. 10: Part 1: Loranthaceae, Viscaceae (1979), Santalaceae, Grubbiaceae, Opiliaceae, Olacaceae, Balanophoraceae, Aristo- lochiaceae, Rafflesiaceae, Hydnoraceae, Polygonaceae, Chenopodiaceae, Amaranthaceae, Nyctaginaceae
Vol. 11: Phytolaccaceae, Aizoaceae, Mesembryanthemaceae
Vol. 12: Portulacaceae, Basellaceae, Caryophyllaceae, Illecebraceae, Cabombaceae, Nymphaeaceae, Ceratophyllaceae (1997 in Bothalia 27: 125-128), Ranunculaceae, Menispermaceae, Annonaceae, Trimeniaceae, Lauraceae, Hemandiaceae, Papaveraceae, Fumariaceae
Vol. 13: Brassicaceae, Capparaceae, Resedaceae, Moringaceae, Droseraceae, Roridulaceae, Podostemaceae, Hydro- stachyaceae (1970)
Vol. 14: Crassulaceae (1985)
Vol. 15: Vahliaceae, Montiniaceae, Escalloniaceae, Pittosporaceae, Cunoniaceae, Myrothamnaceae, Bruniaceae, Hama- melidaceae, Rosaceae, Connaraceae
Vol. 16: Fabaceae: Part 1: Mimosoideae (1975)
Part 2: Caesalpinioideae (1977)
Part 3: Papilionoideae: Fascicle 1: Swartzieae-Robinieae Fascicle 2: Indigofereae Fascicle 3: Desmodieae, Phaseoleae Fascicle 4: Psoraleeae-Galegeae Fascicle 5: Loteae-Liparieae Fascicle 6: Crotalarieae (Aspalathus) (1988)
Fascicle 7: Crotalarieae (Bolusia-Lebeckia)
Fascicle 8: Crotalarieae (Lotononis-Wiborgia)
Fascicle 9: Crotalarieae ( Pearsonia-Argyrolobium ), Genisteae (Cytisus-Ulex)
Vol. 17: Geraniaceae, Oxalidaceae
Vol. 18: Part 1: Linaceae, Erythroxylaceae, Zygophyllaceae, Balanitaceae Part 2: Rutaceae
Part 3: Simaroubaceae, Burseraceae, Ptaeroxylaceae, Meliaceae (Aitoniaceae), Malpighiaceae (1986)
Vol. 19: Part 1: Polygalaceae, Dichapetalaceae
Part 2: Euphorbiaceae, Callitrichaceae, Buxaceae (1996 in Bothalia 26: 3 7 — 40 )
Part 3: Anacardiaceae: Fascicle 1: Rhus (1993)
Fascicle 2: remaining genera Aquifoliaceae (1994 in Bothalia 24: 163-166)
Vol. 20: Celastraceae, Icacinaceae, Sapindaceae, Melianthaceae, Greyiaceae, Balsaminaceae, Rhamnaceae, Vitaceae
Vol. 21: Part 1: Tiliaceae (1984)
Malvaceae, Bombacaceae, Sterculiaceae
Vol. 22: Ochnaceae, Clusiaceae, Elatinaceae, Frankeniaceae, Tamaricaceae, Canellaceae, Violaceae, Flacourtiaceae, Turneraceae, Passifloraceae, Achariaceae, Loasaceae, Begoniaceae, Cactaceae (1976)
Vol. 23: Geissolomaceae, Penaeaceae, Oliniaceae, Thymelaeaceae, Lythraceae, Lecythidaceae
Vol. 24: Rhizophoraceae, Combretaceae, Myrtaceae, Melastomataceae, Onagraceae (1997 in Bothalia 27: 149-165), Trap- aceae, Haloragaceae, Gunneraceae, Araliaceae, Apiaceae, Comaceae
