Alstroemeria: A Charming Gcnus

Andrca l'vlariel Sanso1, Marta Camargo de Assis~ and Cecilia Carmen Xifrcda3

ICOJ\ICET. Insti tuto de Botánica Darwinion. Labardcn 200. C.C. 22. 13 16-CHYD San Jsidro. Buenos Aires - Opto. de Ecologia y Genética Evoluti ,·a. facultad de C it:ncias Exactas y Naturalcs. Universidad de Buenos Aires. Argentina ~DIBRAPNCNPM (Centro Nacional de Pesquisa em Monitoramento por Sat.:lite) Av.

Dr. Júlio So;~res de Arruda 803,13088-300 Campinas. SP, Brazil ·'C! C. Laboratorio de Etnobotánica y Botánica Aplicada. Facultad do: Ci.:ncias ?\atur:1ks v

t-.Iuseo. Univer<idad de La Plata. Paseo dei Bosque s/n. La Plata. Argentina ·

Kcywords: cut flowers, pot flowers, landscape use, biodi,·ers ity, phylogcny.

Abstract Alstroemeria (Als trocmeriaccae), the Incas lily , an cxclusivcly South

American genus comprises ncar 90 species ranging from Venezuela (3° 1'\orth) to Ticrra dcl Fuego, Argentina (53° South). A/stroemeria species and hybrids havc "aincd world-wide importanee as cut flowcr erop due to its attra ctivc flowers and thc cxccllent kecping qualit)· of thcm. Neverthelcss, they a r e also s uitablc for growing in pots and for garden and landscape uses. Thc rccent ta xo nomic rcvisions of the group for· Brazil (Assis, 2001), Ar·gcntina (Sanso, 1996), and Chile (Baycr, 1987), tire biosystcmatic rcscarch (Aagcsen and Sanso, 2003; Sanso and Xifreda, 1998, 1999, 2001, 2003) and the clrrornosome s tudics (Sanso and Hunziker, 1998; Sanso, 2002) lrave facilitated tir e idcntifica tion of thc taxa and irnprond tir e basic knowlcdgc of tir e genus. Tire nomcnclature of the spccies, their gcographical distribution, ecological clraracteristics and tire interspcci fic relationships are now better understood. Many wild taxa, that are sca rccly known so far, carry genctic traits tlr at open possibilities for improverncnt and devclopment of more attractin new \·arieties. For example, some of tire Brazilia n taxa have a n im portant potcntia l valuc in br·ceding prograrnmcs beca us e of their showy flowcrs and lhe rigid )caves, instcad of tire very tender foliage tira! posscss most of tir e spccics; some P atagonian oncs havc rather· short plant hciglrt, wlrich is important for pottcd pla n ts. Sc\·cr al spccies o f tire wild populations are nowadays tlrrcatcncd and with risk o f cxtinctiorr. Most of tire species occupy reduccd geographical arcas. Specics that inhabi t plain regions are vulnerablc becausc those ficlds are burned and cultivations are introduccd. P a tagorrian spccies cspccially necd to bc protccted , against thc grazing o f tire animais. Tire cultivation of tir e rr a tive plants would hclp to thc ex situ conscr·vatiorr. Rcintr·oduction in some natural arcas might also play an impo1·tant role in ordcr to maintain thc landscape valuc.

Rcsumcn .-Jf.çfml'mrrio (i\lstmrrnrriarrar), <'i lírio clr los In cas. cs rxclusi,·amcntc

sudarrrcrica no y compr·ende cer·ca de 90 cspccics distl"ibuidas desde Ven ezuela (3° Li'i) hasta Tierra dei Fucgo, Argentina (53° LS). L as cspecics c híbridos lran ganado importancia mundial corno flo r de corte dcbido a sus atr·acti,·as norcs y a la excelente calidad de las mism as. Sin embargo, cll as sou también ap rop iad as para ser utilizadas cn cultivos cn macctas y cn jardincría y paisajismo. Las rccientcs rcvisioncs taxonómicas d e! grupo par·a Brasil (Assis, 2001 ), Argentina (Sanso, 1996) y Chile (Baycr, 1987), los cstudios biosistcmát icos (Aagcscn y Sanso, 2003; Sanso y Xifrcda, 1998, 1999, 2001, 2003) y cromosómicos (Sanso y Hunzikcr, 1998; Sanso, 2002) lran facilitado la identificación de los taxa y contribuído ai conocimicn to b:ís ico de! géner·o. La nomenclatura de las espccics, sus distribucioncs gcogníficas y cara cterís ticas ecológicas, as í corno la s relaciones intcrcspccífi cas son a hora mcjor comprcndidas. Muchos taxa silvestres, hasta ahora poco conocidos, portan rasgos genéticos que amplían las posibilidadcs de mcjorarnicnto y dcsarrollo de nucns y

Pro\. \·Ih IS on Ncw Flor. Crops Eds: A .EC. Tombokuo and G.~l. Di:ls-Taglia(.·oao .-\CIJ ll,>n 68.1. ISHS 2005

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m:ís at ractivas variedades. l'or ejemplo, algunos de los taxa l> ras ilcros ticncn un importante va lo•· potencial en programas d.e mejora miento debido a sus .vistosas flores y a sus hoj as rígidas, en lugar dcl fo ll:tj e n~uy fino que P.osee 1:• mayona de las cspecies; algunas especies pa tagónicas tienen. bap altur~, caracter tmportante para su cultivo en macctas. Diversas poblaciones stlvestres es tan actualmcnte amenazadas y en ricsgo de extinción. La mayor p~ rtc de las especies ocupa áreas ge.ográficas rcducidas. Aq uellas especies qu~ ha bttan . !lanUJ·as ~on vulnerable~ debtdo .a. las quemazones de esos campos y ~a mtroduccwn de cultivos. ~as espectes p~tagomcas neccsitan ser protegidas, cspccta lmentc ~ ~ I past~rco de a n_unales. E! .cultivo de las plantas nativas ayuda ría a la conscn ·acwn ex sttu. La rcmtrod~cc!on en algunas áreas naturales podría jugar un rol importante para cl mantenmu ento dcl valor paisaj ístico.

