Morphological and chromatographie investigation on the species of the series conicibaccata of the genus solanum
Texto completo
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(5) • CONmNTS. Acknoli'ledgements I. Introduction. II. Li terature. III. Series. IV. l"aterials. Conicibaccata and its. species. and methods. a.. l{¡aterials. b.. Msthods. • v. revieu. 1.. Gormina t ion. 2.. Morphology. 3.. Cariological studies. 4.. Chromatograpby. Results l.. Germination. 2.. Morphological observations on field and herbarium specimens. 3.. Chromoso¡;¡e counting. 4.. Cr~omatography. I. VI. Discussicn. of phenolic compounds in leaves. and ccnclusion /. VII. Bibliography. -ii-. /.
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(8) ;rOHPIIOLOSICAL ARO CIIROM!\.TOGRAP'JIC n!V',S'l'IGATION ON '~. A ,. I. li.. A thesis presen ted to the }'~culty 01' Se i.ltce in pc'rtÍ'll fulfi1ment "f the requirülnents for the deg"l'ee of M2st'3r oí'" Sciencr;l. by. Depélrtment of BO'Célny UnivarsHy of Bir;n:lng-ham -. í-. ,. /?; í1 >n i /1 'í./J4n ' 3eptember 1972. ------.
(9) -.( 'i. l I. • ACIQIO~'¡LEDGEr.ISNTS. 'fhe author gratefully acknOlüedges th0 <3ncouragement, guidance and facili ties providad by Professor J.O. Hawkes during tha tonure of this study.. Thanks are also extended to Dr. R.N.Lester for suggesting this theme and for his instructions on chromatographic techiliques and. to. tilo British Council for financially supporting the author's stay in tllis country. Tho permission grantad to tho author by tha Instituto Colombiccno Agropecuario (ICA) to undertake tila. ].l. Se. • degree eourse in. tlle DElpartment of Botany, University 01' Birmingrram is also. •. grateful1y acknouledged •. /. I,, 4. ¡ I. -i-.
(10) - 1 1. IN'rRODUCTIOlT. 'l'he serics CO:rrCIBACCI,l'A of the sub-seetion HyperbasaTthrurn in. th'3. eenus. SOls.L1Ul'l. is a vTidely diú Gributed series in the ArJaricas. ....... ..... __ .......... -_.~..;....:. _.~-,:¡. ~. in advanc e as : Sok'1.num a{)Timonifo1iu:n. from r"lex.ico. ,2.. eolcnoic_num. from Colombia. ~.. from Ecuador. p~uciju~u~. S. urubambae. from Peru. Tue species belonging to this series. a~e ~ather. easily. recognisable on account of thell' berrics vhich are lon¡;-ovoid. 01'. eonical-elongate. '1'ho habit is highly variable, i.e.; species such as. s.. •. JlGucijugum. colombia. nu:n. 2nd.§... ~[1hu?tensG. Can reach 2 metres, ancl. violaceimarmoratum. have a low groYrth, T¡fhile. S.. s.. urub3,mb2.0, and. s.. may reaeh 3 m. in height, in thoir natural habitat.. !\Iost of the spGcies are hcrb.:.ceous, areot ascending, ueak1y erect or deeumbent. The habi tat is as variv.ble as the habi t too: ,2.. aL""; monifolium frorn Nexico and GU3.tornala, and,2.. eolombio.nurn from Colombia Venezuela prefer a "et, cold 11abí tat. §... urubambae. and. grovs very well in. Urubarnba l,hieh is a warm valley at 2,200 m. above sea level near. Cuzco in Peru. S.. violaceim2.rmor~. has be en found among bushes. and forest at 3,000 m. a.s.l. in Bolivia. The corulla shapc in almost all the spooios of the serios is generally rotate to pentcgonal; fe,; of thon aro sub-stellate, and occasionally. stellate as Corroll (1962) points out. The colour of.. the coralla varies from white to violet or not a good charaoter to assign. p~ple.. Tho loaves are. species to the series duo to tilo. enormous variation in shape and. siza intho different sp3cies and.
(11) - 2 -. ,. ¡. 1. •. 'l,. , f. ! j. !,, 1. ,I í. •. AG¡:". Pie. 1.. K305·. GOD.2ral fo:--::; o::' pot(:-~to lq2f aG Q3ts.blish8d b.y Iie1~~kos .:::.nd i{jcrtin2: (1969). Loo.1' of 3.~:;,"-,:J:1i..:;~olit1n H 305.. ir. '1-¡or:¡~ínc'1.1. L 2. Priírl:'.:',,! 1,:~~;o:'Ql ·ScCOj~~clr~,:."y. 3. ~oJ.."'ci;:n:'y. lccd'lct. lo,:";.flot· l .. ,:tcro.l ,-:..n.¿ l:::.to::-c:.l ir'. . tc:,j'3ctod. 18.:J.fle-t. A "C::-08COp:'C ln.cflo·~ S3 30mi-J~:::;i:,;copic lo~~f'13t p ·j'otiolG. PL TI. Potiolulo R¡l['~chio. /.
(12) - 3 evo n ',ri thin tho. S:inlO. c90cio3. Thoy nre odd-pinné1te a.nd tho numbcr of. pnil's uf ln.teral lcnflots (Fisure 1) is 2 to 7, occacion:üly 9, with. numer-ous to no intG;.:'stitial leaflets.. pubescent. All of them ").'e tubor-bearing, though thiz chc.racter is not ahrays As. 1<1011. Sh01/S. numbGr is 24, 48 8,nd '(2.. mn:rked. The chI'omosome. by tho variation of the morphological characters,. Conicibaccat,,, 1s a difficuI t series to evaIuate taxonomico,lly. It has been a matter o:' controvcrsy foI' scientists to put soms of i ts species in the right taxonomio. posi'~ion.. boen olassi!'ied on a morphological. J.!ost of the spocias have. basis anO. i t l/as thought that. bicchemical characters coulo. be successfully spplied in heIping. to. cIarify the taxonomic position of the doubtful speciee. LIston anO. '1'=ne1' (196:». •. pointed ou~ the.t bioclwmical comp=isone coulo. be a. gooo. dO. in clearing. up o.il'i'iculties. in botanical. taxa •. 1!any stuo.ies using bioohemical techniC[ues such as eIectrophoresis, immu.'1oelectrophoresis. and chromato,gTaphy. have been. successl'uly applied, coveriJUJ o.ifferent aspects of Solanum taxono:ny. The present .ork is a study of most. of the species ol' series. Conicibaccata by: (i) morphological observation on availP.ble herbc:rium materi.ü,. (ii) morphological comparisons on living plants. erovrn undcr standard. environmental conditions anO. (iii) comparisono of the wpeoies using Chromdographio patterns, (thin layer ollrom2.tograpl¡y) as taxonomio characters (prosencejabsenoe of spots) froro leaf samples..
(13) •. 4 -. '1 "}. • 1 J. 11. 1. LI:I\2.LiATURE REVI3t,[. i:l2.ny scicntists are involvcd in the t;;;.xonor:ty of So12,nu:n spccies.. ,. J. !. aG on. otbar Gspects oí tbe potato, suob as breeding anO. conservation. An important stud,y oí almost all tbo species cf tuber-bearing So1anurn 1ms 'ill"de by Corren (1962), wbilst studio:: oí spocies from Peru ;¡ore made. by Ocnoa (1962),::ü.d Vc:rgc.z (1949),al I.d. from nolivia. by A. C&.:cdenas. (1966).. The botanist Ugont from UnHed States bas colleoted tTild speoies in Soutb America anO. especial1y in 1-1exico and bas boen interested in taxonomic problems and morpbologica1 changos in Solanum. species. (1967,1968,1970). l.!orphological chaI'acters h:wo been vridoly usad in tbe taxcnolt.y of the species but recent e.dvances in comparative plant biochemistry. •. bave p:rovided nelI teoh.lliques for tbo ::;eparation oí specios not only in. ~2~~~. but in other genera sucb as Phaseolus, Triticum ,etc.. These techniques aro electropboresis,. imm~~oelectrcphoresis. and. chromatography. Sava:t'al works bavo been achieved en Solanum species. appl;v.h~. disc-electrcphoresis to 3ee the variation and distributicn of esteras03. f~'om. Sol"num tubel's; foL' separating proGeins anO.. fOl'. determining species by their pattorns of soluble ,tubor proteins as 'Oell as for identification cf cemmercia1 varie'tios oí' pctato (Desbcrough and Pe10Quin 1966, 1967, 1969). lforks on the tuber-bearing species of Sole.num using immunoelectrophoresis include that of Goll, Hawkes anO. ¡frigh~ (1960) ~Tho. ¡'Tere searching for re1ationsbips betvreen uild );!exiCiln anO. South. American speoios. Their results inrluced them to divide tho lf,exican species into thrae groups by testing the species against. 2- tuberosum. antiserum. Tile South American spocies aud certain Hexican ones gava.
(14) - 5iclentlco.l irnmunoloeic3.1 p-atterns to. ~.. tub·:::r-03um. The ovidence. obtained allowod them to recommend this techniquG as very useful in mod.ern taxonomic studies. ).lore recently Lestor (1965) confirmed the results of Gell Gt 21. (1960), obtaining similar p.s:.ttorns ¡-ri th South Americ.::o..n sl1ecies. using. douale diffusion GxperimentG wi th S. eC8.ule end S. tuberosun. c.ntiserum. Ho¡.¡ever, "í'rh:en he used an. a different. spectru~. e~ctract. of S" cO;7lr.lersonii he got. of moxe lines.. Furthel' immuno10gica1 studios W'ere carried OClt on S. bulOocD.Gto.num High variation Has found in. sambucinum, tUG i",munologicill. patterns confi.l:'ming its na.tural hybrid origin between S. uinne.tisectum. and·.2._ cél,rdiouhyllum subsp. ehrer!,.bcrgii" :Many otheL" studies have. dGffionstratod that immunologicé\l techniques may prove the hybrid. •. nature of many plants. Dii'ferent types of chrome.tcgraph;i:c such as gas, papel'. ",nd thin 1ayer, havEl been ,ridely applied for taxonomic puxposes in several kinds oí plante. Alstotr anO. Turner (1963), Stebgins. ~!1.. (1963), J,IcClure anO. !üston (1966), Smith and Levin (1963), B~ (1966), Ising and Frost (1969) have successfully used leaf or flo;¡ar pheno1ic cor. '.s in the identification of species or the perants. of. resPQct" to. 110X\. ~mplished. Dod~. )55). inh ino.ep,. I. .---c010ur. chrom'"tograph;i:~on. Solanuffi species, much. using ,üld and cu1tivateo. epecies., 2.. ir',. anO. Dodds anO. Paxman (1962) studied the in dip10id potatoes. They founo. that three. df!t "10oi (P,R and 1) ere concerned uith the prGSenCe 01' laclt. of certain anthocyanic pigmonts. Thoy uesd extracted pigmonts froro tuber skin, Spl'outs "nO. i'lol,ers .::.nd sepiJ-rated them by papel'. .. chrowatography. The anthocyanic pigments in. ~uestion. e.re Polargonidin,. Cyanidin, Peonidin, De1phinidin, Petunidin and malvidin, but the stl'ucture of each ene ,"ras not inve5tigated.. !.
