Seaweeds of the South African South Coast

Book 4: Phylum Rhodophyta (Red Seaweeds)
Order Gigartinales to Order Rhodymeniales

By

Robert James Anderson
University of Cape Town and Department of Agriculture, Forestry and Fisheries, South Africa

Herre Stegenga
Naturalis, Netherlands

John Joseph Bolton
University of Cape Town, South Africa

2020

Herre Stegenga's line drawings from Seaweeds of the South African West Coast
(Contributions from the Bolus Herbarium, Vol 18, 1997) are reproduced with permission.  

Copyright © 2020 Robert Anderson et al. All rights reserved.

Copyright © 2020 by Robert Anderson et al.
All rights reserved. No part of this publication
may be reproduced, distributed, or transmitted in any
form or by any means, including photocopying, recording,
or other electronic or mechanical methods,
without the express written permission of the authors
except for the use of brief quotations in a book review.

Printed in the Republic of South Africa

First Edition, 2020

ISBN 0-0000000-0-0

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Private Bag X3,
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Ordering Information:
This eBook can be downloaded free from: southafrseaweeds.uct.ac.za/

INTRODUCTION TO THE RHODOPHYTA (red seaweeds)

The Phylum Rhodophyta is commonly known as the ‘red algae’. Most species of red algae are seaweeds, but there are also many different forms, including single-celled organisms on soil and various forms in rivers and lakes. A fossil red alga from Canada, Bangiomorpha pubescens, is the oldest known record of eukaryotic life on earth, dated at around 1200 million years old. Yoon et al. (2017) list 7 classes of Rhodophyta, although almost all the seaweed species here are in the class Florideophyceae, with very few exceptions such as species of Porphyra (Bangiophyceae) and Stylonema (Stylonematophyceae). Red seaweeds grow in all marine environments where there is sufficient light, from the intertidal to over 200m depth in very clear tropical seas. The larger species are smaller than the largest brown algae, with a maximum of around 3m worldwide. The largest recorded in South Africa are specimens of Pachymenia orbitosa on the west coast, which grow to around 2m. On the south coast, the largest species are not more than around 50cm long. Red seaweeds make up ca. 70% of the seaweed species present in South Africa.

The common name ‘red algae’ can be confusing for some members of this group. Most species of red algae look various shades of red (from dark burgundy to pink), due to the dominance of the red-coloured pigment phycoerythrin. The Rhodophyta, however, contain other pigments including chlorophylls (green), carotenoids (yellow to brown) and phycocyanin (blue-green). Sometimes these other pigments dominate in the plant or species, for example many freshwater ‘red algae’ are blue-green in colour. Among south coast seaweeds, one of the most abundant , Hypnea spicifera, is often almost completely green, with a few darker reddish parts where it is shaded at the base. High intertidal species such as Porphyra and Bostrychia often appear blackish. Also, some bladed shallow water species, which occur in high light, appear more brownish than reddish. It should be remembered that most species of seaweeds on this coast are red algae, and if plants are reddish in some small part they are likely to be members of the Rhodophyta.

The red seaweeds of the South African south coast are divided here into four ebooks based on their taxonomic position (hence evolutionary relationships), using data from Algaebase.org (Guiry and Guiry 2016).This series contains keys to the species within each genus, but no key or picture guide to higher-level groups of taxa, because this is often not possible by simple observation. The high-level taxa (sub-phyla, classes, orders) were traditionally defined using microscopic characteristics, and nowadays mostly by DNA sequencing. In marine guidebooks containing a small selection of common species, they are often arranged in ‘form groups’, which although useful, do not follow the evolutionary relationships between them (e.g. Branch et al. 2017). Such guidebooks remain a useful adjunct to the present series.

The forms of red seaweeds range from small threadlike filaments, one to many cells wide, to various sizes of branched or unbranched, cylindrical to leafy plants. Most of the species diversity in the red algae on the south coast is comprised of small filamentous species, many of which grow attached to larger algae (epiphytes). These are mostly in the families Ceramiaceae (e.g. Ceramium), Rhodomelaceae (e.g. Polysiphonia, Pterosiphonia, Herposiphonia) and Callithamniaceae (e.g. Callithamnion). A few are crusts, which form a very thin layer growing closely attached as a thin ‘skin’ over the surface of rocks (e.g. Hildenbrandia rubra). A particularly important form group of red seaweeds is the coralline red algae, which deposit significant amounts of calcium carbonate in their cell walls, forming a hard, internal skeleton. Many of these exists as crusts, and these have been fairly well studied in South Africa (Maneveldt et al. 2016). The crustose coralline red algae require specific techniques to collect, identify and study, including chiselling them off the rock and dissolving away the calcium carbonate with dilute acid to reveal the critical microscopic structures. They are not included in these ebooks. The other form group of coralline algae is known as geniculate (articulated or jointed) corallines and they exist as upright seaweeds (see Book 4 in this series). They consist of calcified segments (genicula), separated by non-calcified tissue (intergenicula) which allows flexibility (e.g. species of Amphiroa, Arthrocardia, Jania). Many of these geniculate corallines are particularly difficult to identify to species.

Red seaweeds are ecologically dominant on many local seashores. Porphyra is not as abundant on the south coast as the west coast, but patches of Porphyra agulhensis, where present, are the uppermost seaweeds in the south coast intertidal, with Bostrychia intricata in upper shore crevices. Many south coast seashores tend to be animal dominated (limpets and barnacles) in the mid-intertidal, with most seaweed diversity in rock pools. In the lower intertidal and shallow subtidal there is often dominance of a wide variety of red seaweeds. In most western areas of the south coast, this are often various species of Laurencia, Hypnea (especiallyHypnea spicifera) and Plocamium (especially Plocamium corallorhiza, one of the most abundant red seaweeds in South Africa). Further east there is an increasing dominance of geniculate coralline red seaweeds (species of Amphiroa, Arthrocardia etc.: Kogame et al. 2016).

Red seaweeds have the most complicated life cycles in the algae, and the reproductive structures are often important in species identification. Most species exist as three different types of plants on the seashore:

- Sporophytes: In the Florideophyceae usally called ‘tetrasporophytes’ as they mostly produce tetraspores in packets of four (see Callithamnion tripinnatum). A few produce more than four, such as the polysporangia of Pleonosporium filicinum.

- Female gametophytes: A few larger seaweeds have visible rounded structures on the surface called ‘cystocarps’, which produce carpospores (e.g. Martensia elegans or Sarcodia dentata). More often these structures can only be seen by microscopic examination (e.g. Platysiphonia delicata), sometimes after cutting a cross-section of the tissue in bladelike species.

- Male gametophytes: These are often known only as the plants without tetraspores or cystocarps, as the male reproductive structures can be difficult to see even with a compound microscope. In some small species, they are easier to see under the microscope, but may not be easy to find (e.g. Ceramium glanduliferum).

The three different plants with their respective reproductive structures are clearly shown in the micrographs of Lophosiphonia capensis. Many species of red algae have populations which are dominated by tetrasporophytes, and a few (especially members of the Gigartinaceae) are dominated by gametophytes.

Some forms of red algae, particularly fine filamentous seaweeds and some leaflike blades, cannot be identified to species without microscopic study at 400x magnification. The descriptions here thus contain microscope pictures of filaments or sections of blades wherever possible.

SYSTEMATIC CLASSIFICATION OF SOUTH COAST RHODOPHYTA

Note: the list below of taxonomic placing of genera was revised in November 2016, using AlgaeBase. The higher-level taxonomic placing of certain genera may differ from that shown under each genus or species on this website, because some of these have not yet been updated. When in doubt see AlgaeBase.org

BOOK 3

CLASS BANGIOPHYCEAE

Sub Class Bangiophycidae

Order Bangiales

FAMILY BANGIACEAE
      Porphyra capensis Kützing
      Porphyra sp. 1

CLASS STYLONEMATOPHYCEAE

Order Stylonematales

FAMILY STYLONEMATACEAE
      Stylonema alsidii (Zanardini) Drew
      Stylonema cornu-cervi Reinsch (see under S. alsidii)

CLASS COMPSOPOGONOPHYCEAE

Order Erythropeltidales

FAMILY ERYTHROTRICHIACEAE
      Erythrotrichia bertholdii  Batters
      Erythrotrichia platyphylla Stegenga, Engledow, Bolton & Anderson
      Erythrotrichia welwitschii (Ruprecht) Batters
      Membranella africana Stegenga, Bolton & Anderson
      Porphyrostromium boryanum (Montagne) P.C. Silva in Silva, Basson & Moe
      Sahlingia subintegra (Rosenvinge) Kornmann

CLASS FLORIDEOPHYCEAE

Sub Class Hildenbrandiophycidae

Order Hildenbrandiales

FAMILY HILDENBRANDIACEAE
      Hildenbrandia lecanellierii Hariot
      Hildenbrandia rubra (Sommerfelt) Meneghini

Sub Class Nemaliophycideae

Order Acrochaetiales

FAMILY ACROCHAETIACEAE
      Acrochaetium brebneri (Batters) G. Hamel
      Acrochaetium catenulatum M.A.Howe
      Acrochaetium hlulekaense Stegenga
      Acrochaetium moniliforme (Rosenvinge) Børgesen
      Acrochaetium reductum (Rosenvinge) G.Hamel
      Acrochaetium tenuissimum (F.S.Collins) Papenfuss

Order Colaconematales

FAMILY COLACONEMATACEAE
      Colaconema caespitosum (J.Agardh) Jackelman, Stegenga & J.J.Bolton
      Colaconema codicola (Børgesen) H.Stegenga, J.J.Bolton, & R.J.Anderson
      Colaconema daviesii (Dillwyn) Stegenga
      Colaconema dictyotae (Collins) I.-K.Hwang & H.-S.Kim
      Colaconema endophyticum (Batters) J.T.Harper & G.W.Saunders
      Colaconema interpositum (Heydrich) H.Stegenga, J.J.Bolton & R.J.Anderson
      Colaconema nemalionis (De Notaris ex L. Dufour) Stegenga
      Colaconema pectinatum (Kylin) J.T.Harper & G.W.Saunders
      Colaconema seriatum (Børgesen) Stegenga
      Colaconema spongicola (Weber-van Bosse) Woelkerling

Order Nemaliales

FAMILY GALAXAURACEAE
      Dichotomaria diesingiana (Zanardini) Huisman, J.T. Harper & G.W. Saunders
      Dichotomaria hommersandii L. Liu & S-M Lin
      Dichotomaria obtusata (J. Ellis & Solander) J.V. Lamouroux
      Dichotomaria tenera (Kjellman) Huisman, J.T. Harper & G.W. Saunders

FAMILY HELMINTHOCLADACEAE
      Helminthocladia australis Harvey
      Helminthora furcellata (Reinbold in Tyson) Martin

FAMILY SCINAIACEAE
      Nothogenia erinacea (Turner) Parkinson
      Nothogenia ovalis (Suhr) Parkinson
      Scinaia capensis (Setchell) Huisman
      Scinaia salicorniodes (Kützing) J. Agardh

Order Gelidiales

FAMILY GELIDIACEAE
      Gelidium abbottiorum R. Norris
      Gelidium arenarium Kylin
      Gelidium capense (S.G. Gmelin) P.C. Silva in P.C. Silva, E.G. Meñez, & Moe
      Gelidium crinale (Hare ex Turner) Gaillon
      Gelidium declerckii E. M. Tronchin in Millar and Freshwater
      Gelidium foliaceum (Okamura) E. M. Tronchin
      Gelidium isabelae W.R.Taylor
      Gelidium micropterum Kützing
      Gelidium pristoides (Turner) Kützing
      Gelidium pteridifolium R.E. Norris, Hommersand and Fredericq
      Gelidium reptans (Suhr) Kylin
      Pterocladiella caespitosa (Kylin) Santelices
      Pterocladiella caloglossoides (M.A. Howe) Santelices
      Ptilophora diversifolia (Suhr) Papenfuss
      Ptilophora pinnatifida J. Agardh

Order Palmariales

FAMILY RHODOTHAMNIELLACEAE
      Rhodothamniella floridula (Dillwyn) Feldmann in T.Christensen

FAMILY MEIODISCACEAE
      Meiodiscus concrescens (K.M.Drew) P.W.Gabrielson in Gabrielsen et al.

Sub Class Rhodymeniophycidae

Order Bonnemaisoniales

FAMILY BONNEMAISONIACEAE
      Asparagopsis armata Harvey
      Asparagopsis taxiformis (Delile) Trevisan de Saint-Léon
      Delisea flaccida (Suhr) Papenfuss

Order Peyssonneliales

FAMILY PEYSSONNELIACEAE
      Peyssonnelia atropurpurea Crouan & Crouan
      Peyssonnelia replicata Kützing
      Sonderopelta capensis (Montagne) D.M. Krayesky

Order Halymeniales

FAMILY HALYMENIACEAE
      Codiophyllum natalense J. E. Gray
      Cryptonemia natalensis (J. Agardh) Chiang
      Grateloupia capensis O. De Clerck ex De Clerck, Gavio, Fredericq, Barbara & Coppejans
      Pachymenia cornea (Kützing) Chiang
      Pachymenia ulvoidea (F. Schmitz) L. Russell in Russell, Hurd, Nelson & Broom
      Polyopes constrictus (Turner) J. Agardh
      Prionitis filiformis Kylin
      Prionitis nodifera (Hering) Barton

Order Gracilariales

FAMILY GRACILARIACEAE
      Gracilaria aculeata (Hering) Papenfuss
      Gracilaria beckeri (J.Agardh) Papenfuss
      Gracilaria capensis F. Schmitz ex Mazza
      Gracilaria denticulata F. Schmitz ex Mazza
      Gracilaria gracilis (Stackhouse) M.Steentoft, L.M.Irvine & W.F.Farnham
      Gracilariopsis funicularis Iyer, Bolton & Coyne
      Gracilariopsis longissima (S.G.Gmelin) M.Steentoft, L.M.Irvine & W.F.Farnham

FAMILY PTEROCLADIOPHILACEAE
      Gelidiocolax margaritoides (M.T.Martin & M.A.Pocock) K.-C.Fan & Papenfuss

BOOK 4

Order Gigartinales

FAMILY KALLYMENIACEAE
      Callophyllis variegata (Bory de Saint-Vincent) Kützing
      Pugetia harveyana (J. Agardh) R.E. Norris
      Pugetia porphyroidea (F. Schmitz ex Holmes) R.E. Norris
      Thamnophyllis pocockiae R.E. Norris

FAMILY GYMNOPHLOEACEAE
      Schizymenia apoda (J. Agardh) J. Agardh

FAMILY PHACELOCARPACEAE
      Phacelocarpus oligacanthus Kützing
      Phacelocarpus tortuosus Endlicher & Diesing

FAMILY SOLIERIACEAE
      Callophycus densus (Sonder) Kraft
      Callophycus tridentifer Kraft
      Rhabdonia natalensis Reinbold

FAMILY ARESCHOUGIACEAE
      Erythroclonium corallinum (Suhr) Holmes

FAMILY CAULACANTHACEAE
      Catenella nipae Zanardini
      Caulacanthus ustulatus (Turner) Kützing
      Heringia mirabilis (C. Agardh) J. Agardh

FAMILY CYSTOCLONIACEAE
      Calliblepharis fimbriata (Greville) Kützing
      Hypnea arenaria Kylin
      Hypnea intricata Kylin
      Hypnea musciformis (Wulfen) J. V. Lamouroux
      Hypnea rosea Papenfuss
      Hypnea spicifera (Suhr) Harvey
      Hypnea tenuis Kylin
      Hypnea viridis Papenfuss
      Rhodophyllis reptans (Suhr) Papenfuss

FAMILY RHIZOPHYLLIDACEAE
      Portieria hornemannii (Lyngbye) P.C. Silva in P.C. Silva, Meñez & Moe
      Portieria tripinnata (Hering) De Clerck in De Clerck, Tronchin & Schils

FAMILY PHYLLOPHORACEAE
      Gymnogongrus glomeratus J. Agardh
      Gymnogongrus polycladus (Kützing) J. Agardh
      Gymnogongrus tetrasporifer Papenfuss in R.J.Anderson & Bolton

FAMILY GIGARTINACEAE
      Chondracanthus acicularis (Roth) Fredericq in Hommersand, Guiry, Fredericq & Leister
      Gigartina insignis (Endlicher & Diesing) Schmitz in Barton
      Gigartina minima Kylin
      Gigartina paxillata Papenfuss
      Gigartina pistillata (S.G. Gmelin) Stackhouse
      Gigartina polycarpa (Kützing) Setchell & Gardner

Order Plocamiales

FAMILY PLOCAMIACEAE
      Plocamiocolax papenfussiana Martin & Pocock
      Plocamium beckeri Simons
      Plocamium corallorhiza (Turner) J.D. Hooker & Harvey
      Plocamium cornutum (Turner) Harvey
      Plocamium glomeratum J. Agardh
      Plocamium maxillosum (Poiret) J.V. Lamouroux
      Plocamium rigidum Bory de Saint-Vincent in Bélanger & Bory de Saint-Vincent
      Plocamium suhrii Kützing
      Plocamium telfairiae (W. J. Hooker & Harvey) Harvey ex Kützing

FAMILY SARCODIACEAE
      Sarcodia dentata (Suhr) R. E. Norris
      Trematocarpus flabellatus (J.Agardh) De Toni

Order Rhodymeniales

FAMILY RHODYMENIACEAE
      Botryocladia beckeriana (Holmes) Papenfuss
      Botryocladia madagascariensis G. Feldmann
      Botryocladia pienaarii RE Norris
      Ceratodictyon variabile (Greville ex J. Agardh) RE Norris
      Erythrymenia obovata Schmitz ex Mazza
      Rhodymenia capensis J. Agardh
      Rhodymenia holmesii Ardissone
      Rhodymenia natalensis Kylin

FAMILY CHAMPIACEAE
      Champia compressa Harvey
      Champia parvula (C. Agardh) Harvey
      Chylocladia capensis Harvey

FAMILY HYMENOCLADIACEAE
      Hymenocladia kallymenioides (Holmes) F. Schmitz

FAMILY LOMENTARIACEAE
      Lomentaria amplexans RE Norris
      Lomentaria corallicola Børgesen
      Lomentaria diffusa Stegenga, Bolton & Anderson

BOOK 5

Order Ceramiales (in part)