INTRODUCTJON Alstroemeria (Aistroemeriaceae), the Incas lily, i~ an cxclusively S~uth A!11crican

genus, rcprcscntcd by around 90 spccics th:ll occupy a d_tvcrsc :angc o f bal>ttats, lrom.sca levei to 4500 m altitude. They are herbaccous pen:nnmal~, wt th flcshy-ro_o_t tubers. Thc !caves are frcqucntly rcsupinatc, thc inflorcscenccs tcnmnal and umbch torm and the flowers usually zygomorphic with pattcrned tepals. • . . .

Thc rcccnt taxonomic revisions of the group lar Braztl, Argcntma and Clule, thc biosystematic research (Aagcsen a~1d Sanso, 2003; Sanso_ and Xifreda, 1998, 1999, 200 I , 2003) and the chromosome studtcs (Sanso and Hunztker, . 1998; Sanso, 2002) havc fac ilitated the identilication of the taxa and tmproved the baste knowkdge of thc genus. The ·nomenclaturc of the species, their gcographical distributions and ecological characteristics and the interspecific relationships are now bctter u~dcrstood. The Cl~ilcan species monographed by Bayer (1987) are 31. _Sanso ( 1996) .rcvte>~ed the Argentme~n Alstroemeria spccies and reco&nized I O ta_xa w~th _one en~emtc spectes. Rec~ntly, Assts (200 1) distinguished for Braztl , 38 spectes dtstnbuted m the castern reg_10ns of the country; 35 of them occur exclusivcly in Brazil and ~en o f them, wcre descnbed as ne~v spccies (Assis and Mello-Silva, 2003). The spectes from Paraguay occur also 111

Argentina and Brazil (Sanso and Xifreda, in prep.). . . Because o f their showy flowers and the excellent keepmg qualtty o f them, thcse

plants have been successfully introd ~ced into cultivation a~d are used as vase flowers. In Europe, werc introduced as early as m the. century XVI. Smcc those_ days, severa! works of improvement have been carried out wluch has bccome the Inca lt ly m a very popular cut flower. These cut flowers crops has been developed through hybridization, primarily among the Chilcan group, particularly A. aurea Graham, A_. ligtu L: and A. pelegrina L. and one Brazilian species, A. ps illacina Lch_m. In Hollat~d ts one ol t!1c most marketal>le 11owcrs (Griflin, 1995; Buitendijk, 1998), JUSI as that m Japan (Mtyake et ~1. , 1989). Recently an intcrest in growing Alstroemeria as a pot plant has appcared tn U.S.~ .• lf0lland, Dcnmnrk. 1t should hc possihlc to incrcase the variation and enhance the quallly of the commcrcial cultivars by including a largcr part ol thc gcncttc vanattun luunJ among wild specics. Many othcr wild taxa. that are scarcely known so far, an: a_lso of potential use in l>recding programmes. Emphasts has bccn takcn _to take spcc1cs of Brazilian origin with potcntial ornamental value and plants of low hctght for use ns ncw pot plants.

RESUL TS ANO DISCUSSION

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i j

f ~ ' l ~

Distribution . . . l lts northern limits are southcm Venezuela and north-west Brazi l (3-4° N) wlule ti J

cxtcnds as far south as Ticrra de! Fuego in Argentina (52-53° S), with two main ccnters o f ., distribution, onc in Chile (extending into Peru,_ Bolívia a~d Arge~tina) and the s~cond throughout Brazil, Paraguay and Argentina (F tg. 1). _Nattve spcctcs_ ~ccupy a clt_verse range ofhabits from high Andes to marshy lands. In Chtle, Alstroemerw ts geograplucally

6-l

distri llll tcd from a lmi tud~ of_22• S, near :rocopilla (Region LI) in thc non h (.·1. r.::.:>acula Phil.). whe:e thc. vegctatton ts sparse, wtth ,·ario_us spccics o f cacti and othcr :-;,·[,,ohqic p! Jnt~. to ) I • ~ 11~ th~ ~~uth (.-1. patago~n~·a l~lul.). lhe central ~c!:!ion ,,f Chi!,· b3s · thc grcat~stmJm~cJ oi spc~J~s~ South,of the__,6 la~lludc thc numbcr oi spcc1c:s is limitd to A. aurea. thc r .ost \\ tdc spt .:a~ . spccJes m Chtle, and A. preslwna 1-Ierb.(Or,:.urd and Kristi:tnsen, 1998). The Braztltan spectes occur at di fferent habitats: forc:st. s:n·;:~nJ . hioh lields . n~arsh, "campos rupcstres" and '_'caatinga", from a height o r 300 !ll in th,· .·\maz~n up w 2.J00 m ~~ Sara do ltattata. Wllh excepuon of A. tsabellan:t Herb ... -1. -,:.:rdneri Bakc:r. :1. apertiflvrt~ Bakcr, _A. cunha V c! L. A. srenopetala Schenk. A. psi((acf~:: 1 anel A. /vngil'l.l"ia Schcnk, nwst spcctcs h ave a restnctcd distri bution (Assis. 2001 ).

