Historical biogeography of Asteraceae from Tandilia and Ventania mountain ranges (Buenos Aires, Argentina)

21 Pages·2001·3.5 MB·English

by  V., Jorge Crisci

Caldasia 23(1): 21-41
HISTORICAL BIOGEOGRAPHY OF THE
ASTERACEAE FROM TANDILIA AND VENTANIA
MOUNTAIN RANGES (BUENOS AIRES, ARGENTINA)
A nuestra amiga Pilar
JORGE CRISCI- V.
Departamento Científico de Plantas Vasculares, Museo de La Plata, Paseo del Bosque s.n.,
1900. LaPlata, Argentina. [email protected]
SUSANA FREIRE-E.
Departamento Científico de Plantas Vasculares, Museo de La Plata, Paseo del Bosque s.n.,
1900. LaPlata, Argentina.
GISELA SANCHO
Departamento Científico de Plantas Vasculares, Museo de La Plata, Paseo del Bosque s.n.,
1900. LaPlata, Argentina.
LILlANA KATlNAS
Departamento Científico de Plantas Vasculares, Museo de La Plata, Paseo del Bosque s.n.,
1900. LaPlata, Argentina.
ABSTRACT
Tandilia and Ventania arethe only systems ofmountain ranges situated inagrassy
steppe or"pampas" inthe political province ofBuenos Aires inArgentina. Tandilia
and Ventania have ahigh taxa diversity and endemicity. Ahistorical biogeographic
analysis was carried out on the basis of distributional patterns of species and
infraspecific taxa ofAsteraceae inhabiting Tandilia andVentania inorderto establish
therelationships ofthese mountain rangeswithotherareas.Twomethods were applied
fortheanalysis:panbiogeography usingthecompatibility trackmethod andparsimony
analysis ofendemicity (PAE). Thirteen areas were delimited forthe study: Southern
North America and Central América, 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 not
included). Bothmethods connectsouthern Brazil,Uruguay,Pampa, Tandilia, Ventania,
and Sierras Pampeanas, showing that the Asteraceae biota ofTandilia and Ventania
have closer relationships with the biota ofthese areasrather than with that ofSierras
Subandinas, North Andean, Chaco, Patagonia, Mahuidas, and Central Chile. The
Asteraceae ofCentral andNorth America appear lessrelated toTandilia and Ventania
inregard ofthe restofthe areas.The evolution ofAsteraceae inTandilia 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, isexplained principally byvicariance under dry conditions.
Keywords. Asteraceae; biogeography; Tandilia; Ventania.
RESUMEN
Las sierras septentrionales, odeTandilia ylasaustrales, ode Ventania, caracteriza-
daspor sugran diversidad orgánica ysualto número deendemismos, están ubicadas
en la planicie bonaerense depastizales denominada "pampa". Con elobjeto deesta-
blecer lasrelaciones entre Tandilia yVentania con otras áreas, seanalizaron lasdis-
tribuciones de lostaxones específicos einfraespecíficos de Asteraceae que habitan
estas sierras. Sedelimitaron trece áreas: surdeAmérica delNorte yAmérica Central,
surde Brasil, Uruguay, Pampa, Tandilia, Ventania, Chaco, Sierras Pampeanas, Sie-
rras Subandinas, Mahuidas, Patagonia, Chile Central, yAndes delNorte. Lasunida-
des de estudio fueron 112taxones de Asteraceae presentes en Tandilia yVentania
(taxones endémicos, adventicios ynaturalizados nofueron incluidos enelanálisis).
Las distribuciones de estos taxones fueron analizadas utilizando dos métodos:
panbiogeografía (compatibilidad detrazos) y análisis desimplicidad deendemismos.
Ambos métodos conectan surdeBrasil, Uruguay, Pampa, Tandilia, Ventania ySie-
rrasPampeanas. LasAsteraceae deTandilia yVentania estarían másrelacionadas con
estas áreas que con lade lasSierras Subandinas, Andes delNorte, Chaco, Patagonia,
Mahuidas yChile Central. Elárea deAmérica delNorte yAmérica Central aparece
menos relacionada aTandi1iayVentania enrelación alasotrasáreas. Laevolución de
lasAsteraceae deTandilia yVentania ylasáreas relacionadas hasidoprobablemente
afectada por loseventos geológicos delosperíodos Terciario yCuaternario. Ladis-
continuidad actual en el patrón de distribución de las Asteraceae, restringidas en
general a las zonas elevadas, es explicada principalmente por eventos vicariantes
debidos acondiciones áridas.
Palabras clave. Asteraceae; biogeografía; Tandilia; Ventania.
INTRODUCfION mountains rangesor"sierras", namely Tandilia
andVentania.
