Speciation and radiation of the Espeletiinae in the Andes - Cuatrecasas 1986 by JPZ.

~peciation

an d Radiation of the Espeletiinae in the

An des JOS E CUA TRE CAS AS

CHAR ACTE RfZA TfOf\ J OF THE ESPE LETII NAE

The Espe letiin ae (A stera ceae . tribe Helia nthea e) comp rise a group of abou t 130 species of highly e volve d fto\ve ring plant s. They are geographically and ecolo gical ly highly speci alized to th e pecul iar life cond itions of the north ern sectio n of the tro. pic:.ll Ande s. Deta iled chara cteriz ation s are available in the origin al descr iption of the subtr ibe (Cuat recas as, 1976) , and in the autho r's n1onograph . \vhich is in final prepa ratio n. At this time it may suffice to point ou·t the mo{.)t" outstan~ing differ ential featu res. of the subtr ibe Esperetiiriae \Vithin· the· · Heliantfle ~i-e··: ( T) spira l · phyllotaxis~ · (2) obpy ran1idal shape of the .ache nes, \vhich are , in addit ion , smoo th, \vitho ut striat ions, glabr ous . and epap-pose (with a n1inin1al excep tion); (3) xerom orphi c structu~e and organ izatio n of the bioty pes, inclu ding head s and leaves; (4) fertile fernale rav. flowers and functionally male disc fto\ve rs; (5) basic chro n1oso rne nurnb er /\' = 19. Thes e chara cters separ ate the Espel etiin ae from other group s of the tribe such as tvfelan1podiinae and Polvm niina e, as descr ibed in other \vork s (Ben tha1n , 1873; Hoff mann _, 1890-1894~ S.tuessy, 1973 , 1977; Robi nson , 1981). ~

basis. (_)ther impo rtant taxo nomi c fcciturcs are found in the inflor escen ces and gro\vth forms. Some bf these struc tural \·aria tions undo ubted ly repre sent evolu tiona ry trend s that are influe nced by the extre me envir ontne ntal facto rs prevailing in the north . . ern tropic al Ande s . Thre e n1orphoiogical featu res seem most signif icant in the differentia tion and e\ft®lution of the Espel etiina e: leave s, inflor escen ces, and biotypes. '-

·Leaves

The leave s are usually large . up to 50 X L2 (70 x 25) em, the blade s vario usly oblon g. elliptic to linea r, only slightly atten uate at the base, there fore se ssile~ or atten uate and proxirnally contracte d into a tnore or less distin ct petio le. The tnargin is revol ute. entire or minu tely retno te tooth ed; the venat ion is pinn a te ~ usually pron1inent with a robus t midd le nerve . The sheat hs are usually e nlarged~ ampl ectan t or semia tnple ctant to the stem. mostl v- imbri cated on iC they. are eithe r open and flat, or curvj ng only to embr ace the stem with free marg ins. or encircling the stem and tubul ar. The bl ades mav be . mode rately -trtibesc-enror m-ore·usTJally·a·e nse1Y'iesfe·a ;·-a:rTec.ist on the lo\ve r side , the vestu re is frequ ently very thick , the blade s Janate or seric eous. The sheat hs are usually serice ous on the abaxi al side. Th~ great range of varia tion indic ated in the shape and size of the blade s. venat ion, and qual- · ity and densi ty of indun 1entu m is impo rtant \vhen consi derin g speci ation . The struc tiJre of the lea f base has in1po rtanc e at the gener ic level ; e.g., tubul ar sheat hs chara cteriz e L ibano rha1nnus and Carranzbo a, open sheat hs the other gener a (Fig. ll-1) . Impo rtant trend s of evol~tion in the Espel etijnae are on one hand the trend that has. favor ed the tubul ar type of sheat hs an q, on the other , a -

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CHAR ACTE RIZA TION OF INFR A SUBT RIBA L TAXA

Varia tions in nurnber:- shape , size. color . and kind or densi ty of tricho rnes for every given part provi de floral or veget ative chara cteris tics of taxonon1ic value at the sp~.cific ]eve] and to some gener ic level. Varia tions in leaf oute :-o-~n t at the ..... line and type of indum entum are basic consi deration s for specit!c distin ction s. Some aspec ts of th e leaf struc ture and archi tectu re are impo rtant in gen eric differentiation , and phylo genet ic trend s in the Espe letiin ae may be traced on that

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sid~n}d~. (1) the to\v~r, ve,gefatfve,,"leafy part, ,:and . favoritn::g' HJe {2} the: upper,. fe:rtJle,, floriferous part. vYithin the· .iii1.'btict~t~ kind. . ;o f cover -that ~rote:c.ts the body of monotelic· synfto.r:e.scences' in the :Espeleti.ina.~ I the C(~Lilitosulas;•..Another trend ·is the progressiv e r¢tt~r6ti~on .. oflirHf ·pseudo;petie>le---and·· co:nse.quent :. .,. ·_cdi$tfn·g:tri:s;h ·· two ...,f;){pes:.-·-.(-.J.}··· d.icha~,kll.. .synftqf.:~-s~ · . ;and . centes~ with ·opp:n.si te- bran ch.es·~;··= b.racts.~ ·J?-rot.tcie=fl:;in·E: ot: the Jan1ina base:, be.cominit st~s:s:He·. .

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·i nt1oresce:n te structural system in the. E's:peletilr;q;t~· i~ nlo)toteLi.c, u.Slia.Jiy ccunp.ound ·or, as

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by W:·:· TtoJJ'-' a S:btnllorescenc.e. The t.erm nio.nQ.titliC. \vas· establish·ed by w·! ·Troll for the kin·d of i nfloresCe~·Clt:s· ch.<.tracteri·z ed ey the ··1 '· \""'"' . ;. 't"~'rmi·( l· ·.1 (.. ' 11. m :J. t:•::.d .._,LTJ~()Wtb- ,..) _f: tilt~- '··l'·v.-e.s·. _,.,,:-.h:t":c;t:-11 "tre nate d. or closed by an ap-ical tiower. These .at~ the inftores-cenc,e s us~tally te,-r:zn-e.d definite .· deter.rni1 xi • T ·r· o1· ··- ~ ·;vtt close' . .';!:. ...! : ·' · .Jf\.;:"7 ··::.., . .·...:·1 ·.·,,VY ,c .j .::.. ~~··.. :-~c·)~·s· <2~ · : ·. cv.n :. .. d ..ar1·· na·t,,.,~ ·S.J:..~ ~ ~~ ... 19t , ~)'· • ·.. :t·,r··e·c·asa;j ; t :l ,. ~ ~ C\.. C~,. u~ct. ·1 ::r-~l"T--:~: u ::~ , . frd·. enee tVvo · ZQn.es can be conIn a svnfloresc -. .,

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leaves. · They are :\veil deve.lopecl in E~f)peletia . .fF~g. ) 1,.,;2·A.,) ~ .(.2:J morroGh.a·si'a} ·~·:Y.n.ti.ore.scen~~'CS1 vvith,aJf$:>rrrat.e.:,..br~tfflG'rt:es.:· itJ:t;aets~ und l:eaves. 'fhe-y are· fol1_nd in othq:r gener~l ·\\-·it'h tn'any variati{),n'S., e ..g., EspeletiOtJ~:}is :and Liban.o!lu.un.tnts. This-ty-pe b h . h . . ,... d . .. . . . : J.S· :Constite·~red der;tv;e ·.- . fr.o::m ·r ;e :tb.nn.e.r ...v- t! : .e alterrra.te developtn:ent oJ only .:orre.: of the;: t\vo. elett1ents (Fi·g 11-Z.B ). latetal o~npos'ite . r:' These t\v-o kin.cf~;. of sy;;_n.fior~sce.~J.T9:es are ft.rrrd.~lmenrally· distinct h::r the E.speJetfitHl~. and htiv e ecological a.nd ta·xQ:ITOmic implicatiot1s. The ·p erfect th_yQ.r~ticaJ d.ichasiaJ '·model can be 'h . :o:.f • p . . ".. . . .f ' ·E" :ei:eti/tL~: .: 1 - l)ut t . e .tn.u:tfound Jn rrl~ny ~m:¢,:~-leS .o;. ,.£S,fJ. vid uals ir1 ftiet. o.fte.n. fail t.O develop S'Ome parts .o f the svnflotescence. Variations that do not affect -~·

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iridivi.ct·u~i in ia.te:florescence. Terminal FIG~ J 1--3.' A·.B ...Monocarpic caulirosulas of Rui!ope~ia figueirasii~· A. S·tefi1e individual: rti· ll. ~y.ntlores-· :synfl.:oresc(!nce'" C.-Polycarp"ic··c~u.liros.u.ln:.of£spe/edp ,grall;<lijloiq;.·dhtgranJI11~t'i<:..'in. 1ongti'rudin·ril s·e2tion: Axifhtf~rthite .·. . --

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CUAT RECA SAS

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its basic struc ture and svtnn .. 1etrv., are those relate d to the numb er of pairs of' leave s or pairs of brai1 ches or pedun cles. Othe r types of varia tions are asyrn metri cal r~g':l~.~i q ifs_,._ _ \Yl:l~.~ll. t_ak~ _p.lace _by. ..- slipprcssion -- (_1i . displ acem ent of one . or the two leave s or one of the branc hes or pedu ncles in a g.i.ven pair, .\:vhich then becom e · altern ate. Significa nt evolu tiona ry trend s in dichasia! synno resarc Espef ctia.. by ex e1nplif1.~d cencc s, ( 1) reduc tion of branc hing and hence of the n u n1 be r of cap it u 1a to a small n urn be r, do vy n to th e singie head; (2) reduc tion of the lengt h of the . intern odes, pedun cles, and pedic eJs leadi ng to the conce ntrati on of a numb er of capit ula into a _ more or less comp act glom erule ; (3) reduc tion of · the numb er of steril e leave s or bract s of the vegetativ e sectio n of the synfl oresc ences (the final step being a nake d steril e part of the axis); ~1nd (4) asy1nrnetrical reduc tion of branc hes, pedu nc{es~ bract s and steril e leave s . .\vith partia l. effec t ·rhis- kind of varia tion is of altern ate branc hing. ._.. p:rese nt in some speci es (e.g. ... E. schu!tzii), but neve r affec ts the whol e svn flores cence .:

Life Form s -'

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The Espe letiin ae ha ve diversifjf:-d ..grn\v th forrn\; __adapt ed. -to--vari o u-;; --extrer ne· en\ ~ronrrh:~ n tal ·con-- dition s of the high tropic al /\ nde:;. 'T he xen)rnor·· phic ch aract eristics of rnost of th~ p<Srarna ph~nt s 1 · t· 1·;~ oener ai J' r e·r our·ti-,:.:.c.t 'tl1L1 h· r \-' t-=~ l: r· ..,.I, ~,..·J . ,. ...- (·: ·· ;b,·~a· l' \ 1al' r -r -, c . '9~J !) i ,.::.i.--~ . ·· 9 I "'·J .,-. • . I',:.: It , Ll ·· r ( rc . . ' n ' J. \#v' Is f.l> ... (.. .) '\ ,_.. . . ( ( "'-' '-" • \._ ·. .. v• .~ !' t). ..,./~);. ,_)·) <.>t i a 1.. r.11. ·~ -:t"~ ::. s t...~ h• 1. , l {71-7 (.i .·· ,,' -· 1;C--. C.. t' . -1':/; • • ' vl •d r e. 1<.J. :~... !-;;Jd·, The Esoe ietiin cH:O. in addit ion to Inanv of tho~e g enera l veEe ._, tativc anato .n1ic featu res, have dev·eJ._ oped a few excep tiona l and distinctive grovv th forn1s (with coord inate d stru ctura l featu res that prote ct the ir biolo gical cycl ~ ·agrrinsr coJd and physi ologi cal dryne ss}. \vhich ha ve . been extra - ordin arily :.;uccessfuL SonH~, such .-as the cauJirosul ate Espe!etia a n d Coespe!etia·., duxni nate exten sjve stress ful param o regio ns. 'This type of adap tation - is except.io.rud anJon g c!icory'ledons .. altho ugh conve re.en t -for n1s· are \veli :kno\vn-·fronJ the high n1oun tains of tropic al East i\.frica with 1964·: benL Dend rosen ecio and Lobe lia (Hed :._. . Mabb erley ., -1973; J-i.edberg and I-ie dhe.rg:, 1979; " see . Chap . 4) and . fro m·· I-favvaii \\'it:h: \Vilkesia. Argy roxip hhun ~ and Duba utia ( l(eck ~ 1936; Carl-.q uist ~- 1974). . · Lateral and terminal synf-lorescences In the -~speletiinac \Ve_ find grQ\vth forrns from ·- ·-· ,.rhe rn·ono chasi aJ-syn ftotcs cence s also vary w'ith.......·-·:. .~.r~-~~ . .tf.~_es....C~Y.Q.Q.d.y p_l~tn ts . \vi.r.h... a.. .m ~~-~:q ... mass iveiy · respe cr to the· exte·nr of branc hing ·arid. heii-ce -·t() _·_. v~oody trunk undiv ided in the~ lo-\ver h~~lf to one the numb er o.f capit ula, \vhich in rare cases are _· fourt h, and profu sely branc hed upv{ard), .reachreduc ed to a single one. The ::Hnou nt of foliag e of : ing 20 n1 high, to small . sess ile roset te plants no more than 20 cn1 bro ~~d. They can be c1assit1ed as the veget ative porti on also varie s und rrtay sho\v follo\vs: 2n evolu tiona ry tende ncy to\va rd reduc tion to a nake d axis (Fig . 11:-2). 1. Trees. \\'ood y plant ·\Vith an erect distin ct In many cases the synft oresc ences are dowe rtrunk branc hed profu sely at a consi derab le ·dising branc~es axilla ry to the medi an leave-s of the roset tes. The centr al or apica l bud ( rneris ten1) of · -tance .froin the groun d (Fig . .11-4 ). a. rvronopodiaJ trees. ·\\. it!; laten:d buds and the roset te js conti nuou sly active for n1any years , axillary dicha sial synftGres.cences (mod ef: ('arthe single. St_en1 grO\Ving vertic ally \Vithout iilterrupti on. 'T'be axill? ry (later al) synft oresc ences are ·. ra.nzboa trujiilen.'i-is). T'he n1ain trunk and produ ced perio dical ly; in the Espe letiin ae they . branc hes exhib it rnono podia L indefinite~ apica l can be of the dicha sial type (Esp eletia~ Carratr~.- - . grow th: t~~- 1~-~Y~.~-~-D_g__Q.f_~I~:~;h~.~. slre:_ .s.pically.. aLt.er.~---·-boa) --( F-ig:-- Ji-..3€)·. or. nTonachasi ·~1l tEspe/etiofis·zJr--;. --. ~-':i1 ate ~--tfie leave s are usual ly large and broad . and green \vith tubul ar she aths . mostly cro\v ded at the Coespeletia).. Term inal svnft oresc ences are produ ce·d direct1v : end of the branc hes (Fi£. 11--+B). by the apica l ~eristem of a stem or branc h; this · :--> - b. Basic ally mono podia } trees, with latera l impl-ies the. end· of the life of the stem or branch ~:·...~. · buds and branc h es and termi nal mono chasi al syn(Fig. 11-3.B ). I_-n the Espeletiinae~ the termi nal , - ~ _··florescences (n1odel: Libar iotha 1nnus n eri~folius). synfl oresc ences are ahvays of the mono chasi al ~·~· ,._ Leaves are spira lly a lterna te, -,~.~-it h tubul ar type, usual ly corym biforr n and a bund antly flori- -·-.· .. sheat hs, the Iamin'as oblon g. coriacequs~ and xer-. .:.. · o-t norph ic; they are us_u ally .clust ered at the t::ild of ... ferou s.. ·-the branc hlets . The n1 a in trunk is rno nopo dial. 1 'he ultim ate tern1i nal inflor escen ce in fact is The n1ain branc hes are spiral ly altern ate by often the capit ulum , · \Vhich in Co1n posita e plays the pseudover~icils are produ ced by t\YO to five role the fto\ver has in other families. \'aria tions in _size and shape of capit ula partia lly r_eficct trend s : branc hes arisin g very close ly to each other ;· these of _e_yolution and often are coord inate d \vith other ... .-.pseud overt icils may appe ar only once or sever al 5--l in Cuatr ecatin1.es over the san1e trunk (fig. ...... featu res. •

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RAD IATIO N OF ESPE LETI INAE

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.. Pseud odich oonii. chard nia Tanza of Tree B, al). (parti ensis bocon var. (J/ius nt!riij nnus othcu Liban of tre7 a in hing Branc A. II-+. . FIG , totnic branc hing, symp odial . C, Tree of Carrarnboa trujillensis. Mono podia ! altern ate branc hing. .

sas, 1979 a); main bran ches · and bran chle ts are also mon opod ia!. In regu larly deve lopi ng trees , the dista l bran ches \Vith tenn inal synf lores cenc es have furth er syn1 podi ai g:ro\vth by \Vay of suba pical axill ary shoo ts. lf.l shor t, the basic cent ral arch itect ure of such trees is mon opod iaL but the

to the -.- m.o nopo dial cent ral bran chin g can vary cons ider ably due to chan ges of envi ronn 1ent al facto rs that affe ct its appa rent arch itect ure. Th·is has been espe ciall y noti ced in popu latio ns of L. occultus. in \vhic h the peri pher al · sym podi al. bran chin g

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, ----___ -.. g.-.... chin 5ran oiall iopo nioi d -uce ~·-rea --_ : r-, o.aia p m sy· . -is o-\vn cr. e -oiTh tureife-c afch .Eir f .e' _p_e.r1'f5h c. Tree s have mon opod ia} trun ks, \Vith ·bran ch(Fig. ll-4 A). . and al, podi sym ~nd · us omo chot dodi pseu ing . a of type l tura struc a ribe desc to g tryin Whe n sial ocha mon and inal term ces scen tore synf with · zed reali be t mus it ies, spec or IS gen-t: ar icul part that the tree s are not alwa ys allow ed to gro\v and -· (Fig . ll-4 C) (mo del: Tanzania chdr doni i; fig. 5-2

in Cua trec asas , 1979 a).- Leav es are spira lly _a lter.deve lop fre~ly in natU re. The re are irreg ular envi ronm enta l facto rs in a part icul ar loca lity, .· . ·nat e, with open sheaths~ and the lami nae are coriace ous and dens ely vest ed .ben eath ;· leav es are such ?S the- lack of nece ssar y shad~ fo..r .·young mor e or less cro\ yded . in term inal clus~e~s. The plan ts, exce ssive vege tatio n and cotnpeti~ion, and mat ure trun k is rrtassi\-'el y wo-ody ~ The arc hi tee- · nt utila tion s, ofte n prod uced by anim als incid en: ture of su_c h tree s agre es \vith the '"Le uwe nber g tally or by thei r eatin g the tend er shoo ts. Exam inmod er' of Halle and Olde maii . (i970). Ta-,nania . atio n of man y trees of Liba notf zatn nus indic ates·· COn side rable Vari ation in the numbe~ of sing le . deve lops an· initia l.. robu st, erect Stem \Vith ·bra nche s and pseu dove rtici llate bran cqes on· the · .· - spar se, . spira lly alte rnat e leav es, f9rm ing a larg e te efini d in: r ~~fte y. inall terrn tte ro~e or· ter clus ... of unt amo The tree. le sing .a in and. ies sam e spec smeri inal 'term the s, year ral seve ably prob , time n ortio prop in tree le . sing ·a in g chin bran al podi syn1 .