Char:u:ters 1. Sul!tcrranca n Organs. In Alstroemeria, the specics have ci ther a slwn. nodosc rh!zonH.:, as for cxat~!p~c. tn :1. pyg_maea I krb. or: more frcquc n_tl y, lhe rhi/om.: is Jong (F1g. 2A). l_n some ~~~.:c tes t!le rhtzomcs run honzontally, and 111 cach !!rowi1;,: ,cason produc:c: aerwl veg_c wuvc lcaly shouts and, usua lly, onc nowcring stcn1• ·tho.:so.: ;[,,lúns or runncrs ~llow raptd shoot. growth by means. o f vcgctativc rnultiplication. Tr:tJ ition,lllv propagattOn o f Al.w~oemena for horttculturc ts clone by division o f thc rhizomc;. Young roo_ts an: thtn and l1brous but some of thcse t!1ickcn apical ly to bccomc: cylin.:lrical 0~ fustform. >~ hHc. o r cream-colourcd, starch contammg storagc organs. Thc tl.:sh,· rc'Ois can be cnlargc~ as til ~ .• pslflacma, ± cnlarged as in A. aurea andA. paragonica (Fi~ . 28) or slcndcr as mA. pltshana_ . . Thcy may bc dJrcctl y grouped next to thc rhizomc: :ts in A. ape:tiflora (Sanso and Xtlrcda_ 2001: Fif\. 2A) andA. pygmaea (op. cit.: f i:: . IDJ or dtstmctly separate fron_1 the rluzome as tn A. patagonica Phil. (J7ig. 20) (S:u~so and Xlfrcda, 200 I). Accordmg to !3aycr ( 1998), A. gram!nea Phil., an annual plant. d,'CS not d~vel_op storage r~ots. Me_erow and ai. ( 1999) descnbed a ncw species from o,, i ás and Dtstnto Fed~ral tn Braztl, . A. hygrophila Meerow, Tombo!. & r. \ \". Me,·. ,\·hich appa~cnt ly dtd not form thtckcncd roots cither but !ater it was synomyzcd. \\·ith A. /ongtst;4a and obsen:cct that the roots ~re also cnlargcd (Assis, 2001 ). 2 .. Foltaro and Caulmar AJ_Ja~omy. 1 he st_em pres:nts a single-laycred epidcnnis with th!ckenm9 s o f ou ~cr and pencltnal walls, thmncr anttclmal oncs and sunkcn stotmta \\·ith thick cullcu l ~r nd~es. Closed and eollateral vascular bundlcs are arrangc..: in an atactostclc, \\ 1th 1-J large metaxylem vessels surroundcd by parenchYnHl. Th~ central bundles are larger than the outer ones (Fig. 3A). Mechanical tissuc· as a ú'ntinuous schlcrenc_h):matous sheatl!. bounds to the cortex, prcscnting gcncralh· 3-6 !J\·.:rs of charactensttc short cells (1-tgs. 38 , C). · ·

Resupinatc kaves are characteristic for ne~r ly ali mcmbers of thc !.!Cnus. The !caves are t\nstc~ nc:ar the ~asc or along thc lamina. hcnce thc aba."\ial lcaf surfacc becomcs the functtOnally adaxtal surface. A comparativc sun·cy o f leaf anatom\· ha; bccn pe_rformed by Sanso and Xtfreda ( 1998; 2003 ). Thc m~~or charactcristics ar.:: bnina th tc~ness betwecn 4 to_! O mesophyll ccll la\·ers, two typcs o f eh lorcnch,·ma disirít-ution Uúi'~J.r~nlJai 111 _mu~t ~> l th~ ~1/sirucmeria spcci<.:~ anJ isulateral in sckctc:(i on•> a; ~0111~ Braztl!an .spectes \l·nh ng1d l ~aYes. Few species exhibit irregularly shapcd. lobed chlo~cnch) ma ,cells. and many tnte_rccllular spaccs as in A. iyabel/mw (Figs . .J.-\-C), f>?SStb!~ an :v.aptalion to hydrophily. Closcd vascula r bundles organiz..:d in ;mnllcl dtsposttton are. usually surroundcd bt a distinct parenchymatic shcath l'r with schlercnchymattc cells extendmg as gt rdcrs IO\I·ards both cpidcrmiscs. 1\nc'mOc\·tic stomata, usually su n~en, occur on onc or both surfaccs. ldioblasts with oxalatc rnt,hid.:s in lhe mesophyll are WJdesprcad in thc genus. ' ~· lnflorcsccncc and Flowcrs. In few taxa. there is an unique flower. c.g .. ·1. pah:gonica. n~f!gmaea o~. two oncs, e. g. A. amima Phil . subsp. \"CJ!ustula Ehr. Baycr (J7ig. 2E ). Thc te al~rs are eptgynous a~d are_ ncarly ah~ays zyg~~JOrphtc due to the posnton o f th<" inner . Ph · I hc z~ gomorpl_ l ~ ts remforced b) the postt10n o f the lilaments. whtch are located m t c lmrcr t f ti 11 · d . I . (f. (lig. , ~ar o_ 1c 011cr_ unng a1:t lCStS -~ g. 2F). In , J. pygmaca :tnd .-1 . pawgonica

