Caldasia 23(1): 21-41 HISTORICAL BIOGEOGRAPHY OF THE FROM TANDILIA AND VENTANIA MOUNTAIN RANGES (BUENOS AIRES, )

A nuestra amiga Pilar

JORGE CRISCI- V. Departamento Científico de Plantas Vasculares, Museo de La Plata, Paseo del Bosque s.n., 1900. La Plata, Argentina. [email protected]

SUSANA FREIRE-E. Departamento Científico de Plantas Vasculares, Museo de La Plata, Paseo del Bosque s.n., 1900. La Plata, Argentina.

GISELA SANCHO Departamento Científico de Plantas Vasculares, Museo de La Plata, Paseo del Bosque s.n., 1900. La Plata, Argentina.

LILlANA KATlNAS Departamento Científico de Plantas Vasculares, Museo de La Plata, Paseo del Bosque s.n., 1900. La Plata, Argentina.

ABSTRACT Tandilia and Ventania are the only systems ofmountain ranges situated in a grassy steppe or "pampas" in the political province ofBuenos Aires in Argentina. Tandilia and Ventania have a high taxa diversity and endemicity. A historical biogeographic analysis was carried out on the basis of distributional patterns of species and infraspecific taxa of Asteraceae inhabiting Tandilia and Ventania in orderto establish the relationships ofthese mountain ranges with other areas. Two methods were applied for the analysis: panbiogeography using the compatibility track method and parsimony analysis of endemicity (PAE). Thirteen areas were delimited for the study: Southern North America and Central América, Southern , , 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 not included). Both methods connect southern Brazil, Uruguay, Pampa, Tandilia, Ventania, and Sierras Pampeanas, showing that the Asteraceae biota ofTandilia and Ventania have closer relationships with the biota ofthese are as rather than with that ofSierras Subandinas, North Andean, Chaco, Patagonia, Mahuidas, and Central Chile. The Asteraceae ofCentral and North America appear less related to Tandilia and Ventania in regard ofthe rest ofthe areas. The evolution of Asteraceae in Tandilia and Ventania and related areas is hypothetized to have been affected mainly by Tertiary and Asteraccae from Tandilia and Ventania Quaternary geologic events. The discontinuous pattern found, occurring mostly in elevated areas, is explained principally by vicariance under dry conditions.

Key words. Asteraceae; biogeography; Tandilia; Ventania.

RESUMEN Las sierras septentrionales, o de Tandilia y las australes, o de Ventania, caracteriza- das por su gran diversidad orgánica y su alto número de endemismos, están ubicadas en la planicie bonaerense de pastizales denominada "pampa". Con el objeto de esta- blecer las relaciones entre Tandilia y Ventania con otras áreas, se analizaron las dis- tribuciones de los taxones específicos e infraespecíficos de Asteraceae que habitan estas sierras. Se delimitaron trece áreas: sur de América del Norte y América 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 endémicos, adventicios y naturalizados no fueron incluidos en el análisis). Las distribuciones de estos taxones fueron analizadas utilizando dos métodos: panbiogeografía (compatibilidad de trazos) y análisis de simplicidad de endem ismos. Ambos métodos conectan sur de Brasil, Uruguay, Pampa, Tandilia, Ventania y Sie- rras Pampeanas. Las Asteraceae de Tandilia y Ventania estarían más relacionadas con estas áreas que con la de las Sierras Subandinas, Andes del Norte, Chaco, Patagonia, Mahuidas y Chile Central. El área de América del Norte y América Central aparece menos relacionada a Tandi1ia y Ventania en relación a las otras áreas. La evolución de las Asteraceae de Tandilia y Ventania y las áreas relacionadas ha sido probablemente afectada por los eventos geológicos de los períodos Terciario y Cuaternario. La dis- continuidad actual en el patrón de distribución de las Asteraceae, restringidas en general a las zonas elevadas, es explicada principalmente por eventos vicariantes debidos a condiciones áridas.

Palabras clave. Asteraceae; biogeografía; Tandilia; Ventania.

INTRODUCfION mountains ranges or"sierras", namely Tandilia and Ventania. Since Darwin'sjourney in Argentina in 1835 From a geological point ofview, Tandilia and the geology ofTandilia and Ventania and their Ventania do not appear to be very closely inhabiting biota have attracted several related (Rolleri 1975). Tandilia, southeastern generations of geologists and biologists (e.g., of Buenos Aires province, constitutes the Darwin 1846, Alboff 1895, Hauthal190 1, Ca- oldest nucleus in the country originated in brera 1938, Sota 1967, Teruggi & Kilmurray the Proterozoic age. Ventania is located 1975). The political province ofBuenos Aires, southwest of the province and has a more situated in central eastern Argentina, is covered recent origin dated from the Paleozoic age. in most of its surface by an herbaceous, grassy Gondwanaland and Pre-Gondwanaland steppe, ca\led "pampas". The pampas are a plain events seem to have played a major role in the situated at the sea level, or even below the sea building ofthese mountain ranges. level (like the big depression ofthe Salado river), The biodiversity ofTandilia and Ventania is with the exception of two major systems of particularly interesting because oftheir high

22 Crisci el al. number of endemic plantae and animal taxa areas. In order to achieve this goal, we apply (Table 1), and led to consider the ranges as herein two methods: a panbiogeographic "orcgraphic islands" (Kristensen & Frangi approach (compatibility track analysis) and 1995). When compared with the remaining parsimony analysis ofendemicity (Morrone & vegetation of Buenos Aires, Tandilia and Crisci 1990, 1995, Crisci et al. 2000) based on Ventania have not only a characteristic distributional patterns exhibited by Asteraceae. petrophylous flora, but they are the on ly regions in the province with endemic MATERIAL AND METHODS taxa (Parodi 1947). There have been numerous floristic (Alboff 1895, Spegazzini 1896, 1901, Areas Cabrera 1938, 1940, 1963) and ecological (Frangi 1973, Ponce 1982, 1986, Kristensen & Except for a few weedy introductions (e.g., Frangi 1995) studies in the area. The different Facelis retusa, Gamochaeta subfalcatai, the communities were explained by the variation Asteraceae of this region shows no links in the local climate and soil, which is caused outside of the Americas. For these reason, by aspects of the topography of the ranges areas such as Palearctic, Afrotropical, Orien- (Kristensen & Frangi 1995). tal, Australian, Cape subregions, were not Within the diverse plant families inhabiting taken into consideration for the delimitation Tandilia and Ventania, Asteraceae have the areas. second place (after Poaceae, the grass family) On the basis of several over1apping taxa in number of genera and species represented distribution and phytogeographical and in the area (Parodi 1947). Approximately 30 % geological criteria the areas delimited here are (112 species and infraspecific taxa) out of the (Fig.I): 356 Asteraceae known from the Buenos Ai- res province (Cabrera 1963, Zuloaga & (1) Central Chile (CHI): Chile between 30° Morrone 1999, Cabrera et al. 2000) occur in and 37° south latitude (Cabrera & Willink 1973, Ventania and Tandilia. Furthermore, there have Morrone et al. 1997). been found until the present four species of (2) Central and North America (CNA): Asteraceae endemic to Tandilia, three species Southern North America and Central America. and two varieties endemic to Ventania, and (3) Chaco (CHA): xerophyllous forest one species endemic to both, Tandilia and extending from southern Bolivia, western Pa- Ventania. raguay, and northern and central Argentina The current understanding of the biotic (Cabrera & Willink 1973). history ofTandilia and Ventania is based on (4) Mahuidas (MA): group of mountain several lines of evidence. Systematic studies ranges in the political province ofLa Pampa, have documented much of the taxonomic in central Argentina. They are 300-500 m high, diversity and distribution patterns of ferns and they are dated from the Cambric age (500 (Sota 1967, 1972, 1973, 1985) and scorpions m. y. ago)(Zambrano 1980). (Maury 1973) and some hypotheses were (5) Northern Andes (NA): corresponds to the postulated to explain those distributions. Paramo and Puna provinces, and comprises the highlands over 3000 m altitude from Venezuela It would be interesting to seek the historical to northwestern Argentina (Morrone 1994). explanations that led to the high diversity and (6) Pampa (PAM): grassy steppe ranging from endemicity of Asteraceae in Tandilia and southern Brazil, Uruguay, and eastern Argen- Ventania and the relationships of the biota tina, from 30° south latitude to 39° south inhabiting these mountain ranges with other latitude (Cabrera & Willink 1973).