(15) ,¡. 6 llaroorne (1960a, 1960b, 19620.) continueú studine tha pheno1ic co:~pounds. of SoL,nur.1 species. The chemica1 nature of the main pheno1s. and the distribu-tion in f'¡"'YI . -. 7'\T'0V-i.-lA -~. ,. A. ~'Tild. -. J.:rl-·(~8.t. and cu1 ti vateél: species hAll"). in. separa ted by chromatograllrty.. !. ¡ !. !. •. /. .. 7;1.16. ,T1!S. id.anti:;:'l.ca.t~on. achieved. His. ox- pnenols.
(16) - 7-. Th€l sories Conicibaccz.ta ,;;ras originally esto.blished by Ilitterin 1912 enJ renemad b:r RYdbarg in 1924. ?ollOHing tha 1aw of priori ty thc. i'~irst. neme is validly estaülished. 3.nCL. tt';,e second l'elegated to. synonyny (Ha,rkes, 1944). The species rEJcognizec. by Heltkes Corren (1962) undar series Conicibuccé1ta are. &.S. p.0l";-nmS. •. follo>fs: C01T'lELL. a~yirTlOnifoliuíil. 1. §... 1. S. DIT;:imor¡ifclium. Z. .§.. colombianu!.l Dun.. 2. .§.. C010Clbi "llnm. ~. ". .§... ffi0SGOD2nlJ71'I. Ha,rkes. 3. s.. 4. §... oxy,:prOUtn. Schieda. 4. S. oxyca:r'pum. 5. §.. s.'.ntolal1¿le. 5. .§... 6. .§.. urubambae. 7. ~::. ~. ". viol3.ceimnrt'J.oratum Bitt.. 6. §... viol,,=:ceimarr!1ore.tum. 8. .§... éty2cucho~. 7. .§.. ayacuclroense. 9. .:2... buesíi. 8. .§.. bU0Sii. 10. .§.. jaenense. 11. ~. lexissiT!1um. 9. s.. 12. ~. 13. .§.. neov2,rgasii. 14. .§.. paucijul{Um. 10. .§.. u;:;:·uc i .iu1um. 15. S. -oillahu::l.tense. Vargas. 11. §.. ni11ahnatensa. 16. .§.. villusnetalum. Vargas. 17. §.. Hoodsonii. 12. .§.. lroodsonii. 13. S. límcYls0 Cor:r: •. 14. S. hUé"!l1cabambense Ochoa. 15. S. cl·::.viforn:e. Corro. 16. .§.. f12,he.ul tii. Bitt •. 17. S. otites. 18. .:2... ~. 'lydb.. Vargas. mOGco-panttm. ~~tolallae. Juz •. Ochoa. Vargas. Oohoa Eitt.. . mnl t iflorum. 1:3xiss"t.mum. Vargas Ochoa Bitt.. COl'r.. /. .. (1963) and. Dun.. filamcntu:n. Corr..
(17) - 8 19. s.. aC~88co~~. 20. S.. ~uconditum. Ochoa Corr. ll. species,is re1egated by Corre1l (1962) to a synonym of S. violacoim.:1rmorntl1m.. 1.. j:J,cnenSG. as. ~,.¡ell. &8. ~.. neovo.rr;Ctr::ii , both. described by Ochoa, are not menti0!13d by Correll. S.. Jr.u1tif1~~,. published by Vargas (1956), i3 placed by Correll in series Transaequatorialia, though considerad intermediate betueon Vi. olac(3Í'11ormora'tum ana. -S. -=:...=::.= villusDetalum. .;..:;.;~=~-~="-='. species. ~ight. and. .2.-. ;¡i th the comment "this. eventually prove te be referable to. sel"i0S. CO?licib.'l.cc3.ta'.;'. . ".:Cum 1s also placed in Tramf-ec¡uatorialia by Correll. Ocho a (1962) hOlfever placed i t in s0ries Conicibaccata. Certain species 1'1hich are considered. valid by Correll are. classed by Hawkes (1963) as follo;¡s:. •. .§.. limen38. Synonym oí .[. ch2.COenS8 ( series Commersoniana) §... hua~6abambense.. Ser íes. Pi urana. .§... cl;:J.vix'orme. Synonyrn. ef §... §.. fl2.haultiL Synonym. oí' §.. pDuciju'jum;. otites. s.. ~tolallaei. o colombianum; und §.. filameni:um. Synonyrns oí' .;:... acroscopicum. Series. Tuberoso.. ~. recond:i. tum. Synonym of S. iO~8t{;;.lum, (serios. Demisa.). Ocho" (1952) stated that §.. Umenso lYas based en material of. §.. ehaceense iml'orted in to Lima. í'rom Are;(mtina. He placed the next .'. three species. as follo"¡s:. §.. huancabaT:1bense. Series. /. Tuberos2.•. §.. claviforme. Synonym of .§.. ~.. •. acroscouicum. Series. 12.xis8i~.. Tuberosa •.
(18) - 9 IV. a). JLWE?.IALS. Mm. !,ffiT¿ODS. f'lA TERIJ.LS -, ,,;> ,.. u..:... m&.ter:Lal usad in. -'~ lo. surveY. uresen:tsd.. ~n. Table 1, was provided by Professor Hawkes froTa his collection which is ke9t at. ~'lintorbaurne. Gardens of the Universi ty of Eirminghz.m .So;ne. accessions appear 1'lith tho prefix H.Some. P 1 indicating. that tboy are. f~'om. others aro prefixed. xith. the Interregj.onal Potato. Colleation of Sturgeon Eay in ;risconsin U.S.A. ; other s&mp1os were o1'ow¿ht from the Co1occion Central Colombiana Instituto ¿gropeouario in Colombia. o',rn prefix. (c.c.c. l·. of potatoes at. The samples ',rhich appear ,-Ti th ¡;¡y. (L.1. l are recent collections uilich ha.ve not yet been. registered in C.C.C. b). KZTHODS. l). •. Gerllina tion Ten seads from ec.ch accGssion (Table 1) ;rero so~ra. "i thout pretreatment. at tilo beginin,g of April in plastic seed trays. 1dth John Innes seed compost in the glasshouse at !finterbourne. As sorne accessiol1n did not germinate aftor 4 w0"ks, a néHr sOiring. was. made, Id th the resul ts lfhich appear later on T2ble 2. '¡,¡hen more or lesa one inch hieh. 2-ik. ~he. seedlings. 1'1'01"e. }?o·t;ted in. inch diameter clay pots l1itil Johu Innes compost ¡{o.l, and ,;-hen. thoy llere 6 inch tall, they l,ere repotted. in 5 ineh plnstie posts· in. John Innes coopost No.2; The maximU¡¡L temperature recorded in the glnsshouse durin',¡ 8 weeks after the so,ring "Ias 36 C. a.no. the minimum l;as 6 C.; tila humidi ty fluotuated betlJeen 361; eeeasionéüly loss, and. 30'¡&. As is 1'7011 known, high temperaturas are not eonvonient for the production of flmlers llnd fruits,especially in ,ril;l Sole.num speoies Yrhieh are fror,) cold arens ,rhere tho m::tximum tem;;>erature is 15 C. '1'11is circumstance. made it nocessary to groif. nODIO snffiples in tilo field.
(19) 1. !. - 10 -. .-. Tabla l. list of a.cces3iolls. in grsenhouse at Hinterbourne. Go~;n'l. 6-4-72. No.of. -- - ....... '..J";". 1 2 3 4 5 6 7 8 9 10 11. •. •. 12 13 14 15 16 17 18 19 20 21 22 23 24 25 26 27 28 29 30 31 32 33 34 35 36 37 38 39 40 41 42 43. SP¿;CI_~S. J,.GCES.3ION. ~. saunes. Hi1.RV ~s:rIUG n!-,rjll~. j;-....... \....~_. AGF COL COL COL COL. H 305 709 757 H 758 H. 2544 H H. Hai~·kes. collection. 11. 11. " ". 11. 11. Colombia 11. COL COL FLH. L.L. L.L .. L.1.2196. 1(3 1,'(3. H 734. ¡.:SP. MSP. mc. OXC OlCC PCJ PCJ PLL PLL PLL SAN. mm. VIO VIO VIO VIO VIO CHM CHli!. cmr. TUQ TUQ TUQ TUQ 1M].! 10P 10P IOP 10P 10P IOP IOP. " n. " Hmrkes col1ection Sturgeon tay. P.I 283083 • 11 11 P.1. ?30462 11 n P.L 230517 H 98x H 313 Hawkes col1oction 11 H 958 "11 11 H 960 Stlu'geon :eay P.I 246971 P.I.246971.2 " " L.L.1676 Colombia 1676.3xl676.6 Colombia 5103 Ha.;¡k0S collection H 663 ., 11 H 712 4436 4474 5040 5042 H 767 Hawk.es col1ectiol1 H 101 H 686 " "11 H 723 "11 H 759 " Colombia C.C.C.2547 C.C.C.2547.1 " C.C.C.2547.10 " Hal1kes collection H 703 11 H 316 " 11 H 317 "11 H 318 " /' " H 319 " 11 H 320 "11 2292 "11 11 H 351. 1959 1960 1995 1965 1971 1970 1971 1971 1971 1964 1964 -1964 1961 1964 1964 1971 1971 1971 1971 1971 1960 1962 1971 1971 1971 1971 1971 1964 1964 1964 1965 1970 1970 1970 1963 1962 1962 1963 1964 1962 1962. S:3SD3 S(ldN. 30 10 10 10 10 10 10 10 10 10 30 10 10 10 10 10 10 10 10 10 10 10 10 10 10 10 10 10 10 10 10 10 10 10 10 10 10 10 10 10 10 10 10.
(20) ¡. ¡. I•. l. - 11 in ord.er te get flol'el's fer roorphologioal description3, corolla preparations and th0 collection of. herbariQ~ s~ucimens.. July a plant. r:as placed in a phytotron of. At tha end oí. ¡. j. frora each étccession. ~'rhich. xespectiv81y. This is about equiv(:.lent to the natural enVirOll.11antal. conditions oí the pl,c.nts in question. Lightwas regulated at 1.000 Lumens. oí intensity at three feet aboye the ground giving 18 houro. dayUght.. 2). l:oruhologY Herbarium specimens l'¡ere taken from seedlings. transplanted to the field and phytotron. Thase samples 'rere used as a basis for the description of chars.ctE1I'S ando a comparison \,ith living plants remaining in the field and pr.ytotron; the living plants wera. •. used in addition for the description oí characters not easily discernable in the herbal'ium specimens. Corolla preparations ,rere made fron! the plante froro ;¡hich the herbarium spacimens were taken, ,.hich ware grO¡ffi for this purpose. Complementary information was taken from herbarium specimens kept in tha herbarium of Professor Havrkes, and froro Correll (1962) and Ochoa (1962). The oharaoters Hhich Hore mainly oonsidered. on each species,. such as habit, fIauer sha.pe, aizo and colour, pedicel artioulation, :. lateral, terminal, and intersti tial.leaflets etc. lfere taken from, living and herbarium specimens ,[hen literature when certain. 3). they "ere avai1able ar froro the. organs Here missing.. Caryolor;ica1 studies . Dodds (1965) pointed out that. •. has 20 zp8cies froro which two are diploid, six /. Conicibaccata serios are. totraploids, one.