- _!)i thc llOI\Crs are actmomorph1c but wHh some ones shO\\·mg some degrcc o f

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zygomorphy. i\lost of the tloral characters havc taxonomic importancc. s~veral of !hem ar~ also of ornamental wcight as thc basic colour, which varies from ycllow, orangc through r~d-purplc and violei to _pink and whitc, thc disposition of it, thc form of thc· tepals and lhe prescncc or not ol spots or streaks on them. A largc variation rcgarding flower colours and patterns is represented. Some speeies have tiny flowers with a diametcr o f about I em, e.g. A. re1·oluta Ruiz et. Pav .. while other sue h as A. magnijica Hc:rb. andA. pelegrína L., the type species o f the genus. have largc showy flowers with a diameter of up 7 em. Some Brazilian specics, as A. ísabellana (Fig. 2C), have more tubular flowers with narrower perianth segments. -t. Pollcn. The pollcn o f ali species investigated is ellipsoid (boat-shaped). heteropolar and monosulcate (Figs. 5A, C, D). Symmetry is bilateral. Size ranges from 50-120 flll1, long axis, x 25-62 pm, short axis. Since pollcn size in genera\ ranges from c. 20- 150 ftm, these polkn grains would be considered to be largc to very largc. Thc cxinc is brcvi-striato­rcticulatc (Fig. 58). The striate muri are arrangcd in a para llcl to sub-parallcl pattern. In some species small, interconnecting muri are apparent between striae. This structurc rcsults in thc roundish to clongatc perforations and givcs a scmitectate cxinc structure. Thc pcrforations are fine, for example in A. gramínea, A. angustifolía subsp. angustifolía and A. 1rerdermanii Ehr. 8ayer, in A. ísabellana, A. pseudospathulaw Ehr. 8ayer and A. versícolor Ruiz et Pav. the perforations are much coarser. The ultrastructurc of the sporodenn is detai led in Sanso and Xifreda (200 1). 5. Fruit Morphology and Dchisccncc. The fntit of Alstroemería is a 3-valvcd loculicidal capsule. In superior view it is rounded, while in lateral view it is ovoid, 6-ribbed, slightly or notably umbonate towards the apex (Figs. 2G, H), coriaccous and bright light brown when mature. When unripe the fruits are greenish, with six longitudinal ribs. which are joined by a transverse rib below the high, widely conical apex, which is notably marked and more angular than in the mature fruit. So111etimes the ri bs are purple (A. presliana). At maturity, the fruits are rounded, the colour turns golden or clear brown and the texture is papiraceous-coriaceous (Sanso and Xifrcda, 200 I: Fig. 7 A). In A. pygmaea and A. patagoníca, the inmature fruits are covered by leaves. In some spccies, the umbo (apex) is short, 0.5-3 m111 in length, as in A. aurea, A. andína, A. ísabe/lana (fig. 2H) while in other spccies it is more pronounced, it 111ay be as much as 20 111111 in length, as in A. pseudospathulata (Fig. 2G). A. pelegrína, A. presliana, A. pulchra Si111s. The dehiscence is explosive, the capsule splits from the base into three parts. The tluee valves, together with the secds, are thrown a considerable distance. Dehiscence takes place when the fruit is dry. The ste111 remains with three (to six) o f the longitudinal ribs more o r less remaining in the shape o f a claw. Rarely, when the tension is not too grcat, the capsule is not totally dcstroyed, and the 3 valves do not complctely separate from the stcm (Sanso and Xifreda, 200 I). 6. Sccd. In Alstroemería the seeds per capsule are numerous. Thc size ranges from 2-4 mm in diameter. They are yellowish-brown. tan or brown, globose or subglobose (Figs. 21-J, 6A- E). The seed coat is dry and hard. Thc primary sculpturing consists of conspicuous protub..:ranccs, which gi v.:s tit..: sc.:ds a peita te thg. 61'), papillosc o r verrucose appcarancc. In severa! specics the testa is colliculate, for example in A. aurea and A. preslímw. 1-ligh magnification with SEM shows the individual units of the secd surface sculpturing, although basically com•ex, to have considcrable morphological variation betwcen species, some are conical with a wid.: base, for example in A. amlína subsp. renustula, whilc in olhe r spcch:s very specialized structurcs occur. In A. psíllacína. a central cell is surroundcd by 9-1 O pcriphcral cclls (Fig. 6C). The cclls that comprisc these ornamcntalunits. and give roughncss o f the surface o f the sced coat are thick-walled and pitted. The surface of the outer periclinal cell wall may also exhibit micro­ornamcntation and sccondary sculpturing. for cxa111ple, irregular hollows or wrinkles (A. andína subsp. venustula, Sanso and Xifreda, 200 I: Figs. 8E, H), micro-papillate (verrucate) (A. gramínea, op. cit. : figs. 98-C) or S111ooth (A. aurea. A. presliana). A dark brownish-black discoid appcndage, arising fro111 the chalazal region occurs in the seeds of Alstroemería (Figs. 68, E) and the raphe is a shallow longitudinal rib. The thick secd coat

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'sts of 1·ust a f..:,v ccll lavcrs. The seeds are dispersed by a ballistic mechnni s111 . Thc const . • ffi · · l 1 · · 1 r alrnost pcrfect sphcrical sh:~pe guamntc~s c tclcnt propngauon l)' t 1c ccntntup ,,1rcc ' . tcd on th.:: sccd as it ts expdled lrom thc capsulc. In some cases thc ~cc:J coat excr · · 1 1· ·rotubcrances with large surf~ce cells, may have an adhe~I\'C propcrty. r\ stra1g ll. mear ~nd short cmbryo is encloscd 111 the endo~perm near the lll!Cro~tlc (o.c.).

