is oe me see i “ tue . a ean? Tat =) Ses Ne ee ie a - ; = o9 - Pee ee he ees q ic Sins - ~Aoey LA Ree "ali iz Digitized by the Internet Archive in 2016 with funding from South African National Biodiversity Institute Libraries https ://archive.org/details/journalofsouthaf1i9unse | VOLUME XIX. LIBRARY | BIVIBIGN OF BOTANY @me : PLANT PATHSLSCY ‘ § Catulegue No...... Sakesess il 3% i -12-4959 SiSLIOTEEK Zr ihe ae ENT ICURBE : 1953. a a a rae TRC re et oeeereonecor THE JOURNAL OF SOUTH AFRICAN BOTANY PUBLISHED UNDER THE AUTHORITY OF THE TRUSTEES OF THE NATIONAL BOTANIC GARDENS OF SOUTH AFRICA KIRSTENBOSCH, NEWLANDS CAPE PROVINCE EDITOR R. H. COMPTON, M.A. (Cantab.), F.R.S.S.Af., Hon. F.R.H.S. ACTING DIRECTOR OF THE NATIONAL BOTANIC GARDENS, PROFESSOR EMERITUS IN THE UNIVERSITY OF CAPE TOWN. THE JOURNAL OF SOUTH AFRICAN BOTANY. VotumME XIX, 1953. CONTENTS. PAGE Four New ALOEs FRoM Kenya Cotony. By Dr. G. W. Reynolds, Hon. D.Sc. (Cape Town), F.L.S. (With Plates I-VII) .. ee ue Bi : 1 Tue IpeENTITY OF ALOE SECUNDIFLORA ENGLER. By Dr. G. W. Reynolds. (With Plates VITI-X) .. : : 5 : 13 A New ALOE FROM SOMALILAND PRrotecTorRATE. By Dr. G. W. Reynolds (With Plate IX) : 5 he a 21 Tue IpeENnTITY OF ALOE JOHNSTONH Bak.: A New Synonym. By Dr. G. W. Reynolds. (With Plates XII-XIIT) .. an ; 25 A Note on Sex in RoyeNna GuaBra L. (EBENACEAE). By Capt. T. M. Salter, R.N. (Retd.) Be ae a ss 29 KaRYOLOGICAL STUDIES AND CHROMOSOME NUMBERS IN HYPARRHENIA AUCTA anp H. nirta. By Dr. Stefan Krupko, Ph.D. (Warsaw). (With Plates XIV-XV) of #: EF: ay oe ey. te a 31 South AFRicaN SEAWEED VEGETATION AND FuTURE INVESTIGATIONS IN THIs FieLp. By Professor W. Edwyn Isaac, B.Sc.. Ph.D., F.R.S.S.Af. -. 59 Tar MorpHoLocy, GEoGRAPHICAL DIsTRIBUTION AND EcoLtocy oF HyPNEA SPICIFERA (SUHR) Harv. By Professor W. Edwyn Isaac and Miss Florence Hewitt. (With Plates XVI-XVII) a st: as 73 SEAWEED Resources oF SoutH Arrica. By Professor W. Edwyn Isaac and C.J. Molteno .. be a of. isis ae ve ov: te 85 On THE History oF PLanT STUDY UPON THE WITWATERSRAND. By Dr. H. B. Gilliland, D.Se. .. be ae 93 Puantar Novak ArricaNak. SeRtES XXXI. By Professor R. H. Compton, M.A., F.R.S.S.Af. (With Plate XVIII) a a oe ~ ae LOS A New Svuccunent EvuPHORBIA FROM THE WESTERN CAPE Province. By ; Dr. R. A. Dyer, M.Sc., D.Sc., F.R.S.S.Af. (With Plate XIX) .. .. 135 TuREE SmatL Spectus or OrHonna. By Professor R. H. Compton. (With re Plate XX) A New Crassuna FROM THE RicuTERSvELD. By H. Herre. (With Plate XXI) 146 SIIcULARIA. A NEW GENUS OF CRUCIFERAE. WITH NOTES ON RELATED GENERA. By Professor R. H. Compton NoTEs oN NOMENCLATURE IN Licutrootis. By Professor R. 8. Adamson. M.A.. D.Sc., F.B.S.S.Af. ALTEN PLANTS GROWING WITHOUT CULTIVATION IN THE SOMERSET WEST NEIGHBOURHOOD. By R. N. Parker ADDENDA ET CORRIGENDA TO THE Woopy PLANTS OF THE BECHUANALAND Protectorate. By O. B. Miller. L.S.0.. F.LS. REVIEWS: L. E. W. Codd. Trees and Shrubs of the Krger National Park. lees R. H. Com pton.) H. N. and A. L. Moldenke, Plants of ‘he Bible. (Pusan Lig tile ¢ om io C. yan Dillwijn, Botany of Sugar Cane. (Professor R. H. Compton) J.J. Finan, Maize in the Great Herbals. (Professor R. H. Compton) R. Story, A Botanical Survey of the Keiskammahoek District. ee I? 8. Adamson) 42 E. M. van Zinderen Bakker, South African Pollen Gee and Spores, Part J. (Mr. 8. Garside) . R. Good, The Geoeree Ly of the Flowering Plants: aa I. (Professor Ro: Compton) W. Crocker and L. V. Barton. Physiology of seas (Profesor W. Baym Isaac) 2A L. M. Cranwell. New Zealand Pollen Studies. The Monomegieiode ot r. S. Garside) .. ; ahs IspEx oF Prant Names, Vou. XIX PAGE 147 ISSUED FEB 18 19653 JOURNAL OF SOUTH AFRICAN BOTANY VOL. XIX. FOUR NEW ALOES FROM KENYA COLONY By G. W. REYNOLDS. (With Plates I—VII.) With the passing of Mr. H. Basil Christian, who contributed so much to the advancement of knowledge of the Aloes of Tropical Africa, and with the Aloes of South Africa having been monographed, it is now my hopeful ambition to investigate and eventually to monograph, the Aloes of Tropical Africa. The descriptions of most of the earliest described species of Aloe from East Africa are incomplete, vague, and sometimes misleading, while there are no figures of most of them. Some type material in Berlin was, unfortunately, destroyed during the war, while such type material as does exist today, is mostly incomplete, fragmentary, and more exasperating than helpful. In some cases, leaves and habit of growth were unknown. It soon became clear, that nothing less than a personal visit to the actual type localities, would ever enable me to solve the problem of the identity of most of these early Aloes—or at least to attempt it. Consequently, at the invitation of Mr. P. R. O. Bally, Botanist at the Coryndon Museum, Nairobi, I flew to Nairobi in March 1952, and together we travelled 4,300 miles in six weeks, visiting all the Aloe type localities in Northern Tanganyika, i.e. Kilimanjaro, the Northern and Southern Pare Mountains, the Western and Kastern Usambara Moun- tains, and several in Kenya and Uganda. In Kenya we travelled from Mombasa to Witu beyond the Tana River, returned to Nairobi, then went through the Rift Valley to Nakuru, Thomson’s Falls, Nanyuki and beyond Isiolo, thence westwards via Tororo and Jinja to Entebbe in Uganda, where A. dawei Berger, the last of the early species needed to complete the investigations, was found. 1 bo The Journal of South African Botany. These journeys were highly successful, and all but two of the earliest described Aloes in the area covered. can now be identified with reasonable certainty, while several new species of Aloe were found. Many photo- graphs were taken, much herbarium material gathered, and copious field notes written up, almost all of which have been sent to the National Herbarium, Pretoria, and the Royal Botanic Gardens, Kew. I am grateful to Mr. Bally for his invaluable help, to Dr. L. 8S. Leakey, Curator of the Coryndon Museum, Nairobi, and to Mr. P. Greenway, O.B.E., for much assistance. Papers establishing the identity of some of the earliest described Aloes are being prepared for publication, while several new species await description, of which four are described hereunder. B. J. Aloe ballyi Reynolds. Spec. nov. Nulli alii arcte affinis; forsitan sectionem propriam constituere debet. Planta succulenta; truncus simplex, glaber, gracilis, aetate 6—8 met. altus, 1O—15 cm. crass. Folia ca. 25, dense rosulata, usque ad 90cm. longa, basi 14cm. lata, longe-attenuata, profunde canaliculata, griseo- viridula, glabra, immaculata, dentibus marginibus albidis 4—5mm. longis, 10—l5mm. distantibus armata. Inflorescentia multi-ramosa, ca. 60cm. longa et lata. Racemi ca. 14cm. longi, subsecundi, sublaxe ca. 20-floribus. Bracteae ovato-acutae, 5mm. longae, 5mm. latae, scariosae, sordide-albidae. ca. 5-nervatae. Pedicelli 10mm. longi. Perigonium carminatum, cylindricum, 33mm. longum, basi 9mm. diam.; segmenta exteriora per 22mm. libera. Antherae 4—5mm. exsertae. Stigma demum 5mm. exsertae. Ovarium 6mm. longum, 4mm. diam. [Plates I, I1.]} Hab. Kenya Colony: Indense bush 23 miles west of Mwatate, 20 miles West of Voi (approx. 3°30’S 38°23’E), alt. 3,000 ft., fl. 31 March 1952, Reynolds 6378! (type) in Nat. Herb. Pretoria, with type number in Herb. Kew, and Cor. Mus. Herb. Nairobi. Tanganyika Territory: S. Pare Mtns., Mamba Track, 17 June 1942, Greenway 6470! in E.A. Herb. Nairobi, and Nat. Herb. Pretoria; S. Pare Mtns., Kisiwani, 2 Feb. 1936, P. J. Greenway 4573! in Cor. Mus. Herb. Nairobi, “whole plant smells strongly of mice’. Our new species is named after Mr. P. R. O. Bally, Botanist, Coryn- don Museum, Nairobi, who first found plants near Makania on the western side of the S. Pare Mtns. in September 1934, and again in December of that year near the top of the Manyara Escarpment beyond Lake Manyara and Mtuwambu on the road westwards from Arusha via Makuyuni to Ngorongoro Crater, in Tanganyika Territory. He has also Four New Aloes from Kenya Colony. 3 photographed plants on the Nguruman Escarpment near Oldonyo Sambu, north of Mount Meru, in Tanganyika Territory. The writer found it near Kisiwani, at the northern end of the S. Pare Mtns. in dense bush and in almost impenetrable thickets of Sansevieria and Euphorbia, growing socially with A. deserti Berger, Kisiwani being the type locality of the latter. Mr. P. J. Greenway records that A. ballyi is “locally common all down the eastern side of the S. Pare Mtns. on the foot slopes between 4,000—5,000 ft.” A. ballyi is a very distinctive species with a tall, slender, simple stem reaching 20 feet and more in height. The leaves are deeply channelled and recurved, while old dried leaves fall, giving the stem a clean smooth appearance. The leaves and sap give off a permeating, very decided odour of rats or mice, on which account this species has for some time been known as the “Rat Aloe’’. The inflorescence is produced sub-obliquely, with the reddish-orange flowers more or less secund, depending on the angle of the raceme; the more horizontal the raceme, the more secund the flowers. On Mr. H. A. Delap’s Estate “Kayata”’, about 50 miles east of Nairobi, and east of Donyo Sabuk in dense bush near the Athi River, the writer noticed many tall simple-stemmed plants, and also several which branched from the base and formed large shrubs; but this is a form not typical of the species as a whole. The flowering period appears to be from December to June, depending on the rains, and the locality. ; Description: Stem slender, simple, 5—8 met. high, 10—l5em. thick, glabrous, the old leaves not persistent. Leaves about 25, densely rosulate at apex of stem, lanceolate-long attenuate, up to 90cm. long, 14cm. broad at base, the youngest leaves spreading, the oldest much recurved, deeply canaliculate, and “U” shaped in cross-section; both surfaces grey-green, glabrous throughout, without spots or lineation; margins armed with laterally compressed deltoid, pungent, white teeth averaging 4—5mm. long, 10—l5mm. distant, more crowded nearer base of leaf, more distant upwards, the teeth sometimes forward-uncinnate, the interspaces straight. Inflorescence a much branched panicle, about 60cm. long and broad, produced sub-obliquely. Peduncle biconvex and 20mm. diam. at base, terete upwards, with about 20 branches from low down, the lowest re-branched, and sub- tended at base by a thin, dry, white bract about 18mm. broad, 14mm. long, about 7-nerved, the branches mostly spreading with the racemes oblique. + The Journal of South African Botany. Racemes up to 14cm. long, rather laxly about 20-flowered, the buds dull scarlet, tipped bluish-green, with a slight bloom, the open flowers carmine to reddish-orange, the pedicels upturned with the buds and flowers mostly subsecund to secund, depending on the degree of obliquity of the racemes. ; Bracts clasping the pedicels, ovate-acute, 5mm. long, 5mm. broad at base, thin, scarious, dirty white, about 5-nerved. Pedicels lowest 10mm. long, mostly up-curved. Perianth carmine to reddish-orange, greyish-tipped, 33mm. long, cylindric, 9mm. diam. near base, exceedingly slightly constricted on under-side above the ovary, very slightly curved, the mouth wide open; outer segments free for two-thirds their length (tube 1lmm.), distinctly 3-nerved from base to apex, the nerves dull pale reddish, the apices subacute, slightly spreading: inner segments themselves free but dorsally adnate to the outer for one-third their length, broader than the outer, with their brownish-tipped apices more obtuse and more spreading than the outer, and with a prominent median nerve throughout. Filaments pale lemon, filiform-flattened. the 3 inner narrower and lengthening in advance of the 3 outer. with their anthers in turn exserted 4—_ 5mm. Style pale yellow, filiform, with the stigma at length exserted 5mm. Ovary pale green. 6mm. long, 4mm. diam., finely 6-grooved. Capsule for some time enwrapped with the dried remains of the perianth: mature fruits and seeds not seen. Aloe kedongensis Reynolds. Spec. nov. in Sect. EvAaLor, subsect. Prolongatae. Planta succulenta, fruticosa. Caules graciles usque ad 4 met. longi, 4cm. crass. Folia ex vaginis striatulis lineari-lanceolata, sensim attenuata, 30cm. longa et 3.5em. lata, patula vel recurvula, immaculata, canali- culata, dentibus marginibus 2—3mm. longis, 10mm. distantibus armata. Inflorescentia 50cm. alta, 2—4-ramosa. Racemi 10—12cm. longi, ca. 8cem. lati. Bracteae ovato-acutae, 5mm. longae et latae, 3-nervatae. Pedicelli 20—25mm. longi. Perigonium coccineum, 35mm. longum; segmenta exteriora per l4mm. libera. Antherae 2—3mm. exsertae. Stigma demum 4mm. exserta. Ovarium 7mm. longum, 3mm. diam. [Plates IIT, IV.] Hab. Kenya Colony: Rift Valley, near Lake Naivasha, 2 miles south of Lake Hotel. 6 miles south of Naivasha, approx. 0°48’S 36°25’E, alt. 6,100 ft., fi. 19 April 1952, Reynolds 6546! (type) in Nat. Herb. Pretoria, with type number in Herb. Kew, and Cor. Mus. Herb. Nairobi: Gilgil Four New Aloes from Kenya Colony. 5 Escarpment, 3 miles north-west of Gilgil, fl. 19 April 1952, Reynolds 6552! in Nat. Herb. Pretoria, Herb. Kew, and Cor. Mus. Herb. Nairobi; near top of Kedong Escarpment, 24 miles north-west of Nairobi, fl. 19 April 1952, Reynolds 6544! in Nat. Herb. Pretoria, and Herb. Kew. A. kedongensis is named after that part of the Kenya portion of the Great Rift which appears to be the specific centre. It occurs in dense bush from near the top to near the foot of the Kedong Escarpment, 24—28 miles north-west of Nairobi, also on Mt. Longonot nearby, where Mr. P. R. O. Bally has collected it. At the type locality, which is 2 miles south of the Lake Hotel (6 miles south of Naivasha), near the south- eastern shore of Lake Naivasha, A. kedongensis occurs in flatter country in dense masses, sometimes forming untidy impenetrable thickets up to 12 feet high and several yards across. On dry slopes of volcanic rock near the southern extremity of Lake Naivasha, it is also found in large numbers growing socially with masses of A. secundiflora Engler. Although both species were seen in flower (April) no trace of hybrids was found. A. kedongensis also grows in large numbers on rocky slopes of the Gilgil Escarpment, overlooking Lake Elmenteita, 2—3 miles north-west of Gilgil, and was also observed 10 miles south of Nakuru. During the last war Major A. G. McLoughlin sent plants from the Gilgil Escarpment, with photographs, to the Division of Botany, Pretoria. Characters which distinguish A. kedongensis from its nearest allies, include slender spindly stems up to 4 met. long, and narrow leaves up to 30cm. long, but not exceeding 3}cm. broad at base, the leaves being unspotted and having small marginal teeth only 2—3mm. long. Until such time as the Aloes of Africa have been investigated, and sections and groupings revised, A. kedongensis can be placed in the section HKuALOog, subsection Prolongatae. A. rabaiensis Rendle, which I have seen from near Mombasa to near Nairobi, differs in having thicker stems, much larger greyer leaves, and smaller almost capitate racemes, and could never be confused. A. nyeriensis Christian, which occurs near Nanyuki and southwards along the Nyeri road, has much larger leaves, and much thicker stems which are slenderer near base becoming thicker upwards. A. pole-evansii Christian from near Kisumu has thicker stems, larger leaves, and different racemes. A. dawei Berger, which I have seen near Entebe in Uganda, is a totally different species with thicker stems about | met. or a little more long, and much larger different leaves. Description. Plants succulent, fruticose, forming shrubs varying from 6 The Journal of South African Botany. 2 met. high to dense thickets 3—4 met. high and sometimes several metres across. Stems slender, spindly, about 4cem. thick, up to 4 met. tall, mostly branched from the base, sometimes with few to many shoots produced at random, the terminal 30—60cm. of main stems sublaxly foliate, with the internodes 2—3cm. distant, the sheaths obscurely striate; the apical portion of stems usually with old dried leaves persistent for about 1 met., nude downwards. Leaves dull greyish-green to dull yellowish-green, basally sheathing, 3.5em. broad at base, 30cm. long, gradually narrowing from base to apex, varying from spreading to recurved: upper surface canaliculate, lower surface rounded: both surfaces unspotted except sometimes in young shoots, the spots disappearing with age; margins armed with pale deltoid teeth with reddish-brown apices, the teeth sometimes forward-uncinnate and averaging 2—3mm. long, 10mm. distant. Inflorescence 50cm. tall, 2—4-branched from about the middle, the branches mostly arcuate-ascending. Peduncle brown with a slight bloom, basally plano-convex and 12—15mm. diam., terete upwards, the lowest branch subtended at base by an ovate-cuspidate, thin, scarious, 5-nerved bract 6mm. long, 12mm. broad at base. Racemes broadly cylindric-acuminate, the terminal 10—12cm. long, about 8cm. diam., laterals a little smaller. Bracts ovate-acute, 5mm. long, 5mm. broad at base, thin, scarious, about 3-nerved. Pedicels 20—25mm. long, the colour of the perianth. Perianth scarlet. cylindric, slightly trigonous, averaging 35mm. long, shortly stipitate. obtusely tapering into the pedicel, 7mm. diam. over the ovary, thence slightly curved and very slightly constricted on the under- side only, the mouth wide open: outer segments free for 14mm. paler at their margins, 5-nerved, the apices subacute, spreading; inner segments themselves free but dorsally adnate to the outer for 20mm., broader than the outer and with more obtuse more spreading brownish apices. Filaments pale-yellow, filiform-flattened, the 3 inner narrower and lengthening in advance of the 3 outer with their anthers in turn exserted 2—3mm. Style lemon, filiform, with stigma at length exserted 4mm. Ovary green, 7mm. long, 3mm. diam., finely 6-grooved. Capsule 22mm. long, 9mm. diam., broadly 3-angled. A. ngobitensis Reynolds. Spec. nov., in Sect. EvAaLor, subsect. Prolongatae. Planta succulenta, fruticosa, ca. 2 met. alta et lata. Four New Aloes from Kenya Colony. a Folia griseo-viridia, 35cm. longa, 5em. lata, sensim attenuata, patentia vel recurvula, dentibus marginibus-deltoideis 3—4mm. longis, 8—l12mm. distantibus armata. IJnflorescentia 80cm. alta, 3—5-ramosa. Racemi subdensi, 15cm. longi, Tem. diam. Bracteae ovato-acutae, scarlosae, 5mm. longae, 4mm. latae, 5-nervatae. Pedicelli 20mm. longi. Perigonium laete aureo-coccineum 36mm. longum, basi 7mm. diam; segmenta exteriora per 15mm. libera. Antherae per 3—4mm. exsertae. Stigma demum 5mm. exserta. [Plate V.] Hab. Kenya Colony: Bushy slopes near Ngobit Bridge, 31 miles north-west of Nyeri on direct road to Mutara Police Post and Thomson’s Falls, approx. 0°5’S 36°50’E. Fl. 22 April 1952, Reynolds 6579! (type) in Nat. Herb. Pretoria, with type number in Herb. Kew and Cor. Mus. Herb. Nairobi; 18 miles east of Thomson’s Falls, fl. 22 April 1952, Rey- nolds 6559! in Nat. Herb. Pretoria, and Herb. Kew. A. ngobitensis is named after the locality which appears to be its specific centre, namely, the valley of the Ngobit River, near Ngobit Bridge, 31 miles north-west of Nyeri, on the direct road from Nyeri to Mutara Police Post and Thomson’s Falls, Kenya Colony. For a mile or more each side of Ngobit Bridge, large numbers of plants were found on bushy slopes in stony ground, forming shrubs about 6 feet high and broad, and mostly branched from the base with the main stems slightly divergent. The most striking feature of A. ngobitensis, and one which immediately distinguishes it from all other shrubby Aloes known to me in Kenya and elsewhere in Africa, is the remarkable colour of the flowers which are a striking and most attractive bright sheeny orange-scarlet, the peri- anth being usually minutely white-spotted. It is also distinguished by having rather slender peduncles and branches, and subdensely flowered somewhat narrow racemes about 15cm. long. Stems branch at ground level, very slightly increasing in thickness upwards, which is also a character of A. nyeriensis found further east near Nanyuki and for some miles along the road southwards towards Nyeri, but the latter is a larger plant with thicker stems, larger leaves, and different racemes with dull scarlet flowers, and could hardly be confused. Near Ngobit Bridge, numbers of A. ngobitensis were seen growing socially with large numbers of A. secundiflora. Both were flowering in April, but no crosses were noticed. At the Suguroi River 14 miles northwest of Ngobit Bridge (which locality is 32 miles west of Nanyuki and 28 miles east of Thomson’s Falls), A. ngobitensis was found among similar Aloe shrubs having slightly shorter denser racemes of dull scarlet flowers. Whether the latter are 8 The Journal of South African Botany. colour forms of A. ngobitensis, or more probably forms of A. nyeriensis extending westwards from Nanyuki, requires further investigation. At a point 18 miles east of Thomson’s Falls, an occasional plant of A. ngobitensis with its bright sheeny orange-scarlet flowers stood out vividly among numbers of Aloe shrubs with dull scarlet flowers. On account of its most attractive colour of flowers, A. ngobitensis is a plant well worth cultivating. Description. Plants succulent, fruticose, forming shrubs about 2 met. high and broad. Stems 2 met. tall, 4—5cem. diam., mostly branched from base, the terminal portion sublaxly foliate for 30—50cm., the old dried leaves persistent, with the lower half of stems nude. Leaves 5cem. broad at base, gradually attenuate and 35cm. long, basally sheathing with the internodes 2—3cm. distant, the sheathing portion obscurely lineate: upper surface flat to slightly canaliculate; lower surface rounded; both surfaces grey-green, without lines or spots except sometimes in young shoots, the spots disappearing with age; margins armed with teeth the colour of the leaf at base, paler upwards and pale brown at apex, averaging 3—4mm. long, 8—l2mm. distant, the interspaces rounded. Inflorescence 80cm. high, rather compactly 3—5-branched from about the middle. Peduncle flattened at base, terete upwards, with the branches slender. Racemes cylindric-acuminate, the terminal licm. long. 7cm. diam., subdensely flowered. Bracts ovate-acute, 5mm. long, 4mm. broad at base, thin, scarious, about 5-nerved. Pedicels the lowest of terminal racemes 20mm. long. Perianth bright sheeny orange-scarlet, greenish tipped, 36mm. long, cylindric slightly trigonous, basally obtusely tapering into the pedicel, 7mm. diam. over the ovary, thence slightly constricted on the underside, enlarging slightly to the mouth; outer segments free for 15mm., paler at the edges, with the apices subacute, slightly spreading; inner segments themselves free but dorsally adnate to the outer for two-thirds their length, broader than the outer, and with more obtuse more spreading apices. Filaments pale lemon, filiform-fiattened, the 3-inner narrower and lengthening in advance of the 3 outer, with their anthers in turn exserted 3—4mm. Style pale yellow, filiform, with stigma at length exserted 5mm. Ovary pale olive-green, 5mm. long, 3mm. diam., finely 6-grooved. Four New Aloes from Kenya Colony. 9 Aloe graminicola Reynolds. Spec. nov. in ser. Saponariae, A. lateritia Engler affinis, sed floribus gracilioribus differt. Planta succulenta, acaulis, dichotome divisa. Folia ca. 16, dense rosulata, lanceolata, patentia, usque ad 27cm. longa, basi 7em. lata; swpra obscure viridia, planiuscula, maculis oblongis lateraliter saepe confluentibus trans- verse seriatis picta; swhtus convexa, maculis transverse picta, ad margines sinuato-dentata, dentibus deltoideis 4—5mm. longis et 10—l5mm. inter se distantibus. Inflorescentia 1 met. alta, 3—5-ramosa. Racemi dense capitati, 5cm. longi, 8cm. diam. Bracteae anguste deltoideo-acuminatae, scariosae, 12mm. longae, 2mm. latae, 3—5-nervatae. Pedicelli 20mm. longi. Perigonium coccineum (interdum luteum), 33mm. longum; tubus circa ovarium globoso-inflatus et 8mm. diam., deinde constrictus et 5mm. diam., hine decurvatus et paullum ampliatus; segmenta exteriora per 8mm. libera. Antherae 1—2mm. exsertae. Stigma demum 2—3mm. exserta. Ovarium 6mm. longum, 24mm. diam. [Plates VI, VII.] Hab. Kenya Colony: In open grassland 8 miles south of Nanyuki (116 miles north of Nairobi), approx. 0°5'S 37°4’E, alt. 6,200 ft., fl. 22 April 1952, Reynolds 6576! (type) in Nat. Herb. Pretoria, with type number in Herb. Kew, and Cor. Mus. Herb. Nairobi; } mile north of Nanyuki, alt. 6,300 ft., fl. 22 April 1952, Reynolds 6574! in Nat. Herb. and Herb. Kew; flat grasslands at Gilgil (74 miles north-west of Nairobi), 0°30’S 36°18’E, alt. 6,580 ft., fl. 19 April 1952, Reynolds 6551! in Nat. Herb. and Herb. Kew; flat grasslands near Ol Joro Orok, 33 miles north of Gilgil, 0°4’S 36°22’E, alt. 7,780 ft., fl. 20 April 1952, Reynolds 6554! in Nat. Herb. Pretoria, and Herb. Kew; Naro Moru (14 miles south of Nanyuki), Aug. 1932, E. R. Napier 2187! in E.A. Herb. and Cor. Mus. Herb. Nairobi. Our new species is allied to A. lateritia Engler—which I have studied at the type locality, Rombo, on south-eastern foothills of Kilimanjaro— but differs from it, inter alia, in having smaller, denser, capitate racemes of much narrower more curved flowers. In A. graminicola the basal swelling of the perianth is also much smaller and the two species could hardly be confused. Plants were found near Timau (14 miles north of Nanyuki), at Nan- yuki, and in large numbers 8 miles south of Nanyuki (type locality) in open grasslands, while yellow-flowered forms were noticed further south near Naro Moru. Considerable quantities occur for 30 miles north- westwards from Nyeri, near Thomson’s Falls, and from there to Ol Joro Orok. It was also seen in the Rift Valley near Nakuru, Gilgil and Nai- vasha, flowering in April. 10 The Journal of South African Botany. In all localities visited, A. graminicola was found growing in grasslands, which suggested the specific epithet. I did not see this species on rocks, or in dry, arid thorn country. Altitudes from 6,000 ft. to 7,800 ft. in the Kenya Highlands, seem to suit it. Between Njoro and Elburgon, some 20 miles west of Nakuru on the road to Mau Summit, it seems that A. graminicola grades through inter- mediates into A. /ateritia. Beyond the Mau Range, especially for 12 miles before reaching Eldoret, only robust forms of A. /ateritia were noticed. Beyond Eldoret, near Hoey’s Bridge, Kitale and to the north, there appear to be at least two more distinctive undescribed species in the Series Saponariae which require further investigation. Description. Plant succulent, solitary, or forming small groups by division, acaulescent. Leaves about 16, densely rosulate, erectly spreading, lanceolate- attenuate, up to 27cm. long. 7em. broad at base, the apex dried and twisted: upper surface flat or slightly canaliculate, dull green with numerous dull white “H’-shaped spots, the spots irregularly scattered or sometimes more or less arranged in interrupted undulating transverse bands; lower surface paler green, with fewer spots, the spots irregularly scattered or more or less arranged in interrupted undulating transverse bands: margins sinuate-dentate, with horny edge, armed with pale brown, pungent, deltoid teeth 4—5mm. long, 10—l5mm. distant, the interspaces rounded and the colour of the leaf. (Sap dries yellow.) Inflorescence a branched panicle up to 1 met. high. Peduncle brown with a grey powdery bloom, flattened and 2cm. diam. low down, terete upwards, 3—5-branched from the middle or higher, the lowest branch subtended at base by a rather fleshy many-nerved bract up to 3cm. long, 12mm. broad at base. Racemes capitate, densely flowered, the terminal averaging 5cm.long, 8cm. diam., the lateral a little smaller. Bracts narrowly deltoid-acuminate, thin, scarious, 12mm. long, 2mm. broad at base, 3—d-nerved. Pedicels averaging 20mm. long. Perianth scarlet to orange-scarlet (sometimes yellow), paler to yellowish at mouth, averaging 33mm. long, slender, with basal swelling 8mm. diam., constricted above the ovary to 5mm. diam., thence decurved, laterally compressed and enlarging towards the throat, outer segments free for 8mm., paler at the margins, the apices subacute, slightly spreading; inner segments broader than the outer, with broader thin white borders and more obtuse spreading apices. Four New Aloes from Kenya Colony. 11 Filaments almost white, filiform-flattened, the 3 inner narrower and lengthening in advance of the 3 outer, with their anthers in turn exserted 1—2mm. Style pale-yellow, filiform, with the stigma at length exserted 2—3mm. Ovary pale green, 6mm. long, 24mm. diam., finely 6-grooved. Capsule 32mm. long, 16mm. diam. at the middle. :ojoy) ‘A104 Joy, Beytvurs L “NO TA “FIN JO Y yu ‘nqureg OAUOP|G wou “PFGE, toqutoydog Y JOUNDY YOlBy 1 “IF OF UBT *] :030Yq) “AuojoH vAUEy “1OA JO 4SOM SoU QZ ‘oPVZVMI JO Ysom soprUt ous 2/770q a0) *f stawTg @ Ol *T *SIy Bie. 1: Fie. 2. PxLate II: A. ballyi Reynolds. Fic. §: Flowers 1/1 from bud to post-pollination stage. with one flower partly laid bare. (Photo: G. W. Reynolds.) Fic. 2: Inflorescence, approx. 75, with sunbird. (Photo: P. R. O. Bally.) Both from a plant near Mwatate. 20 miles west of Voi, Kenya Colony. FI. 31 March 1952. ‘spjousoy sisuahuopey 20)7 2] ALVId G ‘esuys uoleulyjod-ysod 0} pnq uwloay [/[ SHOMOTY 2% “OI “Auojoy) vAUoy ‘BYSVAIBN JO YYNOS SoU g “OOH OYVT JO 43 GI SQOMOIUY, 27] DIT ‘AOTIVA WEL ygnos separ Z ‘“ZeG] [dy 61 PB “USsty BOM jod-ysod Oy png utod O wAuO yy “TEST VW R WUT FY] Hy | SHOMO[y[ FZ YLy] it ‘spjouAey sesuayqobu ao)p :\ ALVIg ‘asR4s uoeurjod-ysod 09 pnq utody [/] SdoMOT WT 2% “DIST GEG [dy cz ‘py ‘Auojog ekuoyy ‘0AN Jo ysoM-YQI0U soul TE ‘esplg qIqosN xreou ‘YsIq “ay g QURTG :T ‘DIT 3) Us| ‘spjousey vpoouunsb 20)f 2, A ALVId ‘adeys uoleurjod-ysod 03 pnq wo [/] SIOMOL 2% “OI “AUOTOD eduay ‘ynduen jo ygnos sopiur g ‘eg, [dy ZZ ‘“P SqURid FT “OI % “OLY ‘SppouAoY] PpoovUuUDLD “yr SPP MLV ‘o@ugs uoLRurpod-ysod oF png worl |) | stoMmofy, $% “Mlyy *AUOJOY) wAuO yy, AOUPA WET USI VE SEG L Md y GLH SUL FL Oto % “OIil THE IDENTITY OF ALOE SECUN DIFLORA ENGLER By G. W. REYNOLDs. (With Plates VITI— xX.) The type of Aloe secundiflora is Volkens 530, in Herb. Berlin, a photograph of which has been sent to me by Prof. G. Werdermann, Dahlem. The type locality is (from the German): “Kilimanjaro District: At the foot of the Rhino Hills (Nashornhugels), in stony ground with Kuphorbia trees; leg. Volkens, 4 July 1893”. The present place-name is Kifaru, which, in Swahili means Rhinoceros. Kifaru is situated at the northern extremity of the North Pare Mountains at approximately 3°35'S 37°33’E which is 9 miles south of Himo, Himo being 18 miles east of Moshi, Tanganyika Territory. I visited the Rhino Hills in April 1952 (with Mr. P. R. O. Bally, botanist at the Coryndon Museum, Nairobi) for the purpose of investi- gating A. secundiflora at its type locality. Large numbers of plants were found at Kifaru in arid country, with tree Euphorbias, Acacia xantho- phloea, and Dum Palms (Hyphaene) nearby. The identity of A. secundiflora can now be established with certainty, but two other species became involved, namely A. engleri Berger, and A. floramaculata Christian. A. engleri was collected by Engler in 1902 and described by Berger in Engler Bot. Jahrb. 36: 60 (1905), the type locality being (from the German) “Kilimanjaro District: Between Taveta and the Bura Hills”. Mr. Bally and I journeyed from Taveta (in Kenya Colony, near the Tanganyika border) eastwards to the Bura Hills, which are at the western end of the Teita Hills, to search for A. engleri. Large numbers of plants were found—many flowering—near Maktau, 34 miles east of Taveta, 38 miles west of Voi, and west of the Bura Hills, in Kenya Colony, at approximately 3°26’S 38°9’H. Prof. G. Werdermann has also kindly sent me a photograph of the type material of A. engleri in Herb. Berlin. From this photograph, and from the description and locality, there can be no doubt whatever that the Maktau plants belong to A. engleri, but an examination revealed that they were, in all characters, identical with A. secundiflora at Kifaru. Consequently, A. engleri, being a later publication, cannot be upheld, and must go into synonymy. Berger’s Fig. 94 in Engler Pflanzenr. Liliac.-Aloin. 252 (1908) of a whole plant of A. engleri shows the rosette of leaves approximately 13 14 The Journal of South African Botany. correctly, but the inflorescence far from agrees with the type material, it does not fit the description, and is in all ways entirely misleading; in fact, it could only have been founded on imagination. Berger’s figure of a flower natural size is also wrong, and far from resembles the actual flower. A difficulty with Berger’s figures is that, with most of the Tropical Aloes, he has obviously founded those of flowers natural size, on dried pressed Herbarium material. and has depicted shapes which he imagined the flowers had when alive, with the result that such figures are mostly inaccurate and misleading. Aloe species which can readily be separated when alive often look much the same when pressed and dried. With Aloe, no sooner are racemes of flowers scalded, pressed and dried in the field, than some distinguishing characters are irretrievably lost. This applies especially to the small white specks or spots found in the perianth of both A. secundt- flora and “A. engleri’”, a character hitherto unrecorded in these two species. The spotting of the perianth is one of the main characters noted by the late Mr. H. Basil Christian when describing his A. floramaculata from the Makumba Valley (south of Mwanza and Lake Victoria), Shin- yanga District, Tanganyika Territory. Mr. Christian submitted material to Dr. J. C. Hopkins, Chief Botanist and Plant Pathologist, Dept. of Agriculture, Salisbury, who reported: ““The pale specks on flowers are not due to a pigment. but are pale translucent spots or stripes caused by the presence of bubbles of gas in the sub-epidermal tissue. The gas does not appear to be enclosed in any special structure.” Since the character “spotted perianth’’ was not mentioned in the description of A. secundiflora, Mr. Christian could hardly have been expected to know of it, while an examination of the actual type material could not have revealed it, since the minute gas-bubble spots disappear when flowers are pressed, and do not re-appear when dried flowers are boiled up. I studied and photographed the type plant of A. floramaculata at Ewanrigg when flowering there in June 1939, and in the light of further knowledge, it now proves to be conspecific with A. secundiflora. Subjoined is the synonymy, with a description based on personal observations of large numbers of plants over a wide area in parts of Tanganyika and Kenya. A. secundiflora Engler in Pflanzenwelt Ostafrikas 140 (1895); Baker in Th. Dyer Fl. Trop. Afr. 7: 459 (1898); Berger in Engler Pflanzenr. Liliac.-Aloin. 267 (1908). The Identity of Aloe secundiflora Engler. 15 —A. engleri Berger in Engler Bot. Jahrb. 36: 60 (1905), in Engler Pflanzenr. Liliac.-Aloin. 252 (1908). —A. floramaculata Christian in Journ. S.A. Bot. 6: 182 (1940). Plants acaulescent or very shortly caulescent; mostly solitary, or in small groups, sometimes soboliferous. Leaves about 20, densely rosulate, erectly spreading, slightly recurved near apex, ovate-lanceolate-attenuate, averaging 45cm. long, 12—14cm. broad at base, dull green, rather glossy, without spots: wpper surface flat low down, slightly canaliculate towards apex; lower surface rather flat low down, more rounded upwards; margins obtuse, sinuate-dentate, armed with laterally compressed deltoid teeth which are pungent, brown with paler tips, straight or hooked forward, averaging 4mm. long, 15mm. distant, the teeth isolated in younger leaves, sometimes joined by a horny brown marginal edge in older leaves. Sap dries yellow. Inflorescence a divaricately branched panicle 1—1.75 met. high, sometimes 2 simultaneously, branched from below the middle with about 10—12 slender oblique branches, sometimes producing 50 and more racemes per inflorescence, the lowest branch subtended at base by a broadly ovate, thin, scarious, pale-brown, 7-nerved bract about 12mm. long, 20mm. broad at base. Peduncle brown with a grey powdery bloom, plano-convex and 35mm. diam. at base, terete upwards, the branches slender, oblique. Racemes more or less oblique, 15—20cm. long, rather laxly about 18-flowered, the flowers more or less secund, the buds rose-pink and greyish tipped, more clearly spotted, the open flowers usually less clearly spotted; the buds suberect and grouped along the top of branch, open flowers cernuous to pendulous, after pollination the drying perianth and fruit becoming erect. Bracts ovate-acute or deltoid, thin, scarious, about 4—5mm. long, 4—5mm. broad at base, 3—5-nerved. Pedicels 8—10mm. long. Perianth rose-pink to dull scarlet-red, paler at mouth, obscurely to clearly minutely white-spotted, averaging 35mm. long, cylindric, basally flat and not at all stipitate, 9mm. diam. over the ovary, slightly constricted above the ovary, thence slightly trigonous and slightly enlarging to the open mouth; outer segments free for half their length, with paler margins and subacute spreading apices, obscurely 3-nerved, the nerves turning brownish at apex; inner segments themselves free but dorsally adnate to the outer and with broad pale border, with 3 congested nerves forming a keel throughout, the apices more obtuse and more spreading than the outer. 16 The Journal of South African Botany. Filaments palest lemon, filiform-flattened, the 3 inner narrower and lengthening in advance of the 3 outer, with their anthers in turn exserted omm. Style lemon, filiform, with stigma at length exserted 6mm. Ovary pale green, 6mm. long, 3mm. diam., finely 6-grooved. Note.—The soboliferous form occurs in the localities furthest west, i.e. the Shinyanga and Biharamulo Districts. In the Rift Valley (Lake Naivasha area) small groups are found, increasing by division and not from suckers. In localities furthest east, i.e. Kifaru, Maktau, Kinango, Mariakani, plants are mostly solitary and not in groups. DISTRIBUTION. A. secundiflora appears to be one of the most frequent and most widely distributed species of Aloe in the northern portion of Tanganyika Territory, and in parts of Kenya Colony. It is found from a few hundred feet above sea level to over 6,000 ft. in the Rift Valley and near Nanyuki, on the Equator. From a few recorded localities, but mostly from personal observations, the following is the distribution as at present known: Tanganyika Territory: From Biharamulo in the north-west, east- wards through the Shinyanga, Singida and Kondoa Districts to Arusha, Moshi, Lake Chala, and north of Kilimanjaro; from the northern end of the North Pare Mountains southwards to Kisangiro, and the Mwembe and Vudea valleys on the western side of the Southern Pare Mountains; also 18 miles north of Tanga. Kenya Colony: Abundent near Kinango (36 miles west-south-west of Mombasa) and for the next 30 miles along the Tanga road; from Maria- kani (20 miles north-west of Mombasa) along the road to Voi, thence westwards to Maktau and Taveta; Sultan Hamud, occasional on the Kapiti Plains; Machakos; at the foot of Lukenya Hill; abundant at the Athi River road bridge, Stony Athi (20 miles south-east of Nairobi); east of Thika, near Donyo Sabuk; in the Kedong Valley (north-west of Nairobi); Mt. Margaret; near Mt. Longonot and at the southern end of Lake Naivasha; near Elmenteita; southwards along the Rift Valley to Olorgesailie Prehistoric Site; Kajiado; near Nanyuki (122 miles north of Nairobi); tremendous quantities 34 miles west of Nanyuki near the Mutara Police Post where, in April, the countryside is pink with flowers for a few miles; in the Ngobit valley 16 miles south-east of Mutara. Flowering period varies from April to August, depending on the rains, and the locality. ‘Io[suq diopr~punoas 20)F *TITA ALVTd -gsuqs uolRurjod-ysod 04 pnq Ulody *[/] SHOMOTY 2% “OL “ALOPLAIDT, VBYTAUBTUR J, ‘Bysnay JO 4SOM SoplUT 77 ‘TCG L YOMBIN SZ U IURT FT Ol ‘G “Ol LWT ‘opus uorunjod-ysod oy pug Woda [/ | SHOMO[oL Fe “OL 4QsSRO ‘(aotuogy aajbuo “pr =) “Auojopn vAuoy “VQorny, JO SOU FE “NeIYRT Uwou “ZEG_ Youeyw OF “VU WTC +1 Ole amay ty w “BEET onl: Ze oY ‘Aaogtase J, “elsopoyy, “Gg ‘Aanqsipeg reo “SdWUBM GL vyrAursuey, “ysiq woURAUIYg Utorf WeIYStIy—) KF jo gued odAy, :z “914 6L lady 61 8 Weld tL OW pIpnowDULDLoy * -Auojog vAuey ‘AoTTeA Wry “VYysBAleN oYe'T JO pus UTEyANOS Iv0U a Gg ‘lopsug, Duoyipunoas "Pix ULV I Na . I Pe c ee eee ee BOOK REVIEWS. TREES AND SHRUBS OF THE KruGcER NationaL Park. By L. E. W. Codd. Memoir no. 26, Bot. Survey, Div. of Botany and Plant Patho- logy, Dept. of Agriculture, Pretoria. 1951. 7s. 6d. This book should receive a very warm welcome, both for its own sake and as, we hope, the forerunner of a series of such books on the other biological groups—mammals, birds, reptiles, etc.—in the Kruger National Park. It may be regarded as the first serious attempt to educate the ordinary visitor to the Park in the living organisms which are so lavishly displayed around him. It is fitting that the vegetation of the Park should be the first to receive attention, as it is fundamental to all other forms of life: and we trust that corresponding zoological handbooks will soon follow this admirable exemplar. For the visitor the illustrations in the book are of prime importance: no keys are provided for identification—such would be almost impos- sible—and the amateur’s method of matching with pictures is the only means available of getting to know and discriminate between the host of trees and shrubs. For this purpose the large number of photographs and drawings in the book are very well suited: and having made identi- fications, the descriptions, notes, popular names (African and European), etc., given in the text will increase the interest of the intelligent visitor very greatly. These descriptions are such as to be understood by persons without special botanical knowledge, and at the same time are accurate by botanical standards. Certain groups, e.g. the Acacias and Albizzias, present special difficulties to the amateur (as to the professional botanist) and the book gives greater assistance in such cases. The text and illustrations are printed in a uniform sepia ink on slightly tinted paper and the general effect is soft and pleasing. Six charming colour-plates by Miss Cythna Letty adorn the book. The photographs of lion and elephant are perhaps added to suggest that botanising should be done with due awareness of other forms of life. The Government Printer has gone unusually gay in the paper cover, and is indeed to be congratulated on the attractiveness of the production. Dr. Codd deserves cordial thanks for a very noteworthy contribution to knowledge of the Park’s flora and to the pleasure and interest of its visitors whether botanical or otherwise. R. H. Comerton. 18 The Journal of South African Botany. PLANTS OF THE BrBte. By Harold N. Moldenke and Alma L. Moldenke. xx + 364 pp. 95 illus. Waltham, Mass., U.S.A.: Chronica Botanica Co. 1952. $7.50. London, W.C.2. Wm. Dawson & Sons, Ltd. To find a comprehensive book on Bible plants written by competent botanists from a critical scientific point of view is a quite unusual pleasure. As Dr. and Mrs. Moldenke remark: “Anyone delving even very super- ficially into the literature of Bible plants will be impressed at once by the amazing discrepancies, contradictions, palpable mis-identifications, erroneous statements and general confusion which exist there’. They think however that “it would be more surprising to find accuracy in botanical matters than not to find it” in the Scriptures themselves and in subsequent commentaries thereon. The Authorised Version itself is responsible for the perpetuation and origination of many of the mis- identifications of biblical plants with common English ones. Now that the flora of the Middle East—though greatly changed since biblical times—is reasonably well known, it is possible to correct many of the popular misconceptions current among biblical students. This the Moldenkes have done in their highly efficient, thorough and at the same time readable treatise. Not only are the plants identified with reasonable certainty, but an enormous amount of relative comment is added, thus casting a most valuable and entrancing light on the whole subject of the vegetational background of the scriptural narratives. The illustrations are a most attractive feature of the book: some are photographs or drawings of the plants themselves, others illustrate biblical incidents or legends. Most charming of all is perhaps a reproduction of the elder Cranach’s Garden of Eden: and there are several other representations of the “‘apple” legend. (For “apple” should probably be read “apricot”’.) It is unfortunate that the confusion with regard to Aloe succotrina, a South African plant, as the source of Socotra Aloes (A. Perryi or A. vera), is perpetuated. (See Reynolds in Journal S.A. Botany XIV, p- 1, 1948.) The inclusion among “Bible plants” of the organisms causing plague, ringworm, vinegar fermentation, etc., is a little surprising, but adds greatly to the general interest of the book. “Plants of the Bible’, says the publishers’ note, “has been prepared carefully to be serviceable to interested lay- and churchmen, Bible lovers of Catholic, Jewish and Protestant faiths, scientists as well as nature- and garden-lovers.”’ In this comprehensive aim it should be eminently successful: and it is easy to prophesy a very wide appeal for what must certainly be regarded as the outstanding, because wholesomely scientific, book on the subject. R. H. Compton. Book Reviews. 19 Botany oF SuGar Cane. By C. van Dillewijn. Pp. xxiv + 371. 229 line drawings, photographs and diagrams. Waltham, Mass., U.S.A.: Chronica Botanica Co. 1952. $6.00. London, W.C.2. Wm. Dawson & Sons, Ltd. This fine monograph brings into volume form the copious work that has been done on every aspect of the structure, physiology and chemistry of the Sugar Cane—a plant to which, as the author says, perhaps more research has been devoted than to any other commercial crop, largely on account of the splendid work of governmental and private experiment stations, the latter entirely run by the sugar industry. It deserves much fuller notice, in a country where sugar cultivation is one of the chief agricultural industries, than can be given here, and other journals will undoubtedly commend it to growers and manufacturers. It is written throughout with an eye to the improvement of sugar culture, and emphasis is laid on the discrepancy between efficiency in factory and in field. “Whereas in the factory,” says Dr. van Dillewijn, “every effort is made to recover as much sugar as possible, in the field only a fraction of the economical yield is obtained ...” ““The greatest possibilities of reducing the cost price of sugar are to be found in the field...” “In many cane-growing countries field practice has failed to keep pace with scientific progress, and in fact is lagging far behind.” How far these trenchant remarks apply to South Africa is not stated, and can best be judged by the very able research workers attached to the sugar industry here. In its dignified style and the quality of its binding, paper, print and illustrations this is perhaps the most noteworthy volume that the Chronica Botanica Co. has yet produced: Dr. van Dillewijn’s deep know- ledge of his subject has been presented in a worthy form. R. H. Compton. MaizE IN THE GREAT HERBALS. By John J. Finan. Waltham, Mass., U.S.A.: Chronica Botanica Co. 1950. $3.00. London, W.C.2. Wm. Dawson & Sons, Ltd. This elegantly-produced volume is mainly a reprint from the Annals of the Missouri Botanical Garden, vol. 35, 1948, handbound as designed by Dr. Frans Verdoorn, Editor of Chronica Botanica, and with a fore- word by Dr. Edgar Anderson. In South Africa we are so familiar with the maize plant that we forget what an extraordinary grass it is. This book enables us to recapture some of the wonder of the renaissance herbalists as reflected in their woodcuts, engravings and descriptions of this marvellous plant. Mr. Finan has brought together all the early references to it, collated them, 20 The Journal of South African Botany. reproduced the illustrations and written a very interesting account of the early history of the plant thus revealed. It would appear that at least two distinct types of maize were introduced to Europe. One was the kind brought back from the Caribbean by Columbus and _ his successors. The other presents an unsolved problem: it is probably the plant grown by the eastern North American Indians, but the herbalists believed it to have come to Europe from Asia. Apart from being occasionally repetitive, Mr. Finan’s study is a very attractive piece of work on a limited problem, and is perhaps specially valuable in America where the Herbals are less familiar than in the European countries. It is interesting to note that it was financed by a commercial company selling hybrid corn, in spite of the fact that, as Dr. Anderson points out, the subject is of no conceivable practical value. — R. H. Comerton. JOURNAL OF SOUTH AFRICAN BOTANY VOL. XIX. A NEW ALOE FROM SOMALILAND PROTECTORATE. By G. W. REYNOLDs. (With Plate XT.) Aloe jucunda Reynolds. Species nova. Forsitan in Sect. EUALOE Subsect. Parvae pertinet; nulli alii arcte affinis; sectionem propriam constituere debet. Planta parva, sobolifera, mox caespitosa. Folia ca. 12, dense rosulata, ovato-acuminata, 4 cm. longa, 2.5 em. lata, patentia vel apice recurvula, swpra viridula, parce maculata; subtus convexa, maculis numerosis parvis picta; marginibus dentibus 2 mm. longis, 3—4 mm. distantibus armata. Inflorescentia simplex, 33 cm. alta. Racemus 13 cm. longus, 5 em. diam., sublaxe ca. 20-floribus. Bracteae ovato-acutae, 5 mm. longae, 3 mm. latae, scariosae, 3-nervatae. Pedicelli 7 mm. longi. Perigoniwm roseum, cylindricum, 20 mm. longum; segmenta exteriora per 7 mm. libera. Antherae per 2 mm. exsertae. Stigma demum 2—3 mm. exserta. Ovarium 5 mm. longum, 2.5 mm. diam. Hab. Somaliland Protectorate: Gaan Libah, ca. 9° 55’N 44° 18’E, coll. P. R. O. Bally, no. 7157, cult. Johannesburg, fl. 31 Jan. 1953, Reynolds 6223 (type) in Nat. Herb. Pretoria. This charming and very distinctive little Aloe was collected by Mr. P. R. O. Bally, botanist at the Coryndon Museum, Nairobi, on 2nd May 1949, at Gaan Libah, Somaliland Protectorate. This locality is at the western end of the Golis Range about 5 miles east of Mandera (which is on the Berbera-Hargeisa road), and about 25 miles west of Upper Sheik (on the Berbera-Burau road). Mr. Bally found numbers of plants forming large clumps, 1—2 ft. across, commonly 21 22 The Journal of South African Botany. hidden in bushes in open places on the upper edge of a dry, windswept escarpment, flowering on 2nd May 1949. A plant sent to Kew flowered there in May 1951, when the inflorescence was preserved in liquid, but no description was drawn up at the time. When Mr. Bally’s plants flowered in Nairobi on 6th October 1951, he made an excellent life-size painting of a flowering plant and kindly sent the painting to me, together with two plants. These plants are now well established in Johannesburg, and when one of them flowered on 31st January 1953, photographs were taken, a full description drawn up, and the type material prepared for the National Herbarium, Pretoria. A.jucunda is a most distinctive little species with rather squat, pale rose-pink flowers which, in shape and size, are allied to those of A .mcloughlinii Christian (see Fl. Plants Afr. 28: Pl. 1112 (1951)), but the latter, which comes from Diredawa in Eastern Ethiopia, is a considerably larger plant with a branched inflorescence, and entirely different leaves. At a quick glance the clustered rosettes and leaves of A.jucunda bear a slight resemblance to those of Haworthia tesselata. The leaves of A.jucunda are flat at base with the marginal teeth more or less in the same plane, while in the upper two-thirds, the margins are somewhat involute with the marginal teeth more or less standing up at right angles to the median plane of the leaf. When the Aloes of Tropical Africa have been more fully investigated, it is clear that affinities and groupings will have to be drastically revised. In the meantime, A.jucunda could be included under Berger’s subsect. Parvae, merely on account of its size. Description: Plant succulent, acaulescent or with very short stem, soboliferous, forming dense groups up to 50 cm. and more across, the rosettes averaging 8—9 cm. diam., with cylindric roots. Leaves about 12, densely rosulate, broadly ovate-acuminate, spreading to recurved, 4 cm. long, 2-5 cm. broad at base, rather fleshy; wpper surface dark green and flat low down, turning brownish and channelled in upper half, with numerous pale-green to dull-white spots throughout, the spots lenticular, sometimes confluent, smaller and more crowded low down, larger and fewer upwards; lower surface rounded, deep green, copiously spotted throughout, the spots smaller and much more numerous than on upper surface; margins armed with horny, reddish-brown, pungent, deltoid teeth up to 2 mm. long, 3—4 mm. distant, the interspaces rounded; the teeth smaller near base of leaf, largest near the middle and smaller towards apex, with a few minute spines dorsally in median line of leaf near apex. Inflorescence simple, slender, 33 cm. high. A New Aloe from Somaliland Protectorate. 