(21) - 12 iG hexClploid :in,l (llevon ."re unlmo:m, b:.¡t ho did not montion an;¡ apooies. ;,mik05 (1963) indic:lted two spooies (~.smtol!lllae, and S. Vi. olí.,ceiDE.rmoraturn) o.s diploid, three spccies (.2.a.~ir;lOnifoliurtl ,. ,. .~. , .' '" ",:",. ~. ~>.-. hexaploid. Although actu'llly ths clu'omosome number oí' mO:Jt oí' th" ;:;pecies is already kno1>'n, i t ¡¡as deoided. to verií'y i t on the available. material. The technique oí' Tarn ( Cribo 1970 ) was follo,red for root tip chrornosome counting. Root tips uere taksn í'rom seedlings, grown in the glasshouse, pretreated in cool water with ice overnight, then fixed in Jacobsen's fixative (j0;:\ etnanol and lactic acid 99:1) at OoC. for two weeks, but could be longer before obsorvati:m. The root tips ,rero hydro1isod in 1O~ HCl fer ten minutes, washed in distilled. •. water and stained in. l"cto-prepienic ercein for up to one hour. Tuo mm of the root 'tip ;rere squashed prepared. in 45% acetic acid. The lacto-propionic orcein "ras. as follows: one gr. oí' natural orcein ¡¡as dissol ved in a. mixturo of 23 m1.of l"Ctio. ~cid. and 23 ml.oí' propionie neid at room. temporaturo. The mutu¡'e was made up to 100 ml.with distilled water, sh"km and fil'tered, (Haskell and llills 1968). This stain gave oettor resul ts than. Feu1gen ¡·t¡ioh was also. tested.. 4). ChrO¡;w.to¡<;rauhy The glass plates (20x20) for chromatography were. ooated with a suspension of 15 gr. microcellulose. pOl,der grado 300 in. 100 mI. of distilled ¡,ater mued in D.n electrie blender, 10 seeonds at 101·r speed, thGn a minute at high spGed repeatsd twiee, proeessing only fouc: glasses eaeh time. The spread glass plates. "ere plr.ced il1. o. a drying rack and áied ovornight in tho ovon at 60 C. Threo or four loavos from the same plant of each aeeession gro}rn in the glasshouse.
(22) J. - 13 -. 1 1. .. 45. '.{ere h'crv8'3ted '<,no. dried in a fore8o. c,ir oven at 'l'h3 dricd le. lf. l~mirn ',fOCS. 1. plcce3o. in ..... t- __. .. ...... "". I. 1% HCI in methanol. I¡. 1'01'. 1· -. _" ...... :1. _ .". ~u.v. C.. 1'01'. 72 hours.. glCt3s vLl Dnrl ground to. •. :1. fine. ml. o:{. ......... '-' ...... e:~ch 200 l1',g:¡.'. of erushed leéd anO. 10ft in the. dark overnight to extre,ct. The extracted solution ;ms applied on glass pla te (tilin layer ehromatograpily) at :, cm from two edges ;1i th. 1. 3.. fine. pipette, repeating st:;von times ando d.ryinc; after sach .::.pplic.:::.tion getting a spot of abeut l em in diameter. The. t~{O. dimensional chroffiatograms ¡'Tere ru.."l in Pangla,s Shandon. Chrom2.tographic t.:nks, pütt.l.Xlg' fo'U.l' glass plates in o.:1oh tunk. The first diraension ,ras run in butanol 1-01, aeetic aeio., and wat'er (BAW) 4:1:5 resneetivel:r during three hours. :\fter dr;yting one hour in an oven at. 45 c.. the secono. dirnension. 1mS. the pIe.tes for run in. 210. aeetio aoid. After drying the em:-ornatograms uere observed unden'. •. d3.ylight and, ultra-violet light u1th and ;rithout ammonia (NH3) vapoUl' • Tile spots ;rere reeorded en polY:;;il:;lene plastic and then on paper by their respective colour. A number .'¡as assigned to each spot identifying ene by in eacn ehromatogram. NOTE: It should be made clear that the glasshouse, in whieh the Selanu1l! species for the present study ,¡ere fumigated ui th nieoti.ne.. /. grOl-ffi,. uas regularly.
(23) - 14 -. •. 'v. 1). RSSULTS. Germination ¡~,. ,..,-, ---. ~,-·'·,.v,,·,·,'. -.;--. ~. -. c.I,,,-,,,. ~*-~. ,-, •. , ... -. ~. ~-.-,.-,-",. 2.:ppeers in Table l. Ii'rom the -43 accessions. SQ1rn. •. Yf,. r-"" ,-,.. -~--~. -.. ! ....... -¡-. .:... ..... only 28 appea"r on. T.::..ble 2, beca.use of tus failure in gel"min¿tion of the other ¿,..;cessions. The species S.. OXYC[l.rpum. was. so';~'n. t¡.¡ice but in both attempts,. germination >las unsuccess:t'ul. Other accessions germinated too late and :t'or that reason they are missing in Table 2.. 2). Moruholo,o;ical observations on 1'ield "-nd herbarium. s1J8cimens Although i t lfas hoped that all the species 01' series. .. Conicibaccata might be included in this study, thi8 vras not po"sible dua to the lack of seeds oí the species S. oxycST1JUm, ~. bU8Sii,. l2... 2. .. 2~'y2.cuchocnse,. jé'.en9nse, S. lllU 1 tiflorum, S. noov2.rga,sii, : .. viII uoneta.lum,. and S. 1foodsoniii, "hich are sp:; cies accepted by Hó.,¡kes (1963). An abstract of the morphologic2.1 observo.tions on eo.ch described species, Ó.ppears on Tabla 3. in conjuntion with sorne species of the series l'iurana and Domissa which are 0103 o to sories Conioibaocata. SoLo.num a,c;rimonifolium PLANT. herbc,coous, erGot or. in natural. Fig. l.. Rydb. asce',1din~. to 0.30. m~. tall, but up to 2 m.. h;~bi ta:t.. s'rsu robust, (lngular, green to pUl'ple-tinged, branched or simple. LEAF 18.4 om long (l!'ig. 1) x 8.5 cm broz-d, g1abrous; la.teral leaflets 8 pairs, elliptic-lanceola te to lanceolate, apica11y acute. to ". acuminate and With roundGd obligue bose, sossilo, 2.8-4.5 cm long x 1.0-1.5 cm broad; the 101fermost pair of lateral loaflets aro vory roduced; interjected le,~lets are n~~erous 0.2-1.0 cm long x 0.1-1.7 cm broad, ovate to ovato elliptic..
(24) j 1. ¡. l· J. - 15 -. •. Ta.b1e 2.. list of accessionn tre'3ted in chror.la.tography, and. their chromoGome number.. ~. ..r~U.:. .... J'-'-.-.) 11. 40. 2. COL. H709. 48. 3. COL. IT758. 48. vlILiJ POP;\yAN. 72. ~~. 4. .. m:n:BSR. .. ,. •. C1-301,~OSO¡~ "\. 5. COL. PASTO SIB.. 48. 6. FLH. L.L.2196. 24. 7. LXS. PI283088. 24. 8. MSP. PI230:517. 72. 9. PCJ. PI246971. 24. 10. PCJ. PI 246971. 2. 24. 11. PLL. LL1676. 24. 12. PLL. LL1676.3X1676.6. 24. 13. PLL. 5103. 24. 14. SAN. H663. 24. 15. URU. H712. 24. 16. VIO. 4436. 24. 17. VIO. 4474. 24. 18. VIO. 5040. 24. 19. VIO. 5042. 24. 20. CHI.!. H101. 24. 21. Cm.!. H686. 24. 22. CHM. H723. 23. TUQ. H759. 48. 24. TUQ. 10. 48. 25. TUQ.. C.C.C.2547. 48. 26. TUQ. C.C.C. 2547.1. 48. 27. IMM. H703. 24. 28. ClR. L.1.. 24. ./. 24.
(25) •. - 16 Observation on herbarium specimens. PL)J!T up to 2.5 m. tallo. lonG, glabrous. to pubsscent. Lateral leafleto 7-9 pairs, elliptic-. l;cnceolate to lanceolate, apically acute to acuminate a.nd ,Ti th base rounded to cune,,,te, oblique, 2.8-12.0 cm lono; a.ud 1.0-4.0 cm broad,. sessile or very short petiolate. The lOHermost. pair is rnilch reduced;. the terminal le"flet ls similar to the :.laterals but gonorally broe.der. Few to numerous interjected lea1'lets, ovate to ovate elliptic,. 0.2-2.5 cm long x. O~1-1~5. cm broad, sccoile.. rNFLOR3SCZNCE pseudo terminal ; peduncles 8-16 cm long, pedice1 articulation at the middle or aboye the middlo; flowers purple.to violet, COROLLA rotato, 2.0-3.0 cm. •. diameto:~.. BClrries g-.'Clen, long oonical, up. to 4.5 cm •. Herbe..rium SD8cimens. oX8.minod:. liÍexico: Chiapas State,·Se.n Cristobal, Las Casas, Corro . .,.. Zohtehui-~z,. path to Los Angeles, at tho highestpoint, 2.800 m. alt., July 13, 1949, J.G. Ha..¡!ces 1019, Herb. J.G. Hal'¡!WS. Guatomctla: Dept. San Marcos, aldea San Andres Chapil, ruta 12 from San !l!arcos te Tejutla at Km,255 frero Guatemala and 8 Km. from San Xarcos. BoloIl c:liffs in almost constant spray from a cascado. P1ants intermittent1y along stolons conicCll, 4.5 x 1. 7x 1.3. SolClnum colombianum. up to 2 m. tall; tubers produced. (c.f.S. nolyadenium); berrios long. CI:lS t1iO. groovedj 2150. TIl.. alt., Nov. J, 1958,. Dun.. PLANT herbaceous branched., T;Teakly erect or spre3.ding, 0.,60 m. te.ll,. 1.20-2.00 in natural habitat, tubor':"bearing. ST,i;¡;¡ round to triangu1cr,green to palo pUt'ple, spcrsely puboscent, straight to undulate narro\', ¡-Tings..
