proyecto final artículo historical biogeography asteraceae_panbiogeography.pdf

8/11/2019 Proyecto Final artculo Historical biogeography Asteraceae_Panbiogeography.pdf

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Caldasia 23(1): 21-41

HISTORICAL BIOGEOGRAPHY OF THEASTERACEAE FROM TANDILIA AND VENTANIA

MOUNTAIN RANGES BUENOS AIRES, ARGENTINA

A nuestr a am iga P i l ar

JORGE CRISCI- V.D epar tamento C ientfi co de P lantas Vascular es, M useo de L a P lata , P aseo del B osque s.1 9 00 . L a P l a ta , A rg en ti n a. c r i sc i@ m u seo j cn ym .u n lp .ed u .a r

SUSANA FREIRE-E.

D epar tam ento C ientfi co de P lantas Vascular es, M useo de L a P lata, P aseo del B osque s.1 90 0. L a P l ata , A r gen ti na .

GISELA SANCHO

D epar tam ento C ientfi co de P lantas Vascular es, M u seo de L a P lata, P aseo del B osque s.1 90 0. L a P l ata , A r gen ti na .

LILlANA KATlNAS

D epar tam ento C ientfico de P lantas Vascular es, M u seo de L a P la ta , P aseo del B osque s.1 90 0. L a P l ata , A r gen ti na .

ABSTRACTTandilia and Ventania are the only systems ofmountain ranges situated in a grassysteppe or pampas in the political province ofBuenos Aires in Argentina. Tandiliaand Ventania have a high taxa diversity and endemicity. A historical biogeographicanalysis was carried out on the basis of distributional patterns of species andinfraspecific taxa of Asteraceae inhabiting Tandilia and Ventania in orderto establishthe relationships ofthese mountain ranges with other areas. Two methods were appliedfor the analysis: panbiogeography using the compatibility track method and parsimonyanalysis of endemicity (PAE). Thirteen areas were delimited for the study: SouthernNorth America and Central Amrica, Southern Brazil, Uruguay, Pampa, Tandilia,Ventania, Chaco, Sierras Pampeanas, Sierras Subandinas, Mahuidas, Patagonia, Cen-tral Chile, and Northern Andes. The units of the study were 112 taxa inhabiting

Tandilia and Ventania (endernic, naturalized, and adventicious species were notincluded). Both methods connect southern Brazil, Uruguay, Pampa, Tandilia, Ventania,and Sierras Pampeanas, showing that the Asteraceae biota ofTandilia and Ventaniahave closer relationships with the biota ofthese are as rather than with that ofSierrasSubandinas, North Andean, Chaco, Patagonia, Mahuidas, and Central Chile. TheAsteraceae ofCentral and North America appear less related to Tandilia and Ventaniain regard ofthe rest ofthe areas. The evolution of Asteraceae in Tandilia and Ventaniaand related areas is hypothetized to have been affected mainly by Tertiary and


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Asteraccae from Tandilia and Ventania

Quaternary geologic events. The discontinuous pattern found, occurring mostly inelevated areas, is explained principally by vicariance under dry conditions.

Key words. Asteraceae; biogeography; Tandilia; Ventania.

RESUMENLas sierras septentrionales, o de Tandilia y las australes, o de Ventania, caracteriza-das por su gran diversidad orgnica y su alto nmero de endemismos, estn ubicadasen la planicie bonaerense de pastizales denominada pampa . Con el objeto de esta-blecer las relaciones entre Tandilia y Ventania con otras reas, se analizaron las dis-tribuciones de los taxones especficos e infraespecficos de Asteraceae que habitanestas sierras. Se delimitaron trece reas: sur de Amrica del Norte y Amrica Central,sur de Brasil, Uruguay, Pampa, Tandilia, Ventania, Chaco, Sierras Pampeanas, Sie-rras Subandinas, Mahuidas, Patagonia, Chile Central, y Andes del Norte. Las unida-des de estudio fueron 112 taxones de Asteraceae presentes en Tandilia y Ventania(taxones endmicos, adventicios y naturalizados no fueron incluidos en el anlisis).Las distribuciones de estos taxones fueron analizadas utilizando dos mtodos:panbiogeografa (compatibilidad de trazos) y anlisis de simplicidad de endem ismos.Ambos mtodos conectan sur de Brasil, Uruguay, Pampa, Tandilia, Ventania y Sie-rras Pampeanas. Las Asteraceae de Tandilia y Ventania estaran ms relacionadas conestas reas que con la de las Sierras Subandinas, Andes del Norte, Chaco, Patagonia,Mahuidas y Chile Central. El rea de Amrica del Norte y Amrica Central aparecemenos relacionada a Tandi1ia y Ventania en relacin a las otras reas. La evolucin delas Asteraceae de Tandilia y Ventania y las reas relacionadas ha sido probablementeafectada por los eventos geolgicos de los perodos Terciario y Cuaternario. La dis-continuidad actual en el patrn de distribucin de las Asteraceae, restringidas engeneral a las zonas elevadas, es explicada principalmente por eventos vicariantesdebidos a condiciones ridas.

Palabras clave. Asteraceae; biogeografa; Tandilia; Ventania.

INTRODUCfION

Since Darwin'sjourney in Argentina in 1835the geology ofTandilia and Ventania and theirinhabiting biota have attracted severalgenerations of geologists and biologists (e.g.,Darwin 1846, Alboff 1895, Hauthal190 1, Ca-brera 1938, Sota 1967, Teruggi Kilmurray1975). The political province ofBuenos Aires,situated in central eastern Argentina, is coveredin most of its surface by an herbaceous, grassysteppe, ca\led pampas . The pampas are a plainsituated at the sea level, or even below the sealevel (like the big depression ofthe Salado river),with the exception of two major systems of

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mountains ranges or sierras , namely Tandiliaand Ventania.From a geological point ofview, Tandilia andVentania do not appear to be very closelyrelated (Rolleri 1975). Tandilia, southeasternof Buenos Aires province, constitutes theoldest nucleus in the country originated inthe Proterozoic age. Ventania is locatedsouthwest of the province and has a morerecent origin dated from the Paleozoic age.Gondwanaland and Pre-Gondwanalandevents seem to have played a major role in thebuilding ofthese mountain ranges.The biodiversity ofTandilia and Ventania isparticularly interesting because oftheir high