23 Peduncle plano-convex and 5 mm. broad at base, gradually becoming terete upwards; clothed with about 6 sterile-bracts, the lowest deltoid- acute, 9 mm. long, 4 mm. broad at base, thin, dirty-white, subscarious, 5-nerved, the margins of upper half with a few minute soft teeth. Raceme 13 cm. long, 5 cm. diam. low down, sublaxly about 20-flowered, the youngest buds suberect, grey-green striped in upper third and more crowded, the older buds spreading, with the open flowers nutant and more laxly disposed (10—15 mm. distant). Bracts lowest ovate-acute, 5 mm. long, 3 mm. broad at base, thin, scarious, dirty-white, 3-nerved. Pedicels lowest 7 mm. long, of a deeper pink colour than the base of the perianth. Perianth 20 mm. long, pale rose-pink (nearest coral-pink R.C.S. 13), cylindric, basally flat or exceedingly shortly stipitate, 7 mm. diam. around the ovary, thence very slightly narrowed and slightly trigonous, the throat paler to whitish, the mouth wide open; outer segments free for 7 mm., white, with 3 pink nerves, the apices subacute, spreading; inner segments free but dorsally adnate to the outer for two-thirds their length, broader than the outer, white, with 3 congested nerves throughout, the nerves deep pink turning greenish at apex, the apices more obtuse and more spreading than the outer. Filaments lemon-yellow, filiform-flattened, the 3 inner narrower and lengthening before the 3 outer. Anthers in turn exserted 2 mm. Style filiform, yellow, with the stigma at length exserted 2—3 mm. Ovary pale olive-green, 5 mm. long, 24 mm. diam., obtusely tapering into the style. Fig. 2. Fig. 3. Fig. 4. Fig. 1. Plant coll. P. R. O. Bally, Gaan Libah, Somaliland Pro- tectorate, cult. Johannesburg, fl. 31 Jan.1953, x 1/3 approx. Fig. 2. Leaf 1/1, upper surface. Fig. 3. Leaf 1/1, lower surface. Fig. 4. Flower and pedicel, 1/1. Puate XI. Aloe jucunda Reynolds. THE IDENTITY OF ALOE JOHNSTONII BAK. —A NEW SYNONYM. By G. W. REYNOLDs. (With Plates XII and XIII.) Aloe johnstonii Bak. was described in Trans. Linn. Soc. Ser. II, Bot. ii, 351, ¢. 63 (1887), and is included under “Enumeration of the plants collected by Mr. H. H. Johnston on the Kilima-njaro Expedition 1884, by Prof. D. Oliver, F.R.S., F.L.S., and the Officers of the Kew Herbarium”. The locality cited was “‘Kilima-njaro, 2,000—5,000 ft.’’, without date, or material stated. H. H. (later Sir Harry) Johnston, in his book The Kilima-njaro Expedition published in 1886, mentions various Aloes in the neighbourhood of Kilimanjaro, and from my knowledge of that region, he had clearly seen the species later described as A. Volkensii Engler, A. lateritia Engler and A. secundiflora Engler. He does not mention a small grasslike Aloe, but he published the “Enumeration of the plants collected by Mr. H. H. Johnston on the Kilima-njaro Expedition 1884’, wherein, on page 347, is included “‘Aloe (Eualoe) Johnstoni, Baker—Kilima-njaro 2,000—5,000 ft.”. No mention of Taveta is made in Baker’s or Johnston’s account. In his second publication, Baker in Fl. Trop. Afr. 7: 456 (1898) omits Kilimanjaro, and gives “British East Africa: Taveta, 2,000 ft., Johnston !” Berger saw the Johnston sheet at Kew, and in Engler Pflanzenr. Liliac.-Aloin. 167 (1908) he quotes ‘Kilimanjaro: Without precise locality, 2,000—5,000 ft.’’. Mr. E. Milne-Redhead informs me that there is no date stated on the sheet, that the locality given is “Taveta, 2,000 ft.”, and that there are no other notes on the sheet. From his text and maps, it is clear that Johnston had been to Rombo, and was also at or very near the places now known as Marangu, Himo and Moshi (2,700 ft.), as well as Taveta (2,525 ft.) which is in Kenya Colony, 2 miles east of the Tanganyika border, 9 miles east of Himo and 26 miles east of Moshi. It seems that A. johnstonii could have been collected somewhere between Moshi and Rombo, on lower grassy slopes of foothills lying to the south-east of Kilimanjaro, rather than near Taveta which is in drier country with less grass and more thorn. I have not seen specimens from that region, but I have examined material collected by Mr. P. R. O. Bally, botanist at the Coryndon 25 26 The Journal of South African Botany. Museum Nairobi, on the nearby Chyulu Hills to the north-east of Kilimanjaro. This proved to be conspecific with the South African species A. myriacantha (Haw.) R. & S. from the Zuurberg, near Grahamstown, in the Eastern Province of the Cape. On 6th May 1952, accompanied by Mr. Bally, I examined numbers of flowering plants in wet grasslands near the eastern boundary of the Royal Nairobi Golf Course, and more near Langata, about 5 miles south- west of Nairobi. These plants were unquestionably conspecific with the Chyulu Hills species, and in no characters could they be separated from A. myriacantha of the Cape. For these reasons A. johnstonii cannot be upheld as being specifically distinct from the earlier described A. myriacantha and consequently should go into synonymy. In the South African and East African plants the narrowly linear leaves are dorsally obtusely keeled, which is a character unfortunately omitted from the description in Aloes S. Afr. 116 (1950). It is not necessary to describe the species again, but the synonymy will now read: A. myriacantha (Haw.) R. & S. in Syst. Veg. 7: 704 (1829); Kunth. Enum. pl. 4: 516 (1843); Baker in Journ. Linn. Soc. 18: 156 (1880), in Th. Dyer FI. capens. 6: 306 (1896): Schonland in Rec. Alb. Mus. 1: 35 (1903); Berger in Engler Pflanzenr. Liliac.-Aloin. 166 (1908); Reynolds in Journ. S.A. Bot. 13: 100 (1947), in Aloes 8. Afr. 116 1950). ial Bobi myriacantha Haw. in Phil. Mag. 122 (1827). — Leptaloe myriacantha (Haw.) Stapf in Bot. Mag. t. 9300 (1933); Christian in Fl. Pl. S. Afr. 20: Pl. 799 (1940). — Aloe johnstoni Bak. in Trans. Linn. Soc. Ser. 2, Bot. 2, 351, t. 63 (1887), in Th. Dyer Fl. Trop. Afr. 7: 456 (1898); Durand et Schinz Consp. Fl. Afr. 5: 308 (1893); Engler Pflanzenw. Ost-Afr. 140 (1895); Berger in Engler Pflanzenr. Liliac.-Aloin. 167 (1908); Reynolds in Journ. S.A. Bot. 13: 100 (1947). The following is the distribution as at present known: South Africa: Grahamstown, Port Alfred, East London. Zululand: near Richard’s Bay. Southern Rhodesia: Inyanga Downs. Tanganyika Territory: Kilimanjaro; Bukoba Dist., Kiboroga steppe north of Kagera (c. 1° 10’S 31° 20’E). Kenya Colony: Chyulu Hills, Machakos, Nairobi, Ngong. Uganda: The following material is in Herb. Kew.: Ankole: Gayaza. 5,000 ft., May 1929, J. D. Snowdon 1912; E. Ankole, Sanga Rest Camp, 4,400 ft., Oct. 1932, W. J. Eggeling 593; Ankole, Mbarara, 5,000 ft., The Identity of Aloe johnstonii Bak.—A New Synonym. 27 March 1939, J. W. Purseglove 654. This represents a range of almost 3,000 miles, and is the widest at present known for any species of Aloe in Africa. In South Africa A. myriacantha flowers once a year, in April. In East Africa and Uganda plants flower after the rains, from March until October according to locality, and it appears sometimes more than once a year. Two plants collected near Nairobi when flowering there on 6th May 1952, flowered in Johannesburg in November 1952, and again in March 1953. In am indebted to the Director and to Mr. E. Milne-Redhead of Kew for a duplicate illustration of the drawing of A. johnstonii used for t. 63 in Trans. Linn. Soc., for notes on the type material of A. johnstonii, and for the Uganda citations. ‘ wi deer: nt af! ip 7 - H lh } eS : Dis “rarie F Huo ohial iS ny or id. ; rf via AL ve t as a Vy DM tty 7 Be ian na rf af ond ail Aut lal Trans Lise Sot. Sen 2 Bot You th & 83 DNagre Fak oe Baker ALDE JDHMSTON: . Fig. 1. Prate XII. Aloe myriacantha (Haw.) R. & S. ). north-east of Kilimanjaro, Kenya Colony. 7 2, t. 63 (188 Bot. s. Linn. Soc. Ser. 2, | Fig. 1. The figure of A. johnstonii Bak. in Tran April, 1938, on the Chyulu Hills, aterial collected by Mr. P. R. O. Bally, Photo P. R. O. Bally. 2. M Mig. Fig. 2. Prate XIII. A. myriacantha (Haw.) R. & 8. Fig. 1. Plant « 1/3, fl. 6th May, 1952, Royal Nairobi Golf Course. Fig. 2. Raceme 1/1, from a plant fl. 6th May, 1952, Langata, near Nairobi. Fig. 3. Flowers 1/1, from bud to fruit stage, Langata, near Nairobi. Hig. 3. A NOTE ON SEX IN ROYENA GLABRA L. (EBENACEAB). By Captain T. M. Satter, R.N. (RET.). The small shrub Royena glabra L. is widely spread in the Western Province and is particularly common on the slopes in the northern part of the Cape Peninsula. In the description given by Hiern in Vol. IV § 2, p. 457 of the Flora Capensis the flowers of this plant are said to be hermaphrodite or polygamous; the anthers about ;’5 in. (approximately 2 mm.) long, not always fertile; the fruit globose or oblong, 4 to 3 in. (8—17 mm.) long, purple or reddish, and the seed spheroidal, black. I was recently asked to collect some seeds of this Royena for growing in the famous gardens of Tresco Abbey in the Scilly Isles and was at once struck by the fact that only a small proportion of the bushes bore fertile fruit. These fertile fruits are oblong or oblong-ellipsoid, usually 13—18 mm. long and 6—9 mm. in diam., at first green, then turning to deep reddish plum colour and are finally black and wrinkled before falling. Very rarely a deformed fertile fruit may be found which approaches spherical in shape. The seed, freed from the pulp, is ellipsoid or oblong- ellipsoid, extremely hard and shining chestnut brown, usually solitary, often 1—1-5 cm. long and 4—5 mm. in diam., but occasionally found in adpressed pairs, with the adjacent faces flattened. Most of the bushes bear no fruits at all, while others produce a few, or more rarely many small abortive, almost spherical fruits, usually 4—7 mm. long, which produce no seeds. These abortive fruits sometimes persist on the plants during the following flowering season. Both abortive and fertile fruits are puberulous. These observations suggested that the flowers of this species were possibly dioecious for, though a large number of bushes were examined in the neighbourhood of Kirstenbosch and Wynberg Hill, no fertile fruits could be found on the suspected male plants, although they were usually copious on the supposed female plants. Later, in the flowering season, the flowers of certain marked plants were examined and it was found that the anthers of the supposed ¢ plants, including those bearing abortive fruits, varied from 2—3 mm. in length and bore copious pollen, whereas the anthers of the plants which bore fertile fruits were only 1 mm. long or less, and entirely devoid of pollen. Furthermore, the flowers of many more plants examined in the field showed that the sex could be at once determined by the marked difference in the size of the anthers. The comparatively few, suspected as 2 from the small anthers, have been found later to be producing fertile fruits. The ovary 29 30 The Journal of South African Botany. of the 3 contains + ovules and appears to be indistinguishable from that of the flowering 2. The styles, too, in both seem to be identical. At this stage, therefore, the flowers might easily be taken for hermaphrodite. An examination of bushes on the slopes near the northern boundaries of Kirstenbosch in February 1953 revealed that out of 19 adult bushes examined, only 2 bore fertile fruits. To avoid any suspicion that perhaps a local form or variant of R. glabra may occur in the limited area investigated, the flowers of 16 herbarium specimens from the Provinces of Caledon, Ceres, George, Knysna, Paarl, Piketberg, Tulbagh and Worcester, were dissected, and inspection of the anthers showed that 13 were male and only 3 female. SUMMARY. It seems. therefore. beyond question that the flowers of Royena glabra are dioecious and that the bushes are definitely either 3 or 9, the 3 plants predominating. The important discrepancies in the description in the Flora Capensis may be summarised as follows:— Organ. As described in Flora As observed in many living Capensis. and dried plants. Flowers .. xs .. | Hermaphrodite or poly- | Dioecious. gamous. Anthers .. . | About 2 mm. long, not | g 2—3 mm. long, fertile. always fertile. 2 0-7—1 mm. long. sterile. Fruits 8—17 mm. long 3g 6—7 mm. long, abortive, rarely developed. 2 13—18 mm. long, fertile. Seed ze So .. | Spheroidal, black .. | Ellipsoid or oblong-ellipsoid, shining, brown. It is presumed that the descriptions of Royena given by Mr. Hiern in the Flora Capensis were all made from the study of dried specimens only, and the facts here recorded from field study reveal the difficulties with which the purely herbarium systematist has to contend. In the case of R. glabra, he has cited 37 collectings and apparently not been able to arrive at the true facts. It is noteworthy, also. that of the other 16 species described in the Flora Capensis, only one collecting is cited in the case of 6 species and only two or three for 4 others. While it is true that in 4 of these cases the sex of the flowers is not specified definitely and the term subdioecious is used freely in this respect. it seems that full field observa- tion might reveal much that is at present unknown about the sex question in this genus. The casual collecting of individual specimens, whether in flower or in fruit. proves to be of little help in this direction. ACKNOWLEDGEMENT. I gratefully acknowledge the help given to me by Dr. M. R. Levyns in the microscopic examination of the anthers. KARYOLOGICAL STUDIES AND CHROMOSOME NUMBERS IN HYPARRHENIA AUCTA AND H. HIRTA. By Steran Krupxko, Ph.D. (Warsaw). (With Plates XIV, XV.) INTRODUCTION. The area of distribution of Hyparrhenia hirta is extremely extended. It covers the whole continent of Africa, especially its eastern side, with only a few breaks. It passes even to Western Asia. In the Union it was recognised quite long ago as an important species for soil conservation. However, the use of this grass in eroded areas has been restricted because of the low germinability of its seeds. Plant ecologists and grass breeders found that the above species includes two or three forms without sharp and distinct differences amongst them. The present paper deals mostly with these two problems. Its aim is to find the causes of the partial sterility in Hyparrhenia hirta and reveal the karyological differences, if any are present, among the morphological forms. TECHNIQUE. In order to have material for cytological investigations one first must get seeds for germination. In the beginning of this work soaking of collected seeds in 95 per cent Alc. was often used in order to separate easily empty scales and florets from fertile seeds. To check the influence of this treatment on root mitosis and on the germination of seeds, a small series of experiments was carried out with different times of treatment compared with untreated control samples of the same samples of seeds. It was found that 5’ or even 10’ treatment of florets with 95 per cent Alc. had no influence on the root mitosis, but definitely affected the germination. Thus to be on the safe side when this treatment was used, the material was soaked for 5’ and afterwards the florets were transferred to water for 15’—30’. Subsequently fertile seeds were selected under a dissecting microscope using needles. To obtain adventitious roots from mature plants dug from the soil the following method was used. The leaves and stems were cut to a height 31 32 The Journal of South African Botany. of 10—15 em. above soil level and smaller roots were also cut. The plants were then soaked overnight or for 24 hours in a Hortomone solution of the concentration: 14 cc. of Hortomone to | pint of water. Next the plants were placed between sterilised wet blotting paper in sterilised dishes and the dishes then placed in a thermostat at + 40°C. On the tenth day usually the first adventitious roots appeared. It was found that the treatment with Hortomone did not affect the mitosis. Shorter soaking, for example for 4 hours also had no effect on germination, but stronger concentrations of Hortomone gave less satisfactory results. Often plants were potted out in soil after such treatment and gave a good crop of young roots in 1—2 weeks’ time. Most of the root tips for microtome technique were fixed mainly in two fixatives: I. Navashin fixative, Stockholm modification (see Maheshwari, 22). Liquid A: Chrom. acid—1l gm., glace. Acet. ac.—l0 cc., water— 65 ce. Liquid B: Commercial formalin—40 cc., water—35 cc. Before use equal volumes of A and B are mixed. Il. Levitsky fixative (Avduloy, 2). The best results were obtained with a modification of the above fixative. Equal parts of the following solutions were mixed: (a) 4 per cent formol; (6) 2 per cent Chrom. acid. The material was infiltrated for 15’ with this mixture. using a pump. For staining microtomed material Newton’s Crystal Violet stain was used after Navashin’s fixative. Adventitious root sections were stained for 30’ and differentiated in Lugol for 30”. Primary root sections were stained for 20’ and differentiated in Lugol for 35”. These times of staining were the best to bring out chromosome morphology, but for counting deeper staining was preferred. and the time required for differentiation was shorter—20”. Another modification of value for morphology was: staining 15’, Lugol 30—32”. For the above staining techniques the most satisfactory thickness of sections was 10 mu. After the treatment with Levitsky fixative the material was stained with Iron Hematoxylin. Before staining the mordants used were: (a) three days’ treatment in | per cent Chrom. acid followed by short washing in distilled water and transferring to (6) 6 per cent alum solution for 24 hours. The thickness of sections here was 7—8 mu. FEULGEN METHOD. MopDIFIED: After the use of Navashin fixative the Feulgen method of Warmke’s (45) modification for squashing of isolated sections and isolated cells with metaphase was used. Slides with root tip sections (thickness 8 mu.) were hydrolised in N HCl for 45’ at 60°C. Next followed staining in Karyological Studies and Chromosome Numbers in 33 Hyparrhenia aucta and H. hirta. leucobasic fuchsin for 3 hours, then—mounting in dilute balsam. Squashing of the sections was advisable only on the second to third day after mounting. With this method it was possible to squash sections and to spread the chromosomes in 3 slides of H. hirta (7.VI1.48—11.X.48). The counting was 2n = 44. This squashing method was a difficult and laborious process. It was noted that: (a) squashing was impossible on the sixth or seventh day after mounting, and (b) that in a few months’ time the chromosomes lost their colour. SeuasH Merrnops: There are some difficulties in the staining of pollen grain mother cells which are not easy to explain. The best results were achieved in the following procedure with the anthers: 1. fixation in 1 p. of glacial acetic acid and 5 p. of absolute alcohol. 2. 3 days in aceto-carmine. 3. mounting in 45 per cent acetic acid, with slight heating, followed by squashing and immediate mounting in Euparol. With direct squashing in aceto-carmine it was difficult to obtain good spreading as well as good staining. It was helpful to use aceto-carmine mixed with 4 per cent Iron Acetate as follows: 1 drop of Iron Acetate added to 10 cc. of aceto-carmine. The mixture must be prepared fresh nearly every day. Sometimes good results were obtained by using carmine saturated in glacial acetic acid instead of solution 1. Snamt SromacH Juice (Fabergé, 10): The sections prepared from fresh root tips were fixed (acetic alcohol) and then passed to 70 per cent ale. Next, the sections were passed through alcohol and water changes and left in snail stomach juice for 5} hours at + 40°C. Afterwards sections were washed in water, mounted in a drop of aceto-orcein, heated and squashed. They had been left in aceto-orcein for 20’. Sometimes it is necessary to use an additional 1—4” of heating during the staining. The active substance here is probably cytase and the best source of it is the edible snail—Helix pomatia. We used the South African relatives of this genus with very good results. The process of dissection of the snails and of extraction of juice is, however, troublesome and time devouring. ProtinoL MetHop (MacKey and Clarke, 21): Pectinol in 1 per cent watery solution may be used at room temperature or at temperatures not exceeding 49°C. Entire roots may be treated or 34 The Journal of South African Botany. sections of root tips previously fixed, passed through the alcohols and washed carefully in water. Then the above sections were transferred to 1 per cent pectinol for 5—6 hours at 40°C., or for 15—17 hours at room temperature. Finally they were washed in water, mounted in a drop of aceto-orcein (or aceto-carmine), squashed and stained. The roots preserved for a long time in 70 per cent alcohol require more complicated treatment as follows: 1. prepare hand sections, 2. transfer for 1—14 hours to the Warmke liquid (equal parts of N HCl and 95 per cent alcohol) at 58°—60°C., 3. pass sections quickly through 95 per cent alcohol, and then gradually bring down to water, 4. change water, 5. leave sections in 1 per cent pectinol for 24 hours at room temperature, 6. pass through water, squash and stain with aceto-orcein. GERSTEL METHOD (12): The original Gerstel method, when properly modified for difficult grass roots, gave quite good results with regard to counting of the chromosomes. Here again entire root tips or sections can be treated. With Hyparrhenia hirta it was found preferable to prepare sections from the fresh living roots which were immediately fixed. The procedure was as follows: 1. fixation in cold 10 per cent N HCl and infiltration for 15’, 2. transfer sections in the 10 per cent N HCl to the thermostat at 58°C. (or 60°C.), for 1 hour and 30 minutes, 3. 95 per cent alcohol for 4—1 minute and transfer to aceto-orcein mixed with HCl (in the proportion of 1 ee. N HCl + 10 cc. aceto-orcein), 4. aceto-orcein + N HCl for 1 hour, 5. transfer to a drop of pure aceto-orcein, heat and squash. This procedure was found to be necessary for difficult roots like those of H. hirta. Applied to other plants, the procedure may be simplified when, for example, the transfer to 95 per cent alcohol may be omitted, aceto-orcein may be used without HCl, and the times may be shortened. O'Mara MetuHop (32): This method gives very distinct staining of resting nuclei (chromo- centers) and prophase stages. Metaphase chromosomes were, however, rather difficult to stain properly. Fresh root tips or sections of them are treated as follows: 1. 3 per cent methanol for 3 hours, which is then replaced by the following fixative: methanol—65 parts; chloroform—5 parts; glacial acetic acid—30 parts; 2. leave in the above fixative for 13 days (other plants require only 3 days, e.g. Kniphofia roots): 3. transfer to a drop of aceto-orcein (which may be used with N HCl as described under Gerstel’s method) or to fresh aceto-lakmoid; 4. squash and heat at least twice; 5. stain 5—10’. Karyological Studies and Chromosome Numbers in 35 Hyparrhenia aucta and H. hirta. Leucopasic Fucusin SquasH MeErHop: Root tips are fixed in acetic alcohol for not less than 2 hours. Two modifications were used: A. Sections of the fixed material were treated as follows: 1. In 1 per cent pectinol for 4 hours at 40°C.; 2. quick washing in water; 3. hydrolysis in N HCl at 60°C. for 1 hour; 4. long staining in LB-fuchsin (up to 12 hours); 5. squashing in a drop of LB-fuchsin. B. Sections were treated as follows: 1. In 4 per cent NH,OH at 60°C. for 15’; 2. transferred to N HCl for 14 hours at 58°C. or 60°C.; 3. long staining in LB-fuchsin (from 3—12 hours); 4. squashing in a drop of LB-fuchsin (or a drop of 45 per cent acetic acid). Many modifications of above squash methods are useful only for counting the chromosomes. One must always remember that aceto-orcein causes much more swelling of the chromosomes than aceto-carmine. With the LB-fuchsin squash method chromosomes are not swollen but they stain rather faintly. For the morphology of chromosomes in Hyparrhenia hirta the squash technique is useless. The chief aim of this chapter is to save grass cytologists much time and trouble as it is the writer’s hope that his experience will be useful for future grass investigations in South Africa. MeEIotTic CHROMOSOMES, NUMBERS AND BEHAVIOUR. The investigation of meiotic chromosomes (bivalents) in both 1-meiotic metaphase and in diakinesis was done mainly in two ways: (a) with varia- tions of the squash method and (6) with material fixed in Navashin liquid and Carnoy and sectioned on the microtome. The details of the techniques and difficulties met with have been described in the previous chapter. The results will now be given. From the commencement of this work with Hyparrhenia hirta (strain 2n = 44) in early 1944, it was clear that the plant is extremely variable in number of chromosomes in pollen grain | mother cells and rich in many irregularities during disjunction. The species always gives difficulties in squashing and staining with aceto- carmine and with other aceto-methods. It was first thought that the variable numbers in the haploid set might have been due perhaps to very hot and dry weather in April, May and June, 1944, but when material was collected in a wet season in the following year the results were the same. It was soon found that observations on the valents association in diakinesis are very uncertain due to the smallness and indistinctness of bivalents. 36 The Journal of South African Botany. TEXT FIGURE 1. Fic. 1.—H. aucta. Root tip metaphase with 2n = 20. Fixation—Levitsky (5:5) with 4% formol, 1% chrom. acid 3 days. Stained—iron haematox. Thickness 7 p. B. & L. Fl. oil immers. x 98 and Comp. oc. x 10. Magnification + 1400 x. Fic. 2 to Fic. 5.—Drawn with B. & L. Fl. oil immers. x 98 and comp. ok. x 15. Magnification + 2800. Fic. 2.—H. hirta, root tip metaphase with 2n = 44. Fix—Levitsky (5:5) with 10% formol, 1% Ac. Chrom.—3 days. Stained—iron haematox. Thickness 7 p. Fic. 3.—H. hirta from U.S.A., adv. root tip metaphase with 2n = 30. Fix.— Navashin. Staining—Crystal Violet. “‘S’’-like chromosome present. Fic. 4.—H. hirta from U.S.A. 2n = 30. All conditions as above. “S’’-like chromo- some absent. Fic. 5.—Schizochyrium semibarbe from Frankenwald (from the field numbered in 1945—P4). Fix.—Levitsky (5:5) with 10% formol. Stained—iron haematox. Thickness 8 p. Root tip metaphase, 2n = 50. Fic. 6.—Optics as Fig. 7. Tubes 0. Decreased-magnification + 750 x. Stained as Fig. 7. Enormously big diads. Resting nuclei. Laggers in cytoplasm still persistent. Fic. 7.—H. hirta, 2n = 44. P.gm.c. Acetocarmine squash. Leitz, big microscope. B. & L. oil Immers. x 97, comp. ok. x 10. Tubes 160. Magnification + 1700 x. First metaphase; n = 22. In December, 1945, 22 bivalents (Pl. XIV, fig. 4, Text-Fig. 1: 7) were obtained in about half the number of pollen grain mother cells investigated, both in freshly squashed and in microtomed material. It was, however, impossible to state anything definite on their morphology. The rest of the metaphases showed the following numbers: 20, 18 (Pl. XIV. fig. 5), 16 (very rare), 14 (Pl. XIV, figs. 2 and 3 give the same diakinesis taken in two levels). Even numbers were usually found in these observations and odd numbers, e.g. 21 and 17, were rarely found. From the beginning of these observations many laggers (Pl. XIV, photo. 6) were evident in Karyological Studies and Chromosome Numbers in 37 Hyparrhenia aucta and H., hirta. anaphases. They were found in every season, in material collected in moderately warm weather in December (PI. XIV, fig. 8), as well asin a very hot period during February and March. It was thought important to determine exactly whether the laggers were included later in the daughter nuclei or whether they persisted in the cytoplasm during the telophase and what was the final fate of these bodies. In anaphase they were found in various positions in relation to the poles and the equator, as well as in very variable numbers. Numerous small laggers were present on the equator (Pl. XIV, fig. 6) in the beginning of anaphase. They were also present in the middle telophase (Pl. XIV, fig. 7) approaching one of the daughter nuclei, sometimes from two directions. In some cases they were arranged in one line between two daughter nuclei during late telophase (Pl. XIV, fig. 8). In squash aceto-carmine slides they could be found in different stages, for example in diads (Text-Fig. 1; 6). In tetrads they were very distinct, either like separate chromosomes (Pl. XIV, fig. 9), considerably decreased in size, or organised into dwarf nuclei (Pl. XIV, fig. 11). In very young pollen grains with | nucleus, before the formation of the exine, they have been seen either single (Pl. XIV, fig. 10) or even as many as four (Pl. XIV, fig. 16). This was the last stage in which they were observed and in older pollen grains they are not present. Probably they are later absorbed completely by the cytoplasm as they are never found subsequent to the 2-nucleated stage in pollen grain development. Nevertheless, the accumulated evidence shows clearly that abundant lagging can be the cause of variable and small chromosome numbers in first meiotic metaphase. Persistence of the laggers through all the stages of pollen grain development including 1-nucleated pollen grains shows that deficient chromosome numbers can not be readjusted and that they are present in pollen grain nuclei as well. In the light of the above facts we can consider the lagging of chromo- somes in the first meiotic metaphase as a chief cause of deficient chromo- some numbers in male gametes and as a chief cause of partial sterility of seeds in this strain of Hyparrhenia hirta. Mature PoLueEN GRAINS AND THEIR GERMINATION. The first attempt to investigate mature pollen grains of Hyparrhenia hirta (strain 2n = 44) showed that the task would not be easy. From the beginning of this work in 1944 and 1945 a high percentage of degenerated and shrunken pollen grains were found; these had an extremely distorted abnormal shape and in many cases it was clear that they were empty, containing neither cytoplasm nor nuclei. The same has been recently 38 The Journal of South African Botany. observed in Anthoxanthum by Lima de Faria (20), who considered it to be an indication of a high degree of disturbance in meiosis and a sign of extreme degeneration. It was estimated that in 1944 and 1945 there were approximately 20 per cent of degenerated pollen grains in each anther. An attempt was made to germinate the pollen grains using different con- centrations of saccharose with agar and gelatine according to H. E. Newcomer's prescription (31). Different types of water were used in the preparation of the media: (a) distilled water: (6) Johannesburg Munici- pality tap water and (c) well water from Frankenwald. Keeping in mind Brink’s observation (4) that pH —7 is optimal for the germination of pollen grains, the pH of Johannesburg tap water was determined, and was found to be 8-0. Accordingly some of the media were acidified slightly but no success was obtained. Swelling of pollen grains was observed, and sometimes they actually exploded, but usually not much development was seen. After application of a 40 per cent concentration of saccharose a few tubes were observed, but they were usually very short and soon died. Artschwaeger and Maguire (1) have recently used the same concentra- tion with success for pollen grains of Sorghum vulgare. In this species the pollen grains started to germinate 15’ after sowing and the growth of the tubes continued for 3 hours. H. hirta, in spite of being a member of the same tribe Andropogoneae, behaved quite differently. Nevertheless, from these unsuccessful preparations of the tubes some important observations were made, not on germinated pollen grains but on those swollen enough to give a clear picture of their nuclei after staining by Newton’s crystal violet method. The mature pollen grain is 3-nucleated, as is usual in the Gramineae (except in the case of degenerated pollen grains). Pollen grains with 4 nuclei (Pl. XIV, fig. 12) were found three times on slides stained with crystal violet and twice in temporary aceto-carmine squashes (5.6.44). Once even a 5-nucleated pollen grain was seen (Pl. XIV, figs. 14 and 15). This matter has been given a fair amount of attention in recent literature. Thus de Mohl (24) described 4, 5, 6 and even 7-nucleated pollen grains in Hyacinthus plants, the bulbs of which had been accelerated in their growth. Apart from these there were some giant pollen grains which the author considered to be diploid. According to de Mohl pollen grains with supernumerary nuclei obtained their additional nuclei from the division of generative nuclei. In his opinion they are supernumerary male gametes and are fertile and compatible. The writer feels, however, that there is not sufficient evidence for this view. On the other hand, La Cour (18) in Tradescantia bracteata, and K. Sax in Tradescantia virginica (38) found pollen grains with supernumerary nuclei after special ID1 OGRAM OF HYPARRHENIA CHROMOSOMES. ~ x2800. H. aucta. JU UU vu ve ow LA ffs eas Hoh. 2n=44 WY UL uu UU vw vy ow we Sf WU Cee fe a I. 2. Fist h 5. 6. 7. 8. g. ioe > ik 12. 3. \%, IS. 6. (7 18. 19. 20. 1 a © © H.h. 2n= Jo. VV UY VY wu wy UY KA Ll the ge 6 ee ii aé a6 i a. : 7. 8. lo. it. Bo TB. 1% 15. 7. 2o. — Karyological Studies and Chromosome Numbers in 39 Hyparrhenia aucta and H. hirta. treatment with heat. These investigators considered they were derived from the additional mitosis of generative nuclei and supposed them to be fertile. The former author also counted the normal number of chromo- somes in supernumerary nuclei (n = 6). Lima de Faria (20, p. 540—543) found pollen grains with supernumerary nuclei in Anthoxanthum and proved that they were derived from tube nuclei (except in one case in which there were two additional mitoses in both tube and generative nuclei). He considers the possibility that they are the result of degeneration. Artschwaeger and Maguire (l.c., p. 664) found supernumerary nuclei in pollen grains of Sorghum vulgare, which they claimed were derived from generative nuclei. K. Sax (39) in Tradescantia and Upcott (43) in Belladonna found some pollen grains with supernumerary nuclei from tube nuclei in triploid strains. They regard this occurrence as a result of unbalance in chromo- some numbers in meiosis of triploids and suppose them to be infertile. In all the five described cases of supernumerary nuclei in H. hirta there was no distinct evidence as to the source from which the additional nucleus of the pollen grain is derived, as mitosis has not been observed. The only evidence of their nature could be found in their different ability to take up stains as compared with normal pollen grains, as they stain like abnormal nuclei. Summing up the observations on the mature pollen grains the two following causes of partial sterility in H. hirta should be added to those given: (a) degeneration of pollen grains; (b) occurrence of pollen grains with supernumerary nuclei. In a test of the viability of pollen grains with N.R. (neutral red— A. Guilliermond) in general all except degenerated and deformed ones were found to be still alive in spite of their inability to germinate in artificial media. The normal mature pollen grains of H. hirta are very rich in starch as shown by their reaction with Lugol. CHROMOSOME MorRPHOLOGY AND CHROMOSOME SET. The following method was used to investigate the morphology of chromosomes in detail. Five to ten of the best metaphase plates in root tips were selected and drawn as exactly as possible with a Zeiss drawing apparatus and at the same magnification, approximately 2,800 x. Next the chromosome drawings were cut out with scissors and arranged in a row on a sheet of cardboard in such a way that the bases of the chromo- somes were on the same level. Homologous pairs were arranged according 40) The Journal of South African Botany. to their heights in descending order so that the longest pair in each group was on the left and the shortest on the extreme right of each group. The conventional grouping was followed, namely, on the left the “V”- shaped chromosomes: next to them the “L-shaped ones and lastly the rod-shaped pairs. In this way all homologous pairs of the same height were arranged approximately on the same vertical line. Having drawn all the best metaphase chromosomes and placed them on one sheet of paper the average diagrammatic pair was prepared from them for each vertical row of pairs. The diploid set of such diagrammatic pairs forms an idiogram of the chromosome set (a term introduced by S. Navashin (27)) and its method of use can be found in older literature (vide L. Delaunay’s paper on Muscari (8) and more recently Taylor’s paper on chromosomes of Aloe, Gasteria. Haworthia (41, 42) and M. Navashin on Crepis (28)). Having prepared idiograms for H. aucta and for two strains of H. hirta the task of comparing them became much easier (see idiograms Text figure 2.) The diploid set of chromosomes in H. aucta was 20 (Pl. XIV, fig. 1, 17: Text-fig. 1; 1). The chromosomes were bigger and a little wider than the chromosomes in both strains of H. hirta. In H. aucta the haploid number was 10 (counted by Mrs. E. Gluckmann, unpublished). In meiosis of H. aucta no abnormalities were observed and there is no difficulty in assuming a basic number of 10 in H. aucta as this is a rather frequent basic number in the Gramineae. We can easily see from the idiogram (see Text-fig. 2) of the diploid set of H. aucta that it consists of the following kinds of chromosomes: (a) One pair—the largest chromosomes—“V’’-shaped with unequal arms and with two constrictions; one on the larger arm and the second on the tip of the “V” angle; (6) three “V-shaped pairs with equal arms; (c) one “V’’-shaped pair with unequal arms, a single median con- striction and a very obtuse angle. This pair might as well be described as “L’’-shaped: (d) three “L’’-shaped pairs; (e) one “S’’-shaped pair; (f) one short rod-shaped pair with median constriction. A more detailed analysis of this idiogram shows three “V-shaped pairs with equal arms consisting of different morphological pairs, having in common only the shape similar to a “V” and equality of the arms. The second pair from the left on the idiogram has two constrictions; one median (on top of the “V” angle. the kinetochore), and the second constriction on the arm. Also this pair is the largest amongst the “V-shaped with the equal arms mentioned under (8). Karyological Studies and Chromosome Numbers in 41 Hyparrhenia aucta and H. hirta. The next pair on the right side is a much smaller one but is dis- tinguished from the preceding pair not only by its size, but also by having only one median constriction. The next pair on the right (the fourth on the idiogram from the left side, text figure 2.) is smaller again, with the single median constriction. It is distinguished from the two pairs mentioned previously under (+) by its very obtuse angle. The three ““L’’- shaped pairs mentioned above under (d) also show variation in their characters. It is possible to distinguish fairly easily (1) one big “L’’- shaped pair with the 2 constrictions, both on the longer arm; (2) two much smaller ““L’’-shaped pairs with the median constrictions. Summing up the morphological features of the chromosome set in H. aucta we can say that it is easily recognisable not only by the number of 20, but also by the presence of such types of pairs as the “‘S’’-shaped pair, the big “V’’-shaped with unequal arms, and a single short rod-shaped pair with median constriction. In H. hirta the author found two strains respectively with 2n — 30 and 2n = 44, growing sometimes very near each other. The diploid set of 2n = 44 may be described first (see the idiogram text fig. 2, Pl. XIV, fig. 18, 19, and Text-fig. 1; 2, 3, 4, in this connection). Here we can recognise the following groups: (a) Seven “V”-shaped pairs with equal arms; (6) one “V’’-shaped pair (N2) with unequal arms and two constric- tions, one on the bigger arm and a median one; (c) one pair “S’’-shaped (N9); (d) seven “L’’-shaped pairs; (e) six rod-shaped pairs. In describing in detail the morphology of the above set it will be noticed that amongst these ““V’’-shaped pairs with equal arms the following types will be found: (1) the largest “V-shaped pair which has either two or sometimes even three constrictions. In the majority of metaphasic plates the constrictions were two in number with usually one on the bigger arm and one (probably the kinetochore) in the median position ; (2) a smaller pair than the previous one (N3) with two constrictions on both arms; (3) two still smaller pairs (N4 and N5) with median constrictions but differing from each other in size; (4) three pairs, again smaller (N6, 7, 8) differing from the above described groups under (1) to (3) by their distinctly obtuse angle. For example, pair N8 could be described as a small “L”- shaped pair because the chromosomes have a very obtuse angle. 42 The Journal of South African Botany. In the group of “L-shaped chromosomes (mentioned under (d)) there are the following types: (1) four “L’’-shaped pairs (N10, 11, 12, 15) differing markedly in size with two constrictions each. The secondary constriction is usually found on the bigger arm while the kinetochore usually has a median position. The pair N11 sometimes shows 2 con- strictions on one homologous chromosome and one on the other. This is due, possibly, to insufficient differentiation: (2) three smaller “L”’-shaped pairs (N13, 14, 16) have one median constriction. They differ less in size amongst themselves than those of the former group. Under heading (e) six rod-shaped pairs have been listed. Amongst them we can distinguish two types: ~ (1) three rod-shaped pairs with one subterminal constriction (N17, 20, 22); and (2) three rod-shaped pairs with one median constriction (N18, 19. 21). The morphology of the last group was very difficult to determine due to the smallness of the chromosomes. The pairs number 20, 21 and 22 are the smallest of the set and it was sometimes difficult to decide whether the constriction was really subterminal or median. The H. hirta strain with 2n = 30 chromosomes has a diploid set consisting of the following groups: (a) five “V-shaped pairs (N1, 4, 6, 7, 8) with equal arms; (6) one “V”’-shaped pair with unequal arms and two constrictions; (c) six “L”-shaped pairs; and (d) three short rod-shaped pairs. In comparing this set with the former strain (2n = 44) we can note without difficulty a striking similarity between the two sets. The 2n = 30 strain simply has a set of 2n = 44 strain chromosomes less seven definite pairs. These missing pairs are: (a) N3 and N5 pairs from “V”’-shaped groups with equal arms; (5) the “S’’-shaped pair: (c) the smallest N16 pair from the “L’’-shaped group; (d) three pairs N18, 19, 21 are absent from the short rod group. In other words, all the chromosomes in the above group which have median constrictions are absent, and all the short rod-shaped chromosomes with subterminal constrictions are present. As a result of this study it was found easy to recognise roots of H. hirta, 2n = 30 strain, without counting the chromosomes. The metaphase plates were better spread and the chromosomes less crowded; the absence of the “S’’-shaped pair (N9) was easily noted, Karyological Studies and Chromosome Numbers in 43 Hyparrhenia aucta and H. hirta. also the “V-shaped pair with unequal arms; and finally the presence of only 3 short rod-shaped pairs—these are the leading distinctive features. The “‘S’’-shaped pair is such a characteristic feature that it deserves some additional remarks. Many dozens of metaphases of the strain with 2n = 30 from 3 localities were investigated but the “S’-shaped pair was found only once (see Text-fig. 1; 3). In this metaphase it was not very distinctly expressed. The number of chromosomes was the same, 30, so the only explanation of its appearance was to suppose that it had developed from some pair of “L’’-shaped chromosomes. As this occurred only in one single metaphase it could not be decided which pair of “L’’- shaped chromosomes has this tendency of transformation into ‘‘S’’- shaped chromosomes. Probably the “‘S” shape of this chromosome has a mechanical condition such that it lies in the metaphase plate more frequently with one of the short arms perpendicular to the plane of drawing. This would mean that it could be taken for a “L’’-shaped or even for a rod-shaped chromosome. The single occurrence in the slides of the 2n = 30 strain would probably represent a 1 per cent frequency or perhaps even less. With this reservation the absence of “S’’-shaped chromosomes in 2n = 30 strain can still serve as a useful differential character, but it must be compared with the other two mentioned above, especially when only 3 short rod-shaped pairs are present. It is rather difficult to define distinct morphological differences between the strains 2n = 30 and 2n = 44. The most distinct ones are perhaps: (1) a bluish colour of the leaves (as opposed to green in the strain 2n = 44), (2) white nerves on the leaves (as opposed to green nerves in the 2n = 44). On the other hand it must be remembered that the chromosomic races and strains do not necessarily differ in morphological characters. The observation on Bromus inermis recently published by Hill and Meyers (5) in which there are many chromosomic but only two morphological strains is an illustration of such behaviour. An even more striking example is provided by Poa pratensis with numerous strains ranging from 2n = 28 to 2n = + 124. Amongst all these strains conspicuous morphological distinctions do not exist. They differ rather in physiological properties (Kramer, 17). A final illustration is provided by Meyers and Hill (23) whose estimation of the number of ‘intraspecific’? chromosomic races in the Gramineae is as high as 99 (from this estimate Poa pratensis was excluded). : DISCUSSION. Hyparrhenia aucta was included in cytological investigations of Hyparrhenia hirta mainly for the reason that this species commonly te The Journal of South African Botany. grows nearby or mixed with H. hirta. (For the same reason a count of the chromosomes of Schizachyrium semiberbe (Text-fig. 1: 5) is included as well. In this species 2n = 50 was found in adventitious root tips.) There is, however, no close karyological similarity between these two Hyparrhenia species. Firstly, the chromosome numbers differ widely; secondly, the diploid chromosomes of H. aucta are longer and a little wider than those of Hi. hirta. (Compare Text-fig. 1; 1 with 1; 2. 