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CUA TRE CAS AS

272

A plan t rr1ay cont inuo usly prnduc~~~ -~nnorcstern goes into the repr oduc tive phas e anJ cenc es, altho ugh usually the florescen~e is inter deve lops a larg e robu st synf1orescence \vho se axis is the cont inua tion of the main stem of the tree. ___ rr~i-~tenr: _)-b~r~ ca.fl __b_e, _m~ny c_yct~.s in_ a y.~ar. and_ the_-timing is not yet weJi kno\Vn _ (Stu dies on flo"At - t he ~-·ame""'tirne-, Olle _o_r_ mor-e d()'r'ma-nr axilfary ration cycl es have been carr ieJ out in Venezuela by buds~ loca ted imm edia telv belo w the infio resMonastc_rio [ 1979J; see also i\. P. Srnrth 119751; cenc e. begi n to deve lop into shoo ts. The se shoo ts rv~d that ir\.,_dry perio ds obse been has It 3) hap. (~ of th gro\v he t ir1ue cont that ches bran the n1e beco ·inf1or~scenccs _ arc rare ant!.in rainy peri6 Js they are the tree~ __u~ually fon11i ng a f~d?e dich otoin y. profusely (often continuously) produced:-after frucOcc asio naJi y n1ore than t\vo l'uds n1ay deve lop at J

t.'.t

the top of the n1ain sten1, prod ucin g a pseudop!ciochas iai branchin1r. 2. C'au!irosufas. Fler e \Ve see mult ifoli ate rosimbr icate ette·s \\:ho se leav es have enla rged ._,. shea ths arra nged in n1uitispiral cir~les at the end of a long or sbor t pach ycau l stem_ The tenn inaL -

1.-

tification the int-lorescences die out but usually remain attac hed to the trun k. hang! ng an1ong the rnarcescent leav es ( rr:odel: 1:..-speletin grandijiora l-I.

and B.) (Fig . ll-3 C). Polycarpi..: caul irosu las are

esse ntial ly unbr anch ed (mon ucau lous ). but due to acci dent al injur y of the apical bud, fe\v branches may repla ce the other\yis_c single axis. Rarely... true apic al, vege tativ e bud cont inuo usly prod uces nevv to ec[ resp \N.ith . uced prod s. i . g chiri bran ous tane spon ·are ~_s leav l in1a prox r olde the e \Vhil leav es, has form t ocau tnon its re~ tectu archi g chin bran the g ajnin ren1 ·and : ering \Vith ng, dryi ely essiv succ been !')uperficially plac ed into - the ·~sensu an1o .. lo'' attac hed to the stein . en1hracing it vvith the Corn er mod el grou p of Halle .and OJden1an (197 0:) clos ely imbr icate shea ths. Gen erali ties about cauclass ifica tion. Hovvcvcr, Espe !etia caulirosula ·has liros ula gro'vvth forn1 have been given in '~arlier . main the : ches bran tive oduc repr iated rent diffe ~ 1968 b, 194S> ~ 1934 as~ s _ reca publ icati ons ( Cuat ary, axill the reas whe is lotax phyl l spira has k .trun -ure itect ·arch its of tlnt acco dged abri an and 1976 ) later al inflo resc ence s have dichasial branching vvith and of the diffe rent type s . vvith bibl iogr aphy , \\:·as decu ssate phyllotaxis; · this plac es Espe letia into pres ente d by the auth or at the 197?3 1-\arhus Symanot her arch itect ural type apar t fron1 the <:.~orner posi ~In ( Cuat reca sns" ' 1979 a). --r he an.at oiny of the ---------- el. mod apic al mer istem and its func tion \Veje stud ied in Th-e fo'llow"ing fo ur vari ation s of polv caro ic cau- deta il by W·eber··{I9.56)-~fnd Rc>ck --(r972-, -·T981). Jirosulas can be cons ider ed: The foJio\ving is a brie f revie\v of the vario us are e 'fhes er. prop la irosu caul ic carp Poly (i)_ nae letii Espe ~he in d foun ias irosu caul kind s of caul irosu las 'rlith an elev ated n1onopodial taJl (Fig. 11-3). ..... stem~ typif ied by Esr;cleria. It is repr esen ted by a. Polycar[Jic caulirosula. The usually large - mos t spec ies of Espe letia , C'oespeletia~ and £sperose ttes term inat e- a~ erec t, sin1 pie, strai ght stern co1n pone nt of the ~~caulirosu the is it psis; letio arise s ence resc Inflo . or trun k 10 cn1 to 15 rn hitdl ...... And leta ., form atio ns that don1 inate large . - ean ....... from the axi1s of the rose tte leav es. '"fhe leaf e essiv impr their by d rjze acte char s, area mo para er rath , rigid or ible 11ex us~ ~ceo cori blad es are -pec ulia r physiogn_o my. oblo ng, xeron1o_rphic~ and cove red at leas t on the (ii) f )o(vcarJJic caulirosula .. sessile. 1... hese are unde rsur face by a thick v.;hite or yello\vish induthe caui irosu las \Vith shor t stem s- (not -gro wing n entu m ...rhe Large _shea ths, as \Vide as or \Vider r_ high er than 20 em) , encl osed by their O\vn rose tte than the lami na, :are closely in1bricated and and the corn pact pack of n1arcescent leav es. appr esse d, tigh tly surr ound ing the sten1. The •

··n1arcescent·-Ie-av·e s -·can ---·per sisr-z·--:--Atrhough--fltey-a-o-·r:or -gr6\-i:1IY1ieiglif--tfiey___c~an bly idera cons g asin incre old, ly eme extr me beco the on ring cove y bulk a ing form ly~ indefinite the thick ness of the \voo d cylin der. Due to the stem . The trun.k appe ars to be seve raj times _thick er than the real sten1 and had a colu 1nna r - relat ive shor tiles s of the sten1 and the bulk of the a.s pect . The sten1~ usua lly.\ vith a diam eter of 6-12 - .. rose ttes and the mar cesc ent leaf laye rs, the rose tte seem s to be sess ile. They are · ofte n des(~on1etitnes 15) ~n:1, has a very thick cent ral soft are h \Vhic nt, lesce acau or t u1a: liros acau as ed crib _ _ ular -vasc dy v;oo , rigid , hard a by ded oun pith surr irocaul le sessi the ong 1-\m s. term ate opri appr not _ . bark t tigh a and nder cyli sula s can be .men tion ed Espeletia argentc;a, E.: _ 1n this kind of caul irosu la the apic al rner.i~t~zn bo.,vacensis, Espe letio psis rnui ska, -Espe letio psis rem ai.ns inde finit ely activ e, pion 1oti ng the con~ Cau a). 1979 , asecas uatr C in 5-11 (fig. osa pann -_ _ of \val rene and _ ~tern the of vth gro-\ ard _stan t up\v d calle be cat~th gro\v slo\v of letia Espe ulan liros of way by rsoccu nce esce Flor es. leav _the rose tte ier stag es . earl r in ile sess _· ... of axils the _ fron1 ches bran ce .. -thei scen lore . !ate raJ svnf .,. 'rhe c. rnati rhizo ~ ula_ iros caul ic· carp Poly ) --(iii -, --tte rose .lL\ . tte rose the of es !eav lt adu_ ian rned _the rna y prod uce man y synf lores cenc es at -one tirne , - real stem is subt erra nean and the rose tte sessile .. sub\ ·crti cilla tely, spira lly arra nged like the leav es. - The rhiz ome can be unbr anch ed or sp~1ringiy --old er-;·--- -·h-angin-g -~

\,_..:

RAD IATI ON OF ESP ELE TJJN AE

273 ·

the by ered cov kly thic are es nch bra the ; \vth gro tte. rose a with ing end bra nch ed, eac h · bra nch t, ben cum pro and , aths she or es leav t cen ces mar thae mar ia elet Esp like ts, plan The se are small -_· rest ~.tt~ ~ . l._to mina te.r vy hea : and e arg h.aJ wit h eac ... . . : . .. as, s .ca_ atre (C.u . del) (.mo asii cald . .--an d .Esp elet iops is. red o\ve lyft rich a1, !1in .terf The . und gro th~ on ing 4.) 1979a .. fig. 5-1 that axis k thic a with ct ere ds stan nce esce ftor syn ·stem The . rose tube , ul~z iros caul oic car: (iv) Po!y n., atio tific fruc At . stem the of_ n atio 'inu con~ a is irre or id pso .elli ally usu er, tub ean rran te sub is a the who fc rose tte dies. Th e bus hy_cha ract er is not gula rly cyli ndr ical and thick. The ro s~tte is sesln~rge the . of h eac e hus bec. nt, are app ally usu . -· as ue~ -s ts\plan ll sma ally sil e . The se are ·usu lly usua , und gro the on be to ears app ttes rose ~ved ia elet Esp , del) (mo tr. Cua Esp elet ia batata a in nies colo e larg ; \vay ous gari gre a in d \vde cro n nlt> com The tr. Cua a nan E. ancJ ~ dellii Sch. Bip. -. sup als vidu indi y man by e mad are ion ulat pop ca~ trc Cua ( to bata n /ej6 ji·ai is e typ bio nam e of this mod el The . ttes rose eral sev to one from g tin por 2). 5-1 , 5-2 . figs . 9a sas. 197 atre (Cu ii jahn ezia ilop J?u is form \vth grb this of lircau of type s Thi . b. 1\1onocarpic cau liro sula casas~ 197 9a, fig. 5-5 ). osu la has a veg etat ive stru ctur e sim ilar to the pol yca rpic caulir6su1a typified by the Esp elet ia mod el. The b~ological cycle is diff eren t. Aft er a . . tim e. usually man y yea rs · of_veg etat ive gro\ vth" THE ORIGIN OF THE COMPOSITAE the apic al n1eristem abr upt ly cha nge s from a veg(HE LIA NTH·EAE } etat i ve to a sex ual phase, bringing forth a large, te-rininal syn ftor esce nce , the \:vhole plan t dying and e tim the ut abo tion cula spe h muc is re The c anti a?< hap , rpic oca mon is it on~ afte r fructificati n bee had It . itae pos Com the of in orig of e plac zia lope Rul is el mod The B). ~ A . 3 (Fig . 11the in ed inat orig they t tha med assu ly eral gen figs. 9a, 197 2; fig. 6, 197 ~ asas trec (Cua sii · figu eira Low er Oli goc ene or the Lat e . Cre tace ous 5_,.3 , 5-1 4, p. 402). The · foll owi ng four variants can il foss able tion ues unq iest earl The 7). 197 r" rne (Tu _ . . _ be considered ~__ ~1iocene, the from y onl s date r, eve how , ord rec · are _ se: The . per pro sula liro cau ic (i) lvfo noc arp ·. the i fron fial \v.--t shq data ical .Iog O n_ paly ent rec and the with eing agre , _ ~I __r~_l_l __ ~~-uHI9~-~.l-~-~~---·D1~~!!-9Pqq~ -~uri'ount --of het --ent, s e' r j)' the 'to_ "erie c ' 0 M1' . . dd1e ~~1'i · '{$ · i~gnch b·ra ited Lim ier. earl . ed mod el des crib ease s incr nts ime sed the in len pol itae pos Com , cies spe er Oth t. iden acc by_ y on1 ed pro duc ller , Mu and g, pin Hop d, eraa rll) (Ge ly ical mat dra mar R. ii, ruiz zia lope Rui are , el mod the ides bes 4; 197 , men Ham deivan 0; 197 , ller. .tvtu ; 8. 196 . ides onio palt R. and i, cesc ens , R. lope z-pa laci ly, nate ortu Unf 7}.p. Cha see 7; 197 , essy Stu 9; 197 itec arch ed plac be can Thi s cau liro sula biot ype ion inct dist h muc \v allo not s doe still ogy nol paly .and le Hal of el mod ttum Hol tura lly und er the , eae erac Ast the in _ra gene of ups gro or era gen of . 70) Old ema n (19 for ortn unif er rath g bein y log pho mor len pol the The . atic o1n rhiz , sula liro ·cau rpic oca (ii) J\1on the larg est par t of the fam ily. Act uall y such uniros ette is sess ile; -the stem for the mos t par t is sub of th you the of on cati indi r the ano be · may ity form rs bea t tha tion por al dist terr ane an, exc ept for the the Ast era cea e. the term inal rose tte. It can pro duc e sub axil lary al. et aad mer Ger by . nt eme stat ng owi foll The es nch bra into elop dev may ch whi bas al sho ots , r rlie '"Ea : still d vali red side con is ) 283 p. 8, (196 s case h suc In . ttes rose inal term e larg top ped by s of the basi the on t. tha 5) 195 al., et yl (Ku s view . rpic oca mon also are tte rose its and nch bra eac h I

~erg e-·~s ~-one-e. . . ae race -sfe a~-·A e-coi r r-· gica o -n-oT pafy -· -'fiio tfilsf eTo d mofie1s-·t urea p pui iro tl R.!l!Topeiza-

nts \Vi thin the · ang iospme elo de~ st nge you the of . · -_ . · ·· . . . e. typ is in orig of e plac The _ ed. firm con s thu are ms per The . . rose tube , sula liro cau pic • (iii) lv/ono car e hav l \yel · very may and r, eve how e, cur obs still an, ane terr sub is dex cau the ile; sess ros ette is ti~e of the , belt ical trop the side out ated situ n~ bee --· . . life ive etat veg long a ~ith . se ero tub and k. thic orig in presum~bly bein g mid - Ter tiar y., . At the tim·e of flor esce ·n ce it pro duc es a ·larg e and n disc uss . ~ve Of ine erm det to ible po~s not 'is It · all ts aus exh that _ nce esce sho\vy term inal synflor inal orig the be ht mig e trib ch \Vhi e ·tim this at _ · le \vho tne and , stem us ero rese rve s of the ~ub the of ive ntat rese rep e itiv prim re mo the or up gro sis nen fabo zia lope Rui are es mpl pla nt ~ies. Exa the tedtula pos 5) (195 ist nqu Cro e. cea era Ast . . · · . cosa floc R. and ta~ gari tnar R. (rno del ), the of up gro e itiv prim t mos the be to ae the lian ·He or re 1no and lo\tv , sula (iv) lV/onocarpic cauliro . in, orig of e plac the as l \Vei as ~ ory the s Thi ily. ·fam ed mm ·ste rt' sho are ttes less bra nch ed. The rose st~ lqui (Car s rker \Vo eral sev by d· usse disc n bee has : , rt.Iy sh:o b~ can t plan The and sess-ile or subsessile. ~.nRob ;. 7 197 , ner Tur 7; 197 , essy Stu 6; .197 6=196 . · ; ~-. : r_ alte are · s che bran the· ed; nch mo noc aul - or bra · ·" . ·, . . . . ·. -. ·. .. .. ./ _. _ _ I). 198 son~ . d---ifie ram re mo or ent tqu infr and rt sho n ofte nat e. entl y curr itae pos Cqm ·of ety vari the ing erv Obs · _ . . _ of ner man ate tern or · mic oto dich udo pse a \Vith

274

CUAT RECA SAS

distrib uted throu ghou t trooic al Amer ica revea ls PLACE OF ORIG IN OF THE E8PE LETll N,4.F the amaz ing capab ility of the earlie r Aster aceae -to -d iffere ntiate--into -m.an y h-i-g h categ ory .grou psThe subtr ibe h as abou t 130 speci es in 7 uener a orese rving the funda ment al struc ture and organ distri buted in an area appro xiiria tely 1700 .kzn izati on of the fa1niJy embo died in the capitu lurn. long , from north ern Ecu ado r (Lian gana ti s., TunThey have the flexibility to .adapt to the· n1 ost gurah ua) to th e iittora l moun tain ·c hain . of E·J diverse kinds of habita-ts and the abilit y to trans·Jorn1 their veget ative - struc tures as prote ctive 1~ vii a ( Ve nczuc la). It ·~i s -o hvio us, luoki ng ·(1' th~~ distri butio n (Fig. 11-5)~ that the r11ajor conce ndevi'c es again st stress ful envir onme ntal co ntratio n of taxa occur s in the oro graph ic n1assif of '(Jltions. ·Ever ythin g prove s the genet ic capac ity of the Sierr a Nev,t da de IVIGriua \Vith its vario us thi:: A stera ceae for vi.rtuaJiy unlim ited kinds ·of exten sions north east\v ard and . south \vest\ vard. n1utations that provi de oppo rtuni ties for over-rhis Ver.1ezuelan branc h C?f ~he North ern Ande s is coini ng Inost obsta cles to survival posed by consepar ated by the iV'laracaibo .B asin and the Cucu ta tinuo us ecolo gical ch anges . Depr essio n from the Easte rn Colo mbia n Cordi lSuch a ·stron g biological '' drive " of the Aster alera endin g in the Parar no de -ranla -El Cobr e ceae has produ ced spect acul.a r gro\v th in - a Massif; fro In this point to its extren 1e north., at numb er of taxa and individuals., hence of the Parar nos de Nepe s and de Barb.aco.as .(Tr.uj iHopopu lation s of this famil y, durin g. a rel.atively Barinas)~ this orogr aphic unit . is abou t · 300 krn ~~hort perio d of _geolo gic tin1e-. The tribe I-Ielia ntheae , \vith 265 gene ra and abou t 3000 .species--_. -long ; the rnaxi mum \vidth of the large st central-sectio n at 3000 .m is 50 krn: its. base at 1000 In is accor ding to recen t revision (Rot?inson, 1S)81.).. abou t 77 krn broad . of t 2ixa and life est varie tv the and · \Vith ., __, ,G!:reat . . ln.. this Vene zuela n segm ent . of the Ande s .are. fonns-, are heavi ly conce ntrate d in the Neotr opics found 6 gene ra and ·63·endenTic.species of Espel eand are clear ly Neot ropic al in origin and devel optiinae . All 6 gene ra are \vide spre ad in this . rnent . The tribe excee ds -o ther majo r tribes in the g their maxi m urn havjn an range't 4 of them .Ande ae, . J~.re_a _..( e_~_g:. '· ~-~ry-~_ cjQn~ae-~ _t'\stereae, Eupa torie .. ... .. .ancL ~'lutisiea-e ) . in . ·nu_rnbers ~and . _cU.y~x~.i.ty_~. T.b~ ------·q-~y~.l_.qp_r.n~nt_ i!1 . JP.G ...~J.~fl:~ __T\VQ __gerJera ((arrCff!J-. boa and Coespeletia) tire ende mic~ Ruilo pezia is sub tribe E.spe ietii.n ae origi nated in -this evolu tion_nearl y enden1ic~ for or· its 21 . speci es only . 1 is ary conte xt, \Vith the abilit y to adop t a varie ty of found outsi de the area (alth ough very close to it). morp hological and repro ducti ve strate gies that Of the t\VO other gene ra . . Espe.fetiopsis i$ widely in1pro ved their abilit y to survive and penn itted distri buted in the Colo. mbian _O rienta l c.hain and then1 to play a predo mina nt role amon g the overLibanothcunnus, conc entra ted in the Vene zuelan all veget ation of large areas in the high Ande s. ....