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number of endemic plantae and animal taxa(Table 1), and led to consider the ranges as orcgraphic islands (Kristensen & Frangi1995). When compared with the remainingvegetation of Buenos Aires, Tandilia andVentania have not only a characteristicpetrophylous flora, but they are the on lyregions in the province with endemic planttaxa (Parodi 1947). There have been numerousfloristic (Alboff 1895, Spegazzini 1896, 1901,Cabrera 1938, 1940, 1963) and ecological(Frangi 1973, Ponce 1982, 1986, Kristensen &

Frangi 1995) studies in the area. The differentcommunities were explained by the variationin the local climate and soil, which is causedby aspects of the topography of the ranges(Kristensen & Frangi 1995).Within the diverse plant families inhabitingTandilia and Ventania, Asteraceae have thesecond place (after Poaceae, the grass family)in number of genera and species represented

in the area (Parodi 1947). Approximately 30 %(112 species and infraspecific taxa) out of the356 Asteraceae known from the Buenos Ai-res province (Cabrera 1963, Zuloaga &Morrone 1999, Cabrera et al. 2000) occur inVentania and Tandilia. Furthermore, there havebeen found until the present four species ofAsteraceae endemic to Tandilia, three speciesand two varieties endemic to Ventania, and

one species endemic to both, Tandilia andVentania.The current understanding of the biotichistory ofTandilia and Ventania is based onseveral lines of evidence. Systematic studieshave documented much of the taxonomicdiversity and distribution patterns of ferns(Sota 1967, 1972, 1973, 1985) and scorpions(Maury 1973) and some hypotheses werepostulated to explain those distributions.

It would be interesting to seek the historicalexplanations that led to the high diversity andendemicity of Asteraceae in Tandilia andVentania and the relationships of the biotainhabiting these mountain ranges with other

Crisci el al .

areas. In order to achieve this goal, we applyherein two methods: a panbiogeographicapproach (compatibility track analysis) and

parsimony analysis ofendemicity (Morrone &Crisci 1990, 1995, Crisci et al. 2000) based ondistributional patterns exhibited by Asteraceae.

M A T ERI A L A ND M ET H ODS

Areas

Except for a few weedy introductions (e.g.,

F a cel is r etu sa, G a mo ch aeta su bfa lca ta i,theAsteraceae of this region shows no linksoutside of the Americas. For these reason,areas such as Palearctic, Afrotropical, Orien-tal, Australian, Cape subregions, were nottaken into consideration for the delimitationareas.On the basis of several over1apping taxadistribution and phytogeographical and

geological criteria the areas delimited here are(Fig.I):

(1) C entr al C hi le(CHI): Chile between 30and 37 south latitude (Cabrera & Willink 1973,Morrone et al. 1997).(2) C entr al and N or th A mer ica(CNA):Southern North America and Central America.(3) C haco (CHA): xerophyllous forest

extending from southern Bolivia, western Pa-raguay, and northern and central Argentina(Cabrera & Willink 1973).(4) M a huidas (MA): group of mountainranges in the political province ofLa Pampa,in central Argentina. They are 300-500 m high,and they are dated from the Cambric age (500m. y. ago)(Zambrano 1980).(5) N or ther n A ndes(NA): corresponds to theParamo and Puna provinces, and comprises thehighlands over 3000 m altitude from Venezuelato northwestern Argentina (Morrone 1994).(6 ) P am pa(PAM): grassy steppe ranging fromsouthern Brazil, Uruguay, and eastern Argen-tina, from 30 south latitude to 39 southlatitude (Cabrera & Willink 1973).

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Asteraccac frorn Tandilia and Ventania

Fig Areas of endemism considered in the historical biogeographic analysis: CHA = Chaco;CHI = Central Chile; MA = Mahuidas; NA = Northern Andes; CNA = Central and North America;PAM = Pampa; PAT = Patagonia; SA = Sierras Subandinas; SB = Southern Brazil; SPA =Sierras Pampeanas; TA = Tandilia; UR = Uruguay; VE = Ventania.

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Asteraceae from Tandilia and Ventania

Table 1. Taxa endemic to Tandilia and Ventania with the main references.

Taxa Tandilia Ventania References Tuxa Tandilia Vcntania References

I'LANTAE IRIDACEAE

ASTERACEAE C y p ct to h er b er t l (Lindl.) Herb X X Ravenna 1968B a cc ha ns r u fesc cn sSprcng val' X Cabrera 1963 subsp. l1'O/jjhucj. ,eli(1 Iauman)vcntctnculaCabrera RavennaB a c ch a n s t o nd t ten s sSpeg. X Cabrera 1963 Sisyrinchinrnjunccnmi:. i\1ey X X Ravenna 1968l I c r ct c n m b u r l c ar t i Sleumer X X Cabrera 1963 subsp. la inezi {Hicken ) RavennaH c r c t ci n m c h ac o en se(Zahn ) X Cabrera 1963Sleumet PLANTAGINACEAE

H c r o c i nm t a nd t te nscSleumer X Cabrera 1963 l/al/lago btsmarcktNiederl. X Pontiroli 1965,)'1.'11(,(,;0 tcncopcptusCabrera X Cabrera 1963 J/onfuj. ,() tundilcnsis(Pilg.) X Pontiroli \065S t a nd t tc n s sCabrera X Cabrera 1963 Rahn.'J'. twavcnsisCabrera X Cabrera 1963 POACEAES v cn a nc nsi sCabrera X Cabrera 1963 B r o m u s b o na r cn si sParodi J. X Cmara HStcvict satureiafot a(Lam.) Lam X Cabrera 1963 A.Cmara 1970vnr. \'CI1ICII/{.:nsisCabrera Fcstnca ventanico/a Speg X Cabrera 1970BRASSICACEAE l ptochact umca/J cSCL I1S Parodi X Tones 1970a ?o ;ppa vcntanensisCabrera X Iloelcke 1967Stsvmbr iurn ventancnscBoelcke X Boelcke 1967 t o a i r t df ol i a Ilauman X X Torres 1970b