3). (Note: The magnification of fig. 1 is + 1,400x and fig. 2 and 3 + 2.800x.) If one ignored these essential differences for a moment, one would find some similarities between the H. hirta strain 2n = 44 and H. aucta in the following pairs of chromosomes: (1) The “S” chromosome pair N9 of H. hirta is similar to H. aucta N7; (2) the biggest “L” of H. hirta (N10) is comparable with H. aucta N6, with one important reservation, viz. that the distribution of 2 constrictions is not completely identical: (3) the single short rod pair with median constriction in H. aucta is similar to the corresponding pair of H. hirta, probably to N18. Moreover, there are some similarities between two pairs of “L”’- shaped chromosomes with median constriction to pairs cor- responding in height (N16 and 14) of H. hirta. Regarding this comparison with critical reserve it would appear that only the similarities between “‘S”’- and the short rod-shaped chromosomes would stand, though they differ very much in size. A distinct similarity in only two pairs is certainly not enough to establish affinity in morphology in complements of chromosomes. Gluckmann confirmed the writer’s finding of a haploid number of ten in H. aucta but did not observe any irregularity in meiosis of P.M.C. (unfortunately her drawings were not preserved). These facts probably are sufficient to establish that H. aucta is a piploid species with the basic number ten. This would be in accord with the general estimation of basic chromosome numbers in Gramineae and especially in the tribe Andropogoneae. Avdulov (2, p. 82—83) expressed as his opinion that the basic numbers in Andropogoneae are 9 and 10. In his diagram of the frequency of basic numbers in the mono- cotyledons the basic number 10 has the highest frequency: it occupies the second place (Avdulov. p. 328) after the basic number of 12. In Darlington and Janaki-Ammal’s Chromosome Atlas (6) this numbe> is given for many Andropogoneae. The acceptance of 10 as a basic number for H. aucta will involve some inconvenience in regard to H. hirta chromo- somic valency. Karyological Studies and Chromosome Numbers in 4 Hyparrhenia aucta and H. hirta. 1 The existence of 2 forms in H. hirta in the Transvaal and on the Rand was known long ago to Prof. Phillips and his co-workers. It was difficult to define the morphological differences between the two strains as there are overlapping forms. For this reason both forms have been fixed from the inception of this work but unfortunately the fixed material of the 2n = 30 strain, which was collected at the Frankenwald Experiment Station and which was provisionally described as “‘tall blue from Frankenwald”’’, has been spoilt by accident in two consecutive years in an overheated thermostat. For this reason the first count concerned the strain 2n = 44. On account of this occurrence Garber’s (11) publication achieved priority. The above author mainly described chromosome numbers of different species of Sorghum and only incidentally included H. hirta as a member of the same tribe of Andropogoneae by way of illustration. He investigated meiosis only and found the haploid number to be 15. His publication reached South Africa in 1947 and soon afterwards the count of 2n = 30 in root tip metaphases of the “tall blue” strain was confirmed. The American author was asked to send seed of his strain and to give any information about its origin. In due course information from Mr. B. P. Robinson was received, who kindly supplied a sample of seeds. From this correspondence it appeared that the strain of H. hirta was introduced to the U.S.A. from two institutions in the Union of South Africa in 1941, namely: (1) from the McGregor Museum, Kimberley; (2) from Frankenwald University Experimental Station. Mr. Robinson wrote that it was impossible to state from which of the two places the strain, which Garber and he himself investigated cytologically, originated. There was no evidence in the U.S.A. of differences registered between them. Thus far it has not been possible to obtain seed or further informa- tion about this strain from the McGregor Museum, Kimberley. Neverthe- less, it has been thought reasonable to assume that the Kimberley strain also has a number 2n = 30. If this is not true, there should be some evidence in American institutes of registered differences, or of the failure of germination of the seeds. Soon afterwards good root tip material of both strains was obtained from the U.S.A. and from the “‘tall blue” form from Frankenwald. It appeared that both strains are karyologically identical, as the diploid number in both was the same (2n = 30). Morphologically both strains also were identical. In a quick investigation of meiosis of the “tall blue” form no irregularities were found. 46 The Journal of South African Botany. Another strain of H. hirta, investigated from two localities (1. Wit- watersrand University: 2. Frankenwald Experiment Station) was, however, karyologically quite different. The root tip metaphases were very crowded: the chromosomes overlapped frequently and counting was extremely difficult. The diploid number consistently was 44 from all investigated localities (see the list of strains given later). There were few rare instances in which the number found was 44 + 1 fragment. This indistinct fragment was small, without any constriction. No relationship could be found between it and the other chromosomes. Moreover, during the investigations of meiotic metaphases and diakinesis in the anther (see former chapter) no body corresponding to the above fragment was found in spite of the great instability in the number of bivalents. As to the nature and importance of the fragments we first may briefly consider the general position of this question in the world cytological literature where it has been discussed for about 30 years. Broadly speaking, there are three types of supernumerary chromosomes and fragments: (1) The supernumerary B-chromosomes, described by Ostergren (33) and Lima de Faria (20), both in Anthoxanthum; (2) “Standard” fragments described in Rye by Miintzing (26). These are fragments of the size of the smallest normal chromosomes, in any case they are no bigger: (3) “iso-fragments” (Miintzing, 26), much smaller than the smallest of the normal chromosomes and usually smaller than the “standard fragments’. The B-chromosomes are usually heterochromatic and when present in some plants they appear during mitosis in every organ. The “standard fragments” in rye are smaller than the smallest of normal chromosomes. They have distinct subterminal kinetochores. They are separated in meiosis and later exhibit non-disjunction, which means both daughter fragments go to one cell. The “‘iso-fragments” are the smallest but they still have a distinct kinetochore and are easily recognised in rye by a delay in the separation of the chromatids, which have four free ends during mitotic metaphase. The iso-fragments have little genetical effect in rye when they are single or double, but in plants in which they accumulate in larger numbers they produce quite distinct negative effects (Miintzing, 25, Popoff, 34). In particular they are able to cause a decrease in fertility, as shown both in a reduced ability of seeds to germinate and in a smaller output of seeds. Miintzing (1.c.) has shown their deleterious effect on pollen grain germina- tion. In view of the above remarks in connection with the rare fragments in H. hirta strain 2n = 44 it is clear that the Hyparrhenia fragments do not fit the above types. Karyological Studies and Chromosome Numbers in 47 Hyparrhenia aucta and H. hirta. Firstly, they are never heterochromatic; secondly, they are deprived of a kinetochore; thirdly, they are not constant, and fourthly, their participation in mitosis has not been proved: As to the origin of the supernumerary chromosomes and fragments there are two prevailing views. The first attempts to explain their appearance by duplication of one of the normal chromosomes or one of its arms, for example, Zea mays (Miintzing, 25). The second view is based on the fragmentation of the normal chromo- somes as an origin of fragments. This view has its extreme expression perhaps in Chakravartis’ paper (5), where this author explains the origin of many new species, especially those with aneuploid numbers, in the families Cucurbitaceae, Musaceae, Zingiberaceae, Cannaceae, Scitamineae, by transverse fragmentation of normal chromosomes in place of secondary constriction. It is quite possible that the rare fragments in H. hirta have a similar origin. This can, however, be expressed only as pure conjecture as there is no evidence either of shortening of normal chromosomes or of direct observation of fragmentation. The nature of H. hirta fragments and their origin are quite obscure, but it is not thought that they play any important part in the heredity of Hyparrhenia. From the above it seems extremely improbable that H. hirta and H. aucta should have the same basic number, namely ten, and still more improbable is Garber’s proposition of a basic number of five. Thus far no distinct affinity between chromosomes of H. aucta and H. hirta has been found, which may be due to the fact that the plants have different basic numbers. We can reasonably assume a basic number of 10 for H. aucta and of 15 for both H. hirta strains. The basic number 15 is one of least frequent numbers in Monocotyledons (see Avdulov, p. 398, and his discussion of frequency of chromosome numbers). It has a fourth place in rarity, after ““4’’ as the rarest number, and 17, 18. On the other hand, its frequency is about the same as the basic number 5 proposed by Garber (l.c.).. We have already examples in Gramineae of a basic chromosome number of 15 in the tribes Paniceae (Pennisetum 2n = 45), Oryzeae (Darlington and Janaki-Ammal, p. 325) and in Stipeae (L.c. p. 350—351). There are further not less than 7 species with 2n = 44. For the latter the author proposes a basic number of 11. This, however, is not acceptable for H. hirta for the following reasons: Firstly, it would not explain the diploid number 30 in the strain “‘tall, blue” from Franken- wald of the same species; secondly, it would mean that the 2n = 44 strain of H. hirta is tetraploid. The low fertility in the above strain and the frequent variation in numbers of bivalents in meiosis do not support this 48 The Journal of South African Botany. presumption. Apparently, there is no agreement on the question of the most frequent basic chromosome number in Gramineae. Wanscher (44) considers the most frequent basic number in this family to be n = 7, which is present in 89 species. Next in order of frequency comes n = 14 with 76 species, n = 10 in 10 species and n = 5 is present only in | species. The tribe Andropogoneae according to Nielson (30, p. 370—3871) definitely has a prevailing basic number n = 10 with few exceptions, but n — 59 is not even considered at all by this author. In the writer’s opinion the only reasonable choice is to assume 15 as a basic number, at least temporarily until such time as present in- adequate knowledge of the chromosomes of the genus Hyparrhenia has been enriched by further investigations. On the basis of this assumption we can come to two further conclusions, viz.: (1) In the genus Hyparrhenia there are probably two lines of development; one with a basic number of 10 and the other with a basic number 15: (2) in H. hirta there are two strains; one diploid, with high fertility and 2n = 30 and one triploid monosomic, much less fertile, with 2n — 44. The weights of mature seeds give some support to the above assumption. It has been generally stated that the seeds of polyploids are heavier than the seeds of diploids. Thus it was noted on the 16th of June, 1944, that the average weight of 100 seeds of the 2n = 30 strain was 60 mgm. and of 100 seeds of the 2n = 44 strain 75 mgm. The mature seeds for this test had been collected 2 months earlier and selected by hand using a magnifier. The increase in weight represents a difference of 20 per cent. In spite of great variations in weight over the seasons a one- year estimation could serve as a useful illustration of the polyploid nature of the 2n = 44 strain. The figures for germinability of the seed also fit in well with the above assumption. In germination experiments with the H. hirta strain 2n = 44 rather low percentages were obtained. The highest for many years was 54 per cent, though the usual rate lay between 30 per cent to 40 per cent and very often was less than 30 per cent. The germination capacity of the H. hirta strain 2n = 30 in the U.S.A. is much higher. As stated by Robinson and Potts (37) it is as high as 85 per cent and the average is 75 per cent. Germination tests of the pure seeds of 2n = 30 strain from Frankenwald never gave results as high as the foregoing. Following on some observations on the time of opening of florets and shedding of pollen grains the following tentative explanation of this behaviour is put forward: Both strains of H. hirta are growing at Frankenwald in close proximity. The flowers open and shed pollen about the same time and thus the chances of inter-strain pollination equal those of intra-strain pollination. These two kinds of pollen grains are completely different in their chromosomic content and fertilisation value. Practically all pollen grains NATIONAL E L NSTITNUTE PRIVA r = AO PRETC 04 EPUBLIC OF SOUTH AFRICA Karyological Studies and Gireniesome Wayilbore Ho 49 Hyparrhenia aucta and H. hirta. of the strain 2n = 30 have 15 chromosomes and are highly compatible in fertilisation, while of the pollen grains coming from the 2n = 44 strain none has a compatible number of chromosomes. Nearly half of them have n = 22, and then follow the numbers 24, 18, 16 and 14. All these are even numbers with few exceptions. To be compatible for the female gamete in the plant with 2n = 30, the foreign pollen grain should at least have an odd number. Rich co-pollination with the pollen grains shed by 2n = 44 plants with the 15 chromosome type is probably the main cause of the lower fertility of this strain under local conditions as compared with high per- certage germination of the same strain in the U.S.A. The strain of H. hirta 2n = 30 introduced from South Africa to the U.S.A. found itself there artificially isolated from “‘contamination” by related pollen, as there are no indigenous Hyparrhenia species in the U.S.A. Both strains introduced were apparently of the same chromosomic value (2n = 30) and were therefore pollinated only with pure pollen grains containing n = 15 chromosomes. This then is the chief cause of a much higher percentage of germination and higher fertility. In connection with the above assumption it would be of interest to know something more definite about the geographical distribution of both strains in the Union of South Africa. Unfortunately, the writer can make only a very modest contribution to this question. Strains from the following localities were the only ones investigated :— 1. From the Experimental Station, Rietvlei, near Irene, Transvaal &,4 ae ae Be emo ot 2. Grootfontein Reperanen Station, Middelburg, Cape... 2n = 30 3. Experiment Station, Dohne .. ss <3 .. mn=44 4. From Jaegerbush, Losberg se ae .. g2n=44 5. From Jaegerbush, Losberg, 1 plant one on ee Tie — eo) 6. From the Frankenwald Experiment Station .. eerie 130) 7. From the Frankenwald Experiment Station .. 2n = 44 8. Witwatersrand University, Milner Park, J ohenmesbure 2n = 44 In addition to the above list of localities there is reasonable evidence to suppose that the Kimberley strain sent in 1941 to the U.S.A had probably 2n = 30 chromosomes. (See above for the reasons for this supposition. ) The above findings are certainly not sufficient for making any assumptions about the geographical distribution. It is not even known which strains are predominant in the various localities, but it could be supposed that the samples were sent to the author on the ground that one type of Hyparrhenia appeared to differ from the common type or because it was dominant in the region. 50 The Journal of South African Botany. In view of this scarcity of information the author feels it would be wise to limit his statements only to the conditions prevailing in the Trans- yvaal, especially around Johannesburg, where both strains grow practically in the same habitats. The 2n = 44 strain of H. hirta is much more frequent, while the H. hirta 2n = 30 strain prevails rather on old vleis. In addition to the above some details of the technique of counting should perhaps be added. From each locality not less than 10 roots usually belonging to 10 different plants were investigated. In this way each statement on chromosome numbers from different localities is based on not less than 10 slides, with numerous metaphases. The roots of H. hirta 2n = 44 from the University and Frankenwald and the strain 2n = 30, the “‘tall blue’, from Frankenwald and that from the U.S.A. were exceptions to this rule. as with different methods of squashing, staining and fixation more than 100 roots of the strain 2n = 44 and 30 to 40 roots of the 2n = 30 strain were studied. In spite of these numbers the amazing uniformity, with one single exception, was striking. Only in the plants from Jaegerbush, Losberg, a metaphase with 2n = 56 was found on one occasion. This root belonged to the plant with 2n = 44 chromosomes. Unfortunately, from one crowded metaphase nothing could be learnt about the morphology of the chromo- somes. That was probably a case of a mutant root. Mention has already been made of the crowdedness of metaphases in the H. hirta strain 2n = 44 and the difficulty of counting them. In order to spread the chromosomes better and make the counting easier experiments were undertaken with pre-treatment of the roots with cold treatment before fixation (see Darlington and La Cour (7), Delaunay, 9). There was also another reason for trying this method. In the course of previous years there was some evidence in cytological literature that cold pre-treatment would reveal a third kind of constriction in chromo- somes (Resende and Cabral, 35). which would be constant for given species and given chromosomes. One of the investigators—F. Resende—even proposed to give them the special term “Olistherozones”’ (Nature, v. 144, p- 481 and Acta Portugaliae Biologica, Vol. 1, fase. 2, 1945). For these two reasons methodical pre-treatment at a temperature of 4°C. was given to root tips before fixation. The roots were fixed at intervals of one hour starting with a treatment of one hour and increasing up to 24 hours. The shortening and spreading of metaphasic chromosomes was at a maximum after 4—6 hours pre-treatment. However, the added advantage of better spreading and shortening of chromosomes was con- spicuously reduced in value by the peculiarity that they were now dis- tributed in many planes, sometimes very distant ones. The danger of omission in the counting of some chromosomes occurring in different Karyological Studies and Chromosome Numbers in 51 Hyparrhenia aucta and H. hirta. planes is obvious. With longer pre-treatment the chromosomes became crowded, and with still longer treatment sticking occurred, while after 24 hours the metaphases were shrunk, exhibiting indistinct clumps of chromatin with frequent bridges in anaphases. With regard to the new tertiary constrictions, the so-called ‘‘Olis- therozones’’, the results of our experiments were completely negative. This is in line with previous experiments on Senecio isatideus (Krupko and Goldsmith, 13). In this instance constrictions which were visible without pre-treatment disappeared completely. This was not the case, however, with H. hirta. The kinetochores and secondary constrictions are clearly seen when maximum spreading has been achieved, i.e., after 4—6 hours of treatment, but new constrictions were not revealed. The temperature of 4°C. was chosen for these experiments because Delaunay claimed that he obtained the best results at that temperature with the majority of plants. It is quite possible that H. hirta would respond differently at different temperatures. For the same purpose soaking in cold water before fixation was also tried. This gave some improvement in the spreading of chromosomes after 2—3 hours of treatment (see also Goldsmith and Krupko, 13) which was just the reverse of Gluckmann’s results with Themeda (unpublished). Considering the H. hirta strain 2n = 44 as a monosomic triploid plant we could expect that the majority of chromosomes, in somatic metaphases at least, would be found not in homologous pairs but in homologous triplets. In order to check this aspect of the karyogram an attempt was made many times to arrange the drawings of morpho- logically similar chromosomes, cut out of the paper in triplets, but this invariably failed. Only 6 rod-shaped pairs could sometimes be arranged in triplets, but this was without significance because of the smallness of the rod shaped chromosomes. Also there was great uncertainty in deter- mining the nature of the constrictions in these chromosomes. It has been pointed out that it was often difficult to decide which type of con- striction was present, namely, subterminal or median. It is usually easy to arrange all other chromosomes in pairs. It is supposed that this aspect of the karyogram has some theoretical significance with regard to the character of triploidy. It probably means that triploidy in this strain was achieved very long ago. The H. hirta strain 2n = 44 is an old triploid species in which triplets have had enough time to adapt themselves to the diploid physiological function, and become pairs. It is possible that in early age this strain was a simple triploid with a full complement of 45 chromosomes. For some reason unknown at present one chromosome of the complement was gradually excluded 52 The Journal of South African Botany. in disjunction, and the strain finally became a monosomic triploid. If this is so the fragment mentioned previously which occurs sometimes in root metaphases may be the relic of the odd chromosome. ADDITIONAL OBSERVATIONS. During this work on Hyparrhenia hirta some additional observations were made not directly connected with the cytology of the plant, but probably being of some interest in the expansion of our knowledge of South African grasses. First may be mentioned observations on the dichogamy of Hyparrhenia florets, which were carried out on the H. hirta strain from the Grootfontein College of Agriculture which has 2n = 30 as described above. The spike- lets and florets were marked with coloured wool threads. In this way the observation was soon made that there is no protandric behaviour at all, contrary to the prevailing statements of German ecologists in many text- books of Ecology. Nearly all of them state that protandry is the prevailing condition in the Gramineae family (in Europe). Apparently it is not the case in H. hirta, at least in the strain from Grootfontein. In this strain protogyny is the prevailing condition. The stigmas appeared early in the morning, sometimes between 8—10 hours. They first had a white colour which was maintained during the first day, but the next day changed to a light yellow. On the second morning the stigmas were coloured a deep red and this colour was maintained, gradually changing to violet-red and ultimately to black-red. Quite possibly this change of colour is connected with pollination or even with the fertilisation process. Similar observations of protogyny were made by Artschwaeger and Maguire (1) in Sorghum vulgare, though they did not mention the colour of the stigmas in the plants. A remarkable fact is that the florets remain open for the very short time of 30—60 minutes. In addition to the protogynic behaviour, cases of simultaneous appearance of stigmas and anthers were noted in H. hirta. The second observation is connected with the number of xylem strands in the primary roots. In H. /irta the vascular bundles are all pentarch, without any exception. It may be of value to compare the statements on the same subject in current anatomical literature. Mostly this item is ignored, or there are only observations on cereals, from which a general idea of the structures of other grasses are drawn. For example, Avery (3) mentioned that the primary roots of Triticum are hepta-octarch and according to this author maize roots have even more “‘Tays”’. The third observation concerns the structure of growing points of adventitious roots. In H. hirta all four histogens are present (see Pl. XIV, fig. 15, Pl. XV, fig. 21). It exhibits a classical picture of the Monocot Karyological Studies and Chromosome Numbers in 53 Hyparrhenia aucta and H. harta. structure: the calyptrogen gives rise to the calyptra, the dermatogen to the epidermis, the periblem to the cortex, and the plerome to the central cylinder. As said before, there is a tendency amongst grass growers to see the structure of cereals as a model for all grass structures. In this case such a generalisation would be completely wrong. According to Percival Nelson (29), Hector (14) and Standford (40) the cereals have only three histogens, cortex and epidermis having one common histogen for both tissues. In this respect again H. hirta differs from the cereals. Avdulov (2, pp. 324—325) attached special attention to the shape and to the position of the first leaf in the grasses. In his opinion Hackel’s (1889) statement that a lanceolate or elliptic first leaf in the Andropogoneae with a more or less horizontal position is a primitive character as com- pared with linear or awl-shaped leaves, as prevailing amongst other grasses, should be extended to the other tribes. He found (l.c., p. 325) the same shape and character of the first leaf in all 6 tribes of Series A in the Gramineae (according to Hackel’s system), which he calls the subfamily Sacchariferae. Moreover, when this shape occurs in other species of Series B of the family it is invariably connected with primitiveness of characters or with the relic nature of that species. In the light of the above, Avdulov’s interpretation is useful perhaps in giving a picture (PI. XV, fig. 24) of the first leaf of H. hirta. It is distinctly elliptical and it keeps a practically horizontal position, at least in the beginning. Its margin is ciliated. Unfortunately, the photograph included fails completely to reproduce ciliation of the margin, which however can easily be seen with a xX 10 magnifier. In this respect the first leaf of H. hirta agrees with Hackel’s rule for the tribe Andropogoneae and with Avdulov’s extension of that rule to the subfamily Sacchariferae. The fifth observation is connected with the great difficulty experienced in squashing fresh root tip sections of H. hirta for quick staining and counting of chromosomes. Schlerenchymic tissues in root tips were sought in vain but very soon it was apparent that resistance to the squashing force is due to the thick external wall of the epidermis (see Pl. XV, fig. 22). The thickness of this wall is frequently equal to the diameter of the lumen of the epidermis cells and sometimes it is twice as thick as the diameter. This external wall stains distinctly with 1 per cent Congo Red, revealing its cellulosic nature. The same wall may he stained pink with ruthenium red, the colour appearing evenly on the whole thickness of the wall without striations or layers appearing. The impression was gained that the external layer of the wall is a solid cellulosic membrane. Below this wall there is probably a dense elastic layer of cellulosic and pectic slime, highly resistant to pressure. This extremely thick wall covers only the meri- 54 The Journal of South African Botany. stematic region and the more differentiated parts of the root have an epidermis with a wall of solid cellulose of usual thickness. RECOMMENDATIONS. Having in mind the ecological importance of the genus Hyparrhenia in Africa and the possibility of the use on a wide scale of some of its species in soil conservation measures, the author regards it as very important that the following lines of investigation will be carried out: (1) Counting of chromosome numbers and analysis of meiosis of all species of Hyparrhenia in Africa; (2) chromosome counting and chromosome morphology of H. hirta strains from different countries (up to the Red Sea); (3) collection of facts on the geographical dis- tribution of chromosomie strains of H. hirta in the Union of South Africa, based on direct counting of chromosomes in plants from different localities, but not based on the external morphological characters; (4) experimental plots of the diploid strain of H. hirta should be arranged in different parts of the country with strict isolation of the plots from pollination by other strains of H. hirta or by other Hyparrhenia species. It is the writer’s hope that these plots would prove the greater fertility of the diploid strain in isolated conditions. SUMMARY. There are probably at least two lines of development in the genus Hyparrhenia: one with a basic chromosome number = 10, and the other with a basic number = 15. Hyparrhenia aucta has the following chromosome numbers: 2n = 20; n = 10. Its basic number is probably 10. Hyparrhenia hirta has two strains, diploid with 2n = 30, n = 15, monosomic triploid with 2n = 44. The latter has various numbers of bivalents in pollen grain mother cell meiosis: 22, 20, 18, 16, 14. Laggers are frequent and persisting up to the 1-nucleated stage of the young pollen grain. The basic number of chromosomes in H. hirta is probably 15. Of the mature pollen grains of the above-mentioned strain + 20 per cent is degenerated. Normal looking pollen grains germinate very rarely and with difficulty. The tubes are short, growing slowly and are short lived. There are some mature pollen grains with supernumerary nuclei. The above disturbances is pollen grain development and the unbalanced number of chromosomes in male gametes are probably the chief causes of low fertility of the monosomic triploid strain. Schizochyrium semiberbe has 2n = 50. Karyological Studies and Chromosome Numbers in 55 Hyparrhenia aucta and H. hirta. ACKNOWLEDGMENTS. From the time of my first appearance at the Witwatersrand University I was received with great hospitality. Many marks of cordial sympathy and of friendly help have been shown to me by my scientific colleagues, both in the botany department and in other specialities. It is my pleasant duty to thank them all here, both those whom I still see and remember and those who are now working at other places and whom I perhaps have forgotten. I am deeply indebted to Professor J. V. F. Phillips, former head of the Botany Department, University of the Witwatersrand, for suggesting this investigation on grass cytology. During the war years there were many difficulties and technical shortages which threatened to handicap con- siderably this kind of work, but Professor Phillips’ energy and constant support overcame all obstacles. His enthusiastic approach to the prob- lems of soil conservation and grass investigations in this country was a constant source of inspiration. I am greatly indebted also to the Department of Agriculture for research grants, which were received through the Frankenwald Botanical Research Station. My special thanks are further due to the following persons: Professor N. P. Badenhuizen, the present Head of the Department of Botany, for his support in my research endeavours and his interest in them; Mrs. M. Moss, retired Senior Lecturer of the Botany Department, for cordial help in my first days in this University and for constant encourage- ment in my teaching work at this Department; Dr. D. B. D. Meredith for his keen interest in my work on grasses and for his help in arrangement of experiments at Frankenwald and collection of material, for supplying me with current literature and lastly and perhaps most important, for helping to correct the present paper; Dr. H. B. Gilliland, Senior Lecturer of the Botany Department, for naming of many grasses and for useful advice; Dr. E. R. Roux, Senior Lecturer of the Botany Department, for much useful advice; Dr. R. L. Davidson, Lecturer of the Botany Depart- ment and Deputy-Director of Frankenwald Experimental Station, for help in the arrangement of my experiments, for collection of material and for assistance in preparing photographs; Mrs. Crook (formerly Miss Chippindall) for the naming of many grasses. Many thanks for preparing microphotographs are due to Mr. Armando Salbany from Mozambique, to Dr. Poultney, formerly at Frankenwald and to Mr. G. Stiven. I thank also Dr. R. E. Altona of African Explosives and Chemical Industries, Limited, for the care of my experiments at Frankenwald. The Journal of South African Botany. The generous contribution made by the University of the Witwatersrand to the cost of printing this paper is gratefully acknowledged. Finally, I thank all these Experimental Stations, Government Institutions, Agricultural Officers, here in the Union and in other countries in Africa, who kindly sent me seeds and samples of grasses. Botany Department, University of the Witwatersrand, Johannesburg. BIBLIOGRAPHY. ARTSCHWAEGER, E. anp R. C. Macurre. 1949. Cytology of reproduction in Sorghum vulgare. (Jour. of Agricultural Research, vol. 57, No. 12, pp. 659—673). AvputLov, N. P. 1931. Karyo-taxonomical investigations in Gramineae. (In Russian). Leningrad. (Suppl. 34th to the Bull. of Applied Botany, Genetics and Plant Breeding). Avery, G. 8. 1930. Comparative anatomy and morphology of embryos and seedlings of maize, oats and wheat. (Bot. Gaz. vol. LXX XIX, No. 1). Brink, R. A. 1925. The influence of hydrogen ion concentration on the pollen tube of the sweet pea, Lathyrus odoratus. (Amer. J. of Botany, vol. XXX, p. 149). CHAKRAVORTI, A. K. 1948. Theory of fragmentation of chromosomes and evolution of species. (Science and Culture, vol. 13, No. 8). DaRLiIncton, C. D. anp JANAKI Ammat, BE. K. 1945. Chromosome Atlas. London. 1945. Darineton, C. D. anp La Cour, L. 1940. Nucleic starvation of chromosomes in Trillium. (Jour. of Genetics, vol. 40, No. 1—2). DeLaunay. L. 1926. Zeitschrft. Zellf. Mikr. Anat. v. 4, 338—364. Denaunay, L. N. 1931. Das verkurzen der chromosomen involge der Abkithlung. (Ber. Inst. Pflanzenzticht Maslovka, 4). Faperceé, A. C. 1945. Snail stomach cytase, a new reagent for plant cytology. 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Genetical effects of duplicated fragment chromosomes in rye. (Hereditas, vol. 29). Munrzine, A. 1946. Cytological studies of extra fragment chromosomes in rye. III. The mechanism of non-disjunction at the pollen mitosis. (Hereditas, vol. XXXII, 97—119. 1946). Nawasuin, 8. 1915. (Bull. Acad. Imp. Sci. St. Petersburg, v. 9, pp. 1821—1834). Nawasuin, M. 1925. (Genetics, vol. 10, 583—592.). Newtson, A. 1946. Principles of agricultural botany. Thos. Nelson & Sons, Ltd., London & Edinburgh. Nietsen, E. L. 1939. Grass studies III: Additional somatic chromosome complements. (Amer. J. of Bot., vol. 26). Newcomer, H. EH. 1938. A procedure for growing, staining and making permanent slides of pollen tubes. (Stain Technology, vol. 13, No. 3. July). O'Mara, I. G. 1948. Acetic acid methods for chromosome studies at pro- phase, metaphase in meristems. (Stain Technology, vol. 23 (4); 201—204. 1948). OSTERGREN, G. 1947. Heterochromatic B-chromosomes in Anthoxanthun. (Hereditas, vol. XX XIII, p. 261). Poporr, A. 1939. Untersuchungen uber den formenreichtum und die Schartigkeit des roggens. (Angewandte Botanik, Bd. 21, H.4., pp. 325—356). RESENDE, F. anp A. Capray. 1944. Sur la structure de chromosomes dans les mitoses des meristemes radiculaires III. L’action de la temperature sur le structure chromosomique. (Acta Portugaliae Biologica, vol. 1, f. 1, 1944). RESENDE, F. 1945. Heterochromatine. (Acta Portugalae Biol. vol. I, f. 2., 1945). Rosrtnson, B. P. ano R. C. Ports. 1950. Seed setting and germination in Hyparrhenia hirta (L.) Stapf (South African blue stem) as affected by nitrogen, phosphorus and potassium. (Amer. J. Agron., Feb., 1950). Sax, K. 1935. The effect of temperature on nuclear differentiation in micro- spore development. (Jour. Arnold Arboretum, vol. 16, 301—310. 1935). Sax, K. 1937. Chromosome behaviour and nuclear development in Tradescantia. (Genetics, vol. 22: 523—533). Stanrorp, E. E. 1939. Economic Plants. (New York: London, 1939). Appleton-Century Co. Taytor, W. R. 1925. Chromosome constructions as distinguishing characteristic in plants. (Amer. J. Bot., vol. 12, p. 238). Taytor, W. R. 1926. Cytological studies on Gasteria II: A comparison of the chromosomes of Gasteria, Aloe and Haworthia. (Amer. J. Bot., vol. 12). Urcotr, M. 1939. The external mechanics of chromosomes. Its abnormal mitosis in pol. grain. (Chromosoma, vol. | (2): 178—196). WanscuHer, J. H. 1934. The basic chromosome number in higher Plants. (The New Phytologist, vol. XX XIII). Warmke, H. E. 1941. (Stain Technology). OL The Journal of South African Botany. PLATE XIV. Fic. 1 anp Fic. 17.—H. aucta, root tip metaphase—2n = 20. Fix. Levitsky (5:5) with 4% formol. Stained—Iron haematox. Thickness—7 pp. Photo. taken with B. & L. apochr. x 90 oil immers. and Comp. oc. x 10. Yellow filter. Magnification + 1150 x. Fies. 2 To 7, 6 To 11, 16.—H. hirta, strain 2n = 44. P.g.m.c. Fix.—Carnoy. Stained—Iron haematox. Thickness 8 ». Taken with B. & L. apochromat. < 90 oil immers. and Comp. oc. x 12. Magnification + 1000 x. Fies. 12 ro 14.—H. hirta. Similar strain. Smears, fixed in Nawashin, stained with Crystal Violet after Newton. Mature p. grain. Fie. 1.—Metaphase, 2n = 20. Fie. 2.—P.g.m.c. diakinesis with n = 14. Taken in two levels. For continuation see Fig. No. 3. Seen 5 bivalents and nucleolus. Taken with green filter No. 58 Wratten. Exposure 17”. Fie. 3.—The same pollen grain m.c. another level. Here is visible continuation of nucleolus and of one bivalent indistinct above it. Beneath the nucleolus two bivalents (not seen in Fig. 2). All together one can see 9 bivalents not seen in Fig. 2. Fic. 4.—P.g.m.c. Ist meiotic metaphase with 2n = 22. Taken with green filter No. 58 Wratten. Exposure 20”. Fic. 5.—P.g.m.c. Ist meiosic metaphase withn = 18. Filter the same. Exposure 25”. Fic. 6.—P.g.m.c. Ist meiotic anaphase with numerous laggers. Green filter. Exposure 15”. Fic. 7.—P.g.m.c. Ist meiotic middle telophase with 4 laggers. Taken without filter. Exposure 4”. Fic. 8.—H. hirta, strain 2n = 44. Acetocarmine squash. P.g.m.c. Ist meiotic telophase. Laggers. Taken with green filter No. 58 Wratten. Exposure 23”. Optics and magn. like above. Fie. 9.—Old tetrads with the laggers. Taken without filter. Exposure 4’. Fie. 10.—V.y.p.gr. (gones), 1-lagger. Taken without filter. Exposure 10”. Fic. 11.—The same. Dwarf nucleus—dw. n. nucleus—n. without filter. Exposure 3”. Fie. 12.—Mature p. gr. with 4 nuclei, taken in 2 levels; for continuation see Fig. 13. Yellow filter. Exposure 22”. Fic. 13.—Continuation of Fig. 12. Exposure 5”. Fic. 14.—Mature p.g. with 5 nuclei; two of them indistinct—slide fading. Yellow and green filters. Exposure 35”. Irie. 15.—H. hirta, strain 2n = 30, from Frankenwald. Fix.—Nawashin. Stained— Haematox. + Fast green. Thickness 8 p. Root tip—longitudinal, showing 4 histogens. Taken with B. & L. obj. « 20 Comp. oc. * 10. Magnification + 250 x. Fic. 16.—V.y.p.g. (gones). Four laggers. Without filter. Exposure—10’. Fria. 17.—Like Fig. No. 1. Metaphase in root tips epidermis with numerous ““V’’- shaped chromosomes and “S’’-shaped one. Fic. 18.—H. hirta, strain 2n = 30. Root tip metaphase. Fixation Nawashin. Staining—Crystal Violet. Thickness 10 ». B. & L. Fl. oil immers. x 98,Comp. oc. < 10. . Taken in two levels. Exposure: upper photo 8”, lower one 16”. Magnification + 1200 » Fic. 19.—H. hirta, strain 2n = 44. Root tip metaphase. Fix.—Levitsky (5:5) with 10% formol. 1% Chrom. acid—8 days. Staining—iron haematox. Thickness 7 ». Optics as in Fig. 18. Exposure 16”. Magnification + 1200 x. Fic. 20.—H. hirta, strain 2n = 44, root tip, metaphase. Fix. and staining as in Fig. 18. B. & L. oil immers. x 97. Comp. oc. x 7:5. Final magnification + 3300 x. PLATE XV. Fie. 21.—H. hirta, strain 2n = 30 from Frankenwald. Root tip longitudinal, showing growing point, 4 histogens. Fix.—Nawashin. Staining—iron haematox +- fast green. Thickness 8 p. Taken with B. & L. obj. x 20. oc. « 20., enlarged + 4 times. Total magnification + 1000 x. Fic. 22.-—H. hirta, adv. roots. Growing point longitudinal sect. showing unusually thick ext. wall of epidermis cells. Stained: Congo red + haematox. Thickness 8 wp. Taken with B. & L. obj. 60, ocul. x 10; exposure 2”. Magnification +- 500. ep. = epidermis cell, w = external wall, ca = calyptra cells. Fic. 23.—H. hirta, strain 2n = 44. Enlargement of Fig. 19, Pl. XIV + 3300 x, to show more distinctly the shape of chromosomes. Fra. 24.—H. hirta, strain 2n = 44. Seedlings. PLATE XIV. Karyology of Hyparrhenia. PLATE XV. Karyology of Hyparrhenia. SOUTH AFRICAN SEAWEED VEGETATION AND FUTURE INVESTIGATIONS IN THIS FIELD. By Wm. Epwyn Isaac. (Botany Department, University of Cape Town.) Paper read at the First International Seaweed Symposium, Edinburgh, July, 1952. CONTENTS. PAGE Introduction Bh Bf “is sa0 Gy) Ocean Currents and Sea Renmpenature condinions si 59 General characteristics of the Seaweed vegetation and Aor of ate South African Coasts .. : ae ie de oc cauctOrl The Marine Algae of the West Gees an ar i a: -. 63 The Marine Algae of the East Coast .. si ‘is bit .. 65 Future jou se op x Sg te i ueeOG Summary - wi = af St aa oo OY) Wein uledvementh me ae hs iis an oe se oY) Literature : a “if Bee 0) Appendix: Raxonornie motes with Pines merorentees 56 wa Ol INTRODUCTION. Seaweed investigation presents a more complex and varied field in South Africa than in Britain. In the first place the coastline of South Africa is much more extensive and many parts are not easily accessible. Secondly, and still more important, the range of physical conditions is greater due to the influence of two ocean currents. Two other considerations may be mentioned: the seaweed flora is less well known and only scant information is available concerning sub-littoral seaweeds, especially east of Cape Agulhas. This last consideration is important in relation to the examina- tion of drift material since there must often be uncertainty in sorting out what has been thrown up locally from deeper waters and what may have been carried by currents for a longer or shorter distance. OcEAN CURRENTS AND SEA TEMPERATURE CONDITIONS. The west coast is subjected to the influence of water of sub-antarctic origin which is referred to as the Benguela Current while the east and south coasts are influenced by the Agulhas Current, a warm water current originating about latitude 11°S. off the coast of East Africa, The Agulhas 59 60 The Journal of South African Botany. PORT MOLLOTH PORT ST. JOHN'S EAST LONDON Q-LANGEBAAN LAGOON PORT ALFRED > PORT ELIZABETH CAPE POINT a) FALSE BAY CAPE AGULHAS Fre. 1. Outline of Union coastline indicating the chief localities mentioned in the text. Current flows southwards and westwards until it reaches the Agulhas Bank, which is the only extensive part of the continental shelf around the coasts of Southern Africa. The Agulhas Bank deflects most of the warm water eastwards again but at a higher latitude. Branches of the Agulhas Current, however, move westwards (von Bonde, 1939). Coastal sea temperatures increase in an easterly and northerly direction from Cape Agulhas but they fall in a northerly direction along the west coast of the Union. On the west coast, north of the Union, coastal sea temperatures rise again but even at Walvis Bay just within the tropic of Capricorn, sea tempera- tures are lower than on the east coast. Thus, the average annual tempera- ture of the sea at Durban (21-8°C) is 5-7°C higher than that of Walvis Bay (16-1°C) although Durban is 7° latitude further south (Isaac, 1937a). Between Cape Agulhas and Cape Point the coastal waters are not directly affected by the ocean currents and temperatures are much affected by local factors such as depth of water and topography especially in relation to the south-east winds of summer. Temperature conditions in False Bay are abnormally high in relation to its geographical position. If we except False Bay, however, the region from Cape Point to Agulhas is best regarded as an extension of the west coast. South African Seaweed Vegetation and Future 61 Investigations in this Field. For convenience, in the remarks which follow, the coast west of Cape Agulhas! has been referred to as the west coast and the coast to the east as the east coast, although it would be more correct to distinguish between the south coast from Agulhas to about Port Elizabeth and the east coast, north of Port Elizabeth. By “cold” or ‘“‘colder’”’ waters is meant the waters off the west coast as defined above but excluding False Bay and ‘‘warm’’ or “warmer” waters refer to the seas east of Cape Agulhas. The terms “warm” or “warmer” waters are also applicable to the waters of the northern part of False Bay. Clearly such terms in reference to sea temperatures are relative; this is especially the case with regard to ‘“‘cold’”’ waters (Isaac, 1951). It is inevitable that the existence of contrasting east and west coast sea temperature gradients will result in at least two groups of seaweed floras and vegetation. The contrasts and changes do not present a simple picture since there is overlapping in distribution of the flora and vegetation characteristics between one part of the coast and the next. This results from the fact that species differ in the temperature ranges under which they flourish and in their tolerance to temperatures under which they may occur. In this connection it is well to remember that there is no direct relationship between the range of temperature tolerance and the extent of coastal distribution expressed in miles. A species occurring in a region of relatively rapid temperature change may show a wider temperature tolerance than a species distributed over a longer length of coastline where temperature changes are more gradual. The temperature data available indicate that in general a stretch of west coast will show less temperature change than a corresponding stretch of east coast. GENERAL CHARACTERISTICS OF THE SEAWEED VEGETATION AND FLORA oF SoutH AFRICAN COASTS. In spite of the contrast which exists between the seaweed flora and vegetation of the east and west coasts, the South African seaweed vege- tation and flora have a few general characteristics. A first general characteristic is the relatively limited vertical extent of the inter-tidal seaweed vegetation. This is due chiefly to the limited tidal range and to the paucity of algal growth at higher inter-tidal levels which is in turn associated, at least to some extent, with the climatic conditions of the land (Isaac, 1938). Along the coasts of South Africa the tidal range from low water to high water spring tides varies from about 5 to 6 feet (Bauer, 1933; Admiralty charts). Except locally little 1 Cape Agulhas is the most southerly point of the African continent. 62 The Journal of South African Botany. or no emersed algal vegetation occurs above high water level of neap tides. On the west coast the prominent alga at and some distance below high water neaps is Porphyra capensis. This species is found from Port Nolloth (Bright, 1938) to Port St. Johns but east of Agulhas it becomes very sporadic in distribution, smaller in size, paler in colour and forms less dense and much smaller communities than on the west coast. On the west coast it frequently forms a dense, more or less continuous girdle* persisting throughout the year while at Port St. Johns an hour’s search may yield only a few individuals. The dense and extensive Porphyra communities of the west coast are not replaced by other corre- sponding communities on the east coast which thus presents an impression of even more limited inter-tidal seaweed vegetation. Locally, especially where spray is hurled upwards by rough seas, small pale plants of Porphyra occur above high water of neap tides and even above high water level of spring tides. Again, in damp shaded places a dense growth of various other algae especially Bostrychia occurs at high inter-tidal levels. Such occurrences tend, however, to emphasise the general barrenness above high water neap tides on the west coast, and east of Agulhas the general barrenness above about mid-tide level. In rock pools, especially where fresh water runs into the sea, Ulva and particularly Enteromorpha spp. may be prominent above high water neaps. A second general characteristic of the inter-tidal algal vegetation of South African coasts is the predominance of Rhodophyceae. As: far back as 1893 Murray had drawn attention to the relative richness of species of Rhodophyceae in the seaweed flora of South Africa. Few of the South African inter-tidal Rhodophyceae are a bright red colour although such are not lacking, e.g. Plocamium corallorhiza. The promi- nent warm water species Hypnea spicifera and the much less common H. viridis are usually bright green while the common west coast Aeodes orbitosa is usually brown. Other species (Gigartina radula, G. stiriata, cold water Porphyra capensis) are usually dark, almost blackish-red. The Rhodophyceae, however. over the greater extent of the coast dominate the vegetation at every inter-tidal level. Unlike the coasts of Western Europe. the Fucales are not a prominent feature of the vege- tation. The outstanding exception in this respect is Bifurcaria brassicae- formis which dominates a lower inter-tidal level from Cape Town to Cape Agulhas (Isaac, 1951). Sargassum longifolium is locally common or even, very locally, dominant, in the upper sub-littoral and lowermost littoral levels between Cape Point and Cape Agulhas (Isaac, 1947) and at Dwesa on the Transkei coast. Other species of Sargassum are common in the > The term “girdle” as recommended by Feldmann (1951. p. 321) is used here instead of “zone”. South African Seaweed Vegetation and Future 63 Investigations in this Field. warmer waters east of Agulhas but these plants do not dominate inter- tidal girdles although locally they may form clans of emersed plants as well as sometimes being common in the upper sub-littoral and in rock pools, the vegetation of which they sometimes dominate. Another member of the Fucales, Carpophyllum scalare is abundant or dominant very locally at Port Alfred and at Dwesa. In warm waters, members of the Caulerpaceae may dominate the vegetation or cover areas to the exclusion of other plants in shallow sea or at low inter-tidal levels. This is the case with Caulerpa filiformis in shallow water, usually in sand, in places at Port Alfred (The Kowie) and to a lesser extent in localities from East London to Port St. Johns, as well as in places on the north-west coast of False Bay (part of the St. James coast); and at lower inter-tidal levels Cawlerpa racemosa var. zeyheri occurs in moderately sheltered places where there is swell, for example at Port Alfred, East London and Port St. Johns. Still, such occurrences as we have cited can be regarded as local intrusions into levels which are chiefly dominated by Rhodophyceae along the entire length of the coastline of the Union of South Africa. Brief attention may here be called to those ubiquitous genera Ulva and Hnteromorpha. The distribution of these plants is largely related to salinity variations and they flourish under conditions of both low and high salinity. Thus along the coastline of the Union Ulva lactuca and species of Enteromorpha occur in rock pools high in the inter-tidal area as well as in places where there is a mingling of fresh and salt water. Plants of these genera also occur in places where they remain emersed by the falling tide although they rarely show signs of drying out since they occur in habitats where there is spray or where water is retained due to the presence of numerous small depressions of the rock surface. Of the emersed Ulva, U. lactuca is more characteristic of cold waters, U. rigida flourishes characteristically east of Cape Agulhas, and U. minuta is characteristic of warm waters. The emersed micro-species of Hnteromorpha are a feature of the warmer waters. THe Marine ALGAE OF THE WEST COAST. The contrast between the seaweed vegetation of the west coast north of Cape Town and of the east coast north of Port Elizabeth is obvious and striking but the contrast is less marked for the region between Cape Agulhas and Port Elizabeth (South Coast) and either the west coast or the east coast north of Port Elizabeth. Clearly the region east of Agulhas needs to be subdivided, but before this can be done satisfactorily more data are needed for the stretch of coast between Agulhas and Port Elizabeth. The west coast can be subdivided into two:— 64 The Journal of South African Botany. I. Port Nolloth to Cape Town. II. Cape Town to Cape Agulhas but excluding False Bay. Locally, conditions may deviate to a considerable extent from the con- ditions prevailing in neighbouring districts. This is pre-eminently the case with False Bay which needs to be considered on its own. It is also the case with the Saldanha Langebaan lagoon. The most obvious characteristic of the west coast is the presence of an extensive sub-littoral growth of kelps (Isaac, 1937b, 1938, 1942). These beds extend to depths of at least 25 to 30 feet since the stipes of the largest species, Ecklonia maxima attain these lengths. The fronds and sporophylls of Ecklonia toss about at the surface of the water. Laminaria pallida is a second common species of kelp. Laminaria is a smaller plant than Ecklonia and it lacks a float and hollow stipe which are found in the latter but it may well extend into deeper waters. Precise data on this matter are lacking. A third west coast kelp is a species of Macrocystis which in the past has been named M. pyrifera: the specific identity of this plant, however, is somewhat uncertain.