--~

---------------------------E8 · 70 -~--7'(. -----------------------------------------------------7~

FTG.

al center. of the. _. _ 1.1-5 Origin ....... ~

Espele tiinae . 'le-nez uelan Andea n :. branch \Vith 65 specie s and 7 gener a. 4 of them endcn 1ic . .The specie s are . all endem ic.

•

275

RADIATION OF -ESPELETII NAE

colder or drier, and \Vith less co_mpctition . from other groups. ',--,·--\vas -a -. -. . . -anc-estor;- -or--·'' proto ...E spe Ietunae Th ts perennial, evergreen , woody plant occurring in a ._" tropical, -rather mesic, cl i rna te favoring u nin terrupted gro'vvth during a long life span. It was shrubby and branched· \V!_th xeromorp hic foJi~~ge ~ capable of evolving into diversified gro\vth fonn s. These initial shrubs vvere distribute d in an earlv, _, . probably sen1iopcn, rnontane forest alternatin g -i n a_mosaic vvith patches of scrubhy vGgetation and associated with trees of l~odocarpus . vVeintnannia, !vlyrica ~ Lauraccae , 1-/ypericrtrn. etc., known to be inhabitant s of the region since the Late Pliocene (van der Hamrnen , 1973, 1974~ 1979a,b). The area vvas at an altitude similar to that of the present Andean forest1 greatly extended horizontally and therefore easily expos~d to climatic variTHE ANCESTO R OF THE ESPELETIINAE, THE ables caused by the topograph ic limits on the one PROTO-E SPELETII NAE hand and glac-ial fluctuation s on the other. This Andean zone was · ecological ly intern1edi ate bet\veen humid and xeric and between tenlperThe Heliantf?eae., pre~ominant1y opposite leaved, ate-cold and cold habitats, conditions nTodified by probably \vere already establishe d in northern intermitte nt local changes of climate reflected in South Atnerica in Miocene time (e.g., Turner.. glaciation s. Stebbins (1977) \vrote that "'~volu1977). During the Pliocene, acomn1on ancestor of tiDn in the Composit ae . . . has bet?n a s·uccession -th~ P-f~~ent ~~p~_r.jg_~_ );~p~l~_t~_in?_~ prqp~f?ly devel. .. op.e.d along the.Venez ueJan An_d es during the last _____ _ o_f__<:)d_aptiY.~ ... rf!_d.i~tjQ_n_s__.t!J.th~r.. . ~h~q __ '1 --~-t~J~~s _.Qf trends in any paricular directio-n~ . and also that phases of their uplift. This ancestor had already the adaptive radiation ·goes .fr9m seinixeric habi-_ attained a stable· type of floral structure and capitutats in one direction to specialize d mesic or even _Iurn organizati on. It \vas an heterogarn ous capitulum with ray female flowers and functional ly mitle · hydric conditions . Radiation and evolution of the Espeletiin ae have certainly foHe~~ed such a specdisk flowers., the primitive pappose achene \VOuld . · trun1. evolve into an epappose ·one in response to the The hypothetic al pre-Espel etiinae shrub would inefficienc y of a pappus in the non windy environdiversify in different ways and adapt to _mesic or ment of the tropical mountain forests. The achene humid-ha bitats on the one hand, or to xeric and \vas prismatic or pyratn.idal 3-4-angul ar \Vith a cold or very cold (paramos or superpara mos) on smooth surface. The involucra! phyllaries would the other hand. In this process.. shrubs would keep the fruits confined in the heads for son1e time, evolve into mesophyt ic or - ~erophytic trees~ and the receptacle \Vas _k ept chaffy, the.paleas pro~ 'unbranch ed caulirosu1as, or sessile caulirosul as; viding protection for the t1owers against de.ssicato polycarpic from diversify also \:vould they JioQ_apq__~gJ9.~ . ·-· -- . .. -···- ·-··----· ·····--· ·- ________.. _ -· - -- ..-----·-· . The basic chromoso me ·number vvas already 19, .- monocarp ic gro\vth forms. \vhich indicates a primitive constituen cy for the sub tribe. The \voody basic habit of the Espeletiinae coinci~es with the general acceptanc~ o.f t~e LIVING TAXA OF THE ESPELETIINAE idea that \Voodiness is· an' ancestral feature ·of the Asteracea e , especially of the Helianthe ae (CarlThe folloVv'ing section .describes the possiblequist, 1976; Cronquist , 1977; Robinson , 1981). I consider the \Voody structure · to be· a common · -_·-·course of adaptive differentia tion and evolution that had given \Vay to the pr~sent living taxa of _feature of the dicots growing in humid or p1esic · · · · the Espeletii~ae (Fig. · 1 f-6). . troptcs. Derived from some older ancestor-\vith decussate leaves . the immediate ancestor of the EspeTamania letinae had evolved into a form with alternate spiral phyllotaxy . This .leaf disposition probably Tamanias are trees of medium size \Vith flexible arose in response to strong local climatic changes coriaceou s leaves and ter1ninal infloresce nces; the and diverse conditions encounter ed in its. emiflo\vers have a caducous pappus. The gro\vth grations- into more _open areas, usually ._n1uch Andes. radiat~s- to neighborin g mountains ofCoiombia. Only one genus! E·speletia. is spread . th-rouQ:hou-t Co-lombia -alona-i-ts t\VO· princi-pal Ger-dilleras southward to Ecuador .. to stop at lcs·. If \V e include the Venezuela n border of the Colombia n Eastern Corllillera (Paramo de Tanui and Sierra de Perij a). a total of 71 .species and all 7 gene ra of the- Espcl c tiinae are found in · this area. 'The 1nonotypi c genus Tanzania is restricted to the mountains around the Paramo de Tan1a-El c:obre massif. This unbalance d distributio n of taxa \Vith r:najor concentra tion in the Venezuela n Andes suggests this region as the original place of fornlation and Jiversifica tion of the Espeletiin ae. ~

---·

. .M

. .

•

...

Venezuela Colotnb!a . Merida ,Aricagua

Granates

Jabon-Cende

Napes

Coastal Range

Sto. Domingo S. Nev~da San Jose Quirora . .

Bocono TuYiame Guirigay

CristaHna

Negra

•

Bata.lldn

Sta. Mana

Zumbador Tama

Perija S. Nevada

Ocaiia

------------------·---------------~----~----------------------------------------------------------------~----~--------------~----~-·.

~ f 15 )

(F 1l""'J

l ,...

( 13

...1

r--..,

.,.'

r....,

I 6 J

( 18

\..--r-./ .

_../

(s) '

r..... ( 19 ) '-...../

3600

2900 m

••

;

L ~-- eft

. \

occultus

'-- ·---~.~---

., I 1 J /'

t 4 )

2800 m

'-.,./

2ooo m .,. ·. :

. -·-------~~------~--~----_.~·--~~--~----------~._-----------------------.------~ '

...

L. neriifolius

. -

Libanotharnnus ~-----------------~

P roto-Esp e letiinae '

FIG.

•

~---------------------1 .

11-7. Phylogenetic tree of LibtlllOIIJan?'tus Ernst. •

RAD IAT ION· OF ESP ELE TIIN AE . · 279 ·

Ma rta . Lik ew ise , it cou ld - hav e crp sse d the Ch am a Ri ver to the nor the rn end of the Me rid a Cor di1 lera and to the cer ros of Tru jiii o Sta te; etc. .Successive gla cia l rec ess ion , foll ow ed by asc ensio.ri of tiii.s Llbcin othcimr£us· -hig her··ph its of the · mo unt ain s and val ley s, led to ne\ v pop ula tion s iso late d fro m eac h o the r by the rou gh rno unt ain -

nor the rn end of the Ve nez uel an Co rdi ller a dur ing a col d Ple isto cen e per iod \vh ich fac ilit ate d mig rati on acr oss som e dep res sio ns in the mo un·tain Tan ges .. Th is sp,eci.es. also. pas.s~.d acro~~._.th~ ~ T ach ira De pre ssi on to the .C olo mb ian Co rdi ller a Ori en tal and fro m the re to the end of the Sie rra d e Va lled upa r and Sie rra Ne vad a de San ta

70 ~

I I

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-68

66 l

--·~ - .~

··--

.... -.- .

-- .l.' j

8 ·

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iI I

I :.

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11-8. Pro bable dist ribu tion of Lib ano tha mnu s in the Lat e Pleistocen e ~ bef ore the Las t Gla cial . FI G .

Prob able distr ibuti on of Liba noth amn us duri ng the late Pleis toce ne, befo re the Gre ar Last G,lacial .

•

r • i

i _ __ ro r~·.~ I

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,-------~--~~~~~~---+~

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. . . ... ·-·· -·

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-.·.-·~··....:, ... '

.-.

11-9 . Pro bab le dist ribu tion of Libano thcunn us in the late r par t of th~ ·Las t Glacial, + 14.000-1 3.00 0 :£?.P. (Su saca Inte rsta dial ). FIG.

. ..

•

! '

•

. .. . '

. I

FIG.

11-10. Pre sen t dist ribu tion of

Liba no(h arnJ zus .

!•

280

CUA TRE CAS AS

orep.:the and e tativ vege the both of on trati This on. regi the of c risti acte char hy grap ous topo by ishcd n1pl acco is d tren This s. part ive duct ing lead n iatio rent diffe ed allow , time in t isolation' fev~' . · a to g chin bran e tativ vege the of n ctio redu ltus occu L. , way this In ies. spec mic ende y tnan to . lii.i() es. ieav theo(· n ratio ccn.t co"n ··and s nth~ brci ·· . · ·· L: it~· bert hunl -~· L · ·to t~y direc less or· more gave r-ise larg e term inal rose ttes; in addi tion , the · ~ores banksiaefolius, L. granatesianus, L. lucid us in the cenc es, ah.vays term inaL are redu ced in nu.m ber main 1'11erida n1assif, L. parv ulus and L. cristabut incr ease d in size . The most prim it.ivG, form in nzontis in the Cen de-B arba coas rang e. L. arborthis serie s is the branch~d caul iros ula usu~~ Iy w·ith eus and /.J. griff inus in the Guir igay -Boc on6 a by cJ inate tcrn1 each . ches bran trate pros fl!\V et.lu Vall anfa Scrr · the in nsis orie divis regi on . L. nebrol /?. ii. jahn R. .. e.g , tte rose pic ocar 1non n uela l!nez an-Y n1bi Colo the at anus tan_z L. par, lioides (Cua trcc asus , 1~7~Ja :- Figs. 5~-3, 5-4) . ·rhc bord er of Tam<L and L. glos soph yllus in the tren d tovv<.lrd simpJification .in this line of evol uSier ra Nev ada de Sant a 1\ltarta. In shor t, 10 or 12 tion is towa rd the sing le mon ocar pic cau.Jirosi1Ia spec ies \vere diffe rent iated along seve ral sect ions with a single cent ra] inflo resc ence ; iris achi eved of the cord iller a (see Fig. 11-1 0). Besi des, fe\v by the mon ocau l sess ile cau1 irosu la (\vhe ther rhivari eties or subs peci es have been dete cted . tuberose~ e.g ... or . urea purp atro R. e.g." atic, zom iorig ably prob us amn noth Liba , zing tnari Sun1 nate d in. the main · Ven ezue lan And es, prob ably . R. jabo nens is) and by the tall n1onocarpic cauJ irosula (e.g ., R. palt onio ides . R. ruizii, R. figu eiranot far sout h of the pres ent rvierida loca tion , dursii) (Cua trec asas , .1976, ·fig. 2; 1979a . figs. 5--3 . ing Late Plio cene or Earl y Pleisto~ene time ~ \vhen the Cord iller a had not yet reac hed its pres-· 5-4, 5-5) (Fig . ll-4 /\~ 8.). ia, opez Ruil in la irosu caul pic ocar mon The . , us ifoli neri L. ies, spec l initia The . ation ent elev d broa a \vith d bine com tall~ or ile sess ther \Vhe kly, quic very ad spre it, to e clos very her anot or rang e of vari atio ns in the xero mor phic stru ctur e mos tly ho~izontally., alon g the whole Cord iller a at of the leav es, was an adeq uate gro\ vth forn1 to a vari able altit ude in a prim itive mesic, rnon tane adap t tQ difficult cond ition s such as long peri ods fore st; its abili ty to thriv e unde r relat ively broa d of seas onal dryn ess and lo\v tern.pera ture . The extr eme s of tem pera ture and dryn ess allo\ ved the _speci es to cove r a great. area . From L. neriifolius . hars hnes s · of the c1irnate, and in addition the li-·· ···· stress ·intlicted on the plan ts by con tin Cfotis tnig:rc.t".: . .s.toc k, e.volving__into .a t.yp.e.-fit.ted .for-.cold er-·c • tory osci llati ons caus.ed by alter natin g glaciations, mate : eme rged a seco nd species, like L. occult us~ gene rate d serio us difficulties for the surv ival of mor e marked~y psyc hrom orph ic. This ne\v spethe species duri ng the Plei stoc ene, ~specially in cies esta blish ed itsel f para llel to the first.. form ing the nort heas tern part of the '/ene zuel an And es. · a ne\v popu latio n belt of L.ibanothamnus at an Ruil opez ia evol ved a serie s of strat egie s to over irreg ular cold er clirnati~ zone a~ a high er altit ude ornon a of tion adop The es. culti diffi e thes e corr1 Both a. iller c·ord the of n atio elev the by caus ed the to n tatio adap an \Vas e cycl l gica biolo ic carp both and ible poss as far as nded exte ies spec extr eme seas onal mac rocl irna te ofte n prev ailin g reac hed the east ern front of the Colo mbi an Corin the area . Her e som e of the sessile caul irosu las dille ra Orie ntal , \vhe re they pers isted . The grow in sem iope n loca tion s in the upp er mon tane seco nd spec ies, or L. occultus, split into loca l ; zone o aram subp y bush open the in and st fore · by drate sepa r~, ille Cord the g alon ns popu latio of that to lar simi e ctur stru leaf lly 'vvoo \Vith rs othe lt resu the es; ridg y rock p stee and ys valle deep open to ted adap ectly perf me beca letia Espe e som ies spec ne\V of n iatio rent diffe and tion isola 'vvas ..~!1~--~-~~~R~<;:~.~§.~ J~-"Y~~- pr.Q_R.~.l?ly_9uring_.th.e__noJ:th=-·---···gra ss ..pa±-amGs-{€ .g~-,-·· ·R-=--flecees a,·· ·R :·-jab-o-ttensis)':··· The deri vativ es with tall caul irosu las \vere perern And ean Last Glac ial that lowe ring of vegefectly fit to grow amo ng the .tree s in the And ean tatio n belts facil itate d the emigrat~on of L.·· fore st, espe ciall y at its upp er level. Som e of the· occultus north'vvard to the Si~rra de Perija~ and take to bility flexi the had ies spec ed inat orig ly nc:w ra Sier the ·to ion ress Dep r Cesa deep the ss acro adva ntag e of the new open area s, \Vhether at higp Nev ada de Sant·a ~'l?rta, \Vhere the ne\v_popu laelev atio ns (e.g ., R. jabonensis) or lo\ver, \Vhere tion , in isolation~ evol ved into the poly mor phou s the tree s for sorn e reas on rece ded or \Ve.re .cesand mor e mar kedl y :xero mop hous spec ies, L. troy ed (e.g ., R. pa/t onio ides , R. figueirasii). glos sopl zyllu s (Figs. 11-7 , 11-8 , 11-9, 11-1 0). Ho\v ever , the oppo site also took plac e; dur_ing the peri ods of recession of the glaci_ers, \vith elevatio n of the thern 1al belts . the mon tane or Ruilopezia And ean fore st spre ad up,v ard. forcing the subpara n1o· and para mo spec ies up to the . sum mits The genu s Ruil opez ia prob ably orig inate d frorn the hypo thet ical prim itive shru bby ance stor .. \vith subs eque nt redu ctio n of · thei r area. This redu ctio n . so.m etim es (esp ecia lly in the periC?d invo lving an evol utio nary tren d to\vard conc en-

RADIAT ION OF ESPELE TIINAE

281

atropu rpurea /~. hracteosa) or stipitat e. The rnajori ty of the living species of the genus are tall . cauli.r.osulas .growin g in .the upper :?;9.11.~ p( ~p~ _ for.~ . . est at tirnber line or in the subpar amo zone. Altitudinal lines for the total geogra phic area are difficul t to trace beca~ se of the irregul arity o f the region, and becaus e the zonatio n lines varv from one rn o u n t a i n to the o t h·e r. Three of the eight tall cauliro s ulan Ruilop ezia 1

FIG.

