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STRELITZIA

A series of occasional publications on southern African flora and vegetation, replacing Memoirs of the Botanical

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A palaeoflora on a pattern comparable to that of the Flora of southern Africa. Much of the information is pre-

sented in the form of tables and photographic plates depicting fossil populations. Now available:

Molteno Formation (Triassic) Vol. 1. Introduction. Dicroidium, 1983, by J.M. & H.M. Anderson.

Molteno Formation (Triassic) Vol. 2. Gymnosperms (excluding Dicroidium), 1983, by J.M. & H.M.

Anderson.

Prodromus of South African Megafloras. Devonian to Lower Cretaceous, 1985, by J.M. & H.M. Anderson.

Obtainable from: A. A. Balkema Marketing, Box 317, Claremont 7735, RSA.

Towards Gondwana Alive. Promoting biodiversity and stemming the Sixth Extinction, 1999, by J.M.

Anderson (ed.).

Heyday of the gymnosperms: systematics and biodiversity of the Late Triassic Molteno fructifications,

2003, by J.M. Anderson & H.M. Anderson. Strelitzia 1 5.

Brief history of the gymnosperms: classification, biodiversity, phytogeography and ecology, 2007, by

J.M. Anderson, H.M. Anderson & C.J. Cleal. Strelitzia 20.

Molteno ferns: Late Triassic biodiversity in southern Africa, 2008, by H.M. Anderson & J.M. Anderson.

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A JOURNAL OF BOTANICAL RESEARCH

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Editorial Board

D.F. Cutler

B.J. Huntley

P.H. Raven

M.J.A. Werger

Royal Botanic Gardens, Kew, UK

South African National Biodiversity Institute, Cape Town, RSA

Missouri Botanical Garden, St Louis, USA

University of Utrecht, Utrecht, The Netherlands

Acknowledgements to referees

Allemann, Dr J. University of the Free State, Bloemfontein, RSA.

Archer, Mrs C. South African National Biodiversity Institute, Pretoria, RSA.

Archer, Dr R. South African National Biodiversity Institute, Pretoria, RSA.

Briggs, Dr B. Royal Botanic Gardens, Sydney, Australia.

Brummitt, Dr R.K. Royal Botanic Gardens, Kew, England, UK.

Burrows, J.E. Box 710, 1120 Lydenburg, RSA.

ClaBen-Bockhoff, Prof. R. J. Gutenberg-Universitat Mainz, Mainz, Germany.

Craib, C. Box 67142, Bryanston, 2021 Sandton, RSA.

Crouch, Prof. N.R. South African National Biodiversity Institute, Durban, RSA.

Ellery, Prof. W.N. Rhodes University, Grahamstown, RSA.

Feuillet, Dr C. Smithsonian Institution, Washington DC, USA.

Forster, Dr PE Queensland Herbarium, Brisbane Botanic Gardens, Australia.

Glen, Dr H.F. South African National Biodiversity Institute, Durban, RSA.

Goldblatt, Dr P. Missouri Botanical Garden, St Louis, USA.

Grobler, Mrs A. South African National Biodiversity Institute, Pretoria, RSA.

Hill, DrT.R. University of KwaZulu-Natal, Pietermaritzburg, RSA.

Hind, Dr D.J.N. Royal Botanic Gardens, Kew, England, UK.

Klopper, Mrs R.R. South African National Biodiversity Institute, Pretoria, RSA.

Lammers, Prof. T.G. University of Wisconsin, Oshkosh, USA.

Lavranos, J. Apartado Postal 243, 8100 Louie, Portugal.

Leistner, Dr O.A. 194 Griselda Rd, Murrayfield, 0184 Pretoria, RSA.

Manning, Dr J.C. South African National Biodiversity Institute, Cape Town, RSA.

Mfundisi, Dr K. Harry Oppenheimer Okavango Research Centre, Maun, Botswana.

Retief, Dr E. South African National Biodiversity Institute, Pretoria, RSA.

Roux, Dr J.P. South African National Biodiversity Institute, Cape Town, RSA.

Schrire, Dr B. Royal Botanic Gardens, Kew, England, UK.

Smithies, Mrs S.J. South African National Biodiversity Institute, Pretoria, RSA.

Snijman, Dr D. South African National Biodiversity Institute, Cape Town, RSA.

Steiner, Dr K.E. California Academy of Sciences, San Francisco, USA.

Van Jaarsveld, E.J. South African National Biodiversity Institute, Cape Town, RSA.

Van Wyk, Prof. B-E. University of Johannesburg, RSA.

Vlok, J. Regalis Environmental Services, P.O. Box 1512, 6620 Oudtshoorn, RSA.

Welman, Ms W.G. South African National Biodiversity Institute, Pretoria, RSA.

Werger, Prof. M.J.A. Department Plant Ecology and Evolutionary Biology, Utrecht, The Netherlands.

Yatskievych, Dr G. Missouri Botanical Garden, St Louis, USA.

Yoshida, DrN. Miyazaki University, Miyazaki, Japan.

Date of publication of Bothalia 39,2: 6 October 2009.

CONTENTS

Bothalia 40,1

1. A generic classification of the Restioneae (Restionaceae), southern Africa. H.R LINDER and C.R. HAR-

DY 1

2. New synonyms and a new name in Asteraceae: Senecioneae from the southern African winter rainfall

region. J.C. MANNING and R GOLDBLATT 37

3. New taxa of Babiana (Iridaceae: Crocoideae) from coastal Western Cape, South Africa. R GOLDBLATT

and J.C. MANNING 47

4. Notes on African plants:

Arecaceae. Livistona chinensis, a first record of a naturalized palm in South Africa. S.J. SIEBERT,

A.M. ZOBOLOand J.L. DOWE 55

Asphodelaceae. Occurrence of Haworthia bolusii var. blackbeardiana in the Free State, South Af-

rica. P.C. ZIETSMAN and G.F. SMITH 58

Asphodelaceae. Inclusion of the genus Jodrellia in Bulbine (Asphodeloideae). J.S. BOATWRIGHT

and J.C. MANNING 59

Asphodelaceae: Alooideae. Reinstatement of Aloe spectabilis. R.R. KLOPPER and G.F. SMITH .... 91

Asphodelaceae: Alooideae. Aloe neilcrouchii , a new robust leptaloe from KwaZulu-Natal, South

Africa. R.R. KLOPPER and G.F. SMITH 93

Boraginaceae. Nomenclatural notes on Echium fmticosum var. major and var. minor. M.H. BUYS

and B. NORDENSTAM 90

Bruniaceae. New species of Thamnea and Brunia from Western Cape, South Africa. A.V. HALL,

E.G.H. OLIVER and R. CLAI3EN-BOCKHOFF 96

Fabaceae. Pearsonia mbabanensis , an overlooked synonym of P. sessilifolia subsp. marginata

(tribe Crotalarieae). J.S. BOATWRIGHT 83

Hyacinthaceae. Drimia cooperi in KwaZulu-Natal, and the ethnomedicinal trade. N.R. CROUCH,

V.L. WILLIAMS, T.J. EDWARDS and V.J. BRUETON 75

Iridaceae. Reappraisal of Ixia maculata with I. calendulacea sp. nov., and an earlier name for /.

lutea P. GOLDBLATT and J.C. MANNING 59

Lamiaceae. Rediscovery in South Africa of the neglected African vegetable Plectranthus esculen-

tus. N.R. CROUCH and D.G.A. STYLES 65

Passifloraceae. First description of female flowers of the dioecious Adenia fruticosa subsp. trifo-

liolata. N.R. CROUCH, A. BEAUMONT and G.F. SMITH 78

Pteridophyta. New distribution records and noteworthy collections of pteridophytes in KwaZulu-

Natal. R.R. KLOPPER and N.R. CROUCH 68

Pteridophyta. Notes on some naturalized ferns of the Eastern Cape and KwaZulu-Natal. N.R.

CROUCH and R.R. KLOPPER 71

Pteridophyta. Cheilanthes perrieri J.P.Roux, nom. nov. (Pteridaceae), correcting a nomenclatural

error. J.P. ROUX 81

Pteridophyta. Range extension records from the southern Drakensberg, Eastern Cape, South Africa.

R.R. KLOPPER, S.P. BESTER and G.F. SMITH 82

Pteridophyta. The correct author citation for Cheilanthes marlothii (Sinopteridaceae). J.P. ROUX .... 84

Pteridophyta: Polypodiaceae. The status of x Pleopodium in Africa. N.R. CROUCH, R.R.

KLOPPER and H.F. GLEN 101

Rubiaceae. First record of Geophila in southern Africa. N.R. CROUCH and R. EDWARDS 70

Scrophulariaceae. Two new species of Limoselleae from western South Africa: Trieenia occulta

and Zaluzianskya regalis. J.C. MANNING and P. GOLDBLATT 84

5. Pollen and reproductive morphology of Rhigiophyllum and Siphocodon (Campanulaceae): two unique genera

of the fynbos vegetation of South Africa. W.M.M. EDDIE, C.N.CUPIDO and J.J. SKVARLA 103

6. Floristic composition of wetlands of the South African section of the Maloti-Drakensberg Transfrontier

Park. E.J.J. SIEBEN, D.C. KOTZE and C.D. MORRIS 117

7. Wetland craft plants in KwaZulu-Natal: an ecological review of harvesting impacts and implications for

sustainable utilization. C.H. TRAYNOR, D.C. KOTZE and S.G. McKEAN 135

8. Obituary: Santiago Castroviejo Bolibar (1946-2009). G.F. SMITH 145

New combinations, genus, names, species, statuses, subgenera, subspecies and tribe in Bothalia 40,1 (2010)

Aloe neilcrouchii R.R.Klopper & Gideon FSm., sp. nov., 95

Babiana avicularis Goldblatt & J.C. Manning, sp. nov., 47

Babiana ringens subsp. australis Goldblatt & J.C. Manning,

subsp. nov., 50

Babiana teretifolia Goldblatt & J.C. Manning, sp. nov., 51

Brunia compacta A. V.Hall, sp. nov., 97

Bulbine macrocarpa (Baijnath) Boatwr. & J.C. Manning, comb,

nov., 59

Cheilanthes perrieri J.P.Roux, nom. nov., 81

Elegia elephantina H.P.Linder, sp. nov., 12

Ixia abbreviata var. ovata (Andrews) Goldblatt & J.C. Manning,

comb, nov., 64

Ixia calendulacea Goldblatt & J.C. Manning, sp. nov., 60

Othonna daucifolia J.C. Manning & Goldblatt, nom. nov., 37

Othonna undulosa (DC.) J.C. Manning & Goldblatt, comb,

nov., 39

Platycaulos galpinii ( Pillans ) H.P.Linder & C.R.Hardy, comb,

nov., 8

Platycaulos mahonii (N.E.Br.) H.P.Linder & C.R. Hardy subsp.

humbertii (Cherm.) H.P.Linder & C.R.Hardy, comb, nov., 8

Platycaulos mahonii (N.E.Br.) H.P.Linder & C.R.Hardy subsp.

mahonii, comb, nov., 8

Platycaulos mlanjiensis (H.P.Linder) H.P.Linder & C.R.Hardy,

comb, nov., 8

Platycaulos quartziticola (H.P.Linder) H.P.Linder & C.R.Har-

dy, comb, nov., 8

Pleopeltis x simiana (Schelpe & N .C. Anthony) N.R. Crouch &

Klopper subsp. simiana, comb, nov., 102

Restio adpressus (Esterh.) H.P.Linder & C.R.Hardy, comb,

nov., 24

Restio affinis (Esterh.) H.P.Linder & C.R.Hardy, comb, nov., 28

Restio albotuberculatus H.P.Linder & C.R.Hardy , nom. nov., 24

Restio andreaeanus (Pillans) H.P.Linder & C.R.Hardy, comb,

nov., 24

Restio anomalus H.P.Linder sp. nov., 26

Restio asperus (Mast.) H.P.Linder & C.R.Hardy, comb, nov., 21

Restio caespitosus (Esterh.) H.P.Linder & C.R.Hardy, comb,

nov., 27

Restio calcicola H.P.Linder & C.R.Hardy, nom. nov., 24

Restio capensis (L.) H.P.Linder & C.R.Hardy, comb, nov., 30

Restio cedarbergensis H.P.Linder , sp. nov., 29

Restio clandestinus (Esterh.) H.P.Linder & C.R.Hardy, comb,

nov., 22

Restio constipatus H.P.Linder, sp. nov., 27

Restio curvibracteatus (Esterh.) H.P.Linder & C.R.Hardy,

comb, nov., 28

Restio distylis H.P.Linder & C.R.Hardy, nom. nov., 17

Restio durus (Esterh.) H.P.Linder & C.R.Hardy, comb, nov., 24

Restio elsieae H.P.Linder, sp. nov., 28

Restio femineus (Esterh.) H.P.Linder & C.R.Hardy, comb,

nov., 26

Restio hyalinus (Mast.) H.P.Linder & C.R.Hardy, comb, nov.,

21

Restio karooicus (Esterh.) H.P.Linder & C.R.Hardy, comb, nov.,

28

Restio levynsiae (Pillans) LI.PLinder & C.R.Hardy, comb, nov.,

21

Restio longiaristatus (Pillans ex H.P.Linder) H.P.Linder &

C.R.Hardy, comb, nov., 26

Restio luxurians (Pillans) H.P.Linder, comb, et stat. nov., 30

Restio monostylis (Pillans) H.P.Linder & C.R.Hardy, comb,

nov., 17

Restio muirii (Pillans) H.P.Linder & C.R.Hardy, comb, nov.,

24

Restio nanus (Esterh.) H.P.Linder & C.R.Hardy, comb, nov.,

27

Restio nubigenus (Esterh.) H.P.Linder & C.R.Hardy, comb,

nov., 30

Restio nudiflorus (Pillans) H.P.Linder & C.R.Hardy, comb,

nov., 22

Restio paludicola H.P.Linder, sp. nov., 22

Restio papillosus (Esterh.) H.P.Linder & C.R.Hardy, comb,

nov., 26

Restio parthenocarpos H.P. Linder sp. nov., 30

Restio parvispiculus H.P.Linder & C.R.Hardy, nom. nov., 21

Restio pratensis (Esterh.) H.P.Linder & C.R.Hardy, comb,

nov., 26

Restio pulcher (Esterh.) H.P.Linder & C.R.Hardy, comb, nov.,

21

Restio ramosissimus H.P.Linder & C.R.Hardy , nom. nov., 24

Restio rigidus (Mast.) H.P.Linder & C.R.Hardy, comb, nov.,

21

Restio rigoratus (Mast.) H.P.Linder & C.R.Hardy, comb, nov.,

24

Restio rivulus (Esterh.) H.P.Linder & C.R.Hardy, comb, nov.,

26

Restio rudolfii (Pillans) H.P.Linder & C.R.Hardy, nom. nov.,

24

Restio saxatilis (Esterh.) H.P.Linder & C.R.Hardy, comb, nov.,

27

Restio sporadicus (Esterh.) H.P.Linder & C.R.Hardy, comb,

nov., 26

Restio subgen. Calopsis (Beauv. ex Desv.) H.P.Linder &

C.R.Hardy, stat. nov., 20

Restio subgen. Craspedolepis (Steud.) H.P.Linder & C.R. Hardy,

stat. nov., 1 7

Restio subgen. Eremorestio H.P.Linder & C.R.Hardy, subgen.

nov., 20

Restio subgen. Ischyrolepis (Steud.) H.P.Linder & C.R.Hardy,

stat. nov., 24

Restio subgen. Locapsis H.P.Linder & C.R.Hardy, subgen.

nov., 23

Restio subgen. Pendulostemon H.P.Linder & C.R.Hardy,

subgen. nov., 1 7

Restio subgen. Simplicaulos H.P.Linder & C.R.Hardy, subgen.

nov., 16

Restio subgen. Varirestio H.P.Linder & C.R.Hardy, subgen.

nov., 18

Restio tenuispicatus H.P.Linder & C.R.Hardy, nom. nov., 24

Restio unispicatus (H.P.Linder) H.P.Linder & C.R.Hardy, comb,

nov., 3 1

Restio villosus H.P.Linder & C.R.Hardy, nom. nov., 22

Restio wittebergensis (Esterh.) H.P.Linder & C.R.Hardy, comb,

nov., 28

Rhigiophylleae Eddie & Cupido, tribus nov., 113

Rhodocoma foliosa (N.E.Br.) H.P.Linder & C.R.Hardy, comb,

nov., 1 1

Rhodocoma vleibergensis H.P.Linder, sp. nov., 11

Soroveta H.P.Linder & C.R.Hardy, gen. nov., 6

Soroveta ambigua (Mast.) H.P.Linder & C.R.Hardy, comb, nov.,

6

Thamnea matroosbergensis A. V.Hall, sp. nov., 96

Trieenia occulta ./. C. Manning & Goldblatt, sp. nov., 85

Zaluzianskya regalis J.C. Manning & Goldblatt, sp. nov., 87

IV

Bothalia 40,1 : 1-35 (2010)

A generic classification of the Restioneae (Restionaceae), southern

Africa

H.P. LINDER* and C.R. HARDY**

Keywords: Africa, generic classification, Restionaceae, Restioneae, taxonomy

ABSTRACT

We propose a new generic classification of the African Restionaceae, tribe Restioneae (subfamily Restionoideae), based on the

phylogeny and on extensive morphological data. The phylogeny is based on both plastid sequence data and morphological data.

We delimit the genera to be monophyletic, to minimize the nomenclatural changes, and to maximize the ability to diagnose the

genera. We recognize eight genera, one of which with nine subgenera, in the tribe. Of the currently accepted genera, only three

need changes. We provide descriptions for all genera and subgenera, and include a key to them. In this paper we erect one new

genus, Soroveta, redelimit Platycaulos and Restio, and reduce Calopsis and Ischyrolepis to synonomy under Restio. We list the

species which we recognize under each genus, make 37 new combinations, propose eight new names, and also describe eight new

species that belong in these genera.

CONTENTS

Abstract 1

Introduction 1

Delimiting genera 2

Taxonomy 5

Artificial key to genera of Restioneae 5

Soroveta-Platycaulos grade 6

1 . Soroveta 6

2. Platycaulos 6

Staberoha-Elegia clade 8

3. Staberoha 8

4. Thamnochortus 9

5. Rhodocoma 10

6. Elegia 11

7. Askidiosperma 14

Restio clade 15

8. Restio 15

8. 1 subgen. Simplicaulos 16

8.2 subgen . Pendulostemon 17

8.3 subgen. Craspedolepis 17

8.4 subgen. Varirestio 18

8.5 subgen. Calopsis 20

8.6 subgen. Eremorestio 20

8.7 subgen. Restio 21

8.8 subgen. Locapsis 23

8.9 subgen. Ischyrolepis 24

Acknowledgements 31

References 31

Index 32

INTRODUCTION

The African Restionaceae fsubfam. Restionoideae, ±

350 species) comprise one of the dominant elements of

the fynbos vegetation of the Cape Floristic Region (CFR,

* Institute of Systematic Botany, University of Zurich, Zurich 8008,

Switzerland.

** James C. Parks Herbarium, Department of Biology, Millersville

University, P.O. Box 1002, Millersville, PA 17551. USA/Department

of Botany, MRC-166, National Museum of Natural History, Smithso-

nian Institution, P.O. Box 37012, Washington. DC 20013, USA.

MS. received: 2009-09-23.

Goldblatt 1978) (Taylor 1978; Rebelo et al. 2006), and

is regarded as one of the most important clades of the

larger Cape flora (Linder 2003). Within the African Res-

tionaceae, the tribe Restioneae (Briggs & Linder 2009)

includes most species (288), all of which are endemic

to sub-Saharan Africa or Madagascar. A user-friendly,

maximally informative generic classification of the tribe

is thus essential for the effective study of the Cape flora.

The taxonomy of the Restioneae has been investigated

several times over the past centuries, using different data

sets and different taxonomic concepts. Most of the publi-

cations from the 1 9th and 20th centuries were concerned

with describing the many species which were brought to

Europe by early collectors: for a summary, see Linder

( 1985). However, several publications also focused on the

generic delimitations. In the 19th century, the morphol-

ogy of the species was the primary evidence on which

the generic delimitation was based, resulting in the clas-

sification proposed by Masters (1878; 1897) and used by

Pillans (1928). This was based heavily on the gynoecial

characters and remained in use until 1984 (Adamson &

Salter 1950; Dyer 1976). Citing problems with the Mas-

ters classification, Gilg-Benedict (1930) based an alter-

native classification on the culm anatomical data of her

husband, Gilg (1891), but this classification was unfor-

tunately ignored. By 1 980 it was evident that there were

two major problems with the Masters-Pillans generic

classification. Firstly, it did not effectively summarize the

new anatomical and palynological data, and so was not

natural in a phenetic sense. Secondly, many of the gen-

era were clearly based on plesiomorphic features, and so

could not be monophyletic. In 1984, Finder proposed a

new generic classification, incorporating a much wider

data base than had previously been used — including ana-

tomical (Gilg 1891; Cutler 1969), palynological (Chanda

1966; Chanda & Rowley 1967; Finder & Ferguson

1985), and phytochemical (Harborne & Clifford 1969;

Harbome 1979; Harborne et al. 1985) data in addition to

morphology. He also attempted to define monophyletic

genera. However, this classification still contained prob-

lems. Firstly, the data sampling was not complete, and the

eventual classification proposed was based on the congru-

ence of partial data-set-specific cladograms. Secondly, for

Bothalia 40,1 (2010)

2

several genera, no explicit morphological or anatomical

synapomorphies could be identified, and so these genera

could not be justified in a classic Hennigian sense. The

first set of problems was in the Elegia clade, where Chon-

dropetalum lacked synapomorphies. Moline & Linder

(2005) solved this problem by combining Chondropeta-

Iwn , Elegia and Dovea. The next set of problems was

in the Restio clade. Under Restio , Linder noted: 'How-

ever. it is still not possible to demonstrate that Restio , as

delimited here, is monophyletic’. Under Calopsis, Linder

noted: ‘the exact boundary between Restio and Calopsis

is not yet resolved’. These problems remain unresolved.

Recently, Hardy et ai. (2008) published an almost com-

pletely sampled phylogeny of the African Restionaceae

(subfam. Restionoideae), expanding the previously pub-

lished phylogeny for the nested Elegia group (Moline &

Linder 2005) to include a complete species sampling of

the entire tribe Restioneae (Figures 1-5). This study cor-

roborated earlier indications from a much more sparsely

sampled phylogeny (Eldenas & Linder 2000) that several

genera were para- or polyphyletic. It also provided the

sampling density that enabled a test of the monophyly of

all previously recognized genera, as well as the discov-

ery of many new clades, some of which warrant descrip-

tion as new taxa. As such, the objective of this article is

to integrate this new information into a revised generic

classification of the Restioneae. In order to facilitate the

use of the new generic classification, we list the species

accepted under each genus and subgenus, and we simul-

taneously make all necessary new combinations, and for-

mally describe several as yet undescribed species.

DELIMITING GENERA

Theoretical criteria

The criteria for the circumscription and ranking of

genera have received little attention and most authors

simply indicate that they follow the criteria proposed by

Backlund & Bremer (1998). These were expanded by

Humphreys & Linder (2009), who argued for larger gen-

era, more consistent with the generic concept used by

Bentham & Hooker in Genera plantarum. Thus, genera

are not necessarily the smallest monophyletic or diagnos-

able groups. These, they suggest, may be better recog-

nized by formal infrageneric taxa. Our classification that

follows represents our attempt to optimize our adherence

to the following criteria: I, monophyly; 2, diagnosabil-

ity; 3, nomenclatural stability; and 4, informativeness.

1 . Monophyly. All genera and subgenera recognized

here, were resolved as (potentially) monophyletic (i.e. as

clades) by Hardy et al. (2008; summarized in Figures 1—

5). Monophyletic taxa contain a closed segment of evo-

lutionary history; therefore, such taxa can be expected to

have a much greater predictive power than taxa that are

not monophyletic. Furthermore, monophyletic taxa are

more comparable for evolutionary studies or conservation

efforts which seek to account for phylogenetic diversity

in addition to species diversity in management decisions.

However, in this context monophyly refers to the ‘true’

phylogeny. The data collected by phylogeneticists only

allow us to estimate this phylogeny. These estimates con-

tain several sources of error, two of which are pertinent to

our study. Firstly, different genome partitions may reflect

different phylogenetic histories (Doyle 1992; Maddi-

son 1997). Such incongruent phylogenetic histories may

result from the transfer of a chloroplast from one species

to another during hybridization. In this case, a phylogeny

based on the chloroplast plastid genome may not reflect

the phylogeny of the nuclear genome. The second error

may occur when stochastic variation in DNA base pairs

results in the retrieval of clades. In this case, the shared

DNA base pairs that lead to the retrieval of a clade are

not the result of a common history. An extreme, and pos-

sibly quite rare form of this phenomenon results in long

branch attraction (Felsenstein 1978). Much more com-

mon are nodes with relatively little statistical support.

The phylogenetic inferences of Hardy et al. (2008) were

based on both parsimony and Bayesian analysis of a mod-

erately large dataset consisting of plastid DNA sequences

(± 7.3 Kb) and 1 50 morphological and anatomical charac-

ters for all 292 species and subspecies of the Restioneae.

The robustness of these inferences, however, is limited by

the inability of Hardy et al. (2008) to successfully generate

nuclear DNA sequences. As such, the phylogenetic infer-

ences of Hardy et al. (2008) were strongly influenced by

evolutionary history of the plastid genome which, because

of problems associated with phenomena such as introgres-

sive hybridization (e.g. chloroplast capture) or lineage sort-

ing, may not faithfully reflect the organismal phylogeny in

all aspects (Doyle 1992; Maddison 1997). Additionally, a

plastid-specific bias or imbalance in molecular evolution-

ary rates across the phylogeny could result in phyloge-

netic error unchecked by another, unlinked nuclear DNA

dataset (e.g. Felsenstein 1978; Doyle 1992). Although the

morphological and anatomical data are thought to provide

phylogenetic signal under the influence of the nuclear

genome and therefore independent of the plastid dataset,

the relatively large size of the plastid dataset may have had

a disproportionate influence on the resulting phylogenetic

inferences. However, because the addition of our mor-

phological data to the plastid data in a combined analysis

resulted in increased support (via bootstrap and posterior

probabilities) for most clades than did either data set alone

( Hardy et al. 2008), we are confident that the phylogeny is

a good approximation of the species phylogeny.

A second problem with our phylogeny is that several

of the deeper nodes from the strict consensus trees have

very low bootstrap support. This could mean that these

nodes will not be retrieved if a larger dataset were to be

generated.

2. Diagnosability. In addition to monophyletic groups,

we sought to recognize genera that were diagnosable mor-

phologically. Essential and differential morphological

attributes can also be used for the assignation of species

to the genera, and to develop a morphological concept of

a genus. These characters were sought by using the parsi-

mony options in Mesquite (Maddison & Maddison 2003)

to optimize the morphological and anatomical character

matrix used in Hardy et al. (2008; available at http://her-

barium.millersville.edu/pubs-support.php) over the com-

plete cladogram from Hardy et at. (2008). We defined as

diagnostic characters those character states that were opti-

mized to the crown-node (base) of each clade that could

be treated as a genus or subgenus, i.e. we ignored variation

Bothalia 40,1 (2010)

3

within the clade. We then coded these diagnostic charac-

ters (Table 1) for the clades into a new matrix (Table 2)

and then mapped them onto a summary tree of the clades

(Figure 1) using CLADOS (Nixon 1993). Morphological

concepts and states were largely as used by Linder (1984),

and are explained, illustrated, and fully documented in the

interactive key to the species (Linder 2001a) (http://www.

systbot.uzh.ch/Bestimmungsschluessel/Restionaceae.

html). Species for which molecular data were not available

(20 out of 289) were placed on the basis of morphological

and anatomical data, by their inclusion in a cladistic analy-

sis with the full data set. Essential and differential morpho-

logical and anatomical attributes were extracted from the

morphological data set for each postulated clade. Essential

attributes are typical (albeit sometimes with exceptions)

of each clade, but not unique to it. Differential attributes,

often as combinations of characters, are unique to each

clade, and can be used to diagnose the clades. Differential

attributes can be used to assign previously unstudied spe-

cies to their appropriate clades.

A second important component of diagnosability is

the ability of users to be able to assign species to the

right genus without too complex a procedure, ideally

even without having to use a hand-lens. Thus diagnos-

ability that is only possible on anatomical attributes is

not optimal.

3. Nomenclatural stability. Genera should be recog-

nized at nodes that will minimize nomenclatural changes.

Nomenclatural changes fall into three elements. Firstly,

minimizing the number of new combinations: these are

most common when genera are divided into segregate

genera; in these instances the specific epithet stays the

same. Secondly, reducing the number of new names that

need to be proposed: these are most common when gen-

era are combined, and result when a specific epithet is

already occupied; these are much more confusing than

new combinations, as the species now has a completely

TABLE 1. — Selected generic and subgeneric characters mapped onto

trees in Figure 1. Character numbers above branch and state of

character below branch

1 . Culms: round or flattened = 0; square or angular = 1 .

2. Culms branching: simple = 0; branching = 1 .

3. Culms: round = 0; compressed = 1.

4. Sheaths apical half: as lower part = 0; membranous and decaying

= 1.

5. Membranous sheath shoulders: absent = 0; present = 1 .

6. Sheaths: persistent = 0; caducous = 1 .

7. Floral bract apices: like body = 0; with hollow cells = 1 .

8. Floral bracts: overtopping flowers = 0; shorter than flowers = 1 .

9. Floral bracts: chartaceous, cartilaginous or bony = 0; membra-

nous = 1 .

10. Floral bracts transverse pitting: absent = 0; present = 1.

1 1 . Male spikelets: erect = 0; pendulous = 1 .

12. Anthers at anthesis: exserted from flowers = 0; included in the

flowers = 1 .

13. Female tepals (indumentum): glabrous = 0; villous = 1 .

14. Female tepals (texture): papery (flexible, not transparent) = 1;

bony or cartilaginous (stiff) = 2.

15. Female lateral tepals: as odd tepal = 0; conduplicate = 1; winged

= 2. [additive].

1 6. Number of styles: 1 = 0; 2 = 1 ; 3 = 2. [additive].

1 7. Styles: free or on a stylopodium = 0; fused to form a peg = 1 .

18. Number of ovary locules: 1 = 0; 2 = 1; 3 = 2. [additive].

19. Ovary: dehiscent = 0; indehiscent = 1.

20. Central ground tissue cavities: none = 0; single = 1 ; many = 2.

[additive],

different name (both genus and species have changed).

Thirdly, frequent changes in the combinations: this results

when there is a frequent changing of the generic concepts.

In some circumstances no new formal names are required,

but from the user point of view there are still changes.

A second consideration should be not only to mini-

mize the number of nomenclatural changes required,

but also to consider the nomenclatural consequences of

future changes in our understanding of the phylogeny.

Ideally, genera should be erected at nodes which are

likely to remain robust to future additional data, be it

new species or new data sources. Theoretically, at least,

larger genera should be less sensitive to smaller changes

in the phylogenetic hypothesis.

4. Informativeness. Genera should have a morpho-

logical and ecological ‘coherence’, and not consist of

mere collections of species that happen to share a plastid

genome. This maximizes the number of attributes that we

can assign to a genus, and makes the genus concept more

useful.

Genera in the Restioneae

With the cladistic and morphological data available

for the Restioneae, several generic classifications for the

Restioneae are possible. In order to facilitate the discus-

sion here, we refer to the names of clades described later

in the paper.

1. The maximal option is to include the whole sub-

tribe in one genus. This genus would be strongly sup-

ported as being monopliyletic by the plastid data (Eldenas

& Linder 2000; Hardy et al. 2008), but would be difficult

to diagnose, as the morphological characters separating

Willdenowieae and Restioneae are cryptic anatomical and

palynological attributes (Linder 1984), and the clade has

no diagnostic attributes. The characters listed in Figure I

as synapomorphies for the clade (the numbers of styles

and carpels), should actually be listed under the Willde-

TABLE 2. — Matrix-representation of distribution of diagnostic or

distinctive characters from Table 1 for proposed genera and sub-

genera of tribe Restioneae. Polymorphism codes as follows: A =

states 0 and 1 ; B = states 1 and 2; C = states 0 and 2

1 5 10 15 20

0 1 000000000002 0 100 1 100

0 1 0000000 1 000202 0 10 1 00

01 100000000002 A2 010000

0000010100010212020001

00000000001 101 B2 001 1A0

0000010010010212020000

0001000000 1 1022000 1210

00010001001 102020202 A0

1 10000000000 A2 12001 100

0100001000001212010000

0 1 0000000000 Al 12001000

0100 A0 0010001 1111 A0 100

0 1 0000000000 1 1 1 2 001100

0100000000100212010100

010010000000 AB1200A100

0000000000000212010100

010000 A0 0000 A2 120 CO 100

Willdenowieae

Soroveta

Platycaulos

Elegia

Staberoha

Askidiosperma

Thamnochortus

Rhodocoma

Restio subgen. Calopsis

Restio subgen. Craspedolepis

Restio subgen. Eremorestio

Restio subgen. Ischyrolepis

Restio subgen. Locapsis

Restio subgen. Pendidostemon

Restio subgen. Restio

Restio subgen. Simplicaulos

Restio subgen. Varirestio

4

Bothalia 40,1 (2010)

nowieae — the Restioneae have the ancestral condition of

three styles and three carpels. The genus would not be

exceptionally large (some 300 species), but would be mor-

phologically very heterogenous. It would require numer-

ous name changes, and would lose on informativeness, as

several very distinct genera, such as Elegia, Thamnochor-

tus and Staberoha , would be lost. This broad approach has

been followed in several Cape taxa, such as Erica (Oliver

2000) and Disa (Bytebier et al. 2007, 2008).

2. The minimal option is to retain all existing genera

that were retrieved as monophyletic (i.e. Askidiosperma,

Elegia, Ischyrolepis, Platycaulos , Rhodocoma, Staberoha

and Thamnochortus), and split the remainder up into the

smallest number of genera based on clades with at least

70 % bootstrap support (named in Figures 1-5 as Restio

subgen. Calopsis, Restio subgen. Craspedolepis, Restio

subgen. Eremorestio , Restio subgen. Locapsis , Restio sub-

gen. Pendulostemon , Restio subgen. Restio , Restio subgen.

Simplicaulos , Restio subgen. Varirestio and Soroveta ).

Note that Platycaulos is here somewhat unusual, as this

genus receives several species misplaced to Restio in the

previous classifications. The recognition of more than one

genus in the tribe implies that the basal branch leading to

Restio ambiguus has to be recognized at generic level (as

the genus Soroveta ), as the node excluding this species

from the rest of the tribe is very strongly supported. This

results in 16 genera, all of which are strongly supported

as being monophyletic by the combined plastid DNA and

morphological datasets, but not all by the morphological

dataset alone. There are great difficulties in diagnosing

several of these clades. In some instances no essential or

differential characters could be found (e.g. Restio subgen.

Varirestio), in others (e.g. Restio subgen. Craspedolepis,

Restio subgen. Eremorestio , Restio subgen. Locapsis, Res-

tio s.stc , Restio subgen. Simplicaulos ) there are essential

but no differential characters, i.e. they also occur in other

genera albeit relatively rarely. As a result we can have a

vague concept of the genus, but cannot key it out. Predict-

ably, these smaller genera are mostly morphologically

quite homogeneous. This approach requires more than 70

new combinations, but no new names.

The advantage of this approach is that the monophyly

of all genera is strongly supported by the available phy-

logenetic hypothesis. However, there are two problems.

The first is that the generic classification will not be user

friendly, as it will be difficult to assign species to the gen-

era, either in the field or in a herbarium. Secondly, there

is a very heavy reliance on plastid data in the absence

of nuclear DNA data (Doyle 1992). Any mistakes in the

phylogeny will result in generic redelimitation.

3. The intermediate option is to retain all existing

genera that are not proven to be para- or polyphyletic, and

not to start with the assumption that all existing genera that

are retrieved as monophyletic have to be retained. In this

option, the distinctive and well-supported genera Askid-

iosperma, Elegia, Platycaulos, Rhodocoma, Soroveta, Sta-

beroha and Thamnochortus are retained, and a larger Res-

tio sdat. is assembled. The major advantage of this option

is that the two difficult segregates, Eremorestio and Vari-

restio, will not be recognized as distinct genera. Due to the

structure of the phylogeny, in all these solutions, Ischy-

rolepis will need to be included in Restio sdat., together

with most species of Calopsis, as the nodes embedding

these two genera within Restio s.lat. are strongly supported

in the combined analysis. In the morphological phylogeny

of Linder (1984) Ischyrolepis was not included in the Res-

tio-Calopsis clade, due to its very different pollen mor-

phology. A major insight from the molecular phylogeny is

that the pollen data are not phylogenetically conservative,

as a result Ischyrolepis is found to be nested in the Res-

tio-Calopsis clade. Three variants of Restio s.lat. can be

proposed:

3a. Include Calopsis s.str., Craspedolepis, Eremores-

tio, Ischyrolepis, Locapsis, Pendulostemon, Restio s.str,

Simplicaulos and Varirestio in one genus. There is weak

evidence that this grouping is paraphyletic: in the com-

bined phylogeny, but with no bootstrap support, Pendu-

lostemon groups with the Thamnochortus-Rhodocoma

clade, and Simplicaulos with Staberoha). This large

genus is easy to diagnose by exclusion (not Rhodocoma,

Thamnochortus, Soroveta, Platycaulos, Elegia or Askid-

iosperma). It is the most conservative approach, requir-

ing statistically significant evidence to dismantle a genus.

The disadvantage of this classification is that there is no

evidence for the monophyly of the genus Restio s.lat.

(but also no evidence that it is not monophyletic). The

advantage is that the genus is easy to recognize in the

field and herbarium and that it is nomenclaturally con-

servative (if eventually there is sufficient evidence that

to separate Simplicaulos and Pendulostemon will not

require undoing a set of nomenclatural changes). The

strongly supported monophyletic units can then be rec-

ognized as subgenera of Restio s.lat., these should be

used for evolutionary investigations.

3b. As above, but retain Simplicaulos and Penduloste-

mon as distinct genera on the basis that there is weak

evidence linking these to Staberoha and Thamnochortus

respectively. It removes all evidence that Restio s.lat.

could be paraphyletic, but there is also no evidence that

the remaining Restio s.lat. is monophyletic. Penduloste-

mon can be readily diagnosed, but Simplicaulos lacks

differential characters. The most important attribute is

the unbranched culms: this is a highly variable character.

The advantage is that no genera are recognized for which

there is (even poorly supported) evidence for paraphyly.

The disadvantages are two-fold: due to the low support

values, it remains possible that additional datasets will

group the two segregates again with Restio s.lat., leading

to name changes. Furthermore, one of the two segregate

genera cannot be keyed out.

3c. Recognize, in addition to Simplicaulos and Pen-

dulostemon, also Craspedolepis. This leaves the rest of

Restio s.lat. with positive (albeit weak) evidence of its

monophyly, but it adds another genus that is difficult to

diagnose. Frustratingly, Craspedolepis does have a strik-

ing essential character, a band of hollow (concave) cells at

the apical margins of the floral bracts, but this is in some

species poorly developed, and also occurs in a few species

outside the genus. It is thus of little use as a differential

character.

We follow here option 3a, which recognizes eight

genera in the tribe. The eight strictly monophyletic seg-

regates of Restio s.lat. are recognized as subgenera. We

hope that this will combine the need for strictly mono-

phyletic taxa for evolutionary analysis (the subgenera).

Bothalia 40,1 (2010)

5

FIGURE 1. — Cladograms summarizing generic and subgeneric relationships in the Restioneae (Hardy et al. 2008). A, relationships in the strict

consensus tree from the analysis of all 292 species and subspecies. B, relationships in the strict consensus tree from the analysis of 272 spe-

cies and subspecies omitting 20 species/subspecies for which DNA data were lacking. Bootstrap percentage values (in square brackets) are

based on the analysis of Hardy et al. (2008). Characters from Tables 1 and 2 are optimized using the slow (deltran) options in WinClada: the

character number appears above the solid and open squares. The state of the character appears below the solid and open squares. An open

square indicates a character that evolved at least twice, whereas a solid quare indicates a character that evolved only once.

with easy-to-recognize, nomenclaturally stable genera. It

is important, though, that no taxa that are demonstrably

para- or polyphyletic be recognized.

TAXONOMY

The full nomenclature and synonymy of the species

is not repeated: it was published in 1985, consequently

only the accepted species and their place of publica-

tion is given. However, for the new combinations, the

basionym and the type is indicated, as required. Again

no full synonymy is presented. The complete synonymy,

descriptions, illustrations to all species, and an interac-

tive key is available online at http://www.systbot.uzh.

ch/Bestimmungsschluessel/Restionaceae.html, or can be

bought as a CD from the Bolus Herbarium. University

of Cape Town. The species are presented in a ‘natural’

sequence, partially based on the phylogeny, partially

based on their similarity.

Artificial key to genera of Restioneae

la Floral bracts reddish brown, bony, with transverse lacunae 1 . Soroveta

lb Floral bracts various, always without transverse lacunae:

2a Male spikelets pendulous:

3a Ovary indehiscent; diaspore a nut enclosed in a persistent, papery to cartilaginous, perianth, which may be winged:

4a Male spikelets linear-oblong; female flowers with a single style exserted from floral bracts; perianth a firm winged or keeled struc-

ture; sheaths decaying in upper half 4. Thamnochortus

4b Male spikelets ± globose; females with 1-3 styles obscured behind floral bracts; perianth papery, sometimes keeled; sheath apical

margins firm, persistent 3. Staberoha

3b Ovary dehiscent; diaspore a seed:

5a Culms branching; female floral bracts taller than flowers, often with longitudinal striations 8.2 Restio subgen. Pendulostemon

5b Culms simple (except for R. capensis, where culms have whorled branches); female floral bracts shorter than flowers, never with

longitudinal striations 5. Rhodocoma

2b Male spikelets erect:

6a Sheaths falling off, sometimes somewhat tardily so, leaving a distinct dehiscence ring:

7a Floral bracts membranous, much taller than flowers; ovary dehiscent or tardily indehiscent 7. Askidiosperma

7b Floral bracts leathery to bony, rarely taller than flowers, mostly shorter; ovary either dehiscent or indehiscent 6. Elegia

6b Sheaths persistent:

8a Culms laterally compressed (sometimes only at apex); sheaths with stout mucro, both same green colour as culm, usually with

woolly scale in sheath axils 2. Platycaulos

6

Bothalia 40,1 (2010)

8b Culms round; if laterally compressed then sheaths brown, clearly different from culm, and without woolly scales:

9a Female inflorescences of numerous poorly organized spikelets arranged at several nodes; bracts not obscuring flowers; spathes

much overtopping groups of spikelets at nodes 6, Elegia

9b Female inflorescences of one to many well-organized spikelets, either racemosely or paniculately organized; bracts mostly

obscuring flowers; spathes at most as tall as spikelets (Restio s.lat.):

10a Culms square or otherwise angular in cross section; plants 1 m or taller; inflorescences large, paniculate, with very numerous

spikelets 8.5 Restio subgen. Calopsis

10b Culms round, terete or compressed (flattened) in cross section; plants variable in height, but often less than 1 m; inflorescence

rarely very large:

11a Style branches two, fused below into persistent stylar peg; ovary always dehiscent 8.9 Restio subgen. Ischyrolepis

lib Style branches 1, 2 or 3, never fused into persistent stylar peg; ovary either dehiscent or indehiscent:

12a Culms simple 8.1 Restio subgen. Simplicaalos

12b Fertile culms branched:

13a Floral bracts with upper margin markedly different from bract body, consisting of hollow cells; ovary usually bilocular,

dehiscent 8.3 Restio subgen. Craspedolepis

13b Floral bracts without such upper margins; ovary various:

14a Female perianth papery, truncate or rounded; ovary indehiscent, unilocular:

15a Female floral bracts acuminate, apices somewhat recurved, darker than bract body; sheaths always tightly rolled

around culms; nuts triangular, more than 2 mm long 8.6 Restio subgen. Eremorestio

15b Female floral bracts acute, apices erect, same colour as body of bracts; sheaths mostly spreading, sometimes tightly

rolled around the culms; nuts various 8.8 Restio subgen. Locapsis

14b Female perianth coriaceous or bony, acute; ovary indehiscent or dehiscent, with 1, 2, or 3 locules:

16a Ovary with 1 or 3 locules; sheath awn sometimes as long as sheath; sheaths with poorly developed membranous

lobes 8.4 Restio subgen. Varirestio

16b Ovary with 1 or 2, rarely 3 locules; sheath awn shorter than sheath 8.7 Restio subgen. Restio

Soroveta-Platycaulos grade

An expanded Platycaulos and the newly erected

genus Soroveta H.P.Linder & C.R. Hardy, gen. nov.,

comprise a relatively species-poor grade within the Res-

tioneae that is paraphyletic to the remainder of the Res-

tioneae (Figures 1-5). The monotypic Soroveta is rather

isolated and does not share any particularly striking

morphological characteristics with Platycaulos. Perhaps

their greatest resemblances lie in their native habitats,

where both characteristically grow in areas of impeded

drainage or summer rainfall, although the two genera do

not co-occur.

1. Soroveta H.P.Linder & C.R. Hardy, gen. nov., ab

aliis generibus tribus Restioneae bracteis floralibus trans-

versaliter foveatis et combinatione vaginorum persisten-

tium cum tepalis uniformibus distinguendum.

TYPE. — Soroveta ambigua (Mast.) H.P.Linder & C.R.

Hardy.

Plants clumped. Fertile culms sparsely branched,

round; sheaths persistent, closely convoluted, margins

coriaceous, without hyaline shoulders. Male inflores-

cence racemose; spikelets 1-5, erect, several-flowered;

spathes persistent, shorter than spikelets; bracts dark

brown, with transverse pitting; anthers exserted from

(lowers. Female inflorescence and spikelets similar

to males, with 1-5 linear or oblong, acute, 2-8-flow-

ered spikelets; flowers not laterally compressed; tepals

chartaceous or cartilaginous, scabrid all over, outer lat-

eral tepals not differentiated from outer odd tepal; stami-

nodes present. Gynoecium with 3 white, feathery styles

free to base; ovary bilocular, dehiscent. Seed brown,

pitted. Culm anatomy, epidermal cells single-layered,

lateral radial wall straight, unthickened; chlorenchyma

two-layered, stomatal cavities with protective cells;

parenchymatous layer between chlorenchyma and scle-

renchyma of 3-5 cells; central ground tissue with single,

central cavity; tannin and silica absent.

Etymology. Soroveta (f): soror (Latin; f., Ill), sis-

ter; veins (Latin), ancient; referring to the topological

position of the genus, as the sister to the rest of Res-

tioneae (Ligure 2).

Notes', the isolated position of this species was

already recognized by Masters (1897) who gave it the

very prescient specific epithet. Morphologically, the

species shows several unusual features in the male and

female inflorescences, notably resembling Elegia in the

naked female flowers without any lateral (dorsiventral)

flattening, but without the caducous sheaths and other

attributes typical of that clade. Furthermore, the dark

brown floral bracts with transverse pitting are unique in

the subfamily. However, there are no obvious synapo-

morphies for the rest of the Restioneae, consequently the

basal position of this species cannot be detected morpho-

logically. In the absence of molecular data, the species

would probably have been retained as a highly special-

ized species of Restio.

1.1. Soroveta ambigua (Mast.) H.P.Linder & C.R.

Hardy , comb. nov.

Restio ambiguus Mast, in Flora capensis 7: 96 (1897). Type: Cape,

s.loc ., Zeyher s.n. (K, liolo.!; B!).

2. Platycaulos H.P.Linder in Bothalia 1 5: 64 (1984).

Type; Platycaulos compressus (Rottb.) H.P.Linder.