FAMILY CALLITHAMNIACEAE
      Callithamnion collabens (Rudolphi) L.McIvor & Maggs
      Callithamnion cordatum Børgesen
      Callithamnion granulatum (Ducluzeau) C. Agardh
      Callithamnion hookeri (Dillwyn) S.F. Gray
      Callithamnion stuposum Suhr
      Callithamnion tripinnatum C. Agardh
      Callithamnion sp. 1
      Callithamnion sp. 2
      Crouania attenuata (C. Agardh) J. Agardh
      Crouania francescoi Cormaci, G Furnari et Scammacca
      Euptilota mollis (Wollaston) O. De Clerck
      Heteroptilon pappeanum (Kützing) Hommersand in Hommersand, D.W. Freshwater, J. López-Bautista, & S. Fredericq

FAMILY CERAMIACEAE
      Acrothamnion preissii (Sonder) E.M. Wollaston
      Amoenothamnion lycopodioides Stegenga
      Antithamnion diminuatum Wollaston
      Antithamnion hubbsii E.Y. Dawson
      Antithamnionella spirographidis (Schiffner) E.M. Wollaston
      Antithamnionella ternifolia (J.D.Hooker & Harvey) Lyle
      Antithamnionella tormentosa Stegenga
      Antithamnionella verticillata (Suhr) Lyle
      Antithamnionella sp.
      Bornetia repens Stegenga
      Callithamniella capensis  Simons
      Callithamniella tingitana (Schousboe ex Bornet) Feldmann-Mazoyer
      Carpoblepharis flaccida (J.V.Lamouroux) Kützing
      Centroceras spp.
      Ceramium aff. callipterum Mazoyer
      Ceramium arenarium Simons
      Ceramium camoui Dawson E.Y.Dawson
      Ceramium centroceratiforme Simons
      Ceramium glanduliferum Kylin
      Ceramium graecum/lenticulare complex
      Ceramium juliae A.J.K.Millar
      Ceramium obsoletum C. Agardh
      Ceramium planum Kützing
      Ceramium poeppigianum Grunow
      Ceramium tenerrimum (G. Martens) Okamura
      Ceramium sp. 1
      Ceramium sp. 2
      Ceramium sp. 3
      Compsothamnionella sciadophila Stegenga
      Episporium centroceratis Möbius
      Gayliella dawsonii (= Ceramium dawsonii A.B. Joly)
      Gayliella papenfussiana (= Ceramium papenfussianum Simons)
      Gayliella transversalis (Collins et Hervey) T.O. Cho et Fredericq
      Laurenciophila minima Stegenga
      Microcladia exserta  M.J.Wynne
      Microcladia gelidii R.H.Simons
      Perischelia glomulifera (J.Agardh) J.Agardh ex Kylin
      Pterothamnion recurvatum (Wollaston) Athanasiadis & Kraft
      Pterothamnion sp.
      Ptilothamnion codicola (E.Y.Dawson) I.A. Abbott
      Ptilothamnion goukammae Stegenga, Bolton & R.J. Anderson
      Ptilothamnion polysporum Gordon-Mills & Wollaston in Wollaston
      Scageliopsis tsitsikammae Stegenga, R.J. Anderson & J.J. Bolton
      Syringocolax macroblepharis Reinsch
      Zonariophila semiendophytica Stegenga et Kemperman

FAMILY DASYACEAE
      Dasya echinata Stegenga, Bolton & Anderson
      Dasya scoparia Harvey ex J. Agardh
      Dasya sp. 1
      Dasya sp. 2
      Dasya sp. 3
      Dipterocladia pinnatifolia (Suhr) Y.S.M. De Jong
      Heterosiphonia arenaria Kylin
      Heterosiphonia crispa (Suhr) Falkenberg
      Heterosiphonia dubia (Suhr) Falkenberg

FAMILY DELESSERIACEAE
      Acrosorium acrospermum (J. Agardh) Kylin
      Acrosorium ciliolatum (Harvey) Kylin
      Acrosorium maculatum (Sonder ex Kützing) Papenfuss
      Apoglossum ruscifolium (Turner) J. Agardh
      Apoglossum spathulatum (Sonder) Womersley and Shepley
      Bartoniella crenata Kylin
      Caloglossa leprieurii (Montagne) G. Martens
      Gonimophyllum africanum Martin and Pocock
      Holmesia capensis J Agardh
      Hypoglossum imperfectum Stegenga, Anderson and Bolton
      Martensia elegans Hering
      Neoholmesia natalensis M. J. Wynne
      Nienburgia pinnatifida (Suhr) Papenfuss
      Nienburgia serrata (Suhr) Papenfuss
      Papenfussia laciniata (Harvey) Kylin
      Papenfussia minuta Kylin
      Platysiphonia delicata (Clemente y Rubio) Cremades
      Taenioma perpusillum (J. Agardh) J. Agardh
      Valeriemaya seagriefii De Clerck, Wynne and Stegenga

FAMILY INKYULEEACEAE
      Inkyuleea beckeri (F.Schmitz ex Mazza) H.-G.Choi, Kraft & G.W.Saunders

BOOK 6

Order Ceramiales (in part)

FAMILY RHODOMELACEAE
      Acanthophora spicifera (M.Vahl) Børgesen
      Aiolocolax pulchellus M.A.Pocock
      Aphanocladia ecorticata Stegenga, R.J.Anderson & Bolton
      Bostrychia harveyi Montagne
      Bostrychia intricata (Bory de Saint-Vincent) Montagne
      Bostrychia moritziana (Sonder ex Kützing) J.Agardh
      Bostrychia radicans (Montagne) Montagne
      Bostrychia simpliciuscula Harvey ex J.Agardh
      Bostrychia tangatensis Post
      Bostrychia tenella (Lamouroux) J. Agardh
      Chondria armata (Kützing) Okamura
      Chondria capensis (Harvey) Askenasy
      Chondria dasyphylla (Woodward) C.Agardh
      Chondria sp.
      Colacopsis velutina (M.T.Martin & M.A.Pocock) R.E.Norris
      Dasyclonium incisum (J.Agardh) Kylin
      Dasyclonium palmatifidum (Grunow) Scagel
      Dictyomenia stephensonii Papenfuss
      Hawaiia sp.
      Herposiphonia clavata M.J. Wynne
      Herposiphonia didymosporangia Stegenga and Kemperman
      Herposiphonia falcata
      Herposiphonia prorepens (Harvey) F. Schmitz
      Herposiphonia secunda (C.Agardh) Ambronn
      Herposiphonia secunda f. tenella (C.Agardh) M.J.Wynne
      Janczewskia meridionalis M.T.Martin & M.A.Pocock
      Laurencia alfredensis Francis, Bolton, Mattio & R.J. Anderson
      Laurencia complanata (Suhr) Kützing
      Laurencia cf corymbosa J.Agardh
      Laurencia digitata Francis, Bolton, Mattio & R.J. Anderson
      Laurencia flexuosa (Suhr) Kützing
      Laurencia glomerata (Kützing) Kützing
      Laurencia griseaviolacea M.J. Wynne
      Laurencia multiclavata Francis, Bolton, Mattio & R.J. Anderson
      Laurencia natalensis Kylin
      Laurencia pumila (Grunow) Papenfuss
      Laurencia pumila var. dehoopiensis Francis, Bolton, Mattio & R.J. Anderson
      Lophosiphonia capensis (Kylin) R.E.Norris
      Lophosiponia reptabunda (Suhr) Kylin
      Microcolax africanus M.T.Martin & M.A.Pocock
      Onychocolax polysiphoniae M.A. Pocock
      Ophidocladus simpliciusculus (P.L.Crouan & H.M.Crouan) Falkenberg in Schmitz & Falkenberg
      Osmundaria serrata (Suhr) R.E.Norris
      Pachychaeta brachyarthra (Kützing) Trevisan
      Pachychaeta cryptoclada Falkenberg
      Placophora binderi (J.Agardh) J.Agardh
      Placophora monocarpa (Montagne) Papenfuss
      Kentrophora natalensis (J.Agardh) S.M.Wilson & Kraft in R.J.F. Henderson, S.M. Wilson & Kraft
      Neosiphonia sertularioides (Grateloup) K.W. Nam & P.J. Kang
      Polysiphonia foetidissima Cocks ex Bornet
      Polysiphonia incompta Harvey
      Polysiphonia kowiensis Stegenga, J.J. Bolton & R.J. Anderson
      Polysiphonia namibiensis Stegenga et Engledow in Stegenga, Bolton et Anderson
      Polysiphonia scopulorum Harvey
      Polysiphonia sp. 1
      Polysiphonia sp. 2
      Polysiphonia sp. 3 (note only – see under P. incompta)
      Polysiphonia sp. 4
      Polysiphonia subtilissima  Montagne (note only – see under genus description)
      Polyzonia elegans Suhr
      Pterosiphonia cloiophylla (C.Agardh) Falkenberg in Schmitz & Falkenberg
      Pterosiphonia spinifera (Kützing) Ardré
      Pterosiphonia stangerii (J.Agardh) Falkenberg
      Pterosiphonia stegengae Savoie & G.W.Saunders
      Pterosiphonia sp.
      Rhodomelopsis africana M.A.Pocock
      Streblocladia corymbifera (C.Agardh) Kylin
      Streblocladia tenuissima M.A.Pocock
      Streblocladia sp.
      Symphyocladia marchantioides (Harvey) Falkenberg in Schmitz & Falkenberg
      Tayloriella tenebrosa (Harvey) Kylin

FAMILY SPYRIDIACEAE
      Spyridia cupressina Kützing
      Spyridia filamentosa (Wulfen) Harvey in Hooker
      Spyridia horridula F. Schmitz ex J. Agardh
      Spyridia hypnoides (Bory) Papenfuss
      Spyridia plumosa F. Schmitz ex J. Agardh

FAMILY WRANGELIACEAE
      Anotrichium furcellatum (J. Agardh) Baldock
      Anotrichium tenue (C. Agardh) Nägeli
      Dasyphila cryptocarpa (Holmes) F. Schmitz
      Griffithsia confervoides Suhr
      Griffithsia subbiconica Stegenga
      Griffithsia sp.
      Gymnothamnion elegans (Schousboe ex C. Agardh) J. Agardh
      Gymnothamnion elegans var. bisporum Stegenga
      Haloplegma duperreyi Montagne
      Pleonosporium callicladum R.E. Norris
      Pleonosporium filicinum (Harvey ex J. Agardh) De Toni
      Pleonosporium harveyanum (J. Agardh) De Toni
      Shepleya elixithamnia Gordon-Mills & R.E. Norris
      Spermothamnieae indet.
      Spongoclonium caribaeum (Børgesen) M.J.Wynne
      Stegengaea capensis (Stegenga) Alongi, Cormaci & G.Furnari
      Tiffaniella cymodoceae (Børgesen) E.M.Gordon
      Tiffaniella schmitziana (Barton) Bolton et Stegenga
      Vickersia baccata (J. Agardh) Karsakoff
      Wrangelia purpurifera (Harvey) J. Agardh

Sub Class Corallinophycidae

Order Corallinales

FAMILY CORALLINACEAE

Crustose corallines (non-geniculate) species: these are not covered here. See Maneveldt, Chamberlain & Keats 2008. A catalogue with keys to the non-geniculate coralline algae (Corallinales, Rhodophyta) of South Africa. South African Journal of Botany 74(4):555-566.

SUB-FAMILY CORALLINOIDEAE (articulated, geniculate taxa)
      Amphiroa anceps (Lamarck) Decaisne
      Amphiroa beauvoisii  Lamouroux
      Amphiroa bowerbankii Harvey
      Amphiroa capensis Areschoug in J. Agardh
      Amphiroa ephedraea (Lamarck) Decaisne
      Amphiroa rigida J.V. Lamouroux
      Arthrocardia carinata (Kützing) Johansen
      Arthrocardia corymbosa (Lamarck) Decaisne
      Arthrocardia duthieae Johansen
      Arthrocardia flabellata (Kützing) Manza
      Arthrocardia palmata (J. Ellis & Solander) J.E. Areschoug in J. Agardh
      Corallina berteroi Montagne ex Kützing
      Corallina sp. 1
      Jania adhaerens Lamouroux
      Jania capillacea Harvey
      Jania cultrata (Harvey) J. H. Kim, Guiry & H.-G. Choi
      Jania prolifera (J.V.Lamouroux) J.H.Kim, Guiry & H.-G.Choi
      Jania sagittata (J.V.Lamouroux) Blainville
      Jania subulata (Ellis & Solander) Sonder
      Jania verrucosa Lamouroux

SUB-FAMILY MASTOPHOROIDEAE
      Metamastophora flabellata (Sonder) Setchell

Order Hapalidales

FAMILY HAPALIDACAE
      Choreonema thuretii (Bornet) F.Schmitz

Order Gigartinales

Family Kallymeniaceae

Callophyllis variegata (Bory de Saint-Vincent) Kützing 1843: 401

Plants bright to dark red, to about 20 cm tall, with small hapterous holdfast. Blade flat, soft cartilaginous, laciniate, up to about 400 µm thick. Branching in one plane, irregular, alternate or somewhat dichotomous, often with marginal proliferations. Medulla of large colourless cells interspersed with smaller-celled filaments; cortex of 1-2 layers of small, pigmented cells. Tetrasporangia not seen in our specimens. Cystocarps about 1 mm in diameter, scattered near margins of blades and on marginal proliferations.

Collections, ecology and regional distribution

Recorded from a few scattered localities between Saldanha Bay and Waterloo Bay (13-40), apparently subtidal and rare, but may be a little more common in the Agulhas – Arniston area.

World distribution: Alaska, southern South America, New Zealand, some sub-Antarctic islands, Antarctica (MD Guiry in Guiry & Guiry 2015).

Syntype localities: New Guinea and Concepción, Chile (Silva et al. 1996).

Note: Norris (1964) first identified South African specimens as C. variegata, and suggested that the species may be rare here as it might be a “recent arrival’’. South African species are described in detail by Norris (1964), and a fundamental study of the morphology and anatomy of the Kallymeniaceae is that of Norris (1957). Although recorded from a few localities on the west coast, this species was not described or illustrated by Stegenga et al. (1997).

Callophyllis variegata (herbarium specimen).

Order Gigartinales

Family Kallymeniaceae

Pugetia Kylin

Plants with small discoid holdfasts, short stipes and slippery, membranous red fronds with smooth to ruffled margins. Fronds either orbicular or broadly foliose, irregularly lobed or broadly cuneate and somewhat irregularly incised. Medulla of large colourless cells interspersed with pigmented filaments; cortex of several layers of small pigmented cells. See Norris (1964) for a treatment of South African Pugetia.

Key to species

1a. Blades neatly orbicular to ovate

P. harveyana

1b. Blades cuneate to broadly lobed, may be incised

P. porphyroidea

Order Gigartinales

Family Kallymeniaceae

Pugetia harveyana (J. Agardh) R.E. Norris 1964: 119

Plants foliaceous, membranous, slippery, bright red to orange-red, with small holdfast, short stipe (1-4 mm) and broad rounded or lobed blade up to about 15 cm in diameter, with smooth, often ruffled margin. Blade up to 700 µm thick. Medulla of 1-2 layers of large cells loosely interwoven with branched filaments of smaller cells. Cortex of several layers of small cells showing no anticlinal cell rows. Tetrasporangia scattered over thallus, immersed in cortex, 25-30 x 12-15 µm, cruciately divided. Cystocarps large, immersed and visible as dark red circular (to 2 mm diameter) patches, not causing notable swelling.

Collections, ecology and regional distribution

Recorded from Namibia to Tsitsikamma (1-32). Usually epiphytic on other seaweeds (e.g. Gigartina, Trematocarpus), subtidal.

World distribution: Also recorded from Korea (Lee 2008 in Guiry & Guiry 2010).

Type locality: Unknown. Agardh Herb No 24582 Kallymenia harveyana (Norris 1964). bears no information on locality.

Note: South coast specimens of P. harveyana that we have seen do not appear to reach the size of those on the west coast (to 30 cm).

Pugetia harveyana, Tsitsikamma, on Gelidium capense (BOL).

Pugetia harveyana, large west coast specimens from Table Bay (BOL).

Pugetia harveyana, west coast specimens. 1-3. Habit, 2 and 3 with carposporophytes.
Reproduced from Stegenga et al. (1997).

Pugetia harveyana. 1. Cross section. 2. Corss section of tetrasporangial thallus.
Reproduced from Stegenga et al. (1997).

Order Gigartinales

Family Kallymeniaceae

Pugetia porphyroidea (F. Schmitz ex Holmes) R.E. Norris 1964: 113, 115-119, figs 37-45, pl. 7,8

Plants foliaceous, membranous, slippery, bright red, with small membranous holdfast, short stipe (to 4 mm), and blade up to 25 cm long that varies in shape from cuneate to broadly rounded or with several shallow lobes. Margin smooth, often ruffled. Medulla of large cells interspersed with branched filaments of smaller cells. Cortex of 1-2 layers of deeply pigmented, small rounded cells. Tetrasporangia scattered over blade, immersed in cortex, small (19x12 µm), cruciately divided. Cystocarps swollen, up to 1 mm diameter, sometimes distributed randomly over thallus, but often in small groups.

Collections, ecology and regional distribution

Usually epiphytic on other seaweeds, subtidal. Although recorded from the west coast by Norris (1964), Stegenga et al. (1997) noted that they had not seen west coast material and were unsure of its relationship with the south coast entity. Distribution therefore uncertain: either Yzerfontein on the west coast to the Port Alfred area (14-39), or in a restricted area around Port Alfred (39).

World distribution: A South African endemic.

Type Locality: “Cape of Good Hope” (Silva et al. 1996).

Note: P. porphyroidea reaches its largest size on the south coast; west coast specimens (if they are the same species) seem to be smaller than indicated above.

Pugetia porphyroidea, Port Alfred (BOL).

Pugetia porphyroidea, XS of mature blade (re-drawn from Norris 1964) Fig 41.

Order Gigartinales

Family Kallymeniaceae

Thamnophyllis pocockiae R.E. Norris 1964: 105-110, figs 20-26; pls 4,5

Plants foliaceous, membranous, deep red, with a small holdfast and short stipe. Blade expanding sharply from stipe, with several deeply-incised lobes, up to 20 cm long and 40 cm wide. Margin usually dentate or fimbriate, with untidy proliferations and short irregular branches at apices. Medulla of central layer of longitudinally-arranged, smallish cells, between two layers of large rounded cells; stellate cells present in medulla of mature tissues. Outer cortex of 1-2 layers of small round to ovoid cells. Tetrasporangia scattered over thallus, about 18x27 µm, irregularly divided. Cystocarps scattered over thallus, up to 1 mm diameter, bulging more towards one side than the other, ostiolate.

Collections ecology and regional distribution

Recorded from St Helena Bay on west coast to Kei Mouth area (11-43). Subtidal, usually epiphytic on other seaweeds.

World distribution: South African endemic.