11-11. Distribu tion of Ruilopezia and of Tcu_nania. ·'

consec utive·: to the Last Glacia l) \vas conside rable, fragme nting the continu ity of popula tions, creatin g biologi cal islands, some of thern very - Smal1_-,; -W..h.ic.h __in__.fact Were refuges in \Vhich ne\V . trends . . Gf differe ntiatio n ...and.___s.peciatio.n ....might. have taken place. The extrern e reducti on of the ecolog ical areas (island s) also caused extinct ion. of spe~ies popula tions and even of whole species . In a_ny case, the evoluti onary line of Ruifop ezia, .\vith its plastic ity to evolve a broad range of

morph ologica l variati ons,. enable d the group to prevail in a variety of habitat s . along the whole · Venez uelan Cordil lera .in spite of the rough conditions impose d on it during the Pleisto cene (Fig. 11-11). Specia tion in Ruilop ezia, as it is seen today, took place in a relativ ely narro\v alt-itudinal zone, the ecoton e betwee n montan e forest and paramo . --Ihis ...zruie~.p.rob.a.b.ly___was...the._Jno.st_.affe.~ted QY__J_h~... glacial _fluctuations becaus e of its broken physio graphy with many partiall y discon nected hills, m_o untain peaks, and ridges of median altitud e, \vhich have alterna ted ,Petween period s of bioloand,. . commu nicatio n. These gical isolatio . n

grow in the hills of the northe astern part of the Cordil lera; the other five gro\v along the south\Vestern half. The t\VO or three sessile species mentio ned earlier belong also to the northe astern \vherea s the t\vo section (Trujil Jo-Lar a), branch ed ones gro\v in the south\v estern ranges. Anoth e r group of species has special ized by evolvin g ._ a sessile, more or less tubero us, cormus , a dense mass of linear~ \\'hite, \VOolly leaves . and ·a central \vhite inflore scence with bright or pale yellow long rays. This group has evolve d at the upper zone of high param o and at ·presen t it is

represe nted by four known species found on -o pen summi ts (R. jabonensis~ R. margar ita [northe ast], R. leucactina (southw est]. and R. floccosa [central area, Santo Domi~go ]) (Fig. 11-12). Summa rizing, Rt_Lilopezia _probab ly arose durVenezu elan ·Corcene in. the ing . _.......,.... the Earlv Pleisto ... dillera~ an initiar form spread ing fro.m ·Tach ira-to .. Trujill o-Lara , and then differe ntiatin g regiona lly -- into the subpar amo zone. This zone produc ed 17 species , 5 in the northe astern subcen ter of speciation and 12 in the southw estern subcen ter. of speciation . Also in the param o·· zone . four additio nal species \vere produc ed, three of them in the northe astern section (Lara-T rujillo- Merida ), the other in. the south\vestern sectio·n (Merid aTachir a) .· Probab ly severa l more species existed in the past and n1ore might be discov ered in the future. Some of the species live today in very ·restric ted areas of the broken mount ains of the Sierra · Nevad a and its southe rn slopes and · -'·

b-ranches-te.g~-~-R~-}os-eph-e!Tszs ,-R~·-rliizil,R.--oro:. nzelioides, R. cuatrecasasii. R. coloradarum , R.

hanbur iana) , an indicat ion that other species may be found in some of the many quebra das or hills still _unexpl ored. Some of the local endem ics are relicts from larger popula tions occupy ing more repeat ed enviro nmenta l change s probab ly have extens ive param o-subp aramo surface s · that __ beeri r~sponsible for a great amoun t of selectio n in lo\ver glacial and dry pe riods. Anoth er existed · eRuilop in n ntiatio differe ful success by ed follo\v . . exampl~ of a relict species is R. lopez-palacii, . z.ia. From the shrubb y ancesto r of the Espele tii\Vhich is found only at the Param o del Guaram a·nae there develo ped rather lo\v plants \Vith fe\v cal. This "'param o ,, is in fact a subpar amo formabranch es, usually prostra te \Vith large and dense tion of limited size , at the to p of Guaram acal hill ~ rosette s adapte d to gro\v in open paramo with at 3100 m or 3200 m. In this localitv it is associrat_h er rocky ground . Specia lized end produ~ts of ated with · ·R . viridis. and in .a more .open spot this· line of evolut ion are, e.g., R. jahnii a_n d R. there it is.ftank ed by a format ion of R. jabonensis. bro1ne!ioides. J\t'Iost of the species develo ped \Vith But these paramo -subpa ran1o species are found basical ly a single rosette , either sessile (e.g., R. .

Venezuela Colombia ·.

Cristalina Bocono Gnmates Sto. Domingo !Merida Ari-.::agtm San Jo~e Coloradas Balallon Zurnbador Tama -----------------~--------------~-----------------------+------------------------------------------------~----------High altitude (Cacuminale) 3000-380 0 m · Paramo p::oper

Subparamo .silv.a 3500-280 0m . . '

Subparamo 2800-260 0 m ·

•

. '

. :

A. .Sessile, (with basal bud$), corlaceou' s, subobovat e, . gr~enlsh leaves B. Sessile or tall caulirosula , oblong and linear leaves E. Long stem I

F. Sessile . .•

C. S.e&sile. caulirosut a,

•

!. tuberous;

white, lanate leaves

•,

D. Usuaily branched, rosette pl'ostrate shrubs

Ruilopezia; Proto- Esp e letiiha e '·' ' '

FKi.

,1·t-l2. Phyloge:nel.ic tree ·o(f<.ullop ezia Cuatr. •

•

· 283

RADIAT ION OF ESPELE TIINAE

only at the top of the mounta in. Heavy n1ontane (Ande an) forest surrou nds the. ',;paramo~" and covers all the slopes of the mounta in except for __s_<?~~ . secti<?.~~- . !~at hav~-. . ~.e~~ ~P~f?~d ~~~eD~)y a nd in \Vhich the forests are being progressively destroy ed. This ~ ~paramo', is ~ no doubt, a re lict

a nd this summi t· of the Guaram acal hill a refuge .... for the species that in former tirnes occupi ed a larger part of the mou1.1tain.~ \vhich later \Vas invade d by the forest after the recession of the glacier s. . The extrem e diversi fication represe nted by the more than 20 species of Ruifop ezia reflects its

.......

relativ ely long evolut ionary history.

Espefe tiqpsis

-rhe prototy pe of Espelet iopsis seems to have been a sparsel y branch ed shrub or tree \vith axill-

ary inflore scence s, growing in a mesoph ytic·. Andea n forest jn the Venezu elan Andes and Col-:onlbian Easter n Cordill era. The clin1ate probab ly ~?~ _like_ !he..J~~~.s~D.t o.n~'- .rath.~r. .hl:lmi9 w~ll~t P.. short season al subdry period1 favorin g continu·ing growth and po1ycarpism. As in other groups , Espele tiopsis shows fl stron_g tenden cy toward siznplification . adoptin g _• the cauliro sula gro~vth form. E. purpur ascens , gro\ving.in the. Paran1o ue Tam a area of Colom bia. and Venezu ela. and £. insignis from Norte de Santan der are good examp les of tall caulirosulas with naked stems up to 12 m long , thriving in the Andea n n1ontane forest and raising their large sclerop hyJlous rosette to the top of the canopy . In undistu rbed climax forests of the high Chitag a Valley . in 1940, these large palmlik e rosulas of E. insignis could · be seen from a distanc e and from above, scattered in the forest (Figs. 11-13, ll-.l4). Correl ated vvith their ascent to the subp<.1ramo

78 (.

12°t ----- +--- ----- 1000 m 3000 m <(:,! ~,co

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l@i Pciramos 11-13. Present distribu tion of Espeleriopsis. Indicati on of ~ubareas and directio ns of radiatio n. Subarea s or paramo blocks: Aa, Trujillo ; Ab, tv!erida: B. Santand er; C, Boyaca; D. FIG.

Cundinarnarca.

ESPELETIOPSIS 1

o so

100 150 km

~ Subandean and ~ Andean forest

Distribu tion and migrati on directio ns

235

CUATRECASAS

Colon 1bia, \vith the subce nter isl_a nds prod.uc.ing and the p.aramo zones , the cau~irosulas evolv ed, four speci es in Norte de Santa nder- Sahta nder keepi ng their marc escen t leave s, a condi tion found in most of the species. A fit:1al gro~~t.h form . __. (B.o.r:ner~!t-A.hno.r??de r.o.), t~v_o in Boyuc~i., a.n.d on.e .. .in the ·-se.ries . of E:~·{;eierioj; .~: is s~p·e·c~e·s at high alti- in Cund inam arca (Fig. I J--1 4: Table 11-I) . It is in1po rtant to note that E,speleriopsi.Y does tude is the subse ssile q_nd sessil e rhizon1atous caunot occur south of Bogo t a and does not enter the lirosu las. Undo ubted ly, son1e branc hed speci es p{tramo zone of the large . high P c:in.\n1~, de Sunladid sprea d initially throu ghou t the total area, and paz. In fact. th e.'nerrio ra1 speci~.:~s ..can live well in subst! lJUently diversif1ecL but at prese nt there are . atrno spher ically hurni d areas like th e rncso phytic fe\v hranc hed speci es. ·r\vo are kno\vn in Vene ;.\nde an fores t and are found fron1 the fQrcsted zuela , £. crista linens is and £. rneride:nsis, which slope s -of Param o de Tama south \vest\ vard to the are subpa ran1o cauli ros ulas vvith usu ally fe·w hills east of Bogo ta. But the parat no or s.ubp<1rbranc hes and thickly whiti sh lanat e leave s. The arno spec.ies of Espeletiopsis are n1issing in the first speci es exten ds from Truji llo to the midd le true parar no zone of Paran1o de T atna becau se Sierr a N€va da de Merida (sout h). 'vvhereas E"'. they .n1ay need some pecul iar environ1nental con~ tnerid ensis is found in the extre me south'vvest, in dition s proba bly lacki ng in that area, e.g.~ highe r Param os La Negr~~ and Bata116n. T\vo other often ten1p eratu re, diffe rent soil.. rocky groun d, or branc hed speci es are found in Colotnbia~ \Vhich belon g to anoth er group 1 notab le for their rathe r . bette r soil drain age. The absen ce of some of the neces sary cond itions also proba bly preve nts these ·hard sclero phyll ous leaves. Thes e t\vo speci es(£. speci es fro,m migra ting farth er south of Cun.dinagarci ae and £. pleio chasia) belon g to the forest rnarca. The lovvering of the clitna tic zones durin g . zone and occas ional ly to the subpa rarno ; they are glaci ation broug ht the subp{trarno belt to a 1000only spa.rs e!y branc hed or rnono caul. This 1200-m lo""·er 1eve·l and allo\ved the Espe1 etiae,. branching.~ being a primitive- chara cter, contr asts but not Espeletiopsis , to n1igrate southvvard along ;..vith the adva nced natur e of the vegetative p ~trt of the E-~lstern Cord illera . Neve rthele ss, the broad the inflor escen ces of these t\vo speci es: 'vvhich range · o f vari,ition a·nd the high deg-ree specialilack bract s. This 0-bsence of bract s found only in a fevv otp e r ·s peci~s that have cvol ved·~ir1d'epe'i1- -~- .. zati.o n in Espeletiopsis indic ate its consi derab le d-i-e--.. "dc.i'1 t1y (e·.g·. ~ ·E: ·cor.Viiz bb.~ci a{td ·£~· g!ci'i-idulosci) .·· . -· .<:in fiq ui ty . . lts- ·a-r'i"giii.. pro 6-~tb'iy T\vo n1ajo r group s of speci es can be recog nized · Lo\ve r Pleis tocen e. and its developn1ent in Colotnbi a fron1 the !vliddl~ Pleis tocen e . long befor e in Espe letiopsis: the first can be called sclero phylthe Last Glaci al. Ious becau se their leave s are mostlv hard coria ceo us c~nd usu.ally gr~en or g~eenish. '"fhis group. is cente red in Colornbi~, havin g at prese nt abou t 10 Carra mboa speci es distri buted regio nally throu ghou t the Easte rn Cord illera . Oniv., one reach es Vene zuela Carran_zboa has abou t seven arbor eal species (E. purp uiasc ens) <?11 the Paran 1o de Tam a, ~1t the found only in the .iv1erida Andes~ from the north easte rn end of the Color nbian Cord illera Orien ern limits of the state s of Truji llo and Lara to the taL Subc enter s of speci ation are .the param q south ern end · at Param o del Zumb ador in island s of Norte de:Santa nder (4 spp.) , Boya ca (7 Tach ira. The area is StJlit into two sectio ns. The spp.) , and Cund inam arca (2 spp.) . One speci es ( £. gland ulosa ) depar ts consi derab ly from the_ north ern sectio n, in Truji llo State and neigh bor'ing -La·ra~ ·~nas t\vo s·pe-tl es ·grb\v'ir1g mthe. mo·nf cfrie- .. ...<?.the-rs~.. -i-n -the ·· :g-r oup··-by--having ... lin-ear--revot u r prote cted and humi d parts . One rathe in t fores herba and v, ulosit gland of e deg:re high a s, leave _, . of the speci es (C. rruji!lensis) ~ which has been ceous outer invol ucra! bract s. found on both sides of the Serra nia del Jab6n · The secon d group consists of subpa ramo and Cend e, the divid e bet\v een Truji llo and Lara parar no species~ all caulir osula s \Vith \vhite lanat e (Hum o.caro Bajo -Agu a de Obisp o), also gro\vs or seric eous leaves~ and a short or up to 1 m high on the Param o de La Crista lina (Truj illotrunk . In this group , one cente r of diver sity is in Boco no) and on the slope s of Para1no del (}uar athe \/ene zuelan i\nde s, \Vhere there are four sub·rn·acal ( elose to .B ocon 6), bet\v een· 2400 i'H·1.d· 2800. paran 1o speci es vvith yello\v rays (£. jajoensis, E.. D1. 'The other species~ C. H'l.trd ackii. gro\VS in crisrafinensis, E. p_ozoensis, and £. 1neridensis) _lirnited locali ty of Queb rada del I'urm a1, 2400 m and t\VO high param o speci es \Vith \vhite rays (E. altitu de' in the \Vest ern drain age of Param o del pann osa and E·. angusttfolia). Such white -raye d Jab6 n. speci es are exceo tiona l in the genus , the first from The secon d sectio n of the area inclu des both the \lene zuela n centr al massif and North Cord illowe r slope s of the Cham a Valley (ivlerida) and lera, and £~ angustf:folia being mostl y from the the moun tain range s bet\\' een Sierr a Neva da de south ern hills. The secon d cente r is locat ed in

·dates.. . from

te ·

'-""

'-"'

RAD IATIO N OF ESPE LETI INAE

287 -

. leys nort h and sout hwa rd at sorn e\vh at ltnver c·lt~ .. vatio ns. The isoth ermi c and perm anen tly hur-nid gro\ vs alon g fore sted bank s of cree ks of EI Mor ro and ~r-~ca_gu~_ as .~ell .as_i-n .the r~gion of the. Par.-:_. . .. · .con d.itio,n. of- the clou d fo-rest, -whe re · the genu~ : . amo de San Jose , at 2400 -300 0 m altit ude. C. pit- · orig inate d- and deve lope d, prob ably favored the h \vhiL· s. ence resc inflo asial dich the of n ntio rete in d foun is ~ . ies ·spec ad spre wide t tieri, the mos vegnate alter lly spira ked mar the .with rasts cont of es slop the of es ravin deep in y forests~ espe ciall iv1erida and Para mo del Zum bado r. C. badilloi-

of til·c · . c. risti acte char ches bran ·and es leav ve etati de mos p,}ra s. rada Colo Las ' de mo Para the .. Plcist(·>.ccn~:. the of ods peri id hum the In . ribe subt Pico , ter-o Quin .. de a Nles ora, Quir de and ara IY1ij the genu s prob ably spre ad thro ugho ut the arc;t . Hor ma. Para mo del Bata llon , Para mo del Zum but the verti cal osci llati ons of the veu.etation~. hado r (nea r Quc niqu ea) bet\v een 2500 and 2800 caus ed by the alter nati ng peri ods of glac iatio n cfi-. (som etim es 3000 ) m. Oth er spec ies of limi ted dismin a ted it from large part s of the area . ·rht: curtribu tion are C. rodr igue zii, grow ing in the for· , P"rf t grea in t, esen repr s litie loca n know ly rent rds to\va s rada Colo Las de mo Para of este d area and e Stat illo Truj for y ciall espe tats, habi ge refu of is irens tach C. and m, 0 -280 2600 ~ ino El ivlol pt:rs isls ou anzh Carr But s. area d reste defo r othe o, oner Port achu elo de La Grit a to\va rd Preg . v in deep , 1e·ss acce ssibl e valle ys and alon g sn1,tiL ~g00-2950 m altit ude. C. littleii., mos t clos ely rela ted to C. badilloi, grow s in the fore sted area _ still- fore sted cree kban ks. I · have no doub t that ies, spec ed scrib unde be may and , lities loca e mor at te; Nor del 1era J i· Cord the in bio of San Euse th~ rug·of ion orat expl re futu th i w_ d foun be will m. 0 2400-260 ged Ven ezue lan And es (Fig s. 11-6 , 11-15). The genu s Carr arrb oa seem s to have deve loped dire ctly from the con1n1on Espe letii nae ance stor , in tree form . It retai ns the basic spira lly Esp.eletia alte rnat e phyl 1ota xis \Vith usua lly large or very larg e, gree n, spar sely or mod erate ly pube scen t This ge~us prob ably orig inat ed in an earli er glaleav es. The inflo resc ence s are axill ary, \vith.. the or. aps . pcrh or ne toce _ PI.eis er_ LO\V the of od p~_ri cial inal term e finit ·inde ng havi ches bran and main axis the. Upp_ex ..P.Jioqe_n e..:whe n cold ,.. \Y.e.t_.paranl<Lco.u- . gro\ vth. The ...bran chin g .·o f the_...irifloresc.e nces . ·is ditions prev ailed in a zone bet\v een the Wes tern . prhil'ifriry dTchasi~il, · iiavfiig.· i\t ....least.... the. --~~v~,.~r· as Ven ezue lan and East e-rn Colo mbi an Cord iller ' bran ches and brac ts opposite~ \Vhe reas the uppe r as first d ergc m" e · la irosu caul tall The· 6). 11-1 . (Fig that s seem It . nate alter or site ones. may be oppo the biot ype able to thriv e in· the wet subp anun o Carrcunbaa was orig inall y adap ted to a hum id besta s serie this of · k stoc first The . zone b scru . . ally zont hori ad spre y inall orig and st mon tane fore of form the in ent ronm envi that in f itsel d 1ishe valinto and ida Mer de ada Nev ra Sier the arou nd poly carp ic caul irosu las \v.ith inde finit e term inal grow th and axill ary inflo resc ence s, well equi pped .to surv ive in the pere nnia lly cold clim ate, huin id atmo sphe re, and wet grou nd. ·The pr!m'!-rY evol utio nary proc ess that caus ed . Espe~etiin_ae to deve lop its the of stor ance the 1000 m 300 0m _ dire ction som cged chan is lotax phyl l spira c basi ~ .....