Set a r a v a gi n a taSpreng val' X Nicora 1970BROMELlACEAE tcmdtlcnsisNicora

yc k a n st no ti i l o r aOtto DictrX Cabrera 1968 Slipa vcntuncoluCabrera X Cabrera &

Tones TO Tes 1970var. 10I1d/CIIS;S (Speg.) Cabrera

POLYGALACEAE

Titondso hergcri Mcz X Cabrera 1968 P u l y ga t u v c nt a nc n s sGrandona X Fnbris 1965CACTACEAE

(~Vl1ln()caZ)'c;1/11I platense Speg.)X Zuloaga SOLANACEAE

Morrone 1999 N i c r c m b er v i a a nd i l c nst s X X Cabrera 1965Britton Rose var. platense

(Kuntze) CabreraX Zuloaga ANIMALIA

Gyrnnocc t ivc um pla tense(Speg.)Morronc 1999 BOTIIRIURIDAE

Britton & Rose var, vcntantcotuB n th r u r n s v ov anMauri X Maury 1973

(Speg. ) R. KieslingIGUANIDAE

FAIl\CI:AE L o l a en n r s g r a c i ti sspp. X Vega&A d c sm t o t - on a r c n st sBurkart X Burkart 1967 13cllagambaA. pampeana Speg. X Burkart 1967 1990A. pselldogrisea Burk1I1 X Burkart 1967 Lio laemussp X Vega Asfragalus argef1l lIIs X X Burkart 1967 BellagambaManganaro 1990

L c h vr u s h o o kc r iG. Don f. X Burkart 1967 l rvs tidac tylus cc t shna cns s X Kristensen &

otbt t tora (Kuntze) Burkart Frangi 1995M i m osa r oca cLorentz X X Burkart 1967 I.EPTOTYPIILOPI DAENicdcrt. IJeJ7I()~VJ7/loJ7sunoa Orejas X X Vega 8:Vicia muntevidensisVogel X Burkart 1967 Miranda Bcllagambaoblonga Burkart 1990V sc t ot a Kunth Val X X Burkart 1967 M c l n no p h r yn sc u s st c l zn c r X Vega honaricnss Burkart subsp. numtcvidensisPhilippi Bcllagamba

1990

The compatibilitytrack method herein applied,developed by Craw (1988), is based on theconcept of distributional compatibility (Co-

nnor 1988, Craw 1989). In this method, indivi-dual tracks are treated as biogeograph ic

hypotheses of locality or distribution area

relationship. Two or more individual tracksare regarded as being compatible with each

other only if they are the same pairwise

comparison or if one track is a subset ofthe other

(i.e., tracks are either included within or

replicated by one another). It basically consistsof constructing a matrix (areas vs. tracks),

where each entry is 1 or O depending onwhether the track is present or absent in each

area, and using a compati-bility analysis

program to find the largest clique(s) of com-

patible tracks. The method involves finding a

simple form ofspanning tree linking localities

or distribution areas. This tree is constructed


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from the largest clique of compatibledistributions in a distributional compatibilityrnatrixand is identified as a generalized track.If more than one largest clique or severalcliques of considerable size are found, then ahypothesis of existence of several generalizedtracks linking the localities or distributionareas in more than one way can be considered.Alternatively, the intersection (i.e., thosetracks cornrnon to all the largest cliques) ofthe largest el iques can be identified as ageneralized track (Craw 1990). For more details

and other applications ofthis method see Craw(1988,1989),Morrone& Crisci(1990,1995),Crawet al. (1999) and Crisci et al. (2000). The analysisof the data matrix of 13 areas of endemisrn vs.distributional data of I 12 taxa (Table 3) wascarried out with SECANT 2.2 (Salisbury 1999).SECANT isa program for identifying all groups

Crisci el al .

of eladis-tically compatible characters. Wetreated our tracks as binary characters orderedwith absence as the ancestral state for each(= outgroup with all zeros).

Parsimony analysis of endernicity (PAE)

Parsimony analysis of endernicity or PAE(Rosen 1988, Rosen & Smith 1988) is anapproach of historical biogeography that ledto identify the distributional pattern oforganisms. It classifies localities, quadrats or

areas (which are analogous to taxa) by theirshared taxa (wh ich are analogous to characters)according to the most parsirnonious solution(Morrone & Crisci 1995, Crisci et al. 2000).The rnethod was originally proposed by Rosen(1988) using localities as study units.

Fig.2 A-B. Individual tracks. A, A c hy r o c i n e sa tu r ei o i desce), B acch ar i s cr i sp aC-); B, B .ul ic ina(e), C on yza b on ar i en si s ( -) .

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Astcraccae trom Tandilia and Vcntania

Table 2. Species of Asteraceae from Tandilia and Ventania and other areas of endemism consideredin the study. Taxa with the same areas of endemism have been indented. Areas ofendemism foreach taxa are indicated in Table 3.

1 A c hv r oc l i ne sa tn r ei o id cs(Lam.) De.2 . A cm el l a d ec um ben s(Sw.) R. K. Jansen var. aff inis (Hook.

Arn.) R. K. Jansen3 . A s/ er sq ua ma tu s(Spreng.) Hieran.4 . B ac ch ar i s a r tet ni si oi desI I ook. A rn.5. Ji art icutata (Lam.) Pers.6. H crispo Spreng.7. Ji gil l iesi i A. Gray8. J i spicat (Lam.) Bail .9 H sleJ/opl1yl/aAriza10 n t r i angula r i s Hauman11 n t r identcua Vahl var. suboppositc: (DC) Cabrera12. 1It r i tnera (Less.) DC.

13. B . n li ci naHook. Arn.14. lsidens laevis (L.) Britton, Sterns Poggenb. 5 C h up ta li a ex sc ap a(Pers.) Bakcr var. cxscapa16. C. i;;J1ola Burkart17. C. integcrr inta (Vel .) I3urkar11 8. ( I i l l fsel l l f ides(Vahl) Baker19. r : runcinata Kunth20. (' sinucna (Less.) Baker

2 B ac ch ar i s c or d if ol aOC.2 2. C o ny zu su mc ur en s s(Reitz.) E. Walkcr

2 3. C l w(j ui r oK u er i na ceaD. Don subsp. crinacea2 4. C on v: a h on ar i en si s(L.) Cronq. var. bonaricnsis25. e:f lo r ibunda Kunth26. (.'. tnonorchis (Griseb.) Cabrera

2 7. H n l oc het ln s b r asi l ien s s L.) Cabrera2 8. M i k an ia p cr i pl ocfo li aIlook. Arn.2 0. P an ph al ea h et er o ph vl l aLess.3 0. P nd nco ma h ier ac f li a(Poir.) Cass.