(26) .1. ¡. 1 1. 1 ,I. - 17 .-. •. ;. I. J. 1, ,. .1. ¡. , 1 ;. I. ,I. .~. ¡ ¡. 1. ¡. A. 1 1!. ,,. ¡ 1. 8. •. coi H 709. COL. P2!sto Sibundoy. l. : 1. ¡. ",. S ' . ......o num. ~~S. 2 __ caI Oi':10J.. " ' - Q.l..i..' " ~f 1 ..;0. . eren t access:tons. i n ·~n grOj·.... ~h u. ' e greer.nouse.. 1 j. 1. 1l·. !. L.óAF 14.0-25.0 cm long x 9.0-12.0 cm broc.d; petio1e 2.0-3.0 en leng, pubescent to denscly pub0sccnt . .LatB¡'al leafleto. 4-5 jugE:.te, s0ssile. te shortly peti01u1ato, "Uiptic 1D.ncoo1¿to to bro¿,d1y olliptic, apex acu~inate. to obtusa. b~sG. round to oblique, slightly decurrent en to. the rhachis. InteI'jectod leaf'lct::::: fEnr to nur::.erous, sce FiS's. 2:A-B, 3. FLOfJE:tS llhi te to light pur:ple; podicel ru."'ticule.tion inconstantly at. the r:1iddlo, al' aoove. 0:(". valo1'¡ ~. CO~OLLA. =otatc-?8ntugc~al,. 1.5-3.5 cm. dic.r.l8ter. Bcrries. e11ipsoid-. conica1 to ovate·.conic;::;1, a bout 3 cm lone> ecnerz:Uy g2'''''11 • . Threo variotios. are de:;cribod in the ¡¡pecies S. C010!'1bi2,:l1m:.
(27) - 18. •. •. Fie;.3. ~.. colombiD.num va:r. trianao grOlm. in the greenhouse.. va:r. tri,'?n,:-t9 Bi tt.,. petiolul~te. ~las. or. t.¡i th l'~nccolatG. sessile~. lateral leaflots shortly. and whi te ór. 1~~hi tish. flovlors. V,,:"1X. !!le::-idionrtle. cstablíshcd by Hawlwa on plants o,?llccted in Ecuador ati.d. dif:fering from tne typa s-pecially in the minor numbeI" of int0:rjected. le":..flets, l::;rger corolla t 16J:'goX' s.tyle ,uno. simple ctigma Ha~tikes (1944). Varo zin2quirEnuCl Hm-rkcs is common in the middlo of Colombia, mainly. in the d.0partmont of Cu."1dinam'_'trca.. someti:r.os 3-4, with 2. Ol'. Late¡,~o..l. leaflets. moro intorjocted leaflota. 5. paired. 1. bot¡,¡oon 03.ch pair.
(28) j. j. . 1. 1. •. j. - 19 of 1" teral 1edlets; corolla 1arg8r than in the 1;7po, stigma simple. L::.tin dOGcri"tion in "",,,kos (1944).. 1 -1. 1. Val'. onC2,nu'E. lli th e11iptic 19nceo1ato, short to long acuminate. ¡. le.terc.l lea.flets .::.ud feíi ínter jccted laaflets., Flo-'iicrs pale purple". 11. I-~ OCCi.1rS. I. in soma parts of Colombia 2nd o:f Ecuador, as snmfn later ... !. Var. {;\lambianuOl ,'fith broa.d1y ob10ne; torminal loat'lets, attenuate. 1. 1 I. or &cuminate at the apex, fOl; intcrjcctod 1eaflets. F101fers mauve with. J. i. white regions in centre of pota1s. It occurs in Sorne parts of Colombia.. !. Vé.J.'. Jmoescens it differs frou the other varietios in the dense pubescence. It occurs in Cundinamarca, Colombia. I-Ierborium. Suec:..!]9nS. eXé.mined:. So13.nuTil colo;;,bta:r.um ve:r. t""'ianD.G Colombia: Dept. Narino, Near El. Encano, on road from Pasto to Santiago, Santa Maria, 28 Km. from. l; • I. Pasto, 3050 m. aH. Aug. 30, 1965, J.C. Ha"kos 2544 H'3rb. J.C.Hawkes. Dept. Callca, Para::lO 0.0 Moscopan, 22 Km. from Furaco on tho roao. to. 1 1. 1.;oscopan, 3450 m. alto Ju1y 21, 1948, J.C. Ha'llkes 191.E<erb. :r....a.i.'-----~_. j. llar-rkes.. Var. encanum Colombia: Dept. Na:rillO: corca de el Encano 7 km. en la carretora paraJ.iocoé'., 3.000 m.alt., April21, 1949, J.G. HE.Hkes 605 Herb. ;J.G. ¡{¿nrkes; 4. Jan. de El Enc&no en1'\ carretera para Sibundoy, 2.900 m.alt., July 30, 1950, J.G. Havrkes 633, Harb. J.G. Haukes.. Var. guambianu:n Colombia, Dept. Cauce.: by the old mule track from Silvia to the Pararno de 18.s Delioia.s, abovo Cuambi2, 3.000 m alt., July 18, 1948, J.G. Hé'.wkos 162. Harb. J.C. ITa\'fkes •. .. Ecuao.or, Prov. Imbabur¡o". Lngos San Karcos, Cé'.yambe, 11. 200 í't.,.
(29) - 20 -. • f'_'"l. __ .'_." _.. vv .... v,.. ..J_,_~. 1"0. ~.. ___. ;...:!~~~,. : •• -;.,,',-~;·!,"'-O.~. \f0ste-::n slope va.l1ey cl ::tio San a.:;!ncisco, o8b.ir.cl :-,:onz8:':::'2.te, 2. 9D0 c • -Tune 18, 1948, J ~G. F.a1·7kc~. .alt.". v~... 66. Eero. J .G. Hl:nrkes.. zi nnauir2.l1u:n. Colo¡;¡bi['.: Cuncli:cv1m·:Jxca, ne8:!:'. Bo,~ot2..,. iJestern slop8,valley of Rio San .[.'rancisco, :'!olino ¿el 30(:¡uaror~, 2.700.c al t.,. Paura, 2.350. ;n. Jt:..."10. 02.stcrn cordille't'a, b0t~:reon. !,lon50:zrate anO.. 18, 1943, J".G .. Ha,:'¡kes 67 .. Ec:cb.. aH., Nove:;¡ber 10, 1948 J.C. l:!a,:kos 500. Herb. HaJorkos •. •. i i. 4. 1 1 1. Pig.4. (A) ~. flahaultii Eitt. collccted in Colombio., Cunclin;~rr:r.:.rca, "'o-ch-d .. , ú~ ~a·oJ'~ -n. a "s" 1. 0 (~D\ c: ...""U" '''~U'l :s,4t .. , .... "-" ':;',;.;..1 J"r"'~ v...., .... (1[:") - .... _ , 3 . 3Jí) ..... ~J :::..~ -"" 1.·_:...1L'.··. ,: fro1'\ tho Intc:cre Gion;:Jl Pota/co Collcctiol1 of StUl~D'coa J)c.y, :·!iscor. . sin. ,..,¡.::;.. ... U.S./,\".. ..J.\.o. ...
(30) j. ¡• 1. 1. I. - 21 Bitt. PL/~Kll. Fig. 4-A.. herbaceous, tube::c-bearing. 20-60 cm tallo D;:.' e en. hiA:hl.Y branched. erect. 1. flexuous, densely piloso, hairs up to 3 rnJl long; H'hitish, Hin:;less.. l. LE!Lii'. l. jUg:-.tB, donscly pilosa ¡'¡i th long hairs, petiole 1-2 cm lanS 1 as. , I. odd pinnc:;te, pale green, up to 13. CT.1. long x 4-8 cr.¡ broad, 1-2. l. pubescent. J. 2.0-4.5 cm long x 1.0-2.0 cm broad, oblong elliptic, obtuse at the. ,j ,OJ;leJ>:. 1. 7. and narro,fly winged as the rhachis; latoral lS3.flets. cr shortly acmünate, bo.se round to obliCJ.ue, petiolule up to. mm long or sessil::::, terminal leq,flet 2-3 times hieg-er than lateral. ones, p8ti01ule 1.0-2.0 cm long, inter jected leaflets 0-2. ,". H!l'LORESCSt'!CE. lateral or subterminal, 3-7 flowers; peduncle 3-7 -cm. i. 1. 1. ¡. I• I. long,. den~ely. pubescont, bi-forked; pedicel articule.tion at or aboye. the m:.ddle. CALYX. 4-5 mm long, green, }!ubescent; lobes elliptic 3-4 mOl long,. COROLLA rotate-pantagonal, 1.5-2.0 COl violet or mauve. EGrries. I. ovoid conica!. Horbarium s-oecir..ens examined:. The only speoimen observad ,ras obteined from tea accession L .Lopez No. 2196 gro",m in the greenhouse at 'Ifinterbourne.. I 1j ¡. Sol.:::.num laxissimum PLANT. Bitt.. Fig. 5,-A.. herbcceous, laxly ascendont, 0.60 m. tall, but 0.70-2.5 m. as. Ochoa (1962) pointed auto Tuber-bearing. ST";I". robust, angulate, wi th scattered pubescent, subpigmented, wi th. narrow straight to undulate l-rineS. LSP-F odd-pinnate, 10-20 cm long 7.0-15.0 cm broad, 4-5 jug2te, glabrescent; petiole 1.5-5.0 cm long. Latera.l leaflets 4.0-6.0 cm long x 1.5-2.0 C8 broad, ovate lanceolate, apically acute, and base scmicordBte occasionelly obliquo¡ potiolula 0.3-1.5 cm long, Tha two 101·rermost pairs of lateral leaflets smaller. than thC3. other;.
(31) -- 22 -. •. B PLL. LL. 1676. • FiC;. 5. (A) S. l,é~.z:i.ssi:.".um :Si tt. g:;:'o-:-m in gre811~OUSG at Hint0roourne from sGeds ootail'led. i":;':;o::l tha Int0rrc:":.'icnal potr~to CollGction, Stt:.rgcon l3.-:l.y !,'[i3co~13in U.S.A. (13) S. l)illa.:'!u,:o,tGn~;e Vargas, dono.ted by Dr. Vaxgas in Cuzco Peru. - ~ .. torminal. lAa~lot. about the same shape and siza as tha lateral ones,. ,Ji th petiolulo 1. 5-6.0 c;;¡ long. Inter jectGo. lcaflets 3-8, ovatel<3.Y'.. ceolato, 0.3-1.3 cm lon,g x 0.2 x 0.8 cm broad, subsB:ssile ~ pseudostipul,U" le :;.ves soraiov¿;,to, 1.5 cm long. ::fFLOR¡;;SC¡;;;~C3. ter;;¡inal anO. l2.ter,,-1 as Ochoa (1962) ino.icateo., 10-40. flo:-rers, pua.uncle 7-10 cm long, bi-tri-for'wa.j pcd.icels articulate at thc mio.dle. C;\LYX. 7. ffiül. 1':]112:, 101:.:)3 lanccolato, abruptly apiculate..
