A Taxonomic Revision of the Genus Asparagus in South Africa
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A T A X O N O M I C REVISION 0 F T H E G E N U S A S P A R A G U S I N S O UT H A F R I C A B y J. P. JESSOPTown Cape of Being a thesis presented for the degree of Master of Science in accordance with the regulations of the UniversityUniversity of Cape Town. Cape Town November 1964. Town The copyright of this thesis vests in the author. No quotation from it or information derived from it is to be Cape published without full acknowledgement of the source. of The thesis is to be used for private study or non- commercial research purposes only. Published by the University of Cape Town (UCT) in terms of the non-exclusive license granted to UCT by theUniversity author. CONTENTS: Introductory remarks •• •• •• •• •• •• •• 3 Acknowledgements • • •• •• •• •• •• •• •• 4 Taxonomic character,s •• •• •• •• •• •• •• 4 Subgeneric groups • . •• •• •• •• •• •• 8 Proposed sections • • • •• •• •• •• •• •• 11 The morphology of the photosynthetic organs of Asparagus •• . • • • • • • •• •• •• 14 Cytological investigations • • •• •• •• •• 20 The recorded history of the study of Aspar� in South Africa • •• • • • • •• •• • • •• 24 References •• • • • • • • •• • • • • •• •• 31 Introduction to the taxonomic treatment •• •• 34 Key to the species of Asparagus in South Africa 35 Section CAPENSES • • • • • • • •• •• •• •• 40 1. Asparagus ca.rensis • • •• •• •• •• •• 40 2. Asparagus stipulaceus . •• •• •• •• 47 3. Asparagus suaveolens •• •• •• •• •• 50 4. Asparagus glaucus •• •• •• •• •• •• 58 Section AFRICANI •• •• •• •• •• •• •• •• 60 5. Asparag_u.s africanus • • • •• •• •• • • 60 6. Asparagus J21EffiOS� •• • • .. •• . •• 72 7. Aspara_g}!s denudatus • •• •• •• •• •• 75 8, Asparagus virgatus •• •• •• •• •• •• 79 9. Asparagus consangµineus • • • • • •• •• 81 10. Asparagus thunbergianus . .. •• •• •• 84 11. Asparagus mucronatus •• •• . .. •• 88 12. As12aragus macowanii • • • • • • • •• •• go 13. Asparagu_� retrofractus •• •• •• •• •• 92 -2- • •• • • • • • 95 14. Asp2:.ra_fills com.pa,ctus •• • 15. .Asparagus laricinus • • • . • • • • •• 99 16. Asparagus microrhaphis • • •• . •• 102 Section EXUVIllLI . • • . •• • • • • . .. 104 17. As:p8.£§.gu s _ _Eu_yiali s • • • • • •• • • • • 105 Section RACEMOSI .. • • • • . • • • • •• 108 18. A�8:_:r:_a�_s>XX§l.Canthus • • • • • • • • . 108 19. Asp�ragu.Ll_u.p.J�oides • • .. • • • • . 110 20. Asparagus bracteolatus---- . • • 111 21. Asparagus sann.entosus . • • . •• . 114 22. AsparaB1J.S saundersiae . •• 118 23. Asparagus aethj.__2I:_i_2us .. • • 122 24. Asr2aragus falcatus • • • • • • • • . •• 130 25. As12aragus racemosus • . • • •• • • . 133 26. Asparagus krebsianus • • .. .. • • 137 27. As12aragus pubescens • • • . • • . •• 140 28. Asparagus kougaensis • • • • 0 • . •• 142 29. Asparagus nelsii • • • • . • • . • • 144 Section STRIATI • • • • .. • • • • • • . 146 30. As12a:r:_agus striatus . • • •• • • •• •• 147 31. As12aragus subulatus • • • • • . .. •• 151 Section SCAJ'JDEWTES • • • • ... ·• . .. • • 154 32. Aspara_gus scandens . • • • • . 155 33. Asparagus ra.mosissimus . • • . •• 158 Section CRISPI • • . • • .. • • . • • 161 34. Asparagus criSJ2US • • . •• . • • •• 161 Section MYRSIPHYLLI .. • • • • • • • • . 