Plants tufted, tangled or mat-forming. Fertile culms

branching, more or less compressed, though sometimes

only near apex; sheaths persistent, with no abscission

line, closely convoluted, usually densely textured and

same colour as culms, apical margins narrowly membra-

nous, continuing behind the stout, awl-like awn. Male

inflorescence racemose or paniculate with up to 20 sessile

or stiffly erect, often distichous spikelets; spathes persist-

ent or caducous; male bract with upper margin like body

of bract; anthers exserted from flowers. Female inflor-

escence with 1-10 spikelets; spikelets 1-15-flowered,

variously shaped; bracts at least as tall as flowers, often

much taller, mostly cartilaginous, apical margin like rest

of bract; imbricate and obscuring spikelet axis; tepals

chartaceous or cartilaginous, glabrous or keels of lateral

sepals villous; staminodes present. Gynoecium with 3

white, feathery styles free to base; ovary with 1 or 2 fer-

Bothalia 40,1 (2010)

7

Soroveta ambigua

1 001 Platycaulos depauperatus

Platycaulos subcompressus

Platycaulos galpinii (Restio galpinii)

58 1 Platycaulos mahonii subsp. mahonii (Restio mahonii subsp. mahonil)

Platycaulos mlanjlensis (Restio mlanjiensis)

88 1 Platycaulos mahonii subsp. humbertii (Restio mahonii subsp. humbertii) '

Platycaulos quartziticola (Restio quartziticola) *

Platycaulos anceps

Platycaulos compressus

1 00 Platycaulos major

61 | Platycaulos acutus

68 I 88 I Platycaulos callistachyus

84 Platycaulos cascadensis

Restio fuscidulus (Ischyrolepis fuscidula) *

Staberoha stokoei

Staberoha remota

Staberoha aemula

Staberoha cernua

Staberoha distachyos

Staberoha banksii

3 1 nnl 85 | Staberoha multispicula

29 I 88 I Staberoha ornata

Staberoha vaginata

Restio pedicellatus

Restio sp. aff. pedicellatus

Restio stereocaulis *

Restio subtilis

Restio confusus

Restio miser

Restio distylis (Calopsis esterhuyseniae)

Restio monostylis (Calopsis monostylis)

Restio papyraceus

Restio echinatus

80 | Restio nuwebergensis

7« I 80 | Restio bifarius

Restio bifidus

80

82

‘ Restio egregius

‘ Restio micans

Restio subgen. Simplicaulos

Rhodocoma vteibergensis

‘ Rhodocoma alpina

‘ Rhodocoma fruticosa

‘ Rhodocoma foliosa

' Rhodocoma gigantea

' Rhodocoma arida

' Rhodocoma capensis

" Rhodocoma gracilis

' Thamnochortus levynsiae

' Thamnochortus pulcher

' Thamnochortus gracilis

' Thamnochortus nutans

Restio subgen. Pendulostemon

Thamnochortus karooica

Thamnochortus muirii

Thamnochortus paniculatus

Thamnochortus spicigerus

Thamnochortus fraternus

Thamnochortus pluristachyus

Thamnochortus cinereus

Thamnochortus rigidus

Thamnochortus glaber

Thamnochortus amoena *

51 | Thamnochortus papyraceus

40 I 99 j Thamnochortus acuminatus

99 Thamnochortus fruticosus

Thamnochortus guthrieae

Thamnochortus erectus

Thamnochortus insignis

Thamnochortus arenarius

Thamnochortus ellipticus *

Thamnochortus pellucidus

Thamnochortus stokoei

Thamnochortus dumosus

Thamnochortus lucens

Thamnochortus platypteris

Thamnochortus schlechteri

Thamnochortus obtusus

Thamnochortus sporadicus

Thamnochortus bachmannii

Thamnochortus punctatus

FIGURE 2. — Portion of strict consensus tree of total combined analysis of all species by Hardy et al. (2008) containing Soroveta, Platycaulos,

Staberoha, Restio subgen. Simplicaulos, Restio subgen. Pendulostemon, Rhodocoma, and Thamnochortus . Names in parentheses are the now

synonymous names of Linder (2001a) when different from the ones here proposed. A single species of Restio subgen. Ischyrolepis ( Restio

fuscidulus) that was missing many data (including all molecular data), was resolved in this tree as sister to Staberoha, albeit with 3 % boot-

strap support which we do not consider substantial enough to warrant disputing the monophyly of Restio subgen. Ischyrolepis or the genus

Restio as circumscribed here. Bootstrap values above branches indicate support based on a 500-replicate bootstrap analysis that included

20 species for which DNA data were lacking (asterisked taxa). Bootstrap values below branches indicate support based on an analysis that

excluded these morphology-only taxa. Where no value is given below a branch, it indicates that the clade did not appear in the analysis

excluding morphology-only species.

Bothalia 40,1 (2010)

tile, dehiscent locules. Seed brown, black, white or tan,

with brittle white ornamentation or rarely rugose. Culm

anatomy : epidermal cells in 1 layer, lateral wall straight,

unthickened; parenchymatous layer of 1-4 cells; central

ground tissue without cavity; tannin absent or present in

epidermis, sclerenchyma or central ground tissue; silica

absent.

Etymology : Platycaulos (m.): platvs (Greek), broad,

flat; kaulos (Greek), stalk; referring to the compressed

culms typical of this genus.

Notes', the genus was separated from Restio by Linder

in 1984. based on a range of morphological, anatomical

and palynological attributes. At the time, Linder did not

realise that these features were also visible in the tropi-

cal species (now included below via new combinations),

albeit in a less than obvious fashion. Fortunately, the

taxon’s namesake attribute, the more or less compressed

culms, is also found in the tropical species, therefore the

generic name remains appropriate. Almost all species are

found in habitats with impeded drainage or summer rain-

fall.

2. 1 . Platycaulos depauperatus ( Kunth ) H.RLinder in Bo-

thalia 15:436(1985).

2.2. Platycaulos subcompressus (Pil/ans) H.PLinder in Bo-

thalia 15: 347 (1985).

2.3. Platycaulosgalpinii(P/7/<ms) H.PLinder &C.R.

Hardy, comb. nov.

Restio galpinii Pillans in Transactions of the Royal Society of South

Africa 30: 250 (1945). Type: Natal, Underberg, between Cathkin Peak

and Giants Castle, Esterhuysen 8820 (BOL, lecto.! [Linder in Bothalia

15:447(1985)], K!,NBG!, NY).

2.4a. Platycaulos maA\om\(N.E.Br) H.PLinder & C.R.

Hardy subsp. mahonii, comb. nov.

Hypolaena mahonii N.E.Br. in Flora of tropical Africa 8: 265

(1901). Restio mahonii (N.E.Br.) Pillans: 255 (1945). Type: Malawi,

Southern Province, MtZomba, Mahon s.n. (K, holo.!; B!, BOL!).

2.4b. Platycaulos mahonii (N.E.Br.) H.PLinder &

C.R. Hardy subsp. humbertii (Cherm.) H.PLinder & C.R.

Hardy, comb. nov.

Restio madagascariensis Cherm. var. humbertii Cherm. in Archives

de Botanique, Bulletin Mensuel, Caen 7: 85 (1930). R. mahonii

(N.E.Br.) Linder subsp. humbertii (Cherm.) H.PLinder: 103 (1986).

Type: Madagascar, massif de i'Andohahelo, Humbert 6147 (P, holo.!;

B!, K!, NY).

2.5. Platycaulos mlanjiensis (H.PLinder) H.P.Lin-

der & C.R. Hardy, comb. nov.

Restio mlanjiensis H.PLinder in Kew Bulletin 50: 623 (1995).

Type: Malawi, Mt Mlanje, Sombani, Verboom, Pauw & Hooks 1 (BOL,

holo.!; K!, MALI, MO!, NSW!, PRE!, S!, SRGH!, WAG!, Z!).

2.6. Platycaulos quartziticola (H.PLinder) H.P.

Linder & C.R. Hardy, comb. nov.

Restio quartziticola H.PLinder in Kew Bulletin 41:103 (1986).

Type: Zimbabwe, Chimanimani Mts, Whellan 1252 (K, holo.!; BOL!,

BR, LISC, SRGH!).

2.7. Platycaulos compressus ( Rottb .) H.P.Linder in Bo-

thalia 15; 436(1985).

2.8. Platycaulos major (Mast.) H.P.Linder in Bothalia 15:

436(1985).

2.9. Platycaulos nnceps(Mast.) H.P.Linder in Bothalia 15:

436(1985).

2. 1 0. Platycaulos acutus Esterh. in Linder in Bothalia 15;

434(1985).

2.11. Platycaulos cascadensis (Pillans) H.P.Linder in Bo-

thalia 15: 436 (1985).

2. 12. Platycaulos cattistachyus (Kunth) H.P. Linder in Bo-

thalia 15:436(1985).

Staberoha-Elegia clade

Although molecular support for this clade is poor (see

also Hardy et al. 2008), this clade makes sense morpho-

logically, based largely on its combination of glabrous,

conduplicate tepals and, mostly, simple, unbranched

culms. Interestingly, the species in the family with pen-

dulous male spikelets (e.g. Staberoha, Thamnochortus ,

Rhodocoma, Restio subgen. Pendulostemon H.P.Linder &

C.R. Hardy subgen. nov.) all occur in this clade, suggest-

ing that pendulous male spikelets are either plesiomorphic

or have evolved repeatedly in parallel within the clade.

Within this clade, the association between Staberoha

and Restio subgen. Simplicaulos H.P.Linder & C.R. Hardy

subgen. nov. (Figure 2) is weak, and an alternative posi-

tion is for Simplicaulos to be sister to Elegia, and for Sta-

beroha to occupy an isolated position (Hardy et al. 2008).

The two taxa may be linked morphologically by their sim-

ple culms, but that could be plesiomorphic in the whole

clade, with a reversal in Restio subgen. Pendulostemon.

Many species in these two taxa also have large female

bracts that completely obscure the female flowers.

Restio subgen. Pendulostemon, consisting of just two

species, is both phylogenetically and morphologically

isolated in the clade. In particular, the branching culms

of this genus are unusual for the clade, although species

with branching culms also occur occasionally in Elegia

and Thamnochortus.

The Thamnochortus-Rhodocoma subclade was previ-

ously recognized by Linder (1984). The two genera share

pendulous male spikelets (also -present in Staberoha and

Restio subgen. Pendulostemon), scattered cavities in the

central ground tissue, and sheaths of which the upper

half is more or less membranous and soon decays. The

genera have very different gynoecia: in Rhodocoma the

ovaries are dehiscent, and the diaspore is a seed, whereas

in Thamnochortus the ovaries are unilocular and indehis-

cent, and the fruit a soft-walled nut, included in an often

widely winged, persistent perianth.

The E/egia-Askidiosperma subclade also dates back

to the morphological phylogeny of Linder ( 1984), and is

recognized by the caducous, or when persistent, poorly

structured sheaths, and the inflorescences with numerous

partial panicles originating from several nodes along a

central axis. Furthermore, the tepals are quite bony, with

generally no differentiation between the lateral and odd

tepals. The central ground tissue lacks a cavity. Elegia is

distinguished from Askidiosperma by a double-layered

culm epidermis, whereas Askidiosperma is diagnosed

by the very long, membranous bracts, and its more well-

defined spikelets.

3. Staberoha Kunth, Enumeratio plantarum 3: 442

( 1841 ). Lectotype: Staberoha distachyos (Rottb.) Kunth.

Plants caespitose or clumped. Fertile culms unbranched,

round; sheaths persistent, closely or rarely loosely con-

voluted, margins usually like rest of sheath, rarely nar-

Bothalia 40,1 (2010)

9

rowly or broadly membranous, but hyaline shoulders

never developed. Male inflorescence paniculate with I

to several ± globose spikelets, pendulous on flattened,

flexible pedicels, rarely racemose and erect (S. stokoei)',

bracts much taller than flowers and completely obscur-

ing them; tepals papery, lateral tepals keeled; anthers

usually included in flowers at anthesis. Female inflor-

escence racemose with 1-5 linear, oblong, elliptical or

ovate, erect spikelets, each with 6-25 flowers and no

sterile bracts; female bracts at least twice as long as and

obscuring flowers, imbricate. Female flowers laterally

compressed; tepals mostly chartaceous, glabrous, outer

lateral tepals flattened, keeled or winged; staminodes

sometimes present. Gynoecium with 1-3, white, flat-

tened styles obscured behind bracts at anthesis; style

bases usually free or rarely seated on a stylopodium;

ovary unilocular, indehiscent. Fruit a soft-walled nut

dispersed with persistent, often winged, perianth. Culm

anatomy, epidermal cells in 1 layer, lateral wall usually

straight and unthickened, rarely sinuose and thickened;

parenchymatous layer of 1-4 cells; central ground tissue

rarely solid, usually with scattered cavities or a single,

central cavity; tannin usually absent, if present usually

restricted to epidermis; silica absent.

Etymology. Staberoha (f.): named after H. Staberoh, a

medical doctor, pharmacist and experimentalist.

Notes', a very distinctive genus, almost restricted to

the Western Cape, where often several species are found

co-occurring. The ball-like, pendulous male spikelets

and the small female flowers obscured behind the large

female bracts allow an easy recognition of the genus. All

species are found on well-drained soils, both on deep

sandy soils (such as Staberoha distachyos ) and on rocky

slopes (S. remota). S. stokoei is a remarkable exception

to many of the morphological attributes (the male spike-

lets are erect, and almost identical to the female spike-

lets), and is a geographically restricted species in the

Great Swartberg, where it occupies almost vertical rock

ledges on the upper south-facing slopes. It is satisfying

to see that it occupies a topological position sister to the

rest of the genus, indicating that many of the distinctive

attributes of the genus evolved after the origin of the

genus. If subdivision of the genus is appropriate, then S.

stokoei could be regarded as one division, and the rest of

the genus the second.

The pollen of this genus is unusual, with a thickened

annulus, very similar to the pollen typical of the Poaceae

(Linder & Ferguson 1985). Similar pollen is also found

in Restio subgen. Ischyrolepis. In the morphological

analysis by Linder (1984) this pollen played an impor-

tant role in emphasizing the phylogenetically early

branching position of Staberoha , but the molecular anal-

ysis indicates that this curious pollen type most likely

evolved twice, independently, in the Restioneae.

3.1. Staberoha stokoei Pillans in Journal of South Afri-

can Botany 18: 118 (1952).

3.2. Staberoha remota Pillans in Transactions of the

Royal Society of South Africa 29: 351 (1 942).

3.3. Staberoha aemula (Kunth) Pillans in Transactions

of the Royal Society of South Africa 16: 386 (1928).

3.4. Staberoha cernua (L.f.) Dur. & Schinz, Conspectus

florae africae 5: 520 (1894).

3.5. Staberoha distachyos (Rottb.) Kunth , Enumeratio

plantarum 3 : 444 (1841).

3.6. Staberoha banksii Pillans in Transactions of the

Royal Society of South Africa 16: 385 (1928).

3.7. Staberoha multispicula Pillans in Journal of South

African Botany 18: 117 (1952).

3.8. Staberoha ornata Esterh. in Linder in Bothalia 15:

396 (1985).

3.9. Staberoha vaginata (Thunb.) Pillans in Transactions

of the Royal Society of South Africa 1 6: 384 ( 1 928).

4. Thamnochortus RJ.Rergms, Descriptiones plan-

tarum ex Capite Bonae Spei 353 (1767). Type: Thamno-

chortus fruticosus P.J.Bergius.

Plants caespitose or with spreading underground rhi-

zomes. Fertile culms round and, except in two species,

simple; sheaths persistent, closely convoluted, upper

half broadly chartaceous and soon becoming lacerated,

hyaline shoulders absent. Male inflorescence panicu-

late from one or several nodes, with up to 1 00 spikelets;

spathes persistent; spikelets generally lorate to oblong,

pendulous on flexible pedicels; bract margins like body

of bract, much taller than and obscuring papery male

flowers; anthers included in flowers at anthesis. Female

inflorescence racemose or somewhat paniculate, with 1-

100 erect, many-flowered, lorate to obtriangular spike-

lets; bracts taller than flowers with only stigmas visible

at anthesis. Female flowers with glabrous and smooth

tepals; outer lateral tepals strongly keeled or widely

winged; staminodes usually absent. Gynoecium with soli-

tary, white, plumose style; ovary unilocular, indehiscent.

Diaspore an oblong to completely round, wind-dispersed

structure; nut enclosed by persistent perianth. Culm

anatomy, epidermal cells single-layered, lateral walls

usually sinuose and thickened; parenchymatous layer of

1-4 cells; central ground tissue with scattered cavities;

tannin present in epidermis, parenchyma, sclerenchyma

or central ground tissue; silica mostly absent.

Etymology’'. Thamnochortus (m.): thamnos (Greek), bush,

shrub; chortus (Greek), green herbage, grass, fodder.

Notes: mostly associated with more arid areas, espe-

cially in the northwestern Cape, along the coastal plains

( Thamnochortus bachmannii ), Nieuwoudtville ( T. platyp-

teris), but also typical of the wettest fynbos vegeta-

tion (T. cinereus). Also noted for the important thatch-

ing plants ( T. insignis). The plants often form extensive

stands, and can locally dominate the vegetation. Striking

in some species are the widely winged, persistent peri-

anths, making for efficient wind dispersal of the seed.

This genus is very clearly demarcated, and is very

readily diagnosed by the slender, pendulous male spike-

lets, the closely convoluted sheaths with a decaying

upper half, the many-flowered, erect female spikelets,

and the single plumose style. The closest relative is

Rhodocoma, with which it shares features of the culm

anatomy and pollen structure, as well as the pendulous

male spikelets. However, Thamnochortus has an inde-

hiscent, unilocular ovary, whereas Rhodocoma has a

dehiscent, three-locular ovary. In many ways this rela-

tionship is similar to that between Chondropetalum and

Elegia s.str., but in the case of Thamnochortus there

are no morphological intermediates and there is strong

10

Bothalia 40,1 (2010)

molecular evidence for the reciprocal monophyly of the

two genera.

The biogeography and evolution in this genus was

investigated by Linder & Mann (1998). It is possible to

assign the species to five main groups, not all of which

are monophyletic (Figure 2).

a) Basal grade: includes the only two species with

branching culms:

4.1. Thamnochortus levynsiae Pillans in Transactions

of the Royal Society of South Africa 16: 364 ( 1928).

4.2. Thamnochortus pulcher Pillans in Transactions of

the Royal Society of South Africa 29: 350 (1942).

4.3. Thamnochortus nutans (Plumb.) Pillans in Transac-

tions of the Royal Society of South Africa 1 6: 365 ( 1 928).

4.4. Thamnochortus gracilis Mast, in A. DC., Mono-

graphiae phanerogamarum 1: 327 (1878).

b) Limestone clade: all species associated with lime-

stones or coastal dunes, except T. karooica :

4.5. Thamnochortus karooica H.P.Linder in South

African Journal of Botany 56: 450 (1990).

4.6. Thamnochortus muirii Pillans in Transactions of

the Royal Society of South Africa 16: 366 (1928).

4.7. Thamnochortus paniculatus Mast, in Botanische

Jahrbiicher 29, Beiblatt 66: 12 (1900b).

4.8. Thamnochortus spicigerus (Thunb.) Spreng ., Syste-

ma vegetabilium 1: 187(1824).

4.9. Thamnochortus fraternus Pillans in Transactions

of the Royal Society of South Africa 1 6: 370 ( 1 928).

4.10. Thamnochortus pluristachyus Mast, in Botanis-

che Jahrbiicher 29, Beiblatt 66: 12 (1 900b).

c) Big bract group: species with large, often silver-

coloured female bracts:

4.11. Thamnochortus cinereus H.P.Linder in Bothalia

15: 473 (1985).

4.12. Thamnochortus rigidus Esterh. in Linder in Both-

alia 15: 477 (1985).

4.13. Thamnochortus glaber (Mast.) Pillans in Transac-

tions of the Royal Society of South Africa 16: 363 (1928).

4.14. Thamnochortus papyraceus Pillans in Trans-

actions of the Royal Society of South Africa 29: 350

(1942).

4.15. Thamnochortus amoena H.P.Linder in South

African Journal of Botany 56: 451 (1990).

4.16. Thamnochortus acuniinatus Pillans in Trans-

actions of the Royal Society of South Africa 29: 349

(1942).

4.17. Thamnochortus fruticosus PJ.Bergius , Descrip-

tiones plantarum ex Capite Bonae Spei 353 (1767).

d) Thatching reed group: plants up to 2 m tall:

4.18. Thamnochortus erectus (Thunb.) Mast, in Journal

of the Linnean Society, Botany 14: 419 (1874).

4.19. Thamnochortus insignis Mast, in Gardeners’

Chronicle, ser. 3, 25: 242 ( 1 899).

e) Typical group: mostly plants < 1 m tall, diaspores

often round (as broad as long):

4.20. Thamnochortus guthrieae Pillans in Transactions

of the Royal Society of South Africa 16:371 (1 928).

4.21. Thamnochortus ellipticus Pillans in Transactions

of the Royal Society of South Africa 16: 368 (1928).

4.22. Thamnochortus lucens (Poir.) H.P.Linder in Bo-

thalia 15: 475 (1985).

4.23. Thamnochortus arenarius Esterh. in Linder in Bo-

thalia 15: 472 (1985).

4.24. Thamnochortus sporadicus Pillans in Journal of

South African Botany 18: 116 (1952).

4.25. Thamnochortus pellucidus Pillans in Journal of

South African Botany 18: 113 (1952).

4.26. Thamnochortus obtusus Pillans in Journal of South

African Botany 18: 112 (1952).

4.27. Thamnochortus stokoei Pillans in Transactions of

the Royal Society of South Africa 16: 375 (1928).

4.28. Thamnochortus dumosus Mast, in Botanische Jahr-

biicher29, Beiblatt 66: 11 (1900b).

4.29. Thamnochortus platypteris Kunth in Enumeratio

plantarum 3 : 429 (1841).

4.30. Thamnochortus schlechteri Pillans in Transactions

of the Royal Society of South Africa 16: 372 ( 1 928).

4.31. Thamnochortus bachmannii Mast, in Annalen

des Kaiserlich-Koniglichen Naturhistorischen Hofmuse-

ums, Wien 15: 11 (1900a).

4.32. Thamnochortus punctatus Pillans in Transactions

of the Royal Society of South Africa 16: 376 ( 1 928).

5. Rhodocoma Nees in Lindl., A natural system of

botany, edn 2: 450 (1836). Type: Rhodocoma capensis

Nees ex Steud.

Plants caespitose or with straight rhizomes. Fertile

culms round, unbranched or in R. capensis much branched

with branches whorled at each node; sheaths persistent,

upper half chartaceous and soon decaying, leaving a lac-

erated remnant; hyaline shoulders absent. Male inflores-

cence racemose from several nodes, each partial inflor-

escence with several to numerous pendulous spikelets

on flexible pedicels; spikelets ± elliptical; bracts without

distinct membranous upper margin; anthers included in

flowers at anthesis. Female inflorescence similar to male,

but partial inflorescences stiffly erect with few to several

spikelets; spathes sometimes prominent and membranous

or chartaceous; spikelets elliptical to ovate, with 1-5 flow-

ers; bracts shorter than flowers. Female flowers not later-

ally compressed; tepals bony, glabrous and smooth; stami-

nodes present. Gynoecium with 3, white to red, feathery

styles free to base; ovary 3-locular, dehiscent or rarely

falling with seed. Seed brown, grey or tan, colliculate or

rugose. Culm anatomy, epidermal cells 1 -layered, lateral

walls straight and unthickened or sinuose and thickened;

parenchyma of 1-M rows of cells; central ground tissue

with scattered cavities; tannin present in epidermis, paren-

chyma, sclerenchyma and central ground tissue; silica

usually absent, or when present, in parenchyma.

Etymology. Rhodocoma ( f . ) : rhodon (Greek), rosy;

koine (Greek), hair of the head; presumably referring to

the strikingly red or rosy styles in the type species, R.

capensis.

Notes : most of the species are found in the south-

ern Cape, where a clear pattern of ecological and geo-

graphical replacement is evident. Rhodocoma fruticosa

is widespread from the Cape to the Drakensberg, and is

very common over large areas along the eastern margins

of the fynbos.

Bothalia 40,1 (2010)

Superficially the species are similar to Thamnochor-

tus, especially because of the pendulous male spikelets

and the similar culm and sheath construction. However,

they can be separated by the dehiscent female ovaries,

by the short bracts in the females, and less confidently

by the shorter and more elliptical spikelets.

The evolution in this genus has been investigated

twice, first by Linder & Vlok (1991), then by Hardy &

Linder (2007).

5.1. Rhodocoma foliosa (N.E.Br) H.P.Linder &

C.R. Hardy, comb. nov.

Restio foliosus N.E.Br. in Flora capensis 7: 753 (1900). Type: Gar-

cia's Pass, Galpin 4783 (K, lecto.! [Linder in Bothalia 15: 478 (1985)];

B!, BOL!, K!).

5.2. Rhodocoma gigantea (Kunth) H.P.Linder in Both-

alia 15: 479 (1985).

5.3. Rhodocoma capensis Nees ex Steud., Synopsis

plantarum glumacearum 2: 249 (1855).

5.4. Rhodocoma gracilis H.P.Linder & Vlok in Plant

Systematics and Evolution 175: 156 (1991).

5.5. Rhodocoma arida H.P.Linder & Vlok in Plant Sys-

tematics and Evolution 175: 158 (1991).

5.6. Rhodocoma vleibergensis H.P.Linder, sp. nov.,

R. aridae affinis, sed caespitibus pauciculmis, spiculis femi-

neis longioribus (5-7 mm longis), stomatibus superficie-

bus notabilis.

TYPE. — Western Cape, 3319 (Worcester): Ceres, Bavi-

aanshoek, Vleiberg, (-BA), 14-10-1994, H.P.Linder 5882

(BOL, holo.; K, Z).

Plants tufted with few 1.0-1. 5 m tall culms and with

spreading rhizomes. Fertile culms unbranched, smooth,

olivaceous, apical diam. 1.5-2. 5 mm; sheaths closely con-

voluted, 40-60 mm long, apical margins broadly charta-

ceous and soon decaying, hyaline shoulders absent, api-

cally acute; mucro penicillate, straight and erect, 4-10 mm

long. Male inflorescence with up to 500 spikelets, panicu-

late, 80-300 x 30-50 mm; spathes persistent, chartaceous,

upper margins lacerated and largely decayed at anthesis,

shorter than spikelets; spikelets pendulous on flexible

pedicels longer than spikelets, elliptical, rounded, 4.5-

6.5 mm long, 5-7-flowered; bracts shorter than to as tall

as flowers, 1 .5-2.5 mm long, oblong or ovate, rounded or

obtuse, chartaceous. Male flower 2. 0-2. 5 mm long; tepals

all same size, cartilaginous, glabrous, outer lateral tepals

conduplicate; anthers 1. 2-1.4 mm long, included in flow-

ers, pistillode present. Female inflorescence with up to 500

spikelets, paniculate, sparsely branching, stiffly erect, 80-

200 x 10-30 mm; spathes persistent, chartaceous or mem-

branous, longer than spikelets but not obscuring them;

spikelets 1 -flowered, with 3-6 sterile bracts shorter than

flowers, 2-3 mm long, ovate, rounded or obtuse or acute,

chartaceous. Female flower 4. 5-6.0 mm long; tepals bony,

glabrous and smooth, midrib flush with tepal body, apices

acute, inner and outer whorls of same length; staminodes

present. Gynoecium: styles feathery, white, 3, free to base;

ovary with 3 dehiscent locules. Culm anatomy: epidermal

cells single-layered, lateral wall sinuose, thickened, outer

wall thickened, colliculate; glabrous, 1/w ratio 6 : 4, all the

same length; stomatal apparatus superficial, guard cells

seated on top of support cells; chlorenchyma of 2 layers

of cells, 1/w ratio 12:8, inner and outer layers somewhat

dissimilar; protective cells reaching to base of chloren-

chyma layer; parenchymatous layer of 2 or 3 cells, cells

smaller than epidermal cells; sclerenchyma ring of 9-13

layers, without protrusions, cell walls thick; central ground

tissue with scattered cavities; tannin present in epidermis,

sclerenchyma or central ground tissue; silica sand absent.

Flowering time : September, seed release in November.

Etymology ;: Vleiberg, a little known peak in the

Swartruggens, from which this new species was first

recorded.

Distribution and ecology: endemic to the western

margins of the Tanqua Karoo, from the Bonteberg at

Touws River, northwards via Baviaansberg, Vleiberg to

the Bokkeveld Sneeukop. The altitude is from 1 000-1

600 m, the habitat restricted to sandstone- or quart-

zite-derived soils, where populations are found in well-

drained habitats on rocky slopes. The plants are killed by

fire, and populations are re-established from seed.

Discussion: this species is morphologically very simi-

lar to Rhodocoma arida , but with a different 'gestalt' , the

plants being smaller, tattier looking, and spreading with

a more developed rhizome. The spikelets are generally

larger. This applies particularly to the female spikelets.

Furthermore, there is a geographical separation, with R.

vleibergensis found north of the Laingsburg Witteberg,

and R. arida east of this line. This also determines the pro-

portion of summer rain in the area, with R. arida receiving

a substantial proportion of its rain in summer, whereas R.

vleibergensis receives almost no summer rain. Despite the

morphological similarity to R. arida , the phylogeny of

Hardy & Linder (2007) indicates that the species is more

closely related to the R. alpina-R. fruticosa subclade.

Additional collections

WESTERN CAPE. — 3319 (Worcester): Ceres, Baviaanshoek,

Watervalsberg, between top of track and trig beacon, common on stony

soils, (-BA), 14-10-1994, H.P.Linder 5854 (Z, K); Bonteberg, Abra-

hamshoek, summit ridge on well-drained soils, (-BB), 25-10-2000,

H.P.Linder 7103 (Z).

5.7. Rhodocoma fruticosa (Thunb.) H.P.Linder in Both-

alia 15: 478 (1985).

5.8. Rhodocoma aipina H.P.Linder & Vlokm Plant Sys-

tematics and Evolution 175: 156 (1991).

6. Elegia L., Mantissa plantarum altera 162 (1771).

Type: Elegia juncea L.

Chondropetalum Rottb.: II (1772). Type: Chondrope-

talum deustum Rottb. [= Elegia deusta (Rottb.) Kunth],

lecto. [Linder 1984].

Dovea Kunth: 457 (1841). Type: Dovea macrocarpa

Kunth [= Elegia macrocarpa (Kunth) Moline & H.P.

Linder],

Lamprocaulis Mast, in A. DC.: 349 (1878). Type: Lamp-

rocaulis grandis (Kunth) Mast. [= Elegia grandis Kunth],

Plants tufted, clumped or mat-forming, often with

well-developed, spreading rhizomes. Fertile culms mostly

unbranched, in some species branched, and in two spe-

cies branches whorled at each node, mostly round, some-

times ± compressed; sheaths sometimes persistent, but

more commonly caducous with distinct abscission line.

12

Bothalia 40,1 (2010)

when present, most commonly loosely convoluted and

same texture over whole structure, rarely upper margins

membranous or otherwise different from rest of organ.

Male inflorescence of several clusters of partial, paniculate

inflorescences, each with several to very numerous flowers

only poorly organized into spikelets; young partial inflo-

rescences generally hidden behind large spathes, which

drop before anthesis; large spathe-like spathellae occasion-

ally present; male bracts usually shorter than perianth, but

in appearance similar to tepals. Male flowers with undif-

ferentiated, glabrous tepals; anthers included in flowers at

anthesis. Female inflorescence structurally similar to male

inflorescences; each partial inflorescence with 1-many

flowers, these poorly organized into spikelets; spathes

large, usually persistent, often completely obscuring flow-

ers at anthesis; spikelets often clustered in paniculate par-

tial inflorescences, making them difficult to delimit; bracts

mostly shorter than flowers with no differentiation between

body and margin, rarely acuminate and/or lacerated on

margins. Female flowers with undifferentiated, glabrous

tepals; staminodes usually present. Gynoecium with 2 or 3

mostly feathery styles free to base or more rarely seated on

a stylopodium; ovary with 3 usually dehiscent or 1 inde-

hiscent locule. Nuts generally triangular, often black, with-

out elaiosomes, perianth sometimes persistent on nuts but

never winged; seeds from dehiscent ovaries often brown

and smooth. Culm anatomy, epidermal cells almost always

in 2 layers, lateral wall straight, unthickened; parenchyma-

tous layer of 1-6 cells; central ground tissue solid or with

scattered cavities, rarely with a single, central cavity; tan-

nin usually in epidermis and central ground tissue, rarely

in parenchyma or sclerenchyma; silica absent.

Etymology. Elegia (f. ), from the Greek elegos, mean-

ing weeping or lament, and this was then used by Plinius

in the Latinized form of Elegia to refer to a kind of reed.

Notes', this genus contains a remarkably wide range of

variation in morphology, previously recognized as four

genera. However, this variation does not readily group

into reciprocally monophyletic groups, and consequently

the whole group is best treated as one large genus, as sug-

gested by Moline & Linder (2005). This larger clade is

readily diagnosed by anatomical attributes (epidermis in

two layers: Figure 1 ). The morphological attributes (cadu-

cous sheaths; paniculate inflorescences) have numerous

exceptions, which makes them more difficult to use.

The genus contains several well-known species, such

as Elegia tec to rum , widely cultivated, previously widely

used as a thatching reed; E. capensis , widely cultivated,

dominant in many areas in regions with some groundwa-

ter, where the stoloniferous growth allows the species to

take over the vegetation; E. filacea, which has an enor-

mous ecological range, and is found in almost all veg-

etation types.

The species are organized into four clades (Figure 3).

a) Elegia mucronata clade, without obvious charac-

ters:

6.1. Elegia cuspidata Mast, in Journal of the Linnean

Society, Botany 10: 240(1868).

6.2. Elegia mucronata (Nees) Kunth , Enumeratio plan-

tarum 3: 475 (1841).

b) Chondropetalum clade: includes most species pre-

viously included in Chondropetalum. Morphologically

this clade is defined by the dehiscent ovaries. However,

it also includes Elegia verreauxii , which has an indehis-

cent ovary:

6.3. Elegia nuda (Rotth.) Kunth , Enumeratio plantarum 3:

462(1841).

6.4. Elegia tectorum (L.f.) Moline & H.P.Linder in Sys-

tematic Botany 30: 772 (2005).

6.5. Elegia elephantina H.P.Linder , sp. nov., a E.

tectoro caespitibus elatioribus (1.0-2. 5 m), culmis valid-

ioribus (4-6 mm), spiculis maribus maioribus (3. 0-3. 5

mm diametro), tepalis femineis exterioribus interiores

aequantibus recedit.

TYPE. — Western Cape, 3318 (Cape Town): between

Hopefield and Koperfontein, 33° 05' 31"S, 18° 34' 36"E,

plants up to 2 m tall, forming small populations in hol-

lows among the dunes, (-AB), 22-08-2000, H.P.Linder

7072 (Z, holo.).

Evergreen, tufted plants, 1.0-2. 5 m tall. Rhizome

spreading, unbranched or sparsely branched, culms

evenly spaced. Fertile culms unbranched, round, smooth,

olivaceous, 4-6 mm diam. at base, 1.4-2. 5 mm diam. at

apex; sheaths caducous, with an abscission line present,

loosely convoluted, 30-60 mm long, margins entire, red-

dish to dark brown with fine tan speckles and tan margins,

slightly glossy; sheath mucro penicillate, straight and

erect, 6-10 mm long. Male inflorescence with more than

100 spikelets, paniculate, 80-500 x 20-30 mm; spathes

caducous, taller than spikelets, cartilaginous; spikelets

erect on short stiff pedicels, subglobose, 3. 0-3. 5 mm in

diam., 3- or 4-flowered; bracts shorter than flowers, ovate,

1.2- 1. 7 mm long, acute, margins entire, chartaceous or

cartilaginous, bract awn minute or absent. Male flower

2. 3- 2. 9 mm long, glabrous; tepals chartaceous or cartilag-

inous, inner tepals longer than outer tepals, outer lateral

tepals conduplicate; anthers included in flowers. Female

inflorescence with more than 50 spikelets, interrupted,

80^400 x 1 5-20 mm; spathes longer than spikelets, cadu-

cous, bony; spikelets subglobose, 3. 2^1.2 mm long, 4- or

5-flowered; bracts all fertile, shorter than flowers, erect,

1. 5-2.0 mm long, orbicular (at times wider than tall),

rounded or obtuse, cartilaginous (with a sturdy midrib),

apical margin like rest of bract, same as body of bract,

awn minute or absent. Female flowers 2.2-2. 8 mm long,

tepals bony, glabrous and smooth, midrib raised, margins

entire, apices acute, inner and outer whorls equally long;

outer lateral tepals conduplicate, odd outer tepal elliptical,

inner tepals ovate, 1 .8-2.3 mm long; staminodes present.

Gynoecium with 3 feathery styles, bases free; 3-locular,

dehiscent ovary. Seed 0.6-0. 8 x 0.3-0. 5 mm, shape in

side view elliptical, shape round in diam. and rectangular

in cross section, brown. Flowering time : July.

Distribution and ecologyr. endemic to the west coast of

South Africa, from Blouberg to Elands Bay. Local in damp

hollows and seepages on well-leached sand below 200 m.

Etymology, this species is known as olifantsriet on

the west coast. Unfortunately, this name appears to be

applied to any very large member of the Restionaceae,

thus is not very useful. The name is derived from the

Latin for elephant: elephas.

Bothalia 40,1 (2010)

13

1 00r

87

94

70

78

81

83

69

69

73

77

100'

87

89

100

100

1 0Or

100'

74

75

73

74

Askidiosperma esterhuyseniae

Askidiosperma rugosum

Askidiosperma chartaceum

Askidiosperma andreaeanum

66 | Askidiosperma alticolum

64 Askidiosperma paniculatum

Askidiosperma albo-aristatum

Askidiosperma nitidum

Askidiosperma insigne

Askidiosperma longiflorum

46 | Askidiosperma capitatum

50 Askidiosperma delicatulum

' Elegia mucronata

' Elegia cuspidata

94

28

29

100

100

100

16

94

98

' Elegia nuda

| Elegia elephantina

Elegia tectorum

' Elegia marlothii *

53

63

19

16

94

17

51

43

45

56

54

66

41

Elegia decipiens

Elegia hookeriana

Elegia aggregata

Elegia ebracteata

Elegia deusta

Elegia acockii

100

100

Elegia microcarpa

100| Elegia recta

100 Elegia verreauxii

60

37

43

60

' Elegia macrocarpa

99

94

94

99

98

81

81

98

25

25

1 00r

' Elegia esterhuyseniae

| Elegia capensis

Elegia hutchinsonii

' Elegia neesii

Elegia muirii

52 | Elegia grandis

32 Elegia prominens

80

89

100L

48

61

' Elegia galpinii

' Elegia thyrsoidea

' Elegia extensa

92

81

21

28

91

92

68

100.

1001

49

87

95

81

94

99

Elegia asperiflora

Elegia caespitosa

Elegia dregeana

Elegia intermedia

Elegia fastigiata

Elegia fucata

Elegia fistulosa

Elegia coleu ra

36 - -

39

39

33

39

62

94

100

66

83

43

35

100

53

79

75

76

75

Elegia racemosa

Elegia persistens

Elegia altigena *

99 | Elegia stipularis

1 00 Elegia stokoei

Elegia atratiflora

Elegia squamosa

Elegia fenestrata

28 j Elegia juncea

28 Elegia vaginulata

43 | Elegia equisetacea

44 Elegia filacea

FIGURE 3 . — Portion of strict consensus tree of total combined analysis of Hardy et at. (2008) containing Askidiosperma and Elegia sensu Moline

& Linder (2005). Bootstrap values above the branches indicate support based on a 500-replicate bootstrap analysis that included 20 species

for which DNA data were lacking (asterisked taxa). Bootstrap values below branches indicate support based upon a 500-replicate analysis

that included only taxa for which both DNA data and morphology data were available. Where no value is given below a branch, it indicates

that the clade did not appear in the analysis excluding morphology-only species.

14

Bothalia 40,1 (2010)

Discussion: this species is very close to Elegia tector-

um, from which it primarily differs in being bigger in

all dimensions. Furthermore, the inner and outer whorls

of the female tepals are the same length, whereas in E.

tectorum the inner tepals are longer than the outer. Both

species co-occur on the west coast at Blouberg, and no

intermediate plants have been seen.

6.6. Elegia marlothii (Pillans) Moline & H.PLinder in

Systematic Botany 30: 772 (2005).

6.7. Elegia hookeriana (Mast.) Moline & H.PLinder in

Systematic Botany 30: 772 (2005).

6.8. Elegia decipiens (Esterh.) Moline & H.PLinder in

Systematic Botany 30: 771 (2005).

6.9. Elegia ebracteata (Kunth) Moline & H.PLinder in

Systematic Botany 30: 771 (2005).

6.10. Elegia aggregata (Mast.) Moline & H.PLinder in

Systematic Botany 30: 771 (2005).

6.11. Elegia deusta (Rottb.) Kunth , Enumeratio planta-

rum 3: 460 (1841).

6.12. Elegia acockii (Pillans) Moline & H.PLinder in Sys-

tematic Botany 30: 771 (2005).

6.13. Elegia microcarpa (Kunth) Moline & H.PLinder

in Systematic Botany 30: 772 (2005).

6.14. Elegia recta (Mast.) Moline & H.PLinder in Sys-

tematic Botany 30: 772 (2005).

6.15. Elegia verreauxii Mast, in Journal of the Linnean

Society, Botany 21 : 589 (1885).

c) Lamprocaulis clade: includes most species with

branching culms, and which have, except for Elegia

macrocarpa, indehiscent nuts. In several species the

sheaths are persistent:

6.16. Elegia macrocarpa (Kunth) Moline & H.P.Linder

in Systematic Botany 30: 772 (2005).

6.17. Elegia capensis (Burmf.) Schelpe in Journal of South

African Botany 33: 156 (1967).

6.18. Elegia hutchinsonii Pillans in Transactions of the

Royal Society of South Africa 29: 344 ( 1942).

6.19. Elegia esterhuyseniae Pillans in Transactions of

the Royal Society of South Africa 30: 259 (1945).

6.20. Elegia neesii Mast, in Journal of the Linnean Socie-

ty, Botany 1 0: 246 ( 1 868).

6.21 . Elegia muirii Pillans in Transactions of the Royal

Society of South Africa 1 6: 3 1 9 ( 1 928).

6.22. Elegia grandis (Nees) Kunth , Enumeratio planta-

rum 3: 475 (1841).

6.23. Elegia prominens Pillans in Transactions of the Ro-

yal Society of South Africa 16: 320(1928).

d) Elegia clade: includes almost all species of the pre-

vious genus Elegia , characterized by the indehiscent ova-

ries and usually unbranched culms:

6.24. Elegia galpinii N.E.Br. in Flora capensis 7: 754

(1900).

6.25. Elegia thyrsoidea (Mast.) Pillans in Annals of the

Bolus Herbarium 3,3: 146 (1922).

6.26. Elegia extensa Pillans in Transactions of the Royal

Society of South Africa 29: 343 (1942).

6.27. Elegia asperiflora (Nees) Kunth , Enumeratio plan-

tarum 3: 474 (1841).

6.28. Elegia caespitosa Esterh. in Linder in Bothalia 15:

421 (1985).

6.29. Elegia dregeana Kunth , Enumeratio plantarum 3:

469(1841).

6.30. Elegia intermedia (Steud.) Pillans in Transactions

of the Royal Society of South Africa 16: 332 (1928).

6.31. Elegia fastigiata Mast, in Botanische Jahrbiicher

29, Beiblatt 66: 7 (1900b).

6.32. Elegia fucata Esterh. in Linder in Bothalia 15: 423

(1985).

6.33. Elegia fistulosa Kunth, Enumeratio plantarum 3: 467

(1841).

6.34. Elegia coleura Nees ex Mast, in A. DC., Monogra-

phiae phanerogamarum 1 : 358 ( 1 878).

6.35. Elegia rigida Mast, in Journal of the Linnean Soci-

ety, Botany 21: 587 (1885).

6.36. Elegia spathacea Mast, in Journal of the Linnean

Society, Botany 2 1 : 588 ( 1 885).

6.37. Elegia grandispicata H.P.Linder in Bothalia 15:

424(1985).

6.38. Elegia thyrsifera (Rottb.) Pers ., Synopsis planta-

rum 2: 607 (1807).

6.39. Elegia fenestrata Pillans in Transactions of the Ro-

yal Society of South Africa 16: 338 (1928).

6.40. Elegia persistens Mast, in Botanische Jahrbiicher

29 Beiblatt, 66: 8 (1900b).

6.4 1 . Elegia racemosa (Poir.)Pers. , Synopsis plantarum 2 :

607(1807).

6.42. Elegia amoena Pillans in Transactions of the Ro-

yal Society of South Africa 30: 258 (1945).

6.43. Elegia altigena Pillans in Transactions of the Ro-

yal Society of South Africa 30: 257 (1945).

6.44. Elegia equisetacea Mast, in Journal of the Linnean

Society, Botany 2 1 : 583 ( 1 885 ).

6.45. Elegia filacea Mast, in Journal of the Linnean So-

ciety, Botany 21: 589 (1885).

6.46. Elegia atratiflora Esterh. in Linder in Bothalia 15:

420(1985).

6.47. Elegia squamosa Mast, in Journal of the Linnean

Society, Botany 10: 244 ( 1868).

6.48. Elegia stipularis Mast, in Journal of the Linnean

Society, Botany 21: 587 (1885).

6.49. Elegia stokoei Pillans in Transactions of the Royal

Society of South Africa 29: 345 ( 1 942).

6.50. Elegia juncea L., Mantissa plantarum altera 297

(1771).

6.51. Elegia vaginulata Mast, in Journal of the Linnean

Society, Botany 21: 586 ( 1885).

7. Askidiosperma Steud., Synopsis plantarum glu-

macearum 2: 257 (1855). Type: Askidiosperma capita-

turn Steud.

Plants caespitose. Fertile culms unbranched, round;

sheaths dropping off, with an abscission line present,

loosely convoluted or flat and standing free from culm,

margins coriaceous, like rest of body, hyaline shoul-

ders absent. Male inflorescence with I to several partial

inflorescences subtended by large spathes; each partial

inflorescence with 1 to several, many-flowered spikelets;

bracts much taller than flowers, papery or membranous,

margins entire or lacerated. Male flowers not laterally

compressed; tepals glabrous, inner and outer often of

different lengths; anthers included in flowers at anthe-

sis. Female inflorescence , spikelets and flowers similar

to male. Female flowers with staminodes. Gynoecium :

styles 2 or 3, white, feathery, free to base; ovary 1-3-loc-

ular, dehiscent or sometimes dropping with seed. Seed

silvery or more commonly brown and smooth. Culm

Bothalia 40,1 (2010)

15

anatomy: epidermis with a single layer of cells, lateral

walls straight, unthickened; parenchyma of 1-3 layers

of cells; central ground tissue solid, rarely with a single,

central cavity; tannin usually absent, when present in

epidermis, sclerenchyma or (rarely) in central ground tis-

sue; silica usually absent, when present, in parenchyma

or sclerenchyma.

Etymology. Askidiosperma (n.): from Greek askion

or askidion , which is the diminutive of askos , which is

a bag, sac, wine skin or hide; and sperma (n., Greek),

which is a seed. The connection to the genus is unclear.

Notes', this genus shares with Elegia the caducous

sheaths, and the tepals which are all the same (outer lat-

eral tepals are not flattened and keeled). The species had

been included in Chondropetalum by Pillans ( 1928), but

were again separated by Linder (1984), as they differ

from Elegia by the culms with simple (not doubled-lay-

ered) epidermis, and by the floral bracts which are hya-

line-membranous and much taller than the flowers, as

well as by the phytochemical composition (Harborne et

al. 1985).

The species are mostly found in the western moun-

tains, rather rarely in the southern mountains of the

Western Cape, and are absent from the coastal flats

and plains. Species range from seepages over bedrock

( Askidiosperma insigne) to alluvial soils along streams

and on well-drained rocky slopes.

7.1. Askidiosperma chartaceum (Pillans) H.P.Linder in

Bothalia 15: 431 (1985).

7.2. Askidiosperma andreaeanum (Pillans) H.P.Linder

in Bothalia 15: 43 1 ( 1985).

7.3. Askidiosperma aiticolum (Esterh.) H.P.Linder in

Nordic Journal of Botany 21 : 198 (2001b).

7.4. Askidiosperma paniculatum (Mast.) H.P.Linder in

Bothalia 15: 432 (1985).

7.5. Askidiosperma esterhuyseniae (Pillans) H.P.Linder

in Bothalia 15: 432 (1985).

7.6. Askidiosperma rugosum Esterh. in Linder in Botha-

lia 15:432(1985).

7.7. Askidiosperma albo-aristatum (Pillans) H.P.Linder

in Bothalia 15: 431 ( 1985).

7.8. Askidiosperma nitidum (Mast.) H.P.Linder in Bo-

thalia 15: 432 (1985).

7.9. Askidiosperma insigne (Pillans) H.P.Linder in Bo-

thalia 15: 432 (1985).

7.10. Askidiosperma longiflorum (Pillans) H.P.Linder

in Bothalia 15: 432 (1985).

7.11. Askidiosperma capitatum Stead., Synopsis planta-

rum glumacearum. 2: 257 (1855).

7.12. Askidiosperma delicatulum H.P.Linder in Nordic

Journal of Botany 21: 196 (2001b).

Restio clade

This large clade is readily defined by the laterally

compressed female flowers, where the keels of the outer

lateral tepals are more or less villous. In most species

the culms are branching. This clade largely includes

most of the segregates of the genus Restio sensu Linder

(1984) and Calopsis sensu Linder (1984), as well as

Ischyrolepis. Several strongly supported subclades are

recognized: Craspedolepis, Calopsis s.str. , Restio s.str. ,

Locapsis and Ischyrolepis which are characterized by

different apomorphies and apomorphic tendencies:

Craspedolepis'. diagnosed by the bracts with hollow

(deeply concave) cells along the upper margin (honey-

comb cells), and the plants and spikelets tend to be

robust, the ovaries mostly dehiscent, with two locules.

In many species the spikelets are relatively massive.

Calopsis : highly paniculate inflorescences with very

many small spikelets; culms mostly angular or even

square.

Restio: sheaths mostly with large membranous lobes

flanking the mucro; the plants and spikelets tend to be

small, the ovary mostly unilocular and often indehis-

cent.

Locapsis: sheaths mostly spreading and without a dif-

ferentiated apical margin; perianth papery, rounded;

diaspore a nut.

Ischyrolepis: ovaries with two styles which are basally

fused, this fused portion persisting as a peg on the

dehisced ovary.

Varirestio and Eremorestio , however, lack characters,

and are recognized in order to retain monophyly.

Morphologically, Eremorestio would fit with Locap-

sis, while Varirestio could be polymorphic. It remains

possible that the apparent monophyly of these two segre-

gates is an artifact of ancient hybridization, and that the

plastid phylogeny is not tracking the species phylogeny.

However, the phylogenetic structure within the clade

is unclear. There is little molecular support for the group-

ings, and we cannot postulate good morphological char-

acters for the various molecular clades.

Also included in this clade are Pendulostemon and

Simplicaulos. The balance of the evidence places these

two segregates into the Staberoha-Elegia clade, but taxo-

nomically they are better kept in the large Restio s.lat.

8. Restio Rottb., Descriptiones plantarum rariorum

9 (1772). Type: Restio triticeus Rottb. (lecto. McVaugh

1968).

Calopsis Beauv. ex Desv.: 44, t. 3 (1828). Type:

Calopsis paniculata (Rottb.) Desv. (= Restio paniculatus

Rottb.).

Ischyrolepis Steud.: 249 (1855). Type: Ischyrolepis

subverticellatus Steud. (= Restio subverticellatus (Steud.)

Mast.).

Craspedolepis Steud.: 264 (1855). Type: Craspedole-

pis verreauxii Steud. (= Restio filiformis Poir.).