7. Chromosomcs. Alstroemería has spccwlmterest bccJusc ol 1ts largc chromosomcs and · asvm111etric karyotypes. lt may be constdered chroi11?SOI1}ally stable wnh a consta~cy ~~ thé chromosome numbcr, 2n = 16, an unique levei ol pl01dy. 2x. and an asymmctncal complement (Sanso a~d Hunziker, 1998; Sa~so, 2002). A.lth.ough chromosomc s1zcs \·~.ry from specics to spec1es, they are karyotyptcally v~ry sm11lar 111 terms of rclall\e s1ze relationships between the cluo.mos~mes. The proportton ofth: ~otal complcmcnt ?ccup!cd b , the Jargest chromosome p:m vanes bctwccn 21.4 % and <b Vo, wuh thc ~xccpuon o f A. lí~/11 L: subsp. /ígtu (Buitcndijk and Ramanna, 1996). lktwccn spcc1cs. thcr..: JS • a considcrable variation in nuclear DN0 con.tent, thc amount o f C-bandcd hctcrochro111alln (Buitcndijk and Ramanna, 1996; Bllltendijk and ai. •. 199!). and lhe prcscnc~ or lack of satcllitcs on severa! pairs o f cluo111osomcs. Two spectes dtlkn:d by about a lactor o f two in the total chromoso111e length (A. magnífica = 11 6 ~;~m; A. P.l:gmaea =. 22-4 pm). Ho,\·ever, in Alstroemería, bimodal karyotypes occur dcsptte thcsc d1ffcrences tn the total chromosome length. 1t secms that in this group DNA has bccn added to thc complcmcnts without altcring the karyotype morphology too much (Sanso, 2002). . 8. Phylogcny. Combining the morphology, rps1 6 and rbcL data s~ts and analyz~ng u~d~r equal weights, the strict consensus tree shows that A~slr~emerra 1s monophylc11c: ,,·,thm Alstroemería, the Brazilian species form a clade, whtch ts stster group to thc rcst of the genus. Among the Chilean/Argentinean species A. patagoníc~ and ri. pygmal!a 1s thc sister group to the remaining species. Ali other 111inor sp.eetes (A. pse.udo.spallw~ma, A. gramínea, A. dndína, and A. werdermanníí) are .gathered 111 a cladc whtch IS ~hc s1ster group of a cladc consisting of purely Chilean spectes (Aagesen an.d Sanso, 200J). \Vhcn analyzed under i111plied weights the results dtffer fro111 those obtamed .. \\·hen us1ng equal weiohts mainly by placing A. andína, A. gramínea and ..l. 1rerdermann11 as Stster group to the " rest of the genus while the 8razilian spccies are embedded withm the Art!entincan/Chilean clade, still forming a 111onophyletiq;roup. Only when thc rpsl6 data are-analyzcd alone is monophyly of Alstroemería t~ot 111aintained. 13_oth morphology and rbcL data support monophyly of the genus. Nellh~r Als1roen!erw gram111ea nor .·.1. pygmaea e\·er occur in a posi~ion that . 111~kes thctr s~gregalion fro111 Alstrocmerra possible. Monophyly o f thc Brazd1an spcctcs ts supported.m ali analyses. whcrc thc th~ee data scts are combined. Whcn analyzcd under equal wetghts, thc cons1dercd Brazthan species (A . apert!flora, A. í.mbellana, 11. caryophyllaea Jacq. andA. psíllacína) are placcd ncar the root o f the gcnus. The flowcrs o f some Braziltan. A lstrol!m~rw spcc~cs superficially rescmble Bomarea flowers more than the Jlowcrs o( some Chilcan spcc1es do (i.c. smaller and more tubular llowers wtth tht: samc color pattcrn on ali 1nncr tcpals) anel thcre is som.- support fo r thc fkt7.i li;m spccics hrillg plac,-d ncar thc root 0f lhe· f:l'Jl\" whcn ali data sets are combincd as a result or thc influcncc of thc morphologlcal characters. 1-lowevcr, whcn implicd weights are uscd thc 13razilian spccics are placcd among thc Argentincan/Chilcan spccics. which is th~ position f:l\wc:d by rbcL. :'\t prcscnt. monophyly of thc Brazil ian specics seems supportcd. but morphology anel 0:\A scqucncc data disagrce aboutthc position o f the clade (o.c.).

Enluation o f Spccics for Ornamental Purposc . Alstroemería psítwcina, nali\·e from SE 13razi l. i\ E o f Para~uay. L1 mgua~ and NE

of Argentina. A. cmyophyllaea, from 8razil, A. aurea, fro111 Argentma.and Cht!c . ..J. l1gtu. A. rasícolor, A. rel'Oiuta, A. pelegrína, A. vívlacea Pht! ., A. hooken Schultcs. ali fr~m Chik, ha\·e been used, among other, in crossings from ,,·hich sc\·cral commerctal cuJti,·ars ha\·c bccn obtaincd. l'Jc\·crthcless, many othcrs \\·ild taxa. that ar~ sc:Jrc;;ly known so far, are also o f potcntial use in brceding programs. Se\·cral o f thc wtld spec1es carri~s genetic traits that widc-opcn possibilities for improYcment anel dc,·clopmcnt of

67

~1ore attractive new varicties. For example, some o f the Brazilian taxa have rigid !caves, mstead o f the very tender foliage that possess most o f the species; some Patagonian ones have rather short plant height, which is important for pottcd plants.