Vol. 25: Ericaceae
Vol. 26: Myrsinaceae, Primulaceae, Plumbaginaceae, Sapotaceae, Ebenaceae, Oleaceae, Salvadoraceae, Loganiaceae, Gentianaceae, Apocynaceae (1963)
Vol. 27: Part 1: Periplocaceae, Asclepiadaceae (Microloma-Xysmalobium)
Part 2: Asclepiadaceae ( Schizoglossum-Woodia )
Part 3: Asclepiadaceae ( Asclepias-Anisotoma )
Part 4: Asclepiadaceae (Brachystelma-Riocreuxia) (1980)
Asclepiadaceae (remaining genera)
Vol. 28: Part 1: Cuscutaceae, Convolvulaceae Part 2: Hydrophyllaceae, Boraginaceae
Part 3: Stilbaceae, Verbenaceae ( Vitex, 1996 in Bothalia 26: 141-151)
Part 4: Lamiaceae (1985)
Part 5: Solanaceae, Retziaceae
A- 3
Vol. 29: Scrophulariaceae
Vol. 30: Part 1 : Bignoniaceae, Pedaliaceae, Martyniaceae, Orobanchaceae Part 2: Gesneriaceae, Lentibulariaceae
Part 3: Acanthaceae: Fascicle 1: Justiciinae ( Metarungia , Siphonoglossa, Rhinacanthus, Duvernoia, Justicia, Monechma, Adhatoda, Anisotes) (1995)
Acanthaceae (remaining genera), Myoporaceae Vol. 31: Part 1: Fascicle 1: Plantaginaceae, Rubiaceae (Rubioideae — first part)
Fascicle 2: Rubiaceae (Rubioideae — second part): Paederieae, Anthospermeae, Rubieae (1986) Fascicle 3: Ixoroideae, Chinchonoideae Part 2: Valerianaceae, Dipsacaceae, Cucurbitaceae
Vol. 32: Campanulaceae, Sphenocleaceae, Lobeliaceae, Goodeniaceae Vol. 33: Asteraceae: Part 1: Lactuceae, Mutisieae, ‘Tarchonantheae’
Part 2: Vemonieae, Cardueae
Part 3: Arctotideae
Part 4: Anthemideae
Part 5: Astereae
Part 6: Calenduleae
Part 7: Inuleae: Fascicle 1: Inulinae
Fascicle 2: Gnaphaiiinae (first part) (1983)
Part 8: Heliantheae, Eupatorieae Part 9: Senecioneae
FSA CONTRIBUTIONS IN BOTHALIA
FSA contributions 1: Aquifoliaceae. S. ANDREWS. 1994. Bothalia 24: 163-166.
FSA contributions 2: Asphodelaceae/Aloaceae, 1029010 Chortolirion. G.F. SMITH. 1995. Bothalia 25: 31-33. FSA contributions 3: Asphodelaceae/Aloaceae, 1028010 Poellnitzia. G.F. SMITH. 1995. Bothalia 25: 35, 36. FSA contributions 4: Agavaceae. G.F. SMITH & M. MOSSMER. 1996. Bothalia 26: 31-35.
FSA contributions 5: Buxaceae. H.F. GLEN. 1996. Bothalia 26: 37-40.
FSA contributions 6: Orchidaceae: Holothrix. K.L. IMMELMAN. 1996. Bothalia 26: 125-140.
FSA contributions 7: Verbenaceae: Vitex. C.L. BREDENKAMP & D.J. BOTHA. 1996. Bothalia 26: 141-151. FSA contributions 8: Ceratophyllaceae. C.M. WILMOT-DEAR. 1997. Bothalia 27: 125-128.
FSA contributions 9: Onagraceae. P. GOLDBLATT & P.H. RAVEN. 1997. Bothalia 27: 149-165.