\Vhat at a certa in p~int__j_~_ j!?_P_hylg_g~ne_ti.~-~-~vct~---·

·6p_m-en_f_ . Tn-- t-he -:~preceqing

.,..

FIG.

·-

11.- 15. Distr ibuti on of Carra1nboa.

gene ra, -· exce pt for . Carr amb o·a, spira l phyl lotax is and alter nate bran chin g \Vere gene ral in both the vege tativ e and Tepr oduc tive part s. Ho\v ever ~ .in one of the resc ence s reta ined th_e · phyl-etic lines the inflo . prim itive dich asial stru ctur e alon g \Vith decu ssate phyl lotax is. Presumably~ relat ively - perm anen t . hum id at1n osph eric cond ition s have fav:o red the pers isten ce of the dicp asial cond ition in the ephe mer al repr oduc tive bran chin g. This phyl etic line prod uced Carrarnboa :jn. the cool -tem pera te nem oral environment~ and led to Espe letia . a stoc k pote ntial ly best fit to com pete in a _cont i··· nual ly cold , wet, open habi tat . . ' . Espe letia 'is fund ame ntal ly · greg ario us and ·specialized for cold ·' hum id ha~itats \vith hum ifer-

CUA TRE CAS AS

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ESPELETIA 1u"'n • tJ

11-16. Espeletia. Probable area occu pied by the genu s in the Pleist ocene ~ befor e the Last Glaci aL FIG.

The se isola ted com1nunities may deve lop som e ous soil, pref erab ly in wet 9r mar shy para mos , and son1 etim es rock y , very hum id places. Its. .deg ree of diffe rent iatio n and in the long run evol ve into new subs peci es or spec ies. In the prorath er heav y seed s with no disp ersal mechanisms ·-·-are·--p--r-of. usel...\r-p·--r<YdUce"tt·seVetai···-ri!ne s-i·rr a-'J1ea r_ ··-, ---··-cess-o f stow ·~- ..long._ ·.::distance-·-mig._rati-on:, -ada ptati onto vari ous degr ees of hum idity , diffe renc es in but the loca l repr oduc tive rate is lo\v. The greg ardrai nage ,. or chen1ic.al and physical soil conious popu latio ns may cove r seve ral squa re kil9rne ters of pata n1o but they are ofte n less · di tion s, appa rent ly sma ll vari ation s in loca l habi exte.nsiv e beca use of topo grap hic or ecological . tats caus ed or faci litat ed evolutionary chan ges· and spe,c iatio n. Ne\V taxa \Vere prod uced and disobstacles to cont inue d propag(ltion . . Espeietia tribu ted acco rdin g to ecological pref eren ces. _species may form dom inan t com mun ities .arou nd . l::speletia is the mos t · repr esen tativ e genu s of the uppe r part or peak s of mou ntai ns and are the- ecologica1 conc ept of a para mo. Mos t of its. usua lly Ii1nited a't thei r base by And ean fore sts. In spec ies deve lop best in the paramo prop er zone . this \vay popu latio ns on ernerging mou ntain s, The y arc foun d also in the subp aram o zone~ hills~ . or peak s .becotr1e isola ted. Oth er pref erre d onal ecot in or ion ibut distr aic· mos in y ciall espe n1s botto The sts. fore by ted limi also are s ha bitat rsupe the in rare er rath is letia Espe . ities mun corn eramod r o flat and . ys valle 'vet . e -larg ll-to sma of para mo zone , appr oach ing only its lq'rver bord er. tely inclined gras slan ds beco me q.uickJy cove red \lery fe\v species, e.g. , E. cleefii and E. lope zii t by Espeletia com mun ities , like islan ds·co n1pl etely alticola~ may thriv e abun dant ly in rock y ·habi tats surr oun ded ·by heav y And ean forest.

Venezuela Colombia Trujillo

Merida

....

Equador

Venezuela Colombia

Cord. Oriental

Oepressaon

· S. Nevodos

Jobon·Cende Guifigoy

Tuiiome Sto. Domingo Merida . Perijo

Sonlander N. de Santander!

Tomei N. de Santander Son lander

B. Espelita •

Proto-Espeletiinoe FIG.

Cord. Central

ll-17 . Phylog~nclic tree of Esp£'1etia Mut is ex II. an<.f

I~.

Cocuy

Vodohondo

Guonliva La Ru~io Tunjo

D. A. Narrow p.h.yflories B. Broad phyllories · C. lndument tanate D. lndument prostrate E. lnf lorescences foliate ; F. Inflorescences ap~yllous

Mocizo de Bogota

Sumopoz

Pmo Popos Cord. Cent•ol Troncol Sur

Cord. Occidental

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Pciramos .Pl Subandeon an~ . L:J Andean forest 00

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. . Probable distribution during 0 50 100 150 the later part of the Last Glacial km :!_ 14,000-13-,000 S.P. L . 7'...,0

·i.r1-t-he--sttpeq:rar-:am=o--sJopes--ef-t-h-e. -Si-8rra---N-ev-a:da~ del CoClfY (Colombia) and, sporadically, £. serniglobulata and . E'. schultzii in the Sierra N eva9a de Jv1e-rJda, defying the characteristic frost and solifluction cotnmon in that periglacial

area (see Chap. _2).

-t-ula-- a-r€-··9:€-riv.g{;i-"'-phylgtic---·tre-nds -{Fig...-ll~2i~). Pseudopetiolate leaves are primitive and evolutionarily the general trend . is toward reduction the. of th~ pseudopetiolar s.e ction, and elimination . . derived form of lamina _being sessile. Lanate .

indumentum is probably primitive, · becoming more or less dense or thick according to environ_mental situations; divergent phyletic Hnes are

i\.lain Trends in Specific Differentiation l . •'

11-18. Probable distribution of Espeletia during the later part of the Last (J,lacial. 14.000-13.000 B.-P. FIG.

•

The most noticeable trends are related to the size. and kind of ·_.inflorescences, the shape of the: leaves, and type of indumenturn. The large, tall . multicapitulate inflorescences are . primitive; r~duction in total length, reduction and suppres-. sion of bracts of the proximal sterile portion~ and re duction in number and increase in size of capi-

sericeo_us indun1entu~~ or by a -straight . patul0us. velutin.o us t}pe. several groups, tliere is a minor .reduction in the .ray · corollas~ ri:p proaching obsolescence. ·The length of the tube of t~e ray corollas initially increases to ~e>me extent; ho\vever, a major trend to\vard reduction of th_e tube is obvious along the migration route of Espeleria from Boyaca to. the chara.~-~ ~ rizeq ~y .Pn?.~tr~te

LETJJ NAE · 291 RADIA.T.ION OF ESPE .

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11-19 . Presen t areas of distrib_uti<?n of Espeletia.; areas and . subare as: A. Cende-J ab6n; B, Sierra s de f\'1 erida-Trujil lo; C, Tama·-.. -santa nde r ancf"Nortc"ae ·sarHander; D, Sierra de Perija -v'alle dupar ; E. FIG.

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pararnos of Boyaca; F~ pa-ramos of Cundi namar ca ; G. main Centra l Cordi llera of Colombia~ H, Macizo Colom biano nucleu s; I. Tronc al Sur of the Centr al Cordi llera; J , Sierra de Los Llanganatis, Ecuad or; K,L~ northe rn parts of \Veste rn and .. centra l Cordi llera Chaqu iro-Fr ontino , paraunos .

extre me south ern limit of the genu s (Figs. 11-17 , 11-18 . 11:-:-19. 11--:20). TJ,VO ·.

ivlain Direc tions of Radiation

COLbr viBIAN CORD ILLER AS

Cordillera: Esp~letia ·first emig rated hori. zonta lly in t:wo oppo site direction's, one along the Ve~ezue]an Ande s; the other ·from the Vene zuelan bord er to the Easte rn Cord illera of Colombia~ · . T~:J.S.·- . latter stock . \vas · e:xtremely successful in es.tab lishin g popu lation in wet places in. the subparam o zone and even tuany ih the uppe r level of to reste d ·e cotoo al zones , migrating from the present .Cobr e-Ta ma n1assif south ward alona0 the Eastern~ Cord illera of ~olombia throu gh a semioEasu~rn

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(subparamo~ · o:r para__m_Q,$-},..___pre.f.er.a..b.ly __.in. .... tbe.~ :-m.o .r~,.- ..q:u.mid__e_a.s.t~f.n_.

pen or open uppe r belt

drain age. Subs equen tly, colon ies sp-read::aggressively across the heigh ts of the Cord illera and trans versa lly to the \Vester n uppe r sl?pe~ : and at the same. time radia ted south\vard-~(~.ig .. lf-2q ). ·The -Colo mbia n Cordille-ra Orien taC the main Ande an range of north ern South ·Ame rica, is extra ordin arily Iarg€ and co:mp licate d,_· having e·x"tensive areas abov e 3000 ·m up· to 4200. m ·altitude, not to ment ion the n1uch highe r altitudes of partic ular hills .or high indiv idual massifs such as the Sierra Neva da del Cocu y (5493 m) and the · Neva do de Suma paz (4250 m). High er massifs and deep valley s are \vell-kno\vn bar:rie~~ to the disse mina tion of param o plant s but. )n addit ion, .

CUATRECASAS

292

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l ESPELETI A 00 0 Population areas. L___.~...;;..o_l_._oo___,lSO Directions o:f migration-~ .._ . . 11111 km Glacial areas. I

· FI.G .

11~20. ·Direct{ons of !nigration

and radiation in Espe_letia.

innumerable. smaller valleys or breaks, peaks and ridges on uplands co!lstitute physi~graphic bar-

_ specie-s--and -rre:rs-'lo. .The__. migratio1f .9f___ p·a:ramo__

cause isolation \Vith consequen t speciation . Espeletia presumab ly n1igrated across and .alo,z1g th.e Eastern. Cordillera, south\var~ iri an ·early or:. 'n1iddle period of the Ple1stocen e. Long before the Last / Glacial its . popul~ti'ons had reached and establishe d then1selves in Cundinamarca and had succeeded.. in. dominatin g the_parti1ern arno l~ndsc.ape ,oJ the ..region ...B.ut at the sou_ bord.e r of the Paramo de Sum~~paz the ad-v ance of Espeletia vvas stopped by a depression of the ·Cordillera· and bv .- the dense Andean forest that covered the steep slopes and ridges of that sector of the range; moreover , there ·,\vere no paramos from about 3. 7° N south to 2° N \\-'here the Eastern and Central Cordillera s join. In the lo\ver and ¥

•

narro\ver northern .extremes of . the .-Cordillera p_ara.mos_o_r north of Oca~.~-L QDLY._~_w.!!lL.~-~<J..tter~o- _ .subpramo s are found in .the Sierra de vralledupa r and Sierr'"a de Perij a, which apparently -escaped early colonizati on by Espeleria (Fig. ·11-19,D) . - The major part of tbe Cordillera Oriental·, from Ocana to. . Paramo de Tama and frdm there to Paramo de Sumapaz, an area roughly 570 kn1 long, supports 36.recogn ized species of Espeletia., besides two more at the northern end of Sierra de Perjja:- .·T·hese num.q ers indicate a considerab le . ; degree of .speciation for the regi~n.. The ~nudy of species distributio n .t eveals the existence of three rna in-areas or centers of speciation ·on the Cordi I-· niaior orolera Oriental that are coinciden t with ~

graphic nuclei .(Fig. 11-19.~ C, E, - F)~ ·'Jhe -first · area - includ.e s the paramos of Santander and Norte de Santander -Tachira; the second are<~,

RAD IATIO N OF ESPE LETI JNAE

para mos of Boy aca; and the third area , para mos of Cun dina mar ca. Of the species growing in each maj or-· area , ..... som e·-· may- occu r mor e · or ···less ·· thro ugho ut the area , but most are rest ricte d to a sma ller ecol ogic al or topo grap hic rang e inside each area . -rhe sam e cent ers of spec iatio n have alre ady been cons ider ed for othe r grou ps · (e.g ., · · · · Espe letio psis) . The first cent er of radi ation is Sant ande r and Nor te de Sant ande r-Ta chir a. This comprises the para mos of Tan1 a and Cobre~ Fon tib6 n-Pa mpion a, Sant urbu n, Rom eral , Berl in, Oca na, and Alm orza dero in a broa d sens e. The se are mor e or less lirnited para mos , which were to vary ing degr ees con nee-ted, at least in several past peri ods of time. The y supp ort a total of eigh t spec ies. The prim itive type of the genu s (with large pa~iculate, tall inflo resc ence s, and pseudopetiolate-, subl anceol ate leaf lami nas) is repr esen ted by E. ste..vermar kii at the Ven ezue lan side of the Cord iller a and £. stand ley ana in the . lo\ve r part of p·a ram o de Santurb~n-Berlin and east ern slopes of P~1r amo del Alm orza dero . Red ucti on of inflorescenc es \:vith supp ress ion of steri le leav es \Vas .exp erie nced by £. bras sico idea in form.e rly luxuriant com mun ities of the Para mo. de Fon tibo n i.P..4mpJ~Q.oa},. ..nO.\-V. . n.ear...e.x_tin.ction~ .Fur ther reduc-: . tion of inflorescences , in total leng th and num ber of capi tula , lead s to ~. dugandii (litn ited to a section of Para mo del Alm orza dero ), E. cana scen s (top~ 4209 m, of Para n1o del Rom eral ), and E. congloJnerata exte ndin g from Alm orza dero to the para mos of San turb an, Veg as, and Rom eral , and to the heig hts of Para mo de Tam a. Extr eme speciali zatio n was reac hed in this area by £. estanislana . an hygr ophi lous species with single larg e head s, foun d in small redu ced area s in Para mos del Alm orza dero and Berl in .. Rela ted to this is E. roberti, an isola ted relic t at the cacu mina l para mo of Juris dicc ione s (Oca iia). A subc ente r of speciation can be cons ider ed the Sier ra de Per!ja. at the .. nort hern end of the Cord iller a with t\VO . rela ted · ·spe cies , E. tilletii and E. perijaensis. The seco nd cent er of radi atio n is. Bov aca. Thr ee subc ente rs ·of spec iatio n . may be con- · ....

side red:

1. Sier ra Nev ada del Coc uy or Guic a.n: In. this area ther e are four spec ies, all tall -caul irosu ·.· . las. t\VO belo ngin g to the primitive mainstre am of the genu s, E. curialensis and E. c!eefii, and the othe r t\vo, E. ann.emariana and £. lopezi~, repr esen ting a para llel line of evol utio n with aphy llou s inflo resc ence s. E. lope zii form s the larg est com mun ities in the _ area with a broa d rang e of altit udes . £~ cur,ialensis and E. anne1nariana gro\ v in med ian