(32) - 2} -. 1. 1 •. I. CO.'OLV\.. ¡. ~cccrdins. I. both zilles.. 1. substellate, bluish, purple, }.0-3.5 cm diamoter, acum:ms, tc Correll (1962), 0.5-1.0 Cm long, coarsely pubescent o.n. 1. Slla.ringlY pubescent. Style 0.9-1.0 cm lo:.:JS', stigma 3ubglobose.. J. lons-conical, green, 2.5-3.0 cm long, as Ochoa. j. He:-b2rium sU8cimens. !. B8::;ri~z,. (1962) pointed out.. exar.;i~. Only one sPGcimen seen in not very good cond.ition, "ith several. 1. cenic".l berries, 1.5-2.5 cm long x 1.2-1.6 cm diameter, plenty of. !. SG3ds. It is said to be from Peru, Huanuco, Carpish, cumbres cntre. j. Huanuco y Tingo Maria, 2.100- 2.900 m. alt., I·~arch 1947, R. F,erreira. 1942.. 1. ,I i. 1. •. Solanum rcoscopanum. Ha,¡kes. PLANT. decumbent. herbaceous. STZ~í. ascending flexuous, 0.60 m. tall,. 5.0-7.0 mm thick, branched, lons pu::,escent 01.' hairiness up oí the. stem, green, rounded. 1. 1EAlf~:. I. 01.'. tuber-bearing.. 1. II. Fig.6.. ,rith narron straight wings.. odd pinnate, bright dark gTeen, 7.0-12.0 cm la.1.'g x }.5-6.0 cm. broad, 3-4 jUliate,.densely pube3cent especially petiole 0.5-1. 5 cm long, purplish as ·the. tae uppermost. leavos,. rhachis, pubosccnt, lateral. lea.flets 1.0-5.0 cm long x 0.7-2.0 cm broad, elliptic 1,s.nc00late, to semiacute te obtuse at tus apex, round to. obli~ue. 1. oblong-lanceolate. 1. at the base>, shortly petiolu2ate or sessile, the lowcrmost pai1.' of. 1. 1 J. lateral leaflets very small; terminal loaflct tend elliptic, lightly ¡;¡¡;¡. to be oblong-. longer than lateral ones, with petiolule 3.0-12.0. 10ili..:n inter jected leaflets 4-6 on the rhac¡üs and. 0-2 on the. petiolule 01 the termin9.1 leaflet; pseudostipular leallets semicircular, about 1 cm long. Ili?10RESC¡;;HC8. lateral, 5-14 flo-,rered, pedUjlcJ.es 4.0-16.0 cm long,. t ..rice forkad, scarsely pubescont, pala. pigruented., :[ledicels 1.0-2.5.
(33) 24 -. 1 1. ¡ 1 '1. 1 1. 1 1. 1. I• i. MS? el. 1. 230517. ¡ I. ¡. Pig. 6. ~. TL!OGcop"·~num Ha\',~1(e8, eTo~m from seeds obtained f'ron Intorregionul Potato Collection, colleetod by Oenoa in Ecuador.. pubescent, articulated u:t or -belorl the middlo-.. CALYX. purplish.. 7.0-8:0 raro long, lob0s 4.0-5.0 cm long, acumans 1 mm. lon. ~.. COROLLA. rota. te, flattened or eoncave;' purple to light lavendor,. 2.5. cm diumeter.·. ANTHZRS. 6-7. mm long, filamont. Berries ovate conical. less than 3 cm. long. l-lc-.:'Drlrium suocimans oxamined:. Colombia: Dcpt. Cauea, Paramo de !;1ozeollan, S:ln :.!a.rcos, 32 Kr.¡, !':;:om Puraco }!(J.I{kes.. 2.920 m. alto July 21, 1948, J.G. Hal-rlccs 192,193, Herb. J.G. I.
(34) 1. j. !. - 25 SOl,:-~l'l'Uc':1. Pig. 4-B.. Bitt.. herbaceous, 10-40 cm tall, 1l0ak1y erect al' spreading, tuber-. PLJ',..NT. 1. u.J.ucijuc;um. bc;::xinS·. >fhitish hairs, round, green. L3:AVES oud-pinnato, dark-green, 4.0-8.0 cm long x :'.0-5.0 cm 'uroad,. 3-5 jugate, shortly pubescent, petiole 0.7-1.3 cm long, sparse1y pubescent as the rhachis; lateral leaflets 2.0-3.0 cm long x 0.8-1.5 cm broad; elliptic to. elliptic-Ianceo~e,. obtuse at the apex, shortly. petiohüate to sessile, moro or less decurrent on to the rhechis of the leaf, decreasing in sizo to tha lo,¡ermost. j 1 j J. 1 •. leafla'~s,. terminal. leaflet elliptic to broadly oblong, apically acuminata basally cuneate, ' "ritb potiolules 0.3-1.1 cm long, longer tban lateral onos; intorjected leafl~s. 0-8, rounded 1-3 mm long; pseudostipular leaflet elliptic,. oblique, 0.3-0.8 mm long. INFLORSSCZNCS. pseudoterminal al' lateral, 3-6 flowered, paduncles 1-4. cm long, pilose, twioe-forked, green, pedicels 1-2 cm long, articulated at the middle, piloso, slonder and bent. CALYX. 4-6 mm long, subpigmonted, pubescent; lobes olliptic, 4-5 mm. long, aoumens 2 mm long.. COROLLA rotate, violet to purplish, 2.0-2.5 cm dip.ffiotor, acumonz 2-3 mm long, triangular. lanceolate-oonical, 4.0-4.5 cm long, filam0nts 1-2 mm long,. &NTH&~S. glabrous.. glabrous, 6-7 ¡¡m long.. STlLE. I. II ¡ ~. _,/". BSRRIE~. as Correll (1962) pointed out, ovate obtuso at apex, light. green i1ith darker lines, 2.0 cm long. HGrbarium. soecimen exarnined. It "as observed only living specimens.. plants due to tho 13c1<: of herbarium.
(35) - 26 -. •. nillchulltcnso. -3,,1~nuCl. horbaceous, 15-50 cm tall, tubor-beal'ing.. PV,Wl' S~'=l-:. thin, branched, glabrescent, pigmonted, . i th na:l'row straight to. ¡,¡ndulate. w~nes.. L3f.. VES odd--pinnate, dark 2,Teen, 8.0-18.0 cm long, x 4.0-9.0 cm broad,. 4-5 jugate, glahrescent petioles 0.1-1.2 cm long, scarcely puhesccl1t as the rhachis; lateral leaflets 2.0-4.5 cm long x 0.5-1.1 cm broad, oyate lanceolate, apically cuneate; rounded to oblique or semicordate baSe:lly, petiolule 0.2-1.3 cm long; the lC'irermcst t.lO pairs. cf lateral. leaflets are smeller than the uppermost pairs, terminal le::cflet similar to lateral ones, petiolu10 1.2-1.1 cm long; interjected 1eaflots 0-6, oyate 1ancoo1ate; pseudostipulsx 1eaves 1anceo1ate, pseuuostipu1ar leaves 1::cnoeo1ate semiouryed. I¡\?LOc¡ ssc Jo:liCE. •. terminal and lateral, 5-10 flo"ered, 10-18 acoording. to Ocho a (1962); peCluncles 2.0-5.0 cm long, t,rico forked, .. i th sc~ce hairs,. subpigmented; pedice1s up to 1.1 cm lone, articulated at tho middle. 4 [Jm lon&, pig;nented, 1fith lanoeolate greenish lob:es, acumens. CALT,í.. 1-2 mm lon&, glabrous. COl:¡OLLA. rotate-stellate, Hhitish intol'petalar tisuo, star greenish,. lobes lilao, 2.5-2.8 om ¡. ¡ .. diameter, acumens triangular, 3-4 mm long,. glabrescent. ANTHSRS. lanceolate, 6 mm lon&, filamente 1. t.lt.l. long, glabrous.. no exsert on tho anthers, stigma sUbglablose, thicknor than. S'EYL8. tho style. BSRP.IES. oonical, 1.5 cm long as Ochoa pointed out.. The isotypo examined in Professor Haukes's. herbari~~. is oompletely. different from the living plante, thus confusing their idontity, althou¿h tho accession ( one berry ) was given to the prcsent author.
(36) ,. i j. 1. - 27 -. I. 1. •. I. by the author of tua specios in 1Jruba:-:b2., Cuzco, Peru. H,Jrbarium $1)3Cincn eX2.mined. ,. ?cru, Dopt .. Cuzco, prov. P¿uc2.rtarabo, entre paso de Aguila y. ;. --. .-. ~. .. l.- ~ ..l. .J.,.c;;,.ll u.c.... 1. ,j. "' ..... ,. ".... (J,., ..... ""."'-V'-'. ¡~.. ..:... ......... ). ,. -. -. ,-, ,- ,,',",,"':>~. .... ..:.-. ~. Herú. J .G. Hcl'.-rkos. Solé:..nu~. 1I 1. PV,"JT S'l';;;",. s2.ntol:::Ülae. Vargas. herbc:ceousO.60 cm tall, l=ly ascendent, tuber-be3.I'ing. slender, rounded to triangulate, branohed. at the bó:se,glabrescent,. green, m;o.:rbled l1ith purple at the base of branches; 'Tings narrow. undulate to stxaight. LEAF. odd-pinnó:te, dark-green, up to 22 cm long x 4.5-1.3 cm broad,. 3-5 jugate, scú.tter8d pubescence, petiole up to 2.5 cm sp=sely pub.:oscent lateral leaflets 2.0-7.0 cnl long x 1.2-2.3 cm broad, elliptic-lanceolo.te, shortly acuminate at the apax, rounded to oblique 1 I. , I. •. at the bO',se; petiolule 0.4-1.5 om long; termine.l leaflet about the Sé.me as lateral ones lIith petiolule 0.6-3.0 om long, interjected loaflets. 0-7, oblique-elliptic up to 2.2 cm long.. IN?LOR~SCZ¡'¡CE. pseudoterminal and lateral, 7-10 flo1{ered; peduncles. 6.0-8.- cm long, t1'1ice-fork0d, shortly pubesoent, subpigmented; pedicels 1.0-2.0 cm long, Qrticulated at or above th0 middle.. ¡. . COROLL~. rota te-pentagonal , 'fhi te, purplish dorsally.. 1. BER?IES. longly conical pale-green, 2.0-2.5 cm iong.. j. l. ¡ 1 i. The h0rbarium sp9cimen examinad has thin stem without ramification a.nd glabrous; lec.ves.:t. 10 cm long x 5.7 cm broc,d, shortly pubescent; 3-4 jugde, petiolG 1.5-4.5 cm long, lateral leéifJás harro1'11y la.ncoolate long-¿cu::!inate, 2.5-5.0 cm long x 0.6-1. 3 cm broad, petiolules 1-4 mm long, none to 4 interjected le2.flets, Hhite flOI/ers, pedicel articulation at or 'above the middle, conical Berries (1.0 Cm lonG, inmature)..
(37) ,. 1. j-. I 1. ,. 1¡ , 1. 1. ~. 1¡ ,, ,. ,. 1,. j. !I ,.... 1. -~. 1, 1. 1'1. Ii'i,~.. 7.. ~.. sc:.ntolé::ll2.B ¡·lintorbournc •. .. SAN. H 663. Vargas, grOHll in the gX'8oenhouse at.