A-4
ALPHABETICAL ARRANGEMENT OF TAXA LISTED AS ‘PUBLISHED’ IN PLAN OF FLORA OF SOUTHERN AFRICA
* exotic families
Acanthaceae: Justiciinae, Vol. 30, Part 3, Fasc.l (1995) Achariaceae, Vol. 22 (1976)
Adhatoda, Acanthaceae: Justiciinae, Vol. 30, Part 3, Fasc. 1 (1995)
Agavaceae (Bothalia 26, 1996)
Alismataceae, Vol. 1 (1966)
Anacardiaceae: Rhus , Vol. 19, Part 3, Fasc. 1 (1993) Anisotes , Acanthaceae: Justiciinae, Vol. 30, Part 3, Fasc. 1 (1995)
Anthospermeae, Rubiaceae: Rubioideae (second part), Vol.
31, Part 1, Fasc. 2 (1986)
Apocynaceae, Vol. 26 (1963)
Aponogetonaceae, Vol. 1 (1966)
Aquifoliaceae ( Bothalia 24, 1994)
Asclepiadaceae: Brachystelma-Riocreuxia , Vol. 27, Part 4 (1980)
Aspalathus , Fabaceae: Papilionoideae, Vol. 16, Part 3, Fasc. 6 (1988)
Asparagaceae, Vol. 5 (1992)
Asphodelaceae: Chortolirion, Poellnitzia ( Bothalia 25, 1995)
Asteraceae: Inuleae: Gnaphaliinae (first part), Vol. 33, Part 7, Fasc. 2 (1983)
Begoniaceae, Vol. 22 (1976)
Brachystelma , Asclepiadaceae, Vol. 27, Part 4 (1980) Brassicaceae, Vol. 13 (1970)
Bryophyta (three fascicles published 1981, 1987, 1998: see p. 445 of this Fascicle)
Burseraceae, Vol. 18 (1986)
Buxaceae ( Bothalia 26, 1996)
Cactaceae, Vol. 22 (1976)
Caesalpinioideae, Fabaceae, Vol. 16, Part 2 (1977) Canellaceae, Vol. 22 (1976)
Capparaceae, Vol. 13 (1970)
Ceratophyllaceae ( Bothalia 27, 1997)
Charophyta, Cryptogams ‘Vol. 9’ (1978)
Chortolirion , Asphodelaceae (Bothalia 25, 1995) Clusiaceae, Vol. 22 (1976)
Commelinaceae, Vol. 4 (1985)
Crassulaceae, Vol. 14 (1985)
Crotalarieae, Aspalathus , Fabaceae: Papilionoideae, Vol.
16, Part 3, Fasc. 6 (1988)
Cupressaceae, Vol. 1 (1966)
Dracaenaceae, Vol. 5 (1992)
Droseraceae, Vol. 13 (1970)
Duvemoia, Acanthaceae: Justiciinae, Vol. 30, Part 3, Fasc. 1 (1995)
Ebenaceae, Vol. 26 (1963)
Elatinaceae, Vol. 22 (1976)
Eriocaulaceae, Vol. 4 (1985)
Fabaceae: Caesalpinioideae, Vol. 16, Part 2 (1977) Fabaceae: Mimosoideae, Vol. 16, Part 1 (1975)
Fabaceae: Papilionoideae, Crotalariae, Aspalathus, Vol. 16, Part 3, Fasc. 6 (1988)
Flacourtiaceae, Vol. 22 (1976)
Frankeniaceae, Vol. 22 (1976)
Gentianaceae, Vol. 26 (1963)
Gnaphaliinae (first part), Asteraceae: Inuleae, Vol. 33, Part 7, Fasc. 2 (1983)
Holothrix , Orchidaceae (Bothalia 26, 1996) Hydrocharitaceae, Vol. 1 ( 1966)
Hydrostachyaceae, Vol. 13 (1970)
Inuleae, Asteraceae: Gnaphaliinae (first part), Vol. 33, Part 7, Fasc. 2 (1983)
Iridaceae: Syringodea, Romulea, Vol. 7, Part 2, Fasc. 2 (1983)