293

and low para mo and subp aram o zone s, spre adin g alon g the east ern fron t of the Cor.. ·dill e-ra; E .. lope zii and £; ·cleefii dom inate · the · upp er part s of the Coc uy _1v1assif ascending well to the rock y supe rpar amo s. £. lope zii, \Vith shor t nake d inflorescences of thre e larg e head s, form s ..exte nsiv e coin mun ities in the· para mo prop er of the \ves tcrn drai nage , in addi tion to its aggressive pere grin ation to the sout h alon g the para mos of the east ern slop es, e.g. , Para mos de Soch a and Pisb a (Fig. 11-20). 2. Para mos of Gua ntiv a-La Rusi a, A,Jto Las Cruc es (bet wee n Dui tam a and Enci no and Bele n-Su sac6 n-G onza ga, with a max imum altit ude of 4200 m), a grea t n1ass-if with exte nsiv e para mos or v~lleys betw een 3000 and 4000 m: In this regi qp a large vari ety of caul irosu letas co~er larg e flat area s or stee p slop es. In thes e land scap es Espeletia- species form mon ospe cific cons ocia tion s. It is the rich est exis ting cent er o.f Espe letia , with 13 livi.ng species (bes ides ,vari eties and subspec;ies). The tnai nstr eam of the genu s is repr e·anclijlora sent ed here .by a y.ariant 9f £. g!_ (ssp. boyqca na) in _...small ref~ges ~ t .... Jh~. . heig hts of....ra.r.am.os ~.de ....La ;Rus.i.a . .and .Las .. Cruc es. Oth er spec ies \vith leafy inflores_c ence s are £. ~zucarina. and £. congestiflora.; this last, with extr etne ly . thick , com pact , curl ed lana te ,. yello\vish indu men tum , and fe\v capi tula com pact ly glom erat e in a com pou nd head , is a very specialized form suite d to less hun1id habi tats of well -dra ined slop es. Six othe r spec ies belo ng to the P.arallel line of evol utio n \vith . aphy llou s inflo resc ence s, \vith decr ease d num bers of capi tula (E. mur -

illoi, E. ne1nekenei ~ E. incana, E. brachyaxiantha, E. - arbelaezi, a1:1d . E. ·disco idea). E. nen1 eken ei and E. murilloi have 57""11(-17) c~pitula on ve~y_ exse rted inft_g r~sc_~~-~-~.~ ;._.th~.

othe r spec ies, livin g at high er altit udes , have 3-5 capi tula , mos tly . .3,- the inflo resc ence

shor ter than or equa ling the leaves. In . addi tion , E. rositae usually has 5 large· capi t11la, but a vari ety has deve lope d with a sing le head . Ano ther ·spe cies · with welldeve lope d inflo resc ence s, . E. boya cens is, repr esen ts a disti nct line of evol utio n charac-

·by pr9s trate seric eous· indu~ent (sec t. Bon plan dia) . in addi tion to very .larg e, long inflorescences \Vith sma ller head s. 3. An irreg ular serie s of para mos at the· east ern · side of the Co_rdille ra sepa rate d from the Gua ntiv a -regi on by the Chic amo cha Vaiiey: This -is the regi on of Para mos de Soch a and Pisb a, de Vad oho ndo in the valley of the Rio teriz~d

294

CUA TRE CAS AS

the prirnitive characteri~tics ·of ps.e udop etio late Cu.s.iiana, ParairLo del 1\.nii'cal ., Paran1o de la lanc eoJa te lea ves and larg e pani cula te intlo resSarn a near Lagu na de T o ta, and hills arou nd getat1ve ·po~r1ions ~ 0 -t:her spe·~9gcnno.~.o ... ln . this regi on .are tb.r.ee .end em.ic . ·-···- cenc es \Vith leafy - ve· nsis dore mira £. are line etic phyl e san1 e th of cies .. oi nzill jara £. , type. itive prirn spec ies of the in i.ti.es rnun con1 in.g forn1 ila roph tapi £. and ties varie and ~ ana aldi os~v /~. and ~ E"'. tunjana n1aSu de o rn Para er upp the on s area icted restr , var. iana 1nar anne . as£ such ies, spec r othe of in 1Ttg gro\v ies spec ant ort iiT,lp an is ei urib E. paz. cenbo.ra £. ii, £. lope E. ~ tion addi In . srris ril]Je th ~ uppe r zo ne of the And~ an · ro·r est ·and in subsis ,. .and E. 1nurilloi are pres ent in abun paru mo bush es . It is foun d in some open area s as danc e. Also pres e nt is E"'. cong estif!ora. E-. a ren1 nant of dam age d fore sts. The spec ies has tunj ana is a ren1 arka ble spt:c ies beca use of its s genu the for n give cs risti acte char itive prin1 the s. ence resc infto ched Jan!., ....... ~., bran .and in addi tion ca.n deve lop a high stei:n up to 1.4 In shor t , 17 spec ies (p·lus a subs peci es from the m tall und er fore st prot ectio n. and a trun k of 16next area ) .a~e knovvn fro,m the Boyac.a cent er. cm diarn e ter \Vith 1O-mn1-thick cork y bark and a The third cent er of radi ation is Cun dina marc a. thick cylin der of wGo d. A11 this indic ates. that this itive prin1 t n1os e th of one t esen repr n1ay ype biot lv1as apaz Sum otaBog the s i: r e-ntc subc maio The sif in a series o.·f para mos in a larg~· e, _b roke n area . type of Espe letia . Ano ther spec ies clos e to it with simi lar m.or phol ogic al ch~ract~.ristics and ecol ogiA seco ndar y · subc ente r is ·the Zipa quir a-Ne usa cal pref eren ces is £. curia lens is of· the ·eas tern subc ordi llera , west of the Saba n a de Bogota. .slop es of the CordiiJer~l in ·Boy acc:i .' 1. The Zipa quir a-Ne Hsa :area .:·. r·Iere thre e en deThis stud y \Vou ld supp ort- the idea that · the mic spec ies are cons ider ed: E. choc onta na genus orig inate d in the · mor e hum id, clo:u dy and E. , cave tana , beJon~in·g to the folia te environn1ent of the upp er '.VOOdy be·lt ·Ofth e east-gro up th~ sam e phyleti~. li~e as£. gran dtf /ora~ and £. barc laya na of the aphy llou s . ern slop es of the C:ord i11e ra. Red uctio n has occurred wi_th the migratio~ to the open .par amo s .. infJo resc ence serie s. £. graiz diflo ra · is al so T!.vo othe r u;np~rtan.t spec ies of the re~1on ~r~ ?f .:P~~~~I?·t .a_~ jt .. i.s _~l.ll. . aro.u.n.d . the Saba n a de f.> ogot.~1 - as _vv.eJl _as ..£ .__ argentea; bHt thes e- t-\V€> - ··· .._t.h~--· ~lpbyl!ous::tn.florescence . - . st~QC-k-:-- ·- ·E~. - ··· kzlltptt ~ ~tbundant t Tl t~e Gua sca- Bog ota a~ea , prefera~ly species have thei r origin in the large B\.)g ota 111 eastern-dr~unage.,. gras sy, wet para mos , and Its l\11 assif at the east ern side of the Saba n a. ~l?se relative,_ E:_surn rnap acis, a s~?wy spec~es 2. Bog ota Massif-Paramo de Sum apaz : This is a vVIth sno\v-vvhite tnfto.resceTices c?nstste~tly \VIth larg e mas sif re achi ng altit udes abov e 3800 m thre e larg e he~ds and an ende mic of htgh a .r eas at man y poin ts and . ~llso \vith man y -flat or ercabr ~· . apaz Surn ~e mo Para tn m) 000 0--4 (~9? ous vari oes, surfa ned incli ely erat mod enszs fro~ the sout. h\ve.ster n slope~ of P.ara mo de degr ees of humidity from deep swam py to Sum ap:az 1s a mult icap t.tul ate spect~s \VJth aphy ldrai ne d area s, alter natin g \Vith rocky spot s, lous 1nfior.e scen ces, prob ably Inde pend ently cres ted rido es and abru pt brok en cliffs \Vith d~rived from a folia te -typ. e .spe.cies i~ ~he sam e gree n gra~y' ar1d bushy area s. Fore sts, Para m? de S~mapaz .. A GISti~ctrve specLes· of t.he espe ciall y on the east- facin g slopes., may Bog 9_ta ~rea In Cundi_nam~rca, E. ar~ente~ (With cove r the terra in up: to a very high altit ude, an erad iate form }, lS Wid espr ead· .In ·para mos 35()( ) Til and mor e whe re it has not been desbel? w 350? .ITl altit uoe.:. -~J- h.a,s. ___fla.ttened ~. sily.er-Y.,troy ed as h'!ppen~ on stee.p_.rQc..k_y_rid.g e.s . senc eous 1ndu men tum and belongs to an Jnde ···pend ent stoc k (sec t. Bonr )lan dia) that prod uced Eigh t spec ies are foun d in the area~ the m·os t the: alre ady men tion ed E. boya cens is in the ~o.minant of ~vlhich, E. grandiflora~ is found . Boy aca regi on. thro ugh out the who le area from the top of som e Cent ral Cord iller a: The Cen tral.· Cord iller a of hili~ and gras slan ds of Para rno de Sun1 apaz at Colo mbi a that depa rts from the so-c alled Tron cal 4000 m altit ude, ·dO\V h to ne ar the Saba na de rd hwa sout s intte cont h \vhic e~ rang ntain n1oq Sur The s. spot open in rn 2700 at ally adic spor B\og ota a iller Cord the from rges div,e , ador Ecu inro ing fortn m, 3000 e abov best ..... . lops spec ies deve ........ . ·las Papa s ~.JT exte nsi\. e greg ario us caul irosu lan com mun ities in · · Orie ntal ·in the regi on of Para mo 9 N·. It 2 \v bel-o at e\vh som ,, ano, mbi Colo acizo ;~Ivl g pyin occu in gr,:.ss1ands. This spec ies is aggr essiv e is a huge Cord iller a, 420 km long and above 3000 nt\V open area s and can be used as an exaiJ1p1e of est high the of ral seve des inclu h whic ude altit min para the inate don1 to aged tnan ·has leria Espe hovv peak s of Colo mbi a (the Quin dfo ·gro up· of voJc amos of the 'Nho le Cord iller a. Its loca l varia tions noes': Toli ma . 52 15m : Ruiz , 5400 m: Sant a Isab eL and seve ra11 ocal varie ties or subs peci es also indi5100 .t n; Quin dio. 5150 m: N.eYad o del Huf la. cate th~ plas ticit y of the genu s. In this cont exc it is 'NOrth men tion ing that E. gran diflo ra retai ns · 5750 m ~ Pura ce, 4646 m). The Cordillera for-

AE RADIA TION . OF ESPE LETIIN

. 295

tral is very old and the param o envir onme nt and n1erly \vas cover ed by dense forest s on both its veget ation are kno\v n to have been \Veil develslope s and a seri-e s o'f _ahnost unint errup ted pa.raoped and capab Je of suppo rting Espele_tia since mos abov e 3500 m from the south ern Param o de - las· Papits to the- Qtiin dfo Mass if ·a fabbi it so· ID·'-·N. ··-~ -.. the ·Earl y· and Middle ·PJi·o cene.-----But-·the Co lorn...bian Espeletia stock came from the borde r of the Smal ler param os are also scatte red at the north Ve nezue lan and Colo mbia n Ande s. as alrea dy ern lo'.ve r e xtens io n of the Cordillera up to near expla ined, and in its !migr ation along the Colom7'J N . .From the Param o ue Ia$ Pap:as south ward . bian · C o rdille ra Orien!al \Vas halted ·at <the the p{tramos conti nue along the Troncal Cordi llera and its fateraJ ex-tensions and across the ' vol- --. uepre'ss ion' south l)f ·ParanH) de Sunla paz \Vher e the moun tain range drops consi derab l,y \vith no canic regio n of Pasto , Cumb al, Tulc,in.. and param os for 270 km until the rvtacizo- Colom Chile s of south ern Colo mbia and north ern Ecua biano , \vhere both Cord illera s· meet (Fig. 11-16 ). dor south to the Para-mos del Angel to abou t It was only in one of .the last glacial _periods, cer0°30' ~(.Fig. 11-19 ). tainly durin g the \vet Late Glaci aL ca. · 14,00 0/ Only one species ~ E.- hartwegiana, is found in 12~000 B .P .-ca. 10,00 0 B. P. (van de r Hamm en, the great exten sion of _param o surfaces along the 1979a,b; van der Ham men et al., 1980) that the Cord illera Cent ral fron1 Quin dfo to the rviacizo . P'1rat no belt \vas lo\ve red to an exten t that it Col o mbi~no (Fig. 11-1 9,0 ). In the P'iram o de cover ed the south ern section· of the Easte rn Corlas Papa s, anoth er speci es, £. idrr;;boi~ forms dillera (Fig. 11-1 8). It was at this time that Espesmall colonies~ \vhereas E. hart'tvegiana is also found south\.vard .sporadically in Nariiio . letia was able to sprea d farth er south than the Param o de Suma paz and reach the Cord illera (Fig. 11~19 , H). South ward of the Param o de Jas Centr al at -Para mo de las Pap_as in the Macizo Papa s ~ t'vvo other speci es are prese nt: £. schul teColo mbia no. One species of Espeletia close to E. siana aroun 9 the Lagu na de La Cocha't Cerro de grandiflor(z djd reach that' far. £. idroboi is found Patas coy (Putu mayo , 'Narifio ), and £. PJ'Cnoin limite d colon ies at the lv'Ia.cizo Colo mbia no, phyll a (Fig. 11-19 , I). E. hartrvegictna and . E. but two other relate d species havin g ·broa der, sesschul tesian a both have -inflorescences of the leafy sile leave s with shagg ier ., thick er lanat e .inpun1entype · -and --rath er· sessH~}-- -leaves ;- -E.- -(droboi.- had tum-----d€-velo ped-:----~-(-1.) - .E.-... -hart}vegicu1-a ,-~ -~-hich ·pseu dope tiolat e -subl anceo late- -Ieay·e s; ·E. ·pycn omove d mainly to the north , and \Vi·th tretne ndou s · phyll a has nake d inflorescences and broad oboaggressiveness sprea d along the \Vhole: Centr ?l vate- oblon g sessil e leave s. The domi nant speci es c;ord illera , succe eding in domi nating its p~ramos. -are E. hartwegiana along the \Vhole Cordillera The humi d habit at and humi ferou s ·soil in the Cent ral and £. pycno phyll a in the troncal series region facili tated rapid settle ment of the invader; _ of param os exten ding irito Ecua dor do\vn to P~r and (2). E. schultes.i(Jna, \vhich sprea d south'vvard amo.s del Ange l., and farth er south still in the to the Sibun qoy regio n ,- Cerro . Patas coy, Lagu na Llang anati s (or Llang anate s). Anot her species , the uppe r -head\ vater s of the Rio de La Coch a_ occur s in param o-sub param o at the north ern end Guamues~ The most impo rtant speci es in ·this of the Cord illera Centr al in Antio quia: £. occisouth ern regio n is, howe ver, E. pycno phyll a, dentalis ssp. ·antioquensi~ (Fig. 11-19 , K). which can be consi dered a desce ndan t of one of Ther e is a mark ed contr ast betw een the Colthe form er sp;e cies or.of tbejr ances tor,- mainly by ombi an Cord illera s with · respe ct _to speci ation in losing the leaves of the proxi mal parr of the Espeletia: the Cord illera Orien tal has 38 species, --w h.e-reas----th·e----Co-z:diHe:Fa - --C.et=~tr-ai .....has.-----onht..._. 5 . . inflo r.esce nces ._E~ ...p.y_ct~ap hJila._m.ay_d e.veJ op._y_e.ry tali cauli rosul as (up to· 10 ·m) and build \vides..; . altho ugh its whol e exten sion is simil ar (abou t -400 . pre a~ comm unitie s exten ding from the Parai nos km). The contr ast in· sp,e cies diversity is particude Quili nsaya co, del Bord on cillo, del Galeras~ larly notic eable \Vhen individual limited massi fs etc., south ward in Tron cal Sur,. to the large volare comp ared, for exam ple, the_ Neva da del can'ic regio n -at the Colo mbia · and Ecua dor borCocu y-Ch ita·, \vhich has 4 species· of .Espeletia, der, Ot:l wide sprea d grass lands aroun d the \Vher eas the Tolim a-Ru iz J\tlassif \Vith as large or· Chiles~ and Tulca n. From bal. Cum of noes volca m.uch large r param o surfa ces cov~red by .. luxur . here the· param os with E. pycnophyll~ go.Ja rther · iant cauli rosul eta of Esp.eletia, had only 1 -speci es. · abou t to the ·Para m.os d e l Ange ..... l a·n d south \vard .to P~trarno de la Rusia -Gua ntiva .has 10 species.and 0°30~ N ~ At this point the Espeletia popu lation Param o de·:Suma paz 7 species; in contrast'! the stops (Fig. 11-19 ). Neve rthele ss. a long distance Htiila ~1assif in the Cord illera Centr al, &s \veil as from here sudde nly reapp ears·a large comm unity Purac e. has only· I speci €s. Thus ~- the fact that the of · Espel etiii _in the isolat ed · Sacha -Llan ganat is Cord illera Orien tal is \Vider \Vith a large r surface de , at and 3600 m· a'ltitu .2600 eenbet\v is-: -insufficient to. expla in_the great contr ast. \Vhat · moun tains t ' . 1o S in Ecua dor. This ·range is the easte rnmo st of Cencoun ts is the age of the. area. The Cordillera . •

........