23 Asteraccac frorn Tandilia and Ventania

Fig. 1. 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.

24 Crisci el al. (7) Patagonia (PA T): semidesert east of the Taxa analyzed southern Andes to the Atlantic coast in Ar- gentina, approximately between 35° and 55° The units of study (Table 2) were species and south latitude (Cabrera & Willink 1973). infraspecific taxa of Asteraceae inhabiting (8) Sierras Pampeanas (SPA): group of moun- Tandilia and Ventania. The distributional data tain ranges in central Argentina, extending in were taken from the specimens' labels ofthe the political provinces of Córdoba, Santiago herbarium ofDepartamento Científico de Plan- del Estero, eastern Catamarca, and San Luis. tas Vasculares (Herbarium LP), Museo de La They are 3500 m high and their Precambric Plata, Argentina and from the literature. The rocks were uplifted during Tertiary and number of specimens analyzed are ca. 2500. Quaternary times (Gordillo & Lencinas 1980, Each specimen was correctly identified prior Michaut 1980). to the extraction of data. Checklists (Troiani (9) Sierras Subandinas (SA): a group of & Steibel 1999, Zuloaga & Morrone 1999, Ca- mountain ranges in northwestern Argentina, brera et al. 2000, Peter 2000) and floristic extending in the political provinces of Salta treatments of the province of Buenos Aires and Jujuy. They are 3000-5000 m high; the (Cabrera 1963, Cabrera & Zardini 1978), and most ancient rocks be long to the Precambric revisionary studies were taken al so into age (Mingramm et al. 1980) and were uplifted consideration. For nomenclature purposes we in the Tertiary and Quaternary periods. followed Zuloaga & Morrone (1999). Adven- (10) Southern Brazil (SB): subtropical forest ticious (e.g., Anthemis cotula, Matricaria that corresponds to the Paran ense province recutita, Sonchus asper) , cultivated (e.g., ofCabrera & Willink (1973) in southern Brazil Ci-chorium intybus), naturalized (e.g., (ranging west of Serra do Mar to central Rio Artemisia velortorum, Sonchus oleraceous) Grande do Sul), northeastern Argentina, and or cosmo-politan (e.g., Solidago eh ilensis, eastern Paraguay. Taraxacum officinale¡ species were not taken (1 1) Tandilia (TA): or "Sierras Septentriona- into consi-deration. les", a group ofmountain ranges east ofBue- nos Aires province. They are 500 m high, and Panbiogeography (compatibility track represent the most ancient mountains in Ar- method) gentina dated from 2200 m. y. ago, in the Proterozoic (Teruggi & Kilmurray 1975, 1980). Croizat postulated that earth and life evolve (12) Uruguay (UR): prairie with endemic taxa, together (Croizat 1958), meaning that geo- and low mountain ranges. It corresponds to graphic barriers evolve together with biotas. the eastern Uruguayan province (Castellanos The panbiogeographic approach (Croizat 1958, & Pérez Moreau 1944,Chebataroff 1960), situa- 1981) consists of plotting distributions of ted in the south of Brazil (southern Rio Gran- organisms on maps, connecting their separate de do Sul) and Uruguay. distributional areas together with lines called (13) Venlania (VE): or "Sierras Australes" a individual tracks. As an example, some ofthe group of moutain ranges southwest of Bue- 112 individual tracks produced in this study of nos Aires province. They are 900-1000 m high, Asteraceae inhabiting Tandilia and/or Ventania and the geological age corresponds to 490 m. and other areas are illustrated (Figs. 2-3). When y. ago, in the Paleozoic (Llambías & Prozzi individual tracks coincide, the resulting 1972, Harrington 1980). summary lines are considered gene-ralized tracks, which indicate the pre-existence of an- cestral biotas, that become fragmented by tectonic and/or climatic changes.

25 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 Cypctto herbertlí (Lindl.) Herb X X Ravenna 1968 Bacchans rufesccns Sprcng val' X Cabrera 1963 subsp. l1'O/jjhucj.!,eli (1 Iauman) vcntctnícula Cabrera Ravenna Bacchans tondttens¡s Speg. X Cabrera 1963 Sisyrinchinrnjunccnm i:. i\1ey X X Ravenna 1968 l Iícrctcínm burlcartií Sleumer X X Cabrera 1963 subsp. lainezií {Hicken ) Ravenna Hícrctcinm chacoense (Zahn ) X Cabrera 1963 Sleumet PLANTAGINACEAE Hícrocinm tandttensc Sleumer X Cabrera 1963 ¡l/al/lago btsmarckt¡ Niederl. X Pontiroli 1965

,)'1.'11(,(,;0 tcncopcptus Cabrera X Cabrera 1963 !J/onfuj.!,() tundilcnsis (Pilg.) X Pontiroli \065 S tandttcnsís Cabrera X Cabrera 1963 Rahn .'J'. twavcnsis Cabrera X Cabrera 1963 POACEAE S vcníancnsis Cabrera X Cabrera 1963 Bromus bonarícnsis Parodi & J. X Cámara H Stcvict satureiafotía (Lam.) Lam X Cabrera 1963 A.Cámara 1970 vnr. \'CI1ICII/{.:nsisCabrera Fcstnca ventanico/a Speg X Cabrera 1970

BRASSICACEAE l'íptochactíum ca/J'cSCL'I1S Parodi X Tones 1970a !?o";ppa vcntanensis Cabrera X Iloelcke 1967 Stsvmbriurn ventancnsc Boelcke X Boelcke 1967 t'oa irtdfolia Ilauman X X Torres 1970b Setar¡a vaginata Spreng val' X Nicora 1970 BROMELlACEAE tcmdtlcnsis Nicora X Cabrera 1968 Slipa vcntunícolu Cabrera & X Cabrera & ñyck¡a nstnotiilora Otto & Dictr Tones TO!Tes 1970 var. 10I1dí/CIIS;S (Speg.) Cabrera POLYGALACEAE Titíondsío hergcri Mcz X Cabrera 1968 Pulygatu vcntancns¡s Grandona X Fnbris 1965 CACTACEAE X Zuloaga & SOLANACEAE (~Vl1ln()caZ)'c;1/11I platense (Speg.) Morrone 1999 Nicrcmbervia íandilcnsts X X Cabrera 1965 Britton & Rose var. platense (Kuntze) Cabrera X Zuloaga & ANIMALIA Gyrnnocctivcíum platense (Speg.) Morronc 1999 BOTIIRIURIDAE Britton & Rose var, vcntantcotu Bnthríurns vovan Mauri X Maury 1973 (Speg.) R. Kiesling IGUANIDAE FAIl¡\CI:AE Líolaennrs gracitis spp. X Vega& Adcsmto t-onarícnsts Burkart X Burkart 1967 13cllagamba A. pampeana Speg. X Burkart 1967 1990 A. pselldogrisea Burk¡1I1 X Burkart 1967 Liolaemus sp X Vega & Asfragalus argef1lÍ!lIIs X X Burkart 1967 Bellagamba Manganaro 1990 Lcühvrus hookcri G. Don f. X Burkart 1967 l'rvstidactylus cctshnaíícnsís X Kristensen & otbtttora (Kuntze) Burkart Frangi 1995 Mimosa rocac Lorentz & X X Burkart 1967 I.EPTOTYPIILOPI DAE Nicdcrt. IJeJ7I()~VJ7/¡loJ7smunoaí Orejas X X Vega 8: Vicia muntevidensis Vogel X Burkart 1967 Miranda Bcllagamba oblonga Burkart 1990 V scütotía Kunth Val X X Burkart 1967 Mclnnophryníscus stclzncrí X Vega & honaricnsís Burkart subsp. numtcvidensis Philippi Bcllagamba 1990