Sclc~tcd Br~zilian Alstroemeria spccies are briclly summarizcd, thc oncs that have mo~c potent1al us~ 111 breeding programs: A. tsabe~lana (F1g. 2A) is found in Argentina, Uruguay, Paraguay and Brazil. In this count'!', 1s poss1bly the most widely distributed spccics of Alstroemeria, occurring at high • elcvatwn and along the marshs from thc statc o f Rio de Janeriro to Rio Grande do Sul and west to Minas Gerais. This plant can be up to about I ,5 m tall, has rigid I e aves, and pendulous, tubular flowcrs (Fig. 2C), colored pink, orange red, or yellow and tippcd in green. Inncr tepals are stripped and spotted. It flowcrs and fruits in Jan.-Dcc. A. longistaminea Mart. ex Schult. & Schult. f. (Fig. 7A) occurs at semi-shadcd sites in forcsts ~f Pem~mbuco, Sergip~ and Bahia statcs, flowers in May-Aug. and fruits in Aug.­Scp. Th1s spcc1es o f tender fohagc can bc up to I ,5 m ta li . It has pendulous Jlowcrs wilh thc o~ter tepals broader thcn inncr, and its colour varying from yellow grccnish to pink grcemsh. A . sel/owiana. Scub. ex Schenk (Fig. ?~). an odoriferous spccies, 'just as A. cmyophyllaea, }S. shortcr than the other Braz1han specie although can be up to 0,5 m tall ; the leaves are ng1d, narrow and the Jlowers are yellow to orange with dark spots on inncr !c pais. It occurs at marshs o f high fields from Paraná to Rio Grande do Sul states, flowers 111 Nov.-Feb. and fruits in Jan.-Feb. A. foliosa . Ma'!. ex Schult. & Schult. ( (Fig. 7C) is perhaps, one o f thc showicst specics from Brazll. It IS about 0,4-0,6 m tall, the !caves are chartaccus the flowers are about O 4-0,6 m long, and t~e colour varies fr?~l ycllow throught dcep o;angc or red. The tepals ~rc br?adcr tha~ IYJ?ICal of m.ost Braz1han Alstroemeria spccies. It grows in high ficlds of Mmas Gcra1s, RIO de Janc•ro and São Paulo states, and Jlowers in Nov.-Apr. A. speciosa M.C.Assis (Fig. 70) is another showy species that grows on the shadow at forcst of Serra do Mar in São Paulo states, Jlowers in Dec.-Feb. and fruits in Mar. It can be up to I ,5 m ta li; the !caves are Iarge and tender and lhe flowers are also Iarge 4-5 em long., red or yellow colored with the inner tepals dark spotted. ' A. variegata M.C.Assis (Fig. 7E) is another attractive species with crowded Ú:aves and flowers ranging from red to yellowish on outside, and yellow and dark striatcd on inside. This species is endemic o f campos rupestres o f Caldas in Minas Gerais Satates Jlowers in Nov.-Feb. and fruits in Jan. ' A .. plantagi~tea Mart. ex Schult. & Schult. f. (Fig. 7F) grows in "campos rupestres" o f Mmas Germs, São Paulo and Bahia statcs, flowcrs in Dec.-Apr. and fruits in Feb.-Apr. This species o f rigid or chartaceous !caves can be up to I ,2 m tal!. Thc flowers are about 3.5 em long, with the outcr tepals colour ranging from rcd to orangc and thc inner oncs with a yellow background and dark spots. A. gar~lneri (fig. 7G), a planl abOl.'~ 0.3-1,5.m.tall, with ri gid !caves, is v~ry frequcnt al . thc Sa\annahs ofMato Grosso, G01as and Dtstnto Federal, flowcrs and fnuts in.lan.-Dce. I h<: llow<:rs ~izc is abuut 3,5·4,5 Clll long and lhe coluur varies from tkcp rcd tu yd luw rcd on thc outsidc. ycllow and spottcd intcrnally. A . malmeana Kracnzl. (Fig. 71-1) is scarccly found in marshs of Rio Grande do Sul and Paraná states, Jlowc.rs. in Jan . -F~b. and fruits from Jan. This spccics can bc up to 0,9 111 tal! , the leaves. are ng1d, amplex1caulos and the flowers, red, pendulous with the stamcns and the yellowtsh stigma cxposed. A. stenof!h~:l/a M.C:Assis (Fig. 71) is a charming species because of its vegctative charactenst1es. lts he1ght runs abo~t .0,2-0, 7 m tal!. The !caves are narrow linear, rigid or ehartace.us. The flowers are very stmdar to A. gardneri, reddish or orange, 2-3.8 em Iong. and the ~nner tepals are dark spotted. lt grows on the rocky outcrops and on ri verine forest borders 111 the state.s o f Goiás a~d Minas Gerais, flowcrs in Mar.-t,lay and fruits in May.