165 35. As12aragus as12aragoides • • . .. • • 165 36. Asparagus ·-----undula.tus • • • • • . • • • • 179 Summary • • .. • • .. • • •• • • •• . 183 Index • • • • • • • • • •• • • •• . • • ... 184 -3- J{IJ.,:;.:TS: J�'T.:_�O:DUC�ORYw In the J?lora ( lfS 6)' ' forty :five speci·,.:fJ o.: 1-i.:', Jarag11s from the Cane Co1m:.y, Since t'-i.e,1 and 1J/ n.s1,1 s•1ecies. Salter str,essed the ca�e Peninsula in 1�50, the:refo:::'c, decided to 11n.dert e a 8.C'COr:1- ·,r t'ield v1ort. Collections nocossnry, t.f these, ·.n 'l ·,; r, '\T re, y V r..1.� .. u� ... • • -'---- ., __c� , '-' , ,___. __ .. : ,, c• n" a� :..J'TU J ••s O ft e11 11 of di2tri'1ution. has bc:en recordeJ iro,;, i/ie southern C3. 1Je P�covi:;ce 9 1rJ -4- the ea13t coast of Af'ric3 9 a.1:.d accord· w: to :::.2.t:er ( 1875) extends into Asia and Aust7aJia. f"en,-1s:> Asparr:F.us___,.._ �· .... ,_.. __ - occurs t�roushout Af�ica 9 is very cratefully ackn6wledged. have coLi__ect 1 0 In pe:rticnl8r 9 Dr l\. 1 • ?:2.ll must be thanlred for his '1" -:- · ('' -, -L ad vies on cytoloc.icel t-:-:ch::1.:Lcpss9 and Cant. ·_,_: •. -• )cu_ t,er ·,.,-':'ile t111? gre2t 1 a::Cll)(l 9 facilities ·;,ei"e a1so TJ::..novi:ler. ,:r the Curators of The T[oyal The Botanic (ia:rd ens and l'�ll s eur:i.9 -L·O- - , t a"1iscne. ' G-.ar t ·en und _,- uGe-um d er lT11i- versittit, Zurich. The Jepartment of Aeric�ltural Technical Services must also be thanke0 for releasinc; the present author frora hi3 other dtities so as to undort�:c this revision. -----·T.1\XC>CE,IC-�·>O.·--- C�IAf.ACTBRS·----·----"· � Up to the present 9 there has heen no satisfact- ory delimitation of the species. because no-one has undertaken a major revj_sion of t·:e 0 iroup since th0 Flora Capensis in 1C G, 2nd nartly because, in the absence of flc'ral characters, most of -5- ,., the ecies Ill1J_S t 't)e cl ined on 'I t3t e te:rs env c'".!..J. are 0 en SlJ.. SC ible to factors. A list of the characters of UGO i� d below generic level i en notes on ';he gene�al v ue of erent characters. All e s fa ), e members of the ar-:3 Doti.et es sl 1 ff b::r.oad,=.;r the oute::r seg- s, and e distal borders are so�et C i se c}1aJ:ac. ters sre e:..1u viith in. a :': e ter• ised by hav tho d hc1lf of -tepal reflexed. 1,.J� ) ------Stame:'.rn The s s s so es ea pair of minute lateral spurs at a.rie�}ts. is 1 character not b c er� u ::.ie c1 il ter s tin e Flors of the C e P�1 tis valu,2. G V from. yellow to 1 orange ack.:. Wor!::: on r..i.ate:rial 8 '.l.OW that co�our is useful, but on spec ens the colour can usually not be determined. c) Fruit 1 '] l) The iruit is gen -v a ' flesh;y e with two or three s2 and ':io in shape } but dry s e-se berries as in A. scanden occur. eciec', A. C has a persistent perianth enclos e e characters ar• usual eat spec ic level. Fig. 1. Raceme of Asparagus aethiopicus. / -6- d) Peduncles All species have flowers borne solitarily on the ultimate flowering stalks. In A.--·-----·-- stinulaceus·- ..... �. ··-··- -·- ....,._ and 1 .A.