The compatibilitytrack method herein applied, (i.e., tracks are either included within or developed by Craw (1988), is based on the replicated by one another). It basically consists concept of distributional compatibility (Co- of constructing a matrix (areas vs. tracks), nnor 1988, Craw 1989). In this method, indivi- where each entry is 1 or O depending on dual tracks are treated as biogeograph ic whether the track is present or absent in each hypotheses of locality or distribution area area, and using a compati-bility analysis relationship. Two or more individual tracks program to find the largest clique(s) of com- are regarded as being compatible with each patible tracks. The method involves finding a other only if they are the same pairwise simple form ofspanning tree linking localities comparison or if one track is a subset ofthe other or distribution areas. This tree is constructed

26 Crisci el al. from the largest clique of compatible of eladis-tically compatible characters. We distributions in a distributional compatibility treated our tracks as binary characters ordered rnatrix and is identified as a generalized track. with absence as the "ancestral" state for each If more than one largest clique or several (= outgroup with all zeros). cliques of considerable size are found, then a hypothesis of existence of several generalized Parsimony analysis of endernicity (PAE) tracks linking the localities or distribution areas in more than one way can be considered. Parsimony analysis of endernicity or PAE Alternatively, the intersection (i.e., those (Rosen 1988, Rosen & Smith 1988) is an tracks cornrnon to all the largest cliques) of approach of historical biogeography that led the largest el iques can be identified as a to identify the distributional pattern of generalized track (Craw 1990). For more details organisms. It classifies localities, quadrats or and other applications ofthis method see Craw areas (which are analogous to taxa) by their (1988, 1989),Morrone& Crisci(1990,1995),Craw shared taxa (wh ich are analogous to characters) et al. (1999) and Crisci et al. (2000). The analysis according to the most parsirnonious solution of the data matrix of 13 areas of endemisrn vs. (Morrone & Crisci 1995, Crisci et al. 2000). distributional data of I 12 taxa (Table 3) was The rnethod was originally proposed by Rosen carried out with SECANT 2.2 (Salisbury 1999). (1988) using localities as study units. SECANT isa program for identifying all groups

Fig.2 A-B. Individual tracks. A, Achyroc!ine satureioides ce), Baccharis crispa C-); B, B. ulicina (e), Conyza bonariensis (-).

27 Astcraccae trom Tandilia and Vcntania Table 2. Species of Asteraceae from Tandilia and Ventania and other areas of endemism considered in the study. Taxa with the same areas of endemism have been indented. Areas ofendemism for each taxa are indicated in Table 3.

1 Achvrocline satnreioidcs (Lam.) De. 59. Grindelia aegialítís Cabrera 2. Acmella decumbens (Sw.) R. K. Jansen var. affinis (Hook. & 60. U. brachystephana Griseb. Arn.) R. K. Jansen 61. U./JII/chella Dunal var. discoidea (Hook & Am.) A. Bartoli 3. As/er squamatus (Spreng.) Hieran. & Tortosa 4. Baccharis artetnisioides I Iook. & Arn. 62. c. vcntancnsis A. Bartoli & Tortosa 5. Ji articutata (Lam.) Pers. 63. Gutierrezia gilliesii Griseb. 6. H crispo Spreng. 64. Helenium radiotum (Lcss.) Seckt 7. Ji gilliesii A. Gray 65. Hieracium palezienxii Zahn 8. Ji spicat« (Lam.) Bail!. 66. l Iyalis argenten D. Don ex Hook. & Arn, val'. latisouama 9. H. sleJ/opl1yl/a Ariza Cabrera 10. n triangularis Hauman 67. Trichoctine sínucüa (D. Don) Cabrera 11 n tridentcua Vahl var. suboppositc: (DC) Cabrera 68. Hypochaeris pctiolaris (Ilook. & Arn.) Griseb, 12. 1I tritnera (Less.) DC. 69. Hystenonicojasionotdcs Willd. 13. B. nlicina Hook. & Arn. 70. Cony:a bnnoríens¡s (L.) Cronq. var. allglfsli/olia (Cabrera)

14. lsidens laevis (L.) Britton, Sterns & Poggenb. l. Cabrera 15. Chuptalia exscapa (Pers.) Bakcr var. cxscapa 71.I(vs/eriolliea pinifotia (Poir.) l3aker 16. C. i;;J1ola Burkart 72. tIvnochaerís roscnguruii Cabrera var ptnncülfidu 17. C. integcrrinta (Vel!.) I3urkar1 (Speg.) Cabrera 18. (' I'illfselllfides (Vahl) Baker 73. l.ncilia acntifuli« (Poir.) Cass. 19. r: runcinata Kunth 74. Microgyncl!a trifurcata (Less.) Grau 20. (' sinucna (Less.) Baker 75. Noticastrum marginantm (Kunth) Cuatrcc. 21 Baccharis corídifola OC. 76.I'crezia multiflora (Humb. & Bonpl.) Lcss. subsp. sonctnfotiu 22. Conyzu sumcurensís (Reitz.) E. Walkcr (I3aker) Vuillcum. 23. Clw(juiroKu erinacea D. Don subsp. crinacea 77. liupatorintn connnersanii (Cass.) l Iieron. 24. Conv:a honariensis (L.) Cronq. var. bonaricnsis 78. llypuchuerís grisehachii Cabrera 25. e: floribunda Kunth 79. Pluchea sagi//alls (Larn.) Cabrera 26. (.'. tnonorchis (Griseb.) Cabrera 80. ,)'enecio grisebctchii Bakcr val'. grisebachii 27. Hnlochetlns brasiliensís (L.) Cabrera 81. Podocoma hírsrüa (Hook. & Am.) Bakcr 28. Mikania pcriplocífolia Ilook. & Arn. 82. Cony:a hlukei (Cabrera) Cabrera 20. Panphalea heterophvlla Less. 83. Eupatortnm snbhastut um lIook. & Arn. 30. Pndncoma hierac!fália (Poir.) Cass. 84. Noticastrum diffusnn: (Pcrs.) Cabrera 31 Conv:a prinmlifolia (Lam.) Cuatree. & l.ourteig 85. Senecto arechavaletae Bakcr 32. C. serrana Cabrera 86. S bonaricnsis Hook. & Arn.

XL Enpcuoriutn bnníifolirnn I Iook. & Arn. var. Ivnniifolinrn 87. S. griselwchii Baker val'. subincanns Cabrera 34. lo'. bnpleurifolimn DC 88. S osteni¡ Mattf. 35. h'. macrocrphalntn Less. 89. Senecto ntontevidensis (Sprcng.) Bakcr 36. Chevreulia sarmentosa (Pcrs.) S. r. Blake 90. Micropsis australts Cabrera 37. Enpcnoriuin squarrnlosum Hook. & Arn, 91 Sencc¡o panrpeanus Cabrera 38. Bctccharis gnapbcüíoidcs Sprcng. 92. S pulcher Ilook. & Arn. 39. Micropsis spastudata (Pers.) Cabrera 93. liupatoriiun tanacctifolunn Gillics ex Ilook. & Arn 40. Facelís renrsa (Lam.) Seh. Bip. subsp, pcttulü Beauv. 94, Grindelia tniphthahnouk:s DC. 41 Jo'. re/lisa (Larn.) Sch. Bip. subsp. re/lisa 95 l lvpochaeris l'oriegu/o (Lam.) Baker 42. Guilla nt¡« meg apot amtca (Spreng.) Bak cr val' 96. Sonunerfeltia spinulosa (Spreng.) I.css. scabiosoidca (Arn. ex DC) Ilaker 97. Senecto subnlan,» O. Don ex Ilook. & Arn. varo crcctus Ilook. 43. Flavcria hcnunanii Dimitri & Orfila &Am. 44. Stuckcrticlla peregrina Beauv. 98. So/ira pterosperm« (Juss.) Less. 45, Guillardia nusgapotamica (Sprcng.) Baker var. tueg«: 99. Stevia saturciijolia (Lam.) Larn. var. pcuaganica lIieron. po/{flJllca 100. S. sotureütot¡« (Lam.) Lam, varo satureiifolia 46. Berrea gnaphalioídes (Less.) Bcauv. 101 Ihclesperma megajJotmJ/iclflll (Sprcng.) Kuntzc 47. Grnnochaeta americana (Mill.) Wedd. 102. Noticastrum argentinensc (Cabrera) Cuatrcc. 48. (j. argentina Cabrera 103. Vernonia echioides Lcss. 49. (; fatccua (Lam.) Cabrera 104. Actnella decumhcns (Sw.) R. K. Jansen val' decumbcns 50. (i subfulcato (Cabrera) Cabrera 105. líaccharis gel1;sli!o/ia DC. 51. Gienochactofituginea (DC.) Cabrera 106. Criscía stricta (Spreng.) Katinas 52. U plutensis (Cabrera) Cabrera 107. Enpatorinm twccdiamnn Hook. & Arn. 53. r; stachvdifolío (Larn.) Cabrera 108. Hypochacris megapotomíca Cabrera 54, Gnuphctlium cabrcrcu: S. E. Freire 109 Senecto selloí (Spreng.) DC 55. Hypochucrts pcnnpasíca Cabrera 110. Stcvia nntltiüristüta Sprcng. 56, Gnaphaliun) cheironthifolnnn Lam. 111 Vernoniaflexuosa Sims. 57. c. gaudic/wudiomfm DC. \12. Zcxtncniu buphthahniflom (Lorcntz) Ariza 58. (i lelfcopep!tflll Cabrera