Plants caespitose, clumped, tangled or mat-form-

ing. Fertile culms simple or branching, round, or rarely

square or compressed; sheaths persistent, closely or

loosely convoluted, usually with a narrow membranous

margin, often with tall, acute membranous lobes flank-

ing mucro. Male and female inflorescences mostly very

similar; mostly paniculate, more rarely racemose, with

1 to numerous spikelets; spathes usually persistent;

16

Bothalia 40,1 (2010)

bracts usually coriaceous, varying from shorter to taller

than flowers; anthers exserted from flowers at anthesis.

Female inflorescence occasionally with fewer spikelets

than males; spikelets with up to 30 laterally compressed

flowers; tepals chartaceous or cartilaginous, glabrous

or sparsely (rarely densely) villous on keels of lateral

sepals; staminodes present. Gynoecium with 1, 2 or 3

feathery styles free to base, fused into a basal peg, or

rarely seated on a stylopodium; ovary 1-, 2- or 3-locular,

either dehiscent or indehiscent. Diaspore either a seed

or a soft-walled nut; seed variously coloured, usually

smooth, occasionally colliculate; nut dropped with a per-

sistent papery perianth that is sometimes winged. Culm

anatomy, epidermal cells 1 -layered, lateral walls straight

and unthickened or sinuose and thickened; parenchyma-

tous layer of up to 6 cells; central ground tissue either

with no cavity, a single central cavity, or with scattered

cavities; tannin rarely absent, when present then in the

epidermis, sclerenchyma, or central ground tissue; silica

either absent, or when present, in chlorenchyma or paren-

chyma.

Etymology. Restio (m., Latin), a ropemaker; it is

unclear why Linnaeus referred to restios as such, since

they are not to our knowledge useful for the making of

ropes, due to the brittle nodes.

The extensive variation in the genus is best expressed

in the eight subgenera recognized.

8.1. Restio subgen. Simplicaulos H.P.Lincier &

C.R. Hardy, subgen. nov., primo adspectu Restioni, Cras-

pedolepi, Calopsi, Locapsi, Varirestioni et Eremorestioni

culmis rotundis, vaginis persistentibus, spiculis maribus

erectis, staminodiis praesentibus, stylis libris maxime

simile, sed a speciebus olim ad subgenera Calopsem et

Restionem ascriptis culmis simplicibus et inflorescentiis

maribus et femineis valde similibus differt.

TYPE. — Restio bifidus Thunb.

Plants usually tufted. Fertile culms mostly unbranched

or rarely sparsely branched, round or very rarely com-

pressed (Restio subtilis ); sheaths persistent, closely

or loosely convoluted or rarely flat and standing free

from culm (R. papyraceus ); apical sheath margins nar-

rowly or broadly membranous; hyaline shoulders absent

or variously developed. Male inflorescence racemose,

with 1-20 spikelets, paniculate with pendulous spike-

lets only in R. bifarius', spathes persistent; bracts with

upper margin like body of bract, sometimes with hollow

cells, obtuse to acuminate and recurved; anthers usually

exserted from flowers at anthesis. Female inflorescence

always racemose with 1-5 variously shaped, 1-40-flow-

ered spikelets; bracts shorter to much longer than flow-

ers, apical margin like rest of bract, membranous or with

honeycombed cells, imbricate and obscuring spikelet

axis. Flowers with outer lateral tepals usually condu-

plicate, but in three species, undifferentiated; all tepals

cartilaginous, glabrous or keels of lateral sepals villous;

staminodes present. Gynoecium with I, 2 or 3 white,

feathery, free styles; ovary with 1-3 dehiscent or rarely

indehiscent locules. Fruit: in species with indehiscent

ovaries, perianth winged, persistent around ovary, in spe-

cies with dehiscent ovaries, seed silvery, brown or grey;

surface pitted, colliculate or striate. Culm anatomy : epi-

dermal cells in 1 layer, lateral wall straight, unthickened

or rarely sinuose and thickened; parenchymatous layer of

1-3 cells; central ground tissue solid or rarely with scat-

tered cavities or with single, central cavity; tannin absent

or present in epidermis, sclerenchyma or central ground

tissue; silica absent.

Etymolog y: Simplicaulos (m.): simplex (Latin), sim-

ple, unbranched; caulos (Greek), stem; referring to the

usually unbranched culms in this subgenus.

Notes: many species in this subgenus are associated

with impeded drainage, and Restio miser and R. confusus

are often dominants in convex-domed seepages on shale

bands and on sandstone. Restio bifidus is typical of shal-

low seepages, and can be dominant over much of the

upper plateau of Table Mountain. However, many of the

other species are found in well-drained habitats.

There are very distinct morphological groups in this

subgenus. The first group has large, almost flat, bracts

( Restio bifarius, R. bifidus, R. nuwebergensis and R.

papyraceus ), that are quite similar to the female spikelets

of the possible sister clade Staberoha. The second group

has small button-like spikelets, borne on fine-stemmed

tussocky plants, often with only one spikelet per culm,

and round female flowers in which the lateral tepals are

not conduplicate (R. subtilis, R. confusus, R. miser). The

third group has thick spindle-shaped spikelets, and nut-

lets with a persistent perianth forming a wing ( R . mon-

ostvlis, R. esterhuyseniae ). Finally, R. echinatus and R.

pedicellatus have acuminate bracts.

This subgenus lacks any striking synapomorphies.

However, optimization on the phylogeny shows that

the subgenus ancestrally had simple culms, and that

branched culms evolved only in the Restio bifarius

group. Even in this group, branching is often absent,

and when present, is sparse. The unbranched culms are

shared with its possibly nearest relatives, Elegia and Sta-

beroha, but differentiate the subgenus from the rest of

the genus Restio. Although it is easy to differentiate sub-

gen. Simplicaulos from its putative phylogenetic relatives

Staberoha or Elegia by the persistent sheaths, by stami-

nodes still present in the female spikelets, the (mostly)

erect male spikelets and the male flowers aggregated into

clear spikelets, these are all plesiomorphic features. It is

remarkable how variable the subgenus is regarding pre-

viously used generic characters, such as the number of

locules and styles and the type of diaspore (nut or seed).

Not surprisingly, subgen. Simplicaulos contains spe-

cies previously placed in Restio sensu Linder 1985 and

Ca/opsis sensu Linder 1985.

Although the clade is strongly supported, its position

within the Restioneae is not. The plastid data suggests

(weakly) a phylogenetic relationship to Staberoha or

Elegia, but the possibility that it is related to Restio s.lat.

cannot be rejected, even by plastid data.

8.1.1. Restio pedicellatus Mast, in Journal of the Lin-

nean Society, Botany 8: 252 ( 1865).

8.1.2. Restio echinatus Kunth, Enumeratio plantarum 3:

384(1841).

8.1.3. Restio stereocaulis Mast, in Botanische Jahrbii-

cher 29 Beiblatt 66: 1 ( 1900b).

Bothalia 40,1 (2010)

17

8. 1 .4. Restio subtilis Nees ex Mast, in Journal of the Lin-

nean Society, Botany 8: 251 (1865).

8.1.5. Restio confusus Pillans in Journal of South Afri-

can Botany 18: 103 (1952).

8.1.6. Restio miser Kunth, Enumeratio plantarum 3: 392

(1841).

8.1.7. Restio distylis H.P.Linder & C.R. Hardy, nom.

nov. pro Leptocarpus esterhuyseniae Pillans in Transac-

tions of the Royal Society of South Africa 30: 260 (1945).

Calopsis esterhuyseniae (Pillans) H.P.Linder: 466 (1985).

Type: Mitchell’s Pass, Slab Peak, Esterhuysen 6211 (BOL,

holo.!; K!), non Restio esterhuyseniae Pillans.

Named for the unusual condition of having two styles.

8.1.8. Restio monostylis (Pillans) H.P.Linder &

C.R. Hardy, comb. nov.

Leptocarpus monostylis Pillans in Journal of South African Botany

18: 108 (1952). Calopsis monostylis (Pillans) H.P.Linder: 469 (1985).

Type: south slopes of the Langeberg near Riversdale, Esterhuysen

16993 (BOL, holo.!).

8.1.9. Restio papyraceus Pillans in Transactions of the

Royal Society of South Africa 29: 342 (1942).

8.1.10. Restio nuwebergensis Esterh. in Linder in

Bothalia 15: 453 (1985).

8.1.11. Restio bifarius Mast, in Journal of the Linnean

Society, Botany 10: 278 (1868).

8.1.12. Restio bifidus Thunb., Phytographische Blatter,

Gottingen 1 : 7 (1803).

8.2. Restio subgen. Pendulostemon H.P.Linder &

C.R. Hardy, subgen. nov., primo adspectu Restioni, Cras-

pedolepi, Calopsi, Locapsi, Varirestioni et Eremorestioni

culmis rotundis, ramificantibus, vaginis persistentibus,

staminodiis praesentibus, stylis libris maxime simile, sed

a speciebus olim ad subgenera Restionem vel Calopsem

ascriptis combinatione spiculorum marium pendulorum

cum tepalis femineis glabris spiculisque femineis gran-

dibus striis longitalibus instructis differt.

TYPE. — Restio micans Nees.

Plants tufted or clumped. Fertile culms branching,

round; sheaths persistent, closely convoluted, margins

similar to body of bracts or narrowly membranous. Male

inflorescence paniculate, with 6-50 pendulous spikelets;

bracts ± leathery, overtopping flowers. Female inflores-

cences usually racemose, with up to 20 erect, elliptical

or obovate, rounded to acute, several-flowered spikelets;

bracts at least as tall as flowers, imbricate, usually with

longitudinal striations, margins like body or narrowly

membranous. Female flowers laterally compressed; tepals

bony or cartilaginous, glabrous, outer lateral tepals usu-

ally conduplicate and keeled; staminodes present. Gyn-

oecium with 3, white, feathery, free styles; ovary with 2

locules, dehiscent. Seed silvery or pink, smooth or col-

liculate. Culm anatomy, epidermis single-layered, epider-

mal lateral cell walls straight, unthickened; parenchyma

1-5-layered; central ground tissue solid or with single,

central cavity; tannin found in epidermis or sclerenchyma

or central ground tissue; silica absent or present in paren-

chyma.

Etymology’: Pendulostemon (m.): pendu/o (Latin), pen-

dulous; stemon (Greek), stamen; referring to the pendu-

lous male spikelets.

Notes', the two species of this subgenus mentioned

below were previously included in Restio s.str. because

of their dehiscent ovaries, laterally flattened flowers

and branched culms. Their current position is uncertain,

but plastid data weakly indicate a relationship to Tham-

nochortus and Rhodocomct (Ligure 2) with which they

share the pendulous male spikelets. However, a closer

relationship to Restio s.str. cannot be rejected.

Although the two species had not previously been asso-

ciated, they share a number of morphological attributes.

The plants are rather untidy, erect tussocks with robust,

erect, branching culms. The sheaths are coriaceous with-

out a substantial membranous margin. The spikelets, both

male and female, are spindle-shaped and relatively large.

The male spikelets are pendulous. The only unique feature

is the longitudinal striation on the bracts, but this feature

is not always easy to observe. Restio micans is a rare spe-

cies on coastal sands, R. egregius is more widespread in

the wetter mountains of the Western Cape.

8.2.1. Restio egregius Hochst. in Krauss in Flora 28:

337 (1845).

8.2.2. Restio micans Nees in Linnaea 5: 649 (1830).

8.3. Restio subgen. Craspedolepis (Steud.) H.P.Lin-

der & C.R. Hardy, stat. nov.

Craspedolepis Steud., Synopsis plantarum glumacea-

rum 2: 264 (1855). Type: Craspedolepis verreauxii Steud.

(= Restio filiformis Poir.).

Plants caespitose to tangled, without spreading rhi-

zomes. Fertile culms mostly branching, round or rarely

compressed; sheaths persistent, usually closely convo-

luted, apical margin ± membranous, sometimes with large

membranous shoulders flanking rnucro. Male inflores-

cence racemose or paniculate, with up to 10 (rarely more)

usually erect, mostly elliptical, spikelets (2 spp. with pen-

dulous spikelets); male bract with upper margin usually

with honeycombed cells, which eventually decay, taller

than flowers; anthers exserted from flowers at anthesis.

Female inflorescence similar to males, but spikelets are

somewhat bigger, mostly elliptical, with up to 40 flow-

ers; female bracts imbricate, shorter to taller than flowers,

apical margin with honeycombed cells. Female flowers :

outer tepals differentiated into conduplicate lateral tepals

and a flat odd tepal, rarely glabrous and smooth, usually

densely villous on keels of conduplicate tepals, occasion-

ally also on back of outer odd tepal; staminodes present.

Gynoecium with 3, white, feathery styles free to base;

ovary usually with 2, rarely 3 or 1, fertile locules, dehis-

cent. Seed variously coloured, surface smooth or collicu-

late. Culm anatomy, epidermal cells 1 -layered, lateral

wall straight and unthickened or sinuose and thickened;

parenchyma up to 4-layered; central ground tissue usually

solid; tannin usually present in epidermis, sclerenchyma

and central ground tissue; silica usually absent.

Etymology. Craspedolepis (f.): kraspedon (Greek),

edge, border or fringe; lepis (Greek), scale. This name

may be interpreted to refer to the upper margin of the

floral bracts (scales) which differ from the body of the

bract by the honeycombed cells, and are thus bordered.

Notes', this is a very distinctive segregate from Res-

tio s.str. The most notable synapomorphy are the hollow

18

Bothalia 40,1 (2010)

cells (honeycomb cells) that form the upper margins of

the floral bracts. This is absent from one species in the

subgenus, and similar structures are also found in a few

species in two other subgenera. The other characters are

more in the nature of ‘tendencies’: large, elliptical spike-

lets (where Restio insignis and R. strobilifer are extreme

examples); very hairy tepals, with often all three outer

tepals hairy along the keels; almost always with two

fertile locules. This contrasts with Restio s.str. with its

tendency to unilocular, indehiscent or tardily dehiscent

ovaries.

The subgenus is widespread especially in the west-

ern half of the Cape Floristic Region (CFR), and is

almost absent from the eastern parts of the CFR. There

is a remarkable range of seed surface ornamentation,

from colliculate to smooth. The inflorescence structure

is also highly variable, from solitary spikelets, to many-

spikeletted, paniculate structures.

The species are organized into several poorly sepa-

rated subgroups (Figure 4).

a) Restio fusiformis group: seed triangular in cross

section, either smooth or colliculate. The female spike-

lets are rather diverse in appearance:

8.3.1. Restio inveteratus Esterh. in Linder in Bothalia

15:450 (1985).

8.3.2. Restio obscurus Pillans in Transactions of the

Royal Society of South Africa 29: 341 (1942).

8.3.3. Restio fusiformis Pillans in Journal of South Afri-

can Botany 18: 104(1952).

8.3.4. Restio acockii Pillans in Transactions of the

Royal Society of South Africa 29: 339 ( 1 942).

8.3.5. Restio praeacutus Mast, in Flora capensis 7: 84

(1897).

8.3.6. Restio rupicola Esterh. in Linder in Bothalia 15:

459 (1985).

b) Restio perseverans group: inner male tepals shorter

than the outer; seed triangular in cross section, surface

ornamentation smooth:

8.3.7. Restio perseverans Esterh. in Linder in Both-

alia 15: 457 (1985).

c) Restio aureolas group: female flowers only sparsely

hairy, seeds planoconvex in cross section, surface orna-

mentation smooth. This includes R. occultus, with

numerous few-flowered spikelets in paniculate inflores-

cences and richly branched plants.

8.3.8. Restio aureolus Pillans in Transactions of the

Royal Society of South Africa 30: 246(1945).

8.3.9. Restio perplexus Kunth , Enumeratio plantarum 3:

406(1841).

8.3.10. Restio pulvinatus Esterh. in Linder in Bothalia

15: 457 (1985).

8.3.1 1 . Restio cymosus (Mast.) Pillans in Annals of the

Bolus Herbarium 3,2: 85 ( 1 921 ).

8.3.12. Restio capillaris Kunth, Enumeratio plantarum 3:

405 (1841).

8.3.13. Restio patens Mast, in Flora capensis 7:97

(1897).

8.3.14. Restio brachiatus (Mast.) Pillans in Annals of the

Bolus Herbarium 3,2: 85 ( 1921 ).

8.3.15. Restio occultus (Mast.) Pillans in Transactions

of the Royal Society of South Africa 16: 243 (1928).

d) Restio filiformis group: seed triangular in cross sec-

tion, surface ornamentation colliculate. In this group the

female spikelets are compact, globose to elliptical, often

rather few:

8.3. 16. Restio filiformis Poir. in Lam., Encyclopedic me-

thodique. Botanique 6: 173 (1804).

8.3.17. Restio brunneus Pillans in Transactions of the

Royal Society of South Africa 16: 250 (1928).

8.3.18. Restio burchellii Pillans in Transactions of the

Royal Society of South Africa 29: 340 ( 1 942).

8.3.19. Restio insignis Pillans in Transactions of the

Royal Society of South Africa 30: 251 (1945).

8.3.20. Restio pachystachyus Kunth , Enumeratio planta-

rum 3: 399 (1841).

8.3.21. Restio strobilifer Kunth, Enumeratio plantarum

3: 398 (1841).

8.3.22. Restio nodosus Pillans in Transactions of the Ro-

yal Society of South Africa 30: 252 (1945).

8.3.23. Restio bifureus Nees ex Mast, in Journal of the

Linnean Society, Botany 8: 247 (1865).

8.3.24. Restio bolusii Pillans in Transactions of the

Royal Society of South Africa 16: 247 (1928).

8.4. Restio subgen. Varirestio H.P.Linder & C.R.

Hardy, subgen. nov., primo adspectu Restioni, Crasped-

olepi, Calopsi, Locapsi et Eremorestioni culmis rotundis,

ramificantibus, vaginis persistentibus, spiculis maribus

erectis, staminodiis praesentibus, stylis 3, libris, ovario

loculis 1 vel 3, dehiscentibus maxime simile, sed vaginis

interdum mucronibus pro ratione longis (longioribus

quam dimidia pars vaginae) munitis differt et praeterea

positio topologica recognitionem subgeneris cogit.

TYPE. — Restio debilis Nees.

Plants caespitose. Fertile culms branching or rarely

simple, round; sheaths persistent, closely to very loosely

convoluted; apical margins undifferentiated or forming

membranous lobes; mucro sometimes as long as body of

sheath. Male inflorescences racemose to paniculate, with

up to 10 erect spikelets; spathes often overtopping spike-

lets; bracts with upper margins sometimes with honey-

combed cells; anthers exserted from flowers at anthesis.

Female inflorescence basically similar to male, but with

fewer and larger spikelets; spikelets oblong, elliptical,

ovate or obovate, with up to 20 flowers; bracts shorter or

longer than flowers, apical margin sometimes with honey-

combed cells. Female flowers laterally compressed;

outer lateral tepals conduplicate, outer odd tepal flat,

laterals either glabrous or villous; staminodes present.

Gynoecium : with 3, white, feathery styles, free to base;

ovary with 1 or 3 dehiscent locules. Seed variously col-

oured, smooth or colliculate. Culm anatomy, epidermal

cells 1 -layered, lateral wall straight and unthickened or

more rarely sinuose and thickened; parenchyma 1- or

2-layered; central ground tissue solid or with a single,

central cavity; tannin, when present, found in epidermis,

sclerenchyma or central ground tissue; silica absent.

Etymology : Varirestio (m.): vari (Latin), various, varia-

ble; restio (Latin), ropemaker; the name reflects the

extremely heterogeneous nature of this subgenus.

Bothalia 40,1 (2010)

19

‘ Restio inveteratus

' Restio obscurus

‘ Restio fusiformis

" Restio acockii

53 i Restio praeacutus

‘ Restio rupicola *

‘ Restio perseverans

‘ Restio aureolus

' Restio perplexus

‘ Restio pulvinatus

' Restio cymosus

' Restio capillaris

36 I Restio patens

I 29 | Restio brachiatus

32 Restio occultus

‘ Restio fitiformis

Restio brunneus

Restio burchellii

Restio insignis

Restio pachystachyus

Restio strobilifer

Restio nodosus

Restio bifurcus

Restio bolusii

' Restio debilis

‘ Restio distans

‘ Restio ievynsiae (Calopsis levynsiae)

' Restio rigidus (Calopsis rigida)

" Restio paniculatus (Calopsis paniculata)

' Restio quadratus

‘ Restio tetragonus

‘ Restio distichus

' Restio puicher (Calopsis pulchra)

‘ Restio harveyi

" Restio asperus (Calopsis aspera)

' Restio parvispiculus (Calopsis membranacea)

' Restio hyalin us (Calopsis hyalina)

' Resto impolitus (Calopsis impolita)

‘ Restio versatilis

' Restio strictus

' Restio triticeus

' Restio verrucosus

' Restio pumilus

' Restio zwartbergensis

'Restio villosus (Calopsis sparsa) *

' Restio clandestinus (Calopsis clandestina)

' Restio nudiflorus (Calopsis nudiflora)

' Restio pillansii

' Restio corneolus

57 | Restio saroclados

sr 1 48 i Restio ejuncidus

40

‘ Restio raws

' Restio degenerans

3 | Restio singularis *

1 5 i Restio alticola

' Restio scaber ’

" Restio montanus *

‘ Restio multiflorus

Restio arcuatus

Restio inconspicuus

Restio vallis-simius

Restio secundus

Restio decipiens

46 | Restio fragiiis

M I 44 j Restio coiiicuiospermus

45 Restio implicatus

Restio festuciformis

Restio zuiuensis

Restio dodii var. dodii

Restio dodii var, purpurascens

Restio dispar

Restio purpurascens

Restio ieptostachyus

Restio subgen. Craspedoiepis

Restio subgen. Varirestio

Restio subgen. Eremorestio

Restio subgen. Calopsis

Restio subgen. Restio

FIGURE 4. — Portion of strict consensus tree of total combined analysis of Hardy el al. (2008) containing Restio subgen. Craspedoiepis , R. subgen.

Varirestio, R. subgen. Eremorestio, R. subgen. Calopsis, and R. subgen. Restio. Names in parentheses are the now synonymous names of

Linder (2001a) when different from the ones here proposed. Bootstrap values above the branches indicate support based on a 500-replicate

bootstrap analysis that included 20 species for which DNA data were lacking (asterisked taxa). Bootstrap values below branches indicate

support based upon a 500-replicate analysis that included only taxa for which both DNA data and morphology data were available. Where no

value is given below a branch, it indicates that the clade did not appear in the analysis excluding morphology-only species.

20

Bothalia 40,1 (2010)

Notes: recognition of this subgenus is forced by its

topological position, sister to distinctive subgenera such

as Ischyrolepis, Locapsis, Restio and Calopsis. Its four

species appear not to share any unique attributes, and are

morphologically very divergent. There is no morphologi-

cal support for the recognition of this clade. Of the four

species, two have three locules and two just one; two

species have exceptionally long mucros, the others short

mucros. Similarly there is no homogeneity in the anat-

omy or the seed colour or ornamentation. Nor is there a

pattern of variation that would allow the recognition of

two diagnosable taxa.

8.4.1. Restio quinquefarius Nees in Linnaea 5: 639

(1830).

8.4.2. Restio similis Pillans in Annals of the Bolus Her-

barium 3,2: 82 ( 192 1 ).

8.4.3. Restio debilis Nees in Linnaea 5: 641 ( 1830).

8.4.4. Restio distans Pillans in Transactions of the Ro-

yal Society of South Africa 30: 247 (1945).

8.5. Restio subgen. Calopsis (Beauv. ex Desv.) H.P

Linder & C.R. Hardy, stat. nov.

Calopsis Beauv. ex Desv., Annales des Sciences Natu-

relles 13: 44, t. 3 (1828). Type: Calopsis paniculata

(Rottb.) Desv. (= Restio paniculatus Rottb.).

Large plants with large rhizomes. Fertile culms

branching, square or at least in upper parts semicircular,

always somewhere angular; sheaths persistent, with no

abscission line, closely convoluted, apical margins either

coriaceous like rest of body, or with large acute hya-

line shoulders flanking the penicillate mucro. Male and

female inflorescence very similar, racemose to panicu-

late, with very many partial inflorescences terminating

apical part of fertile shoot, each with 1 to many ellipti-

cal to oblong, many-flowered spikelets. Male flowers

with anthers exserted at anthesis. Female spikelets with

up to 14 flowers; bracts shorter than, to as long as, flow-

ers, margins not differentiated; outer lateral tepals con-

duplicate, keels glabrous to sparsely villous; staminodes

present. Gynoecium with 3, white, feathery styles, free to

base; ovary with 1-3 locules, indehiscent or dehiscent, or

sometimes dropping with seed. Seed brown and smooth.

Culm anatomy, epidermal cells single-layered, lateral

walls straight and unthickened; parenchymatous layer of

1 or 2 cells; central ground tissue solid or with single,

central cavity; tannin present in epidermis, parenchyma,

sclerenchyma, or central ground tissue; silica present in

parenchyma or central ground tissue.

Etymology. Calopsis (f.): calos (Greek), beautiful;

opsis (Greek), aspect; they are quite striking plants, con-

sequently the generic name Calopsis (= looking beauti-

ful) is highly appropriate.

Notes : this is a dramatically redefined concept of Calop-

sis, as it excludes the whole genus as currently recognized

except the type species, and includes two species formerly

placed in Restio. However, the three species share a very

similar growth form: one can think of them as being huge

paniculate inflorescences of which the lower portion is

sterile. Segregating them into separate genera was based

on an over-reliance on the ovary dehiscence ( Restio with

capsules, Calopsis with nuts). Furthermore, in all three

species the culms are more or less angular, and in two of

them they are sharply square. All three species are asso-

ciated with ground water, and C. paniculata is a typical

streambank species, widespread in the CFR.

8.5.1. Restio paniculatus Rottb., Descriptions plantarum

rariorum 10 (1772).

8.5.2. Restio quadratus Mast, in Journal of the Linnean

Society, Botany 10: 277 (1868).

8.5.3. Restio tetragonus Thunb., Dissertatione de Res-

tione: 17 ( 1788).

8.6. Restio subgen. Eremorestio H.P.Linder & C.R.

Hardy, subgen. nov., Calopsi sensu Linder (1984) vagi-

nis persistentibus, antheris exsertis, ovariis uniloculari-

bus indehiscentibus affinis, sed combinatione vaginorum

arete convolutorum, cum nuce triquetra, 2-3 mm longa

differt.

TYPE. — Calopsis rigida (Mast.) H.P.Linder.

Plants tufted or mat-forming. Fertile culms branch-

ing, round; sheaths persistent, closely convoluted, upper

half more membranous than lower half, soon decaying.

Male inflorescence racemose or sparsely paniculate with

2-20 ± elliptical spikelets; spathes persistent, sometimes

taller than spikelets; bracts acute to acuminate, slightly

taller than flowers, apices somewhat darker than bod-

ies; anthers exserted from flowers at anthesis. Female

inflorescence similar to males, but always racemose

and with fewer, somewhat larger spikelets; bracts taller

than flowers, bony or cartilaginous, acute to acuminate,

apices recurved somewhat from spikelet, giving it a

bristly appearance; flowers laterally compressed; tepals

chartaceous, rounded, glabrous or keels of lateral sepals

sparsely villous; staminodes present. Gynoecium with

3, white, feathery styles free to base; ovary unilocu-

lar, indehiscent. Diaspore a soft-walled nut enclosed in

perianth, triangular, 2-3 mm long, perianth not winged.

Culm anatomy, epidermal cells 1 -layered, lateral wall

straight, unthickened; parenchymatous layer of 2 or 3

cells; central ground tissue solid; tannin, when present,

in epidermis or central ground tissue; silica absent.

Etymology n Eremorestio (m.): Eremia (Greek), desert;

Restio (Latin), ropemaker; restios living in arid areas,

referring to the habitat of the two species in this genus,

occurring along the arid margins of the Cape Floristic

Region.

Notes: similar to the situation in Varirestio, this sub-

genus of two species is recognized in order to be able

to separate the larger subgenera Restio s.str. and Ischy-

rolepis. Contrary to the situation in Varirestio , however,

there are numerous similarities between the two species.

They have similar diaspores (soft-walled nuts dispersed

in a papery perianth). Such diaspores defined the genus

Calopsis sensu Linder 1984. The papery perianth and

rather brittle bracts are also typical of this group. Mor-

phologically the genus is very close to Locapsis, and it

is not clear whether monophyly would be significantly

rejected if the two were combined. The morphological

distinction between Eremorestio and Locapsis is not easy,

as most of the characteristics of Eremorestio are also

found in the larger subgen. Locapsis. Possibly the best

attribute could be that the tips of the floral bracts in the

Bothalia 40,1 (2010)

21

two species of Eremorestio are darker than the bodies of

the bracts, and these tips are recurved away from the axis

of the spikelet. In Locapsis the bract tips are generally

the same colour as the body of the bracts, and the tips are

closely rolled around the spikelets, resulting in slender

spindle-shaped spikelets. However, it is not clear whether

this attribute will be a useful, consistent key character.

These two species also occupy a typical Locapsis

habitat, along the arid fringes of the Cape. They have a

disjunct distribution, with Restio levynsiae in the Swart-

ruggens Mountains, and R. rigidus from the Voetpads-

berg near Laingsburg to the Great Swartberg.

8.6. 1 . Restio levynsiae (Pillans) H.P.Linder & C.R.

Hardy, comb. nov.

Leptocarpus levynsiae Pillans in Transactions of the Royal Soci-

ety of South Africa 29: 346 (1942). Calopsis levynsiae (Pillans)

H.P.Linder: 467 (1985). Type: Katbakkies, Levyns 1845 (BOL, holo.!).

8.6.2. Restio rigidus (Mast.) H.P.Linder & C.R. Har-

dy, comb. nov.

Leplocarpus rigida Mast, in Botanische Jahrbiicher 29, Beiblatt

66: 10 (1900b). Calopsis rigida (Mast.) H.P.Linder: 470 (1985). Type:

Roue Bokkeveld, Schlechter 8882, 8883 (K, lecto. ! [Linder in Bothalia

15: 470 (1985)]; BM!, BOL!, BR!, K!, P!, S!, WAG!, Z! ).

8.7. Restio subgen. Restio

Plants caespitose, clumped, tangled or mat-form-

ing. Fertile culms branching (with the sole exception of

R. strictus), round; sheaths persistent, closely or loosely

convoluted, usually with a narrow membranous mar-

gin, often with tall, acute, membranous lobes flank-

ing mucro. Male and female inflorescences mostly very

similar; mostly paniculate, more rarely racemose, with 1

to numerous spikelets; spathes usually persistent, often

overtopping spikelets; bracts varying from shorter to

taller than flowers, without honeycombed cells in api-

cal parts; anthers exserted from flowers at anthesis.

Female inflorescence occasionally with fewer spikelets

than males; spikelets with up to 16 laterally compressed

flowers; tepals chartaceous or cartilaginous, glabrous

or sparsely (rarely densely) villous on keels of lateral

sepals; staminodes present. Gynoecium with 3, white,

feathery styles free to base or rarely seated on a stylopo-

dium; ovary mostly unilocular, rarely with 2 locules, only

in R. sejunctus with 3 locules, either dehiscent or inde-

hiscent. Diaspore either a seed or a soft-walled nut; seed

variously coloured, usually smooth, occasionally collicu-

late; nut dropped with a persistent papery perianth that is

sometimes winged. Culm anatomy, epidermal cells 1 -lay-

ered, lateral walls usually straight and unthickened, occa-

sionally sinuose and thickened; parenchymatous layer of

up to 6 cells; central ground tissue either with no cavity,

a single central cavity, or with scattered cavities; tannin

rarely absent, when present then in the epidermis, scler-

enchyma, or central ground tissue; silica either absent, or

when present, in chlorenchyma or parenchyma.

Notes : Restio, as defined here, is still a very broad

subgenus, including species from Calopsis and Restio

sensu Linder 1984. Broadly speaking, it includes the

generally more slender members of Restio s.lat., with a

tendency towards small spikelets, to unilocular and often

indehiscent ovaries, and to more membranous sheaths

and bracts. This contrasts to subgen. Craspedolepis,

which includes the more robust Restio species.

Although there are no clearcut synapomorphies for

the subgenus, there are a number of attributes which in

combination will often lead to a correct identification.

Most species in the subgenus have tall membranous

lobes flanking the mucro on the sheath. These are also

found in many other genera (e.g. subgen. Ischyrolepis),

but are rare in the other segregates of Restio s.lat. In

Restio s.str. they are optimized as being ancestrally

present. As such, this character can be seen as a synapo-

morphy for the subgenus, although it cannot be used to

key out the subgenus, due to the absence of this feature

from many species of Restio, and its presence in many

species in other genera. In most species in the subgenus

there is a single functional locule. Furthermore, in most

cases this locule is also dehiscent. Typical of the subge-

nus are rather small spikelets, often with few flowers,

rather than the more massive spikelets characteristic of

Craspedolepis.

Ecologically most species are found along the wetter

coastal mountains, where they often form an understorey

under other vegetation.

Despite the substantial variation in the subgenus, it is

not possible to formally recognize sections, but to facili-

tate an understanding of the variation in this large taxon,

the species are placed into informal groups.

a) Restio aspera group: mostly ex-Calopsis species,

with paniculate inflorescences, rounded tepal apices,

tepals sometimes winged, and indehiscent ovaries:

8.7.1. Restio distichus Rottb., Descriptiones plantarum

rariorum 11(1 772).

8.7.2. Restio pulcher (Esterh.) H.P.Linder & C.R.

Hardy, comb. nov.

Calopsis pulchra Esterh. in Linder in Bothalia 15: 469 (1985).

Type: along road from Elim to Stanford, Esterhuysen 31255 (BOL,

holo.!; B!, BOL!, C!, E!, F, GRA!, K!, L, LD. ML MO!, NBG!, NY,

PRE!, RSA, S!, STE!, TCD!, UC, US, W!, WAG!).

8.7.3. Restio harveyi Mast, in Journal of the Linnean

Society, Botany 8: 253 (1865).

8.7.4. Restio asperus (Mast.) H.P.Linder & C.R. Har-

dy, comb. nov.

Hvpolaena aspera Mast, in Journal of the Linnean Society, Botany

10: 264 (1868). Calorophus asper (Mast.) Kuntze: 747 (1891). Lep-

locarpus asper (Mast.) Pillans: 345 (1928). Calopsis aspera (Mast.)

H.P.Linder: 465 (1985). Type: Nieuw Kloof, Houw Hoek. Burchell

8069 (K, lecto.! [Linder in Bothalia 15: 465 (1985)]; BOL!, K!, P!);

Hottentots Holland Mtns, Ecklon & Zeyher s.n. (K!); s.loc. Thom 632

(K!); mountains at Grietjiesgat, Ecklon & Zeyher s.n. (K!); Palmiet

River, Ecklon 951 ( B ! ).

8.7.5. Restio parvispiculus H.P.Linder & C.R. Har-

dy, nom. nov.

Hvpolaena burchellii Mast, in Journal of the Linnean Society, Bot-

any 10: 268 (1868). Calorophus burchellii (Mast.) Kuntze: 747 (1891).

Leptocarpus membranaceus Pillans: 346 (1928). Type: Nieuw Kloof,

Houw Hoek Mtns, Burchell 8116 (BOL!, K!); Baviaanskloof Mtns,

near Genadendal, Burchell 7894a (BOL!, K!, P!); 7632 (K, lecto.!

[Linder in Bothalia 15: 469 (1985)]; BOL!), non Restio burchellii Pil-

lans et Restio membranaceus Nees (= Elegia intermedia (Steud.) Pil-

lans.

8.7.6. Restio hyalinus (Mast.) H.P.Linder & C.R.

Hardy, comb. nov.

Hvpolaena hvalina Mast, in Botanische Jahrbiicher 29, Beiblatt

66: 13 (1900b). Leptocarpus hyalinus (Mast.) Pillans: 344 (1928).

Mastersiella hyalina (Mast.) Gilg-Ben.: 25 (1930). Calopsis hyalina

(Mast.) H.P.Linder: 467 (1985). Type: Koude River, Schlechter 10464,

Bothalia 40,1 (2010)

22

10465 (K, Iecto.! [Linder in Bothalia 15: 467 (1985)]; BOLL BR!,

MEL!, MO!, NBGL P!, S!, WAG!, Z!).

8.7.7. Restio impolitus Kunth, Enumeratio plantarum 3:

404(1841).

8.7.8. Restio versatilis H.P.Linder in Bothalia 15: 463

(1985).

b) Restio triticeus group: has dehiscent unilocular

ovaries. Two subgroups are recognized, one of which

has colliculate seed (rather than the smooth seed typical

of the rest of the genus):

8.7.9. Restio strictus N.E.Br. in Flora capensis 7: 752

(1900).

8.7.10. Restio triticeus Rottb., Descriptiones plantarum

rariorum 11 (1 772).

8.7.11. Restio verrucosus Esterh. in Linder in Bothalia

15: 462 (1985).

8.7.12. Restio pumilus Esterh. in Linder in Bothalia 15:

458 (1985).

8.7. 1 3. Restio zwartbergensis Pillans in Transactions of

the Royal Society of South Africa 1 6: 229 ( 1 928).

8.7.14. Restio villosus H.P.Linder & C.R. Hardy,

nom. nov. pro Calopsis sparsa Esterh. in Linder in South

African Journal of Botany 56: 454 (1990). Type: Cale-

don Div., Arieskraal, Esterhuvsen 32751 (BOL, holo.!,

B!, C!, El, K!, L!, Ml, MO!, PRE!, S!, STE!, UPS!), non

Restio sparsus Mast. (= Restio strobilifer Kunth).

The name refers to the uniquely villous flower bases.

8.7.15. Restio clandestinus (Esterh.) H.P.Linder &

C.R. Hardy, comb. nov.

Calopsis clandestina Esterh. in Linder in Bothalia 15: 465 (1985).

Type: mountain above Harold Porter Reserve, Esterhuysen 34146

(BOL, holo.!, B!, C!, E!, F, GRA!, K!, L, M!, MO!, NBGL NY, PRE,

RSA, S!, STE!, TCD!, UC, US, W!, WAG!).

8.7.16. Restio nudifiorus (Pillans) H.P.Linder &

C.R. Hardy, comb. nov.

Leptocarpus nudifiorus Pillans in Transactions of the Royal Soci-

ety of South Africa 30: 261 (1945). Calopsis nudiflora (Pillans)

H.P.Linder: 469 (1985). Type: Somerset Sneeukop, Esterhuysen 8230

(BOL, holo.!; K.!).

8.7.17. Restio pillansii H.P.Linder in Bothalia 15: 457

(1985).

8.7.18. Restio corneolus Esterh. in Linder in Bothalia

15: 443 (1985).

8.7.19. Restio saroclados Mast, in A. DC., Monogra-

phiae phanerogamarum 1 : 291 ( 1878).

8.7.20. Restio ejuncidus Mast, in Botanische Jahrbiicher

29, Beiblatt 66: 2 (1900b).

8.7.21. Restio leptostachyus Kunth, Enumeratio planta-

rum 3: 407 (1841).

c) Restio multiflorus group: bracts ovate, margins cili-

ate or shallowly toothed; ovaries with 2(3) locules:

8.7.22. Restio montanus Esterh. in Linder in Bothalia 15:

453 (1985).

8.7.23. Restio multiflorus Spreng., Systemavegetabilium

I: 187 (1824).

8.7.24. Restio tuberculatus Pillans in Annals of the Bo-

lus Herbarium 3,3: 146 (1922).

8.7.25. Restio scaberulus N.E.Br. in Flora capensis 7:

751 (1900).

8.7.26. Restio sejunctus Mast, in Flora capensis 7: 97

(1897).

8.7.27. Restio stokoei Pillans in Transactions of the Ro-

yal Society of South Africa 16:231 (1 928).

d) Restio dodii group: spathes taller than the spikelets,

plants often very fine and tangled, ovaries unilocular and

dehiscent:

8.7.28. Restio rarus Esterh. in Linder in Bothalia 15: 459

(1985).

8.7.29. Restio degenerans Pillans in Transactions of the

Royal Society of South Africa 30: 246 (1945).

8.7.30. Restio singularis Esterh. in Linder in Bothalia

15:460 (1985).

8.7.31. Restio alticola Pillans in Journal of South Afri-

can Botany 19: 101 (1952).

8.7.32. Restio scaber Mast, in Botanische Jahrbiicher 29,

Beiblatt 66: 1 ( 1900b).

8.7.33. Restio peculiaris Esterh. in Linder in Bothalia

15: 455 (1985).

8.7.34. Restio arcuatus Mast, in A. DC., Monographiae

phanerogamarum 1: 247 (1878).

8.7.35. Restio inconspicuus Esterh. in Linder in Bothalia

15: 449 (1985).

8.7.36. Restio vallis-simius /YRZrWer in Bothalia 15:462

(1985).

8.7.37. Restio secundus (Pillans) H.P.Linder in Bothalia

15: 460 (1985).

8.7.38. Restio decipiens (N.E.Br.) H.P.Linder in Botha-

lia 15:444(1985).

8.7.39. Restio fragilis Esterh. in Linder in Bothalia 15:

447 (1985).

8.7.40. Restio colliculospermus H.P.Linder in Bothalia

15:443 (1985).

8.7.41. Restio implicatus Esterh. in Linder in Bothalia

15:447 (1985).

8.7.42. Restio festuciformis Nees ex Mast, in Journal of

the Linnean Society, Botany 8: 248 (1865), as festucae-

formis.

8.7.43. Restio zuluensis H.P.Linder in Bothalia 15: 463

(1985).

8.7.44a. Restio dodii Pillans in Annals of the Bolus Her-

barium 3,2: 85 (1921) var. dodii.

8.7.44b. Restio dodii Pillans var. purpurascens Pillans

in Transactions of the Royal Society of South Africa 16:

252 (1928).

8.7.45. Restio dispar Mast, in Journal of the Linnean

Society, Botany 8: 246 (1865).

8.7.46. Restio purpurascens Nees ex Mast, in Journal

of the Linnean Society, Botany 8: 249 (1865).

8.7.47. Restio ingens Esterh. in Linder in Bothalia 15:449

(1985).

8.7.48. Restio communis Pillans in Transactions of the

Royal Society of South Africa 16: 264 (1928).

8.7.49. Restio paludicola H.P.Linder , sp. nov., a R.

communis spiculis maribus minoribus (8-10 mm longis),

paucifloris (1 vel 2) differt.

TYPE. — Western Cape, 3419 (Caledon): Caledon

Div., Hermanus Mountain, on rocky banks of a small per-

ennial stream, overlooking coast, growing quite densely

in places, 1-2 ft [0.3-0. 6 m] tall, ± 1000 ft [300 m], (-

AC), 26-08-1973, Esterhuysen 33222 (BOL, holo.).

Bothalia 40,1 (2010)

23

Plants mat-forming or tangled, stoloniferous. Fertile

culms sparsely branched, finely to roughly warty, green

or olivaceous, 0.3-0. 8 m long, 0. 1-0.7 mm diam. at apex;

sheaths closely convoluted, 10-35 mm long, reddish

brown with golden mottling, tuberculate, apical margins

truncate, coriaceous, hyaline shoulders up to half as long

as sheath, acute; mucro penicil late, straight and erect or

recurved, 3-9 mm long. Male inflorescence racemose,

with 2-10 spikelets, 40-80 mm long; spathes charta-

ceous, reddish brown with a golden speckling, persistent,

taller than spikelets; spikelets sessile, obovate (obtrian-

gular), rounded, 8-10 mm long, 1- or 2-flowered; bracts

longer than flowers, 3-6 mm long, linear or oblong, acute,

chartaceous, bract with upper margin like body of bract,

bract awn minute or absent. Male flower 2. 0-3. 5 mm long;

outer tepals cartilaginous; inner tepals shorter than outer,

membranous, outer lateral tepals conduplicate or as other

tepals, glabrous or keel sparsely villous (visible only under

higher magnification); anthers 1.2-1. 5 mm long, exserted

from flowers; pistillode present. Female inflorescence

racemose, with 2-10 spikelets, 30-80 mm long; spathes

longer than spikelets, persistent, cartilaginous; spikelet

sessile, obtriangular, truncate, 9-14 mm long, mostly sin-

gle-flowered, rarely 2- or 3-flowered; bracts at least as tall

as flowers, 5-9 mm long, linear or oblong, acute, charta-

ceous, awn minute or absent. Female flower 3. 5^4.0 mm

long; tepals cartilaginous, glabrous and smooth or keels

of lateral sepals sparsely villous, apices acute, inner and

outer whorls same length. 3. 5-4.0 mm long; odd outer

tepal linear or oblong, inner tepals linear or oblong; stami-

nodes present. Gynoecium : styles feathery, white, 3, free to

base; ovary unilocular, dehiscent. Seed 1.6-1. 7 x 0.9-0.95

mm, elliptical in side view, triangular in cross section, tan

or pink, smooth (shiny). Culm anatomy: epidermal cells

1 -layered, lateral walls straight, unthickened, outer wall

thickened, colliculate, glabrous, 1/w ratio 2.5 : 1.5; sto-

matal apparatus superficial, guard cells seated on top of

support cells; chlorenchyma of 2 layers of cells, 1/w ratio

5 : 3, inner and outer layers similar; protective cells reach-

ing to base of chlorenchyma layer; parenchymatous layer

of 1 or 2 cells, cells smaller than epidermal cells; scleren-

chyma with 4—7 layers of cells, without protrusions; cen-

tral ground tissue with scattered cavities; tannin found in

epidermis, sclerenchyma or central ground tissue; silica

sand in chlorenchyma (where protective cells touch paren-

chyma). Flowering time : March or April; seed release in

August or September.

Distribution and ecology, restricted to the Kleinri-

vierberge behind Hermanus, in the Western Cape, where

it is found at an altitude of 300-700 m, on Table Moun-

tain Sandstone. The species is restricted to wet habi-

tats, in seepages with Roridula , and along stream mar-

gins. These are found on the south-facing upper slopes

of these low mountains, where they receive ample rain

throughout the year. The plants grow into a dense tan-

gled understorey under the other vegetation.

Notes : the new species is very close to Restio communis

from the Cape Peninsula, but it differs in the less robust

spikelets, the more slender flowers, and the smaller female

bracts. The male spikelets are 8-10 mm long (instead

of 15-20 mm), and have only 1 or 2 flowers (instead of

6-12 flowers), and the male flowers are 2. 0-3. 5 mm long

(instead of 3. 6-4.0 mm long). There is a single collection

from the Caledon Swartberg, which might be intermediate,

or which might constitute yet another segregate.

Etymology, paluster (Latin), marshy; cola (Latin), dwel-

ler in; referring to the habitat of the species in boggy

places.

Additional collections

WESTERN CAPE. — 3419 (Caledon): Hermanus Mountain, Rori-

dula Stream, forming a dense undergrowth in wet seepages, (-AC), 16-

05-1999, H.P.Linder 6933 (Z); Caledon, above Vogelklip Kloof, Her-

manus, banks of small open stream, diffuse, amongst shrubs, (-AD),

18-11-1973, Esterhuysen 33352 (BOL); Caledon, Klein River Mtns,

nr Hermanus, Rocklands Peak area, in small marsh, on stream banks,

(-AD), 03-05-1971, Esterhuysen 32572 (BOL); Caledon. Maanschyn

Kop (Rocklands Kop), ± 1 mile [1.6 km] E of summit peak; in small

marsh below upper slopes of ridge, (-AD), 08-10-1967, Esterhuysen

31732 (BOL); Caledon, Maanschyn Kop, between Hermanus and

Stanford, in small marsh on E slopes, (-AD), 22-02-1968, Esterhuysen

31930a (BOL); Caledon, Swartberg, at base of marsh, on N slope, over

ridge, (-AB), 13-03-1976, Esterhuysen 34208 (BOL).

8.8. Restio subgen. Locapsis H.P.Linder & C.R.

Hardy, subgen. nov., a subgeneribus ad Restio ascriptis

combinatione epidermidis culmorum unistratae, cum

vaginis persistentibus, spiculisque maribus erectis, ovari-

oque indehiscenti, spiculisque femineis fusiformibus,

tepalisque chartaceis rotundatis distinguendum.

TYPE. — Restio vimineus Rottb.

Plants mostly tufted, occasionally tending to mat-form-

ing from spreading underground stolons. Fertile culms

mostly branching, round; sheaths persistent, closely or

loosely convoluted or often flat and standing free from

culm, margins generally same texture as body of sheath,

without membranous upper margin or membranous shoul-

ders. Male and female inflorescences similar, racemose or

paniculate, with up to 20 erect spikelets; spathes persist-

ent, very rarely overtopping spikelets; bracts concolorous,

margins same texture and colour as body of bract, usually

cartilaginous, as tall as or somewhat taller than flowers.

Male flowers with exserted anthers. Female spikelet gen-

erally narrowly elliptical, with up to 12 flowers, as flowers

mature bracts usually diverge, transforming spikelet from

slender to bristly. Female flowers laterally compressed;

female tepals chartaceous and apically rounded, keels of

lateral sepals generally ± villous; staminodes present or

absent. Gynoecium with 3, feathery, white to red styles

free to base or seated on a stylopodium; ovary unilocular,

indehiscent. Diaspores small, soft nuts, perianth persistent

and ± enclosing nuts. Culm anatomy, epidermal cells in

I layer, lateral walls straight and unthickened or sinuose

and thickened; parenchyma (1)2(3) cells wide; central

ground tissue with a single central cavity (this lacking in

R. ramosissimus) tannin usually absent, when present, in

epidermis and central ground tissue; silica usually absent,

but present in parenchyma of R. rigoratus.

Etymologer Locapsis (fi), an anagram of Calopsis.

Notes'. Restio subgen. Locapsis contains the largest

fragment of Calopsis sensu Linder (1984). The genus

is readily diagnosed by several attributes. The spread-

ing, concolorous sheaths are not found in all species, but

are quite unique in the tribe. The narrow, usually many-

flowered female spikelets are also distinctive, especially

together with the more-or-less papery, rounded tepals. As

such, the genus is easy to delimit against its phylogeneti-

24

Bothalia 40,1 (2010)

cally closest relatives: Restio s.str. and R. subgen. Ischy-

rolepis (Figure 5). However, it is much more difficult

to delimit the genus from Restio subgen. Eremorestio,

and morphologically and ecologically it would be much

more satisfying to combine these two genera.