296

CUATRECASAS

the con 1ple x of thre e para llel n1ountain rang es call ed the Cor dill era de los Lla nga nati s. The pr.es<:_n~.e. of.E.speletiq .Ln the Sa< ;ha- Llan gan utis , s.o __ far a\~ay from the afo rem enti one d pop ulat ions of the gen us, and its app aren t abs enc e in the t\VO oth er I..la nga nati s Cor dill eras (i\n dra de ~1arin, l<J40) are difficult to explain. It mus t be said that rad e rv·tarfn . .the eas tern mos t of acc ord ing ...... to And the thre e Llan .ganatis~ in con tras t to the oth er t\vo . is extr eme ly wet and is per man entl y cloudy or rain y or foggy: the gro und is svvampy with man y lake s, the terr ain bro ken and rocky, and t orea a with ered cov ion etat the luxu rian t veo b c rnas s of n1osses . liver\vorts, and ~erns. An explanati on that cou l_d be given for the exis tenc e of this isol ated Espeletia pop ulat ion is some sort of longdist anc e disp ersa l. Ho\vever, pro bab ly tnost Es'fJeletia at the end of the Last Glacial spre ad nJuch fart her sou th of the pre sen t limits of the gen us in Car chi Prr:>vince. iV1oving thro ugh the con tinu ous seri es of \vet pan11nos of- that tin1e~ EsiJe!etia \Vas able to reach the Lla nga pati s. Fin ding in th~se mou ntai ns its. perf ect hab itat . Espeletia has rem aine d firrnly esta blis hed up to t~e pre sen t. Aft er glac iati on. \Yith the gen era! lifting .of the par amo belt . .the Llanganatis par arn os \ver e isJo :. ~. ted from tho se of the n1ain Ecu ado .ria n Cor - · di 1Iera. Ft1r thcr.n1ore :·-a perlocC"o}.!n.ie-nse -(fiynt~-ss. in nor the rn Ecu ado r pro bab ly c.au~ed the eliminati on of forn 1er Esp eletia pop ulat ion s ~ituated fart her sou th .of the pre sen t ljJnits of the gen us in ~

C~archi

Pro vin ce.

infrasp~ci!"ic

inlportanc e, suc h as var. galerana of the Volcano Guier~ as (Pa s_to) " ssp.. a.ngelensis, _spre.ad _..mostly .in nor the rn Ecu ado r in c ·arc hi Province at Par amo s del 1-\ngel and bor der ing C'olombia, in the volcan ic regi on \Vith volc ano es Cum baL Chiles~ l"ul c'in .. Tuq uer res. and so on. Alth oug h it is onl y kno \vn from - ori·e collection. the Lia nga .... nati s pt)p ulat ion sho\.vs sma11 differences but they are apfJ aren tly suffici e nt to \varrant subspecific recogniti on (ssp. llanganatensis) (Fig. ll-1 9) .. The nor ther n extr eme of the Cor dill era Cen tral has rath er poo r rep rese ntat ion of Espe!etia pop ulati ons . The spe cies of the area~ £. occidentalis ssp. antioquensis. is very different fron1 E·. hartvv·egiana and is limi ted to the lo\v er pararnos or sub par amo s of tha t region of Ant ioqu ia. The origin of this pop ulat ion rnig ht be different from the mai n pop ulat ion of the Cor dill era. extr en ther nor the in y Onl era. rdill C"'o tern ~Ves . . mity of the \Ve ster n Cor dill era ure ther e par amo s \Vith Espeletia. Thr ee species have been fou nd in . the regi on: E. occidentalis ssp. occidentalis . £. praefrontintL and £. fi·ontinoensis. One hyp othe sis abo ut thei r origin is that they r:1ay have cros sed ove r the Rio Cau ca in Ant ioq uia thro ugh . itig-the Cor dil- • "\r~\lley -~;"epar.iu the· ·of t par i ro\\ riar a .. . . • • .• .. . ' . .. ... . • • •.•• • • .. 0••. • - .. .. . .. 0.. . . .. . -· . . - • •.. . lera s. I-Iere . pas t volcanic activities have fan ned brid ges bet\ vee n the Cor dill eras across the pril nitive rive r basin. The anc esto r could have bee n son1e for1n of E. har twe gian a, giving rise in the nevv hab itat to E. occidentalis and £. praefrontina; fron1 this was deri ved E. ji·ontinoensis (Fig. 11-19, L). reg ion al diff eren tiat ions of

.....

E. hart¥tt·egiana is the only species of the rr1ain bulk of the Cor. dill era Cen tral . · It . seen1s that Espeletia, hav ing inv ade d the Cor dill era from the DILL ERA COR N ELA EZU VEN d .J sou t h ano spr ea · ing to\v ard th.e nor th , has not yet k stoc etia spe! E · first the of nch bra n mai er oth The the so a! ion; ciat spe had the tim e for furt her id hum the in e zon n1o para er lovv the in ed inat orig orn1 unif . ly tive rela a had ~ordillera Cen tral. It era. dill Cor elan ezu Ven the of tion sec tern wes e. cen olo H . the ing dur larly i<;:u en \·ironment., part n mai the ng owi foll n ctio dire rn este th\v nor a k too ulapop in seen be can ns atio Nev erth eles s, vari Val ma Cha the of s side h bot on ge ran n ntai mou _ . that ns atio vari , area the of ts par ous tion s in vari f as-o r e· a"in r calie·r · of ;-pafb " )el; u-pj: -t1ie ig cn!i :rea ·· le-y~ ea liat re1i iff~ fbf"d uf"r l5 ixed f " ·arf)T -see iiflc f be- gehetrc con and o ing Dom to San and ata, CuJ La , rida Me ion ulat pop The us. stat ific eno ugh to \Varrant spec dilCor lo ujil ·'Tr the ss acro ard thw nor ed tinu . nt iste s · con s ure feat has s)if j\rfa of the Rui z-T oli1 na ered suff ly bab pro s ion ulat pop ary prim The ."~ lera cen. (ssp cies spe sub a red side con eno ugh to be and on regi the in s nge cha atic clim stic dra from · raBar de s amo Par of ts plan the 9o troandina) as ther al udin altit the of ions illat osc y man too from · er oth s): ensi rag brtr . (ssp sas rno Her gan and p~riods of hig her the that r clea ms see It s. belt mal . . . (var ores raft Plli de amo Par in nd fou are s iant var er lo\v n fror ia elet Esp ted nina elir re atu per tem .. o, ning Do.I to San de s atno Par vegasana) and in the n ther sDu . the e, anc inst for as, ges ran n ntai mou the n·z), raru rno . (var as Pap de l\~.1oras, and de Las n, 116 Bata del. os ram (Pa era dill Cor the of tion sec pair erlo\v the uce red to y enc tend tatt er \vith the ~arne forces The ). etc. , dor nba Zuz and ra, Neg La interm d tren s Thi ces. cen ores infl of leav es of the facts relic of us stat the to cies spe e som d uce red se The la). h.yi nop pyc £. { cies spe r the ates in ano ing -ext inct ion. In any case~ it is ce.rtain . that the var iati ons are signs of inci pien t spe ciat ion. oseas ng stro and s nge cha t men iron env al tinu con . · read csp .\vid the vvith urs occ A sim ilar situ atio n of ts par t n1os _ in ate clim the of tics eris ract cha nai t n1os the ly bab pro ~ a hyll nop pop ulat ion of E. pyc aeren diff ic m no. taxo rked zna d ore fav on regi the e som ~'s sho It ia. elet Esp rec ent spe cies of

RADIA TION OF ESPE LETIIN AE

297

popu lation s. no\v restri cted to the summ its of tion. This is evide nt in the living Vene zuela n speParam os de Cend e-Jab 6n-L as Rosa s (Fig. ·11-19, cies of Espe/ etia. The speci es close r to and more A). The group of snutll er plant s with tuber ose repre senta tive of the ances tor of the group is £. ·scltultzii, . . ·ordin arily · with · large inflor escen ces, ··· -·corm s .. is conce n-trat ed in p-ar a-mos· -of ~southern Truji llo and Meri da State~ being endem ic to small thick ly \V'hite lanat e leave s and a trunk usual ly no param o areas and adap ted to very wet groun d rnore than 1 m tall:. it thrive s best' in a \\'et '-grass (e.g . ., E. tenor e. f, .. n1arthae)~ some may be more land en vi ron rnen t at high altitu des (e.g.~ 3o00 m). broad ly distri buted bq.t are ecolo gicall y limite d to E-. sch ult:.ii has great vitali ty \Vith a treme ndou s mars hy or less \Vet areas , e.g. , E. batata . E. ·nana, capac ity to accon lnlou atl! to less than optim al and £. ~veddellii. The cente r of speci ation of cond itions . As a resul t, its popu lation s cover E'speletia in the Vene zuela n Cord illera proba bly large expan ses of param o at high and midd le . and was the regio n of Apar tader os, \Vhich is conti nueven lo\v.'C2600 m) , altitu des , and readi ly colon ize ous \vith the param os of Piiian go and Timo tes. A newly open territ ories such as aban done d fields subce nter was the south ern Truji llo regio n of Pttrand clear ed fores tland . The area cover ed by E. amos del Guiri gay-L iano Corr edor . exten ding to schu!Lzii today is much large r than it \voul d be the bord er of Truji llo-M erida . witho ut huma n interf erenc e (Fig. 11-19, B). Of the other nine speci es gro\v ing in this Cordiller a, two stron gly diver ged from the primi tive Coes pelet ia genet ic type by adop ting a distin ctive kind of indun 1entu m. In E. sernJg!obulata the indum enThe genu s Coespeletia is an earlie r stock close to tum is \vh]te and super ficial ly appre ssed. In Espe!etia , also with poJyc arpic cauli rosul an bioaddition~ its heads are small er, and the invol ucres . type, but diver ging by a mono chasi al type of and flo\ve rs also ·show mark ed differ ences . E. inflorescence~ It sprea d throu ghou t the main ariste guieta na has an indum entum uniqu e in the Meri da Ande s and reach ed their sumr nits . adapt genu s, tome ntose -velu tinou s, surel y mark ing a ing'"' .to the extre me minim al therm al condi tions dis-tinctive genet ic line. £. semiglobulata gro\:vs in . te of COm qjned with the mar~~9. S~?.-~Onal clima Strict ly . \Vet edaph ic COnditions at high altitu des - - - . .. -- - . . - {Sier-ra~- Neva da-C ulata Ivtass.if) ;. E.~-aristeg.uietana _______these .para mo...an.d s.upe rpara mo areas . ~fhe . p.rimi- . tive speci es of c·oespeletia' proba bly gro\v ing at is only knO\Vn fronl Truji llo in Cerro s de La Crismedi an altitu des (3100 -3500 m) . had panic ulate tal ina (2600-2800 .m), possi bly a relict of a form er racem iform inflor escen ces. that is , deter rnina te Iarge r area. The other seven speci es· are close ly· racem es with short ly branc hed pedun cles (e.g., relate d to E. schul tzii . shari ng \vith it the san1e C. thyrs iform is). Thro ugh emig ration and settle kind of invol ucre m~de up of narro \v. oblon g, ment at high altitu des (4000 -4500 0 n1), a strict , acute phyll aries. They must have been deriv ed from this speci es or from an ances tor havin g an . simp le-rac emos e type of inflor escen ce \Vas, ho\vever, firmly estab lished . Furth er evolu tiona ry hirsu te recep tacle . Thes e seven speci es all gro\v steps i~volved the reduc tion of the numb er of in humi d true param os or in super param os, most head s in each inflor escen ce comb ined with headof them (five) in Param os de Apar tader ossize incre ase (C. spicata to C. tinzotensis). A final Pinan go-Ti mote s-San to Dom ingo. and in the step resul ted in the mono cepha lous type of C. neigh borin g regio n of ·Truji llo. E. cunic uloru n1, n1oritziana, \vhich has the large st capit ula in the only in Param o de Los Cone jos, has, like the g_~nq?..:. _I~.i-~~j~-~J~9_.~.h.~ .- se~~i~.~--b~?~.~ ~q_~JF.eed to .original _.t.yp.e. .o.f.E~. 5.cbtt!tzii_~ :catb~.r_p_~rf~Gt _cti~l:l~.thriv e in the highe st altitu des of the Vene zuela n sial inflor escen ces but their foliar · struc ture and superpar~mo (dese rt param o in Chap . 3), which , the consi stentl y 'hirsu te recep tacle make a diver besid es being extre mely cold., is a rocky , sandy , gent ende mic type. The tende ncy for reduc tion of or grave lly, arid, and expo sed habit at, \Vith frethe numb er of elem ents, and for asym metry of quen t or regul ar diurn al freez e-tha \v cycle s of the the inflor escen ce, alrea dy stron gly ~eve loped in ' soil, a chara cteris tic of the perig lacial zone E. schul tz ii, has led to a gr~up of speci es of -(Fig. 11-21 ). small er size, with a subte rrane an .. often tuber ose , The heigh ts of the· orogr aphic nucle us of the caud ex, ·\vith very fe\v or single head s · in ~ the Sierr a Neva da de 1v1erida sensu lata, and the · i~nftorescences. The cente r of radia tion of this inter medi ate zone of Apar tader os benv een th~ group is proba bly the param.o regio ns of Tun arneSierr a Neva da of Ivlerida.,· the Sierr a of Santo Guiri gay-N iquita o. The speci es sprea d to Param o Dom ingo and the Sierr a · de La Culat a (del de Los Gran ates- Santo Dorn ingo and to Apar taderos._ and also to the Cord illera de Truji llo · Nort e), \vere the cente r of speci ation of Coespeletia~ One of the more primi tive speci es, C. elon-: north \vard to Param os ·de Cend e-Jab 6n. Toda y gata , sprea d along the Sierr a del Norte , and later E. ulotricha is a relict of once more \Vide sprea d

298

CUATRE CA.SAS

one rnay conclu de that the reprod uctive ~111d rnorpholog ical strateg ies develo ped by ('oespeletia re p:r:~s~::}t the n1ajor success in the evoluti on of ... ... ... . . . ... . . ... .:. C h '1 p • 1 .) . . ..:s f t.· e ··e· ·r· : - ~~- -- ~: .)r.,., < ..... _ ,."J . 1,.,., J.- r!::·~-~ f'-T i .,· 11 ::1 t::

- ~

_,..,..,~

. IA COMP/-\RATJVE DISTRJBUTfON OF ES:0 ELE-:T V\ffTH OTHER TAXON OMIC GROU PS 'The pattern of distrib ution of E.'·;peletia, sho\vin g

an overwheln1ing nun1ber of 38 species in the Easter n Cordill era.. against only 4 spp. in the ! ! Centra l and 3 in the northe rn end. of the -Weste rn Cordil lera (see Table ll-1). compa res well \Vith a fc\v other taxono mic groups . e.g., Puya (Brorn eliaccae ) (Fig. 11.-22). This genus has 8 species in COESPELETIA the Easter n Cordil lera paramo s. only 2 species in Presen t areas... . ;.;.;.,.,. the Centra l Cordill era, and I species in the southern section of the Weste rn Cordill era. 1-\Jthou.....gh the totab.n umber of species. is lo\ver than that of Espeletia, the pattern of distrib ution· is paralle l. FIG. 11-21. Distribu tion of c:oespeletia . . But Puya .has in auditio n 4 species in the Sierra Nevad a de Santa ~·1arta~ . \vhere E"speleria is gave rise to C._spic~zta, C. tbnoten sis ( \Vith forma lacking, and 1 species ( fJ. occidenralis) in the Farlutescens) ~ and c·. moritz iana' which migrat ed in allones .de Cali_, q highlan d param o \Vithout Espeall direction~ horizon_t~lllY and upward ..to the • letia. Beside s, species o( Pu_ya are also presen t in mount ain slopes, settling locally according to ecothe pax~DJ9S ..9.f.CQ~t.a Ri.c.a .and in thos.e .oJ Ecua- . lo.~icai pr&"ferences. Todav , they. domina te the dor and northe rn Peru, where Espeletia is-lacking ~~]inl "con1n1un1tie's and"~the landsca pe of t!1e (Cuatr ecasas , 1.979b; Sn1ith and Do\vns~ 1974). In largest part of the superp ararno zone (or desert predict ing the numbe r of species in paramo paran1 o ). of the central massif of the iVIerida Corislands and in gene raJ in high .A.ndean '~islands,"' diHeras. This zone in great part coincid es \vith the perigla cial region (Schub ert, 1975; 1979). The .. some \VOrkers consid er the .size of the islands and the distanc es separa ting them to be most import area was describ ed by Monas te rio-( 1979, Chap. ant variabl es (Simps on \ l uilleum ier, 1971; Simp3), as '"Paran1o Desertico,-'~ becaus e of the scarson., 1975; Vuilleu mier and E\\·ert, 1978). In _.city of.othe r p!p.nts in ..associa tion \Vith sho\vy fraiorder to unders tand pattern s of speciat ion, radia. lejones . Nevert heless" throug hout the perigla cial tion~ and distrib ution, it is iinport ant also to conzone a natura l landsc ape of cauliro suletum of varsider the kind of 1norph ologica l devices the iable density is \videsp read '·· consisti-ng of a fe\v propag ules have to faci1itate their distribution. species of Coesp.eletia, n1ainly C. tim.otensis, The absolu te lack of any such de\·ices in Espeletia \Vhich is extrem ely irnpres sive and· unique . In restrict s its distrib ution ~~--~-~e_ s!~.Y~- n_a!h _gf _ pr9~ ..... ~pite of the great difficul ties solifluc tion poses for gressive local expans ion of each gregari ous popu-·seedllng'" sur\ii\·ar-{the ···moitaJ.ify--iate-·inay reach lation. The contro l on means of propag ation 1 OO~Io )., the species have manag ed not on]y tq surlimits the area of interbr eeding po pulatio ns , causvive but to domin ate large areas of this hostile ing more freque nt speciat ion. Espeletia- popula habitat . \Veil-b ased estima tes (A . .P. Smith, 1978) l physica or ical ecolog by ted separa tions," . · sis tin1oten C. of uals indicat e that the adult individ . . usually surpas s 100 years of age. The cauliro sulas . barrie~s, even small ones such as a band or girdle of forest, ·or a rocky cresL mav remain fore·v er of (~oespeletia are best equipp ed morpho logical ly isolate d withou t opport unities for interbr eeding ; to gro\v-in this enviro nment. The massiv e, colum·the a~jacent popula tions \Vill continu e to diffe-~· nar trunk has a cover of marces cent,. exceed ingly Convers-e]\·. the seeds of itelv. entiate indefin . tjght Iea\es~ incornp arably thicker , n1ore con1"' . Puya are light and have a memb rC;l nous margin al pact~ and insulat ing than that of any Espele tiinae \ving thfit facilita tes dispers al: . some species, furof the \Vetter Colom bian pararn os. The conside rthermo re, e.g.~ P. goudot iana. may have a long, able thickne ss of these giants of the param o flora gives the impres sion of stabilit y and tnajest y. . thick=' spicifo rm, n1ultiflorous panicle toppin g a \\'hich reicase s .thousa nds robust axis 2-6 rr1 high, ...., Consid ering the comple xity of its morphology. 0

l.::;l

.•

• ; ,. •

- ··

•

•• - · ·

•

OF ESP ELETIIN AE

RADIAT ION

2·99

76° ...