Juncaceae, Vol. 4 (1985)
Juncaginaceae, Vol. 1 (1966)
Justicia , Acanthaceae: Justiciinae, Vol. 30, Part 3, Fasc. 1 (1995)
Justiciinae, Acanthaceae, Vol. 30, Part 3, Fasc. 1 (1995) Lamiaceae, Vol. 28 (1985)
Loasaceae, Vol. 22 (1976)
Loganiaceae, Vol. 26 (1963)
Loranthaceae, Vol. 10 (1979)
Luzuriagaceae, Vol. 5 (1992)
Malpighiaceae, Vol. 18 (1986)
Meliaceae, Vol. 18 (1986)
Metarungia , Acanthaceae: Justiciinae, Vol. 30, Part 3, Fasc. 1 (1995)
Mimosoideae, Fabaceae, Vol. 16, Part 1 (1975)
Monechma, Acanthaceae: Justiciinae, Vol. 30, Part 3, Fasc. 1 (1995)
Moringaceae, Vol. 13 (1970)
Myrsinaceae, Vol. 26 (1963)
Najadaceae, Vol. 1 (1966)
Ochnaceae, Vol. 22 (1976)
Oleaceae, Vol. 26 (1963)
Onagraceae ( Bothalia 27, 1997)
Orchidaceae: Holothrix (Bothalia 26, 1996)
Paederieae, Rubiaceae: Rubioideae (second part), Vol. 31, Part 1, Fasc. 2 (1986)
Passifloraceae, Vol. 22 (1976)
Pinaceae*, Vol. 1 (1966)
Plumbaginaceae, Vol. 26 (1963)
Podocarpaceae, Vol. 1 (1966)
Podostemaceae, Vol. 13 (1970)
Poellnitzia, Asphodelaceae ( Bothalia 25, 1995) Pontederiaceae, Vol. 4 (1985)
Potamogetonaceae, Vol. I (1966)
Primulaceae, Vol. 26 (1963)
Ptaeroxylaceae, Vol. 18 (1986)
Pteridophyta (1986) (for list of families, see p. v of Pteridophyta volume)
A-5
Resedaceae, Vol. 13 (1970)
Rhinacanthus , Acanthaceae: Justiciinae, Vol. 30, Part 3, Fasc. 1 (1995)
Rhus, Anacardiaceae, Vol. 19, Part 3, Fasc. I (1993) Riocreuxia, Asclepiadaceae, Vol. 27, Part 4 (1980) Romulea, Iridaceae, Vol. 7, Part 2, Fasc. 2 (1983) Roridulaceae, Vol. 13 (1970)
Rubiaceae: Rubioideae (second part): Paederieae,
Anthospermeae, Rubieae, Vol. 31, Part 1, Fasc. 2 (1986)
Rubieae, Rubiaceae: Rubioideae (second part), Vol. 31, Part 1, Fasc. 2 (1986)
Rubioideae (second part), Rubiaceae, Vol. 31, Part 1, Fasc. 2(1986)
Ruppiaceae, Vol. 1 (1966)
Salvadoraceae, Vol. 26 (1963)
Sapotaceae, Vol. 26 (1963)
Simaroubaceae, Vol. 18 (1986)
Siphonoglossa, Acanthaceae: Justiciinae, Vol. 30, Part 3, Fasc. 1 (1995)
Smilacaceae, Vol. 5 (1992)
Stangeriaceae, Vol. 1 (1966)
Syringodea, Iridaceae, Vol. 7, Part 2, Fasc. 2 (1983) Tamaricaceae, Vol. 22 (1976)
Tiliaceae, Vol. 21 (1984)
Tumeraceae, Vol. 22 (1976)
Typhaceae, Vol. 1 (1966)
Verbenaceae: Vitex (Bothalia 26, 1996)
Violaceae, Vol. 22 (1976)
Viscaceae, Vol. 10 (1979)
Vitex, Verbenaceae ( Bothalia 26, 1996)
Welwitschiaceae, Vol. 1 (1966)
Xyridaceae, Vol. 4 ( 1985)
Zamiaceae, Vol. 1 (1966)
Zannichelliaceae, Vol. 1 (1966)
Zosteraceae, Vol. 1 (1966)