* The kelps are typically sub- littoral plants but in places they occur at the lower inter-tidal levels. Macrocystis is much less abundant than the other two species, being con- fined to a limited stretch of the southern part of the west coast north of Cape Point but locally it may be abundant. It occurs in situations sheltered from the full force of the sea, as in quiet bays or within a palisade of Ecklonia (Isaac, 1937b). Laminaria pallida is more typically a cold water species than Ecklonia maxima. The former kelp is at best a rare species east of the Cape Peninsula* and becomes relatively more abundant in a northward direction along the west coast. Ecklonia occurs from Port Nolloth (Bright, 1938) to Cape Agulhas and it is a prominent plant over the greater part of this range. At its eastern limits it is a relatively small plant and not abundant. In many localities large quantities of kelp are thrown up on the shore and accumulate above tide level. A second outstanding characteristic of the west coast is the presence of a girdle of Porphyra capensis extending downwards from high water neaps. The growth of Porphyra is often dense, P. capensis forming almost pure communities. On the west coast the plants are large, of a dark purplish red colour and the Porphyra capensis community (association) remains without essential change throughout the year (Isaac, 19375: 1942). Taken as a whole the common and characteristic cold water species are relatively large; such are Champia lumbricalis and Aeodes orbitosa. Other large plants found in cold water are Gigartina radula, G. stiriata ’ A paper on South African Macrocystis is in course of preparation. + Unless it is common in deeper water so as not to be visible at the surface. South African Seaweed Vegetation and Future 65 Investigations in this Field. and Codium fragile. Splachnidium rugosum is a larger plant and more abundant on the west coast than in the warmer waters east of Agulhas. Small species are also found, e.g. Caulacanthus ustulatus, which is widely distributed in cold and warm waters. The outstanding difference between the west coast north of Cape Town and the part from Cape Town to Cape Agulhas is that the “fucoid’’ species Bifurcaria brassicaeformis is a prominent species of the latter where it dominates a girdle in the lower part of the inter-tidal region. It forms dense communities (associations) to the practical exclusion of other plants (Isaac, 1951). Over the greater part of this sub-region Gelidiwm pristoides also occurs. THe Marine ALGAE OF THE Mast Coast. The east coast seaweed vegetation presents a marked contrast to that of the west coast in three outstanding respects:— (1) The absence of large sub-littoral algae reaching the surface of the water. (2) The absence of a Porphyra girdle although, as indicated earlier, P. capensis extends at least as far north-eastwards as Port St. Johns. (3) The seaweeds are in general of smaller size than those of the cold waters and dense mat-like or sward-like mixed communities of small algae are a prominent feature. As a result of these characteristics the coasts of the warmer seas have a much more barren appearance than those of the colder seas at high water of all tides and at low water of neap tides. At low water of spring tides, however, a great and varied wealth of algal growth is seen. Although the large kelps are absent east of Cape Agulhas, two species of Ecklonia are found in warmer waters: EL. richardiana and EL. biruncinata. These plants are most typical of the central part of the warm water region and H#. biruncinata is seemingly far more abundant than H. richardiana but further study of the warm water Ecklonias is needed. E. biruncinata is dominant in places at Port Alfred and East London and shows a very luxuriant growth at Dwesa where over most of the coast it dominates the uppermost sub-littoral. As compared with the cold water kelps, the warm water Ecklonias are small plants usually not exceeding 4 feet in length inclusive of stipe and frond. Since these plants are relatively short the warm-water Ecklonia girdle is relatively narrow as seen at the surface of the sea and thus is never such a spectacular sight as may be presented by the kelp association of cold waters. While there is a smaller bulk of algal tissue between tide levels in warmer as compared with colder waters, the flora is richer in species. 66 The Journal of South African Botany. The increase in the number of species becomes more marked and tropical elements more pronounced in a north eastward direction from Cape Agulhas, these changes being correlated with a gradient of increasing sea temperatures. Of the warm water species, Hypnea spicifera is a widespread .dominant plant at the lowest inter-tidal levels. The species is dealt with in a separate paper (Isaac and Hewitt, 1953). Gelidium pristoides and G. reptans are also worthy of note. The former is essentially characteristic of moderately warm waters but is distributed from the central west coast (Kommetje) of the Cape Peninsula to about 60 miles north of Port St.-Johns. G. reptans extends further into warmer waters (Stephenson, 1947). These species are found at mid-tidal levels. At the lowest tidal levels we find Gelidium cartilagineum and the smaller, slenderer but somewhat similar G. rigidum. From the floristic point of view the warmer seas are characterised by a greater wealth of littoral Chlorophyceae and Phaeophyceae than the colder seas. The characteristic Chlorophyceae belong especially to the Siphonales and Siphonocladiales. Among the former are a number of species of Caulerpa, C. filiformis and varieties of C. racemosa being frequently very abundant. Mention may also be made of Halimeda cuneata, Pseudocodium de-vriesei and several species of Codiwm. Among the Siphonocladiales are those strange algae, Valonia acgagropila and V. macrophysa. Prominent among the Phaeophyceae are Sargassum spp. and. Dictyotales. Sargassum longifolium is exceptional in that it flourishes in places between Cape Point and Cape Agulhas in which localities it grows intermingled with Ecklonia maxima (Isaac, 1947). Of the Dictyotales, Dictyopteris, Zonaria tournefortii and Padina spp. including P. com- mersonit may be listed; all plants found exclusively in warmer waters, The following are some of the Rhodophyceae known only well within the warm water region: Hypnea viridis?, Dictymema stephensonir, Corallopsis aculeata, and Mesotrema elegans, It should be realised that the species named in the foregoing account by no means exhaust even the common algae of the South African coast as no more has been attempted than a general sketch of the most out- standing features of the algal life of South African coasts. FutTuRE INVESTIGATIONS. The early history of the study of South African seaweeds will be published elsewhere and it will be sufficient for our present purpose to 5 Range given by Papenfuss (Papenfuss, 1947) is Natal coast. It was found by the writer growing on the Pondoland coast (Port St. Johns, Mpandi) and on the Transkei coast at Dwesa. South African Seaweed Vegetation and Future 67 Investigations in this Field. draw attention to the fact that until the early thirties only a little work had been done in this field. From the early thirties onwards there was an increase in the bulk and scope of the work published. This work has included ecological studies, life history investigations and taxonomic work. Much of the taxonomic work has been concerned with corrections of previous errors in nomenclature without any change in the delimitation of the species concerned but a number of new species have also been described. Kylin, Levring, Mantza and especially Papenfuss have been active in this work. As a result of the ecological work of Stephenson and his co-workers, who dealt with algae as well as with animal species, and of Isaac, a general picture is available of the geographical distribution of at least most of the chief algal species and of the main features of the seaweed ecology of South African coasts. Further information is needed to complete and clarify this general picture. Thus more studies of particular localities are needed as well as of the algal ecology of the coast between Cape Agulhas and Port Elizabeth. Also, even in terms of a general picture, certain published accounts need amplification and correction, as for example the account given of the algae of Port Elizabeth by Stephenson, Stephenson and Bright (1938). Detailed ecological investigations of particular localities, however, would possibly be more profitably postponed until more taxonomic work has been done. The need of critical taxonomic investiga- tion of certain genera and groups of genera is apparent to investigators of South African seaweed ecology. This applies especially to the algae of warm waters. The conduct of seaweed investigations would be greatly facilitated by the publication of a flora. The time is possibly not ripe for the publication of such a work but the publication of a provisional flora would prove a great boon. Meanwhile there is considerable scope for detailed work on the ecology and physiological ecology of common species and groups of species as well as for preliminary investigations of the floras and ecology of rock pools and of the sub-littoral. The need of further work relating to the general algal ecology of South African shores has been indicated above. In relation to the autecology of common species, it may be pointed out that little or nothing is known on many matters of importance relating to the exploitation of South African seaweeds for industrial and com- mercial purposes. Information is much needed on the following matters:— (1) Estimation of the absolute growth density of individual species in sample areas; density to be expressed in terms of fresh and dry weights per unit area. 68 The Journal of South African Botany. (2) The extent to which perennial species can be cut back and regenerate and studies of the rate of regeneration. (3) The rate of re-colonisation by algal species following the clearing of sample rock surfaces. One study bearing on this subject has already been published (Bokenham and Stephenson, 1938). (4) Seasonal reproductive cycles. Work on item (4) is especially urgent in view of possible commercial exploitation. The investigation of seasonal reproductive cycles involves collection and sorting of adequate samples of a species at regular intervals throughout the year for a few years. It also involves the examination of samples from a number of localities, especially in the case of those species which show a relatively wide tolerance of changes in habitat conditions. Such work will be chiefly directed, in so far as harvesting is concerned, to determining whether there are months of the year during which the plants should not be collected. Provisional advice for a given species might be given after one or two years’ investigation but more certain information will involve work extending over longer periods. With such species as those of the Laminariales and Fucales the problem is relatively simple since what is involved is periodic determination of the proportion of fertile and non-fertile diploid individuals, since the haploid individuals are microscopic and not collected (Laminariales) or absent (Fucales). With the majority of Rhodophyceae, however, sorting of samples into non- fertile, male, female and tetrasporic plants is involved. As these phases of a single species are usually all morphologically similar a much more complex problem is involved. (5) Seasonal changes of vegetative parts of perennial species. This includes for a given species assessment of periods of relative dormancy and of active growth or the establishment of the fact that growth is more or less uniform throughout the year. It also includes a determina- tion of the extent to which an increase in tissue in a particular species is due to the formation of new plants by sexual or asexual reproduction and to what extent it is due to vegetative increase of existing plants. For example, to what extent is the dense growth of the Hypnea spicifera and Bifucaria brassicaeformis associations due to the vegetative growth of new fronds from the creeping rhizomes? The answer to this question will clearly have a bearing on action to be taken in relation to seasonal reproductive cycles. The information obtained in relation to the exploitation of species of economic importance is necessary if exploitation is to be based on a sound conservation basis. In this connection, however, it is fair to point out that in the case of a few species (e.g., Ecklonia maxima and Gracilaria South African Seaweed Vegetation and Future 69 Investigations in this Field. confervoides) exploitation can proceed far by making use of cast up weed. This, however, is not the case with such species as Gelidiwm pristoides, Gigartina radula, Aeodes orbitosa, Hypnea spicifera and Porphyra capensis, to name some of the species of actual or potential economic importance. The information obtained from such studies will be of ecological importance as well as of economic importance. Autecological information, however, is needed for a wider range of species both perennial and annual. For the warmer waters especially, it is necessary to follow out the times of disappearance of certain species as well as of changes in relative abundance and any changes that there might be in inter-tidal level. (6) Seasonal changes in the amounts of the chief organic compounds present. This information will be of value to the biochemist and physiologist as well as to the industrialist. From these remarks it will be clear that while a general picture of seaweed distribution on South African coasts is now available much work remains to be done. While further detailed work in certain directions is held up until further taxonomic investigations are completed, the autecology of the common species and preliminary ecological work on hitherto neglected aspects present fruitful fields for investigation. SUMMARY. The study of South African seaweed vegetation is complex due to the extensive coastline, the range of sea temperatures, and the imperfectly studied algal flora. Along most of the coast the coastal waters are affected by one of two ocean currents; the cold Benguela Current moving northwards along the west coast and the warm Agulhas Current moving southwards and west- wards along the east and south coasts as far as Cape Agulhas, the most southerly point of Africa. Between Agulhas and Cape Point sea tempera- tures are affected by branches of the Agulhas Current, coastal topo- graphy and prevailing winds. False Bay has abnormally high tempera- tures for its geographical position and its algal flora and vegetation must be considered separately. For convenience in a general account, the coasts west and east of Agulhas are called west and east coast respectively. The seaweed flora and vegetation of east and west coasts show marked contrasts although there is overlap, especially at their southern limits. Changes are also evident along each coastline in relation to sea temperature gradients. Certain features are common to both coasts. (a) Limited vertical extent of inter-tidal vegetation due to narrow tidal range (low water springs to high water springs, 5—6 ft.) and paucity of algal growth at 70 The Journal of South African Botany. higher inter-tidal levels. Algae are mostly absent above high water neaps on west coast and above mid-tide level on east coast. (6) Over most of the coast Rhodophyceae dominate every inter-tidal level. Outstanding exceptions are the dominance at a lower inter-tidal level of Bifurcaria brassicaeformis from Cape Town to Cape Agulhas and the local dominance of Caulerpaceae in warm waters. The west coast is characterised by extensive sub-littoral growths of large kelps—EHcklonia maxima, Laminaria pallida and to a much smaller extent, Macrocystis, and by a girdle, frequently dense, of Porphyra capensis extending downwards from high water neaps. East coast species of EHcklonia are small and much less abundant. A Porphyra girdle is absent although small, pale plants of P. capensis occur as far north on the east coast as Port St. Johns. Hast coast seaweeds are mostly smaller and dense mixed communities of small algae are common. The flora is richer and many species are confined to warmer seas. Future investigations are considered, bearmg in mind information needed for seaweed exploitation on a conservation basis. While detailed investigation is held up in certain directions until further taxonomic work is completed, preliminary ecological work on previously neglected aspects and the autecology of common species present fruitful fields of investigation. ACKNOWLEDGEMENT. Some of the observations included in this paper were made while the author was doing field work financed by the South African Council for Scientific and Industrial Research. REFERENCES. Bauer, H. A. (1933): A World Map of Tides, Geogr. Rev. vol. 23, p. 259. BoxenuamM N. A. H. and StepHEenson, T. A. (1938): The Colonization of Denuded Rock Surfaces in the Intertidal Region of the Cape Peninsula, Ann. Natal Mus., vol. 9, p. 113. Bricut, K, M. F. (1938): The South African Intertidal Zone and its Relation to Ocean Currents, III. An area on the Northern Part of the West Coast, Trans. Roy. Soc. S. Africa, vol. 26, p. 67. FELDMANN, J. (1951): Ecology of Marine Algae, Chap. 16, of Manual of Phycology (Edited by G. M. Smith) Waltham, Mass., U.S.A. Tsaac, Wm. Epwyn (1937a): South African Coastal Waters in relation to Ocean Currents, Geogr. Rev. vol. 27, p. 651. (19376): Studies of South African Seaweed Vegetation. I.—West Coast from Lamberts Bay to the Cape of Good Hope, Trans. Roy. Soc. S. Africa, vol. 25, p. 115. (1938): The Geographical Distribution of Seaweed Vegetation in relation to temperature and other factors, with special reference to South Africa, Comptes rendus du Congrés international de Géographie, Amsterdam, 1938, tome ii, Section 7, p. 12, Leiden. (1942): Seaweeds of Possible Economic Importance in the Union of South Africa, Jour. South African Botany, vol. 8, p. 225. (1949): Studies of South African seaweed vegetation. II.—South Coast: Rooi Els to Gansbaai, with special reference to Gansbaai’, Trans. Roy. Soc. S. Africa, vol. 32, p. 125. South African Seaweed Vegetation and Future ol Investigations in this Field. (1951): Observations on the Ecology of Bifurcaria brassicaeformis (Kiitz.) Barton, Jour. Ecology, vol. 39, p. 94. Isaac, Wm. Epwyn & Hewitt, F., The Geographical Distribution and Ecology of Hypnea spicifera, Paper presented to First Seaweed Symposium, Edin- burgh, 1952. (This Journal, to appear, 1953.) Murray, G. (1893): A Comparison of the Marine Floras of the warm Atlantic, Indian Ocean, and the Cape of Good Hope, Phycological Memoirs, part 2, p. 65. Papenruss, G. F. (1947): New Marine Algae from South Africa: I, Univ. Calif. Publ. Botany, vol. 23, No. 1. STEPHENSON, T. A. (1947): The Constitution of the Intertidal Fauna and Flora of South Africa. Part III, Ann. Natal Mus., vol. XI, p. 207. STEPHENSON, A. and Bricut, K. M. F. (1938): The South African Intertidal Zone and its relation to Ocean Currents, IV, The Port Elizabeth District, Ann. Natal Museum, vol. IX, p. 1. Taxonomic NOTES. Chlorophyceae. Ulva minuta Papenfuss, ined. Papenfuss’ authenticated material (L. 131) in Botany Department, University of Cape Town. Phaeophyceae. Ecklonia biruncinata (Bory) Papenfuss. This plant was known earlier as H. radiata (Turn.) J. Ag. Papenfuss (1940) later published the view that the plant should be named £. exasperata (Turn.) J. Ag.; being regarded as a distinct species and not as a variety of H. radiata. Still later (Papenfuss, 1944), this species was renamed E. biruncinata (Bory) Papenfuss since the plant was first described as Laminaria biruncinata by Bory in 1826. Ecklonia richardiana J. Ag. Stephenson (1947) regards the South African plants named #. richardiana by Papenfuss (1940a) as probably variants of H. biruncinata but he gives no reasons for his view. Rhodophyceae. Aeodes orbitosa (Suhr) Schm. In earlier papers the name Iridaca capensis was used to cover both Aeodes orbitosa and Iridophycus capensis. Gelidium rigidum (Vahl) Grey. There is some doubt as to whether the plant known by this name in South Africa is not G. amansii (Lamour) Lamour. Gigartina stiriata (Turn.) Aresch. This includes the tetrasporie plant which is morphologically distinct and was formerly known as G. burmanii (Mert.) J. Ag. CHANGES IN NOMENCLATURE. Chlorophyceae. Caulerpa filiformis (Suhr.) Hering; C. ligulata Harv. ex J. Ag. (Papenfuss, 1940a). Phaeophyceae. Ecklonia maxima (Osbeck) Papenf.; 2. bwecinalis (L.) Hornem. (Papenfuss, 1940b). Stypopodium zonale (Lamour.) Papenf.; S. lobatwm (Ag) Kutz. (Papenfuss, 1940a). Rhodophyceae. Martensia elegans Hering: Mesotrema elegans (Her.) Papenf. (Papenfuss, 1942). REFERENCES. Parenruss, G. F. (1940a): Notes on South African Marine Algae, I., Bot. Notiser, 1940. ———— (1940b): A revision of the South African marine algae in Herbarium Thunberg, Symbolae Botanicae Upsaliensis, vol. 4, No. 3. a (1942): Notes on algal Nomenclature: I. Pollexfenia, Jeanerettia and Mesotrema, Proc. Nat. Acad. Sci. Washington, vol. 28, p. 446. (1944): Notes on Algal Nomenclature, III. Miscellaneous species of Chlorophyceae, Phasophyceae and Rhodophyceae, Farlowia, vol. 1, p. 337. SrepHEeyson, T. A. (1947): The Constitution of the Intertidal Fauna and Flora of South Africa, Part III, Ann. Natal Mus., vol. XI, p. 207. y, i, 5 , eis i 7 ‘oe ag TRADE Bey : et! g i ISSUED IO) IL, JOURNAL OF SOUTH AFRICAN BOTANY VOL. XIX. THE MORPHOLOGY, GEOGRAPHICAL DISTRIBU- TION AND ECOLOGY OF HYPNEA SPICIFERA (SUHR.) HARV. By Wm. Epwyn Isaac and FLrorence Hewirr. (Cape Town and Rhodes Universities.) (With Plates XVI and XVII.) (Paper presented to the First International Seaweed Symposium, Edinburgh, July, 1952.) CONTENTS. PAGE Introduction... : es Ss si Ps bos Be 73 Morphology and Colour: of es ee % ie sy 74 Geographical Distribution Les Be * Hs be 5 Ud Intertidal Position - ee ed ne ee oe nes 81 Keological Status ae ay be fee se ss 81 Occurrence in relation to iSiesaaaiiys a ate i oie a 82 - Summary : mG Ls she a bs Be Be 83 Acknow. ledgers be a a 23 ie a Ce 83 Literature eae i = sh a ate as et 84 This report is presented because Hypnea spicifera is one of the most important ecological species at intertidal levels over a large part of the South African coast, and because of its possible economic importance. The published information relating to it is scattered and insufficient to give a clear picture. It is one of several species of Hypnea occurring in the intertidal zone, but it is much more common than any of the others. The account given here of the morphology and ecology of the species is chiefly based on observations carried out on plants growing from Richmond (Boknes) to the Great Fish River Mouth and is thus to be regarded as a preliminary account of the species in South Africa as a whole. 73 24 1958 74 The Journal of South African Botany. EAST LONDON GR FISH RIVER MOUTH PORT ALFRED | PORT ELIZABETH CP MOP ZOR SOE 4050) Scale in Miles Fic. 1. The coast from Port Elizabeth to East London. Morphology and Colour. There are distinct male, female and tetrasporic plants which are of similar morphology and size. Detailed studies at Port Alfred (The Kowie) over a period of a year showed that in that locality tetrasporic and sexual plants occurred at all seasons, though the sexual plants were always found to constitute a very small proportion of the population. This conclusion was supported by additional observations over a stretch of coast from Diaz Cross (approximately six miles east of Richmond) to Fish River Mouth (Fig. 1). Similar studies are needed over other parts of its dis- tribution range. The plant (Fig. 2) consists of a tuft of erect, cylindrical fronds arising from a creeping prostrate system in character somewhat resembling a bird’s nest. This is further dealt with in considering the formation of communities dominated by Hypnea spicifera. The erect fronds are densely crowded together and often much branched, giving the plant a bushy appearance. The main axes and branches of the upright fronds have straight tips and more or less pointed apices. ° Mature fronds are often 30 to 40 cms. (about 12 to 16 inches) long or more, and about 1-5 mm. in diameter near the base. A length of nearly 60 cm. (nearly 2 ft.) has been recorded. Branching is irregular and in all directions. Of the mature 76 The Journal of South African Botany. Itc. 3. Enlarged view of fertile region of tetrasporic plant of Hypnea spicifera. Stippled areas are fertile parts of stichidia. fronds the fertile are the more conspicuous, since they are usually several ems. longer than the vegetative fronds and often greater in diameter. The upper portions of the fertile fronds are usually densely covered with numbers of short spinulose branchlets which form a stiff bristle-like covering on the main axes and their branches, extending to within a few millimetres of the tips. These specialised branchlets (stichidia) bear the reproductive organs (Fig. 3). Tetraspores occur in zonate groups within the tissues of the stichidia. The antheridia occur in sori in depres- sions on the surface of the thallus. Cystocarpic fronds are readily identifiable to the naked eye by the presence of globular fruits about 1 mm. in diameter and bright pink when ripe. The Morphology, Geographical Distribution and Ecology WY of Hypnea spicifera (Suhr.) Harv. The colour is variable and in the main appears to be related to the level at which the plant grows. Plants in the upper part of the Hypnea girdle have short, succulent upright fronds glossy green in colour, with trans- lucent tips. Plants in the lower portion of the girdle have long dark fronds varying in colour from dark olive green to almost black. In the lowest part where wave action is most severe the long blackish fronds are often wiry and sparsely branched. Mature parts of the prostrate system are dark in colour. The young fronds of both the erect and prostrate systems are a pale, semi-translucent pink colour. Fertile fronds, especially cystocarpic fronds, tend to be a more brownish green than the vegetative fronds of the same plant. Geographical Distribution. More data are needed before a full and exact picture can be given of the distribution of Hypnea spicifera on the coasts of South Africa. The general outlines of the picture, however, are clear on the basis of the data available. Hypnea spicifera is not endemic but the only record of its occurrence outside the Union known to the writers is at Karachi on the north-west coast of India (Bérgesen, 1934). On the coast of the Union it has a very wide distribution (Fig. 4), being one of the most widely distributed sea- weeds on South African coasts. It has been recorded from the northern coast of Natal (Umpangazi) to Buffels River within less than 40 miles south of Port Nolloth (Stephenson, 1947). A bare statement of the geographical range on the Union coasts of such a widely distributed species as Hypnea spicifera gives an erroneous impression since the size of the plant and its prominence in the seaweed vegetation varies much within its distribution range. As an important constituent of the seaweed vegetation it has two centres of distribution (Fig. 4): (a) From about Boknes (about 50 miles east of Port Elizabeth) to the northern coast of Natal. (b) False Bay and the southern end of the west coast of the Cape Peninsula. The first of these is much more extensive geographically than the second. In fact it is or rather, it includes (as will be shown later) the true centre of distribution. The smaller centre is best regarded as derivative or secondary since locally favourable conditions! make it possible for a 1 For comments and references on ocean currents and sea temperature conditions in South African coastal waters see: Isaac, 1937a and Isaac, 1953a. ‘puafroids OSI ool OS (0) neudhz jo UoKyNgIyAStp ey Jo yUosord ye UMOUY st yey Surstreurums dey “p ‘org ERIE) Ble 2122'S SVWHINIV Adv) IWVE ILS UE shsh/ 2) 3} ILel@al SNHOL 39S 1H0d Ayisuaq [PEK GaN Octe9) CIA MYDS) Nf KMARAAAAASSSSS quo ys0dui Ajjo21bo joo Ta Sees] HLOTION Ldod abuoy uorgngr494sig P®P40d2y4 YoRY AYVH (4HNS) WUSSIDIdS WANdAH JO NOILNEIYLSIC The Morphology, Geographical Distribution and Ecology 79 of Hypnea spicifera (Suhr.) Harv. GANSBAAI DANGER Po! sTRUYS BAY CAPE AGULHAS ———————————— ce) 10 20 30 40 Sede m Miles Fic. 5. The Agulhas Peninsula and the coast to the west as far as the eastern end of False Bay. widely distributed species to become ecologically important in localities distant from the main centre of distribution. Except at Rheboksdam Bay (Isaac, 1937b), Hypnea spicifera is not a species of ecological importance on the west coast where it is rare or occasional. From False Bay to and including Port Elizabeth it shows an erratic distribution but additional information is needed relating to its occurrence along this considerable stretch of coast (about 500 miles). In some localities such as Struys Bay and Agulhas (Fig. 5) it is absent or rare; in other localities, while it is not uncommon it is not one of the common, let alone dominant or sub-dominant, species. Thus at Gansbaai it is a constituent of the mixed algal vegetation of which it is not one of the commonest species; further, the mixed algal vegetation is rather poorly represented and is one of four possible types of algal vegetation at low water level of spring tides, the most common community being the Bifurcaria association (Isaac, 1949; 1951). In other localities such as Frikkie’s Baai on the eastern side of Danger Point, Pearly Beach between Quoin and Danger Points, and Hermanus 80 The Journal of South African Botany. (Fig. 5) it is a common or locally prominent species. In the Still Bay region Hypnea spicifera is listed as one of the algae occurring on an isolated rocky mass and in an annotated list it is marked with an asterisk to indicate that it is a fairly or very common species (Stephenson, Stephenson and du Toit, 1937). As far as our evidence goes, nowhere between False Bay and Port Elizabeth does Hypnea spicifera attain the status of a dominant species but this view may have to be modified when more information is available. In the Port Elizabeth district (Port Elizabeth to Chelsea Point), the small amount of this species is a note- worthy feature. It is the converse as it were to the dominance of Corallinaceae and Laurencia spp. at the lowest intertidal levels. This point was not made by Stephenson, Stephenson and Bright (1938). On closer examination of the evidence it becomes clear that Hypnea spicifera is not uniform in its ecological status throughout its two centres of distribution. In the secondary centre, the plant flourishes best along the north-western and northern coasts and the northern half of the eastern coast of False Bay. Over this extent of coastline it is dominant in places at the lowest intertidal levels although the Hypnea girdle may be a narrow one. Along most of the east coast of the Cape Peninsula to the southern end of the west coast (Rheboksdam Bay) the species is more erratic in its ecological importance since it may be rare, occasional and _ local, common or very common to locally dominant. In general, it clearly becomes ecologically more important in a north- ward direction along the east coast of the Cape Peninsula, attaining optimum development in its secondary centre of distribution along the north-western, northern and eastern shores of False Bay. In regard to its major centre of distribution, Hypnea spicifera flourishes best from about Richmond to somewhere between East London and Port St. Johns. At East London this plant is abundant and except in more sheltered situations it is the outstanding dominant species at lower intertidal levels. At Port St. Johns Hypnea spicifera is still frequently a dominant species at lowest intertidal levels but there is evidence that it has already extended beyond its region of optimum development. This is seen in its smaller size which does not usually exceed 43 inches; in a greater degree of inter- mingling with other species; and in its replacement by other species in moderately sheltered as well as in sheltered situations. The account given by Eyre and Stephenson (1938) makes it clear that in the Durban area the plant has shown a further decline from its optimum condition. It is still smaller in size (less than three inches), it is less important in relation to other species growing at the same intertidal level and is less frequently a dominant species. The Morphology, Geographical Distribution and Ecology 81 of Hypnea spicifera (Suhr.) Harv. Intertidal Position. Hypnea spicifera occupies the lower part of the intertidal zone where it is emersed at low water of spring tides, although the greater part of the Hypnea girdle may be regularly washed by the waves. (Plates XVI and XVII.) The Hypnea girdle commonly dominates rocky ridges, the vertical sides of rocky platforms, the walls of gulleys, and the roofs of overhanging ledges. The upper part of the girdle which usually lies above wave action at low water of spring tides may be subjected to constant spray. The lower part of the girdle is almost always subjected to very severe pounding by the waves. Hypnea spicifera grows also at corre- sponding levels on isolated rocks beyond the intertidal belt. It extends into the sublittoral to only a limited extent. It is not typically a rock pool species, but where it occurs it grows just below the surface of the water. It is always attached to a firm substratum, usually to rock, sometimes to mussels. It can withstand being partially buried in sand. Ecological Status. In the region of optimum development Hypnea spicifera not only dominates, but forms almost pure communities. This can be accounted for by certain features of the prostrate system. This system consists of an intricate network of creeping axes, from which numerous branches spread out in all directions covering the substratum. These creeping axes are anchored by minute, more or less circular discs (haptera) which arise at irregular intervals from the branches. (Fig. 6.) Numbers of Fic. 6. Small part of basal region of Hypnea spicifera, A. Bases of erect fronds. B. Sand grains adhering to discs. 82 The Journal of South African Botany. erect fronds develop from the creeping axes. By this form of vegetative growth the Hypnea community not only spreads extensively but. is built up in an almost pure state. It would thus appear that the plant is only partly dependent on reproduction by tetraspores and sexual cells. As explained above, under optimum conditions of sea temperature and exposure, Hypnea spicifera forms dense communities which are almost pure, other species being crowded out. In this respect these communities recall those of Bifurcaria brassicaeformis (Isaac, 1951). Even under optimum conditions there may be some admixture of species at the sea- ward limits of the Hypnea girdle. Thus at East London, one or more of the species Gelidium cartilagineum, G. rigidum* and Ecklonia biruncinata grow alongside Hypnea spicifera at its lowest levels. Again at Port Alfred, clans of Chondria sp. occur in and about the Hypnea girdle as well as to some extent growing among plants of Hypnea spicifera. Where conditions are less favourable, Hypnea spicifera forms a con- stituent of a mixed algal vegetation. It occurs thus at Gansbaai and Hermanus (Isaac, 1949) but in neither locality is it one of the commonest species. At Rooi Els it is sometimes co-dominant with Gelidiwm cartilagineum. In moderately sheltered places at Port St. Johns Hypnea spicifera may occur as one of a number of species. Occurrence in Relation to Exposure. Over most of its geographical range, Hypnea spicifera flourishes best in exposed situations where it withstands considerable battering from the sea. This is most obvious in its major centre of distribution. At East London, Dwesa and Port St. Johns it is chiefly replaced by Caulerpa racemosa var. zeyheri in less exposed situations. At Fish River Mouth, Hypnea spicifera is replaced in more sheltered places by Caulerpa fili- formis. At its western limits of distribution as an ecologically significant species there are indications that Hypnea spicifera favours more sheltered situations or is at least more tolerant of them. In this connection it might be noted that in the very exposed places at Hermanus (Kwaaiwater) it is Plocamium cornutum and not Hypnea spicifera that is the most character- istic species, the latter occurring as a member of the mixed algal vegetation in less exposed situations (Isaac, 1949). Further, most of the secondary centre of distribution occurs in False Bay. It is true that the chief reason why Hypnea spicifera flourishes in the more northern parts of False Bay is the higher sea temperatures but it should be noted that False Bay is also a region of relatively calm waters. * See note, Isaac, 1953a, appendix, The Morphology, Geographical Distribution and Ecology 83 of Hypnea spicifera (Suhr.) Harv. SUMMARY. The male, female and tetrasporic plants of Hypnea spicifera (Suhr) Harv. are morphologically similar. The plant varies from less than three inches to about 16 inches and more in height and from a bright green to dark olive green, almost black colour. Size and colour variations are related to geographical and ecological factors. Fertile plants are somewhat longer than purely vegetative plants and bear numerous small fertile branchlets (stichidia). The plant consists of an extensive, richly branched prostrate part from which numerous upright richly branched fronds arise. The prostrate part bears numerous small discs by means of which the plant adheres to a hard substrate, usually rock. Hypnea spicifera, which is of possible economic importance, is dis- tributed over almost the entire extent of the Union coastline but is of ecological importance only in warmer waters. Except locally at the very southern end, it is a rare or occasional species on the west coast which is subject to the influence of the cold Benguela Current. It is ecologically important from a little east of Port Elizabeth (about Boknes) northwards along the east coast of the Union. Along the whole of this coastline it is subject to the influence of the warm Agulhas Current, sea temperatures increasing in a northward direction. In the southern part of this stretch of coast it shows a luxuriant growth and dominates a low intertidal level to the practical exclusion of other species except in sheltered situations. To the north, while remaining an ecologically important species it becomes smaller in size and less important in the seaweed vegetation. Abnormally high sea temperatures in False Bay immediately east of the Cape Peninsula are correlated with a secondary centre of distribution along the northern and eastern shores of the bay, where Hypnea spicifera again becomes a common or dominant species, although the Hypnea girdle is often narrow. Hypnea spicifera occurs to some extent in pools and in the sublittoral but it is chiefly found at the lowest intertidal levels. Over the greater part of its distribution range it flourishes in exposed situations. Under optimum conditions it forms dense almost pure communities, this being facilitated by the creeping prostrate part of the thallus. Along other parts of the coast it may be locally dominant or it may be a constituent of greater or lesser importance in a mixed algal vegetation. ACKNOWLEDGEMENTS. Some of the data included in this paper are taken from a thesis to be presented for the degree of M.Sc. of Rhodes University by one of the 84 The Journal of South African Botany. authors (Miss F. Hewitt). We wish to thank Rhodes University for permission to use this information. We also wish to thank the South African Council for Scientific and Industrial Research for financial assistance which made it possible to ascertain some of the information contributed by the other author. To Mr. R. Simons we are indebted for Figs. 1, 4 and 5. LITERATURE. Borcesen, F. (1934): “Some Marine Algae from the northern part of the Arabian Sea with remarks on their Geographical Distribution”, Det. Kgl. Danske Videnskabernes Selskab Biologiske Meddelelser, vol. X1, 6. Eyre, J. and StepHenson, T. A. (1938): “The South African Intertidal Zone and its relation to Ocean Currents. V. A Sub-tropical Indian Ocean Shore,” Ann. Natal Museum, vol. IX, p. 21. Isaac, Wu. Epwywn (1937a): “South African Coastal Waters in relation to Ocean Currents,” Geogr. Rev., vol. 27, p. 651. ———— (1937b): “Studies of South African Seaweed Vegetation. I. West Coast from Lambert’s Bay to the Cape of Good Hope,” Trans. Roy. Soc.S. Africa, vol. 25, p. 115. (1949): “Studies of South African Seaweed Vegetation. II. South Coast: Rooi Els to Gansbaai, with special reference to Gansbaai.” Ibid, vol. 32, p. 125. ———— (1951): “Observations on the Ecology of Bifurcaria brassicaeformis (Kutz.) Barton,” Jour. Ecology, vol. 39, p. 94. ——— (1953): “South African Seaweed Vegetation and future investigations in this field.” Journ. S. African Botany, vol. XIX, p. 59. STEPHENSON, T. A. (1947): “The Constitution of the Intertidal Fauna and Flora of South Africa, Part III.”, Ann. Natal Mus., vol. XI, p. 207. STEPHENSON, T. A. and A., and Bricut, K. M. F. (1938): ““The South African Inter- tidal Zone and its relation to Ocean Currents. IV. The Port. Elizabeth District,” Ibid, vol. 9, p. 1. STEPHENSON, T. A. and A., and pu Torr, C. A. (1937): “The South African Intertidal Zone and its relation to Ocean Currents. I. A Temperate Indian Ocean Shore,” Trans. Roy. Soc. S. Africa, vol. XXIV, p. 341. ‘UOISeI PaT[W MA <) Reg 7B a LVI, I 4IOg :(9S9A4 OIMO IMO) TOA UBS 78 + I [pais waudhy * s TAX d oN LA O189 Ve Ros uodo oy 04 ojsun un ye Ayjnd B UL oppald poudA EP “TIAN “havi UOLdOd POLLY WO s(GSOAA OIMOYT) SEAWEED RESOURCES OF SOUTH AFRICA. By Wm. Epwyn Isaac and C. J. Moreno. (Contribution to a discussion on World Seaweed Resources held at First International Seaweed Symposium, Edinburgh, July, 1952.) CONTENTS. PAGE Algin yielding weeds .. be a Se a i ss 85 Agarophytes .. i a, op: bit oe oid ‘ee 87 Species yielding Phycocolloids other than Alginic acid and Agar 88 Miscellaneous .. = he si oe GE - ee 90 Concluding remarks... = ae eo 5 a 90 Summary Si we ne ae ai oe a a 91 Literature He eS bis es eh bi ae Me 91 Precise figures of South African seaweed resources cannot be given since no quantitative surveys have been carried out. Indicative data, however, are available from the records of Vitamin Oils, Ltd., Cape Town. These estimates are based on the amounts of certain species collected at particular localities. Algin yielding weeds. Alginie acid can be obtained from Hcklonia maxima, South African Macrocystis, Laminaria pallida, Bifurcaria brassicaeformis and Sargassum longifolium (Lighthelm, von Holdt and Schumann, unpublished paper). The last named species has the lowest content and is not present in sufficient amount to be of commercial value. The last remark applies also to Macrocystis. As an abundant intertidal species, Bifurcaria brassicaeformis has a restricted geographical range, being abundant only between Cape Town and Cape Agulhas (Isaac, 1951). It is not cast up to any extent and thus would have to be cut from the rocks at low water of spring tides. Laminaria pallida and especially Ecklonia maxima are abundant and are cast up in large quantities along certain parts of the west coast. Ecklonia is cast up throughout the year and being buoyant because of hollow stipe and float it is cast high up on the shore where most of it remains. Laminaria is cast up after heavy storms, mostly in July and 85 FOR A. Gracilaria confervoides B. “Kelps, Gigertina radula, Aeodes orbifosa C. Gelidium pristoides MELKBOSCH CAMPSBAY EXPLOITED 86 The Journal of South African Botany. AREAS SEAWEEDS Scale in M les fe) 50 100 Fic. 1. Map showing areas exploited for seaweeds. August. Lacking the large internal cavities of Ecklonia, it is less buoyant and is cast up lower on the beach, and is removed again to a considerable extent by the outgoing tide. Related to the relatively small seasonal variation in temperature of the colder South African waters in which these kelps grow (Isaac, 1937a, 1937b, 1949) there is no difference in the cast up Ecklonia at different times of year corresponding to the May and winter casts of kelp on British coasts (Newton, 1951). Referring to cast up material of Ecklonia maxima and Laminaria pallida, Shuttleworth (1951, ce, p. 115) writes: “A fairly conservative estimate would seem to put the total quantity of these two weeds annually deposited on our coastline after storms, etc.,at many hundreds of thousands if not millions of tons.” He does not indicate how he arrives at his estimate. Again, there are no critical data on which to base Shuttleworth’s assertion that: “‘Our re- sources are almost certainly greater than those of Scotland”! (Ibid, p. 116.) It is clear, however, that there are large quantities of sublittoral kelp westwards from Cape Agulhas. Thus from five localities only (Melkbosch, Kommetje, Plaatboom Point, Onrust and Gansbaai) 6,000 tons dry weight (about 15 per cent. water) of cast up kelp, mostly Hcklonia, can be collected annually. These localities (Fig. 1) represent but a small part of the total coastline along which kelps grow but much of the west coast is very inaccessible for the commercial collection of cast up kelp. Also the distances and lack of transport facilities involved would make it un- economical to convey seaweed from remoter parts of the coast. 1 Although available data will not allow of a clear refutation, there can be no doubt that Shuttleworth exaggerates the quantities of kelp on South African coasts. KE! RIVER M7 Seaweed Resources of South Africa. 