28 Crisci el al. Craw ( 1988) and Cracraft (1991) presented a from the fíve cliques,as follows (Fig.4): variation of the method using are as of Clique A: ««««(T A, UR), SB), PAM), VE), endem ism as study un its to identify the SPA), CHA), SA), NA), PAT, CHI), supported hierarchical information contained in the by the 30 individual tracks 3, 5, 20, 21,23,25, geographical distribution of organisms to 26,27,28,29,30,36,37,38,40,41,81,82,83,84, establish are a relationships. In this method 87,88,103,104,105,106,107,108,109,110 a data matrix of endemism areas by taxa is (Fig.4A). constructed, and the presence of a taxon in Clique B: ««««(T A, UR), SB), PAM), VE), an area is coded as 1 and its absence as O. A SPA), SA), CHA), NA), CHI, PAT), supported parsimony algorithm is applied to the matrix by the 30 individual tracks 3,5,20,21,23,25, to obtain a cladogram(s). PAE cladograms 26,27,28,29,30,36,37,38,63,64,81,82,83,84, represent nested sets of areas, in which ter- 87,88,103,104,105,106,107,108,109,110 minal dichotomies represent two areas (Fig. 4 B). between which the most recent biotic Clique C: ««««(T A, UR), SB), PAM), VE), interchange has occurred (Morrone & Crisci SPA), CHA), SA), CHI) NA, PAT), supported 1995). In this analysis the same are as and by the 30 individual tracks 3, 20, 21,23,25,26, data matrix (Table 3) used in the 27,28,29,30,36,37,38,40,41,81,82,83,84,87, panbiogeographic approach were taken into 88,98,103,104,105,106,107,108,109,110 consideration. The analysis was carried out (Fig.4C). with PAUP* version 4.0 beta 2 (Swofford Clique O: ««««(TA, UR), SB), PAM), VE), 1999), applying the branch-and-bound SPA), SA), CHA), CHI), NA, PAT), supported option. A strict consensus tree was by the 30 individual tracks 3, 20, 21,23,25,26, constructed based on the results generated 27,28,29,30,36,37,38,63,64,81,82,83,84,87, from PAUP*. As proposed by Rosen (1988), 88,98,103,104,105,106,107,108,109,110 the cladogram was rooted with a hypothetical (Fig.4 O). area coded all zeros. Clique E: ««««(TA, UR), SB), PAM), VE), SPA), SA, CHA) NA, PAT, CHl), supported RESULTS by the 27 individual tracks 3, 20, 21,23,25,26, 27,28,29,30,36,37,38,81,82,83,84,87,88, Pan biogeography (compatibility track method) 103, 104,105,106,107, 108, 109, 110(Fig. 4 E). Based on the data matrix (Table 3), the Mahuidas and Central and North America are combination ofthe individual tracks resulted the only areas that appear in none of these in four largest cliques (generalized tracks) of pattern of relationships in the fíve generalized 30 individual tracks each one (Fig. 4 A-O). tracks obtained. The intersection of 27 individual tracks Oue to the fifth clique (E) represents the (identifíed with the numbers 3, 20, 21,23,25, intersection of27 individual tracks common 26,27,28,29,30,36,37,38,81,82,83,84,87,88, to the four largest cliques, it was selected to 103,104,105,106,107,108,109,110 inTable2) show the taxa supporting the areas common to the four largest cliques is identifíed relationship (Fig. 4 E): as the fifth clique or generalized track (Fig. 4 (1) (TA,UR). Based on Senecio grisebachii E). In this way, six individual tracks (5, 40, 41, varo subincanus (individual track 87) and S. 63, 64, and 98 in Table 2) are combined in ostenii (track 88). groups ofthree with the 27 individual tracks (2) «TA,UR), SB). Based on Eupatorium to form the four cliques of30 individual tracks squarrulosum (track 37), Baccharis gnapha- mentioned above. lioides (track 38), and Micropsis spathulata Fivetrees connecting the areas can be constructed (track 39).

29 Asteraceae from Tandilia and Ventania

Fig.3 A-F. Individual tracks. A, Podocoma hirsuta ce), Senecio arechavaletae C-); B, Hieracium palezieuxii ce), Hyalis argentea var. latisquama C-). e, Hysterionica pinifolia ce), Perezia multiflora subsp. sonchifolia C-); D, Senecio pulcher ce), Sommerfeltia spinulosa C-); E, Senecio bonariensis ce), Senecio pampeanus C-); F, Vernonia echioides ce), Zexmenia buphtalmiflora C-).

30 Crisci el al .

,------CHI .------PAT f------PAT ~------CHI ~------NA .------NA ,------SA .------CHA ....------CHA r------SA ~-----SPA ,------SPA .------VE ,------VE .-----PAM ~---PAM .----SB ~---SB TA TA UR UR

,.------PAT ~------PAT f------NA ~------NA ....------CHI ~------CHI ~------SA ~------CHA ,------CHA ....------SA r------SPA ,------SPA .------VE .------VE ,.----PAM ,.-----PAM ,----SB .----SB TA TA UR UR

INDIVIDUAL TRACKS ....------CHI ~------PAT $ 3.24.31 ~~------NA Q5 r------CHA X 20.21 ~------SA .98 ....------SPA <:> 22.64 ,----- VE • 41.42 .------PAM * 81.82.83.84 ~--SB $ 26.27.28.29.30 TA * 103.104.105.106 E UR 107.108.109.110 * 37.38.39 ~ 87.88

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 taxa names and Table 3 for individual tracks). CHA = Chaco; CHI = Central Chile; NA = Northern Andes; PAM = Pampa; PAT= Patagonia; SA = Sierras Subandinas; SB = Southem Brazil; SPA = Sierras Pampeanas; TA = Tandilia; UR = Uruguay; VE = Ventania.