.,.

l2°t--- ---t--------;. ....'- ---···

I I

·--~-· ,--·--

0'<0~:

I I

'f'A.

; ~,..

I .. .

10°t--- -.._'- - i

•

j !

•

-·--·-·-·6 spp.

10 spp .. -· ,_ __

: :-. ., _ _ _ _ _--H4 0 i •

QJ I

7 spp. 0 'v

., Il!.

•

·-!

- - - - - - - ; . . . ,·

.I I

11-22. Distribu tion of paramo species of Puya \Vi th number of species per paramo block. Compar e with Table 11-1 for figures pertaini ng to Espeletia.

4 spp.

Subandean·and r, Andean forest ·---,_~ l . --------~--------~----------~00 ~---I . ..... . ...,.I

FIG.

t@l Priramos

I I I

78°

of 'vvinged seeds that are easily carried by the :-~!.1.1ds .frQJn _9-D e_.P:~I_<Am.Q__g.t:-~.(1.19...a.n o.th_~r ~ . .£ossib l.x for this reason , P. goudotiana,- with its tall inflorescenc es, is t~e most widesp read species in the param os of the Colom bian Easter n Cordill era, from Bogota to Norte de Santand~r. The tall P. aristeguietae also has a ~ide range, from southe rn Trujill o to tvierida and· Tachir a, to the Sierra de Perija and the Nevad o del Cocuy. Similar ly P. hcunata, 2-4 .in tall, has spread from the Centra l Cordil lera of . Colom bia, southw ard throug h Ecuad or as far as Cajam arca, Peru. The Cordillera Orient al is. obviou sly a center of speciat ion f9r Puya, althou gh most ·of the species have a larger range than do individ ual species ofEspe letia . .·f or examp le, the Cundin amarca area has 11 · species of Espelet ia , most of them \Vith a restric ted subare a, \Vhereas its 7 species of Puya

r-:9 ~

' 1 50 100 0...__......... ~0 · Paramo specie s of PUYA . .__.;_~ km

76°

are more widesp read and at least 5 of them .e.xte.ud__f.a.rthe.r_thro_u.gh ~b_e ..B.o.¥-ac_a.. are.a .and. l.to_ ·Norte de Santan der. Probab ly from this center of speciat ion (Cordi llera Orient al). Puya radiate d to the !Y1erida Cordil lera, to the Serran fa de Valledupar- Perija, to the Sierra Nevad a de Santa· Mart~, to the Wester n Cordil lera of Colom bia and to the paramo s of Costa Rica, either by progressiv e terrest rial propag ation or . by )<?ng-distance transpo rtation , or both. An interes ting . compa rison can be made betwee n the distrib ution of Espele tia .·and Puya~ and that of Diplos tephiun z species in · Colom bia. Displo stephiu m is a. genuin e Neotro pical Andea n genus of Corripo sit,)e \vhose ·maxim um develop~ · ment has been in Colom bia (57 species ) and Ecuado r-Peru (50 species ). The adapta tion to \vind dispersal of the Diplos teplziu m propag ules;

CUAT RECA SAS

300

Which are light achen es \Vith a papp us, has facilitated inter param o diffusion and interc ordill eran trans porta tion .. Diplestephiun1 has thus not. been· exclusively depe nden t on. the lo\vering of temp erature and veget ation belts to expan d· its populations. Long -dista nce wind trans port is a reaso nable expla natio n for the prese nce of Dip/oste:Jhium in the Sierr a l'!eva da de Santa ~-tarta, Cerro s de Tatat na, and Faral lones de Cali (Cordiller a Occid ental ) and the Costa Rican param os, all lackin g Espeletia. It also expla ins the great profu sion of speci es of Diplo steph hun in the Central Cord illera , in contr ast to the li1nited speci es of the non-\ vind- distri buted Espeletia in this area. The main cente r of Diplo steph ilun speci ation in the north ern Ande s is also the Colo1 nbian Eastern Cord illera with 31 speci es, 25 of them endeTABLE

tnic to the area. The Sierra Nevada de Santa Mart a follows as a subce nter of sp~ciation with 12 hi.ch l'l ar'e·e nde.tn ic to· the···. islai1d. · ~· ·species~ of·\v_ The · (~en tral Cord illera has 17 s·pe cies with SJ ende mic to it or exten deu to its conti nuati c,n, the Tron cal Sur (R species. including 1 addit ional ende mic). The vVestern Cor ,:! ill~ra }~lS 6 sp~cies 'Nith ·3 enden1ics. Only 2 speci es are feund in th e rnain Vene zuela n Cordilleras (Nh~rida and Trujiilo) ; the other 6 cited fron1 \/ene zuela (inclu ding 3 ende mics ), orogr aphic ally corre spond to the Vene zuela n easte rn side of the n1ain Colon 1bian Easte rn Cord illera . It indicates that Diplo stephiun l radia ted frorn the Colon~bian cente r of speci ation , from the soutlnvest to the north east .

11-2. Comp arison of the numb er of species and endemics (in bracke ts) of Espelelia and Diplosleplzi!un found in compa rable panun o sectio ns of the t.wo n1aii1 Colom bian Cordil leras.

------------------------------------·-------------------------------------------------------------------------------------------------------------------------------Cordil lera .Ct:n tral Cordil lera Orien tal Diplvs tephil an

Espe!etia

Diplos rephitt fn

Espele tia ~1 acizo

P'='iramos Almo rzader o-Rom eral

11(5)

6(6)

ellipti.c um dentatcun " oblongifoliurn canescan.~· nzu.tiscuanurn brassicoidea tolinzense conglot;:erata juajibioi duga1u.iii lacunosum revolutum ros1narinijoliur glutinosum apiculauun Nevad a del Cocuy 7(0) 4(2) colom bianu m lopezi i lacunosum cleefii revolutum curialensis • ..alveo/atum -annen tar'tan a· rhon'lboidale tenuifoliurn huerrasii ~facizo Bogot a·Sum apaz _10(3) 11 ( 10) oclzraceurn gran.diflora • • rupestre orensz•s 1nrraa ., . pIu.a oida!e f. rho1nb ·. tap;ro . revolutum chocontana a/veolatum ca...-etana phylicoides barcla"vana fosbergii kil!ipii • huertasii sum1napacrs juajibioi ssp. cabrerensis heterophyllu1n uribei argentea estanislana s F:~·nd Ievana

1(-1) hartwegiana ssp. centro andina

Ruiz-Toiima 6{1)

erivphortilri · rupesrre schuli:ii violacf!unz ret:olu tunz

tolilnense

Nevad o del Hufla 7(2)

1( 1)

hartwegiana

var. mora rum

glandulosum ritterbushii v... ./ehJna :ii .... . . .. ... n. . . . . -.-... - .-. .nianul -. - - ·. ---schu!r cinerascens v. puracense pittieri floribundtun bicolor Par amos del Purace 9(0)

1(1)

hartwegiana ssp. harcwegiana

rupesrre ,.,ii hartu:e ~ . spinu.loszun glandulo.nun schulr=ii v. lehnzaniat~urn • scens v. puracens ~ cutera pittieri fioribu ndtun v. puttanayense bi~olor

------------

---------~---~----------------------------------------------------------------------------------------------

:.·... _

but stopp ed abrup tly at the easte rn borde r of the the stron g, sts tha~ Cord illera Oriental~ This sugge . . north easte rn wind s blowing over the Venezueh1ir .· .qfrE~o.s and ag~inst· the· Aride s -are:~-:~_ai~ly ..respon~ .·sHll~e- for the limita tion of the Diplas·tephi~_tnz mig.:·· ration · into . Vene zuela . · .CQn~erseJy~ - the EspeJ etiina e \Vith their heavy, 'b<:i.re vac~enes~ and . . their slo"vv progr essio n on the grour id. \voul d hay<;.~ no difficulty iri · migra.ting in_. ar~,y di_re~ti.on . \v-i_t'h·· . respe ct to \vinds. The capab ility for·. pl'uris pe~ifi. c _·~popula.tiofl·s of th~ . CoJ9rribiaii ·.cordiller?~ Ceo:. · in Diplostephiurn . . .. . .''·· - . . tral is show n i~ . a ~ coinp~lris'orr. 9f·sirrgle _f)aram9 . · areas .in ·bqth C~rd}ll__~r~s (~a pi~~ J t~~ X:·_. ·· As Tabl e I 1--2. indi~~.te~_,;:·' -singl~ p_~-~~-iiio~ n.uclei"' in the .Cord illera Centr al, _populatcd)?i a sing~c\ speci·es of Espeletia, may huve. as -~~ElY species o{ os of }' the DljJlO~)·tephiurn _ as comp arabl e par:a m_ · Cord illera Orien tal. The Cord illera <:;entr.a l par- . amo group s of Ruiz- Tolim a, Htlila and :Purace · (all \vilh ·only E. hartP /egian a) l_l.<ive~ ·respectively, , 6 . 7 . and 9 species of Diplost//pl1iuJrt:, This conipares \Vith the param o group s of Almo rzade r9Rom eraJ, Cocu y _a nd·· Bogota.-Su~apaz:; haying, e.s and 11.~ .. respe ctive ly·, 6~ 4, and. l f Espeletfa. sp~ci . 7, and· 1.0 DijJ!ostep~zi~Ln1. species. No.te.._tJ~~- higher_· numb er of local endemic specie~ o_f Espeletia in r-· numb er 'of era. and ...tbe }O\Ve o.rdi.H the Easte r'n.:C . .. ....-. -··-· ···- - ·-- -- -- ··- - --. . . local DtjJhJstephiLLnl ·endemics;·in:·.p<?th Corc)ilieras: the easie r ditfus ion· ability of the -papp us-be aring DlJJlost~ph/uJn · propa gule·s · expla ins the..r· differ ence. The effec ts of an insul ar situat ion on ·diffe rentiatio n . anci speci a-t ion of Diplostephii.un are extr~mely yisibl e in ..the Sierr a ·Neva da:.de ·.Santa -l\1ar ta (Tab le 11-1) w-here th~. genu·s ·is repr~ . sente d by .11 endemic., .m.uch-ciiyersi.fi·e.d speci es,; in addit ion .. to one (D~ rosmar1nifolj~u1!) rathe r genus., comm.on in · the . Easte rn -Cordillera. . The ...... must be. very ancie nt on that . mountain., but.. th~ o'rigin cann ot be obtai ned ·beca use of the;· date _difti~u.l. ~~:e-~j_pjq~_pt.ifY..iD.g,. Com_QQ~~ ~~a-~__ g~ nera from

30·t .. · ..~ .

';::

.·• '

REFERENCES. .

...

:-... ./!> , .

p·ollen depositions.

A'ristcQ.ui_eta.-L·.

••

•

t .

•

f:ornpositae ·.. In Flora·de Venezuela ..

1964~

· V~L 10~ ·pt-: .i~.;··pp._ · 407~9i"~-~c·a·f.~ca~l -~nstituto B6i(~;- : - ..

nrco. . . Azoca r.· •: A.~ . and11 .M. tvfcHl~sterjo ·. ' 1.97.9·...; Variabi.lidaJ .

· umbic ntal en_. el · P'!ran1o de, ~iucLfF<}jf·...In £/ IV!ccii~· AJ1ibieizte : P!in?"ln o. l\il...' -L.-· Salgadt)~ Ln.bouriau. ed .'~ . pp: . :119~ I:S9~_:r~·'c ,i-r~t?~ls ;·· : EdTc·i.<'5~cS. · Q0ntr~) \Esr~i~)~_: .

;\'\:a. niath>' . .. , s. · . Benth am : G/ I H-73~ Conij-,osh~tc.· f n Geliei·u 1'/antarunz, G

! .

·- ...

.. •

•

Bcnihan'l'.'.-r ani.l. :· J>. Ho.o kcr, .~: cds.: , -· Vbl. . 2~ . · no : ··I ·~··pp .·· J63~53J: -; i~:ondt)d: ·. t~)\'C II . - R:~.cye. ·.' Willi~uns an(L_·. . . te:. . .. i'rorga : .. ... . ...... •

- ..

- .::.. ~

•

· Cark{ liij)t·: S : 1966--: ·'".WoocJ· a:na·t~):my of·G'<(;?nip<)sitac·: A sum-:~·:. In~ry. with c<·)~mcnts o~J~rci.s>r.~:: c<:J~~~§.!.iing w~)?d ·cvo. lut. . .. . -44 ' . i on. Aliso 6:~~25:---: ~: 197-L · /:"ilt~nd · . . hio(ogy::- :N.e\¥:\i'X:~r.k: · · Colum hia. Univcrsi ty·:.·Press· . . .. - - - . 1976. Tribal · intcr.rchitiohships ·.a:rid phyJqg cny· of . .the Astcra ccac.-A/iso ~k ~-65-~92-. · ·C,Jce f, A. N .: 1;?78. Chan1it~ri$rics:_~f' neotn.)picaf r4r,~n1o ons.~ ....In Gif o· c.. rcfat'i .tarcti n· - and - its ~s.uban ve·,.....f!ctalio :"" . . ecological· n4atiQ~rs-· ·:. 'b¢nf;een t/ze.::'s qulh!?r n ,. iemper t.iff:·: · ~one and the· · u~opical:·:·inoLnifatnsor- C.;: Tn>ll:. and \V Laue;r:.. eds. ,- Erd\vis~cnschaftl~ich.~ Forschung~··vol. 'i_f, pp. 365-3 90:-Wiesbhden:: E·ranz S.teTner Verlag. phi·caF pos.ition of ,the ricoe.ogr;a - - - . ·1979\ The phytog . . . tropieafyaS£U1-fi.f-I)ar:H ffiO~- fiDr~ . --.-S\Lith-spechtl .rc:fC.r.cnc.C: ' rdil t~ r a..:o··r-.ie nt aJ. ~ Jif :~Tl~Op i c.al ·-t 6·.':{ t he-- Go 10;in·b.,f~Hl .-G.o 'boti~nr ;· .e.Qs... K:. tarscn a.na Li: .fiolm..:N ielsen'!. pp.; ·.t?~~ f84;rL:o.ndon: Acade·mic ·at~ess.:. •'• •

a-'

•.

• •

•

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__._:·-~ t98I i -'ftfilt~\.~egetatil>tr~of rlie;j)_lir:_~iptQ~,-_(J.f th~ .(1Jionibhur

Cdtdfll.. e'-~~i~D~.;;,;,,,fiL. Disserrati"ones... Boiai1icae: 6.1: ·Vadu·z: Chi'meP. 1 C:r<>nq~is:r_; AJ:,:~t.~.$.o:.Li?hylogenx- and.tai<:ono~y bf_the ·C9ni~ · posiiae ...:A'ine·,;:.....Nf'i)iL 'i'~;tL . 53·:=4.1.'8-;-S_t.i.':. · 1977. ~~ The ···;·(:0-mposi'rae ··: revisited..· -13i·ittp:~lia· · 29: -·3·'•· ;· --· · 1..). .·rj~-7

~ Col' .Ob~:er:vuciones_.geob.ot~hi'~as Gtiatre:.ea~Tas; J.. ·J9.3:~L· ·. . . ombia . Ti·ab. · !Vh.LL 1Vac. Cienc.. ·iVat;,-' Ser. ·Bot. 27:·

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high A·noes~ B·Lill: . [Jhica go·1\ rlu. H/st. ivlus. ·19;)..:::3~ ~--- ·1949a~. Les, espe~ies del genero · ~speletiq.~ BufL. lnsL C.ata!. ·Hi~·t. 1Yat..· 37: 30-.41...

----~ I949b;~:Rosett~. tre·es~;-a .tr'0picafg.rowth-./ortn...·fJuli~

Chicago 'Nar~-:-His_t: ·Lvtus: . 20~· &7 .. · . bia,~ ·fn ·co10m types:· ation~ - - - . 1954a . .·o utline ;. of v~ge.t . . - . "· ·.. .. . Rappo tts·-- Contt-n:· 8~m~....6 ongl:.. /i1t:, -Bot. ·~ ..Sect. ·...VII: 77-78 . Rap~ : _titr. 1954b~ . Distri' bution of · th~ ge.nus>:Esp.ele . . . po.i·ts ·· Cointn'. ~ srmt! Cot1gr..' 1nien1qt~ · .f;J'ot~ Sect,~ IV: • '-

ACKNOWLEDGMENTS

The contin u_e d .suppo rt ot Dr. Dieter \Vasshausen. Chairciated ; as ·appre y, is much en·t 6f Botan ep?trtm man of the:_D . . . . . : ib_ tht:· CQ..ll_e:~b~r.ation of Miss Alice TangerinL \vho prepar ed to. ·Drs. Harol d ul g:ratef am I : ations illustr the. of . most · . .. . . . · R'Ob+nson arid. Richa rd Co,va n for :.their ·,assistance· in the fina] . _ eoitin g of the manuscript. ·Appreciation . is also . · ~xprc sS:e d to Dr~. T. van der Hammen, Leo J. Hicke y, and for valuab le inforin atioh on the '""""g:eolo£....ric A. l\1. Cleef . his. . tory and ecolog y relate d the subJect-'. . '

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b:··. Cienc~ bia·. Rev:. . .r.tcad. :Colorn . -:-i·n: ·tzi,.:,. -268: . . ----- . 19'68. Param o. vegeta t·ion anet its: lif~._.:for;ins, Collo~ · .qLtiu1n · Geograp'fz.icain ..9: ·I63""7'.f.8(j . .

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ED ITE D BY FRAu'J"(:0IS

· LLELTT\1IER

Al'~D

rvi r\X IIv iiN A M ONASTERI0 ·Published by Oxford lJniversity Press and th~

1\n1e.rican Museum. of~ Natural History

Ne \\. York

Oxfo rd

OxfOrd lTniv'ersitv Press .I

1986