Type locality: St Helena Bay, Western Cape (Silva et al. 1996).

Thamnophyllis pocockiae, Tsitsikamma.

Thamnophyllis pocockiae, detail of thallus margins.

Thamnophyllis pocockiae, cross section of thallus.

Thamnophyllis pocockiae, stellate cell detail.

Thamnophyllis pocockiae, carpogonial system.

Thamnophyllis pocockiae, cross section, cortex.

Thamnophyllis pocockiae, cross section through mature cystocarp.

Thamnophyllis pocockiae, 1. Habit. 2. Cross section with tetrasporangia and stellate medullary cell. Reproduced from Stegenga et al. (1997).

Order Gigartinales

Family Gymnophloeaceae

Schizymenia apoda (J. Agardh) J. Agardh 1851: 175

Thallus dull red, flaccid and slippery. Blades foliose, simple, up to 40 cm long, broadly lanceolate, surface slightly rugose, often with marginal proliferations or lacerated at distal end. Blades up to 600 µm thick; medulla of densely interwoven filaments; cortex of anticlinal bundles of filaments interspersed with abundant gland cells up to 70 µm long and 20 µm wide; surface cortical cells 3-6 µm in diameter. Carposporophytes immersed in thallus, with ostiole; tetrasporangia unknown.

Collections, ecology and regional distribution

A subtidal species, common on the west coast but found as far east as the Kei Mouth area (1-43).

World distribution: Also known from Tristan da Cunha (as S. obovata), St. Paul Island, Namibia, Somalia (Silva et al. 1996).

Type locality: Table Bay, Western Cape, South Africa (Silva et al. 1996).

Note: Stegenga et al. (1997) point out that this species (as S. obovata) may have a crustose tetrasporophyte, as is suggested for S. dubyi (Chauvin ex Duby) J. Agardh (Ardré 1980).

Schizymenia apoda, De Hoop.

Schizymenia apoda, xs through cortex and outer medulla.

Schizymenia apoda. 1. Habit. 2-3. Cross sections (note darker gland cells). 4. Carposporophyte in cross section of thallus. Reproduced from Stegenga et al. (1997)(as S. obovata).

Order Gigartinales

Family Phacelocarpaceae

Genus Phacelocarpus Endlicher & Diesing

Plants erect, with serrulate axes, discoid holdfast and short stipe. Thalli armed with teeth formed from short terminal branchlets. Thalli uniaxial; large central filament and medulla of rhizoidal filaments; cortex of isodiametric cells becoming smaller towards periphery. Cystocarps stalked, with multicellular pericarps and transverse (slit-like) ostioles; tetrasporangia zonately divided and formed in nemathecia.

Key to species

1a. Teeth always in spirals, thalli always terete

P. oligacanthus

1b. Teeth formed in two ranks; either along margins (flat branches) or
in spiral (terete branches)

P. tortuosus

Order Gigartinales

Family Phacelocarpaceae

Phacelocarpus oligacanthus Kützing 1868: 25, pl. 71, a,b

Thalli dark red, erect, up to 25 cm or longer, attached by discoid holdfasts. Axes terete, 2-4 mm in diameter (wider at base in old plants), bearing many short lateral branches that arise irregularly. Thallus serrate, with many short branchlets/spines arranged in regular radial pattern; cortication increasing with age of thallus or part of thallus, so that basal parts of thalli or older thalli may lack spines and have smooth but uneven texture; branch apices crowded with spines. Structure of branches uniaxial, in cross-section axial cell distinct and much larger than surrounding medullary cells; cortex of anticlinal rows of cells decreasing very noticeably in diameter towards periphery, outer cortical cells very small (2-3 µm diameter) and darkly pigmented. Dioecious; reproductive structures formed on spiny lateral branchlets. Tetrasporangia zonately divided, arranged around the sides of deep, open channels in the branchlets. Mature cystocarpic branchlets resemble swollen vegetative branchlets with well-developed teeth; carposporangia released though slits. Spermatangia borne in short, spiny, lateral branchlets.

Collections, ecology and regional distribution

Recorded from Cape Agulhas to near Umkomaas on the KwaZulu-Natal coast (22-50), in the lower eulittoral and sublitoral zones (De Clerck et al. 2005).

World distribution: South African endemic.

Type locality: “Cap Bona Spei” (Silva et al. 1996).

Note: For details of morphology and reproductive structure see Searles (1968).

Phacelocarpus oligacanthus, Mkambati (BOL).

Phacelocarpus oligacanthus, Mkambati , detail of spiny branchlets (herbarium specimen, BOL).

Order Gigartinales

Family Phacelocarpaceae

Phacelocarpus tortuosus Endlicher & Diesing 1845: 290.

Thalli dark red, erect, tough, densely branched, up to 30 cm tall; attached by short stipe to discoid holdfast. Axes either flattened, with teeth along margins, or terete, with teeth in spirals around axes; sometimes both types on one plant. Thallus uniaxial; axial cell distinct and surrounded by much smaller medullary cells; cortical cells large, becoming progressively smaller towards periphery. Reproductive structures borne on lateral branchlets; globose tetrasporangial nemathecia borne in or near the sinus between adjacent teeth, zonately divided tetrasporangia arranged in channels below surface of nemathecium; mature cystocarps stalked, globose, bilobed, opening via a slit.

Collections, ecology and regional distribution

Found from De Hoop to the Durban area of KZN (24-51), in the lower eulittoral and shallow sublittoral zones.

World distribution: South African endemic.

Type locality: Port Natal (Durban) South Africa (Silva et al., 1996).

Note: For details of morphology and reproductive structure see Searles (1968).

Phacelocarpus tortuosus, Kenton-on-Sea.

Phacelocarpus tortuosus, detail of spiny branchlets. Fresh specimen, Port St Johns.

Phacelocarpus tortuosus, detail of herbarium specimen (BOL) from Port Alfred.

Order Gigartinales

Family Solieriaceae

Callophycus Trevisan

Thallus erect, branching complanate and pinnate, branches compressed to flat, with or without midrib, margin entire to serrate, holdfast discoid. Structure of loose to fairly dense medulla of longituduinal and rhizoidal filaments clearly demarcated from psudoparenchymatous cortex. Isomorphic gametophytes and tetrasporophytes. Cystocarps immersed in pinnules, ostiolate. Tetrasporangia zonately divided, scattered in outer cortex.

Key to species

Regularly serrate branch margins, branches not tridentate

C. densus

Non-serrate branch margins, branches tridentate (3-pronged)

C. tridentifer

Order Gigartinales

Family Solieriaceae

Callophycus densus (Sonder) Kraft 1984: 58

Thalli red (darkening on drying), about 5 – 20 (30) cm tall, compressed, pinnately branched, margins regularly serrated. Discoid holdfast gives rise to single main axis with prominent midrib. Axes narrow at base, increasing to 2-3 mm width, about 0.5 mm thick. Branching opposite, sub-opposite or alternate. Structure of thallus multiaxial. Medulla of two types of longitudinal filaments: most are thin with 2-3 µm diameter cell lumen, others with cell lumen of about 6 µm diameter and thick walls giving a total cell diameter of about 20 µm. Cortex pseudoparenchymatous, comprising two layers, the inner comprising two layers of large cells (up to 50 µm diameter), the outer a 2-3 layered outer cortex of palisade-like cells 3-5 µm in diameter. Cystocarps scattered singly or in small groups on all parts of thallus, visible as swellings, ostiolate. Tetrasporangia scattered irregularly over blades, immersed in outer cortex, zonately divided, 13-19 µm long, 5-12 µm wide.

Collections, ecology and regional distribution

Subtidal, from Olifantsbos (just west of Cape Point) in drift to southern KwaZulu-Natal (17-57).

World distribution: Also recorded from Australia and Fiji (Guiry & Guiry 2011).

Type locality: Cape York, Queensland, Australia (Kraft, 1984).

Note: Detailed descriptions of this species are provided by Hewitt (1960) and Kraft (1984).

Callophycus densus, Kini Bay, near East London (BOL).

Callophycus densus, specimen from west coast of Cape Peninsula (BOL).

Callophycus densus. 1. Habit. 2. Longitudinal section. 3. Cross section. Reproduced from Stegenga et al. (1997).

Order Gigartinales

Family Solieriaceae

Callophycus tridentifer Kraft 1984: 61,65,67, figs 33-47, 52.

Thallus brownish- to pinkish-red (darkening on drying), erect, up to 20 cm tall, cartilaginous. Discoid holdfast gives rise to a single primary axis bearing up to 4 orders of compound, more-or-less opposite pinnae; fronds slightly compressed, 1-3 mm wide, pinnae and pinnules arising at angles of 30-45o, crowded, apices tridentate (3-pronged). In cross-section of lower axis, medulla 500-600 µm wide, comprising inner zone of sparse rhizoids surrounded by layer of dense rhizoids. Inner cortex 1-2 cells deep, cells isodiametric; outer cortex of 2-3 layers of radially elongate cells, surface cells 7-12 µm long, 5-8 µm wide. Mature cystocarps large, embedded in branches, ostiolate. Tetrasporangia embedded in outer cortex, zonately divided.

Collections, ecology and regional distribution

Found from Port St Johns area to northern KZN, subtidal (46-58).

World distribution: First recorded from Australia (Kraft 1984) and later from South Africa by Norris and Aken (1985).

Type locality: Coffs harbour, NSW, Australia (Kraft 1984).

Note: Detailed description of this species in Kraft (1984); comparisons with South African specimens provided by Norris &Aken (1985).

Callophycus tridentifer, specimen from Trafalgar, southern KwaZulu-Natal (BOL).

Callophycus tridentifer, detail of branching.

Order Gigartinales

Family Solieriaceae

Rhabdonia natalensis Reinbold 1912: 199-200

Thalli red, terete, a few to 30 cm tall, rather lax. Axes slender and up to about 1 mm diameter, but up to 2 mm in lower axes; irregularly dichotomously branched, with up to 10 bifurcations per axis. Axis apices usually gradually pointed but may be rounded in (older?) specimens. Cystocarps ostiolate, forming as distinct globose swellings (about 1.5 mm diameter) just below the apices of terminal branchlets. In many specimens the branch tips remain visible on top of the cystocarps as tiny spines. Tetrasporangia not seen in BOL material.

Collections, ecology and regional distribution

Rather rare, having being found only near Port Alfred and at Cape Morgan (39-43), and apparently only in drift, indicating a subtidal habitat.

World distribution: also recorded from Mauritius (Børgesen 1943).

Lectotype locality: Kowie (Port Alfred), recorded by Papenfuss (1968): a note on Specimen 035362 in BOL, by Papenfuss, denotes it as “type”.

Note: We have only examined pressed herbarium specimens (BOL). R. natalensis has not been recorded in KZN. Norris points out that the epithet natalensis is probably based on Reinbold’s misconception about the location of the collecting site.

Rhabdonia natalensis, Kowie specimen (BOL).

Rhabdonia natalensis, Kowie specimen (BOL), showing apical branches and terminal cystocarps.

Order Gigartinales

Family Areschougiaceae

Erythroclonium corallinum (Suhr) Holmes 1896: 350-351

Thalli red, forming branched chains up to about 7 cm tall, of ellipsoid segments each up to about 3mm long. Branches comprise either wider segments (up to about 1mm; most upright branches) or narrower segments (about 300 um; all prostrate and some upright branches). Branches with narrow segments may develop small discoid holdfasts that attach to any other substrata; branches with broader segments do not appear to do this. Discoid attachments usually form at nodes on narrow branches. Segments often narrower toward distal ends of erect branches and often bear small distal segments with short projections containing axial cells. Nodal regions cylindrical and about 3 cells deep. Branching of axes is sparse and occurs at nodes, usually a single (but up to three) branches being produced. Structure uniaxial, most axial cells bearing two (opposite) branches. Medullary filaments branching 4-5 times in middle of segments and ending in cortical cells up to 25 µm long. Only tetrasporic material seen. Tetrasporangia evenly distributed in cortex, zonately divided; up to 50 x 35 µm when mature.

Collections, ecology and regional distribution

Found from Woody Cape northwards to at least Park Rynie, KZN ((38-55))> Rather rare; intertidal to at least 30 m depth.

World distribution: South African endemic.

Type locality: Kowie (Port Alfred) (Norris, 1988).

Note: Norris (1988) describes this species in detail, and points out that it is the only species of that genus to occur outside Australia, where 4 other species are known to occur.

Erythroclonium corallinum, herbarium specimen from Kowie (BOL).

Erythroclonium corallinum, detail of herbarium specimen from Kowie (BOL).

Order Gigartinales

Family Caulacanthaceae

Catenella nipae Zanardini 1872: 143-145, pl. 6: A, figs 1-7

Thallus brown-red to maroon, up to 1 cm tall, growing in clumps. Branching di-or tri-chotomous, segments 2-4 mm long, swollen conspicuously and constricted at nodes. Branch segments arising from nodes. Holdfasts developing from nodes, cylindrical, with discoid attachments. Branches uniaxial, hollow, filled with mucilage; outer cortical cells elongate, pigmented; inner cortical cells larger, spherical; system of dichotomous filaments connecting inner cortical cells to laxly-arranged medullary filaments orientated transversely to main axial filament. No fertile plants noted in South African material by Lambert et al (1987), but reports from other areas describe swollen, stipitate cystocarps and immersed tetrasporangia scattered in cortical tissue.

Collections, ecology and regional distribution

Port St Johns area (Pondoland) – Mngazana and Mntafufu estuaries eastward to northern KZN (46-58). Found on mangrove substrata at or below water level (Lambert et al., 1987).

World distribution: Also recorded from Tanzania and many localities in the tropical Indo-Pacific, including India, Japan, Indonesia, Australia and New Zealand (Guiry & Guiry 2011).

Type locality: Sarawak, Indonesia (Lambert et al., 1987).

Note: see Lambert et al., 1987 for further details of South African material.

Catenella nipae, showing swollen vertical branches and holdfasts (H). Drawn from photograph in Lambert et al. (1987: Fig. 3 H).

Order Gigartinales

Family Caulacanthaceae

Caulacanthus ustulatus (Turner) Kützing 1843: 395

Thalli forming dense, tangled, brown clumps or turfy patches 1-3 cm high and usually up to several cm in diameter. Axes about 1 mm diameter, frequently and irregularly branched, apices spiny. Axes often secondarily attached to substrata or other axes by pads formed from outgrowths of cortical tissue. Structure uniaxial, each cell of central filament bearing two pericentral cells at more-or-less right angles; these divide dichotomously to produce a filamentous medulla and cortex and a compact epidermis of small (4-6 µm diameter) cells. Tetrasporangia (70 x 25 µm) zonately divided and scattered in cortex. Carposporophytes develop sub-terminally in branch tips; mature cystocarps form globose swellings with latero-apical ostiole.

Collections, ecology and regional distribution

Found around the whole SA coast (1-58). Common in the mid and lower eulittoral, forming brown, mossy clumps; often associated with mussels.

World distribution: Very widespread in temperate, subtropical and tropical waters.

Type locality: Lectotype locality is Cádiz, Spain (Silva et al. 1996).

Note: Searles (1968) details the morphology and reproduction of this species.

Caulacanthus ustulatus, Keurboomstrand, forming typical cushions in upper eulittoral zone.

Caulacanthus ustulatus, detail of branches.

Caulacanthus ustulatus, xs showing anatomy including zonately-divided tetrasporangia (Nature’s Valley material).

Caulacanthus ustulatus. 1. Habit (detail of plant). 2. Longitudinal section near thallus apex. 3. Cross sections (ca = central filament). 4. Tetrasporangia. Reproduced from Stegenga et al. (1997).

Order Gigartinales

Family Caulacanthaceae

Heringia mirabilis (C. Agardh) J. Agardh 1842: 68

Plants erect, bushy, up to 30 cm tall, dark red. Often associated with sponges. Holdfast crustose to discoid, with many erect axes; axes terete to slightly compressed, cartilaginous and wiry, 250 -500 µm in diameter. Main axis indistinct, dichotomies regular, profuse terminal branches. Structure uniaxial; in cross-section central filament distinct, surrounded by parenchymatous medulla of largish cells (12-20 µm diameter), and cortex of small-celled radial filaments imparting almost striate appearance. Epidermal cells 4-8 µm in diameter. Tetrasporangia in raised nemathecia near branch tips, zonately divided and about 30-40 µm long. Cystocarps just below thallus apices, globose, with laterally-displaced ostiole.

Collections, ecology and regional distribution

Recorded from Namibia and around almost the whole coast of South Africa, to near Cape Vidal, KZN. (1-56). Found in the sublittoral fringe and subtidally. On the west coast H. mirabilis is usually epiphytic on Cladophora mirabilis, but on the south and east coasts it is either attached to one of a variety of seaweeds and animals (especially sponges) or is epilithic.

World distribution : southern African endemic.

Type locality: “Cape of Good Hope”, South Africa (Silva at al. 1996).

Note: A detailed description of morphology and development in H. mirabilis is given by Searles (1968).

Heringia mirabilis, De Hoop. Whitish strands of bacterial filaments seen on this specimen.

Heringia mirabilis, detail of branches under dissecting microscope.

Heringia mirabilis, Three Sisters (BOL).

Heringia mirabilis, Kowie (BOL), showing branching pattern.

Heringia mirabilis, habit. Reproduced from Stegenga et al. (1997).

Order Gigartinales

Family Cystocloniaceae

Calliblepharis fimbriata (Greville) Kützing 1843: 404

Thalli complanate, strap-like, up to 25 (30) cm long, flaccid, bright to dark red, branched, markedly frilly. Holdfast bearing short stipe that widens into main axis. Branching opposite, sub-opposite or irregular. Main axis and main branches widest in the middle (usually 8-15 (-20) mm wide), tapering towards both ends, with marginal proliferations or teeth. Structure uniaxial, in cross-section with large un-pigmented medullary cells surrounded by smaller cortical cells. Dioecious, cystocarps in smaller branchlets, protruberant, ostiole absent. Tetrasporangia zonately divided, scattered in outer cortical tissue. This rather pretty species is quite distinctive on account of its soft, almost delicate texture and frilly margins.

Collections, ecology and regional distribution

Recorded from Arniston eastwards as far as the Port Shepstone area in southern KwaZulu-Natal (23-49). A subtidal species, found down to at least 20 m depth.

World distribution: also recorded from Oman and Pakistan (Silva et al. 1996).

Type locality: “Cape” (Kützing 1843).

Calliblepharis fimbriata, Arniston.

Calliblepharis fimbriata, Arniston.

Calliblepharis fimbriata, herbarium specimen (BOL).