3 C o nv :a p r in ml i fo li a(Lam.) Cuatree. l.ourteig32. C. serrana CabreraXL E n pc uo r i ut n b n n if ol i r n nI Iook. Arn. var. Ivnni i fo l in rn34. lo'.bnpleur i fo l imnDC35 h macrocrpha ln tn Less.

3 6. C h ev reu li a sa r men to sa(Pcrs.) S. r. Blake3 7. E n pc no r iu in sq ua r rn lo su mHook. Arn,

3 8. B c tc ch ar i s g na pb c o i dc sSprcng.3 9. M i cr op si s sp astu da ta(Pers.) Cabrera

4 0. F ac el s r en rsa(Lam.) Seh. Bip. subsp, pcttulBeauv.41 J o .re/lisa (L arn.) Sch. B ip.subsp. re/lisa

42. G ui l la nt m eg apot am tca (Spreng.) Bak cr val'scabiosoidca (Arn. ex DC) Ilaker

4 3. F l av cr i a h cn un an iiDimitri Orfila4 4 . Stuckc r t i cl l a peregrina Beauv.

4 5, G ui ll ar di a nusgapotamica (Sprcng.) Baker var. tueg:po/{flJllca4 6. B er r ea g na ph al i o des(Less.) Bcauv.

4 7. G r nn och aeta a mer i ca na(M ill.) Wed d.48. j. argentina Cabrera

49. (; fatccua (Lam.) Cabrera50. i subfulcato (Cabrera) Cabrera

51. Gienochactof i tuginea (DC.) Cabrera52. U pl utensi s (Cabrera) Cabrera53. r ; stachvdifol o (Larn.) Cabrera54, G n up hc tl i um c ab r cr c u:S. E. Freire

55 H y po ch uc r ts p cn np as caCabrera56, G n a ph a li u n) c hei r o nt hi f ol n nnLam.57. c. gaudic/wudiomfm DC.58. lelfcopep tflll Cabrera

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5 9. G r in del ia a eg ia ltsCabrera6 0. U . b r ac hy st ep ha na Griseb.6 1 . U . / JI I / c hel l aDunal var. discoidea (Hook Am.) A.Bartoli

Tortosa62. c vcntancnsis A. Bartoli Tortosa6 3. G uti er r ezi a g il li esi iGriseb.64. H el en ium r adi otum(Lcss.) Seckt6 H i e r ac i um p al ezi en xi iZahn6 6. l I ya li s a rg en tenD. Don ex Hook. Arn, val'. la t isouama

Cabrera6 7. T ri ch octi ne sn uc a(D. Don) Cabrera

6 8. H y po ch aer i s p cti ol ar is(Ilook. Arn.) Griseb,69. Hystenonicojasionotdcs Willd.

7 0. C on y: a b nn or en s s(L.) Cronq. var. a l lg l f s l i /o l i a(Cabrera)l. Cabrera

7 1.I (vs/er io ll iea p in ifo ti a(Poir.) l3aker72. t Ivnochaers roscngurui i Cabrera var ptnnclf idu

(Speg.) Cabrera7 3. l .n ci li a a cn ti fu li (Poir.) Cass.7 4 M i c ro gy ncl a tr i fu rca ta(Less.) Grau7 5. N o ti ca st ru m m ar g in an tm(Kunth) Cuatrcc.7 6.I c r ezi a m ul ti fl or a(Humb. Bonpl.) Lcss. subsp. sonctnfotiu

(I3aker) Vuillcum.7 7. l i up at or i nt n c on nn er sa ni i(Cass.) l Iieron.7 8. l ly pu ch uer s g ri seh ach iiCabrera

7 9. P lu ch ea sa gi //a ll s(Larn.) Cabrera80. ,)'enecio grisebctchi i Bakcr val '. grisebachii

8 1. P od oc om a h r sr a(Hook. Am.) Bakcr82. C ony: a hl ukei(Cabrera) Cabrera83. E upator tnm snbhastut umlIook. Arn.84. Noticastrum diffusnn: (Pcrs.) Cabrera

8 5. Sen ec to a r ec ha va l et aeBakcr86. S bonaricnsis Hook. Arn.8 7. S. g r i sel w c hi iBaker val . subincanns Cabrera

88. S o steni Mattf.8 9. Sen ec to n to nt ev i den si s(Sprcng.) B akcr

90. M i cr op si s austr al tsCabrera9 1 Sen cc o p an rp ea nu sCabrera

92. S pulcher Ilook. Arn.93. l i u pa to r i i un t an ac ct if ol u nnG illics ex Ilook. Arn9 4, G r i nd el i a t ni ph th ah no uk :sDC.95 l lvpochaeris l'oriegu/o (Lam.) Baker9 6. So nu ner fel ti a sp in ul osa(Spreng.) I.css.

9 7 . Se nec t o su b nl a n ,O. Don ex I look . A rn. varocrcctus Ilook.&Am.

98. So/ira pterosperm Juss.) Less.9 9. St ev ia sa tu r ci i jo li a(L am .) L arn.var. pcuaganica lIieron.100. S. so tu r e to t (Lam.) Lam, varosa ture i i fo l ia101 Ihclesperma megajJotmJ/iclflll (Sprcng.) Kuntzc

1 02 . N o ti ca st ru m a r gen ti nen sc(Cabrera) Cuatrcc.1 03 . Ver n on ia ec hi oi desLcss.

1 04 . A c tn el l a d ec um h cn s(Sw.) R. K.Jansen val' decumbcns1 05 . l ac ch ar i sgel1;sli o/ia DC.1 06 . C r isca str i cta(Spreng.) Katinas1 07 . E np ato r in m t wc cd ia mn nHook. Arn.1 08 . H y po ch ac ri s m eg ap oto m caCabrera1 0 9 Sen ec to sello (Spreng.) DC1 1 St cv i a n n tl t i r i s t t aSprcng.