8.8.1. Restio andreaeanus (Pillans) H.P.Linder &

C.R. Hardy, comb. nov.

Leptocarpus andreaeanus Pillans in Transactions of the Royal

Society of South Africa 16: 349 (1928). Calopsis andreaeana (Pillans)

H.P.Linder: 465 (1985). Type: Willowmore Aasvogelberg, Andreae 975

(BOL, holo.!; STE!).

8.8.2. Restio tenuispicatus H.P.Linder & C.R. Har-

dy, nom. nov. pro Hypolaena filiformis Mast, in journal

of the Linnean Society, Botany 10: 267 (1868). Calo-

rophus filiformis (Mast.) Kuntze: 747 (1891). Calopsis

filiformis (Mast.) H.P.Linder: 467 (1985). Type: Houw

Hoek Mtn, Zeyher 4349 (K, lecto.! [Linder in Bothalia

15: 467 (1985)]; BOL!, BR!, MEL!, P!); s.loc., Thom

1031 (K!), non Restio filiformis Poir.

The name refers to the slender culms, also expressed

in the specific epithet filiformis.

8.8.3. Restio calcicola H.P.Linder & C.R. Hardy,

nom. nov. pro Leptocarpus fruticosus Mast, in Bota-

nische Jahrbucher 29, Beiblatt 66: 9 (1900b). Calopsis

fruticosa (Mast.) H.P.Linder: 467 (1985). Type: in hills

near Ratels River, Schlechter 9718 (K, lecto.! [Linder,

Bothalia 15: 467 (1985)]; BM!, BR!, MO!, NBG!, P!,

S!, WAG!), Schlechter 9717 (B!, BM!, BR!, K!, MO!,

NBG!, P!, S!, WAG!), non Restio fruticosus Thunb. (=

Rhodocoma fruticosa (Thunb.) H.P.Linder).

This is a limestone species, hence the specific name.

8.8.4. Restio albotuberculatus H.P.Linder & C.R.

Hardy, nom. nov. pro Leptocarpus burchellii Mast, in

Journal of the Linnean Society, Botany 10: 222 (1868).

Calopsis burchellii (Mast.) H.P.Linder: 465 (1985).

Type: between Vet River and Krombecks River, Burchell

7185 (K, lecto.! [Linder in Bothalia 15: 465 (1985)];

BOL!); foot of Langeberg at Kampsche Berg, Burchell

7146 (BOL!, K!); Malgaten River at Wolf Drift, Burchell

6101 (BOL!, K!), non Restio burchellii Pillans.

Named for the characteristic white tubercles on the

culms.

8.8.5. Restio muirii (Pillans) H.P.Linder & C.R.

Hardy, comb. nov.

Leptocarpus muirii Pillans in Transactions of the Royal Society of

South Africa 16: 353 (1928). Calopsis muirii (Pillans) H.P.Linder: 469

(1985). Type: north end of Garcia’s Pass, Muir 3178, 31 79 (BOL, holo.l).

8.8.6. Restio ramosissimus H.P.Linder & C.R.

Hardy, nom. nov. pro Hypolaena gracilis Nees ex

Mast, in Journal of the Linnean Society, Botany 10: 266

( 1 868). Calorophus gracilis (Mast.) Kuntze: 747 (1891).

Leptocarpus gracilis (Mast.) Pillans: 146 (1922). Calop-

sis gracilis (Mast.) H.P.Linder: 467 (1985). Type: Mui-

zenberg, Zeyher 4347 (K, lecto.! [Linder in Bothalia 15:

467 (1985)]; MEL!, P!, S!, Z!); near Simonstown, Zey-

her 1006 (K!); Wright 500 (P!), non Restio gracilis R.Br.

8.8.7. Restio rigoratus (Mast.) H.P.Linder & C.R.

Hardy, comb. nov.

Leptocarpus rigoratus Mast, in Botanische Jahrbucher 29, Beiblatt

66: 9 (1900b). Calopsis rigorata (Mast.) H.P.Linder: 470 (1985). Type:

in hills near Vogelvlei near Elim, Schlechter 10490 (B, lecto.! [Linder

in Bothalia 15: 470 (1985)]; BOL!, K!, MO!, NBG!, P!, S!, WAG!, Z!).

8.8.8. Restio rudolfii (Pillans) H.P.Linder & C.R.

Hardy, nom. nov. pro Leptocarpus marlothii Pillans in

Transactions of the Royal Society of South Africa 16:

353 ( 1928). Calopsis marlothii (Pillans) H.P. Linder: 467

(1985). Type: Vaalkloof, near Karoopoort, Marloth 9102

(BOL, holo.!; B!, K!), non Restio marlothii Pillans.

Named for Rudolf Marloth.

8.8.9. Restio vimineus Rottb., Descriptiones plantarum

rariorum 10(1 772).

8.8.10. Restio adpressus (Esterh.) H.P.Linder &

C.R. Hardy, comb. nov.

Calopsis adpressa Esterh. in Linder in Bothalia 15: 465 (1985).

Type: hills inland from Pearly Beach, Esterhuysen 32977 (BOL, holo.!;

E!,K!,L, M!,MO!,S!).

8.8.11. Restio durus (Esterh.) H.P.Linder & C.R.

Hardy, comb. nov.

Calopsis dura Esterh. in Linder in Bothalia 15: 466 (1985). Type:

South Cedarberg suurvlakte, between Kaffirkop and Loskop, Ester-

huysen 34251 (BOL, holo.!; B!, C!, E!, F, GRA!, K!, L, LD. M!, MO!,

NBG!, NY, PRE!, RSA, S!, STE!, TCD!, UC, US, W!, WAG!).

8.9. Restio subgen. Ischyrolepis (Steud.) H.P.Lin-

der & C.R. Hardy, stat. nov.

Ischyrolepis Steud., Synopsis plantarum glumacea-

rum. 2: 249 (1855). Type: Ischyrolepis subverticellatus

Steud. (= Restio subverticellatus (Steud.) Mast.).

Plants tufted or tangled, often with spreading sto-

lons, but very rarely with rhizomes. Fertile culms usu-

ally branching, rarely unbranched or branches whorled

at nodes; round or very rarely compressed; sheaths per-

sistent, closely or more rarely loosely convoluted, very

rarely standing free from culm; margins coriaceous or

membranous, sometimes with tall membranous lobes

flanking mucro. Male and female inflorescences basi-

cally quite similar, racemose to paniculate, I to numerous

spikelets, except that females generally have fewer and

larger spikelets than males; in some species male spikelets

tightly clustered, while females tend to have single spike-

lets. Male spikelets sometimes linear and curved, bracts

obtuse to often acuminate and recurved, or extended into

a slender awn; anthers exserted from flowers at anthesis.

Female spikelets with up to 40 flowers; bracts shorter to

taller than flowers, apically obtuse to often acuminate and

recurved, or extended into a fine awn; flowers laterally

compressed; tepals chartaceous or cartilaginous; glabrous

or keels of lateral sepals more or less villous; staminodes

absent. Gynoecium with 2, white to red, feathery styles;

style bases fused to form a pillar; ovary with 2(1) loc-

ules, dehiscent. Seed translucent or variously coloured,

smooth, pitted, colliculate, striate or rarely rugose. Culm

anatomy: epidermal cells 1 -layered, lateral walls straight

and unthickened or sinuose and thickened; parenchyma-

tous layer 1-3 cells deep; central ground tissue solid or

with a single, central cavity; tannin absent or present in

epidermis, sclerenchyma or central ground tissue; silica

absent or present in parenchyma and/or rarely in central

ground tissue.

Etymology: Ischyrolepis ( f . ) : Ischyros (Greek), strong;

lepis (Greek), scale; presumably this refers to the usually

quite tough floral scales which enclose the flowers.

Bothalia 40,1 (2010)

25

Restio andreaeanus (Calopsis andreaeana)

Restio tenuispicatus (Calopsis filiformis)

1001 Restio calcicola (Calopsis fruticosa)

99 l_75_| Restio albotuberculatus (Calopsis burchellii)

76 Restio muirii (Calopsis muirii)

Restio ramosissimus (Calopsis gracilis)

Restio rigoratus (Calopsis rigorata)

120| Restio rudolfii (Calopsis mariothii)

98 Restio vimineus (Calopsis viminea)

88_J Restio adpressus (Calopsis adpressa)

61 Restio durus (Calopsis dura)

Restio anomaius

Restio femineus (Ischyrolepis feminea)

Restio wallichii (Ischyrolepis wallichii)

Restio rivulus (Ischyrolepis rivula)

Restio tenuissimus (Ischyrolepis tenuissima)

Restio iongiaristatus (Ischyrolepis longiaristata)

100| Restio sabulosus (Ischyrolepis sabulosa)

innl 1001 Restio rottboellioides (Ischyrolepis rottboellioides)

10QL 100| Restio paludosus (Ischyrolepis paludosa)

99 Restio papillosus (Ischyrolepis papillosa)

Restio mariothii (Ischyrolepis mariothii)

Restio schoenoides (Ischyrolepis schoenoides)

Restio constipatus

Restio virgeus (Ischyrolepis virgea)

Restio pratensis (Ischyrolepis pratensis)

Restio saxatilis (Ischyrolepis saxatilis) *

Restio sporadicus (Ischyrolepis sporadica)

Restio macer (Ischyrolepis macer)

Restio pygmaeus (Ischyrolepis pygmaea)

Restio caespitosus (Ischyrolepis caespitosa)

Restio cincinnatus (Ischyrolepis cincinnata)

Restio curviramis (Ischyrolepis cun/iramis)

Restio duthieae (Ischyrolepis duthieae)

Restio nanus (Ischyrolepis nana)

Restio vilis (Ischyrolepis vilis)

Restio laniger (Ischyrolepis laniger)

Restio coactilis (Ischyrolepis coactilis)

Restio karooicus (Ischyrolepis karooica)

Restio sieberi (Ischyrolepis sieberi)

Restio monanthos (Ischyrolepis monanthos)

27 | Restio parthenocarpos

Restio cedarbergensis

Restio hystrix (Ischyrolepis hystrix )

Restio esterhuyseniae (Ischyrolepis esterhuyseniae)

‘ Restio fraternus (Ischyrolepis fraterna) *

‘ Restio gossypinus (Ischyrolepis gossypina)

' Restio affinis (Ischyrolepis affinis) *

‘ Restio elsieae

' Restio ocreatus (Ischyrolepis ocreata)

‘ Restio aridus (Ischyrolepis arida)

‘ Restio capensis (Ischyrolepis capensis)

29 I Restio curvibracteatus (Ischyrolepis curvibracteata)

47 Restio wittebergensis (Ischyrolepis wittebergensis)

‘ Restio eleocharis (Ischyrolepis eleocharis)

‘ Restio leptoclados (Ischyrolepis leptoclados)

J Restio subverticellatus (Ischyrolepis subverticellata)

Restio triflora (Ischyrolepis triflora)

1^ | Restio helenae (Ischyrolepis helenae) *

Restio setiger (Ischyrolepis setiger)

' Restio nubigenus (Ischyrolepis nubigena)

100| Restio gaudichaldianus (Ischyrolepis gaudichaudiana) *

' Restio luxurians

' Restio venustulus (Ischyrolepis venustulus)

79 I Restio distractus (Ischyrolepis distracta)

70 Restio unispicatus (Ischyrolepis unispicata)

Restio subgen, Locapsis

Restio subgen. Ischyrolepis

FIGURE 5. — Portion of strict consensus tree of the total combined analysis of Hardy et al. (2008) containing Restio subgen. Locapsis andR. subgen.

Ischyrolepis. A single species of Restio subgen. Ischyrolepis (Restio fuscidulus) that was missing many data (including all molecular data),

was resolved in this tree as sister to Staberoha (see Figure 1), albeit with 3 % bootstrap support which we do not consider substantial enough

to warrant disputing the monophyly of Restio subgen. Ischyrolepis. Names in parentheses are the now synonymous names of Linder (2001a)

when different from the ones here proposed. Bootstrap values above branches indicate support based on a 500-replicate bootstrap analysis

that included 20 species for which DNA data were lacking (asterisked taxa). Bootstrap values below branches indicate support based upon a

500-replicate analysis that included only taxa for which both DNA data and morphology data were available. Where no value is given below

a branch it indicates that the clade did not appear in the analysis excluding morphology-only species.

Notes : Restio subgen. Ischyrolepis is readily diag-

nosed by a basal stylar peg, formed by the fusion of the

two style arms. This persists on the ripe ovary, even after

the seed has been released, as a distinctive peg. This is

unique in the Restioneae. Furthermore, it usually has

two styles (compare to the three or one more typically

found). The species can look superficially very similar to

Restio s.str. or Restio subgen. Craspedolepis. The pollen

type is held in common with Staberoha , but the two gen-

era are not phylogenetically closely related, according to

the molecular data.

It is particularly common in the more arid parts of the

Greater Cape Floristic Region (Born et al. 2007), and is

replaced along the wetter coastal mountains by the sub-

genera Craspedolepis and Restio.

26

Bothalia 40,1 (2010)

There are a number of groups in this large subge-

nus, but it is not possible to clearly diagnose them. Fur-

thermore, the subgenus, as currently defined, is so eas-

ily diagnosed that there is very little sense recognizing

smaller groups at subgeneric level. However, further

research may indicate that these subgroups deserve sec-

tional recognition.

a) Female spikelets small and mostly with single

flowers; styles often red:

8.9.1. Restio anomalus H.P.Linder sp. nov., a R.

femineus marginibus vaginarum late membranaceis, tepa-

lis interioribus florum feminearum 3.0 — 4.5 mm longis,

ovario biloculari statim dignoscenda.

TYPE. — Western Cape, 3419 (Caledon): Pheasants-

hoek near Viljoenshof, in a marshy area near Elegia

fistulosa and Restio festucaeformis, and along a small

stream (or furrow), in peaty soil and on clayish bank,

no male found, (-DA), 15-09-1976, Esterhuysen 34374

(BOL, holo.).

Plants compact, tufted, 0.15-0.25 m tall. Fertile culms

branching, round, solid or with a small central cavity,

finely rugulose or finely warty, green or olivaceous, 0.2-

0.4 mm diam. at apex; sheaths several, persistent, closely

convoluted, 5-13 mm long, acute, margins entire, varia-

ble: greenish/olivaceaous or tan or light to darker brown,

always with brown speckles and tan margins, hyaline

shoulders absent; mucro penicillate, straight and erect or

recurved, 3-7 mm long. Female inflorescence with 2-10

spikelets. interrupted or linear, 5-70 x 3-5 mm; spathes

shorter than spikelets, persistent, coriaceous or cartilagi-

nous; spikelet obovate or triangular with fruit sitting at

base and tepals flaring apart at top, 4. 0-6. 5 mm long,

with 1 flower, with 0 sterile bracts; bracts shorter than

flowers, erect, 2. 6-5.0 mm long, oblong, obtuse, carti-

laginous, awn minute to less than half as long as bract

body. Flower with four tepals; tepals 3 . 0 — 4 . 5 mm long,

undifferentiated, coriaceous or cartilaginous, glabrous

and smooth; margins entire, apices acute; staminodes

absent; ovary with 2 locules, dehiscent; style 1, flattened.

Seed ( 1 . 1 6—) 1 .27(— 1 .44) x (0.68— )0.73(— 0.8 ) mm, in side

view elliptical, in transverse view round, white, pitted.

Flowering time : September.

Distribution and ecology, endemic on the Agulhas

plains between Elim, Viljoenshof and the Soetanysberg,

at an altitude of 50-150 m, on either sandstone or sil-

cretes. The species is locally common on gravelly soils

or damp sand, often with impeded drainage.

Notes : this species is known currently only from

the female material, and since it can be very common

locally, it is presumably apomictic. It is closely related to

Restio femineus , which is also known only from female

material, and which forms similar neat little branching

tufts. However, R. anomalus differs in many details from

R. femineus. The sheath margins are broadly membra-

nous (instead of coriaceus or narrowly membranous),

the spathes overtop the spikelets (instead of being as tall

as the spikelets), the inner and outer tepals are the same

length (instead of the inner tepals being shorter than the

outer tepals), the inner tepals are 3. 0-4. 5 mm long, com-

pared to 2. 2-2. 6 mm in R. femineus , and the ovary has

two locules instead of one. In addition, there are numer-

ous small anatomical differences in the culm.

Etymology, a- (in Greek composition), not; normalis

(Greek), normal; abnormal, referring to the absence of

male plants.

Additional collections

WESTERN CAPE. — 3419 (Caledon): along road from Baardskeer-

dersbos to Elim; on laterite scrape on hill W of Elim. Locally common

on damp sand over laterite, probably seasonally waterlogged conditions.

Erect, tuft-like plants, only females seen, (-DA), 03-07-1999, Linder

6940 (BOL); Farm ‘Brandfontein'; plateau between the sea and Soetanys-

berg. Table Mountain Sandstone (TMS) with sand overlay. In shallow wet

sand on shaved off TMS bedrock, forming small tussocks. No males seen,

(-DD), 25-10-1990, Linder 5163 (BOL); Hangnes, base of Soetanysberg;

in marsh, near windmill, (-DD), 12-04-1978, Esterhuysen 34894 (BOL).

8.9.2. Restio femineus (Esterh.) H.P.Linder & C.R.

Hardy, comb. nov.

Ischyrolepis feminea Esterh. in Linder in Bothalia 15: 404 (1985).

Type: Simonstown, near the Four Seasons, W of Betty’s Bay, Esterhuysen

31662 (BOL, holo.!; Cl, E!, K!, LI, Ml, MO!, NBG!, PRE!, S!, STE!).

8.9.3. Restio wallichii Mast, in Journal of the Linnean So-

ciety, Botany 8: 234 ( 1865).

8.9.4. Restio rivulus (Esterh.) H.P.Linder & C.R.

Hardy, comb. nov.

Ischyrolepis rivula Esterh. in Linder in Bothalia 15: 413 (1985).

Type: Wuppertal, Boboskloof, Esterhuysen 31912 (BOL, holo.!; C!,

E!, F!, K!, L!, LD!, M!, MO!, NBG!,’ PRE!, S!, STE!, TCD!. UC!,

US!. W!, WAG!).

8.9.5. Restio tenuissimus Kunth in Enumeratio planta-

rum 3: 394(1841).

8.9.6. Restio longiaristatus (Pillans ex H.P.Linder)

H.P.Linder & C.R. Hardy, comb. nov.

Ischyrolepis longiaristata Pillans ex H.P.Linder in Bothalia 15:

407 (1985). Type: Van Rhynsdorp, Driekoppen, Gifberg, Esterhuysen

30749 (BOL, holo.!; B!, D!, E!, F!, GRA!, K!, L!, LD!, M!, MO!,

NBG!, NY!, PRE!, RSA!, S!, STE!, TCD!, UC!, US!, W!, WAG!).

8.9.7. Restio sabulosus Pillans in Transactions of the

Royal Society of South Africa 16: 285 (1928).

8.9.8. Restio rottboellioides Kunth, Enumeratio planta-

tion 3: 394 (1841).

8.9.9. Restio paludosus Pillans in Annals of the Bolus

Herbarium 3,3: 142 ( 1922).

8.9.10. Restio papillosus (Esterh.) H.P.Linder &

C.R. Hardy, comb. nov.

Ischyrolepis papillosa Esterh. in Linder in Bothalia 15: 410 (1985).

Type: Simonstown, Kenilworth Race Course, Esterhuysen 30849

(BOL, holo.!; B!, C!, E!, F!, GRA!, K!, L!. LD!, M!, MO!, NBG!,

NY!, PRE!, RSA!, S!, STE!, TCD!, UC!, US!, W!, WAG!).

8.9. 1 1 . Restio pratensis (Esterh.) H.P.Linder & C.R.

Hardy, comb. nov.

Ischyrolepis pratensis Esterh. in Linder in Bothalia 15: 411 (1985).

Type: Cape Town, Paarl Reserve on Paarl Mountain, Esterhuysen 31206

(BOL, holo.!; C!, E!, F!, K!, L!, M!, MO!, NBG!, S!, STE!, UC!).

8.9. 12. Restio macer Kunth, Enumeratio plantarum 3: 390

(1841).

8.9.13. Restio sporadicus (Esterh.) H.P.Linder &

C.R. Hardy, comb. nov.

Ischyrolepis sporadica Esterh. in Linder in Bothalia 15:413 (1985).

Type: Cape Town, Riverlands near Mamre Road, Esterhuysen 34656

(BOL, holo.!; C!, E!, K!, L!, M!, MO!, S!, STE!).

8.9.14. Restio pygmaeus Pillans in Transactions of the

Royal Society of South Africa 30: 253 (1945).

Bothalia 40,1 (2010)

27

8.9.15. Restio cincinnatus Mast, in Journal of the Lin-

nean Society. Botany 8: 240 (1865).

8.9.16. Restio caespitosus ( Esterh .) H.PLinder &

C.R. Hardy, comb. nov.

Ischyrolepis caespitosa Esterh. in Linder in Bothalia 15: 402

(1985). Type: Caledon, Pheasantshoek near Viljoenshof, Esterhuysen

34358 (BOL, holo.!; B!, C!, E!, F!, GRA!, K!, L!, LD!, M!, MO!,

NBG!, NY!, PRE!. RSA!, S!, STE!, TCD!, UC!, US!, W!, WAG!).

8.9.17. Restio saxatilis (Esterh.) H.P.Linder &

C.R. Hardy, comb. nov.

Ischyrolepis saxatilis Esterh. in Linder in South African Journal of

Botany 56: 456 (1990). Type: Stellenbosch, Dragoon Buttress, Ester-

huysen 32360 (BOL, holo.!; K!, M!, MO!, S!).

8.9.18. Restio curviramis Kunth, Enumeratio plantarum

3: 395 (1841).

8.9.19. Restio nanus (Esterh.) H.P.Linder & C.R.

Hardy, comb. nov.

Ischyrolepis nana Esterh. in Linder in Bothalia 15: 409 (1985).

Type: Worcester, lower Wellington Sneeukop, at W base of the shale

band. Esterhuysen 32658 (BOL, holo.!; C!, E!, K!, L!, M!, MO!, S!,

STE!).

8.9.20. Restio duthieae Pillans in Transactions of the

Royal Society of South Africa 16: 287 (1928).

b) Spikelets larger, usually few, many-flowered:

8.9.21. Restio marlothii Pillans in Annals of the Bolus

Herbarium 3,2: 83 (1921).

8.9.22. Restio schoenoides Kunth , Enumeratio planta-

rum 3: 391 (1841).

8.9.23. Restio constipatus H.P.Linder, sp. nov.,

a Restio wittebergensi culmis ramosis, a R. schoenoide

spiculis maribus 3-6 mm longis, spathis femineis spicula

aequantibus, seminibus argenteis recedit.

TYPE. — Western Cape, 3319 (Worcester): sum-

mit of Matroosberg, among rocks, (-BC), 21-01-2001,

H.P.Linder 7139 (Z, holo.; BOL, K, MO, NBG).

Plants clumped or tangled, stoloniferous, forming dense

cushions among boulders, 0.2-0. 5 m tall, or spreading

mats on open slopes. Fertile culms branching, smooth,

green or olivaceous, 0.5-1. 5 mm diam. at apex; sheaths

closely convoluted, 10-25 mm long, dark brown, apical

margins narrowly membranous, hyaline shoulders absent,

apex acute to acuminate; rnucro penicillate, straight and

erect, 2-10 mm long. Male inflorescence with 1 spikelet

(very rarely, in the Swartberg) or 2-10 spikelets, forming a

tight button, 8-25 x 7-25 mm; spathes persistent, as tall as

spikelets, coriaceous, rather similar to lower bracts; male

spikelets sessile, oblong or elliptical, 7-12 x 3-6 mm, 3-

10-flowered; bracts taller than flowers. 5.5-10 mm long,

oblong, acuminate, cartilaginous. Male flower 4. 5-5.0 mm

long, both tepal whorls equally long, hyaline or membra-

nous, linear or oblong, outer lateral tepals conduplicate,

sparsely villous on keels; anthers 2-3 mm long, exserted

from flowers; pistillode absent. Female inflorescence

with 1 spikelet (rarely, then mostly at western and eastern

extremes of range) or 2-10 spikelets, forming a tight but-

ton, 10-25 x 3-25 mm; spathes equalling spikelets, per-

sistent, coriaceous; spikelets sessile, elliptical, obtuse or

acute, 10-20 mm long, 5-1 5 -flowered, with 4 or 5 sterile

bracts; bracts taller than flowers, erect or reflexed, 5.5-10

mm long, oblong, acuminate, cartilaginous, apical margin

like rest of bract, same as body of bract, awn less than halt

as long as bract body. Female flower 5. 0-6. 5 mm long,

tepals cartilaginous, all equally long, outer lateral tepals

conduplicate with sparsely villous keels, odd outer and

inner tepals oblong; staminodes absent. Gynoecium: styles

2, feathery, white; style bases free but adjacent; ovary

dehiscent, bilocular. Seed 1.3-1. 5 x 1 mm, in side view

elliptical, in diam. round, silvery, pitted. Flowering time :

September to December.

Distribution and ecology, widespread on the sum-

mits of the inner ranges of the Cape Fold Mountains,

from Sneeukop in the Cedarberg to Mannetjiesberg in

the Kamannassie and Meiringspoortberg in the Groot

Swartberg. The species is absent from the lower coastal

mountains: the Kogelberg, Kleinriviersberg and the Cape

Peninsula. The altitude range is 1 500-2 100 m, all col-

lections are from sandstone habitats. This is a species of

rocky summits and exposed ridges. On the Matroosberg,

the first author found it growing on shady rock ledges,

well protected from fire, and quite cool, but dry. This

is consistent with most habitat notes, which indicate

cool, dry places, associated with rocks or rocky sites, or

ledges. Possibly these are all fire-protected habitats.

Notes: the new species is similar to Restio witteber-

gensis ; both species have both male and female spikelets

aggregated into button-like heads. However, it differs

from R. wittebergensis by its branching culms (R. witte-

bergensi s has simple culms), and the stoloniferous habit

(R. wittebergensis is caespitose). The stoloniferous habit

and aggregated spikelets suggest a similarity to R. sch-

oenoides, but this species is more slender, carpet-form-

ing rather than tangled. There is a wide range of vari-

ation within R. constipatus , and in its current definition

it might include two entities: the typical boulder-habi-

tat (rocky summit) plants from the western part of the

range, and more mat-forming plants from the Swartberg,

which are very common along the summit ridges of

these inland mountains. The differences in growth form

are corroborated by a subtle difference in the flower size

and a more slender growth form. This needs more, criti-

cal, investigation. These collections have in the past been

assigned to R. schoenoides', however, the latter species

is better understood as a summer rainfall species wide-

spread in the Drakensberg and reaching to the Blouberg

in the Limpopo Province. The interface between R. sch-

oenoides and R. wittebergensis remains taxonomically

difficult.

Etymology: constipata (Latin), crowded together;

referring to the male and female spikelets, clustered

together in capitate inflorescences. The name does not

refer to medicinal use.

Additional collections

WESTERN CAPE. — 3219 (Wuppertal): Cedarberg, kloof above

Crystal Pool, (-AC), 25-06-1942, Esterhuysen 789 7 (BOL); Cedar-

berg, peak near Sneeuwkop, (-AC), 02-01-1942, Esterhuysen 7567

(BOL); Cedarberg, Middelberg, SW slopes, (-AC), 01-03-1940, Ester-

huysen 2524b (BOL); Cold Bokkeveld, Turret Peak, in a cool shady

spot above stream, and at base of rock, in colonies, forming patches,

(-CC), 1 5-10-1972, Esterhuysen 33025 (BOL).

3319 (Worcester): Ceres-Tulbagh, Swartgat Peak, Sneeugat peaks,

on steep upper slopes, SE aspect, (-AA), 02-05-1964, Esterhuysen

30687 (BOL); Worcester, Mosterts Hoek Twins, (-AD), 08-01-1944,

Esterhuysen 9832 (BOL); Worcester, Mostert’s Hoek Twins, S side of

summit at base of low rock cap, in solid masses, (-AD), 23-02-1964,

28

Bothalia 40,1 (2010)

Esterhuysen 30615 (BOL); Worcester, Ridge Peaks, Waaihoek, sum-

mit, 1 800 m, amongst rocks, (-AD), 15-12-1942, Esterhuysen 8372

(BOL); Ceres, Roodeberg, (-BC), 01-01-1940, Esterhuysen 1487

(BOL); Worcester-Ceres, Matroosberg, at rocky summit, in shallow

soil around base of low rock or along cracks or in stony soil, dense

wiry plants, (-BC), 10-10-1962, Esterhuysen 29728 (BOL); Worcester,

Roodeberg, Matroosberg group, on upper rocky slopes, often along step

ledges, at the base of rock, dense growth, (-BC), 05-04-1964; Ester-

huysen 30677 (BOL); Worcester, Matroosberg, on upper rocky slopes,

along the base of low rock, dense masses, (-BC), 08-04-1980, Ester-

huysen 35419a (BOL); Ceres, Roodeberg, (-BC), 01-01-1940, Ester-

huysen 1487 (BOL); Worcester, Pulpit Rock Nek, Hex River Mtns,

in cracks in rock pavement ledges, (-BC), 09-03-1963, Esterhuysen

30076 (BOL); Worcester, Upper Wellington Sneeukop, N aspect, steep,

rocky, well-drained slope, (-CA), 04-10-1964, Esterhuysen 30779a

(BOL); Worcester, Du Toit’s Peak, on rocky upper slopes and sum-

mit plateau, in dense masses, (-CA), 21-12-1975, Esterhuysen 34164

(BOL); Worcester, Du Toit’s Peak, on rocky summit and upper rocky

slopes and ridges, S aspect, varying in height, (-CA), 01-12-1963,

Esterhuysen 30561 (BOL); Worcester, Du Toit's Peak, (-CA), 26-01-

1943, Esterhuysen 8558 (BOL); Worcester, Witteberg, on rocky slopes

near summit, (-CA), 28-02-1965, Esterhuysen 30957 (BOL); Worces-

ter, Upper Wellington Sneeukop, growing against or next to rock, not

seen on the smoother slopes, (- CA), 23-01-1972, Esterhuysen 32798

(BOL); Worcester, Slanghoek Peak, (- CA), 28-12-1975, Esterhuysen

s.n. (BOL); Worcester, Slanghoek Peak, on shale band on S side,

(-CA), 31-12-1972, Esterhuysen 33070a (BOL); Worcester, Slang-

hoek Mtns, ridge S of Grassy Dome, (-CA), 05-06-1966, Esterhuysen

31556 (BOL); Worcester, Du Toit’s Peak, at rocky summit, growing

between low rock, (-CA), 25-11-1962, Esterhuysen 29892 (BOL);

Worcester, Witteberg, above Du Toit’s Kloof, on summit, growing

from between rock and in shallow marshy soil, (-CA), 04-08-1963,

Esterhuysen 30253 (BOL); Worcester, Brandwacht Peak, near summit,

growing between rock surfaces or from crevices on W side, growing

densely, (-CB), 07-04-1963, Esterhuysen 30124\ Worcester, Waaihoek

Peak, on SSE side of rocky summit, and amongst rocks near summit,

common in patches, (-CB), 05-06-1965, Esterhuysen 31063 (BOL);

Kaffir Kop, between Paardekop and Adolph’s Kop, Fransch Hoek

Pass, Villiersdorp, on sloping sandstone pavement, S aspect, (-CC),

02-11-1969, Esterhuysen 32309a (BOL); Caledon, Olifantsberg area

near Genadendal, Boscheveld, rocky, summits, (-CD), 02 Jan. 1965,

Esterhuysen 30895 (BOL); Soutrivier, Koo Rd, ± 14 mis [22.4 km] S

of Matroosberg Station, in Protea mellifera veld, (-DB), 09-02-1964,

Jessop s.n. (BOL).

3321 (Ladismith): Ladismith, Toverkop, Swartberg, on upper

slopes, (-AC), 02-09-1973, Esterhuysen 33231 (BOL); Ladismith,

Toverkop, Swartberg, on ledges and rocky places, N or roughly N

aspect, common in patches, (-AC), 30-03-1964, Esterhuysen 30671

(BOL); Ladismith, from Toverkop to Seven Weeks Poort Berg, com-

mon on upper rocky slopes, (-AC), 30-03-1959, Esterhuysen 28269

(BOL); Prince Albert, Seven Weeks Poort Mtns, (-AD), 01-12-1928,

Stokoe s.n. (BOL); Ladismith, Swartberg, Kouveldberg, (-AD), 30-03-

1 964, Esterhuysen 30666a (BOL); Ladismith, Seven Weeks Poort Berg

to Toverkop, Swartberg, very common on upper slopes forming dense

stands or pure communities in places, in stony, rocky places, on ledges,

1.0-1. 5 ft [0.3-0. 5 m] tall, (-AD), 27-03-1964, Esterhuysen 30656

(BOL); Ladismith, Seven Weeks Poort Berg, common on upper slopes,

N north side and summit, (-AD), 01-04-1975, Esterhuysen 33798

(BOL); Calitzdorp, Gamka Reserve, Snygans-Baakens Kop area, at

base of small cliff, S aspect, below hilltop, and between rock on top of

cliff, (-BC), 22-05-1975, Esterhuysen 33834 (BOL).

3322 (Oudtshoom): Prince Albert, Swartberg, (-AC), 01-10-1949,

Stokoe in SAM74945 (BOL); Prince Albert, Swartberg, (-AC), 01-10-

1949, Stokoe in SAM74944 (BOL); Prince Albert, Tierberg area, Swart-

berg, N slopes, (-AD), 01-06-1962, Esterhuysen 29549 (BOL); Oudts-

hoom, Meiringspoortberg, on S slopes near summit and on N side on

small vlaktc, (-BC), 09-04-1966, Esterhuysen 31494 (BOL); Uniondale,

Mannetjicsberg, S slopes, (- DB), 05-1 1-1941, Esterhuysen 6403 (BOL).

8.9.24. Restio virgeus Mast, in Botanische Jahrbiicher

29, Beibiatt 66: 4 (1900b).

8.9.25. Restio curvibracteatus (Esterh.) H.P.Lin-

c/er & C.R. Hardy, comb. nov.

Ischyrolepis curvibracteata Esterh. in Linder in Bothalia 15; 403

(1985). Type; Worcester, Du Toit’s Peak, Esterhuysen 29894 (BOL,

holo.l; Bl, Cl, El, FI, Kl, LI, Ml, MO!, S!, STE1, UC1).

8.9.26. Restio wittebergensis (Esterh.) H.P.Linder

& C.R. Hardy, comb. nov.

Ischyrolepis wittebergensis Esterh. in Linder in Bothalia 15: 417

(1985). Type: Montagu, upper rocky S slopes of Witteberge, Ester-

huysen 30479 (BOL, holo.!; Cl, El, FI, K!, L!, LD!, M!, MO!. NBG!,

S!, STE!, UC!, WAG!).

8.9.27. Restio setiger Kunth, Enumeratio plantarum 3:

385 (1841).

8.9.28. Restio vilis Kunth , Enumeratio plantarum 3: 389

(1841).

8.9.29. Restio coactilis Mast, in Botanische Jahrbiicher

29, Beibiatt 66: 3 (1900b).

8.9.30. Restio karooicus (Esterh.) H.P.Linder & C.R.

Hardy, comb. nov.

Ischyrolepis karooica Esterh. in Linder in Bothalia 15: 406 (1985).

Type: Worcester, Karoopoort, on mountain slopes above the poort,

Esterhuysen 30458 (BOL, holo.!; BM!, CL E!, F!, GRA!, K!, L!, LD!,

M!, MO!, NBG!, NY!, PRE!, RSAL S!, STE!, TCDL UC!, US!, W!,

WAG!).

8.9.3 1 . Restio hystrix Mast, in Journal of the Linnean So-

ciety, Botany 10: 276 (1868).

8.9.32. Restio esterhuyseniae Pi/Ians in Transactions of

the Royal Society of South Africa 30: 248 (1945).

8.9.33. Restio gossypinus Mast, in Botanische Jahrbii-

cher 29, Beibiatt 66: 3 (1 900b).

8.9.34. Restio affinis (Esterh.) H.P.Linder & C.R.

Hardy, comb. nov.

Ischyrolepis affinis Esterh. in Linder in Bothalia 15: 401 (1985).

Type: Montagu, Langeberg, Kruispad Ridge between Goedgeloof Peak

and Protea Valley, Esterhuysen 35612 (BOL, holo.!; BL CL EL FL

K!, L!, LD!, M!, MO!, NBG!, PRE!, SL STE!, TCD!, UC!, US!, WL

WAG!).

8.9.35. Restio ocreatus Kunth, Enumeratio plantarum 3:

385 (1841).

8.9.36. Restio fraternus Kunth, Enumeratio plantarum

3: 386 (1841).

8.9.37. Restio elsieae H.P.Linder, sp. nov., R. ocre-

atae affinis, sed culmis simplicibus, spiculis femineis

ellipticis vel ovatis, seminibus longioribus (2.2-2. 8 mm)

notabilis.

TYPE. — Western Cape, 3319 (Worcester): Waaihoek

Mtns, between Zebasberg and Mt Superior, alt. I 600 m,

(-AD), 17-03-2002, H.P.Linder 7460 (Z, holo.; B, BOL,

K, MO, NBG, NSW, PRE).

Plants tufted, 0.16-0.45 m tall, without spreading rhi-

zomes or stolons. Fertile culms unbranched, round, solid

or with a small central cavity, smooth, olivaceous, 1.2-

2.0 mm diam. at apex; sheaths several, persistent, loosely

convoluted, 18-36 mm long, orange to reddish brown,

apical margins narrowly to broadly membranous, acute to

acuminate, hyaline shoulders continuing behind mucro;

mucro hair-like, straight and erect or recurved or twisted,

3-8 mm long. Male inflorescence racemose or globose,

15-35 x 9-20 mm wide, with 2-10 spikelets; spathes

persistent, at least as tall as spikelets, coriaceous or carti-

laginous; spikelets erect, elliptical, 7-13 x 2.8^1 mm, 10-

40-flowered; bracts longer than flowers, 5-1 1 mm long,

widely oblong, acuminate, coriaceous, bract with upper

margin membranous, awn recurved, acuminate, at least

half as long as the bract body. Male flower 4.5-6. 0 mm

long; tepals equally long, oblong to elliptical, outer tepals

more rigid than inner tepals, outer lateral tepals condupli-

cate and sparsely to densely villous on keels; anthers 1 .8-

Bothalia 40,1 (2010)

29

3.0 mm long, exserted from flowers; pistillode absent.

Female inflorescence with 1 spikelet; spathes up to as

long as spikelet, persistent, coriaceous or cartilaginous;

spikelet elliptical or ovate, 14-23 mm long, 10-30-flow-

ered; bracts longer than flowers, reflexed, 8-14 mm long,

ovate, acute or acuminate, coriaceous, awn up to as long

as bract body. Female flower 5-7 mm long, tepals carti-

laginous; outer lateral tepals conduplicate, villous along

keels, 5-7 mm long; odd outer tepal oblong or ovate, 5-

7 mm long; inner tepals ovate, 4. 0-6. 5 mm long; stami-

nodes absent. Gynoecium : styles 2, feathery, style bases

fused to form a pillar; ovary dehiscent, bilocular. Seed

(2.24— )2.47(-2.76) x ( 1 .3—) 1 .43(— 1 .53 ) mm, in side view

elliptical, diam. round, silvery or white (and shiny), pit-

ted. Flowering time : January and February; seed release

in March and April.

Distribution and ecology’', endemic in the Hex River

Mountains between Worcester and Ceres, where it is

locally very common between 1 600-2 100 m. It is

found on stony, well-drained mountain slopes. It appears

to occupy the same habitat as Restio virgeus , and the two

species alternate on the stony slopes between Zebasberg

and Mt Superior. The plants appear to be killed by fire,

regenerating from seed.

Notes'. Restio elsieae species is a high-altitude seg-

regate from R. ocreata. It differs by the simple, stout

culms, and smaller growth form. The single female

spikelet with recurved bracts and the unbranched

culms are similar to R. curvibracteatus, but R. elsieae

can immediately be recognized by the flat, spreading

sheaths.

Etymology', the species is named for Elsie E. Ester-

huysen. the foremost collector from the Cape mountains,

who made Restionaceae her speciality. She collected

actively from 1930 to 1990, and contributed more to

our knowledge of the Cape flora than any other collec-

tor. Her meticulous collections, assembled to reflect the

distribution range and morphological variability of each

species, are currently housed at the Bolus Herbarium,

but duplicates of her collections are widely distributed.

Additional collections

WESTERN CAPE. — 3319 (Worcester): Brandwacht Peak, rocky

slopes SW-S aspect, below beacon and at base of summit cap, clumps,

forming dense pure stands in places, erect, stiffly rounded, seed shed,

(-CB), 07-04-1963, Esterhuysen 30123 (BOL); Worcester, Fonteintjies-

berg, rocky places on E slopes of ridge below plateau, local, stiff habit,

dense rounded clumps, stems pointing in all directions, (-CB), 20-10-

1963, Esterhuysen 30423 (BOL); Worcester, Fonteintjiesberg, above

Pulpit Rock Nek, rocky plateau, N and S aspects and steep slope, (-CB),

16-12-1963, Esterhuysen 30588 (BOL); Ceres, Buffelshoek Peak, lead-

ing into Witels, shale band, SW aspect, (—AD), 08-10-1956, Esterhuysen

26361 (BOL); Ceres, Milner Peak, Hex River Mtns, rocky W slopes,

(-AD), 11-11-1960, Esterhuysen 28606 (BOL); Worcester, Waaihoek

Peak, upper rocky slopes, abundant in patches of almost pure stands,

dense, scratchy tufts, stems unbranched, (-AD), 11-02-1964, Ester-

huysen 30612 (BOL); Worcester, Buffelshoek Peak and Buffelshoek

Twins, rocky places, (-AD), 19-09-1965, Esterhuysen 31183 (BOL).

8.9.38. Restio cedarbergensis H.P.Linder, sp. nov.,

a R. ocreatae culmis gracilioribus, bracteis maribus bre-

vioribus (3. 5-5.0 mm longis), floribus maribus breviori-

bus (2. 5-3.0 mm longis), spiculis femineis brevioribus

(5-11 mm longis), floribus spiculis femineis paucibus

(3-8 floribus), seminibus laevibus differt.

TYPE. — Western Cape, 3219 (Wuppertal): Central

Cedarberg, Gabriel’s Pass directly above De Rif, 1 300

m, (-AC), 30-09-2000, H.P.Linder 7090 (Z, holo.; BOL,

K, MO.NBG, PRE).

Plants tufted, 0.3-0. 6 m tall, without spreading rhi-

zomes or stolons. Fertile culms branching, round, smooth

to roughly warty, green or olivaceous, slender, 0.2-0. 5

mm diam. at apex; sheaths loosely convoluted or flat and

standing free from culm, 8-20 mm long, golden brown to

yellow, acuminate, apical margins narrowly membranous,

hyaline shoulders absent; mucro absent or awl- or needle-

shaped, straight and erect, 0. 5-3.0 mm long. Male inflo-

rescence with 6-20 spikelets, racemose or paniculate,

20-55 x 6-12 mm; spathes persistent, at most as tall as

spikelets, cartilaginous, acuminate; spikelets pendulous

on flexible pedicels, linear or ovate, acute, 6-12 x 2-3

mm, 4-15-flowered; bracts equalling flowers, 3. 5-5.0

mm long, ovate, acuminate, cartilaginous, awn minute

or at least less than half as long as bract body, acuminate

recurved apices give spikelet a bristly appearance. Male

flowers 2. 5-3.0 mm long; tepals all equally long, linear

to oblong; outer tepals cartilaginous, laterals conduplicate

and sparsely villous on keels; inner tepals chartaceous;

anthers 1. 5-2.0 mm long, exserted from flowers; pistil-

lode absent. Female inflorescence with up to 5 spikelets,

sparsely paniculate, 5-35 x 4-8 mm; spathes up to as

long as spikelets, persistent, cartilaginous; spikelets ses-

sile, elliptical or obovate, rounded, obtuse or acute, 5-11

mm long, 3-8-flowered; bracts 4. 5-6.0 mm long, ovate,

apiculate, bony or coriaceous, awn less than half as long

as bract body. Female flower 3. 0-4.5 mm long; tepals

all equally long, 2.5 — 4.5 mm long, ovate, acute, bony or

coriaceous, outer lateral tepals conduplicate with densely

villous keels; staminodes absent. Gynoecium : styles 2,

feathery, bases fused to form a pillar; ovary dehiscent,

bilocular. Seed 1.7-2 x 1.1-1. 3 mm, side view oblong

(but distinctly bulged at one end), cross section triangular

(with corners very rounded), grey (often with brown mot-

tling), smooth. Flowering time : April.

Distribution and ecology’', this new species is

restricted to the southern Cedarberg and the northern

Bokkeveld, and has been collected from Bloukop, north

of the Bokkeveld Sneeukop, to Gabriel's Pass in the cen-

tral Cedarberg. In this region, Restio cedarbergensis is

found from 900-1 500 m, on well-drained soils, often

described as rocky. Several collections are from shale

bands, and the first author has seen the species grow next

to renosterveld, where sandstone meets shale, as well as

on typical dry TMS soils. The species appears to pre-

fer hot, dry, well-drained localities. Plants appear to be

killed by fire, regenerating from seed.

Notes'. Restio cedarbergensis was previously included

under R. ocreatus , but it differs by the slender culms,

and much smaller spikelets. On Gabriel’s Pass the two

species co-occur, without intermediates. The new spe-

cies has finer culms and smaller spikelets, and forms

lower, more tangled plants, whereas R. ocreatus is much

stouter, with larger spikelets. Curiously, the molecular

phylogeny does not indicate the relationship to R. ocrea-

tus, but suggests a relationship to the R. sieberi group,

which appears to be unlikely on the basis of the morpho-

logical data.

30

Bothalia 40,1 (2010)

Etymology', the name indicates the distribution range

of the species, from the Cedarberg.

Additional collections

WESTERN CAPE. — 3219 (Wuppertal): Cedarberg, Wolfberg-

Tafelberg area, in rocky places, common (-AC), 28-12-1962, Ester-

huysen 30008 (BOL); S Cedarberg, (-AC), 01-04-1947, Stokoe 9570

(BOL); S Cedarberg, Dwarsrivierberg, rocky slope, (-CA), 01 Apr.

1956, Esterhuysen 25548 (BOL); S Cedarberg, on E spur ol' Hond-

verbrand Ridge, sandy vlakte [flats], (-CB), 21-04-1946, Esterhuysen

12719 (BOL); S Cedarberg, Sandfontein Peak, between rocks, (-CB),

05-04-1947, Esterhuysen 13871 (BOL); S Cedarberg, Sandfontein

Peak, on rocky sandy W slopes, frequent, (-CB), 20-04-1957, Ester-

huysen 27213 (BOL); S Cedarberg, between Sandfontein Peak and

Gideon’s Kop, on lower stony W slopes, (-CB), 18-04-1965, Ester-

huysen 31024 (BOL); N Cold Bokkeveld, Bloukop, on shale band

slopes, locally very common, not seen along rocky sandstone ridge,

(-CB), 07-07-1968, Esterhuysen 31971 (BOL)._

8.9.39. Restio fuscidulus Pillans in Transactions of the

Royal Society of South Africa 30: 249 (1945).

8.9.40. Restio aridus Pillans in Annals of the Bolus

Herbarium 3,2: 84(1921).

8.9.41. Restio capensis (L.) H.P.Linder & C.R. Har-

dy, comb. nov.

Schoenus capensis L., Amoenitates 4: 264 (1755). Restio

dichotomus L.: 735 (1767), nom. illeg. Thamnochortus dichtomus (L.)

Spreng.: 187 (1824), nom. illeg. Ischyrolepis capensis (L.) Linder: 402

(1985). Type: Cape, s.loc., in Herb. Linn. 1164.3 (LINN holo.).

8.9.42. Restio eleocharis Mast, in Journal of the Lin-

nean Society, Botany 8: 238 (1865).

8.9.43. Restio leptoclados Mast, in Journal of the Lin-

nean Society, Botany 8 (1865).

8.9.44. Restio subverticellatus (Stead.) Mast, in Journal

of the Linnean Society, Botany 8: 227 ( 1 865)

8.9.45. Restio triflora Rottb., Descriptiones plantarum

rariorum 10 (1772).

8.9.46. Restio helenae Mast, in Journal of the Linnean

Society, Botany 8: 233 (1865)

8.9.47. Restio sieberi Kunth, Enumeratio plantarum 3:

387(1841).

8.9.48. Restio monanthos Mast, in Journal of the Lin-

nean Society, Botany 8: 238 (1865).

8.9.49. Restio parthenocarpos H.P. Linder sp.

nov., a R. sieberi spiculis femineis floribus 2 vel 3,

bracteis femineis acutis, stylis roseis, a R. monanthe

spiculis femineis 10-15 mm longis, bracteis femineis

10-13 mm longis, tepalis 5. 0-5. 5 mm longis, ab amba-

bus culmis parce ramificantibus, basi styli pubescenti

recedit.

TYPE. — Western Cape, 3219 (Wuppertal): Northern

Bokkeveld at start of pass to Suurvlakte, (-CA), 13-09-

2002, H.P.Linder 7509 (Z, holo.; BOL, K, MO, NBG,

NSW, PRE).