Order Gigartinales

Family Cystocloniaceae

Hypnea Lamouroux

Plants bushy or caespitose; thalli terete or slightly compressed, either attached basally or with hooked branches or apices and entangled in other algae. Apices uniaxial, structure pseudoparenchymatous throughout, central axis obscure except in young axes. Generations isomorphic. Tetrasporangia zonately divided and arranged in nemathecia or sori on ultimate or penultimate branches (stichidia). Cystocarps in ultimate or penultimate branches, protruberant, ostiolate or non-ostiolate, comprising central sterile tissue of large cells with radiating gonomiblasts ending in single carposporangia. Hypnea currently includes 60 species worldwide (Guiry & Guiry 2015), of which seven are recorded in our area. We note that some species are very difficult to tell apart, and often differ only in size: this genus in South Africa is the subject of a current molecular and morphological study.

Key to south coast species (provisional; March 2015)

1a. Thallus of stiff uprights, 1-2 mm in diameter, green to dark purple

H. spicifera

1b. Thallus 1 mm or less in diameter

2

2a. Thallus ca. 180 µm in diameter, less than 5 mm tall

H. arenaria

2b. Thallus 200 µm to 1 mm in diameter, a few to many cm tall

3

3a. Thallus iridescent blue in situ, without hooked tips

H. viridis

3b.Thallus pink, red or brown-yellow, tips sometimes or often hooked

4

4a. Main axes 200-400 µm in diameter, branching mostly second

H. tenuis

4b. Main axes between 400 µm and 1 mm in diameter, branching irregular

5

5a. Thallus brownish-red to straw coloured

H. musciformis

5b.Thallus pink to bright red

6

6a. Main axes about 1 mm in diameter

H. rosea

6b. Main axes about 0.5 mm in diameter

H. intricata

Order Gigartinales

Family Cystocloniaceae

Hypnea arenaria Kylin 1938: 12, fig. 6

Thallus developing creeping axes with erect parts, 3-5 mm high, simple or sparsely branched; axes and branches about 180 µm thick; tetrasporangia in the slightly swollen tips of erect branches. Creeping among fine sand, with tetraspores found in July. (Description as in Kylin, 1938).

Collections, ecology and regional distribution

Recorded from East London area to Kosi Bay, northern KwaZulu-Natal (41-52).

World distribution: South African endemic.

Type locality: Inyone Rocks, near Durban (Silva et al., 1996).

Note: BOL has only three specimens under this name, but they do not fit the type description.

Hypnea arenaria, re-drawn from Kylin (1938).

Order Gigartinales

Family Cystocloniaceae

Hypnea intricata Kylin 1938: 11, figs 4B, C.

“Thallus 2-3 cm tall, rather irregularly branched with long and short axes mixed together; longer axes with hooked apices; terminal axes 0.5 mm thick.” This very brief description is from Kylin (1938).

Collections, ecology and regional distribution

Apparently recorded from Tsitsikamma to the Durban area (32-51). We have only applied the name to a specimen from Addo, and then with some doubt.

Type locality: Port Elizabeth (Kylin 1938).

Note: This species appears to be distinguished from some others (some specimens of H. tenuis, H. rosea, H. musciformis) almost entirely on size, especially axis width. The result is that some specimens cannot clearly be ascribed to any one of these species. 

Hypnea intricata (BOL), smallest scale divisions 1 mm.

Hypnea intricata, re-drawn from Kylin (1938).

Order Gigartinales

Family Cystocloniaceae

Hypnea musciformis (Wulfen) J. V. Lamouroux 1813: 43

Plants reddish brown, a few to about 15 cm long, more or less caespitose, usually entangled with other algae in turfs. No percurrent main axis, major branches irregularly ramified; branch-ends often curved into hooks; apices often with short spinous branchlets; main axes sometimes with very profuse short (1-3 mm ) side branches. Main axes about 1mm diameter, in cross-section central filament not easily visible; medulla comprising relatively thick-walled cells up to 100 µm in diameter. Tetrasporangial stichidia short, branched, spinous.

Collections, ecology and regional distribution

Found in algal turfs in the mid - lower eulittoral zone, and in rock pools and the shallow subtidal zone, throughout the south and east coasts from just west of Cape Agulhas to northern KwaZulu-Natal (23-58). Intertidal plants tend to be shorter, while long specimens may be found in sheltered pools or in the subtidal.

World distribution: Widespread in warm temperate-tropical regions, including the African east coast (Guiry & Guiry 2015).

Type locality: Trieste, Italy (Silva et al. 1996).

Note: A current molecular study indicates that this species may not occur in South Africa at all, and that specimens ascribed to it may be H. rosea (pers. comm. V. Johnson).

Hypnea musciformis, Port Alfred (BOL). Smallest scale divisions 1 mm.

Hypnea musciformis, fresh material, Arniston.

Hypnea musciformis, detail of branch tips, fresh material, Arniston.

Hypnea musciformis, fresh specimen, Hluleka.

Hypnea musciformis, longitudinal section with tetrasporangia. Reproduced from Stegenga et al. (1997).

Hypnea musciformis, habit. Reproduced from Stegenga et al. (1997).

Order Gigartinales

Family Cystocloniaceae

Hypnea rosea Papenfuss 1947: 1-2; fig 1; pl. 2; figs 5-8

Plants bright red, up to 17 cm long, comprising irregularly branched axes usually tangled with other algae, main axis only distinguishable in parts of plant. Axes terete, about 1 mm diameter, set with numerous short spinous ramuli imparting spiky appearance. Terminal apices frequent, spinous, may broaden slightly, often hooked or sickle-shaped (falcate). In cross-section composed of inconspicuous central filament, surrounded by large (up to 250 µm diameter) medullary cells with smaller medullary cells towards periphery, encircled by a cortex of small pigmented cells. Tetrasporangial sori on terminal branchlets. Sexual reproductive structures not seen.

Collections, ecology and regional distribution

Collected from Table Bay eastward, throughout the south and east coasts (15-58).

Common in the lower eulittoral, eulittoral fringe and down to at least several metres depth; entangled in other algae.

World distribution: Also recorded from Mozambique, Kenya and Madagascar (Silva et al. 1996).

Type locality: Mtwalume, KZN, South Africa (Silva et al. 1996).

Hypnea rosea, entangled in rock-pool seaweeds.

Hypnea rosea, detail showing slightly falcate branch-tips (fresh material, Port Alfred).

Hypnea rosea, specimen entangled with Gelidium abbottiorum, from Mbotyi (BOL).

Hypnea rosea. Habit (upper); cross section (below). Reproduced from Stegenga et al. (1997).

Order Gigartinales

Family Cystocloniaceae

Hypnea spicifera (Suhr) Harvey 1847:14

Plants green, purplish-green or purple, erect, a few to 50 cm tall, usually in dense tufts. Rhizomatous base giving rise to stiff, irregularly branched, terete, upright axes up to 3mm in daimeter, upper parts with numerous short spines, hooked apices absent. In cross –section central filament not distinguishable, central medulla with smaller cells (up to 50 µm diameter) than outer medulla (up to 150 µm). Reproductive structures in clustered spinous appendages on upper thallus parts. This species can be very variable in habit, from a short (2-3 cm), bright green turf typical of intertidal platforms, to dark purple stands of long (up to 50 cm ) thalli.

Collections, ecology and regional distribution

Found from Namibia along the whole South African coast into northern KZN (1-56), but typical of the south coast, where plants attain their greatest sizes. This species is typical of the lower eulittoral and eulittoral fringe on wave-exposed shores, particularly those influenced by sand.

World distribution: Widely distributed in the Indian Ocean (Silva et al. 1996).

Type locality: Isipingo Beach, near Durban (Silva et al. 1996).

Note: Further information on the morphology, ecology and population dynamics of H. spicifera in South Africa can be found in Isaac & Hewitt (1953) and van Zyl (1993).

Hypnea spicifera, Arniston.

Hypnea spicifera, subtidal, De Hoop.

Hypnea spicifera, terminal branchlets (purple seaweed in background is a Laurencia).

Hypnea spicifera. 1. Habit. 2. Cross section. 3. Detail of figure 2 showing outer cortical layers. Reproduced from Stegenga et al. (1997).

Order Gigartinales

Family Cystocloniaceae

Hypnea tenuis Kylin 1938: 11-12, fig. 5

Plants small (up to 5 cm), orange-red to dark red, forming loose tangle. Primary attachment often untraceable, secondary attachment to other algae by hooked apices. Axes terete, thin (200-400 µm diameter), delicate, irregularly branched; percurrent axis absent, main axes irregularly to secundly branched. In cross-section main axes with distinct central filament surrounded by medullary cells of similar diameter (40-70 µm); cortex of up to 2 layers of small cells. Tetrasporangial stichidia spindle-shaped, developing from ultimate ramuli.

Collections, ecology and regional distribution

A species of the south and east coasts, recorded from Swartklip (False Bay) to northern Kwazulu-Natal (18-58). Epiphytic on larger algae, especially Gelidium, in the sublittoral fringe and shallow sublittoral zone at wave-exposed sites.

World distribution: Also recorded from Namibia and Mozambique (Silva et al. 1996).

Type locality: Isipingo Beach, KZN, South Africa (Kylin 1938).

Hypnea tenuis, Morgan Bay.

Hypnea tenuis, branch detail.

Hypnea tenuis, BOL.

Hypnea tenuis, re-drawn from Kylin (1938: Fig. 5).

Hypnea tenuis.  Habit (above), cross section (below). Reproduced from Stegenga et al. (1997).

Order Gigartinales

Family Cystocloniaceae

Hypnea viridis Papenfuss 1947: 2, pl. 1; fig. 2; pl. 2: figs 9-11; pl. 3: fig. 14

Plants iridescent blue in situ, forming low cushions up to 2 cm high; attached by small discoid holdfasts; axes terete, up to 1 mm in diameter, branching irregular to pinnate, terminal branches short and spiky, not hooked, up to 5 mm long. In cross-section central axial filament surrounded by 5-6 large medullary cells surrounded by medullary cells decreasing in size towards the periphery: cortex of small, pigmented cells. Tetrasporangia zonately divided and arranged in sori girdling determinate branchlets; sexual reproductive structures not seen.

Collections, ecology and regional distribution

Found from northern Transkei (Pondoland) eastward through KZN (47-58), into Mozambique, in lower eulittoral zone, rock pools and subtidal down to at least 14m deep.

World distribution: A southern African endemic.

Type locality: Umhlali Beach, KZN, South Africa (Silva et al. 1996).

Note: The relationship of this species to the tropical H. pannosa, which it strongly resembles, needs investigation (De Clerck et al. 2005).

Hypnea viridis, showing blueish iridescence.

Hypnea viridis, Palm Beach (KZN) specimens, BOL.

Order Gigartinales

Family Cystocloniaceae

Rhodophyllis reptans (Suhr) Papenfuss 1956: 71-73

Plants bright red, irregularly branched in cervicorn manner, a few to 5 cm tall, lower parts of thallus prostrate. Axes membranous, 0.5 - 2 mm wide and about 200 µm thick, tapering near apices. Thallus uniaxial, with wedge-shaped apical cell, axial filament inconspicuous in older parts. Medulla usually single filamentous layer, surrounded by one or two-layered pseudoparenchymatous cortex and epidermis of squarish cells. Cells of epidermis 10-20 µm in diameter in surface view. Cystocarps scattered along thallus margins, without ostioles. Tetrasporangia scattered among cortical cells, about 60 x 30 µm, zonately divided.

Collections, ecology and regional distribution

Recorded along the west coast to northern Transkei (Pondoland) (1-47). A subtidal species, found to at least 15m depth.

World distribution : also recorded from Namibia, St Paul Island (Silva et al. 1996) and Antarctica (Papenfuss 1964).

Type locality: Algoa Bay, Cape Province, South Africa (Silva et al. 1996).

Rhodophyllis reptans, False Bay, Western Cape (BOL).

Rhodophyllis reptans, detail of branching, fresh tetrasporangial material, Tsitsikamma.

Rhodophyllis reptans, cross section showing zonate tetrasporangia (fresh material, Tsitsikamma).

Rhodophyllis reptans, cross section showing medullary and cortical cells (fresh material, Tsitsikamma).

Rhodophyllis reptans. 1. Habit. 2. Section of vegetative thallus. 3. Section of tetrasporiferous thatllus. 4. Surface view of tetrasporiferous thallus (note “repair”of holes left by released spore. Reproduced from Stegenga et al. (1997).

Order Gigartinales

Family Rhizophyllidaceae

Genus Portieria

Plants to about 20 cm tall, recumbent to erect, comprising compressed or flattened branches with inrolled apices. Branching irregular, bi- or tri-pinnate. Structure uniaxial, axial cell conspicuous; medulla psudoparenchymatous, comprising large cells, surrounded by cortex of smaller cells in anticlinal filaments. Reproductive structures in nemathecia on terminal branches. Seven species are currently recognized worldwide (Guiry & Guiry 2013) of which two are recorded from the south coast. However, recent molecular studies indicate many cryptic species in this genus worldwide (pers. comm., Olivier De Clerck).

Key to species

1a. Plants up to 20 cm tall, branches 1-3 mm wide in mid-thallus

P. hornemannii

1b. Plants up to 5 cm tall, branches 0.5-1.0 mm wide in mid-thallus

P. tripinnata

Order Gigartinales

Family Rhizophyllidaceae

Portieria hornemannii (Lyngbye) Silva in Silva et al. 1987: 129

Plants bright to orange-red, up to 20 cm tall, comprising complanate fronds arising from a small discoid to encrusting holdfast. Thallus pinnately branched, up to 5 (7) orders of alternate or subopposite laterals, indeterminate branches arising irregularly. Tips of indeterminate branches curved. Axes compressed to flattened, 1-3 mm wide, narrowest at tips, about 500 µm thick. Growth pattern uniaxial, in cross- section central axis ovoid, narrow, conspicuous, up to 170 µm in diameter; large-celled pseudoparenchymatous medulla of globose cells ranging in diameter from 17 µm in apical parts to 180 µm in basal parts, decreasing in size towards periphery and surrounded by cortex of small pigmented cells, 3-25 µm long, 3-12 µm wide, disposed in anticlinal filaments. Tetrasporangia in nemathecia concentrated along thallus edge, zonately divided and forming solid surface layer without interspersed sterile cells. Wart-like cystocarps on surface of terminal branches, developing several gonimolobes separated by thin sterile partitions.

Collections, ecology and regional distribution

Found from Table Bay to northern KZN and eastward (15-58). Epilithic, extending from intertidal zone and rock pools into subtidal zone down to at least 20 m.

World distribution: South Africa, Mozambique and widespread in Indo-Pacific (Silva et al. 1996; Guiry & Guiry 2013).

Type locality: Probably the Red Sea, although stated as Scandinavia (Silva et al., 1996).

Notes: 1. This species was formerly known as Chondrococcus hornemannii (Lyngbye) Schmitz (see Silva et al. 1987).

2. De Clerck et al. (2005) point out that tropical specimens of P. hornemannii have narrower axes and a wider branching angle than South African specimens. They also note that the South African entity once known as Chondrococcus lambertii (Grunow) Schmitz, but considered a synonym of P. hornemannii by Papenfuss (see Silva et al. 1996), may require further examination.

Portieria hornemannii, Stilbaai specimen (BOL).

Portieria hornemannii, Natures Valley. Many shallow-water specimens can be bright orange.

Portieria hornemannii, detail of branch apices.

Portieria hornemannii, female plant with cystocarps (Natures Valley).

Portieria hornemannii. 1. Thallus apex. 2-3. Cross sections. Reproduced from Stegenga et al. (1997).

Order Gigartinales

Family Rhizophyllidaceae

Portieria tripinnata (Hering) De Clerck in De Clerck et al. 2005: 192, figs 165, 166.

Plants erect, orange-brown, forming dense tufts up to 5 cm high. Discoid or encrusting holdfast; fronds complanate, compressed to flattened, with recurved tips; branching pinnate, alternate to sub-opposite; axes 0.5-1.0 mm wide in mid-thallus, narrower at both ends; ultimate branches usually < 0.5 mm long, resembling small teeth. Growth pattern uniaxial; in cross-section axial cell ovoid, conspicuous; medulla of un-pigmented globose cells 22-100 µm diameter, decreasing in size towards periphery and grading rapidly into cortex of 2-3 layers of pigmented cells 5-12 µm long and 5- 10 µm wide. Reproductive structures developing in wart-like nemathecia on terminal branches; tetrasporangia zonately divided.

Collections, ecology and regional distribution

Found from Cape Padrone area (east of Algoa Bay) to Mission Rocks, northern KZN (38-51), mostly on rock in the mid-eulittoral or in rock-pools and gulleys.

World distribution: Reported from Mauritius (Børgesen 1943).

Type locality: “Natalpoint” (Durban) Kwazulu-Natal, SA. (Krauss 1846: 209).

Note: We follow Papenfuss (1940) and De Clerck et al. (2005) in regarding P. tripinnata as a species distinct from P. hornemannii, although Silva et al. (1996) regarded it as a synonym of P. hornemannii.

Portieria tripinnata, Hluleka (BOL).

Portieria tripinnata, intertidal rocks, Glen Muir, showing typical recumbent form.

Portieria tripinnata, Haga Haga, branch detail of fresh material.

Portieria tripinnata, Palm Beach, KZN, detail of branching pattern.

Order Gigartinales

Family Phyllophoraceae

Genus Gymnogongrus Martius

Gametophytes erect, leathery or cartilaginous, terete or flattened, more-or-less regularly dichotomously branched. Anatomy pseudo-parenchymatous, with medulla of relatively large cells interspersed with smaller cells and cortex of anticlinal rows of small cells. Sporophytes either separate, crustose thalli or borne on the gametophyte in the form of pustular nemathecia. Tetrasporangia in catenate series, cruciately divided.

Note: Silva and DeCew (1992) erected the genus Ahnfeltiopsis to include species previously assigned to Gymnogongrus and Ahnfeltia that have immersed cystocarps and a crustose tertrasporophyte in their life history. These they distinguished from species with “tetrasporopblastic” nemathecia borne on the gametophyte (which remained in Gymnogongrus). They published new combinations, including the South African species Ahnfeltiopsis complicata, A. vermicularis, A. glomerata, A. polyclada, and A. intermedia. (see also Stegenga et al. 1997). However, a molecular study by Fredericq et al. (2003) showed that some species show both types of life history, and some species with immersed cystocarps are more closely related to some with nemathecia than other “Ahnfeltiopsis” species. They therefore questioned the validity of Ahnfeltiopsis and noted the need for molecular studies of the genus worldwide. On these grounds we retain the original concept of the genus Gymnogongrus, which includes the South African species previously ascribed to Gymnogongrus as well as to Ahnfeltiopsis sensu Silva and DeCew (1992). Anderson & Bolton (1990) report on the life histories of South African species.