111 Ver n on i af lex uo sa Sims.\ 12 . Z cx tn cn iu b up hth ah ni fl om(Lorcntz) Ariza


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Craw ( 1988) and Cracraft (1991) presented avariation of th e method using are as ofendem ism as study un its to identify thehierarchical information contained in thegeographical distribution of organisms toestablish are a relationships. In this methoda data matrix of endemism areas by taxa isconstructed, and the presence of a taxon inan area is coded as 1 and its absence as O . Aparsimony algorithm is applied to the matrixto obtain a cladogram(s). PAE cladogramsrepresent nested sets of areas, in which ter-minal dichotomies represent two areasbetween which the most recent bioticinterchange has occurred (Morrone & Crisci1995). In this analysis the same are as anddata matrix (Table 3) used in thepanbiogeographic approach were taken intoconsideration. The analysis was carried outwith PAUP* version 4.0 beta 2 (Swofford1999), applying the branch-and-bound

option. A strict consensus tree wasconstructed based on the results generatedfrom PAUP*. As proposed by Rosen (1988),the cladogram was rooted with a hypotheticalarea coded all zeros.

RESULTS

Pan b iogeogr ap hy com p at ib il it y t r ack m et hod

Based on the data matrix (Table 3), thecombination ofthe individual tracks resultedin four largest cliques (generalized tracks) of30 individual tracks each one (Fig. 4 A-O).The intersection of 27 individual tracks(identifed with the numbers 3, 20, 21,23,25,26,27,28,29,30,36,37,38,81,82,83,84,87,88,103,104,105,106,107,108,109,110 inTable2)common to the four largest cliques is identifed

as the fifth clique or generalized track (Fig. 4E). In this way, six individual tracks (5, 40, 41,63, 64, and 98 in Table 2) are combined ingroups ofthree with the 27 individual tracksto form the four cliques of30 individual tracksmentioned above.Fivetreesconnecting the areas canbe constructed

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from thefve cliques,as follows (Fig.4):Clique A: (T A, UR), SB), PAM), VE),SPA), CHA), SA), NA), PAT, CHI), supportedby the 30 individual tracks 3, 5, 20, 21,23,25,26,27,28,29,30,36,37,38,40,41,81,82,83,84,87,88,103,104,105,106,107,108,109,110(Fig.4A).Clique B: (T A, UR), SB), PAM), VE),SPA), SA), CHA), NA), CHI, PAT), supportedby the 30 individual tracks 3,5,20,21,23,25,26,27,28,29,30,36,37,38,63,64,81,82,83,84,87,88,103,104,105,106,107,108,109,110(Fig. 4 B).Clique C: (T A, UR), SB), PAM), VE),SPA), CHA), SA), CHI) NA, PAT), supportedby the 30 individual tracks 3, 20, 21,23,25,26,27,28,29,30,36,37,38,40,41,81,82,83,84,87,88,98,103,104,105,106,107,108,109,110(Fig.4C).Clique O: (TA, UR), SB), PAM), VE),SPA), SA), CHA), CHI), NA, PAT), supported

by the 30 individual tracks 3, 20, 21,23,25,26,27,28,29,30,36,37,38,63,64,81,82,83,84,87,88,98,103,104,105,106,107,108,109,110(Fig.4 O).Clique E: (TA, UR), SB), PAM), VE),SPA), SA, CHA) NA, PAT, CHl), supportedby the 27 individual tracks 3, 20, 21,23,25,26,27,28,29,30,36,37,38,81,82,83,84,87,88,103, 104,105,106,107, 108, 109, 110(Fig. 4 E).

Mahuidas and Central and North America arethe only areas that appear in none of thesepattern of relationships in the fve generalizedtracks obtained.Oue to the fifth clique (E) represents theintersection of27 individual tracks commonto the four largest cliques, it was selected toshow the taxa supporting the areasrelationship (Fig. 4 E):

(1) (TA,UR). Based on Senec io gr i sebach i ivaro subincanus (individual track 87) and S.ostenii(track 88).(2) TA,UR), SB). Based on Eupator iumsquarrulosum(track 37), B ac ch ar i s g na ph a-l ioides (track 38), and M i cr o psi s sp at hu la ta(track 39).

29


10/21


Fig.3 A-F. Individual tracks. A, P o do co m a h i r su tac e Senecio arechavaletaeC-); B, Hierac iumpalezieuxii c e H y al i s a rgen teavar. la t isquama C-).e, H y st er i o ni ca p in if ol i ace), Pereziamul t i f lorasubsp. sonchifol iaC-); D, Sen ec io p u lc herce So m mer f el ti a sp i nu lo saC-); E,Sen ec io b on ar i en si sc e Sen ec io p am p ea nu sC- ; F, Ver n on ia ec hi oi desc e Zexmeniabuphta lmif loraC-).

30


11/21

- ---------- CHf---------- PAT

~-------- NA - -------- SA

....-------- CHA~----SPA

.------VE.-----PAM

.--- -SBTAUR

. --------- PATf----------- NA....------ ----- ---CH

~-------- SA - ---------- CHA

r------SPA.------VE

. ----PAM - ---SB

TAUR

INDIVIDUAL TRACKS

3 24 31Q5

X 20 21

98

< :>22 64 41 42

*81 82 83 84 26 27 28 29 30* 103 104 105 106107 108 109 110* 37 38 39

~ 87 88

Crisci el al .

.------------------ PAT~---------------CH

.-----------NA.--------- CHA

r---------- SA - ------- SPA

- ------ VE~--PAM

~---SBTAUR

~-----------PAT~------------NA~--------------CH

~--------- CHA....--------- SA

- ------- SPA.------VE

. -----PAM.--- -SB

TAUR

... .--------------- CH~-------PAT

~~----------- - NAr-------- CHA~------- SA

....------- SPA - ---- VE

.------ PAM~--SB

TAUR

Fig. 4. A-E. Cliques obtained in the compatibility track method after applying SECANT 2.2,

showed as cladograms. A-D, Four largest c\iques each supported by 30 individual tracks. E,Clique obtained by the intersection of27 individual tracks common to the four largest c\iques.The table shows the individual tracks or the sets of individual tracks, represented by syrnbols,that are superimposed onto the cladograms (see Table 2 for numbers corresponding to taxanames and Table 3 for individual tracks). CHA = Chaco; CHI = Central Chile; NA = NorthernAndes; PAM = Pampa; PAT= Patagonia; SA = Sierras Subandinas; SB = Southem Brazil; SPA= Sierras Pampeanas; TA = Tandilia; UR = Uruguay; VE = Ventania.