Plants tufted, compact, 0.25-0.4 m tall, without spread-

ing rhizomes or stolons, all plants female. Fertile culms

sparsely branched, round, smooth, olivaceous, finely warty

or finely rugulose, 0. 7-0.9 mm diam. at apex; sheaths sev-

eral, persistent, closely convoluted, brown, acute, 13-20

mm long, hyaline shoulders absent, mucro awl- or needle-

shaped, straight and erect, 3-8 mm long. Female inflores-

cence with up to 5 spikelets, racemose or spicate, 10-35 x

3-10 mm; spathes like floral bracts, persistent, coriaceous;

spikelets sessile, oblong, elliptical or ovate, rounded to

acute, 10-15 mm long, 2-3-flowered; bracts overtopping

flowers, ovate, 10-13 mm long, acute, coriaceous or carti-

laginous, imbricate and obscuring spikelet axis; bract awn

less than half as long as bract body. Female flower 5-6

mm long; tepals coriaceous or chartaceous, apices acute

to acuminate; outer lateral tepals 5. 0-5. 5 mm long, con-

duplicate with keels densely villous; odd outer tepal ovate,

4.0^4. 5 mm long; inner tepals shorter than outer, ovate,

3 mm long; staminodes absent; styles 2, feathery, pink or

red, with villous bases fused to form a pillar; ovary dehis-

cent, bilocular. Seed 1. 9-2.1 x 1.2-1. 3 mm, in side view

oblong, in cross section round or elliptical, silvery, pitted.

Flowering time'. September and October.

Distribution and ecology, this new, apparently

apomictic species is known from only two localities in

the Cold Bokkeveld, from the Suurvlakte at the north-

ern end of the Cold Bokkeveld, and from the summit

of the Skurweberg Pass at the southern end of the Cold

Bokkeveld. The two locations are at 750-1 100 m, on

sandstone, in deep sand over groundwater. Both loca-

tions showed some disturbance; the Skurweberg Pass

population showed substantial disturbance.

Notes', the affinities of Restio parthenocarpos are

not clear, but it may be related to Restio sieberi and

R. monanthos by the seed surface morphology, seed

shape, and general spikelet construction. It may also be

of hybrid origin. The molecular phylogeny indicates an

affinity to R. sieberi, R. monanthos and R. cedarber-

gensis. It differs from all species to which it might be

assigned by numerous minor characters. The complex

of variation associated with R. sieberi has not been

critically explored, and this possibly apomictic species

appears to be well embedded within this variation.

Etymology: parthenocarpos (Greek), producing seed

or fruit without fertilization; the specific epithet is a noun

in apposition.

Additional collection

WESTERN CAPE. — 3319 (Worcester): Agterwitzenberg Pass at

dam on pass summit, (-AB), 14-09-2002, Linder, Hardy & Moline

7523 (BOL, E, K, MO, NBG, NSW, PRE, Z).

8.9.50. Restio nubigenus ( Esterh .) H.P.Linder & C.R.

Hardy, comb. nov.

Ischyrolepis nubigena Esterh. in Linder in Bothalia 15: 409 (1985).

Type: Worcester, Matroosberg, Esterhuysen 27780 (BOL, holo.!; El,

Kl, LI, Ml, MO!, S!, STE1).

8.9.51. Restio gaudichaudianus Kunth, Enumeratio plan-

tarum 3: 387 (1841).

8.9.52. Restio luxurians (Pillans) H.P.Linder, comb,

et stat. nov.

Restio gaudichaudianus Kunth var. luxurians Pillans in Transac-

tions of the Royal Society of South Africa 16: 280 (1928). Type: Glen

Leith, foot of the Lange Berge (?), Muir 3078 (BOL, lecto.l [Linder

in Bothalia 15: 405 (1985)]; K! ); Worcester, Hex River Mountains,

Bolus 4234, 4235 (BOL!, K!); Bolus 15949, 15950 (BOL!); Wilde

Paarde Berg (?), Stokoe in BOL17665 (BOL!); Montagu, Langeberge

at Montagu, Marloth 3118, 3119 (BOL!); Page in BOL16540 (BOL!);

Mitchell in BOL16626 (BOL!); Ladismith, Zwarteberge, Marloth 3170

(BOL!); Willowmore, Aasvogelberg, Andreae 940 (BOL!); s.loc.,

Zwartwaterpoort, Burchell 3409 (BOL!, K!).

Notes: since this form can always and readily be dis-

tinguished from Restio gaudichaudianus, it is best rec-

ognized at specific level.

Bothalia 40,1 (2010)

31

8.9.53. Restio venustulus Kunth, Enumeratio plan-

tarum 3: 388 (1841).

Notes : although the spikelet structure is very similar

to the hyper-variable (and probably too broadly defined)

species Restio sieberi, the plants can always be distin-

guished by growth form, by the woolly prophylls, and

by the stouter culms and somewhat larger spikelets.

Ecologically it is also distinct, and is always associated

with large rocks, or even more commonly with passages

through the sandstone bedrock, where it often forms

large monospecific stands. These habitats are presum-

ably more shaded and colder than the typically sunny

open habitat of the numerous forms of R. sieberi.

8.9.54. Restio laniger Kunth , Enumeratio plantarum 3:

386(1841).

8.9.55. Restio distractus Mast., Flora capensis 7: 70

(1897).

8.9.56. Restio unispicatus (H.P.Linder) H.P.Linder

& C.R. Hardy, comb. nov.

Ischyrolepis unispicata H.P.Linder in Bothalia 15: 415 (1985). Type:

Wuppertal, Northern Cold Bokkeveld, Vredelus at E base of Schoonge-

zicht Peak, Esterhuysen 29670 (BOL, holo.l; Cl, El, Kl, LI, Ml, MO!,

SI).

ACKNOWLEDGEMENTS

This study was funded by the Swiss Science Foun-

dation grant SNF 31-66594-01, and the field work was

supported by the Claraz Foundation and National Geo-

graphic Society Grant 7289-02. CapeNature provided

collecting permission. We particularly want to thank

Philip Moline (field work, collaboration on the E/egia

complex), the Bolus Herbarium and Terry Trinder-Smith

for handling our collections in Cape Town. Peter Gold-

blatt. Barbara Briggs and Nick Helme are thanked for

critical input into the paper, and Peter Wilson and Otto

Feistner for checking the Fatin.

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BENTHAM, G. & HOOKER, J.D. 1862-1883. Genera plantarum.

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BERGIUS, P.J. 1767. Descriptiones plantarum ex Capite Bonae Spei.

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INDEX

Askidiosperma Steud., 14

albo-aristatum (Pillans) H P. Linder, 15

alticolum (Esterh.) H.P.Linder, 15

andreaeanum (Pillans) H.P.Linder, 15

capitatum Steud. , 1 5

chartaceum (Pillans) H.P.Linder, 15

delicatulum H.P.Linder, 15

esterhuyseniae (Pillans) H.P.Linder, 15

in’signe (Pillans) H.P.Linder, 15

longiflorum (Pillans) H.P.Linder, 15

nitidum (Mast.) H.P.Linder, 15

paniculatum (Mast.) H.P.Linder, 15

rugosum Esterh., 15

Calopsis Beauv. ex Desv., 1 5, 20

adpressa Esterh., 24

andreaeana (Pillans) FI.PLinder, 24

aspera (Mast.) H.P.Linder, 21

burchel/ii (Mast.) H.P.Linder, 24

clandestina Esterh., 22

dura Esterh., 24

esterhuyseniae (Pillans) H.P.Linder, 17

filiformis (Mast.) H.P.Linder, 24

fruticosa (Mast.) H.P.Linder, 24

gracilis (Mast.) H.P.Linder, 24

hyalina (Mast.) H.P.Linder, 21

levynsiae (Pillans) FI.PLinder, 21

marlolhii (Pillans) H.P. Linder, 24

monos tylis (Pillans) H.P.Linder, 17

muirii (Pillans) H.P.Linder, 24

nudifora (Pillans) H.P.Linder, 22

pulchra Esterh., 21

rigida (Mast.) H.P.Linder, 21

rigorata (Mast.) H.P.Linder, 24

Calorophus

asper (Mast. ) Kuntze, 2 1

burchellii (Mast.) Kuntze, 21

filiformis (Mast.) Kuntze, 24

gracilis (Mast.) Kuntze, 24

Chondropetalum Rottb., 1 1

Craspedolepis Steud., 15, 17

Dovea Kunth, 1 1

Elegia L., 1 1

acockii (Pillans) Moline & H.P.Linder, 14

aggregata (Mast.) Moline & H.P.Linder, 14

altigena Pillans, 14

amoena Pillans, 1 4

asperi flora (Nees) Kunth, 14

atrati flora Esterh., 14

caespitosa Esterh., 14

capensis (Burmf.) Schelpe, 14

coleura Nees ex Mast., 14

cuspidata Mast. , 12

decipiens (Esterh.) Moline & H.P.Linder, 14

deusta (Rottb.) Kunth, 14

dregeana Kunth, 14

ebracteata (Kunth) Moline & H.P.Linder, 14

elephantina H.P.Linder, 12

equisetacea Mast., 14

esterhuyseniae Pillans, 14

extensa Pillans , 1 4

fastigiata Mast., 14

fenestrata Pillans, 1 4

Bothalia 40.1 (2010)

33

Elegial. (cont.)

filacea Mast., 14

fistulosa Kunth, 14

fucata Esterh. , 14

galpinii N.E.Br., 14

grandis (Nees) Kunth, 14

grandispicata H.P.Linder, 14

hookeriana (Mast.) Moline & H.P.Linder, 14

hutchinsonii Pillans, 14

intermedia (Stead.) PHlans, 14

juncea L., 14

macrocarpa (Kunth) Moline & H.P.Linder, 14

marlothii (Pillans) Moline & H.P.Linder, 14

microcarpa (Kunth) Moline & H.P.Linder, 14

mucronata (Nees ) Kunth, 12

muirii Pillans, 14

neesii Mast., 14

nuda (Rottb.) Kunth, 1 2

persistens Mast., 14

prominens Pillans, 14

racemosa (Poir.) Pers., 14

recta (Mast.) Moline & H.P.Linder, 14

rigida Mast., 14

spathacea Mast. , 1 4

squamosa Mast., 14

stipularis Mast., 14

stokoei Pillans, 1 4

tectorum (L.f.) Moline & H.P.Linder, 12

thyrsifera (Rottb.) Pers., 14

thyrsoidea (Mast.) Pillans, 14

vaginulata Mast., 14

verreauxii Mast., 14

Hypolaena

aspera Mast., 21

burchellii Mast., 21

ftliformis Mast., 24

hyalina Mast., 2 1

mahonii N.E.Br., 8

Ischyrolepis Steud., 1 5, 24

affinis Esterh.. 28

caespitosa Esterh., 27

capensis (L.) Linder, 30

curvibracteata Esterh., 28

feminea Esterh., 26

karooica Esterh., 28

longiaristata Pillans, 26

nana Esterh., 27

nubigena Esterh., 30

papillosa Esterh., 26

pratensis Esterh., 26

rivula Esterh., 26

saxatilis Esterh., 27

sporadica Esterh., 26

unispicata H.P.Linder, 31

wittebergensis Esterh., 28

Lamprocaulis Mast., 1 1

Leptocarpus

andreaeanus Pillans, 24

asper (Mast.) Pillans, 21

gracilis (Mast.) Pillans, 24

hyalinus (Mast.) Pillans, 21

levynsiae Pillans, 21

membranaceus Pillans, 21

monostylis Pillans, 17

muirii Pillans, 24

nudiflorus Pillans, 22

rigida Mast., 21

rigoratus Mast., 24

Mastersiella hyalina (Mast.) Gilg-Ben., 21

Platycaulos H.P.Linder, 6

acutus Esterh., 8

anceps (Mast.) H.P.Linder, 8

callistachyus (Kunth) H.P.Linder, 8

cascadensis (Pillans) H.P.Linder, 8

compressus (Rottb.) H.P.Linder, 8

depauperatus (Kunth) H.P.Linder, 8

galpinii (Pillans) H.P.Linder & C.R. Hardy, 8

mahonii (N.E.Br.) H.P.Linder & C.R. Hardy, 8

subsp. humbertii (Cherm.) H.P.Linder & C.R. Hardy, 8

subsp. mahonii, 8

major (Mast.) H.P.Linder, 8

mlanjiensis (H.P.Linder) H.P.Linder & C.R. Hardy, 8

quartziticola (H.P.Linder) H.P.Linder & C.R.Hardy, 8

subcompressus (Pillans) H.P.Linder, 8

Restio Rottb., 15

acockii Pillans, 18

adpressus (Esterh.) H.P.Linder & C.R.Hardy, 24

affinis (Esterh.) H.P.Linder & C.R.Hardy, 28

albotuberculatus H.P.Linder & C.R.Hardy, 24

alticola Pillans, 22

ambiguus Mast., 6

andreaeanus (Pillans) H.P.Linder & C.R.Hardy, 24

anomalus H.P.Linder, 26

arcuatus Mast., 22

aridus Pillans, 30

asperus (Mast.) H.P.Linder & C.R.Hardy, 21

aureolus Pillans, 1 8

bifarius Mast., 17

bifidus Thunb., 17

bifurcus Nees ex Mast., 18

bolusii Pillans, 18

brachiatus (Mast.) Pillans, 1 8

brunneus Pillans, 1 8

burchellii Pillans, 18

caespitosus (Esterh.) H.P.Linder & C.R.Hardy, 27

calcicola H.P.Linder & C.R.Hardy, 24

capensis (L.) H.P.Linder & C.R.Hardy, 30

capillaris Kunth, 1 8

cedarbergensis H.P.Linder, 29

cincinnatus Mast., 27

clandestinus (Esterh.) H.P.Linder & C.R.Hardy, 22

coactilis Mast., 28

colliculospermus H.P.Linder, 22

communis Pillans, 22

confusus Pillans, 17

constipatus H.P.Linder, 27

comeolus Esterh., 22

curvibracteatus (Esterh.) H.P.Linder & C.R.Hardy, 28

curviramis Kunth, 27

cymosus (Mast.) Pillans, 1 8

debilis Nees, 20

decipiens (N.E.Br.) H.P.Linder, 22

degenerans Pillans, 22

dichotomus L., 30

dispar Mast., 22

distans Pillans, 20

distichus Rottb., 21

distractus Mast., 31

distylis H.P.Linder & C.R.Hardy, 17

dodii Pillans

var. dodii, 22

var. purpurascens Pillans, 22

durus (Esterh.) H.P.Linder & C.R.Hardy, 24

duthieae Pillans, 27

echinatus Kunth, 1 6

egregius Hochst., 1 7

ejuncidus Mast., 22

eleocharis Mast., 30

elsieae H.P.Linder, 28

esterhuyseniae Pillans, 28

femineus (Esterh.) H.P.Linder & C.R.Hardy, 26

festuciformis Nees ex Mast., 22

filiformis Poir., 18

foliosus N.E.Br., 11

fragilis Esterh., 22

fratemus Kunth, 28

fuscidulus Pillans, 30

fusiformis Pillans, 1 8

galpinii Pillans, 8

gaudichaudianus Kunth, 30

var. luxurious Pillans, 30

gossypinus Mast., 28

harveyi Mast. , 2 1

helenae Mast., 30

hyalinus (Mast.) H.P.Linder & C.R.Hardy, 21

hystrix Mast., 28

implicatus Esterh., 22

impolitus Kunth, 22

inconspicuus Esterh., 22

34

Bothalia 40,1 (2010)

Restio Roltb., (cont.)

ingens Esterh. , 22

insignis Pillans, 18

inveteratus Esterh., 18

karooicus (Esterh.) H.PLinder & C.R.Hardy, 28

laniger Kunth, 3 1

leptoclados Mast., 30

leptostachyus Kunth, 22

levynsiae (Pdlans) H.PLinder & C.R. Hardy, 21

longiaristatus (Pillans ex H.PLinder) H.PLinder & C.R.Hardy, 26

luxurians (Pillans) H.PLinder, 30

macer Kunth, 26

madagascariensis Cherm. var. humbertii Cherrn., 8

mahonii (N.E.Br.) Linder subsp. humbertii (Cherm.) H.RLinder, 8

mahonii (N.E.Br.) Pillans, 8

marlothii Pillans, 27

micans Nees, 1 7

miser Kunth, 17

mlanjiensis H.PLinder, 8

monanthos Mast., 30

monostylis (Pillans) H.PLinder & C.R.Hardy, 17

montanus Esterh. , 22

muirii (Pillans) H.PLinder & C.R.Hardy, 24

multiflorus Spreng., 22

nanus (Esterh.) H.PLinder & C.R.Hardy, 27

nodosus Pillans, 1 8

nubigenus (Esterh.) H.PLinder & C.R.Hardy, 30

nudiflorus (Pillans) H.PLinder & C.R.Hardy, 22

nuwebergensis Esterh., 17

obscurus Pillans, 1 8

occultus (Mast.) Pillans, 18

ocreatus Kunth, 28

pachystachyus Kunth, 1 8

paludicola H.PLinder, 22

paludosus Pillans, 26

paniculatus Rottb., 20

papillosus (Esterh.) H.PLinder & C.R.Hardy, 26

papyraceus Pillans, 1 7

parthenocarpos H P. Linder, 30

parvispiculus H.PLinder & C.R.Hardy, 21

patens Mast., 18

peculiaris Esterh., 22

pedicellatus Mast., 16

perplexus Kunth, 1 8

perseverans Esterh., 18

pillansii H.PLinder, 22

praeacutus Mast., 18

pratensis (Esterh.) H.PLinder & C.R.Hardy, 26

pulcher (Esterh.) H.PLinder & C.R.Hardy, 21

pulvinatus Esterh., 18

pumilus Esterh ., 22

purpurascens Nees ex Mast., 22

pygmaeus Pillans, 26

quadratus Mast., 20

quartziticola H.RLinder, 8

quinquefarius Nees, 20

ramosissimus H.P.Linder & C.R.Hardy, 24

rarus Esterh., 22

rigidus (Mast.) H.P.Linder & C.R.Hardy, 21

rigoratus (Mast.) H.P.Linder & C.R.Hardy, 24

rivulus (Esterh.) H.P.Linder & C.R.Hardy , 26

rottboellioides Kunth, 26

rudolfii (Pillans) H.P.Linder & C.R.Hardy, 24

rupicola Esterh., 18

sabulosus Pillans, 26

saroclados Mast, in A. DC., 22

saxatilis (Esterh.) H.P.Linder & C.R.Hardy, 21

scaber Mast. , 22

scaberulus N.E.Br., 22

schoenoides Kunth, 27

secundus (Pillans) H.P.Linder, 22

sejunctus Mast., 22

setiger Kunth, 28

sieberi Kunth, 30

similis Pillans, 20

singularis Esterh., 22

sporadicus (Esterh.) H.P.Linder & C.R.Hardy, 26

stereocaul is Mast., 16

stokoei Pillans, 22

strictus N.E.Br., 22

strobi lifer Kunth, 18

subgen. Calopsis (Beam, ex Desv.) H.P.Linder & C.R.Hardy, 20

subgen. Craspedolepis (Steud.) H.P.Linder & C.R.Hardy, 17

subgen. Eremorestio H.P.Linder & C.R.Hardy, 20

subgen. Ischyrolepis (Steud.) H.P.Linder & C.R.Hardy, 24

subgen. Locapsis H.P.Linder & C.R.Hardy, 23

subgen. Pendulostemon H.PLinder & C.R.Hardy, 17

subgen. Restio, 21

subgen. Simplicaulos H.P.Linder & C.R.Hardy, 16

subgen. Varirestio H.P.Linder & C.R.Hardy, 18

subtilis Nees ex Mast., 1 7

subverticellatus (Steud.) Mast., 30

tenuispicatus H.P.Linder & C.R.Hardy, 24

tenuissimus Kunth, 26

tetragonus Tlntnb., 20

triflora Rottb., 30

triticeus Rottb., 22

tuberculatus Pillans, 22

unispicatus (H.P.Linder) H.P.Linder & C.R.Hardy, 31

vallis-simius H.P.Linder, 22

venustulus Kunth, 3 1

verrucosus Esterh., 22

versatilis H.P.Linder, 22

vilis Kunth, 28

villosus H.P.Linder & C.R.Hardy, 22

vimineus Rottb., 24

virgeus Mast., 28

wallichii Mast., 26

wittebergensis (Esterh.) H.P.Linder & C.R.Hardy, 28

zuluensis H.P.Linder, 22

zwartbergensis Pillans, 22

Rhodocoma Nees, 1 0

alpina H.P.Linder & VI ok, 1 1

arida H.P.Linder & Vlok, 11

capensis Nees ex Steud., 1 1

foliosa (N.E.Br.) H.P.Linder & C.R.Hardy, 1 1

fruticosa (Thunb.) H.P.Linder, 1 1

gigantea (Kunth) H.P.Linder, 1 1

gracilis H.P.Linder & Vlok, 11

vleibergensis H.P.Linder, 1 1

Schoenus capensis L., 30

Soroveta H.P.Linder & C.R.Hardy, 6

ambigua (Mast.) H.P.Linder & C.R.Hardy, 6

Staberoha Kunth, 8

aemula (Kunth) Pillans, 9

banksii Pillans, 9

cemua (L.f.) Din: & Schinz, 9

distachyos (Rottb.) Kunth, 9

multispicula Pillans, 9

omata Esterh. , 9

remota Pillans, 9

stokoei Pillans, 9

vaginata (Thunb.) Pillans, 9

Thamnochortus P.J.Bergius, 9

acuminatus Pillans, 10

amoena H.PLinder, 10

arenarius Esterh., 10

bachmannii Mast., 10

cinereus H.P.Linder, 10

dichtomus (L.) Spreng., 30

dumosus Mast., 10

elliplicus Pillans, 10

erectus (Thunb.) Mast., 10

fraternus Pillans, 10

fruticosus P.J.Bergius, 10

glaber (Mast.) Pillans, 10

gracilis Mast., 10

guthrieae Pillans, 1 0

insignis Mast., 10

karooica H.P.Linder, 10

levynsiae Pillans, 10

lucens (Poir.) H.P.Linder, 10

muirii Pillans, 10

nutans (Thunb.) Pillans, 10

obtusus Pillans, 1 0

paniculatus Mast., 10

papyraceus Pillans, 1 0

pellucidus Pillans, 10

platypteris AT///7//7, 10

pluristachyus Mast., 10

Bothalia 40,1 (2010)

35

Thamnochortus P.J.Bergius (cont.)

pulcher Pillans, 10

punctatus Pillans, 1 0

rigidus Esterh., 10

schlechteri Pillans, 10

spicigerus (Thunb.) Spreng., 10

sporadicus Pillans, 10

stokoei Pillans, 1 0

Bothalia 40,1: 37^46 (2010)

New synonyms and a new name in Asteraceae: Senecioneae from the

southern African winter rainfall region

J.C. MANNING* and P. GOLDBLATT**

Keywords: Aluka website, Asteraceae, Othonna, Senecio, southern Africa, taxonomy

ABSTRACT

A review of the genera Othonna and Senecio undertaken for the forthcoming Greater Cape plants 2: Namaqualand-south-

ern Namib and western Karoo (Manning in prep.) led to a re-examination of the taxonomic status of several species. This

was facilitated by the recent availability of high-resolution digital images on the Aluka website (www.aluka.org) of the Drege

isotypes in the Paris Herbarium that formed the basis of many species described by De Candolle in his Prodromus systematis

naturalis regni vegetabilis. These images made it possible to identify several names whose application had remained uncer-

tain until now. Each case is briefly discussed, with citation of additional relevant herbarium specimens. The following species

are reduced to synonomy: O. incisa Harv. is included in O. rosea Harv.; O. spektakelensis Compton and O. zeyheri Sond. ex

Harv. are included in O. retrorsa DC.; S. maydae Merxm. is included in S. albopnnctatus Bolus, which is now considered to

include forms with radiate and discoid capitula; S. cakilefolius DC. is included in O. arenarius Thunb.; S. pearsonii Hutch,

is included in O. asperulus DC.; S. parvifolius DC. is included in S. carroensis DC.; S. eriobasis DC. is included in 5. erosns

L.f.; and S. lobelioides DC. is included in S.flavus (Decne.) Sch.Bip. The name 5. panduratus (Thunb.) Less, is identified as

a synonym of S. erosus L.f. and plants that are currently known under this name should be called S. robertiifolius DC. The

confusion in the application of the names O. perfoliata (L.f.) Jacq. and O.filicaulis Jacq. is examined. O. perfoliata is lecto-

typified against a specimen in the Linnaean Herbarium (LINN) with radiate capitula. The name O. filicaulis correctly applies

to a radiate species and is treated as a synonym of O. perfoliata. The vegetatively similar taxon with disciform capitula that is

currently known as O.filicaulis should be known as O. undulosa (DC.) J.C. Manning & Goldblatt, comb. nov. The new name

O. daucifolia J.C. Manning & Goldblatt is provided to replace the later homonym O. abrotanifolia (Harv.) Druce.

INTRODUCTION

The South African species of Othonna L. and Senecio

L. were last revised at a regional level by Harvey (1865).

Since then, the summer rainfall species from KwaZulu-

Natal have been studied intensively by Hilliard (1977)

but many species in the southern African winter rainfall

region remain poorly understood. The recent availabil-

ity of high-resolution digital images of type material of

many African taxa (www.aluka.org), especially the Paris

isotype material of the Drege collections that formed the

basis of a number of De Candolle’s (1838) species, has

made it possible to identify several taxa that were never

seen by Harvey (1865) and whose identity has remained

uncertain. During the preparation of a review of the tribe

Senecioneae (Manning in prep.), various nomenclatural

and taxonomic issues were encountered that can now be

addressed. The application of some of these names and

other taxonomic and nomenclatural issues arising during

the study are dealt with here.

The digital images, or virtual herbaria, available online

through Aluka and other sites such as the Herbarium of the

Linnean Society of London (www.linnean-online.org) are

often adequate for providing confirmation of the identity

of names where the gross morphology of the taxa in ques-

tion is diagnostic but they cannot replace microscopic

examination of actual material for critical features. We

have, therefore, avoided making any taxonomic infer-

ences in instances where the identity of the digital image

depends on examination of such micro-characters. In addi-

* Compton Herbarium, South African National Biodiversity Institute,

Private Bag X7, 7735 Claremont, Cape Town.

** B.A. Krukoff Curator of African Botany, Missouri Botanical Gar-

den. P.O. Box 299, St. Louis, Missouri 63166, USA.

MS. received: 2009-03-31.

tion, digital images have only been accepted as genuine

type material where it is absolutely clear that this is the

case from the collecting number and locality details on the

sheets in question.

OTHONNA

1. The shrublet with dissected leaves and radiate

capitula from the southwestern Cape currently known

as Euryops abrotanifolius (L.) DC. (1838) is based on

Othonna abrotanifolia L. (1753). The same epithet was

also used for Doria abrotanifolia Harv. (1865), a species

with disciform capitula from Namaqualand, which was

later transferred to Othonna as O. abrotanifolia (Harv.)

Druce (1917). Although aware of the existence of Lin-

naeus’ combination, Druce (1917) did not consider it an

impediment to his transfer of the epithet and it escaped

the notice of later authors, who continued to treat the

species under this name (e.g. Welman 2006). However,

the existence of Linnaeus’ earlier name renders Druce’s

combination an illegitimate homonym (McNeil et al.

2006: Art. 53.1 ), and the Namaqualand species therefore

requires a new name, which we provide here, alluding to

the unusually finely dissected leaves.

Othonna daucifolia J.C. Manning & Goldblatt , nom.

nov. pro Othonna abrotanifolia (Harv.) Druce: 638 (1917),

non Othonna abrotanifolia L. ( 1753) [= Euryops abrotani-

folius (L.) DC.]. Doria abrotanifolia Harv.: 324 (1865).

Type: South Africa [Northern Cape], Springbokfontein

[Springbok], without date, Whitehead s.n. TCD0003161

(TCD, holo.-Aluka image!, website accessed 20-03-09).

2. The application of the names currently used for the

two species Othonna filicaulis Jacq. and O. perfoliata

Jacq. (e.g. Goldblatt & Manning 2000) is highly confused

38

Bothalia 40,1 (2010)

(Table 1 ). These vegetatively similar, deciduous geophytes

from the Western Cape are tuberous perennials produc-

ing slender, often scandent, annual stems bearing perfoli-

ate or amplexicaul leaves and terminating in pedunculate

capitula. Their remarkable vegetative similarity led Row-

ley ( 1994: 174) to treat both radiate and disciform plants in

a single, variable species under the name O. filicaalis. He

may be correct, but until the situation is fully investigated

we continue to treat them as distinct.

Othonna perfoliata is distinguished from O. filicau-

lis by its generally fewer phyllaries (8-10 vs 10-12) and

radiate marginal florets, although in some collections the

rays are reduced and rather small. The marginal florets

in O. filicaulis , in contrast, are filiform and rayless, with

the truncate corolla less than half as long as the style. All

fruiting material of O. perfoliata that we have seen has

the mature pappus in the marginal florets 5-8 mm long

and ± as long as the cypselae or only slightly longer,

whereas in O. filicaulis the pappus often elongates greatly,

reaching 10-20 mm long, therefore 2-\ times as long as

the cypselae at maturity. O. filicaulis as currently under-

stood (e.g. Goldblatt & Manning 2000) is widespread

in the southern African winter rainfall region, mainly on

sandy, often coastal flats, from southern Namibia as far

east as Uniondale, whereas O. perfoliata is essentially

restricted to montane habitats in the southwestern Cape,

where it occurs on rocky slopes between the Bokkeveld

Mountains and Caledon (Goldblatt & Manning 2000).

The epithet perfoliata was first used in this context

by Linnaeus f. (1782) in the combination Cineraria per-

foliata L.f., based on material collected at the Cape of

Good Hope by Thunberg. There are no sheets under this

name in the Linnaean Herbarium at Stockholm (S) but

the Herbarium of the Linnean Society (LINN) contains

two sheets labelled as such, one of them a radiate plant

(LINN1000.32) and the other one evidently disciform

(LINN1000.33). The protologue, unfortunately, makes no

mention of the condition of the capitula but the descrip-

tion of the leaves as ovate-cordate and of the peduncles

as elongated, with a solitary capitulum [foliis ovatis cor-

datis amplexicaulibus, pedunculis unifloris elongatis ]

matches the radiate-flowered specimen more closely than

the disciform one, in which the leaves are lanceolate-

undulate and the inflorescence sparsely branched. It is in

the sense of a radiate taxon that Jacquin (1797) illustrated

and described the species, citing C. perfoliata L.f. under

the name Othonna perfoliata , which therefore constitutes

the new combination O. perfoliata (L.f.) Jacq.

Thunberg, however, understood the taxon in the oppo-

site way when he established the genus Doria Thunb.

(1800a) for various Senecioneae with disciform capitula

and transferred the name to that genus as Doria perfo-

liata (L.f.) Thunb. (1800b). This interpretation evidently

stems from the duplicate material in his possession, as

the Thunberg Herbarium (UPS-THUNB) contains a

single, disciform plant {UPS-THUNB 19833) under the

name Doria perfoliata. The name was subsequently used

in this sense as applying to the disciform taxon by both

De Candolle ( 1 838) and Harvey ( 1 865) (although the lat-

ter mistakenly attributed the basionym to Lamarck). As

a result of this, De Candolle (1838) described the new

species O. amplexifolia DC. for the conspicuously radi-

ate taxon, based on material collected by Drege, citing

Jacquin’s illustration and interpretation of O. perfoliata

in the protologue to his O. amplexifolia , but explicitly

excluding the Linnaean basionym Cineraria perfoliata

in keeping with his application of this name to the dis-

ciform taxon (as Doria perfoliata (L.f.) Thunb.). The

name was subsequently transferred to Othonna as O.

perfoliata (L.f.) Sch.Bip. (1844) but this combination is

in any event preoccupied by Othonna perfoliata (L.f.)

Jacq. (1797), making it a later homonym and thus ille-

gitimate (McNeil et al. 2006: Art. 53).

At the time that he figured the radiate taxon under

the name Othonna perfoliata , Jacquin (1797) described

a second, similar taxon under the name O. filicaulis

Jacq. This species was distinguished from O. perfoliata

by the narrower, lanceolate-undulate leaves and by the

smaller, linear rays. The ligulate condition of the mar-

ginal florets is clearly described and is also illustrated in

a detail of a marginal floret but the rays are not evident

in the drawing of the whole plant, which thus appears

to be disciform. Both De Candolle (1838) and Harvey

(1865) treated the species as circumscribed by Jacquin

although neither knew it from actual material. The name

later became associated with the disciform species, pre-

sumably because Jacquin’s painting of the whole plant

appeared to represent a disciform individual, and it is

this application that is current (e.g. Goldblatt & Man-

ning 2000). This transfer in the application of the name

O. filicaulis to the disciform taxon appears to date to the

Flora of the Cape Peninsula (Adamson & Salter 1950),

in which the name O. filicaulis is explicitly applied to the

disciform species with filiform, truncate marginal corol-

las ‘previously misidentified as O. perfoliata ’ (Adamson

& Salter 1950: 820). In any event, the protologue of O.

filicaulis makes it quite clear that the name is properly

applied to a radiate plant.

Examination of available herbarium specimens of Oth-

onna perfoliata shows that the species exhibits a wide

range of leaf shapes, ranging from suborbicular and plane

(typical O. perfoliata ) to lanceolate and undulate (typical

O. filicaulis). The ray florets also vary from well-developed

and oblong to smaller and narrow. We therefore conclude

that O. filicaulis is conspecific with O. perfoliata and place

the name in synonymy under it.

TABLE I. — Application of epithets in Othonna filicaulis-perfoliata group

Epithet Application

Linnaeus f. (1782) Jacquin (1797) Thunberg (1800) De Candolle ( 1 838) Harvey (1865) Adamson & Salter

Bothalia 40,1 (2010)

39

This leaves the disciform species, currently identified

as Othonna filicaulis , without a name. Doria undulosa

DC. (1838) was described for a collection with straggling

stems bearing cordate-lanceolate, undulate leaves and dis-

ciform capitula with short, truncate marginal corollas that

was gathered in the Hex River Mountains by Drege. The

species was imperfectly known to Harvey (1865) but our

examination of the isotype material at Paris convinces us

that it falls within the range of variation of the taxon that

is currently incorrectly called O. filicaulis. The name D.

undulosa is therefore available for the disciform taxon and

the new combination required in Othonna is made here.

Othonna perfoliata (L.f.) Jacq.. Plantarum rario-

rum horti caesarei schoenbrunnensis 2: 61 , t. 240 ( 1 797).

Cineraria perfoliata L.f.: 375 (1782 ‘1781’). Doria per-

foliata (L.f.) Thunb.: 155 (1800b). Othonna perfoliata

(L.f.) Sch.Bip.: 769 (1844), illegit. superfl. name. Type:

South Africa, without locality or date, Thunberg s.n.

(LINN1000.32, lecto., here designated-Linnean image!,

website accessed 20-03-09).

O. filicaulis Jacq.: 62, t. 241 (1797), not of Adamson & Salter

(1950) and later authors [= Othonna undulosa (DC.) J.C.Manning &

Goldblatt], syn. nov. Type: illustration in Jacq.: t. 241 (1797), lecto.,

here designated: original material unknown.

O. amplexifolia DC.: 480 (1838), syn. nov. Type: South Africa,

[Western Cape], Paarl, [31 August 1827], Drege [6063] (G-DC, holo.-

microfichel; K, P-Aluka images!, website accessed 20-03-09).

Representative additional specimens examined

NORTHERN CAPE.— 3119 (Calvinia): Oorlogskloof Nature

Reserve, (-AC), 6 June 1995, Pretorius 245 (NBG).

WESTERN CAPE. — 3118 (Vanrhynsdorp): Matsikamma, (-DB),

12 July 1974, Thompson 2054 (NBG). 3219 (Wuppertal): Cedarberg,

Gonnafontein, (-CB), 23 May 2000, Pond 77 (NBG). 3319 (Worces-

ter): Rawsonville, (-CA), 28 July 1962, Walters 659 (NBG); Voet-

padsberg. (-DA). 8 June 1985, Van Wyk 23 73 (NBG). 3419 (Caledon):

Panorama, (-CB), 4 July 1976, Neethling s.n. (NBG).

Othonna undulosa (DC.) J.C.Manning & Gold-

blatt, comb. nov. Doria undulosa DC.: 472 (1838). Type:

South Africa, [Western Cape], Hexriviersberg, [Hex

River Mountains, 3 June 1820], Drege [289] (G-DC,

holo.-microfiche!; P-Aluka image!, website accessed

20-03-09).

O. filicaulis sensu auct., non Jacq. [= O. perfoliata (L.f.) Jacq.].

Representative additional specimens examined

NORTHERN CAPE. — 2917 (Springbok): bottom of Wildepaarde-

hoek Pass, (-DC), 24 August 1976, Van Jaarsveld 1399 (NBG). 3018

(Kamiesberg): Studer’s Pass, (-AC), 30 August 1975, Oliver 5952

(NBG).

WESTERN CAPE.— 3218 (Velddrif): Rocher Pan Reserve, (-CB),

25 July 1981, Le Roux & Van Rooyen 6 (NBG). 3319 (Worcester):

Karoo Garden veld, (-CB), Peny 165 (NBG). 3420 (Bredasdorp):

Zoetendalsvlei Vallei turnoff from Bredasorp-Agulhas road, (-CA), 6

July 1995. Paterson-Jones 433 (NBG).

3. Othonna lingua L.f. (1782) was based on a collection

made by Thunberg at the Cape of Good Hope. The name

of the taxon has been incorrectly rendered as O. lingua

(Less.) Sch.Bip. (1844) based on the name Doria lingua

Less. (1832) but reference to Lessing (1832: 89) reveals

that he did not in fact publish this name. It first appears

as D. lingua in De Candolle (1838: 471 ), where it is mis-

takenly attributed to Lessing as being based on the basio-

nym Othonna lingua Jacq. What Lessing (1832) actu-

ally published was the combination Doria digitata (L.)

Less., based on Othonna digitata L., for a similar taxon

with toothed leaves, followed by a list of names that he

regarded as conspecific with it. Among them was Othonna

lingua sensu Jacquin (1797). Jacquin (1797) was in any

event also not the author of the name since he cited the

Systema vegetabilium of 1784 (edition 14, Murray 1784),

which in turn refers the name to Linnaeus fil. (1782). This

is in fact the first appearance of the name, which is thus

correctly attributed to Linnaeus f. as O. lingua L.f.

There is no material under the name Othonna lingua in

either LINN or S but there is a specimen under this name

in the Thunberg herbarium (UPS-THUNB20882), which

therefore represents the type. This specimen was cited by

Harvey (1865: 342) under O. tuberosa Thunb. (itself an

illegitimate, superfluous name for O. bulbosa L.), who

followed De Candolle (1838: 471 ) in retaining the name

[Doria lingua (Jacq.) Less.] as applied to a quite differ-

ent taxon with disciform capitula. We have examined

the specimen of Doria lingua in the Thunberg herbarium

and find no reason to disagree with Harvey (1865) that it

represents O. bulbosa L. The name O. lingua L.f. should

therefore be treated as a synonym of O. bulbosa L. and

we make the necessary nomenclatural correction below.

The incorrect application of the name Othonna lingua

by De Candolle (1838) and Harvey ( 1 865 ) arose from an

initial misunderstanding by Jacquin (1797), whose illus-

tration and description apply to an erect-stemmed, tuber-

ous geophyte with petiolate radical leaves, lanceolate

cauline leaves, and disciform capitula.

Linnaeus f. (1782) did not mention the condition of

the marginal florets in the protologue of Othonna lingua

but neither did he mention the florets in any but one of

the remaining 14 species enumerated by him in Othonna.

Since all of these are in fact radiate, this is strong cir-

cumstantial evidence that he had a radiate taxon in mind

when describing O. lingua. There is also no indication as

to whether the epithet refers to the shape of the leaves or

to the presence of rays but the description of the leaves

as ovate-lanceolate and semi-amplexicaul suggests that

it more probably alludes to the latter. In any event, the

Thunberg collection, which is evidently the type, fixes

the application of the name. We accordingly place O.

lingua in synonymy under O. bulbosa.

Othonna bulbosa L., Species plantarum: 1309

( 1 753). Type: illustration in Breyne: t. 66 (1674-1678).

O. lingua L.f.: 387 (1782 ‘1781’), syn. nov.; Jacq.: 60 (1797), name

only, excl. description and figure; Thunb.: 718 (1823); Sch.Bip.: 769

(1844) [as ‘(Less.) Sch.Bip.’]. Doria lingua (L.f.) DC.: 471 (1838).

Type: South Africa, without precise locality, UPS-THUNB20882 (UPS-

THUNB, presumed holo.-microfiche!).

O. bulbosa Thunb.: 720 (1823), illegit. superfl. name. Type: as for

O. bulbosa L.

4. Othonna lingua is a species with radiate capitula

and is, according to our interpretation of the type, conspe-

cific with O. bulbosa (see discussion above). It is, there-

fore, necessary to consider the identity of the disciform

species to which this name has been applied in many her-

baria. The collections under this name represent a tuber-

ous geophyte with disciform capitula and ± erect, annual

stems bearing variously oblanceolate to ovate leaves, the

40

Bothalia 40,1 (2010)

lowermost leaves ± tapering to the base and ± truncate or

sessile, and the upper leaves mostly auriculate.

We have examined herbarium material filed under

the name O. lingua and are able to distinguish two sets

of populations. One represents a western, coastal spe-

cies occurring from Namaqualand to the Olifants River,

mostly with fleshy, oblanceolate leaves with conspicu-

ously revolute margins, capitula in which the phyllaries

are connate in the basal third, and disc cypselae with

short, caducous pappus bristles 1-3 mm long. A sec-

ond series of plants with ± plane leaves occurs inland,

from the Cedarberg eastwards to Port Elizabeth, and is

distinguished by the phyllaries connate for ± half their

length or more, and most strikingly by the disc cypselae

entirely lacking a pappus (very rarely with one or two

short bristles on the outer disc florets in some collections

from the Little Karoo).

Only two species of Othonna are known in which the

disc florets lack a pappus and both were described by De

Candolle (1838). O. gymnodiscus (DC.) Sch.Bip. was

based on a plant collected by Ecklon near Port Elizabeth,

and O. semicalva (DC.) Sch.Bip. on a collection made

by Drege in the Olifant’s River Valley. The two taxa

were distinguished primarily by differences in leaf shape

(respectively oblong-ovate and auriculate vs linear-lan-

ceolate and sessile). Both taxa were known to De Can-

dolle (1838) from the type specimens only. Collections

made since then have filled in the distribution between

the type localities and also suggest that the purported dif-

ferences between them in leaf shape and size of capitula

are not significant. Leaves in herbarium material range

from narrrowly lanceolate to obovate, with the leaf base

narrowed or± auriculate, and the phyllaries vary from 6-

1 0 mm long, sometimes even on the same plant depend-

ing on their stage of development. Our examination of

type material of both names leads us to conclude that they

represent forms of a single species, for which we choose

the name O. gymnodiscus as being most appropriate. The

name O. semicalva is accordingly reduced to synonymy.

The second series of populations from the west coast

with pappus bristles present in the disc florets appears

to represent an undescribed species but further study is

required to assess this.

Othonna gymnodiscus (DC.) Sch.Bip ., Composi-

tae Kraussianae: 769 (1844). Doria gymnodiscus DC.:

473 (1838). Type: South Africa, [Eastern Cape], near

Port Elizabeth, without date, Ecklon & Zeyher (G-DC,

holo.-microfiche!; S-digital image!).

O. semicalva (DC.) Sch.Bip.: 769 ( 1 844), syn. nov. Doria semicalva

DC.: 473 (1838). Type: South Africa, [Western Cape], Langevaley ad

Olifantsrivier, [Rhinosterfontein, 15 July 1830], Drege [2878] (G-DC,

holo.-microfiche!; P-Aluka image!, website accessed 20-03-09).

O. lingua sensu Goldblatt & Manning (2000), non L.f. [= O. bul-

bosa L.].

Selected additional specimens examined

WESTERN CAPE.— 3218 (Clanwilliam): Pakhuis Pass, (-BB),

7 July 1984, Taylor 10985 (NBG). 3219 (Wuppertal): Swartrug-

gens, Knolfontein Farm, (-DC), 7 August 2007, Jardine & Jardine

699 (NBG). 3318 (Cape Town): Malmesbury, Katzenberg, (-DA), 26

June 1976, Andrag & Boucher 5 (NBG); McGregor, Farm Vrolikheid,

(-DD), 1971 without month, Jooste 88 (NBG). 3319 (Worcester):

Rawsonville, (-CA), 28 July 1962, Walters 652 (NBG); Karoo Garden,

(-CB), 12 July 1948, Compton 20525 (NBG); Karoo Garden, (-CB),

30 June 1958, Willems 47 (NBG); Karoo Garden, (-CB), 18 August

1976, Bayer 184 (NBG); Worcester Veld Reserve, (-CB), 13 August

1986, Midgley & Bosenberg 50 (NBG). 3321 (Ladismith): Calitzdorp,

Sandberg, (-DA), 24 July 1953, Compton 24101 (NBG). 3322 (Oudts-

hoom): Robinson Pass, (-CC), 30 July 1947, Compton 19620 (NBG).

5. Only three species of Othonna are known with pur-

ple or magenta rays, all of them tuberous geophytes from

Namaqualand. They are distinguished essentially by the

shape of the leaves, specifically the degree of lobing. At

the time that Harvey (1865) established O. incisa Harv. and

O. rosea Harv. as species distinct from O. cakilefolia DC.,

he knew all three taxa from just a single collection each.

With further collecting it appears that O. incisa and O.

rosea represent extreme forms that grade into one another.

Plants from the Richtersveld southwards to Springbok have

leaves that are entire, sparsely denticulate or dentate (typi-

cal O. rosea), whereas those from the Kamiesberg south-

wards to Nuwerus have sharply pinnatifid leaves (typical

O. incisa). The difference in lobing appears to be one of

degree and not of kind. The phyllaries in O. rosea were

described by Harvey ( 1865) as ‘semiconcrete’ and those of

O. incisa as ‘connate’. In the material of O. rosea that we

have examined, the phyllaries range from basally connate

to joined up to halfway, and a similar variation is evident in

the material of O. incisa ( Hall 4932), in which the degree

of fusion ranges from one quarter to two thirds of the

length of the phyllaries. Although the cypselae of O. rosea

were described as glabrous, this observation was based on

immature fruits, and in such cases the hairs are usually not

readily seen. More recent collections of plants with entire

leaves (e.g. Hall 575 NBG) develop pubescent achenes.

In our opinion, O. rosea and O. incisa represent extreme

forms of a single species and should be treated as one. The

name O. rosea is chosen as being the more descriptive.

Collections of Othonna cakilefolia that we have

examined have leaves very deeply dissected into nar-

row, ± obtuse lobes, thus differing significantly from the

lacerate-pinnatifid forms of O. rosea , which always have

distinctly apiculate lobes.

Othonna rosea Harv. in Harv. & Sond., Flora cap-

ensis 3: 341 (1865). Type: South Africa, [Northern Cape,

Springbok], Modderfontein, without date, Whitehead s.n.

(TCD, holo.-Aluka image!, website accessed 20-03-09).

O. incisa Harv.: 341 ( 1865), syn. nov. Othonna cakilefolia var. lati-

folia DC.: 482 (1838). Type: South Africa, [Western Cape, Nuwerus],

between Uitkomst and Geelbekskraal, without date [August 1830],

Drege (Doc. 95.2) (G-DC, holo.-microfiche!; P-Aluka image!, web-

site accessed 25-1 1-09).

Additional specimens examined

NORTHERN CAPE. — 2817 (Vioolsdril): hills east of Perdewater,

(-CA), 11 July 1997, Bruyns 7268 (NBG). 2917 (Springbok): Harrasberg,

(-AD), 16 July 1995, Bruyns 6344 (BOL); Anenous Pass, (-BA), 30 May

1954, Hall 910 (NBG); west of Bulletrap on road to Waalheuwel, (-BC),

12 May 2005, Narrower 1500 (NBG); 5 miles [8 km], north of Grootmis,

(-CA), 20 August 1952, Hall 575 (NBG); Komaggas, (-CD), 5 July 1991,

Bruyns 4614 (BOL); Spektakel Pass, (-DB), 15 May 2003, Manning 2923

(NBG); 10 km west of fool of Spektakel Pass, (-DA), 4 July 1987, Bru-

yns 2705 (NBG); Messelpad, (-DC), 23 July 1948, R.H. Compton 20672

(NBG); Kourkammaberg, (-DC), 1 1 July 1987, Bruyns 3876 (BOL). 3017

(Hondeklipbaai): Riethuis, (-AB), II July 1989, Bruyns 3860 (BOL).

3018 (Kamiesberg): west slopes of Rusbospoort on Farm Damsland, (-

AC), 23 April 1982, Hall 4932 (NBG); Andrieskloof, (-CB), 24 August

1991, Bruyns 4750 (BOL); Knersvlakte, Bruinberg, (-DC), 10 September

1992, Bruyns 5322 (BOL).

Bothalia 40,1 (2010)

41

6. Othonna retrorsa DC. is an easily recognizable,

cushion-forming perennial with a many-headed caudex

producing rosettes of very distinctive, leathery, reticu-

late-veined leaves. These are oblanceolate with carti-

laginous margins bearing few to many pale, patent or

retrorse denticles. The leaves are persistent at the base

and their dried remains accumulate around the short

stems, gradually decaying into a fibrous mass. One or

more, sparsely branched flowering stems are produced

from each rosette. The species was described by De Can-

dolle (1838) from material collected in the Kamiesberg

by Drege. Plants from further north, around Springbok,

were later treated by Compton (1953) as the separate

species O. spektakelensis. Differences between the two

taxa are slight and are centred on the relatively broader

leaves and more northerly distribution of O. spektakelen-

sis. Other listed differences include the average number

of phyllaries and ray florets but these are inconsistent

even within the type material. The taxon was subse-

quently treated as merely a variety of O. retrorsa (Row-

ley 1990). A collection with smaller, narrower leaves

made by Drege at an unlocalized site in Namaqualand

was described as var. linearifolia DC. (1838). Material

matching the description has subsequently been collected

from Ezelskop near Leliefontein in the Kamiesberg (Le

Roux & Ramsey 760 NBG) and from near Kamieskroon

(. Hall s.n. NBG462/52 NBG). These plants were grow-

ing in cracks in exposed granite sheets, which is con-

sistent with their more compact and dwarf habit. This

habitat struck Compton (1953) as significantly differ-

ent from that of O. retrorsa , which he knew from rocky

slopes in somewhat deeper soils, but subsequent collec-

tions of typical O. retrorsa have been made from bare

rock cracks and sandy depressions on granite ( Oliver

5965 NBG) and the variation in growth form is evidently

purely ecological. The salient differences between the

three varieties are given in Rowley (1994). Additional

collections (Le Roux 3391 NBG) of O. retrorsa from

Ezelsfontein, the type locality of O. spektakelensis, have

the relatively narrow leaves of the type variety and there

seems to be nothing to be gained by continuing to distin-

guish varieties in the species. Indeed, cultivated plants of

var. retrorsa and var. spektakelensis illustrated by Row-

ley (1994: 192) are vegetatively indistinguishable.