31 Asteraccac from TandiJia and Ventania

(3) (((TA,UR), SB), PAM). Based on Vernonia applied in this study, and represented in Fig. echioides (track 103), Acmella decumbens 6 as a generalized track. Both methods show (track 104), Baccharis genistifolia (track 105), that Tandilia is closer to Uruguay, southern stricta (track 106), Eupatorium Brazil, and Pampa than to Ventania. On the tweedianum (track 107), Hypochaeris other hand, PAE and panbiogeography megapotamica (track 108), Senecio selloi (track resulted in different relationships of the 109), and Stevia multiaristata (track 110). remaining other areas with the m a in (4) ((((TA,UR), SB), PAM), VE). Based on distributional pattern. PAE shows that Sie- Conyza monorchis (track 26), Holocheilus rras Subandinas (SA) (Fig. 5 A-E) are related brasiliensis (track 27), Mikania periplocifolia to this pattern, whereas panbiogeo-graphy (track 28), Panphalea heterophylla (track 29), shows that Sierras Subandinas can be directly and Podocoma hieracifolia (track 30). related to it (Fig. 4 B, D) or cannot be directly (5) (((((TA,UR), SB), PAM), VE), SPA). Based related (Fig. 4 A, C); PAE shows Chaco (CHA) on Podocoma hirsuta (track 81), Conyza blakei as not closely related to the pattern (Fig. 5 A- (track 82), Eupatorium subhastatum (track 83), E), whereas panbiogeography shows CHA and Noticastrum diffusum (track 84). connected to it (Fig. 4 A-E); PAE shows Mahuidas (MA) and Central and North Parsimony analysis of endemicity Arnerica (CNA) not closely related to the pattern (Fig. 5 A-E), and panbiogeography The analysis ofthe data matrix (Table 3) with shows these last two areas very distantly PAUP* generated four area cladograms (Fig. related to it (for this reason they are not 5 A-D) with 263 steps, c.i.= 0.426, and r.i. = indicated in the results). 0.663. The strict con sen sus ofthe four trees (Fig. 5 E) shows the following area relation- DISCUSSION ships: (Mahuidas (Patagonia (Chile (Sierras Subandinas (Sierras Pampeanas (Ventania The main distributional pattern connecting (Pampa (Southern Brazil - Tandilia - Uru- Tandilia and Ventania with southern Brazil, guay)))))))). The remaining areas: Chaco Pampa, Uruguay, and Sierras Pampeanas (CHA), Central and North America (CNA), and obtained through the cornpatibility track northern Andes (NA) do not show resolved method and PAE partially agree with others relationships with the other areas in the authors' hypotheses. consensus. A biotic rnigratorial route frorn the Andes to Brazil, through Sierras Pampeanas, sierras of Comparison ofboth methods Buenos Aires, sierras ofUruguay, and Planalto and sierras of Brazil was postulated rnany years The comparison of the results obtained ago by Hicken (1918-1919) and Brade (1942). applying panbiogeography (compatibility Frenguelli (1950) gavethe name of periparnpasic track rnethod) and parsimony analysis of orogenic are (arco peripampásico serrano) to endernicity to the areas under study shows a the biotic corridor constituted by Sierras coincidence in the relationships of Sierras Pampeanas, Mahuidas, and Tandilia. Ringuelet Pampeanas (SPA), Ventania (VE), Pampa (1956) postulated that the province ofBuenos (PAM), Southern Brazil (SB), Tandilia (T A), Aires constitutes a cornposite area due to the and Uruguay (UR). presence oftwo types offauna: one related to The areas relationship PAM-SB-SPA-TA-UR- the Subtropical domain (Guaianian-Brazilian), VE conform a rnain distributional pattern and the other related to the Austral region strongly supported by the two approaches (Andean-Patagonic, or Chilean-Patagonic).

32 Crisci el al.

11 Q) ,.... ,.... ,.... ,.... ,.... o u ,....~l o ,....o o o o o,.... o o,.... ,.... o,.... u f- e o o o o o o o o o ~ Q) o o o o o o o ,.... o o ,.... o ,.... ,.... « VJ ,.... ,.... ,.... ,.... o 6 e, .D 0-, o o o o o o ,.... o o ,.... o ,.... ,.... o (.:j f'? 8 8 8 8 8 8 ,.... 8 8 ,.... 8 •....• ,.... 8 11 « ,.... 'O o o o o o o ,.... o o ,.... ,.... ,.... o. ,.... or, ,.... ,.... o ,.... ,.... o « .s:;j "1' 8 8 8 8 8 8 ,.... 8 8 ,.... 8 ,.... ,.... 8 ::c 6 e ,....o ,....o ,.... ,....o ,.... ,....o ,.... ,.... ,....o e,~ co Ñ,.... 8 8 ,.... 8 ,.... 8 ,.... ,.... 8 o ,.... o ,.... ,.... U ~ o o o o o o o 11 Q) o o o o o o o ,.... o o o ,.... ,.... o ,.... .~ 0-, o o ,.... o ,.... o o ,.... o o ,.... ,.... o ,.... ,.... ,.... ,.... o ,.... ,.... ,.... ,.... ,.... ,.... .~ ;¿ > f'? 8 o ,.... 8 8 8 ,.... ,.... ,.... ,.... ,.... o o o 11 o 'O o o o o o o o o o ,.... o ,.... ,.... ,.... 6 « [..Ll ,.... 'n ,.... ,.... ,.... ,.... Q) e, ...... ,.... -o "1' 8 8 8 8 8 8 8 8 8 ...... 8 ,...... > ...... ,.... ,...... e o. Ñ...... 8 8 8 8 ,....8 8 ...... 8 8...... 8 ...... 8 •....• ...... Q) c-, o o o o o o o o 4- g ~ ...... o :.... ::l o o o o o o o o o ,.... bl) O'. o o o o o o ...... ,.... o ,.... ,...... ,.... VJ bl) o...... ::l f'? •....• ,.... 'Vi S :.... 8 8 8 8 8 8 8...... 8...... 8 8 ,....8 ...... 8,.... >. o 0-, 'O o o o o o o o o o...... ~ 'n ,....o ...... o ...... o ...... o ...... ,.... 11 "1' 8 8 8 8 8 8 ...... 8 8 ...... ,.... ,...... ,.... ~ 11 ...... ,.... ,.... •....• ...... ,.... ¡¡¡ f- Ñ...... 8 8 8 8 8 8 ...... 8 8 ,...... •....• ,.... ,.... >,~ ~ o o o o o o o o ~ o o ...... o o o o ...... o ...... ,...... o § O O'. o ...... o o o o ...... o ...... ,.... ,.... o r ¿ ,.... •....• ,...... VJ .- f!2 ...... •....• ,...... 5 v;-::::: 8 8 8 8 8 8 ...... 8 8 8 8 ...... Vl o o o o -o -o 00 'O o o o ...... o...... o o...... '- 'ro o...... o ...... o ...... ~ e e "1' 8 8 8,.... 8 8 o...... 8 ...... o ,...... •....•8 ...... ,.... 0.« ~ Ñ o o o o o o •....• ...... ,.... Q) f- ...... 8 8 8 8 8 e o o o o o o o o o o o ...... -5 11 ...... •... Q)'- o o ,.... o o o ,.... o o ,...... c: O'. o ...... ,.... o ...... o ...... ,.... o ...... o o ...... t.8 < ...... o ...... t f- f!2 8 8 8 8 8 8 ,.... 8 8 8 ,.... 8 o ...... -o o t..· 'O o o o o o o ,.... o o o ...... o o ,.... Vl oro ...... ,.... e z; ~ ...... ,....o ,....o ...... •....• ...... ,.... ~ e "1' 8 8 ...... 8 8 ,....8 8 ,.... 8...... ,.... VJ ,....o ...... ~ ...... Ñ 8 8 ,....o 8 o 8 ...... o ...... o o 8...... ,....o ,.... ,....8 Vl o.Q) o o o o o o o >. « ...... ,...... o o o o o o ,.... o o •....• o "'§:z 6 O'. o o ...... o o ...... o o o...... o ...... o ...... o ,....o ,....o ~ f!2 ,....o ...... ,....o ,...... ~ Vl e, 8 8 o 8 8 ,.... ce 'O o o ...... o ...... o o o o ...... ,...... ~ VJ ...... u ce 'O 'n ...... ,...... ~ ~ "1' 8 8 8 ,....8 8 ...... 8 ...... 8 ,.... ,....8 ,....8 ,.... .t:: ::l t:: o...... c: ...... Ñ 8 8...... 8 o...... 8 ...... o 8 o...... o o ...... o ...... co .~ o o o o o ,.... .~\ VJ ;¿ o o o o o o o ...... o ,.... ,...... o ...... 11 0-, o o o o o o ...... o ...... ,.... ,.... o ...... :.o 11 N ...... o ...... o ...... o o o ...... ,...... <' f'? 8 8 ...... ,.... o ...... ,.... ,...... ~< e, ~ 'O o o ...... o o o ...... o ...... o ,.... ,.... .D 'ro 'ro ...... o ...... ,...... ;::-;¿ VJ ro 8 8 ...... 8 8 8 ...... o ...... o ,.... "1' o ...... ,.... ,...... o ...... 8o o ,.... § f- Ñ 8 ...... o 8 8 8 ...... o o ...... ~ 'Ñ 4- ...... o ...... o o o o ...... o o ,...... •....• ...... u u ~ o Q) .¡:: :...... Q) o o o o o o o o o ,.... o Q) O'...... ,.... ,.... -5 ca VJ o o o o o o o o o ...... o ,.... ,.... o 6 o f'? ...... •....• ...... '- e•... 8 8 8 8 8 8 ...... 8 8 ...... 8 ...... 8 8 t.8 <' -5 'O o o o ,.... o o o ,.... o ...... "1' 'n ,...... ,.... o ,.... ~ . ..c 8 8 8 o 8 8 ...... 8 ...... o ,...... ,.... VJ "1' ,.... ,.... ,...... ,...... ,.... ,.... ,.... t ~ 8 ...... •....• ,...... ~ g -o Ñ...... 8 ...... 8 ,....o 8 o ...... 8 ...... o ...... 8 ...... o ,.... ,...... u o e o o o ~ Z VJ b o.o o o o o o o o ...... o o ...... o ...... -o 11 0-, o o o o o o ...... o o ,.... o •....• ...... C;; e Vl ...... •....• ,...... ::l ~ ro •...Q) f'? 8 8 8 8 8 8 ...... 8 8 ...... 8 ...... ,...... VJ 'O o o o o o o ,.... o o ...... o ...... ,.... ,.... M ...... ,.... ,.... ,.... ,...... ~ ~ o'- 'n ...... o ...... ,.... VJ 8 8 o ,.... ,.... .2: ~ u "1' ...... o ,.... ,.... o ...... o o ...... -o ~ C"J Ñ 8 8 o 8 8 8 ...... o ,.... ,.... ,...... (jj .::: ...... ,...... ,...... ,...... ::: -o x o o o o o o U e E o o ...... o o o o ...... o ...... ,.... ,.... ,...... vi 4- O'...... ,...... •....• ,.... 11 ~ o o o ,.... o o ...... o...... > .D o f!2 ...... o ,.... o...... o...... o ...... Vl « ::l :.... 8 8 8 8 8 ...... 8 ...... o Q) 'O o o o o o o o o o o o ~ VJ """'1 Ir, ...... ,.... ,.... ,...... Q)•... Z .D ,....O ,....O ,....O ...... O ,.... ,.... ,...... VJ "1' 8 ...... 8...... 8...... ,.... ,....O U •...~ E ...... O...... O...... ~ ;::l ...... Ñ 8 ...... O 8 8 O ...... O ...... O ,....8 ,...... X ~ '- O o· O O O O O .~ z ...... "5 ..c: VJ O O O O O O O ,....O ,.... O O ...... O O...... ,.... ~ O'. O ...... O O O O O ...... O ,....O ...... •....•O ...... U 11 O O...... O O E f'? 8 ...... 8 8 8 ,...... O ...... O O ,.... Cij 'O O O ...... O O O ,...... O O ...... O ...... ~ < Q) ...... ,.... ,...... VJ 'n ...... O ...... ,...... 'i:j b U "1' 8 ,....O ...... 8 8 O,...... ,....8 ,.... ,....O ...... •....• ...... e e ...... o ,.... ,.... ,.... O Q) .~ Q) ...... Ñ 8 o o 8 ...... o ...... o ...... o ...... o ...... o...... o...... Vl o o o o o ,...... -i U e Q) o :.... Q) bl) o. :c ~ ~ "'" E-< ~ ::c 'i:j o 5 ~ ~ ~ ~ ~ § p... p...