These. spcctes havc an 1mportant potential use in brccding programs and deservc much at~entwn.. However, therc are. no avai lable studics about pollination and reproductwn, nellher are therc about 111 sllu and ex si tu, conscrvation or in vitro culturc in

68

Jalion to the Brazil ian taxa. re Plant hcight also varies considerably in thc gcnus but thc spccics with Jo,,·cr height are found among thc chilcan or argcnlinc·chilcan taxa. Some of th.:sc specics with otcntial use as pots plants are:. ~ . .

Pl. 1ygmaea, a.small hcrb, 6-1 :> em tall .. grows at J500-.J~OO m 111 lhe Andcan nH'u.nt:tJns ~f ~crú, Bolívia and north-west Arg~ntma. T!1c flower IS not usually zygomorph1c and bri!!ht yellow. Th~ upper leaves grow 111 thc so!I. _ A. patagonica (Ftg .. 2B), 2-14 ~m t~ll , cxcepttonally can be up to. 2) cnL Thc lea,·es are linear and the tem11nal flower ts bnght yeilow or orange, someumes \\'Jth small purple 5 ots on the te pais. J andina subsp. venustula, also a small hcrb, 5- 16 em .tall, o~curs. at 2800-3700 m abovc sca in Argentina and Clu lc. Thc flowcrs .a_re crcct, "!fundt? uht.orm, Wtth ali thc lepals very reflex (Fig. 2E); the outer ones, whtush to ross1sh •. w1th hlac and t~e ap.cx •rcc:nish thc inncr ones, o f severa! colours, Y. ncar apcx, vwlct-ldae, Y.. lower, whttc wnh ; transv~rsal y~l low band and n~ar - ~he basis, whitc, v!ol.ct spols on ali .lhe surfacc, diminishing in s1zc lowards thc bas1s. I hc anothcr subspccJc 1s also o f low hctght. A. pseudospat/11//a~a, anothc.r trans·Andcan spc.cics of Argentina and Chi le •. inhabits .thc Patagonian meseta 111 Argentina. In some cases ts tal.ler than thc above descn be.d specJCs, bctwcen I 5 and 30 em tal!. The flowers, erect and b1g (F1g. 2F), yciiow, somett mes havc dark spots near thc thickencd apex. . A. spatlwlata Presl. , has spatulate and rather suceulent !caves and the flowers. wlth thickcncd pink te pais, are only raised 4- I 5 em a~ovc thc gravei. . . A. garaventae Ehr. Bayer, 5-20 em tal!, a spce1cs relatcd wtth thc prC\'JOus onc but wllh thc !caves linear to Ianceolate. A. 11111bellata Mcyen, 2-8 em taii, flowers o f medi um sizc, pink, infundibul iform with thc ou ter te pais decp-set in the apcx. . . A. hookeri Loddiges, with four subspccies, betwcen 4-1 O anâ 4-22 em tal!. 1 hc colour ol the flowcrs varies among pink, rossish and white with different pattem of stripes and spots on the inncr te pais. . . . . A. graminea, 2,5- 8 em tall, !caves gramm~fo~n w1 th a soht~ry flo.wer; the outer :md th~ inferior inner tcpals rossish, thc two supcnor mner ones, 1/J mfenor. ross1sh and the 2/J superior yellow with dark spots. This speeics with the two ones dcsen bed belo"· occur at Atacama descri. A ll'erdermannii Ehr. Baycr, 4-8 em tali, with smaii and elliptic lca\·es and vcry n:mow ou ter tepals. . . . . A. po(rphy/la Ph;I, 6-20 em taii, !caves w1dc and sessde, flowcrs of medlllm sJzc, rcd, lilac or white.

Thc variation among and wilhin species and thc many possibil itics for spccics hybridization opcns up lhe possibilily for production of a widc range of ncw cul tivars suitcd for various purposcs, sue h as eut flowcrs, pot plants and gardcn and landscape uses . Besidcs severa! of thc wild populations are nowadays thrcatcned and wi 1h ri sk of , ;,;inct i;,,L I\ lo~; oí lhe spcciC's ocêupics rC'dt•<"rd c,eocr;~phi<"~l ilrc~s . Sp('cics that inhahi1 plain rcgions are vulncrablc bccausc thosc fi clds are burncd anel ctdti \·ations are introduccd. Patagonian spccics likc A. pseudospathulaw or A. pawgonica n.:cd lo bc prolcctcd csp.:cially, from thc grazing of thc animais, whil;h causcs. thcir tkc linJi ion. ThC' cul ti,·alion of thc nativc plants would hclp to lhe ex s1tu conscrvauon. Rcmtroduc110n 111

some natural arcas might also play an importan t role in order to maintain thc landscapc valuc.

ACKI"OWLEDGEMENTS Wc thank Lic. Rafael Urréjola (La Plata Muscum, Argentina) for technica!

assistancc and Sr. Yladimiro Dudás for valuable hclp with the figures.

69

Litcraturc Citcu Aagescn. L. and Sanso, A.M. 2003. Phylogeny of the Alstroemeriaceae, based on

morphology, rps l6 intron, and rbsL sequence data. Syst. Bot. 28 (I): 47-69. Assis, M.C. de. 200 I. Alstroemeria L. (Aistroemeriaceae) do Brasil. Ph.D. Thesis.

Universidade de São Paulo. São Paulo. Brasil. Assis, M.C. de. and Mello-Silva, R. de. 2003. Brazilian Alstroemeria L.

(Aistroemeriaceac): synopsis. Postcr prcscntcd at Monocots Ill, Thi rd lnt l. Conferencc on the Comparative Biol. of the Monocotyledons. California. U.S.A.