87 Agarophytes. Agar can be obtained from the following species: Gelidium cartilagineum, G. pristoides, Gracilaria confervoides and Suhria vittata (Isaac, Finlayson and Simon, 1943). Fox and Stephens (1943) reported agar from these species as well as from Caulacanthus ustulatus and Hypnea spicifera but their paper contains no details regarding the products extracted. Isaac et al. (1943) showed that the Hypnea spicifera extractive is not a true agar. It was also reported that the agar extracted from Gelidiwm pristoides was of inferior quaility, but subsequent work at Vitamin Oils, Ltd. has shown that a high quality agar is obtainable from this plant. Agar might be extracted from other South African species, including species of Gelidium, but these plants are either too small or not present in sufficient amounts to make them valuable commercially. Gelidiwm corneum was listed for South African coasts by Delf and Michell (1921) and Fox and Stephens (1943) reported that there is some evidence that it may occur on the coasts of the Eastern Province. Papenfuss (1943) refers to a species of Gelidium from the east coast of South Africa which “has been referred to Gelidiwm corneum by some authors and to G. rigidum (now Gelidiella acerosa) by others, but the writer is still uncertain of its identity” (p. 91, footnote 6). Newton (1951, p. 118) wrongly reports Isaac as indicating that G. corneum occurs on South African coasts. Thus it is still uncertain whether the species occurs on these coasts and in view of investigations carried out over an extensive range of South African coasts it is clear that it is not a common species. Caulacanthus ustulatus, Gelidium cartilagineum and Suhria vittata do not occur in sufficient amounts to be commercial sources of agar. G. cartilagt- neum is a smaller and less luxuriant plant on South African coasts than on Californian coasts and diving operations have shown that it does not extend into the sublittoral in the way that the Californian plant does. On South African coasts it is also frequently found growing alongside the species which has been known here as G. rigidum? with which it can sometimes be confused by harvesters. @. rigidum yields only a small amount of low grade agar. Thus as the picture is at present known, the two commercially important agarophytes are G. pristoides and Gracilaria confervoides. The latter gives in some respects a relatively inferior agar but has the advantage of being cast up in very large amounts in the Saldanha—Langebaan area. About a thousand tons of dry weed is cast up annually in this region. It occurs also at Hout Bay and at Luderitz but in much smaller amounts. Thus the first area named was, and remains, the source of the commercial 2 The South African plant known as Gelidiwm rigidum (now Gelidiella acerosa; see Papenfuss, 1943) seems to be Gelidium amansii. (Papenfuss, 1951). 88 The Journal of South African Botany. seaweed. Originally South African agar was obtained exclusively from this species but since 1951 agar has been obtained commercially from this species and from Gelidium pristoides*. The latter species is not cast up and has to be picked from mid-tide levels. It is widely distributed on South African coasts in moderately warm waters. It has been collected commercially from Hamburg to Kei River mouth (about 70 miles of coastline) at the rate of about 80 to possibly 100 tons dry weight per annum. Unfortunately no data are at present available regarding its rate of growth and seasonal reproductive cycle although work in regard to the latter is in hand. The full story of South African Gracilaria confervoides has not been elucidated but Shuttleworth’s account is not correct. Shuttleworth (195lb) seems to assume that the account given for North Carolina material (Causey et al., 1946) is true for South African Gracilaria but this isnot so. Apart from cast up weed this plant occurs at Saldanha—Lange- baan floating in the water and on sand banks where it is weighted down by sand or entangled with mussels. A fuller account will be published later but the chief results obtained are indicated below: (a) Only small amounts of attached plants have been found?. (6) Freshly cast up material, sandbank material and floating material will grow vegetatively in frequently changed sea water in the laboratory. (c) Fertile as well as sterile material is used for agar manufacture. It may be noted that only sterile plants of Gracilaria confervoides are used for the manufacture of agar in eastern U.S.A. (Causey et al., 1946). Species yielding phycocolloids other than Alginic acid and Agar. The chief South African sources of phycocolloids other than alginic acid and agar at present known are Aeodes orbitosa, Gigartina radula and Hypnea spicifera. The properties of Hypnea spicifera extractive have been discussed by Shuttleworth (195la) and the distribution of this abundant South African seaweed has been outlined by Isaac and Hewitt (1953). More research, however, needs to be carried out on the extractive and the technology of extraction. Aeodes orbitosa is found in cold and moderately cold water. It occurs in the lower parts of the mid-tidal level. It is often a relatively small plant but it may attain lengths of 2—3 feet. In some localities (Melkbosch, Kommetje, Camps Bay, Gansbaai) it is a common or abundant species. 3 For reasons not connected with the quality of the agar, this species is not being harvested at present. No attached plant or basal attaching dise had been found when this paper was presented in July, 1952. Fic. 2 A. Aeodes orbitosa showing holdfast. Vy, SS \ Ds / =\ sot \ | Ne ae PY \ \ 7 ‘Tg i [ e 20 a L TINCH > Seole Fig. 2B. Iridophycus capensis showing holdfast and stipe. 90 The Journal of South African Botany. Its general appearance is similar to that of Iridophycus capensis, which seems to have a similar vertical and geographical distribution. Iridophycus tends to be darker in colour and does not grow to the maximum dimensions attained by Aeodes orbitosa. The two plants can be distinguished by their basal parts. Aeodes orbitosa has a simple, relatively large and strong hold- fast so that the plant is not easily detached from the rock (Fig. 2A). Iridophycus has a smaller holdfast which in general grips the rock less firmly and it also has a more or less distinct short stipe which is more or less distinctly canaliculated (Fig. 2 B). A stipe is completely absent in Aeodes. Except locally, Aeodes is much more abundant than Iridophycus. More information, however, is needed on the ecology of these two plants. Aeodes has been used successfully for copper fining of beer but it is not sufficiently abundant to be of industrial importance. Dried Gigartina radula is also excellent material for beer fining and has already been exported for this purpose. It occurs at lower intertidal levels in places where the coast is sheltered from the full force of the sea and is found on the west coast and in localities as far east as Danger Point. Although it may be absent on many parts of the coast where the seaweeds are exposed to rough seas, it is common in other places. An indication of its abundance is afforded by the claim that up to 100 tons dry weight (20—30 per cent. water) per annum can be obtained from the Peninsula coasts. Miscellaneous. The possible use of South African seaweeds as food and as fertilizer was discussed in a previous paper (Isaac, 1942). Concluding Remarks. An attempt has been made to give an indication of seaweed resources available for industry in South Africa. More exact surveys are clearly required to give a fuller picture. Where estimates have been given they have been conservative. Fig. 1 helps to put the matter into perspective since it makes clear that only a small part of the total extent of coastline has been exploited for seaweeds. From the data presented it is obvious that considerably more seaweed would be available by collecting over a greater extent of coastline. Information is needed, however, not only of the amounts available at any given time but also of the rates of growth since it is possible that growth rates will be higher for example than those of British seaweeds. Further, where attached seaweed is harvested, information relating to the annual reproductive cycle is essential if exploitation is to be on a con- servation basis. Seaweed Resources of South Africa. 91 SUMMARY. Precise information cannot be given of the seaweed resources of South Africa as no quantitative surveys have been made. Indicative information, however, is available from commercial records of amounts of certain species collected and from known distribution and ecological status of the seaweeds concerned. The coastline of South Africa is very extensive and only small stretches of it are at present being exploited. The long distances and lack of adequate transport facilities make the exploitation of more inaccessible parts of the coast uneconomical. A further consideration that must be stressed is that more information regarding growth rates and a knowledge of the seasonal reproductive cycles of the exploited species are necessary if exploitation is to be on a sound conservation basis. Of the algin yielding species large quantities of Laminaria pallida and Ecklonia maxima are available, especially of the latter which is cast up throughout the year. In five localities only, 6,000 tons dry weight of cast up kelps can be collected annually. Gracilaria confervoides is a commercially important agarophyte. About 1,000 tons dry weight of the species is cast up annually in the Langebaan —Saldanha area. Gelidiwm pristoides is also possibly a commercially important agarophyte. There is no clear evidence of Gelidiwm corneum on South African coasts. Aeodes orbitosa and Gigartina radula have been successfully used in the copper fining of beer and the latter species has been exported for this purpose. These species are not present in large amounts. ACKNOWLEDGEMENTS. Thanks are due to Mr. R. Simons for drawing Figs. 1 and 2 and to Dr. R. F. Milton for his comments on this paper. LITERATURE. Causey, N. B., PryrHercs, J. P., McCasxi11, J., Hum, H. J. and Wo tr, F. A. (1946): “Influence of Environmental factors upon the growth of Gracilaria confervoides,” Duke University Marine Station, Bull. No. 3, p. 19. Detr, E. M. and MicuHett, M. R. (1921): “The Tyson Collection of Marine Algae,” Ann. Bolus Herbarium, vol. 3, p. 89. Fox, F. W. and Strepuens, E. (1943): “Agar from South African Seaweeds,” S. Afr. Jour. Sci., vol. 39, p. 147. Tsaac, Wm. Epwyn (1937a) “South African Coastal Waters in relation to Ocean Currents,” Geogr. Rev., vol. 27, p. 651. (1937b): “Studies of South African Seaweed Vegetation. I. West Coast from Lambert’s Bay to the Cape of Good Hope.” Trans. Roy. Soc. S. Africa, vol. 25, p. 115. 92 The Journal of South African Botany. (1942): ““Seaweeds of possible economic importance in the Union of South Africa,” Jour. S. Afr. Bot., vol. 8, p. 225. (1949): “Studies of South African Seaweed Vegetation. II. South Coast: Rooi Els to Gansbaai, with special reference to Gansbaai,”’ Trans. Roy. Soc. S. Africa, vol. 32, p. 125. (1951): “Observations on the Ecology of Bifurcaria brassicaeformis (Kiuitz.) Barton,” Jour. Ecol., vol. 39, p. 94. Isaac, W. E., Frntayson, M. H. and Simon, M. G. (1943): “Agar from South African Seaweeds,” Nature, vol. 151, p. 532. Isaac, W. E. and Hewirt, F.: “The morphology, geographical distribution and ecology of Hypnea spicifera (Suhr), Harv.” This Journal. vol. 19, p. 73. LiGHTHELM, S. P., von Hoxp7, M. M., and Schumann, H. I.: ‘““The Composition of some South African Phaeophyceae,” paper presented to First International Seaweed Conference, Edinburgh, 1952. Newton, L. (1951): Seaweed Utilization, London. Papenruss, G. F. (1943): “Notes on South African Marine Algae, II,” Jour. South African Bot., vol. 9, p. 79. (1951): ““Notes on South African Marine Algae, III,” Ibid, vol. 17, p. 167 SHUTTLEWORTH, R. G. (1951): “The Industrial potentialities of South African Sea- weed,” S. Afr. Industrial Chemist, vol. 5. (a) “Part 2, Carrageenin and related products,” p. 76. (b) “Part 3, Agar,” p. 88. (c) “Part 4, Algin,” p. 115. ON THE HISTORY OF PLANT STUDY UPON THE WITWATERSRAND. By H. B. Giniimanp. Department of Botany, Witwatersrand University. The history of the study of the botany of an area begins with the identification and study of the plants which grow there. Should fate decree that the area should subsequently be industrialised, it is but natural that interest should swing to those plant activities which best serve the needs of the industrial community. The march of events upon the Wit- watersrand has followed this course, and it is time that the early botanical events of this region were recorded. The Witwatersrand area may be variously defined; most simply as the watershed between the Indian Ocean drainage (the Limpopo River) and the Atlantic Ocean drainage (the Vaal River) of southern Africa; most conveniently as the region of the gold mines of the Rand. It is approximately 2,500 sq. miles. Specifically Pretoria in the north, Vereeniging in the south and Heidelberg in the south-east are excluded. First settled by a European population in the early 40’s of the nineteenth century, it remained a sparsely peopled farming area until the finding of gold and the founding of Johannesburg in 1886 (Gray, 1937). Thereafter its history botanically falls into four periods; the Republican period terminating in the Anglo- Boer War; the colonial period terminating in the formation of the Union of South Africa; the early period of Union, culminating in the First World War, and the local development period whose first chapter may be said to have concluded with the Second World War. The Early Period (pre-1886). No botanical collector seems to have investigated the Witwatersrand in this period. To the north of the escarpment in the valley of the Jukskei River it seems probable that Capt. Cornwallis Harris hunted in pre- republican days. In 1911 Mr. W. Nelson records the finding of Bowiea volubilis Harv. at Blaauwbank near Krugersdorp “35 years before’, i.e. in 1876 and there is a specimen collected by him in the herbarium of the Transvaal Museum, Pretoria, labelled ‘““Nelson 514. Yukskyt River. Dec. 1877. Limpopo Sources” of Potamogeton nodosus Poir. Since local tradition associates the Witpoortjie stream and falls with the “Limpopo 93 94 The Journal of South African Botany. Sources” this, in all probability, was in our area. Nelson subsequently became a nurseryman operating the Nelsonia Nursery in Booysens, Johannesburg. The Republican Period. The early gold discoveries led on to the find of the conglomerates of the Reef, and it soon became clear that gold mining on the Witwatersrand was an enterprise of considerable importance—slowly it began to be realised of some permanence. The early mining town gradually became transformed into the embryo of the city of today and in the process a variety of services grew up, notably those of banking, law and explosives manufacture. I have singled out these three more especially because it is from their ranks that the persons with the interest to make the first serious botanical collections upon the Rand emerged. The Galpin-Gilfillan Family. The first Witwatersrand plant of which I can find mention in the literature is an orchid, Habenaria galpini Bolus (H. fetrapetala var. galpini Bolus) described by Dr. H. Bolus in Part I of his first volume of “TIcones Orchidearum Austro-africanarum t. 17 (1893)”. The published notes read, “Transvaal Republic: rocky places near Johannesburg, alt. about 1850 m.; fl. March. E. E. Galpin, 392”. Mr. Galpin was a bank manager from Queenstown who was early transferred to the Transvaal. The story is best described in the words of his son, Mr. E. A. Galpin of Mosdene, Transvaal, who writes to me as follows:— “Dr. Galpin was a banker by profession and had 40 years’ service mostly as manager in various parts of the country, in the Oriental Banking Corporation until it was absorbed by the Bank of Africa, and throughout this Bank’s complete history until it was in turn absorbed by the National Bank, retiring from this bank in 1918. Thus a great deal of his collecting was confined entirely to ‘leisure’ hours. He was always a very busy man with a tremendous capacity for work. “Tn 1887 he was manager of the Bank of Africa in Grahamstown and was transferred to their Johannesburg branch as sub-manager, arriving there in 1888. He lived in a tin shack on the present site of the New Law Courts. Owing to the shortage of trained bookkeepers and accountants on the Rand, there was a heavy loss of bank staff to the mining houses, stock-brokers and others, resulting in a constant drain on their supply. This placed a severe burden of work on the remaining staff. The Stock Exchange was very active and as it only closed after normal banking hours, my father as sub-manager was in charge of the scrip and had to On the History of Plant Study upon the Witwatersrand. 95 remain in office until dark, clearing the Stock Exchange business. This was all done in cramped temporary quarters. It was only after he had left Johannesburg that the Commissioner Street Central Branch of the Bank was built. “This meant that he had very little time in which to botanise, and most of his collecting was done on Sunday afternoons along the Parktown and Kensington hills which were then virgin veld. He spent only about six months in Johannesburg and was sent off at a moment’s notice to take over the managership of the Barberton branch, where there had been staff trouble. “In 1892 he married Marie Elizabeth de Jongh, in Pretoria. He had met her in Barberton. She was sister to Jim de Jongh, one of the earliest chairmen of the Stock Exchange and a quondam owner of that well-known residence, “Saratoga Villa’, Doornfontein. She was the Marie E. Galpin. Her sister was married to Douglas Flemmer Gilfillan, a well-known Johannesburg lawyer and pioneer and a member of the Reform Committee who was imprisoned after the Jameson Raid. My father interested D. F. Gilfillan in botanical collecting, of which he did quite an appreciable amount in the environs of Johannesburg, sending the material to my father who recorded it as collected by D. F. Gilfillan. Apart from an occasional holiday visit, during which a few hours might be spent in collecting, my father had no further connection with Johannesburg.” Actually Dr. E. E. Galpin’s collections upon the Witwatersrand were not numerous, but he was carefully compiling an herbarium in which he preserved, as his son’s letter shows, plants collected by D. F. Gilfillan from the Johannesburg area. His wife also collected extensively upon the “ridge above Jeppe’s Town”. Happily, the Galpin herbarium was donated to the Government after Union and formed the basis of what has now become the National Herbarium at the Union Buildings, Pretoria, and in it Gilfillan’s and Mrs. Galpin’s plants are preserved. Mr. Gilfillan is also survived by a son, who has likewise very kindly sent me a letter with pertinent matter. “Dr. and Mrs. Galpin were my uncle and aunt, my mother being a sister of Mrs. Galpin. My father, who is the Gilfillan you refer to, was always a keen naturalist and his brother-in-law, Dr. Galpin, caused him to become interested in botany. “My father was a solicitor by profession, and he came through to Barberton in 1888, where he practised as a solicitor. In those days, Johannesburg was thought to be a flash in the pan, and Barberton was believed to be the real goldfield. I think he was only in Barberton for three years, and consequently came to Johannesburg in about 1891 or 1892. At that time Dr. Galpin was either Manager or Accountant at the 96 The Journal of South African Botany. Standard Bank, in Johannesburg. My father reached considerable eminence in his profession and was a partner in the firm of Bowman and Gilfillan, which was established before the Boer War. Since his youth my father was always interested in nature including plants and was a keen observer. He became interested in botany due to his friendship for his brother-in-law, Dr. Galpin. They remained life-long friends and went on many botanical expeditions together. I believe that in many respects my father acted as a collector for Dr. Galpin and handed over his specimens for inclusion in Dr. Galpin’s collection. “T do not think that any of his correspondence on botanical matters survives but he kept a diary during the Boer War and ever since, till the date of his death at the age of 83, in 1948. The copies of the diary during the Boer War were lost, but the balance are in my possession. I have only read a few of the more recent sections, but I hope some day to investigate whether anything can be made of it. In his diary he recorded all matters which interested him and I feel sure that he would have recorded his botanical work. As a matter of interest my father was one of the founders of the Wild Life Protection Society and was for many years its President. He was a member of the National Parks Board of Trustees and was a keen ornithologist, on which subject he wrote many papers and articles.” This family group then provides us with the first records of actual Johannesburg plants. Dr. Paul Conrath. North-east of Johannesburg there was established very soon after the discovery of the Reef, as a State Monopoly, the “Zuid-Afrikaansche Fabriek voor Ontplofbare Stoffen” which supplied the mines with explosives. To this in 1895 came Paul Conrath as Assistant Manager. By the chance absence of the Manager on leave just before the Anglo- Bore War, he found himself in charge during the occupation of the Transvaal. This may perhaps have led to a period of unexpected leisure: at any rate he then made extensive collections of plants on the factory estate which became in consequence the locus classicus for many a species described from the Transvaal. This is indeed fortunate, for the estate of the factory, which is also an explosives magazine, is required by law to be extensive, and has also been enlarged with the progress of the factory, which is now one of the principal production centres of Messrs. African Explosives and Chemical Industries Ltd. This well-known firm has always adopted an enlightened policy towards Wild Life, and the estate is carefully conducted as a Nature Reserve. Conrath left the Transvaal On the History of Plant Study upon the Witwatersrand. 97 in 1902, but continued to occupy his leisure with his collections, and to describe new species from his residence in Germany. (Conrath, 1906, 1914.) Other Items of the Period. Joubert Park, now buried in the heart of the City, was at that time on its outskirts—a block away from the famous old Wanderers Club which has now been incorporated in Park Station—and its first curator, Mr. R. W. Adlam, was a correspondent of Dr. Medley Wood of Durban and sent him many seeds. Several of his gatherings, some from Joubert Park itself, are likewise preserved in the National Herbarium. Botany had its commercial aspect on the Rand even at that early date for someone prepared a collection of plants, styled the “East South Central African Herbarium”, which eventually found its way to Kew, as an extract from a letter from Dr. Turrill, Curator of the Kew Herbarium, shows:— “. . . Kew purchased the E.S.C.A. (East South Central African) Herbarium from O. Weigel, Leipsig, in February, 1895. It consisted of 614 specimens—‘5 centuries’. Weigel got it from Laidley & Co., about whom I can find nothing. They may have collected the specimens or had them collected. There are 8 specimens of Cyperaceae at Kew from the E.S.C.A. Herbarium, but not one of these bears a date. There are printed and stamped data on the labels plus C. B. Clarke’s determinations. The style of printing suggests a date past the middle of last century . . .” This note was prepared by E. Nelmes, who conducts the study of the Cyperaceae at the Kew Herbarium, on 7 February, 1951. By the time the new regime of the Transvaal Colony had become established then, several hundred plants had been collected upon the Wit- watersrand and quite a large number had been described as new by various authors. Other collectors of this period, of whom I can find no further informa- tion, were Barker, J. Hall, and a reference to specimens from Max Leitchlin, a well-known nurseryman of Baden-Baden. The Colonial Period. With the establishment of Colonial Government, an official botanist was soon appointed, and thus we find Dr. Joseph Burtt Davy entering upon the Transvaal scene in 1903. Burtt Davy made no very extensive collections upon the Rand but specimens of his gathering in 1905 (Germiston, Observatory Ridge) and 1908 (Ormonde, Booysens) survive. The first Check-list of Transvaal plants (Burtt Davy and Pott-Leendertz, 1912) resulted from his enthusiasm, and the first Flora of the Transvaal 98 The Journal of South African Botany. (Burtt Davy, 1926), unhappily never completed, was due to his subsequent labours at Oxford. Dr. I. B. Pole Evans, fortunately still with us, was soon thereafter appointed mycologist to the Transvaal. His prodigious energy ultimately led to the development of the Division of Plant Industry, and that beautifully illustrated “Flowering Plants of South Africa” in 1921, in which not a few Witwatersrand plants have received notice for the first time. During his period as Chief of the Division, the National Herbarium in its present form took shape. He made personal collections upon the Witwatersrand as early as 1907. However, it was not to the official botanists to whom we owe the first appearance of accounts of Johannesburg’s flora. Capt. H. T. Ommanney collected in the Johannesburg area in 1901/02 and his plants were listed from the British Museum in the “Journal of Botany” for 1902 (Moore, 8., 1902). Ipomaea ommanneyi Rendle was there described for the first time together with identifications of the collection. Very shortly afterwards the aptly named Dr. R. F. Rand, who had already contributed the first botanical notes concerning the Mashonaland Flora, arrived upon the Witwatersrand, and with Johannes- burg as his headquarters, began to study the local flora. His “Wayfaring Notes in the Transvaal” (Rand, R. F., 1903) began to appear in 1903, and contain the first notes on the fertilisation of the flowers of local plants—notes which were not supplemented until the work of Davidson in 1941. In the course of his description, Dr. Rand mentions the tree fern Cyathea dregei Kunze along the stream at Witpoortjie. Alas, this plant no longer occurs there or indeed anywhere on the Rand. As far as I know, it, together with the white Arum (Zantedeschia albomaculata Baill.), has been exterminated from the local flora during the sixty-four years of the Witwatersrand’s industrialisation. The first plant to bear the name of the City of Johannesburg, Senecio johannesburgensis S. Moore (1903), was a gathering of Dr. Rand’s No. 1135 “Johannesburg, Klipriviersberg to southward”’. Miss Alice Pegler, who contributed so much to our knowledge of the Transkeian flora, visited and collected plants on the Rand in 1904. Next followed a period when the botanical giants of those days visited the Rand. Dr. Medley Wood, of “‘Natal Plants” fame was here in 1903, though he did not publish anything on his finds. In 1904 Dr. H. Bolus gathered plants in the south of Johannesburg and his daughter-in-law Dr. L. Bolus has very kindly furnished me with extracts from his diary which are of interest. “I have been looking at Dr. Harry Bolus’ diary and find he and his son Frank (who later was my husband) were in Johannes- burg on 17th and 19th February, 1904. The entry reads:—‘Feb. 17 Wed. arr. Johannesburg, 5689 feet, 11 a.m. Victoria Hotel . .. Walked about the On the History of Plant Study upon the Witwatersrand. 99 town—rained aftn., changed our pants ... Hermann Mundt came to hotel and had a long talk. Feb. 18. Thurs. Went by ’bus to Booysens some 4 miles south of the town—just beyond plantations and enclosures. Gathered on open grassy hills—say “‘in graminosis apertis ca. 5900 ped.” .. . Thunderstorm came on just as we reached the starting place of the ‘bus on return—and continued with heavy rain—In the ’bus met one Krohn who .. . told me of Nelson, the collector, who lived still as a nursery- man at Nelsonia—a mile from where we had been, who would, K. said, have been very glad to see me. At Johannesburg still raining . . . and then to our hotel. (On our trip to Booysens lost our best frame with paper.)’ I have copied the above entries from the diary as giving all the information available in connection with Dr. Bolus’ visit to Johannesburg. I do not know of any other visit...” Within a few months the South African Association for the Advance- ment of Science, holding that year a joint meeting with the British Association for the Advancement of Science, held its business meeting in Johannesburg in July and Dr. R. Marloth, an active member of Council, was present. This visit was commemorated botanically in the well- known Aloe marlothii Berg, whose flowers were “collected by Marloth on the Klip River Hills south of Johannesburg” (Reynolds, 1950, p. 480). At about the same time, and probably in connection with the visit of the British Association, Prof. A. Engler of Berlin visited Johannesburg and made a small collection. This is commemorated in Hulophia engleri Rolfe “‘S. Africa. Transvaal; amongst stones at Klip River Mountains, 1830 m. Engler, 2745.” Dr. Engler also honoured the City botanically for his Hermannia johannesburgiana Engl. appeared in 1907. Dr. Engler re-visited the City in 1910. Other notable personalities to gather botanical specimens on the Witwatersrand during this period were Dr. Haagner, Director of the Zoological Gardens, Pretoria, and Mrs. Pott-Leendertz, noted earlier as the co-author of the first Transvaal Check-list, who was the Botanist to the Transvaal Museum. Gatherings were also made by Sampson, Agronomist in the Department of Agriculture on the East Rand, and Preddy in the Germiston area, and more generally by W. McDonald who was Director of Agriculture for the Transvaal, and edited the Transvaal Agricultural Journal. The Ist Union period (1910-1918). Apart from the official botanists who of course continued with their duties, working upon the Witwatersrand as occasion demanded, those who made local plant studies included several interesting personalities. It was during this period that Prof. C. E. Moss first arrived upon the Rand, 100 The Journal of South African Botany. but as his work properly belongs to the next period we here simply note his arrival. Among those of the community he found were interested in plants, the most famous was perhaps Archdeacon F. A. Rogers, who made such extensive collections all over southern Africa. Rogers started collecting on the Rand in 1915, and in 1918 he and Prof. Moss joined forces and made a very extensive collection together. Prof. Moss’ fore- sight resulted in the preservation of the collections of Mr. G. T. Weeks in what has now become the Moss Herbarium. Mr. Weeks was ““Town Botanist”’, with vaguely defined duties, including the inspection of abattoirs and the judging of animals, but he made fine specimens of local plants and we owe to him, as well as to Adlam mentioned earlier, what we know of the plants that grew wild in Joubert Park in its earlier days. Another notable personality, happily still with us, who started her botanical collections in those times is Miss Gwen Edwards. Other names of the period are those of McIntosh, Baker, Kling, Hohlen in the City area, Halford at Bedford Farm, Sykes at Natal Spruit, C. H. Fenn at Germiston and in Johannesburg, whilst Mr. F. Frith, the well-known succulent grower, started making his collections, and alas his all too few specimens, in this period. The Inter-war period. (1918-1939). Despite the work which had gone before, this is actually the most important period in the growth of the study of plants on the Rand, for it saw the broadening of the field of interest from the purely nomenclatural cataloguing of what grows here to the experimental study of the behaviour of the plants in relation to Agriculture and Industry. During this period Prof. Moss’ basic taxonomic work was succeeded by Prof. Phillips’ ecological studies of grassland and Mrs. Rebecca Lurie Brown’s establish- ment of the Mycological section of the Timber Research Laboratories. Prof. C. E. Moss was appointed to the chair at the South African School of Mines and Technology in 1917, then in the process of developing into the University College, Johannesburg, and subsequently into the University of the Witwatersrand. He very soon saw that progress in the development of knowledge of Witwatersrand plants, and of their behaviour, depended upon the compilation of a Flora, with a detailed record of the plants that grew here, and set about the gargantuan task of preparing one, virtually single-handed. For the purpose he began the careful and systematic collection of specimens of all local plants and inspired many others to assist him. During his lifetime some 18,000 specimens were amassed, critically named and arranged, and formed the solid foundation of the present Moss Herbarium. Moss spent most of a On the History of Plant Study upon the Witwatersrand. 101 his long vacations at Kew critically naming his material, but did not live to complete his self appointed task. The material exists for its completion. Prof. Moss, who in earlier days had carried out a notable ecological survey of the Peak District in Britain, did not overlook ecological matters and one of his students, Miss Dora Weintroub (1933), fortunately still with us, published the first ecological account of vegetation upon the Rand. Although collecting extensively over the length and breadth of the Rand, Prof. Moss had his favourite sites, notably the Witpoortjie Gorge and Falls and the Houghton Estate, while he naturally collected regularly at Milner Park, the present site of the University. He died in 1930. In the Moss Herbarium are preserved many specimens collected by Prof. Moss’ students, while at the National Herbarium in Pretoria are many records of his and of others, perhaps stimulated by his enthusiasm. Mr. E. G. Bryant, who conducted a mining and engineering business, and moved to Prieska in 1920, made an extensive collection, principally at Turffontein and on Hospital Hill in 1918-19; Miss Sydney M. Stent brought back an extensive collection of grasses from Henley-on-Klip from a visit in 1922; Mr. and Mrs. R. G. N. Young made collections widely on the Rand commencing about 1926; Miss J. J. Gilmore, of Jeppe High School for Girls, added to the record in 1926-28; Mr. Flugge- de Smidt commenced his work at Geduld in 1929 and even Sir Julius Jeppe evinced interest in lawn grass mixtures in 1925. Two notable overseas visitors at this time were Prof. A. S. Hitchcock, of the United States Department of Agriculture, who made a wide collection of grasses during a brief late winter visit in 1929, and Dr. J.S. Hutchinson of Kew, who made a quick dash towards Johannesburg from Trene during his extensive South African tour in 1929. (Hutchinson, 1946.) This period also saw the commencement of the notable work of Prof. J. M. Watt who was appointed to the Chair of Pharmacology at the Witwatersrand University in 1920 and published his well-known “The Medicinal and Poisonous Plants of Southern Africa” in 1932. Prof. Moss was succeeded by Prof. John Phillips who, his interests lying primarily in the ecological sphere of botany, proceeded with the aid of Dr. T. D. Hall, Agricultural Adviser to Messrs African Explosives and Chemical Industries Ltd. to develop the estate Frankenwald, be- queathed by Alfred Beit to the “University of the Transvaal’’, and which had come into the possession of the Witwatersrand University. By a strange coincidence the estate lies immediately adjacent to the Modder- fontein Estate, scene of Dr. Paul Conrath’s collections of thirty years before. Professor Phillips gave one of the earliest accounts of the vegetation 102 The Journal of South African Botany. of the Rand (1933), and made Frankenwald a centre of research upon ecological problems, more especially those appertaining to pasture manage- ment and conservation. His students hold posts in the field services, from the Cape to the Sudan and the Gold Coast, and his contribution in this way alone is impressive, including as it does the 100 or so students trained in Soil Conservation from the returning veterans of the second World War. Frankenwald remains a notable centre of ecological research, where the biotic community, climatic and edaphic factors, and the impact of fire, grazing, and protection on native grassland are studied, together with the more directly agronomic problems of fertiliser application and chemical weed control, which continue under the aegis of Messrs African Explosives and Chemical Industries Ltd. One of Prof. Phillips’ students, Dr. P. E. Glover, contributed the first phenological study of Witwaters- rand plants (Glover, 1937), and another, Miss E. Cohen (now Mrs. Gillon), took up the study of the native flora of the Houghton Kopjes, a study which was, in due course of time, to lead to the development of that famous municipal garden ““The Wilds”’. Meantime the curatorship of the Moss collections passed to Mrs. M. Moss, who valiantly maintained and developed them in the intervals of a heavy teaching programme for the remainder of this period. While the Moss Herbarium, now in existence, provided a nucleus around which the botanical work of the Witwatersrand proceeded steadily, there were others working independently. Thus on the East Rand Flugge-de Smidt quietly continued with those studies which were to result in “Flowers by the Roadside” in 1947, the Rev. Father Murray collected extensively along the Reef and R. D. Bradfield collected in Benoni in 1934 for the Albany Museum in Grahamstown. Luckily Bradfield’s material was sufficient for a duplicate set of specimens to be sent to the National Herbarium where they may now be readily consulted. Prof. Phillips’ activities as a member of the Botanical Survey of South Africa led him to make a large “Botanical Survey Collection’, at first maintained separately, but subsequently incorporated in the Moss Herbarium. Dr. Elaine Young (now Mrs. Laughton), appointed to Prof. Moss’ staff in 1920, early set about teaching mycology. Amongst her students was Miss Rebecca Lurie (now Dr. Rebecca Lurie Brown) who, becoming interested in the fungi which caused the decay of wood, soon set in motion those researches, which, with the generous backing of the Chamber of Mines, have led, not only to the substantial saving in costs for the Mining Industry in timber and ventilating apparatus but also to the establishment of the Timber Research Laboratories in Johannesburg. In concert with Miss Dora Weintroub, mentioned earlier, Dr. Brown traced On the History of Plant Study wpon the Witwatersrand. 103 a sporotrichosis, which threatened to cause considerable unrest, to a fungus and thus averted a strike. So valuable did her work appear to the Chamber of Mines that on her retirement from active participation in the work of the laboratory, the Chamber honoured her service by founding the Rebecca Lurie Brown Prize in Botany at the Witwatersrand University. Dr. Elaine Young, mentioned earlier, had in collaboration with Miss C. G. Crocker (1933), commenced the study of cytology with an investi- gation of the chromosomes of Digitaria in 1933, while Dr. H. Weinmann’s notable researches in plant physiological chemistry commenced towards the close of the period. (Weinmann, 1940.) This was the stage of development reached with the outbreak of the Second World War, a solid foundation had been laid and the stage had been set for the further specialisation and ramification which were to follow. ACKNOWLEDGEMENTS. Much of my information concerning Dr. Paul Conrath I owe to Mr. D. Allen of Johannesburg, and I have to record with gratitude the help of Mrs. M. Moss, lately senior lecturer in Botany of the Witwatersrand University, and of Miss R. H. Gunn, Librarian, Division of Botany, Department of Agriculture, in compiling these notes. REFERENCES. Bonus, H. (1893): “Ieones Orchidearum Austro-africanarum,” pt. i, t. 17. Burtt Davy, J. i (1926); ii (1931): “A Manual of the Flowering Plants and Ferns of the Transvaal with Swaziland.” Burtr Davy, J. and Porr-LEENDERTZ, R. (1912): “A first check-list of the Flowering Plants and Ferns of the Transvaal and Swaziland,” Ann. Transvaal Mus., IDOL, 83, WiN@, ConratH, P. (1906) in Vierteljahrsschr. Nat. Ges. Zur., LI, 131; and in Kew Bull., 135, 1914, and ete. See also Baker, J. G. in Schinz, Beit.z.Kent.d.Afri- eanischen Flora, XVI. Bull. Herb. Boiss., iv, 1904. Davipson, R. L. (1941): “‘A note on the anthesis of some common grasses near Johannesburg and the relation of anthesis to collection of pollen for medical purposes,” Jour. S. Afr. Bot., 145. Eneuer, A. (1907) in Engl. Jahrb., xxxix, 585 and see Rolfe in Kew Bull., 282, 1910. Guover, P. E. (1937): “A contribution to the ecology of the Highveld Flora,” S.A. Jour. Sci., xxxiii, 443. GRAY, J. (1937): “Payable Gold,’ C.N.A., Johannesburg. Hutcuinson, J. (1946): “A Botanist in Southern Africa,” p. 384. Martoru, R. (1904): see Rep. 8. Afr. Ass. Adv. Sci. Johannesburg, p. 11. Netson, W. (1911) in South African Gardening and Agriculture, Ist June, 1900. OMMANNEY, H. T. (1902): see Moors, S. in Jour. Bot. xl, 380. Puaiwuies, J. F. V. (1933): “The Flora of the Golden City,” in Macmillan, “The Golden City,” Collingridge. 104 The Journal of South African Botany. Ranp, R. F. (1903) in Jour. Bot., xli, 52, etc.; also Moorg, S., 1.c., 131, ete. REYNOLDS, G. W. (1950): “The Aloes of South Africa,” p. 480. WEINMANN, H. (1940): “Storage of Root Reserves in Rhodes Grass,’ Plant Phys. 15. 467. Werntrous, D. (1933): “A preliminary account of the aquatic and sub-aquatic vegetation and flora of the Witwatersrand,” Jour. Hcol., xxi, 44. Youne, E. M. and Crocker, C. G. (1933): “Chromosome studies in Digitaria sp. I,” S. Afr. Jour. Sci., xxx, 258. BOOK REVIEWS A BoraNicaL SURVEY OF THE KEISKAMMAHOEK District. By R. Story. Bot. Surv. 8. Afr. Mem.27. 1952. pp. xi & 184. 85 photos, | map, 5 graphs. 10s. So much of the vegetation of the country has undergone or is under- going change as the result of man’s activities that detailed studies of any part are of importance both as a record and still more as a guide to the proper procedures for preservation. This memoir is a case in point. It deals with an area which supports a considerable population and has a vegetation that has undergone a good deal of change but change not yet carried to the extent of obscuring the original relationships. It is an area with considerable diversity in topographic and climatic conditions. After a rather full account of the climatic conditions, the descriptions of the vegetation are based on the existing communities. While inter- relationships to one another and to habitat are discussed there is no attempt to force any rigid doctrine. For descriptive purposes a primary classification into “woodland” and “grassland”? is adopted, a scheme which really does not have much more than its essential simplicity to recommend it. “Woodland” comprises communities with scattered trees or bushes, true forest, and communities composed wholly of bushes. Six types are recognised. Five of these form a series ranging from Acacia scrub through different types of more closed scrub to dry and finally moist forest. These correspond to successive zones of increasing altitude and rainfall. The sixth type is the so-called ““macchia” which occupies a zone above that of the moist forest, but which is so distinct both in structure and composition that it has to be treated separately throughout. Incidentally it is a little unfortunate that an alien local term be used for vegetation that has little similarity either in habitat or composition with its Mediterranean prototype. Local terms are used for other communities so why not here? The noncommittal “fynbos” is available. Grassland is divided into “‘sweetveld”’ at the lower and drier levels, and “‘sourveld” at the upper and moister. Each is subdivided. The treatment of the various types is somewhat unequal. Acacia scrub and “macchia” are dealt with in some detail whereas forests are treated much more shortly. In each case the descriptions are followed by a discussion of relationships and problems of usage, preservation and treatment. The various forest and scrub types are looked on as being true climatic climaxes for the different zones. “Macchia” is also a climax in the zone 105 106 The Journal of South African Botany. above the limits of forest though it is successional to and replaced by forest at lower levels. Within the area grassland is not a true climax though it does become a stable community where local conditions are unfavour- able to the full development of the succession. At the higher levels the sourveld is maintained by destructive burning of the “macchia’”’. The mountains here are not of sufficient altitude to reach the zone of mountain grassland. Changes brought about by burning or overgrazing in the various types are discussed together with methods of reclamation or stabilisation. The whole survey is one that has been carefully worked out and which has yielded results of definite practical importance. The book, is however, not one that is easily read owing to a certain verbosity of style and to the inclusion of a large amount of rather extraneous detail. Editorial pruning would have assisted the average reader. The book is illustrated by no fewer than 85 photographs. These have been carefully selected to illustrate the various points. Unfortunately in many cases the repro- duction does little justice to the originals. The vegetation map is on a large scale but owing to a similarity in the symbols used for different communities is not very clear at first glance. The volume concludes with an alphabetical list of plants, an appendix with some useful hints on the use of meteorological instruments, and a bibliography. The last which extends over five and a half pages has a number of striking omissions and contains a number of items of little obvious value. R. S. ADAMSON. SourH AFRICAN POLLEN GRAINS AND Spores. By EH. M. Van Zinderen Bakker, Phil. Nat. D. (Amsterdam). Part I. Pp. 72, 16 plates. Published by A. A. Balkema, Amsterdam and Cape Town, 1953. 15s. The publication of Part I of Dr. Van Zinderen Bakker’s work on the pollen grains and spores of South African plants is an event to which local students of palynology have looked forward for some time, as this is likely to be a work of the greatest importance to both taxonomy and pollen analysis in South Africa. The pollen grains described and figured are arranged according to Engler’s system of classification, and Part I includes the Gymnosperms, Monocotyledons and the Dicotyledons up to the family Hydnoraceae. Not all the South African genera are included, but only those which are likely to be of most value to the palynologist interested in pollen analysis or in allergy, and it is to be remarked that some introduced genera, such as Pandanus, Saccharum, Casuarina, Populus and Cannabis have been for various reasons included. Book Reviews. 107 It is unfortunate that the endemic family Grubbiaceae, as well as the Santalaceae, which ought to be in Part I, have been omitted, but we may hope that this deficiency will be made good in a supplement. A complete description is given of each grain figured, and in many cases, additional species belonging to the same genus are described, although not figured. Great precision is given to the author’s work by the citation of the herbarium specimens from which the material was taken, and also the herbarium in which the specimens are to be found. Mention is made of the general distribution in South Africa of most of the species described. The sixteen plates included in this Part are reproductions by photo- lithography of pencil drawings made by the author, and the figures, almost all drawn from acetolysed material, have a delicate beauty unsur- passed by any figures of pollen which have been published since the copper-plates of Hugo von Mohl’s “Pollenkérner”’ (1834). A bibliography and a convenient and useful index to all the genera and species described and figured in Part I are included, as well as a glossary of most of the special terms used in the body of the work. The excellent paper, clear typography and beautiful plates are a credit to the publisher, and all palynologists will eagerly await the four subsequent parts of this useful work. S. GARSIDE. din Wee ae a i ililyGig ae MR) AD lan eT EE a henna rma: Guar Me ipaaruee:: Jasoetg ty ISSUED DEC 22 i953 JOURNAL OF SOUTH AFRICAN BOTANY VOL. XIX. PLANTAE NOVAE AFRICANAE. Ex Africa semper aliquid novi.—Pliny. SERIES XXXI. By R. H. Compton. (With Plate XVIII.) Podalyria insignis Compton sp. nov. Frutex robustus, foliis orbiculatis, coriaceis, obtusis, rugosulis, dense griseo-tomentosis. Injflorescentiae omnino velutinae, dense paniculatae, terminales et laterales. Corolla conspicua, glabra, flava. Habitat. Cape Province. Montagu Division: Dobbelaar’s Kloof, 4000 ft. alt., 25 Sept. 1946, Compton 18441. (Type, in Herb. Nat. Bot. Gdns.) May 1940, Esterhuysen 1869, Bond 337, Walgate s.n. Kiesiesberg, 3500 ft. alt., 27 Sept. 1946, Compton 18495. A robust evergreen shrub, up to 2 m. high, with long leafy rod-like branches. Young stems terete, up to 6 mm. diam., densely grey-tomentose. Leaves ovate-orbicular, coriaceous, entire, cordate, obtuse, mucronate, densely tomentose on both surfaces, especially below, tending to become rusty with age, rugulose above, the whole venation very prominent below but only the midrib prominent above, up to 5 cm. long and broad, on a stout petiole c. 4 mm. long. Inflorescences terminal and subterminal, pyramidal, up to 10 cm. long and broad, the flowers densely massed and very conspicuous. Peduncles, bracts, pedicels and calyces densely rusty- velvety-pubescent. Bract c. 10 mm. long, 14 mm. wide. Pedicel c. 15 mm. long. Calyx truncate or slightly intruse at the base, c. 9 mm. long, variable in lobing: anterior sepal almost free, ovate-lanceolate, c. 8 mm. long, 4 mm. wide; the two posterior sepals united nearly to the apex; the lateral sepals sometimes free, sometimes united to the posterior sepals. Corolla completely glabrous, distinctly veined, bright yellow. Veaxillum with claw 109 110 The Journal of South African Botany. c. 3 mm. long, the lamina orbicular, obtuse, 16 mm. long, 25 mm. wide: alae with claws c. 5 mm. long, the laminae obliquely ovate, obtuse, c. 13 mm. long, 10 mm. wide: carina with claws c. 5 mm. long, the lamina ce. 13 mm. long, 9 mm. wide. Stamens free almost to the base, filaments c. 13 mm. long, broadening towards the base. Ovary densely hirsute, with several ovules. Fruit about 4 cm. long, 1 cm. wide, densely velvety, the valves twisting spirally on dehiscence. A magnificent plant, well distinguished from all other known species of Podalyria except P. chrysantha Adamson by the yellow colour of its corolla, by the dense paniculate terminal and subterminal inflorescences and by many details of leaves and flowers which are larger in all dimen- sions. It is so far only known from two localities in the Montagu Division where it occurs on the south side of mountains bordering the Karoo, near their summit. It has not so far been brought into cultivation, but it richly deserves it. DYMONDIA Compton gen. nov. Compositae—Arctotideae—Arctoteae. Capitulum sessile, subterminale, solitarium. Involucrumcampanulatum, bracteis multiseriatis, non appendiculatis. Receptaculum leviter convexum, alveolarum marginibus dense hirsutis. Flosculi radii feminei, ligulati; disci hermaphroditi, tubulares. Achaenia glabra. Pappus paleolis hyalinis, biseriatis, brevibus. Styli rami breves, radii ovati, disci truncati. Genus adhuc monotypicum. So far monotypic. Capitulum solitary, subterminal, sessile. Involucre campanulate, the bracts multiseriate, without appendages. Receptacle slightly convex, the margins of the alveolae densely hirsute. Ray florets female, ligulate: disc florets hermaphrodite, tubular. Achenes glabrous. Pappus short, of two rows of hyaline scales. Style branches short, those of the ray florets ovate, of the disc florets truncate, long-exserted. Dymondia Margaretae Compton sp. nov. Fruticulus caulibus subterraneis implectentibus, ramulis brevibus erectis, foliosis. Folia explanata, lineari-oblanceolata, parum sinuato- dentata, supra viridia, glabra, subter dense albo-sericeo-lanata, margini- bus leviter inflexis. Involucrum c. 12 mm. long, 10 mm. lat. Flosculi radu 8—13, ligulati; disci numerosi, tubulosi; omnes flavi. Aliter ut in generis descriptione. Habitat. Cape Province. Bredasdorp Division: Flats south of Bredas- dorp, M@. Dryden-Dymond, Nat. Bot. Gdns. 1710/33. (Type, in Herb Nat. Bot. Gdns.) Bontbok National Park, in shallow dry pan, 8 October 1950, Compton 22178. Plantae Novae Africanae. 