Key to species

1 a. Axes always terete

G. polycladus

b. Axes flattened or compressed

2

2 a. Axes flattened (strap-shaped); when fertile, cystocarps immersed

G. glomeratus

b. Axes compressed; when fertile, pustule-like external nemathecia present

G. tetrasporifer

Order Gigartinales

Family Phyllophoraceae

Gymnogongrus glomeratus J. Agardh 1849: 88

Gametophytic thalli stiff, erect, flattened, 10-40 mm tall, 1-2 (3) mm wide, several times dichotomously or irregularly branched, brownish-red to dark red. Several to many axes arising from a common crustose holdfast, often giving appearance of a rounded tuft. Distal branches often incurved. Carposporophytes immersed, cystocarps hemispherical and projecting from the thallus surface, with one microscopic, terminal ostiole (when mature).

Sporophytes crustose, comprising a monostromatic basal layer that produces erect filaments dorsally (Anderson & Bolton 1990). Tetrasporangia cruciately divided, developing in a chain-like series several cells below the surface of the crust. Crusts red, closely adherent to substratum.

Collections, ecology and regional distribution

Found from Namibia to Goukamma (1-29). Often occurs in sand-affected areas: crustose holdfasts and tetrasporophytes shown to survive long periods of sand-burial (Anderson et al. 2008).

World distribution: Also found in Namibia: Southern African endemic.

Type locality: Cape of Good Hope (Silva et al. 1996).

Notes: 1. South Coast specimens appear to be smaller in general than many on the west coast of South Africa (as described in Stegenga et al. 1997).

2. Based on a DNA study, Fredericq et al. (2003) considered Ahnfeltiopsis glomerata to be synonymous with Gymnogongrus dilatatus. However, we maintain G. glomeratus as a separate species because all the S coast specimens we have found clearly fit the description of this species, and specimens fitting the description of G. dilatatus do not appear to occur here.

Gymnogongrus glomeratus, with Ulva sp in the background.

Gymnogongrus glomeratus, XS of thallus showing small cortical and large medullary cells.

Gymnogongrus glomeratus, XS of blade and two cystocarps.

Gymnogongrus glomeratus, XS of outer region of male thallus showing peripheral layer of colourless spermatangia.

Gymnogongrus glomeratus, crusts of young sporophytes grown in culture (coin ca 15 mm in diameter).

Gymnogongrus glomeratus, cross section of cultured tetrasporophytic crust showing adherent filaments of tetrasporangia.

Gymnogongrus glomeratus, habit. Reproduced from Stegenga et al. (1997)(as Ahnfeltiopsis glomeratus).

Order Gigartinales

Family Phyllophoraceae

Gymnogongrus polycladus (Kützing) J. Agardh 1851

Gametophytic thalli stiff, erect, from a few to 10 cm tall, branching up to 6 times. Axes terete, dichotomies in the upper half fairly regular. Older axes often with some short, rather pointed lateral branches in secund or opposite series. Axes in small specimens less than 1 mm in diameter, in larger specimens up to 2 mm diameter. Apices tapered. Carposporophytes immersed, visible as swellings in main axes and sometimes in short side-branches. Sporophytes crustose (see Anderson & Bolton 1990 or description of G. glomeratus).

Collections, ecology and regional distribution

Found from False Bay eastwards to Cape Morgan (17-43), with the smaller “G. intermedius” form (see note below) recorded from Jongensfontein, Keurboomstrand and Port St Johns. Typically found in sand-influenced lower sublittoral zone.

World distribution: South Africa, Tristan da Cunha.

Type locality: Kalk Bay, False Bay (Silva et al. 1996).

Note: This species includes the former Gymnogongrus intermedius (Ahnfeltiopsis intermedia), which on the basis of DNA comparisons was shown to be a small form of G. polycladus (Fredericq et al 2003).

Gymnogongrus polycladus, herbarium specimen from Kowie (BOL).

Gymnogongrus polycladus, cross section of female thallus showing ostiole between the cystocarp and exterior.

Gymnogongrus polycladus, habit. Reproduced from Stegenga et al. (1997) (as Ahnfeltiopsis polycladus).

Order Gigartinales

Family Phyllophoraceae

Gymnogongrus tetrasporifer Papenfuss in R.J.Anderson & Bolton 1990: 383, fig. 6

Thalli erect, 4-6 cm tall and up to 1 mm diameter, slightly compressed, regularly dichotomously branched. Axes often irregularly constricted, with rounded apices. Hemispherical nemathecia up to 1.5 mm in diameter occuring on the most distal 3 orders of branches. Nemathecia comprising filaments of cells with cruciately-divided tetrasporangia forming distally.

Collections, ecology and regional distribution

Found from Kowie to northern KwaZulu-Natal ((39-58).), in intertidal rockpools and in sublittoral down to at least 6m depth.

World distribution: South African endemic.

Note: The name Gymnogongrus tetrasporifer was used by Papenfuss in an unpublished manuscript, and it is illustrated and referred to by Anderson and Bolton (1990) and De Clerck et al. (2005), but awaits more comprehensive molecular studies of the genus before it can be validated (see Fredericq et al. 2003).

Gymnogongrus tetrasporifer, Haga Haga.

Gymnogongrus tetrasporifer, showing distinctly protruding nemathecia near branch apices (BOL, Kowie specimen).

Gymnogongrus tetrasporifer, nemathecia on fresh material.

Gymnogongrus tetrasporifer, Cape Morgan, det by Papenfuss (BOL).

Gymnogongrus tetrasporifer, XS through nemathecium (circle of loose tissue on left) and axis (dark-outlined tissue on right) (re-hydrated BOL specimen).

Order Gigartinales

Family Gigartinaceae

Chondracanthus acicularis (Roth) Fredericq in Hommersand, Guiry, Fredericq & Leister 1993: 117

Thalli prostrate or erect, firm to cartilaginous, dark reddish-purple, sometimes with white or green spots, up to about 100 mm long, but often smaller. Holdfast gives rise to several fronds, branching pinnate to irregular, branches 1 (-2) mm wide, cylindrical to compressed, curved, branch tips very pointed; Branches may attach secondarily to substratum. Holdfast initially discoid but may become stoloniferous. Structure multiaxial; medulla of compact thick-walled filaments; cortex of radial filaments of small cells (about 6 µm in diameter). Spermatangia not seen. Cystocarps forming swellings near apices. Tetrasporangia embedded in inner cortex in sori, producing dark slightly swollen areas, cruciately divided, 20-40 µm long.

Collections, ecology and regional distribution

Found from De Hoop to Cape Morgan in the Eastern Cape (24-40). Often associated with sand.

World distribution: widespread in tropical and temperate seas.

Type locality: Adriatic Sea (Dixon & Irvine 1977).

Note: 1. The genus Chondracanthus was resurrected by Hommersand et al. (1993); this species was previously known as Gigartina acicularis (Roth) Lamouroux. According to Hommersand et al. (1993) the centre of distribution of the genus Chondracanthus is the eastern North Pacific Ocean.

2. There are several specimens in BOL from the Kowie and labeled Gigartina teedei but later annotated “G. acicularis” by G.F. Papenfuss. These specimens fit the description of C. acicularis, and closely resemble European and larger local specimens of C. acicularis. We conclude that Chondracanthus teedei does not occur on the south coast.

Chondracanthus acicularis, De Hoop intertidal zone.

Chondracanthus acicularis, detached clump of thalli.

Order Gigartinales

Family Gigartinaceae

Gigartina Stackhouse 1809: 55

Thalli usually erect, ranging in shape from foliose to narrowly pinnate. Plants may be large and foliose or small and procumbent, with creeping hapterous holdfasts. Thalli often bear superficial branchlets or papillae. Growth is by meristems located in apices or thallus margins. Cystocarps are conspicuous and found in papillae or swellings on the surfaces and/or margins of thalli. A filamentous sheath surrounds the carposporophyte and carpospores are released through ostioles. Tetrasporangia are formed in sori at the inner cortex/medulla boundary, from inner cortical cells. Tetraspores are released in a gelatinous matrix by excision of soral tissue.

According to Guiry & Guiry (2012), 38 species are currently recognized in the genus Gigartina, which is distributed throughout cool to cold temperate seas. South Africa (along with Chile, Japan, New Zealand and California) is recognized as a centre of rich species diversity.

Key to species

1a. Axes terete or almost terete

2

1b. Axes compressed or flattened (blades)

4

2a. Thallus irregularly branched, small (diameter ca.1 mm, up to 1.5 cm high)

G. minima

2b. Thallus dichotomously branched, large (diameter 2-3 mm, up to 20 cm high)

G. pistillata

3a. Blades with smooth surfaces but small branchlets along margins

G. insignis

3b. Blade surfaces rough in mature thalli, with papillae or peg-like outgrowths

4

4a. Blades ovate, covered in papillae

G. polycarpa

4b. Blades wedge- or strap-shaped, with peg-like superficial outgrowths

G. paxillata

Order Gigartinales

Family Gigartinaceae

Gigartina insignis (Endlicher and Diesing) Schmitz in Barton 1896: 459

Thalli dark to blackish-red, leathery, branched, up to 20 (-40) cm tall. Holdfast a small disc; main axes compressed, up to about 2 (-3) cm wide, branching irregular to dichotomous, branches with numerous opposite or sub-opposite laterals one to a few mm long and about 1-2 mm wide. Thallus multiaxial; in cross-section medulla a network of branching cells, cortex of anticlinal filaments of small cells. Cystocarps visible as globose swellings about 2 mm in diameter, in lateral proliferations. Tetrasporangia developing in inner cortex, visible in fresh blades as small dark spots.

Collections, ecology and regional distribution

Collected from False Bay (Muizenberg) to Port Alfred, but apparently extends to at least Durban(18-51), where the type specimen was collected. There are many large specimens in BOL that were collected in drift, indicating a mainly subtidal vertical distribution.

World distribution: Endemic to South Africa.

Type locality: Port Natal (Durban), South Africa (Silva et al. 1996).

Note: This Gigartina species was not examined by Hommersand et al. (1993, 1994).

Gigartina insignis, female thallus, Kowie (BOL).

Gigartina insignis, female thallus, detail of margins showing papillae.

Gigartina insignis, male or vegetative thallus with no marginal papillae, Kowie (BOL).

Gigartina insignis, young plant from East London (BOL).

Gigartina insignis. 1. Habit. 2. Cross section. Reproduced from Stegenga et al. (1997).

Order Gigartinales

Family Gigartinaceae

Gigartina minima Kylin 1938: 13, fig. 11.

Thallus reddish-brown, 0.5 – 1.5 cm high, sparely and irregularly branched, axes 600-900 µm diameter, tapering towards apices and bearing thorn-like branchlets. Thallus multiaxial; in cross-section medulla a network of branching cells, cortex of anticlinal filaments of small cells. Cystocarps almost spherical, borne singly on branchlets. Tetrasporangia developing in cortical layer.

Collections, ecology and regional distribution

Found from Goukamma to near Cape St Lucia in KZN (29-55). Lower eulittoral, usually as a rather cryptic component of mixed turfs.

World distribution: apparently a South African endemic.

Type locality: Isipingo Beach, near Durban (Kylin, 1938).

Gigartina minima, from Palm Beach, southern KZN.

Gigartina minima, cross section.

Order Gigartinales

Family Gigartinaceae

Gigartina paxillata Papenfuss 1947: 2-3, pl. 1: fig. 3; pl. 3: fig. 15; pl.4

Plants dark red to blackish-red, erect to loosely caespitose, with simple or forked blades bearing peg-like branchlets from surface and margins. Stipes terete to slightly compressed, to 1.5 cm long and 2 mm diameter, widening into blades up to about 20 cm long and 4 cm broad, 750-1250 µm thick, forked once or twice, bearing simple or forked marginal branchlets and simple or divided terete to slightly compressed papillate to elongated superficial and marginal outgrowths up to 1.3 cm long and 1.2 mm diameter. Cystocarps terminal or lateral on the marginal peg-like branchlets, 1-2 mm diameter, with distinct pericarp and terminal ostiole. Tetrasporangia in raised, lens-like sori formed in the cortex-medulla boundary, 150-1500 µm in diameter, on both the superficial and marginal outgrowths and the faces of fronds. Tetraspores formed by cruciate cleavage and released in gelatinous matrix.

Distribution and ecology

Recorded from De Hoop to Cove Rock, near East London (24-41), in the lower eulittoral zone.

World Distribution: South African endemic.

Type locality: Storms River, Tsitsikamma, South Africa (Papenfuss 1947).

Notes: 1. G. paxillata was not included in the species of Gigartinaceae examined by Hommersand et al. 1993. However, Levitt (1998) showed that this species is correctly placed within the genus Gigartina because it has “mature carposporangia in grape-like clusters and broadened pit-connections in the placental tissue, and “ tetrasporangia formed progressively at the cortex-medulla boundary and tetraspores released by excision of the entire sorus” unlike Chondracanthus, where tetrasporangia are formed within the cortex and tetraspores via a pore.

2. The specimen “Gigartina stiriata” from Port Elizabeth illustrated by Kylin (1938: fig. 12) appears to be G. paxillata.

Gigartina paxillata, Natures Valley.

Gigartina paxillata, tetrasporangia visible in fronds as dark dots.

Gigartina paxillata, single female frond (Natures Valley).

Gigartina paxillata, cross section.

Gigartina paxillata, cross section showing outer layer of medulla and the cortex.

Order Gigartinales

Family Gigartinaceae

Gigartina pistillata (S.G. Gmelin) Stackhouse 1809: 74

Thalli usually dark- to blackish-red but sometimes green or greenish-purple, terete basally to slightly flattened distally, irregularly dichotomously branched, up to 20 cm tall. Holdfast a small disc or compound disc giving rise to many uprights; axes 2-3 mm diameter, tough, branched more frequently and sometimes very densely towards distal end. Structure multiaxial; medulla of thick-walled filaments surrounded by narrow layer of cortical filaments in a mucilaginous matrix. Female thalli with fairly numerous small lateral branches that come off mainly in two opposite rows running down opposite sides of the flattened distal axes. Lateral branches bearing one to more than 5 globose cystocarps (2-3 mm diameter), each with a distinct ostiole. Tetrasporangial sori immersed in inner cortex along margins of distal axes, forming dark longitudinal swellings; tetrasporangia cruciately divided, 25-40 µm long and 10-25 µm wide.

Distribution and ecology

Found from False Bay eastwards to the Port Alfred area (17-39), in the sublittoral fringe and shallow sublittoral.

World distribution: Found on the warm temperate coasts of Europe and at the Canary Islands.

Type locality: Unknown, for information on type see Dixon and Irvine 1977: 239.

Note: This species was retained in the genus Gigartina by Hommersand et al. (1993, 1994).

Gigartina pistillata, green form from Three Sisters, near Port Alfred.

Gigartina pistillata, habit. Reproduced from Stegenga et al. (1997).

Order Gigartinales

Family Gigartinaceae

Gigartina polycarpa (Kützing) Setchell and Gardner 1933: 295.

Thalli foliose, reddish or yellowish-brown to black, up to 40 cm long but usually shorter, covered with roughish papillae. Holdfast a small disc or compound disc from which many uprights arise; stipes short, terete to slightly compressed, bearing one to several blades, usually only one or two main blades that are ovate and larger than the others. Surfaces of blades (particularly main blades) distinctly papillate; cystocarpic specimens with well-developed globose papillae. Specimens from the Eastern Cape generally more densely papillate than those from the Western Cape (Stegenga et al. 1997).

Distribution and ecology

Found from the west coast around to Three Sisters, near Port Alfred (17-39), in the lower eulittoral zone. Generally not as abundant on the south coast as on the west coast where this species often dominates parts of this zone.

World distribution: the South African entity is probably endemic (Stegenga et al. 1997).

Note: the South African entity was previously included in G. radula, the name of which was changed to G. polycarpa (Hommersand et al. 1994). However, the type of G. radula is Australian, and the SA entity was not examined by Hommersand et al. (1994), and requires further study.

Gigartina polycarpa.

Gigartina polycarpa, yellowish-brown plants, Arniston.

Gigartina polycarpa, habit. Reproduced from Stegenga et al. (1997; as G. radula).

Order Gigartinales

Family Plocamiaceae

Plocamiocolax papenfussiana Martin & Pocock 1953: 50

Plants parasitic on Plocamium corallorhiza, forming dense whitish pustular tufts up to 6 mm in diameter, usually on reproductive sori of the host, but sometimes on margins or bases of fronds; individual thalli colourless to whitish, comprising stalk-like portion bearing a number of irregular branches, ultimate branches with acute to sub-acute tips; colourless rhizoidal filaments penetrating host tissue. Reproductive structures resemble those of the host but are colourless; tetrasporangia in forked stichidia, zonately divided and up to 20 x 40 µm in size; cystocarps up to 400 µm in diameter, globose; spermatangia not seen, but species reported to be monoecious (Martin & Pocock 1953).

Collections, ecology and regional distribution:

Reported from Melkbosstrand, north of Cape Town, to East London (15-41).

World distribution: South African endemic.

Type locality: Strandfontein, False Bay, Western Cape (Martin & Pocock 1953).

Plocamiocolax papenfussiana on branches of Plocamium corallorhiza.

Plocamiocolax papenfussiana. 1. Microscopic habit of female plant with cystocarps, on section of Plocamium corallorhiza (below). 2. Carposporophyte. 3. Detail of fertile tetrasporophyte. Reproduced from Stegenga et al. (1997).

Order Gigartinales

Family Plocamiaceae

Plocamium Lamouroux

Plants with a more-or-less-developed holdfast of prostrate axes bearing several erect axes which may be terete, compressed or flattened; erect axes often complanate. Branching is sympodial, producing an alternate series of simple and/or compound laterals (branchlets), compound laterals repeating the branching pattern. The number of simple and /or compound laterals is generally characteristic of each species. Growth is apical, the central filament of the uniaxial thalli remaining visible in some species. The central filament lies within a pseudoparenchymatous medulla of large cells and a cortex of smaller cells. Reproductive structures develop on the ultimate and penultimate laterals or on adventitious branches in the axils of laterals. Tetrasporangia develop on modified branchlets (stichidia) in double rows, and are zonately divided. Cystocarps spherical, without ostiole.

The genus is cosmopolitan, and 39 species are recognised worldwide, among which 10 occur in South Africa (Guiry & Guiry 2009). We have recorded 8 species on the South African south coast.