Since Darwin'sjourney inArgentina in 1835 From ageological point ofview, Tandilia and
thegeology ofTandilia andVentania andtheir 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 incentral eastern Argentina, iscovered recent origin dated from the Paleozoic age.
inmost of itssurface byanherbaceous, grassy Gondwanaland and Pre-Gondwanaland
steppe,ca\led"pampas". Thepampas areaplain events seem tohave played amajor role inthe
situated atthe sea level, oreven below the sea building ofthese mountain ranges.
level(likethebigdepressionofthe Saladoriver), 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 etal.2000) based on
Ventania have not only a characteristic distributional patterns exhibited byAsteraceae.
petrophylous flora, but they are the only
regions in the province with endemic plant MATERIAL AND METHODS
taxa(Parodi 1947).There havebeennumerous
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 inthe area. The different Facelis retusa, Gamochaeta subfalcatai, the
communities were explained bythe variation Asteraceae of this region shows no links
inthe local climate and soil, which iscaused 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, thegrass family) On the basis of several over1apping taxa
innumber of genera and species represented distribution and phytogeographical and
inthearea(Parodi 1947).Approximately 30% geological criteria theareasdelimited here are
(112 species and infraspecific taxa) outofthe (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 and37°southlatitude(Cabrera &Willink 1973,
Ventania andTandilia. Furthermore, therehave Morrone etal. 1997).
been found until the present four species of (2) Central and North America (CNA):
Asteraceae endemic toTandilia, three species Southern North America andCentral 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 isbased on (4) Mahuidas (MA): group of mountain
several lines of evidence. Systematic studies ranges inthe political province ofLa Pampa,
have documented much of the taxonomic incentralArgentina. They are300-500 mhigh,
diversity and distribution patterns of ferns andthey aredated 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. Paramoand Punaprovinces, andcomprises the
highlands over3000maltitudefromVenezuela
It would be interesting to seek the historical tonorthwestern Argentina (Morrone 1994).
explanations that ledto 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 ofendemism considered inthe historical biogeographic analysis: CHA =Chaco;
CHI=Central Chile;MA=Mahuidas; NA =Northern Andes; CNA=Central andNorth 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 (PAT): semidesert east of the Taxaanalyzed
southern Andes to the Atlantic coast in Ar-
gentina, approximately between 35° and 55° The unitsofstudy (Table 2)were species and
south latitude (Cabrera &Willink 1973). infraspecific taxa of Asteraceae inhabiting
(8)Sierras Pampeanas (SPA):group ofmoun- Tandilia and Ventania. The distributional data
tain ranges incentral Argentina, extending in were taken from the specimens' labels ofthe
the political provinces of Córdoba, Santiago herbarium ofDepartamento Científico dePlan-
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 belong to the Precambric revisionary studies were taken also into
age(Mingramm etal. 1980)andwere uplifted consideration. Fornomenclature purposes we
inthe 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 Paranense province recutita, Sonchus asper), cultivated (e.g.,
ofCabrera &Willink(1973) insouthern 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 ehilensis,
eastern Paraguay. Taraxacum officinale¡ species were nottaken
(11)Tandilia (TA): or"Sierras Septentriona- into consi-deration.
les", agroup ofmountain ranges east ofBue-
nosAires province. They are500 mhigh, 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): prairiewithendemic 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 Thepanbiogeographic approach (Croizat 1958,
&PérezMoreau 1944,Chebataroff 1960),situa- 1981) consists of plotting distributions of
ted inthe south of Brazil (southern RioGran- organisms onmaps, connecting their separate
de do Sul) and Uruguay. distributional areas together with lines called
(13) Venlania (VE): or "Sierras Australes" a individual tracks. As anexample, some ofthe
group of moutain ranges southwest of Bue- 112individual tracks produced inthisstudy of
nosAiresprovince. They are900-1000 mhigh, Asteraceae inhabitingTandilia and/orVentania
and thegeological age corresponds to490 m. andotherareasareillustrated (Figs.2-3). When
y. ago, inthe Paleozoic (Llambías & Prozzi individual tracks coincide, the resulting
1972,Harrington 1980). summary lines are considered gene-ralized
tracks, which indicate thepre-existence ofan-
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 Cypcttoherbertlí (Lindl.) Herb X X Ravenna 1968
Bacchans rufesccns Sprcng val' X Cabrera 1963 subsp. l1'O/jjhucj.!,eli(1Iauman)
vcntctnículaCabrera Ravenna
Bacchans tondttens¡s Speg. X Cabrera 1963 Sisyrinchinrnjunccnm i:.i\1ey X X Ravenna 1968
lIícrctcínmburlcartiíSleumer X X Cabrera 1963 subsp. lainezií {Hicken )Ravenna
Hícrctcinmchacoense (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 Bromusbonarícnsis Parodi &J. X Cámara H
Stcvictsatureiafotía (Lam.) Lam X Cabrera 1963 A.Cámara 1970
vnr. \'CI1ICII/{.:nsisCabrera Fcstncaventanico/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'oairtdfolia Ilauman X X Torres 1970b
Setar¡avaginata Spreng val' X Nicora 1970
BROMELlACEAE tcmdtlcnsisNicora
X Cabrera 1968 Slipa vcntunícolu Cabrera & X Cabrera &
ñyck¡a nstnotiiloraOtto & 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
Gyrnnocctivcíumplatense (Speg.)