111 A much-branched sub-shrub with a network of rhizome-like branches at or just below the surface of the ground, with numerous short erect shoots bearing leaf-rosettes, the whole forming a dense mat completely covering the soil. Horizontal rhizomes up to 8 mm. diam. Leaves numerous in each rosette, spreading horizontally, up to 5 cm. long and 5 mm. wide, usually less, narrowly linear-oblanceolate, obtuse, sinuate-dentate with 2 or 3 shallow obtuse teeth on each side, the margins somewhat incurved, showing the white underside (more so when dry): upper surface dark green and glabrous, almost glossy,.lower surface with densely felted white silky-woolly indumentum. Capitula not numerous, solitary, subterminal, (growth being apparently sympodial), sessile and closely clasped by the surrounding foliage leaves. Involucral bracts multiseriate, the outermost longest, c. 13 mm. long, linear, acute, their distal halves having lanate indumentum: median bracts broadly oblong, c. 8 mm. long, 3 mm. broad, with a narrow subapical dorsal patch of indumentum: the innermost rather shorter and narrower. Receptacle somewhat convex, the insertion of the dise florets surrounded by a ring of numerous white hairs. Ray florets 8—13, female: ovary glabrous, turbinate: tubular portion of corolla 6 mm. long, lamina elliptical, 12—14 mm. long, 3 mm. wide, yellow, minutely fibrollose and with a dark median stripe beneath, with about 4 veins: style branches short, ovate, lobulate: pappus of two rings of short broad laciniate hyaline scales. Disc florets c. 30, hermaphrodite; ovary, basal hairs and pappus as in the ray florets; corolla tubular, the basal portion slender, 2 mm. long, the upper portion campanulate, 2 mm. long, the lobes 1 mm. long, the margins slightly inrolled, hooded and minutely lobulate; anthers shortly sagittate; style branches short and truncate, long-exserted. The genus and species are named in memory of the plant’s first collector, Miss Margaret E. Dryden-Dymond, gardening apprentice and improver at the National Botanic Gardens, Kirstenbosch from 1927 to 1932, when she became a member of the staff: she resigned in 1937 to take up teaching work and died in June 1952. The plant was first collected by Miss Dymond on a Kirstenbosch expedition to the Bredasdorp District. Only one patch was seen, a few square yards in extent, by the roadside. Living material was brought back and propagated; and it has been extensively used at Kirstenbosch ever since in rock-work and for covering small areas of bare ground, a purpose for which it is eminently suited. In spite of repeated search, the original spot was never located subsequently, and no further records of the plant were made until 1949, when recognisable fragments were brought to the Kirstenbosch Herbarium by Mr. H. David, who had collected them in flat ground at the foot of the Potteberg, Bredasdorp 112 The Journal of South African Botany. District. In October 1950 it was found uniformly covering a large shallow dry pan in the Bontbok National Park. (Compton 22178.) Dymondia Margaretae does not fit into any genus hitherto described. Its nearest affinities are with Landtia and Arctotheca, from one or both of which it differs in its peculiar woody-rhizomatous habit, its sessile capitula, non-appendiculate multiseriate involucral bracts, densely hairy alveolar margins and biseriate pappus scales. Its peculiar habit of growth and the attractive appearance of its vegetative parts make it a valuable garden plant, even though its flowers, which may be produced at almost any season of the year, are never profuse. It can be propagated with great ease by breaking up into pieces. Its adaptability to climates other than that of the Cape coastal belt has not been determined, but it can withstand a light frost. Cenia expansa Compton sp. nov. Species nova, C’. Duckitteae L. Bolus affinis, sed floribus ligulatis parum paucioribus minoribusque, flavis non aureis, et aliter differt. Habitat. Cape Province. Malmesbury Division: Kotze (mouth of Berg River), 2 Sept. 1944. Compton 15936 (T'ype, in Herb. Nat. Bot. Gdns.) Piquetberg Division: Near Sauer, 10 Sept. 1949. Barker 5778. Specimens cultivated at Kirstenbosch, source unknown, in Herb. Nat. Bot. Gdns. (24 Sept. 1941, Walgate 31 Oct. 1941, Henderson 2168, 2 Nov. 1944.) A soft strongly growing annual, branching mainly from the base, the branches decumbent and leafy below, becoming erect and pedunculoid above, up to 50 cm. long. Stems pale, terete, hirsute throughout. Basal = LSS Fie. 1. Cenia expansa Compton. 1, Fragment of stem and cauline leaves, nat. size. 2.3. 4. Capitulum from above, below and in section, nat. size. 5. Outer involucral bract x 5. 6.7.8. Marginal, ligular and disc florets x 5. Hort. Nat. Bot. Gdns., 31 Oct. 1941. (Del. M. Walgate.) Plantae Novae Africanae. 113 leaves up to 7 cm. long, 2 em. wide, 2—3-pinnatifid in the upper half, the cauline leaves progressively smaller and less divided: segments linear- lanceolate, channeled, appressed-hirsute, the apex setose-apiculate- Capitulum solitary, diameter up to 3 cm. in luxuriant specimens, often less. Imvolucre inflated, broadly obconic, glabrous, glaucous, the segments 16—25 in two rows, broadly deltoid with a scarious margin and sometimes a blunt, elongated, scarious apex, many-veined, glabrous except for a few appressed hairs near the tip. Marginal florets few, without corolla, but with a bilobed style and producing an achene. Ligulate florets c. 25, yellow, the tube very short, the lamina c. 5 mm. long, 2 mm. wide, the small inner lobes recurved. Disc florets very numerous, the corolla 2mm. long, tubular below, campanulate above, 4-lobed, yellow. Marginal achenes compressed, scarious-margined below, obovate, emarginate, glabrous, 2 mm. long, 1.5 mm. wide. A frequent plant in the western sandveld. It is very variable in size according to soil and season, and it can be a showy and conspicuous plant with bright yellow capitula up to the size of a florin. It is, however, less handsome than the other large-flowered species, C. Duckitteae L. Bolus, from the Darling district (e.g. at Bok Bay, Compton 6780, Kies 154): this has more numerous and much longer ligulate florets which are of a deep orange colour; the involucral bracts are rather narrower and have longer appendages, and the whole plant, including the involucral bracts, is less hirsute. Gnaphalium alticolum Compton sp. nov. Compositae—Senecionideae—Gnaphalieae. Herba perennis, caulibus decumbentibus, foliis omnibus utrinque lanatis, basalibus obovato-spathulatis, obtusis, caulinis angustioribus subacutis, capitulis numerosis in capitibus hemisphericis dense glomeratis, bracteis membranaceis, erectis, obtusis, stramineis, floribus ¢. 55, quorum dimidiis filiformis, femineis. Habitat. Basutoland: Phutha, near Mokhotlong, c. 10,000 ft. alt., 26 Feb. 1949, Compton 21570. (Type, in Herb. Nat. Bot. Gdns.): Machaba Peak, 21 March 1936, Galpin s.n. Griqualand East: Mt. Fletcher District, summit of Satsanna’s Peak, 9,550 ft. alt., 17 March 1904, Galpin 6670. A small perennial plant with a woody rootstock from which arise numerous herbaceous stems, prostrate below, becoming suberect below the inflorescence, loosely lanate. Leaves soft, entire, loosely lanate on both sides, the basal ones obovate-spathulate, obtuse, c. 3 cm. long, | cm. wide, the cauline leaves gradually diminishing in size and changing in shape to those below the inflorescence which are lanceolate, subacute and somewhat undulate. Capitula numerous, subsessile and aggregated 114 The Journal of South African Botany. in a subsessile dense terminal rounded head up to 2 cm. in diameter, matted together at their base by long white woolly hairs arising from the short peduncles and the bases of the outermost bracts, c. 4 mm. long, 3 mm. diam. Involucral bracts all erect, membranous, straw-coloured, the outer ones ovate, subacute, the others subspathulate, obtuse, slightly longer than the florets. Florets c. 55, of which rather more than half (the outermost) are filiform females, the others hermaphrodite. All florets have a white pappus of about 10 very fine caducous hairs, barbellate towards the ends, c. 3 mm. long. A species on the borderline between Gnaphalium and Helichrysum, only separated from the latter genus (with which it has, perhaps, more real affinity) by the large number of peripheral filiform female florets, which are in several rows and are, in fact, more numerous than the central perfect florets. Macowania sororis Compton sp. nov. Compositae—Inuloideae. Species M. pulvinari affinis, sed foliis validioribus, obtusioribus, capitulis et eorum partibus majoribus, et aliter differt. Habitat. Cape Province. Griqualand East: Mount Currie, 7000 ft. alt., among rocks, April 1938, Sister Mildred 8. (Type, in Herb. Nat. Bot. Gdns.) Basutoland: Machaba Peak, highest point, 21 March 1936, Galpin s.n. 714088. Natal: Mont-aux-Sources, 9500 ft. alt., Jan. 1894, Flanagan 1879. A dense cushion-like shrub up to c. 25 cm. high, with numerous forking and diverging densely leafy branches ending in solitary capitula. Stems and leaf-bases loosely lanate when young. Leaves linear, suberect when young, patent and somewhat reflexed when mature, obtuse or bluntly apiculate, rigid, densely glandular-hispid on the upper surface, the mar- gins thick and reflexed almost covering the lanate lower surface, with a narrow median groove above, 1.2—1.5 cm. long, 2 mm. wide. Capitula solitary, terminal, pedunculate, erect, exserted just beyond the leaves. Peduncle ec. 8 mm. long, terete, bractless, lanate. Involucre c. 1 em. long, 8 mm. diam., the scales rigid, light coloured, scarious, glandular and lanate, multiseriate, gradated, the longest lanceolate, slightly carinate, acute, c. 7 mm. long, 1.5 mm. wide. Ray florets c. 16, female, ligulate, yellow, ovary c. 2 mm. long, terete, pubescent, tube 4 mm. long, puberulous, lamina c. 7 mm. long, 2 mm. wide, apex shortly 3-lobed, style-branches slender, c. 1.3 mm. long, pappus bristles c. 18, stiff, straight, uniformly barbellate, c. 6 mm. long, a little longer than the corolla tube. Disc florets c. 40, hermaphrodite, ovary as in ray florets, corolla tubular, yellow, c.6 mm. long, the upper half wider than the puberulous lower half, anthers Plantae Novae Africanae. 115 <a Sill (> iy aay QS pet a 14 10 Fic. 2. Macowania sororis Compton. 1. Portion of plant, nat. size. 2, 3. Leaves, upper and lower surfaces, nat. size. 4. Leaf, transverse section x 4. 5, 6. Inner and outer involucral scales « 2. 7. Capitulum, longitudinal section x2. 8. Ray floret x 8. 9. Dise floret « 8. 10. Corolla of dise floret spread out (distal portion only) x 8. 11. Stamen x 8. 12, 13. Part of style and branches from ray and dise florets x8. 14. Part of pappus bristle x16. Sister Mildred 8. Del. M. Walgate. with slender basal awns appressed to filament, pappus as in ray florets, about equalling the corolla tube. As compared with M. pulvinaris N. E. Brown (type Galpin 2258!) which it most resembles, this is a more robust plant with stouter and more obtuse spreading leaves and larger capitula whose parts are also larger and more numerous. (For descriptions and drawings of previously recog- nised species of Macowania—except M. (Homochaete) conferta (Benth.) Phillips—see C. A. Smith in Bothalia II, p. 347, 1927.) Othonna Barkerae Compton sp. nov. Frutex sublignosus, omnino glaber. Folia alterna, herbacea, parum incrassata, oblanceolata, base gradatim longe attenuata, irregulariter 116 The Journal of South African Botany. dentata, acuta, margine coriacea. Inflorescentiae terminales vel subtermi- nales, laxe corymbosae, pedunculis elongatis, vix folia excedentes. Involucrum campanulatum, profunde 8-lobatum, herbaceum, marginibus anguste scariosis. Flosculi radii ligulati, feminei, fertiles, laminis elongatis, flavis; disci masculini, tubulares, ovariis glabris. Achaenia dense appresse sericeo-hirsuta. Pappi capilli numerosi, rigidi, albi. Habitat. Cape Province. Ladismith Division: Seven Weeks Poort, 6 August 1949, Barker 5453. (Type, in Herb. Nat. Bot. Gdns.) Marloth 2884: Wurts 1108, 22 Aug. 1953, 2500 ft., frequent. An erect soft-wooded shrub c. 100 em. high, glabrous throughout (except the achenes), slightly glaucous. Stems of current year’s growth up to 5 mm. or more in diameter, slightly ridged by the decurrent leaf veins and margins: older stems with prominent leaf scars and wrinkled bark. Leaves alternate, clustered towards the ends of the branches, erecto-patent, oblanceolate, tapering gradually below to a narrowly winged base, the apex acute, the distal part of the lamina with a few irregular acute teeth, the proximal part usually entire or sometimes denticulate, the margin coriaceous: texture herbaceous, somewhat fleshy: lateral veins immersed, becoming more noticeable in the dry state: overall length ce. 8—12 cm., width 1.5—2.5 em. Inflorescences loose few-headed corymbs, terminal or subterminal, slightly exceeding the leaves, Peduncles pale: bracts small, linear. Capitula borne singly on elongated pedicels. Involucre campanulate, of about 8 lanceolate bracts which are uniseriate, separate almost to the base, herbaceous with narrow scarious margins, c. 7 mm. long. Ray florets c. 8, ligulate, yellow, the tubular portion of the corolla 5 mm. long, the ligular portion c. 2 em. long, 4 mm. wide, 5-veined, the ovary turbinate, hairy, 2 mm. long. Disc florets numerous, male, the corolla slender, tubular, 4 mm. long, the lobes small, yellow, the ovary 3 mm. long, glabrous. Achenes c. 5 mm. long, densely appressed-silky- hairy. Pappus of many stiff white minutely barbellate bristles, c. 7 mm. long. This new Othonna shows some affinity, judging from the rather inade- quate descriptions by De Candolle and Harvey, to O. osteospermoides DC., collected by Drége in the Zwanepoelpoortsberg (that is, near Miller) and, fide De Candolle, in the Zwart Ruggens (that is, near Jansenville) though Drége does not mention this latter locality in his index. Material of this species was, however, not available for comparison. Its locality lies 150 miles in a direct line from Seven Weeks Poort. Another, more remote, affinity is with O. quinquedentata of the Cape Peninsula ete. A collection recently received from Rev. T. Wurts, cited above, is apparently a luxuriant form with broader leaves than usual, more numer- Plantae Novae Africanae. 117 EX VR z Ww Fie. 3. Othonna Barkerae Compton. 1. Branch, nat. size. 2. Capitulum x 2. 3. Ray floret x 4. 4. Dise floret x 4. 5. Corolla of dise floret laid out x 4. 6. Three stamens x 4. 7. Style of disc floret x 4. 8. Achene x 4. 9. Achene after soaking in hot water (pappus shortened) x 4. Barker 5453. Del. W. F. Barker. 118 The Journal of South African Botany. ous capitula, thicker involucral bracts and wider but slightly shorter ray florets. What may be starved forms of the species have been collected near the summit of the Zwartberg Pass, c. 5000 ft. alt.: Dyer 92 in Herb. Albany Mus., Pocock S. 197 in Herb Bolus. It is strange that this rather conspicuous plant has not been collected more often, as according to Mr. Wurts it is frequent in Seven Weeks Poort: perhaps its superficial resemblance to the widely spread Chrysanthemoides moniliformis Norlindh may account for this neglect. Othonna spektakelensis Compton sp. nov. Species O. retrorsi affinis, sed caulibus minus lanatis, foliorum denti- culis paucioribus, basibus angustioribus, laminis latioribus, involucrum bracteis paucioribus, plerumque 5, radii flosculis paucioribus, distincta. Habitat. Cape Province. Namaqualand: Near Spektakel, 3500—3800 ft. alt., Sept. 1883, Bolus 9596. Spektakel, 25 Aug. 1941, Compton 11466, Esterhuysen 5732. Eselsfontein (between Springbok and Spektakel), Aug. 1949, Hall, Nat. Bot. Gdns. 771/49; 8 Sept. 1950, Hall 141. (Type, in Herb. Nat. Bot. Gdns.) Spektakel Hill, 4 Sept. 1951, Maguire 1010. Plant perennial with a woody obconical rootstock tapering quickly to a long tap-root and bearing a few densely leafy abbreviated aerial shoots. Leaves varying much in size up to c. 6 em. long, 1.5 em. wide, suberect, oblanceolate, tapering to the base, of thin texture, glabrous, rather glaucous, the apex obtuse or subacute, the margins denticulate, the teeth varying in size, callous, about 8 to the centimetre, venation prominent on both sides when dry, reticulate, the veins joining to 5 parallel veins in the thicker sub-petiolar base which eventually splits into persistent bristles: a small amount of woolly packing is present between the leaf bases. Inflorescences axillary, several from each crown, up to 35 cm. high, often less, almost bractless, the slender peduncle forking a few times high above the base and bearing the small capitula on slender pedicels in a loose panicle, glabrous throughout. Involucre obconic, bracts 5, rarely 6, obovate-lan- ceolate, free nearly to the base, glabrous, glaucous, narrowly membrane- edged, 5 mm. long. Ray florets 5—7, usually 5, rarely 8, ligulate, female, fertile, corolla tube 1.5 mm. long, lamina 7 mm. long, 3 mm. wide, usually 5-veined, yellow: style branches linear: ovary pubescent on the ridges. Disc florets 15—30, usually about 20, male: corolla tubular, obconic, yellow, c. 3 mm. long: ovary slender, glabrous c. 1.5 mm. long. Achenes densely hairy, 1.5 mm. long: pappus white, minutely barbellate, 3 mm. long. Allied to O. retrorsa DC., but differing in its general aspect and in a number of more or less quantitative details. It is less robust, the leaves Plantae Novae Africanae. 119 are fewer, narrower at the base and broader in the lamina, with fewer and coarser denticles, and their bases are more inclined to break up into fibrous tufts and are less copiously packed with wool. The peduncles and pedicels are more slender. The involucral bracts are normally fewer (usually 5), and the ray florets are also almost always less than 8. The area of distribution is also somewhat different, lying between Springbok and Spektakel, whereas O. retrorsa occurs mainly in and to the south of the Khamiesberg. The first collection of O. spektakelensis was by Harry Bolus in 1883: his specimens in the Bolus Herbarium (Bolus 9596) bears a label “‘O. retrorsa” in N. E. Brown’s handwriting. De Candolle’s description of O. retrorsa, founded on Drége’s Khamiesberg collections, indicates a different plant, however, from which, in my opinion, the Spektakel plant should be distinguished. Other gatherings of true O. retrorsa are Compton 17304 (10 miles south of Khamieskroon), Lewis in Herb. S.A. Mus. 57547 (5 miles south of Khamieskroon), Leighton 1244 (quartzite koppie north of Darter’s Grave), Schlechter 11193 (between Garies and Springbok), Hutchinson 840a (Khamieskroon). Another species related to O. retrorsa is O. Zeyher (Sond.) Harv., (Fl. Cap. III, p. 339), founded on imperfect specimens, Zeyher 996, from Kammapus. (Zeyher’s journey was “von Lieslap uber Kammapus bis Hofkraal an Hartebeest Rivier’’: recent maps show “Kamaboos” or “Gannabos”’ in this area which lies to the south-east of the Khamiesberg near Bokkraal). This is probably identical with a plant collected by Hall (Nat. Bot. Gdns. 462/52) on granite boulders at Khamieskroon, and with other plants from the Khamiesberg (e.g. Pearson, Sladen Expedition 5808) placed as O. retrorsa var. linearifolia Harv. It is a cushion-like plant often many inches in diameter, occurring in rock crevices—a distinct habitat from either true O. retrorsa or O. spektakelensis. EREMIELLA Compton gen. nov. Ericaceae—Ericoideae. Genus adhuc monotypicum. Folia 4-nata. Flores 4-nati, terminales, sessiles. Bracteae, sepala, corollaque hirsutae. Bracteae 3, scariosae. Sepala 4, valde discreta, scariosa. Corolla cyathiformis, 3-lobata, hirsuta, lobis obtusis. Stamina 6; antherae inclusae, basifixae, cellulis base discretis, muticae. Ovariwm 3-carpella- tum, carpellis uniovulatis, ovulis pendentibus. Stylus filiformis, stigmate capitellato, exserto. Leaves 4-nate. Flowers 4-nate, terminal, sessile. Bracts, sepals and corolla hirsute. Bracts 3, scarious. Sepals 4, separate to the base, scarious. 120 The Journal of South African Botany. Corolla cyathiform, 3-lobed, hirsute, the lobes obtuse. Stamens 6, the anthers included, basifixed, muticous, the cells separate to the base. Ovary of 3 carpels which are uniovulate, the ovules pendent. Style filiform, stigma capitellate, exserted. Eremiella outeniquae Compton sp. nov. Fruticulus ramosus. Caules hirsuti. Folia patentia, supra glabra, infra hirsuta. Aliter ut in generis descriptione. Habitat. Cape Province. Mossel Bay Division: Ruyterskop, 4500 ft. alt., 26 Nov. 1949. Compton 21818. (Type, in Herb. Nat. Bot. Gdns.) A small erect much branched shrub, a few inches to a foot or more high. Stems roughly hirsute with hairs of different lengths. Leaves 4-nate, petioles appressed, laminas patent, limear-oblong, obtuse, sulcate, almost glabrous above, hirsute with simple and gland-tipped white hairs on margin and under surface, glabrescent with age, 1.5—2 mm. long. Flowers sessile in numerous small terminal clusters, 4-nate, erect or nodding, ec. 4 mm. diam. Bracts and sepals closely appressed to one another and to the corolla. Bracts 3, oblong-lanceolate, the lowest a little broader than the others, 1.5 mm. long, scarious, tinted on margins and under surface, darker, obscurely keeled and with a few long hairs at the apex. Sepals 4, free to the base, linear-oblong, otherwise like the bracts. Corolla cyathi- form, 3-lobed to about the middle, 2 mm. long and in diameter, dull pink, hirsute, the lobes obtuse and rounded. Stamens 6, filaments filiform, ] mm. long, slightly thicker towards the top, bearing two quite separate anther- cells 0.6 mm. long, which are basifixed, glabrous, obovate, obtuse, muti- cous and dark coloured, and open by oval pores about half their length. Ovary globose, dark coloured, on a dark inconspicuous disc, glabrous or minutely pubescent on the upper half, 3-carpellary, each carpel containing one apically attached flattened ovule. Style slender, 2 mm. long. Stigma capitellate, exserted. Mature frwit with a coriaceous wall, globose, 1.5 mm. diam. The addition of a new genus to the existing list of Ericoideae is a matter for some hesitancy, but there can be little doubt that this very distinct plant merits generic rank as much as other usually accepted genera. Though it has so far only been collected on one occasion, several plants were then obtained: these were uniform among themselves and showed no signs of being abnormal or a chance hybrid. The remarkable circumstance that with a 4-sepalate calyx, the corolla should be 3-lobed, the androecium of 6 stamens and the ovary of 3 uniovulate carpels, is sufficient to distinguish it from other Ericoid genera: and the cyathiform corolla, included anthers and other features are such as to separate it from most of them. Plantae Novae Africanae. 12] 18 2. 2. Flower from above x 20. Fie. 4. Hremiella outeniquae Compton. 1. Branch x 3. Flower, side view x 20. 4. Flower from below x 20. 5. Corolla x 20. 6, 7, 8. Bracts x 20. 9, 10, 11, 12. Calyx segments x 20. 13, 14. Stamens, side and front view x 20. 15. Gynaecium x 20. 16. Ovule x 20. 17. Ripe capsule x 20. 18. Seed x 20. 19. Leaf from below x 20. 20. Leaf, side view x 20. Compton 21818. Del. W. F. Barker. 122 The Journal of South African Botany. If we accept the reduction of all the South African Ericoideae to seven genera, as proposed by Dr. E. P. Phillips (Journ. 8. Af. Bot. X, p. 69,1944; Genera of S. Af. Flowering Plants ed. II, p. 558, 1951) this would fall into his enlarged and very aggregate genus Eremia. In my opinion, however, it is doubtful whether such a wholesale reduction of genera is justified, or whether it does not increase the difficulties of analysing this complex group: it might well be argued that, on the other hand, the great genus Erica deserves generic subdivision to bring it more into line with its allies. The only reason for retaining it as a single genus is that its floral formula is practically uniform throughout, and this is the reason why, I think, the “minor Ericoideae” are best divided into a rather large number of genera. While Eremiella is distinct enough from all other genera, its nearest affinities are with Eremia as described by N. E. Brown in the Flora Capensis, IV. i, 1905/9. A comparison may be made with the little-known Eremia brevifolia Benth. (DC. Prodr. VII, p. 700; Fl. Cap. IV. i, p. 334), a plant of similar general appearance, only known from the original gather- ing, Masson 57, from Attaqua’s Kloof—a locality not far removed from Ruyterskop. From this plant, however, Hremiella outeniquae differs in its trimerous corolla, androecium and gynaecium, its hairy corolla, glabrous and less manifest anthers, wider calyx lobes, coarsely hairy leaves and other minor features. Acrostemon Barkerae Compton sp. nov. Ericaceae—Ericoideae. Fruticulus erectus, ramosus. Caules puberuli. Folia 3-nata, erecto- patentia, juventute linearia, albo-pilosa, demum ovata, glabra. Flores in capitulis terminalibus aggregati. Pedicelli, bracteae et calyx longe albo- pilosi. Bracteae 3, approximatae. Calyx profunde lobatus. Corolla ovoidea, glabra, ore angusto. Antherae terminales, exsertae, angustae, muticae, medio divisae, fuscae. Ovarium globosum, 4-angulatum, glabrum, 4-carpellatum, carpellis uniovulatis. Stylus longe exsertus, stigmate parvo. Habitat. Cape Province. Worcester Division: Scherpenheuvel, hillsides, 29 Sept. 1951. Barker 7523. (Type, in Herb Nat. Bot. Gdns.) Clayish foothills on east side of Stettynsberg near Villiersdorp, Hsterhuysen 15584. (A. Zeyher s.n. plant named A. equisetordes Kl. by Reichenbach f., from Worcester, of which there are a few fragments in the Bolus Herbarium, may be the same.) A densely branched erect shrublet. Young stems slender, puberulous. Leaves 3-nate, erecto-patent, petiole 0.5 mm. long, lamina 2—3 mm. long, linear when young becoming ovoid, obtuse, sulcate, at first white- Plantae Novae Africanae. 123 yY VF 8S Ws Wi <=. Fic. 5. Acrostemon Barkerae Compton. 1. Branch, nat. size. 2. Flower x 10. 3. Corolla « 10. 4,5, 6. Bracts x 10. 7. Calyx segments « 10. 8. Three calyx segments, inner view * 10. 9. Stamen, front view 10. Stamen, side view 10. 11. Gynaecium = 10. 12. Ovule x 10. 13. Young leaf x 10. 14. Old leaf x 10. Barker 7523. Del. W. F. Barker. pilose especially towards the tip, soon becoming glabrous, the margins with minute sessile glands when young. Flowers numerous in terminal subglobose clusters of about 8. Pedicel, bracts and calyx covered with long white spreading simple hairs. Pedicel ec. 1.2 mm. long. Bracts approximate, median bract lanceolate, c. 1.2 mm. long, lateral bracts linear, c. 1 mm. long. Calyx divided to near its base, segments lanceolate, c. 1.5 mm. long, 0.5 mm. wide. Corolla c. 4 mm. long, ¢. 2 mm, diam., pink, glabrous, ovoid, the mouth very narrow, the lobes very small. Stamens 4, filaments slender, anthers long-exserted, terminal, muticous, dark brown, narrow, divided about half way down, the lobes parallel, c. 2 mm. long, 0.5 mm. wide. Ovary globose, 4-angled, glabrous, c. 1 mm. long and wide, 4-locular, each cell containing one slender pen- dulous ovule. Style filiform, ce. 7 mm. long, far-exserted, the stigma minute. 124 The Journal of South African Botany. An attractive species with apparently a very restricted distribution. Its nearest affinity is probably with A. equisetoides Klotzsch from Tulbagh Waterfall. I name it after Miss W. F. Barker, whose excellent material in the Kirstenbosch Herbarium establishes its distinctness, and to whom I owe many admirable drawings of Ericaceae and other plants described in the volumes of the Journal of South African Botany. Erica monantha Compton sp. nov. § Ceramia. Fruticulus diffusus vel suberectus, internodiis elongatis. Folia 3-nata, patentia, mucronata, juventute hirsuta, glandulifera, demum glabra. Flores axillares, solitarii. Bracteae basales. Pedicelli angusti, elongati. Sepala acuta, longe mucronata, marginibus reflexis. Corolla cyathiformis, subquadrangulata, puberula, segmentibus erectis, obtusis vel parum emarginatis. Filamenta angusta, glabra. Antherae inclusae, oblongae, muticae, glabrae. Ovarium depresso-globosum, hirsutum, lobatum, stylo recto vel curvato, stigmate subcapitato, exserto. Habitat. Cape Province. Ceres Division: Wabooms River (Cold Bokkeveld) 3000 ft. alt., 25 Sept. 1936. Compton 6641. (Type, in Herb Nat. Bot. Gdns.). Clanwilliam Division: Elands Kloof, 25 March 1951. Compton 22690. Fic. 6. Erica monantha Compton. 1. Portion of plant, nat. size. 2. Fragment of stem and leaves x 5. 3. Flowers x 5. 4. Flower cut lengthwise x 5. 5. Sepal x 10. 6. Stamens x 10. 7. Ovary x 10. 8. Transverse section of ovary x 5 Compton 6641. Del. M. Walzate. A small diffuse or suberect shrub, the internodes slender, elongated, glandular-pubescent. Leaves 3-nata, occasionally 4-nate, patent, oblong, sulcate, the lamina 2—3 mm. long, clothed with long soft white hairs and shorter glandular hairs and tipped with a long stiff mucro, finally glabrous. Flowers axillary, solitary. Bracts like the leaves, basal. Pedicels slender, sub-cernuous, thinly glandular-pubescent, 5—7 mm. long. Sepals ovate, acute, 1.2 mm. long, long-mucronate, glandular-hairy, the margins reflexed. Corolla cyathiform, subquadrangular, puberulous, Plantae Novae Africanae. 125 pale rose-coloured, 3 mm. long and broad; segments erect, obtuse or slightly emarginate, 1 mm. long, 2 mm. wide. Stamens included, filaments slender, glabrous: anthers muticous, oblong, 0.8 mm. long, the pore small. Ovary hirsute, depressed-globose, lobed, the style straight or slightly curved, the stigma subcapitate, shortly exserted. A distinct species of the § Ceramia, its nearest allies being EL. leptoclada van Heurck and Mull.-Arg., H. myriocodon Guthrie and Bolus, and E. cryptanthera Guthrie and Bolus. From £. leptoclada (including the var. aristata Bolus) it differs in its hirsute ovary, its long pedicel and in the shape of its corolla. From EF. myriocodon it differs in its muticous anthers, long pedicel, leaves mostly 3-nate, the shape of the leaves and their long mucro. From E. cryptanthera it differs in its muticous anthers, hirsute ovary, sulcate leaves, rosy corolla, the hairiness of its leaves, sepals, etc. Its locality is very different from that of its nearest allies. The following specimens may also belong to this species, though they have open-backed leaves and sepals, a white corolla and slightly larger sepals and corolla. They come from sheltered positions, which may account for the differences from the type as described above. The bracts may be borne in varying positions on the pedicel—in some cases near the base, in others half way up, and occasionally near the top. Esterhuysen 14719: Ceres Division: Kloof 8S. W. side of Schurfteberg, on sheltered rather dry earthy cliff face, side of kloof, 7 Dec. 1948. Hster- huysen 15202: Ceres Division: Mitchell’s Pass, crevices in overhanging rock where moisture seeps through, afternoon sun only, 27 March 1949- Esterhuysen 16612: Piketberg Division: Rocky plateau on Twenty-four Rivers Mts., 3000—3800 ft. Above Porterville, on shaded sandy floor of overhanging rock-shelter. 16 Dec. 1949. Erica Porteri Compton sp. nov. § Pleurocallis. Species LH. Thomae affinis, sed bracteis sepalisque brevioribus, corolla angustiore bicolorata, antheris brevioribus muticis inclusis, foliis breviori- bus patentibus, et aliter distincta. Habitat. Cape Province. Caledon Division: Buffels Kloof, near Pringle Bay, 500 ft. alt., 25 March, 16 June, 17 August 1953. H. Porter s.n. (Type, in Herb. Nat. Bot. Gdns.) An erect rigid rather virgate shrub, about 50 cm. high. Stems grey- pubescent. Leaves numerous, 6-nate, erect at base, spreading horizontally and curving upwards at the tips, stiff, linear, sulcate, acute, glabrous, 7—12 mm. long, 1 mm. wide. Flowers axillary, borne densely along about 2—3 em. of the stem near its apex. Pedicels erecto-patent, c. 7—8 mm. long, puberulous. Bracts 3, appressed to the pedicel, one just below its 126 The Journal of South African Botany. A h = | 8 9 ! BAN x 71 i 12 13 Fie. 7. Erica Porteri Compton. 1. Branch, nat. size. 2. Flower « 2. 3. Calyx seg- ment x 2. 4. Upper bract x 2. 5. Lower bract x 2. 6. Gynaecium x 2. 7. Stamen, side view x 2. 8, 9, 10. Anthers, side, front and back views x 10. 11, 12. Leaves, lower and upper views x 4. 13. Leaf, side view x 4. H. Porter 8.n. in Herb. Nat. Bot. Gdns. Del. W. F. Barker. \\ ay ZPD ° GW \ SS Z middle, the others c. 2 mm. below the calyx, deltoid, acute, viscid, 1.5 mm. long. Sepals green, glabrous, viscid, erect, closely appressed to the corolla, linear-lanceolate, coriaceous, c. 5 mm. long, 1.2 mm. broad. Corolla tubular, somewhat viscid, the basal 2/3 bright rose-red shading to white in the distal 1/3, slightly curved downwards, glabrous but with a rugulose surface, c. 25 mm. long, gradually widening from 3 mm. diam. Plantae Novae Africanae. 127 at the base to 5 mm. diam. at the mouth, the lobes erect, orbicular, obtuse, 2mm. long. Filaments filiform, pink to white. Anthers included, linear, smooth, muticous, dark brown, 2 mm. long, the pore 1 mm. long. Ovary c. 1.5 mm. long, 8-lobed, the apex intruse, glabrous. Style exserted, deflexed outside the corolla: stigma capitate. Related to EH. Thomae L. Bolus, but differing much in general appear- ance. The leaves are 6-nate, are only about half as long and spread stiffly from the stem. The flowers are smaller, both in length and diameter, and are strikingly bicoloured. The anthers are included, their tips being about 3 mm. below the apex of the corolla, and are less acute at the apex, truncate and muticous at the base. The calyx and bracts are only half as long, and the position of the bracts (judging from the drawings in Ann. Bolus Herb. IV, Pl. I.1, 1925) is different. This beautiful Heath was discovered by Mr. Harold Porter in abun- dance in one small locality, two or three acres in extent, to which he con- ducted me. The plants were quite uniform in appearance and showed no ordinary indication of hybridity. Towards the periphery of the patch it thinned out to scattered plants, the impression given being that of a “distribution circle” of a recently evolved species. I have pleasure in naming the species in honour of Mr. H. Porter, an Erica enthusiast, who is a member of the Council of the Botanical Society of S. Africa and who, as chairman of the Hangklip Beach Estates, is making every effort to conserve the exceptionally rich flora of the area controlled by his syndicate. Erica subimbricata Compton sp. nov. § Geissostegia. Species H. Sonderianae affinis, sed pedicellis longioribus, bracteis remotioribus, non viscidis, sepalis corollaque majoribus, non viscidis, antheris basifixis longioribus, et aliter distincta. Habitat. Cape Province. Ceres Division: Elands Kloof Pass, 2500 ft. alt., sandy ground, 23 Sept. 1936, Compton 6650 (Type, in Herb. Nat. Bot. Gdns.) Piquetberg Division: Piquetberg, Sept. 1937, Pillans 7728. A small erect shrub. Stems puberulous. Leaves 3-nate, imbricate, erect or slightly incurved, sulcate, oblong, obtuse, glaucous, ciliolate at the base when young, 4—5 mm. long. Flowers terminal, 3-nate, mostly on short erecto-patent lateral branches, horizontal or deflexed. Pedicels pubescent, 6—7 mm. long. Bracts white, rigid, 2 mm. long, keel-tipped, the lowest c. 2 mm. below the flower, the other two 1 mm. below the flower and clasping its base. Calyx white, rigid, 3 mm. long, the sepals broadly ovate, shortly keel-tipped, glabrous, minutely denticulate, the base coriaceous, the midrib somewhat wrinkled transversely. Corolla white, tubular, slightly wider in the middle, c. 3.5 mm. long including the 128 The Journal of South African Botany. 14 3 8 9 Fie. 8. Erica subimbricata Compton. 1. Branch, nat. size. 2. Flower x 5. 3. Corolla x 8. 4.Sepal x 8. 5, 6, 7. Anthers, side, front and back views x 10. 8. Stamen, side view x 5. 9, Gynaecium x 8. 10, 11,12. Bracts x 8. 13. Whorl of leaves x 5. 14. Leaf from below x 5. Compton 6650. Del. E. Wasserfall. obes which are broadly orbicular, erect, obtuse, minutely denticulate, overlapping one another at the edges, 2 mm. diam., fading brown. Fila- ments slender, straight, c. 4.5 mm. long, passing smoothly into the anthers which are well-exserted, dark brown, linear, subacute, muticous, glabrous, c. 2 mm. long, the pore 1.5 mm. long. Ovary glabrous, c. 1.5 mm. long, including a ribbed obtuse beak. Style 5 mm. long, straight, stigma clavate, projecting slightly beyond the anthers. An affinity of #. Sonderiana Guthrie and Bolus, but differing suffici- ently to warrant separation as a species. The pedicels are about three times as long, the bracts are further below the flower and are not viscid, the sepals and corolla are rather longer and are not viscid, the anthers are twice as long, slender and borne continuously with the filaments. Phillips from Sneeuwgat, 4—4500 ft. alt., Nov. 1916, in Herb. Kew., may be the same. Plantae Novae Africanae. 12) Erica syngenesia Compton sp. nov. § Pachysa. Species H. spectabili affinis, sed floribus grandioribus, filamentis latiori- bus, antheris inter se adhaerentibus, et aliter differt. Habitat. Cape Province. Ladismith Division: Seven Weeks Poort 2500 ft. alt., 8 July 1953, 12 Aug. 1953, Wurts 1016. (Type, in Herb. Nat. Bot. Gdns.) A stout erect branching shrub. Stems puberulous with minute reflexed hairs. Leaves 3-nate, erecto-patent or slightly inflexed, imbricate, linear- oblong, sulcate, very minutely puberulous with reflexed hairs, c. 5 mm. long, 1 mm. wide. Flowers terminal, 3-nate, erect or nodding, the pedicels, bracts, calyx and corolla white. Pedicels c. 5 mm. long, minutely puberu- lous and with a few sessile glands. First bract median, scarious, sulcate- LY Fie. 9. Hrica syngenesia Compton. 1. Branch, nat. size. 2. Flower x 5. 3. Bract x 5. 4,5. Sepals x 5. 6. Androecium and gynaecium x 5. 7. Three stamens, front view x 5. 8. Stamen, side view x 5. 9. Stamen, front view x 5. 10. Stamen, back view x 5. 11. Gynaecium x 5. 12. Leaf, side view x 5. 13. Leaf, lower view < 5. Wurts 1016. Del. W. F. Barker. 130 The Journal of South African Botany. keeled, green-tipped; other bracts a little higher on the pedicel, sulcate- keeled: all c. 3 mm. long, 2 mm. wide. Sepals broadly ovate narrowing to a sulcate-keeled obtuse apex, c. 3 mm. long, 2 mm. wide, minutely puberu- lous, ciliolate towards the base and with a few marginal sessile glands. Corolla broadly obovoid-urceolate, somewhat contracted below the mouth, the tube c. 5 mm. long and in diameter, distinctly veined, the lobes erect, obtuse, c. 1 mm. long. Filaments white, ribbon-shaped with a distinct mid-vein, 1 mm. wide in the middle, narrower above and below, sharply bent down and up again below the anther. Anthers dark-coloured, of two distinct cells, c. 1.5 mm. long, minutely hispidulous, the pore almost as long as the cell, aristate, the awns adnate to the filament, their subulate tips diverging widely: each anther-cell when young adheres firmly to the adjoining cell of the next anther, thus forming a united anther-ring around the style: later the anthers break apart to reveal their pores and shed their pollen. Ovary inserted on a dark 8-lobed disc, globular, glabrous, 4-lobed with a depressed apex. Style c. 2.5 mm. long: stigma included, conical. E. syngenesia is related to EL. spectabilis Klotzsch, a species which occurs on the coastal limestone hills of the Riversdale and Bredasdorp Divisions— a very different and rather remote habitat. The present species is a taller but rather less floriferous shrub, the individual flowers being rather larger in all parts. Somewhat similar plants occur in other inland localities, e.g. on the Witteberg, Laingsburg Division. The most noteworthy feature of EH. syngenesia is the way in which the rather large hispidulous anthers adhere to one another, especially in the young state, so that the cells of adjacent anthers appear more closely connected than the two separate cells of the same anther: the filaments are much broader, especially in the middle, than in HE. spectabilis, the anther itself is larger, and there are other minor differences. Erica tenuipedicellata Compton sp. nov. § Chlorocodon. E. coarctata affinis, sed pedicellis multo longioribus, patentibus, corolla alba, foliis angustioribus, ceterisque differt. Habitat. Cape Province. Caledon Division: Flats near Hawston, 23 Aug. 1946, Compton 18170. (Type, in Herb. Nat. Bot. Gdns.) A small dense virgately branched shrub. Stems slender, erect, flexuous, minutely puberulous. Leaves 3-nate, erect, imbricate, linear, straight or slightly incurved, acute, glabrous, 5—6 mm. long. Flowers numerous, singly or in pairs at the end of very short axillary shoots, forming a long pseudo-raceme. Pedicels filiform, erecto-patent or spreading widely from the main stem, glabrous, 5 mm. long: one bract basal, the others about Plantae Novae Africanae. 131 1 mm. above the base, clasping the pedicel, c. 0.5 mm. long. Flowers borne far beyond the leaves, mostly nodding when mature. Sepals scarious, keeled, c. 2 mm. long, about equalling the corolla. Corolla cyathiform, glabrous, white, c. 2 mm. long, the segments nearly as long as the tube, obtuse, very minutely denticulate. Filaments slender, c. 1 mm. long. Anther cells distinct, muticous, acute at the apex, c. 1 mm. long, the pore c. 0.6 mm. long, manifest or slightly exserted, the pollen dust-like. Ovary glabrous, c. 0.5 mm. long. Style c. 1.5 mm. long, often somewhat deflexed, the stigma exserted, obconic-peltate. A distinct little Heath, related to H. coarctata Wendl., but of quite different aspect, mainly on account of its much longer spreading pedicels, the narrower, straighter and more acute leaves and the white corolla. So far only collected on one occasion. Erica zebrensis Compton sp. nov. § Polycodon. Frutex erectus, ramosus. Folva 3-nata, erecta, linearia, acuta, glabra. Flores terminales, 3-nata, subcernui, omnino colorata. Pedicelli elongati. Bracteae 3, quarum 2 approximatae, 1 parum remotiora. Sepala ovata, concava. Corolla late cyathiformis, glabra. Antherae manifestae, obtusae, base breviter apiculatae. Ovarium globosum, minute puberulum. Stylus exsertus, stigmate capitulato. Habitat. Cape Province. George Division: Zebra, north of Outeniqua Pass, 2000 ft. alt., 3 Dec. 1951. Compton 23087. (Type, in Herb. Nat. Bot. Gdns.) An erect much-branched shrub. Stems slender, puberulous with re- flexed simple hairs. Leaves 3-nate, erect, glabrous, sulcate, acute, equal- ling the internodes or somewhat imbricate, c. 3 mm. long. Flowers 3-nate, numerous, terminating slender erecto-patent lateral branches, more or less cernuous. Pedicels, bracts, sepals and corolla magenta-coloured, Pedicels 3 mm. long, thinly puberulous. Bracts lanceolate, two close beneath the calyx, the other about 1 mm. below, 1 mm. long. Sepals scarious, ovate, concave, somewhat keeled, 1 mm. long. Corolla broadly cyathiform, glabrous, 2 mm. long, the tube equalling the broad obtuse lobes, c. 3 mm. diam. at the mouth. Filaments and anthers of about equal length; anthers manifest, dark brown, 0.6 mm. long, oval, obtuse at apex with two short basal points, the pore c. 0.3 mm. long. Ovary globose, flattened and minutely puberulous at the top, dark coloured. Style slender, 2.5 mm. long, exserted; stigma capitulate. An attractive species on account of its graceful form and the brightly coloured perianth contrasting with the dark manifest anthers. Perhaps related to H. nemorosa Klotzsch, H. floribunda Lodd. and EL. Pearsoniana 132 The Journal of South African Botany. L. Bolus, but distinct on account of the long pedicels, the simple not floccose hairs, the well-spaced flowers, approximate bracts, broader- shaped corolla and manifest anthers. Paranomus micranthus (Schltr) Compton comb. nov. (Nivenia micrantha Schlechter in Engl. Bot. Jahrb. XXVII, p. 107 1900.) While studying an unusually small and distinct-looking Paranomus from the Potteberg (on the boundary of the Swellendam and Bredasdorp Divisions) with a view to describing it as a new species, I found that a plant from the Elim Hills (Bredasdorp Division) had been described by its collector, Schlechter, under the name Nivenia micrantha, and that this had been sunk by Phillips and Hutchinson (in Fl. Cap. V.i. p. 716) in Nivenia Lagopus R. Br., a species whose area of distribution, viz. Tul- bagh and further north, is remote from Elim. Nivenia micrantha was recognised by Schlechter as being related to N. Lagopus, but he pointed out that the small flowers and conspicuously dark stigmas are good dis- tinguishing characters and that the perianth segments are rosy-red inside. The Potteberg plant is, I consider, conspecific with the Elim plant, though its inflorescences are often even shorter and practically spherical, and it remains to place it under Paranomus, a change not made by N. E. Brown in his article on the Nivenias. (Trans. Roy. Soc. 8. Af. X XI, pp. 262, 264, 1934.) Bolus 7863 is another collection of the same plant from the Elim locality: in fact both Schlechter 9640 and Bolus 7863 are labelled “in collibus prope Elim.” Phillips and Hutchinson, however, (l.c. p. 717) cite the Bolus plant under NV. spicata R. Br:! In restoring Schlechter’s species to independent rank I cite the follow- ing specimens: Schlechter 9640 (type): in collibus prope Elim. Bolus 7863: in collibus prope Elim. Pillans 9417: hills north of Elandspad Farm (i.e. on the south side of the Potteberg) 15 Oct. 1940. Compton 19538: Albertsdal, north side of the Potteberg, 31 May 1947. Lachnea ruscifolia Compton sp. nov. Thymelaeaceae. Frutex erectus. Caules pubescentes. Folia alterna, ovata, integra, coriacea, supra plana, infra valde venosa. Flores axillares, sessiles, solitarii. Perianthii lobi utrinque tubusque albi, pubescentes; twbus angustus, ore dense pilosa; lobi patentes, lanceolati. Stamina 8, inaequales, filamentis longis, exsertis. Squamae 8, flavae, lineares. Stigma exsertum, penicilla- tum. Plantae Novae Africanae. 133 Fie. 10. Lachnea ruscifolia Compton. 1. Branch, nat. size. 2. Flower from above x 5. 3. Flower, side view x 5. 4. Perianth lobe and part of tube with scales and stamens attached x 5. 5. Seale x 10. 6. Long stamen x 10. 7. Short stamen xX 10. 8. Gynaecium x 5. 9. Young fruit x 5. 10. Young seed x 5. 11, 12. Bracteoles x 10. 13. Bud with bracteoles attached « 5. 14,15. Young leaves, lower and upper views x 5. 16,17. Old leaves, lower and upper views x 5. Wurts 1036. Del. W. F. Barker. Habitat. Cape Province. Ladismith Division: Elandsberg, Klein Zwartberg Range, 2000 ft. alt. July/August 1953, 7. Wurts 1036. (T'ype, in Herb. Nat. Bot. Gdns.): Paardeberg, at head of Huis River Pass, south side of ridge, 23 May 1950, Hsterhuysen 17108: South slope of Roodeberg, 24 May 1950, Hsterhuysen 17193. Erect, sparsely branched shrub, up to 60 cm. high. Stems terete, pubescent when young, dark coloured, with conspicuous leaf scars. 134 The Journal of South African Botany. Leaves alternate, spreading, coriaceous, glabrous, ovate to orbicular, entire, subacute, concave above, narrow-based, smooth above, minutely punctulate, with about 8 prominent ridges below which converge to base and apex, the margin indurated, pilose when young; c. 6—10 mm. long, 4—7 mm. wide. Flowers solitary, axillary, sessile. Bracteoles minute, scarious. Perianth white, the tube and both surfaces of the lobes densely pubescent: tube slender, c. 4 mm. long, slightly dilated at the base around the ovary, expanding abruptly just below the lobes: lobes 4, spreading, narrowly lanceolate, 5—6 mm. long, 1.5—2 mm. wide, marked with a few faint veins: throat with a dense tuft of white hairs. Stamens 8, all exserted, the filaments slender, the longest over 2 mm., the shortest c. 1 mm.: anthers yellow. Scales 8. alternating with the stamens and at the same level, linear, laterally flattened, 0.7 mm. long, yellow. Ovary glabrous, style lateral, stigma shortly exserted, capitate-penicillate. Fruit 2 mm. long, enclosed in the persistent base of the perianth tube. The flowers are those of a Lachnea, but their position singly in the leaf- axils is unusual in the genus, though it occurs in L. micrantha Schltr. and L. axillaris Meissn. From these species L. ruscifolia differs in its much larger and broader leaves and larger pubescent flowers. The leaves are distinctive but of the same general type as in UL. stricta Meissn. and L. elegans Compton, recalling a Ruscus or some Cliffortia sp. b | Prare XVIII. Dymondia Margaretae Compton. 1, Portion of plant, nat. size. 2,3, Capitula, nat. size. 4, Median involueral bract <x 3. 5, Ray floret x 4. 6, Dise floret x 4. 7, Stamen x 10. 8, Stigma of disc floret x 10. 9, Ovary x 10. 