Another similar species, Othonna zeyheri Sond. ex

Harv., was based on Zeyher 996 collected near Kliprand

at the southeastern end of the Kamiesberg. The status of

this taxon in relation to O. retrorsa was questioned by

Harvey himself (1865), and Rowley (1994) later ten-

tatively included it in O. retrorsa var. spektakelensis.

Compton (1953), however, was convinced that it was

a good species, equating it with Hall’s collection from

near Kamieskroon mentioned above. Examination of the

ample isotype material shows that this taxon matches

typical O. retrorsa and cannot be retained at any level.

Othonna retrorsa DC., Prodromus compositae

6: 479 (1838). Type: South Africa, [Northern Cape],

Kasparskloof, 19 August 1839, Drege 2726 (G-DC,

holo.-microfiche!; P-Aluka image!, website accessed

25-11-09).

O. retrorsa var. linearifolia DC.: 479 (1838), syn. nov. Type: South

Africa, Namaqualand without precise locality, Drege (G-DC, holo.-

microfiche!).

O. zeyheri Sond. ex Harv.: 339 (1865), syn. nov. Type: South

Africa, [Western Cape, Kliprand], between Lieslap and Hofkraal, 1846

without month, Zeyher 996 (S, holo.; BOL!, BM-Aluka image!, web-

site accessed 25-11-09, P, SAM!).

O. spektakelensis Compton: 118 (1953), syn. nov. O. retrorsa

var. spektakelensis (Compton) G.D. Rowley: 282 (1990). Type:

South Africa, [Northern Cape], Ezelsfontein, between Springbok and

Spektakel Pass, 8 September 1950, H. Hall 141 (NBG, holo.!).

Additional specimens examined

NORTHERN CAPE. — 2917 (Springbok): Eselsfontein, between

Springbok and Spektakel, (-DA). 21 August 1953, Hall NBG771/49

(NBG); Farm Ezelfontein, between Tierhoek and Sandhoogte, (-DA),

3 September 1985, Le Roux 3391 (NBG); Spektakel Mountain, (-DA),

26 September 1974, Goldhlatt 2805 (NBG); Komaggas, Koufontein-

berg, (-CD), October 1933, Herre 2929 (NBG). 3017 (Hondeklipbaai):

Grootvlei, (-BB), 26 August 1954, Barker 8413 (NBG); Kamieskroon,

(-BB), July 1952 [fruiting]. Hall NBG462/52 (NBG); 10 miles [16

km] south of Kamieskroon, (-BB), 7 September 1945, Compton 17304

(NBG). 3018 (Kamiesberg): west side of Eselkop Mountain, (-AC),

13 October 1981, Le Roux & Ramsey 760 (NBG); Kamiesberg, Farm

Welkom, (-AC), 31 August 1975, Oliver 5965 (NBG); Boplaas Farm,

near top of Studer’s Pass, (-AC), 30 August 2004, Helme 3258 (NBG).

7. Cacalia rigida Thunb. is a thorny shrublet with obo-

vate, sparsely denticulate leaves and shortly pedunculate,

disciform capitula (Thunberg 1823). The species was

transferred to Kleinia by De Candolle ( 1 838) on the basis

of the description alone, with the result that the taxon

was redescribed by him in the same publication under the

name Doria spinescens DC. from a collection of Drege’s

from the Sneeuberg. In a reversal of the situation, Har-

vey (1865), who had access to the Thunberg herbarium

but not to the Drege specimen, was nevertheless confi-

dent that they represented the same species and therefore

transferred Thunberg’s name to Doria, placing De Can-

dolle’s name in synonymy under it. We have been able to

examine both relevant types for the first time and confirm

Harvey in his decision. Schultz (1944), who included

Doria in Othonna, later transferred D. spinescens to Oth-

onna under the new name O. rhamnoides Sch.Bip. as De

Candolle’s epithet was preoccupied in the genus. Thun-

berg’s Cacalia rigida is actually the earliest name for the

taxon but this epithet is also preoccupied in Othonna, and

Schultz’s new name is thus the correct name for the spe-

cies in Othonna. The species, which is rarely collected,

has been overlooked in southern African checklists for

the family under any of its available names (Welman

2006). We provide the complete nomenclature here.

Othonna rhamnoides Sch.Bip., Compositae Kraus-

sianae: 769 (1944) [replacement name for Doria spines-

cens DC.: 470 ( 1838), non Othonna spinescens DC.: 332

(1838)]. Type: South Africa, [Eastern Cape], Sneeuberg,

[September 1829], Drege [619] (G-DC, holo.-micro-

fiche!; K-Aluka image!, website accessed 25-1 1-09).

Doria rigida (Thunb.) Harv.: 322 (1865), non Othonna rigida DC.:

476 (1838) [= Othonna amp/exicaulis Thunb.]. Cacalia rigida Thunb.:

624 (1823). Kleinia rigida (Thunb.) DC.: 338 (1838). Type: South

Africa, without precise locality or date, UPS-THUNBI8765 (UPS-

THUNB, holo.-microfiche!).

SENECIO

8. Senecio albopunctatus Bolus (1887) was described

from a single gathering of several plants collected in

Namaqualand at Klipfontein, west of Steinkopf on the

old Steinkopf-Port Nolloth railway. S. albopunctatus

42

Bothalia 40,1 (2010)

is a subshrub with terminal tufts of pinnatifid leaves, the

lobes tipped with characteristic pale thickenings or calli

that give the species its name, and solitary, radiate capitula

on long peduncles. The plant was described as glabrous but

careful examination of the type material shows the leaves

and peduncles to be scantily clad in minute, sessile glands.

No similar plants with radiate capitula have been collected

since then but several collections have been made of plants

that are vegetatively identical to S. albopunctatus but differ

from it in having discoid capitula. The first of these collec-

tions was made in 1935 by R.H. Compton at Klipfontein

hill, west of Steinkopf (Compton 5442 NBG), at the type

locality of S. albopunctatus. ft was identified tentatively by

Compton as S. albopunctatus following careful comparison

with the type, with the comment that apart from the slightly

less indurated leaflet tips, he could find no significant dif-

ferences between the two species and that he, therefore,

interpreted his material as a rayless variant of S. albopunc-

tatus. Since then several additional collections of this ray-

less form have been made in the Richtersveld, all of them

agreeing exactly with Bolus’s and Compton’s original gath-

erings and also proving very good matches with the type of

S. albopunctatus , apart from the clear absence of rays.

The occurrence of both discoid and radiate forms

in a single species is rare in Senecio but not unknown.

Among species from the Cape Floristic Region (CFR) it

has been recorded in S. agapetes C. Jeffrey and S. cris-

pus Thunb. (Goldblatt & Manning 2000), and in several

species from KwaZulu-Natal, including S. conrathii

N.E.Br., S. hypochoerideus DC., S. polyodon DC. and

S. poseideonis Hilliard & B.L.Burtt (Hilliard 1977).

Such species are scattered throughout the genus. In most

instances one of the forms is dominant, with the other

rare or occasional, but in a few species both forms are

common. A strikingly similar example is provided by S.

erosus L.f., in which both radiate and discoid plants have

been collected near Mooreesburg (Helme 2131 [dis-

coid] and Helme 2339 [radiate] NBG) in this otherwise

entirely radiate species. We have no hesitation, therefore,

in following Compton in treating both discoid and radi-

ate forms as a single species.

However, it is now clear that the discoid material of

Senecio albopunctatus from the Richtersveld is indistin-

guishable from S. maydae Merxm. (1960), which was

based on several collections from the Huib Hoch Plateau

and adjacent hills in southern Namibia (Dinter 1932).

Of particular significance is the presence of unusual,

sessile glands on the leaves, peduncles and phyllaries

that render the surface of the plant sticky, evidenced in

some specimens by a conspicuous load of adhering sand

grains. The name S. maydae is accordingly placed in

synonymy under S. albopunctatus .

Senecio albopunctatus Bolus in Botanical Jour-

nal of the Linnean Society 24: 177 (1887). Type:

South Africa, [Northern Cape], Namaqualand minor

[Namaqualand], Klipfontein, September 1883, Bolus 423

(BOL, holo.!; K, P-Aluka images!, website accessed 25-

11-09, SAM!).

S. maydae Merxm.: 608 (1960) [replacement name for S. longipe-

dunculatus Dinter: 93 (1932)], syn. nov. Syntypes: Namibia, [Liider-

itz], Garub, Tigerberg, 17 October 1922, Dinter 4107 ( WIND, SAM!);

Namibia, Bucluiberge [Boegoeberg], 28 June 1929, Dinter 6464

(NBG!, PRE, SAM!, WIND).

Additional specimens examined

NORTHERN CAPE. — 2816 (Oranjemund): Richtersveld, Hels-

kloof area northeast of Khubus, (-BD), 29 August 1977, Oliver, Tolken

& Venter 348 (NBG); Vandersterrsberg, (-AC), 30 July 1933, Van der

Wait 296 (NBG). 2817 (Vioolsdrif): Kodaspiek, (-AA), 2 Septem-

ber 1977, Oliver, Tolken & Venter 450 (NBG); south of Van Zylsrus,

(-CB), 4 September 1977, Thompson & Le Roux 324 (NBG). 2917

(Steinkopf): Klipfontein koppie, (-BA), 29 August 1935, Compton

5442 (NBG).

9. Among the 20 annual species of Senecio recog-

nized by Harvey (1865), are five with mauve or pur-

ple ray florets. Two of these species are well known: S.

elegans L. is essentially a species of sand dunes along

the western and southern Cape coast, from Saldanha in

Western Cape to Port Alfred in Eastern Cape, whereas

S. arenarius Thunb. is widely distributed on sandy and

gravelly flats and in washes along the west coast and

interior, from central Namibia through much of the west-

ern half of South Africa as far south as Agulhas in West-

ern Cape (Goldblatt & Manning 2000). Both are ± glan-

dular-pubescent herbs, extremely variable in leaf form

[polymorphous is how they are described by Harvey

(1865)] but readily separable by their different involu-

cres: ± cylindrical and with a few subulate bracteoles

in S. arenarius ; ± campanulate and closely enveloped at

the base by several imbricating, lanceolate, black-tipped

bracteoles in S', elegans. The remaining three species in

the group were known to Harvey from the type speci-

men or description alone and they remain poorly known

today.

One of them, Senecio cakilefolius DC., was based on

a collection made by Drege at Silwerfontein, southeast of

Springbok. This material was not seen by Harvey ( 1 865),

who relied entirely on De Candolle’s (1838) description.

The species was distinguished from S. arenarius by its

glabrous stem and leaves, and supposedly larger capitula

but examination of the type collection confirms that the

plants are actually sparsely but quite evidently glandular-

pubescent and the capitula are no larger than commonly

encountered in S. arenarius. The name has subsequently

been applied rather indiscriminately to any arenarius-

like plants with less than the usual pubescence. Subgla-

brous or thinly pubescent plants of the cakilefolius type

are common between Springbok and Kamiesberg but

also occur further south near Clanwilliam and through

the Tanqua River Basin to Whitehill, whereas more

densely pubescent plants of the arenarius type are wide-

spread. With a full range of material from Namaqua-

land and the West Coast now available, it is clear that

there are any number of intermediate conditions from

almost glabrous plants to those with sparsely glandular-

pubescent stems and leaves to densely glandular-pubes-

cent plants. The size of the capitula also varies greatly

and independently of the vestiture. Populations from

Namaqualand and the Bokkeveld Plateau have slightly

larger capitula with phyllaries 5-7 mm long compared

with those from the Cold Bokkeveld and Little Karoo, in

which the phyllaries are 4-6 mm long, but this small size

difference is not correlated with other differences, and

is not uncommonly encountered in other species in the

genus. With the ample collections now at our disposal,

it appears to us that the concepts of S. arenarius and S.

cakilefolius represent the extremes of a continuous range

of variation and we thus treat them as a single species.

Bothalia 40,1 (2010)

43

Senecio arenarius Thunb., Prodromus plantarum

capensium: 158 (1800b). Type: South Africa, without

precise locality or date, Thunberg UPS-THUNB 19545

(UPS-THUNB, holo.-microfiche!).

5. cakilefoliits DC.: 408 (1838), syn. nov. Type: South Africa,

[Northern Cape], Silverfontain [Silwerfontein, September 1830],

Drege [ 2816] (G-DC, holo.-microfiche!; P-Aluka image!, website

accessed 25-11-09).

10. Senecio pearsonii Hutch. (1917) was based on a

collection from the Kamiesberg and diagnosed against

5. hypochoerideus DC., from which it was distinguished

by its more finely serrate leaves and striate-papillate, as

opposed to uniformly pubescent, achenes. Actually, the

achenes of S. hypochoerideus, like those of most species

of Senecio , are striate-hispid (Hilliard 1977). Examina-

tion of the type of S. pearsonii confirms Hutchinson’s

opinion of the Kamiesberg material in respect of S. hypo-

choerideus but shows his species to be a perfect match

for the closely allied S. asperulus DC. This species dif-

fers primarily from S. hypochoerideus in its narrower

leaves 2-10 mm wide, with more finely serrate margins

with simple teeth vs broader, doubly-serrate leaves 10-

40 mm wide in S. hypochoerideus (Hilliard 1977). At

the time of Hilliard’s (1977) treatment of the genus in

Natal, S. asperulus was known from the escarpment of

the Eastern Cape northwards along the Drakensberg into

the Highveld but recent collections have extended its

range westwards through the Nieuweveld Mountains and

along the western escarpment as far as the Hantamsberg.

The inclusion of S. pearsonii in S. asperulus is a natural

extension of the range along the western escarpment to

the Kamiesberg.

Senecio asperulus DC., Prodromus systematis

naturalis regni vegetabilis 6: 386 (1838). Type: South

Africa, [Eastern Cape], Albany, Ecklon [53] (G-DC,

lecto., here designated-microfiche!; P-Aluka image!,

website accessed 25-11-09).

S. pearsonii Hutch, in Pearson & Hutchinson: 398 (1917), syn.

nov. Type: South Africa, [Northern Cape], Khamiesberg [Kamiesberg],

southwest of Leliefontein, 16 January 1911, Pearson 6310 (K, holo-

Aluka image!, website accessed 25-11-09).

Additional specimens (Greater CFR) examined

NORTHERN CAPE. — 3119 (Calvinia): Hantamsberg, Van Rhyns-

hoek Farm. (-BD), 18 August 1976, M.F. Thompson 2349 (NBG); 10

October 1983, M. Thomas 41a (NBG). 3220 (Sutherland): Sutherland,

(-BC), 8 July 1968, F. Stayner s.n. NBG87.402 (NBG). 3221 (Merwe-

ville): Nuweveldberge, Bok se Plaas, (-BA), 26 February 1986, Mof-

fett & Steensma 401 7 (NBG).

11. Senecio carroensis DC. (1838) was based on a

collection made by Drege along the southern margin

of the Great Karoo at Kendo [Kendouslaagte] between

Klaarstroom and Willowmore. The species is a slen-

der, laxly branched shrublet with deeply incised, almost

bipinnatisect leaves, the lobes narrow and deeply

toothed. The sparsely woolly or subglabrous branches

are characteristically yellowish striate when young,

often flushed purple when older. The radiate, yellow

capitula are in lax corymbs, with glabrous phyllaries.

Ample material of the taxon has now been collected

from the drier mountains of the Little Karoo westwards

to Karoopoort and thence northwards through the Cold

Bokkeveld and Swartruggens as far as the Bokkeveld

Mountains. This material displays significant variation

in the shape and size of the leaf lobes, a feature that was

already evident to De Candolle. Although mostly nar-

rowly oblong or linear and 2-7 mm long, the lobes in

some collections are much reduced, almost quadrate, and

1-2 mm long. These plants are an exact match for Sene-

cio parvifolius DC., another of Drege’s collections, from

the Kamiesberg in Namaqualand, and which was distin-

guished from S. carroensis essentially by its smaller leaf

lobes. Further collections from the Kamiesberg confirm

the general constancy of this leaf character among the

Namaqualand plants but also include plants in which

the leaf lobes are longer and narrower and thus indistin-

guishable from more xeromorphic forms of S. carroen-

sis. With this larger range of material now available, we

conclude that these two species represent extreme leaf

forms in a single species.

Senecio carroensis DC., Prodromus systematis

naturalis regni vegetabilis 6: 396 (1838). Type: South

Africa, [Eastern Cape], Cairo [Willowmore], Kendo, [22

June 1829], Drege [5910] (G-DC, holo.-microfiche!; K,

P-Aluka images!, website accessed 25-1 1-09).

S. parvifolius DC.: 396 (1838), syn. nov. Type: South Africa,

[Northern Cape], Kamiesberg, Modderfontein, [4 November 1830],

Drege [2820] (G-DC, holo.-microfiche!; K, P-Aluka images!, website

accessed 25- 1 1 -09, SAM ! ).

Additional specimens examined

NORTHERN CAPE.— 2917 (Springbok): Brakdam, (-BC), 24

July 1941, Compton 11085 (NBG). 3018 (Kamiesberg): Kamiesberg,

Farm Welkom, (-AC), 16 October 1954, Esterhuysen 23680 (NBG);

koppie east of Rooiberg, (—AC), 14 October 1981, Le Roux & Ramsey

791 (NBG); between Garies and Leliefontein, (-AC), November 1939.

Esterhuysen 1399 (NBG). 3118 (Vanrhynsdorp): Gifberg, (-DC), 2

September 1948, Compton 20777 (NBG); Nardouw, (-DC), 6 Sep-

tember 1951, Compton 22833 (NBG). 3119 (Calvinia): Oorlogskloof

Nature Reserve, (-AC), 19 September 1996, Pretorius 353 (NBG); 20

September 2000, Pretorius 508 (NBG); top of Botterkloof Pass, (-CD),

24 August 1950, B. Maguire 188 (NBG).

WESTERN CAPE. — 3219 (Wuppertal): Cedarberg, Matjiesrivier

Reserve, (-CB), 4 October 1997, Lechmere-Oertel 749 (NBG); Swar-

truggens, Knolfontein, (-DC), 1 5 September 2006, Jardine & Jardine

448 (NBG). 3319 (Worcester): Karoopoort, (-BA), 28 September

1951, Compton 22932 (NBG). 3320 (Montagu): Laingsburg, Witte-

bergskloof, (-BC), 16 July 1923, Compton 2502 (NBG); eastern end of

Anysberg, (-DA), 6 October 1982, Van Zyl 33 72 (NBG); Touwsberg,

(-DB), 17 September 1993, Snijman 1336 (NBG); 7 October 1993,

McDonald 2442 (NBG). 3321 (Merweville): Ladismith, Seweweeks-

poort. (-AD), 13 September 1938, Compton 7877 (NBG).

12. Among the taxa with yellow, radiate capitula that

were included in Harvey’s (1865) Sinuosi are two spe-

cies characterized by a short, vertical rhizome closely

covered by imbricating leaves, with the base of the peti-

oles expanded and encircling the rhizome. The invest-

ing leaf bases form an almost bulb-like structure, with

their inner faces densely covered with woolly hairs that

also run up along the margin of the sheath and onto the

petiole for a short distance. These woolly rhizomes set

these two species apart from others in the group but the

difference between them is not clear. Senecio eriobasis

DC. (1838), based on a collection from Worcester, was

distinguished from S. erosus L.f. (1782) by its glabrous

vs scabro-pubescent leaves and involucre. Like many

other species of Senecio, S. erosus is extremely variable

in the degree of development of the vestiture, varying

from sparsely to densely pubescent, with no clear dis-

tinction between the two conditions. More significantly,

however, examination of the type of S. eriobasis shows

44

Bothalia 40,1 (2010)

the leaves and peduncles to be distinctly pubescent and

quite indistinguishable from those of S. erosus , and there

is no doubt that Goldblatt & Manning (2000) were cor-

rect in their conclusion that the two could not be sepa-

rated. We accordingly formally include S. eriobasis in

the synonomy of S. erosus. The species is distributed

from Namaqualand to the southern Cape.

Examination of the type of Doria incisa Thunb.,

treated as a synonym of Senecio erosus by Harvey

(1865), confirms that it has radiate capitula, despite its

initial placement in Doria , and that it cannot be distin-

guished from S. erosus. Harvey (1865) also included in

S. erosus the species described by Thunberg (1823) as

Cineraria pandurata Thunb. (later transferred to Sene-

cio as S. panduratus (Thunb.) Less.), but he explicitly

excluded De Candolle’s (1838) application of the name.

De Candolle (1838) based his interpretation of S. pandu-

ratus on several specimens in G-DC. These represent a

taxon very similar to S. erosus in leaf and inflorescence

but differing from it in the rootstock, which is a ± hori-

zontal rhizome bearing a lax tuft of leaves that are not

strongly widened at the base and lacking the distinctive

woolly vestiture in the axils and along the base of the

petiole that is such a striking feature of S. erosus. Harvey

(1865) considered that De Candolle’s plants represented

S. hastatus but this is not the case (see below under S.

robertiifolius).

There are two specimens labelled Cineraria pandu-

rata in UPS-THUNB. One of them, a complete plant

with the diagnostic erect, villous rhizome of Senecio ero-

sus, is also labelled Doria incisa and constitutes the type

of that name. The second specimen, securely identified

by the locality data as the type of Cineraria pandurata ,

comprises just an inflorescence and a single, unattached

basal leaf. This incomplete specimen, in our opinion,

might equally be assigned either to S. erosus or to the

taxon that is currently known under the name S. pan-

duratus but Thunberg ’s (1823) application of the name

is quite clear. His phrase ‘ caulis erectus, striatus, villo-

sus ’ is essentially identical to his description of the stem

of Doria incisa as ‘'caulis ... flexuoso-erectus, simplex,

teres, striatus, pubescens' and clearly indicates S. erosus.

We thus have no hesitation in following Harvey (1865)

in treating S. panduratus as a synonym of S. erosus.

Although mostly radiate, occasional plants have dis-

coid capitula, sometimes with both forms in the same

population (e.g. Helme 2131, 2339 NBG).

Senecio erosus L.f, Supplementarum plantarum:

370 (1782 ‘1781’). Type: South Africa, without local-

ity or date, Thunberg 446 (LINN996.72, holo.-Linnean

Society of London image!, website accessed 25-1 1-09).

Cineraria incisa (Thunb.) Willd.: 2074 (1803). Doria incisa

Thunb.: 156 (1800b). Type: South Africa, without precise locality or

date, Thunberg UPS- T H UNB 1 983 0 (UPS-THUNB, holo.-micro-

fichel).

S. panduratus (Thunb.) Less.: 392 (1832). Cineraria pandu-

rata Thunb.: 672 (1823). Type: South Africa, [Western Cape], Kam-

tous [Gamtoos] River, December without year, Thunberg UPS-

THUNB19934 (UPS-THUNB, holo.-microfichel).

S. eriobasis DC.: 388 (1838), syn. nov. Type: South Africa, [West-

ern Cape|, Worcester, without date, Ecklon [1901] (G-DC, holo-

microfiche; P-Aluka image!, website accessed 25-1 1-09).

Selected additional specimens examined

NORTHERN CAPE. — 3017 (Hondeklipbaai): Kamieskroon,

(-BB), 29 August 1937, Compton 6799 (NBG); Grootvlei, (-BB), 7

September 1945, Compton 17284 (NBG); Darter’s Grave, (-BD), 2

September 1951, Maguire 970 (NBG). 3018 (Kamiesberg): Welkom,

near Caries, (-AB), 16 October 1954, Esterhuysen 23666 (BOL) [dis-

coid capitula]. 3119 (Calvinia): top ofVanrhyn’s Pass, (-AC), 1 Octo-

ber 1947, Taylor 2864 (NBG); west of Nieuwoudtville on road to Van-

rhyn’s Pass, (-AC), 23 August 1950, Barker 6453, Midddlemost 1603

(NBG); Nieuwoudtville Reserve, (-AC), 8 September 1983, Peny &

Snijman 2333 (NBG); Oorlogskloof Nature Reserve, (-AC), 19 Sep-

tember 1995, Pretorius 298 (NBG); Karigabosch Fountain, (-DD), 20

August 1975, Thompson 2468 (NBG); Kareehout River, south of Per-

dekloof, (-DD), 21 August 1975, Thompson 2520 (NBG). 3220 (Suth-

erland): 11 km east of Sutherland, (-BD), 22 September 1985, Mof-

fett 3774 (NBG). 3319 (Worcester): Tweedside, (-AB), 25 September

1932, Compton 4000 (NBG).

WESTERN CAPE. — 3118 (Vanrhynsdorp): Nardouw, (-DC), 6

September 1 95 1 , Johnson 242 (NBG). 3217 (Vredenburg): Vredenburg,

Trekossenkraal, (-DD), 28 September 2002, Boucher 6992 (NBG).

3218 (Clanwilliam): near Clanwilliam, (-BB), 20 July 1941, Compton

10996 (NBG). 3219 (Wuppertal): Pakhuis, (-AA), 18 September 1937,

Compton 6951 (NBG); 13 September 1947, Barker 4693 (NBG); Nieu-

woudt’s Pass, (-AC), 3 September 1982, Le Maitre 322 (NBG); Matjies-

rivier, (-AD), September 1943, Wagener 183 (NBG); Kromrivier Farm,

(-CA), 15 September 1984, Taylor 11050 (NBG); Gonnafontein Farm,

(-CB), 2 September 2000, Pond 125 (NBG); Swartruggens, Knolfon-

tein, (-DC), 15 September 2006, Jardine & Jardine 452 (NBG). 3318

(Cape Town): Mooreesburg, (-BA), [discoid capitula], 31 August 2001,

Helme 2131 (NBG); 12 September 2001, Helme 2339 (NBG); Stellen-

bosch, (-DD), 13 September 1989, Buys 30 (NBG). 3319 (Worcester):

Worcester, (-CB), 23 August 1976. Bayer 202 (NBG); near Rusbos,

(-CC), 29 August 1979, Hugo 1824 (NBG). 3320 (Montagu): Bantams,

(-BA), 27 October 1941, Compton 12216 (NBG). 3321 (Ladismith):

Gamka Mountain Nature Reserve, (-CB), 15 August 1983, Cattell 291

(NBG). 3322 (Oudtshoom): George, (-AC), September 1932, Four-

cade 4709 (BOL). 3418 (Simon’s Town): Cape of Good Hope Nature

Reserve, (-AD), 17 September 19709, Taylor 7792 (NBG); Greyton

Nature Reserve, (-BA), 19 September 1980, Rycroft 3365 (NBG). 3420

(Bredasdorp): De Hoop, (-AD), 14 September 1979, Burgers 2242

(NBG). 3421 (Riversdale): Reisiesbaan Siding, (-AB), 6 August 1983,

Bohnen 8231 (NBG). 3424 (Humansdorp): Kromme River, Company’s

Drift, (-BA), September 1922, Fourcade 2318 (BOL).

13. The name Senecio panduratus (Thunb.) Less, was

misapplied by De Candolle (1838) to plants that resem-

ble S. hastatus L. in general appearance but which are

distinct from it in their sparsely branched corymbs with

much larger capitula. Both S. hastatus and S. panduratus

sensu DC. have a short, ± horizontal rhizome, and radi-

cal leaves with long petioles and inciso-pinnatifid blades,

and the stems and leaves are thinly or densely pubescent

with a mix of short, glandular hairs and longer eglandular

hairs. True S. hastatus is characterized by few- to well-

branched corymbs of up to 20, relatively small, cylindri-

cal capitula, 9-12 x 5-7 mm, with 12-14 phyllaries. It

is widely distributed in moister situations from Ceres in

the southwestern Cape eastwards to Lesotho and the free

State (Hilliard 1977; Goldblatt & Manning 2000). The

taxon currently known under the name S. panduratus ,

in contrast, has sparsely branched corymbs of ( 1— )3— 1 0,

larger, campanulate capitula, 10-12 x 8-12 mm, with 20-

24 phyllaries. It has a more restricted distribution in the

interior southwestern Cape, in more arid environments.

The name S. hastatus is in fact a synonym of S. erosus

(see above) and the plant currently known under that

name thus requires a new name. Examination of the type

of S. robertiifolius DC. (1838), a poorly known taxon

based on a collection made in the Kamiesberg, confirms

that it precisely matches the plants currently identified as

S. hastatus and this name is therefore available for use.

Bothalia 40,1 (2010)

45

Senecio robertiifolius DC., Prodromus systema-

tis naturalis regni vegetabilis 6: 384 (1838). Type: South

Africa, [Northern Cape], Camisberg [Kamiesberg], Mod-

derfontein. [4 November 1830], Drege [5901] (G-DC,

holo.-microfiche!; P-Aluka image!, website accessed

25-11-09).

S. panduratus sensu DC., non (Thunb.) Less.

Additional specimens examined

NORTHERN CAPE. — 2917 (Springbok): Steinkopf, Besonder-

meid, (-BC), 27 September 1933, Herre s.n. (NBG). 3017 (Honde-

klipbaai): Kamieskroon, (-BB), 29 August 1937, Compton 6810

(NBG). 3018 (Kamiesberg): Welkom, near Garies, (-AB), 16 Octo-

ber 1954, Esterhuysen 23683 (BOL). 3119 (Calvinia): Niewoudtville

Reserve, (-AC), 7 September 1983, Perry & Snijman 2297 (NBG);

Oorlogskloof Nature Reserve, (-AC), 28 September 2000. Preto-

rius 538 (NBG); Zoetwater, west of Calvinia, (-BC), 24 September

1952, Maguire 1933 (NBG); Akkerendam Nature Reserve, (-BD),

6 August 1993, Jooste 1 (NBG); behind Hantam, Farm Saaihoek, (—

BD), 7 September 2002, Rosch 124 (NBG). 3220 (Sutherland): 11 km

east of Sutherland, (-BD), 22 September 1985, Moffett 3 772 (NBG);

Komsberg, Farm Klein Jakhals Valley, (-DA), 6 September 1986,

Cloete & Haselau 237 (NBG).

WESTERN CAPE. — 3219 (Wuppertal): Pakhuis, (-AA), 29

September 1940, Compton 9559 (NBG). 3318 (Cape Town): Rie-

beek-Kasteel, (-BD), 14 September 1941, Compton 11723 (NBG).

3319 (Worcester): Tweedside, (-AB), 25 September 1932, Compton

4011 (NBG); Whitehill, (-BA), 17 September 1945, Compton 17398

(NBG); Whitehill Ridge, (-BA), 20 September 1943, Compton 14900

(NBG); Bonteberg, (-BD), 20 September 1931, Compton 3780 (NBG);

Rabiesberg, (-DA). 26 September 1935, Compton 5835 (NBG). 3321

(Ladismith): west of Seweweekspoortberg, (-AD), 3 February 1992,

Oliver 9998 (NBG); Klein Swartberg, east of Kouveldberg, (-AD), 23

December 2006, Helme 4370 (NBG). 3322 (Oudtshoom): Kamanassie,

Mannetjiesberg, (-DB), 1 February 1941, Compton 10563 (NBG).

14. Senecio lobelioides DC. (1838) was based on

material collected by Drege on the Farm Silwerfontein,

midway between Springbok and Kamieskroon. It was

not seen by Harvey (1865), who merely repeated De

Candolle’s description. Examination of isotype material

at Paris confirms that it is readily recognized among the

other annual species by being completely glabrous, and

with characteristic leaves, the lower conspicuously petio-

late with ovate-reniform blades, and the upper sessile and

cordate-amplexicaul. Although becoming progressively

smaller towards the end of the branches, the upper leaves

retain their distinctive shape, giving the inflorescences a

rather leafy character. Another distinctive feature of the

species is the narrowly cylindrical capitula, which are

obscurely radiate with very short rays barely exceeding

the involucre. Drege’s material of .S’, lobelioides , how-

ever, is indistinguishable from collections at SAM that

have been identified as Senecio flavus (Decne.) Sch.Bip.

subsp. flavus by both Merxmiiller (1967) and later, in

1988, by Aaron Lister (now Department of Botany and

Plant Pathology. Oregon State University). Although we

have not been able to examine type material of S. flavus,

the protologue (Decaisne 1834), as well as the descrip-

tion and accompanying illustration in Boulos & Hind

(2002), give us no reason to doubt this opinion and we

accordingly treat S. lobelioides as a synonym of S. fla-

vus.

Senecio flavus (Decne.) Sch.Bip. in Webb. & Berth.,

Histoire naturelle des lies Canaries 2: 3 1 9, t. 1 07 ( 1 845).

Crassocephalum flavum Decne.: 265 (1834). Type: Ara-

bia [Egypt], Sinai, without date or collector (P). Illus.:

Boulos & Hind: 264 (2002).

S. lobelioides DC.: 382 (1838), syn. nov. Type: South Africa, [North-

ern Cape], Silverfountain [Silwerfontein], [31 August 1830], Drege

[2730] (G-DC, holo.-microfiche!; P-Aluka image!, website accessed

25-9-09).

Additional specimens examined

NAMIBIA. — 2426 (Maltahohe): Bullspoort, (-AB), July 1947,

R.G. Strey 2175 (NBG).

NORTHERN CAPE. — 3120 (Williston): Wolwe Rivier, (-CA), 26

July 1941, R.H. Compton 11144 (NBG).

15. The replacement name Senecio odontophyllus

C.Jeffrey (1992) that was published for the Indian spe-

cies until then known under the illegitimate later homo-

nym S’, linifolius (DC.) C.B. Clarke, was applied in error

by Goldblatt & Manning (2000) to the Eastern Cape spe-

cies that is correctly known as S. linifolius L.

Senecio linifolius L., Systema naturae 2: 1215

(1759). Senecio odontophyllus sensu Goldblatt & Man-

ning (2000) [non C.Jeffrey: 95 (1992) (= Senecio linifo-

lius (DC.) C.B. Clarke: 202 (1876)].

ACKNOWLEDGEMENTS

We thank Anne-Lise Fourie of the Mary Gunn Library,

SANBI for making the G-DC microfiche available to us

for study, Mary Stiffler of Missouri Botanical Garden

for assistance with older literature, and Dr Arne Ander-

berg and Dr Johannes Lundberg for arranging for a dig-

ital image of the S isotype of Doria gymnodiscus. Two

anonymous referees provided valuable insights and com-

ments. for which we are most grateful.

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Bothalia 40,1: 47-53 (2010)

New taxa of Babiana (Iridaceae: Crocoideae) from coastal Western

Cape, South Africa

P. GOLDBLATT* and J.C. MANNING**

Keywords: Babiana Ker Gawl., Iridaceae, southern Africa, systematics, winter rainfall region

ABSTRACT

Discovery of populations south of Elandsbaai of a small-flowered plant closely allied to Babiana ringens led to a critical

re-evaluation of this sunbird-pollinated Western Cape species. We conclude that these populations represent a new species,

B. avicularis, recognized by long, arching, subterete leaves, and flowers with the lower part of the perianth tube sigmoid and

± 4 mm long, a dorsal tepal 1 5-1 8 mm long, pale green lower tepals directed forward, and a style dividing below the bases

of the anthers. In addition, the southern coastal populations of B. ringens merit recognition as a separate subsp. australis,

recognized by the smaller flower, filaments not reaching the apex of the dorsal tepal and the style dividing at or below the

bases of the anthers. Field work along the Western Cape coast also resulted in the discovery of a new species, B. teretifolia,

allied to the distinctive B. brachystachys but differing from that species in the linear, spreading, twisted tepals, filaments 12

mm long, white anthers 5. 5-6.0 mm long, and the style dividing opposite the anther tips, with branches ± 5 mm long and

notched at the tips.

INTRODUCTION

The southern African and largely winter rainfall genus

Babiana Ker Gawl. was recently revised (Goldblatt &

Manning 2007a), at which time 88 species were recog-

nized. but novelties have continued to be added. Just a

year later, two new species were described (Goldblatt

et al. 2008), both found while the revision was in press.

In the spring of 2008, two more novelties came to our

attention. Neither species was to our knowledge repre-

sented in any herbarium but a careful search revealed

that one of them, the red-flowered B. avicularis, had

been collected twice before but had been identified as B.

ringens (L.) Ker Gawl., largely because of the red flower

and unusual sterile main stem. We found B. avicularis

in strandveld south of Elandsbaai but earlier records are

from the Olifants River Mountains north of Clanwilliam

and near Leipoldtville. Although resembling B. ringens

in its sterile main stem axis with flowers carried on

branches borne close to ground level, B. avicularis dif-

fers in the long, subterete, arching leaves and smaller

flowers with inter alia the lower part of the perianth tube

sigmoid and only ± 4 mm long, dorsal tepal 15-18 mm

long, and style dividing below the bases of the anthers.

The discovery of this species prompted us to review the

variation in B. ringens and closely allied B. hirsuta and

we conclude that the southern populations of the former

merit recognition as a separate subspecies that we name

subsp. australis. This taxon has smaller flowers than

subsp. ringens , with a dorsal tepal only 1 8-30 mm long

vs 25^45(-50) mm in subsp. ringens, filaments not

reaching the apex of the dorsal tepal and, like B. avicu-

laris. a style dividing below the bases of the anthers.

The second new species, Babiana teretifolia, from

sandveld on the farms Kommandokraal and Skilpadvlei,

* B.A. Krukoff Curator of African Botany, Missouri Botanical Garden,

RO. Box 299, St. Louis, Missouri 63166, USA. peter.goldblatt@mobot.

org.

** Compton Herbarium, South African National Biodiversity Institute,

Private Bag X7, 7735 Claremont, Cape Town, [email protected].

MS. received: 2009-04-21.

northwest of Vredendal in southern Namaqualand, is

allied to B. brachystachys (Baker) G.J. Lewis, a species

centred in coastal Namaqualand but extending south to

Lambert’s Bay in Western Cape, the latter a range exten-

sion for the species. Both B. brachystachys and B. tereti-

folia have ± terete leaves with 5 or 6 narrow longitudi-

nal grooves, the main and lateral spikes decumbent and

usually held at ground level, and flowers with an elon-

gate perianth tube exceeding 60 mm. B. teretifolia is dis-

tinctive in its linear, channelled, slightly twisted tepals

26-30 mm long, spreading at right angles to the tube,

with the dorsal tepal barely differing from the others in

size, shape and orientation. In addition, B. teretifolia has

filaments exserted ± 9.5 mm, white anthers 5. 5-6.0 mm

long, and longer style branches ± 5 mm long compared

with filaments exserted up to 3 mm, anthers 4-5 mm

long, usually mauve to violet, and style branches ± 2.5

mm long in B. brachystachys. With the two new species

described here, Babiana now comprises 92 species.

1. Babiana avicularis Goldblatt & J.C. Manning,

sp. nov.

Plantae ad 120 mm altae foliis exclusis, axe princi-

pal! sterili velutino, ramis ad basem productis, foliis 6-8

linearibus ad 300 x 2-3 mm arcuatis laevigatis, spicis

horizontalibus subsecundis usitate 8-12-floris, bracteis

minute scabridis viridibus apicibus brunneis mucronatis,

bractea externa 14-22 mm longa interiora fere ad basim

divisa, floribus zygomorphis rubro-carneis in fauce flavis

tepalis inferioribus pallide viridibus in medio atroviridi-

oribus, tubo perianthii compresso-infundibuliformi parte

inferiori cylindrico ± 4 mm longo, parte superiori 12-17

mm longa, tepalis inaequalibus dorsale 15-18 mm longo,

filamentis 25-32 mm longis, antheris 3. 5-5.0 mm longis

atroviolaceis ovario laevigato, stylo infra basim anther-

arum diviso, ramis 4-5 mm longis.

TYPE. — Western Cape, 3218 (Clanwilliam): coast

south of Elands Bay, ± 30 km north of Rocher Pan,

sandy flats, (-AD), 1 1 September 2008, Goldblatt &

Porter 13109 (NBG, holo.; K, MO, PRE, S, iso.).

48

Bothalia 40,1 (2010)

Plants up to 120 mm high, excluding leaves, with

main axis suberect and sterile, usually with two decum-

bent, fertile branches borne close to ground level,

minutely velvety on aerial axes. Leaves 6-8, linear, up to

300 x 2-3 mm, firm but arching toward ground, ± oval

in cross section and closely plicate, usually with 2 or 3

folds on each surface, glabrous. Fertile spike horizon-

tal, subsecund with flowers in two ranks, mostly 8-12-

flowered; bracts minutely scabrid, green with dry brown

apices, outer bracts 14-22 mm long, bearing a prominent

mucro, inner ± half as long as outer, divided almost to

base and diverging distally, joined by transparent tis-

sue at ovary level. Flowers zygomorphic, facing spike

apex, predominantly reddish pink, paler in throat, lower

part of tube yellow, lower five tepals with pale green

limbs sometimes fading to pink near tips, darker green

in midline, unscented; perianth tube compressed-funnel-

shaped, lower part ± 4 mm long, cylindric, sigmoid dis-

tally then abruptly expanded into flared upper part 12-17

mm long; tepals unequal, outer whorl prominently mucr-

onate, dorsal 15-18 x 2. 5-3. 5 mm, ascending, chan-

nelled below with margins curving inward and enclosing

filaments and style for most of their length, expanded

to 3 mm wide and recurved in upper 5 mm, lower three

tepals united with upper laterals for ± 4 mm in promi-

nent lip, free parts directed forward, lower and upper

laterals ± 8 x 1.5 mm, lower median 3-7 x 1-2 mm. Sta-

mens unilateral; filaments straight, suberect, 25-32 mm

long, enclosed below by dorsal tepal, exserted ± 16 mm

and reaching apex of dorsal tepal; anthers 3. 5-5.0 mm

long, purple. Ovary smooth; style dividing shortly below

base of anthers, style branches 4-5 mm long, recurved.

Capsules barrel-shaped, ±11 x 8.5 mm, showing outline

of seeds. Mature seeds not known. Flowering time : mid-

August to late September. Figure 1A-D.

Distribution and ecology : restricted to the West Coast

and near interior of Western Cape (Figure 2). Just four

populations of Babiana avicularis are known: two are

from sandy flats between Elands Bay and Rocher Pan, a

short distance inland from the densely vegetated coastal

dunes adjacent to the beach; a third is from the Olifants

River Valley north of Clanwilliam, also on sandy ground;

and the fourth, the earliest record, is from coastal fynbos

close to Leipoldtville. The four records fall within three

quarter-degree squares of latitude and longitude.

Diagnosis and relationships', flowers of Babiana avic-

ularis resemble those of the other two predominantly red-

flowered species of the genus, B. hirsuta (Lam.) Gold-

blatt & J.C. Manning (= B. thunbergii Ker Gawl.), and B.

ringens, except that they are almost half the size (Figure

1). Like B. ringens , the main axis is sterile and we infer

that, as in B. ringens , it serves as a perch for birds tak-

ing nectar from the flowers (Anderson et al. 2005) but

the flowers differ from that species in some important

respects apart from their smaller size (Table 1 ). Flowers

of B. ringens subsp. ringens have a perianth tube 28-40

mm long, a dorsal tepal 25^45(— 50) mm long, enlarged

lower lateral tepals ± 20 mm long, and the upper tepals

are recurved (Figure 1H). In addition, the filaments are

37-70 mm long, the anthers 5. 0-7. 5 mm long, and the

style divides opposite the upper half of the anthers or

shortly beyond their tips. Subsp. australis has slightly

smaller flowers with the dorsal tepal 18-30 mm long

and filaments 24-34 mm long. In contrast, flowers of B.

avicularis have a tube 16-21 mm long, a dorsal tepal

1 5-1 8 mm long, and all three lower tepals pale green and

directed forward. In both species, as well as in B. hir-

suta, the margins of the dorsal tepal curve together and

loosely enclose the filaments and style, and only the dis-

tal third or quarter is expanded and slightly recurved. The

filaments of B. avicularis are 25-32 mm long, the anthers

3. 5-5.0 mm long and the style divides below the bases

of the anthers. The leaves of B. ringens are lanceolate to

sublinear but firm to rigid in texture with almost pungent

tips, in contrast to the longer, linear, fairly soft-textured

leaves of B. avicularis that arch outward gracefully.

The flowers of Babiana avicularis resemble those of

B. hirsuta more closely than B. ringens in their propor-

tions but in B. hirsuta the upper and lower lateral tepals

are recurved distally and extended outward, and the low-

ermost tepal is substantially larger than the lower laterals

(Figure 1E-G), the reverse of the situation in B. avicularis.

The lower tepals of B. avicularis are pale green, sometimes

pink at the tips, lanceolate and directed forward and the

perianth tube has one feature not found in its two allies: it

is slightly constricted at the apex of the short, narrow cylin-

drical part and forms an S-shaped bend before expanding

into the flared upper part of the tube.

The floral bracts of Babiana avicularis are notable

in two respects: they are rust-tipped and prominently

TABLE 1. — Comparison of important taxonomic features of Babiana avicularis, B. ringens and B. hirsuta. Data were taken from new collections

and well-pressed herbarium specimens

Bothalia 40,1 (2010)

49

FIGURE 1. — A— D, Babiana avicularis, Goldblatt & Porter 13109 (NBG); E— G, B. hirsuta, Goldblatt & Porter 13110; H, B. ringens subsp. ringens,

found at Atlantis, Schnitzler & Manning 13 (NBG); I, J, Babiana ringens subsp. australis, De Waal s.n. (NBG). A, whole plant; B, half-

flower; C, outer (left) and inner (right) bracts; D, capsule; E, flower, side view; F, half-flower; G, outer (left) and inner (right) bracts; H,

flower; 1, flower; J, outer (left) and inner (right) bracts. Scale bar: 10 mm. Artist: John Manning.

mucronate, and the shorter inner bracts are divided upper third to half and do not notably diverge. The two

for about three-quarters of their length, with the two coastal populations of B. avicularis are sympatric and

halves strongly divergent, whereas in both B. hirsuta co-blooming with B. hirsuta but we found no sign of

and B. ringens , the inner bracts are divided in the hybrids at either site.

50

Bothalia 40,1 (2010)

FIGURE 2. — Known distribution of Babiana avicularis , A; B. brachys-

tachvs, ■ ; B ringens subsp. ringens, • (Graafwater populations

arrowed); B. ringens subsp. australis , O; and B. teretifolia, □.

We assume from the floral morphology that, like its

two relatives, Babiana avicularis is adapted for pollina-

tion by sunbirds. Flowers produce 3. 8-5.0 pi of nectar of

relatively low sugar concentration (24 % ±1.6% sucrose

equivalents; n = 5). Flowers of B. hirsuta produce up to

30 pi of nectar of ± 25 % sucrose equivalents and in B.

ringens up to 28 pi nectar has been recorded of ± 23 %

sucrose equivalents (Goldblatt et al. 1999; Goldblatt &

Manning 2007b). The nectar in B. avicularis is therefore

similar in sucrose concentration but far less in quantity

to that offered by B. hirsuta and B. ringens , a reflec-

tion of the smaller size of the flower and much shorter

perianth tube. If B. avicularis proves to be pollinated by

sunbirds, it will be among the smallest-flowered species

of the Iridaceae with this pollination system.

Additional specimens examined

WESTERN CAPE. — 3218 (Clanwilliam): 19 km north of Clanwil-

liam, sand dunes, (-BB), 23 August 1974, Nordenstam & Lundgren

1505 (MO, NBG, S); west coast south of Elands Bay, ± 30 km north of

Rocher Pan, sandy flats, (-AD), 24 September 2008, Goldblatt & Por-

ter 13161 (MO, NBG, PRE); 2.5 miles [3.5 km] SW of Leipoldtville,

coastal fynbos on white sand, ± 400 ft [640 m], (-AB), 23 August

1958, Acocks 19677 (K, PRE).

2. Babiana ringens subsp. australis Goldblatt &

J.C. Manning, subsp. nov.

Plantae ad 120 mm altae, spicis 2-4-floris; bractea

externa 18-30 mm longa, tubo perianthii 27-35 mm

longo, tepalis inaequalibus tepalo dorsali 18-30 mm

longo, filamentis 24-34 mm longis, antheris 4. 0-5. 5 mm

longis, stylo infra basim antherarum diviso, ramis 2-4

mm longis.

TYPE. — Western Cape, 3419 (Caledon): Bredasdorp,

Frikkiesbaai, (-CB), 24 August 1946, Compton 18185

(NBG).

Like Babiana ringens in general aspect, subsp. aus-

tralis has a sterile stem up to 120 mm long and spikes

with 2-4 smaller flowers. Flowers with tepals oriented as

in subsp. ringens ; perianth tube 27-35 mm long, slender

lower part 15-20 mm long and upper wider part 12-15

mm long; dorsal tepal 18-30 mm long, upper lateral and

lower median tepals narrowly lanceolate, 20-25 mm long,

attenuate and mucronate, lower lateral tepals clawed with

ovate blade, 22-25 x 7.5 mm. Stamens unilateral; fila-

ments 24-34 mm long, exserted 15-22 mm, not reaching

apex of dorsal tepal; anthers 4. 0-5. 5 mm long, reaching

or sometimes exceeding apex of dorsal tepal. Style divid-

ing at or 1-2 mm below base of anthers; style branches

2^4 mm long. Capsules and seeds unknown. Flowering

time : mid-July to late August. Figure 1 1, J.

Distribution and ecology. Babiana ringens subsp.

australis extends along the southern coast of Western

Cape, from Albertinia in the east to the Agulhas Penin-

sula, with outlying populations in the southern Cape

Peninsula south of Scarborough (Figure 2). Plants occur

in sandy ground in coastal fynbos and are often more

prominent after fire.