(38) 29 nerb~'.rit:m S12.CCirlH~n. cxa¡;;ined.. Peru, Dept. Cuzco, prov. ¡. i. ¡.;:w. P~uc~tambo,. Pillahuata, 2.700 m. alt.,. 9-10, 1945, Vargas 51090.. Flora ol' Peru.. PU¡·iT. herbaceous, 0.60. ffi.. tnll, 'V.:p. ~o. 3 m. es ill8nticned by Ochoa. (1962). STZI'¡. thick, to 20 mm diameter, J'lexuos asce::lding, branched, pubescent,. v.'een to pale purple pigmented, ,.ingless. LEP~. 1-2 jugate, up to 25.0 cm long x 22.0 cm broad, bright. densely pubeGcent, petiole 2.0-8.0 cm long pubescent as the. d~~-green,. rhachisj. lateral leaflets 4.5-10.0 cm long x 2-4 cm broad, acute at :the apex; base rounded, oblique, oblong-ovate to oblong-elliptic, petiolule. 0.2-1.0 mm 101',<;; termina.l leaflet oí' the SE,me size as .J. late~'al. ones, or. l=ger, uith petiolule 0.4-2.6 cm long, without interjec"i;ec1 leaflets • IwnORSSCE:i'iCE. lateral to terminrll, 10-14. floHe~'ed;. peduncle 5.0-,8.5. cm:,densely pubescent, twice-l'orked; pedicels. pubescent, up to 3,5. "'. C"1. articulated aboye, ªt or uelow the middle. 6 mm long, lobes 5-6 mm long, oblong to elliptic, acumens. CALTÁ. triangular, 1-2 mm long, purple, pube"cent. COROLIA rotate, 2.5-3.5 cm diameter, purple, acumens pubescent with whitish hairs • 1U~THERS. S'I'YLE. 7-8 mm long, J'ilaments glabrescent, 0.7 mm long.. 9 mm long, barely eJCsertedj stigma globcse, slightly thicker. !. than the style.. ,i. BERRIES. i. \. ¡. ,I. ¡, !. ¡• I. I. ,/. ovoid-conical. Herbarium s"O.ecimGns exa.mined. Peru: Dept. Cuzco, prov. Urubamba, km 94-104 Pampackahua to Sta. Rita, bordering reilrOad. tracks, 2.200 m. alt., !.íarch 9, 1944,. C. Vargas 4158, Herb. J.G. Hauke3..
(39) - 30 -. .. 1. V lO 4436 B. l' l. }lir;. 8.. ". , i. ,!. ¡,. s.. VIO 4474. violn.coim:'!.rmora.tum Bit-:;, g¡."o1,m in the grocnhouno at Z00d.G colleotcd by HaHkes 0-: 2011. in Bolivia, Dopt.. ~'¡i~tGrbo~no fro:n. Cochabamba .. prov. Ch"'pare, Kr.l 104 on road :'rom Cochabamba to Chapare, aud Km 100 for A snd 3 rospectivoly.. :! :. Fig.8. PL!üIT. herbaceous 0.60-1. 20 ro; tall, up to 3 ro., as refered by Cor:¡,'ell. (1962), tubor-bearing. ST~:,:. j. sléndG:;:", flexuous. --. ... ascend" n.o:, 4-8 nu~ ~lab~osce~t , ' "-:-,n'.; - - c'.'c, round ,,,,.'.
(40) - 31 -. I. /. @;r~-. . F'. A. I. Jl. .,. \. 9". §.... VIO. 5042. l· I. vio12"Ceií;V.:~::'f,iO~~"2.t:ln -Bi tt", g-rO¡,:n. •i. in the graenhouse at i'ro¡:c S00QS collected by H.?lrk8s ct a11. in Bolivia, Deüt. ¡:'a Paz, prov.Nor Yun::;as, road fron Unduavi Coroico. (A) anormal plant, (:5) froEl norm&l planto. j. "pUJ.~ple ~iG'mented. Fia". ~lintGrbournG. l·. to. to t1é..rbled, wi thout. fro;;¡-. ~.¡ir..gs.. I. LSAVES odd-pinnate, dar-green, 10.0 ...18.0 cm long x 6.0-10.0 cm broc.d,. I. pubescent as on tho rhachis, lateral loaflets 2.0-5.0 cm long. 1. 3-4 jugate slightly pubascent, rctioles 2.0-5.0 cm long, short. x 1.2-2" 2 era bro,;.'.d, bro.:1dly-elli ptic to ovate-elliptic, acu.:.inG. to z..t the apoX,. ~.¡ith. corda.to to. terminal loaflet sligh,tly. o~liquo ~'TidGr. base, potiolule 0.3-1.2 cm. tl1an' lo.tera.l ones or the. vrri th petiolule up to 1.9 cm. lar.,:;; ínter jactad. le~.flets. lo:-;g~. S:J.mo SiZ0,. 1-5, brOQ.dly.
(41) - 32 oblol'.g, pseudostipular leaves scmiovate, cbout 1.5 cn long. IH~lLO~-{~SCE:JC:S. ,. i'. ¡. ~. up to 10.0. .,. CJ:\. lateral and pseudoterminel, 8 or mo.:-e flol'rered; pcduncle. long, t1·rice-forke:l., subsla;,rous, purplish; pedicals. 1.0-2.0 cm long, shortly pubascont, articulatad at Ol.' balow tha middle, gradually sl1011en to tha calyx. 6-7 mm long, pigmentad, pubarulent, lobes elliptic aoortly. CALYX. acuminate.. ¡ , ! , i, ,i. COROLLA. i. rotate pentagonal, víolet, 2.0-2.5 cm dlam., acumens. trían¡:;ular, 3 mm long shortly puboscent. 6 mm lone, f:iJam,mt 1. mm lone.. ANTHr:ns. STY1E. slendel.' 8-9 mm long, stigrna short, thickor than style.. B&~IE3. conical olongated, 2.5 cm long, as Correll (1962) poínted out.. Two of the. • -j. ., l. i. accessions (VIO 4436,4474) are abTee with the. descl'iption. The other accessions (VIO 5040,5042, and 5042 anormal) are smaller plants, stemangulate, straight to undulate winged; leav.es dense~y. pubescent, corolla rotate-stellate, calyx three and twíce-. labiate, appearing quite diffel.'ent. to considere toem as the same. species.. :!. ¡,. ~. AH of them were collected by Ha1·rkes. 1. ,!. ,í '1. ,. ~. ill.. as fol101its: VIO. 4436, Bolivia, Dept. Cochabamba, prov. Chapare, Km 104 on road from. I. ,. Cochabam'oa to Chapare and Villa Tumari, 20 m.. i. alt, February 17, 1971. VIO 4476, as for 4436 'out at Km 100; 2.550. :i. be~ore. Hotel, 3025 m.. ;¡. i. 1. 1. I 1 iI ,¡ 11 1I. I !I i. ;I. I. Iq •. m. alt., February 19, 1971.V10 5040, Bolivia, Dept. La Paz, prov. Nor Yungas, road from Unduavi to Coroioo at 6 Km. froo Unduavi¡ 3275' m. alt., i\pril 4,1971. VIO 5042,as fol.' 5040 but 10 Km Unducvi; 3.225. ffi.. alt., April 3, 1.971.. HcrbC',rium sD8ci.mens eXDinined. .§.. vi012.ceimccrr;¡oratum Bitt. var uani1losu¡;¡, Eolivia, Dept. Cochaba.mba, Llanta. . hduana, neo.r Coloni, 3.000 m. alt., February,. •1. ,1. 1946, Cardenas 3637, Herb. J.G. HaTikes.. /.
(42) ,. !. ¡ 1. 1!. •. J\. T2.o1e 3. Korphologic8.1 cha:racters of. 30m3. of the species. 1,. oí series. Conicibaccata and . B3ABING. relat~d. series.. 'T'''~-'''''. ".~.', ~\.. AGF. 'Erec::;.. or o.scending. S~-L.';'P3. SIZ~-cliatl.. yes. Rotate. 2.0-3.0. Bright purple to violet. yes. Rotatepentagonal. 1.5-3.5 cm. ;;hite to light purple. 0.25-2.50 m.. COL. ':feakly erect or sprcading. COLQUR. 0.60-2.0 m. Erect flBxuous. yes. 0.20-0.60 L'CS. La..'{ly é15cendAnt. yes. 0.60-2.50 m. MSP. PCJ PLL. Flexuous ascending or decum'oent. yes. l'feakly aract or spre5.ding. yes. Laxly ascendent. yes. Laxly ascendent Flexuou3 ascendent. Rote.te. Rotate. yes yes. Rotatestellate Rotatepentagonal Rotate. ,. Flexuous ascendent. cm. 3.0-3.5 cm. Violet or mc.uve Bluish purple. 2.5 cm or more. Pale blue to lig:ht lavender. 2.0-3.0. Violet to purplish. 2.5-2.8 cm. j;jauve .Star lrhitish. cm. :'ihi te Do:::,sally Purplish. 2·5-3·5. Purple. 2.2-5.0. Illue purple to violet. 1.8-3.0. cm. 0.60-3.00 m. VIO. 1.5-2.0. cm. 0.60 m. URU. Rot2te substellate :i;lattened o:::, concave lobes. 0.30-0.70 m. SAU. Rotatepente.gonal. yes. Rotatepentagonal. cm. CID;. Eract, ascending branched 0.80-1.00 m.. yes. Rotate to rotate Stellate. 2.0-4.0. Purplish or violet. TU:?. Erect. yes. Rotate. 2.0-3.0. Pale to deep purple. Rotatepantat;onal. 1.5-3.0. ;rhi te. Rotate to rotute-substellate. 2.5-3.5. 01'. decumbent. 0.30-0.80 m.. pent2_gonal.. nm. Erect or ueakly ascending 0.40 m.. yes. IOP. Erect 0.80 m.. yes. cm Purple.
(43) • ----------~------------------------~-~~--------------------------. PLInS Ol?. L3jüI'L~'r. LA'V:,,,AL. SIL~r~. Cent:=c:..l. 0'"1: aboVH. Zlli~Jcóic. lnnceolate to. 7-9. l:?~nceol.3.te,. e. . cute to long c:.cumin2.te. Similar or slishtly. Feif to numarous. broador. CentrC\.l, above O:' boloN. EllipC~ic. lrmceolat>;> to broad.ly elliptic, acute to obtuse. 4-5. Largor to sli,:sht largor. Nona to numerous. 2 sizes. Above centre. Oblon¿:¡ elliptic. 1-2. Largor. Absent, rarely 2. C.8tlt:r-al or aboye. Oyate-lanceolate. 4-5. Similar. 3-8. Cantral above or balo1v. Elli~tic-lanceolate. 3-5. I,arger. 4-8. 3-5. Larger. 0-8. •. Central'. to oblong-lanceolate. . Slliptic to lanceolate, obtuse apex. Central. Ovate-lanceolate. 4-5. Similar. 0-6. Central or. Ovate-lanceolate. 3-5. occasionally larger. 0-7. Oblone oyate to oblongelliptic. 1-2. Similar. O. centre. Central or belo,1. Broadly elliptic to oyate elliptic. 3-4. Smilar or sligh i;ly lGI'ger. 1-5. l'.bove ce'ntre. Oyate elliptic to elliptic lanceolate. 3-4. Similar. Absent or several. Aboye centre. Ovate lanceolate to elliptic lanceolate. 3-4. Usually larger. Few. ContrGl or aboye. OVGte lancoolate to elliptic-lanceolate. 4. Smilar. Few. Ccntrnl or cbovo. ElJlptic. 3-4. Larger. Absont Ol' vory fe.. aboye. Aboy" or belo::;,.;. to many.