Key to species

1a. Erect axes 5 mm wide or wider

P. corallorhiza

1b. Erect axes 3 mm wide or narrower

2

2a. Pectinate series of laterals comprising up to 6 (-10) branchlets, lower branchlet often compound

P. glomeratum

2b. Pectinate series of laterals comprising no more than 4 branchlets

3

3a. Erect axes terete throughout, pectinate series of laterals comprising 1 simple and 1 compound branchlet

P. cornutum

3b. Erect axes flattened or compressed throughout or at least in distal parts

4

4a. Erect axes rather membranous, up to 3 mm wide, pectinate series of laterals comprising 1 simple and 2-3 compound branchlets, stichidia (if present) terminal on compound branchlets

P. maxillosum

4b. Erect axes less than 3 mm wide

5

5a. Pectinate series of laterals generally comprising 2-4 branchlets (1 simple and 1, 2 or 3 compound)

6

5b. Pectinate series of laterals consisting of 2 branchlets (1 simple, 1 compound)

7

6a. Plants delicate, less than 0.5 mm wide, axes zig-zagging somewhat

P. beckeri

6b. Plants rather robust, to 1 mm wide, profusely branched, young laterals slightly

P. rigidum

7a. Simple lateral branchlets in pectinate series narrowly triangular, less than 0.5 mm wide at base, with smooth margins

P. telfairiae

7b. Simple branchlets broadly triangular, often more than 1 mm wide at base, outer edge sometimes slightly serrated

P. suhrii

Order Gigartinales

Family Plocamiaceae

Plocamium beckeri Simons 1964: 185

Plants rather fine and delicate, bright red to pink, largely complanate, up to 15 cm tall; holdfast rhizomatous, intertwined; axes terete basally, becoming compressed distally, up to about 400 µm in diameter, usually with slight zig-zag along their length; branching prolific, distichous, alternate; a simple lateral alternates with a group of 2 (-3) compound laterals; simple lateral awl-shaped, up to 1 (-1.5) mm long and 100-300 µm wide at base, margins smooth. Tetrasporangial stichidia developing from modified laterals, clavate, simple or compound, often cruciform; tetrasporangia 65 x 40 µm, zonately divided.

Collections, ecology and regional distribution

Recorded from Struisbaai all along the south coast and as far as Black Rock, northern Kwazulu-Natal (23-58), in mid to lower eulittoral tide pools and the shallow sublittoral zone.

World distribution: Also recorded from Angola (Stegenga et al. 1997).

Type locality: Port Edward, Kwazulu-Natal, South Africa (Simons 1964).

Plocamium beckeri, Arniston.

Plocamium beckeri, branch detail.

Plocamium beckeri, detail of stichidial branch.

Plocamium beckeri, stichidia on lower branches.

Plocamium beckeri, stichidium.

Plocamium beckeri, tetrasporiferous thallus apex. Reproduced from Stegenga et al. (1997).

Order Gigartinales

Family Plocamiaceae

Plocamium corallorhiza (Turner) J.D. Hooker & Harvey 1845: 542

Plants erect, bright pink sometimes with bluish cast, up to 30 cm tall, robust; well-developed stoloniferous holdfast bearing several to many erect axes; erect axes complanate, foliaceous, up to 7 mm wide, branching distichous, alternate; Series of laterals comprising simple lateral alternating with a compound lateral, simple laterals incurved when young, triangular when mature, up to 5 mm wide at base, abaxial margin distinctly dentate. Tetrasporangial stichidia densely clustered in the axils of laterals, cylindrical, simple or once-forked, about 120 µm in diameter; tetrasporangia zonately divided, about 65 µm long.

Collections, ecology and regional distribution

Recorded from Yzerfontein on the west coast to northern Kwazulu-Natal (14-58). This species is often common and abundant, forming stands in the shallow sublittoral.

World distribution: Also recorded from Namibia, Cape Verde Islands, Madagascar, Mozambique, and Amsterdam and St. Paul Islands (Guiry & Guiry 2009).

Type locality: Cape of Good Hope (Silva et al. 1996).

Note: Plocamium robertiae F. Schmitz ex Mazza may be a valid species, although it is currently regarded as a synonym of a P. corallorhiza. (Simons 1964; Stegenga et al. 1997). However, DNA sequences of the two species need to be compared.

Plocamium corallorhiza, Arniston, showing stichidia (dark spots) in axils of branchlets.

Plocamium corallorhiza, showing stichidia in branch axils. Plants often have a bluish iridescence.

Plocamium corallorhiza. 1. Habit (with the parasite Plocamiocolax papenfussianum – arrowed). 2. Thallus apex. 3. Cross section. 4. Tetrasporangial stichidia. Reproduced from Stegenga et al. (1997).

Order Gigartinales

Family Plocamiaceae

Plocamium cornutum (Turner) Harvey 1849 (1847-1849): 123

Plants reddish-brown, fairly robust, up to 20 cm tall, often in clumps or swards. Holdfast stout, branched, stoloniferous; erect axes terete, up to 1.5 mm in diameter, not complanate; main axes sparsely branched, alternating pectinate series of laterals comprising one simple and one compound branch; lower lateral awl-shaped, simple, up to 1 mm wide at the base and to 1 cm long, juveniles rather incurved; upper laterals compoundly branched; laterals somewhat crowded, overlapping, appearing to arise on all sides of axes. Reproductive structures clustered in axils of laterals.

Collections ecology and regional distribution

Recorded from Namibia along the west coast and as far as the Kei Mouth area on the south coast (1-43). Epilithic in the mid to lower eulittoral on wave-exposed shores.

World distribution: A southern African endemic.

Type locality: Cape of Good Hope, South Africa (Turner 1819).

Plocamium cornutum, Arniston, in lower eulittoral zone.

Plocamium cornutum.

Plocamium cornutum, detail of frond.

Plocamium cornutum. 1. Habit. 2. Thallus apex. 3. Cross section of main axis. 4. Detail of medullary cells (note numerous sedcondary pit connections). Reproduced from Stegenga et al. (1997).

 Order Gigartinales

Family Plocamiaceae

Plocamium glomeratum J. Agardh 1851: 397

Plants bright to dark red, up to 10 cm tall. Axes complanate, flattened, up to 1 mm wide, branching subdichotomous to irregular; alternating series of laterals comprising up to 6 (-10) branchlets, all compound or proximal branchlet at first simple and later branching; young branchlets somewhat incurved, older branchlets almost straight and with acute apex. Tetrasporangial stichidia developing from apical parts of plant, becoming crowded and almost ball-like.

Collections, ecology and regional distribution

Recorded from Namibia to Still Bay (Simons 1964) (1-26). Epilithic in the subtidal fringe and in tidal pools, often associated with sand.

World distribution: Also recorded from Madagascar and Mozambique (Silva et al. 1996).

Type locality: Cape of Good Hope, South Africa (Silva et al. 1996).

Plocamium glomeratum. 1. Thallus apex. 2. Thallus apex, with tetrasporangial stichidia. Reproduced from Stegenga et al. (1997).

Order Gigartinales

Family Plocamiaceae

Plocamium maxillosum (Poiret) J.V. Lamouroux 1813: 138

Plants bright red, up to 20 cm tall, complanate. Axes compressed, up to 3 mm wide, rather membranous. Branching fairly regular, groups of laterals comprising one simple and two or sometimes three compound branchlets; simple branchlets triangular, up to 1 mm wide at the base, sometimes slightly serrated along the abaxial margin. Tetrasporangial stichidia borne on ends of compound branchlets, not notably crowded; young stichidia stout, maturing into cylindrical 1-3 times forked structures of about 100 µm diameter.

Collections ecology and regional distribution

Recorded from the Orange River to Tsitiskamma (1-32). Apparently a sublittoral species, to at least 24 m depth.

World distribution: Also recorded from Namibia (Rull Lluch 2002) and Mozambique (Silva et al. 1996)

Type locality: “Cape of Good Hope”, South Africa (Silva et al. 1996).

Plocamium maxillosum, west coast specimen (Cape Point).

Plocamium maxillosum, slightly narrower form (Knysna Heads).

Plocamium maxillosum, tettrasporangial stichidia.

Plocamium maxillosum, thallus apex. Reproduced from Stegenga et al. (1997).

Order Gigartinales

Family Plocamiaceae

Plocamium rigidum Bory de Saint-Vincent in Bélanger & Bory de Saint-Vincent 1834: 164

Plants red to reddish-brown, up to 20 cm tall, largely complanate, profusely branched, rather cartilaginous. Axes terete to slightly compressed, up to 1 mm wide. Groups of laterals comprising one simple and one or two compound branchlets; young laterals somewhat incurved, simple branchlets awl-shaped, up to 0.5 mm wide at the base. Tetrasporangial stichidia borne on ultimate or sometimes penultimate compound branchlets, simple and rather stout when young, with one or two bifurcations when mature, up to 100 µm in diameter.

Collections ecology and regional distribution

Recorded from Namibia, along the South African west and south coasts, to Palm Beach (Port Edward area) in southern Kwazulu-Natal (1-48). Common in the sublittoral fringe, rock pools and the shallow sublittoral zone.

World distribution: Also recorded from Namibia and from Amsterdam Island in the Indian Ocean (Silva et al. 1996).

Type locality: “Cape of Good Hope” (Silva et al. 1996).

Plocamium rigidum, Struisbaai.

Plocamium rigidum.

Plocamium rigidum, branch detail.

Plocamium rigidum. 1. Thallus apex with tetrasporangial stichidia. 2-3. Tetrasporangial stichidia, in ventral and lateral view respectively. Reproduced from Stegenga et al. (1997).

Plocamium rigidum, thallus apex, vegetative. Reproduced from Stegenga et al. (1997).

Order Gigartinales

Family Plocamiaceae

Plocamium suhrii Kützing 1849: 886

Plants bright to pale red, up to 15 cm tall, complanate, profusely branched. Erect axes compressed, 1.5 – 2 mm wide, groups of lateral branchlets comprising one simple and one compound branchlet; simple branchlets triangular, up to 4 mm long and 1.5 mm wide at the base, sometimes with slightly serrated outer margin. Tetrasporangial stichidia 2-3 times branched, developing laterally on adventitious branchlets borne in axils of lateral branchlets.

Collections, ecology and regional distribution

Recorded from Stilbaai to Mission Rocks in northern Kwazulu-Natal (32- 56). Found in lower-eulittoral rock pools in the shallow sublittoral.

World distribution: Also recorded from Angola, Namibia, Madagascar and Amsterdam and St Paul Islands, as well as from Korea (Guiry & Guiry 2009).

Type locality: “Cape of Good Hope” (Silva et al. 1996).

Note: Stegenga et al. (1997) point out that there is some confusion about the identity of specimens from False Bay that have been referred to this species. We regard the distribution of this entity on the South African coast as lying to the east of Cape Agulhas.

Plocamium suhrii, Arniston.

Plocamium suhrii, branch detail.

Plocamium suhrii, stichidia.

Plocamium suhrii, thallus apex. Reproduced from Stegenga et al. (1997).

Order Gigartinales

Family Plocamiaceae

Plocamium telfairiae (W. J. Hooker & Harvey) Harvey ex Kützing: 1849: 885

Plants reddish-pink, up to 9 cm tall, somewhat membranous, rather triangular in outline. Erect axes slender, compressed, 1 (-2) mm wide, lateral branchlets in alternating series of one simple, entire, narrowly triangular branchlet and one compound branchlet. Tetrasporangial stichidia stellate or tree-shaped, replacing a compound branchlet.

Collections, ecology and regional distribution

Recorded from Port Alfred to northern Kwazulu-Natal (39-58). Found in intertidal pools and the sublittoral zone (to at least 20 m depth in Kwazulu-Natal).

World distribution: Also recorded from Ghana, Mozambique, and many other warm temperate and tropical localities (Guiry & Guiry 2009). Curiously, recorded from Antarctica by Papenfuss (1964).

Plocamium telfairiae, BOL.

Plocamium telfairiae, specimen from northern KZN. Note stichidial branchlets lower down on thallus.

Plocamium telfairiae, with terminal stichidia on branchlets.

Order Gigartinales

Family Sarcodiaceae

Sarcodia dentata (Suhr) R. E. Norris 1987: 152-154, figs 1-8

Plants with a small discoid holdfast and a dark red, cartilaginous, upright thallus of strap-like, pseudodichotomously branched blades, often with irregularly dentate margins and sometimes with marginal proliferations towards apices. Thalli up to 40 (-45) cm tall, blades usually a few cm wide, and about 1 mm thick. Growth multiaxial; medulla filamentous, interspersed with denser stellate cells, towards exterior rapidly changing to large-celled pseudoparenchymatous cortex; outer cortex of small cells in anticlinal rows. Tetrasporangial plants smooth, tetrasporangia scattered in outer cortex, zonately divided, about 50 x 22 µm. Female plants covered with conspicuous, protuberant cystocarps which may be elevated on stalks. Male plants smooth, with light, slightly raised areas of spermatangial sori.

Collections, ecology and regional distribution

A subtidal species, found from just west of Agulhas (Die Walle) to southern KZN (Palm Beach) (18 -48). There is, however, one specimen from Swartklip, False Bay, in the Pocock collection (GRA).

World distribution: Australia and NZ (Papenfuss 1964), Mauritius, Indonesia (Silva et al. 1996).

Type locality: Algoa Bay (Silva et al. 1996).

Sarcodia dentata, Kenton-on-Sea.

Sarcodia dentata, cystocarpic blades (BOL).

Sarcodia dentata, smooth (male?) blades, Port Alfred (BOL).

Sarcodia dentata, XS showing filamentous medulla, large-celled inner cortex and small-celled outer cortex.

Sarcodia dentata, outer cortex with zonately divided tetrasporangia.

Sarcodia dentata. 1. Habit. 2. Cross section with tetrasporangia. Reproduced from Stegenga et al. (1997).

Order Gigartinales

Family Plocamiaceae

Trematocarpus flabellatus (J.Agardh) De Toni 1900: 418-419

Plants red, erect, stiffly cartilaginous, up to about 6 cm tall, several to twelve times dichotomously branched. Axes flattened, in cross-section semicircular to elongate ellipsoidal, 2-3 mm wide and up to 1 mm thick. Medulla pseudoparenchymatous with rounded cells up to about 250 µm in diameter and well-developed intercellular spaces; outer cortical cells small, elongated radially. Tetrasporangial sori on one or both sides of blade, sporangia immersed, up to 70 x 25 µm. Cystocarps scattered, lateral on blades in subterminal sections of the thallus.

Collections, ecology and regional distribution

Recorded along the entire west coast to as far east as Kini Bay, near East London (1-41). Epilithic, found in rock pools and from the lower eulittoral to subtidal zone to at least 10 m depth; sometimes forming a dense band just above LWS.

World distribution: Also recorded from Namibia (Rull Lluch 2002) and Amsterdam and St Paul Islands in the southern Indian Ocean (Silva et al. 1996).

Type locality: Cape of Good Hope, South Africa (Sea Point, Cape Town) (Simons 1983).

Trematocarpus flabellatus, Noetsie, De Hoop, exposed at low tide.

Trematocarpus flabellatus, XS showing internal structure.

Trematocarpus flabellatus, 1 Habit. 2 Cross section. Reproduced from Stegenga et al. (1997).

Order Rhodymeniales

Family Rhodymeniaceae

Botryocladia Kylin

Thalli consisting of solid, short or long, upright or creeping axes bearing hollow, mucilage-filled vesiculate branches. About 30 species currently recognized worldwide, mostly from tropical and warm-temperate shores (Guiry & Guiry, 2009). See Norris (1989) for an account of KwaZulu-Natal species, including south coast taxa.

Key to species

1a Main axes flattened, 3-8 mm wide

B. beckeriana

1b Main axes terete to slightly compressed, less than 2 mm wide

2

2a Thalli stiffish, vesicles 4-10 mm long, ovate, slightly compressed

B. madagascariensis

2b Thalli lax, vesicles mostly 1-2 mm long, usually unilateral on axes

B. pienaarii

Order Rhodymeniales

Family Rhodymeniaceae

Botryocladia beckeriana (Holmes) Papenfuss 1958: 107

Plants up to 70 cm long or longer, comprising a thick, compressed axis (to 8 mm wide) bearing two opposite rows of small stalked vesicles; axis pseudodichotomously branched, tapering gradually, apex acute and terminating in two vesicles; vesicles stalked, 1-2 (-5) mm long, usually crowded but rarely sparse or absent. Axes in transverse section with medulla of round to ovate hyaline cells about 300-400 µm in diameter, interspersed with smaller cells and surrounded by cortex of small pigmented cells; vesicles hollow, wall comprising 3 (rarely to 5) layers of cells; inner medullar layer of round to elongated cells 50-70 µm in diameter, bearing groups of 2-3 secretory cells that protrude into central cavity; outer cortical cells about 8-10 µm in size. Tetrasporangia spheroidal, tetrahedrally divided, about 15 µm in diameter, in sori covering all but tips and bases of vesicles; cystocarps (up to 6 on same vesicle) immersed in vesicle wall, up to 700 µm in diameter. Male plants unknown.

Collections, ecology and regional distribution

Recorded from Port Alfred to northern Kwazulu-Natal (39-58). Found in the sublittoral fringe and deeper.

World distribution: South African endemic.

Type locality: Kowie (Port Alfred) (Silva et al. 1996).

Botryocladia beckeriana, Cape Vidal.

Order Rhodymeniales

Family Rhodymeniaceae

Botryocladia madagascariensis G. Feldmann 1945: 57-59, figs 3-5

Plants erect, up to 6 cm tall, comprising irregularly branched solid axes bearing two rows of vesicles, marked with blue iridescence when fresh. Axes terete, 1-2 mm in diameter, stiffish; vesicles compressed, alternately distichously arranged, elliptical to obovate, 4-10 mm long and 3-5 mm wide, with short (to 1 mm) stalk. Axes in cross section with medulla of hyaline, roundish cells of various sizes (from about 10 to 60 µm diameter) and a small-celled outer cortex. Vesicles hollow, with wall of 4-6 cell layers; outer cortex of small spheroidal to ellipsoidal cells about 8 µm in diameter, grading into large (to 100 µm in diameter) inner medullary cells; secretory cells, sometimes in short chains, borne internally on medullary and special bearing cells. Tetrasporangia ellipsoidal, borne in irregular sori on vesicles, cruciately (rarely tetrahedrally) divided, about 40 x20 µm, immersed in outer cortex and surrounded by elongated sterile paraphyses; cystocarps up to 500 µm diameter, embedded in vesicles, 3-6 on the same vesicle, often in a centrally-placed ring; male plants with spermatangial sori near centre of vesicles.

Collections, ecology and regional distribution

Recorded from De Hoop to the Durban area (24-51). Found in the sublittoral fringe on wave-exposed shores, usually under ledges and in crevices.