Diagnosis', subsp. australis stands out among the

numerous collections of Babiana ringens , a fairly com-

mon coastal and near-interior species that is especially

frequent in the western half of Western Cape, extending

from Albertinia to the Cape Peninsula and north to Lam-

bert’s Bay, in its generally smaller size, 2-4-flowered

spike and markedly smaller flowers, with tube 27-35

mm long, and smaller tepals, the dorsal 1 8-30 mm long

(Tables 1; 2). In comparison, subsp. ringens has spikes

of (3 — )6— 1 0 flowers, a perianth tube 28-40 mm long, and

a dorsal tepal 25^45(-50) mm long. In keeping with the

longer perianth tube, the filaments are 37-70 mm long,

and the anthers are 5. 0-7. 5 mm long vs filaments 24-34

mm long and anthers 4. 0-5. 5 mm long in subsp. austra-

lis. The style of subsp. australis divides at or below the

base of the anthers and the style branches are relatively

short, 2-4 mm, compared with a style usually dividing

beyond the anther tips (rarely opposite the middle of the

anthers) in subsp. ringens and the style branches are 4-5

mm long. Both subspecies occur on the Cape Peninsula,

with subsp. australis recorded from Scarborough in the

southwestern Peninsula and subsp. ringens north of Fish

Hoek; therefore, as far as we can determine, their ranges

abut but do not overlap.

Two northern populations of Babiana ringens from

the immediate vicinity of Graafwater ( Goldblatt 3628

MO; Schlechter 8518 MO) (Figure 2) have unusually

small flowers for the subspecies (Table 2), with a peri-

anth tube 31-40 mm long and a dorsal tepal ± 25 mm

long, but unlike subsp. australis they have anthers 5.0-

5.5 mm long, a style dividing opposite the middle to

upper third of the anthers, and style branches 4-5 mm

long. These populations occur close to others with larger

flowers, typical of subsp. ringens, and may represent a

local race or even introgressed hybrids with B. avicularis.

Additional specimens examined

WESTERN CAPE. — 3418 (Simonstown): Cape Peninsula, Schus-

terskraal, after lire, (-AB), 10 October 1945, Barker 3885 (NBG);

Scarborough, Red Hill trail, (-AB), 4 August 2009, De Waal s.n.

(NBG). 3421 (Riversdale): 2 km west of Dekriet, west of Albertinia,

(-AB), 26 July 1979, Bolmen 6101 (NBG); 10 km west of Albertinia,

(-AB), 22 September 1949, Sidey 1762 (MO); Stilbaai, Groot Jongens-

fontein, (-AD), 23 August 1978, Bolmen 3968 (NBG); Victoriasdale,

sandy slopes of Brandkop, 600 ft [190 m], (-AC), 6 September 1975,

Bothalia 40.1 (2010)

51

Oliver 5996 (NBG); Albertinia commonage, next to graveyard, sandy

hillside, (-BA), 20 August 1971, Thomas s.n. (NBG92938).

3. Babiana teretifolia Goldblatt & J.C. Manning,

sp. nov.

Plantae acaulescentes ad 300 mm altae foliis inclusis,

cormo 25-30 mm diam., foliis subteretibus ±2x1 .3-1.6

mm diam., anguste 3-sulcatis, spica decumbenti congesta

subsecunda spica principali ad 10-flora, bracteis viridi-

bus apicibus atrobrunneis acuto-attenuatis, 18-25 mm

longis, bractea interiora ad apicem 3^1 mm furcata, tubo

perianthii cylindrico elongato 65-70 mm longo pauciter

curvato, tepalis subaequalibus linearibus torsivis paten-

tibus tepalo dorsali ± 30 x 4 mm alliis ± 26 x 2.5 mm,

filamentis ± 12 mm longis ± 9.5 mm exsertis, antheris

5. 5-6.0 mm longis albis, stylo ad apices antherarum vel

ultra diviso, ramis ± 5 mm longis ad apicem bifurcatis.

TYPE. — Western Cape, 3118 (Vanrhynsdorp): south-

ern Namaqualand, Farm Kommandokraal, NW of Vre-

dendal. (-AC), 10 September 2008, Goldblatt & Porter

13101 (NBG, holo.; MO. iso.).

Plants acaulescent, up to 300 mm high including

leaves; corm deeply seated, up to 200 mm below ground,

globose, 25-30 mm diam., outer tunics dry and brown,

extending upward with dry remains of leaf sheaths in col-

lar around stem base; stem usually 1- or 2-branched at

ground level, glabrous or sparsely pubescent. Leaves ± 8,

± oval to round in section, ±2 x 1.3-1. 6 mm. narrowly

2-grooved on wider surfaces, 1 -grooved on narrow sur-

faces, margins sparsely hairy proximally. Spike decum-

bent, crowded, secund with flowers ± 1.5 mm apart in 2

ranks, main spike with up to 10 flowers, branches usually

with fewer flowers; bracts 1 8-25 mm long, acute-attenu-

ate, green below and rust-coloured at tips, inner ± as long

as outer, forked apically for 3^1 mm. Flowers zygomor-

phic, predominantly pink, beige toward bases of tepals;

lower 3 tepals with narrow diamond-shaped markings in

lower midline, red in throat; perianth tube cylindric, slen-

der, 65-70 mm long, hollow, slightly curved; tepals nar-

rowly linear and twisted, spreading with dorsal one held

slightly apart from upper lateral, dorsal ± 30 x 4 mm.

upper lateral and lower tepals ± 26 x 2.5 mm. tepals ±

uniformly pink outside, tube translucent, light purple dis-

tally, fading to beige proximally. Stamens unilateral; fila-

ments ± 12 mm long, inserted ± 2.5 mm below mouth of

tube, therefore exserted ± 9.5 mm; anthers 5. 5-6.0 mm

long, white, pollen white. Ovary smooth; style arching

over stamens, dividing at or shortly beyond anther tips.

branches ± 5 mm long, notched apically. Capsules and

seeds unknown. Flowering time : September. Figure 3.

Distribution', known from a single extended popula-

tion on the farms Kommandokraal and Skilpadvlei in

southern Namaqualand, in deep sand in coastal sandveld

(Figure 2).

Diagnosis and relationships'. Babiana teretifolia

superficially appears to be merely a variant of B. brachy-

stachys , a species centred in coastal Namaqualand and

extending south to Fambert’s Bay in Western Cape, in

its subterete leaves and crowded, decumbent spikes of

extremely long-tubed flowers (Fewis 1959). The flow-

ers themselves are. however, different in several respects,

especially in the remarkably narrow, spreading and

twisted tepals and the dorsal tepal hardly differentiated

from the remaining five. The tepals are linear, ± chan-

nelled, and twisted with the dorsal tepal ± 30 x 4 mm and

the remaining tepals ± 26 x 2.5 mm, whereas typical B.

brachystachvs has oblong-lanceolate, ± plane tepals, the

dorsal tepal 20-24 x 6-8 mm and the lower three tepals

15-18 x 4-6 mm (e.g. the type, Mader s.n. K; Goldblatt

& Manning 9997 K, MO. NBG). These differences are

combined with striking differences in the stamens: in B.

teretifolia the filaments are ± 12 mm long and exserted

± 9.5 mm from the tube and the anthers are 5. 5-6.0 mm

long and white, as is the pollen. Populations of B. brachv-

stachys have filaments 4-5 mm long, exserted 1-3 mm,

and anthers 4-5 mm long and are usually pale violet to

purple when fresh, as is the pollen (anthers in a popula-

tion north of Fambert’s Bay discovered in September

2009, Goldlbatt & Porter 13321 MO, NBG, have white

anthers and pollen). In addition, the style of B. teretifolia

divides opposite the anther tips so that the style branches,

± 5 mm long, arch above the anthers and can readily be

seen to be notched at the tips. In B. brachystachvs, the

style divides opposite the upper third of the anthers and

the style branches are only ± 2.5 mm long. Added to

the floral differences are the sparsely hairy leaf sheaths

and flowers spaced ± 1.5 mm apart, in contrast with the

woolly leaf sheaths and flowers set 3-6 mm apart in most

populations of B. brachystachvs. The extraordinarily

long, linear twisted tepals, compared to the more conven-

tional oblong-lanceolate tepals of B. brachystachvs , mark

B. teretifolia as a derived, geographic segregate of its

already highly specialized sister species.

Babiana teretifolia , like B. brachystachvs , has the

stereotypical adaptations for pollination by the long-

TABLE 2. — Comparison of important taxonomic features of population sets of Babiana ringens

Population sets: Graafwater vicinity. Central: central Cape Peninsula (Fish Hoek) to Lambert’s Bay; Worcester District, Clanwilliam to Botterkloof.

Southern: Cape Peninsula (Scarborough) to Albertinia (Figure 2).

52

Bothalia 40,1 (2010)

FIGURE 3 . — Babiana teretifoliu. Goldblatt & Porter 13101 (NBG). A, whole plant; B, flower, front view; C, half- flower; D, outer (left) and inner

(right) bracts. Scale bar: 10 mm. Artist: John Manning.

Bothalia 40,1 (2010)

53

proboscid fly, Moegistorhynchus longirostris (Manning

& Goldblatt 1997; Goldblatt & Manning 2000), viz.

beige-pink tepals marked with red towards the base, and

an elongate perianth tube. Other species in the guild of

species adapted for pollination by Moegistorhynchus

that co-occur with B. teretifolia, include Geissorhiza

exscapa (Thunb.) Ker Gawk, Pelargonium appendicu-

latum (L.f.) Willd., and (flowering slightly later in the

season) Lapeirousia anceps (L.f.) Ker Gawl. and L. fab-

ricii (D.Delaroche) Ker Gawl. We did record Moegis-

torhynchus longirostris visiting one member of the

guild, G. exscapa in September 2009, but saw no visi-

tors to B. teretifolia during three days in the field in the

same month in 2008. perhaps due either to cool, windy

weather that is less than ideal for fly activity.

REFERENCES

ANDERSON, B„ COLE, W.W. & BARRETT, S.C.H. 2005. Special-

ized bird perch aids cross-pollination. Nature 435: 41. 42.

GOLDBLATT, P„ BERNHARDT, P. & MANNING, J.C. 1999. Evidence

of bird pollination in the Iridaceae of southern Africa. Adansonia,

ser. 3,21:25-40.

GOLDBLATT, P & MANNING, J.C. 2000. The long-proboscid fly polli-

nation system in southern Africa. Annals of the Missouri Botanical

Garden 87: 146—170.

GOLDBLATT, P & MANNING, J.C. 2007a. A revision of the south-

ern African genus Babiana, Iridaceae: Crocoideae. Strelitzia 18.

South African National Biodiversity Institute, Pretoria and Mis-

souri Botanical Garden, Missouri.

GOLDBLATT, P. & MANNING, J.C. 2007b. Floral biology of Babiana

(Iridaceae: Crocoideae): adaptive floral radiation and pollina-

tion. Annals of the Missouri Botanical Garden 94: 709-733.

GOLDBLATT, P„ MANNING, J.C. & GEREAU, R.E. 2008. Two new

species of Babiana (Iridaceae: Crocoideae) from western South

Africa, new names for B. longiftora and B. thunbergii, and com-

ments on the original publication of the genus. Bothalia 38:

49-55.

LEWIS, G.J. 1959. The genus Babiana. Journal of South African Bot-

any. Suppl. 3.

MANNING, J.C. & GOLDBLATT, P. 1997. The Moegistorhynchus lon-

girostris (Diptera: Nemestrinidae) pollination guild: long-tubed

flowers and a specialized long-proboscid fly pollination system in

southern Africa. Plant Systematics and Evolution 206: 5 1-69.

Bothalia 40,1: 55-102 (2010)

Notes on African plants

VARIOUS AUTHORS

ARECACEAE

LIVISTONA CHINENSIS , A FIRST RECORD OF A NATURALIZED PALM IN SOUTH AFRICA

Seven genera of the Arecaceae occur in southern

Africa and southern tropical Africa, comprising 18

indigenous species (Table 1 ) (Glen 2000; Leistner 2005).

Of these, only six palm species are indigenous to South

Africa, probably due to prevailing climatic conditions

that are unfavourable for tropical taxa. and it is therefore

not surprising that no non-indigenous member of the

Arecaceae has previously been regarded as naturalized

or invasive in southern Africa (Henderson 2006).

The aim of this paper is to report on the first palm to

have become semi-naturalized in South Africa, namely

Livistona chinensis (Jacq.) R.Br. ex Mart. This palm

is also considered as naturalized in some parts of the

United States of America (Butts 1959; Oppenheimer

2003) and Mauritius (Maunder et al. 2002). The palm

was introduced to South Africa as an ornamental dur-

ing the early 1900s and has hitherto been widely planted

throughout the eastern coastal regions of South Africa

due to its availability, vigour and hardiness (Esterhuyse

et al. 2001 ).

Although one of the most commonly cultivated orna-

mental palms in the world (Dowe 2009), Livistona

chinensis is now ‘Vulnerable’ in its indigenous Japan,

Taiwan and China (Hainan Island), where it occurs

mostly in coastal forests on various soils, often in sand,

sometimes in dense monospecific colonies, or isolated

groups or individuals, up to 100 m in altitude. The

species was first described as Latania chinensis (Jacquin

1801), from plants cultivated in Mauritius and brought

to Schoenbrunn Gardens, Vienna in 1788. The species

name was taken from that used for the palm in Mauritius,

‘ Latanier de la Chine'. Bretschneider (1898) provided

some evidence to suggest that the naturalist and traveller

TABLE 1. — Palm genera and no. species per genus occurring naturally

in southern Africa and southern tropical Africa respectively and

in totality

Pierre Poivre was responsible for introducing the palm,

during the mid- 1700s, to Mauritius where it became

naturalized. The common name, Chinese fan palm,

came into use very early (Jacquin 1801) and was based

on the Latin name as well as subsequent knowledge of

its distribution in eastern Asia (Yoshida et al. 2000). In

South Africa it is also known as the Chinese fountain

palm. An extensive search in southern African herbaria

for Livistona chinensis revealed only two voucher

specimens, both of garden origin, which were made by

H.F. Glen on 23 March 1991 at Vaalwater, Limpopo, and

by J.C. Scheepers in April 1992 in Pretoria, Gauteng.

Arecaceae (alt. Palmae), subfamily Coryphoideae,

tribe Trachycarpeae, subtribe Livistoninae (Dransfield et

al. 2008).

Livistona chinensis (Jacq.) R.Br. ex Mart., Historia

Naturalis Palmarum 3,7: 240 (1838). Latania chinensis

Jacq.: t. 11, fig. 1 (1801). Saribus chinensis (Jacq.)

Blume: 49 (1838). Livistona sinensis Griff: 131 (1845),

ortho, var. Type: without locality and collector [lecto.,

illustration in Jacquin: tab. 11, fig. 1 (1801), designated

by Moore (1979)].

Latania borbonica auct. non Lam.: 427 (1792).

Livistona mauritiana Wall, ex Mart.: 240 (1838), nomen.

Chamaerops biroo Siebold: 11 (1 830). Type: Japan, Siebold s.n. [L,

lecto.!, designated by Dowe (2009)].

Livistona olivaeformis (Hassk.) Mart.: 319 (1850); Miquel: 59

(1855); Miquel: 13 (1868). Saribus olivaeformis Hassk.: 176 (1842).

Latania olivaeformis (Hassk.) Devansaye: 34 (1875), nom. illeg. Type:

Indonesia, cultivation, Bogor Botanic Gardens, ‘nel viale presso la

chieta’. May 1878, Beccari s.n. [FI, lecto.!, sheets 1131, 1131 -B &

1 131 -C, designated by Dowe (2009)].

Livistona subglobosa (Hassk.) Mart.: 319 (1850); Miquel: 59

(1855); Miquel: 13 (1868); Nakai: 224 (1935). Saribus subglobosiis

Hassk.: 177 (1842); Hassk.: 65 (1844). Livistona chinensis var.

subglobosa (Hassk.) Becc.: 16 (1921). Type: Indonesia, cultivation,

Bogor Botanic Gardens, May 1878, Beccari s.n. [FI, lecto.!, sheets

11333 & 11333-B, designated by Dowe (2009)].

Livistona japonica Nakai ex Masam. (as Livistonia japonica ): 50

(1929); Masamune & Suzuki: 73 (1933). Type: not designated.

Diagnostic characters', monoecious hermaphroditic

tree, single trunk up to 15 m tall, enveloped by dry leaf

sheaths, dense crown of numerous divided, palmate and

pendulous pale green leaves, up to 1.8 m long, promi-

nent undivided central area and numerous deeply bifid

segments, tips pendulous. Petioles armed with stout

prickles. Flowers sessile, pale yellow, carried in clus-

ters of up to seven, borne in 5-7 inflorescences up to 1

m or more long, of several branches along a single main

rachis, each 2 or 3 times divided into rachillae; bracts

56

Bothalia 40,1 (2010)

FIGURE 1 . — Livislona chinensis. A, habit; B, trunk showing dry leaf sheaths and prickles on petioles; C, base of stem; D, leaf. Photographs: H.

de Wet.

brown tomentose. Fruit ellipsoid to subglobose or pyri-

form, 15-26 x 9-18 mm; bright green to bluish green,

china-blue-grey with age, in dense clusters. Flowering

time'. December to February. Figure I .

Distribution and habitat. Livistona chinensis has

become semi-naturalized in swamp forest in the sub-

tropical coastal region of KwaZulu-Natal. Current,

confirmed populations include three localities in the

Empangeni-Richards Bay area, with sight records for

the Durban area (Figure 2). In Zululand it occurs in for-

est dominated by the trees Bridelia micrantha , Phoenix

reclinata, Syzygium cordatum and Voacanga thouarsii ,

and an understorey dominated by the ferns Microsorum

scolopendria and Nephrolepis biserrata.

Studies by Siebert (2009) could not confirm any natu-

ral distribution vectors, but found that the masses of seed

produced by these palms end up in garden dumping sites

during autumn. Naturalized populations are only associ-

ated with garden refuse in swamp forest patches, as the

prevailing microclimate here, which is predominantly

warm and moist, is probably most suitable for germina-

tion. Siebert (2009) has reported that in these favourable

habitats, single individuals reach maturity over time, but

in turn these successful individuals produce masses of

Bothalia 40,1 (2010)

57

FIGURE 2. — Naturalized distribution of Livistona chinensis in South

Africa. Voucher records. •; sight record. ▲.

seed that lead to mass seedling establishment through

gravitational dispersal. Although this only leads to local-

ized, controllable invasions, it should serve as a warning

sign, and calls for an urgent action to assess the invasive

status and potential of cultivated palms in South Africa.

Voucher specimens

KWAZULU-NATAL. — 2831 (Nkandla): KwaDlangezwa, swamp

forest along Unthlatuze River, (-DD), Siebert 2456 (NH, PRE, ZULU);

Ngwelezana, swamp forest along Umhlatuze River, (-DD), Siebert

3188 (ZULU). 2832 (Mtubatuba): Richards Bay, swamp forest along

Thulazihleka Pan, (-CC), Siebert 3499 (ZULU).

LIMPOPO. — 2427 (Vaalwater): cultivated. Waterberg, (-BB), Glen

2628 ( PRE).

GAUTENG. — 2528 (Pretoria): garden, Menlo Park, (-DB),

Scheepers s.n. (PRE).

ACKNOWLEDGEMENTS

Ms Lesley Henderson, Agricultural Research Coun-

cil, commented on an earlier version of the manuscript.

Mr Barry Long, Durban Botanic Gardens, confirmed

the identity of the palm. Dr Hugh Glen, South African

National Biodiversity Institute (SANBI), confirmed the

occurrence of naturalized individuals of the palm in Dur-

ban. Ms Hester Steyn, National Herbarium. SANBI, is

thanked for the production of the distribution map.

REFERENCES

BECCARI, O. 1921. Recensione delle Palme del vecchio mondo

appartenenti alia tribu delle Corypheae, con descrizione della

specie e varieta nuove che vi appartengono. Webbia 5: 5-70.

BLUME. C.L. 1838. Saribus. Rumphia 2: 48-51.

BRETSCHNEIDER. E. 1898. History of European botanical discover-

ies in China. Press of the Imperial Russian Academy of Sciences,

St. Petersburg.

BUTTS, E.H. 1959. Livistona chinensis naturalized in Florida. Princi-

pes 3: 133.

DEVANSAYE, A. DE LA. 1875. Palmiers. Les Coryphinees. Revue

Horticole 47: 31-35.

DOWE, J.L. 2009. A taxonomic account of Livistona R.Br. (Arecaceae).

Gardens Bulletin Singapore 60: 185—344.

DRANSFIELD, J„ UHL, N.W., ASMUSSEN, C.B., BAKER. W.J.,

HARLEY, M.M. & LEWIS, C.E. 2008. Genera Palmarum:

evolution and classification of the palms, edn 2. Royal Botanic

Gardens, Kew.

ESTERHUYSE, N„ VON BREITENBACH, J. & SOHNGE, H. 2001 .

Remarkable trees of South Africa. Briza Publications, Pretoria.

GLEN, H.F. 2000. Arecaceae (Palmae). In O.A. Leistner, Seed plants

of southern Africa: families and genera. Strelitzia 10: 580-582.

National Botanical Institute, Pretoria.

GLEN, H.F. 2003. Arecaceae. In G. Germishuizen & N.L. Meyer, Plants

of southern Africa: an annotated checklist. Strelitzia 14: 973.

National Botanical Institute, Pretoria.

GRIFFITH. W. 1845. The palms of British East India. Calcutta Journal

of Natural History 5:31 1-355.

HASSKARL, C. 1 842. Plantarum rariorum. Tijdschrift voor Natuurlijke

Geschiedenis en Physiologie 9: 176.

HASSKARL, C. 1 844. Catalogus plantarum in Horto botanico Bogo-

riensi cultarum alter. Typis Officinae Publicae, Bataviae.

HENDERSON, L. 2006. Comparisons of invasive plants in southern

Africa originating from southern temperate, northern temperate

and tropical regions. Bothalia 36: 201-222.

JACQUIN, N.J. VON. 1801. Latania chinensis. Fragmenta botanica

16.

LAMARCK, J.B.A.P.M. 1792. Encyclopedic Methodique, Botanique

3: 427.

LEISTNER, O.A. 2005. Seed plants of southern tropical Africa: families

and genera. Southern African Botanical Diversity Network

Report No. 26: 382, 383. SABONET, Pretoria.

MARTIUS, C.F.P. VON. 1 838. Livistona. Historia naturalis palmarum,

edn 1,3,7:238-242. Leipzig.

MARTIUS, C.F.P. VON. 1 850. Livistona. Historia naturalis palmarum,

edn 2, 3,9: 239-242, 319. Leipzig.

MASAMUNE, G. 1929. A preliminary report on the vegetation of the

island ofYakushima. Kagosima.

MASAMUNE, G. & SUZUKI, S. 1933. A list of plants collected in

Kizan. Annual Report of the Taihoku Botanic Garden 1933 3:

49-75.

MAUNDER. M„ PAGE, W„ MAUREMOOTOO, J.. PAYENDEE, R..

MUNGROO, Y„ MALJKOVIC, A., VERICEL. C. & LYTE. B.

2002. The decline and conservation management of the threat-

ened endemic palms of the Mascarene Islands. Oryx 36: 56-65.

MIQUEL, F.A.W. 1855. Licuala-Livistona. Flora van Nederlandh

Indie 3: 51-60.

MIQUEL, F.A.W. 1868. Palmis archipelagi indici observationes novae.

C.G. v.d. Post, Amsterdam.

MOORE, H.E. Jr. 1979. Order Arecales. Fam. 39. Arecaceae. Flora

Vitiensis Nova 1: 392M38.

NAKAI, T. 1935. Species generis Livistonae in Imperio Japonico sponte

nascentes. Journal of Japanese Botany 1 1 : 217-224.

OPPENHEIMER, H.L. 2003. New plant records from Maui and Hawaii

Counties. Bishop Museum Occasional Papers 73: 3-30.

SIEBERT, S.J. 2009. Livistona chinensis, a semi-naturalised palm of

swamp forest in subtropical South Africa. Palms 53: 193-196.

SIEBOLD. P.F. VON. 1830. Palmae. Synopsis plantarum oeconomica-

rum universi regne Japonici. Ter Lands Drukkerij, Batavia.

YOSHIDA, N„ NOBE, R„ OGAWA, K. & MUROOKA, Y. 2000. Ori-

gin of Livistona chinensis var. subglobosa (Arecaceae) on the

‘Islet of the Gods’ : Aoshima, Japan. American Journal of Botany

87: 1066, 1067.

S.J. SIEBERT*, A.M. ZOBOLO** and J.L. DOWE+

* Author for correspondence: A.P. Goossens Herbarium, School of En-

vironmental Sciences and Development, North-West University, Private

Bag X6001, 2520 Potchefstroom, South Africa.

** Department of Botany, University of Zululand, Private Bag X1001,

3886 KwaDlangezwa, South Africa.

t Australian Centre for Tropical Freshwater Research, James Cook

University, Townsville, Qld 4811, Australia.

MS. received: 2009-04-14.

58

Bothalia 40,1 (2010)

ASPHODELACEAE

OCCURRENCE OF HAWORTHIA BOLUSII VAR. BLA CKBEARDIANA IN THE FREE STATE, SOUTH AFRICA

The genus Haworthia Duval, which consists of ±

85 species and 200 taxa, is endemic to southern Africa

(Bayer et al. 1999). It occurs predominantly in the West-

ern and Eastern Cape Provinces, with outliers extending

into Namibia and Mozambique (see Smith & Van Wyk

1991 for a distribution map of the genus).

To date, the climatically severe grasslands of the

Free State have been known to harbour only a single

species of Haworthia , viz. H. venosa (Lam.) Haw.

subsp. tessellata (Haw.) M.B. Bayer (Scott 1985: xxvii;

Smith et al. 2000). However, the occurrence of a sec-

ond species of Haworthia , H. bolnsii Baker var. black-

beardiana (V.Poelln.) M.B. Bayer, has been suggested

for the Free State, but confirmation of this has thus far

remained elusive (Bayer 1999: 45). In a comprehensive

interpretation of the H. cooperi Baker and H. bolusii

var. blackbeardiana complexes, Bayer (2002) concen-

trated his discussion on their Eastern Cape distribu-

tion ranges. Haworthia bolusii var. blackbeardiana is

a miniature (the size of a golf ball), rosulate plant with

flatfish, incurved leaves that are adorned with hair-like

marginal spines.

Haworthia bolusii var. blackbeardiana was recently

found in the Rouxville District, Free State at the follow-

ing locality:

FREE STATE. — 3027 (Zastron): Brandwacht 190, Rouxville, veg-

etative plant, 1 640 m, S 30° 30' 11.3" E 27° 03’ 26.0", (-AC), 23-05-

2008, PC. Zietsman 4410 (NMB).

This locality falls within the vegetation unit Besem-

karee Koppies Shrubland (Gh 4) of the Grassland Biome

(Mucina & Rutherford 2006).

The plants were found on the western slope of a koppie

(hill). Most of them grow tucked away in cracks between

the dolorite rocks and are well protected from direct sun-

light during the early morning and late afternoon. They

are fully exposed for most of the day (Figure 3).

REFERENCES

BAYER, M.B. 1999. Haworthia revisited. A revision of the genus.

Umdaus Press, Hatfield.

BAYER, M.B. 2002. Haworthia update. Essays on Haworthia, vol. 1.

Umdaus Press, Hatfield.

BAYER, M.B., EGGLI, U., VAN JAARSVELD, E„ SMITH, G.F. &

SUPTHUT, D.J. 1999. From Adrian Haworth to Haworthia.

Haworthia and related South African succulents. Haworthia

Society, St Michaels on Wyre.

MUCINA, L. & RUTHERFORD. M.C. (eds). 2006. The vegetation of

South Africa, Lesotho and Swaziland. Strelitzia 19. South Afri-

can National Biodiversity Institute, Pretoria.

SCOTT, C.L. 1985. The genus Haworthia (Liliaceae). A taxonomic revi-

sion. Aloe Books, Johannesburg.

SMITH, G.F. & VAN WYK, B-E. 1991. Generic relationships in the

Alooideae (Asphodelaceae). Taxon 40: 557-581.

SMITH, G.F., ZIETSMAN, P.C., STEYN, E.M.A. & BREUER, 1.

2000. The distribution of Haworthia venosa subsp. tessellata in

southern Africa. Haworthiad 14,2: 40 — 43.

P.C. ZIETSMAN* and G.F. SMITH**

* National Museum, P.O. Box 266, 9300 Bloemfontein, South Africa

/ Centre for Environmental Management, University of the Free State,

P.O. Box 339, 9300 Bloemfontein, South Africa. [email protected].

** Biosystematics and Biodiversity Collections, South African

National Biodiversity Institute, Private Bag XI 01, 0001 Pretoria, South

Africa / Acocks Chair, H.G.W..I. Schweickerdt Herbarium, Depart-

ment of Botany, University of Pretoria, 0002 Pretoria, South Africa.

[email protected].

MS. received: 2009-03-25.

FIGURE 3. — Haworthia bolusii Baker

var. blackbeardiana in its natu-

ral habitat near Rouxville in the

Free State. Photograph: P.C.

Zietsman.

Bothalia 40,1 (2010)

59

ASPHODELACEAE

INCLUSION OF THE GENUS JODRELLIA IN BULBINE (ASPHODELOIDEAE)

The genus Jodrellia Baijnath was described to accom-

modate three white-flowered species from tropical Africa,

two of which were formerly included in Bulbine Wolf (Baij-

nath 1978). These species were separated from Bulbine

based on a combination of characters, namely radiate roots,

inflorescences shorter than the leaves, a w hite perianth with

three- to five-nerved outer tepals, and obscure stigmatic

papillae (Baijnath 1978). These characters, with the excep-

tion of the three- to five-nerved outer tepals, are also found

in species of Bulbine and other genera of the Asphode-

laceae, thus reducing their significance as generic charac-

ters. Radiate roots, which are situated on a small vertical

rhizome in Jodrellia (Baijnath 1978), occur sporadically in

other genera of the Asphodelaceae, notably Trachyandra

Kunth, and are not unique to Jodrellia. The underground

parts of Bulbine species are variable but normally either

tuberous or a horizontal rhizome. White flowers, as are

found in Jodrellia, occur in forms of Bulbine frutescens

(L.) Willd. (which some authors regard as a separate spe-

cies, i.e. B. triebneri Dinter; Van Jaarsveld 2001; Klopper

et al. 2008; Klopper & Smith 2009), while some forms of

B. abyssinica A. Rich, have inflorescences that are shorter

than the leaves. The outer tepals in Bulbine species are nar-

rower than the inner ones, whereas the reverse is found in

Jodrellia. The outer tepals are wider than the inner ones

and accessory venation has developed, possibly as a result

of the wider lamina. The obscure stigmatic papillae of

Jodrellia , although shorter and fewer, are similar to those

of Bulbine. Both genera share the hairy filaments, which is

the most striking feature of the genus Bulbine and is unique

to these two genera within the Asphodelaceae.

Molecular sequence data from chloroplast (rbcL, niatK.

and ndhF) and nuclear (ITS) markers confirm the close

relationship between the two genera, indicating that Jodrel-

lia is embedded within Bulbine with strong support in both

the analysis of chloroplast and nuclear regions (Devey et

al. 2006). The genus Bulbine is thus paraphyletic without

the inclusion of Jodrellia, and Devey et al. (2006) have no

hesitation in recommending the transfer of Jodrellia to Bul-

bine. This is necessary in order to preserve the monophyly

of the genus. Species of Jodrellia should thus be viewed

as representing a small, specialized lineage within Bulbine

rather than as a genus distinct from it. This interpretation is

supported by the weak morphological differences between

the two, essentially the development of accessory veins in

the outer tepals in Jodrellia. The necessary nomenclatural

changes are presented below.

Bulbine Wolf, Genera plantarum: 84 (1776). Type

species: B. frutescens (L.) Willd. Jodrellia Baijnath: 574

(1978), syn. nov. Type species: Jodrellia macrocarpa

Baijnath.

Bulbine fistulosa Chiov. in Annali di Botanica

(Roma) 9: 143 (1911). Jodrellia fistulosa (Chiov.) Baijnath:

576 ( 1978). Type: Ethiopia, Chiovenda 557 (FI, holo.).

Bulbine macrocarpa (Baijnath) Boatwr. & J.C. Man-

ning, comb. nov. Jodrellia macrocarpa Baijnath in Kew

Bulletin 32: 574 (1978). Type: Kenya, Gillett 13141 (K,

holo.).

Bulbine migiurtina Chiov. in Lavori eseguiti

presso il Reole Istituto Botanico di Catania 1: 5 (1928).

Jodrellia migiurtina (Chiov.) Baijnath: 577 (1978). Type:

Somaliland, Puccioni & Stefanini 777 [860] (FI, holo.).

ACKNOWLEDGEMENTS

This work formed part of a Post-Doctoral Fellowship

of the first author at the Compton Herbarium and the

University of Cape Town. The Parker family from the

Elandsberg Nature Reserve is kindly thanked for gener-

ously providing funding for the Fellowship.

REFERENCES

BAITNATH, H. 1978. Jodrellia , a new genus of Liliaceae from tropical

Africa. Kew Bulletin 32: 571-578.

CHIOVENDA, E. 1911. Plantae novae vel minus notae e regionae

Aethiopica. Annali di Botanica (Rome) 9: 143.

CHIOVENDA, E. 1928. Plantae novae vel minus notae ex Aethiopia.

Lavori eseguiti presso il Reole Istituto Botanico di Catania 1 : 5.

DEVEY, D.S., LEITCH, I., RUDALL, P.J., P1RES, J.C.. P1LLON, Y. &

CHASE, M. 2006. Systematics of Xanthorrhoeaceae sensu lato,

with emphasis on Bulbine. Aliso 22: 345-351.

KLOPPER, R.R., KLOPPER, A.W.. BAIJNATH, H. & SMITH, G.F.

2008. Bulbine triebneri , an earlier name for Bulbine alba , as well

as additional and new localities in Eastern and Northern Cape,

South Africa. Bothalia 38: 67-69.

KLOPPER, R.R. & SMITH, G.F. 2009. Formalizing the synonymy of

Bulbine triebneri. Bothalia 39: 100, 101.

VAN JAARSVELD, E. 2001. South African succulent plants: two new

species and two new combinations. Haseltonia 8: 37-41 .

WOLF, N.M.VON. 1776. Genera plantarum vocubulis chracteristicis

definita : 84. Danzig.

J.S. BOATWRIGHT** and J.C. MANNING* *

* Compton Herbarium, South African National Biodiversity Institute,

Private Bag X7, 7735 Claremont, Cape Town. E-mail: s.boatwright@

sanbi.org. za

* Department of Botany, University of Cape Town, Private Bag, 7700

Rondebosch, Cape Town.

MS. received: 2009-03-09.

IRIDACEAE

REAPPRAISAL OF IXIA MACULATA WITH /. CALENDULACEA SP. NOV., AND AN EARLIER NAME FOR I. LUTEA

A serendipitous discovery of two varieties of Ixia

maculata L., I. maculata var. fusco-citrina (Desf. ex

DC.) G.J. Lewis and I. maculata var. intermedia G.J.

Lewis, growing and flowering in close proximity in

September 2007, led us to review the taxonomy of the

species, which was treated by Lewis ( 1962) and De Vos

(1999) as comprising three varieties. Ixia maculata is a

member of section Ixia of the genus, which is endemic to

60

Bothalia 40,1 (2010)

the winter rainfall zone of southern Africa and comprises

over 70 species at the latest count (Goldblatt & Manning

2008a, b). Section Ixia includes 19 species with brightly

coloured flowers with a narrow, subfiliform perianth

tube, non-decurrent filaments inserted at or close to the

top of the tube, and conduplicate style branches (Lewis

1962). Most species of the section have flowers with

contrasting dark central marks, now believed to repre-

sent beetle marks, and indications are that most species

are pollinated by hopbine beetles (Scarabaeidae: Hop-

liini) (Goldblatt et al. 2000).

Ixia maculata sensu G.J. Lewis stands out in sect. Ixia

in its bright orange or yellow perianth with a dark central

eye, dry and crinkly floral bracts that are usually partly or

entirely brown, basally to completely united filaments, and

the style dividing opposite or above the base of the anthers.

These features are shared in the section only with I. curia

Ker Gawk, which differs from I. maculata in its strongly

dark-cuspidate bracts and its conns. The corms of /. macu-

lata were described by Lewis (1962) as typically having

submembranous or rarely fibrous tunics and producing sto-

lons from the base, whereas corms of I. curta have persist-

ent, fibrous, netted tunics, and cormlets, when present, are

borne at the base of the corm, which is the more common

condition in Ixia. Only I. dubia Vent., I. lutea Eckl. and

I. polvstacliya L. of sect. Ixia also have yellow or orange

flowers, but none have conns producing stolons, and all

three have free filaments and a style usually dividing below

the level of the anthers, except in I. polystachva var. lutea,

which may prove to be a separate species. Ixia lutea and I.

polystachva more often have white or pink flowers and /.

dubia, which is the taxon most often confused with I. macu-

lata, has pale, translucent floral bracts, sometimes suffused

with pink. All these species share a range in Western Cape,

and were once common from Cape Town northward to the

Olifants River.

Lewis (1962) and De Vos (1999) distinguished three

taxa within Ixia maculata primarily using features of the

floral bracts, degree of filament union, and perianth tube

length but a degree of overlap in these characters led to

segregation of the known populations as varieties, rather

than as distinct species. Our observations show that the

nature of the stolons, corm tunics, colour of the filaments,

and the shape and markings of the tepals are equally, if not

more important in distinguishing populations (Table 2).

Some of these features are correlated with ecological dif-

ferences and have convinced us that the present taxonomy

does not adequately represent the biology of the I. macu-

lata complex. We present a revised taxonomy in which

we recognize var. intermedia as a separate species, I. cal-

endulacea. Whereas vars. maculata and fusco-citrina have

yellow filaments, and tepals with a concave base to which

the dark brown or black pigmentation is restricted. I. cal-

endulacea has dark brown filaments (drying violet), and

the tepals are plane, thus lacking a concave base, and the

brown to dull red centre is edged with a halo of translucent

red. In addition, populations of I. calendulacea grow in

deep, well-drained sandy soils, whereas typical I. maculata

grows in sandy or granitic habitats that are always water-

logged in the growing season.

The differences between Ixia maculata var. maculata

and var. fusco-citrina remain more or less as described

by Lewis (1962) but the two taxa are insufficiently

resolved. We note the potential taxonomic significance

of the short stolons bearing up to three small cormlets

that are produced in var. maculata and the long stolons

of var. fusco-citrina which bear a single cormlet but

additional field work is needed to assess the status of

these two taxa.

Ixia calendulacea Goldblatt & J.C. Manning, sp.

nov.

/. maculata var. intermedia G.J. Lewis in Journal of South Afri-

can Botany 27: 142 (1962). Type: Western Cape, 3218 (Clanwilliam):

Redelinghuys, 29 September 1943, IV. F. Barker 2591 (NBG, holo. ! ).

Plantae 200-500 mm altae usitate eranrosae, corrno

depresso-globoso 12-16 mm diam. tunicis fibrosis sobolis

horizontalibus ± 1.5 mm diam., ad 120 mm longis praedito,

foliis usitate 4-6 raro 3, usitate (8-) 12-20 mm latis lanceo-

latis saepe supra torsivis, spica flexuosa usitate 5-8-flora,

bracteis siccis albo-transparentibus infra brunneis supra,

bractea externa ± 8 mm longa ± truncata vel biloba cuspi

brevi centrali, interna ad apicenr furcata in dua cuspibus

attenuata 2 mm longa, floribus vadosuis cupulatis calen-

dulaceis centra brunneo vel rubro ± 15 mm diam., usitate

nrargini rubro-translucenti, tubo perianthii 5— 8(— 1 0) mm

longo cylindrico, tepalis subequalibus ascendentibus pau-

citer imbricatis ( 1 8— )22— 25 * 12-14 mm, extemis ± 2 mm

TABLE 2. — Characteristics of Ixia calendulacea compared with those of I. maculata var. maculata and var. fusco-citrina. Because there is doubt

about the status of cultivated plants referred in herbaria to var. fusco-citrina, we have not included measurements from these specimens.

Observations for taxonomically important features are taken only from well-preserved specimens, bearing in mind that floral features may

shrink up to 20 % of original size, depending on the care with which specimens are prepared. We did not use Lewis’s (1962) or De Vos’s

(1999) measures for any taxa because we apply some names in different ways

Bothalia 40,1 (2010)

61

latior quam intemis, filamentis ± 4 mm longis connatis in

pars dimidio vel omnino raro liberis atrobrunneis, antheris

9-1 1 mm longis, stylo inter basem et inferiori tertio anther-

arum diviso, ramis 3^1 mm longis.

TYPE. — Western Cape, 3318 (Cape Town): gran-

ite-topped hill east of Langebaan, in deep sand, (-AA),

19 September 2008, Goldblatt & Porter 13152 (NBG,

holo., K. MO, PRE, iso.).

Plants 200-500 mm high; conn depressed-globose,

12-16 mm diam., outer tunics fibrous with fibres mostly

oriented vertically, sometimes accumulating, producing

long horizontal stolons up to 120 mm long, ± 1.5 mm

diam.. each bearing a terminal cormlet 8-9 mm diam.;

stem usually unbranched, robust plants often with a

node in upper third bearing a short branch or dry, attenu-

ate scale, sheathing only at base and often curved back

against stem. Leaves (3)4-6, lowermost largest, decreas-

ing in size above, uppermost ± entirely sheathing, 2/3 to

3/4 as long as stem, lanceolate, lowermost mostly (8 — ) 12—

20 mm wide, often twisted in one or two rotations. Spike

flexuose, mostly 5-8-flowered; bracts dry, white-trans-

lucent below, brown in upper half or third, outer bract ±

truncate or bilobed with a short central tooth, (6— )8— 1 2

mm long, becoming ± lacerate, inner ± as long as outer,

forked in upper 2 mm into attenuate cusps. Flowers shal-

lowly cupped when fully open, orange with brown or dull

red central marking, 15-20 mm diam.. usually with trans-

lucent reddish halo; perianth tube cylindric, 5 — 8(— 1 0) mm

long, ± 2 mm diam. in lower part, expanded in upper 1

mm; tepals subequal, ascending and overlapping slightly

when fully open, ( 1 8— )22— 30 x 11-15 mm, outer ± 2 mm

wider than inner. Stamens with filaments 3-4 mm long,

united in lower half or entirely (rarely ± free), dark red-

brown (often drying violet but ultimately losing colour);

anthers ascending, 9-11 mm long, yellow. Style dividing

between base and lower third of anthers, branches (3— )4

mm long. Capsules and seeds unknown. Figure 4A-F.

Distribution and biology: largely a coastal species, Ixia

calendulacea has a scattered distribution along the West-

ern Cape Coast and near interior, from the Cape Peninsula

and Saldanha Bay northwards to the Natnaqualand coastal

plain, as far north as the Groen River (Figure 5). Plants

occur in sandy habitats, usually deep sands but also in areas

of limestone substrate. Plants are most often associated

with strandveld and sandveld vegetation but inland popu-

lations in the Olifants River Valley and the hills west of

Piketberg, occur on stony sandstone slopes and flats in dry

fynbos. When treated by Lewis (1962) and De Vos (1999)

as 7. maculata var. intermedia , its range was known from

Saldanha to the Olifants River Valley. Later collections,

notably Reid 1291 from Namaqualand. near Kotzesrus, and

from the Western Cape west coast northwest of Vredendal,

Goldblatt & Manning 12876 (not in bloom but with the

characteristic long stolons and relatively broad leaves of 7.

calendulacea ), document the presence of the species north

of the Olifants River. A very recent collection from a rem-

nant patch of Cape Flats Sand Fynbos in the southeastern

suburbs of Cape Town, Dorse sub Manning 3231, is the

first record of the species from the Cape Peninsula. There

is no indication that this population, which has flowers with

a large, translucent red centre and dark filaments connate to

the tip, is not native to the Peninsula (Dorse pers. comm.).

The population, which occurs in a military area inacces-

sible to the public, is heavily infested with alien acacias,

and flowered after a fire, six months earlier, burned off the

canopy and woody undergrowth.

Observation of flowering plants from the type local-

ity near Langebaan ( Goldblatt & Porter 13152) confirm

that Ixia calendulacea is visited by small brown hopbine

beetles. These insects become covered in pollen as they

crawl across the open perianth and readily transfer pol-

len as they fly from one open flower to another. This is

consistent with what is known about pollination in sev-

eral other species of section Ixia (Goldblatt et al. 2000).

Diagnosis and relationships : Ixia calendulacea is recog-

nized by the relatively large, bright orange flower with a

brown to dull red central eye with a translucent red halo,

and brown filaments that are partly to entirely united, rarely

± free (Figure 4A, C, D). Plants have fairly large conns,

12-16 mm diam., with fibrous tunics and bearing long, flat-

tened stolons up to 120 mm long (Figure 4B). They typi-

cally have four to six basal leaves with lanceolate blades

twisted in the upper half. As in I. maculata , the bracts

are dry. finn and crinkled and brown at least in the upper

part and the style divides above the base of the anthers.

When compared directly with living I. maculata, the dif-

ferences between the two come into shaip focus (Figure 4;

Table 2). Ixia maculata has a particularly crowded spike,

and the flower, viewed from the side, shows that the tepals

form a shallow cup in the proximal third (the brown part

of the tepals), whereas the distal, orange or yellow por-

tions spread horizontally (Figure 41, J). The tepals are ±

narrowly oblong-ovate with nearly parallel sides, ± 22 x 9

mm, do not overlap, and are tapered below so that the cup

is narrowly windowed. In the southern form of the species

the windows together with orange margins of the cupped

portion of the tepals provides the star-like pattern in the

brown cup. In typical I. maculata (as defined by Lewis

1962) the stolons are short and somewhat twisted and bear

more than one (up to three) small cormlets (Figure 4H) and

these contrast markedly with the long, flattened stolons of

7. calendulacea (Figure 4B). Plants corresponding to var.

fusco-citrina have similar long, flattened stolons and this

feature may prove to be an important distinction between

that and var. maculata. Unfortunately, too few collections

have conns well enough preserved to allow examination of

the stolons; indeed many collections lack conns altogether.

As we noted above, 7. calendulacea and 7. maculata some-

times grow in close proximity but in different habitats, the

latter always in seasonally wet sites. Their ranges, however,

overlap very little (Figure 5), 7. maculata being restricted

to the southwestern Cape between Paarl and the western

end of the Piketberg, whereas 7. calendulacea extends from

Langebaan and Porterville to central Namaqualand. An

annotation on a collection from near Leipoldtville (De Wet

00903 ) indicates a diploid chromosome number of 2 n = 42,

which suggests the species may be polyploid (Goldblatt &

Manning in prep.). Basic chromosome number in Ixia is x

= 10 (Goldblatt & Takei 1997). The count is unpublished.

A collection from near Porterville, Goldblatt 2745, is

somewhat unusual in having narrow leaves (resembling

those of Ixia maculata) but the flowers have united fila-

ments and pale bracts, 9-1 0 mm long, with only the tips

turning brown, thus corresponding with 7. calendulacea.

We regard this collection as best referred to the latter and

it constitutes the southernmost record of the species.

62

Bothalia 40,1 (2010)

FIGURE 4. Ixia calendulacea: A, flowering stem; B, corm; C, two flowers; D, half-flower; E, outer (left) and inner (right) bracts; F, details of

stamens and style. Ixia maculata: G, flowering stem; H, corm; I, flower; J, half-flower; K, outer (left) and inner (right) bracts; L, detail of

stamens and style. Scale bar: A-E, G- K, 10 mm; F, L, 2 mm. Artist: John Manning.

Bothalia 40,1 (2010)

63

FIGURE 5. — Known distribution of Ixia calendulacea, O; and I. macu-

lata including var .fusco-citrina, •.

Key distinctions between Ixia calendulacea and /.

maculata

la Tepals forming concave cup in central dark zone and spread-

ing horizontally in distal part; tepals mostly 15-25 x 8-12

mm, not or hardly overlapping one another toward base;

central eye 6-12 mm diam., dark brown, often with star-

shaped mark within; filaments yellow; stolons either long,

up to 100 mm long, bearing a single terminal cormlet or

short, up to 20 mm long, usually more than one cormlet,

each up to 3 mm diam I. maculata

lb Tepals when fully open forming shallow cup and curved

uniformly from base to apex; tepals mostly 16-28 x 9-14

mm, the inner obviously overlapping the outer in lower

half; central eye of perianth ± 1 5 mm diam., glossy brown

with pale reddish halo or dull, translucent red; filaments

dark red-brown (violet when dry); stolons extending hori-

zontally for up to 120 mm and bearing single terminal

cormlet 7-9 mm diam I. calendulacea

Representative specimens

NORTHERN CAPE. — 3017 (Hondeklipbaai): Farm Hardekoppie

NW of Kotzesrus, (-DC), 29 September 1987, Reid 1291 (PRE).

WESTERN CAPE. — 31 18 (Vanrhynsdorp): Farm Graafwater, W of

Koekenaap, (-AC), August 2007 (sterile), Goldblatt & Manning 12876

(NBG); Nardouw, (-DC/DD), 22 September 1937, Barker 272 (NBG).

3217 (Vredenburg): Witteklip rocks, Vredenburg, (-DD), 18 Septem-

ber 1980, Goldblatt 5845 (MO). 3218 (Clanwilliam): Lambert’s Bay

road at Leipoldtviile turnoff, (-BA), 8 October 1965, De Wet 00903

(PRE); Farm Sandberg, sandy slope burned two years ago, (-BC),

27 September 1995, Goldblatt & Manning 10324 (MO, NBG); Farm

Nooitgedacht, N of Vredenburg, (-CC), 9 October 1985, De Vos 2624

(PRE); Piketberg, Farm Weltevrede, sandy slope, (-DA), 19 September

2007, Goldblatt & Manning 13011 (MO); sandveld between Porterville

and Piekeniers Kloof Pass, (-DD), 24 September 1974, Goldblatt 2745

(MO, NBG. PRE); 'Posberg Reserve’ (ex Darling Flower Show), (?-

AA), 19 September 1986, be Vos 2665 (NBG). 3418 (Simonstown):

Cape Town, Youngsfield Military Base, (-BA), 13 October 2009,

Dorse sub Manning 3231 (NBG).