(44) • -,'-~~------. GL,rjrous. -----------. l'Ja-cro",,¡-conical. 3.0-4.5 cm 19.. glJ.brescent. Broadly ovataconical, 3.0 cm lone;. Shaegy. Ovoid-conical. Dense to. 0u083cent. .J. •. Sp[;..rsely to e;labrescent. Lone;. conice.l 2.5-3 cm 19.. Sp¿:rsely to denso. BI'oadly ovate conical, less than 3 cm long. Short. Blunt ovoid obtuso at apex. Sparce. 1.5 cm long ovoid-conical. Scattered. 2.0-2.5 cm long oV2to-conical. Dense,short, shinillG. Ovold-conical. Sligllt. Conical-elongated-. General1y 2l1abrous ,. Ovoid to ovoidellipsoid. Sc-c,tterad. Ovo id. about 3 cm long.. hc~ir$. Glabrous oI' sli8htly piloso. Ovate-oblcng. G120rcscent to -"iloso. Conical. ./.
(45) - 33 -. 3) t. GhrO';',10S0l?e. cour.:tin:;;. All the accassions .-rere examined in their chromosome. ¡. number erith the results. J. .. i. 1,hich appe,-..r on Table 2.. Chro!:n:to,r.::r.:::ohv of nhe11012.c cOE'!l')ov.tlds in lCG.ves. !. A mast.er chrou1é>.:"'og:c-am of' p'b.enolio extracts from leaves Ol some 01 the species of the series Conicibaccata and related series is given in Fig.10. All the spots >lere recordad by means oí a numbar,. assi~led. ultraviolet. in sequence, and identified by their colour under. light. (uv) and ultraviolet +ammonia vapcur(Wv+}TH3) a.nd. .. their position in each glass plateo A total oí 43 spots were. registered in the species Ol the series studied, froru which 33 >lere found in Conicibaccata., 3 new on8S in PiurllUa, 5 in llemissa, and 2 in. eircaeifolia. The grand total Ol spots >rith their respective colour is reoordad in Tabla 4. The presonce oí spots are represented by + and the absenca. .1. I. by _. The colour of each observad spot. is represented by the. f0110lfing letters: BL = blue, BN. = hr01m,. D. = dark,. FL " fluorescent, GR = ,greenish,. L = light, pur)? " purple, Y " yelo'iT. Tus spots were not cuemically identificd. Soma of the cOffil'ounds aJ:'e prosent in almost all tue samplas,. ! -i. ¡ :t :I I. !. I. such as spo·l; numbcr onCl 1'¡hich appcars in 32 of the somples,. ,. 01'. 94~b. oí the "hole chromatoé';roms; as ~lell as piule spat (No.5) uhieh is registered in 28 cases, 82';';. Others are present in only one oí the samples. .§!..z..,. spot nu;¡¡bers 19,29,33,34,35,41,42, ond 43. The purple. spot numbcr 21 oí the spocies. ~.1Jauci,iUF(u:n,. aud. ~.pillahu,,,tense,. 34. aud 35 af the species S.chomatophUum (H 101), were observad on daylight but missing under UV and 15'11' + Thi¡ last. mB.. tuo eolumnes oí tho Tablc 4. indicate tlle total oí.
(46) - 34 every pos:itive spot occurring in tila 34 chromatograms, and the p ..]rcentage, cO~3idGr'i~'lg 34 spots as 100%.. The chroID\::.tog-.cam3 wore run 2 or 3 times using. ne",{ le:.vcz frola. or ::-cfuting tilO clouotful sperts .It . . ¡as intaresting to nota. tnat the. main spots. (well-marked) tlere pe esent and the s8condary ( weak), ,lere present or mis3ing fro:n one or e.nother cb.rometogram. It. }lE.S. alzo noted. that chromatograms froro difforent plants oí tho S3.me accession prod.uce idontical or different p2.tterns. It may dopend on the degres oí inbreeding 01' eaen aeeession. From Ta.ble 4 i t is found that ~.. choClatouhilum. H 101 and H 723 have only 5 positive mat.ching and. 10 difíersnt spots, whoso Coafficient oí similarity (Cs) isO.33.This resul t seems to be very spacies.. ~.. 101!. sinee both s&mples are. 01'. the sama. tuauerronse gave aproximately uniform patterns in eaeh. runo 'rhe samples 01'. ~.. iOUQtG.lum gave similar or dissimilar. chromatographic patterns aecor:iing to t he morphological difforentiation bet¡.¡een them. To \L'lderstand tho relationship within and. bet¡'leen the speeies of the Table 4, the Coeffieient of similarity (Cs) was ealculate for sample pairs, expr6ssed using tilo formula: es: ~ p + d explained by Sokal and Sneath (Ising and Fr~st, (1971) "rhere p= positive matehes (number of spots(+) pr83ent in both taxa or ehromatograms). d = number of Spo"", not ComIDOl1 (difx'erent matches). ~.&.. Comparing the chrom2.togl'aJ1lc... 1 uith 2 (COL H 758 and COL Pasto. Sibundoy respeetively ),1'10 find that p= 13,d = 1 henc8 p. +n d. 13. = 14""=0.9 2 •. The 11011 mo.rked spots as ITall as the weak, h2.ve bee!'. treated in. the same way to simplify thoir analysis to get the Coeffieients of similari ty. In~der. en. to illustrate tho results, a dendrogram was construetod. based Coefficients of simi12rity. in Table. 5..
(47) ., , '. j ;. - 35 .-. ,. 1 ,. ,. ,1. .. ,. (). ,,. J : ;. i. .¡ ¡ ,;. ¡. J. I. ¡1¡. 8 lw. 2. ". ... U. o;j". c'. ~ q. ca. 22. • :I Fig. 10. l~D.G t c:'. ¡¡cetíc. acid. 1. ~ .... ama t 0"'·-'''' IJ.r· 0.1.: 0 .lo> ""h ,J o ..." ")'-.'-"'01' ... ... ..... "~'" .l.. J.ro"', ,. 1 or·vo"· ~ .... <..J.... .leon:Lci"o.:lcc"4.~ oxt·,J.,(.~ ... ,..,c-,Lo..;! .t:'> ... -.... o :.o:. ::::0:.10 / I,¡ e 8-:100-;0" oP t."lO $o::::"ioc , .... ,,(.ó. nnQ rQl~tcd <:'!'":'r; e~. e .,. t". ,.. .... ...--. ..... 0::...
(48) - 36. •. Table 4.. Chl'oraato~Tnms. ol leo,f lohenolic Gxtractions. of SoLmum spccies.. .. N O. .¡.> {I). 1:'rl lA O. ~. O. p:=! ••. u.v.. 2. + + +. + + + +. 4. 5 6 7. L. B1.. Pink. 11. BL. L. E1. EL. D. 3L. L •. 3L.. 12. L., EL.. 13. L. BL. 3L. BU. B1. L. Y. D. D. L.B1. Purp. L. BL. EN. 31. 3L. EL. EL.. 18 19 20 21 22 23 24 25 26 27 28 29 30 31 32 33 34 35 36 37 39 40 41 42. 43. ,. 1. D.EL. Pink L.BL. FL. B1. L.BL.. 38. .L. O. 'OT u.u.. 17. 1. FP.3. L.BL. FL. + L.E1. FL. +. 14 15 16. •. • ~. O'M Otll. L. :aL. L. EL. L. 3L.. 10. -;~. C() ' . ' 1, "\. 0('... 1 2 3. 8 9. ,. 0'. O::'::. COLOUU. D.. Y. D. D. D. D. Purp. Red EL. EN. EL. L. Y. D. L. Y. D. D.. GR.,. El". D.B1. L.3L. L.31. L.31. D.B1. LBN. D.BL.. :J. : r). 01:'-. +. +. u~. + + + +. il. + + + + + +. + + + + + + +. + + + + + + +. "'" ·,ó"' u>o. .'. ~. C\. ~. (1). (\j ~'. '0. '". C\ r-,. o. r',. 3. 4. 5. 7. 8. +. + +. + + + + +. +. +. + + +. + + +. + +. 1--1. P<P<. +. +. +. +. + + +. + + +. +. +. +. + +. + + + + +. Y. 31. 31. 3L. 3L. Y.EN. Y. LEN.. + + +. + + +. Y. Y.EN. Elack ./. N. OH. ~ ~. P<P<. + + + + + + + +. I. •.BL.. Y.. ,". ::¡. +. BN. Y.EN L.:8L. FL.. L.Y. 3N. BN.. .C-!. rl 1:'-. Ap.... ~n:. + + +. + +. Yo. D.3L. Y.1li~ D.3L. Y.. n,. ." N. 'd rl. rl t'-.
(49) -. 37 -. T·:¡ble 4. (Continued. ). .. '0. -o ::0. 'D. ~. ., .. .,". >-'ji><. ~(;). P<,-¡. 9. 10. ca. In. z(d Ul N <'-D XH tI)~. COLOUR. u.v" 1 2 3 4 5 6 7 8 9 10 11. ti. ,.. "j. "i. 12 13 14 15 16 17 18 19 20 21 22 23 24 25 26 27 28 29 30 31 32 33 34 35 36 37 38 39. 40 41 42 43 I. ¡. • r ,. ¡ I. !. ,. -,.. 1m3. C'-. rl ..... ...J ~-,. ,-.;:.. , ,-,. rl. ~~. ~;:¡. '" p..lr\. ~~ p..lr\. H. 11. 12. 13. 14. 15. 16. -O. '". '!). O. O r;,. O-~. ~""" ~~ -"". L. BL. L. BL. L. EL.BL.. L.BL. FL. L.BL. FL. GR. D.BL.. +. +. +. +. +. +. +. +. Pink. Pink. +. + + + +. + +. +. + +. +. L. BL. BL. L. BL. EL. D. BL. L. BL. L. BL. L. BL. BL. BN. BL. L. Y. D. D. L. BL.. L.BL. FL. BL. L.BL. BL. D.BL. L.BL. L.BL. L.BL.. +. +. + + +. + + +. + +. + +. + +. + +. TI.EL.. Y.K", D.BL. Y. BU. Y.BN. L.BL. FL.. + +. + +. +. +. mT •. BL. BL. BL. BL. D. Y. D. D. D. D.. D.BL. Y.BN. D.BL. Y. Y. L. Y. EN~. BU.. +. +. +. +. +. '. -. + +. + +. +. +. + +. + +. +. + + + +. + +. +. + +. +. + +. +. +. +. + + +. + + +. + + +. +. +. +. + + + +. Pill'p.. Red BL. EN. EL. L. Y. D. L. Y. D. D.. +. ,-. +. +. L.BL. Y. BL. BL. BL. BL. Y.13N. Y. Y.BN. Y. Y.BN. Black. +. + +. +. Purp.. L. BL.. + +. + + + +. ..,.. ,,. -.