34 Crisci el al.

OUT OUT CNA CHA NA CHI CHA PAM CHI UR PAM - r--r- S8 UR -~ TA VE S8 r-e- TA ,.-- SPA VE '-- lr= SA SPA PAT '-- SA MA PAT NA A MA B CNA

OUT ,.------OUT CNA ,------CHA NA .------CHI CHA ,-----PAM CHI S8 PAM UR S8 TA UR L-.-----VE TA L------SPA VE L------SA SPA L------PAT SA L-.------MA PAT L------NA D L-.. CNA MA

OUT CHA CHI PAM S8 TA UR VE SPA SA PAT MA E CNA NA

Fig. 5. A-E, Cladograms obtained after applying parsimony analysis of endemicity after applying PAUP*. 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 Asteraceae from Tandilia and Ventania --1 -J_o'\

\~

re \

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

\ PAM

--\\ \ \ \

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

36 Crisci el al, In surnmary, it is probable that the evolution BR¡\DE,A. C. 1942. A composicáo da flora do of Asteraceae in Tandilia and Ventania was Itatiaia, Rodriguesia 6 (15): 29-43. mainly affected by Tertiary and Quaternary BURK¡\RT,A. 1967. Leguminosae. Vol. 4 (3), pp. geologic events. As a result, this biota shows 394-647 en: A. L. Cabrera (ed.). Flora de la a high level of diversity, many endem ic taxa, provincia de Buenos Aires. Colección and closer relationships with southern Brazil, Científica INTA, Buenos Aires, Argenti- Uruguay, Pampa, and Sierras Pampeanas na. rather than with other arcas of América. The C¡\BRER¡\,A. L. 1938. Excursión botánica por discon-tinuous pattern, i.e., occurring in las Sierras Australes de la provincia de elevated areas surrounding a core area ofCha- Buenos Aires. Revista del Museo de La co and Monte vegetation, is what one would Plata (nueva serie) Sección Oficial: 60-69. expect under vicariance due to dry conditions, C¡\BRERA,A. L. 1940. Notas sobre la vegeta- although dis-persal is taken also as a common ción del parque provincial Sierra de la Ven- and impor-tant distributional event. Further tana. Anuario Rural de la Dirección de correlations with analyses of other plant and Agricultura, Ganadería e Industria 8: 3-16. animal taxa will determine the generality of C¡\BRERA,A. L. 1963. Compuestas. Vol. 4 (6), the pattern shown by Asteraceae. pp. 1-443 en: A. L. Cabrera (ed.). Flora de la provincia de Buenos Aires. Colección ACKNOWLEDGEMENTS Científica INT A, Buenos Aires, Argenti- na. We thank Paul Berry, John Grehan, Edgardo C¡\BRER¡\,A. L. 1965. Solanaceae. Vol. 4 (5), Ortiz Jaureguizar, Mario Teruggi, Rosendo pp. 190-250 en: A. L. Cabrera (ed.). Flora Pascual, and Elías de la Sota for helpful de la provincia de Buenos Aires. Colec- comments on the manuscript. We thank Jorge ción Científica INT A, Buenos Aires, Ar- Frangi for checking the Iist of species. Special gentina. thanks to Benjamin A. Salisbury for his gene- C¡\BRER¡\,A. L. 1968. Bromeliaceae. Vol. 4 (1), rous assistance with his program SECANT. pp. 448-459 en: A. L. Cabrera (ed.). Flora de This study was supported by grant 5776-96 la provincia de Buenos Aires. Colección of the National Geographic Society. Support Científica INT A, Buenos Aires, Argentina. 11'0111the Consejo Nacional de Investigaciones Cien- C¡\BRERA,A. L. 1970. Festuca L. (Poaceae). tíficas y Técnicas (CONICET), Argentina, towhich Vol. 4 (2), pp. 125-128 en: A. L. Cabrera the authors belong, is gratefully acknowledged. (ed.). Flora de la provincia de Buenos Ai- res. Colección Científica INT A, Buenos Ai- L1TERATURE ClTED res, Argentina. C¡\BRER¡\,A. L. & M. A. TORRES.1970. Stipa L. AI.BOIF,N. 1895. Rapport préliminaire sur une (Poaceae). Vol. 4 (2), pp. 255-290 en: A. L. excursion botanique dans la Sierra Ventana. Cabrera (ed.). Flora de la provincia de Bue- Revista del Musco de La Plata 7: 183-187. nos Aires. Colección Científica INTA, AXFLROU,D. i. M. T. K¡\L1NARROYO& P. H. Buenos Aires, Argentina. R¡\ VEN. 1991. Historical development of C¡\BRERA,A. L. & A. WILLlNK. 1973. Biogeo- temperate vegetation in the Americas. Revista grafía de América Latina. Monografía 13, chilena de Historia Natural 64:413-446. Serie de Biología, OEA, Washington, D. C. BOLLCKE,O. 1967.Cruciferae, Vol. 4 (3), pp. 281- C¡\I3RER¡\,A. L. & E. M. Z¡\RDINI.1978. Manual 371 en: A. L. Cabrera (ed.). Flora de la pro- de la flora de los alrededores de Buenos vincia de Buenos Aires. Colección Cientí- Aires. Editorial ACME, S.A.C.I., Buenos fica INT A, Buenos Aires, Argentina. Aires, Argentina.