AN EARLIER NAME FOR IXIA LUTE A

While examining the protologues of synonyms of Ixia

maculata listed by Lewis (1962) in order to determine

if any might apply to I. calendulacea, we found that /.

abbreviata Houtt. (1780) (Figure 6) was misplaced in

the synonomy of this species. The type, an illustration

(Figure 7), shows an Ixia- like plant with large flowers in

which the style divides below the level of the anthers.

The description, remarkably detailed for the time,

emphasizes the short style, noting that this feature dis-

tinguishes the plant from most other species [of Ixia-like

plants]. Houttuyn (1780) described the flowers as aris-

ing from transparent sheaths (i.e. bracts), sulphur yellow

with a bluish central eye, anthers longer than the fila-

ments and with a perianth tube almost half an inch long.

(gj) Ixia die getroste Bloemen heeft , met zecr xxkvl *

korte Stylen. '

De kortKeid van den Styl onderfcheidt deeze^fLAM

,t__xxxviir.

van Fig. 3.,

(34) Ixia Umbella bifidt Raccraofl. Bujrm. Prodr, 2.

(35) Ixia FloiibiisRaccmofia, Piftillo brcyisfim 0, HOUTT.

C 5

(I. Peel. XII* Stub,

( 4 » D R I E at A N N I G JJ LeLIE-

V, van de meefte anderen. Haar Bladen zyn LI.

Avdeel. niaal.LaDcetvormig , bekleedende om laag de

Hoof'd- Stengel, die een Voet lang is , rond en dun,

stub, op ’1 end een Trosjc hebfiende vaq zes Zwa«

velgeele Bloemen , taamelyk grooc. Zy komen

vqort uit Vliezige doojfc^ypspde tweekleppige

Scbeedjes , wier korefte of kleinfle Lip twee

t lange pun(en heeft. Dunne Steeltjes van een

half Duimi langtc , draagen deeze Bloemen

die byna een Duim lang zyn , beftaande uic

zes ovaalaqhuge , fyn geaderde Blaadjes. De

Meelknopjes , taamelyk dllc , langer dan de

Draadjes , JLiniaal of oyeral evep breed , ko»

men uic den Stoel der Bloem , die wac blaauw

achtig is , voort ; naby den oirfprong der drio

Stempels ; i welke Jangwerpig , 'dik en omge-

kromd , naauwlyks eenigen Styl hebben, ztt-

tende op den bodem van de Bloem* Ik hebzo

00k die blaauw zyn van KleUr.

xxxvi. (3 6) I>(?a met getroste Bloemen , die Klokvor*

■ mik zyn, enGrasaclnige Bladen.

Klokbioc- ,

Da figifur der Bloemen , \veik,a naav die der

xxxviii.' Klokjes gelyken , onderfcheidt deeze zo zeev

*• van’de voorgaande niet, als deiangte van den

Styl, waar van de lange dunn'e Stempels boven

de Meclknopjes zig yerhefFei). p.e piaejen zyn

- • Gras.

( 3« ) Ifia IlQtibus Ra,ccmofis , Carapaijifpunib^s , Fol* '

Gtjmjricia. Hoyrr,

FIGURE 6. — Protologue of Ixia abbreviata Houtt.: 41 , 42 ( 1 780). Translated

by J.R Roux: (35) Ixia with flowers in bunches and with short styles /

The short style separates this [species] from most others. The leaves

are linear-lanceolate and fonn a sheath around the stem, which is

one foot long, round and slender, and bears at the tip a single group

of six, reasonably large, sulphur-yellow flowers. They [the flowers]

arise from membranous, transparent, two-valved sheaths [bracts], of

which the lip of the shortest or smallest has two long points. Slender

stalks [tubes] half an inch long bear the flowers, which are nearly

one inch long and consist of six elliptic, finely veined tepals. The

relatively thick, linear or parallel-sided anthers are longer than the

stamens and originate from the base of the flower [mouth of the

tube], which is bluish, and near the origin of the three stigmas [style

branches], which are elongated, thick and recurved. The blue style is

almost absent and seated at the base of the flower.

64

Bothalia 40,1 (2010)

PliAAT LXXVIII.

There are few yellow-flowered species of Ixia section

Ixia and the one that matches the description best is the

plant currently called /. lutea. Ixia polvstachya L. some-

times has pale yellow flowers, but typically has anthers

shorter than, to as long as the filaments (rarely longer).

Yellow-flowered I. curta and I. maculata have a very

dark central eye, a style dividing above the anther base

and brown rather than translucent floral bracts. Lastly, I.

chibia Vent, has a small central dark eye and deep yellow

to orange flowers usually red on the outside, and does

not accord with Houttuyn’s plant.

Ixia abbreviata Houtt ., Natuurlijke historie 12:

41, t. 78 f. 3 ( 1780). Type: South Africa, without precise

locality or collector, illustration in Houttuyn (1780).

I. lutea Eckl.: 24 (1827), syn. nov. Type: South Africa, without pre-

cise locality, cultivated in Cape Town, Ecklon s.n. (S, holo.).

We provisionally follow De Vos (1999) in recognizing

a second variety of the species, providing the new combi-

nation for what she called var. ovata (Andrews) B.Nord.

Ixia abbreviata var. ovata (Andrews) Goldblatt

& J.C. Manning, comb. nov. Ixia capitata var. ovata

Andrews, The botanist’s repository 1: t. 23 (1798). I.

lutea var. ovata (Andrews) B.Nord.: 284 (1972). Type:

South Africa, without precise locality or collector, illus-

tration in Andrews: t. 23 ( 1 798).

ACKNOWLEDGEMENTS

We thank the curators of the following herbaria,

BOL, K, MO, NBG, PRE, and SAM (acronyms follow-

ing Holmgren et al. 1990) for permitting access to their

collections or for the loan of material for extended study.

Field work was supported by grants 7799-05 and 8248-

07 from the National Geographic Society (United States).

Collecting permits were provided by the Nature Conser-

vation authorities of Western Cape. We thank Lendon

Porter for his assistance and companionship in the field.

FIGURE 7. — Original illustration of

Ixia abbreviata. Plate 78, fig.

3 (Houttuyn 1780).

Koos Roux for his translation of Houttuyn’s text, and

Michelle Smith for assistance in preparing the images.

REFERENCES

ANDREWS, H. 1798. Ixia capitata var. ovata. The botanists 's reposi-

tory 1: t. 23.

DE VOS, M.P. 1999. Ixia. In M.P. de Vos & P. Goldblatt, Iridaceae;

Ixioideae: Ixieae (first part); Ixiinae, Tritoniinae. Flora of south-

ern Africa 7, part 2, fascicle 1 : 3-87. National Botanical Insti-

tute, Pretoria.

ECKLON, C.F. 1827. Topographisches Verzeiclmiss der Pflanzen-

sammlung von C.F. Ecklon. Reiseverein, Esslingen.

GOLDBLATT, P„ BERNHARDT, P. & MANNING, J.C. 2000. Adap-

tive radiation of pollination mechanisms in Ixia (Iridaceae: Cro-

coideae). Annals of the Missouri Botanical Garden 87: 564—577.

GOLDBLATT, P. & MANNING, J.C. 2008a. Systematics of the south-

ern African Ixia subgenus Morphixia (Iridaceae). 1. The I.

rapunculoides complex. Bothalia 37: 1-23.

GOLDBLATT, P. & MANNING, J.C. 2008b. Systematics of the south-

ern African genus Ixia (Iridaceae). 2. The filiform-leaved I. cap-

illaris group. Bothalia 38: 115-124.

GOLDBLATT, P. & MANNING, J.C. in prep. Systematics of the south-

ern African genus Ixia (Iridaceae). 3. Sections Hvalis and Mor-

phixia. Bothalia.

GOLDBLATT, P. & TAKEI, M. 1997. Chromosome cytology of Iridace-

ae, base numbers, patterns of variation and modes of karyotype

change. Annals of the Missouri Botanical Garden 84: 285-304.

HOLMGREN, P.K., HOLMGREN, N.H. & BARNETT, L.C. 1990.

Index Herbariorum, part. 1 : the herbaria of the World. New

York Botanical Garden, New York.

HOUTTUYN, M. 1780. Natuurlijke historie 12. Amsterdam.

LEWIS, G.J. 1962. South African Iridaceae. The genus Ixia. Journal of

South African Botany 27: 45-195.

NORDENSTAM, B. 1972. Types of Ecklon ’s 'Topographisches

Verzeiclmiss’ in the Swedish Museum of Natural History in

Stockholm. Journal of South African Botany 38: 277-298.

P. GOLDBLATT* and J.C. MANNING**

* B.A. Krukoff Curator of African Botany, Missouri Botanical Garden,

P. O. Box 299, St. Louis, Missouri 63166, USA.

** Compton Herbarium, South African National Biodiversity Institute,

Private Bag XI, 7735 Claremont, Cape Town.

MS. received 2009-06-29.

Bothalia 40,1 (2010)

65

LAMIACEAE

REDISCOVERY IN SOUTH AFRICA OF THE NEGLECTED AFRICAN VEGETABLE PLECTRANTHUS ESCULENTUS

Plectranthus esculentus N.E.Br. was rediscovered in

habitat during 2005, in rocky grassland on a hillside at

Inanda. inland of Durban (Figure 8). Herbarium records

reveal that this species has not been encountered in the

wild by botanists in southern Africa for over 30 years,

with several records reflecting collections from cultiva-

tion sites made predominantly during the early decades

of the 20th century (e.g. Van Warmelo TRV3617 PRE;

Gerstner 5436 PRE). Subsequent popularity of this

crop in South Africa has evidently waned considerably,

although limited use in Mpumalanga is reported to per-

sist (Allemann 2002).

The rediscovery in South Africa of this taxon has

significant implications for the strengthening of efforts

to reintroduce, for household food security, a neglected

African vegetable which is well adapted to areas of low

agricultural potential. Only one other South African

genotype (from Limpopo Province) is presently known,

and is represented in the holdings of the Agricultural

Research Council (ARC) at the Roodeplaat Vegetable

and Ornamental Plant Institute (J. Allemann pers. comm.

FIGURE 8. — Plectranthus esculentus in habitat, Inanda, KwaZulu-

Natal. Photograph: N. Crouch.

2007). The current find at Inanda is of particular impor-

tance as the formation of tubers has been observed at

latitude > 29.5°S (Crouch 1237 NH) (Figure 9), some-

what beyond the range (15 °N-28 °S) determined for this

species as a crop (Allemann & Hammes 2006). As such,

this collection may represent a photoperiodic ecotype

of agronomic consequence. Plectranthus esculentus is

characterized by finger-like edible tubers (Figure 10) and

bright yellow flowers (Pooley 1998) presented in short

pseudoracemes during spring, usually after the leaves

have been shed (Codd 1985). This geophyte produces

several lax stems which trail amongst grasses and root

at the nodes, thereafter seasonally producing stem tubers

(Allemann et al. 2003). Plants at the Inanda site were

found to not regenerate well from aerial parts, a charac-

teristic earlier documented by Burkill (1995). This fea-

ture is shared with tuberous forms of P. hadiensis (For-

ssk.) Schweinf. ex Spreng. var. hadiensis which occur

in grassland, the stems of which do not strike as well as

those of genus members found in more mesic habitats.

Success with striking of cuttings may relate to the timing

of tuber initiation, which appears to retard aerial growth

(J. Allemann pers. comm. 2008).

The vegetation in which plants may be encoun-

tered at Inanda is referred to as KwaZulu-Natal Sand-

stone Sourveld (SVs 5) by Rutherford et al. (2006)

who describe it as ‘short, species-rich grassland with

scattered low shrubs and geoxylic suffrutices’. The

underlying geology is Ordovician Natal Group sand-

stones. This vegetation type is considered Endangered,

with only 0.2 % statutorily conserved and some 68 %

already transformed (Rutherford et al. 2006). The habi-

tat of Plectranthus esculentus here comprises shallow

soil amongst rocks, on the edge of, and above steep

cliffs and escarpment edges at an altitude of ± 700 m.

Small aggregations of fewer than ten individuals occur

at scattered points on dry, northerly aspects over a dis-

tance of ± 500 m. The dominant grass amongst which

this Plectranthus species grows is Aristida junciformis

subsp. junciformis, which although a typical element

in KwaZulu-Natal Sandstone Sourveld, also prolifer-

ates in response to overgrazing and overburning. Other

associates include Aca/ypha glandulifolia, Gymnospo-

ria woodii, Pentanisia prunelloides, Phymaspermum

pinnatifidum , Plectranthus hadiensis var. hadiensis , Tet-

raselago natalensis and Thunbergia atriplicifolia. The

site is neither suitable for, nor gives indication of prior

cultivation by earlier inhabitants; accordingly the plant

appears here to be native rather than naturalized. Simi-

larly, Angolan subpopulations have been observed ‘in

[a] perfectly wild state’ (Good & Taylor 1931) and in

Zimbabwe within Julbernardia and Brachvstegia wood-

land (Wild et al. 1972).

Elsewhere in South Africa, particularly in the vicin-

ity of Nelspruit and Barberton, annotated herbarium

specimen labels (e.g. Lavranos 4681 PRE; Repton 647

PRE; Thorncroft 353 NH) indicate its natural occur-

rence. Wood ( 1896) noted that the species ‘[ Plectranthus

esculentus , or umbondwe \ is cultivated [around Durban]

66

Bothalia 40,1 (2010)

FIGURE 9, — Reported geographical distribution of Plectranthus escu-

lentus in FSA region based on specimens at BOL, NBG, NH,

NU, PRE and SAM, ■; extant subpopulation at Inanda, A.

by the natives, who use the tuber as a vegetable. I have

not seen it in a wild state, but a closely allied species (P.

floribundus, N.E.B.) is occasionally met with’. Gerst-

ner (1938) similarly claimed that amongst the Zulus, P.

esculenlus is not found wild but that it had been planted

‘since ancient times’. Forester Tustin of Ngome Forest

in KwaZulu-Natal, in correspondence with his superiors

during 1923 noted that ‘I really do not know if the plant

is indigenous to this part, or if it was previously brought

here. It is chiefly found growing in old lands’ (Tustin s.n.

PRE39880). According to oral Zulu tradition, P. esculen-

tus (as umhlaza ) and Colocasia esculenta (F.) Schott (as

amadumbi ) were brought south of the Umfolozi River

by a chief called Fanga. He entered what was later to

become Zululand from the direction of Swaziland, some

ten generations before Tshaka (Webb & Wright 2001).

Accordingly, P. esculenlus may have been introduced to

the region in the mid- to late 16th century.

Based on his field observations. Wood (1896) evi-

dently considered Plectranthus esculentus distinct from

P. floribundus N.E.Br., a taxon described from an Inanda

collection of his. Accordingly, it is likely that the Zulus

at Inanda were at that time cultivating at least one mor-

phologically distinct landrace of P esculenlus. Fox &

Norwood Young (1982) recorded that cultivation of dif-

ferent varieties was once commonplace in the Msinga

District on the middle Thugela. Such local diversity may

further be inferred from the variety of isiZulu names

for this species — no fewer than sixteen are documented

(Wood 1896; Bryant 1908; Gerstner 1938; Fox & Nor-

wood Young 1982; Allemann 2002). After 15 years

of ethnobotanical experience, the first author is yet to

encounter this edible lamiate in cultivation; as elsewhere

(Burkill 1995), this starch-rich and otherwise nutritious

crop (Allemann & Hammes 2003) has been displaced

by less labour-intensive and sometimes higher-yielding

introductions. These include Ipomoea batatas (F.) Earn,

(sweet potato) and maize ( Zea mays F.) from the New

World, and the Old World Colocasia esculenta (taro,

idumbe) from Asia. By the late 1 9th and early 20th cen-

turies, these crop species were well established amongst

the Zulus (Wood 1896; Bryant 1908). During the last

century, cultivation of the New World starch crop Sola-

num tuberosum F. (potato) has further marginalized

Plectranthus esculentus. As P. esculentus has been co-

dispersed synanthropically, its natural distribution is

imprecisely known, although this may at one time have

extended from Senegal in Equatorial Africa broadly

southwards to coastal KwaZulu-Natal (Codd 1975;

Burkill 1995). The original site of domestication and dis-

persal is uncertain, with various authors proposing West

(Purseglove 1976), Central (Porteres 1962), South-cen-

tral or East (Greenway 1944) Africa. It has reasonably

been surmised that domestication occurred independ-

ently in different regions across its wide range (Shaw

1976). Whereas several centres of cultivation are known

from various Central African countries, e.g. Nyanga ter-

races in eastern Zimbabwe (Sutton 1984), some user

groups eat only wild-sourced material, and then just as

a supplement or famine food (Burkill 1995). This may

reflect social stigmas which have led to preferences for

exotic crops (Kyesmu 1994).

Plectranthus esculentus and P. floribundus were

described synchronously by Brown (1894) who distin-

guished them on account of the latter species bearing

taller, more erect stems, and closely sessile leaves with

FIGURE 10. — Stem tuber cluster of a Plectranthus esculentus plant

sourced from Inanda, KwaZulu-Natal. Photograph: N. Crouch.

Bothalia 40,1 (2010)

67

broader, rounded bases, more prominent reticulation and

a rougher surface. Good & Taylor (1931) subsequently

placed P. floribundus in synonymy under Coleus escu-

lentus (N.E.Br.) G.Tayl., so allowing for a circumscrip-

tion that accommodates the wide diversity of cultivars

known, as well as natural variation across its range. Sub-

sequent workers on African Lamiaceae have accepted

this broader species concept (Codd 1975, 1985; Van

Jaarsveld 2006).

Rutherford et al. (2006) observed that most of the

remaining areas of KwaZulu-Natal Sandstone Sourveld

are subjected to grazing pressures and fire frequencies

that are not conducive to the recruitment of seedlings.

This is evident at the Inanda site where the leafless shoots

of plants have, for three consecutive years, been burned

off by intentional fires set during the winter months. This

has resulted in non-flowering and a lack of seed set. The

grassland in which Plectranthus esculentus occurs, still

retains a fair diversity of forbs and geophytes, particu-

larly in the rockiest parts. However, without respite from

these impacts and encroaching urban sprawl, the trend

over time will be towards increased degradation and loss

of species diversity. P. esculentus is a rare species within

its habitat, and is therefore likely to become even more so

in future. In view of the above, further collections from

the last-known South African locality of P. esculentus

should be genebanked as a matter of urgency, if residual

germplasm diversity is to be conserved.

Specimen examined

KWAZULU-NATAL. — 2930 (Pietermaritzburg): Inanda. in grass-

land along rocky ridge, 715 m, S 29° 36' 19.18"; E 30° 49' 34.01",

(-DB), 08^04-2009, Crouch 1237 (NH).

ACKNOWLEDGEMENTS

Dr J. Allemann of the Department of Soil, Crop and

Climate Sciences at the University of the Free State is

thanked for helpful discussions; Mr R. Edwards for

growing the original gathering made by D. Styles; Ms

H. Steyn and Ms E. Fouche for preparing the map; the

Curators of BOL, NBG. NH, NU, PRE and SAM for use

of their specimens; the staff of the Mary Gunn Library

and Dr E. Retief of PRE for facilitating access to litera-

ture, and the Data Section of the National Herbarium for

providing PRECIS data.

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tuberization in the Livingstone potato ( Plectranthus esculentus

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of Plectranthus esculentus N.E.Br. (Lamiaceae) are stem tubers.

Field Crops Research 83: 35-39.

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KYESMU, P.M. 1994. Plectranthus esculentus N.E.Br. A minor tuber

crop in dire need of rescue from extinction. Lamiales Newsletter

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N.R. CROUCH*t and D.G.A. STYLES**

* Ethnobotany Unit, South African National Biodiversity Institute, P.O.

Box 52099, Berea Road 4007, Durban.

+ School of Chemistry, University of KwaZulu-Natal, 4041 Durban.

** P.O. Box 50030, Musgrave 4062, Durban.

MS. received: 2009-06-18.

68

Bothalia 40,1 (2010)

PTERIDOPHYTA

NEW DISTRIBUTION RECORDS AND NOTEWORTHY COLLECTIONS OF PTERIDOPHYTES IN KWAZULU-NATAL

Rediscovery > o/Didymoglossum erosum

During a field trip to Ngoye Forest in Zululand in

early April 2009, a colony of Didymoglossum erosum

(Willd.) Beentje (syn. Trichomanes erosum Willd.) was

encountered on a dry vertical rock face in deep shade.

This small, easily overlooked, filmy fern (Figure 11 ) had

not been gathered in the FSA region since John Med-

ley Wood (1907, 1908) initially collected material from

Ngoye in 1 887.

Didymoglossum erosum (Flymenophyllaceae) is eas-

ily distinguished from most other filmy ferns by the

simple fronds. The only other species in southern Africa

with simple fronds is D. reptans (Sw.) C.Presl (syn. Tri-

chomanes reptans Sw.) from which it differs by being

variably pinnatifid and having sori that do not protrude

from the lamina but are rather winged by it. Furthermore,

they lack marginal hairs, and possess a false marginal

vein (Burrows 1990). The most recent literature considers

D. erosum as a single taxon (Beentje 2008; Roux 2009),

whereas historically, distinction has been made between

Trichomanes erosum var. erosum and T. erosum var. aer-

ugineum (Bosch) C.Chr. ex Bonap. (Schelpe 1970; Bur-

rows 1990; Roux 2001). This was based on morphologi-

cal differences, reinforced by habitat preferences (Schelpe

1970; Burrows 1990). Didymoglossum erosum is a rare

fern that occurs from the eastern highlands of Zimba-

bwe through eastern and central Africa (Figure 12) and

into western Africa to Sierra Feone and Guinea, as well

as Madagascar and the small islands of Annobon, Bioko,

Pemba, Principe, Reunion and Sao Tome (Burrows 1990;

Roux 2009). In South Africa this fern is only known from

the Ngoye Forest Reserve (specimens earlier ascribed

to Trichomanes erosum var. aerugineum , Schelpe &

Anthony 1986), where it has eluded re-collection, despite

ongoing botanical survey work (Huntley 1965).

In recent years, links in the pteridophyte flora of Ngoye

and the rainforests of eastern Zimbabwe have also been

highlighted: Christella buchananii (Schelpe) J.P.Roux and

Christella hispidula (Decne.) Holttum have recently been

collected from Ngoye (Burrows & Burrows 2001), both

of which are also known from the forests of southeastern

Zimbabwe, as is Asplenium blastophorum Hieron. (Bur-

rows 1990).

Specimens examined

Didymoglossum erosum

KWAZULU-NATAL. — 2831 (Nkandla): Ngoye Forest Reserve, ±

800 m northwest of Zululand Birding Route prefabricated accommo-

dation, in valley below, colony growing in deep shade on dry verti-

cal rock face in view of a perennial stream 1 5 m distant. Growing in

close proximity to, but separate from colonies of both Didymoglossum

replans and Crepidomanes melanotrichum , 280 m, (-DC), 0 1 -04-2009,

N. Crouch 1231 (NH, PRE); Ngoye Forest, (-DC), 23-03-1907, J.

Medley Wood 11947 (BOL; NBG; NH; PRE); Ngoye, (-DC), 26-03-

1907, J. Medley Wood 11948 (K).

New distribution record for Amauropelta oppositiformis

Amauropelta oppositiformis (C.Chr.) Holttum [The-

lypteridaceae, syn. Thelypteris oppositiformis (C.Chr.)

Ching] is a rare fern found along fast-flowing mountain

streams in high altitude grassland in full sun or light

shade of evergreen forest margins. It has tufted fronds

that are narrowly elliptic in outline and 2-pinnatifid,

with the basal pinnae gradually decreasing in size. The

pinnae are opposite and the pinnae lobes acute, with

the basal pair of veins not meeting below the sinus. It

is easily confused with A. bergiana (Schltdl.) Holttum

[syn. T. bergiana (Schltdl.) Ching], but is distinguished

from this species by the presence of small golden to

red glands and the lack of hooked hairs on the lower

surface of the frond (Burrows 1990). Within southern

FIGURE 11.— Detached fronds of

Didymoglossum erosum , Ngo-

ye Forest Reserve. Photograph:

N. Crouch.

Bothalia 40,1 (2010)

69

FIGURE 12. — Distribution of Didymoglossum erosum, adapted from

Burrows (1990), •.

and Central Africa it has been collected at Die Berg in

Mpumalanga, South Africa. Nyanga Mountain in Zim-

babwe, and the Zomba Plateau in Malawi. It occurs

further into eastern, central and western Africa as far

north as Ethiopia, Sudan, the DRC and Nigeria, and

is also present on Madagascar (Burrows 1990; Roux

2001, 2009). The current record extends its distribu-

tion into the southern Drakensberg range from Mpu-

malanga by 500 km and is a first record for KwaZulu-

Natal (Figure 13).

Specimen examined

KWAZULU-NATAL. — 2929 (Underberg): Cobham Forest Reserve,

200 m northwest of Pinnacle Rock, growing on side of sinkhole in mon-

tane grassland, 2 000 m, (-CB), 19-10-2008, N. Crouch 1241 (PRE).

First record o/Amauropelta bergiana van calva in South

Africa

Amauropelta bergiana (Schltdl.) Holttum is a fairly

commonly encountered fern in deeply shaded forests

associated with mountainous regions receiving high

rainfall. It grows along permanent or seasonally moist

streambanks, riverine scrub and earthbanks in forests

(Burrows 1990; Roux 2001). Amauropelta bergiana var.

bergiana occurs throughout the Afromontane regions of

southern and eastern Africa, from the Cape Peninsula,

through Central Africa to Ethiopia, as well as on Bioko,

Madagascar and Reunion (Burrows 1990; Roux 2001,

2009). Two other varieties of A. bergiana were described

by Holltum (1974): A. bergiana var. calva Holttum

occurs in West Africa, the Comoro Islands and Reunion;

while A. bergiana var. tristanensis Holttum is confined

to the islands of Gough, Inaccessible and Tristan da

Cunha (Roux 2009).

Amauropelta bergiana is a rather large fern with

tufted, deeply 2-pinnatifid fronds of up to 1 m long. The

basal pinnae gradually decrease in size with a pair of

very small, vestigial pinnae usually present. Basal veins

of the pinnae do not unite below the sinus between each

lobe. The typical variety of this species is characterized

by the presence of hooked hairs on the abaxial surface

of the pinnae, and very small indusia. It can easily be

confused with A. knysnaensis (N.C. Anthony & Schelpe)

Parris (endemic to the Knysna area); this latter species

possesses larger indusia set with minute stalked glands,

and lacks the characteristic hooked hairs of A. bergiana

var. bergiana. Another close relative is A. oppositiformis,

which is distinguished by the absence of hooked hairs

and the presence of small golden to red glands on the

lower surface of the frond (Burrows 1990).

On a recent trip to the Umtamvuna Nature Reserve

in southern KwaZulu-Natal, a plant was found that

matched the description of A. bergiana. However, on

closer inspection, it was found that the hairs on the

abaxial surface were not hooked. Furthermore, the plant

lacked glands on either indusia or lamina surfaces, so

eliminating the two aforementioned related species. This

represents the first record of A. bergiana var. calva in

South Africa (Figure 13). This Umtamvuna taxon has

previously only been recorded on mainland Africa from

Cameroon, and as far south as Reunion in the Indian

Ocean (Roux 2009). It is characterized by the presence

of straight hairs on the rachis, costae and costules and

the absence of hairs on the abaxial surface of the lamina

between the veins. It is without hooked hairs (Holttum

1974).

Specimen examined

KWAZULU-NATAL. — 3130 (Port Edward): Umtamvuna Nature

Reserve, growing at the base of a waterfall tumbling into the Bololo

River, 225 m, (-AA), 15-03-2009, N. Crouch 1228 (PRE).

ACKNOWLEDGEMENTS

Ms Sharon Fouw and Mr Richard Nxele of Ezimvelo-

KZN Wildlife are thanked for kindly facilitating field

work; Dr Hugh Glen for the Fatin translation; Mr John

Burrows for giving permission to use distribution maps

of these species from his book Southern African ferns

and fern allies (1990); Ms Hester Steyn, National Her-

barium, SANBI. Pretoria, for producing updated distri-

bution maps.

FIGURE 13. — Distribution of Amauropelta oppositiformis, adapted

from Burrows (1990),#; new locality in KwaZulu-Natal, A.

Distribution of A. bergiana var. calva in the FSA region, ■.

70

Bothalia 40,1 (2010)

REFERENCES

BEENTJE, H.J. 2008. Hymenophyllaceae. In H.J. Beentje & S.A.

Ghazanfar, Flora of tropical East Africa. Royal Botanic Gar-

dens, Kew.

BURROWS, J.E. 1990. Southern African ferns and fern allies. Frand-

sen, Sandton.

BURROWS. J.E. & BURROWS, S.E. 2001. New distribution records

of southern African Pteridophyta. Bothalia 3 1 : 205-207.

HOLTTUM, R.E. 1974. Thelypteridaceae of African and adjacent

islands. Journal of South African Botany 40: 1 23-168.

HUNTLEY, B.J. 1965. A preliminary account of the Ngoye Forest

Reserve, Zululand. Journal of South African Botany 31: 177—

205.

ROUX, J.P. 2001 . Conspectus of southern African Pteridophyta. South-

ern African Botanical Diversity Network Report No. 13. SAB-

ON ET, Pretoria.

ROUX, J.P. 2009. Synopsis of the Lycopodiophyta and Pteridophyta

of Africa, Madagascar and neighbouring islands. Strelitzia 23.

South African National Biodiversity Institute, Pretoria.

SCHELPE, E.A.C.L.E. 1970. Pteridophyta. InA.W. Exeil& E. Launert,

Flora zambesiaca. Crown Agents for Oversea Governments and

Administrations, London.

SCHELPE, E.A.C.L.E. & ANTHONY, N.C. 1986. Pteridophyta. In

O.A. Leistner, Flora of southern Africa. Botanical Research

Institute, Pretoria.

WOOD, J.M. 1 907. A handbook to the flora of Natal. Bennett & Davis,

Durban.

WOOD, J.M. 1908. Revised list of the flora of Natal. Transactions of

the South African Philosophical Society 18,2: 121-280.

R.R. KLOPPER* and N.R. CROUCH**

* Biosystematics Research and Biodiversity Collections Division,

South African National Biodiversity Institute, Private Bag X 1 0 1 , 0001

Pretoria.

** Ethnobotany Unit, South African National Biodiversity Institute,

P.O. Box 52099, Berea Road, 4007 Durban / School of Chemistry,

University of KwaZulu-Natal, 4041 Durban.

MS. received: 2009-06-25.

RUBIACEAE

FIRST RECORD OF GEOPHILA IN SOUTHERN AFRICA

Geophila D.Don, a small herbaceous genus of forest

floor perennials, has previously not been recorded from the

Flora of southern Africa (FSA) region (Leistner 2005), the

closest known locality being in eastern Zimbabwe (Gov-

aerts et al. 2009). The genus comprises more than 20 spe-

cies from both the Old and New World (Mabberley 2008;

Govaerts et al. 2009), with only three taxa reported in the

Flora zambesiaca (FZ) region (Verdcourt 1989).

In July of 2008, red-fruiting plants of Geophila were

encountered in shade alongside the road traversing Ngoye

Forest in Zululand (Figure 14). This low-growing herb

occupied the poorly developed field layer community

of climax forest (Huntley 1965) in vegetation classified

recently as Scarp Forest (FOz5) (Rutherford et al. 2006).

Material was grown on to flowering in Everton near Durban

and a voucher specimen prepared. A return trip to Ngoye

Forest in late May 2009 enabled the gathering of fruiting

material from the colony, which occupies an area of ± 12

nr. Comparison with literature revealed this collection to

be Geophila repens (L.) I. M. Johnston, a pantropical herb

of evergreen forest floors, widespread in both the Old and

New World (Verdcourt 1989). The nearest known locality

of this species lies some 940 km north of Ngoye Forest

in Zimbabwe’s Chirinda Forest (Drummond & Mapaure

1994), a medium altitude rainforest (Muller 1999) within

the Chimanimani-Nyanga Centre (CIC) of Endemism (Van

Wyk & Smith 2001). These Geophila repens records rep-

resent a significant range extension, and they also serve

to confirm the close floristic relationship of the rainforests

of eastern Zimbabwe with Ngoye Forest — as part of the

Maputaland-Pondoland Region of Floristic Endemism,

approximating the Tongaland-Pondoland Regional Mosaic

of White (1983) (Muller 1999). Various pteridophytes (Bur-

rows & Burrows 2001), lianes ( Urera trinervis), shrubs

( Pseuderanthemum subviscosum) and grasses ( Olyra latifo-

lia ) exhibit similar disjunctions. The distributions of certain

fauna such as the butterfly Euriphene achlys further reflect

this pattern (Swanepoel 1953). Whereas some such taxa

remain regionally localized within Ngoye Forest (Klopper

& Crouch 2010), a number extend their range southwards

to Pondoland (Huntley 1965; Van Wyk & Smith 2001).

The Geophila species occurring nearest to Ngoye For-

est is Geophila obvallata (Schumach.) F.Didr., which was

recorded from the Maputo region of Mozambique (Verd-

court 1989), somewhat closer than Chirinda Forest. This

species is associated with a much broader range of habi-

tats than G. repens , which is restricted to evergreen forest.

Geophila repens is distinguished from G. obvallata by its

bright red or orange rather than black, purple or blue ber-

ries. Furthermore, the inflorescences of G. repens are usu-

ally 1-flowered in our region and without an involucre,

whereas those of G. obvallata are always several-flowered

and subtended by a distinct involucre comprising separate

bracts. The style of G. repens is between 3. 5-7.0 mm long

vs not more than 1 mm in G. obvallata (Verdcourt 1989).

The full synonymy for Geophila repens , with species

description and illustration, is provided by Verdcourt

(1989).

FIGURE 14. — Distribution of Geophila repens in the FSA region, •.

Bothalia 40,1 (2010)

71

Specimens examined

KWAZULU-NATAL. — 2831 (Nkandla): Ngoye Forest Reserve,

± 3 km west of gauge weir in forest, on northern roadside verge in

shade, 440 m, grown on to flowering in Everton, (-DC), 20-05-2009,

R. Edwards 1 (PRE), fruiting, 24-05-2009, R. Edwards 2 (PRE).

ACKNOWLEDGEMENTS

Mr D. Styles is thanked for helpful discussions

regarding the identity of the discovery; Mrs H. Snyman,

SANBI. for preparing the map; and the staff of the Mary

Gunn Library, SANBI, for facilitating access to litera-

ture. This note is dedicated to the late Mr Ian Garland

who for many years recognized the uniqueness of Ngoye

Forest and surrounds, and actively supported its botani-

cal exploration.

REFERENCES

BURROWS. J.E. & BURROWS, S.E. 2001. New distribution records

of southern African Pteridophyta. Bothalia 3 1 : 205-207.

DRUMMOND, R.B. & MAPAURE, I. 1994. List of flowering plants

and ferns. Appendix 1. In J. Timberlake & P. Shaw, Chirinda For-

est— a visitor s guide. Forestry Commission, Harare, Zimbabwe.

GOVAERTS, R„ RUHSAM, M„ ANDERSSON, L„ ROBBRECHT,

E., BRIDSON, D.. DAVIS, A., SCHANZER, I. & SONKE, B.

2009. World checklist ofRubiaceae. The Board of Trustees of the

Royal Botanic Gardens, Kew. Website: http://apps.kew.org/wcsp

(accessed 10-09-2009).

FIUNTLEY, B.J. 1965. A preliminary account of the Ngoye Forest Reserve,

Zululand. Journal of South Afiican Botany 31:1 77-205.

KLOPPER, R. & CROUCH, N.R. 2010. New distribution records

and noteworthy collections of pteridophytes in KwaZulu-Natal

(Pteridophyta). Bothalia 40: 68-70.

LEISTNER, O.A. 2005. Seed plants of southern tropical Africa: fami-

lies and genera. Southern African Botanical Diversity Network

Report No. 26. SABONET, Pretoria.

MABBERLEY, D.J. 2008. Mabberleys plant-book: a portable diction-

ary of plants, their classification and uses, edn 3. Cambridge

University Press.

MULLER, T. 1 999. The distribution, classification and conservation of

rainforests in Zimbabwe. In J. Timberlake & S. Kativu, African

plants: biodiversity, taxonomy and uses: 221-235. Royal Botan-

ic Gardens, Kew.

RUTHERFORD, M.C., POWRIE, L.W., LOTTER, M.C., VON

MALTITZ, G.P., EUSTON-BROWN, D.I.W., MATTHEWS,

W.S.. DOBSON, L. & McKENZIE, B. 2006. Afrotemperate,

subtropical and azonal forests. In L. Mucina & M.C. Ruther-

ford, The vegetation of South Africa, Lesotho and Swaziland.

Strelitzia 19: 584-614.

SWANEPOEL, D.A. 1953. Butterflies of South Africa. Maskew Miller,

Cape Town.

VAN WYK, A.E. & SMITH, G.F. 2001 . Regions offloristic endemism in

southern Africa. A review with emphasis on succulents. Umdaus

Press, Hatfield, Pretoria.

VERDCOURT, B. 1989. Rubiaceae. Flora zambesiaca 5, 1 : 1-210.

WHITE, F. 1983. The vegetation of Africa: a descriptive memoir to

accompany the UNESCO/AETFAT/UNSO vegetation map of

Africa. Natural Resources Research, XX. UNESCO, Paris.

N.R. CROUCH* and R. EDWARDS**

* Ethnobotany Unit, South African National Biodiversity Institute, P.O.

Box 52099, 4007 Berea Road. Durban / School of Chemistry, Univer-

sity of KwaZulu-Natal, 4041 Durban.

** 18 Acutts Drive, 3610 Everton, KwaZulu-Natal.

MS. received: 2009-05-26.

PTERIDOPHYTA

NOTES ON SOME NATURALIZED FERNS OF THE EASTERN CAPE AND KWAZULU-NATAL

BLECHNACEAE

Doodia caudata

There has been some confusion as to the exact iden-

tity of the Doodia species (Figure 15 A) naturalized in

KwaZulu-Natal. South Africa. Specimens from Pieter-

maritzburg collected in the mid-20th century were first

identified as Doodia media R.Br. by Prof. E. Schelpe.

Subsequently, Burrows (1990) referred this taxon to

Doodia caudata (Cav.) R.Br., and more recently it has

been assigned to Doodia squarrosa Colenso (Roux 2001.

2009). The fact that no single identification key to all

Doodia species exists has confounded attempts to verify

the identity of the South African naturalized species.

There are separate keys available to the New Zealand

(Parris 1972) and Australian (Parris 1998) taxa of Doo-

dia. However, D. caudata is an Australian entity (Parris

1980, 1998), whereas D. squarrosa is confined to north-

ern New Zealand (Parris 1972). Although D. caudata was

initially considered to occur also in New Zealand (Parris

1 972), plants so associated with this name were described

subsequently as a New Zealand endemic, Doodia mollis

Parris (Parris 1980). Accordingly, the two available keys

are not definitive in respect of resolving D. squarrosa and

D. caudata.

There is a high level of morphological variation in

most species of Doodia; accordingly D. caudata and

D. squarrosa are apparently often confused in herbaria.

Both have pinnate dimorphic fronds with discrete to

confluent sori, and pinnae that are auriculate or stalked

to partly adnate in the middle and adnate to decurrent

in the upper part of the frond (Parris 1972, 1998). The

fronds of D. caudata are 57^-60 mm long with the long-

est pinnae 2-110 x 1-4 mm (Parris 1998), whereas the

fronds of D. squarrosa are of similar maximum length,

140—450 mm long, but with the longest pinnae only 15-

60 x 1-2 mm (Parris 1972). Frond and pinnae length are

not always useful distinguishing characters especially

since they are frequently environmentally influenced.

South African material was compared to the lectotype

of Doodia squarrosa ( Colenso 9/83 , frond second from

right, K) and a high resolution image of the holotype of

D. caudata (L.Nee s.n. MA). It was found to match the

material for D. caudata more closely. During subsequent

correspondence with Dr B. Parris, she confirmed the

identity of the South African material as D. caudata , the

most variable species in the genus and the one that has

become the most widely naturalized. According to Par-

ris (pers. comm.) the South African material approaches

D. media R.Br. var. moorei Baker in having rather longer

fertile pinnae than the typical form; this aforementioned

variety intergrades with the typical form in Australia and

72

Bothalia 40,1 (2010)

FIGURE 15. — A, Doodia caudata , habit: B, Phlebodium aureum , habit. Photographs: N.R. Crouch.

cannot be upheld. It has consequently been placed in

synonymy under D. caudata (Parris 1998).

Accordingly, the Doodia species that has escaped

from cultivation in KwaZulu-Natal is D. caudata, the

small rasp fern.

Doodia caudata (Cav.) R.Br. in Prodromus florae

Novae-Hollandia: 151 (1810); Burrows: 338(1990). Type:

Nova Hollandia [Australia], L.Nee s.n. (MA, holo.).

D. squarrosa sensu Roux: 153 (2001) et sensu Roux: 105 (2009),

non Colenso: 382 (1881).

For a complete list of synonyms see Parris (1998).

Rhizome erect, up to 0. 1 m tall, to short- or long-

creeping; rhizome scales brown. Fronds markedly

dimorphic; stipe slender, short, dark towards base and

without tubercules. Fertile lamina generally more erect

and scabrous, longer and broader than sterile ones,

57-460 x 5-180 mm, with a long apical segment (V3-)

V —L of lamina length, 1 -pinnate; pinnae more distant

and narrower than sterile ones, simple, oblong to linear-

lanceolate, margins sharply toothed, more than 2 pairs

of pinnae stalked in basal third of lamina, stalked to

partly adnate to decurrent in middle third, partly adnate

to decurrent in distal third, longest pinnae 2-110 x 1-4

mm, rachis and pinna midvein without tubercules. Ster-

ile lamina decumbent, I -pinnate, pinnae simple, oblong,

apices rounded, margins toothed. Venation with some

anastomoses forming areoles on either side of costa. Sori

in I (rarely partial second) row on each side of costa,

0.8-10 (or more) x 0.8- 1.5 mm, often laterally confluent

when mature, confluent across costa, nearer to midvein

than margin; indusium linear, entire to repand, opening

towards the costa (Andrews 1990; Parris 1998).

Doodia caudata is a native of Australia where it

occurs in eastern South Australia, eastern Queensland,

New South Wales, southern Victoria, northern Tasmania

and Lord Howe Island (Parris 1998). It has naturalized

in KwaZulu-Natal, South Africa (Burrows 1990; Roux

2001, 2009), California (LISA), as well as the Azores,

Madeira and in Sri Lanka (Burrows 1990; Parris 1998).

It can locally be found growing exposed or in deep shade

on seasonally moist earthbanks, roadside cuttings and

forest margins (Crouch 1994; Roux 2001).

An attempt has been made to trace the original source

of the Doodia material imported into South Africa. Since

the South African material most closely resembles the

form described as D. media var. moorei which was col-

lected from the Richmond River north of Sydney in New

South Wales, the most likely source would be what is

today known as the Royal Botanic Garden of New South

Wales, Australia (B.S. Parris pers. comm.). Reports of

the Natal Botanic Gardens in Durban from a century

ago, reveal that its Curator, J.M. Wood was regularly

exchanging plant material with the Botanic Gardens in

Sydney, and from the accession register of ‘Overseas and

Tropical African specimens ex Wood Herbarium. Vol. I’

it is apparent that J.H. Maiden of the Sydney Gardens

provided Wood with plants of D. caudata for cultivation.

A voucher (J.M. Wood 5766 NH) at the KwaZulu-Natal

Herbarium indicates that this species was first grown in

the Natal Botanic Gardens (Durban Botanic Gardens)

prior to the mid- 1890s. Notably, this particular speci-

men was correctly named D. caudata and has not been

subject to re-determinations, likely due to its placement

in the cultivated section of the NH collection. As Wood

also exchanged live material with the Pietermaritzburg

Botanic Garden at that time, plants from the Durban

facility may have been transferred to the Pietermaritz-

burg Garden, from where they escaped.

Although Doodia caudata is locally common across

the mistbelt zone of Pietermaritzburg (Figure 16), from

Sunnyside to Ferncliffe ( 1 000 m), this species has

apparently not naturalized elsewhere in the Midlands,

Bothalia 40,1 (2010)

73

FIGURE 16. — Distribution in FSA region of Doodia caudata, • ; and

Phlebodium aureum, A.

besides Howick (Crouch 1994) — this since its docu-

mented escape from cultivation over half a century ago.

It was somewhat surprising then to encounter this taxon

growing in a remote site near the Transkei coast under

different climatic conditions, indicating that it may now

be an emerging weed. This record extends the previous

distribution range southwards some 170 km (Figure 16).

Specimens examined

KWAZULU-NATAL. — 2930 (Pietermaritzburg): Sunnyside, Pie-

termaritzburg, (-CB), 01-03-1953, A. Dohse & L. Lindahl 102 (NH);

Maritzburg District, 4000 ft [1 200 m], (-CB), 16-08-1956, ACS 10

(NU); Femclifife Nature Reserve, Pietermaritzburg, 1 000 m, (-CB),

23-02-1992, N.R. Crouch 561 (NU); Femcliffe Nature Reserve, earth

bank adjacent to parking area, 1 000 m, (-CB), 30-04-2007, N.R.

Crouch 1153 (PRE); KwaZulu-Natal National Botanical Garden, grow-

ing alongside path on banks, in indigenous forest section on far side of

river in garden, (-CB). 03-09-2008, N.R. Crouch 1172 (PRE); oppo-

site 1 1 Tanner Rd, Wembley, Pietermaritzburg, colony on earth bank,

(-CB), 06-01-2008, N.R. Crouch 1145 (NH). 2931 (Stanger): Natal

Botanic Garden, in cultivation, (-CC), J.M. Wood 5766 (NH).

EASTERN CAPE. — 3130 (Port Edward): Lukabem stream junc-

tion with Mtentu River, 30 m upstream of high tide mark, small colony

growing on earthbank in coastal forest, 4 m, (— AA), 04-01-2008, N.R.

Crouch 1143 (NH).

POLYPODIACEAE

Phlebodium aureum

Phlebodium aureum (L.) J.Sm. (Figure 15B) is com-

monly cultivated in South Africa and has escaped from

cultivation in the vicinity of Durban, KwaZulu-Natal and

Port St Johns in the Eastern Cape (Burrows 1990). It has

also been reported to have become naturalized in Zimba-

bwe (Roux 2009). This fern species was documented in

cultivation in the Durban Botanic Gardens in 1941 (C.

Kent 16 NH). and to have started escaping in the immedi-

ate vicinity of the gardens by 1968 ( R.G . Strey 8071 NH).

It has more recently been collected in Kloof some 25 km

distant, and has been sighted and photographed growing

on a decaying log in remnant indigenous riverine scrub,

at Inanda to the north of the metropol (Figure 16).

The golden polypody or golden serpent fern is so

named because of the golden scales on its creeping rhi-

zome. Its arching to pendent fronds are deeply lobed

(Nauman 1993) and very similar to those of Microsorum

scolopendria (Burm.f.) Copel. It is distinguished from

this indigenous fern by the venation of the lamina (fewer

included veinlets), its very dense tomentum of golden

scales clothing the rhizome, the close spacing of the

pinnatifid segments and the sori which are not sunken

into the lamina (Schelpe & Anthony 1986). It has been

reported to grow epiphytically in moist evergreen river-

ine forests and forest margins (Roux 2001, as Polypo-

dium aureum L.), and is also found in Durban growing

as a lithophyte in more exposed situations, and on build-

ing ledges in a manner reminiscent of Pteris vittata L.

Phlebodium aureum is native to Central America

from Florida and Georgia in the USA, through Mexico

to Brazil in South America and the West Indies (Burrows

1990; Nauman 1993; Roux 2001). Plants are report-

edly intolerant of heavy frosts which would account

for limited naturalization of P. aureum along the warm

eastern seaboard. Additionally, although usually ever-

green, plants may be briefly deciduous during periods of

drought stress, so allowing them to invade relatively dry

areas such as man-made structures. Although it is recog-

nized as a garden escape, no herbarium specimens docu-

menting it as a truly naturalized species could be traced

for South Africa; this report with associated voucher cor-

rects this insufficiency.

Phlebodium aureum (L.) J.Sm. in Journal of Bot-

any 4: 59 (1842). Polypodium aureum L.: 1087 (1753).

Type; Herb. Linn. No. 1251.10 (LINN, lecto.), desig-

nated by Proctor (1977: 334).

Rhizome creeping, ± 8— 1 5(— 30) mm diam., densely

scaled; rhizome scales reddish to golden, long attenuate,

1 0-20 mm. Fronds bright green or glaucous, arching to

pendent, 0.3-1. 3 m; stipe 150-500 mm, smooth, with a

few scales near base; lamina deeply pinnatifid, 300-800

x 100-500 mm, glabrous, terminal segment conform;

pinnae up to 35, lanceolate to elliptic, or linear-lanceo-

late to linear, 60-200 x 10-40 mm, margins entire, occa-

sionally undulate. Sori in a single line on each side of

costae, occasionally a 2nd row present, terminal or at a

plexus of included veinlets; exindusiate (Nauman 1993).

Specimens examined

KWAZULU-NATAL. — 2930 (Pietermaritzburg): Krantzkloof Na-

ture Reserve, Kloof, at eastern end of reserve, immediately above The

Splash, growing as a lithophyte adjacent to Molweni River in full sun,

218 m, (-DD), 28-04-2009, N.R. Crouch 1243 (PRE); Kloof, opposite

Maytime Centre on ledge of pedestrian bridge across the M 1 3 highway

on eastbound carriageway, 450 m, (-DD), 27-04-2009, N.R. Crouch

1244 (PRE). 2931 (Stanger): Durban Botanic Gardens, (-CC), 06-