(50) .. 38 'l'ab1o 4. (Continucd). (\1.-j. (T. ,,. g. >:.). g¡:;. '-". :;:l?\. -- -'.~. :>. 17. 18. 19. 20. 21. L.BL. FL. L.BL. F1. GR. D.EL.. +. +. +. +. +. Pink. Pink. +. 1. EL. B1. L. B1.. 1.B1. FL. BL. L.B1. BI" D.B1. 1.B1. 1.B1. 1.BL. D.BL. Y.BN. D.BL.. --- -;..!. '-'. ,-\. r.J. o. ,~. n,J .,~. >l?\. d. ,~. ... .~. J.'~. ::;¡. ';. L ..... ,~ ~.. :...< (:; r.:-;. rl. >:.)~. - ~.",\. ,. o;:. l. ..... o~. ... .¿:. rü. '.. ~. •. H~. (J', , •. ~.. ". ~. l-t. lo. tI:. ". c~. "~=>. U,\. 8(\1. COLOUR. Ir¡:¡ 3. U? V.. 1 2 3 4 5 6 7 8 9 10 11. , ~J.. • ",. 12 13 14 15 16 17 18 19 20 21 22 23 24 25 26 27 28 29 30. 3:l32 33'. .,. • ,. L,. :L. BL. 1. B1. L. E1. E1.. El,.. D. B1. 1. 3L. 1. B1. L. B1. EL. BN. BL. 1. Y. D. D. 1. BL.. Y. D.. D. D. D.. 34. Purp.. 35 36 37 38 39 40 41 42 43. Red BL. BN. BL • L. Y. D. L. Y. D. D•. 23. 24. 25. +. + +. + +. +. +. +. +. + +. +. +. +. T. +. +. + + +. + + +. +. .,. + +. +. +. +. +. + + + + + + +. + + + + + + +. +. + + +. + +. +. + + + + +. -!-. +. + + +. + +. + + +. + +. Y. BN.. Y. BN • T. u.. B1. FL.. Purp.. 1. BL. BN. E1. E1. EL. B1. D.. 22. -. 1.B1.. Y. E1. EL. EL. EL.. Y. EN. Y. Y. BN. Y. Y. EN.. +. + +. + +. + +. +. + +. E1ack. '. + +. D.BL. Y.BN. D.BL.. Y. Y. L. Y. EN. Elf.. ,,. +. +. +. +. +. +. +.
(51) - 39 ~Pablo 4~. (Continued). o o", ;:'j. ,,". i:OÜ\. [-j 0J. ce. 00"'. C'r1 rl\. ~.. ~-. "' ,""' ,"...... o H. o oH oH H. C'-. O.-lN\. " ".. r ic, o ~. 0J H. (7\. N\ ~. id. >-". r1 lf\ ¡.~"\. ,>.. o. H. COLOUR U. V.. 1. L. EL.. 2 4. L. EL. L. :3L. EL.. L.EL. FL. L.EL. FL. GR. D. 13L.. 5. Fink. Pink:. L. :3L.. L.EL. FL. BL. L.BL. EL. D. 13L. L.EL. L.BL. L.Bl.. :> 6 7 8 9 10. ¡. '*. 1. 1. f¡. 1 ,. ,j. ,i. ; i, i I ; 1 .¡. '. ,. :¡ '\. :\ !!. ¡¡. !1 ¡ I. ", I 1j \1. 11 !I :I, ,. d.. lL. EL.. L. EL. J3L. D. 131.. L. EL. 12 L. BL. 13 L. BL. 14 BL. 15 B1~ • 16 BL. 17 L.Y. 18 D. 19 D. 20 L. 131. 21 .Purp. 22 L. EL. 23 BU. 24 BL. 25 BL. 26 BL. 27 13L. 28 D. 29 Y. 30 D. :'>1 D. 32 D. 33 D. 34 Purp. 35 Red 36 Ill. 37 BN .. 38 BL. 39 L. Y. 40 D. 41 L. Y. 42 D. 43 D. 11. ,J. HH3. D.. BL.. Y.BN. D.EL. Y. E¡'¡ • Y.13N. 1.131. FL. L.BL. Y. BL. EL. IlL. 131. Y. BN.. Y. Y. BN. Y. Y.BN. E1ack. 26. 27. 28. 29. 30. + +. + +. + +. + +. + +. + +. +. +. + + + + +. + + + + +. + + + + +. + + + +. + + +. + + +. +. + + + + +. +. +. +. +. +. +. +. +. +. +. +. D.BL. Y. Y. 1.Y. BN. BN.. ,.... H o. :'>1 32 33 34 +. +. + + + + + + +. +. + + + ~. +. +. +. +. +. +. :. ~. E-<. ~. 32 16 8 13 28 14 23 10 20 26 24 24. +. 22 9 6 3 4 1 9. +. 5 +. +. 9. +. +. -. 2 2 5 13 14 6 1 9 4 5 1 1 1 2 5. + '+. + +. +. +. +. +. .... +. + +. + + + +. + + +. % 94 47 23 38 82 41 67 29 58 76. 11. +. +. D.Ill. LEN.. H. +. 4. 4 3. + +. 1 +. +. 1 1. I. 70 70 32 64 26. 17 08 11. 02 26 14 26 05 05 14 38 41 17. 02 26 11. 14 02 02 02 05 14 11 11. 08 02 02 02.
(52) ~~~~~~lA",~,~ "---~. '-...... IIJOTECAAGROI'tC\l\R:A DE COUlMlI1.. Tablo ":,>.. 01' tb-J ;3P0Cio;1. o. "1. 1,. coof'ficlvnt; oí' 31e11' rley l'or ~:;oma o:t' ~-:.~rio~J e::>nJ.oiü:l.Cr)',.t8 i,ald rol"tlvo s~:::r 10;J •. n-:~tl"i7; ~l". 2 -~.. tb.:). O ----. ;2. ;¡J. 0.92 ") ()'>. 0.46 0.,16 ". (. 0.66 0,::/) 0.64 0.29 0.41 0 .. 35 O.2'~ o. :n 7 o~ 33 0.33 o.;?'¡ o.:n 0.31 8 0.27 O.tiO 0.23 0.19 0.20 9 o.. ,:!~~ 0.50 o....wJ o.:n 0.:26 10 o.n 0.5> ol!.·:~7 0.33 o./n. 0.5. 11 O.Z:? O.2,} 0.:23 12 0.11- 0.19 0.15 13 0,,,6 0.33 0.:~3 14 0.14 0.17 0 .. 14 15 O.~10 0.47 0.41 16 0.·12 0.47 (1.41 11 O<il/~6 0.)7 0.35 ¡ ..) 18 ü ....,<,. 19. 0 • .:10 Ü·.3d. 20. .... )'-'. {\. '. ¡.. ',). "" ,""' 0. ')-j. 0.25. 0.13 0.23 0.1;'. 21 0.35 D.47 22 O.i~l 0.50 OSI 23 O. :):} 0.50 0,,[)1 24 O~35 0.33 O,37 25 ().35 0.43 0.:17 ,.,.0.33 O.l~l 0 .. 33 .,.{ 0;33 O.Y) O#,}l ". 2'0 () .¡~(j 0.50 1\ -. . ;1 ,") " 29 O.¡~() 0.5,) 0.,;-) 30 0.33 O.;~.\) 8.35 31 0.43 0.55 \.') .,.,.'17 32 0.30 0.26 'd,. -L'~,J. .~I"-. 0.33. 0.35 0.22 0.37 0.;)0 0)7 " 0.15 0.:;;0 o~23 0.26 0,13 0.19 ,,~. O.G:). 0.56. OS} () ~ ',jO. 0.43 0.46 0.42. O$4;..~. 0.40. 0.3J. 0.53. 0.:.11. o.~~. O+~j 0.33 o.,f) 0.35 U.19 0.33. 0.22. O.4~2 0.30 0.13 OS3 0.33 0.33 0.30 O. :.0 0.513 0.55. 0.33. 0.29 o. ,':3 0.:23 0.')7 \;.35. O,,4?. o_ ;20 o,n. 04;>J. 0.1,1. {'. 'J~. ,,/. :.,,1 Uli35 0.35 0.37 0.15 O••:~Q c-l.J.\lI ..e_,", !\ """J9 O./r) ,) . '7 .... ') a 32 Oe~~l 0.31 O.:::) 0.35 0.35 V'*o'Jl 0.16 Uo·,:J , 0.3) 0.33 O.-;1 0.37 0.19 0,33 O.3G 0.·,,7 0.}0 ;) q :':0 0.22 09¿o;~3 ,; . O.. :.~5 D.~.~3 a.3a ¡} .. 5~~) 0.53 ('oF. "7 O. jG 0.33 0.5') O~;2G 0.;;:2 0.23 ~. ~,-. ."~. ... ry. ~. 0.42. O.5~~. O.;~7. 0 .. 24. O.:''{. 04~-::"O. o. 33 34. O.,U 0 •.\3. o. COL 1:709. 0"1.6 0.43 0.2} ;)" "i1 0 • .36. 0.31 0.1'/. 0.3'). 0.30 O ,. i_, ')').. 0.20 O.::!,) O•.U 0.10. 0.3:). o• .3:). O. y). \,) 11. :> .. "le,? PI 230517 4 ... n~p. ,IUd PO'Q:'y,,<n. 5 .. PUl LL 2196 7 o PCJ PI 246971. 8 .. PCJ .PI 246971,2 9 .. SAl[ n663 10 ". L\~J. PI 28308:3. 11 '" PL1 Lr_ 16'{6.JÚ676.6 12 .. PU, LL 1676 /. 0.13 0.13 0.25 Ü .. :~~3. o.:?6. o. :~{:~ U.U. O,lO 0.1; 04;~O. '7 OolllG. ).."). 1 '" CO.!.. H753. 0,03,3. 1 .. (;0. r) • '# t:'. i~ .....'. lO. -'i_~ J. (j~;-. 0.:/0 0.:)3 0.13 ~). :::) 0.12 o.. jO O.:>;) . . ,t' 0.33 0.33 0.43 0.5:\ O.}7 O• ..\1 l'·J."-O 0.44 ..... ,c;.,) 'c'" 0.29 0 • .11 0.11 0.26 o.:n 0.12 0.16 0",05 ('1. Ooa'5<1. 0.53 0»'33 o.::..\) ;).53 O.:)) 0.2) O .. ~22 0.31 u.50 :). )0 D.37 Ojl.}7 v. ~" ,'j';) 0.;:1 \.,, _/~'7 jQ 0,23 o.,iQ ü,33 0.53 0115:· 0.33 r\ '-J'" 0.;,0 0.16 0.18 o. :~?5 0.,)1 VIilI'--,V 0.16 O.:~l 0,0';) ,).43 0 •. \) 0.16 OG1:J 0,.26 0.15 '.).23 ()"j1 0.:22 o.f.~6 0,','6 0.,;29 0.23 O~31 0.22. 0.53 0.46 o. ;~3 0.53 0.46 0.2). 0.56 0.46 0,5:> 0.57 O.YI 0,:0 O.-'l2 O./¡G 0.,)1. '''''''''''----'--. 12. o.Y?. 3 4 5. •. ll.. 0.71. 2. •. :r.. ...'i. -1. 1. o.?O. ;).19 0.17 0.17 ü,;~l. 0.10 0.13. 0.10 ),10.
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