37 Asteraccae lrom Tandilia and Ventania CABRERA,A. L., 1. V. CRISCI,G. DELUCCHI,S. E. CRAW,R. c., 1. R. GREHAN& M. 1. HEADS.1999. FREIRE, D. GIULlANO, L. rHARLEGIII,L. Panbiogeography: Tracking the his-tory KATlNAS, A. SÁENZ, G. SANCHO & E. oflife. Oxford University Press, New York, URTUBEY.2000. Catálogo ilustrado de las Oxford. compuestas de la provincia de Buenos Ai- CRISCI,1. V., L. KATINAS& P. POSADAS.2000. res. COBrOBO (Comisión de Biodiver- Introducción a la teoría y práctica de la sidad Bonaerense). Convenio Secretaría biogeografía histórica. Sociedad Argenti- de Política Ambiental-UNLP, Provincia de na de Botánica, Buenos Aires, Argentina. Buenos Aires, Argentina. CROIZAT,L. 1958. Panbiogeography. Vols. 1,2a, CÁMARA HERNÁNDEZ,J. 1970. Bromus L. and 2b. Published by the author, Caracas, (Poaceae). Vol. 4 (2), pp. 85-101 en: A. L. Venezuela. Cabrera (ed.). Flora de la provincia de Bue- CROIZAT,L. 1981. Biogeography: past, present, nos Aires. Colección Científica INT A, and future. Pp. 501-523 in: G. Nelson & D. Buenos Aires, Argentina. Rosen (eds.), Vicariance biogeography: A CASTELLANOS,A. & R. PÉREZMOREAU.1944. critique. Columbia University Press, New Los tipos de vegetación de la República York. Argentina. Universidad de Buenos Aires DARLlNGTON,P. 1. JR. 1968. Biogeography of Publicaciones 3. the southem end of the world. Harvard CONNOR,E. F. 1988. Fossils, phenetics and University Press, Cambridge, Massachu- phylogenetics: inferring the historical ssets. dynamics ofbiogeographic distributions. DARWIN,C. 1846. Geological observations of Pp. 254-269 in: J. K. Liebherr (ed.), South America, being the third part ofthe Zoogeography of Caribbean insects. geology ofthe voyage ofthe Beagle during Comell University Press, Ithaca, New York. 1832-1836. London. CHEBATAROFF,J. 1960. Algunos aspectos evo- FABRIS,H. A. 1965. Polygalaceae. Vol. 4 (4), lutivos de la vegetación de la Provincia pp. 58-69 en: A. L. Cabrera (ed.). Flora de Fitogeográfica Uruguayense. Revista Na- la provincia de Buenos Aires. Colección cional (Uruguay) 20 1: 3-18. Científica INT A, Buenos Aires, Argenti- CRACRAFT,J. 1991. Patterns of diversification na. within continental biotas: hierarchical FRANGI,1. L. 1973. Las comunidades vegeta- congruence among the are as of endemism les de las Sierras de Tandil (provincia de of Australian vertebrates. Australian Buenos Aires). Tesis, Universidad Nacio- Systematic Botany 4: 211-427. nal de La Plata, La Plata, Argentina. CRAW, R. C. 1988. Continuing the synthesis FRENGUELLI,1. 1950. Rasgos generales de la between panbiogeography, phylogenetic morfología y la geología de la provincia systematics and geology as illustrated by de Buenos Aires. Serie 2, n. 33, pp. 72. L. studies on the biogeography of New E. M. 1.T., La Plata, Argentina. Zealand and the Chatham islands. FUNK,V. A., H. ROI3INSON,G. S. MCKEE& 1. F. Systematic Zoology 37: 291-310. PRUSKI. 1995. Neotropical montane CRAW, R. C. 1989. Quantitative panbio- Compositae with an emphasis on the An- geography: introduction to methods. New des. Pp. 451-471 in: S. P. Churchill, H. Zealand Journal ofZoology 16: 485-494. Balslev, E. Forero & J. L. Luteyn (eds.). CRAW,R. C. 1990. New Zealand biogeography: Biodiversity and Conservation in a panbiogeographic approach. New Zea- Neotropical Montane Forests. The New land Journal ofZoology 16: 527-547. York Botanical Garden, New York.

38 Crisci el al. GORDILLO,C. E. & A. N. LENCINAS.1980. Sie- Curculionidae) and the historical rras Pampeanas de Córdoba y San Luis. relationships ofthe Andean province. Glo- Vol. 1, pp. 577-638 en: Segundo Simposio bal Ecology and Biogeography Letters 4: de Geología Regional Argentina. Córdo- 188-194. ba, Argentina. MORRONE,J. J. & J. V. CRISC!. 1990. Panbio- HARRINGTON,H. J. 1980. Sierras Australes de geografía: fundamentos y métodos. Evo- la provincia de Buenos Aires. Vol. 2, pp. lución Biológica (Bogotá) 4: 119-140. 967-983 en: Segundo Simposio de Geolo- MORRONE,J. J. & J. V. CRISC!.1995. Historical gía Regional Argentina. Córdoba, Argen- biogeography: introduction to methods. tina. An nu a l Review of Ecology and HAUTHAL,R. 190 l. Contribución al conocimien- Systematics 26: 373-40 l. to de la geología de la Provincia de Bue- MORRONE,J. J., L. KATINAS& J. V. CRISC!.1997. nos Aires. I. Excursión de la Sierra de la A c1adistic biogeographic analysis of Cen- Ventana. II: Apuntes geológicos de las Sie- tral Chile. Joumal ofComparative Biology rras de Olavarría. Publicaciones de la Uni- 2 (1): 25-41. versidad Nacional de La Plata, Facultad Nrconx, E. G. 1970. Setaria Beauv. (Poaceae). de Ciencias Naturales y Museo l. Vol. 4 (2), pp. 439-460 en: A. L. Cabrera HICKEN,C. M. 1918-1919. Lamigración de los (ed.). Flora de la provincia de Buenos Ai- helechos en la flora de Tucumán. Pp. 187- res. Colección Científica INT A, Buenos 209. Primera Reunión Nacional de la So- Aires, Argentina. ciedad Científica Argentina de Ciencias PARODl,L. R. 1947. La estepa pampeana. Vol. Naturales, Tucumán, Argentina. 8, pp. 143-207 en: L. Hauman, A. Burkart, KRISTENSEN,M. J. & J. L. FRANG!.1995. La Sie- L. R. Parodi & A. L. Cabrera (eds.), Lave- rra de la Ventana: una isla de biodiversi- getación de la Argentina, Geografía de la Re- dad. Ciencia Hoy (Argentina) 5(30): 25-34. pública Argentina. Buenos Aires, Argentina. LLAMBÍAS,E. & C. E. PROZZ!.1972. Ventania. PETER,G. 2000. Las especies silvestres de la Pp. 79-10 1. Relatorio del Sexto Congreso familia de las Compuestas que habitan en de Geología Argentina. Buenos Aires, Ar- el partido de Saavedra (provincia de Bue- gentina. nos Aires). Tesina, Universidad Nacional MAURY,E. A. 1973. Los escorpiones de los del Sur, Bahía Blanca, Argentina. sistemas serranos de la provincia de Bue- PONCE,M. M. 1982. Morfologíaecológicacom- nos Aires. Physis (Buenos Aires) 32 (85): parada de las Filicópsidas de las Sierras 351-371. Australes de Buenos Aires (República Ar- MICHAUT,H. N. L. 1980. Sierras Pampeanas del gentina). Boletín de la Sociedad Argenti- norte de Córdoba, sur de Santiago del Es- nade Botánica21 (1-4): 187-211. tero, borde oriental de Catamarca y ángu- PONCE,M. M. 1986. Morfología ecológica de lo sudeste de Tucumán. Vol. 1, pp. 293-347 plantas saxícolas del Cerro de la Ventana, en: Segundo Simposio de Geología Regio- Prov. de Buenos Aires, Argentina. Darwi- nal Argentina, Córdoba, Argentina. niana27 (1-4): 237-271. MINGRAMM,A., A. Russo, A. Pozzo & L. PONTIROLl,A. 1965. Plantaginaceae. Vol. 4 (S), CAZAU.1980. Sierras Subandinas. Vol. 1, pp. 331-342 en: A. L. Cabrera (ed.). Flora pp. 95-137 en: Segundo Simposio de Geo- de la provincia de Buenos Aires. Colec- logía Regional Argentina, Córdoba, Ar- ción Científica lNT A, Buenos Aires, Ar- gentina. gentina. MORRONE,J. J. 1994. Distributional patterns PRADO,D. E. & P. E. GIBBS. 1993. Patterns of of species of Rhytirrhinini (Coleoptera: species distributions in the dry seasonal