World distribution: Mediterranean, Canary Islands, Madagascar (Guiry & Guiry 2009).

Type locality: Madagascar (Silva et al. 1996).

Botryocladia madagascariensis, De Hoop.

Order Rhodymeniales

Family Rhodymeniaceae

Botryocladia pienaarii RE Norris 1989: 132-136, figs 1-15, 43

Plants red, rather delicate, up to 15 cm long, comprising pseudodichotomously branched terete axes usually bearing unilateral rows of round to ovoid hollow vesicles; axes about 1 mm diameter, often with a single larger vesicle in the axil of branches; vesicles almost always borne only on one side of an axis, usually 1-2 mm long (rarely to 5 mm), with short stalks. Axes in cross section with cortex of 1-2 layers of small cells overlying a solid medulla of ellipsoidal cells (about 200 x 50 µm) arranged with their long axes perpendicular to the main axis; vesicles in cross section with wall of 2-3 cell layers, some of the medullary cells with parts of their walls exposed to exterior, except on basal parts of vesicle where cortex is continuous; outer cortical cells about 10 µm diameter, inner medullary cells about 50 µm diameter; series of up to 4 secretory cells borne on inner medullary or special bearing cells, protruding into central cavity. Tetrasporangia scattered over vesicles, embedded in cortex, ellipsoidal to flask-shaped, cruciately divided and up to 40 µm long. Cystocarps about 500 µm in diameter, 1-6 per vesicle, embedded in vesicle walls and protruding internally and externally.

Collections, ecology and regional distribution

Found in the shallow sublittoral from Port Alfred to Richards Bay in northern Kwazulu-Natal (39-54).

World distribution: South African endemic.

Type locality: Kowie (Port Alfred) (Norris 1989).

Note: Norris (1989) reports some morphological differences between winter and summer forms of this species in Kwazulu-Natal: in winter vesicles are small (to 2 mm long) and lie in alternate unilateral rows along axes, while in summer vesicles are larger (to 5 mm long) and sparse on the axes.

Botryocladia pienaarii, re-drawn from Norris (1989: Fig 4).

Order Rhodymeniales

Family Rhodymeniaceae

Ceratodictyon variabile (Greville ex J. Agardh) RE Norris 1987: 243-245, figs 1-9, 12, 13

Plants dark red to blackish, erect, bushy, to about 10 cm tall, comprising slender, terete to slightly flattened, irregularly-branched uprights arising from prostrate stoloniferous system attached by small discs; uprights 350-500 µm in diameter. Structure pseudoparenchymatous; in transverse section inner medulla of narrow cells (about 20 µm diameter), outer medulla of slightly larger cells (about 35 µm diameter) grading into cortex of small cells (outer cells about 10 µm long). Tetrasporangia in sori in swollen distal tips of usually unbranched axes, cruciately divided, surrounded by small-celled interstitial filaments; cystocarps immersed in enlarged, warty branch tips.

Collections, ecology and regional distribution

Eulittoral fringe, Hluleka to northern Kwazulu-Natal(45-58) and East Africa.

World distribution: widespread in many temperate and tropical waters (as Gelidiopsis variabilis), including tropical West Africa (Guiry & Guiry, 2009).

Type locality: Madras, India (Silva et al. 1996).

Note: This species was formerly known as Gelidiopsis variabilis (J. Agardh) Schmitz, but Norris (1987) combined the genus Gelidiopsis with Ceratodictyon.

Ceratodictyon variabile, Hluleka (BOL).

Ceratodictyon variabile, re-drawn from Norris (1987) Fig. 1.

Ceratodictyon variabile, cross section of mature branch, re-drawn from Norris (1987) Fig. 3.

Order Rhodymeniales

Family Rhodymeniaceae

Erythrymenia obovata Schmitz ex Mazza 1921: 107-111

Plants erect, stiff, orange-red with striking blue-green or golden iridescence when fresh, comprising one to several thick, obovate blades tapering basally to a stipe and crustose holdfast; blades up to 10 cm (rarely to 20 cm) long, 1.5 to 5 cm (rarely to 7 cm) wide and from about 400 µm thick near the apex to about 700 µm thick near the base; margins spinose except when worn smooth, blade surfaces smooth in young plants, with stout spinose protrusions in older plants. Structure of a compact medulla initially of only large hyaline cells (80-125 µm in diameter) that subsequently become interspersed with smaller secondary cells; cortex of 3-5 layers of pigmented cells that become smaller towards the surface, to outer cells 2.5-5 µm in diameter. Plants dioecious; cystocarps immersed in both sides of blades; spermatangia in sori covering most of thalli; tetrasporangia about 60 µm long, cruciately to irregularly divided, arranged in nemathecia.

Collections, ecology and regional distribution

Recorded from Port Alfred to Shelly Beach in southern Kwazulu-Natal (39-49). Found in the sublittoral fringe on wave-exposed shores.

World distribution: South African endemic.

Type locality: Port Alfred (Sparling 1957).

Note: E. obovata is described in detail by Sparling (1957).

Erythrymenia obovata, Kenton-on-Sea.

Erythrymenia obovata, Kenton-on-Sea, spiny protrusions and golden iridescence on blade surface.

Order Rhodymeniales

Family Rhodymeniaceae

Rhodymenia Greville

Plants foliaceous, with small to prominent stipe, discoid or stoloniferous holdfast and flat, complanate blades with or without surface proliferations. Structure multiaxial, with small-celled cortex and larger-celled medulla. Reproductive structures scattered on blades, concentrated on blade apices, or restricted to surface proliferations. Tetrasporangia immersed in outer cortex, cruciately divided. Cystocarps projecting from surface, hemispherical and ostiolate.

Note that Womersley (1996) includes Epymenia in Rhodymenia, pointing out that the character formerly separating the genera (reproductive structures in surface leaflets in the former and in the blade surfaces in the latter) does not always hold.

Key to species

1a Blades irregularly branched, often cuneate to fan-shaped, arising from rhizomatous holdfast system

Rhodymenia
holmesii

1b Blades narrowly linear, dichotomously branched, with slender stipe arising from small discoid holdfast

2

2a. Reproductive structures borne on small leaflets/rosettes arising from surfaces of

Rhodymenia
capensis

2b Reproductive structures borne directly on the blades

Rhodymenia
natalensis

Order Rhodymeniales

Family Rhodymeniaceae

Rhodymenia capensis J. Agardh 1894: 58

Plants pale to bright red, 10-15 (-20) cm tall, with a terete stipe and flat, subdichotomously branched blade. Holdfast rhizomatous or a small disc; stipe a few mm to a few cm long, branched or unbranched; blades 1-4 mm wide, apices tapering, rounded or bilobed. Structure pseudoparenchymatous; medulla comprising several layers of larger cells (to about 60 µm diameter and 2-3 times as long) and cortex of 2-3 layers of small cells (about 5 µm diameter at surface). Reproductive structures borne on small proliferations on surfaces of blades; proliferations simple or tufted, round, wedge- or heart-shaped, up to 10 mm long; cystocarps along margins of proliferations; tetrasporangia in sori covering most of the surfaces.

Collections, ecology and regional distribution

Recorded from Olifantsbos on the west coast of the Cape Peninsula to the Port Alfred area (17-39). Found in the sublittoral fringe and subtidal zone to at least 25m depth. Specimens without reproductive proliferations are often impossible to distinguish from Rhodymenia natalensis, so that the distribution of these two species needs further study.

World distribution: South African endemic.

Type locality: Cape of Good Hope (Silva et al. 1996).

Note: This entity was previously known as Epymenia capensis. (see Womersley 1996). The species requires further study: south and east coast specimens usually have narrower blades (1-4 mm) than those from the west coast (5-8 mm) (see Stegenga et al. 1997). Womersley (1996) includes Epymenia in Rhodymenia, making the new combination Rhodymenia obtusa (Greville) Womersley, but he does not mention Epymenia capensis. We follow Guiry & Guiry (2009) in using R. capensis.

Rhodymenia capensis, Three Sisters (BOL).

Rhodymenia capensis, habit. Reproduced from Stegenga et al. (1997).

Rhodymenia capensis, cross section. Reproduced from Stegenga et al. (1997).

Order Rhodymeniales

Family Rhodymeniaceae

Rhodymenia holmesii Ardissone 1893: 682

Plants bright red, up to about 4 cm tall; extensive holdfast system of irregularly branched terete stolons bearing erect blades with short stipes; blades up to 3 cm long, narrowly cuneate at base, subdichotomously branched, often fan-shaped; blades from 1.5 to 5 mm wide, thickness from less than 100 µm at margins to more than 200 µm in centre. Structure of 3-4 layered medulla of large cells (to 60 µm diameter) surrounded by cortex of smaller cells; outer cortical cells 3-5 µm in diameter. No reproductive structures found.

Collections, ecology and regional distribution

Collected from the Cape Peninsula to Tsitiskamma National Park. (17-32).Found in the eulittoral fringe and sublittoral zone to at least 24m depth.

World distribution: also recorded from Namibia and Angola and Eastern Atlantic warm temperate waters (UK, Ireland, France, Portugal, Spain) and Atlantic islands (Azores, Canaries, Ascension) (Guiry & Guiry 2009).

Type locality : Sussex, England (Guiry & Guiry 2009).

Rhodymenia holmesii, Tsitsikamma (BOL). Scale divisions 1 mm.

Rhodymenia holmesii. 1. Habit. 2. Cross section. Reproduced from Stegenga et al. (1997).

Order Rhodymeniales

Family Rhodymeniaceae

Rhodymenia natalensis Kylin 1938:13-14, pl. 5, fig. 13

Plants erect, 8 (-12) cm tall, bright red, with terete stipes and dichotomously branched flat blades. Small discoid holdfast bears longish (10-12 mm) terete stipe, often with horizontal stolons giving rise to further upright axes; blades flat, 1-2 (-3) mm wide, repeatedly dichotomously branched, about 200 µm thick, apices rounded to tapering. Structure pseudoparenchymatous; in cross-section stipe with medulla of round cells 30-50 µm in diameter, cortex of radiating rows of smaller (4-5 µm) cells; blade with medulla of about 5 layers of rounded hyaline (clear) cells up to 80 µm diameter and cortex of 2-3 layers of much smaller cells (about 5 µm diameter at surface). Tetrasporangia cruciately divided, about 20-25 x 10-20 µm in size, in sori in cortex near apices of blades.

Collections, ecology and regional distribution

Found along the southern African coast from Namibia (Wynne 1986) to northern KwaZulu-Natal (1-58 in South Africa) and into southern Mozambique (Isaac 1956). Epilithic subtidally and in the sublittoral fringe, where it often forms patches on steep or overhanging rocks.

World distribution: southern African endemic.

Type locality: Isipingo, Kwazulu-Natal (Silva et al. 1996).

Note: This species is variable in size and shape, from plants a few cm long and 1mm wide to others 12 cm long and 2-3 mm wide. The similar entity identified from Namibia as R. linearis J.Ag. by Wynne (1986) and from Cape Hangklip on the South African west coast (Jackelman et al. 1991) is considered by Stegenga et al. (1997) to be R. natalensis.

Rhodymenia natalensis, Kenton-on-Sea.

Rhodymenia natalensis, Mntafufu, Transkei (BOL).

Rhodymenia natalensis, cross section.

Rhodymenia natalensis. 1. Habit. 2. Section of ‘stipe’. 3. Section of tetrasporangial sorus. Reproduced from Stegenga et al. (1997).

Order Rhodymeniales

Family Champiaceae

Thalli hollow, with cortex of several cell-layers lined internally by medulla of longitudinal filaments; central cavity partitioned by transverse cellular diaphragms. Carpogonial filament four-celled, carposporangia terminal. Tetrasporangia or polysporangia scattered in cortical layer, tetrasporangia tetrahedrally divided.

Champia Desveaux

Thalli hollow, terete or compressed, central cavity partitioned into segments by diaphragms of single cell layer. Structure multiaxial. Thallus walls one to several cell-layers thick, lined internally with longitudinally arranged medullary filaments which connect septa and bear gland cells. Tetrasporangia scattered in cortex, and especially in thin-walled species protruding into thallus cavity; tetrahedrally divided. Spermatangia excised from outer cortical cells in irregular superficial areas. Cystocarps semi-globose to urn-shaped, projecting from thallus surface and with conspicuous ostiole. Carposporophyte possessing central fusion, carposporangia terminal on branched gonimoblasts.

Key to species

1a. Plants a few to 10 cm tall, axes compressed, main axes up to 3 mm

Champia compressa

1b. Plants 1-4 cm tall, axes terete to slightly compressed, main axes
1-1.5 mm in diameter

Champia parvula

Order Rhodymeniales

Family Champiaceae

Champia compressa Harvey 1838: 402

Plants pale to bright red, usually with bright blue or green iridescence when fresh, creeping and branching to form clumps to a few cm tall (rarely as individual tufted plants growing to up to 10 cm tall); axes compressed segmented, pinnately to irregularly branched from margins, up to 3 mm wide and 0.5 mm thick; laterals may be alternate, sub-opposite or regularly bipinnate in arrangement, narrowing towards bases and apices. Thallus hollow with central cavity containing mucus, divided by septa at short intervals, axes slightly constricted at septa; thallus wall of single layer of almost isodiametric cells, 40-50 x 45-55 µm, interspersed with scattered much smaller cells at the outer surface; medulla with filaments (about 10 µm diameter) bearing spherical gland cells of about 15 µm diameter. Tetrasporangia excised from cells of thallus wall and projecting into central cavity, tetrahedrally divided, up to 100 µm diameter. Cystocarps sessile, about 2 x 1.6 mm when mature, conical to urn-shaped.

Collections, ecology and regional distribution

Recorded from the west coast near Cape Columbine, along the south coast and Kwazulu-Natal (11-58) and into East Africa. Common in turfs in pools and from eulittoral to at least 35 m depth.

World distribution: Indian Ocean shores and islands, Philippines, Korea, Australia and New Zealand (Guiry and Guiry 2009).

Type locality: Muizenberg, False Bay, South Africa (Silva et al. 1996).

Note: South coast specimens are often larger than those recorded on the west coast (Stegenga et al. 1997).

Champia compressa, Arniston, without iridescence.

Champia compressa, Kenton-on-Sea, with some iridescence.

Champia compressa, thallus detail.

Champia compressa, cross section showing outer layer.

Champia compressa. 1. Habit. 2. Thallus top, surface view. 3. Cross section. 4. Inside view of hollow thallus, showing septum (s) and tetrasporangia and gland cells (gl). Reproduced from Stegenga et al. (1997).

Order Rhodymeniales

Family Champiaceae

Champia parvula (C. Agardh) Harvey 1853: 76

Plants soft, red to brownish-red, 1-4 cm high, erect or creeping; axes segmented, terete to slightly compressed, main axes 1-1.5 mm diameter, branchlets 0.5 – 1 mm diameter, apices tapering, holdfast discoid. Thallus hollow, filled with mucus, divided by septa at regular intervals; thallus wall of cells of up to 40 µm diameter interspersed with scattered much smaller surface cells; inner wall with longitudinal filaments about 10 µm in diameter bearing occasional spherical to ovoid gland cells; surface cuticle 10-20 µm thick, clear. Tetrasporangia scattered, 60-120 µm diameter, tetrahedrally divided; cystocarps ovoid to urn-shaped, scattered, 0.7-1 mm in diameter, with pericarp and ostiole.

Collections, ecology and regional distribution

Recorded from the East London area through Kwazulu-Natal (41-58). Epilithic or epiphytic in the lower eulittoral to shallow sublittoral.

World distribution: apparently a cosmopolitan species (Guiry & Guiry 2009).

Type locality: Cadiz, Spain (Guiry & Guiry 2009).

Champia parvula on coralline seaweed, Port St Johns.

Champia parvula on coralline seaweed, Port St Johns.

Champia parvula under microcope (stained).

Order Rhodymeniales

Family Champiaceae

Chylocladia capensis Harvey 1847: 79

Plants dark red to brownish, sometimes iridescent, with more-or-less extensive holdfast and several erect axes, some curved and secondarily attached by small pads; erect axes from a few to (rarely) 20 cm tall (often smaller on the south coast), terete, regularly segmented; segments up to 5 x longer than broad in older parts, as long as broad in younger parts; branches (up to three orders) arising in whorls at constrictions in axes. Structure of axes hollow, with monostromatic septa at constrictions; thallus wall with 2-3 cortical layers comprising an epidermis of small (to about 35 µm diameter) almost isodiametric cells and inner 1-2 layers of loose, axially elongated cells; inner medullary later of thin filaments (10-20 µm diameter) bearing scattered small gland cells.

Polysporangia scattered, immersed in cortex, globose, up to 100 µm diameter, usually divided into 8 spores; cystocarps globose, without ostiole, projecting from thallus surface.

Collections, ecology and regional distribution

Found along the west coast and south coast as far as the Port St Johns area. (1-46). Epilithic in lower eulittoral and eulittoral fringe or in rock pools.

World distribution: South African endemic.

Type locality: Table Bay, Cape Province, South Africa (Silva et al. 1996).

Note: Stegenga et al. (1997) point out that this entity includes the South African record of Gastroclonium reflexum (Chauvin) Kützing recorded from Bird island, Algoa Bay in Anderson and Stegenga (1989).

Chylocladia capensis in compact turf, Keurboomstrand.

Chylocladia capensis in rock pool, Arniston.

Chylocladia capensis,  part of individual thallus.

Chylocladia capensis. 1. Habit. 2. Habit of small female plant with cystocarps. 3. Longitudinal section through thallus wall, showing polysporangia and gland cells. Reproduced from Stegenga et al. (1997).

Order Rhodymeniales

Family Hymenocladiaceae

Hymenocladia kallymenioides (Holmes) F. Schmitz 1896: 8-13

Plants dark- to brownish-red, comprising broad, flat, often incised blade to 30 cm long, with short stipe (0.5-1.0 cm) and discoid holdfast. Blade flat, simple, becoming irregularly palmately incised and sometimes perforated, midrib and veins absent, margins entire or toothed; blades usually with scattered dark patches and striae, and often with light patches. Stucture of large clear medullary cells and very small outer cortical cells; clear gland cells visible in inner cortex. Medulla of 1-3 layers of large (to 150 µm diameter) cells, interspersed with smaller secondary cells in older thallus parts; cortex of several layers (2-3 or more in older parts) of smaller pigmented cells becoming very small (abut 5-8 µm long) towards surface. Tetrasporangia about 60 µm long (visible with naked eye), submerged in cortex and extending into medulla, tetrahedrally divided. Cystocarps visible to naked eye as pustule-like swellings (up to1 mm diameter), scattered on one surface of blade; developing between medulla and cortex, with carpostome; carpospores about 25 µm in diameter. Male structures not seen.