31


12/21

Asteraccac from TandiJia and Ventania

(3) (((TA,UR), SB), PAM). Based on Vernoniaechioides (track 103), A cm el la d ecu mb en s(track 104), Bacchar i s gen i st i fo l i a(track 105),C ri scia str icta (track 106), Eupator ium

tweedianum (track 107), Hypochaer ismegapotamica(track 108), Senecio sel loi(track109), and Stev ia mul t ia r i st at a(track 110).(4) ((((TA,UR), SB), PAM), VE). Based on

C onyza m onor chi s(track 26), Holochei lusbrasiliensis(track 27), M i k an i a p er i p l oc if ol i a(track 28), Panpha lea heterophyl l a(track 29),and P od oc om a h ier a ci fo li a(track 30).(5) (((((TA,UR), SB), PAM), VE), SPA). Based

on P o do co ma h ir su ta(track 81), Conyza b lakei(track 82), Eupa to r ium subhasta tum(track 83),and N o ti ca st r u m d i ff usu m(track 84).

Parsimony analysis of endemicity

The analysis ofthe data matrix (Table 3) with

PAUP* generated four area cladograms (Fig.5 A-D) with 263 steps, c.i.= 0.426, and r.i. =

0.663. The strict con sen sus ofthe four trees

(Fig. 5 E) shows the following area relation-ships: (Mahuidas (Patagonia (Chile (Sierras

Subandinas (Sierras Pampeanas (Ventania

(Pampa (Southern Brazil - Tandilia - Uru-

guay)))))))). The remaining areas: Chaco(CHA), Central and North America (CNA), and

northern Andes (NA) do not show resolved

relationships with the other areas in the

consensus.

Compar ison ofboth methods

The comparison of the results obtained

applying panbiogeography (compatibility

track rnethod) and parsimony analysis ofendernicity to the areas under study shows a

coincidence in the relationships of SierrasPampeanas (SPA), Ventania (VE), Pampa(PAM), Southern Brazil (SB), Tandilia (T A),

and Uruguay (UR).

The areas relationship PAM-SB-SPA-TA-UR-

VE conform a rnain distributional pattern

strongly supported by the two approaches

32

applied in this study, and represented in Fig.

6 as a generalized track. Both methods showthat Tandilia is closer to Uruguay, southern

Brazil, and Pampa than to Ventania. On the

other hand, PAE and panbiogeographyresulted in different relationships of the

remaining other areas with the m a in

distributional pattern. PAE shows that Sie-

rras Subandinas (SA) (Fig. 5 A-E) are related

to this pattern, whereas panbiogeo-graphy

shows that Sierras Subandinas can be directly

related to it (Fig. 4 B, D) or cannot be directlyrelated (Fig. 4 A, C); PAE shows Chaco (CHA)

as not closely related to the pattern (Fig. 5 A-

E), whereas panbiogeography shows CHA

connected to it (Fig. 4 A-E); PAE shows

Mahuidas (MA) and Central and North

Arnerica (CNA) not closely related to the

pattern (Fig. 5 A-E), and panbiogeography

shows these last two areas very distantly

related to it (for this reason they are notindicated in the results).

DI SCUSSION

The main distributional pattern connecting

Tandilia and Ventania with southern Brazil,

Pampa, Uruguay, and Sierras Pampeanasobtained through the cornpatibility track

method and PAE partially agree with others

authors' hypotheses.

A biotic rnigratorial route frorn the Andes to

Brazil, through Sierras Pampeanas, sierras of

Buenos Aires, sierras ofUruguay, and Planalto

and sierras of Brazil was postulated rnany yearsago by Hicken (1918-1919) and Brade (1942).

Frenguelli (1950) gavethe name of periparnpasic

orogenic are ( ar c o p er i p am p si co ser r a n o)tothe biotic corridor constituted by Sierras

Pampeanas, Mahuidas, and Tandilia. Ringuelet(1956) postulated that the province ofBuenosAires constitutes a cornposite area due to the

presence oftwo types offauna: one related tothe Subtropical domain (Guaianian-Brazilian),

and the other related to the Austral region

(Andean-Patagonic, or Chilean-Patagonic).


13/21

Crisci el al .

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Q g c-,. . . . . . o o o o . . . . o . . . . . . . . . . . . o o . . . . . . .. . . o . . . . . .

4 ~o :.... ::l o o o o o o o . . . . . . . . . . . . o . . . . . . . . . . . . o . . . . . . . . . .VJ bl bl) O'. o o o o o o

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8 8. . . . . . . . . . . . . . . . . . . . . . . .

f- ~. . . . . . . . . . . . . . .. . . . . . . . . . . . .

>,~. . . . . . o o o o o o . . . . . . o o . . . . . . . . . . .. . . . . . . . . . .

O ~ o o . . . . . . o o o o . . . . . . o . . . . . . . . . . . . . . . . . . . . . . . . . . . . o O'. o . . . . . . o o o o . . . . . . o . . . . . . . . . . . . . . . . . . . . . . . . . . orf 2 8 8 8 8 8 8

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. . .. . . . . . . . . . . VJ .- . . . . . . . . .. . . . . . . . . . .

Vlv ;

O o o o o o o . . . . . . o o o o . . . . . . . . . . . . . . . . . . - -o -o 00 'ro 8 8 8 8 8

. . . . . . o 8. . . . . . o . . . . . . 8 o

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t f- f 2 8 8 8 8 8 8. . . . . .

8 8 8 o 8. . . . . . . . . . . . . . . . . . . . o . . . . . .-o o

Vl t . . O o o o o o o . . . . o o o . . . . . . o o . . . .

e z; oro 8 8 8 8 8 8. . . . . .

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~ ~ 1 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .VJ

e 8 8. . . . . .

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8. . . . . . o 8

. . . .~ o o o o o o . . . . . . . . . .