39 Asteraccac Irom Tandilia and Vcntania

forests of South America. Annals of the SOTA, E. A. DE I.A. 1972. Sinopsis de las Missouri Botanical Garden 80: 902-927. Pteridófitas del Noroeste de la Argentina, RAMos, V. A. 1989. The birth ofSouthern South I. Darwiniana 17: 11-27. America. American Scientist 77 (5): 444- SOTA, E. A. 1)1' LA. 1973. La distribución geo- 450. gráfica de las Pteridófitas en e Icono RAVENNA,P. F. 1968. Iridaceae. Vo\. 4 (1), pp. sur de América meridional. Boletín de 539-565 en: A. L. Cabrera (ed.). Flora de la la Sociedad Argentina de Botánica 15 provincia de Buenos Aires. Colección Cien- (1): 23-34. tífica INT A, Buenos Aires, Argentina. SOTA,E. A. DELA. 1985. Las Pteridófitas de la RINGUELET,R. A. 1956. Panorama zoogeográ- provincia de La Pampa, Argentina. Revis- fico de la provincia de Buenos Aires. No- ta de la Facultad de Agronomía, Universi- tas del Museo de La Plata 18: 1-15. dad Nacional de La Pampa I (1-2): 23-34. ROLLERI, E. O. 1975. Provincias geológicas SPEGAZZlNI,C. L. 1896. Algunas observacio- bonaerenses. Pp. 29-53 en: Relatorio del nes sobre la flora de la Ventana. Revista Sexto Congreso de Geología Argentina. de la Facultad de Agronomía y Veterinaria Buenos Aires, Argentina. de La Plata 2 (23-24): 347-349. ROSEN,B. R. 1988. Frorn fossils to earth history: SPIGA77INI, C. L. 1901. Contribución al estu- applied historical biogeography. Pp. 437- dio de la flora de Tandil. Ed. Sessé, 481 in: A. A. Myers & P. S. Giller (eds.). Larrañaga y Renovales, La Plata y Bue- Analytical biogeography: an integrated nos Aires, Argentina. approach to the study of animal and plant SnJESSY,T. F., T. SANG& M. L. DI' VORE. 1996. distributions. Chapman & Hall, London & Phylogeny and biogeography 01' the NewYork. subfamily Barnadesioideae with impli- ROSEN, B. R. & A. B. SMIlII. 1988. Tectonic cations for early evolution of the Com- frorn fossils? Analysis of reef-coral and positae. Vo\. 1, pp, 463-490 in: D. 1. N. Hind sea-urchin distributions from late & H. J. Beentje (eds.). Compositae: Cretaceous to Recent, using a new Systematics. Proceedings of the Interna- method. Geological Society Special tional Compositae Conference Kew, 1994. Publication n° 37, pp. 275-306 in: M. G. Royal Botanic Gardens Kew. Audley & A. Hallam (eds.). Gondwana and SworFORD, D. L. 1999. PAUP*: Phylogenetic Tethys. Oxford Univ. Press, Oxford. analysis using parsimony. Version 4.0 beta 2. SAUSllURY,B. A. 1999. SECANT: strongest evi- Sinauer Associates, Sunderland, Massachusetts. den ce compatiblity analytic too\. Version TAYLOR, D. W. 1991. Paleobiogeographic 2.2. DepartmentofEcology and Evolutionary relationships of Andean angiosperms of Biology, Yale University, New Haven. Cretaceous to Pliocene age. Palaeogeo- SIMPSON,B. B. 1975. Pleistocene changes in graphy, Palaeoclimatology, Palaeoecology the flora of high tropical Andes. Paleo- 88: 69-84. biology 1: 273-294. TERUGGI, M. E. & 1. O. KILMlJRRAY. 1975. SIMPSONVUILlXUMIER,B. B. 1971. Pleistocene Tandilia. Pp. 55-77 en: Relatorio del Sexto changes in the fauna and flora of South Congreso de Geología Argentina. Buenos América. Science 173:771-774. Aires, Argentina. SOTA, E. A. DELA. 1967. Composición, origen TERlJGGI,M. E. & J. O. KILMURRAY.1980. Sie- y vinculación de la flora pteridológica de rras septentrionales de la provincia de las sierras de Buenos Aires (Argentina). Buenos Aires. Vo\. 2, pp. 918-965 en: Se- Boletín de la Sociedad Argentina de Botá- gundo Simposio de Geología Regional Ar- nica I1 (2-3): 105-128. gentina, Córdoba, Argentina.

40 Crisci el al. TORRES, M. A. 1970a. Piptochaetium Presl VEGA, L. & P. BELLAGAMBA.1990. Lista co- (Poaceae). Vol. 4 (2), pp. 234-255 en: A. L. mentada de la herpetofauna de las Sierras Cabrera (cd.). Flora de laprovinciade Bue- de Balcarce y Mar del Plata, Buenos Ai- nos Aires. Colección Científica INT A, res, Argentina. Cuadernos de Herpetología Buenos Aires, Argentina. 5(2): 10-14. TORRES,M. A. 1970b. roa L. (Poaceae). Vol. 4 ZAMBRANO,1. 1. 1980. Comarca de la cuenca (2), pp. 102-125 en: A. L. Cabrera (ed.). Flo- cretácica de Colorado. Vol. 2, pp. 1033-1070 ra de la provincia de Buenos Aires. Colec- en: Segundo Simposio de Geología Regio- ción Científica INTA, Buenos Aires, Ar- nal Argentina, Córdoba, Argentina. gentina. ZUIOAGA, F. O. & O. MORRONE(eds.). 1999. TROfANf,H. O. & P. E. STEIBEL.1999. Sinopsis Catálogo de las plantas vasculares de la de las compuestas (Compositae Giseke) República Argentina 11. Acanthaceae- de la provincia de La Pampa, República Ar- Euphorbiaceae (Dicotyledoneae). gentina. Revista de la Facultad de Agrono- Monographs in Systematic Botany from m ía de La Pampa, serie supl. 1, 10: 1-86. the Missouri Botanical Garden 74: 1-621.

Recibido: noviembre 1/2000 Aceptada: febrero 2012001

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