__ cayensis these stsll::s are re< uced to discc,. As all leaves are reducei to scales, it is �ot strictly corr- ect to ref er to umbels or rac ee1e<:,. J3y treatin;'. the cladodes as leaves, the 11se of the convec:1.tional term- inology ·Nill be usefr'.l. The tern 12e(uncle ,;j ill be used for the flower stalk excent in the section RACLLiOSI v;here the teTm -�---·--�··nedicel v1ill 1:ie used for the final flower stalks and 2eduncle for the main axis of the inflor2scence. Inflorescences ( termed rac�m_e� heTe) consisting of branches bearing only bracts and ( g ) _pedicels occur. Fi . l. :Jovvev er, in species prod- ucing racemes, cladodes sometimes an�enr at the bases of the pedicels. In some species, both terminal 2nd axilJ.a:rJ flov1ers occur as L-:i. A_. africanus, but ir· others terminal, as. in _·1 •.cs:2._e�'.tsj _,s_, or axiLLor:y ned- uncles, as in A. scandens, are diaGnostic. The uobels� referred to by Baker, are clusters of terminal flowers, and are rarely found as the only type of inflorescence on a rlant. However, exceutions to this occur in the section STRIATI and i.11 A. nlurno s1-1s o 1 I'he ped:..1ncles may be solitary, paired or nore -r,umerous, and these characters are sor!letir,-1.es useful at specific level. In all s�ecies the peduncles are articulated, and in A. nelsii·-·- some\imes tDice. e) Roets The roots may be tu�erous or fi�rous. Cla::rn- ification cannot usually be based on furti1er subdiv ision of tu�er type as the tubers are particularly suscevtible to environmental conditio�s. (Discussed. Fig. 2. Wrinkled surface of aerial stem of Asparagus denudatus. Fig. 3. Grooved surface of aerial stem of Asparagus compactus. Fig. 4. Ridged surface of aerial stem of Asparagus aethiopicus. Fig . 5. Peeling outer layers of aerial stem of Asparagus exuvialis. -7- in more detail under 1::.. • . asparaRoid_es.) In most E:ipecies there is a single root-stock bearin� lateral tube�s, but in several species of the section R.ACE>OSI, as in A._ aeth�jcus, there are several lateral roots each bearing lateral tµbers. f) Stems Roots and aerial stems are produced fro� perennial rhizomes in all species. An:1.ual and p Pren ial aerial stems occur, but the duration of the life of the aerial stem may vary under diflerent conditions. In A. exuvialis the aerial stem is perennial, tut the cladodes are deciduous, and after a oonth or two are re})laced. Some sDecies are predor1inantly clisibers and others are erect. In es.ch of thes·' g2.�oups exceptions may be produced by the effect of the environnent, part icu:i_arly the presence or absence of support ard the amount of light re2,ching the nlc:-mt. The aerial stem. may have a smooth/' v1rinkled (FiG. 2.), grooved (rig. 3i)i ·ridged (Fig.·4�) or hispid surfac�. These charac- terc are sometimes useful. In A. exuvia].is and possibly other snecies 1 the outer layers of the aerial stems and tl1eir branches peel. (Fir;. 5.) In several species, including .A. com12actus and A. retrofJ:'actus, the grooved, white outer- ·layers disintegrate, leavin.z the stem smooth and brown. Another character of occasional value is aorial stem colour, as in A. namaensis which has a green aerial stem. g) Branchlets Branchlets, which are ultimate, specialised branches bearing cladodes and flowers, occur in some s11ecies. (Especially the section CAPENSES, where they are fascicled.) (Fig. 6. ) Fig. 6. Branchlets of Asparagus capensis. Fig. 7. Fascicled spines of Asparagus capensis. -8-, h) Cladodes For a discussion of the use of the te:rm 2J._a0p_d�.