Vl Q. . . . . . o o . . . . o o o . . . . . . . . . . . . o . . . . . . . . . . . . . . . . . .o

>. o. :z 6 o o

. . . . . . o o o o . . . . . . . . . . . . o . . . . . . o . . . . o . . . . . .O'. o o o o o o . . . . o o o o . . . . o o~ f 2

8o . . . . . .

8. . . . . .

8o . . . . . . . . . . . . . . . . . . . . . . . .

8 . . . . . . . .

~ Vl e, . . . . . . . . . .

o . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .

~ c e VJ O O o o . . . . . . o . . . . . . o o o o . . . . . . . . . . . . . . . . . . . . . . . . . . . .~ 8 8. . . . . .

8 8 8. . . . . . . . . . . .

8 8. . . . . .

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. . . .8

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11o o o o o o o . . . . . . o . . . . . . . . . . . . . . o . . . . . . . . . . . .

: .o 11 N 0-, o o o o o o . . . . . . o . . . . . . . . . . . . . . o . . . . . . . . . . . .< f ? 8 8 . . . . . . o . . . . . . o . . . . . . o o o . . . . . . . . . . . . . . . . . . . . . ..~< ~ . . . . . . . . . . o . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .e, O o o . . . . . . o o o . . . . . . . . . . . . o . . . . . . . . . . . . o . . . . . . . .; : : - ;VJ D 'ro 'ro 8 8

. . . . . .8 8 8

. . . . . . . . . . . . o . . . . . . . . . . . . 8 . . . . . . . . . .

r o 1 . . . . . . . . . . . . . . . . . . . . . . . .o . . . . . . o . . . . f- 8 o

. . . . . .8 8 8

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6 f ? 8 8 8 8 8 8 8 8 8. . . . . .

8 . . . . . . . .

8- e o . . . . . . . . . . . . . . . .t .8 < . . .-5 O o o o . . . . o o . . . . . . o . . . . . . . . . . . . . . . . . . . . . . o . . . . . .

~ 1

8 8 8 . . . .

8 8. . . . . .

8. . . . . . . . . . . . . . . . . . . . . . o . . . .~ . ..c

8 1 o. . . . . . . . . . . . . . . . . .

o . . . . . . . . . . . . . .

VJ t 8 8 . . . . 8 . . . . 8 . . . . 8 . . . . . . . . . . . . . . . . . . . . . . . . . . . .~ g -o o o o . . . . . . . . . . . . o . . . . . . . . . . . . . .u o e . . . . . . o . . . . . . . . . . o o . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .~ Z VJ ob -o 11 o. o o o o o o o . . . . . . o o . . . . . . o . . . . . . . . . . . . . . . . . .0-, o o o o o o . . . . . . o o . . . . o . . . . . . . . . . . . . . . .C ; ; e ro Vl f ? 8 8 8 8 8 8. . . . . .

8 8. . . . . .

8 . . . . . . . . . . . . . .

Q::l ~ VJ . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .~ - O o o o o o o . . . . o o . . . . . . o . . . . . . . . . . . . . .~ o M 8 o

. . . . . .8

. . . . . . . . . . . . . . . . . . . . . .. . . . . . . . . . . . . . . . . . . . . . . . . . . . 2 : ~

VJ u 1 . . . . . . . . . . . . o . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .~ 8 8. . . . . .

8 8 8 o . . . . . . . . o . . . . . . o o . . . . . .-o .::: C J o

. . . . . . o . . . . . . . . . . . .. . . . . . . . . . . . . . . . . ..::: (jj x

. . . . . . o . . . . o o o o . . . . . . o . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .U -o Evi e o o . . . . . . o o o o . . . . . . o . . . . . . . . . . . . . . . . . . . . . . . .. . . . . .

1 1 ~ 4 O'. o . . . . . . o . . . . . . o . . . . . . . . . . o o . . . . . . . . . . . . . . . . . . . . o> D o f 2 8 8 8 8 8 o . . . .

8 o. . . . . . o o . . . . . . . . . . . .

Vl ::l :.... . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . o . . . . . .~ O o o o o o o . . . . . . o o o o . . . . . . . . . . . . oQ Z VJ

Q 1 r

8 O O O 8. . . . . . . . . . . .

8 O. . . . . . . . . . . . . . . . . .. . . . . . . . .

VJ D

1 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .. . . . . . O~ U ~ E 8

. . . . . .8 8

. . . . . . . . . . . . . . . . . . . . . . . . . . . . . .O . . . . 8 O . . . . . . . O O . . . . . . . . . . . . O . . . . . . . . . . . . O . . . . . . . . . . . .

X ~ -;::l . . . . . .

O. . . . . .

o O O O. . . . . .

O. . . . . . . . . . . .

O . . . . . . . . . . . . . .

.~ z. . c : VJ O O O O O O O O . . . . . . O O . . . . . . O O . . . . . .~ U O'. O O O O O O . . . . . . . .O O . . . . . . O . . . . . . . . . .E 11 f ? 8 . . . . . . O 8 8 8 . . . . . . O O . . . . . . . . . . . . . . . . . . . .OC ij

.. . . . . . . . . . . . . . . . . . . . . . . . . . . O . . . . . . O O . . . .~ < Q . . . . . . O O O . . . . . . O O O . . . . . . . . . . O O . . . . . . O . . . . . . . . . . . . i:j b VJ

8. . . . . .

8 8 O . . . . . . . .

8. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .

U 1 O . . . . . . O . . . . . . . . . . O . . . . . . . . . . . . . . . . . . . . . . . .O e .~

e 8 . . . . . . . . . .

8 o . . . . . . . . . . . . . . . . . . . . . .. . . . . . . . . . . . . . . . . . . . . .

Q Q o o . . . . . . o o o . . . . o . . . . . . . .o o. . .-iU e Vl

. . . . . . o o o o o . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .Q

Q o :....