Collections, ecology and regional distribution

Found from Addo to Mkambati (38-47) in the sublittoral fringe and below.

World distribution: Also recorded from Madagascar (Silva et al. 1996).

Type locality: Mouth of Kowie River, Port Alfred (Silva et al. 1996).

Note: Details of the structure and reproduction in this species are provided by Sparling (1957). She notes that the dark patches in the thalli are caused by pigmentation in some of the cortical cells, rather than by separate (e.g. reproductive) structures.

Hymenocladia kallymenioides, Kowie (BOL).

Hymenocladia kallymenioides, Kowie (BOL). Detail of blade showing dark spots and striae.

Hymenocladia kallymenioides, XS of young part of blade (re-drawn from Sparling (1957; Fig. 15 e).

Hymenocladia kallymenioides, section of blade with tetrasporangia (re-drawn from Sparling (1957: Fig 15 a).

Hymenocladia kallymenioides, fresh, smallish specimens.

Order Rhodymeniales

Family Lomentariaceae

Lomentaria Lyngbye

Thalli hollow, anatomy of pseudoparenchymatous cortex lined internally with filamentous medulla. Septa, where present, cellular, partitioning the central cavity and dividing axes into segments. Tetrasporangia tetrahedrally divided, arranged in sori sunken in cavities in the cortex. Carpogonial filaments 3-celled, almost all cells of the carposporophyte developing in to carposporangia.

Key to the species

1a. Axes divided into segments by regular constrictions; axes with hooked tips

Lomentaria amplexans

1b. Axes without regular constrictions and without hooked tips

2

2a. Mature axes up to 750 µm in diameter

Lomentaria corallicola

2b. Mature axes 1-1.5 mm in diameter

Lomentaria diffusa

Order Rhodymeniales

Family Lomentariaceae

Lomentaria amplexans RE Norris 1987: 36, figs 1-4

Plants dark red, up to 5 cm long, cylindrical, mostly prostrate, epiphytic, with constricted segments, erect branches usually with slightly swollen hooked tips; hooked tips tending to wrap around host branches and produce lateral branches of similar dimensions to main axes that closely encircle the host and attach by rhizoidal discs. Segments hollow, up to 2 mm long and 750 µm wide, separated by septa comprising several layers of cells that are ovate to irregular in shape and up to 40 µm diameter; cortex of 2-3 layers of cells, cells of inner layer angular to ovate, up to 40 µm long; outer cortical cells ovate, variable in size, up to 25 µm long; medulla of longitudinally arranged filamentous cells of about 8 µm diameter, often with laterally-attached gland cells. Tetrasporangia in sunken sori, mainly proximal on axes, tetrahedrally divided, up to 45 µm in diameter. Male and female plants not found.

Collections, ecology and regional distribution

Recorded from Mkambati to northern Kwazulu-Natal. (47-58). Found from the lower eulittoral to shallow sublittoral, epiphytic on larger algae (often Amphiroa ephedraea or Gelidium spp.), the branches of which may be bound together by the hooked axes.

World distribution: South African endemic.

Type locality: Rocky Bay, Kwazulu-Natal (Norris 1987).

Note: See Norris (1987) for further details and illustrations.

Lomentaria amplexans, habit on left with arrow showing hooked axis binding thallus to an articulated coralline seaweed. Lower right, XS of thallus through tetrasporangial sorus (re-drawn from Norris, 1987 Figs 1 and 3).

Order Rhodymeniales

Family Lomentariaceae

Lomentaria corallicola Børgesen 1939: 113-116, figs 30-32

Plants red, prostrate but with erect branches to 7 mm tall, about 750 µm in diameter, epiphytic in seaweed turfs, axes cylindrical, constricted only at their bases. Septa present only at point of origin of major segments; cortex of 2 layers of cells; outer cortical cells up to 45 µm long x 15 µm wide, sparsely interspersed with smaller cells; inner cortical cells up to 90 x 30 µm; medulla of filamentous cells 75-100 µm long x 10 µm in diameter, lining central cavity and bearing small (5 µm) gland cells. Tetrasporangial sori immersed in thallus cavity, with small pore, tetrasporangia tetrahedrally divided, up to 40 µm in diameter. Cystocarps urn-shaped, about 500 µm in diameter, borne externally and laterally, usually on upright branches.

Collections, ecology and regional distribution

Recorded from Tsitsikamma to northern Kwazulu-Natal (32-58). Epiphytic in the sublittoral fringe and probably below.

World distribution: Seychelles, Iran, Western Australia, some Pacific islands and Colombia (Guiry & Guiry 2009).

Type locality: Iran (Børgesen 1939).

Lomentaria corallicola, habit (re-drawn from Norris, 1987 Fig. 5).

Lomentaria corallicola, habit , XS of thallus through tetrasporangial sorus (re-drawn from Norris, 1987 Fig. 8).

Lomentaria corallicola, cystocarp (specimen from northern KwaZulu-Natal).

Order Rhodymeniales

Family Lomentariaceae

Lomentaria diffusa Stegenga, Bolton & Anderson 1997: 373-374, plate 138

Plants bright red, forming dense turfy clumps, up to several cm in height and diameter, consisting of interwoven branching prostrate and erect axes. Axes terete, hollow, constrictions and septa present only at bases of lateral branches; branching irregular or in well-developed erect axes with lateral in double row on outside of the curved main axis; axes to 1.5 mm in diameter. Structure of axis walls comprising about 5 layers of pseudoparenchymatous cells, cells of outer layer smallest, becoming progressively larger to the inside; medulla of a network of filamentous cells, some bearing spherical to ovate gland cells. Tetrasporangia in sori lining deep depressions in thallus wall, projecting into central cavity, globose to slightly ovate, up to about 70 µm diameter, tetrahedrally divided. Cystocarps ovate to urn-shaped, projecting from thallus surface, ostiolate.

Collections, ecology and regional distribution

Recorded from Langebaan Lagoon on the west coast to Tsitsikamma (12-32). Epilithic or epiphytic in the sublittoral fringe.

World distribution: South African endemic.

Type locality: Kraalbaai, Langebaaan Lagoon (Stegenga et al. 1997).

Lomentaria diffusa, 1. Habit, with cystocarps. 2. Tetrasporiferous branch. Reproduced from Stegenga et al. (1997).

Lomentaria diffusa, 1. Cross section. 2. Surface view. 3. Tetrasporangial sorus in cross section. Reproduced from Stegenga et al. (1997).

REFERENCES

References to South Coast Rhodophyta - Book 4

Agardh, J. G. 1842. Algae maris Mediterranei et Adriatici, observations in diagnosin specierum et dispositionem generum. Paris, 164 pp.

Agardh, J.G. 1849. Algologiska bidrag. Öfversigt af Kongl. Vetenskaps-Adademiens Förhandlingar, Stockholm 6: 79-89.

Agardh, J.G. 1851. Species genera et ordines algarum, seu descriptiones succinctae specierum, generum et ordinum, quibus algarum regnum constituitur. Volumen secundum: algas florideas complectens. Part 1. pp. [i]-xii, [1]-336 + 337-351 [Addenda and Indices]. Lundae [Lund]: C.W.K. Gleerup.

Agardh, J. 1851. Species genera et ordines Floridearum 2(1). Lund 1-351.

Agardh, J. 1894. Analecta Algologica. Cont. II. Lunds Universitetets Arsskrift, Afd. 2, 5 (7); 1-98, 1 pl.

Anderson, R. J., Anderson, D. R. & Anderson, J.S. 2008. Survival of sand-burial by seaweeds with crustose bases or life-history stages structures the biotic community on an intertidal rocky shore. Botanica Marina 51: 10-20.

Anderson, RJ. & Bolton, J.J.1990. Reproductive morphology and life histories of southern African Gymnogongrus species (Rhodophyta, Phyllophoraceae). British Phycological Journal 25: 381-390.

Anderson, R.J. & Stegenga, H. 1989. Subtidal algal communities at Bird Island, Eastern Cape, South Africa. Botanica Marina 32: 299-311.

Ardissone, F. 1893. Nota alla phycologia Mediterranea. Rendiconti del reale Instituto Lombardo di scienze e letter, Serie II , 26: 674-690.

Ardré, F. 1980. Observations sur le cycle de développement du Schizymenia dubyi (Rhodophycee, Gigartinales) en culture, et remarques sur certains genres de Némastomatacées. Cryptogamie: Algologie, 1 (2): 111-140.

Barton, E.S. 1896. Cape Algae. Journal of Botany, London, 35: 193-198, 458-461, 5 figs.

Bélanger, C. & Bory de Saint-Vincent, J.B.G.M. 1834. Voyage aux Indes-Orientales. Botanique 2. Cryptogamie. Paris, 192 pp., 16 pls.

Børgesen F. 1939. Marine algae from the Iranian Gulf especially from the innermost part near Bushire and the Island Kharg. Danish Scientific Investigations in Iran, Part 1, pp. 47-141, Copenhagen.

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GLOSSARY

Bisexual:

having both sexes.

Blade:

a flattened, leaf-like part of the thallus.

Calcar:

spur subtending a cystocarp.

Carpogonium (pl. carpogonia):

the female sex structure in red algae.

Carpospore:

a spore, usually diploid, produced in the carpogonium. In red algae, it grows into a free-living tetrasporophyte.

Cerata:

dorsal appendages on some opisthobranch mollusks (sea slugs) that contain diverticula of the digestive gland. Serve as gills.

Chloroplast:

a cell organelle specialized for photosynthesis, the production of carbohydrates from carbon dioxide, water in the presence of light energy.

Coenocytic:

referring to a multinucleate cell, the coenocyte, derived when the nucleus divides but the cytoplasm does not.

Conceptacle:

a fertile cavity, often with a pore.

Cortex:

tissue external to the medulla, often an outer covering, of a blade.

Cryptostomata:

minute cavities in the outer cortex of Fucales containing hairs.

Cystocarp:

a structure in the female gametophyte of red algae consisting of diploid carposporophyte surrounded by gametophytic tissue. It may be a pigmented bump with or without an ostiole.

Dichotomous:

branching in pairs.

Dioecious:

bearing male and female reproductive structures on separate individuals.

Diploid:

having a normal (double or 2n) set of chromosomes.

Discoid:

disk shaped. Usually referring to the shape of holdfasts.

Distichous:

arranged in two rows on opposite sides of an axis.

Distromatic:

having two layers of cells.

Gametangium (pl. gametangia):

a structure that produces gametes.

Euryhaline:

able to withstand a wide range of salinities.

Eurythermal:

able to withstand a wide range of temperatures.

Filamentous:

a row of cells arranged end to end forming a hair-like thallus.

Gamete:

a sexual cell, male or female (e.g., sperm and eggs).

Gametophyte:

the haploid and sexual phase in the life history of an alga; produces gametes.

Haploid:

having a single set of chromosomes - half the normal number (1n).

Herbivory:

feeding on plants.

Holdfast:

the structure that attaches the alga to its substratum.

Intercallary:

occurring anywhere along the thallus except at the apex.

Intergeniculum (pl. intergenticula): a

calcified segment between the uncalcified joints of coralline algae.

Isomorphic:

having the same shape; looking alike.

Lanceolate:

shaped like a lance; several times longer than wide.

Laminarin:

a polysaccharide carbohydrate energy storage product in Laminaria.

Meristem:

a region of rapidly dividing, undifferentiated cells; an area of growth.

Midrib:

the middle rib of a blade.

Monoecious:

bearing male and female reproductive structures on the same individual.

Monostromatic:

having one layer of cells.

Oogonium:

a female reproductive cell (gametangium) that produces one or more female gametes (eggs).

Ostiole:

a pore.

Ovate:

egg-shaped.

Papilla (pl. papillae):

small rounded or pimple-like projections on a surface

Pericentral:

surrounding central axial cells.

Pinnate:

having similar parts arrange on opposite sides of an axis, like a feather.

Plastid:

a pigmented cell organelle (e.g., a chloroplast which contains chlorophyll).

Pneumatocyst:

a gas bladder or float of brown algae.

Receptacle:

the inflated tip of a branch that holds conceptacles (e.g. the swollen tip of Fucus).

Rhizoid:

a unicellular or filamentous structure for attachment.

Rhizome:

a horizontal, shoot-producing stem growing within the substrate.

Saxicolus:

growing on rocks.

Spermatangium (pl. spermatangia):

a cell in red algae that produces a single, nonflagellated, male gamete (the spermatium).

Spore:

a motile or nonmotile, asexual reproductive structure.

Sporophyll:

the part of the alga, usually a blade, which bears sporangia, the structure that produces spores.

Sporophyte:

the diploid phase in the life history of the alga; produces spores.

Sorus (pl. sori):

a cluster of reproductive structures.

Stipe:

the stem-like part of the thallus between the holdfast and the blade(s).

Stolon:

a runner growing out of the base of the alga capable of producing offshoots.

Substratum:

the material (such as rock) on which the alga is growing (substrate).

Tetraspore:

a spore, grouped in fours, produced by tetrasporophytes

Tetrasporophyte:

a phase in the life history of red algae that produces tetraspores.

Tertiary:

referring to a third stage.

Thallus:

the body of the alga.

Uniseriate:

having a single row of cells.

Utricle:

the inflated part of Codium that contains plastids.

Vegetative:

not associated with reproductive cells or sexual reproductive capacity.

Zygote:

a fertilized egg cell.

INDEX TO GENERA AND SPECIES

FAMILY KALLYMENIACEAE

Callophyllis variegata

Pugetia harveyana

Pugetia porphyroidea

Thamnophyllis pocockiae

FAMILY GYMNOPHLOEACEAE

Schizymenia apoda

FAMILY PHACELOCARPACEAE

Phacelocarpus oligacanthus

Phacelocarpus tortuosus

FAMILY SOLIERIACEAE

Callophycus densus

Callophycus tridentifer

Rhabdonia natalensis

Erythroclonium corallinum

FAMILY CAULACANTHACEAE

Catenella nipae

Caulacanthus ustulatus

Heringia mirabilis

FAMILY CYSTOCLONIACEAE

Calliblepharis fimbriata

Hypnea arenaria

Hypnea intricata

Hypnea musciformis

Hypnea rosea

Hypnea spicifera

Hypnea tenuis

Hypnea viridis

Rhodophyllis reptans

FAMILY RHIZOPHYLLIDACEAE

Portieria hornemannii

Portieria tripinnata

FAMILY PHYLLOPHORACEAE

Gymnogongrus glomeratus

Gymnogongrus polycladus

Gymnogongrus tetrasporifer

FAMILY GIGARTINACEAE

Chondracanthus acicularis

Gigartina insignis

Gigartina minima

Gigartina paxillata

Gigartina pistillata

Gigartina polycarpa

FAMILY PLOCAMIACEAE

family_plocamiaceae

Plocamiocolax papenfussiana

Plocamium beckeri

Plocamium corallorhiza

Plocamium cornutum

Plocamium glomeratum

Plocamium maxillosum

Plocamium rigidum

Plocamium suhrii

Plocamium telfairiae

FAMILY SARCODIACEAE

Sarcodia dentata

Trematocarpus flabellatus

FAMILY RHODYMENIACEAE

Botryocladia beckeriana

Botryocladia madagascariensis

Botryocladia pienaarii

Ceratodictyon variabile

Erythrymenia obovata

Rhodymenia capensis

Rhodymenia holmesii

Rhodymenia natalensis

FAMILY CHAMPIACEAE

Champia compressa

Champia parvula

Chylocladia capensis

FAMILY HYMENOCLADIACEAE

Hymenocladia kallymenioides

FAMILY LOMENTARIACEAE

Lomentaria amplexans

Lomentaria corallicola

Lomentaria diffusa

The Authors

From left: Rob Anderson, Herre Stegenga and John Bolton.

Brief biographies

Rob Anderson completed a PhD at the University of Cape Town in 1982, just after joining the Seaweed Unit in the SA Government fisheries department. Although much of his work involved commercial seaweed resources, he maintained a strong interest and has published in many aspects of seaweed biology and systematics. He has collaborated extensively with John Bolton, Herre Stegenga and others. Recently retired, he continues to surf, dive and enjoy the ocean.

Herre Stegenga completed his Ph D ("doctoraat") in 1979 at the Free University of Amsterdam. He published a Flora of the Dutch Seaweeds in 1983 (with Ivo Mol) and spent the next ten years mainly in South Africa (University of Cape Town, Rhodes University in Grahamstown), resulting in the publication of the Seaweeds of the South African West Coast (Stegenga, Bolton & Anderson 1997). After 1992 he revisited the RSA several times to collect material from the South Coast. Retired in 2012, he is still active in western European seaweed identification.

John Bolton is Emeritus Professor and Senior Research Scholar at the University of Cape Town, where he taught all aspects of phycology (the study of algae) for 35 years. His research interests include the diversity and biogeography of seaweeds, seaweed use and aquaculture, and the integrated aquaculture of seaweeds with marine animals. He is a former President of the International Phycological Society, and was awarded the Gilchrist Memorial Medal in 2017 in recognition of his contribution to South African marine science.

Acknowledgments

We are deeply indebted to Professor Michael Guiry and his colleagues who built and continue to maintain the incomparable algal website AlgaeBase.org – without it we would never have attempted this guide.

Chris and Derek
Our heartfelt thanks to Chris Boothroyd and
Derek Kemp for their enthusiastic and invaluable
technical help throughout the project.

We thank our colleagues and post-graduate students whose knowledge has been of enormous help, especially, Olivier De Clerck, Frederik Leliaert, Heroen Verbruggen and Eric Coppejans (Ghent University, Belgium), Enrico Tronchin (UCT) and others.

This work was funded from several sources over many years. FRD/NRF Post-doctoral bursaries sustained Herre Stegenga at Rhodes University and at UCT when he made many collections of specimens and microscope slides. DAFF (Department of Agriculture, Forestry and Fisheries) supported the participation of Anderson, Kemp and Boothroyd. SANPAD (South Africa Netherlands research Programme on Alternative Development) grants allowed us to train students and improve our collections from the south coast between 2001 and 2004. Various grants from NRF, UCT and DAFF all contributed through various projects. Most importantly, NRF funding of the SeaKeys Programme supported the development of this website. SANPARKS allowed us access to marine parks where important records were found. Finally, we thank the curators of the BOL and GRA herbaria for access to collections.

Colophon

Published by Seaweed Research Unit, Department of Biological Sciences, University of Cape Town and the Department of Agriculture, Forestry and Fisheries of South Africa.
Cape Town, South Africa

Typesetting: Seaweed Research Unit, Department of Biological Sciences, University of Cape Town, Rondebosch 7701, South Africa.