Morronc 1999 BOTIIRIURIDAE
Britton & Rose var, vcntantcotu
Bnthríurns vovan Mauri X Maury 1973
(Speg.) R.Kiesling
IGUANIDAE
FAIl¡\CI:AE Líolaennrsgracitis 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
Lcühvrushookcri 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/¡loJm7sunoaí Orejas X X Vega 8:
Vicia muntevidensis Vogel X Burkart 1967 Miranda Bcllagamba
oblonga Burkart 1990
Vscütotía 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). Itbasically consists
concept of distributional compatibility (Co- of constructing a matrix (areas vs. tracks),
nnor 1988,Craw 1989). Inthis 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 ifone track isasubset 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 inadistributional compatibility treated ourtracks asbinary characters ordered
rnatrix and isidentified asageneralized track. with absence asthe "ancestral" state for each
If more than one largest clique or several (=outgroup with allzeros).
cliques ofconsiderable size are found, then a
hypothesis ofexistence ofseveral generalized Parsimony analysis ofendernicity (PAE)
tracks linking the localities or distribution
areasinmore than oneway canbeconsidered. 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 eliques can be identified as a to identify the distributional pattern of
generalizedtrack(Craw 1990).Formoredetails organisms. Itclassifies localities, quadrats or
andotherapplications ofthis method seeCraw areas (which are analogous to taxa) by their
(1988,1989),Morrone&Crisci(1990,1995),Craw sharedtaxa(whichareanalogous tocharacters)
etal.(1999)andCriscietal.(2000).Theanalysis according to the most parsirnonious solution
ofthedata matrix of 13areas ofendemisrn vs. (Morrone & Crisci 1995, Crisci et al. 2000).
distributional data of I12taxa (Table 3) was The rnethodwasoriginally proposed byRosen
carriedoutwithSECANT 2.2(Salisbury 1999). (1988) using localities as study units.
SECANTisaprogramforidentifying allgroups
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.SpeciesofAsteraceae fromTandiliaandVentania andotherareasofendemismconsidered
inthe study. Taxa with the same areas ofendemism have been indented. Areas ofendemism for
each taxa are indicated inTable 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 IIook. & 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. lIyalis 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.) lIieron.
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 IIook. &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 llvpochaeris 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.DonexIlook. &Arn. varocrcctus 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, varosatureiifolia
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 fromthefívecliques,asfollows(Fig.4):
variation of the method using areas of Clique A:««««(T A, UR), SB), PAM), VE),
endem ism as study units to identify the SPA),CHA), SA),NA), PAT,CHI), supported
hierarchical information contained in the bythe 30individual 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 area 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 iscoded as 1and itsabsence as O. A SPA),SA),CHA),NA), CHI, PAT),supported
parsimony algorithm isapplied tothe matrix bythe30individual 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, inwhich ter- 87,88,103,104,105,106,107,108,109,110
minal dichotomies represent two areas (Fig.4B).
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 areas and bythe30individual 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 bythe30individual 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*. Asproposed by Rosen (1988), 88,98,103,104,105,106,107,108,109,110
the cladogram was rooted with ahypothetical (Fig.4 O).
area coded all zeros. Clique E:««««(TA, UR), SB), PAM), VE),
SPA), SA, CHA) NA, PAT, CHl), supported
RESULTS bythe27individual tracks 3,20,21,23,25,26,
27,28,29,30,36,37,38,81,82,83,84,87,88,
Panbiogeography (compatibility track method) 103,104,105,106,107, 108,109,110(Fig.4E).
Based on the data matrix (Table 3), the Mahuidas and Central andNorth America are
combination ofthe individual tracks resulted the only areas that appear in none of these
infour largest cliques (generalized tracks) of pattern ofrelationships inthe 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 withthe 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, itwas selected to
103,104,105,106,107,108,109,110 inTable2) show the taxa supporting the areas
common tothefour largestcliques isidentifíed relationship (Fig.4E):
asthe fifth clique orgeneralized track (Fig. 4 (1) (TA,UR). Based on Senecio grisebachii
E).Inthisway, sixindividual tracks (5,40,41, varosubincanus (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
toformthefour cliques of30 individual tracks squarrulosum (track 37), Baccharis gnapha-
mentioned above. lioides (track 38), and Micropsis spathulata
Fivetreesconnectingtheareascanbeconstructed (track39).
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