10, Achene x 10. Photographed by the University of Cape Town Photographic Department from coloured drawings by Miss M. Walgate, August 1938, by per- mission of the Curator of the Bolus Herbarium. A NEW SUCCULENT EUPHORBIA FROM THE WESTERN CAPE PROVINCE. By R. A. Dyer. (With Plate XIX.) Euphorbia hallii R. A. Dyer (Euphorbiaceae) Affinis EH. monteiroi Hook. habitu minore foliis pedunculorum angustioribus pedunculis ex axillis tuberculorum 1—5 gracilioribus differt. Planta succulenta perennis usque 40 cm. alta. Radix tuberosa sparse ramosa, radicibus tenuibus paucis instructa. Caulis non vel sparse ramosus, cylindricus, 3—4 cm. crassus, tuberculatus, ramis 2 em. crassis. Tubercula basi plus minusve 6-angulata, 1—2 cm. lata et longa. Folia tuberculis emittentia subrudimentaria, lineari-lanceolata, recurva, supra concava, usque | cm. longa, decidua. Pedunculi 1—5 ex axillis tubercu- lorum producti, plus minusve 4 cm. longi, bracteati; bracteae foliiformes, lineares, usque 10 cm. longae, 3—4 mm. latae; bracteolae infra cymam 2—3, lineari-lanceolatae. Cyma cyatheis bisexualibus 1—3. Involucrum usque | cm. diam. glabrum, glandulis 5 et lobis 5 late ovatis 4 mm. longis 3 mm. latis minute ciliatis; glanduli margine in lobos ramosos 3—5 producti. Ovarium glabrum gynophorio 5—7 mm. longo exsertum; styli 7—8 mm. longi, infra medium connati, apice bifidi; capsula plus minusve 7 mm. diam. obtuse trilobata, semina, ovata, angulata, 5 mm. longa, 3 mm. lata, rugosa. Habitat. Cape Province; Calvinia District, Botterkloof, H. Hall cult. in Nat Bot. Gdns. Kirstenbosch 903/52; May, H. Hall in National Herbarium, Pretoria 28532 (type); Dyer and van Breda 5434. A perennial succulent plant up to about 40 cm. tall with spreading thickened roots and only a few rootlets; main root 2—3 cm. thick and becoming thinner gradually, tough and fibrous with a soft pith. Stem simple or sparsely branched, thickened slightly from the base and becom- ing cylindric above, 3—4 em. thick, tubercled; branches 1—2 em. thick. Tubercles about 6-angled at base but not constant in shape, 1—2 cm. broad and long, uppermost part of margin variously indented and subtending old and young peduncles; young tubercles at the apex of stems producing linear-lanceolate, recurved, channelled leaves up to about 1 em. long, which are early deciduous and leave a central scar on the tubercle. Peduncles from axil of tubercles, 1—several together with the younger ones produced from below the older ones in successive seasons; the first peduncle from a tubercle often sterile, bearing leaf-like bracts, drying with age and per- 135 136 The Journal of South African Botany. sisting for some time but later disintegrating or being eaten off by animals leaving bare stems; Bracts leaf-like, linear, up to 10 em. long and 2—4 mm. broad, with sides folded up and recurved at tip, glaucous-green; bracts subtending the cyme shorter, 2—3, linear-lanceolate. Cymes with 1—3 bisexual cyathia; involucre up to 1 cm. diam., with 5 glands and 5 lobes: glands red, separate, about 0.3 mm. broad and with 3—5 branched processes 2—4 mm. long from the outer margin; lobes broadly ovate, 4 mm. long, 3 mm. broad, minutely ciliate. Stamens dark purple on exserted pedicels. Ovary small, on a gynophore 5—7 mm. long, exserted; ovules filling the ovary cells and attached at the apex within a small hood: styles dark red, united into a column 4.5 mm. long with branches about 3 mm. long and with small bifid tips; capsule about 7 mm. diam., obtusely trilo- bate, seeds ovate, angled, 5 mm. long, 3 mm. broad, rugose. This previously undescribed species was kindly brought to my notice by Professor R. H. Compton, who first examined the specimens collected by Mr. H. Hall of the Kirstenbosch staff. The fact that Mr. Hall found the plants growing in the open near the roadside, surprises one that such a strange looking species was not recorded earlier. It is very distinctive among Cape Province species of Huphorbia and its nearest affinity is with E. monteiroi, which occurs some considerable distance further north in South West Africa, Transvaal and Rhodesia. The tubercled stem is not unlike that of F. restituta N. E. Br., and, in the peculiar feature of pro- ducing several peduncles from a single flowering eye in successive seasons, it shows some relationship to the equally strange looking species, E, schoenlandii Pax. The red glands, ovary, style and filaments make it conspicuous when in flower. It is evident that the peduncles dry and persist for some time if allowed to do so, but plants in the wild state are mostly denuded either by the elements or, more likely, by grazing animals. 8 ra " ie Ons 7 ieee 7 . is'ds de’ sO - Sit Malt if i OW miter irene bla La a rrretrery Le TTEEET EE beer ae teen lm Hatters inka Fatetet Sesthiait Bods freee anole PuaTE XIX. Euphorbia hallii R. A. Dyer. Phot. H. King. THREE SMALL SPECIES OF OTHONNA. By R. H. Compton. (With Plate XX.) The genus Othonna includes, inter alia, a considerable number of spe- cies in the Western Cape Province and adjoining regions—many very imperfectly known—showing enlarged perennial storage organs. In some cases these are subterranean tubers from which rise shoots bearing leaves and flowers in winter and spring which disappear with the onset of the dry summer. In other species there is a thickened and somewhat branched aerial stem, often covered with a horny bark which bears a limited number of more or less abbreviated seasonal leaf- and flower-bearing shoots, often with a basal tuft of woolly hairs. The three species which are the subject of this paper are the dwarfest examples of this latter type. They are plants very liable to escape notice in the field owing to the fact that their very short brown stems hardly rise above the surface of the ground and are often covered by blown soil, and because of their small leaves and flowers and the short season during which they are in evidence. Two of them—0O. cacalioides L.f. and O. minima DC.—have long been on record though very seldom collected since their first discovery: the third species, to be described below, has only recently been brought to light. All three species grow within 50 miles of one another, and all are found in similar situations, viz. in shallow depressions or crevices in hard level or gently sloping quartzitic rocks, in a winter rainfall of perhaps 12—15 inches per annum, and exposed to full sunshine. They are clearly adapted to strongly seasonal conditions involving long aridity and high temper- tures. They are associated in their habitat with other small succulents and bulbous plants: one of them, O. cacalioides, is found in close association with the smallest known Conophytum, the smallest known Anacampseros, and one of the smallest known species of Crassula:* and the other species also grow in proximity to Conophytums. They form a closely related group, obviously derived from common ancestry, showing extreme adaptation to their special conditions of life, and so limited in distribution as to suggest recent evolution. In cultivation they show little change of form. * These three species I had the good fortune to find in one day in 1926 on the rocks near the summit of van Rhyn’s Pass: Conophytum Comptonii N. E. Br., Anacampseros Comptonii Pillans, Crassula Comptonii Hutchinson and Pillans: but I missed the Othonna. (See Hutchinson, Botanist in South Africa, p. 149, 1946.) 137 138 The Journal of South African Botany. Othonna cacalioides L.f. In his brief description the younger Linnaeus (Suppl. p. 388, 1781) gives the locality of O. cacalioides simply as “in rupibus Cap. bonae spei. Thunberg”’. In the Schultes edition of Thunberg’s Flora Capensis (p. 719, 1823), however, the following is given: “Crescit in Bockland’s summo monte in rupibus denudatis planis. Floret Octobri, Novembri.” Harvey (Fl. Cap. III p. 339, 1864) says “Hab. Bocklandsberg, Thunberg, (Herb. Sond.).” The locality, Bocklandsberg, is closely fixed by Thunberg’s Travels (See M. C. Karsten in Journ. 8.A. Bot. V, p. 147, 1939): the route being up the valley of the Doorn River (a tributary of the Zout River) about 20 miles north of van Rhyn’s Pass; the mountain on the south side of this valley being to-day known as the Bokkeveldberg, and being part of the same Bokkeveld Mountains escarpment up which van Rhyn’s Pass climbs. Thunberg’s type is at Uppsala, where it is labelled “‘Cap. b. spei. Thunberg”’. I am indebted to Dr. C. G. Alm for a photograph of the sheet. The isotype in the Stockholm Botanical Museum was lent me through the kindness of Dr. T. Norlindh, and was photographed by the University of Cape Town Photographic Department (Plate XX 1.). This sheet bears on the back a copy of the original L.f. description ete., with the addendum Fie. 1. Othonna cacalioides L.f. 1. Plant, life size: 2. Capitulum x 5. 3. Involucral bract <x 5. 4. Ray floret x 10. 5. Corolla of ray floret x 10. 6. Dise floret x 10. 7. Stamens of disc floret x 20. 8. Pollen presenter of dise floret x 10. 9. Ray achene x 10. G. G. Smith, Nat. Bot. Gdns. 101/46. (Del. W. F. Barker.) Three Small Species of Othonna. 139 “Specimen inde missit Exp. D.D. Thunberg anno 1775. Herb. Montin.”’ Mr. A. A. Bullock informs me that there is no specimen of O. cacalioides at Kew (apart, as we shall see, from “O. Pallansii Hutchinson’’). Mr. N. S. Pillans, who visited van Rhyn’s Pass in 1928, found an Othonna which he regarded as a new species, and on 11th October 1928 he pointed it out to Dr. J. Hutchinson, who gathered specimens for the Kew Herbarium (Hutchinson 781), and described it as O. Pillansii. (Botanist in 8S. Africa, p. 150, 1946.) In 1946 it was collected on the same spot by Mr. G. G. Smith (Nat. Bot. Gardens 101/46) and was grown at Kirstenbosch where it flowered next year and afterwards. (Specimen in Kirstenbosch Herbarium, 11 Sept. 1947.) O. Pillansii is undoubtedly conspecific with O. cacalioides. and Hutchinson’s name must therefore unfortunately disappear. Comparison of the recently collected van Rhyn’s Pass specimens with the isotype material and photographs establish the identity. Drawings of Hutchinson’s plant accompany his description, and further drawings, for which I have to thank Miss W. F. Barker, made from G. G. Smith’s plant, are published herewith. (Fig. 1.) Othonna minima DC. This species was described by De Candolle (Prodr. VI, p. 479, 1837), who stated that it was collected on the Giftberg by Drege. It is mentioned by Drége (Zwei pflgeog. Dok. p. 71, 1843) as being collected on the Giftberg, 1500—2000 ft. alt. in November. In the Flora Capensis (III, p. 339, 1864) Harvey cites a specimen in Sonder’s Herbarium and states that it was “raised in Hamburg Bot. Garden from Drége’s seeds”. Mr. A. A. Bullock (27 April 1948) informs me that Kew has material of O. minima DC. which “‘consists of a single inflorescence and half a dozen detached leaves of a specimen collected by Drége, Giftberg 1500—2000 ft in November 1839* and is evidently part of the type.” By the kindness of Dr. Norlindh I was sent the specimen of O. minima in the Stockholm Botanical Museum, and this has been photographed by the University of Cape Town Photographic Department. (Plate XX. 2). The sheet bears the inscription “Othonna minima DC. Hort. Hamb. 1839 e semina Dregeana enata. 43”’. The Giftberg was little visited botanically during the century since Drége made his collection, and the next record of O. minima was in Sep- tember 1948 when I found it growing in fair quantity on quartzitic pave- ment formation, in crevices and rock-pans at about 3000 ft. alt. (Compton pr ne ee ee ee ee *Drége, however, had left South Africa in 1834. The date 1839 is the same as that on the Stockholm specimen (see below) grown in Hamburg and evidently several years old. 140 The Journal of South African Botany. Fic. 2. Othonna minima DC. 1. Plant with leaves and flowers, nat. size. 2. Plant with leaves only, nat. size. 3, 4, 5. Capitulum, side view, apical view, cut lengthwise, x 5. 6, 7. Ray florets x 10. 8. Corolla of ray floret x 10. 9. Style and stigmas of ray floret x 10. 10. Disc floret x 10. 11. Four stamens of disc floret x 20. 12. Pollen presenter of disc floret x 20. 13. Involucral bract x 10. 14—18. Leaves, nat. size. Compton 20844. (Del. J. Elffers.) Three Small Species of Othonna. 14] 20844). Material was obtained for Kirstenbosch and was grown there and produced leaves and flowers abundantly without apparently changing its habit: specimens of these plants are also in the Kirstenbosch Herbarium. (Nat. Bot. Gdns. 1313/48.) The correspondence between my plants and those of Drége is exact. Miss J. Elffers made drawings of the wild material and these are reproduced herewith. (Fig. 2.) Mr. A. A. Bullock, writing in April 1948 on the subject of the Kew material of these small Othonnas—O. minima and O. Pillansii (i.e. O. cacalioides) remarks that “on the evidence of these specimens the two species must be maintained, but further material will probably show that they are conspecific.” In my opinion, however, they are distinct enough to be regarded still as separate species—to which I now add a third. O. minima differs from O. cacalioides in its more cylindrical (or button- shaped) stem-tips, its entire or only slightly denticulate leaves, its more elongated and slenderer peduncles which sometimes branch slightly, its longer and narrower ray ligules, the broader bell of its disc corollas, and the longer pappus hairs in both its ray and disc florets. Othonna pygmaea Compton sp. nov. Fruticulus parvus, caulibus brevibus incrassatis, foliis spathulatis, . integris vel leviter denticulatis, glabris, pedunculis simplicibus vel paulo ramosis. Capitulum parvum, involucri bracteis 5, late ovatis, obtusis, breviter apiculatis, minute ciliolatis. Radii flosculi 5, flavi, femini, breviter ligulati, parum involucri bracteas excedentes, ovario dense sericeo-pubescente, pappi setis uniseriatis, tenuibus: Disci flosculi c. 13, masculini, corollis flavis, e base anguste campanulatis, pappo nullo. Habitat. Cape Province. Clanwilliam Division: Klipfonteinrand, east of Brandewyn River, on sloping rocks, 1800 ft. alt., 29 April 1947 (young flowers and leaves), Compton 19472 (Type, in Herb. Nat. Bot. Gdns.): Grown at Kirstenbosch 207/47. Barker 4717, 13 Sept. 1947 (mature leaves): grown at Kirstenbosch 554/47. A small depressed shrublet, slightly branched, the branches contiguous, remaining short, obconical, | cm. or less in length, circular in section, the apex truncate, 1.5 em. in diameter or less, bearing one or more clusters of leaves with a few short brown woolly hairs at their base, hard in consis- tency and covered with a smooth brown horny bark which peels off and disappears. Leaves beginning to appear at flowering time, growing as the flowers fade, finally reaching 2.56—3.5 em. long, 1.0—1.5 em. wide in well grown plants, softly succulent, glabrous, glaucous, nerveless when fresh, 3-nerved when dry, flat, broadly spathulate, usually obtuse, occas- ionally retuse or subacute, entire or slightly lobulate-crenate, often having a few distant denticles, deciduous. Inflorescences one or few, erect, arising 142 The Journal of South African Botany. 19 Fic. 3. Othonna pygmaea Compton. 1. Plant in flower, nat. size. 2. A smaller speci- men, nat. size. 3. Plant in leaf, nat. size. 4, 5, 6. Capitulum, side view, apical view, cut lengthwise x 5. 7, 8. Ray florets x 10. 9. Corolla of ray floret x 10. 10. Style of ray floret x 10. 11. Dise floret « 10. 12. Four stamens from disc floret 20. 13. Pollen presenter of dise floret x 20. 14. Involucral bract x 10 15—19. Leaves, nat. size. 20. Achene x 10. Compton 19472 and Nat. Bot. Gdns. 207/47. (Del. W. F. Barker.) Three Small Species of Othonna. 143 from the stem apex, 2—5 em. long, simple or slightly panicled, the peduncle terete, glabrous, c. 0.7 mm. thick, bearing one or a few minute subtending bracts. Capitulum c. 5 mm. diam. when fully expanded. Involucre broadly campanulate, slightly fleshy, green, consisting of 5 bracts, slightly connate at the base, valvate in the bud, broadly ovate, obtuse, shortly apiculate, minutely denticulate, 3 mm. long, 1.8 mm. broad. Ray florets 5, opposite the involucral bracts, shortly ligulate, female, the corolla yellow, 1.5 mm. long, 1.0 mm. wide, scarcely exceeding the involucre; ovary densely silky-pubescent, stigmas filiform. Disc florets c. 13, male, corolla yellow, campanulate from a short slender base, 1.5 mm. long, the 5 lobes about equal in length to the tube, ovary rudimentary, style with a clavate pollen presenter. Pappus of ray florets of numerous delicate hairs in one series, 0.5 mm. long, absent in the disc florets. An inconspicuous plant growing in shallow pan of coarse sandy detri- tus in gently sloping quartzite pavement rocks of the Table Mountain Sandstone formation. Most of the plants have only one or a few stems, but some reach a diameter of 5 cm. and may have as many as 15—20 branches, which however remain short and in the resting state hardly project above the level of the sand. From 0. cacalioides it differs in its usually paniculate inflorescence, the more slender peduncles, the smaller capitula, the broader shape of the involucre, the much shorter ray florets, the fewer disc florets and the absence of pappus in the disc florets: the growth of its stem tends to be more depressed and the leaves spread more widely from the stem apices. From O. minima it differs in its usually freely paniculate inflorescence, _the smaller capitula with fewer disc florets, the shorter ray florets, the shorter pappus in the ray florets and its absence in the disc florets: the stems are of less definite, more or less obconic shape. rs — 7 na abrsy yh yang! 2 = Sai aby a pt deten ma 4 ish Pi 7 we. “Ay pha ai _ . ay aa! al a ti tauoth, en =} peu 4 THI ie toe z PLATE XX. 1. Othonna cacalioides L.f. Thunberg, 1775. Isotype in Botanical Museum, Stockholm. Nat. size. 2. Othonna minima DC. Grown from Drége’s seeds in Hamburg Botanic Garden, 1839. Isotype in Botanical Museum, Stockholm. Nat. size. 3. Othonna pygmaea Compton. Klipfonteinrand, 29 April 1947. Compton 19472 in Herb. Nat. Bot. Gdns. (Type.) % nat. size (linear). Photographs by University of Cape Town Photographic Department. A NEW CRASSULA FROM THE RICHTERSVELD. By H. Herre. (With Plate XXI.) Crassula fusca Herre sp. nov. Herba perennis, glaber 20—30 em. alta. Caulis ramique teretes, internodiis viridibus vel roseis 4—5 mm. longis. Folia sessilia connata basi, 2—4 cm. longa, 9—15 mm. lata, acuta, superi- ore plana vel subconcava, dorso convexa, ad margines minutissime papil- losa, faciebus alba-lineata. Habitat prope Numees et Hellskloof, Richters- veld, folia fusca similia Aloé Pearsonii Schoenl. in cultu +. viridia. Inflorescentia (dichasium) multiflora terminalis ex axillis foliorum superi- orum 8—12 cm. longa et lata. Pedunculi 2—3 cm. longa: bracteis quam folia multo minoribus, floribus pedicillatis, pedicellis glabris, 2—3 mm. longis. Sepala 5, sublibera, lanceolata, acuminata, dorso leviter convexa, viridia, rufo-cincta, glaber, ca. 2 mm. longa. Petela 5, juvenilia alba, adulta pallidissima rosea, sublibera, lanceolata, acuminata, apice leviter recurva, 3—3.25 mm. longa, 1—1.25 mm. lata. Carpidia ca. 2 mm. longa. Stamina 5, ca. 2.5—3 mm. longa, filamentis albis, antheris oblongis, luteis. Squwamae obcuneatae, luteae, ca. 0.5 mm. latae. Perennial, glabrous, stems ascending, about 20—30 em. long with an additional 8—12 cm. for the inflorescence. The stem is about 5 mm. in diameter. Internodes start with 4—5 mm. in length while the last one is about 1—2 cm. long. All are greenish to reddish. Leaves 2—4 cm. long, 9—15 mm. broad, amplexicaul, connate at their base, acuminate, _ sometimes recurved at the apex, fleshy, slightly concave on the inner side, slightly convex on the outer, glabrous, margined with tiny white papillae, thus forming a white edge; green, densely punctate. In its home the leaves are rust-red like those of Aloé Pearsonii Schoenl. and the name is derived from these most beautiful coloured leaves. In cultivation they are more or less green, especially when kept in shade. The inflorescence is a many flowered dichasial cyme about 8—12 cm. long and broad. The floral bracts are like the foliage leaves but much smaller. The cyme branches are glabrous, 2—3 mm. long and at least one of the flowers is almost sessile. The 5 sepals are almost free, pale-green or tinged reddish, lanceo- late, acute, dorsally slightly convex, glabrous, about 2 mm. long. Petals 5, white, when older light pink, almost free, lanceolate, acuminate, re- curved at the apex, about 3—3.25 mm. long and 1—1.25 mm. broad. Stamens 5, about 0.5—1 mm. longer than the pistil, which is about 2 mm. long. Filaments white, anthers yellow. Carpels slender, colourless. Squamae yellowish, obcuneate, about 0.5 mm. broad. 145 146 The Journal of South African Botany. This fine species is nearly related to Crassula MacOwaniana Schoenl. et Bak-f. but differs from it in forming smaller bushes, which have much smaller leaves of another shape, which also show their fine colour especi- ally in their natural environment. The inflorescence and the inner parts of the flower differ too. Numees, Hellskloof, Richtersveld. H. Herre. Sept. 1929. Oct. 1930. Oct. 1933. Oct. 1939. P. van Heerde, June 1949. University Botanic Garden Stellenbosch No. 6077. Flowering: Nov. 1932; Dec. 1933; Nov. 1937: Dec. 1939; Nov./Dec. 1949. As can be made out from the collecting dates given above, this species was collected for the first time by myself during my first visit to the Richtersveld in 1929, but as the plant was at first thought to be a variety or form of C. Macowaniana no description was written down. But the following two collections showed that our plant had nothing to do with the latter species, but seemed to be a good new one. Unfortunately the plants died again and only a bit of them had been preserved in alcohol, but this was not sufficient for a good description. When I collected the plant again in 1939 I sent material of it to the Nestor of the genus Crassula, Dr. 8S. Schoenland at Grahamstown. It was only some weeks before his death that he let me know, that after his careful identification it was a good, new species and he suggested that I describe it as I knew it best in its natural home as also in cultivation. I tried to do so, but before it flowered again, the plants died. It was nearly 10 years later when Mr. P. v. Heerde of Springbok, the well-known succulent collecsor of that area succeeded to get this plant again after I had told him where to look for it. Now I was able to draw up a proper description and to illustrate it by a good photograph of a flowering branch. Hence after all these misfor- tunes this fine species, which is probably the last new one of this large genus which its careful worker, the late Dr. S. Schoenlind, has had in his hands, can be published and no doubt will make friends when intro- duced into cultivation. As regards this, it is not more difficult than other species. It likes a resting period after the flowering time. but as it grows near the Hellskloof which catches quite a lot of clouds and rains with its high mountains, it must not be cultivated too dry, otherwise it collapses, which happened with us before. On the other hand it never wants to be too wet. In its natural environment it grows either alone among stones and crevices or together with other shrubs in the shade or half-shade of higher ones. The plants surely become bigger than those shown in our photographs. Fic. Fic. Fie. PTATE XOXT Crassula fusca Herre. A plant growing in the Richtersveld (Numees, Hell’s Kloof). A plant grown in a rock-garden at Stellenbosch University Botanic Garden. A pot-plant at Stellenbosch University Botanic Garden. SILICULARIA, A NEW GENUS OF CRUCIFERAE, WITH NOTES ON RELATED GENERA. By R. H. Compton. In the mountains of the western Cape Province, and particularly in the Cederberg and Cold Bokkeveld areas of the Ceres and Clanwilliam Divisions, there occur four shrubby genera of Cruciferae characterised by their slender virgate, sparingly leafy, genistoid growth. Three of these genera, viz. Brachycarpaea, Cycloptychis and Schlechteria, have been described up to the present, and a fourth genus, Silicularia, is now added. The similarity in habit of these plants (which is also shared to some extent by certain species of Heliophila) does not extend to their fruits and seeds, on which basis, as is usual in the Cruciferae, the various genera are mainly separated. To the description of Silicularia sigillata below are appended notes on the other genera, as well as on the related genus Thlaspeocarpa. SILICULARIA Compton gen. nov. Genus Cruciferarum-Isatidearum adhue monotypicum. fructus orbiculato-ellipticus, planus, valvis concavis, reticulato-rugosis, erostratus, eseptatus. Semen unum, orbiculatum, complanatum, exalatum. Cotyle- donae transverse conduplicatae. Distinguished from Cycloptychis by the siliculae which contain only one central seed, though the ovary is normally biovulate, and are devoid of both beak and septum. Silicularia sigillata Compton sp. nov. Fruticulus anguste virgatus, omnino glaber. Caules teretes, rigidi. Folia pauca, linearia vel angustissime spathulata, integra. lores numerosi, in racemis ebracteatis pedicellati. Sepala erecta, convexa, oblonga, obtusa. Petala explanata, late spathulata, obtusa. Antherae obtuse sagittatae. Ovariwm lanceolatum, 2-ovulatum, stylo breve, stig- mate globoso. Silicula orbiculato-elliptica, valvis externe concavis, reticulato-rugosis: desunt septum rostrumque. Semen solitarium, com- planatum, exalatum. Habitat. Cape Province. Ceres Division: Ceres Wild Flower Show, Oct. 1932, Compton 4144: De Keur, 27 Sept. 1936, Compton 6491; 1 Oct 1944, Compton 16221. (Type, in Herb. Nat. Bot. Gdns.): top of Gydo Pass, 27 Sept. 1936, Lewis s.n.: Ertjesland Kloof, 28 Sept. 1944, Compton 16092. 147 148 The Journal of South African Botany. 3 Fie. 1. A. Silieularia sigillata Compton. 1. Branch, nat. size. 2. Long stamen x 5. 3. Short stamen x 5. 4. Petal x 5. 5. Gynaecium with one carpel wall removed xX 5. 6. Gynaecium x 5. 7.Sepal x 5. 8. Fruit x 5. 9. Fruit with one valve removed x 5. 10. Embryo x 5. Compton 16221. B. Thlaspeocarpa capensis C. A. Smith. 1. Fruit x 24. 2. Fruit with one valve removed xX 24. 3. Seed x 5. 4. Embryo x 5. Compton 3755. C. Schlechteria capensis Bolus. 1. Fruit x 5. 2. Fruit with one valve removed x 5. 3. Embryo x 5. Esterhuysen 21730. (Del. W. F. Barker.) Silicularia, A New Genus of Cruciferae, 149 with Notes on Related Genera. A slender virgate shrublet, branching mostly at the base, glabrous in all parts, 30—50 cm. high. Stems terete, rigid, smooth, striatulate. Leaves 25 mm. long, 1.5—2 mm. wide, the apex and margins slightly incurved. few, scattered, erect, linear or very narrowly spathulate, entire, 15 Fluwers in a long lax ebracteate raceme up to c. 15 em. long, erect, spreading or deflexed. Pedicels slender, ebracteate, curved, c. 5 mm. long. Sepals erect, ec. 5 mm. long, convex externally, oblong, obtuse, the antero-posterior pair slightly gibbous at the base, the lateral pair obtuse, cucullate, membranous at the tip. Petals spreading, white or pale pink, membranous, broadly spathulate, obtuse, c. 8 mm. long, 4 mm. wide. Filaments 3 mm. long, tapering slightly from base to apex. Anthers obtusely sagittate. Ovary lanceolate, somewhat flattened, 2 mm. long, 1 mm. broad, 2-ovulate: style short, stigma globuse. Fruit orbicular- elliptic, c. 4 mm. long, 3.5 mm. wide, flattened, the valves concave, reticulate-rugose, without septum, containing one seed and one sterile ovule. Seed with long funicle attached to the edge of the valve, orbicular, flattened, wingless, 2 mm. diameter, the testa membranous, the embryo with transversely conduplicate cotyledons. The fruit has vestiges of a very thin septum around the inner edges of the sutures, though the middle is quite open. It may be that the septum is continuous in the very young stages of the ovary, and that it ruptures as the fruit enlarges, leaving only a marginal fringe; but of this I have not been able to satisfy myself. This broom-like Crucifer has only been found in a limited area of the Cold Bokkeveld, viz. at De Keur, near the top of the Gydo Pass and in the adjoining Ertjesland Kloof (the locality of the Ceres Wild Flower Show specimens not being exactly known, though within the Ceres Divi- sion). It grows in crevices and on ledges of Table Mountain Sandstone rocks at an altitude of about 3000 ft. It frequently fails to produce fruits, and in this condition can easily be mistaken for a Cycloptychis or Brachycarpaea. It has an attractive appearance owing to the large number of flowers which are open simultaneously. The corollas are white or pale pink. It flowers in September and October, the fruits, when formed, being mature in November: they are probably indehiscent, though the valves can be split apart with a needle. The specific name refers to the thick valves of the silicula with their curiously impressed pattern. Brachycarpaea DC. Little will be said here about this genus which is the most widely spread of the group. It is commonly understood to consist of two species: B. varians DC., mainly occurring in the western Cape Table Mountain 150 The Journal of South African Botany. Sandstone formation, is a very attractive plant, grown at Kirstenbosch and well worth more extended cultivation: B. axa Sond. has a much wider range, extending from Namaqualand through all the coastal Divisions and some inland ones as far as Bredasdorp, and showing a greater degree of variability, as might be expected. Both species were divided into varieties by Sonder (Flora Capensis), but further study, especially of B. laxa, in relation to habitat is necessary. The form from the Southern Cape Peninsula (which Sonder apparently did not see), should almost certainly be distinguished from those of Montagu, Laingsburg, Clan- william, van Rhynsdorp and Namaqualand. Cycloptychis HK. Mey. Dr. E. P. Phillips (Gen. S. A. Fl. Plants, ed. II, p. 347, 1951) states that this genus comprises three species. Of these C. virgata EK. Mey is the type. The others are presumably C. polygaloides Sond. which I believe to be a Heliophila (see below), and C. Marloth O. E. Schultz, on the type material of which Phillips has written “not genus Cycloptychis”’, but which I think is best retained in that genus. Sonder’s type of “C. polygaloides” (Fl. Cap. I, p. 34, 1859) was collected by Zeyher at Tulbagh in November. I have not seen the type specimen, but the plant Sonder describes is very unlike C. virgata in habit, leaf characters etc., and though the ovary is said to be rostro-acuminate, the ripe fruit, on which generic discrimination must chiefly depend, was un- known. There is nothing in Sonder’s description, in fact, to separate the plant with certainty from Heliophila, a more or less pronounced beak being frequent in that genus. Schlechter (Engl. Jahrb. 27,136, 1899) described a Heliophila which he had collected on the Cold Bokkeveld at 4000 ft. alt. on 9 Sept. 1896 (Schltr. 8929) as H. nubigena, and this plant has been collected on many occasions since. One of the gatherings is by A. Bolus (Bolus 6349) from the slopes of the Winterhoek Mts., Tulbagh, 5500 ft., Oct. 1896—that is, reasonably close to Zeyher’s Cycloptychis polygaloides locality. The same plant was collected by Miss E. Hsterhuysen (1305) in Duivel’s Kloof, Great Drakenstein Mts. 2/3000 ft. on 6 Oct 1935, by myself (Compton 9758) on the Audensberg, Worcester Division, 6 Oct. 1940 and by Miss Esterhuysen on Audensberg summit on the same day, on a Mountain Club Meet. Since then it has been collected repeatedly by Miss Esterhuysen in surrounding mountain localities, viz. Chavonnesberg, 4 Oct 1942 (8187); Keeromsberg 5/6000 ft., 8 No. 1943 (9300); Milner Ridge Peak 4/6000 ft., 11 Nov. 1943 (9335); Brandwacht Mt. 6000 ft., 26 Nov. 1944 (11015); Duivel’s Kloof (again) 14 Sept 1947 (14043); Silicularia, A New Genus of Cruciferae, 151 with Notes on Related Genera. Fie. 2. D. Cycloptychis Marlothii O. E. Schultz. 1. Septum x 5. 2. Outer carpel wall, from inside x 5. 3. Outer carpel wall, from outside x 5. 4. Embryo x 5. Esterhuysen 12716. E. Brachycarpaea varians DC. 1. Fruit x 5. 2. One carpel with one wall removed x 5. 3. Embryo x 5. Henderson 1006. F. Cycloptychis virgata E. Mey. 1. Septum x 5. 2. Outer carpel wall, from inside x 5. 3. Outer carpel wall, from outside x 5. 4. Seed x 5. 5. Embryo x°5. Leipoldt 3900. (Del. W. F. Barker.) 152 The Journal of South African Botany. Chavonnesberg (again), foot of cliffs. 5000 ft., 3 Oct. 1948 (14595): Wemmershoek Peak 5500 ft.. 3 Nov. 1951 (19199a). Schlechter’s H. nubigena has a villous pedicel, calyx and fruit. In his description he states that the ovary is 1—2-ovulate. In fruits which I have dissected there are 4 ovules, of which only one enlarges: a well- marked continuous septum is present, and this is left spread between the persistent margins of the fruit when the two valves fall off. A drawing by Miss M. Walgate seems to indicate that occasionally, at any rate, two ovules may mature. Material is scanty and I have been unable to deter- mine the form of the mature embryo. though I should be greatly surprised if it were conduplicate. These plants agree closely with Sonder’s description of Cycloptychis polygaloides, and I am convinced of their identity. If I am right. and if the plant belongs rather to the genus Heliophila, its name would become Heliophila polygaloides (Sond.) Compton. If this be accepted, Schlechter’s name H. polygaloides for a different, though related plant, Schltr. 8900, must be abandoned and a new name found: I propose to call it Heliophila nubigenoides Compton nom. nov. The nomenclature thus becomes: Heliophila polygaloides (Sond.) Compton non Schlechter, comb. nov. (=Cycloptychis polygaloides Sond.) Heliophila nubigenoides Compton nom. nov. (=H. polygaloides Schltr.) There remains one other plant which has been placed in Cycloptychis, where I propose to leave it. in spite of Phillips’ doubts on the matter. This is C. Marlothii O. E. Schultz (in Engl. Bot. Jahrb. 66, p. 97, 1933), founded on Marloth 3362. collected in the Cold Bokkeveld. 1000 m. alt., in flower, in October 1903. Through the kindness of Dr. R. A. Dyer, chief of the Division of Botany and Plant Pathology, I have been enabled to examine Marloth’s type specimen in the National Herbarium, Pretoria. As far as I can ascertain it has only been collected once since, viz. by Miss Esterhuysen, who obtained fruiting specimens on the east spur of the Hondvyerbrand Ridge, Southern Cederberg. on 21 April 1946. There are no flowers on these specimens (Hsterhuysen 12716 in the Kirstenbosch and National Herbaria, 12719a@ in the Bolus Herbarium), and the fruits are slightly narrower than in the type, but I cannot doubt their specific identity. Schlechteria Bolus. This monotypic genus was founded (Bolus in Engl. Bot. Jahrb. 24, p. 455, 1897) on Schltr 8765 from the Koudeberg, near Wupperthal. 30 August, 1896, the single species being named S. capensis Bolus. I have Silicularia, A New Genus of Cruciferae, 153 with Notes on Related Genera. examined the type in the Bolus Herbarium, which is accompanied by Dr. Bolus’ original MS. description and drawings. Fifty years elapsed before this plant was again collected, when it was found by Miss EK. Hsterhuysen 13130 at Duivelsgat in the Sneeuwberg area of the Cederberg on 11 October 1946: her material shows young and older fruits and only one flower, rather larger than described by Bolus, the whole plant being more leafy and better grown. Next year she collected it again (Hsterhuysen 13803) in a shady rock gully in the Cederberg Tafel- berg at 6000 ft. on 26 Feb. 1947: in this gathering also there were young and older fruits but no flowers. While this paper was being written Miss Esterhuysen again brought in specimens (Hsterhuysen 21730) from Pakhuis Peak, Cederberg, between rocks at 3500 ft. alt, 6 Sept. 1953: this bore mature but still green fruits with fully formed seeds: a very few flowers were also present. It seems clear from the records that the plant has no well-defined flowering season and that the perianth and androecium must be very fugacious. Miss Esterhuysen’s material enables corrections to be made in the original description of the plant. The fruits in Schltr. 8765 are now seen to be immature, and the embryo had not attained its full size and final type of folding. Bolus’ description reads “embryo curvus: cotyledones accumbentes, lineares’. The Esterhuysen plants carry fully developed seeds, flattened like a biconvex lens, about 4 mm. in diameter, completely filled with a conduplicately folded embryo. In another important respect the printed description of Schlechteria requires correction. It is there stated that the ovary is 3-locular—a very unusual feature in the Cruciferae. In Bolus’ original manuscript, confirmed by the drawings, however, it is described as 2-locular, the printed version being obviously in error. The mistakes in the printed description have been perpetuated by Phillips (Genera Fl. Plants ed. I, p. 353), resulting in Schlechteria being placed away from its true affinity. Bolus’ drawings accompanying the type specimens include a “‘diagram- matic section” of the ovary showing a very thin septum with two ovules on each side of it. The description reads “‘siliqua septo membranaceo tenuissimo obliterato 1-locularis abortu l-sperma’’. I have been unable to find a septum in any of the fruits examined, and if a vestigial membrane should be present in the young state it must disappear very early in development. Thlaspeocarpa C. A. Smith. In the Flora Capensis vol I, p. 35, 1859, Sonder described a monotypic genus under the name Palmstruckia with the intention of preserving 154 The Journal of South African Botany. Palmstruck’s name, which had been abandoned in the case of a Scrophu- lariaceous genus to which it had originally been applied. International Rules, however, prohibit recourse to this method of commemoration, and a new name had to be found for the Cruciferous plant in question: this was done by Mr. C. A. Smith (in Kew Bull. 1931, p. 156) who chose the name Thlaspeocarpa, the single species becoming 7’. capensis. This plant, destined to considerable confusion as regards name and identity (see Smith loc. cit.) was collected by Thunberg in November 1774 on the “Onderste Roggeveld”. Smith states that this is “near the southern limits of the Calvinia Division”: but Thunberg had mentioned that this district is so called, not because it lies lower than the other Roggevelds, but because it is furthest from the Cape. (See M. C. Karsten, Journ. S.A. Bot. vol V, p. 148, 1939, whose map shows it east of the Hantam Mountains, across the Fish River, this corresponding with the maps of Lichtenstein, Barrow and Burchell.) Thunberg’s plant was confused with Peltaria capensis Lf. in the Schultes Fl. Cap. p. 490, but was recognised as distinct by Sonder. After Thunberg’s original discovery it remained botanically unnoticed until Bolus collected a Namaqualand specimen in October 1883 “in stony places, Kasteel Poort, near Klipfontein, 3000 ft.” On this specimen (Bolus 6502 in Herb. Bolus) he wrote—“Thunberg’s type of Peltaria capensis was sent from Upsal to Kew for the express purpose of comparing with this in 1885. Mr. N. E. Brown reports that he has no doubt they are the same. Thunberg’s specimen had only filiform leaves, but his specimen consisted only of the upper part of the plant and the leaves were reduced to bracts. The fruit in his was mature and } an inch in diameter.” Thlaspeocarpa was again collected by J. T. Anderson at Klipfontein, Namaqualand in August 1890 (Guthrie no. 2325 in Herb. Nat. Bot. Gdns., determined by Bolus). It has been gathered twice near Nieuwoudtville— a locality not so remote from the Roggeveld as is Namaqualand—by Leipoldt (889, Karoo soil near Willems River 3200 ft., not very common, Sept., in Herb. S.A. Museum), and by Miss M. Lavis (in Herb. Bolus 19828, Klipkoppies about 6 miles N.E. of Nieuwoudtville, c. 2500 ft., in red clay, Sept. 1930). In Sept. 1931 I found it growing in sandy soil at Constable, Laingsburg Division, 3000 ft. (Compton 3755 in the Kirsten- bosch and Bolus Herbaria.) It may be possible to distinguish two species among these widely separated gatherings. Compton 3755 agrees with Thunberg’s plant in the dimensions of the fruit as given by Sonder: the Lavis plant also agrees with these in size of fruit and width of leaflets: Leipoldt’s plant is appar- ently the same. Bolus 6502 has narrower leaflets and smaller fruits than Thunberg’s, which Bolus apparently attributes to immaturity. Silicularia, A New Genus of Cruciferae, 155 with Notes on Related Genera. The Compton plant has a pilose calyx and pedicel and its colour is reddish: other gatherings have glabrous calyx and pedicels and white petals. Eventually it may be found that the Namaqualand plants belong to a distinct species, while the Lavis, Leipoldt and Compton gatherings (neglecting differences of colour and hairiness) are the true 7’. capensis. It may be remarked that considerable differences in habitat occur amongst them, some growing in stony places, some in red clay and some in sand. More material of this curious Crucifer, with its striking Lunaria-like fruits and its rather wide distribution, is very desirable. NOTES ON NOMENCLATURE IN LIGHT FOOTIA. By R. S. ApDAMson. I. L. subulata. There has existed a certain amount of doubt as to the correct name to be used for the species variously known as L. subulata L’Hérit., L. sessili- flora Sond. and L. fruticosa Druce. The last of these which was the one adopted in the Flora of the Cape Peninsula, was based on Campanula fruticosa L., a name that has been the cause of some confusion and which since the early part of last century has been looked upon as belonging to a Lightfootia. The original diagnosis (Sp. Pl. 168. 1753) which was repeated un- changed in 1763 (Sp. Pl. ed. 2. 238), was: “CO. capsulis quinquelocularibus columnaribus, caule fruticoso, foliis lineari- subulatis, pedunculis longissimis. C. Africana, ericae folio, flore coeruleo patulo. Herm. afr. 5.” This is a diagnosis not strictly applicable to any species of either Lightfootia or of Prismatocarpus. As there is no specimen in the Linnaean herbarium, the only means to determine the real identity is the Hermann specimen quoted which can be looked upon as the type of Linné’s species. This is now in the British Museum, London, and Dr. G. Taylor, Keeper of Botany, has kindly examined it for me and reports that there are three specimens so named all of which are without doubt Prismatocarpus. It seems probable that L’Heéritier had seen this specimen when he described Prismatocarpus fruticosus (Sert. Ang. 2. 1788), quoting C. fruticosa L. as a synonym, and rightly transferred the specific epithet to his new genus. His view was followed by later writers, e.g. Gmelin (Syst. 1. 352. 1791), even though the species was retained in Campanula. Doubt about the identity of Linné’s plant seems to have started with Willdenow (Sp. Pl. 2. 912. 1799) who was evidently impressed by the phrase “‘capsulis quinquelocularibus” and expressed doubt as to the genus under which the species should be placed. Roemer & Schultes (Syst. 5. 152. 1819) quote Willdenow’s note and further state that the specimens so named that they had seen were refer- able to Lightfootia. A short time afterwards Sprengel (Syst. i. 809. 1825) went further and regarded C. fruticosa L. as a synonym of L. subulata L’Her. A. DeCandolle in his monograph (Monog. Camp. 108. 1830) discussed, under L. subulata, the application of the name C. fruticosa L., but left it as being of great uncertainty. 157 158 The Journal of South African Botany. The whole difficulty seems to have arisen from attempts, without reference to specimens, to identify plants from an inadequate and incorrect diagnosis. The description of the capsule as 5-chambered seemed to rule out any applicability to Prismatocarpus. The specimens, however, abolish all doubt and completely support the views adopted by L.Héritier. The combination made by Druce (Rep. Bot. Ex. Club Brit. Is. 3.5. 420. 1914) is hence illegitimate. Of the other names, there is no doubt that C. sessiliflora L.f. is the same species but Sonder’s combination is antedated by L. sessiflora Spreng. which is synonymous with Prismatocarpus campanuloides (L.f.) Sond. The correct synonymy is: Lightfootia subulata L’Hérit. Sert. Ang. 1. 1788 non Engler Bot. Jahrb. 40. 48. 1908. Campanula subulata Spreng. Syst. 1. 728. 1825 non Thunb. Prod. Pl. Cap. 38. 1794. C. sessiliflora (L.f.) Supp. Pl. 139. 1781. L, sessiliflora Sond. Fl. Cap. 3. 555. 1865. non Spreng. Syst. 1. 809. 1825. L. fruticosa Druce Rep. Bot. Ex. Club Br. Is. 3.5 420. 1914. Il. L. ciliata Sond. Sonder’s name (Fl. Cap. 3. 561. 1865) is antedated by L. ciliata Spreng. Syst. 1. 809. 1825, which is synonymous with Prismatocarpus fruticosus L’Herit. Sonder’s epithet was taken from Campanula ciliata Thunb. FI. Cap. ed. 1. 2. 24. 1818 (ed. Schult. 172. 1823) but this was itself antedated by C. ciliata Stev. Mem. Soc. Nat. Mose. 3. 256. 1812. and the name for the S. African species was changed to C. Thunbergii by Roemer & Schultes. It was later transferred to Roella by A. DeCandolle. Thunberg’s species was rather meagrely described and as both Roemer and Schultes and A. DeCandolle repeat it unchanged, it would appear that neither were familiar with the plant. A. DeCandolle saw a specimen named by Thunberg himself but it was of a quite different plant. There is no extant type specimen. Taken together the vague description and the lack of a type seem ground for dropping the name. This is especially the case as the combination under Lightfootia might easily be a source of confusion with L. Thunbergiana Buek. (cf. Intern. Code Bot. Nomen. Ine, 383%, 18%.) In describing his species Sonder quoted L. tenella Lodd. Bot. Cab. t. 1038. 1825. as a synonym. This is based on a plate with a very short description. There is no specimen. The plant seems to be more like L. Buekii Sond. than that under discussion. Loddiges’ plant was re- Notes on Nomenclature in Lightfootia. 159 named L. Loddigesii A.DC. Monog. Camp. 114. 1830 but this is a super- fluous name. Buek in 1837 described independently five species, L. divaricata, L. intermedia, L. mucronulata, L. thymifolia, and L. uitenhagensis, all of which can be regarded as belonging to the species in question. Of these L. mucronulata is that form described by Sonder as L. ciliata var. debilis. The specimens issued by Ecklon & Zeyher of L. thymifolia are mixed and represent more than one species. Even though its type is not the commonest form of the species, it is suggested that the name L. divaricata Buek be adopted. The synonymy 1S: L. divaricata Buek in E. & Z. Enum. 375. 1837. L. ciliata Sond. Fl. Cap. 3. 561. 1865 non Spreng. Syst. 1. 809. 1825. Campanula ciliata Thunb. Fl. Cap. ed. 1. 2. 24. 1818 non Stev. Mem. Soc. Nat. Mose. 3. 256. 1812. C. Thunbergu Roem. & Schult. Syst. 5. 135. 1819. Roella Thunbergui A. DC. Monog. Camp. 174. 1830. LI. intermedia Buek. 1. c. 374. L. wtenhagensis Buek 1. c. 375. L. thymifolia Buek 1. ce. 373 (p. pte.). var. debilis (Sond.) comb. nov. L. ciliata var. debilis Sond. Fl. Cap. 3. 561. 1865. L. mucronulata Buek. 1. c. 375. ALIEN PLANTS GROWING WITHOUT CULTIVA- TION IN THE SOMERSET WEST NEIGHBOURHOOD. By R. N. Parker. On coming to Somerset West in 1939 the large number of exotic plants established in the country seemed to me very noticeable. A number of these are described in the Flora Capensis. In the following list of aliens I have observed, those not described therein are marked with an asterisk. Some of these additional species had doubtless been overlooked or not collected at the time the Flora Capensis was written, but many appear to be recent introductions. In the CRUCIFERAE of 12 species only 2 are described in the Flora Capensis published in 1859—60. In the LEGUMINOSAE, of 24 species 6 are described in the Flora Capensis published in 1861—62. In the GRAMINEAE, of 44 species 37 are described in the Flora Capensis published in 1899—1900. It seems therefore that the alien flora is steadily increasing. The country of which these aliens are native has been indicated as far as possible. Of the 215 species listed, 137 are from Europe or the Mediterranean region, approximately half from each of these two areas, which however cannot be sharply distinguished. This amounts to 63 per. cent of the total. The next largest contribution has come from tropical and sub-tropical America, approximately 15 per cent. W. W. Robbins (Bulletin 637, University of California, Berkeley, California, 1940) lists 526 species of aliens in California of which 72 per cent are from Europe and Western Asia and 10 per cent are from S. America. The alien floras of California and the Winter Rainfall area of S. Africa are remarkably similar. More than half of the species in the following list are also naturalized aliens in California. The similarity extends to the families as well as to the species. The families best repre- sented and the number of species in each are: South Africa California Gramineae .. 44 Gramineae .. 111 Compositae .. 24 Compositae .. 96 Cruciferae .. 12 Cruciferae .. 