13ayer, E. 1897. Die Gattung Alstroemeria in Chile. Miit. der Botanisehen Staatssam. Miinehen 24: 1-362.

13ayer, E. 1998. Ta/ta/ia- eine ncuc Gattung in der Familie der Alstroemeriaceac. Sendtnera 5: 5-14.

Buitendij k. J.H. 1998. A cytological characterization' of gcnomcs of Alstroemeria, the. production of interspccilic hybrids, and th.:ir performance during micropropagation. Ph.D. Thesis. Wageningen.The Nethcrlands.

Buitcndijk, J.H. and Ramanna, M.S. 1996. Gicmsa C-bandcd karyotypcs o f cight specics o f Alstroemeria L. and some o f thcir hybrids. Annals ofl3otany 78: 449-457.

Griffin, M. 1995. Problemas para as Tulipas. Agroanalysis 15 (9): 12-14. Hunziker, A.T. 1973. Notas sobre Alstroemcriaceae. Kurtziana 7: 134. Hunziker, J.l-1. 1991. Protandry in Alstroemeria psillacina (Aistroemeriaccae). Polisl\

Botanical Studies 2: 195-198. Hunzikcr, J.H. and Xifreda, C.C. 1991. Chromosome studies in Bomarea and

Alstroemeria (Aistrocmeriaeeae). Darwiniana 30 (1-4): 179-183. Meerow, A.W., Tombolato, A.F.C. and Meyer, F. 1999. Two new species of Alstroemeria

L. (Aistroemeriaceae) from Brazil. Brittonia 51: 429-444. Miyake, I. 1989. Breeding spotless Alstroemeria in Japan. Herbertia 45: 40-44. 0ogaard, M. and Kristiansen, K. 1998. Alstroemeria in Chile. Herbertia 53: 29-39. Sanso, A.M. 1996. El género Alstroemeria (Aistroemeriaceae) en Argentina. Darwiniana

34: 349-382. Sanso, A.M. 2002. Chromosome studies in Andean taxa of Alstroemeria

(Aistroemeriaccae). Bot. Journ. of Linn. Soe. 138: 451- 459. Sanso, A.M. and Hunziker, J.l-1. 1998. Karyological studies in Alstroemeria and Bomarea

(Aistroemeriaceae). Hereditas 129:67-74. Sanso, A.M. and Xifreda, C.C. 1998. Comparative foliar anatomy in Alstroemeriaceac.

Poster prcsented at Monocots 11, Second lntl. Symposium on the Comparative Biol. of the Monoeotylcdons. Sydney. Austral ia.

Sanso, A.M. and Xifreda, C.C. 1999. The synonymy of Schickendantzia with Alstroemeria (Alstrocmeriaceac). Systematics and Geography o f Plants 68: 315-323.

Sanso, A.M. and Xifreda, C.C. 2001. Generic delimitation between Alstroemeria and Bomarea (Alstroemeriaceae ). Annals o f Botany 3 8: I 057-1 069.

Sanso, A.M. and Xifreda, C.C. 2003. Comparative folia r anatomy in Alstroemeriaeeae. Submi ltcd to Bol. Journ. Linn. So.:.

Sanso, A.M. and Xifreda, C. C. Alstroemeriaceae. Flora de Paraguay. Edil. Conservatoirc • el Jardin Botaniques de la Ville de Géncve- Missouri Botanical Gardens. In preparat ion.

70

o·

J

' .. ..... \ .. ~.- I

' ' ~: _,.. .. ~.l ._ ... /

'· I ' I ' ' I . ' ,_, ...... ~f

' .'

., ., .... ,.."),.

Fig. I. Distri bution o f the genus Alstroemeria.

O 500 km

'===---'

71

sl - J

' 'f' G -·-

B

Fig. 2. General aspect. A-B, subtcrrancan organs, A, vegetative shoot, B, reproductivc and Yegetative shoots; C-F, flowcrs, C, tubular perianth, D, more or lcss actinomorph; E, reflex pcrianth, F, zygomorphic,; G-H, fruits. G, with a long umbo, H, with a short umbo; I-J, sccds. A, C, H, I, A. isabel/ano; B, D, J, A. patagonica; E, A. andina subsp. \'enuslllla; F, G, A. pseudospatlwlata. Drawings from Sanso ( 1996).

72 73

..... À

..... 'J•

Fig. 4. Foliar anatomy. A-C, leaf transversal section, general view, A. isabel/ana.

I -- . I J . P . I + r<~,.,__., WJG + IUffP<<cR<~

Fig. S. SEM micrographs of'lolkn. A, pollen grains on stigmat ic surface, 13, dctail ofthc striatc- rcticubte sl!xine. C, D, general vicw or polkn grains. A-C. A. i.wbellana, D, A. psillacina .

Fig. 6. SEM microoraphs o f sccd. A-E, general view, B, E. chalazal region, D, micropilar ~ region; F, detail of ornamental units. A, A. patagonica, B, A. pres/iana, C, A.

psillacina; D-F. A. isabel/ema.

76

Fig. 7. Photograph~ o f some Alstrocmeria species frÓm Braz i i. A . . ·1. longistamin<!a: 13, A. sclloH"iana: C. A.foliosa: D. A. speciosa: E. A. ,·ariegata; F. .·1. plantaginea: G, A. garJneri; H. A. malmeww: I. ..I. stc:noph_dla.