:c bl o. 5 ~~ E -


14/21

Asteraceae frorn Tandilia and Vcntania

Sota (1967,1972, 1973, 1985) in his analyses ofthe ferns growing in Tandilia and Ventania,concluded that the pteridological flora ofthesemountain ranges have four origins: Austral-

Antarctic, Andean-Pampean, Austrobrazilian,and Endemic. According to his studies the Bue-nos Aires' mountain ranges constitute anintermediate orofilous station in the migratorialroutes between the Andean-Pampean or theAustral-Antarctic flora, and the Austrobrazilianflora. Maury (1973) established, on the basis ofthe scorpions inhabiting Tandilia and Ventania,nearly the same kind ofrelationships with other

areas as showed by Sota: with the Subtropicaldomain, Central domain (Sierras Pampeanas),and Patagonian and Central Chilean domains.The studies based on ferns and scorpionsshow a link between Tandilia and Ventaniawith the Austral region, a kind ofrelationshipnot found here for Asteraceae. Althoughsorne Asteraceae inhabiting Tandilia andVentania occur in Patagonia, Central Chile andMahuidas, they do not constitute or areincluded in a generalized track. It is possiblethat different taxonomic groups such asscorpions, ferns and Asteraceae haveundergone different historical processes thatled to their actual distributional pattern.The relationships among the areas ofthe maindistributional pattern found in this studyshow that Tandilia is closer to Uruguay andsouthern Brazil than to Ventania. AlthoughTandilia and Ventania are geographically closeto each other, the geological evidencesuggests that both mountain ranges resultedfrom independent geological processes atdifferent geological times. According to Ra-mos (1989), Ventania was the result of thecollision of a Patagonian block that driftedfrom the south and accreted to Gondwanaland(ca. 360-440 m.y. ago). Tandilia, on the otherhand, was a part of the Ro de La Plata craton(together with northeast Argentina, east Pa-raguay, Uruguay and southeast Brazil) beforethe end ofthe Precambrian (530-570 m.y. ago).Due to the family Asteraceae is believed to

34

have originated in South America in theTertiary, during the Oligocene (38 m.y.ago)(Stuessy et al. 1996), two geological eventsmay have played a significant role in the

Asteraceae evolution and distribution inTandilia and Ventania and related areas, theuplift of the Andes and the Pleistoceneglaciations.During late Cenozoic (10-5 m.y. ago)the upliftof the Andes together with the developmentof cold offshore currents have beenassociated with a diversification of habitatsand evolutionary opportunities for the biota

(Darlington 1968, Axelrod et al. 1991)especially for the Asteraceae (Funk et al. 1995,Stuessy et al. 1996).From the Miocene (25-5 m.y. ago) to Pliocene(5-1 m.y. ago) the climate was drying in southemSouth America caused by the slowly raising ofthe Andean chain and the cold Humboldtcurrent that intensified aridity. The last phaseofthe Andean orogeny in the upper Pliocenecaused additional uplift ofSierras Pampeanasand Sierras Subandinas (Taylor 1991), that hadstarted their uplift with the climax of theAndean cordillera raising during the Tertiary.Finally, during the Pliocene and Pleistocene inthe Quaternary period, glaciations caused dryand wet cycles that caused fragmentation anddifferentiation of the populations (SimpsonVuilleumier 1971, Simpson 1975). Semiaridvegetation extended into the continent, e.g., tothe south Patagonia, north of Atacaman desert,Chaco, and Monte, in the core of SouthArnerica, and the cerrado and caatingas incentral and northeastern Brazil, respectively(Prado & Gibbs 1993). This may resulted in aneventual separation of populations in moreelevated regions, restricted now to rocky andloose soils. The main distributional patternfound in this study connecting the mountainranges ofsouthern Brazil, Uruguay, Tandilia,Ventania and Sierras Pampeanas, and theAsteraceae endemicity found in Tandilia andVentania could be explained as a consequenceof these events.


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OUTCNANA

CHACHIPAMS8UR

TAVESPASA

DATMA

OUTCHACHI

PAMS8TA

UR

VESPASAPAT

E MACNANA

OUTCNANACHACHIPAMURS8TAVE

SPASA

A PATMA

Crisci el al .

r--r- ~

lr=-e- . -- --

--

OUT

CHACHIPAM

URS8TAVE

SPASAPATMANACNA

. OUT CHA

. CHI PAM

S8

URTA

L-.-----VE

L SPAL SA

L PATL . MA

L NAL-. . CNA

Fig. 5. A-E, Cladograms obtained after applying parsimony analysis of endemicity after applyingPAUP*. A-O, Four area cladograms. E, Strict consensus ofthe four cladograms. CHA = Chaco;CHI = Central Chile; MA = Mahuidas; NA =Northem Andes; CNA = Central and North America;PAM = Pampa; PAT = Patagonia; SA = Sierras Subandinas; SS = Southem Brazil; SPA =Sierras Pampeanas; TA = Tandilia; UR = Uruguay; VE = Ventania.

35


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- -1 \

- J _ o

\~re \

\

\\- -\---5PA----

\ PAM

\--\\\\

Fig. 6. Distributional pattern of relationships among areas based on Asteraceae taxa, andrepresented as a generalized track. PAM = Pampa; SS = Southern Brazil; SPA = SierrasPampeanas; TA = Tandilia; UR = Uruguay; VE = Ventania.

36


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In surnm ary, it is probable that the evolutionof Asteraceae in Tandilia and Ventania wasmainly affected by Tertiary and Quaternarygeologic events. A s a result, this biota show sa high level of diversity, many endem ic taxa,and closer relationsh ips w ith south ernBrazil,Uruguay, Pampa, and Sierras Pampeanasratherthan with other arcas ofAmrica.Thediscon-tinuous pattern, i.e., occurring inelevated areas surrounding a corearea ofCha-co and Montevegetation, is what one wouldexpect under vicariance due to dryconditions,although dis-persal is taken also as a comm onand impor-tant distributional event. Furthercorrelations w ith analyses of other plantandanimal taxa will determine the generality ofthe pattern shown by Asteraceae.

ACKNOWLEDGEMENTS

We thank Paul Berry, John Grehan, E dgardo

O rtiz Jaureguizar, M ario Teruggi, RosendoPascual, and Elas de la Sota for helpfulcomments on the manuscript. We thank JorgeFrangi for checking the Iist of species. Specialthanks to Benjamin A . Salisbury for his gene-rous assistance with his program SE CANT.This study was supported by grant5776-96of the National Geographic Society. Support11'0111heConsejoNacional de Investigaciones Cien-

tf icas y Tcnicas (CONICET), Argentina, towhichthe authors belong, is gra tefu lly acknowledged.

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