43 Leguminosae 24 Leguminosae 28 Caryophyllaceae 10 Caryophyllaceae 21 Chenopodiaceae 9 Chenopodiaceae 17 161 162 The Journal of South African Botany. Robbins suggests that the large proportion of European weeds in California is due to the human population having mainly come from Kurope and to the similarity of the climates of the Mediterranean region and of California. Neither explanation seems to me to cover the facts completely as regards the alien flora of South Africa. Australia, which has provided 18 species, has only contributed one annual (A piwm lepto- phyllum F. v. M.), a South American plant said to be also indigenous to Australia. Similarity of climate explains why Australia should have provided over 40 per cent of the introduced trees and shrubs but leaves the absence of annuals unexplained. The tropical and sub-tropical American area, which has made practically no contribution to the human population, is nonetheless an important source of weeds. The introduced herbaceous plants are mostly weeds of cultivation. Agriculture started somewhere in the area of the Eastern Mediterranean, including Egypt and Western Asia. It also arose independently at a later date in the highlands of America from Mexico to Peru. Certain plants in these areas, finding conditions suitable, or, perhaps, becoming adapted to conditions produced by agricultural operations, became weeds of culti- vation. When agriculture spread to new areas these weeds would find conditions equally suitable. The Mediterranean area and Europe, being the oldest and largest continuous area of cultivation, would naturally produce the greatest number of weeds. The highlands of America from Mexico to Peru, being a later agricultural area and also more disconnected, provide a much smaller, though still important, number of weeds. Aus- tralia with no indigenous agriculture, contributes only one weed which may have come to South Africa direct from South America. This list of aliens was started before the Flora of the Cape Peninsula was published and most of the species enumerated will be found in that work, as well as many others which do not seem to have reached as far inland as Somerset West. As aliens enter a country mainly through the ports and take time to spread, it is natural that more should be found near the ports than further inland. I wish to acknowledge assistance in identification to the late Drs. Fernald and Pennel, and to Dr. Piper Smith, C. E. Hubbard and to my nephew, P. Brenan, Kew. (Note.—The numbers in the following list refer to specimens collected and sent to the Bolus Herbarium, Cape Town University.) RANUNCULACEAE. *Ranunculus muricatus Linn. 4464. A garden weed in Somerset West. Mediterranean. Alien Plants Growing without Cultivation 163 in the Somerset West Neighbourhood. PAPAVERACEAE. *Argemone mexicana Linn. Mexican Poppy. Roadside weed. Not common. Tropical and sub-tropical America. *Fumaria capreolata Linn. 4818. Ramping Fumitory. In woods and shady places. Mediterranean. Fumaria muralis Sond. Fumitory. A common weed of gardens and cultivation. Mediterranean. *Papaver dubium Linn. Long-head Poppy. On a farm near Somerset West. Europe. Two other species of poppy (P. rhoeas Linn and P. hybridum Linn). are found in cornfields in the Malmesbury division and are likely to occur. CRUCIFERAE. Alyssum maritimum (Linn.) Lamk. Sweet Alison. On sand dunes at Strand and occasionally on road sides as a garden escape. Mediter- ranean. *Brassica Tournefortii. Gouan. 3594. Common along the coast from Strand to Gordons Bay, both as a garden weed and on the sand dunes. Mediterranean. Coronopus didymus (Linn) Sm. 3769. Wart Cress. Common as a garden weed and on waste ground. South America, now cosmopolitan. *Diplotaxis muralis (Linn.) DC. 4587. Wall- or Sand-mustard. On sand dunes at Strand. Europe. *Lepidium bonariense Linn. 4673. On sand dunes at Strand. South America. *Matthiola incana. R. Br. Stock. On the sea shore at Strand. A garden escape doubtfully established. Mediterranean. *Raphanus raphanistrum Linn. 3568. Wild Radish. A very common weed. Europe. *Raphanus sativus Linn, 4507. Radish. On roadsides and waste places, occasional. Europe. *Rapistrum rugosum (Linn.) All. 3945. A common ruderal usually with one-seeded fruits. Mediterranean. *Sinapis arvensis Linn. 4508. Charlock. Waste places, not common. Europe. *Sisymbrium officinale Linn. 4275. Hedge Mustard. In cultivation, pasture and on road-sides, frequent. Europe. *Sisymbrium orientale Linn. 4127. A common road-side weed in Gordons Bay. Mediterranean. 164 The Journal of South African Botany. PITTOSPORACEAE. *Pittosporum undulatum Vent. Common along the Lourens River and occasionally found near other streams in moist shady places. Australia. VIOLACEAE. *Viola odorata Linn. Sweet Violet. In moist woods near Stellenbosch. Europe, Asia, N. Africa. CARYOPHYLLACEAE. Cerastium viscosum Linn. 3906. Mouse-eared Chickweed. A common weed of cultivation and gardens. Europe. Polycarpon tetraphyllum Linn. 3555. All-seed. Common on paths and dry disturbed ground. Europe. *Sagina apetala Linn. 3941. Common Pearlwort. A weed of moist places such as seed boxes, flower pots and well watered lawns. Europe. *Sagina procumbens Linn. In similar places to the above. Europe. Silene gallica. Linn. 3898. Windmill Pink. A common weed of cultiva- tion. Europe. *Silene latifolia (Mill.) Brit. and Rendle. Bladder Campion. Jonkershoek, not far from habitation. Europe. Asia. Spergula arvensis Linn. 3894. Spurry. Weed of cultivation on sandy soil, frequent. Europe. Spergularia rubra (Linn.) Presl. 4793. Sand Spurry. On sandy soil at Strand. Not common. Europe. Stellaria media (Linn.) Vill. Chickweed. A very commen garden weed. Europe. “Tunica prolifera (Linn.) Scop. 4268. Kohlrauschia prolifera (Linn.) Kunth. Local and common where it occurs. Strand. Mediterranean. PORTULACACEAE. Portulaca oleracea Linn. 3552. Purslane. A common weed of cultivation. Europe. MALVACEAE. *Lavatera arborea Linn. Tree Mallow. Occasional both on the sea shore and inland on road-sides. Mediterranean. *Lavatera cretica Linn. 4674. Cretan Mallow. Common in weedy places, especially near the sea. Mediterranean. Malva parviflora Linn. 4015. Mallow. A common weed of road-sides and cultivation. Southern Europe, Western Asia. Alien Plants Growing without Cultivation 165 in the Somerset West Neighbourhood. ZYGOPHYLLACEAE. Tribulus terrestris Linn. 4563. Waste places, not common. Known in California as the Puncture Vine as the fruits readily puncture the tyres of bicycles ridden across country on footpaths. Mediterranean, but now cosmopolitan in warm climates. GERANIACEAE. Erodium malachoides (Linn.) Willd. In shady places near the sea. Kogel Bay. Mediterranean. Erodium moschatum L’Herit. 3903. Heron’s Bill. A very common weed of gardens and waste places. A good fodder plant. Mediterranean. Geranium dissectum Linn. 4443. Cut-leaved Crane’s Bill, A common garden weed. Europe. Geranium molle Linn. 4082. Dove’s Foot Crane’s Bill. Shady places at Jonkershoek. Europe. OXALIDACEAE. Oxalis corniculata Linn. A weed of gardens, not very common. Possibly originally from Europe but now cosmopolitan. *Oxalis latifolia H.B.K. A garden weed not well established as yet but imported from time to time with plants from Natal. Mexico. TROPAEOLACEAE. *Tropaeolum majus Linn. Nasturtium. Frequent in moist shady places along roads or under trees. South America. LEGUMINOSAE. *Acacia Baileyana F. v. M. Occasionally self-sown but not aggressive. Australia. *Acacia cyanophylla Lindl. The commonest of the naturalized species, often found in pure stands of considerable extent. Australia. *Acacia cyclops A. Cunn. Common especially near the sea on sand. Australia. *Acacia decurrens Willd. Several forms of this variable species are seen in cultivation one of which var. mollis Lindl. is naturalized. Australia. *Acacia longifolia Willd. Often common along streams. Australia. *Acacia melanoxylon R. Br. Blackwood. In wooded areas having spread by root suckers from planted trees. Australia. *Acacia pycnantha Benth. 3517. Golden Wattle. Found in several places but not aggressive. Australia. 166 The Journal of South African Botany. *Albizzia distachya (Vent.) MacBride 4206. Likes moist ravines where it sometimes grows in dense thickets. A. Lophantha (Willd.) Benth. Australia. *Cytisus monspessulanus Linn. Freely self-sown on a farm on the Helder- berg. Mediterranean. *Lathyrus latifolius Linn. 4356. Perennial Pea. In moist shady places near Stellenbosch. Mediterranean. *Lupinus angustifolius Linn. 3525. Blue Lupin. Common on road-sides and near cultivation. Mediterranean. *Lupinus luteus Linn. Yellow Lupin. In similar places to the above, but not as common. Mediterranean. *Medicago Aschersoniana Urb. 4806. On sand at Strand, not common. N. Africa. Medicago hispida Gaertn. 3767. Medick. A very common weed. Europe. Medicago sativa Linn. Lucerne. An occasional escape from cultivation. S. Europe. Melilotus indicus All. Melilot. A common weed originally European, now cosmopolitan. Trifolium angustifolium Linn. Common and locally abundant on road- sides and waste ground. Europe. *Trifolium campestre Schreb. Hop Clover. Common. Europe. *Trifolium dubium Sibth. 3880. Lesser Yellow Clover. A common weed of lawns, also in pasture land. Europe. *Trifolium glomeratum Linn. 3611. Clustered Clover. In similar places to the above and almost as common. Europe. *Trifolium subterraneum Linn. 4528. In farmland on the Helderberg, spreading into indigenous vegetation. Europe. Vicia atropurpurea Desf. 3600. Purple Vetch. On road-sides and in hedges, common. HKurope. *Vicia hirsuta (Linn) S. F. Gray. 3730. Tiny Vetch. A common garden weed. Europe. Vicia sativa Linn. 3602. Common Vetch. In cultivated ground and pas- ture, sometimes growing in dense masses. Europe. ROSACEAE. *Cydonia oblonga Mill. 3839. Quince. Established in shady places near Somerset West and Stellenbosch. Persia to Turkestan. *Pyracantha angustifolia (Franch.) Schneid. Fire Thorn. In shady woods near Somerset West. China. MYRTACEAE. Eucalyptus. Most of the species of this genus commonly planted will regenerate if the ground nearby is ploughed and left fallow or if Alien Plants Growing without Cultivation 167 in the Somerset West Neighbourhood. the vegetation near seed bearing trees is burnt. Mrs. Levyns in Guide to the Flora of the Cape Peninsula mentions E. globulus Labill. and E, Lehmannii Preiss. as naturalized in the Cape Peninsula. Both these may be found growing spontaneously in Somerset West, but the following have better claims to be considered naturalized: *Eucalyptus cladocalyx F. v. M. Sugar Gum. Often seen growing in dense thickets. S. Australia. *Kucalyptus grandis Hill. E. saligna Auct. Flooded Gum. Prefers moister sites than the Sugar Gum and is found along the upper reaches of the Lourens river and in ravines of the Helderberg. E. Australia. *Eucalyptus occidentalis Endl. Flat-topped Yate. Not much planted but where it has been it regenerates very freely after fire. Often flowers when quite small. W. Australia. *Eugenia Jambos Linn. Rose Apple. Common along a stream at Sir Lowry’s Pass village. In swampy places at Somerset West. Burma to Java. *Leptospermum laevigatum F. v. M. Australian Myrtle. Occasional on sandy soil. Australia. ONAGRACEAE. *Oenothera affinis Cambass. 4390. (O. Berteriana Spach.) Fairly common on road-sides near Faure, between Somerset West and Stellenbosch and near Paarl. 8S. America. Oenothera biennis Linn. Evening Primrose. In fields near Stellenbosch, not common. N. America. *Oenothera indecora Cambass. 4839. A garden weed in Faure and Somer- set West. S. America. *Oenothera rosea Ait. 3539. A common weed in gardens, less frequent outside gardens as it needs more moisture than the other introduced species. Central America. *Oenothera stricta Ledeb. 4849. Occasional, on road-sides. S. America. LYTHRACEAE. Lythrum Hyssopifolium Linn. 3773. A weed of cultivation especially in moist places. Hurope (and possibly indigenous in 8. Africa.) PASSIFLORACEAE. *Passiflora caerulea Linn. 4130. Along a stream in Gordons Bay, common but quite local. S. America. 168 The Journal of South African Botany. UMBELLIFERAE. *Apium leptophyllum (Pers.) F. v. M. 4393. Marsh Parsley. A garden weed especially on well watered lawns. S. America, Australia. Now cosmopolitan. *Conium maculatum Linn. 4678. Hemlock. Near cultivation, Jonkers- hoek. Europe. Daucus carota Linn. 3783. Carrot. Occasional on farm land. Sometimes in heavy grass growth. Europe. Foeniculum vulgare Hill. Fennel. Common on road-sides and waste ground near habitation. Europe. CAPRIFOLIACEAE. *Lonicera japonica Thunb. 3751. Japanese Honeysuckle. In shady woods, sometimes abundant, Somerset West, Stellenbosch. Japan. *Lonicera sempervirens Linn. 3750. Trumpet Honeysuckle. In similar places to the above but requires moister conditions and is not so common. United States. RUBIACEAE. *Sherardia arvensis Linn. 4823. Field Madder. In open woods near the Lourens River. Europe. COMPOSITAE. *Ambrosia artemisiaefolia Linn. Ragweed. Near habitation, Somerset West, not common. N. America. *Anthemis Cotula Linn. 4549. Stinking Chamomile. A garden weed, not common. Europe. *Aster exilis Ell. 3777. A garden weed. Seems to be a recent introduction and is spreading. Southern United States. Bidens pilosa Linn. 3788. Hairy Bur Marigold. A weed of fairly moist places. Tropical and sub-tropical America, now cosmopolitan. Centaurea melitensis Linn. 3622. Malta Thistle. A weed of waste ground and cultivation. Mediterranean. Chrysanthemum carinatum Schousb. 4542. Marigold. On road-sides, sometimes growing in dense patches. Morocco. Erigeron canadense Linn. Canadian Fleabane, Horse weed. Occasional, in moister places than the following species. N. America. Erigeron linifolium Willd. 3549. Flax-leaved Flea-bane. A very common summer weed. Conyza ambigua DC. Vide Kew Bull. Misc. (1921), p. 282. Probably a native of the East Mediterranean and West Asia, now cosmopolitan. *Erigeron mucronatum DC. A common garden plant, sometimes found as a weed in lawns. Mexico. Alien Plants Growing without Cultivation 169 in the Somerset West Neighbourhood. *Galinsoga parviflora Cav. A weed of moist places especially gardens and irrigated cultivation. S. America. Gnaphalium luteo-album Linn. Jersey Cudweed. Fairly common in waste places. A cosmopolitan weed of uncertain origin. *Helianthus annuus Linn. Common Sun Flower. An occasional garden escape. N. America. *Helminthia echioides (Linn.) Gaertn. 3628. Bristly Ox Tongue. Appears to be a recent introduction, which has spread rapidly in gardens and waste places. Picris echioides Linn. 8. Europe. *Inula graveolens (Linn.) Desf. 3492. Khaki weed. A very common sum- mer weed apparently of recent introduction. When the coast road from Steenbras to Klemmond was constructed, this plant at once took possession of the loose soil on either side of the road. S. Europe. *Lactuca scariola Linn. 3778. Prickly Lettuce. Not common and only the form with lobed leaves seen. S. Europe. *Schkuhria pinnata (Lamk.) Thell. A common weed on a farm at Welling- ton. §. bonariensis. Hook and Arn. S. America. Senecio vulgaris Linn. Groundsel. On vacant building plots and as a garden weed in Somerset West. Europe. *Silybum Marianum Gaertn. Milk Thistle. Abundant in Sir Lowry’s Pass village growing in nearly pure stands. Occasionally seen elsewhere. Mediterranean. Sonchus asper Hill. 3771. Prickly Sow Thistle. In similar and usually moister places to the following but not nearly as common. Europe. Sonchus oleraceus Linn. Sow Thistle. An abundant weed of garden and cultivation. Europe. *Tagetes minuta Linn. Khaki weed. A summer weed of moist places. Not common. 8. America. *Tragopogon porrifolius Linn. Salsify. Road-sides in Somerset West. Apt to be a troublesome weed on lawns as it is deeprooted. S. Europe. Urospermum picroides (Linn.) F. W. Schmidt. A very common weed. S. Europe. *Xanthium orientale Linn. f. 4169. Cocklebur. Along the railway at Firgrove and elsewhere. Probably originally S. American, but now widely distributed as a weed. ERICACEAE. *Arbutus Unedo. Linn. Strawberry Tree. Planted along a stream in Parel Vallei, Somerset West, and natural seedlings frequently appear in moist shady places. The growth of such seedlings is very slow and it 170 The Journal of South African Botany. is often several years before they reach a height of one foot. S. W. Europe. PLUMBAGINACEAE. *Limonium sinuatum (Linn.) Mill. 4816. On the sea front at the Strand, not common. Mediterranean. PRIMULACEAE. Anagallis arvensis Linn. Pimpernel. A very common weed. Flowers red or blue. Europe. APOCYNACEAE. *Vinca major Linn. Periwinkle. Sometimes abundant in moist shady places. Appears to spread by vegetative means and not by seed. S. Europe. ASCLEPIADACEAE. *Araujia sericifera Brot. 4396. In woods on either side of the Lourens river. Common in a hedge near Stellenbosch. Appears to be spreading. S. America. BORAGINACEAE. Echium violaceum Linn. 4376. Vipers Bugloss. Common and often abun- dant on road-sides and waste places. S. Europe. CONVOLVULACEAE. Convolvulus arvensis Linn. 3789. Bindweed. A weed of road-sides, hedges and cultivation. Abundant in the Strand. Europe. *Ipomoea Learii Paxt. Blue Dawn Flower. Occasional along streams in Stellenbosch and Somerset West. Tropical and sub-tropical America. SOLANACEAE. Datura Stramonium Linn. Thorn Apple. A common weed of gardens and cultivation. Probably indigenous to 8. Europe, but now cos- mopolitan. Nicotiana glauca Grah. Tree Tobacco. Occasional on road-sides. S. America. Solanum pseudocapsicum Linn. Winter Cherry. In shady woods, Somer- set West. Possibly indigenous to Madeira, now cosmopolitan. Lycopersicum esculentum Mill. Tomato and Physalis peruviana Linn. Cape Gooseberry are occasionally seen in rubbishy places near gardens. Alien Plants Growing without Cultivation 171 in the Somerset West Neighbourhood. SCROPHULARIACEAE. Linaria spuria Mill. 3776. Rough-leaved Toadflax. A very common garden weed. S. Europe. *Mimulus moschatus Doug]. 4036. On damp sand in the bed of the Lourens river. N. America. Verbascum virgatum Stokes 4482. Mullein. Jonkershoek. Mediterranean. BIGNONIACEAE. *Campsis radicans Seem. 3639. Trumpet Creeper. A very troublesome weed when established on heavy clay soil. The roots are deep and run horizontally some two feet below the surface. Any pieces of root left in the soil grow again. The plant spreads rapidly by root suckers and. by the long shoots which in the absence of support lie upon the ground and root readily at the nodes. It can be kept in check by heavy grazing. Somerset West, locally abundant. Southern United States. ACANTHACEAE. Thunbergia alata Boj. Black-eyed Susan. In open woods near the Lourens river. Tropical Africa, Natal. MYOPORACEAE. *Myoporum serratum R. Br. 3902. Manitoka. Occasionally self-sown and spreads to some extent by root suckers. Australia. VERBENACEAE. Lippia reptans H. B. K. Occurs occasionally as a lawn weed. Tropical America. Verbena bonariensis Linn. 4037. Road-side ditches, frequent. S. America. Verbena officinalis Linn. Vervain. A weed of gardens and waste places. Europe. LABIATAE. *Mentha Requienii Benth. Abundant as a lawn weed in a garden at Hels- hoogte. Corsica. Stachys arvensis Linn. 4450. Field Woundwort. A garden weed. Europe. PLANTAGINACEAE. *Plantago Coronopus Linn. 4367. Crow’s Foot Plantain. Common near the sea at Strand. S. Europe. Plantago lanceolata Linn. Ribwort. Common on road-sides and waste ground. Europe. 172 The Journal of South African Botany. AMARANTHACEAE. *Amaranthus deflexus Linn. 3793. A fairly common weed in Somerset West. S. America. *Amaranthus hybridus Linn. 3470. A weed of cultivation, common in places. Tropical America. CHENOPODIACEAE. *Atriplex hortensis Linn. 4711. Orach. The red form, an escape from gardens, appeared in some abundance on the sea front at Strand but was not seen the following year. Asia. *Atriplex Muelleri Benth. 3651. Common on road sides at Strand, less frequent inland. Australia. *Atriplex semibaccata R. Br. 3558. Salt Bush. On road sides at Strand. Australia. *Boussingaultia basselloides H. B. K. In moist shady places, Somerset West. S. America. Chenopodium album Linn. 3569. Goosefoot. A common weed from Europe, now cosmopolitan. Chenopodium ambrosioides Linn. 3557. Mexican Tea. Waste places, frequent. Tropical America. Chenopodium murale Linn. Nettle-leaf Goose-foot. A common weed of cultivation and waste places. Europe. Roubieva multifida (Linn.) Mog. 3775. Common on waste land. S. America. *Salsola kali Linn. 3462. Russian Thistle. Common along the sea shore at Strand and apt to be troublesome to bathers as it is very prickly when old and dry. E. Europe. PHYTOLACCACEAE. Phytolacca americana Linn. 4028. Pokeweed. Near the Lourens river at Vergelegen, not common. N. America. POLYGONACEAE. Polygonum aviculare Linn. 3784. Knotweed. A summer weed, common. Europe. Polygonum Convolvulus Linn. 3782. Black Bindweed. An occasional garden weed. Europe. Rumex Acetosella Linn. 3785. Sheep Sorrel. Often abundant on poor sandy soil. Europe. Rumex crispus Linn. 3761. Curled Dock. A weed in vineyards, not common, Somerset West. Europe. Rumex puleher Linn. 3760. Fiddle Dock. A common and troublesome garden weed. S. W. Europe. iN) Alien Plants Growing without Cultivation 1 in the Somerset West Neighbourhood. LAURACEAE. *Cinnamomum Camphora Nees & Eberm. Camphor Tree. Grows in dense thickets on low lying ground near the Lourens river at Vergelegen. China and Japan. PROTEACEAE. *Hakea acicularis Knight. 4084. Scattered along the Helderberg and Hottentots Holland mountains. Liable to spread rapidly after fire. Repeated burning will not control it as one year old seedlings can flower and ripen seed. Australia. *Hakea suaveolens R. Br. Not common and not aggressive. Australia. EUPHORBIACEAE. Euphorbia Helioscopia Linn. Sun Spurge. In waste places, not common, Stellenbosch. Europe. Euphorbia hypericifolia Linn. 4846. A weed on farm land between Sir Lowry’s Pass and Gordons Bay. Tropics. *Kuphorbia maculata Linn. 3790. Spotted Spurge. A weed of garden paths, flower pots and road sides. N. America. Euphorbia Peplus Linn. 3774. Petty Spurge. A common weed of gardens and shady places. Europe. *Euphorbia serpens H. B. K. 3792. In similar places to, and often growing with E. maculata. Tropical America. *Kuphorbia terracina Linn. 4850. On the sea shore at Strand. Mediter- ranean. Ricinus communis Linn. Castor Oil Plant. Occasional on road sides and waste places. Tropical Africa, but now cosmopolitan. MORACEAE. *Fiecus Carica Linn. Fig. Occasional in woods along the Lourens river. Asia Minor. URTICACEAE. *Helxine Soleirolii Req. Mother of Thousands. An abundant weed in a garden at Helshoogte. Corsica. Urtica urens Linn. 3805. Stinging Nettie. Weed of rubbish heaps near habitation. Europe. FAGACEAE. *Quercus robur Linn. Oak. Frequently self-sown in moist places and along streams. Hurope. 174 The Journal of South African Botany. SALICACEAE. *Populus canescens Sm. 3823. Gray Poplar. Only the male tree has been introduced. It spreads readily by root suckers and is often very common along streams. Europe. PINACEAE. *Pinus pinaster Ait. Cluster Pine. Common on mountain slopes and spreads readily by wind-borne seeds. Mediterranean. *Pinus pinea Linn. Stone Pine. Common on flats and lower slopes Regenerates freely but only near seeding trees as the seeds are not wind-borne. Mediterranean. *Pinus radiata D. Don. Monterey Pine, Insignis Pine. Regenerates in moist ravines and on river banks. Pinus insignis Dougl. California. LILIACEAE. *Allium triquetrum Linn. 4091. On a road-side near Somerset West. Mediterranean. COMMELINACEAE. *Tradescantia fluminensis Vell. Wandering Jew. Forms the main ground cover in a poplar wood at Somerset West. S. America. JUNCACEAE. Juncus bufonius Linn. Toad Rush. A weed of wet places. Widely spread in the Northern hemisphere and probably introduced in South Africa. CYPERACEAE. Cyperus esculentus Linn. 4168. A garden weed of wet places. Both this and the following species are pests difficult to eradicate. Introduced from the north. Cyperus rotundus Linn. Nut Grass. In similar places to the above and much commoner. Introduced from the north. GRAMINEAE. Agrostis verticillata Vill. 3529. Water Bent. Common along streams, irrigation channels, etc. Mediterranean. Aira caryophyllea Linn. 4448. Hair-grass. A garden weed in Somerset West. Mediterranean and mountains of tropical Africa. Anthoxanthum odoratum Linn. 4803. Sweet Vernal-grass. In moist places, Jonkershoek. Europe. Avena barbata Brot. 4225. Slender Oat. Common on road-sides and waste places. Mediterranean. Avena fatua Linn. 4254. Wild Oat. In similar places to the above. Some- times abundant as a weed of cultivation. Europe. Alien Plants Growing without Cultivation 175 in the Somerset West Neighbourhood. Avena sativa Linn. 4246. Oat. Occasional on farm land. Europe. Brachypodium distachyum (Linn.) Beauv. 3612. Sand dunes at Strand and a garden weed in Somerset West. Mediterranean. Briza maxima Linn. Large Totter Grass. Common in waste places and as a garden weed. Mediterranean. Briza minor. Linn. 3918. Small Totter Grass. In similar places to the above. Mediterranean. Bromus Gussonii Parl. A weed of waste ground. Mediterranean. Bromus japonicus Thunb. 4918. In pasture land and waste places; common. Mediterranean. *Bromus madritensis Linn. 4825. In weedy places, Somerset West, not common. Mediterranean. Bromus molliformis Lloyd. A garden weed. Mediterranean. Bromus unioloides H. B. K. 3766. Common in waste places and as a garden weed. S. America. Chloris Gayana Kunth. Rhodes Grass. Occasional on road-sides. Tropical Africa. Dactylis glomerata Linn. 4835. Cock’s Foot. In moist shady places, not common. Europe. Digitaria sanguinalis (Linn.) Scop. 3770. Crab Grass. A common summer weed in gardens. Mediterranean. Echinochloa crus-galli (Linn.) Beauv. 3786. In moist places with rich soil, occasional. Europe. Eleusine indica (Linn.) Gaertn. Crow’s Foot Grass. A common summer weed in the Strand. Introduced from the north. *Eragrostis abyssinica (Jacq.) Link. Teff. On farm land, not common. N. E. Africa. Eragrostis cilianensis (All.) Link ex Lutati. 4031. Dry sandy soil, not common. E. major Host. Mediterranean to India. *Gastridium ventricosum (Gouan). Schinz & Thell. 3961. Nit Grass. A grass appearing late in spring or early summer. Stellenbosch, Somerset West. Mediterranean. Hordeum murinum Linn. 3942. Wild Barley. Common on sandy soil, especially near the sea. Europe. Koeleria phleoides (Vill.) Pers. 4274. Frequent on road-sides and waste ground. Mediterranean. Lagurus ovatus Linn. 4297. Hare’s Tail Grass. Common along a foot- path extending far into indigenous vegetation at Rooi Els. Mediter- ranean. Lolium multiflorum Lamk. Italian Rye Grass. Common on road-sides and in pasture. Europe. 176 The Journal of South African Botany. Lolium rigidum Gaudin. 3535. On sand dunes at Strand. Mediterranean. Lolium temulentum Linn. Darnel. Common in waste places and as a garden weed. Europe. Oryzopsis miliacea (Linn.) Aschers. & Schweinf. 4397. Rice Grass. Near the Eerste River in Stellenbosch. Mediterranean. *Paspalum dilatatum Poir. 3765. Common in fairly moist places. S. America. Paspalum distichum Linn. 3625. P. digitaria Poir. Growing in water or very wet soil, Somerset West. America. *Pennisetum clandestinum Hochst. Kikuyu Grass. Extensively planted as a fodder grass and for lawns. Spreads readily by vegetative means, but no seedlings seen. East Africa. *Pennisetum setaceum Choir. 4579. P. Ruppellii Steud. Fountain Grass. On the road sides, not common. Somerset West. Abyssinia. Pennisetum unisetum Benth. 4594. In pasture land in moist places. Not common. Introduced from the north. Phalaris minor Retz 3759. Mediterranean Canary Grass. Common on vacant building plots in Somerset West. Mediterranean. Poa annua Linn. 3570. Winter Grass. Very common on road sides and as a garden weed. Europe. now cosmopolitan. *Poa pratensis Linn. Meadow Grass, Kentucky Blue Grass. Sometimes used for lawns and occasionally persisting as a garden weed. Europe. Polypogon maritimus Willd. 4960. On the sea shore at Strand. Europe. Polypogon monspeliensis (Linn). Desf. 4696. Fairly common in damp places. Possibly indigenous, but more probably introduced from Europe. Polypogon tenuis Brongn. 4386. In similar places to the above. but not so frequent. Europe. Scleropoa rigida (Linn.) Griesb. 4548. A weed of farm lands and gardens, not common. Mediterranean. Setaria verticillata Beauv. 4829. A weed in a nursery garden at Strand. Introduced from the north. ; Vulpia bromoides (Linn.) S. F. Gray. 4826. Frequent, often a weed in gardens. Mediterranean. Vulpia myuros (Linn.) Gmel. 4804. In similar places to the above but not as common. Mediterranean. ADDENDA ET CORRIGENDA TO THE WOODY PLANTS OF THE BECHUANALAND PROTECT- ORATE. (Vol. XVIII pp. 1—100, 1952.) By O. B. Miter. Besides some additional localities and native names, 16 fresh species are now recorded, bringing the total, exclusive of varieties, to 580. For the identification of these new plants and for certain corrections to previous determinations I am indebted to the Chief, Division of Botany & Plant Pathology, Union of South Africa and particularly to Miss Rosemary Robertson of the National Herbarium, Pretoria; to the Director of the Royal Botanic Gardens, Kew, and Mr. E. Milne-Redhead, and to the Director of the Imperial Forestry Institute, University of Oxford and Mr. F. White. Preface; last sentence: after ““Mr. F. White.’’, add “Also to Mr. H. L. Dunkley for corrections made at Oxford and Kew.” p. 8, 1. 3, for “Rhus” read “Combretum.” p. 10, 1. 29, delete “F. sp.” and substitute “F. capraeifolia Del.” p. 11, 1. 15, for “464” read “164”. p- 13, 1. 6, for “leucrura”’ read “‘leucura’’. pli owdeletesssp=mrar Aes p. 16, |. 7, for “cfr. C. oligantha Gilg. & Benn.” read “nr. elaeagnoides Gilg.”’. Pa l6s lines Ié215 for Benn, read) Bent . 17, 1. 19, after “in N.B.P.” insert “An evergreen which however loses many of its leaves in winter. Flrs. (Sept.-Oct.) pale yellow. Frts. Aug.-Sept.” - 19, 1. 29; for “Burtt Davy read “Glover”. 21, 1. 19, between “mosu” and “‘(Mbuk.)” insert “and muthu’’. . 35, after ““(Mbuk.)” add “mbola (Kalaka)”’. . 21, after “(teste Curson)”’ add “mothwakedja (Taw.) . 30, for “‘fassogiensis Klotz.” read “‘fassoglensis Kotsch.”’. 28, |. 32, for ““‘macracantha”’ read “macrantha’’. 30, l. 4, for “COLOPHOSPERMA” read “COLOPHOSPERMUM”’. . 30, 1. 16, insert “‘C. flavicarinata Bak. f. Lax shrub of Baikiaea forest, Serondela. Base only of stem is woody. Flrs. (Dec.) white and yell. Miller B/1261.” ue} oe) ” fel el) hic) (eo) Is bo ie 2) PT 178 The Journal of South African Botany. p. 30, after line 22 insert “C. sp. Lowly woody plant on termite nest nr. Jolley’s Pan, dist. Chobe. Firs (May) yell. Miller B./1327.” p. 30, 1. 32, after “(Mbuk.)’”? add “mogamane (Taw.)”. p- 30, 1. 33, after “Frt 1-seeded” add “Firs. (June) whitish.”’. p- 30, 1. 34, after “B/645” add “B/1332”. p. 31, after 1. 19, insert “DOLICHOS L. II L. D. sp: Small woody climber. Flrs. (May) pale blue to white with distinctly blue keel. Lys. sparse, 3-foliolate. Nr. Deka. Miller B/1325.” page 32, 1. 20, after “J. Leonard” add “II”. p. 33, 1. 8, for “Bek.” read “Bak.” p. 33, 1. 9, after “3-foliolate” insert “A procumbent woody plant. Flrs. (Aug.) pink turning pale blue.” p.33, 1. 10, after “Mrs. Lugard 190” insert “Miller B/1336”. p. 33, delete lines 20 and 21, and substitute “Julbernardia (syn. Isoberlina Craib and Stapf) 1. J. globiflora’”’. p. 34, 1. 4, for “(Sept.)” read “(Aug.-Sept.)” p. 34, 1. 5, after “‘tree-like” add “but very occasionally 3-foliolate, the terminal lft much the largest’. p. 34, after 1. 10, insert “MELOLOBIUM E. & Z. III 1. M. sp. — makgonatsotlhe (Kal.) Spiny shrublet 6 to 8 ins. high. Lys. 3-foliolate. Flrs. (Sept.) yell. Bed. of Kuwe Pan, 8.W. Kalahari. Miller B/355.” p. 37, delete lines 20 & 21 and substitute “Citropsis Swingle & Kellermann 1. C. sp. nov. (Kew det.)”. p. 37, l. 28, after “hut pole” insert “Frts (Apl-May) globose, 12 mm. diam.” and after “B/1110” insert “B/1172”. p. 38, |. 6, for “betchuanica” read ““‘betschuanica”. p. 38, 1. 37, delete “Ngamiland, Curson 117, 792.” and insert ““mkomoto (Mang.). Tree 25 ft. high. Frts. Feby. Bark not papery but flaking green. Lys. pinnate. Tati and Ngamiland. Curson 117, 792. Miller ‘B/MBIg p- 39, after 1. 19, insert “39. C. sp. (possibly C. parvifolium Dinter). Lax shrub. Inflor. (Nov.) capitate, very small heads, pale yell. Serondela. Miller B/1199.” p. 40, delete lines 37 to 39. p. 41, after |. 9; insert “2: T. sp: segelobe (Ngwak.) ~ ke} Ino} Ine} Ine) . 49, after 1. 29 insert “‘simammungwa (Kol.) . 49, 1. 30, delete and substitute “Liane 30 to 40 ft. long, 6 to 7 ins. Addenda et Corrigenda to the Woody Plants 179 of the Bechuanaland Protectorate. Not matched at Kew. Lax shrub with twining brchlts. Lvs. opp. or whorled, minutely hairy. Flrs. (Jany.) canary yell. Hab. among rocks. Miller B/978.” . 44, after 1. 21, insert ““E. matabelensis Pax Shrub to 6 ft. high. Stem not fleshy. Juice milky, copious. Lys. opp. or whorled. Flrs. (May-July) precocious, yell.-green. Brchlts. in 3s. equidistant and in same plane. Bark pale, smooth but corky at base of stem. Lys. linear. Chobe dist., common round Deka, rare elsewhere. Miller B/1320, B/1329, B/1330.” . 44, after 1. 31 insert “maladitapi (Taw.)”’. . 45, after 1. 3, insert “3. P. sp. (possibly P. loandensis Welw., ex descr.) Woody plant to | ft. high. Firs. (Aug.) axillary, minute. Frts. (Aug.) globose, opening by 3 slits. Bark red-brown, stringy. Lvs. sparse or wanting. Hab. destroyed Baikiaea forest, Serondela. Miller B/1335.” 47, AS for ‘guenzu’ read “‘guemzii. . 47, after 1. 11 insert “R. lucens Hutchins Small tree. Frts. (May) flattened. Deka area, dist. Chobe. Miller B/1328.”’. ” diam. Flrl. (Sept.), green, disc blue-green. Male firs. have brick- red anthers. Serondela. Miller B/1334. Rob. & Ellfers 82.”. . 49, 1. 35, for “parvifolia” read “parviflora N. E. Br.”’. . 51, |. 16, for “Zizyphus” read “Ziziphus”. . 52, 1. 3, between “‘precocious” and “‘Matetsi” insert ““Brchlts. ten- drilled. Nr Topsi Siding and” . 53, |. 16, after ““Curson” add “‘ntewa (Kalaka)”’. . 54, 1. 23, delete ““B/476” and transfer to Grewia retinervis. . 54, delete lines 24 and 25, transfer “‘Miller B/538” to Grewia flavescens. . 54, after 1. 28, insert “ Triumfetta L. ]. T. pentandra A. Rich. Slender shrub 3 ft. high. Firs. (Apl.), calyx green, corolla yell. Frts. (Apl.) ovate or globose, echinate, 3 mm. diam. Bark strong, stripping readily. Lvs. simple. The whole plant hairy. Serondela. Miller B/1167, B/1314.”. . 54, after 1. 37 (bottom of page) insert “A. ramosum G. & P. Shrub 4 ft. high. Lys. sparse, whorled. Flrs. (July) yell. Frts. 6-locular, roughly hairy. Serondela. Miller B/1334.” 55, 1. 8, after “its nest.” insert “This is denied by ornithologists who say the bird is parasitic.” 180 The Journal of South African Botany. p- Ko) Sit Bd td 55, delete lines 21 to 23 and substitute ““Thin-stemmed plant to 3$ ft., only lower part woody. Whole plant with small but sharp prickles. Firs. (June-Aug.) ‘yell. with maroon eye’, Mrs. Lugard. Bracts 25 mm. x 1.5 mm. Lys. and stipules digitate. Chobe and Ngamiland. Mrs. LIugard 118. Miller B.1078, B/1079, B/1313. Rob. & Ellfers 79.”. . 56, |. 16, for “H. dinteriana”’. read ““S. dinteriana’’. . 57, 1. 15, after “Melhania” insert “M. burchellii DC. Recorded by Burtt Davy (4). Zeyher 2417.” . 58, |. 26, after “edible” insert “Firs. (Aug.) cauliflorous; male firs. o green with yell. stamens.” after “Miller B/68.” add “B/1338.” . 59, 1. 26, after “yell.” insert ““Frts. (June) + globose, 2.5 cm. diam. Seeds bright yell. with honeycomb surface.”’. . 59. after 1. 32. insert “Oncoba Forsk. 1. O. spinosa Forsk. Small branchy tree with spinose brchlts. “Flrs. about 4—d cm. across, showy white, fragrant’ (4). Frts. (May) globose, 4—5 cm. diam. shell hard with dark longitudinal markings. Lys. serrulate. Baikiaea forest. Serondela. Miller B/1317.”’. . 60, after line 19, insert LYTHRACEAE Nesaea Comm. 1. N. sp. Shrublet 18 ins high. Lys. decussate or verticillate. Flrs. (March) reddish, stamens yell. Frts. (March) black with persistent calyx. Pharing. Miller B/496. . 60, 1. 38, bottom of page, after ““Ngamiland” insert “Flrs. (Aug.) + precocious, pale yell.” 61, after line 4 insert “moswapeba (Taw.)”’. 61, 1. 22, C. holosericium is a syn. of C. gueinzii Sond. 62, 1. 33, after “whorls of 3” insert “(21)”. 62, 1. 35, delete ““(21)’’. 63, 1. 4, after “(Mbuk.)”’ add ““mswaswe (Kalaka)”. . 63, after line 14, insert ““C. triacanthum Fresn. mfula (Rozi). Tree 12 ft. high in Baikiaea forest, Serondela. Lys. 3 em. x 1.5 cm., margins distinctly, midrib slightly hairy. Miller B/1318.” . 64, 1. 31, after “N. B. P.” insert “Flrs. (Jany.) white. giving the tree a hawthorn-like appearance.” . 64, 1. 33, delete ~“B/127” and transfer to Terminalia randii. . 64. 1. 36, delete ““B/411” and transfer to Swartzia madagascariensis on p. 36. . 67, delete 1. 2 and substitute “2. M. kirkii Bak. (Kew det.)’’. . 67, l. 5, delete “B/1161” and substitute “B/1176.” Pp. Pp. . 69, delete 1. 20 and substitute “1. S. trichoclada Welw. (syn. S. buchananii p Addenda et Corrigenda to the Woody Plants 181 of the Bechuanaland Protectorate. 68, 1. 18, after “(G.S.P.)” add “ntshekesane (Kalaka)”’. 68, 1. 19, delete “B/149” and transfer to Euclea divinorum. Bak.)”. p. 70, 1. 18, after “muyimbili (Kol.)” insert “gamgcwana (Sarwa). Tree Pp. 14 ft. high. Frts. (June) 10 cm. diam., said to be neither poisonous nor edible. S. of Deka, dist. Chobe. Miller B/1331.”. . 71, lL. 16, after “Frts.” insert ““when dry and opened”’. . 74, lines 31, 32, after ““Kwebe Hills” insert “and Tati dist. Flrs. Jany.” After “Mrs. Lugard 98,” insert “Miller B/1146.”. . 76, 1. 3, after “Miller B/3” add “Miller B/1319, taken from coppice growth, has 3-foliolate lvs. Frts. (May) 1.5 to 2 cm. diam. and decussate brchlts.”’. . 78, after 1. 14 insert “6. S. sp. Unarmed shrub 2 to 3 ft. high. Flrs. (Nov.) greenish, incon- spicuous. Serondela. Miller B/1197. Ts Se De Shrub 2 to 3 ft. high. Flrs. (Aug.) blue with bright yell. stamens. Miller B/1337.” . 80, 1. 29, after “van Son H28685” add Miller B/1342 identified as B. eenii is herbaceous. Flrs. (July-Aug.) dark yell.”. . 85, after 1. 25, insert “V. randii S. Moore. ngone (Kalaka). Tree 18 ft high. Bark pale, smooth. Frts. (Jany.) white, globose, edible. Tati dist. Miller B/1290.”. . 85, 1. 28, after “Miller B/98” add “B/1206.”. . 86, 1. 26, for “‘(sub.)” read ‘“‘(Sub.)’’. . 87, 1. 15, after “white balls.” insert “Specimens of this sp. gathered in S. B. P. have a hairy receptacle resembling a pappus.”’. . 87, |. 25, after “(21)” insert “Shrub 3 ft. high. Flrs. (May) in heads, white. Nr Cream of Tartar Pan, dist. Chobe (Miller). and after “Lugard 249” insert “Miller B/1321.” 90, in glossary, after “stipitate”’ insert ““Sub.; language of the Masubia’’. pp. 91 to 97, place the following additions and corrections alphabetically: banyo. Brdens schimperi, “Black Jack’. under “‘chitamatazua”’ for ““Mimusops sp.*” read “Mimusops kirkit*’’. gamgewana. Strychnos spinosa* litiela. Tacazzea kirkii* maladitapi. Fluggea virosa* mbola. Albizzia struthiofolia* mogamane. Dialiwm simii* 182 The Journal of South African Botany. mohalahalane. Dicliptera micranthus* mokaoke. Schrebera trichoclada* mokauka. a mokomoto. (Commiphora edulis*) and C. mollis* mophani. for “Colophosperma”’ read ““Colophospermum”’. morutse. for “plan” read “‘plant’’. mositsane. to “Elephantorrhiza spp.” add asterisk. moswapeba. to “Combretum spp.” add asterisk. for “mothwakeya” read ““mothwakedja’”’. motlhakutsane. for ““angloensis” read “‘angolensis”’. msusu. Terminalia sp*. mswaswe. Combretum transvaalense*. mfulu. Combretum triacanthum* mumu. to “Ficus spp.” add asterisk. muphane. for “Colophosperma” read “Colophospermum” muthu. Acacia giraffae* mutswara tshu kuduh=—hartebeest’s horns. Gardenia resiniflua* ngone. Vangueria randii* ntshekesane. Huclea undulata* ntewa. Grewia monticola* nzikwa dombo. Canthium sp.* segelobe. after ““Bulbostylis coleotriche” add “‘and Triaspis sp.*” simamamungwa. Hippocratea nitida* tedu tsa banna. delete “Triumfetta pentandra” and substitute “Centamopsis biflora” tsaudi. for “Guibortia” read ““Guibourtia’’. tulwa. Solanum spp.* tulwatulwani. do. Index, pp. 98—100. Insert the following corrections and additions: ANNONACEAE for ANONACEAE. Bolusanthus, p. 29 for p. 27. Citropsis Swingle & Kellermann p. 37. Colophospermum for Colosphosperma. Dicliptera Juss. p. $1. Dolichos L. p. 31. Julbernardia, p. 33. LYTHRACEAE. p. 60. Manilkara, p. 66 for p. 67 Nesaea. p. 60. Oncoba Forsk. p. 59. Triumfetta L. p. 54. Ziziphus. p. 51. BOOK REVIEWS. THE GEOGRAPHY OF THE FLOWERING PLants. By Ronald Good. (London: Longmans Green & Co.) Second Edition, 1953, xiv + 452, with 75 line drawings, 9 maps in colour, 16 photogravure plates. Geographical Distribution is perhaps the most complicated branch of biological science: it is certainly one of the most enthralling. To follow the migrations of plants, each species with its own individual potentiali- ties, each species undergoing evolutionary changes and ramifications through all the changes of earth’s climate and terrain, some gradual, some almost catastrophic, when almost the only facts available are the end-terms of the whole age-long process, and when there are many gaps in our knowledge of these and endless differences of opinion as to their status and relationships, this is surely a task requiring a vast store of information and the ability and imagination needed to estimate the weight and effect of the many variable factors involved. To this task Professor Good has applied himself in masterly fashion, and his book, now available in a new and revised edition, is to-day certainly the most adequate, balanced and stimulating presentation of the facts and factors of the distribution of flowering plants over the globe. Detailed criticism of such a book is impossible, but one or two aspects of Professor Good’s treatment of the subject may be noted. The first half of his book presents the facts to be recorded and, as it were, faced— their complexity, their apparent anomalies and freakishness: one’s com- prehension of this is assisted by maps of outstanding instances of distri- bution of species and groups, these providing the most valuable pointers to past geographical changes and floristic history. The second half of the book deals with the various factors which have come into play in the production of to-day’s distribution and which may be expected to continue into the indefinite future. “One of the most perplexing things about Angiosperm geography is its inherent chaos” says Professor Good; “‘which expresses itself in the difficulty of accounting satisfactorily for any appreciable portion of the total facts by any one single theoretical explanation of the more detailed sort.” It is interesting to note that the author, in contrast to much— though not all—geological opinion, throws a good deal of weight on con- tinental displacement as a factor. Indeed much of the plant-geographical work of the future will no doubt be concerned with a study of the problems of the “tolerance” and rapidity of migration of the flowering plants in relation to the gradual severing of their continental areas: and of the time problem as to the epoch of origin, primary evolution and distribution of 183 184 The Journal of South African Botany. the more primitive groups in relation to the period and rapidity of conti- nental split. It is pleasant for botanists in the southern hemisphere to note that Professor Good has no narrow northern outlook in his attack on distri- butional problems. Although plant distribution in the northern hemi- sphere has been subject to violent catastrophes in the form of recent glacial periods, which have brought about great changes from which, as Professor Good says, they are clearly recovering at the present day, yet the distribution of plants in the southern hemisphere, with its widely spaced land masses, is the result of far more fundamental world-factors than those caused by the recent ice-encroachments of the north. While Professor Good devotes chapters to plant distribution in that botanically well known country Great Britain, and even to the study of one particular county, Dorset, this serves the purpose of illustrating the vastness of the whole field. In his treatment of the wider problems he shows a breadth and range which are eminently satisfactory. The book is illustrated by several admirable plates, among which it is interesting to notice a reproduction of Schimper’s magnificent classical photograph of Java forest vegetation, and also Mrs. Levyns’ photograph of Karoo succulents first published in the Journal of the Botanical Society of 8. Africa: there are also photographs of Protea cynaroides at the Cape, Central African Lobelias and Senecios, a Transvaal EKuphorbia—in fact Africa is well treated in this respect. The numerous line drawings, maps and diagrams are an important and illuminating feature of the book. Professor Good’s book can be warmly recommended for its broad and at the same time detailed presentation of a stupendous and fascinating subject, “that almost keystone of the laws of creation” as Charles Darwin called it. R. H. Compton. PHYSIOLOGY OF SEEDS by W. Crocker and L. V. Barton. (Waltham, Mass.: The Chronica Botanica Co.; London, W.C.2: Wm. Dawson and Sons, Ltd.) Price, $6.50. The study of seeds and seedlings has a great deal of fascination and enters into a wide range of topics, even when the field is restricted, as with the book under review, to seed physiology, which necessarily includes seedling physiology. Such a book may be expected to appeal to a large number of botanists, including those primarily concerned with horticulture and agriculture. Physiology of Seeds has been written by two investigators who have carried out a considerable amount of research on seeds and seedlings at the Boyce Thompson Institute for Plant Research. Beacause of the sub- Book Reviews. 185 ject and the standing of the two authors the reviewer looked forward with much pleasure to the reading of this book. It must be admitted that taken as a whole the book was not of the quality which was anticipated. In places the style is somewhat clumsy and lacks lucidity, and the English is sometimes bad. Parts of the book make heavy reading, and it is difficult to avoid the impression that in dealing with certain topics the authors have not fully digested their material, and hence their treatment of parts of the subject assumes rather the character of a compilation. This applies especially to the earlier chapters. Additional diagrams would also have been advantageous. A glance at the contents of the book indicates the wide range of topics considered. These include a brief account of certain structural features (Chap. 1); factors relating to seed germination (Chap. 2); chemical composi- tion of seeds (Chaps. 3 and 4); permeability of seed membranes (Chap. 5); respiration of seeds (Chap. 6); factors affecting germination (Chaps. 7 and 8); dormancy in seeds (Chaps. 9 and 10); storage and life span of seeds (Chap. 11); chemical changes during seed development, after- ripening and vernalization, and chemical metabolic changes during ger- mination (Chaps. 12 and 13); the enzymology of seeds and germination (Chap. 14); vernalization and embryo culture (Chaps. 15 and 16); transmis- sion and control of seed borne diseases (Chap. 17). Wide as is the range of topics, it does not cover the whole field. Of this the authors are aware, and they call attention in the preface to one such omission, seed dispersal —a subject which is usually regarded as outside the field of physiology. Two other omissions might be noted. A short connected account of poly- ploidy in relation to size and germination of seeds and in relation to seed yield might have been included, especially as references are made to genetic factors. Under conditions affecting germination some account might also have been given of ethylene effects, extended possibly to des- cribe the use of seedlings as means of detecting traces of the gas. Space could have been found for these topics by shortening the treatment of some other themes, such as the chemical composition of seeds, and pos- sibly by omitting certain elementary statements on physiological processes which are scattered throughout the text. Notwithstanding the criticisms which have been made, this is a valuable book, of interest to botanists in general as well as to the plant physiologists, horticulturists, foresters and agronomists. It brings to the notice of the reader a number of references which are readily over- looked as they are not always easily available and appear in publications which are unfamiliar to many botanists. Also, both explicitly and im- plicitly, leads are given to further research, which should make the book of value to other investigators of seed and seedling physiology. 186 The Journal of South African Botany. Much space is devoted throughout the book to practical applications, and this should make the book useful to nurserymen and others concerned with the growing of plants, provided they have a background of physio- logical knowledge. The later chapters are as a whole better than the earlier ones and mention may be made of the useful chapters on vernalization and embryo culture, and of the stimulating chapters on dormancy and storage and life span of seeds. Although the authors do not comment on the matter the parallels between seed and fruit storage and post-storage behaviour are of interest. In both cases low temperatures are of importance, and on withdrawal from store there is danger of rapid deterioration in both cases, if sufficient attention has not been paid to providing the best storage conditions. Of course, seeds can be kept at lower temperatures than fruits, and the storage moisture relationships are different. The book can indirectly make a contribution to South African botany both “‘pure and applied’. Little is known concerning the seed and seed- ling physiology of our native plants, and this book should stimulate a number of interesting and fruitful investigations bearing on the natural history and ecology of South African plants. The book will also be of value to those in South Africa concerned, directly or indirectly, with the commercial handling of seeds, in which activity there is room for much improvement. Wm. Epwyn Isaac. New ZEALAND PoLLEN StupDIEs. THE MonocotyLepons. By Lucy M. Cranwell. Bulletin of the Auckland Institute and Museum. No. 3. Pp. 1—91. Photographic frontispiece, 10 photographic plates, 66 text-figures and 1 map. (Harvard University Press, 1953.) Miss Cranwell’s “‘Pollen Studies’ contains a very full account of the pollen of New Zealand Monocotyledons and is well illustrated with very numerous text-figures and photographic plates. A somewhat unusual but very welcome feature of the work, and one which the South African palynologist will find of great value, is the general introduction containing a discussion of the basic and aberrant types of pollen-grain found in the Monocotyledons in general. There is a section on morphology and method, with a useful glossary of terms